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TULANE STUDIES IN
ZOOLOGY AND BOTANY ,

Volume 29, Number 1 July 15, 1993

»

ETHEOSTOMA COLOROSUM AND E. BELLATOR,

TWO NEW DARTERS, SUBGENUS ULOCENTRA,

FROM SOUTHEASTERN UNITED STATES

ROYAL D. SUTTKUS

Tulane University Museum of Natural History
Belle Chasse, La 70037

AND

REEVE M. BAILEY

Museum of Zoology, The University of Michigan
Ann Arbor, Mi 48109 1079

TYPE SPECIMENS OF FISHES IN THE

TULANE UNIVERSITY MUSEUM OF

NATURAL HISTORY

HENRY L. BART, JR.

AND

MICHAEL S. TAYLOR

Tulane University Museum of Natural History

Department of Ecology, Evolution, and Organismal Biology

Tulane University, New Orleans, La 70118

TYPE SPECIMENS OF VASCULAR PLANTS AT

TULANE UNIVERSITY, WITH A BRIEF HISTORY

OF THE TULANE HERBARIUM

ANNE S. BRADBURN
AND

STEVEN P. DARWIN

Department of Ecology, Evolution, and Organismal Biology
Tulane University, New Orleans, LA 70118

Tulane
University

New Orleans

TULANE STUDIES IN ZOOLOGY AND BOTANY

ISSN 0082-6782

Department of Ecology, Evolution, and Organismal Biology
Tulane University, New Orleans, Louisiana, 70118

Steven P. Darwin, Editor

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ETHEOSTOMA COLOROSUM AND E. BELLATOR,

TWO NEW DARTERS, SUBGENUS ULOCENTRA,

FROM SOUTHEASTERN UNITED STATES

Royal D. Suttkus

Museum of Natural History, Tulane University
Belle Chasse, LA 70037

Reeve M. Bailey

Museum of Zoology, The University of Michigan
Ann Arbor, MI 48109-1079

Abstract

Two new darters, subgenus Ulocentra, are described from Alabama and west-
ern Florida. Both are members of the Etheostoma duryi species group with the
premaxilla free from the upper lip, and the vomer usually or consistently with
one or a few teeth. Both usually have five branchiostegal rays and lack an orange-
red band in the anal fin. The coastal darter, E. colorosum, is the only species of
Ulocentra inhabiting the coastal streams of southern Alabama and northwestern
Florida between the Perdido and Choctawhatchee rivers. It lacks a red ocellus
anteriorly on the first dorsal fin, with red or orange color restricted to the last
three to five membranes, and has other characters suggesting close affinity with
E. tallapoosae and other as yet unnamed Ulocentra species from the Gulf Coastal
Plain. Nuptial males off. colorosum have a diagnostic series of small red or orange
punctulations along the side and additionally have other associated pigment
features. It is a slightly smaller species than E. tallapoosae and the scales are
larger. The Warrior darter, E. bellator, is discontinuously distributed in the Black
Warrior River system above the Fall Line in Alabama. Nuptial males have a red
ocellus in the first membrane of the spinous dorsal fin. There are two series of
elongate blotches on the midside separated by a pale area, and the bright orange
stripe on the lower side and caudal peduncle is narrow. Two other species of
Ulocentra also live in the upper Black Warrior system, E. zonistium with a charac-
teristically banded first dorsal fin and E. chermocki, confined to the Turkey Creek
drainage, the latter species most closely related to E. bellator but distinguished
by different lateral pigmentation and a broad ventrolateral stripe in nuptial
males.

Introduction

Members of Etheostoma subgenus Ulocentra occur in Gulf Coastal drainages
east of the Mississippi River, several eastern tributaries to the lower Mississippi
River, the Tennessee and Cumberland river systems, and southern tributaries
to the Ohio River. The presence of darters of this subgenus south of the Tennes-
see River system (Mississippi River drainage) has been known for many years.
Etheostoma coosae was the first species to be described from the southern zone of
the subgenus (Fowler, 1945). Four decades later, Bailey and Etnier (1988) de-
scribed E. zonistium, which is distributed primarily in western, lower tributaries
to the Tennessee River but also occurs in a few headwater branches of the Sipsey

2 Tulane Studies in Zoology and Botany [Vol. 29

Fork of the Black Warrior River system, Alabama. These headwater tributaries
are just across the divide from the upper part of Bear Creek, a tributary to the
Tennessee River. Suttkus and Etnier (1991) described two more species {E.
tallapoosae and E. brevirostrum) of the subgenus. In early 1992 one of us (R. D.
S.) and H. T. Boschung, independently discovered and studied an isolated form
of subgenus Ulocentra in the Turkey Creek system, a tributary to the Locust Fork
of Black Warrior River system, Jefferson Co., Alabama. It was described as E.
chermocki by Boschung et al. later in 1992. Some of the data compiled by R. D.
Suttkus are incorporated in this paper under their name, Etheostoma chermocki.
The darters described herein were first recognized many years ago and have
become common knowledge. The details of their geographic distributions and
their relationships to other close geographical forms, until recently, have re-
mained poorly understood. The subgenus Ulocentra, as here interpreted, follows
Bouchard (1977) and Bailey and Etnier (1988).

Materials and Methods

Specimens of the new species and comparative material used are primarily
from the collections of Academy of Natural Sciences of Philadelphia (ANSP);
Auburn University (AUM); Cornell University (CU); Tulane University Museum
of Natural History (TU); University of Alabama (UAIC); University of Florida
including former Florida State University specimens (UF); University of Michi-
gan Museum of Zoology (UMMZ); National Museum of Natural History (USNM);
and University of Tennessee (UT).

Counts and measurements were made as described in Hubbs and Lagler
(1958) except as follows. Transverse body scales were counted from the origin
of the anal fm diagonally upward to the base of the spinous dorsal fm. Gill
rakers, counted on the anterior arch of either the left or the right side, include
both dorsal and ventral rudiments. Measurements were made with needle-point
dial calipers and recorded to the nearest 0.1 mm. Trans-pelvic width was meas-
ured between the outer bases of the pelvic spines. Our measurements of the
length of the spinous dorsal, soft dorsal and anal fins are the total lengths of
these fins when depressed and not the length of the fin base. Names used for
associated fish species follow Robins et al. (1991).

In the listing of type material, each catalog number is followed by the number
of specimens examined and the range of standard length (SL) in millimeters,
e.g., (10, 27—52). In addition to standard compass directions (with the following
"of deleted), the following abbreviations are used: Cr. = Creek, R. = River,
mi = mile(s), airmi = airmile(s), trib. = tributary, Fk. = Fork, hwy = highway,
rd = road, Co. — County, T = Township, R = Range, Sec = Section. In lists
of materials not designated as types, the catalog number is followed by the
number of specimens examined, enclosed in parentheses.

Etheostoma colorosum, new species

Coastal Darter

Figures 1, 8, and 12

Etheostoma simoterum. Gilbert, 1891: 159 (misidentification, Sandy Cr. record). Stevenson, 1976: 37
(in key, characters, orange-sided darter). Mettee, O'Neil, and Harris, 1983: 24, 88 (in part;
Hall Cr., Little Escambia Cr., Pollard Oil Field, AL).

Etheostoma sp. Bailey, Winn, and Smith, 1954: 143 (Orangestripe darter, Escambia R. records).

1993] New Species oi^ E the ostoma 3

Etheostoma (Ulocentra) sp. Smith-Vaniz, 1968: 136 (in part, reports from ("onecuh and (^hoctawhatrhee
River systems). Jenkins, 1976 (in part, undescribed species, distribution). Kuehneand Barbour,
1983: 97-98, pi. 12 (in part, "(iuifcoast Snubnose Darter," characterized, range map in part).
Gilbert and Walsh, 1991 : 16 ("Gulfcoast snubnose darter," probable source of color illustration
in Kuehne and Barbour).

Coastal Plain Darter, Etheostoma sp. Page and Burr, 1991: 30? (in part, pi. 43 [not coastal darter],
map 346, in part).

Holotype: Adult male, TU 162508, 50.1 mm standard length (SL), Pine
Barren Creek, tributary to Escambia River at Still Road (T5N, R32W, Sec 20),
Escambia County, Florida, 29 March 1992, R. D. Suttkus.

Paratopotypes: tu 162509 (47, 34—54), collected with holotype are distrib-
uted as follows: TU 162509 (42), ummz 221365 (5); tu 152006 (36, 33-50), 22
April 1988, distributed as follows: tu 152006 (32), ansp 169684 (2), cu 73644
(2); TU 154470 (27, 36-54), 21 March 1989, distributed as follows: tu 154470
(19), UAic 10506.01 (4), UT 91.4202 (4); tu 157537 (24, 31-45), 3 March 1990,
distributed as follows: tu 157537 (18), up 92633 (4), usnm 324501 (2).

Other Paratypes: tu 151947 (28, 34—51), Pine Barren Creek at Fla Hwy
99 (T5N, R32W, Sec 28), Escambia Co., Florida, 21 April 1988, distributed as
follows: TU 151947 (24), ummz 221366 (4); and aum 26759 (1, 47), Pine Barren
Creek at Escambia Co. Rd 4, 2.3 mi W Bratt, Escambia Co., Florida.

Additional material examined but not designated as types: Perdido River drainage,
Florida. Escambia County: aum 26016 (8), Brushy Cr. at Escambia Co. Rd 99, 2 mi W Walnut
Hill (T4N, R33W, Sec 4), aum 26742 (6), and TU 162538 (9).

Escambia River drainage, Florida. Escambia County: uf 54776 (2), Prittchett Mill Branch,
3.9 mi S Century at US Hwy 29 at Bluff Springs (T5N, R31W, Sec 24); uf 54817 (1), trib. to Canoe
Cr. at Fla Hwy 4, 5.5 mi W junction with US Hwy 29 (T5N, R31W, Sec 8); uf 54835 (4), Canoe Cr.
at Fla Hwy 4, 6.1 mi Wjunction with US Hwy 29 (T5N, R31W, Sec 8); uf 55915 (2), Mitchell Cr.,
trib. to Escambia R., 0.4 mi N McDavid at US Hwy 29 (T4N, R31W, Sec 10). Alabama. Escambia
County: ummz 163552 (3), Big Escambia Cr. at Flomaton (TIN, R8E, Sec 33), ru 16518 (1); aum
5506 (4), Cedar Cr., 3.9 airmi NE East Brewton (T2N, RIOE, Sec 13); aum 1076 (2), Escambia R.
NE Atmore at Interstate Hwy 65 (T2N, R7E, Sec 4); aum 26780 (4), Burnt Corn Cr. at Ala Hwy
41 N Brewton (T2N, RIOE, Sec 20). Conecuh County: usnm 43494 (3), Sand Cr., 2.8 mi E
Evergreen (T6N, RUE, Sec 36); ummz 165987 (8), Murder Cr. at Castleberry (T4N, RIOE, Sec 24);
UMMZ 155519 (6), Bear Cr., trib. to Murder Cr. at US Hwy 31, 2 mi S Castleberry (T4N, RIOE, Sec
35); TU 159841 (72), Jordan Cr. at US Hwy 31, 4.9 mi S Fairview (T5N, RIOE, Sec 35), and ru
162528 (34); ru 159829 (17), Spring Cr., 2.5 mi NW Centernary Methodist Church (T7N, RIOE,
Sec 9). Butler County: ummz 88738-39 (3), trib. to Persimmon Cr., 1 mi W Greenville at Ala
Hwy 10 (TION, R14E, Sec 22); ummz 128770 (1), Hawkins Cr., trib. to Persimmon Cr., 2.9 mi S
Greenville (T9N, R14E, Sec 3); usnm 43527 (4), Greenville, AL (TION, R14E); ummz 139155 (20),
Rocky Cr., trib. to Persimmon Cr., 1 mi N Georgiana and 2.75 mi S Chapman (T8N, R13E, Sec
13). Crenshaw County: tu 2581 (4), trib. to Patsaliga Cr., 20 mi W Luvergne at Ala Hwy 10
(T9N, R18E, Sec 30). Pike County: tu 2605 (1), trib. to Indian Cr., trib. to Conecuh R., 8.4 mi
W Troy at Ala Hwy 10 (TION, R19E, Sec 25). Covinoton Couniy: aum 21082 (3), trib. to Blue
Cr., trib. to Conecuh R., 8.6 airmi NW Wing (T3N, R14E, Sec 31).

Blackwater River drainage, Florida. Santa Rosa County: uf 54713 (22), Big Juniper Cr.,
2.0 mi W Munson at Fla Hwy 4 (T4N, R27W, Sec 12), tu 31731 (13), i u 33842 (2), ru 56741 (72).
and TU 106620 (12); UF 54781 (13), East Fork Coldwater Cr., 5.7 mi WNW Munson at Fla Hwy 4
(T4N, R27W, Sec 5), and TU 20940 (10); ummz 212001 (15), W Branch Juniper Cr., 2.1 mi S
Munson, a campground off Hwy 191 (T4N, R27W, Sec 24, SW 1/4); UF .54978 (14), Sweetwater
Cr., 1.3 mi E Munson at Fla Hwy 4 (T4N, R26W, Sec 16), ummz 163530 and 169783 (16); ru 57082
(116), Bear Cr., trib. to Sweetwater Cr., 1.3 mi E Munson at Fla Hwy 4 (RN, R26W, Sec 16), ru
151585 (22), TU 154458 (67), tu 157516 (22), ru 157816 (2), and ru 162518 (10); uf 54962 (4),
Big Juniper Cr., 5 airmi S Munson (T3N, R27W, Sec 12); uf 58237 (1 ), Alligator Cr., trib. to Juniper
Cr., 6.0 airmi N Harold (T3N, R26W, Sec 30); tu 151591 (4), Big Juniper Cr., 1.0 mi E McLelian
(T6N, R26W, Sec 31); tu 151595 (9), Turkey Cr., trib. to Big Juniper Cr., 3.2 mi NW Munson

Tulane Studies in Zoology and Botany [Vol. 29

(T4N, R27W, Sec 2); UF 58223 (2), Pond Cr., 2 mi WSW Milton at US Hwy 90 (TIN, R28W, Sec
9); UMMZ 165103 (6), Pond Cr., 3.9 mi W Milton along Fla Hwy 10 (TIN, R29W, Sec 1); UF 58196
(1), Pond Cr., 9.8 airmi NW Milton (T3N, R29W, Sec 33); TU 88877 (4), Pond Cr., 7.0 mi NW
Milton at Fla Hwy 191 (T2N, R29W, Sec 15), TU 92033 (7), TU 94767 (11), TU 98571 (10), TU
100739 (4), TU 103115 (11), tu 105584 (16), tu 111392 (2), tu 116023 (5), tu 117581 (4), tu
120137 (3), and tu 124295 (2); UF 58351 (2), Blackwater R., 7.0 airmi NE Milton, 0.5 mi NE mouth
Coldwater Cr. (T2N, R27W, Sec 16); tu 20524 (7), West Fork Big Coldwater Cr., 3.9 mi E Jay at
Fla Hwy 4 (T5N, R29W, Sec 14); tu 27399 (9), Manning Cr., trib. to West Fork Big Coldwater Cr.,
14 mi N Milton at Fla Hwy 87 (T3N, R28W, Sec 4); tu 152863 (7), West Fork Big Coldwater Cr.
at Fla Hwy 87, 1 1.8 mi N Milton (T3N, R28W, Sec 10). Okaloosa County: uf 54982 (1), Beaver
Cr., trib. to Blackwater R., 5.4 mi E Munson at Fla Hwy 4 (T4N, R25W, Sec 18); UF 54838 (3),
Blackwater R. at Fla Hwy 4, 4.5 mi NW Baker (T4N, R25W, Sec 22), and TU 23682 (6); UF 56182
(2), Narrows Cr., trib. to Panther Cr., 1.7 mi W Cannontown, about 10 airmi NW Crestview (T4N,
R25W, Sec 12). Alabama. Escambia County: aum 25995 (1), Blackwater R. at Escambia Co. Rd
4, 1 mi E Bradley (TIN, R13E, Sec 25), and aum 26728 (2).

Yellow River drainage, Florida. Okaloosa County: tu 3719 (17), Big Horse Cr., trib. to
Yellow R., 2.7 mi E Blackman (T5N, R24W, Sec 24); TU 3151 (3), Horsehead Cr., trib. to Yellow
R., 2.9 mi E Fla Hwy 85 at Fla Hwy 2 (T5N, R22W, Sec 15); uf 54992 (7), Silver Cr., 5.0 airmi SE
Crestview (T2N, R23W, Sec 2), and uf 54738 (11); uf 54754 (4), Turkey Gobbler Cr., 8.2 airmi
SW Crestview (T2N, R24W, Sec 17); uf 56167 (1), Gum Springs Cr., 3.5 mi N Milligan ( T4N,
R24W, Sec 34); UF 55832 (1), Silver Cr. public lake drainage, 7 mi N Crestview (T4N, R24W, Sec
24); UF 57156 (5), Turkey Hen Cr., 6.5 mi S Crestview (T2N. R23W, Sec 8), and UF 54733 (3); tu
24085 (21), Laird Mill Cr., 100 yds. before entering Shoal R. (T3N, R22W, Sec 1); tu 24695 (4),
trib. to Shoal R. from westside, 0.2 mi below railroad trestle (T3N, R23W, Sec 23); TU 24076 (3),
Shoal R. cutoff, 6 mi upriver from Fla Hwy 85 bridge (T3N, R23W, Sec 35); tu 24653 (4), Yellow
R. at mouth of nameless creek (T4N, R24W, Sec 35); tu 24647 (3), Horse Cr. at entrance to Yellow
R. (T5N, R23W, Sec 29); TU 72819 (5), Yellow R., 4.6 mi W Fla Hwy 85 at Fla Hwy 2 (T5N, R23W,
Sec 20), TU 82630 (6), tu 79807 (5), and tu 79701 (2); tu 80486 (4), Yellow R., 0.2 mi downriver
from Fla Hwy 2 (T5N, R23W, Sec 20); tu 83099 (1), Yellow R., 3.5 mi SW Crestview at US Hwy
90 (T3N, R24W, Sec 15), tu 101923 (2), and tu 102315 (2); tu 81258 (4), Yellow R. at end of Fla
Hwy 189, 2.9 mi S Holt (T2N, R25W, Sec 16); tu 80536 (1), Mill Cr., trib. to Yellow R., 1.1 mi N
Milligan (T3N, R24W, Sec 15); tu 81660 (1), Horsehead Cr., trib. to Pond Cr., 3.2 mi SE Laurel
Hill (T5N, R22W, Sec 15); aum 14934 (3), Yellow R. between Oak Grove and Milligan (T4N, R24W,
Sec 24); TU 102333 (3), Pond Cr., trib. to Shoal R., 1.0 mi NE Dorcos, Okaloosa Co. Rd 373 (T4N,
R22W, Sec 26). Walton County: tu 1749 (24), Pine Log Cr., 9.1 mi E Fla Hwy 85 at Fla Hwy
2 (T5N, R21W, Sec 28), and TU 82525 (2); uf 54362 (9), Pine Log Cr., 6.4 mi S Clear Springs (T4N,
R21W, Sec 5); uf 54206 (17), Shoal R. at Fla Hwy 285, 4.3 mi S Stella (T4N, R21W, Sec 36), tu
111466 (4), UF 56066 (26), tu 46112 (8), uf 57297 (7), tu 113873 (4), and tu 129426 (4); tu
46398 (1), Turkey Cr., trib. to Shoal R., 1 mi W junction Fla Hwy 2A an US Hwy 331 (T4N, R20W,
Sec 16), TU 46454 (3), and tu 67413 (2); uf 54134 (3), Long Cr., 10.3 mi S Clear Springs (T4N,
R21W, Sec 21); UF 56225 (3), Big Swamp Cr. at US Hwy 331 at Liberty, 9.5 airmi NW De Funiak
Springs (T4N, R20W, Sec 26); tu 24101 (2), Bottle Branch, trib. to Shoal R. (T3N, R21W, Sec 4);
tu 24766 (4), Wolf Cr., trib. to Shoal R. (T3N, R21W, Sec 6). Alabama. Covington County:
TU 72895 (17), Yellow R. at Covington Co. Rd 4, 12 mi W Florala (TIN, R15E, Sec 34); tu 72974
(38), Yellow R., 9.4 mi NNW Florala, Covington Co. Rd 55 (T2N, R16E, Sec 34); tu 73017 (25),
Five Runs Cr., trib. to Yellow R., 15 mi NW Florala (T2N, R15E, Sec 26); TU 73180 (5), Yellow R.,
7 mi N Babbie (T5N, R17E, Sec 27); TU 73162 (12), Yellow R., 10 mi SW Opp (T3N, R17E, Sec
21); TU 82612 (3), Yellow R., 4.6 mi SE Wing at Covington Co. Rd 4 (TIN, R15E, Sec 34).

Choctawhatchee River drainage,. Florida. Walton County: uf 54149 (2), Eight Mile Cr., 2.7
mi WNW Gaskin (T6N, R19W, Sec 31); uf 54228 (3), Natural Bridge Cr. at Natural Bridge, 4.5
mi WNW Gaskin (T6N, R20W, Sec 26). Holmes County: tu 182 (2), Parrot Cr., trib. to Choc-
tawhatchee R., 4.7 mi SSW Geneva, AL (T6N, R17W, Sec 25); tu 1587 (20), Mill Cr., trib. to
Choctawhatchee R. at southern limits of Ponce de Leon (T3N, R17W, Sec 28); TU 2285 (4), Wrights
Cr., 3.5 mi NW junction Fla Hwy 79 and Fla Hwy 177 at Hwy 177 (T4N, R15W, Sec 1), tu 45652
(1), and tu 46130 (3); tu 46268 (1), Wrights Cr., trib. to Choctawhatchee R., 6 mi S junction Fla
Hwy 2 and Fla Hwy 79 at Hwy 79 (T5N, R14W, Sec 32), TU 46434 (6), and TU 46663 (1); UF 54128
(2), Wrights Cr., 6.1 mi W Graceville railroad station at Fla Hwy 2 (T7N, R14W, Sec 35), ummz
166309 (1); tu 2475 (3), Little Cr., trib. to Ten Mile Cr., 6.5 mi NNW junction Fla Hwy 79 and Fla
Hwy 177 (T5N, R15W, Sec 27); uf 51645 (1), Ten Mile Cr., 7.0 mi NNE Bonifay at Fla Hwy 177
(T6N, R15W, Sec 35); uf 54409 (2), East Pittman Cr., 0.8 mi N Pittman (T5N, R16W, Sec 11).
Holmes-Washington County: uf 51250 (3), Holmes Cr., 4.6 mi W Chipley at US Hwy 90
(T4N, R14W, Sec 2). Washington County: uf 57945 (2), trib. to Holmes Cr., 6 airmi SW Chipley

1993] New Species of £//i^05toma

at Washington Co. Hwy 280 (T4N, R14W, Sec 22). Alabama. Geneva County: ru 2418 (2), Spring
Cr., trib. to Choctawhatchee R., 1.8 mi N Black (11 N, R23F„ Sec 27), and rv 16374 (10); ru 2516
(5), Panther Cr., trib. to Little Choctawhatchee R., 3 mi W Pinckard Farm Rd and US Hwy 84 ( r3N,
R25E, Sec 7). Dale County: aum 27121 (2), Claybank Cr. at Ala Hwy 27 (T5N, R25E, Sec 2).

Diagnosis: Etheostoma colorosiim is a member of subgenus Ulocentra as diag-
nosed by Bouchard (1977) and Bailey and Etnier (1988). Further, E. colorosiim
lacks a premaxillary frenum and in this respect is typical of members of the E.
duryi species group of Bailey and Etnier (1988). The vomer is feebly toothed or
may lack teeth. Etheostoma colorosum lacks a red ocellus in the first membrane of
the spinous dorsal fin, as do a few other Ulocentra species from the Gulf Coastal
Plain. The unique diagnostic feature of E. colorosum is a series of small orange
to red spots or punctulations along the side of the nuptial male. This series of
punctulations extends from base of caudal anteriorly to above pectoral fin base
and is immediately dorsal to the mid-lateral brown blotches that are usually
centered on the lateral line. Most nuptial males have a few additional orange or
red punctulations along the lateral line between the lateral blotches. The lateral
blotches are of moderate size and are ovoid to quadrate in shape. The red
coloration in the spinous dorsal fin is restricted to the posterior three to five
interradial membranes and thus does not form a complete band across the fin.
An incomplete band of black blotches extends from the first to the sixth or the
seventh or eighth interradial membrane. The posterior black blotches occur on
the fin proximally to the anteriormost red blotches. Both the spinous and the
soft dorsal fins are noticeably elevated.

Description: Etheostoma colorosum reaches a maximum of 48 mm SL
(females) to 58 mm SL (males). Sexual dimorphism and apparent sexual maturity
occur at the end of the first year at a minimum of 28 mm SL (females) and 30
mm SL (males).

Frequency distributions of scale and fin ray counts are presented in Tables
1 and 2. Number of vertebrae, branchiostegal rays, gill rakers, and preoper-
culomandibular pores are given in Table 3. The lateral line is usually complete
with (39-) 41-47 (-50) scales, occasionally the last scale is unpored. Transverse
scales 10(1 specimen), 11(48), 12(158), 13(158), 14(75), 15(18), or 16(4). Trans-
verse scales from origin of second dorsal fin to anal fin (an alternative count,
see Bailey and Etnier 1988: 30) 9(1), 10(1 1), 1 1(65), 12(30), 13(5); mean of 1 12
counts, 1 1.2. Caudal peduncle scale rows number 12(2), 13(1 1), 14(79), 15(274),
16(147), 17(50), or 18(6) in 569 specimens. Dorsal fin has (9-) 10-1 1 (-12) spines
and (10-) 1 1-12 (-13) soft rays. Anal fin has 2 spines and (6-) 7-8 (-9), modally
7, soft rays. Pectoral fin has (13-) 14-15 (- 16) rays. Branchiostegal rays number
4_5(2), 5—5 (534), 5—6 (12), 6—5 (5), or 6—6 (6) in 559 specimens; vertebrae
number 37(5), 38(104), 39(104), or 40(6), and gill rakers on first arch number
6(9), 7(53), 8(36), or 9(2). Number of vertebrae (Bailey and Gosline, 1955) and
gill rakers are quite consistent within species groups. The sum of left and right
preoperculomandibular pores is nearly constant at 18 (in 180 of 190 specimens),
the remaining ten have 16, 17, or 19 pores. The opercle, cheek, nape, and
prepectoral area typically are covered with exposed scales. The scales on the
cheek usually are smaller than those on the opercle, and frequently there is a
mixture of exposed and embedded scales. The belly varies from being completely
covered with exposed scales to being naked on the anterior half. Females more
frequently than males have the anterior part of the belly naked and in some of

Tulane Studies in Zoology and Botany

[Vol. 29

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these individuals the scales diminish in size anteriorly and may be partially em-
bedded. The breast is consistently scaleless.

In 10 specimens dissected, the vomer is toothless in four, has a single weak
tooth in four, and has two small teeth in two; the palatines are uniformly toothless.

Proportional measurements appear in Tables 4 and 5. Twenty of the 25
males were nuptial specimens taken between the 2nd and the 21st of March.
The remainder of the specimens came from a November collection. The two
lots of five each of females were taken from a March 2 1 and an April 8 collection.

Colors of live and freshly preserved nuptial specimens, collected in March
and April, are described from Pine Barren Creek of the Escambia R. drainage
(Figure 1) and Bear, Sweetwater, and Big Juniper creeks of the Blackwater R.
drainage. Nuptial males have an orange stripe on the lower side. This orange
stripe usually narrows posteriorly and is partially obliterated by an overlay of
turquoise color immediately ventral to the last three lateral blotches. The tur-
quoise extends from the ventral edge of a lateral blotch on one side of the body
to the ventral edge of the corresponding blotch on the opposite side. The lateral
body blotches are dark brown, oval to rounded quadrate in shape and number
seven to nine. These blotches are centered along the lateral line or quite fre-
quently extend farther below the lateral line than above it. One of the diagnostic
color features of Etheostoma colorosum is the "string" of orange to red-orange
small spots that extends along the side above the brown lateral blotches. This
series of spots or punctulations extends from the base of the caudal fin anteriorly
to the shoulder region just above the base of the pectoral fin. The spots are
arranged in linear groups of two to four (or five) closely spaced spots, and the
groups are separated by somewhat wider interspaces. Most nuptial males have
additional orange spots along the lateral line between the brown lateral blotches.
The number of spots between the blotches is usually one or two, or occasionally
three. There are some irregularly shaped brown blotches (more linear than the
lateral blotches) on the dorsal-lateral area of the body. The upper edges of some
of these are connected with the median dorsal blotches (typically eight in number).
The blotch on the nape, and the dorsal blotches 7 and 8 on the caudal peduncle,
are overlaid with turquoise color. The top of the head is dark brown. The ventral
surface of the head from the breast to the anterior tip is pale with an overlay
of turquoise color. The entire breast, gill membranes, lower part of opercle,
cheek, and side of snout are turquoise in color. There is a dark brown preorbital
bar, a dark brown suborbital bar, and several brown blotches along upper margin
of the opercle. The iris is dark with a golden edging around pupil.

The pectoral fins are the only ones that lack turquoise in the nuptial male.
The pelvic fins and all the median fins have some turquoise coloration. The
pectoral fins appear to be clear to the unaided eye, however, magnification
reveals considerable dark pigmentation along the rays and that only the interra-
dial membranes are clear. Nearly the entire anal and pelvic fins are turquoise.
The tips of the spines and soft rays of anal and pelvic fins are milky white. After
the blue color has faded, both the fin rays and the interradial membranes are
found to be densely pigmented with melanophores. The margins of the caudal,
first and second dorsal fins are turquoise. Some males at the peak of nuptial
coloration have more than half the caudal fin turquoise. There are two orange
to red-orange blotches at the base of the caudal fin and, immediately posterior,
a small central pale area. The rest of the caudal fin is turquoise with the more
intense coloration along lower and upper procurrent margins. The marginal
and submarginal areas of the spinous dorsal fin are turquoise. Proximally from

10

Tulane Studies in Zoology and Botany

[Vol. 29

Figures 1 — 3. Etheostoma species from Alabama and Florida. I.E. colorosum, TU 162508, holotype,
adult male, 50 mm SL. Pine Barren Cr., trib. to Escambia R., Escambia Co., FL (photo by R. D.
Suttkus). 2. E. bellator, TU 151837, paratype, adult male, 43 mm SL. Murphy Cr. along US Hwy 31,
2.0 mi SW Blount Springs, Blount Co., AL (photo by J. M. Pierson). 3. E. bellator, TU 163053,
paratype, adult male, 48 mm SL. Murphy and Mill Cr. confluence, 1 mi SW Blount Springs, Blount
Co., AL (photo by R. D. Suttkus).

Figures 4 — 7. Etheostoma species from Alabama. 4. E. bellator, TU 163053, paratype, adult male,
46 mm SL. Murphy and Mill Cr. confluence, 1 mi SW Blount Springs, Blount Co., AL. 5. E. bellator,
TU 163017, adult male, 39 mm SL. Fivemile Cr. at McCalla, Jefferson Co., AL. 6. E. chermocki, TU
163026, adult male, 57 mm SL. Trib. to Turkey Cr. near Ala Hwy 79, 1 mi N Pinson, Jefferson
Co., AL, 28 April 1992. 7. E. chermocki, TU 163045, adult male, 48 mm SL. Turkey Cr. at Ala Hwy
75, 1 mi N Pinson, Jefferson Co., AL, 28 April 1992. Photos by R. D. Suttkus.

1993]

New Species of Etheostoma

11

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Tulane Studies in Zoology and Botany

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1993] New Species of £^/i^05/07nfl 15

the turquoise there are two distinct patterns of color. Posteriorly in the mid-por-
tion of the fin there is a series of red blotches that cover most of the interradial
membranous areas in the posterior part of the fin but diminish abruptly in size
in the successive membranes anteriorly. The number of red blotches varies from
three to six; each is margined or haloed with mixtures of white, cream or yellow.
The other pattern of color is a series or row of black blotches that extend from
the first interradial membrane to the seventh or eighth. These black blotches
occur in the basal half of each interradial membrane. There is a slight distal
arching of the blotches. Small nuptial males usually have fewer black blotches.
The anterior blotch is typically well developed whereas the posterior ones are
diminished in size and intensity of pigmentation. The posterior one or two black
blotches are proximal to the anterior one or two red blotches (Figure 1). Proximal
to the lower ends of the black blotches there is a pale area and beneath that is
a series of small black blotches at the base of the fin. These small blotches also
are centered on the interradial membranes.

The second dorsal fin has a moderately broad margin of turquoise. Across
the middle of the fin is a diagonal band of red color that tapers in width at
either end. The red blotches that form the band are located on the interradial
membranes. The red band quite frequently does not extend anteriorly to the
first or anteriormost interradial area. The next more proximal series (black
blotches) is greatly reduced in comparison to that described for the spinous
dorsal. The series in the second dorsal is reduced to two to four black blotches.
The posterior one to three black blotches are immediately proximal to the an-
teriormost red blotches described above. There are a few small black blotches
at the base of the fin that are similar to those described for the spinous dorsal.
The black blotches and the melanophores underlying the turquoise, red, and
orange-colored areas remain distinct for a long time in preservative, long after
the turquoise, red, and orange coloration has faded.

Nuptial females have some bright coloration although greatly reduced in
coverage compared to the nuptial male. The red blotches in the central posterior
part of the spinous dorsal are small and number three or four. There are four
to six small faint red blotches in a diagonal band across the second dorsal fin.
There are a few orange fiecks and spots on the body but these spots are scattered
and do not form a continuous series as described for the nuptial males. The
lower of the two basal caudal orange blotches is present in some females. The
spinous rays in the first dorsal and the soft rays in the second dorsal fin are
checkered with golden and dusky coloration. There are three or four golden
spots along each soft ray. The golden areas alternate with dusky areas along the
rays producing a checkered pattern. There is also a row of golden spots, each
of which is centered on the interradial membrane along the base of the spinous
dorsal fin. The blotches on the upper part of the body, on the top of the head
and on the dorsum of the body are pale to medium brown. Those on the upper
side are pale brown whereas most of the dorsal blotches are medium brown.
The lateral blotches (usually eight) are dark brown and are centered below the

Figures 8 — 1 1. Etheostoma species from Alabama and Florida. 8. E. colorosum, TU 162509, paratype,
adult female, 44 mm SL. Pine Barren Cr. at Still Road, Escambia (^o., FL. 9. E. bellator, rv 163053,
paratype, adult female, 43 mm SL. Murphy and Mill Cr. confluence, 1 mi SW Blount Springs,
Blount Co., AL. 10. £. bellator, TV 163017, adult female, 37 mm SL. FivemileCr. at McCalla, Jefferson
Co., AL. 11. £. chermocki, TU 163027, adult female, 48 mm SL. Trib. to Turkey Cr. near Ala Hwy
79, 1 mi N Pinson, Jefferson Co., AL, 28 April 1992. Photos by R. D. Suttkus.

16 Tulane Studies in Zoology and Botany [Vol. 29

lateral line. Most of the ventral lateral area below the lateral blotches is immacu-
late, except for a few scattered flecks of color. The pectoral and caudal fin rays
have dusky and pale areas along their lengths and collectively the dusky areas
form wavy vertical bars. There is some yellow along the rays of pectoral, pelvic,
and caudal fins. There is a faint wash of turquoise on the anal fin and along
upper and lower procurrent margins of the caudal fin. There is a brown prepec-
toral blotch, a brown suborbital and preorbital bar, and these brown blotches
contrast with the pale ventral parts of the head and breast.

Distribution: Etheostoma colorosum is confined to the coastal drainages of
southern Alabama and the western part of the panhandle of Florida (Figure
12). One site is known in the Perdido River drainage and numerous sites to the
east in the Escambia R., Blackwater R., Yellow R., and Choctawhatchee R. drain-
ages. The species is not known to occur in the tidal areas around the heads of
Perdido, Escambia, Blackwater, and Choctawhatchee Bays. Kuehne and Bar-
bour's (1983) distribution map (page 97) of the "Gulfcoast Snubnose Darter"
includes additional species in the Alabama and Tombigbee River drainages, and
Page and Burr's (1991) map for their "Coastal Plain Darter" also represents a
complex of species and does not include the Choctawhatchee River drainage
(see page 302, map 346). Page's (1983) plate 16G of the "Coastal Plain darter"
and Page and Burr's (1991) plate 43, bottom, of the "Coastal Plain Darter" are
not referable to Etheostoma colorosum. No other species of subgenus Ulocentra
occurs within the range of E. colorosum.

Habitat and Biology: In general the description of the habitat by Kuehne
and Barbour (1983) for the "Gulfcoast Snubnose Darter" and that by Page and
Burr (1991) for the "Coastal Plain Darter" apply to the habitat off. colorosum.
In addition to the sand and gravel substrates mentioned by the above authors,
there are a few sections of hard clay, limestone, or sandstone in the Blackwater
River drainage of Florida and in the upper reaches of the Escambia and Yellow
River dainages in Alabama that apparently serve as suitable habitats, especially
where there are some nearby patches of sand or gravel or rooted aquatic plants.

Based on recent collections, spawning occurs from early March to late April.
Some late April collections indicated the near end to spawning or included a
mixture of spent and nearspent individuals. The sex ratio is skewed toward more
females. Based on five collections of types (holotype, four series of paratopotypes,
and one series of paratypes from Pine Cr. at Fla. Hwy 99) taken over several
years (1988-1992) in March and April, the 163 specimens include 58 (35.6%)
males and 105 (64.4%) females. In one collection of 116 specimens taken on 8
March 1969 from Bear Cr., trib. to Sweetwater Cr. (Blackwater R. drainage)
there are 47 (40.5%) males versus 69 (59.5%) females. The 58 males in the five
type series range from 33.7 to 54.2 mm SL with a mean of 43.7 mm; the 105
females in the same five series range from 31.2 to 48.1 mm, mean 39.1 mm.
The 47 males in the series from Bear Cr. range from 30.1 to 53.2 mm, mean
39.1 mm; the 69 females from 28.2 to 40.4 mm, mean 34.2 mm.

The species associates of Etheostoma colorosum at the Still Rd and at the Fla
Hwy 99 sites in the five collections of types, are as follows: Ichthyomyzon gagei,
Erimyzon tenuis, Minytrema melanops, Moxostoma poecilurum, Cyprinella venusta,
Notemigonus crysoleucas, Notropis buccatus, N. hypselopterus, N. signipinnis, N. texanus,
N. welaka, Opsopoeodus emiliae, Semotilus thoreauianus, Noturus leptacanthus, Fundulus

1993] New Species of £^A^05toma 17

olivaceus, Gambusia holbrooki, Aphredoderus sayanus, Ambloplites ariommus, Lepomis
macrochirus, L. megalotis, L. punctatus, Micropterus salmoides, Etheostoma davisoni, E.
edwini, and Percina nigrofasciata.

Variation: Counts of lateral-line scales, scales around caudal peduncle,
and transverse scale rows average higher in the Yellow and Choctawhatchee
River drainages than in the three drainages to the west (Table 1). Variation in
fin-ray counts is considerably less than in scale counts. Fin-ray counts are slightly
higher in the three eastern drainages (Blackwater, Yellow, and Choctawhatchee)
than in the Perdido and Escambia River drainages (Table 2).

Comparisons: Etheostoma colorosum males differ notably from E. bellator males
in that they lack a red ocellus in the first membrane of the spinous dorsal fin;
red coloration is restricted to the last few membranes in the spinous dorsal fin
whereas the red forms a submarginal band across the entire fin in E. bellator.
Etheostoma colorosum males (Figure 1) have an incomplete series of black blotches
covering the lower one-third to one-half of the anterior membranes of the spinous
dorsal fin whereas E. bellator males have a more complete narrow band of black
across the lower part of the fin (Figures 2-5).

Etheostoma colorosum has lower scale counts (lateral-line, scales around caudal
peduncle, transverse scale rows), and lower gill raker counts than doesf. bellator
(Tables 1,3).

In general E. colorosum is more slender than E. bellator. This body shape is
best shown by comparison of body depth versus body length and caudal peduncle
depth versus caudal peduncle length. Also E. colorosum has longer soft dorsal,
caudal, and pectoral fins in both sexes than does E. bellator (Tables 4, 5), but the
anal fin is appreciably longer only in males. The spinous and soft dorsal fins
are higher in E. colorosum than in E. bellator in males and females, but the first
anal spine in E. bellator is longer than that of E. colorosum in both sexes.

Based on the similarity of the coloration of the spinous dorsal fin of males,
Etheostoma colorosum is more similar to E. tallapoosae than to any other described
member of subgenus Ulocentra. Both species lack a red ocellus in the first mem-
brane and both have the red coloration restricted to the posterior membranes
(Figure 1; Suttkus and Etnier, 1991: Figure 1). The red coloration is more
extensive in E. tallapoosae than in E. colorosum and this red coloration shades into
brown anteriorly. Together the red and the brown form a broad, more or less
pronounced band, whereas in E. colorosum the red coloration ends abruptly and
is replaced in part by black blotches that extend distally in each of the anterior
membranes (Figure 1). The small orange to red spots or punctulations described
in the diagnosis are unique to E. colorosum. The lateral blotches are smaller in
E. colorosum than in E. tallapoosae.

There are some differences in the morphometries of the two species. Based
on 25 males of each species, the soft dorsal fin length and the longest dorsal
soft ray show no overlap (Table 6). Proportional measurements of soft dorsal
fin length are 281-338 (x = 308) in E. colorosum and 242-276 (x = 260) in E.
tallapoosae. Measurements of longest dorsal soft ray are 150-191 (x = 166) in
E. colorosum and 114—146 (x = 134) in E. tallapoosae. In addition, E. colorosum
has longer caudal, pectoral, and anal fins; the longest dorsal spine and longest
anal soft ray average is greater in E. colorosum than in E. tallapoosae (Table 6).
Thus, the fins are longer and more elevated in E. colorosum than in E. tallapoosae.

18 Tulane Studies in Zoology and Botany [Vol. 29

Etymology: The name colorosum, a Latin adjective in neuter form, is from
color, color, and -osus, denoting usually fullness or abundance of, in reference
to the high color especially of nuptial males. We recommend the vernacular
name coastal darter in allusion to the coastwise distribution.

Etheostoma bellator, new species

Warrior Darter

Figures 2-5, 9-10, 13

Etheostoma (Ulocentra) sp. Smith-Vaniz, 1968: 136 (Black Warrior R.). Dycus and Howell, 1974: 23,
27 (Black Warrior R. system records). Jenkins, 1976 (undescribed species, distribution).

Warrior darter. Page, 1983: pi. 16C, 16D. Page and Burr, 1991: 303, pi. 43, map 346.

Etheostoma sp. B. Mettee, 1978: 149 (habitat, distribution). Mettee, et al. 1989a: 71-76, 189, map
(distribution, species associates, habitat). Mettee, et al. 1989b: 153 (distribution).

Etheostoma sp. Warrior snubnose darter. Boschung, et al. 1992: fig. 1, C and D; 15-20 (morphometric
data, counts, descriptive comparisons, localities).

Holotype: Adult male, tu 162448, 44.0 mm standard length, Murphy
Creek along US Hwy 31, 2.0 miles SW Blount Springs (TBS, R3W, Sec 14),
Mulberry Fork to Black Warrior River system; Blount County, Alabama, 17
April 1988, R. D. Suttkus, M. F. Mettee, P. E. O'Neil, and J. M. Pierson.

Paratopotypes: tu 151837 (76, 25-49), collected with holotype are distrib-
uted as follows: tu 151837 (70), ummz 221367 (6); tu 162671 (4, 34-43),
Murphy Cr. along US Hwy 31, 1.6 mi SW Blount Springs (T13S, R3W, Sec 11),
9 April 1992, Blount Co., AL; tu 62656 (73, 23-58), Murphy Cr. at confluence
with Mill Cr. along US Hwy 31, 1 mi SW Blount Springs (T13S, R3W, Sec 12),
19 December 1969, tu 77091 (179, 27-54), 4 April 1972, distributed as follows:
TU 77091 (155), ANSP 169685 (2), cu 73645 (2), uaic 10507.01 (4), uf 92634
(4), UMMZ 221368 (4), usnm 324502 (4), and ut 91.4203 (4); tu 89606 (89,
21-52), 5 August 1974; tu 125438 (13, 31-44), 15 May 1982; tu 151827 (175,
27-51), 17 April 1988; tu 162665 (2, 32 and 36), 9 April 1992; and tu 163053
(21, 30-48), 29 April 1992.

Other Paratypes: ummz 88848-49 (44, 19-42), Blount Springs Cr., along
US Hwy 31, 10 mi N Warrior, trib. to Mulberry Fk., Blount Co., AL; ummz
168582 (2, 33 and 36), trib. to Mulberry Fk., 5 mi SW Blountsville (Tl IN, RIW,
Sec ca. 32), Blount Co., AL; ummz 168592 (1, 26), trib. to Mulberry Fk., 2 mi
N Blountsville (Tl IN, RIW, Sec 12), Blount Co., AL; ummz 177736 (30, 27-46),
trib. to Blount Springs Cr., S Blount Springs (T13N, R24, Sec ca. 18), Blount
Co., AL; USNM 166360 (32, 21-48), and usnm 166003 (27, 27-42), trib. to
Mulberry Fk., along US Hwy 31, about 9 mi N Warrior, Blount Co., AL; and
USNM 36736 (8, 32-47), Blount Springs, Blount Co., AL.

Additional material examined but not designated as types: Black Warrior River
system, Alabama. Jefferson County: tu 62670 (4), trib. to Valley Cr., 4.7 mi SW Bessemer at
US Hwy 1 1 (T19S, R5W, Sec 36); tu 127804 (2), Fivemile Cr., trib. to Valley Cr. at McCalla (T19S,
R5W, Sec 36), TU 140953 (7), tu 140976 (9), tu 162681 (42), tu 163017 (49), tu 163187 (9); tu
61944 (8), Gurley Cr. at Ala Hwy 79, 7.7 mi N Pinson (T14S, RIW, Sec 30). Winston County:
UMMZ 168565 (2), W Fork Sipsey Fk., 6 mi E Double Springs (TION, R6W, Sec 30). Lawrence
County: tu 163061 (1), Borden Cr. at forest road (TBS, R8W, Sec 28). Blount County: ummz
158288 (2). west flowing trib. to Locust Fk., 3 mi NNE Oneonta, Hwy 32 (T12N, R2E, Sec 9).

1993] New Species of £</i^o5/oma 19

Diagnosis: Etheostoma bellator is a member of the subgenus Ulocentra as
defined by Bouchard (1977) and Bailey and Etnier (1988). Further, E. bellator
lacks a premaxillary frenum and has vomerine teeth and in these respects is
typical of members of the E. diiryi species group of Bailey and Etnier (1988).
Etheostoma bellator has a red ocellus in the first membrane of the spinous dorsal
fin in adult males. Moreover, most females of both the Murphy O. and Fivemile
Cr. populations have a red ocellus and a complete red, submarginal band in the
spinous dorsal fin. In addition, the nuptial males display two distinctive color
patterns. There are two series of more or less elongate blotches along the mid-lat-
eral area of the body and these are separated by a pale area, particularly on the
anterior half of the body. The other pattern is the narrowness of the bright
orange stripe along the lower side.

Description: Etheostoma bellator reaches a maximum of 47 mm SL (females)
to 58 mm SL (males). Sexual dimorphism and apparent sexual maturity occur
at the end of the first year at a minimum of 25 mm SL (females) and 27 mm
SL ((males). A few of the small males did not have fully developed nuptial color
patterns such as were displayed by others of the same size.

Frequency distributions of scale and fin ray counts are presented in Tables
1 and 2. Number of vertebrae, branchiostegal rays, gill rakers and preoper-
culomandibular pores are presented in Table 3. The lateral line is usually com-
plete with (41—) 45—51 (—55) scales. Occasionally the last scale is unpored. Trans-
verse scales number 12 (3 specimens), 13 (42), 14 (106), 15 (73), or 16 (16).
Transverse scales from origin of second dorsal fin to anal fin (an alternative
count, see Bailey and Etnier, 1988: 30) 11(3), 12(20), 13(46), 14(8); mean of 77
counts, 12.8. Caudal peduncle scale rows number 15 (4), 16 (19), 17 (169), 18
(65), 19 (55), 20 (1), or 21 (1). The dorsal fin has (9-) 10-11 spines and (9-)
10-12 soft rays. The anal fin has 2 spines and 6-7 (-8), modally 7, soft rays.
The pectoral fin has 13-15 rays. Branchiostegal rays number 5 — 5 (290), 5 — 6
(1), 6 — 5 (5), 6 — 6 (6) in 302 specimens; vertebrae number 37 (1), 38 (57),
or 39 (9) and gill rakers on first arch number 7 (6), 8 (35), 9 (27), or 10 (2). The
sum of left and right preoperculomandibular pores is nearly constant at 18 (in
124 of 132 specimens), the remaining eight have 17 or 19 pores. Typically the
cheek, opercle, and nape are covered with exposed scales. Occasionally the oper-
cle is not completely covered and also there may be a mixture of exposed and
embedded scales. Infrequently there is a small median naked area on the nape
just behind the occiput. The scales on the cheek are usually smaller than those
on the opercle and frequently some are embedded. The prepectoral area ranges
from being naked to completely covered with scales. The breast is usually naked
but occasionally has one or a few scales, and the belly is completely covered with
exposed scales.

All 10 specimens examined for dentition have a small cluster of one to four
or five, usually two or three, teeth on the head of the vomer; there are no
palatine teeth.

Proportional measurements appear in Tables 4 and 5. Eighteen of the 25
males were nuptial specimens taken between 4 April and 17 April. The two lots
of five each of females were collected on 4 April 1972 and 17 April 1988.

Colors of live and freshly preserved nuptial specimens, collected in April
during different years, are described from Murphy Cr. and Fivemile Cr. speci-
mens (Figures 2-5, 9-10). During the height of the breeding time the nuptial
male is brightly colored with turquoise on snout, lips, preorbital area, cheek.

20

Tulane Studies in Zoology and Botany

[Vol. 29

Table 6. Measurements in thousandths of standard length for Etheostoma colorosum and E. tallapoosae
males.

E. colorosum (N = 25)

E. tallapoosae C^^^ 2b)

Range

X

S.D.

Range

X

S.D.

Standard length (mm)

40.6—53.5

47.9

3.5

43.2 61.2

48.8

3.8

Head length

242—272

257

8.3

236—262

249

8.1

Body depth at dorsal origin

177—224

194

9.8

191—220

203

8.6

Snout length

66—81

73

3.3

66—79

72

4.0

Orbit length

63 75

68

2.8

57 67

63

2.6

Spinous dorsal fin length

273—319

300

12.3

265 308

290

10.6

Longest dorsal spine

133—182

154

11.1

99 142

120

10.9

Soft dorsal fin length

281—338

308

16.1

242—276

260

9.3

Longest dorsal soft ray

150—191

166

10.9

114—146

134

7.7

Caudal peduncle length

291—317

305

7.2

293—328

310

9.0

Caudal peduncle depth

92—112

98

5.0

99—113

106

3.7 ■

Anal fin length

264 307

280

10.5

227—276

246

11.4

First anal spine length

50 91

72

8.5

61—76

66

3.7

Longest anal soft ray

134—168

146

8.0

111—144

129

7.8

Caudal fin length

210 245

228

9.6

176—216

202

10.9

Pectoral fin length

260—343

290

18.7

221—271

250

14.3

Pelvic fin length

200—250

221

11.6

194—245

220

10.6

Trans-pelvic width

77—93

84

4.1

74—87

82

2.7

Maximum body width

123—156

137

7.2

126—164

142

7.4

Interorbital width

39—50

44

2.7

41—48

44

1.9

opercle, gular area, breast, prepectoral area, and top of head. The anterior two
median dorsal saddle blotches and the two posterior ones on the caudal peduncle
are dark turquoise as are the upper and lower procurrent margins of the caudal
fin, the entire anal fin, the pelvic fins except for a pearly white marginal fringe,
a narrow marginal fringe on the spinous dorsal and a broad marginal band on
the soft dorsal fin. The spinous dorsal fin has a bright red ocellus in the first
membrane that is followed by red blotches of less intensity in the succeeding
membranes. All together these red blotches form a submarginal band across the
entire fin. Proximal to the red band is a narrow, pale, cream-colored band (with
yellowish fringe) across the fin and proximal to that is a narrow band of small
black blotches. Another cream-colored band is immediately proximal to the black
band. The median-dorsal saddle blotches seemingly have extensions of pigment
onto the adjoining membranes of the fin. The soft-rayed dorsal fin has a broad
marginal band of turquoise that is separated from the broad median band of
red by a narrow, cream-colored area and, as described for the spinous dorsal,
there is another cream-colored area proximal to the red band. There are exten-
sions of black pigment onto the membranes of the fin at the two median-dorsal
saddle blotches at the base of the fin. The lower half of the pectoral fin is nearly
clear whereas the upper part has cream to yellow pigment along the rays.

The two series of more or less elongate blotches along the mid-lateral area
of the body are bright orange or red posteriorly and russet anteriorly. The lower
series is aligned along the lower margin of the lateral-line. The upper series is
more or less fused with the lower series at its posterior extremity on the caudal
peduncle but is noticeably separated from the lower series in its anterior half by
a pale area. The two series of elongate blotches plus the cream- to pale rose-col-
ored intermediate areas tend to obliterate the underlying brown lateral blotches.

1993]

New Species of Etheostoma

21

88^

87^

86

31

31

88^^

87^

86^

Figure 12. Distribution oi Etheostoma colorosum (solid circles; type locality solid star in open circle).

During the winter when the bright nuptial colors have faded, the brown lateral
blotches (quadrate or diamond-shape) are more pronounced and usually extend
somewhat below the lateral-line. Nuptial males have a narrow orange stripe along
the lower side. This orange stripe is separated from the lower ends of the brown
lateral blotches. The double series of more or less elongate blotches separated
by a pale area and a narrow stripe versus a broad band of orange along the
lower side are diagnostic characters of E. bellator.

Nuptial females from both Murphy Creek and Fivemile Creek (Valley Creek
system) populations have a submarginal red band in the spinous dorsal fin
(Figures 9 and 10). Similar to the nuptial male, the anterior blotch (ocellus) is
bright red whereas the blotches in the remainder of the band are subdued.
Females of both populations exhibit some small flecks of chromatic color on the
body. In addition, many Murphy Creek females have a red band across the
middle of the second dorsal fin. This band is reduced in size and intensity of
pigmentation in comparison to the respective red band in the nuptial male.

Distribution: Etheostoma bellator is confined to the Black Warrior River
system of northcentral Alabama (Figure 13). There are several disjunct popula-
tions. Our description is based on the Murphy Creek and the Valley Creek
populations.

22 Tulane Studies in Zoology and Botany ^ [Vol. 29

Habitat and Biology: Etheostoma bellator inhabits bedrock pools and race-
ways, rubble-strewn pools and fast water, riffles and moving water over gravel-fil-
led depressions in bedrock pools of small to medium streams.

Based on collections made between 1972 and 1992, spawning extends from
March through mid-May with the peak occurring in April. Four collections of
types (all nuptial specimens) from Murphy Creek and five collections of nuptial
specimens from Fivemile Creek (trib. to Valley Cr.) were analyzed for sex ratio,
size range, and average size of nuptial specimens: a collection made on 4 April
1972 has 68 males and 110 females; another made on 17 April 1988 has 60
males and 1 14 females; a second collection made on 17 April 1988 has 39 males
and 38 females; and a collection made on 29 April 1992 has 11 males and 10
females. The total for the four collections of types from Murphy Creek is 450
specimens, 178 males (39.5%) versus 272 females (60.4%). The males ranged in
SL from 26.8 to 53.8 mm, mean 36.6 mm; the females ranged from 25.4 to 46.9
mm, mean 34.6 mm. The five collections from Fivemile Creek total 116 speci-
mens; 44 males (37.9%) versus 72 females (62.1%). Males range from 32.9 to
54. 1 mm SL, mean 39.6 mm, the females from 3 1 .0 to 45.3 mm, mean 36.6 mm.

The species associates oiE. bellator at the type locality area (Murphy and Mill
Creeks) are as follows: Hypentelium etowanum, Campostoma sp., Cyprinella callistia,
C. venusta, Notropis asperifrons, Semotilus atromaculatus, Fundulus olivaceus, Lepomis
cyanellus, L. macrochirus, L. microlophus, Micropterus coosae, M. salmoides, Etheostoma
whipplei, Percina nigrofasciata, and Cottus carolinae.

Variation: There are no dramatic differences in scale or fin-ray counts
between the Murphy Creek and Valley Creek populations and there is no con-
sistency in the slight differences that could be attributed directly to a north-south
cline. The Valley Creek population, the more southern location, has a slightly
higher lateral-line scale count, about the same transverse scale row and caudal
peduncle scale counts, slightly higher dorsal spine count, about the same dorsal
soft ray count, higher anal soft ray count, but a somewhat lower pectoral fin-ray
count. The red band in the spinous dorsal and in the soft dorsal of some Murphy
Creek males is wider than any observed in Valley Creek specimens. Females of
both populations have a red band in the spinous dorsal fin, but only Murphy
Creek females were observed to have a red band in the soft dorsal fin (Figures
9-11); samples from the two populations were not compared morphometrically
and so the absence of discussion on this aspect does not automatically imply that
the two populations are similar or alike in this respect.

Comparisons: Etheostoma bellator is more similar to E. chermocki, a recently
described species (Boschung, Mayden, and Tomelleri, 1992), than toE. zonistium
(Bailey and Etnier, 1988). All three species are inhabitants of the middle or
upper portions of the Black Warrior River system, above the Fall Line. We could
not confirm all statements made by Boschung et al. (1992) in their diagnosis of
E. chermocki and their comparisons of E. chermocki and the 'Warrior darter' [ =
bellator]; we suspect this is because they did not fully appreciate the amount of
variation in color intensity and pattern that is correlated with size of nuptial
males off. chermocki and the amount of variation in color pattern in both nuptial
males and females of £. bellator (Figures 2-7, 9-1 1). We preface our remarks by
noting that we have no color slides, prints, or recent color notes for nuptial
males from Gurley Creek, the upper part of Sipsey Fork, or from the extreme

1993] New Species of Etheostoma 23

upper part of either the Mulberry or the Locust Fork. Our comparisons are
based on recent color photos and color notes, recorded at time of collection, for
E. chermocki from Turkey Creek and of £. bellator from Murphy Creek and Valley
Creek populations.

A tabular comparison of salient characteristics of nuptial males oi E. bellator
and E. chermocki is presented in Table 7. We consider our specimen (Figure 6)
and other males of about 55 mm or greater in SL to be in the larger male
category for E. chermocki; our specimen (Figure 7) and the Boschung et al. ( 1 992)
specimen (Figure 1, A, male, illustrated by painting), to be in the smaller male
category.

The red band in the spinous dorsal of nuptial males of the two species is
distinctly different, but not always in the details illustrated and discussed by
Boschung et al. (1992). Our observations indicate that the near-basal black band
in the spinous dorsal fm is quite similar in width in the two species. We agree
that the red band in the soft-rayed dorsal fin is typically broader in E. chermocki
than in E. bellator, but find no significant differences in coloration of caudal,
anal, and pectoral fins (in their diagnosis), nor are any portrayed in their Figures
1 , A and C. Two statements in their diagnosis are perplexing: "The vermilion-col-
ored venter extends dorsally and is adjacent to [emphasis added] lateral band;"
and "Lateral blotches and brick-red coloration 2iYe separated from [emphasis added]
the ventral vermilion coloration by straw-colored halos." None of our specimens,
small or large, had straw-colored halos separating the red color of the ventral-lat-
eral body area from the ventral margins of the lateral blotches. None of the
females off. chermocki in our collections taken with nuptial males had any chroma-
tic coloration in the spinous dorsal such as displayed by female E. bellator from
Murphy and Fivemile Creeks (Figures 9-11). Also, most females of £. chermocki
had jet-black pigmentation in the posterior membrane of the spinous dorsal fin.
A few E. bellator females from Fivemile Creek had a dusky posterior membrane.

Boschung et al. ( 1 992 : 1 3) stated that the nape is naked in E. chermocki whereas
our observations reveal that most specimens have the nape completely covered
with scales; a rare specimen has a narrow patch of naked skin immediately behind
the occiput. Our gill raker counts range from 7 to 10 whereas they give a range
of 5 to 7.

Comparison of data for E. chermocki and E. bellator males (Table 4) shows
slight differences in the majority of the proportions, however, there are a few
morphometries that show moderate average differences: longest dorsal spine,
longest dorsal soft ray, caudal peduncle length, and caudal fin length. Thus, in
consideration of the four morphometries there is a range in absolute separation
from 15.8% to a maximum of 69.5%. In terms of overlap, the minimum is 30.5%
to the maximum of 84.2% overlap.

Etheostoma bellator differs from E. zonistium in having fewer bands of color
across the spinous dorsal fin and in having a narrow rather than broad stripe
of orange or red on the lower side. The stripe of orange is well separated from
the lateral-band pigmentation whereas in nuptial male E. zonistium the entire
lower side is red, from the lower margin of the lateral stripe to the venter.

Status of Habitat: Mill Creek just above its confluence with Murphy
Creek is being modified by a residential development. One of the initial steps
of the project was to raise the height of the low dam on Mill Creek. The developer
seems to be making an effort to retain some of the natural beauty of the area;

24 Tulane Studies in Zoology and Botany [Vol. 29

during a visit to the area in spring of 1992 there was no evidence of siltation
below the dam. Fivemile Creek is moderately impacted by the residences in and
around the town of McCalla. The continued maintenance of bridges and small
roads, as well as an Interstate Highway that crosses the stream in the vicinity of
McCalla, periodically contributes to disturbance of the stream bed and its banks.

Etymology: The name bellator, Latin masculine noun, a warrior, is for the
Black Warrior River drainage system to which the species is confined. We use
the common name Warrior darter, following precedent of others who have
referred to the species.

Acknowledgments

A list of all individuals who contributed toward the collecting of study material
during the last five decades would be nearly endless. The following is a partial
list of those who helped collect a few to many series: Marian K. Bailey, James
M. Barkuloo, Henry L. Bart, Jr., William L. Brudon, John H. Caruso, Robert
C. Cashner, Glenn H. Clemmer, Charles D. Hancock, Julian M. Humphries,
Clyde Jones, Rudolf Meier, Maurice F. Mettee, Patrick E. O'Neil, J. Malcolm
Pierson, Susan Pollock, John S. Ramsey, Dawn Remington, Steven Rohmann,
Gerald R. Smith, Diana W. Stein, Jamie Thomerson, Bruce A. Thompson, Ver-
onica Trau Colbert, and Ralph W. Yerger. We are grateful to M. F. Mettee, P.
E. O'Neil, J. M. Pierson, and B. A. Thompson for their contributions of color
slides. Paul R. Nichols, C. Lavett Smith, and Teruya Uyeno participated in the
assembling of data, and Dr. Smith unselfishly withdrew from a planned descrip-
tion of E. colorosum initiated long ago. Several years ago R. D. Suttkus m.ade a
cursory examination of the University of Alabama's holdings of E. bellator and
E. colorosum through the courtesy of H. T. Boschung and later additional lots
were examined through the courtesy of R. L. Mayden. Although none of the
specimens was used to determine meristics or morphometries for this study, we
have taken the liberty of adding some of the (UAIC) locality data to the distribu-
tion maps. More recently M. F. Mettee offered new records of £. colorosum that
he and his staff at the Geological Survey of Alabama accumulated during the
past two years; all new localities were added to our distribution map. To all of
these individuals we express our deep appreciation.

We are especially indebted to John S. Ramsey for copies of his field notes,
his detailed color notes of specimens in nuptial condition, his shared thinking
about the relationships of the various species, and his preliminary key to all
described and undescribed species of Ulocentra that were known during the early
1970's.

We thank Ralph W. Yerger for the early loan of the extensive collections of
E. colorosum formerly housed at Florida State University. Too, we thank George
Burgess for his prompt response to our request for University of Florida (UF)
catalog numbers for the respective Florida State University series. We thank
Henry L. Bart, Jr. for extending the loan of materials from Auburn University,
and we are indebted to him and his staff for the many courtesies extended to
us at the Tulane University Museum of Natural History, and to two anonymous
reviewers.

1993]

New Species of Etheostoma

25

88'

34^

33^

87'

•)• S.Se//aUt

86

34°

33°

88°

87°

86'

Figure 13. Black. Warrior River system, Alabama, showing distribution oi Etheostoma bellator (solid
circles; type locality solid star in open circle), and E. chermocki (small solid triangles).

26

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28 Tulane Studies in Zoology and Botany

Literature Cited

Bailey, R. M. and D. A. Etnier. 1988. Comments on the subgenera of darters (Percidae) with

descriptions of two new species of Etheostoma (Ulocentra) from southeastern United States. Misc.

Publ. Mus. Zool. Univ. Michigan 175: 1-48.
Bailey, R. M. and W. A. Gosline. 1955. Variation and systematic significance of vertebral counts

in the American fishes of the family Percidae. Misc. Publ. Mus. Zool. Univ. Michigan 93: 1-44.
Bailey, R. M., H. E. Winn, and C. L. Smith. 1954. Fishes from the Escambia River, Alabama and

Florida, with ecologic and taxonomic notes. Proc. Acad. Nat. Sci. Philadelphia 106: 109-164.
BOSCHUNG, H. T., R. L. Mayden, and J. R. Tomelleri. 1992. Etheostoma chermocki, a new species

of darter (Teleostei: Percidae) from the Black Warrior River drainage of Alabama. Bull. Alabama

Mus. Nat. Hist. 13: 11-20.
Bouchard, R. W. 1977. Etheostoma etnieri, a new percid fish from the Caney Fork (Cumberland)

River system, Tennessee, with a redescription of the subgenus Ulocentra. Tulane Stud. Zool. Bot.

19: 105-130.
Dycus, D. L. and W. M. Howell. 1974. Fishes of the Bankhead National Forest of Alabama.

Alabama Dept. Cons. Natur. Resources, Div. Game & Fish, 51 pp.
Fowler, H. W. 1945. A study of the fishes of the southern Piedmont and Coastal Plain. Monogr.

Acad. Nat. Sci. Philadelphia 7: 1-408.
Gilbert, C. H. 1891. Report of explorations made in Alabama during 1889, with notes on the

fishes of the Tennessee, Alabama, and Escambia rivers. Bull. U. S. Fish Commission (1889) 9:

143-159.
Gilbert, C. R. and S. J. Walsh. 1991. Provenance and disposition of fish specimens appearing in

color plates of Kuehne and Barbour's book, The American Darters. Proc. Southeastern Fishes

Council No. 24: 12-19.
HUBBS, C. L. and K. F. Lagler. 1958. Fishes of the Great Lakes region. Cranbrook Inst. Sci. Bull.

26, 213 pp.
Jenkins, R. E. 1976. A list of the undescribed freshwater fish species of continental United States

and Canada, with additions to the 1970 checklist. Copeia 1976: 642-644.
Kuehne, R. A. and R. W. Barbour. 1983. The American Darters. Univ. Kentucky Press, Lexington,

201 pp.
Mettee, M. F. 1978. The fishes of the Birmingham-Jefferson County Region of Alabama with

ecologic and taxonomic notes. Geol. Survey Alabama Bull. no. 115, 182 pp.
Mattel, M. F., P. E. O'Neil, and S. C. Harris. 1983. A biological inventory of streams draining

the Citronelle, Pollard and Gilbertown Oil Fields in Alabama. Geol. Survey Alabama Circular

108, 101 pp.
Mettee, M. F., P. E. O'Neil, J. M. Pierson, and R. D. Suttkus. 1989a. Fishes of the Black

Warrior River system in Alabama. Geol. Survey Alabama Bull. no. 133, 201 pp.
Mettee, M. F., P. E. O'Neil, J. M. Pierson, and R. D. Suttkus. 1989b. Fishes of the western

Mobile River basin in Alabama and Mississippi. Geol. Survey Alabama Atlas no. 24, 170 pp.
Page, L. M. 1983. Handbook of Darters. T. F. H. Publications, Inc. Neptune City, New Jersey, 27 1 pp.
Page, L. M. and B. M. Burr. 1991. A Field Guide to Freshwater Fishes, North America, North of

Mexico. Houghton Mifflin Co., Boston, Mass., 432 pp.
Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B.

Scott. 1991. Common and scientific names of fishes from the United States and Canada (5th

ed.). Amer. Fish. Soc. Spec. Publ. no. 20, 183 pp.
Smith-Vaniz, W. F. 1968. Freshwater Fishes of Alabama. Auburn Univ. Agric. Exper. Sta.,21 1 pp.
Stevenson, H. M. 1976. Vertebrates of Florida, Identification and Distribution. University Presses

of Florida, Gainesville, 607 pp.
Suttkus, R. D. and D. A. Etnier. 1991 . Etheostoma tallapoosae and E. brevirostrum, two new darters,

subgenus Ulocentra, from the Alabama River drainage. Tulane Stud. Zool. Bot. 28: 1-24.

TYPE SPECIMENS OF FISHES IN THE
TULANE UNIVERSITY MUSEUM OF NATURAL HISTORY

Henry L. Bart, Jr. and Michael S. Taylor

Tulane University Museum of Natural History

Department of Ecology, EvolutioJi, and Organismal Biology

Tulane University, New Orleans, LA 70118

Introduction

This type list is being published in accordance with Recommendation 72G
of the International Code of Zoological Nomenclature (3rd Edition), which en-
courages institutions housing name-bearing types to periodically publish listings
of those types. This is the second listing of types of fishes in the Tulane University
Museum of Natural History, and is intended to update the first list, published
23 years ago (Suttkus, 1970). That first list included a history of the Tulane fish
collection, which is briefly updated here to provide information on the status of
the collection and its curation.

At the time of publication of the first type list, the fish collection, including
the type collection, was in the process of being moved to the F. Edward Hebert
("Riverside") Research Laboratories on the west bank of the Mississippi River
near Belle Chasse (15 miles from Tulane's main campus in New Orleans). The
Systematics and Environmental Biology Laboratory, the facility that initially in-
corporated the fish collection, was formally established in 1971. In 1976, Royal
D. Suttkus, the collection's founder and first Curator, convinced the Tulane
University administration to designate systematic collections at Riverside as the
Tulane University Museum of Natural History.

Suttkus retired from Tulane University in 1990, but continues to work in
the collection in his present capacity as Emeritus Curator. Also in 1990, the
Biology Department at Tulane was reorganized. The Museum is presently ad-
ministered through the Department of Ecology, Evolution, and Organismal Biol-
ogy, Milton Fingerman, Chair. The senior author was hired as Curator of
Fishes and Director of the Museum in 1992. The junior author was hired as
Collection Manager also in 1992.

Zoological collections in the Museum of Natural History are currently housed
in four of 26 World War Il-vintage ammunition bunkers on the Riverside cam-
pus. The fish collection is largely contained in Bunker A-3 near the rear of the
complex, but overflows into parts of two additional bunkers (A-2 and A- 14).
The fish collection has been expanding at a rate of 4500 lots and 200,000 speci-
mens per year. As of August 1992, it contained 6,137,652 fluid-preserved speci-
mens cataloged in 163,469 lots (an average of 37.5 specimens per lot). In addition,
there is an osteological collection with 1352 specimens, of which 638 are cleared
and stained preparations; there are 2010 radiographs representing 21,988 speci-
mens; and there are 2428 scale and otolith samples

Type Collection

Type material is housed in a separate 104 sq. ft. room in the main fish
collection bunker. The arrangement of material on the shelves follows Berg

29

30 Tulane Studies in Zoology and Botany [Vol. 29

(1940). At present, the type collection includes 33 holotypes and 38,835 paratypes
representing 125 fish species. All but the largest type specimens are stored in
glass jars of the highest quality. The necks of jars containing holotypes are
painted red, those of jars containing paratypes are painted blue. Types of
Scaphirhynchus suttkusi are kept in a stainless steel tank in the type room. Paratypes
of Ophichthus rex are stored in an eight ft. wooden tank in the fish range. Also
in the type room is a file of all original literature, and a color-coded card file
with data on types and original literature arranged chronologically and
phylogenetically. The type room also houses two specimens of the extinct harelip
sucker, Lagochila lacera, on permanent loan from Cornell University, as well as
specimens of certain rare or Federally endangered species.

Types of the Tulane fish collection are listed below in Berg family order.
Each listing includes the original species name, author(s), year of publication
and page on which species name appears, and list of types. Complete citations
for the original descriptions are provided in the literature cited. The holotype
is listed first if present. Paratypes collected with the holotype are always the first
entry in the paratype listings; remaining paratypes are listed by increasing catalog
number, which approximates a chronological listing. The complete collection
record is provided in all instances except listings of paratypes collected with
listed holotypes. Any corrections, information of the status of types, and other
information updating Suttkus (1970) are given in parentheses at the end of the
listing. Institutional abbreviations follow Leviton et al. (1985).

List of Type Specimens
Acipenseridae

Scaphirhynchus suttkusi Williams & Clemmer, 1991: 19.

HOLOTYPE: TU 135000, from Alabama, Monroe Cnty., oxbow lake of
Tensaw R. just above Little R., 18 March 1977, GHC 1411, collected by W. E.
Smith.

PARATYPES: TU 50553, 1 specimen from Alabama, Monroe Cnty., Alabama
R. at Bear Cr., river mi. 112.8, 2 March 1968, RDS 4295, collected by J. Gaddy
(fisherman); TU 52868, 1 specimen from Alabama, Wilcox Cnty., Alabama R.
at Clifton Ferry Landing, river mi. 137.3, 29 June 1968, collected by R. D.
Suttkus; TU 64937, 1 specimen from Alabama, Perry Cnty., Cahaba R. near
Marion fish hatchery, 1 mi. W of Sprott, 30 April 1966, collected by Green; TU
135001, 1 specimen from Alabama, Monroe Cnty., overflow lake of Tensaw R.
just above mouth of Little R., 5 April 1977, GHC 1412, collected by W. E. Smith
(incorrectly listed as TU 350001 in description).

Clupeidae

Brevoortia gunteri Hildebrand, 1948: 31.

PARATYPE: TU 10792, 1 specimen from Texas, Aransas Cnty., Aransas
Bay, collected by G. Gunter (formerly USNM 129802).

1993] Type Specimens of Fishes 31

Sudidae (= Chlorophthalmidae)

Chlorophthalmus brasiliensis Mead, 1958: 362.

PARATYPES: TU 18374, 4 specimens from Adantic Ocean, 01°49'N,
46°48'W, 225 fms., 17 November 1958, Oregon Sta. 2083, collected by crew of
M/V Oregon (inadvertently omitted from Suttkus, 1970).

Alepisauridae

Alepisaurus brevirostris Gibbs, 1960: 2.

PARATYPE: TU 22990, 1 specimen from Atlantic Ocean, 37°44'N, 65°42'W,
1957, collected by crew of M/V Delaware.

Characidae

Astyanax armandoi Lozano & Contreras, 1990: 97.

PARATYPES: TU 157268, 2 specimens from Chiapas, Mexico, unnamed
creek flowing through Penjamo, near Palenque, 7 May 1980, MLLV 80-33,
collected by M. L. Lozano (removed from UANL 6202).

Hjsteronoftis mjersi Thomerson &: Weitzman, 1970: 140.

PARATYPES: TU 56456, 2 specimens from Huanuco, Peru, descendents of
specimens collected from a small creek near Tournavista, 23 August 1964, col-
lected by J. E. Thomerson and party.

Cyprinidae

Algansea aphanea Barbour 8c Miller, 1978: 21.

PARATYPES: TU 105478, 10 specimens from Jalisco, Mexico, Rio de Ayutla
6.1 mi. W of junction Mascota Rd. and Mexico Hwy. 80 and 3 mi. SE of Ayutla,
22 April 1969, collected by C. D. Barbour and R. J. Douglas (removed from
UMMZ 192196).

Algansea monticola Barbour 8c Contreras, 1968: 101.

HOLOTYPE: TU 40869, from Zacatecas, Mexico, Rio Juchipila near town
of Jalpa, at Mexico Hwy. 70 bridge, one mi. E of the intersection with Mexico
Hwy. 4 1 , 30 June 1 963, CDB 63-27, collected by C. D. Barbour and S. Contreras.

PARATYPES: TU 30611, 49 specimens collected with holotype; TU 30608,
76 specimens from Zacatecas, Mexico, Rio Juchipila at Mai Paso (a little walled
town) about 16 mi. E of Ciudad Garcia Sahnas, 28 June 1963, CDB 63-23,
collected by C. D. Barbour and S. Contreras; TU 30609, 131 specimens from
Zacatecas, Mexico, tributary to Rio Juchipila, about 5 mi. S of V^illanueva on
Mexico Hwy. 41, 28 June 1963, CDB 63-24, collected by C. D. Barbour and S.
Contreras; TU 30610, 18 specimens from Zacatecas, Mexico, Rio Juchipila at
Tabasco, upstream of bridge at south end of town, 29 June 1963, CDB 63-25,
collected by C. D. Barbour and S. Contreras.

Algansea monticola avia Barbour 8c Miller, 1978: 10.

32 Tulane Studies in Zoology and Botany [Vol. 29

PARATYPES: TU 30816, 20 specimens from Nayarit, Mexico, Rio Tepic at
Mexico Hwy. 15, 27 mi. SE of Tepic, 1957, M57-12, collected by R. R. Miller
and assistants; TU 30817, 13 specimens from Nayarit, Mexico, Arroyo de Maza-
tan at Mazatan, 1955, M55-25, collected by R. R. Miller and assistants; TU 30818,
15 specimens from Nayarit, Mexico, Rio Compostelajust S of Compostela, 1955,
M55-40, collected by R. R. Miller and assistants.

Catnpostoma pauciradii Burr &: Cashner, 1983: 105.

HOLOTYPE: TU 117030, from Georgia, Hall Cnty., Candler Cr., 1.1 mi.
W of Gillsville, Georgia Hwy. 52, 24 April 1980, RDS 7314, collected by R. D.
Suttkus, D. C. Scott, F. Myers, and S. Larson.

PARATYPES: TU 117031, 5 specimens collected with holotype; TU 26138,
1 specimen from Georgia, Gwinnett Cnty., Yellow R. 7 mi. E of Duluth, Georgia
Hwy. 120, 20 April 1962, RDS 3057, collected by R. D. Suttkus, J. S. Ramsey,
J. E. Thomerson, and C. D. Barbour (incorrectly listed as TU 25138 in descrip-
tion); TU 30489, 5 specimens from Georgia, Hall Cnty., North Oconee R. 5.9
mi. NE of Gainesville, U.S. Hwy. 23, 22 September 1963, JSR 63-47, collected
by J. S. Ramsey; TU 30499, 3 specimens from Georgia, Hall Cnty., East Fork
Oconee R. about 2 mi. N of Gillsville, Georgia Hwy. 52, 22 September 1963,
JSR 63-48, collected by J. S. Ramsey; TU 41351, 10 specimens from Georgia,
Hall Cnty., Flat Shoals Cr., at Georgia Hwy. 52 crossing, 1 mi. W of junction of
Georgia Hwy. 323, 4 May 1966, GHC 440, collected by G. H. Clemmer and J.
Reed; TU 114877, 25 specimens from Georgia, Hall Cnty., Candler Cr., 1.2 mi.
W of Gillsville, Georgia Hwy. 52, 12 October 1979, RDS 7157, collected by R.
D. Suttkus, R. C. Cashner, R. Gilbert, and A. R. Heidt; TU 116945, 1 specimen
from Georgia, Hall Cnty., Candler Cr., 1.1 mi. W of Gillsville, Georgia Hwy. 52,
1 April 1980, RDS 7307, collected by R. D. Suttkus and B. J. Freeman; TU
116956, 3 specimens from Georgia, Hall Cnty., Candler Cr., 2 air mi. NW of
Gillsville, 1 April 1980, RDS 7308, collected by R. D. Suttkus and B.J. Freeman;
TU 116964, 5 specimens from Georgia, Hall Cnty., North Oconee R., 2.8 mi.
W of U.S. Hwy. 23, along Whitehall Rd., 1 April 1980, RDS 7309, collected by
R. D. Suttkus and B. J. Freeman; TU 117042, 13 specimens from Georgia, Hall
Cnty., North Oconee R., 2.8 mi. W of U.S. Hwy. 23, along Whitehall Rd., 24
April 1980, RDS 7315, collected by R. D. Suttkus, D. C. Scott, F. Myers, and S.
Larson.

Dionda tnandibularis Contreras & Verduzco, 1977: 259.

PARATYPES: TU 94115, 2 specimens from San Luis Potosi, Mexico, Arroyo,
7 km SSE of the town of Rioverde, 14 July 1968, SCB 68-16, collected by S.
Contreras and party (removed from UANL 1032).

Gila bicolor eurysoma Williams & Bond, 1981: 223.

HOLOTYPE: TU 116230, from Nevada, Washoe Cnty., Fish Cr., approxi-
mately 0.8 km upstream from Andy's Place, 1 1 August 1978, J W 78-32, collected
by J. E. Williams and C. Williams.

PARATYPES: TU 116231, 11 specimens collected with holotype.

1993] T^yp^ Specimens of Fishes 33

Gila boraxobius Williams Sc Bond, 1980: 292.

PARATYPES: TU 116232, 8 specimens from Oregon, Harney Cnty., Hoi
Lake (= Borax Lake), 5 mi. N of Fields, 30 July 1972, collected by C. E. Bond
(removed from OS 4138).

Hybopsis lineapunctata Clemmer & Suttkus, 197 1 : 22. ( = Notropis lineapunctatus)

HOLOTYPE: TU 53405, from Alabama, Clay Cnty., Enitachope Cr., tribut-
ary to Hillabee Cr., 2.9 mi. SW of Ashland, Alabama Hwy. 9, 13 April 1966,
RDS 3877, collected by R. D. Suttkus and G. H. Clemmer.

PARATYPES: TU 40643, 46 specimens collected with holotype; TU 12085,
8 specimens from Alabama, Elmore Cnty., Gold Branch, 6 mi. N of Eclectic,
Alabama Hwy. 63, 5 October 1955, RDS 2525, collected by R. D. Suttkus; TU
15281, 156 specimens from Alabama, Elmore Cnty., Gold Branch, tributary to
Channahatchee Cr., 6 mi. N of Eclectic, Alabama Hwy. 63, 16 March 1957, RDS
2592, collected by R. D. Suttkus, R.J. Miller, H. V. Miller, and J. DeAbate; TU
29884, 1 specimen from Alabama, Clay Cnty., Enitachope Cr., tributary to Hill-
abee Cr., 3.2 mi. SW of Ashland, Alabama Hwy. 9, 21 September 1963, JSR
63-46, collected by J. S. Ramsey; TU 32729, 8 specimens from Alabama, Clay
Cnty., Enitachope Cr., 2.9 mi. S of Ashland, Alabama Hwy. 9, 1 June 1964, RDS
3462, collected by R. D. Suttkus and J. S. Ramsey; TU 40668, 12 specimens from
Alabama, Randolph Cnty., tributary to Little Tallapoosa R., 2.9 mi. W of inter-
section of U.S. Hwy. 431 and Alabama Hwy. 82, 13 April 1966, RDS 3878,
collected by R. D. Suttkus and G. H. Clemmer; TU 41118, 28 specimens from
Alabama, Elmore Cnty., Gold Branch, tributary to Tallapoosa R., 6 mi. N of
Eclectic, Alabama Hwy. 63, 12 April 1966, RDS 3873, collected by R. D. Suttkus
and G. H. Clemmer; TU 41132, 1 specimen from Alabama, Elmore Cnty., Chan-
nahatchee Cr., tributary to Tallapoosa R., 2.5 mi. S of Red Hill, Alabama Hwy.
229, 12 April 1966, RDS 3874, collected by R. D. Suttkus and G. H. Clemmer.

Nocomis asper Lachner & Jenkins, 1971b: 3.

PARATYPES: TU 48547, 9 specimens from Arkansas, Benton Cnty.,
Spavinaw Cr. hole no. 3, 10 June 1966, University of Arkansas graduate students;
TU 48613, 28 specimens from Arkansas, Benton Cnty., Spavinaw Cr. hole no.
2, 10 June 1966, University of Arkansas graduate students; TU 48658, 14 speci-
mens from Arkansas, Benton Cnty., Spavinaw Cr. hole no. 4, 10 June 1966,
University of Arkansas graduate students.

Nocomis effusus Lachner &: Jenkins, 1967: 560.

PARATYPES: TU 19488, 2 specimens from Tennessee, Rutherford Cnty.,
Stones R., 6.5 mi. N of Murfreesboro, U.S. Hwy. 231, 7 September 1958, RDS
2758, collected by R. D. Suttkus and M. Andersson; TU 32838, 5 specimens
from Tennessee, Fentress Cnty., tributary to Wolf R., 9.8 mi. N of Jamestown,
U.S. Hwy. 127, 3 June 1964, RDS 3474, collected by R. D. Suttkus and J. S.
Ramsey; TU 33115, 2 specimens from Tennessee, Cheatham Cnty., Harpeth R.,
8 mi. E of White Bluff, U.S. Hwy. 70, 15 July 1964, JSR 64-60, collected by J.
S. Ramsey and D. W. Walton; TU 33125, 1 1 specimens from Tennessee, William-
son Cnty., South Harpeth R., 4.2 mi. E of Fairview, Tennessee Hwy. 96, 15 July

34 Tulane Studies in Zoology and Botany [Vol. 29

1964, JSR 64-61, collected by J. S. Ramsey and D. W. Walton; TU 33171, 21
specimens from Tennessee, Rutherford Cnty., West Fork Stones R. at W edge
of Murfreesboro, Tennessee Hwy. 96, 15 July 1964, JSR 64-63, collected by J.
S. Ramsey and D. W. Walton; TU 33194, 1 specimen from Tennessee, Rutherford
Cnty., Stones R. at S limit of Walterhill, U.S. Hwy. 231, 16 July 1964, JSR 64-64,
collected by J. S. Ramsey and D. W. Walton.

Nocotnis leptocephalus interocularis Lachner & Wiley, 1971: 6.

PARATYPES: TU 12231, 6 specimens from Georgia, Rabun Cnty., Timspon
Cr., 5.4 mi. W of Clayton, U.S. Hwy. 76, 8 October 1955, RDS 2533, collected
by R. D. Suttkus and party; TU 26178, 11 specimens from Georgia, Madison
Cnty., South Fork Broad R., 1 mi. W of Comer, Georgia Hwy. 72, 21 April 1962,
RDS 3061, collected by R. D. Suttkus, J. S. Ramsey, J. E. Thomerson, and C. D.
Barbour; TU 26181, 19 specimens from Georgia, Madison Cnty., South Fork
Broad R., 1 mi. S of Carlton, 21 April 1961, RDS 3062, collected by R. D. Suttkus,
J. S. Ramsey, J. E. Thomerson, and C. D. Barbour; TU 26190, 30 specimens
from South Carolina, Anderson Cnty., tributary to Savannah R. 5 mi. W of Iva,
South Carolina Hwy. 184, 21 April 1962, RDS 3063, collected by R. D. Suttkus,
J. S. Ramsey, J. E. Thomerson, and C. D. Barbour; TU 26955, 12 specimens
from South Carolina, Oconee Cnty., Little R. 8 mi. NE of Walhalla on South
Carolina Hwy. 11, 5 July 1962, JSR 62-117, collected by J. S. Ramsey, W. S.
Woolcott, and H. H. Howell; TU 26971, 5 specimens from South Carolina,
Oconee Cnty., Thompson R., tributary to Whitewater R., 1.4 mi. S of South
Carolina-North Carolina state line in Sumter National Forest, 10 July 1962, JSR
62-122, collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 26974,
5 specimens from South Carolina, Oconee Cnty., Chauga R. 15.9 mi. S of Georgia-
South Carolina state line on South Carolina Hwy. 28, 13 July 1962, JSR 62-136,
collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 26979, 1
specimen from Georgia, Rabun Cnty., and South Carolina, Oconee Cnty., Chat-
tooga R. at Georgia-South Carolina state line, South Carolina Hwy. 28 above
junction with West Fork, 13 July 1962, JSR 62-135, collected by J. S. Ramsey,
W. S. Woolcott, H. H. Howell, and D. H. Howell; TU 26996, 6 specimens from
South Carolina, Pickens/Oconee Cnty. line, confluence of Horsepasture and
Toxaway rivers, 1.5 mi. S of North Carolina-South Carolina state line, 21 July
1962, JSR 62-152, collected by J. S. Ramsey and H. H. Howell; TU 27001, 34
specimens from North Carolina, Transylvania Cnty., Bearcamp Cr., tributary
to Horsepasture R., 4.0 mi. S of Oakland, 24 July 1962, JSR 62-155, collected
by J. S. Ramsey and H. H. Howell; TU 28017, 2 specimens from North Carolina,
Jackson Cnty., tributary to Chattooga R. 1.7 mi. S of Cashiers, North Carolina
Hwy. 107, 4 June 1962, JSR 62-67, collected byj. S. Ramsey and W. S. Woolcott;
TU 28023, 15 specimens from North Carolina, Transylvania Cnty., Horsepasture
R., tributary to Toxaway R., 0.5 mi. N of North Carolina-South Carolina state
line, 6 June 1962, JSR 62-72, collected by J. S. Ramsey and W. S. Woolcott; TU
29261, 5 specimens from North Carolina, Transylvania Cnty., Bearwallow Cr.
near confluence with Toxaway R., 1.9 mi. due N of North Carolina-South
Carolina state line, 7 June 1962, JSR 62-73, collected by J. S. Ramsey and W. S.
Woolcott; TU 29288, 11 specimens from North Carolina, Transylvania Cnty.,
Bearwallow Cr. at confluence with Toxaway R., 2 mi. due N of South Carolina
state line, 10 June 1962, JSR 62-79, collected byJ. S. Ramsey and W. S. Woolcott;
TU 29303, 5 specimens from North Carolina, Transylvania Cnty., Horsepasture

1993] Type Specimens of Fishes 35

R. 1.1 mi. S of Oakland on Sapphire Rd., 14 June 1962, JSR 62-81, collected by
J. S. Ramsey and W. S. Wolcott; TU 29309, 21 specimens from North Carolina,
Transylvania Cnty., Toxaway R. above Lake Toxaway, 2.6 mi. W of Lake Toxa-
way (town) on gravel road, 15 June 1962, JSR 62-88, collected by J. S. Ramsey
and W. S. Woolcott; TU 29314, 10 specimens fron: North Carolina, Transylvania
Cnty., Horsepasture R., 2.5 mi. S of Oakland on Whitewater Rd., 18 June 1962,
JSR 62-90, collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU
29319, 7 specimens from North Carolina, Transylvania Cntv., Horsepasture R.,
3 mi. S of Oakland on gravel road, 18 June 1962, JSR 62-91, collected bv J. S.
Ramsey, W. S. Woolcott, and H. H. Howell; TU 29322, 9 specimens from Georgia,
Habersham Cnty., Panther Cr., tributary to Tallulah R. 3.9 mi. SW of Tallulah
Falls, U.S. Hwy. 441, 19 June 1962, JSR 62-92, collected by J. S. Ramsey, W. S.
Woolcott, and H. H. Howell; TU 29329, 24 specimens from Georgia, Rabun
Cnty., Tallulah R. 1.5 mi. S of Wiley, U.S. Hwy. 441, 19 June 1962, JSR 62-93,
collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 29391, 12
specimens from North Carolina, Transylvania Cnty., Bearcamp Cr., tributary
to Horsepasture R., 4.0 mi. S of Oakland, 25 June 1962, JSR 62-103, collected
by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 29420, 8 specimens
from North Carolina, Transylvania Cnty., Bearcamp Cr., tributarv to Horsepas-
ture R., 0.2 mi. N of South Carolina state line, 28 June 1962, JSR 62- 107, collected
by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 29422, 2 specimens
from North Carolina, Transylvania Cnty., Bearwallow Cr., tributary to Toxaway
R., below 60 ft. falls, 1.7 mi. N of South Carolina state line, 29 June 1962, JSR
62-108, collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 29436,
3 specimens from North Carolina, Transylvania Cnty., Horsepasture R. between
Drift Falls and Rainbow Falls, 2.6 mi. SW of Oakland, 3 July 1962, JSR 62-1 10,
collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 29449, 6
specimens from North Carolina, Transylvania Cnty., Toxaway R. just below
Toxaway Falls, Hwy. 64, 7 July 1962, JSR 62-118, collected by J. S. Ramsey and
W. S. Woolcott; TU 29450, 1 specimen from North Carolina, Transylvania Cnty.,
Toxaway R. above Lake Toxaway, 2.6 mi. West of Lake Toxaway (town) on gravel
road, 2 July 1962, JSR 62-119, collected by J. S. Ramsey and W. S. Woolcott; TU
29461, 2 specimens from South Carolina, Oconee Cnty., Bearcamp Cr. nearjunction
with Horsepasture R., 0.5 mi. SE of North Carolina state line, lOJulv 1962, JSR
62-123, collected by J. S. Ramsey, W. S. Woolcott, and H. H. Howell; TU 29534,
11 specimens from North Carolina, Transylvania Cnty., Toxaway R. above Lake
Toxaway, 2.6 mi. W of Lake Toxaway (town) on gravel road, 14 July 1962, JSR
62-138, collected by J. S Ramsey, H. H. Howell, D. H. Howell, and W. C. Howell;
TU 29585, 13 specimens from South Carolina, Pickens-Oconee Cnty. line, Toxa-
way R. 0.25 mi. S of junction with North Carolina state line, 23 July 1962, JSR
62-154, collected by J. S. Ramsey and H. H. Howell; TU 29599, 27 specimens
from North Carolina, Transylvania Cnty., Toxaway Cr., tributary to Toxaway
R., about 3 mi. due SW of Rosman, gravel road, 31 July 1962, JSR 62-157,
collected by J. S. Ramsey and H. H. Howell; TU 29652, 5 specimens from North
Carolina, Jackson Cnty., Chattooga R. 2.7 mi. due S of Cashiers, gravel road, 7
August 1962, JSR 62-169, collected by J. S. Ramsey; TU 29784, 12 specimens
from North Carolina, Transylvania Cnty., Bearcamp Cr., tributary to Horsepas-
ture R., 4.0 mi. S of Oakland, 27 August 1962, JSR 62-198, collected by J. S.
Ramsey; TU 37510, 65 specimens from Georgia, Banks Cnty., Hudson R., tribu-
tary to Broad R. about 2 mi. N of Homer, U.S. Hwy 441, 24 April 1965, RDS

36 Tulane Studies in Zoology and Botany [Vol. 29

3656, collected by R. D. Suttkus and F. L. Rose; TU 38588, 9 specimens from
Georgia, Columbia Cnty., Little Kiokee Cr., 10.5 mi. N of Evans, 27 June 1965,
FLR 1, collected by F. L. Rose and J. L. Rose; TU 38597, 3 specimens from
Georgia, Richmond Cnty., Rays Cr. at crossing of Walton Way extension in
Augusta, 5 July 1965, FLR 2, collected by F. L. Rose and J. L. Rose; TU 38605,
2 specimens from Georgia, Richmond Cnty., Rays Cr. at crossing of Barkman
Rd. in Augusta, 6 July 1965, FLR 3, collected by F. L. Rose, J. L. Rose, and W.
T. Rose; TU 38613, 15 specimens from South Carolina, McCormick Cnty., Ste-
vens Cr. 1.4 mi. N of Parksville, Cnty. Rd. S33-21, 7 July 1965, FLR 4, collected
by F. L. Rose, J. L. Rose, and W. T. Rose; TU 38623, 4 specimens from South
Carolina, Edgefield Cnty., tributary to Stevens Cr., 4.6 mi. E of Parksville, Cnty.
Rd. S33-138, 7 July 1965, FLR 5, collected by F. L. Rose, J. L. Rose, and W. T.
Rose; TU 39477, 9 specimens from South Carolina, Edgefield Cnty., Log Cr. at
U.S. Hwy. 25 crossing, 4 mi. N of U.S. Hwy. 25-25A intersection, just N of
Edgefield, 25 October 1965, GHC 366, collected by G. H. Clemmer and F. L.
Rose; TU 39486, 2 specimens from Georgia, Columbia Cnty., Little Kiokee Cr.,
10.5 mi. N of Evans on dirt road, 26 October 1965, GHC 367, collected by G.
H. Clemmer and F. L. Rose.

Nocomis platyrhynchus Lachner & Jenkins, 1971a: 37.

PARATYPES: TU 25902, 17 specimens from North Carolina, Ashe Cnty.,
South Fork New R., 4 mi. SW of Scottville, U.S. Hwy. 221, 19 June 1962, RDS
3109, collected by R. D. Suttkus and M. M. May.

iVocomis ra/ieji Lachner & Jenkins, 1971a: 17.

PARATYPES: TU 25885, 3 specimens from Virginia, Roanoke Cnty.,
Roanoke R., 8 mi. SW of Salem, U.S. Hwy. 11, 18 June 1962, RDS 3107, collected
by R. D. Suttkus and M. M. May.

Notropis aguirrepequenoi Contreras &: Rivera, 1973: 259.

PARATYPES: TU 94116, 20 specimens from Tamaulipas, Mexico, Rio Pilon
at Mexico Hwy. 85, 21 May 1966, SCB 66-8, collected by S. Contreras and party
(removed from UANL 1137).

Notropis ammophilus Suttkus & Boschung, 1990: 50.

HOLOTYPE: TU 151630, from Alabama, Dallas-Wilcox Cnty. line,
Chilatchee Cr., 0.4 mi. N of Alberta, Alabama Hwy. 5, 9 April 1988, RDS 9534,
collected by R. D. Suttkus.

PARATYPES: TU 151631, 124 specimens collected with holotype; TU 60871,
1 147 specimens from Alabama, Dallas- Wilcox Cnty. line, Chilatchee Cr., 0.4 mi.
N of Alberta, Alabama Hwy. 5, 18 December 1969, RDS 4709, collected by R.
D. Suttkus and R. C. Cashner; TU 121386, 129 specimens from Alabama, Dallas-
Wilcox Cnty. line, Chilatchee Cr. 0.4 mi. N of Alberta, Alabama Hwy. 5, 15 May
1981, RDS 7553, collected by R. D. Suttkus; TU 125388, 292 specimens from
Alabama, Dallas-Wilcox Cntv. line, Chilatchee Cr. 0.4 mi. N of Alberta, Alabama
Hwy. 5, 15 May 1982, RDS' 7857, collected by R. D. Suttkus and D. W. Stein;
TU 140934, 207 specimens from Alabama, Dallas- Wilcox Cnty. line, Chilatchee

1993] Type Specimens of Fishes 37

Cr. 0.4 mi. N of Alberta, Alabama Hwy. 5, 24 May 1985, RDS 8770, collected
by R. D. Suttkus; TU 145519, 1 17 specimens from Alabama, Dallas-Wilcox Cnty.
line, Chilatchee Cr. 0.4 mi. N of Alberta, Alabama Hwy. 5, 20 May 1986, RDS
9030, collected by R. D. Suttkus; TU 153083, 98 specimens from Alabama,
Dallas- Wilcox Cnty. line, Chilatchee Cr. 0.4 mi. N of Alberta, Alabama Hwy. 5,
2 August 1988, RDS 9656, collected by R. D. Suttkus; TU 153185, 296 specimens
from Alabama, Dallas-Wilcox Cnty. line, Chilatchee Cr. 0.4 mi. N of Alberta,
Alabama Hwy. 5, 3 August 1988, RDS 9661, collected by R. D. Suttkus; TU
154399, 25 specimens from Alabama, Dallas-Wilcox Cnty. line, Chilatchee Cr.
0.4 mi. N of Alberta, Alabama Hwy. 5, 3 February 1989, RDS 9768, collected
by R. D. Suttkus; TU 154734, 19 specimens from Alabama, Dallas- Wilcox Cnty.
hne, Chilatchee Cr. 0.4 mi. N of Alberta, Alabama Hwy. 5, 9 May 1989, RDS
9799, collected by R. D. Suttkus.

Notropis asperifrons Suttkus 8c Raney, 1955c: 4.

PARATYPES: TU 2974, 22 specimens from Alabama, Wilcox Cnty., Gravel
Cr. tributary to Pursley Cr., 6.3 mi. S of Camden, Hwy. 11, 3 June 1951, RDS
2140, collected by R. D. Suttkus and C. D. Hancock; TU 3063, 5 specimens from
Alabama, Wilcox Cnty., tributary to Pursley Cr., 1.8 mi. E of Camden, Alabama
Hwy. 10, 3 June 1951, RDS 2138, collected by R. D. Suttkus and C. D. Hancock;
TU 3426, 8 specimens from Alabama, Wilcox Cnty., Pursley Cr., tributary to
Alabama R., 3.4 mi. SW of Camden, Hwy. 11, 3 June 1951, RDS 2139, collected
by R. D. Suttkus and C. D. Hancock; TU 4251, 28 specimens from Alabama,
Shelby Cnty., tributary to Waxahatchee Cr., 4.7 mi. SW of Columbiana on
Alabama Hwy. 25, 15 June 1952, RHG 356, collected by R. H. Gibbs and P. P.
Caswell; TU 97113, 5 specimens from Georgia, Murray Cnty., tributary to Holly
Cr. at Ramhurst, 8 mi. N of the Murray Cnty. line, U.S. Hwy. 411, 12 June
1952, RHG 342, collected by R. H. Gibbs and P. P. Casell (removed from CU
28263).

Notropis atrapiculus Snehon, 1972: 58. (= Lythrurus atrapiculus)

PARATYPES: TU 2509, 36 specimens from Alabama, Houston Cnty.,
Panther Cr., tributary to Little Choctawhatchee R., 3 mi. west of Pinckard Farm
Rd. and U.S. Hwy. 84, 29 May 1951, RDS 2108, collected by R. D. Suttkus
(incorrectly listed as Pinehard Rd. in original description).

Notropis baileyi Suttkus & Raney, 1955a: 71.

PARATYPES: TU 2637, 29 specimens from Alabama, Monroe Cnty., tribu-
tary to Alabama R., 17.3 mi. S of Camden, Hwy. 11, 3 June 1951, RDS 2141,
collected by R. D. Suttkus and C. D. Hancock (destroyed); TU 2965, 4 specimens
from Alabama, Wilcox Cnty., Gravel Cr., tributary to Pursley Cr., 6.3 mi. S of
Camden, Hwy. 11,3 June 1951, RDS 2140, collected by R. D. Suttkus and C.
D. Hancock; TU 3066, 41 specimens from Alabama, Wilcox Cnty., tributary to
Pursley Cr., 1.8 mi. E of Camden, Alabama Hwy. 10, 3 June 1951, RDS 2138,
collected by R. D. Suttkus and C. D. Hancock; TU 3195, 4 specimens from
Alabama, Butler Cnty., Pine Barren Cr., 2.5 mi. S of Forest Home, 2 June 1951,
RDS 2 1 36, collected by R. D. Suttkus and C. D. Hancock; TU 3430. 75 specimens
from Alabama, Wilcox Cnty., Pursley Cr., tributary to Alabama R., 3.4 mi. SW

38 Tulane Studies in Zoology and Botany [Vol. 29

of Camden, Hwy. 11, 3 June 1951, RDS 2139, collected by R. D. Suttkus and
C. D. Hancock; TU 97110, 66 specimens from Alabama, Macon Cnty., tributary
to Sawachlahatchee Cr., 1.7 mi. W of Society Hill on U.S. Hwy. 80, 12 June
1949, RDS 1051, collected by R. D. Suttkus, C. F. Cole, and R. H. Gibbs (formerly
CU 16020).

Notropis bellus alegnotus Snelson, 1972: 37. (= Lythrurus bellus alegnotus)

PARATYPES: TU 58810, 10 specimens from Alabama, Jefferson Cnty., Five
Mile Cr., tributary to Valley Cr., U.S. Hwy. 1 1 bridge, 5.0 air mi. SW of junction
Alabama Hwy. 150 & U.S. Hwy. 11 bridge in Bessemer, 16 May 1969, FFS 69-7,
collected by F. F. Snelson and B. W. Menzel.

Notropis cahabae Mayden 8c Kuhajda, 1989: 3.

PARATYPES: TU 156264, 3 specimens from Alabama, Bibb Cnty., Cahaba
R. at mouth of unnamed tributary, 1.1 km downstream from mouth of Little
Ugly Cr . , 7 February 1 985 (incorrectly listed as TU 156263 in original description)
(removed from UAIC 8578.02).

Notropis candidus Suttkus, 1980: 3.

HOLOTYPE: TU 103415, from Alabama, Wilcox Cnty., Alabama R. at Yellow
Jacket Bar, river mi. 129.8, 26 June 1969, RDS 4566, collected by R. D. Suttkus,
G. E. Gunning, and R. C. Cashner.

PARATYPES: TU 57763, 84 specimens collected with holotype; TU 40313,
25 specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar,
river mi. 129.8, 7 April 1966, RDS 3858, collected by R. D. Suttkus and G. E.
Gunning; TU 40903, 26 specimens from Alabama, Wilcox Cnty., Alabama R. at
Evans Upper Bar, river mi. 135.8, 28 June 1966, RDS 3918, collected by R. D.
Suttkus, G. E. Gunning, and G. H. Clemmer; TU 40928, 13 specimens from
Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.8, 28
June 1966, RDS 3919, collected by R. D. Suttkus, G. E. Gunning, and G. H.
Clemmer; TU 41414, 35 specimens from Alabama, Wilcox Cnty., Alabama R.
at Yellow Jacket Bar, river mi. 129.8, 1 July 1966, RDS 3924, R. D. Suttkus and
Environmental Biology class; TU 47903, 109 specimens from Alabama, Wilcox
Cnty., Alabama R. at Evans Upper Bar, river mi. 135.8, 26 September 1967,
RDS 4202, collected by R. D. Suttkus, G. E. Gunning, and A. Kuris; TU 52832,
37 specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar,
river mi. 129.8, 29 June 1968, RDS 4375, collected by R. D. Suttkus and G. E.
Gunning; TU 52854, 23 specimens from Alabama, Wilcox Cnty., Alabama R. at
Evans Upper Bar, river mi. 135.8, 29 June 1968, RDS 4376, collected by R. D.
Suttkus and G. E. Gunning; TU 76231, 132 specimens from Alabama, Wilcox
Cnty., Alabama R. at Evans Lower Bar, river mi. 133, 16 March 1972, RDS 5193,
collected by R. D. Suttkus and B. A. Thompson; TU 76253, 203 specimens from
Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.8, 16
March 1972, RDS 5194, collected by R. D. Suttkus and B. A. Thompson; TU
80311, 250 specimens from Alabama, Wilcox Cnty., Alabama R. at Evans Lower
Bar, river mi. 133, 6 December 1972, RDS 5350, collected by R. D. Suttkus and
R. C. Cashner; TU 86774, 212 specimens from Alabama, Wilcox Cnty., Alabama
R. at Evans Lower Bar, river mi. 133, 19 March 1974, RDS 5590, collected by

1993] Type Specimens of Fishes 39

R. D. Suttkus and B. A. Thompson; TU 96458, 139 specimens from Alabama,
Wilcox Cnty., Alabama R. at Evans Lower Bar, river mi. 133, 18 December 1975,
RDS 6087, collected by R. D. Suttkus and G. E. Gunning.

Notropis cardinalis Mayden, 1988: 156. (= Luxilus cardinalis)

PARATYPES: TU 145026, 3 specimens from Oklahoma, Ottawa Cnty., Five
Mile Cr., T29N, R24E, Sec. 22, 9 May 1984, collected by F. B. Cross and R. L.
Mayden (removed from KU 20963).

Notropis edwardraneyi Suttkus & Clemmer, 1968: 19.

HOLOTYPE: TU 49485, from Alabama, Wilcox Cnty., Alabama R. at Yellow
Jacket Bar, river mi. 129.8, 8 March 1967, RDS 4097, collected by R. D. Suttkus
and G. E. Gunning.

PARATYPES: TU 44028, 7324 specimens collected with holotype; TU 33381 ,
1613 specimens from Alabama, Dallas Cnty., Alabama R. at Watts Bar, 3.5 mi.
upstream from Cahaba, river mi. 204.5, 29 June 1964, RDS 3515, collected by
R. D. Suttkus and Environmental Biology class; TU 35243, 236 specimens from
Alabama, Dallas Cnty., Alabama R. at old ferry landing across river from Cahaba,
27-28 June 1964, RDS 3508, collected by R. D. Suttkus and Environmental
Biology class; TU 35269, 314 specimens from Alabama, Dallas Cnty., Alabama
R. at Cahaba where Cahaba R. enters Alabama R., 28 June 1964, RDS 3513,
collected by R. D. Suttkus, G. H. Clemmer, J. D. Archer, and J. V. Conner; TU
35323, 28 specimens from Alabama, Clarke Cnty., Alabama R. along left bank
across river from Choctaw Bluff, 2 July 1964, RDS 3519, collected by R. D.
Suttkus and Environmental Biology class; TU 40293, 297 specimens from
Alabama, Wilcox Cnty., Alabama R. at Evans Upper Bar, river mi. 135.8, 7 April
1966, RDS 3857, collected by R. D. Suttkus and G. E. Gunning; TU 40303, 1020
specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river
mi. 129.8, 7 April 1966, RDS 3858, collected by R. D. Suttkus and G. E. Gunning;
TU 40320, 491 specimens from Alabama, Wilcox Cnty., Alabama R. at Tait Bar,
river mi. 122.4, 7 April 1966, RDS 3859, collected by R. D. Suttkus and G. E.
Gunning; TU 40335, 243 specimens from Alabama, Wilcox Cnty., Alabama R.
at Wilcox Bar, river mi. 120.3, 7 April 1966, RDS 3860, collected by R. D. Suttkus
and G. E. Gunning; TU 40900, 289 specimens from Alabama, Wilcox Cnty.,
Alabama R. at Evans Upper Bar, river mi. 135.8, 28 June 1966, RDS 3918,
collected by R. D. Suttkus, G. E. Gunning, and G. H. Clemmer; TU 40925, 85
specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river
mi. 129.8, 28 June 1966, RDS 3919, collected by R. D. Suttkus, G. E. Gunning,
and G. H. Clemmer; TU 40940, 226 specimens from Alabama, Wilcox Cnty.,
Alabama R. at Tait Bar, river mi. 122.4, 28 June 1966, RDS 3920, collected by
R. D. Suttkus, G. E. Gunning, and G. H. Clemmer; TU 40950, 109 specimens
from Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar, river mi. 120.3, 29 June
1966, RDS 3921, collected by R. D. Suttkus, G. E. Gunning, and G. H. Clemmer;
TU 41400, 390 specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow
Jacket Bar, river mi. 129.8, 4.6 mi. E of Yellow Bluff Post Office, 1 July 1966,
RDS 3924, collected by R. D. Suttkus and Environmental Biology class; TU
41608, 284 specimens from Alabama, Wilcox Cnty., Alabama R. at Evans Upper
Bar, river mi. 135.8, 4 October 1966, RDS 401 1, collected by R. D. Suttkus and
G. E. Gunning; TU 41618, 33 specimens from Alabama, Wilcox Cnty., Alabama

40 Tulayie Studies in Zoology and Botany [Vol. 29

R. at Yellow Jacket Bar, river mi. 129.7, 4 October 1966, RDS 4012, collected
by R. D. Suttkus and G. E. Gunning; TU 41632, 92 specimens from Alabama,
Wilcox Cnty., Alabama R. at Reeves Bar, river mi. 128.5, 4 October 1966, RDS
4013, collected by R. D. Suttkus and G. E. Gunning; TU 41639, 187 specimens
from Alabama, Wilcox Cnty., Alabama R. at Tait Bar, river mi. 122.5, 4 October
1966, RDS 4014, collected by R. D. Suttkus and G. E. Gunning; TU 41655, 1 19
specimens from Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar, river mi.
120.5, 5 October 1966, RDS 4015, collected by R. D. Suttkus and G. E. Gunning;
TU 41666, 147 specimens from Alabama, Wilcox Cnty., Alabama R. at Ohio
Bar, river mi. 1 1 1.6, 5 October 1966, RDS 4016, collected by R. D. Suttkus and
G. E. Gunning; TU 41670, 301 specimens from Alabama, Monroe Cnty., Alabama
R. at Stein Island, river mi. 107.5, 5 October 1966, RDS 4017, collected by R.
D. Suttkus and G. E. Gunning; TU 41695, 83 specimens from Alabama, Wilcox
Cnty., Alabama R. at Evans Upper Bar, river mi. 135.8, 5 August 1966, RDS
3945, collected by R. D. Suttkus and G. E. Gunning; TU 41711, 29 specimens
from Alabama, Wilcox Cnty., Alabama R. at Evans Lower Bar, river mi. 133, 5
August 1966, RDS 3946, collected by R. D. Suttkus and G. E. Gunning; TU
41726, 16 specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket
Bar, river mi. 129.7, 5 August 1967, RDS 3948, collected by R. D. Suttkus and
G. E. Gunning; TU 41732, 19 specimens from Alabama, Wilcox Cnty., Alabama
R. at Reeves Bar, river mi. 128.5, 5 August 1966, RDS 3949, collected by R. D.
Suttkus and G. E. Gunning; TU 41745, 63 specimens from Alabama, Wilcox
Cnty., Alabama R. at Tait Bar, river mi. 122.5, 5 August 1966, RDS 3950,
collected by R. D. Suttkus and G. E. Gunning; TU 41755, 12 specimens from
Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar, river mi. 120.5, 5 August
1966, RDS 3951, collected by R. D. Suttkus and G. E. Gunning; TU 41761, 9
specimens from Alabama, Wilcox Cnty., Alabama R. at Ohio Bar, river mi. 1 1 1.6,
5 August 1966, RDS 3952, collected by R. D. Suttkus and G. E. Gunning; TU
41772, 151 specimens from Alabama, Monroe Cnty., Alabama R. at Stein Island,
river mi. 107.5, 5 August 1966, RDS 3953, collected by R. D. Suttkus and G. E.
Gunning; TU 41791, 22 specimens from Alabama, Monroe Cnty., Alabama R.
at St. James Bar, river mi. 104, 5 August 1966, RDS 3954, collected by R. D.
Suttkus and G. E. Gunning; TU 41797, 4 specimens from Alabama, Monroe
Cnty., Alabama R. at Bates Bar, river mi. 99, 5 August 1966, RDS 3955, collected
by R. D. Suttkus and G. E. Gunning; TU 41813, 10 specimens from Alabama,
Monroe Cnty., Alabama R. at Silver Cr. Bar, river mi. 87.6, 6 August 1966, RDS
3957, collected by R. D. Suttkus and G. E. Gunning; TU 41823, 1 specimen
from Alabama, Monroe Cnty., Alabama R. at mouth of Limestone Cr., river mi.
80.1, 6 August 1966, RDS 3958, collected by R. D. Suttkus and G. E. Gunning;
TU 42731, 370 specimens from Alabama, Wilcox Cnty., Alabama R. at Evans
Upper Bar, river mi. 135.8, 19 December 1966, RDS 4065, collected by R. D.
Suttkus and G. E. Gunning; TU 42737, 5630 specimens from Alabama, Wilcox
Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.5, 19 December 1966,
RDS 4066, collected by R. D. Suttkus and G. E. Gunning; TU 42746, 687 speci-
mens from Alabama, Wilcox Cnty., Alabama R. at Tait Bar, river mi. 122.5, 19
December 1966, RDS 4067, collected by R. D. Suttkus and G. E. Gunning; TU
42759, 900 specimens from Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar,
river mi. 120.5, 19-20 December 1966, RDS 4068, collected by R. D. Suttkus
and G. E. Gunning; TU 44011, 175 specimens from Alabama, Wilcox Cnty.,
Alabama R. at Evans Upper Bar, river mi. 135.8, 8 March 1967, RDS 4096,
collected by R. D. Suttkus and G. E. Gunning; TU 44029, 94 specimens from

1993] T^ypc Specimens of Fishes 41

Alabama, Wilcox Cnty., Alabama R. at Tail Bar, river mi. 122.5, 9 March 1967,
RDS 4098, collected by R. D. Suttkus and G. E. Gunning; TU 44045, 561 speci-
mens from Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar, river mi. 120.5,
9 March 1967, RDS 4099, collected by R. D. Suttkus and G. E. Gunning; TU
46783, 202 specimens from Alabama, Wilcox Cnty., Alabama R. at Evans Upper
Bar, river mi. 135.8, 31 May 1967, RDS 4143, collected by R. D. Suttkus and G.
E. Gunning; TU 46796, 395 specimens from Alabama, Wilcox Cnty., Alabama
R. at bar above Yellow Jacket Bar, 31 May 1967, RDS 4144, collected by R. D.
Suttkus and G. E. Gunning; TU 46802, 223 specimens from Alabama, Wilcox
Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.8, 31 May 1967, RDS
4145, collected by R. D. Suttkus and G. E. Gunning; TU 46819, 43 specimens
from Alabama, Wilcox Cnty., Alabama R. at Tait Bar, river mi. 122.5, 31 May
1967, RDS 4146, collected by R. D. Suttkus and G. E. Gunning; TU 46830, 32
specimens from Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar, river mi.
120.5, 31 May-1 June 1967, RDS 4147, collected by R. D. Suttkus and G. E.
Gunning; TU 47346, 64 specimens from Alabama, Wilcox Cnty., Alabama R. at
Evans Upper Bar, river mi. 135.7, 7 August 1967, RDS 4177, collected by R. D.
Suttkus and G. E. Gunning; TU 47361, 49 specimens from Alabama, Wilcox
Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.8, 7 August 1967, RDS
4178, collected by R. D. Suttkus and G. E. Gunning; TU 47374, 86 specimens
from Alabama, Wilcox Cnty., Alabama R. at Reeves Bar, river mi. 128.5, 7 August
1967, RDS 4179, collected by R. D. Suttkus and G. E. Gunning; TU 47387, 168
specimens from Alabama, Wilcox Cnty., Alabama R. at Tait Bar, river mi. 122.5,
7-8 August 1967, RDS 4180, collected by R. D. Suttkus and G. E. Gunning; TU
47396, 33 specimens from Alabama, Wilcox Cnty., Alabama R. at Wilcox Bar,
river mi. 120.6, 8 August 1967, RDS 4181, collected by R. D. Suttkus and G. E.
Gunning; TU 47408, 475 specimens from Alabama, Wilcox Cnty., Alabama R.
at Ohio Bar, river mi. 111.7, 8 August 1967, RDS 4182, collected by R. D.
Suttkus, G. H. Clemmer, and G. E. Gunning; TU 47420, 173 specimens from
Alabama, Monroe Cnty., Alabama R. at Stein Island, river mi. 107.5, 8 August
1967, RDS 4183, collected by R. D. Suttkus and G. E. Gunning; TU 47435, 229
specimens from Alabama, Monroe Cnty., Alabama R. at Bates Bar, river mi.
99.4, RDS 4184, collected by R. D. Suttkus and G. E. Gunning; TU 47452, 134
specimens from Alabama, Monroe Cnty., Alabama R. at Haines Island, river mi.
96, 8 August 1967, RDS 4185, collected by R. D. Suttkus and G. E. Gunning;
TU 47477, 89 specimens from Alabama, Wilcox Cnty., Alabama R. at Evans
Lower Bar, river mi. 133, 9 August 1967, RDS 4186, collected by R. D. Suttkus
and G. E. Gunning; TU 47491, 24 specimens from Alabama, Monroe Cnty.,
Alabama R. at St. James Bar, river mi. 1 04, 1 0 August 1 967, RDS 4187, collected
by R. D. Suttkus and G. E. Gunning; TU 47499, 16 specimens from Alabama,
Monroe Cnty., Alabama R. at Silver Cr. Bar, river mi. 87.7, 17 August 1967, RDS
4188, collected by R. D. Suttkus and G. H. Clemmer; TU 47515, 306 specimens
from Alabama, Wilcox Cnty., Alabama R. at Hobbs Bar, river mi. 149.5, 18
August 1967, RDS 4189, collected by R. D. Suttkus and G. H. Clemmer; TU
47762, 143 specimens from Alabama, Wilcox Cnty., Alabama R. at Lower Canton
Bar, river mi. 156.7, along left bank, 18 August 1967, RDS 4190, collected by
R. D. Suttkus and G. H. Clemmer; TU 47781, 9 specimens from Alabama, Wilcox
Cnty., Alabama R. at Lower Canton Bar, river mi. 156.7, along right bank, 18
August 1967, RDS 4191, collected by R. D. Suttkus and G. H. Clemmer; TU
47822, 85 specimens from Alabama, Wilcox Cnty., Alabama R. at Hurricane
Island, river mi. 166.5, 19 August 1967, RDS 4193, collected by R. D. Suttkus

42 Tulane Studies in Zoology and Botany [Vol. 29

and G. H. Clemmer; TU 47838, 75 specimens from Alabama, Wilcox Cnty.,
AlabamaR. atSt.JohnBar, rivermi. 164.8, 19 August 1967, RDS 4192, collected
by R. D. Suttkus and G. H. Clemmer; TU 47909, 152 specimens from Alabama
R. at Evans Upper Bar, river mi. 135.7, 26 September 1967, RDS 4204, collected
by G. E. Gunning and A. Kuris; TU 47924, 105 specimens from Alabama, Wilcox
Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.8, 26 September 1967,
RDS 4203, collected by R. D. Suttkus and G. H. Clemmer; TU 47936, 39 speci-
mens from Alabama, Wilcox Cnty., Alabama R. at Reeves Bar, river mi. 128.5,
26 September 1967, RDS 4204, collected by R. D. Suttkus and G. H. Clemmer;
TU 47968, 230 specimens from Alabama, Wilcox Cnty., Alabama R. at Tait Bar,
river mi. 122.5, 27 September 1967, RDS 4207, collected by R. D. Suttkus and
A. Kuris; TU 47980, 108 specimens from Alabama, Wilcox Cnty., Alabama R.
at Wilcox Bar, river mi. 120.6, 27 September 1967, RDS 4208, collected by R.
D. Suttkus and A. Kuris; TU 47994, 65 specimens from Alabama, Wilcox Cnty.,
Alabama R. at Ohio Bar, river mi. 1 1 1 .7, 27 September 1967, RDS 4209, collected
by R. D. Suttkus and A. Kuris; TU 48012, 232 specimens from Alabama, Monroe
Cnty., Alabama R. at Stein Island, river mi. 107.5, 27 September 1967, RDS
4210, collected by R. D. Suttkus and G. H. Clemmer.

Notropis euryzonus Suttkus, 1955: 85.

PARATYPES: TU 10700, 8 specimens from Alabama, Russell Cnty., tributary
to Little Uchee Cr., 1.1 mi. E of Crawford, U.S. Hwy. 80, 17 September 1955,
RDS 2520, collected by R. D. Suttkus; TU 10718, 60 specimens from Alabama,
Russell Cnty., tributary to Uchee Cr., 3.2 mi. W of Crawford, U.S. Hwy. 80, 17
September 1955, RDS 2521, collected by R. D. Suttkus.

Notropis gihhsi Howell & Williams, 1971: 55. (= Cyprinella gibbsi)

PARATYPES: TU 61102, 28 specimens from Alabama, Randolph Cnty.,
Wedowee Cr. on Alabama Hwy. 48, 0.5 mi. E of Wedowee, T20S, RUE, Sec.
2, 14 August 1964, collected by J. D. Williams and J. C. Hall.

Notropis hubbsi Bailey & Robison, 1978: 3.

PARATYPES: TU 13956, 7 specimens from Texas, Harrison Cnty., Caddo
Lake, 1955; TU 96238, 31 specimens from Arkansas, Calhoun Cnty., Locust
Bayou at Arkansas Hwy. 4 bridge, 0.5 mi. W of Locust Bayou, 20 May 1975,
HWR 75-47, collected by H. W. Robison and S. Pelt.

Notropis hypsilepis Suttkus & Raney, 1955b: 161.

PARATYPES: TU 8883, 4 specimens from Georgia, Fulton Cnty., Vickery
Cr. at junction with Chattahoochee R. at Roswell, U.S. Hwy. 19, 30 March 1950,
ECR 1765 and RDS 1765, collected by E. C. Raney, R. D. Suttkus, C. R. Robins
and R. Backus (removed from CU 17138); TU97111, 8 specimens from Georgia,
Habersham Cnty., Grant Cr., tributary to Soque R. near its junction with Chat-
tahoochee R. near Cornelia, June 1949, collected by J. W. Colston (removed
from CU 18102).

1993] Type Specimens of Fishes 43

Notropis imeldae Cortes,, 1966: 187.

PARATYPES: TU 55375, 5 specimens from Mexico, Oaxaca, Rio Atoyac at
Juchatengo, 16°20'N, 97°05'W, 16 April 1965, collected by I. Martinez and L.
Gonzales (removed from series collected with holotype).

Notropis mekistocholas Snehon, 1971: 451.

PARATYPES: TU 63272, 7 specimens from North Carolina, Chatham Cnty.,
Rocky R. at North Carolina Hwy. 902 bridge, 7.0 air mi. SWof center of Pittsboro,
2 June 1968, FES 68-31, collected by E. E. Snelson and W. M. Palmer (removed
from NCSM 4928).

Notropis rafinesquei Suttkus, 1 99 1 : 1 .

HOLOTYPE: TU 157264, from Mississippi, Carroll Cnty., Teoc Cr. at Mis-
sissippi Hwy. 35, R3E, T20N, Sec. 21, 12 May 1985, RDS 8758, collected by R.
D. Suttkus and V. T. Colbert.

PARATYPES: TU 140774, 309 specimens collected with holotype; TU
139610, 750 specimens from Mississippi, Carroll Cnty., Teoc Cr. at Mississippi
Hwy. 35, R3E, T20N, Sec. 21,17 February 1985, RDS 8661, collected by R. D.
Suttkus and K. G. Goodfellow; TU 141302, 50 specimens from Mississippi,
Carroll Cnty., Aiken Cr. (at gravel road), R3E, T20N, Sec. 17, 22 June 1985,
RDS 8795, collected by R. D. Suttkus; TU 150052, 97 specimens from Mississippi,
Carroll Cnty., Teoc Cr. at Mississippi Hwy. 35, T20N, R3E, Sec. 21,27 September
1987, RDS 9398, collected by R. D. Suttkus; TU 155847, 165 specimens from
Mississippi, Carroll Cnty., Teoc Cr. at Mississippi Hwy. 35, 8 August 1989, RDS
9897, collected by R. D. Suttkus; TU 158025, 346 specimens from Mississippi,
Carroll Cnty., Teoc Cr. at Mississippi Hwy. 35, 17 May 1990, RDS 10084, collected
by R. D. Suttkus.

Notropis signipinnis Bailey & Suttkus, 1952: 1.

PARATYPES: TU 1133, 164 specimens from Mississippi, George Cnty.,
Maletts Spring Branch, tributary to Rocky Cr., tributary to Escatawpa R., 6.6
mi. SE of Lucedale, 4 June 1951, RDS 2147, collected by R. D. Suttkus and C.
D. Hancock.

Notropis uranoscopus Suttkus, 1959: 7.

HOLOTYPE: TU 15289, from Alabama, Bibb Cnty., Cahaba R., 2.2 mi. N
of Centerville, 17 March 1957, RDS 2594, collected by R. D. Suttkus, R.J. Miller,
H. V. Miller, and J. DeAbate.

PARATYPES: TU 18551, 18 specimens collected with holotype; TU 18569,
2 specimens from Alabama, Bibb Cnty., Cahaba R., 2 mi. S of Centerville, 5 July
1954, University of Alabama Number 406, collected by H. H. Howell. B. Brothers,
R. Chermock, A. F. Hemphill, and H. T. Boschung; TU 18570, 8 specimens
from Alabama, Bibb Cnty., Cahaba R., 2 mi. S of Centerville, 5 July 1954,
collected by J. Mallory; TU 18674, 8 specimens from Alabama, Bibb Cnty.,
Cahaba R., 8.5 mi. N of Centerville, Hwy. 27 crossing, 8 July 1958, RDS 2738,
collected by R. D. Suttkus, M. Andersson, S. Salisbury, and J DeAbate; TU 19410,

44 Tulane Studies in Zoology and Botany [Vol. 29

21 specimens from Alabama, Bibb Cnty., Cahaba R., 8.5 mi. N of Centerville,
Hwy. 27 crossing, 8 September 1958, RDS 2760, collected by R. D. Suttkus and
M. Andersson.

Phoxinus cumberlandensis Starnes &: Starnes, 1978: 509.

HOLOTYPE: TU 103312, from Kentucky, Bell Cnty., Brownies Cr. along
Kentucky Hwy. 987 at Cubage, 14 May 1975, collected by G. W. Wolfe and J.
Lowe.

PARATYPES: TU 103313, 11 specimens from Kentucky, Bell Cnty., Brow-
nies Cr. along Kentucky Hwy. 987, 8.5 km E of Cubage, 3 April 1976, collected
by W. C. Starnes and L. B. Starnes; TU 103314, 6 specimens from Kentucky,
Whitley Cnty., Archers Cr. at Cumberland Falls road, 5.4 km W of Kentucky
Hwy. 204, 6 September 1976, collected by W. C. Starnes and L. B. Starnes; TU
103315, 1 specimen from Kentucky, Knox Cnty., Little Poplar Cr. at Cnty. Rd.
1809, 13.7 air km S of Barbourville, 24 September 1976, collected by T. Partin;
TU 103316, 2 specimens (cleared and stained) from Kentucky, Bell Cnty., Brown-
ies Cr. along Kentucky Hwy. 987, 8.5 km E of Cubage, 3 April 1976, collected
by W. C. Starnes and L. B. Starnes; TU 103317, 3 specimens (cleared and stained)
from Kentucky, Whitley Cnty., Youngs Cr. at gravel road, 0.8 km W of U.S.
Hwy. 25 W, 10.7 air km NW of Williamsburg, 6 September 1976, collected by
W. C. Starnes and L. B. Starnes.

Phoxinus tennesseensis Starnes & Jenkins, 1988: 518.

HOLOTYPE: TU 97993, from Tennessee, Blount Cnty., spring seep tribu-
tary to Reed Cr., unnumbered cnty. rd., 5 mi. E of Tennessee Hwy. 73 (Walland),
6 June 1976, collected by W. C. Starnes, L. B. Starnes, and J. A. Loudon.

PARATYPES: TU 97994, 6 specimens collected with holotype; TU (OC) 575,

4 specimens from Tennessee, Roane Cnty., tributary to Bear Cr. at Tennessee
Hwy. 95, 10 road km N of Clinch R. crossing, 15 February 1975, collected by
W. C. Starnes and L. B. Starnes (cleared and stained).

Rhinichthys osculus moapae WiWiams, 1978: 513.

HOLOTYPE: TU 103139, from Nevada, Clark Cnty., Moapa R. at Anderson
dairy farm, 1 May 1965, Nev. SoF 712, collected by J. E. Deacon.

PARATYPES: TU 103140, 3 specimens collected with holotype; TU 103141,

5 specimens from Nevada, Clark Cnty., Moapa R. at Lowhead Dam, 25 February
1967, collected by Vertebrate Zoology class.

Semotilus lumbee Snelson & Suttkus, 1978: 2.

HOLOTYPE: TU 101095, from North Carolina, Moore Cnty., fork of Aber-
deen Cr. at culvert on U.S. Hwy. 1, 0.5 air mi. SW of center of Aberdeen, 8
May 1975, WMP 75-2, collected by W. M. Palmer, A. L. Braswell, and J. E. Cooper.

PARATYPES: TU 101096, 10 specimens collected with holotype; TU 97877,
13 specimens from North Carolina, Moore Cnty., fork of Aberdeen Cr. at culvert
on U.S. Hwy. 1, 0.5 air mi. SW of center of Aberdeen, 12 May 1976, RDS 6148,
collected by R. D. Suttkus and party.

1993] T^ypc Specimens of Fishes 45

Ictaluridae

Ictalurus serracanthus Yerger 8c Relyea, 1968: 373. (= Ameiurus serracanthvs)

PARATYPES: TU 22592, 4 specimens from Florida, Gadsden Cnty., Lake
Talquin just W of mouth of Little R., 20 October 1959, JMB 78, collected by J.
M. Barkuloo and party; TU 22642, 1 specimen from Florida, Leon Cnty., Ochloc-
konee R. at Jackson Bluff, Florida Hwy. 20, 14-15 November 1959, RDS 2861,
collected by R. D. Suttkus, J. M. Barkuloo, and E. Grover; TU 37226, 3 specimens
from Florida, Wakulla Cnty., Sopchoppy R., 2.4 mi. S of Sopchoppy, U.S. Hwy.
319, 27 May 1964, collected by K. Relyea.

iVohirus a/bater Taylor, 1969: 144.

PARATYPES: TU 10180, 1 specimen from Arkansas, Newton Cnty., Big
Buffalo R., 15.8 mi. W of Jasper, Arkansas Hwy. 21, 30 April 1955, RDS 2485,
collected by R. D. Suttkus, A. Harper, W. Hildebrand, and H. D. Hoese; TU
12240, 15 specimens from Arkansas, Washington Cnty., West Fork White R.,
6.1 mi. S of Fayetteville, U.S. Hwy. 71,2 September 1955, RDS 2509, collected
by R. D. Suttkus; TU 12258, 85 specimens from Arkansas, Washington Cnty.,
White R., 8.3 mi. E of Springdale, Arkansas Hwy. 68, 3 September 1955, RDS
2512, collected by R. D. Suttkus; TU 16553, 35 specimens from Arkansas,
Washington Cnty., White R., 8.3 mi. E of Springdale, Arkansas Hwy. 68, 4
October 1957, collected by R. D. Suttkus, C. F. Cole, J. DeAbate, S. Harvey, M.
Andersson, and and B. I. Sundararaj; TU 16575, 12 specimens from Arkansas,
Washington Cnty., White R., 8.3 mi. E of Springdale, Arkansas Hwy. 68, 5
October 1957, RDS 2667, collected by R. D. Suttkus, C. F. Cole, J. DeAbate, S.
Harvey, M. Andersson, and B. I. Sundararaj; TU 22401, 7 specimens from
Arkansas, Madison Cnty., War Eagle Cr., 5.4 mi. N of Huntsville, Arkansas Hwy.
23, 25 October 1959, RDS 2856, collected by R. D. Suttkus, M. Andersson, and
B. L Sundararaj; TU 22504, 37 specimens from Arkansas, Newton Cnty., Buffalo
Fork of White R., 15.5 mi. W of Jasper, Arkansas Hwy. 74, 24 October 1959,
RDS 2853, collected by R. D. Suttkus, M. Andersson, and B. I. Sundararaj; TU
22514, 2 specimens from Arkansas, Newton Cnty., Buffalo Fork of White R.,
15.5 mi. W of Jasper, Arkansas Hwy. 74, 25 October 1959, RDS 2854, collected
by R. D. Suttkus, M. Andersson, and B. I. Sundararaj; TU 22709, 9 specimens
from Arkansas, Cleburne Cnty., Devils Fork Little Red R. between Stark and
Edgemont, 23 October 1959, RDS 2852, collected by R. D. Suttkus, M. Andersson,
and B. I. Sundararaj.

N oturus elegansT dcyXor, 1969: 150.

PARATYPES: TU 19390, 12 specimens from Kentucky, Green Cnty., Green
R., 3.8 mi. ENE of Donnansburg, Kentucky Hwy. 88, 7 September 1958, RDS
2756, collected by R. D. Suttkus and M. Andersson; TU 19460, 16 specimens
from Kentucky, Barren Cnty., tributary to Barren R., 2.6 mi. SW of Lucas, U.S.
Hwy. 3 IE, at old mill dam, 7 September 1958, RDS 2757, collected by R. D.
Suttkus and M. Andersson.

JVorurus 7achneri Taylor, 1969: 54.

PARATYPES: TU 7074, 1 specimen from Arkansas, Garland Cnty., 7.7 mi.
SSW of Owensville, 3 mi. SE of U.S. Hwy. 70, 7 August 1953, EL 94, collected

46 Tulane Studies in Zoology and Botany [Vol. 29

by E. Liner; TU 7077, 2 specimens from Arkansas, Garland Cnty., 7.7 mi. SSW
of Owensville, 3 mi. SE of U.S. Hwy. 70, 7 August 1953, EL 94, collected by E.
Liner.

Noturus tnunitus Suttkus & Taylor, 1965: 171.

HOLOTYPE: TU 26250, from Mississippi, Marion Cnty., Pearl R., 2.6 mi.
E of Sandy Hook, 14 November 1955, collected by R. D. Suttkus and K. T.
Nielsen.

PARATYPES: TU 11311, 195 specimens collected with holotype; TU 1844,
23 specimens from Mississippi, Marion Cnty., Pearl R., 2.3 mi. E of Sandy Hook,
24-25 November 1951, RDS 2207, collected by R. D. Suttkus and P. K. Anderson;
TU 3557, 8 specimens from Mississippi, Marion Cnty., Pearl R., 2.3 mi. E of
Sandy Hook, 16 February 1952, RDS 2238, collected by R. D. Suttkus, P. K.
Anderson, H. K. Suzuki, and T. Cain; TU 3627, 73 specimens from Mississippi,
Marion Cnty., Pearl R., 4 mi. NE of Sandy Hook, 13 January 1952, RDS 2217,
collected by R. D. Suttkus, P. K. Anderson, and W. D. Stone; TU 3951, 11
specimens from Mississippi, Marion Cnty., tributary to Pearl R., 4.8 mi. SE of
Columbia, Mississippi Hwy. 13, 23 May 1952, RDS 2297, collected by R. D.
Suttkus, T. Cain, and F. Sogandares-Bernal; TU 4872, 1 specimen from Missis-
sippi, Marion Cnty., tributary to Pearl R., 4.3 mi. S of Foxworth, 19 August
1952, RDS 2173, collected by R. D. Suttkus, C. D. Hancock, and J. Black; TU
7369, 1 specimen from Louisiana, Washington Parish, Pushepatapa Cr., 0.8 mi.
S of Varnado, 19 January 1952, RDS 2222, collected by R. D. Suttkus and H.
K. Suzuki; TU 8886, 4 specimens from Mississippi, Marion Cnty., tributary to
Pearl R., 0.5 mi. S of Hub or 8.5 mi. S of Columbia, 28 October 1950, RDS
2013, collected by R. D. Suttkus and C. L. Smith; TU 9709, 3 specimens from
Mississippi, Marion Cnty., tributary to Pearl R., 5 mi. S of Columbia, 30 June
1954, RDS 2456, collected by R. D. Suttkus; TU 10462, 3 specimens from
Louisiana, Washington Parish, Pushepatapa Cr., 0.8 mi. S of Varnado, 29 June
1954, RDS 2455, collected by R. D. Suttkus and K. T. Nielsen; TU 11588, 1
specimen from Louisiana, Washington Parish, Pushepatapa Cr., 8.2 mi. N of
Bogalusa, 7 May 1956, RDS 2556, collected by R. D. Suttkus and Field Zoology
class; TU 13954, 214 specimens from Mississippi, Marion Cnty., Pearl R., 2.3
mi. E of Sandy Hook, 5 October 1956, RDS 2565, collected by R. D. Suttkus, R.
J. Miller, and R. K. Chipman; TU 14930, 67 specimens from Mississippi, Marion
Cnty., Pearl R., 2.3 mi. E of Sandy Hook, 12-13 January 1957, RDS 2576,
collected by R. D. Suttkus and R. J. Miller; TU 14938, 82 specimens from
Mississippi, Marion Cnty., Pearl R., 2.6 mi. E. of Sandy Hook, 6 January 1957,
RDS 2574, collected by R. D. Suttkus and R. J. Miller; TU 15073, 2 specimens
from Louisiana, Washington Parish, Pushepatapa Cr., 1 mi. S of Varnado or 7
mi. N of Bogalusa, 13 January 1957, RDS 2578, collected by R. D. Suttkus and
R. J. Miller; TU 15171, 92 specimens from Mississippi, Marion Cnty., Pearl R.,
2.3 mi. E of Sandy Hook, 22-23 January 1957, RDS 2580, collected by R. D.
Suttkus and N. C. Negus; TU 15465, 2 specimens from Louisiana, Washington
Parish, Pushepatapa Cr., 7 mi. N of Bogalusa, Hwy. 21,21 April 1957, RDS
2600, collected by R. D. Suttkus, R. H. Gibbs, and party; TU 16148, 6 specimens
from Mississippi, Marion Cnty., Ten Mile Cr., tributary to Pearl R., 9.7 mi. N
of Sandy Hook, 7 August 1957, RDS 2641, collected by R. D. Suttkus and R.J.
Miller; TU 17472, 1 specimen from Louisiana, Washington Parish, Pushepatapa
Cr., 0.8 mi. S of Varnado, Louisiana Hwy. 21, 20 March 1958, RDS 2692,

1993] Type Specimens of Fishes 47

collected by R. D. Suttkus, M. Andersson, and S. Harvey; TU 17721, 4 specimens
from Mississippi, Simpson Cnty., Strong R., 2 mi. W of Pinola, 3 April 1958,
RDS 2697, collected by R. D. Suttkus, J. S. Suttkus, and M. Andersson; TU
17951, 1 specimen from Mississippi, Marion Cnty., Ten Mile Cr., 10.4 mi. S of
Columbia, Mississippi Hwy. 35, 3 May 1958, RDS 271 1, collected by R. D. Suttkus,
M. Andersson, and S. Simpson; TU 18739, 2 specimens from Mississippi,
Simpson Cnty., Strong R., 2 mi. W of Pinola, Mississippi Hwy. 28 (formerly Hwy.
20), 3 July 1958, RDS 2735, collected by R. D. Suttkus, M. Andersson, and J. S.
Suttkus; TU 18963, 60 specimens from Mississippi, Marion Cnty., Pearl R., 2.3
mi. E of Sandy Hook, 3 November 1957, RJM 1053, collected by R. J. Miller,
H. V. Miller, M. Andersson, and D. Boyer; TU 19781, 9 specimens from Missis-
sippi, Simpson Cnty., Strong R., 2 mi. W of Pinola, Mississippi Hwy. 20, 8 May
1959, RDS 2774, collected by R. D. Suttkus and J. S. Suttkus; TU 19825, 52
specimens from Louisiana, Washington Parish, Bogue Chitto R., 1 mi. W of
Warnerton, 2 May 1959, RDS 2770, collected by R. D. Suttkus, D. Landwehr,
and L. Redman; TU 19880, 3 specimens from Louisiana, Washington Parish,
Bogue Chitto R., 1 mi. W of Warnerton, 3 May 1959, RDS 2773, collected by
R. D. Suttkus and Vertebrate Natural History class; TU 22824, 38 specimens
from Mississippi, Marion Cnty., Pearl R., 2.3 mi. E of Sandy Hook, 26 October
1959, RDS 2857, collected by R. D. Suttkus, M. Andersson, and B. L Sun-
dararaj; TU 23127, 1 specimen from Mississippi, Marion Cnty., Pearl R., 2.3
mi. E of Sandy Hook, 24 March 1960, RDS 2897, collected by R. D. Suttkus
and G. E. Gunning; TU 23310, 1 specimen from Louisiana, Washington Parish,
Bogue Chitto R., 1 mi. S of Enon, Louisiana Hwy. 437, 3 July 1960, RDS
2913, collected by R. D. Suttkus and J. S. Ramsey; TU 23351, 1 specimen
from Louisiana, Washington Parish, Bogue Chitto R., 2 mi. W of Warnerton,
Louisiana Hwy. 438, 3 July 1960, RDS 2914, collected by R. D. Suttkus and
J. S. Ramsey; TU 23518, 24 specimens from Mississippi, Copiah Cnty., Copiah
Cr., 2.4 mi. S of Georgetown, Mississippi Hwy. 27, 22 July 1960, RDS 2935,
collected by R. D. Suttkus and J. S. Ramsey; TU 23701, 1 specimen from
Mississippi, Marion Cnty., tributary to Pearl R., 0.5 mi N of Marion Cnty.
line, 21 July 1960, RDS 2928, collected by R. D. Suttkus and J. S. Ramsey;
TU 23869, 3 specimens from Mississippi, Copiah Cnty., Copiah Cr., 2.1 mi.
S of Georgetown, Mississippi Hwy. 27, 25 November 1960, RDS 2940, collected
by R. D. Suttkus and P. Menzies; TU 26657, 87 specimens from Mississippi,
Marion Cnty., Pearl R., 2.3 mi. E of Sandy Hook, 20 October 1961, JSR 61-95,
collected by J. S. Ramsey and J. E. Thomerson; TU 26695, 1 specimen from
Mississippi, Marion Cnty., Holiday Cr., 0.5 mi. S of Goss, Mississippi Hwy.
13, 21 October 1961, JSR61-98, collected byJ.S.RamseyandJ.E. Thomerson;
TU 26759, 11 specimens from Mississippi, Lawrence Cnty., Fair R., 4.5 mi.
N of Monticello, Mississippi Hwy. 27, 3 November 1961, JSR 61-104, collected
by J. S. Ramsey and J. E. Thomerson; TU 26878, 1 specimen from Mississippi,
Marion Cnty., Silver Cr. at N limits of Morgantown, Mississippi Hwy. 587, 9
December 1961, JSR 61-116, collected by J. S. Ramsey and J. E. Thomerson;
TU 27137, 5 specimens from Mississippi, Lawrence Cnty., Pearl R., 2.5 mi. S
Oakvale, 29 August 1962, RDS 3166, collected by R. D. Suttkus and J. S. Suttkus;
TU 27172, 18 specimens from Mississippi, Marion Cnty., 1.5 mi. SE of Morgan-
town, 30 August 1962, RDS 3168, collected by R. D. Suttkus and J. S. Suttkus;
TU 27299, 12 specimens from Mississippi, Lawrence Cnty., Bahala Cr., 6 mi. S
of Rockport, Mississippi Hwy. 27, 28 October 1961, JSR 61-103, collected by J.
S. Ramsey and J. E. Thomerson; TU 28034, 87 specimens from Mississippi,

48 Tulane Studies in Zoology and Botany [Vol. 29

Marion Cnty., Pearl R., 1.5 mi. SE of Foxworth, 1 December 1962, RDS 3187,
collected by R. D. Suttkus and J. S. Suttkus; TU 28272, 47 specimens from
Mississippi, Marion Cnty., Pearl R., 1.5 mi. SE of Foxworth, 17 October 1962,
RDS 3172, collected by R. D. Suttkus and C. D. Barbour; TU 28303, 4 specimens
from Mississippi, Marion Cnty., Pearl R., 3 mi. SE of Foxworth, 18 October
1962, RDS 3173, collected by R. D. Suttkus and C. D. Barbour; TU 28316, 13
specimens from Mississippi, Marion Cnty., Pearl R. above mouth of Ten Mile
Cr., 6 mi. SE of Foxworth, 18 October 1962, RDS 3174, collected by R. D. Suttkus
and C. D. Barbour; TU 28345, 10 specimens from Mississippi, Marion Cnty.,
Pearl R., 4 mi. NNE of Sandy Hook, 0.5 mi. upstream from mouth of Hurricane
Cr., 18 October 1962, RDS 3175, collected by R. D. Suttkus and C. D. Barbour;
TU 28373, 7 specimens from Mississippi, Marion Cnty., Pearl R., 2 mi. NE of
Sandy Hook, 18 October 1962, RDS 3176, collected by R. D. Suttkus and C. D.
Barbour; TU 28391, 23 specimens from Mississippi, Marion Cnty., Pearl R., 1.5
mi. NE of Sandy Hook, 19 October 1962, RDS 3177, collected by R. D. Suttkus
and C. D. Barbour; TU 28403, 16 specimens from Mississippi, Marion Cnty.,
Pearl R., 2.3 mi. ESE of Sandy Hook, 19 October 1962, RDS 3178, collected by
R. D. Suttkus and C. D. Barbour; TU 28429, 24 specimens from Louisiana,
Washington Parish, Pearl R., 4 mi. E of Angie, 19 October 1962, RDS 3179,
collected by R. D. Suttkus and C. D. Barbour; TU 28534, 240 specimens from
Mississippi, Marion Cnty., Pearl R., 1.5 mi. SE of Foxworth, 19-20 December
1962, RDS 3188, collected by R. D. Suttkus and W. T. Mason; TU 28719, 30
specimens from Mississippi, Marion Cnty., Pearl R., 2.3 mi. E of Sandy Hook,
21 October 1961, JSR 61-97, collected by J. S. Ramsey and J. E. Thomerson;
TU 28768, 129 specimens from Mississippi, Marion Cnty., Pearl R., 2.3 mi. E
of Sandy Hook, 28 December 1962, RDS 3191, collected by R. D. Suttkus and
J. S. Ramsey; TU 28820, 14 specimens from Mississippi, Simpson Cnty., Strong
R., 2 mi. W of Pinola, Mississippi Hwy. 28, 29 December 1962, RDS 3193,
collected by R. D. Suttkus and J. S. Ramsey; TU 29942, 5 specimens from
Louisiana, Washington Parish, Pearl R. just below sill at Pools Bluff, 4 mi. S of
Bogalusa, 30-31 August 1963, collected by R. D. Suttkus and G. E. Gunning;
TU 29962, 1 specimen from Louisiana, St. Tammany Parish, Pearl R. at river
mi. 52, 0.2 mi. below mouth of McGehee Cr., 10 September 1963, RDS 3326,
collected by R. D. Suttkus, J. S. Ramsey, and G. E. Gunning; TU 29975, 1
specimen from Louisiana, St. Tammany Parish, Pearl R. at river mi. 46, 10
September 1963, RDS 3328, collected by R. D. Suttkus, J. S. Ramsey, and G. E.
Gunning; TU 29989, 1 specimen from Louisiana, St. Tammany Parish, West
Pearl R., 5 mi. SE of Tahsheek, 11 September 1963, RDS 3332, collected by R.

D. Suttkus, J. S. Ramsey, and G. E. Gunning; TU 30010, 17 specimens from
Louisiana, Washington Parish, Pearl R. just below sill at Pools Bluff, 4 mi. S of
Bogalusa, 29 September 1963, RDS 3342, collected by R. D. Suttkus, J. S. Ramsey,
and G. E. Gunning; TU 30025, 2 specimens from Louisiana, St. Tammany
Parish, Pearl R., 0.2 mi. below mouth of McGehee Cr., 10 October 1963, RDS
3343, collected by R. D. Suttkus and G. E. Gunning; TU 30040, 2 specimens
from Louisiana, Washington Parish, Pearl R. just below sill at Pools Bluff, 4 mi.
S of Bogalusa, 25 October 1963, RDS 3353, collected by R. D. Suttkus and G.

E. Gunning; TU 30149, 32 specimens from Mississippi, Simpson Cnty., Strong
R. at rapids upstream of Mississippi Hwy. 28 bridge, 2 mi. W of Pinola, 6
November 1963, RDS 3357, collected by R. D. Suttkus and J. S. Ramsey.

1993] Type Specimens of Fishes 49

Noturus phaeus Taylor, 1969: 100.

PARATYPES: TU 3037, 4 specimens from Mississippi, Lafayette Cnty , tribu-
tary to Yocona R., 5.1 mi. E of Yocona or 14.7 mi. E of Oxford, 23 May 1952,
collected by R. D. Suttkus, T. Cain, and F. Sogandares-Bernal; TU 3124, 4
specimens from Mississippi, Lafayette Cnty., tributary to Yocona R., 8 mi. E of
Oxford, Mississippi Hwy. 6, 24 May 1952 (incorrectly listed as TU 3134 in
original description); TU 3668, 3 specimens from Mississippi, Lafayette Cnty.,
tributary to Yocona R., 7.0 mi. E of Oxford, "old" Mississippi Hwy. 6, 24 May
1952, collected by R. D. Suttkus, T. Cain, and F. Sogandares-Bernal (inadvert-
antly omitted from first type list); TU 14014, 1 specimen from Mississippi,
Lafayette Cnty., Yocona R., 7.9 mi. SE of Oxford, Mississippi Hwy. 334, 12 July
1956, collected by G. H. Penn and J. B. Black; TU 14024, 1 specimen from
Mississippi, Alcorn Cnty., 3 mi. N of Biggersville, U.S. Hwy. 45, 12 July 1956,
collected by G. H. Penn and J. B. Black; TU 14036, 2 specimens from Mississippi,
Benton Cnty., 0.3 mi. W of Michigan City, Mississippi Hwy. 7, 11 July 1956,
collected by G. H. Penn and J. B. Black.

Noturus stanauli Etnier & Jenkins, 1980: 19.

HOLOTYPE: TU 110929, from Tennessee, Hancock Cnty., Clinch R. at
Frost Ford, river mi. 181.1, 11.8 air km WSW of Kyles Ford, 21 April 1978,
collected by D. L. Batch, B. H. Bauer, B. A. Branson, R. T. Bryant, D. A. Etnier,
J. L. Harris, J. A. Louton, and M. G. Ryon.

PARATYPES: TU 110930, 5 specimens collected with holotype.

Noturus stigmosus Tay\ov, 1969: 173.

PARATYPES: TU 19217, 4 specimens from Michigan, Washtenaw Cnty.,
Huron R. at North Territorial Rd., 4 mi. NW of Dexter, 31 August 1958, RDS
2750, collected by R. D. Suttkus, C. R. Gilbert, and W. Davis; TU 19246, 6
specimens from Michigan, Washtenaw Cnty., Huron R. at North Territorial Rd.,
4 mi. NW of Dexter, 3 September 1958, RDS 2752, collected by R. D. Suttkus,
M. Andersson, and T. Poulson.

Noturus taylori Douglas, 1972: 785.

PARATYPES: TU 69700, 10 specimens from Arkansas, Montgomery Cntv.,
South Fork of the Caddo R., 1 mi. SE of Hopper and 0.5 mi. S of Arkansas
Hwy. 240, 5 March 1971, collected by N. H. Douglas, D. W. Fruge, D. Heard,
and J. Lindley (missing) (removed from NLU 18098).

Prietella phreatophila Carranza, 1954: 132.

PARATYPES: TU 10810, 4 specimens from Coahuila, Mexico, "EI Potrero"
de Doha Mariana, Municipio de Muzquiz, at the base of Sierra de Santa Rosa,
29 August 1954, collected by J. Carranza and C Bolivar (formerly in the series
retained by J. Carranza at time of publication, subsequently sent to Tulane as a
gift; one specimen removed, skeletonized and recataloged as TU (OC) 178).

50 Tulane Studies in Zoology and Botany [Vol. 29

Pimelodidae

Rhamdia luigiana ViWa, 1977: 133.

PARATYPES: TU 102026, 3 specimens from Nicaragua, Depto. Granada,
Lake Nicaragua between Ometepe and Zapatera Is., 11m deep, 6 November
1976, JV 76-132, collected by J. Villa and crew of M/V Caspar (removed from
series collected with holotype).

Muraenidae

Gymnothorax kolpos Bohlke &: Bohlke, 1980: 223.

PARATYPES: TU 82455, 2 specimens from Gulf of Mexico, 28°28'N,
90°00.3'W, 49 fms., 2 April 1973, JHC 73-315, collected by crew of R/V Oregon
II cruise 44, W. W. Forman and J. H. Caruso.

Ophichthidae

Ophichthus rex Bohlke & Caruso, 1980: 239.

PARATYPES: TU 82453, 1 specimen from Gulf of Mexico, 28°28'N,
90°02'W, 45 fms., 2 April 1973, JHC 73-314, collected by crew of R/V Oregon
cruise 44, W. W. Forman and J. H. Caruso; TU 88498, 1 specimen from Gulf
of Mexico, Grand Isle Block 43, 28°59'N, 89°59'W, 96 ft., 1 March 1969, BAT
69-29, collected by B. A. Thompson; TU 88499, 1 specimen from Louisiana, St.
Bernard Parish, Gulf of Mexico, East Bay area. East Delta Block 34, 135 ft., 3
August 1969, RJI 69-73, collected by R. J. Irwin; TU 88500, 1 specimen from
Louisiana, Gulf of Mexico, 0.2 mi. W of Southwest Pass, 135 ft., 16 August 1969,
RJI 69-79, collected by R. J. Irwin; TU 88501, 1 specimen from Louisiana, Gulf
of Mexico, Grand Isle Block 43AA, 11 April 1970, JHC 70-14, collected by J.
H. Caruso and B. A. Thompson; TU 88502, 1 specimen from Gulf of Mexico,
29°17.1'N, 88°58.9'W, 12 fms., 28 March 1973, JHC 73-286, collected by J. H.
Caruso, W. W. Forman, and crew of R/V Oregon cruise 44; TU 88505, 1 specimen
from Gulf of Mexico, 28°56.5'N, 88°05'W, 36 fms., 28 March 1973, JHC 73-289,
collected by J. H. Caruso, W. W. Forman, and crew of R/V Oregon cruise 44;
TU 88506, 1 specimen from Gulf of Mexico, 28°57'N, 89°39.2'W, 31 fms., 4
April 1973, JHC 73-322, collected by J. H. Caruso, W. W. Forman, and crew of
R/V Oregon cruise 44; TU 88507, 1 specimen (skull only) from Gulf of Mexico,
29°09.2'N, 88°16.8'W, 200 fms., 8 May 1973, JHC 73-361, collected by J. H.
Caruso and crew of R/V Oregon cruise 45; TU 88508, 1 specimen from Gulf
of Mexico, 27°52.8'N, 90°53.1'W, 144-178 fms., 16 May 1973, JHC 73-286,
collected by J. H. Caruso and crew of R/V Oregon cruise 45; TU 88510, 1
specimen from Gulf of Mexico, 28°21'N, 9r28'W, 34 fms., 7 December 1973,
Oregon Sta. 14300, collected by Gutherz and crew of R/V Oregon cruise 48.

Cyprinodontidae
Cualac tessellatus Miller, 1956: 9.

PARATYPES: TU 6557, 32 specimens from San Luis Potosi, Mexico, La
Media Luna, about 7 mi. SSW of settlement of Rioverde, collected by R. T.
Gregg (removed from UMMZ 170948).

1993] Type Specimens of Fishes 51

Cyprinodon labiosus Humphries &: Miller, 1981: 55.

PARATYPES: TU 111575, 6 specimens from Mexico, Lake Chichancanab,
north end of North Pond, Yucatan, 27 July 1974, collected by J. M. Humphries
and M. M. Stevenson (removed from UMMZ 201749).

Cyprinodon maya Humphries 8c Miller, 1981: 55.

PARATYPES: TU 111576, 5 specimens from Mexico, Lake Chichancanab,
lagoon on north side of road at Esmerelda, Yucatan, 27 July 1974, collected by
J. M. Humphries and M. M. Stevenson (removed from UMMZ 201761).

Cyprinodon nazas Miller, 1976: 72.

PARATYPES: TU 30616, 26 specimens from Durango, Mexico, tributary to
Rio del Pehon de Covadonga about 2 mi. SW of Pehon Blanco, 22 June 1963,
CDB 63-10, collected by C. D. Barbour and S. Contreras.

Cyprinodon pecosensis Echelle & Echelle, 1978: 573.

HOLOTYPE: TU 97127, from New Mexico, Chaves Cnty., oxbow on Pecos
R. at Bitter Lakes Wildlife Refuge, 4 April 1972, collected by A. A. Echelle and
A. F. Echelle.

PARATYPES: TU 97128, 80 specimens collected with holotype; TU 97026,
248 specimens from Texas, Reeves Cnty., Salt Cr. at Farm Rd. 652 bridge, 16
August 1971, collected by A. A. Echelle and A. F. Echelle; TU 97030, 12 speci-
mens from New Mexico, Chaves Cnty., Salt Cr. Wilderness Area, single large
sunken pothole, 2 June 1971, collected by A. A. Echelle and A. F. Echelle; TU
97033, 30 specimens from New Mexico, Chaves Cnty., Comanche Draw, just
inside entrance to Bottomless Lakes State Park, 2 April 1972, collected by A. A.
Echelle and A. F. Echelle; TU 97037, 46 specimens from Texas, Reeves Cnty.,
Salt Cr., U.S. Hwy. 285 bridge N of Orla, 28 March 1972, collected by A. A.
Echelle and A. F. Echelle; TU 97039, 88 specimens from New Mexico, Chaves
Cnty., Figure-eight Lake, Bottomless Lakes State Park, April 1972, collected by
A. A. Echelle and A. F. Echelle; TU 97042, 81 specimens from New Mexico,
Chaves Cnty., Bitter Lakes National Wildlife Refuge, pothole no. 2, 30 October
1970, collected by A. A. Echelle and A. F. Echelle; TU 97043, 51 specimens
from New Mexico, Eddy Cnty., Pecos R. about 5 mi. SE of Malaga, 31 March
1972, collected by A. A. Echelle and A. F. Echelle; TU 97048, 86 specimens
from New Mexico, Chaves Cnty., mouth of Lost Cr., Bitter Lakes National
Wildlife Refuge, unit no. 4 near spring, 4 March 1971, collected by A. A. Echelle
and A. F. Echelle; TU 97049, 23 specimens from New Mexico, Chaves Cnty.,
Cottonwood Spring, Bitter Lakes National Wildlife Refuge, 15 August 1971,
collected by A. A. Echelle and A. F. Echelle; TU 97051, 37 specimens from New
Mexico, Chaves Cnty., Cottonwood Spring, upper end unit no. 4, Bitter Lakes
National Wildlife Refuge, 18 November 1971, collected by A. A. Echelle and A.
F. Echelle; TU 97053, 91 specimens from New Mexico, Chaves Cnty., Bitter
Lakes National Wildlife Refuge, last sinkhole with fish along road, 1 June 1971,
collected by A. A. Echelle and A. F. Echelle; TU 97056, 130 specimens from
New Mexico, Chaves Cnty., isolated pool near spring at upper end unit no. 4,
Bitter Lakes National Wildlife Refuge, 1 June 1971, collected by A. A. Echelle

52 Tulane Studies in Zoology and Botany [Vol. 29

and A. F. Echelle (incorrectly listed as TU 97057 in description); TU 97066, 41
specimens from New Mexico, Chaves Cnty., spring no. 1, upper end unit no. 4,
Bitter Lakes National Wildlife Refuge, 15 August 1971, collected by A. A. Echelle
and A. F. Echelle; TU 97070, 50 specimens from Texas, Crane/Pecos Cnty. line,
Pecos R., Farm Rd. 1053 N of Imperial, 26 March 1972, collected by A. A.
Echelle and A. F. Echelle; TU 97081, 120 specimens from New Mexico, Chaves
Cnty., Bitter Lakes National Wildlife Refuge at upper end of unit no. 4, 1 June
1971, collected by A. A. Echelle and A. F. Echelle; TU 97085, 67 specimens
from Texas, Pecos Cnty., ditch below Orient Reservoir dam, ranch road 2593,

26 March 1972, collected by A. A. Echelle and A. F. Echelle; TU 97087, 63
specimens from New Mexico, Chaves Cnty., Bottomless Lakes State Park, 28
October 1970, collected by A. A. Echelle and A. F. Echelle; TU 97092, 83
specimens from New Mexico, Chaves Cnty., Lazy Lagoon, Bottomless Lakes
State Park, April 1972, collected by A. A. Echelle and A. F. Echelle.

Cyprinodon simus Humphries & Miller, 1981: 60.

PARATYPE: TU 111577, 1 specimen from Mexico, Lake Chichancanab,
lagoon on north side of road at Esmerelda, Yucatan, 27 July 1974, collected by
J. M. Humphries and M. M. Stevenson (removed from UMMZ 201762).

Fundulus bifax Cashner &: Rogers in Cashner, et al., 1988: 675.

PARATYPES: TU 15280, 3 specimens from Alabama, Elmore Cnty., Gold
Branch, tributary to Channahatchee Cr., 6 mi. N of Eclectic, Alabama Hwy. 63,
16 March 1957, RDS 2592, collected by R. D. Suttkus, R.J. Miller, H. V. Miller,
and J. DeAbate; TU 41114, 9 specimens from Alabama, Elmore Cnty., Gold
Branch, tributary to Channahatchee R., 6 mi. N of Eclectic, Alabama Hwy. 63,
12 April 1966, RDS 3873, collected by R. D. Suttkus and G. H Clemmer.

Fundulus euryzonus Suttkus & Cashner, 1981: 2.

HOLOTYPE: TU 116631, from Louisiana, Tangipahoa Parish, East Fork
Big Cr., tributary to Tangipahoa R., at Louisiana Hwy. 1054, 4 March 1980,
RDS 7283, collected by R. D. Suttkus, R. Reynolds, and J. Reynolds.

PARATYPES: TU 116632, 44 specimens collected with holotype; TU 83182,
40 specimens from Louisiana, Tangipahoa Parish, East Fork of Big Cr. 4.7 mi.
E of Areola, Louisiana Hwy. 1054, 10 March 1972, RDS 5180, collected by R.

D. Suttkus and party; TU 106576, 22 specimens from Louisiana, Tangipahoa
Parish, East Fork Big Cr., tributary to Tangipahoa R., at Louisiana Hwy. 1054,

27 March 1978, RDS 6672, collected by R. D. Suttkus, E. C. Beckham, W. M.
Howell, and A. Black; TU 106601, 58 specimens from Louisiana, Tangipahoa
Parish, East Fork Big Cr., tributary to Tangipahoa R., at Louisiana Hwy. 1054,
6.8 roadmi. E of Areola, 27 March 1978, RDS 6673, collected by R. D. Suttkus,

E. C. Beckham, W. M. Howell, and A. Black; TU 116357, 21 specimens from
Louisiana, Tangipahoa Parish, East Fork of Big Cr. 4.7 mi. E of Areola, Louisiana
Hwy. 1054, 4 October 1974, RDS 5763, collected by R. D. Suttkus and R. Odom;
TU 116504, 22 specimens from Louisiana, Tangipahoa Parish, East Fork of Big
Cr. 4.9 mi. ENE of Areola at Louisiana Hwy. 1054 bridge, 24 February 1980,
RDS 7273, collected by R. D. Suttkus and R. C. Cashner; TU 116597, 42 speci-
mens from Louisiana, Tangipahoa Parish, East Fork of Big Cr. 4.9 mi. ENE of

1993] T^ypc Specimens of Fishes 53

Areola at Louisiana Hwy. 1054 bridge, 4 March 1980, RDS 7281, collected by
R. D. Suttkus, R. Reynolds, and J. Reynolds.

Fundulus julisia Williams & Etnier, 1982:11.

PARATYPES: TU 33495, 1 specimen from Tennessee, Coffee Cnty., tribut-
ary to tributary to Hickory Cr., tributary to Collins R., 13.1 mi. SW of
McMinnville, Tennessee Hwy. 55, 17 July 1964, JSR 64-69, collected by J. S.
Ramsey and D. W. Walton; TU 33511, 3 specimens from Tennessee, Coffee
Cnty., Little Duck R. at east limit of Manchester, Tennessee Hwy. 55, 17 July
1964, JSR 64-70, collected by J. S. Ramsey and D. W. Walton.

Xiphophorus marmorata Obregon 8c Contreras, 1988: 95.

PARATYPES: TU 157267, 2 specimens from Coahuila, Mexico, El Socavon,
5.2 km SW of Muzquiz, Coahuila, 2 June 1978, SCB 78-18, collected by S.
Contreras (removed from UANL 7275).

Caproidae

Antigonia combatia Berry & Rathjen, 1958: 255.

PARATYPES: TU 18373, 2 specimens from Atlantic Ocean, 07°3rN,
54°16'W, off Surinam, 125 fms., 8 November 1958, Oregon Sta. No. 2013,
collected by crew of M/V Oregon.

Atherinidae

Atherinella ammophila Chernoff 8c Miller, 1984: 4.

PARATYPES: TU 134878, 3 specimens from Veracruz, Mexico, Rio de La
PalmaatLaPalma, 1 January 1978, collected by W. W. Forman and M. Konikoff.

Chirostotna aculeatum Barbour, 1973: 130.

HOLOTYPE: TU 40889, from Guanajuato, Mexico, south shore of Laguna
de Yuriria about 1.5 mi. E of Yuriria, 1 August 1963, CDB 63-71, collected by
C. D. Barbour and S. Contreras.

PARATYPES: TU 40864, 11 specimens collected with holotype.

Chirostotna riojai Solorzano & Lopez, 1965b: 145.

PARATYPES: TU 40173, 2 specimens from Mexico, Mexico, Laguna de
Santiago Tilapa, 31 October 1963, LB 1820, collected by A. Solorzano, R. Cruz,
R. Aguilar, and F. Carmona.

Poblana ferdebueni Solorzano 8c Lopez, 1965a: 209.

PARATYPES: TU 38631, 3 specimens from Mexico, Puebla, Laguna de
Almoloya o Chignahuapan, 22 May 1959, LB 951, collected by A. Solorzano and
J. Obregon.

54 Tulane Studies in Zoology and Botany [Vol. 29

Serranidae

Pikea mexicanus Schultz, 1958: 323. (= Bathyanthias mexicanus)

PARATYPES: TU 11004, 6 specimens from Gulf of Mexico, 27°10'N,
96°20'W, 150 fms., 5 June 1954, Oregon Sta. 1094, collected by crew of M/V
Oregon (gift to USNM); TU 11008, 1 specimen from Gulf of Mexico, 27° lO'N,
96°20'W, 150 fms., 5 June 1954, Oregon Sta. 1094, collected by crew of M/V
Oregon; TU 12972, 11 specimens from Gulf of Mexico, 28°47'N, 85°19'W, 64
fms., 40 ft. flat trawl, 7 March 1953, Oregon Sta. 895, collected by crew of M/V
Oregon; TU 12985, 1 specimen from Gulf of Mexico, 29°19'N, 86°04'W, 82
fms., 40 ft. trawl, 31 October 1953, Oregon Sta. 864, collected by crew of M/V
Oregon; TU 14739, 1 specimen from Gulf of Mexico, 27°40'N, 95°45'W, 100
fms., 80 ft. balloon trawl, 7 May 1956, Oregon Sta. 1510, collected by crew of
M/V Oregon.

Centrarchidae

Ambloplites ariomtnus Viosca, 1936: 37.

PARATYPES: TU 2856, 14 specimens from Louisiana, St. Tammany Parish,
Little Bogue Falaya "Falia" Cr., 3 mi. N of Covington, 19 May 1935, collected
by P. Viosca and G. H. Penn (formerly in the private collection of the late Percy
Viosca; given to Tulane in 1955; specimens discolored green by copper eyelet
of original label).

Ambloplites constellatus Cashner & Suttkus, 1977: 148.

HOLOTYPE: TU 90909, from Arkansas, Marion Cnty., Buffalo R. at mouth
of Rush Cr., 5 mi. E of Arkansas Hwy. 14, 12 May 1972, RCC 72-492, collected
by R. C. Cashner and J. M. Humphries.

PARATYPES: TU 78124, 5 specimens collected with holotype; TU 43520,
2 specimens from Arkansas, Baxter Cnty., Backwater Slough, 1 mi. below mouth
of Buffalo R. on White R. side, 29 July 1965, VII-WR-65-1A, collected by R. C.
Cashner and crew; TU 44085, 8 specimens from Arkansas, Baxter Cnty., White
R., 0.5 mi. up from Buffalo City, 27 July 1965, XVII-WR-65-1B, collected by R.
C. Cashner and crew; TU 45418, 1 specimen from Arkansas, Buffalo R. (general
collection number 4), 21 July 1965, XXXI1I-WR-65-1B, collected by R. C. Cashner
and crew; TU 46559, 2 specimens from Arkansas, Searcy Cnty., Buffalo R. 1
mi. down from U.S. Hwy. 65 bridge, 19 August 1965, XXXV-WR-65-1A, collected
by R. C. Cashner and crew; TU 46611, 11 specimens from Arkansas, Marion
Cnty., Buffalo R., 1.5 mi. up from launch, 6 August 1965, XXXVIII-WR-65-3A,
collected by R. C. Cashner and crew (incorrectly listed as 6 specimens in original
description); TU 49574, 1 specimen from Arkansas, Baxter Cnty., Buffalo R. 3
mi. above its mouth near Buffalo City, 29 July 1966, VII-WR-66-15A, collected
by R. C. Cashner and crew; TU 51019, 1 specimen from Arkansas, Baxter Cnty.,
Buffalo R., just above second riffle from confluence with White R., 6 November
1965, BR-P-IL, collected by Brown and party; TU 51231, 1 specimen from
Arkansas, Baxter Cnty., 150 yards above second riffle up Buffalo R. from mouth,
right side, 5 March 1966, BR-P-IR, collected by Brown and party; TU 52445, 2
specimens from Arkansas, Baxter Cnty., just above second riffle up Buffalo R.
from its mouth, left side facing upstream, 13 August 1966, BR-P-IL, collected

1993] T^ype Specimens of Fishes

DD

by Brown and party; TU 52564, 1 specimen from Arkansas, Baxter Cnty., just
above second riffle up Buffalo R. from its mouth, left side facing upstream, 12
September 1966, BR-P-IL, collected by Brown and party.

Elassoma boehlkei Rohde & Arndt, 1987: 66.

PARATYPES: TU 144145, 10 specimens from North Carolina, Brunswick
Cnty., Juniper Cr. at North Carolina Hwy. 1340, 1.6 km E of Makatoka, 17.8
km NW of Supply, 13 February 1984, collected by F. C. Rohde.

Elassoma okatie Rohde & Arndt, 1987: 77.

PARATYPES: TU 144146, 11 specimens from South Carolina, Jasper Cnty.,
Causeway Swamp Cr. on U.S. Hwy. 17, 9.9 km N of Hardeville, 19 March 1984,
collected by F. C. Rohde, R. G. Arndt, and S. A. George.

Sparidae

Calamus nodosus Randall & Caldwell, 1966: 36.

PARATYPE: TU 37765, 1 specimen from Adantic Ocean off Georgia Coast,
33°42'N, 77°1 rW, 12-20 fms., 9 December 1959, Silver Bay Sta. 1507, collected
by crew of M/V Silver Bay.

Percidae

Ammocrypta hifascia Williams, 1975: 8.

HOLOTYPE: TU 82632, from Florida, Holmes Cnty., Choctawhatchee R.
approximately 1.5 mi. W of Pittman, Florida Hwy. 2, T5N, R16W, Sec. 9, 5 July
1973, collected by J. D. Williams.

PARATYPES: TU 82633, 25 specimens collected with holotype; TU 2278,

4 specimens from Florida, Holmes Cnty., tributary to Choctawhatchee R., 3.5
mi. NW of junction Florida Hwys. 79 and 177, 28 May 1951, RDS 2100, collected
by R. D. Suttkus and party (missing); TU 20405, 1 specimen from Florida,
Holmes Cnty., Wrights Cr., 6.4 mi. N of Bonifay, Florida Hwy. 177, 12 July
1959, RDS 2804, collected by R. D. Suttkus, D. Landwehr, and J. S. Suttkus;
TU 20816, 8 specimens from Florida, Holmes Cnty., Choctawhatchee R., 3 mi.

5 of Browns, Florida Hwy. 2, 24 July 1959, RDS 2821, collected by R. D. Suttkus,
J. M. Barkuloo, E. Grover, and K. Byrd; TU 37399, 2 specimens from Alabama,
Coffee Cnty., Cripple Cr., tributary to Pea R. 1.1 mi. N of Kinston, Alabama
Hwy. 189, 23 April 1965, RDS 3653, collected by R. D. Suttkus and F. L. Rose;
TU 46129, 2 specimens from Florida, Holmes Cnty., Wrights Cr., 3.8 mi. N of
junction of Florida Hwys. 79 and 177 on Florida Hwy. 177N, 1 May 1967, LMJ
67-47, collected by L. M.Jones and party; TU 46435, 1 specimen from Florida,
Holmes Cnty., Wrights Cr., 6.9 mi. N of junction of Florida Hwy. 79 and U.S.
Hwy. 90, on Florida Hwy. 79N, 30 April 1967, LMJ 67-46, collected by L. M.
Jones and party; TU 82174, 1 specimen from Florida, Holmes Cnty., Pittman
Cr. on Florida Hwy. 177A, 2 mi. N of Florida Hwy. 2, 17 May 1973, RCC 565,
collected by D. C. Heins, J. M. Humphries, and R. C. Cashner.

56 Tulane Studies in Zoology and Botany [Vol. 29

Ammocrypta meridana Williams, 1975: 21.

PARATYPES: TU 29137, 1 specimen from Alabama, Bibb Cnty., Cahaba R.
about 7 mi. NE of Centerville, Hwy. 27 crossing, 17-18 July 1962, RDS 3132,
collected by R. D. Suttkus and W. T. Mason; TU 29914, 7 specimens from
Alabama, Perry Cnty., Cahaba R. about 1 mi. W of Sprott, Alabama Hwy. 14 or
5 mi. E of Marion, 15 April 1963, RDS 3232, collected by R. D. Suttkus, J. S.
Ramsey, and M. D. Dahlberg; TU 30080, 1 specimen from Alabama, Perry Cnty.,
Cahaba R. about 5 mi. E of Marion, Alabama Hwy. 14 or 1 mi. W of Sprott, 2
November 1963, RDS 3355, collected by R. D. Suttkus and J. S. Ramsey; TU
32565, 2 specimens from Alabama, Monroe Cnty., Limestone Cr., tributary to
Alabama R. 3.3 mi. NW of Monroeville, Alabama Hwy. 41, 30 May 1964, RDS
3454, collected by R. D. Suttkus and J. S. Ramsey; TU 32590, 1 specimen from
Alabama, Monroe Cnty., Robinson Cr., tributary to Flat Cr. 3.9 mi. N of Tunnel
Springs, Alabama Hwy. 47, 30 May 1964, RDS 3455, collected by R. D. Suttkus
and J. S. Ramsey; TU 35115, 1 specimen from Alabama, Perry Cnty., Cahaba
R. 2 mi. SW of Sprott, 26 June 1964, RDS 3505, R. D. Suttkus and Environmental
Biology class; TU 40420, 7 specimens from Alabama, Monroe Cnty., Flat Cr.,
tributary to Alabama R., 8.7 mi. NW of Monroeville, Alabama Hwy. 41,9 April
1966, RDS 3862, collected by R. D. Suttkus and G. H. Clemmer; TU 41702, 1
specimen from Alabama, Wilcox Cnty., Alabama R. at Evans Lower Bar, river
mi. 133, 5 August 1966, RDS 3946, collected by R. D. Suttkus and G. E. Gunning;
TU 44436, 23 specimens from Alabama, Monroe Cnty., Flat Cr., 8.6 mi. NW of
Monroeville, Alabama Hwy. 41, 18 March 1967, RDS 4107, collected by R. D.
Suttkus and Ichthyology class; TU 44499, 12 specimens from Alabama, Wilcox
Cnty., Turkey Cr. 1.0 mi. S of Kimbrough, Alabama Hwy. 5, 18 March 1967,
RDS 4 1 08, collected by R. D. Suttkus and Ichthyology class; TU 52497, 1 specimen
from Alabama, Perry-Dallas Cnty. line, 4.4 mi. N of junction of Alabama Hwys.
14 and 219 on Alabama Hwy. 219, 6 June 1968, collected by S. P. Marynick and
M. C. Martin; TU 52516, 1 specimen from Alabama, Perry Cnty., Oakmulgee
Cr., 9.9 mi. N of junction of Alabama Hwys. 14 and 219 on Alabama Hwy. 219,
E 0.9 mi. on dirt road, 6 June 1968, collected by S. P. Marynick and M. C.
Martin; TU 53491, 3 specimens from Alabama, Wilcox Cnty., Alabama R. at
Evans Upper Bar, river mi. 135.7, 24 September 1968, TMB 78, collected by G.
E. Gunning and T. M. Berra; TU 58622, 1 specimen from Alabama, Wilcox
Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.7, 11 August 1969, BAT
69-84, collected by G. E. Gunning and B. A. Thompson; TU 58708, 1 specimen
from Alabama, Monroe Cnty., Alabama R. at mouth of Limestone Cr., 14 August
1969, RDS 4594, collected by R. D. Suttkus and G. E. Gunning; TU 64600, 17
specimens from Alabama, Wilcox Cnty., Alabama R. at Evans Upper Bar, river
mi. 135.7, 24 September 1970, RDS 4846, collected by R. D. Suttkus and B. A.
Thompson; TU 65063, 16 specimens from Alabama, Wilcox Cnty., Alabama R.
at Evans Upper Bar, river m.i. 135.7, 27-28 August 1970, RDS 4822, collected
by R. D. Suttkus and R. C. Cashner; TU 65080, 1 specimen from Alabama,
Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river mi. 129.7, 28 August 1970,
RDS 4824, collected by R. D. Suttkus and G. E. Gunning; TU 69108, 3 specimens
from Alabama, Bibb Cnty., Cahaba R., 8.4 mi. NNE of Centerville, Hwy. 27, 9
April 1971, RDS 4976, collected by R. D. Suttkus and party; TU 70627, 1 speci-
men from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river mi.
129.7, 1 August 1971, RDS 5051, collected by R. D. Suttkus and R. C. Cashner;
TU 70639, 1 specimen from Alabama, Wilcox Cnty., Alabama R. at Tait Bar,

1993] T^ype Specimens of Fishes 57

river mi. 122.4, 1 August 1971, RDS 5052, collected by R. D. Suttkus and R. C.
Cashner; TU 71602, 6 specimens from Alabama, Wilcox Cnty., Alabama R. at
Evans Upper Bar, river mi. 135.7, 17 September 1971, RDS 5087, collected by
R. D. Suttkus and B. A. Thompson; TU 71616, 1 specimen from Alabama,
Wilcox Cnty., Alabama R. at Evans Lower Bar, river mi. 133, 17 September 1971,
RDS 5088, collected by R. D. Suttkus and B. A. Thompson; TU 71646, 1 specimen
from Alabama, Wilcox Cnty., Alabama R. at Tait Bar, river mi. 122.4, 17 Sep-
tember 1971, RDS 5090, collected by R. D. Suttkus and B. A. Thompson; TU
78624, 1 specimen from Alabama, Monroe Cnty., Alabama R. just below Haines
Island, river mi. 94.8, 15 August 1972, RDS 5297, collected by R. D. Suttkus
and G. E. Gunning; TU 78647, 1 specimen from Alabama, Monroe Cnty.,
Alabama R. at mouth of Flat Cr., river mi. 81.5, 16 August 1972, RDS 5299,
collected by R. D. Suttkus and G. E. Gunning; TU 79017, 1 specimen from
Alabama, Wilcox Cnty., Alabama R. at Evans Lower Bar, river mi. 133, 14
September 1972, RDS 5312, collected by R. D. Suttkus and B. A. Thompson
(incorrectly listed as 790 1 6 in description) ; TU 79037, 7 specimens from Alabama,
Wilcox Cnty., Alabama R. at Evans Upper Bar, river mi. 135.7, 14 September
1972, RDS 5311, collected by R. D. Suttkus and B. A. Thompson; TU 79046, 2
specimens from Alabama, Wilcox Cnty., Alabama R. at Yellow Jacket Bar, river
mi. 129.7, 14 September 1972, RDS 5313, collected by R. D. Suttkus and B. A.
Thompson; TU 80065, 1 specimen from Alabama, Wilcox Cnty., Alabama R. at
Evans Upper Bar, river mi. 135.7, 6 December 1972, RDS 5349, collected by R.
D. Suttkus and R. C. Cashner; TU 81501, 1 specimen from Alabama, Wilcox
Cnty., Alabama R. at Tait Bar, river mi. 122.4, 22 March 1973, RDS 5381,
collected by R. D. Suttkus and party.

Etheostoma aquali Williams & Etnier, 1978: 464.

HOLOTYPE: TU 105479, from Tennessee, Bedford Cnty., FlatCr., tributary
to Duck R., at Tennessee Hwy. 64, 1 mi. SW of Shelbyville, 13 April 1967,
collected by J. D. Williams and J. G. Armstrong.

PARATYPES: TU 105480, 1 specimen taken with holotype and cataloged
as allotype; TU 105481, 3 specimens collected with holotype and cataloged as
paratypes; TU 32989, 6 specimens from Tennessee, Bedford Cnty., Duck R.
13.4 mi. NW of Tullahoma, U.S. Hwy. 41A, 17 July 1964, JSR 64-72, collected
by J. S. Ramsey and D. W. Walton (designated by Raney and Zorach (1967) and
listed in Suttkus (1970) as paratypes of Etheostoma microlepidum); TU 95729, 1
specimen from Tennessee, Humphries Cnty., Duck R. just above mouth of
Buffalo R., 10 September 1975, RDS 6024, collected by R. D. Suttkus and party.

Etheostoma blennius sequatchiense Burr, 1979: 198.

PARATYPE: TU 33461, 1 specimen from Tennessee, Sequatchie Cnty.,
Sequatchie R. 3.4 mi. S of Dunlap, U.S. Hwy. 127, 16 July 1964, JSR 64-67,
collected by J. S. Ramsey and D.W. Walton.

Etheostoma boschungi Wall & Williams, 1974: 173.

HOLOTYPE: TU 79424, from Alabama, Lauderdale Cnty., Lindsey Cr. at
junction Cnty. Rds. 15 and 6, 0.8 mi. N of Alabama Hwy. 20 at Central Heights,
TIS, R12W, Sec. 3, 22 November 1969, collected by C. Gooch and J. D. Williams.

58 Tulane Studies in Zoology and Botany [Vol. 29

PARATYPES: TU 79425, 6 specimens collected with holotype; TU 79426,

6 specimens from Alabama, Madison Cnty., Copeland Branch, tributary to Briar
Fork of FHnt R., TIS, RIW, Sec. 26, 24 October 1970, collected by B. R. Wall
and party; TU 79427, 15 specimens from Alabama, Lauderdale Cnty., Lindsey
Cr. at junction Cnty. Rds. 15 and 6, 0.8 mi. N of Alabama Hwy. 20 at Central
Heights, TIS, R12W, Sec. 3, 22 December 1971, collected by G. H. Clemmer
and J. D. Williams.

Etheostoma brevirostrum Suttkus &: Etnier, 1991: 15.

HOLOTYPE: TU 158216, from Alabama, Cleburne Cnty., Shoal Cr. at Pine
Glen Recreation Area, 10 May 1989, RDS 9801, collected by R. D. Suttkus and
J. M. Pierson.

PARATYPES: TU 154765, 6 specimens collected with holotype; TU 56167,
2 specimens from Alabama, Cleburne Cnty., Shoal Cr. at Pine Glen Recreation
Area, 8. 1 mi. N of Cleburne (U.S. Hwy. 78), 26 January 1969, RDS 4494, collected
by R. D. Suttkus and B. A. Thompson; TU 57896, 1 specimen from Alabama,
Calhoun Cnty., Shoal Cr. 2 mi. E of White Plains, 17 June 1969, RDS 4559,
collected by R. D. Suttkus and J. S. Suttkus; TU 59644, 44 specimens from
Alabama, Calhoun Cnty., Shoal Cr. tributary to Choccolocco Cr. on gravel road,
2.1 mi. E of White Plains, 14 October 1969, RDS 4641, collected by R. D. Suttkus,
G. E. Gunning, B. A. Thompson, and R. C. Cashner; TU 60351, 5 specimens
from Alabama, Cleburne Cnty., Shoal Cr. at Pine Glen Recreation Area, 8.1 mi.
N of Cleburne (U.S. Hwy. 78), 16 October 1969, RDS 4648, collected by R. D.
Suttkus, G. E. Gunning, B. A Thompson, and R. C. Cashner; TU 60687, 2
specimens from Alabama, Calhoun Cnty., Shoal Cr. at Whitesides Mill, 7 mi.
NE of Choccolocco, October 1966, collected by W. M. Howell; TU 66257, 5
specimens from Alabama, Calhoun Cnty., Shoal Cr. tributary to Choccolocco
Cr., 2.3 mi. E of White Plains on gravel road, 15 December 1970, BAT 70-33,
collected by G. E. Gunning and B. A. Thompson; TU 66556, 1 specimen from
Alabama, Calhoun Cnty., Choccolocco Cr. at Alabama Hwy. 9 bridge, 1 mi. S
of White Plains, 15 December 1970, BAT 70-332, collected by G. E. Gunning
and B. A. Thompson; TU 68585, 3 specimens from Alabama, Calhoun Cnty.,
Shoal Cr. 2.3 mi. E of White Plains, 17 March 1971, BAT 71-359, collected by
G. E. Gunning and B. A. Thompson; TU 74352, 1 specimen from Alabama,
Calhoun Cnty., Shoal Cr., tributary to Choccolocco Cr., 2.3 mi. E of White Plains,

7 December 1971, RCC 71-459, collected by R. C. Cashner and G. E. Gunning;
TU 152214, 2 specimens from Alabama, Cleburne Cnty., Shoal Cr. at Pine Glen
Recreation Area, 13 May 1988, RDS 9580, collected by R. D. Suttkus.

Etheostoma chlorobranchium Zorach, 1972: 431.

PARATYPES: TU 26212, 25 specimens from North Carolina, Swain Cnty.,
Oconaluftee R. at Ela, U.S. Hwy. 19, 19 July 1962, RDS 3138, collected by R.
D. Suttkus, J. S. Ramsey, and W. T. Mason.

Etheostoma collettei Birdsong & Knapp, 1969: 106.

PARATYPES: TU 52876, 5 specimens from Louisiana, Union Parish, Merid-
ian Cr., 1 mi. E of Conway, T22N, RIE, Sec. 17, 21 May 1956, collected by W.
R. Taylor and L. W. Lowe. (A series of 10 uncataloged paratypes, with the same
data as CU 52108, were apparently lost in transit to Tulane University).

1993] Type Specimens of Fishes 59

Etheostoma crossopterum Braasch & Mayden, 1985: 15.

PARATYPES: TU 139109, 2 specimens from Tennessee, Wilson Cnty.,
Stones Cr. at gravel road just off Interstate 40 (Mt. Juliet exit), 12 May 1973,
collected by D. A. Etnier and party (removed from UT 91.838).

Etheostoma ditrema Ramsey & Suttkus, 1965: 66.

HOLOTYPE: TU 35703, from Georgia, Chattooga Cnty., a spring flowing
into a tributary to Mills Cr., tributary to Chattooga R., 4.3 air mi. due W of
Lyerly, 18 July 1962, RDS 3134, collected by R. D. Suttkus and W. T. Mason.

PARATYPES: TU 29153, 21 specimens collected with holotype; TU 26086,

8 specimens from Georgia, Chattooga Cnty., spring tributary to Mills Cr., tribut-
ary to Chattooga R., 5.3 mi. W of Lyerly, 19 April 1962, RDS 3052, collected
by R. D. Suttkus, J. S. Ramsey, J. E. Thomerson, and C. D. Barbour; TU 27566,

9 specimens from Georgia, Chattooga Cnty., spring tributary to Mills Cr., tribut-
ary to Chattooga R., 5.3 mi. W of Lyerly, 30 May 1962, JSR 62-57, collected by
J. S. Ramsey; TU 32762, 34 specimens from Georgia, Chattooga Cnty., spring
tributary to Chattooga R. 5.3 mi. W of Lyerly, 1 June 1964, RDS 3466, collected
by R. D. Suttkus and J. S. Ramsey (inadvertendy omitted from first type list);
TU 32981, 43 specimens from Georgia, Chattooga Cnty., spring tributary to
Mills Cr., tributary to Chattooga R., 5.3 mi. W of Lyerly, 23 June 1964, RDS
3499, collected by R. D. Suttkus and Environmental Biology class.

Etheostoma etnieri Bouchard, 1977: 107.

HOLOTYPE: TU 83147, from Tennessee, White Cnty., Cherry Cr. at Ten-
nessee Hwy. 84, 18 March 1972, collected by R. W. Bouchard, J. D. Way, F. L.
Oakberg, and B. E. Oakberg.

PARATYPES: TU 83148, 1 specimen collected with holotype and cataloged
as allotype; TU 83149, 11 specimens collected with holotype and cataloged as
paratypes; TU 30316, 2 specimens from Tennessee, Van Buren Cnty., Rocky
R. 15 mi. E of McMinnville, Tennessee Hwy. 30, 11 April 1963, RDS 3220,
collected by R. D. Suttkus, J. S. Ramsey, and M. D. Dahlberg; TU 33451, 1
specimen from Tennessee, Van Buren Cnty., Caney Cr., tributary to Caney Fork
5.4 mi. E of Spencer, Tennessee Hwy. 30, 16 July 1964, JSR 64-66, collected by
J. S. Ramsey and D. W. Walton; TU 33479, 5 specimens from Tennessee, Warren
Cnty., Collins R. 5.7 mi. ESE of McMinnville, Tennessee Hwy. 8, 17 July 1964,
JSR 64-68, collected by J. S. Ramsey and D. W. Walton; TU 83150, 81 specimens
from Tennessee, White Cnty., Wildcat Cr. at Cnty. Rd. 4396 off Tennessee Hwy.
84, E of Sparta, 7 October 1972, collected by J. Dewees.

Etheostoma flavum Etnier 8c Bailey, 1989: 3.

PARATYPES: TU 30312, 66 specimens from Tennessee, Coffee Cnty., Duck
R. 2 mi. NW of Manchester, U.S. Hwy 41,11 April 1963, RDS 3219, collected
by R. D. Suttkus, J. S. Ramsey, and M. D. Dahlberg; TU 139774, 15 specimens
from Tennessee, Houston Cnty., Big Richland Cr. at unnumbered gravel road,
5.8 km N of Waverly, 1 2 September 1981, collected by D. A. Etnier and party.

Etheostoma microlepidum Raney 8c Zorach, 1967: 94.

PARATYPES: TU 19479, 19 specimens from Tennessee, Rutherford Cnty.,
East Fork of Stones R., 6 mi. N of Murfreesboro, U.S. Hwy. 231, 7 September

60 Tulane Studies in Zoology and Botany [Vol. 29

1958, RDS 2758, collected by R. D. Suttkus and M. Andersson; TU 32983, 71
specimens from Tennessee, Cheatham Cnty., Harpeth R., 8 mi. E of White Bluff,
U.S. Hwy. 70, 15 July 1964, JSR 64-60, collected by J. S. Ramsey and D. W.
Walton.

Etheostoma moorei Raney & Suttkus, 1964: 131.

PARATYPES: TU 22697, 41 specimens from Arkansas, Cleburne Cnty.,
Devils Fork, Little Red R. between Stark and Edgemont, Arkansas Hwy. 16, 23
October 1959, RDS 2852, collected by R. D. Suttkus, M. Andersson, and B. I.
Sundararaj; TU 26227, 22 specimens from Arkansas, Van Buren Cnty., Middle
Fork, Little Red R. at Shirley, 17 August 1962, RDS 3157, collected by R. D.
Suttkus and W. T. Mason; TU 26249, 21 specimens from Arkansas, Van Buren
Cnty., South Fork, Little Red R., 9 mi. SW of Clinton, Arkansas Hwy. 95, 17
August 1962, RDS 3158, collected by R. D. Suttkus and W. T. Mason.

Etheostoma neopterum Howell & Dingerkus, 1978: 14.

PARATYPES: TU 102024, 7 specimens from Alabama, Lauderdale Cnty.,
small tributary to Litde Buder Cr., about 2 mi. W of Pruitton, 3 April 1976,
GD-AL-27-76, collected by G. Dingerkus, W. M. Howell, and A. Black.

Etheostoma nigripinne Braasch 8c Mayden, 1985: 28.

PARATYPES: TU 139110, 3 specimens from Tennessee, Decatur Cnty.,
Whites Cr., 4 mi. N of Bath Springs, 4 April 1978, collected by M. E. Braasch
(removed from KU 20895).

Etheostoma nuchale Howell & Caldwell, 1965: lOL

PARATYPES: TU 34591, 20 specimens from Alabama, Jefferson Cnty., Glen
Spring at Bessemer, T19S, R4W, Sec. 17, NE 1/4, along Cnty. Hwy. 20, 9 Sep-
tember 1 964, RDS 3582, collected by R. D. Suttkus, J. S. Ramsey, and F. L. Rose.

Etheostoma olivaceum Braasch & Page, 1979: 2.

PARATYPES: TU 101391, 6 specimens from Tennessee, Putnam Cnty., Rock
Springs Branch, 2 km N of Buffalo Valley, 11 December 1976, collected by B.
M. Burr and L. M. Page (removed from INHS 75576).

Etheostoma pyrrhogaster Bailey & Etnier, 1988: 31.

PARATYPES: TU 139772, 4 specimens from Tennessee, Henry Cnty., Clear
Cr., tributary to North Fork Obion R., at unnumbered cnty. rd., 1.7 air mi. S
of Cnty. Rd. 8172, 3.2 air mi. SW of Puryear, 27 April 1972, collected by J. E.
Deck, W. C. Dickinson, and J. Vaughn.

Etheostoma radiosum cyanorum Moore & Rigney, 1952: 10.

PARATYPES: TU 10564, 9 specimens from Oklahoma, Johnston Cnty., Blue
R. at bridge, Oklahoma Hwy. 99, 14 June 1959, collected by G. A. Moore (for-
merly OAM (OSUS) 2913).

1993] Type Specimens of Fishes 61

Etheostoma rubrum Raney &: Suttkus, 1966: 95.

PARATYPES: TU 30171, 47 specimens from Mississippi, Copiah Cnty.,
Bayou Pierre, 8.6 mi. SW of Utica, Mississippi Hwy. 18, 7 December 1963, JSR
63-58, collected by J. S. Ramsey and M. D. Dahlberg; TU 31341, 2 specimens
from Mississippi, Hinds Cnty., White Oak Cr., tributary to Bayou Pierre, 2.7 mi.
S of Utica, Mississippi Hwy. 18, 1 February 1964, RDS 3378, collected by E. C.
Raney and R. D. Suttkus; TU 32264, 1 1 specimens from Mississippi, Copiah
Cnty., Bayou Pierre 18.7 mi. ENE of Port Gibson, Hwy. 18, 17 May 1964, RDS
3435, collected by R. D. Suttkus and J. S. Ramsey (inadvertently omitted from
Suttkus (1970)); TU 37305, 4 specimens from Mississippi, Copiah Cnty., Bayou
Pierre, 8.6 mi. SW of Utica, Mississippi Hwy. 18, 18 March 1965, RDS 3643,
collected by R. D. Suttkus and L. Ogren; TU 37451, 53 specimens from Missis-
sippi, Copiah Cnty., Bayou Pierre, 8.6 mi. SW of Utica, Mississippi Hwy. 18, 28
April 1965, RDS 3661, collected by R. D. Suttkus and F. L. Rose; TU 40368, 10
specimens from Mississippi, Copiah Cnty., Bayou Pierre, 8.6 mi. SW of Utica,
Mississippi Hwy. 18, 2 April 1966, RDS 3855, collected by R. D. Suttkus and G.
H. Clemmer.

Etheostoma smithi Page & Braasch, 1976: 2.

PARATYPES: TU 95964, 13 specimens from Kentucky, Livingston Cnty.,
Ferguson Cr., 3 km E of Smithland, 27 May 1971, collected by L. J. Stannard
and L. M. Page.

Etheostoma striatulum Page & Braasch, 1977: 1.

PARATYPES: TU 97493, 10 specimens from Tennessee, Bedford Cnty.,
Wartrace Cr., 1 mi. E of Bell Buckle, 7 November 1975, collected by L. M. Page
and M. E. Braasch (removed from INHS 75037).

Etheostoma tallapoosae Suttkus 8c Etnier, 1991: 3.

HOLOTYPE: TU 158215, from Alabama, Elmore Cnty., Gold Branch, tribut-
ary to Channahatchee Cr., 6 mi. N of Eclectic, Alabama Hwy. 63, 16 March
1957, RDS 2592, collected by R. D. Suttkus, R. J. Miller, H. V. Miller, and J.
DeAbate.

PARATYPES: TU 15275, 92 specimens collected with holotype; TU 12084,
4 specimens from Alabama, Elmore Cnty., Gold Branch, 6 mi. NE of Eclectic,
Alabama Hwy. 63, 5 October 1955, RDS 2525, collected by R. D. Suttkus; TU
29891, 30 specimens from Alabama, Clay Cnty., Enitachope Cr., tributary to
Hillabee Cr., 3.2 mi. SW of Ashland, Hwy. 9, 21 September 1963, JSR 63-46,
collected by J. S. Ramsey; TU 32737, 21 specimens from Alabama, Clay Cnty.,
Enitachope Cr. 2.9 mi. S of Ashland, Alabama Hwy. 9. 1 June 1964, RDS 3462,
collected by R. D. Suttkus and J. S. Ramsey; TU 40656, 57 specimens from
Alabama, Clay Cnty., Enitachope Cr., tributary to Hillabee Cr., 2.9 mi. SW of
Ashland, Alabama Hwy. 9, 13 April 1966, RDS 3877, collected by R. D. Suttkus
and G. E. Gunning; TU41116, 45 specimens from Alabama, Elmore Cnty., Gold
Branch, tributary to Channahatchee Cr., 6 mi. N of Eclectic, Alabama Hwy. 63,
12 April 1966, RDS 3873, collected by R. D. Suttkus and G. H. Clemmer; TU
41137, 3 specimens from Alabama, Elmore Cnty., Channahatchee Cr., tributary

62 Tulane Studies in Zoology and Botany [Vol. 29

to Tallapoosa R., 2.5 mi. S of Red Hill, Alabama Hwy. 229, 12 April 1966, RDS
3874, collected by R. D. Suttkus and G. H. Clemmer; TU 41160, 12 specimens
from Alabama, Clay Cnty., Hillabee Cr., tributary to Tallapoosa R., 1 mi. SW
of Millerville, Alabama Hwy. 9, 13 April 1966, RDS 3876, collected by R. D.
Suttkus and G. H. Clemmer; TU 152230, 7 specimens from Alabama, Elmore
Cnty., Gold Branch, 0.9 mi. NW of Jordon, Alabama Hwy. 63, 14 May 1988,
RDS 9581, collected by R. D. Suttkus; TU 152243, 15 specimens from Alabama,
Elmore Cnty., Gold Branch, 1.7 mi. SE of Jordon, T20N, R21E, Sec. 29, 14 May
1988, RDS 9582, collected by R. D. Suttkus; TU 157616, 1 specimen from
Alabama, Tallapoosa Cnty., Josie Leg Cr., 3.9 mi. SW of Perryville, Hwy. 22, 23
March 1990, RDS 10050, collected by R. D. Suttkus and J. M. Pierson; TU
157633, 11 specimens from Alabama, Elmore Cnty., Gold Branch, 6 mi. N of
Eclectic, Alabama Hwy. 63, 23 March 1990, RDS 10051, collected by R. D. Suttkus
and J. M. Pierson; TU 157646, 18 specimens from Alabama, Elmore Cnty., Gold
Branch at Cnty. Rd. 357, 1.7 mi. SE of Alabama Hwy. 63 at Jordon, 23 March
1990, RDS 10052, collected by R. D. Suttkus and J. M. Pierson; TU 157781, 2
specimens from Alabama, Clay Cnty., Enitachope Cr., 2.8 mi. SSW of Ashland,
Alabama Hwy. 9, 12 April 1990, RDS 10063, collected by R. D. Suttkus and J.
M. Pierson; TU 157793, 19 specimens from Alabama, Elmore Cnty., Gold Branch
1.8 mi. E of Alabama Hwy. 63, 12 April 1990, RDS 10064, collected by R. D.
Suttkus and J. M. Pierson; TU 157803, 4 specimens from Alabama, Elmore
Cnty., Gold Branch, 6 mi. N of Eclectic, Alabama Hwy. 63, 12 April 1990, RDS
10065, collected by R. D. Suttkus and J. M. Pierson; TU 158249, 8 specimens
from Alabama, Coosa Cnty., Oakachoy Cr. at Alabama Hwy. 259, T21N, R20E,
Sec. 2, 9 August 1988, MP 88-79, collected byj. M. Pierson and C. Stephenson.

Etheostoma wapiti Etnier &: Williams, 1989: 989.

PARATYPES: TU 30271, 1 specimen from Tennessee, Lincoln Cnty., Elk
R. at Fayetteville, 0.25 mi. below U.S. Hwy. 231, 11 April 1963, RDS 3217,
collected by R. D. Suttkus, J. S. Ramsey, and M. D. Dahlberg (missing); TU
148010, 2 specimens from Tennessee, Giles Cnty., Richland Cr. at Cnty. Rd.
4209 crossing, 1 1 .2 air mi. SSE of Pulaski, 7 June 1 982, collected by D. A. Etnier.

Etheostoma zonistium Bailey & Etnier, 1988: 39.

PARATYPES: TU 139773, 7 specimens from Tennessee, Hardin Cnty.,
Walker Branch, tributary to Mud Cr. at unnumbered cnty. rd., 3.1 air mi. SSW
of Savannah, 16 May 1978, collected by D. A. Etnier and party.

Percina antesella Williams & Etnier, 1977: 7.

HOLOTYPE: TU 94031, from Tennessee, Bradley Cnty., Conasauga R. at
Tennessee Hwy. 75, 12.9 mi. SE of Cleveland, 9-10 April 1970, collected by D.
A. Etnier and party.

PARATYPES: TU 58938, 7 specimens from Tennessee, Bradley Cnty., Con-
asauga R. at Tennessee Hwy. 75, 12.9 mi. SE of Cleveland, 17 October 1969,
RDS 4657, collected by R. D. Suttkus, B. A. Thompson, and R. C. Cashner; TU
58968, 3 specimens from Tennessee, Bradley Cnty., Conasauga R. at Tennessee
Hwy. 75, 12.9 mi. SE of Cleveland, 19 October 1969, RDS 4660, collected by R.
D. Suttkus, B. A. Thompson, and R. C. Cashner; TU 59427, 7 specimens from

1993] Type Specimens of Fishes 63

Georgia, Murray Cnty., Conasauga R. 2.3 mi. W of junction Georgia Hwys. 225
and 286 on Georgia Hwy. 286, 19 October 1969, RDS 4661, collected by R. D.
Suttkus, B. A. Thompson, and R. C. Cashner: TU 65937, 7 specimens from
Tennessee, Bradley Cnty., Conasauga R. at Tennessee Hwy. 75, 12.9 mi. SE of
Cleveland, 29 June 1970, RDS 4777, collected by R. D. Suttkus and R. C. Cashner;
TU 69143, 8 specimens from Tennessee, Bradley Cnty., Conasauga R. at Ten-
nessee Hwy. 75, 12.9 mi. SE of Cleveland, 15 April 1971, RDS 4989, collected
by R. D. Suttkus and R. C. Cashner; TU 78368, 1 specimen from Tennessee,
Bradley Cnty., Conasauga R. at Tennessee Hwy. 75, 12.9 mi. SE of Cleveland,
1 July 1972, BAT 72-451, collected by B. A. Thompson, D. A. Etnier, and
University of Tennessee graduate students.

Percina aurolineata Suttkus 8c Ramsey, 1967: 129.

HOLOTYPE: TU 39573, from Georgia, Gilmer Cnty., Coosawattee R. about
200 yards below the mouth of a small spring tributary, 4.2 mi. SW of the center
of Ellijay, 18 June 1966, RDS 3910, collected by R. D. Suttkus and Environmental
Biology class.

PARATYPES: TU 41049, 3 specimens collected with holotype; TU 38320,
5 specimens from Georgia, Gilmer Cnty., Cartecay R., tributary to Coosawattee
R., 2.9 mi. SE of Ellijay, along Georgia Hwy. 52, 18 June 1965, RDS 3695,
collected by R. D. Suttkus and Environmental Biology class; TU 38507, 15 speci-
mens from Georgia, Gilmer Cnty., Coosawattee R., 4.2 mi. SW of Ellijay, 22
June 1965, RDS 3707, collected by R. D. Suttkus and Environmental Biology
class; TU 40714, 4 specimens from Georgia, Gilmer Cnty., tributary to Cartecay
R., 2.9 mi. SE of Ellijay, along Georgia Hwy. 52, 15 April 1966, RDS 3881,
collected by R. D. Suttkus and G. H. Clemmer; TU 41053, 6 specimens from
Georgia, Gilmer Cnty., Cartecay R., 3.1 mi. SE of Ellijay, along Georgia Hwy.
52, 17 June 1966, RDS 3909, collected by R. D. Suttkus, G. H. Clemmer, and
Environmental Biology class.

Percina caprodes fulvitaenia Morris & Page, 1981: 96.

PARATYPES: TU 110587, 2 specimens from Missouri, Osage Cnty., Maries
R., 2 km S of Westphalia, 1 1 October 1969, collected by P. W. Smith and L. M.
Page (removed from I NHS 75634).

Percina gymnocephala Beckham, 1980: 1.

HOLOTYPE: TU 106911, from North Carolina, Ashe Cnty., South Fork of
New R. at E crossing of Cnty. Rd. 1181, 7.0 air mi. SE of West Jefferson, 22
September 1977, ECB 344, collected by E. C. Beckham and E. B. Beckham.

PARATYPES: TU 105137, 3 specimens from North Carolina, Ashe Cnty.,
South Fork of New R. at E crossing of Cnty. Rd. 1181, 7.0 air mi. SE of West
Jefferson, 22 September 1977, ECB 344, collected by E. C. Beckham and E. B.
Beckham; TU 70059, 3 specimens from Virginia, Grayson Cnty., Fox Cr. along
Cnty. Rd. 711, 1.4 mi. above Fox, junction Cnty. Rd. 711 and U.S. Hwy. 58, 13
June 1971, BAT 71-379, collected by B. A. Thompson and K. Thompson; TU
70102, 12 specimens from Virginia, Grayson Cnty., Fox Cr. at U.S. Hwy. 58
bridge, 3.0 mi. NE of [town of] Mouth Of Wilson, 12 June 1971, BAT 71-377,

64 Tulane Studies in Zoology and Botany [Vol. 29

collected by B. A. Thompson and K. Thompson; TU 71144, 3 specimens from
Virginia, Grayson Cnty., Fox Cr. along Cnty. Rd. 711, 1.2 mi. NW of Fox (U.S.
Hwy. 58), 13 June 1971, BAT 71-381, collected by B. A. Thompson and K.
Thompson; TU 105084, 2 specimens from Virginia, Grayson Cnty., Fox Cr. at
U.S. Hwy. 58, 3 mi. E of [town of] Mouth Of Wilson, 20 September 1977, ECB
339, collected by E. C. Beckham and E. B. Beckham; TU 105124, 1 specimen
from North Carolina, Ashe Cnty., South Fork of New R. at North Carolina Hwy.
163, 7.5 air mi. SE of West Jefferson, 22 September 1977, ECB 343, collected
by E. C. Beckham and E. B. Beckham; TU 106912, 2 specimens from North
Carolina, Ashe Cnty., South Fork of New R. at Cnty. Rd. 1105 bridge, 0.1 mi.
S of Cnty. Rd. 1106, 24 March 1978, HLB 18, collected by H. L. Bart, R.
Holzenthal, J. M. Grady, and G. Laiche.

Perciiiajeii/cinsi Thompson, 1985: 4.

HOLOTYPE: TU 106075, from Tennessee, Bradley Cnty., Conasauga R. at
Tennessee Hwy. 74 bridge, 12.6 mi. SE of Cleveland, 1 July 1972, BAT 72-451,
collected by B. A. Thompson, D. A. Etnier, and University of Tennessee graduate
students.

PARATYPES: TU 78370, 2 specimens collected with holotype (incorrecdy
listed as TU 78369 in original description); TU 69140, 6 specimens from Ten-
nessee, Bradley Cnty., Conasauga R., 12.6 mi. SE of Cleveland, Tennessee Hwy.
74, 15 April 1971, RDS 4989, collected by R. D. Suttkus and R. C. Cashner; TU
80506, 1 specimen from Tennessee, Bradley Cnty., Conasauga R., 12.9 mi. SE
of Cleveland, Tennessee Hwy. 74, 29 June 1970, RDS 4777, collected by R. D.
Suttkus and R. C. Cashner; TU 80507, 6 specimens from Tennessee, Bradley
Cnty., Conasauga R., 12.7 mi. SE of Cleveland, Tennessee Hwy. 74, 17 October
1969, RDS 4657, collected by R. D. Suttkus, B. A. Thompson, and R. C. Cashner;
TU 80508, 8 specimens from Tennessee, Bradley Cnty., Conasauga R., 12.7 mi.
SSE of Cleveland, Tennessee Hwy. 74, 19 October 1969, RDS 4660, collected
by R. D. Suttkus, B. A. Thompson, and R. C. Cashner.

Percina lenticula Richards 8c Knapp, 1964: 695.

PARATYPES: TU 15291, 7 specimens from Alabama, Bibb Cnty., Cahaba
R., 2.2 mi. N of Centerville,just off Alabama Hwy. 5, 17 May 1957, RDS 2594,
collected by R. D. Suttkus, R. J. Miller, H. V. Miller, and J. DeAbate.

Percina tanasi Etnier, 1976: 473.

HOLOTYPE: TU 90858, from Tennessee, Loudon Cnty., Little Tennessee
R. at shoal area out from mouth of Coytee Springs, 6 November 1973, BAT
73-564, collected by B. A. Thompson, D. A. Etnier, W. C. Starnes, G. F. Boronow,
B. F. Clark, and R .L. Hensen.

PARATYPES: TU 83994, 2 specimens collected with holotype; TU 90859,
1 specimen from Tennessee, Loudon Cnty., Little Tennessee R. at shoal area
out from mouth of Coytee Springs, 6 November 1973, BAT 73-564, collected
by B. A. Thompson, D. A. Etnier, W. C. Starnes, G. F. Boronow, B. F. Clark,
and R. L. Hensen.

1993] Type Specimens of Fishes 65

Poecilichthys whipplii radiosus Hubbs Be Black, 1941 : 10. (elevated to Etheostoma
radiosum by Moore and Rigney, 1952)

PARATYPES: TU 52871, 1 specimen from Arkansas, Garland Cnty., North
Fork, Ouachita R., 2.5 mi. W of Buckville, 1 June 1939, collected by T. Holder;
TU 52872, 5 specimens from Arkansas, Garland Cnty., Cataract Branch, 2.75
mi. N of Cedar Glade, 2 May 1 939, collected by T. Holder; TU 52873, 2 specimens
from Arkansas, Garland Cnty., Big Branch, 2.25 mi. W of Buckville, 16 May
1939, collected by T. Holder. (These eight paratypes apparently had been re-
tained at the University of Arkansas by John D. Black subsequent to the descrip-
tion. During 1967, a large part of the University of Arkansas fish collection was
given to Tulane University and thus the transfer of the type material).

Lutjanidae

Symphysanodon berryi Anderson, 1970: 338.

PARATYPES: TU 57443, 2 specimens from Atlantic Ocean, 20°45'N,
73°33'W, 217-200 fms., 24 May 1965, Oregon Sta. 5414, collected by crew of
M/V Oregon (removed from series of 43 specimens from same station as
holotype).

Cichlidae

Cichlasoma pantostictum Taylor & Miller, 1983: 15.

PARATYPES: TU 5640, 3 specimens from Tamaulipas, Mexico, Laguna de
Chairel at the mouth of Rio Tamesi at Tampico, 29 December 1952, RMD X-23,
collected by J. H. Darnell and E. Liner.

Clinidae

Starksia y-Iineata Gilbert, 1965: 2.

PARATYPES: TU 36401, 2 specimens from Caribbean Sea, Grand Cayman
Island, "Ironshore" formation in front of Seaview Lodge, S side of Georgetown,
depth 2-12 ft., collected by C. R. Gilbert and J. C. Tyler.

Microdesmidae
Microdesmus cam* Gilbert, 1966: 326.

PARATYPES: TU 36400, 2 specimens from Atlantic Ocean, Costa Rica,
Limon Province, E side of Tortuguero Lagoon, near mouth, 7 August 1964,
collected by D. P. Kelso (removed from UF 1 1297).

Microdesmus suttkusi Gilbert, 1966: 329.

HOLOTYPE: TU 25110, from Costa Rica, Puntarenas Province. Boca de
Rio Baru at Dominical, 28 January 1961, RDS 2961, collected by R. D. Suitkus,
A. Smalley, and party.

PARATYPES: TU 25130, 1 specimen from Costa Rica, Puntarenas Province,
Rio Baru, 1 km N of Dominical, 28 January 1961, RDS 2962, collected by R. D.

66 Tulane Studies in Zoology and Botany [Vol. 29

Suttkus, A. Smalley, and party; TU 25262, 1 specimen from Costa Rica, Pun-
tarenas Province, at La Barca, 27 February 1960, collected by S. Jimenez-C.

Fierasferidae ( = Carapidae)

Echiodon dawsoni Williams 8c Shipp, 1982: 846.

PARATYPES: TU 122841, 2 specimens from Gulf of Mexico, 29°39.8'N,
87°16.2'W, 75.6 m, 7 February 1978, collectors unknown (removed from USAIC
6006).

Draconettidae

Draconetta oregona Briggs & Berry, 1959: 129.

PARATYPES: TU 18831, 3 specimens from Adantic Ocean, 02°04'N,
47°00'W, 125 fms., 17 November 1957, Oregon Sta. 2080, collected by crew of
M/V Oregon (incorrectly listed as TU 188831 in description) (removed from
series collected with holotype).

Gobiidae

Gobiosoma evelynae Bohlke &: Robins, 1968: 103.

PARATYPES: TU 48782, 2 specimens from Adantic Ocean, Virgin Islands,
Anguilla Island, Bowlings Shoal, W of Crocus Bay off N end of Sand Island,
17 July 1965, collected by J. C. Tyler and W. N. Eschmeyer (formerly ANSP
110549).

Gobiosoma genie Bohlke & Robins, 1968: 109.

PARATYPES: TU 48781, 1 specimen from Atlantic ocean, Bahama Islands,
Hog Island, north shore (station 122), 15 March 1953, collected by C. C. G.
Chaplin.

Triglidae

Bellator ribeiroiMWXer, 1965: 259.

PARATYPES: TU 37821, 2 specimens from Adantic Ocean off Colombia,
08°50.5'N, 76°53.5'W, 37-40 fms., 27 May 1964, Oregon Sta. 4899, collected by
crew of M/V Oregon.

Peristedion greyae Miller, 1967: 27.

PARATYPES: TU 33302, 2 specimens from Atlantic Ocean off North
Carohna, 34°15'N, 75°54'W, 190-200 fms., 9 June 1962, Silver Bay Sta. 4159,
collected by crew of M/V Silver Bay.

Cottidae
Cottus carolinae infernatis Williams &: Robins, 1970: 371.

PARATYPES: TU 32497, 5 specimens from Alabama, Clarke Cnty., Big

1993] Type Specimens of Fishes 67

Reedy Cr., tributary to Alabama R., 7 mi. NE of Carlton, 28 May 1964, RDS
3450, collected by R. D. Suttkus and J. S. Ramsey.

Cottus p7g7naeus Williams, 1968: 334.

PARATYPES: TU 34396, 47 specimens from Alabama, Calhoun Cnty., Cold-
water Cr., just below Coldwater Springs at town of Coldwater, 31 August 1964,
RDS 3571, collected by R. D. Suttkus and J. S. Ramsey.

Tetraodontidae

Sphoeroides parvus Shipp &: Yerger, 1969: 478.

PARATYPES: TU 9381, 1 specimen from Louisiana, St. Tammany Parish,
Lake Ponchartrain, 2 mi. W of south drawbridge of Louisville and Nashville
Railroad, 30°10'N, 89°55'W, 5 November 1954, M/V Anna Inez Station 547,
collected by R. D. Suttkus and party; TU 19038, 2 specimens from Louisiana,
Cameron Parish, at jetties along west bank of Calcasieu R. at Cameron, 28 April
1957, RES 7, collected by R. E. Smith and M. Guidry; TU 22573, 14 specimens
from Louisiana, Jefferson Parish, Gulf of Mexico, 100 yards off Grand Terre
Island near Grand Isle, 12 December 1959, collected by D. Landwehr and R.
D. Lumsden.

Batrachoididae

Porichthys bathoiketes Gilbert, 1968: 710.

PARATYPES: TU 18572, 9 specimens from Atlantic Ocean, 1 r27'N,
83°11'W, 135 fms., 9 September 1957, Oregon Sta. 1902, collected by crew of
M/V Oregon; TU 18573, 1 specimen from Atlantic Ocean, 16°07'N, 81°05'W,
130 fms., 24 August 1957, Oregon Sta. 1891, collected by crew of M/V Oregon;
TU 18574, 3 specimens from Atlantic Ocean, 16°38'N, 81°39'W, 150 fms., 22
August 1957, Oregon Sta. 1879, collected by crew of M/V Oregon; TU 19928,
2 specimens from Atlantic Ocean, 16°39'N, 81°43'W, 125 fms., 22 August 1957,
Oregon Sta. 1878, collected by crew of M/V Oregon.

Lophiidae

Lophiodes bruchius Caruso, 1981: 539.

PARATYPES: TU 94331, 3 specimens from Hawaii, Pacific Ocean,
2r03.5'N, 156°29.rW, 274-318 m, 25 November 1968, TC 40-92, collected by
crew of R/V Townsend Cromwell; TU 94332, 2 specimens from Hawaii, Pacific
Ocean, 21°04.5'N, 156°27.5'W, 337-340 m, 22 November 1968, TC 40-74, col-
lected by crew of R/V Townsend Cromwell.

Lophiodes reticulatus Caruso & Suttkus, 1979: 491.

PARATYPES: TU 2733, 1 specimen from Gulf of Mexico, 29°49'N, 85°45'W,
1 12 fms., 24 February 1950, Oregon Sta. 278, collected by crew of M/V Oregon;
TU 90215, 1 specimen from Gulf of Mexico, 29°15.7'N, 88°09'W, 70 fms., 3
May 1973, JHC 73-344, collected by crew of R/V Oregon II cruise 45 and J. H.
Caruso; TU 94027, 3 specimens from Gulf of Mexico, 28°47'N, 85°19'\V, 64
fms., 7 March 1953, Oregon Sta. 895, collected by crew of M/V Oregon.

68 Tulane Studies in Zoology and Botany [Vol. 29

Chaunacidae

Chaunax suttkusi Caruso, 1989: 160.

PARATYPES: TU 2811, 2 specimens from Gulf of Mexico, 29°20'N, 87°25'W,
315 fms., 28 April 1951, Oregon Sta. 319, collected by crew of M/V Oregon;
TU 10947, 5 specimens from Gulf of Mexico, 24°20'N, 83°24'W, 180 fms., 13
April 1954, Oregon Sta. 1007, collected by crew of M/V Oregon; TU 12660, 12
specimens from Gulf of Mexico, 26°24'N, 83°45'W, 210 fms., 19 June 1956,
Oregon Sta. 1556, collected by crew of M/V Oregon; TU 35837, 6 specimens
from Gulf of Mexico, 24°28'N, 83°27'W, 205 fms., 6 August 1963, Oregon Sta.
4371, collected by crew of M/V Oregon and M. D. Dahlberg; TU 90284, 1
specimen from Gulf of Mexico, 29°16.rN, 87°38'W, 250 fms., 4 May 1973, JHC
73-348, collected by crew of R/V Oregon II cruise 45 and J. H. Caruso; TU
90324, 2 specimens from Gulf of Mexico, 29°14.5'N, 87°44'W, 270 fms., 5 May
1973, JHC 73-351, collected by crew of R/V Oregon II cruise 45 and J. H.
Caruso; TU 90381, 1 specimen from Gulf of Mexico, 29°01.8'N, 88°14.9'W, 400
fms., 6 May 1973, JHC 73-356, collected by crew of R/V Oregon II cruise 45
and J. H. Caruso; TU 90461, 2 specimens from Gulf of Mexico, 24°28.7'N,
83°32.5'W, 200fms.,21 May 1973, JHC 73-371, collected by crew of R/V Oregon
II cruise 45 and J. H. Caruso; TU 90473, 1 specimen from Gulf of Mexico,
24°23.4'N, 83°24.1'W, 220 fms., 21 May 1973, JHC 73-372, collected by crew
of R/V Oregon II cruise 45 and J. H. Caruso.

Acknowledgments

We thank Royal D. Suttkus for valuable information used in preparing the
type list, Jan D. Hunter for assistance with the manuscript, and John Caruso
and an anonymous reviewer for comments that improved the manuscript.

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Suttkus, R. D. and H. T. Bosc:hung. 1990. Notropis ammophilus, a new cyprinid iish Ironi south-
eastern United States. Tulane Stud. Zool. Bot. 27: 49-63.
Suttkus, R. D. and R. C. Cashner. 1981. A new species of cyprinodoniid fisli, genus Fimdiilus

(Zygonectes), from Lake Pontchartrain tributaries in Louisiana and Mississippi. Bull. .Alabama

Mus. Nat. Hist. 6: 1-17.
Suttkus, R. D. and G. H. Clemmer. 1968. Notropis edwardraneyi. a new cyprinid fish from the

Alabama and Tombigbee River systems and a discussion of related species. Fulane Stud. Zool.

Bot. 15: 18-39.
Suttkus, R. D. and D. A. Etnier. 1991. Etheostoma tallapoosae and E. hrei'irostrum, two new darters,

subgenus Ulocentra, from the Alabama River drainage. lulane Stud. Zool. Bot. 28: 1-24.

72 Tulane Studies in Zoology and Botany

SuTTKUS, R. D. and J. S. Ramsey. 1967. Percina aurolineata, a new percid fish from the Alabama

River system and a discussion of ecology, distribution, and hybridization of darters of the subgenus

Hadropterus. Tulane Stud. Zool. 13; 129-145.
SuTTKUS, R. D. and E. C. Raney. 1955a. Notropis baileyi, a new cyprinid fish from the Pascagoula

and Mobile Bay drainages of Mississippi and Alabama. Tulane Stud. Zool. 2: 69-86.
SuTTKUS, R. D. and E. C. Raney. 1955b. Notropis hypsilepis, a new cyprinid fish from the Apalachicola

River system of Georgia and Alabama. Tulane Stud. Zool. 2: 159-170.
SuTTKUS, R. D. and E. C. Raney. 1955c. Notropis asperifrons, a new cyprinid fish from the Mobile

Bay drainage of Alabama and Georgia, with studies of related species. Tulane Stud. Zool. 3: 1-33.
SuTTKUS, R. D. and W. R. Taylor. 1965. Noturus munitus, a new species of madtom, family

Ictaluridae, from southern United States. Proc. Biol. Soc. Washington 78: 169-178.
Taylor, J. N. and R. R. Miller. 1983. Cichlid fishes (genus Cichlasoma) of the Rio Ranuco basin,

eastern Mexico, with description of a new species. Occ. Pap. Mus. Nat. Hist. Univ. Kansas 104: 1-24.
Taylor, W. R. 1969. A revision of the catfish genus Noturus Rafinesque with an analysis of higher

groups in the Ictaluridae. Bull. Nat. Mus. 282: 1-315.
Thomerson, J. E. and S. H. Weitzman. 1970. A new species of glandulocaudine characid fish,

Hysteronotus myersi, from Peru. Proc. California Acad Sci., 4th Series, December 31, 1970.
Thompson, B. A. 1985. Percina jenkinsi, a new species of logperch (Pisces: Percidae) from the

Conasauga River, Tennessee and Georgia. Occ. Pap. Mus. Zool. Louisiana State Univ. 61: 1-23.
Villa, J. 1977. A new species of pimelodid catfish of the genus Rhamdia from Nicaragua, Central

America. Brenesia 12/13: 133-142.
Viosca, P., Jr. 1936. A new rock bass from Louisiana and Mississippi. Copeia 1936: 37-45.
Wall, B. R., Jr. and J. D. Williams. 1974. Etheostoma boschungi, a new percid fish from the

Tennessee River drainage in northern Alabama and western Tennessee. Tulane Stud. Zool. Bot.

18: 172-182.
Williams, J. D. 1968. A new species of sculpin, Cottus pygmaeus, from a spring in the Alabama River

basin. Copeia 1968: 334-342.
Williams, J. D. 1975. Systematicsof the percid fishes of the subgenus Ammocrypta, genus Ammocrypta,

with descriptions of two new species. Bull. Alabama Mus. Nat. Hist. 1: 1-56.
Williams, J. D. and G. H. Clemmer. 1991. Scaphirhynchus suttkusi, a new sturgeon (Pisces: Acipen-

seridae) from the Mobile basin of Alabama and Mississippi. Bull. Alabama Mus. Nat. Hist. 10:

17-31.
Williams, J. D. and D. A. Etnier. 1977. Percina (Imostoma) antesella, a new percid fish from the

Coosa River system in Tennessee and Georgia. Proc. Biol. Soc. Washington 90: 6-18.
Williams, J. D. and D. A. Etnier. 1978. Etheostoma aquali, a new percid fish (subgenus Nothonotus)

from the Duck and Buffalo rivers, Tennessee. Proc. Biol. Soc. Washington 91: 463-471.
Williams, J. D. and D. A. Etnier. 1982. Description of a new species, Eundulus julisia, with a

redescription of Eundulus albolineatus and a diagnosis of the subgenus Xenisma (Teleostei: Cyp-

rinodontidae). Occ. Pap. Mus. Nat. Hist. Univ. Kansas 102: 1-20.
Williams, J. D. and C. R. Robins. 1970. Variation in populations of the fish Cottus carolinae in the

Alabama River system with description of a new subspecies from below the Fall Line. Amer.

Midi. Nat. 83: 368-381.
Williams, J. E. 1978. Taxonomic status of Rhinichthys osculus (Cyprinidae) in the Moapa River,

Nevada. Southwest. Nat. 23: 511-518.
Williams, J. E. and C. E. Bond. 1980. Gila boraxobius, a new species of cyprinid fish from southeast-
ern Oregon with a comparison to G. alvordensis Hubbs and Miller. Proc. Biol. Soc. Washington

93: 291-298.
Williams, J. E. and C. E. Bond. 1981. A new subspecies of tui chub (Osteichthyes: Cyprinidae)

from Guano Basin, Nevada and Oregon. Southwest. Nat. 26: 223-230.
Williams, J. T. and R. L. Shipp. 1982. A new species of the genus Echiodon (Pisces: Carapidae)

from the eastern Gulf of Mexico. Copeia 1982: 845-851.
Yerger, R. and K. Relyea. 1968. The flat headed bullheads (Pisces: Ictaluridae) of the southeastern

United States, and a new species oi Ictalurus from the Gulf Coast. Copeia 1968: 361-384.
Zorach, T. 1972. Systematics of the percid fishes, Etheostoma camurum and E. chlorobranchium new

species, with a discussion of the subgenus Nothonotus. Copeia 1972: 427-447.

TYPE SPECIMENS OF VASCULAR PLANTS AT TULANE UNIVERSITY,
WITH A BRIEF HISTORY OF THE TULANE HERBARIUM

Anne S. Bradburn and Steven P. Darwin

Department uj Ecology, Evoluiion, and Organismal Biology,
Tulane University, New Orleans, LA 70118

The Tulane University Herbarium is one of the principal herbaria of the
Gulf South region, and was designated a National Resource Collection by the
American Society of Plant Taxonomists in 1974. The University holds specimens
of fungi, algae, bryophytes, and vascular plants, all housed in Dinwiddle Hall
on the main campus in New Orleans. The collections of algae are maintained
primarily for teaching purposes. The bryological collections (ca. 2500 specimens)
are mainly of Gulf Coast mosses and liverworts representing valuable historical
collections that are heavily annotated. The mycological herbarium includes
roughly 1 1,000 fungal specimens, most of them collected from land areas around
the Gulf of Mexico and the Caribbean Sea. The vascular plant herbarium pre-
sently includes 110,000 specimens of worldwide representation, but is strongest
in the flora of the southeastern United States, southern California, the Rocky
Mountain region of Colorado, and northern Latin America, especially the Yuca-
tan Peninsula of Mexico. Important special collections include the type specimens
listed in this report, early plant collections from the lower Mississippi River
drainage, Arizona and California ferns of J. G. Lemmon, and North American
Gramineae and Cyperaceae of Asa Gray. Recent accessions include collections
made by Tulane faculty and students, and exchange material regularly received
from some thirty institutions worldwide. Herbarium loans support a wide variety
of monographic, floristic, ecological, and ethnobotanical studies. Research in
systematic and taxonomic botany is further supported by the Minna F. Koch
Library of Botany, a collection of more than 1000 titles housed adjacent to the
herbarium and rich in historical botany and plant illustration, as well as worldwide
floras, monographs, dictionaries, and essential reference works.

Historical Sketch

The University Herbarium was established after the close of the New Orleans
World's Industrial and Cotton Centennial Exposition of 1884-1885. Initially, all
plant collections were part of the University Museum, which also had custody
of a small herbarium belonging to the New Orleans Academy of Sciences founded
in 1 853. The Academy's herbarium includes some of the oldest existing Louisiana
plant collections, these mosdy gathered by early faculty of the Medical College
of Louisiana (now Tulane University) in the 1840's. The most significant collec-
tors who contributed to the Academy's and later the University's herbarium are
given below.

Josiah Hale, M.D. (1791-1856) was a Virginia native who settled near Alexan-
dria, Louisiana, in 1825, after taking a medical degree at Transylvania College,
there receiving botanical training from Rafmesque. He retired from medical
practice in 1834, devoting full energy to botany. Torrey and Gray, in their Flora
of North America (1838), frequendy cite specimens obtained from Hale. Among

73

74 Tulane Studies in Zoology and Botany [Vol. 29

his other correspondents were Riddell, Engelmann, Darlington, Leavenworth,
Eaton, and Durand. Hale served as first President of the New Orleans Academy
of Sciences, and the Hale specimens at Tulane, about one hundred, are from
the Academy's herbarium (J. Ewan,J. Soc. Bibliogr. Nat. Hist. 8: 235-243. 1977).

John Leonard Riddell, M.D. (1807-1865), physician, scientist, and intellectual,
arrived in New Orleans in 1836 as Professor of Chemistry at the Medical College
of Louisiana. He subsequently held various professorships, including Pharmacy
and Materia Medica. In 1839 he was appointed Melter and Refiner at the United
States Branch Mint in New Orleans, and in 1860 Federal Postmaster, a position
he held throughout the Civil War period. Riddell's interests extended to botany,
mathematics, and invention, including an early typewriting machine, and the
first practical binocular dissecting microscope. He was a charter member of the
American Association for the Advancement of Science, and the New Orleans
Academy ofSciences(K.Riess, Tulane Stud. Geol.Paleontol. 13: 1-110. 1977).

Riddell began accumulating a herbarium in 1830 while at Marietta, Ohio,
and contributed his first botanical paper in 1832. He had taken the bachelor's
degree in 1829 at Rensselaer School with Amos Eaton, and yet favored the
taxonomic system of Jussieu over that of Linnaeus. He experimented with new
methods of plant collecting, drying, and preservation; he exchanged specimens
with Rafinesque and offered herbaria for sale. Riddell corresponded with the
leading North American botanists, including Torrey, Engelmann, and Short.
His paper "A synopsis of the flora of the western states" appeared in 1835,
followed by "A supplementary catalogue of Ohio plants" in 1836. The "Catalogus
florae ludovicianae," abridged from a larger manuscript, appeared in 1852.

Riddell's specimens are widely distributed among North American and Euro-
pean herbaria; his personal herbarium, much damaged and depleted, was ac-
quired by Charles Mohr, and is now in the U. S. National Herbarium, Smithsonian
Institution. The Riddell specimens held at Tulane, mostly from Ohio and
Louisiana, and numbering about 125, were part of the New Orleans Academy
of Sciences herbarium, and frequently bear specimen labels in Riddell's hand.

William Marbury Carpenter, M.D. (1811-1848) was born in West Feliciana
Parish, Louisiana. He attended West Point and served as Professor of Natural
Sciences at Centenary College before joining the faculty of the Medical College
of Louisiana in New Orleans as Professor of Materia Medica. In 1846, Carpenter
and Charles Lyell made geological excursions in Louisiana, especially studying
the Mississippi River delta. Carpenter's herbarium, except for specimens sent to
Torrey and others, passed to Riddell, who donated a few dozen to the New
Orleans Academy of Sciences (R. S. Cocks, Tulane Graduates' Mag. 3: 122-127.
1914).

Joseph Finleyjoor, M.D. (1848-1892) was a native of Louisiana. He practiced
medicine in New Orleans and Thibodaux, and then in Harrisburg and Birdston,
Texas, before becoming Assistant Commissioner for Texas to prepare a botanical
exhibit for the 1884-85 Cotton Centennial Exposition at New Orleans. In 1888
he assumed charge of Tulane University's Museum, first as Assistant Curator,
and then Curator and Professor of Botany. A few hundred Joor specimens were
incorporated into the New Orleans Academy of Sciences herbarium, of which
he was also curator. His personal herbarium, which was not large, passed to the
Missouri Botanical Garden in 1897. Tulane's Joor specimens, largely undupli-
cated at Missouri, are particularly rich in southern grasses, many specimens
annotated by George Vasey (J. B. S. Norton, Bot. Gaz. 26: 271-274. 1898).

1993] Type Specimens of Vascular Plants 75

Reginald W. S. Cocks, M.A. (1863-1926) was a native of Worcestershire,
England. He took the master's degree at Trinity College, (Cambridge, with first
honors in classics. He was for a year Professor of Botany at Louisiana State
University before moving in 1908 to assume the newly created Ida A. Richardson
Chair of Botany at Tulane University. At about that time, Cocks began a corre-
spondence with Charles Sprague Sargent, first Director of the Arnold Arboretum
of Harvard University. In Cocks, Sargent found an able botanist and field com-
panion on whom he could rely for data concerning the woody flora of the (iulf
States. From Cocks's specimens, Sargent eventually recognized five new species
or varieties of Carya, as well as new species of Quercus, Crataegus, and Tilia.

Cocks may have started plant collecting as early as 1892, but in 1898 he
initiated the first of many annual explorations that would, over a period of
twenty-five years, take him to every part of Louisiana. The result was a sizeable
herbarium and series of published inventories. For a few species. Cocks specimens
are still the only Louisiana record. Many of his specimens carry identifications
communicated by Sargent in a regular correspondence amounting to 302 letters
now preserved in the University's archive (J. Ewan, J. Arnold Arbor. 46: 1-44,
122-159, 324-361, 411-439. 1965).

Cocks was replaced on the Tulane faculty by William T. Penfound, Ph.D.
(1897-1984). Although primarily a plant ecologist, Penfound and his graduate
students gathered a number of Louisiana plant specimens, especially from local
wetland habitats where ecological studies were being conducted. Those specimens
now represent wetland vegetation existing before urban expansion in southern
Louisiana, and voucher the numerous ecological studies published by Penfound
and his co-workers.

When Joseph Ewan, A.B. (1909-) came to Tulane in 1947, he brought with
him a personal herbarium of some 32,000 specimens, probably three or four
times as many specimens as were already present in the University's collection.
The Ewan herbarium is largely responsible for the wide geographical coverage
of the University's present collection, as well as many of its type specimens. In
addition to his own collections, mostly from southern California, the Rocky
Mountain region, and South America, Ewan's herbarium also included specimens
gathered by L. M. Booth (southern California), I. Clokey (Nevada), D. Keck
{Penstemon), J. G. Lemmon (California and Arizona ferns), F. W. Pierson
(Cahfornia), Y. Mexia (Latin America), and the Gray Herbarium exsiccatae of
the Fernald period. Specimens oi Delphinium and Vismia, Ewan's own taxonomic
specialties, are also well represented. The Tulane herbarium benefitted from
Ewan's interest in botanical history and bibliography; a number of specimens
were acquired that had been collected by well-known exploring expeditions of
the nineteenth century, these mosdy duplicates from European herbaria, particu-
larly the British Museum (Natural History) and the Conservatoire et Jardin
Botaniques, Geneva. A nearly complete set of Asa Gray's North American
Gramineae and Cyperaceae was also added. All collections were accommodated
in modern steel cases, and curated by Nesta Dunn Ewan, who worked as a
volunteer for thirty years.

Since 1977, the present curators have attempted to enlarge the holdings of
Louisiana plants, and acquire material from Mexico, particularly the Yucatan
Peninsula, where studies are being carried out by Tulane faculty and students.
To that end, a number of new exchange agreements were established with Latin
American botanical institutions. Increased emphasis has also been placed upon

76 Tulane Studies in Zoology and Botany [Vol. 29

cultivated plants in New Orleans. Present staff includes Steven P. Darwin, Ph.D.,
Curator and Director (taxonomy and phylogeny of angiosperms, tropical
Rubiaceae, flora of Louisiana); Anne S. Bradburn, M.S., Assistant Curator (bar-
rier island vegetation of the Gulf Coast, flora of Yucatan, ethnobotany); Arthur
L. Welden, Ph.D., Curator of Fungi (neotropical fungi, especially
Thelephoraceae); and Joseph A. Ewan, A.B., Sc.D., Curator Emeritus (bibliog-
raphy of American natural history).

Type Collections

The following list documents 148 vascular plant specimens in the Tulane
University Herbarium (NO) known to be holotypes, isotypes, lectotypes, or isolec-
totypes of validly published names of species or infraspecific taxa. The list is
alphabetical by family and then species, beginning with pteridophytes. For each
name typified by a Tulane specimen, we provide reference to the original place
of publication, locality information as it appears on the specimen label, and the
location of other type specimens (e.g., holotype or other isotypes), if that infor-
mation is given by the publishing author. We also cite current taxonomic place-
ment of species names when known.

PTERIDOPHYTA

ASPLENIACEAE

Asplenium curtissii Underwood, Bull. Torrey Bot. Club 33: 194. 1906.

Curtiss 3728. United States of America. Florida: Citrus County at head of
Lake Tsala Apopka. April. Holotype: NY; isotype: NO.

BLECHNACEAE

Blechnum mexiae Copeland, Univ. Calif. Publ. Bot. 17: 32, pi 7. 1932.

YnesMexia 4237. Brazil. Minas Geraes: Districto Carangola, about 12 km along
trail from Areponga to Fazenda de Grama, elevation 970 meters. January
27, 1930. Holotype: UC 419577; isotype: NO.

= Struthiopteris mexiae (Copeland) Ching, Sunyatsenia 5: 243. 1940.

CYATHEACEAE

Cyathea albidopaleata Copeland, Univ. Calif. Publ. Bot. 17: 25, pi 2. 1932.

Ynes Mexia 4868. Brazil. Minas Geraes: Vigosa, Agricultural College, Corrida
Reberro; elevation 675 meters. July 18, 1930. Holotype: UC 466100;
isotype: NO.

Cyathea anacampta Alston, J. Wash. Acad. Sci. 48: 230. 1958.

Ewan 16801. Colombia. Putumayo: Cordillera Oriental, east slope, Rio
Ticuanayoy at confluence of Rio Caqueta, elevation 1100 meters. January
20, 1945. Holotype: BM; isotype: NO.

1993] Type Specimens of Vascular Plants 77

Cyathea catacampta Alston, J. Wash. Acad. Sci. 48: 231. 1958.

Ewan 16153. Colombia. Narino: Between Rio Miraflores and Rio San Martin,
Volcan de Cumbal region, elevation 2680 meters. September 12-13, 1944.
Holotype: BM; isotype: NO.

Cyathea ewanii Alston, J. Wash. Acad. Sci. 48: 231. 1958.

Ewan 16729. Colombia. Putumayo: Cordillera Oriental, east slope, between
Mocoa and Urcusique, en route to Umbria, elevation 900 meters. January
9, 1945. Holotype: BM; isotypes: NO, US.

Cyathea leucolepismata Alston, J. Wash. Acad. Sci. 48: 231. 1958.

Ewan 16784. Colombia. Putumayo: Cordillera Oriental, east slope, near San
Diego de Colorado, and tributary of Rio Putumayo, between Umbria and
Puerto Asis. January 16, 1945. Holotype: BM; isotypes: NO, US.

Cyathea mexiae Copeland, Univ. Calif. Publ. Bot. 17: 30, pi. 4. 1932.

YnesMexia 4882. Brazil. Minas Geraes: Vigosa, Agricultural College, hill south-
east of main buildings, elevation 700 meters. July 21, 1930. Holotype: US
466093; isotype: NO.

DRYOPTERIDACEAE

Dryopteris arguta (Kaulf.) Watt, forma bella Ewan, Amer. Fern J. 34: 1 12. 1944.

Fosberg8127. United States of America. California: Los Angeles County, Santa
Catalina Island, Big Wash Canyon, Avalon. May 6, 1932. Holotype: COLO;
isotype: NO.

Dryopteris arguta forma flocculosa Ewan, Amer. Fern J. 34: 1 12. 1944.

Grant, Ballou ^ Ewan 7908. United States of America. California: Santa Bar-
bara County, San Miguelito Creek, 3 miles above Lompoc. May 30, 1933.
Holotype: COLO; isotype: NO.

Dryopteris arguta forma nudata Ewan, Amer. Fern J. 34: 111. 1944.

Campbell 755. United States of America. California: Mariposa, Merced River
below Yosemite, elevation 3800 feet. June 14, 1929. Holotype: COLO;
isotype: NO.

Dryopteris normalis C. Christensen var. lindheimeri A. Br. ex C. Christensen,
Kongel. Danske Vidensk.-Selsk. Skr. VIL 10(2): 182. 1912.

F. Lindheimer 742. United States of America. Texas. 1847. Holotype: B;
isotypes: NO, W.

= Dryopteris normalis C. Christensen; D. S. Correll in C. Lundell (ed.). Flora
of Texas 1: 106. 1942.

Polystichum munitum subsp. curtum Ewan, Amer. Fern J. 33: 99. 1942.

Ewan 5159. United States of America. California: San Bernardino, Sawpit
Canyon, San Bernardino Mountains, elevation 4200 feet. September 7,
1931. Holotype: NO; isotypes: DS, US.

78 Tulane Studies in Zoology and Botany [Vol. 29

Woodsia obtusa Torr. var. nana Lemmon, Fern Bull. 15: 126. 1908.

Lemmon s.7i. United States of America. Arizona: Deer Park on summit of Santa
Catalina Mountains. (?)1880. Isotype: NO.

= Woodsia pusilla Fourn. var. glandulosa (Eaton 8c Faxon) T. Taylor, Amer.
Fern J. 37: 86. 1947.

ISOETACEAE

Isoetes hieroglyphica A. A. Eaton, Fernwort Papers 10. 1900.

Pringle s.n. United States of America. Maine: St. Francis Lakes. July 31, 1880.
Holotype: Herbarium of A. A. Eaton; isotype: NO.

=Isoetes macrospora Durieu forma hieroglyphica (A. A. Eaton) Pfeiffer, Ann.
Missouri Bot. Card. 9: 194, p/. 19. 1922.

POLYPODIACEAE

Microgramma acatallela Alston, J. Wash. Acad. Sci. 48: 232. 1958.

Ewan 16706. Colombia. Putumayo: Cordillera Oriental (east slope), Rio Mocoa
drainage, 3 km above Mocoa, along trail to San Antonio, elevation 600
meters. January 8, 1945. Holotype: BM; isotypes: NO, US.

PTERIDACEAE

Eriosorus ewanii A. F. Tryon, Contr. Gray Herb. 200: 165. 1970.

Ewan 16316. Colombia. Naririo: north slope of Volcan de El Galeras above
Pasto, elevation 3500 meters. October 18, 1944. Holotype: GH; isotypes:
NO, S, US.

Notholaena californica Eaton subsp. nigrescens Ewan, Amer. Fern J. 32: 93,

tab. 10, fig b. 1942.

Wheeler 878. United States of America. California: Los Angeles County, San
Gabriel Mountains, 6 miles from mouth of San Gabriel Canyon, elevation
1000 feet. June 26, 1932. Holotype: COLO; isotypes: NO, LAM.

=Aleuritopteris cretacea subsp. nigrescens (Ewan) Munz, Aliso 4: 87. 1958.

Pomatophytum pocillatum M. E.Jones, Contr. W. Bot. 16: 12. 1930.

Jones 24690. United States of America. Arizona: Ramsey Canyon, Huachuca
Mountains. September 28, 1929. Isotype: NO.

= Cheilanthes lendigera (Cav.) Sw.; R. & A. Tryon, Ferns Allied PI. 249. 1982.

SCHIZAEACEAE

Anemia tripinnata Copeland, Univ. Calif. Publ. Bot. 17: 24, pi. 1. 1932.

Ynes Mexia 5627 . Brazil. Minas Geraes: Corinto, near Fazenda do Diamante,
base of Serra de Anjico, elevation 600 meters. April 15, 1931. Holotype:
UC 466099; isotype: NO.

1993] Type Specimens of Vascular Plants 79

SPERMATOPHYTA

ACANTHACEAE

Dicliptera ewanii Leonard, Contr. U. S. Nad. Herb. 31: 356, /?^. 131. 1958.

Ewan 15628. Colombia. Cundinamarca: Cordillera Oriental, 7 km east of Hilo,
valley of QuebradaChiniata, elevation 2430 meters. May 1, 1944. Holotype:
NO; isotypes: K, US.

Pseuderanthemum ewanii Leonard, Contr. U. S. Nad. Herb. 31: 302, fig. 111.
1953.

Ewan 15629. Colombia. Cundinamarca: Cordillera Oriental, valley of Queb-
rada Chiniata, 7 km east of Hilo, elevation 2430 meters. May 1, 1944.
Holotype: NO.

AGAVACEAE

Pleomele hawaiiensis O. & L Degener, Fl. Hawaiiensis, Fam. 68. 1980 [Publ.
privately].

Degener ^ Ordonez 12550. United States of America. Hawaii: Makawao, Maui.
August 3, 1939. Isotypes: NO(2).

ANNONACEAE

Malmea macrocarpa R. E. Fries, Ark. Bot. (n.s.) 4: 23, tab. 1. 1957.

Asplund 17590. Ecuador. Province of Guayas: vicinity of Guayaquil, Cerro
Azul forest. September 10, 1955. Holotype: S; isotype: NO.

APIACEAE (UMBELLIFERAE)

Sanicula odorata (Raf.) Pryer & Phillippe, Canad. J. Bot. 67: 703. 1989.

Darwin &' Sundell 1379. United States of America. Mississippi: Wilkinson
County, vicinity of Pinckneyville. April 28, 1979. Neotype: NO.

ARACEAE

Peltandra tharpii Barkley, Madrono 7(5): 131, lab. 21. 1944.

Barkley 13424. United States of America. Texas: Robertson County, near
Newbaden. August 8, 1943. Holotype: TEX; isotype: NO.

ARALIACEAE

Schefflera sachamatensis Cuatrecasas, Revista Acad. Colomb. Ci. Exact. 8: 315.
1951.

Ewan 16678. Colombia. Putumayo: Cordillera Oriental (east slope), along trail
from San Francisco to Sachamate, upper Rio Mocoa, 3 km above Sachamate,
elevation 1700 meters. January 6, 1945. Holotype: NO.

80 Tulane Studies in Zoology and Botany [Vol. 29

ASTERACEAE (COMPOSITAE)

Antennaria plantaginifolia (L.) Richards var. monocephala Torrey & A. Gray,
Fl. North Amer. 2: 431. 1843.

Carpenter s.n. United States of America. Louisiana: Felicianas. Syntype: NO.

=Antennaria solitaria Rydberg, Bull. Torrey Bot. Club 24: 304. 1897.

Antennaria soliceps Blake, Proc. Biol. Soc. Wash. 51:7. 1938.

Clokey ^ Bean 7459. United States of America. Nevada: Clark County, Char-
leston Mountains, elevation 3350 meters. July 18, 1937. Holotype: US
1699460; isotype: NO.

Bacc/iaris pedersenii Cabrera, Darwiniana 16(1-2): 410. 1970.

Pedersen 2642. Argentina. Mburucuya: Province Corrientes at Estancia "Santa
Teresa." March 18, 1954. Isotype: NO.

Bidens heterodoxa (Fernald) Fernald &c St. John var. interstes Fassett, Rhodora
26: 178. 1924.

Fassett 852. United States of America. Maine: Kennebec County, Kennebec
River, Gardiner. September 18, 1923. Holotype: GH; isotype: NO.

=Bidens eatonii Fernald var. interstes (Fassett) Fassett, Rhodora 27: 142. 1925.

Crepis nana Richards subsp. ramosa Babcock, Univ. Calif. Publ. Bot. 22: 542,
fig. 155. 1947.

Thompson 7883. United States of America. Washington: Jefferson County,
above Lake Constance, elevation 6000 feet. August 11, 1931. Holotype:
UC 470750; isotypes: NO, DS, MO, Blake herbarium.

Eupatorium cyrilli-nelsonii A. Molina, Ceiba 22: 39. 1978.

Cirilo Nelson 3912. Honduras. Comayagua: Villa de Taulabe; quebrada La
Caliche, elevation 600 meters. February 19, 1977. Holotype: TEFH;
isotypes: NO, US 2851248.

=Peteravenia cyrilli-nelsonii (A. Molina) King & Robinson, Phytologia 44: 86.
1979.

Grindelia camporum Greene var. interioris (Jepson) Steyermark forma foliacea
Steyermark, Ann. Missouri Bot. Card. 21: 538. 1934.

Howell 5201. United States of America. California: Solano County, 1 mile
from Vacaville on road to Elmira. May 30, 1930. Holotype: CAS 179831;
isotypes: NO, GH, POM.

Mikania antioquiensis Hieron var. subcuneata Robinson, Contr. Gray Herb.

104: 32. 1934.

Ynes Mexia 6492. Peru. Loreto: Iquitos, trail to San Juan, elevation 105 meters.
February 7, 1932. Holotype: GH; isotype: NO.

= Mikania banisteriae DC; W. Holmes [herbarium annotation, 1976].

1993] Type Specimens of Vascular Plants 81

Mikania rimachii Holmes & McDaniel, Phytologia 41: 195. 1978.

McDaniel et al. 21646. Peru. Loreto: Maynas, 3 km up Rio Amazonas across
from Indiana. May 26, 1978. Holotype: IBE; isotypes: NO, AMAZ, F, MO,
NATC, NY, USM.

Mutisia ochroleuca Cuatrecasas, Feddes Repert. Spec. Nov. Regni Veg. 55: 122.
1953.

Ewan 16089. Colombia. Narino: Volcan de Chiles, northwest shoulder, eleva-
tion 3100 meters. September 6-8, 1944. Holotype: F; isotype: NO.

Pyrrhopappus georgianus Shinners, Field and Lab. 21: 93. 1953.

Cronquist 4276. United States of America. Georgia: Glynn County, St. Simon's
Island, near Brunswick. April 11, 1947. Holotype: SMU; isotypes: NO,
GA, MO.

=Pyrrhopappus carolinianus var. georgianus (Shinners) Ahles, J. Elisha Mitchell
Sci. Soc. 80: 173. 1964.

Senecio andersonii Clokey, Bull. S. Calif. Acad. Sci. 37: 10. 1938.

Clokey ^ Anderson 5638. United States of America. Nevada: Clark County,
Charleston Mountains, Lee Canyon, elevation 2550 meters. August 5, 1935.
Holotype: Clokey herbarium; isotype: NO.

Senecio chingualensis Cuatrecasas, Feddes Repert. Spec. Nov. Regni Veg. 55:
137. 1953.

Ewan 16212. Colombia. Narino: Cordillera Oriental, Rio Chingual at San Jose,
below La Victoria, elevation 2650 meters. September 25, 1944. Holotype:
F; isotype: NO.

Vernonia angustifoUa Torrey & A. Gray var. texana A. Gray, Syn. Fl. N. Amer.
1 (2): 91. 1884.

J. Hale s.n. United States of America. [Louisiana: vicinity of Alexandria (?),
without date]. Lectotype: NY; possible isolectotype: NO.

=Vernonia texana (A. Gray) Small, Fl. Southeastern U. S. 1 160. 1903.

Zexmenia hispida A. Gray var. ramosissima Greenman, Field Mus. Nat. Hist.,
Bot. Ser. 3: 127,^/. /. 1904.

Gaumer ^ sons 826. Mexico. Yucatan. Holotype: F; isotype: NO.

=Wedelia acapulcensis var. ramosissima (Greenman) Strother, Syst. Bot. Monogr.
33: 55. 1991.

BRASSICACEAE (CRUCIFERAE)

Cardamine constancei Detling, Madrofio 3(4): 176, pi. 9. 1935.

Constance et al. 1108. United States of America. Idaho: Idaho County, 4 miles
below Lowell, on Three Devils Creek at Middle Fork of Clearwater River,
Selway National Forest. June 2, 1935. Holotype: DS 229378; isotype: NO.

82 Tulane Studies in Zoology and Botany [Vol. 29

Lesquerella lyrata Rollins, Rhodora 57: 252, tab. 1209. 1955.

Rollins et al. 5599. United States of America. Alabama: Franklin County, near
Richardson's Crossing, 7 miles eastof Russellville. April 16, 1955. Holotype:
GH; isotype: NO.

Lesquerella perforata Rollins, Rhodora 54: 190. 1952.

Rollins ^ Rollins 5207. United States of America. Tennessee: Wilson County,
five miles north of Lebanon. March 30, 1952. Holotype: GH; isotype: NO.

CAMPANULACEAE

Centropogon minimus McVaugh, Brittonia 6: 467. 1954.

Ewan 15699. Colombia. Antioquia: Paramo de Sonson, above Sonson, Cordill-
era Central, elevation 2743 meters. May 23, 1944. Holotype: NA; isotype:
NO.

Siphocampylus buUatas McVaugh, Brittonia 6: 457. 1949.

Ewan 16611. Colombia. Narifio: West side of Volcan Dona Juana, elevation
3200-3400 meters. December 16, 1944. Holotype: US; isotype: NO.

CARYOPHYLLACEAE

Silene andersonii Clokey, Bull. S. Calif. Acad. Sci. 38: 2. 1939.

Clokey 7514. United States of America. Nevada: Clark County, Charleston
Mountains, Lovell Canyon, elevation 2600 meters. August 9, 1937.
Holotype: Clokey herbarium; isotypes: NO, CAS.

=Silene verecunda Wats, subsp. andersonii (Clokey) Hitchcock & Maguire, Univ.
Wash. Publ. Biol. 13: 43. 1947.

CLUSIACEAE (GUTTIFERAE)

Clusia chusqueae Ewan, Nat. Hist. Misc. 88: 6. 1951.

Ewan 15741. Colombia. Antioquia: Cordillera Central, Alto Capiro, above
Sonson- Abejorral camino, elevation 2800 meters. May 26-28, 1944.
Holotype: NA; isotypes: NO, COL.

Vismia buchtienii Ewan, Contr. U. S. Nad. Herb. 35(5): 353. 1962.

Buchtien 2022. Bolivia. San Antonio, Mapui region, elevation 850 meters.
December, 1907. Holotype: US 1159313; isotypes: NO, L.

Vismia cuatrecasasii Ewan, Contr. U. S. Natl. Herb. 35(5): 331. 1962.

Cuatrecasas 16687 . Colombia. Dept. del Valle: Rio Calima (region del Choco);
entre La Herradura de Ordonez y Peha de Campotriste, elevation 5-10
meters. March 3, 1944. Holotype: US 1950013; isotype: NO.

Vismia guianensis subsp. persicoides Ewan, Contr. U. S. Natl. Herb. 35(5): 342.
1962.

1993] Type Specimens of Vascular Plants 83

Schultes ^ Cabrera 12570. Colombia. Amazonas-Vaupes: Soratama, Rio
Apaporis, entre el Rio Pacoa y el Rio Kananari, elevation 250 meters. June
15, 1951. Holotype: NO; isotype: US.

Vismia laxiHora Reichardt in Martius, Fl. Bras. 12(1): 203. 1878.

Schomburgk 837. British Guiana [or Venezuela? - vide Ewan, Contr. U. S. Natl.
Herb. 35: 361. 1962]. Isotype: NO.

Vismia rufa Cuatrecasas, Revista Acad. Colomb. Ci. Exact. 7: 41 , fig. 1. 1946.

Cuatrecasas 17449. Colombia. Dept. del Valle: Costa del Pacifico; Rio Cajambre,
elevation 5-80 meters. May 5-15, 1944. Holotype: Colecion de la Comision
de Botanica del Valle; isotype: NO.

Vismia urceolata Ewan, Contr. U. S. Nad. Herb. 35(5): 324. 1962.

E. L. Little & R. R. Little 8275. Colombia. Puerto Lopez: Intendencia del Meta,
elevation 240 meters. July 28, 1944. Holotype: US 2 166544; isotype: NO.

CYCLANTHACEAE

Sphaeradenia stylosa Harling, Acta Horti Berg. 18(1): 352, fig. 94g-h, tab. 85.
1958.

Ewan 16085. Colombia. Nariiio: Volcan de Chiles, northwest slope, Mayasquer
trail, elevation 2560-3350 meters. September 6-8, 1944. Holotype: US;
isotype: NO.

CYRILLACEAE

Purdiaea stenopetala Grisebach, Abh. Konigl. Ges. Wiss. Gortingen 9: 45. 1 860.
C. Wright 3 4 l.Cnhdi. Orientali. 1859-1860. Isotypes: NO, MO, GH, NY, UC.

ERICACEAE

Arctostaphylos pinetorum Rollins, Rhodora 39: 462. 1937.

Rollins 1586. United States of America. Colorado: Mesa County, Uncompahgre
Plateau, T 50 N, R 16 W, 3 miles west of Cold Spring Ranger Station,
elevation 7500 feet. September 1, 1936. Holotype: GH; isotype: NO.

=A. parryana var. pinetorum (Rollins) Wieslander & Schreiber, Madrono 5: 46.
1939.

EUPHORBIACEAE

Ditaxis diversiHora Clokey, Bull. S. Calif. Acad. Sci. 37: 6. 1938.

Clokey 7576. United States of America. Nevada: Clark County, Charleston
Mountains, Charleston Park, elevadon 2280 meters. May 24, 1937.
Holotype: Clokey herbarium; isotype: NO.

£up/iorWa iO[Taci7ior Cronquist, Castanea 14: 102. 1949.

84 Tulane Studies in Zoology and Botany [Vol. 29

Cronquist 5334. United States of America. Georgia: Tattnall County, 2 mi.
northwest of Reidsville. June 16, 1948. Holotype: US; isotypes: NO, GA,
GH, MICH, MO, NY, PH, SMU, UC.

= Euphorbia exserta (Small) Coker; M.J. Huft [herbarium annotation, 1979].

FABACEAE (LEGUMINOSAE)

Cynometra insularis A. C. Smith, Sargentia 1: 38. 1942.

Degener 15491. Fiji. Viti Levu: Ra Province, Vatundamusewa, near Rewasa.
June 6, 1941. Holotype: A; isotype: NO.

Erythrina elenae Howard & Briggs, J. Arnold Arbor. 34: 183. 1953.

Howard, Briggs, et al. 377. Cuba. Las Villas: Hillside slope 0.25 mile west of
spot where Camino de la Sur crosses Rio San Juan along north slope of
Trinidad Mountains. July 1-20, 1950. Holotype: GH; isotype: NO.

Galactia bahamensis Urban, Symb. Antill. 2: 331. 1900.

Hitchcock s.n. Bahamas. Crooked Island. November, 1890. Holotype: MO
21394; isotype: NO.

RhynchosiarojasiiHsiSsler, Feddes Repert. Spec. Nov. Regni Veg. 7: 77. 1909.

Hassler 14024. Paraguay. Sierra de Amambay [without date]. Isotype: NO.

Trifolium grantianum Heller, Muhlenbergia 1: 136. 1906.

G. B. Grant 6343. United States of America. California: Mt. San Gorgonio,
San Bernardino Mountains. July 23, 1904. Holotype: POM; isotypes: NO,
UC 127247.

=Trifolium monanthum A. Gray var. grantianum (Heller) Parish, Plant World
20: 220. 1917; Zohary & D. Heller, Genus Trifolium, 544. 1984.

FAGACEAE

Quercus x ganderi C. B. Wolf, Proc. Calif. Acad. Sci., Ser. 4, 25: 178. 1944.

Wolf ^ Everett 9543. United States of America. California: San Diego County,
between Lake Henshaw and Santa Ysabel, just 300 yds. above entrance to
Volcan Indian School, elevation 3 100 feet. October 20, 1939. Isotype: NO.

Quercus grandidentata Ewan, Bull. Torrey Bot. Club 64: 512. 1927.

Ewan 8487. United States of America. Cahfornia: Los Angeles County, San
Gabriel Mountains, Monrovia, 2 mi. above A. T. and S. F. railroad tracks
at mouth of Sawpit Canyon (Monrovia Canyon). December 27, 1934.
Holotype: UC 533189; isotype: NO.

Quercus stellata Wangenh. var. araniosa Sargent, Bot. Gaz. 65: 441. 1918.

Palmer 8770. United States of America. Louisiana: Natchitoches Parish, Grand
Ecore. October 2, 1915. Holotype: A; isotype: NO.

1993] Type Specimens of Vascular Plants 85

= Quercus stellata Wangenh. var. margaretta (Ashe) Sargent; Little, Checklist
U. S. Trees (U. S. Forest Serv. Agric. Handb. vol. 541), 242. 1979.

GENTIANACEAE

Lisianthius peduncularis L. O. Williams, Fieldiana, Bot. 31: 408, fig. 1. 1968.

Allen 3410. Panama. Code: El Valle de Anton, elevation 1000 meters. March
16, 1946. Holotype: MO; isotype: NO.

[The generic name was spelled '' Lisianthus" in the species protologue; this was
corrected in Ann. Missouri Bot. Card. 62(1): 71. 1975.]

HYDROPHYLLACEAE

Nemophila menziesii Hook. & Arn. subsp. australis A. Brand var. minima A.
Brand, Pflanzenreich IV. Fam. 251 (Heft 59): 50. 1913.

M. E. Jones 3037 . United States of America. California: Los Angeles County,
Pasadena. February 25, 1882. Holotype: B; isotype: NO.

JUNCACEAE

Juncus duranii Ewan, Rhodora 47: 186. 1945.

J. ^ N. Ewan 10060. United States of America. California: Los Angeles County,
San Gabriel Mountains, Lilly Springs, north slope of Mt. Hawkins. August
14, 1936. Holotype: COLO; isotype: NO.

LAMIACEAE (LABIATAE)

Monardella lanceolata A. Gray var. glandulifera I. M.Johnston, Bull. S. Calif.
Acad. Sci. 18: 20. 1919.

Johnston 2139. United States of America. California: San Antonio Mountains,
Brown's Flats, Upper Sonoran Zone, elevation 4300 feet. September 1,
1918. Holotype: POM; isotypes: NO, DS, UC.

Salvia camosa Dougl. subsp. gilmanii Epling, Ann. Missouri Bot. Card. 25: 132.
1938.

Epling ^ Gilman s.n. United States of America. California: Inyo County,
Panamint Mountains above Pifion Mesa, Wild Rose Canyon. July 7, 1937.
Holotype: LA; isotypes: NO, CAS, LA, MO, UC.

-Salvia dorrii (Kellogg) Abrams var. dorrii; ]. L. Strachan, Brittonia 34: 160.
1982.

LAURACEAE

Cryptocarya degeneri A\\en, Sargentia 1: 34. 1942.

Degener 15412. Fiji. Viti Levu: Ra, Mataimeravula, vicinity of Rewasa, near
Vaileka, elevation 50-200 meters. June 3, 1941. Holotype: A; isotypes: NO,
BISH, K, NY, US.

= Cryptocarya hornei Gillespie; A. C. Smith, Fl. Vit. Nov. 2: 129. 1981.

86 Tulane Studies in Zoology and Botany [Vol. 29

LILIACEAE

A/Zium eurotop/ii7uni Wiggins, Contr. Dudley Herb. 1: X^"^, pi. 12, fig. 1. 1933.

Wiggins <y Demaree 5040. Mexico. Baja California: Sierra San Pedro Martir,
elevation 2800 meters. September 24, 1930. Holotype: DS 194171; isotype:
NO.

Calochortus rhodothecus Clokey, Bull. S. Calif. Acad. Sci. 37: 1. 1938.

Clokey 7479. United States of America. Nevada: Clark County, Charleston
Mountains, Charleston Park, elevation 2270 meters. July 21 [July 12 in
protologue], 1937. Holotype: Clokey herbarium; isotype: NO.

MAGNOLIACEAE

Magnolia acuminata (L.) L. var. ludoviciana Sargent, Bull. Torrey Bot. Club
55: 464. 1928.

R. S Cocks s.n. United States of America. Louisiana: West Feliciana, Catalpa.
October 15, 1915. Lectotype: NO.

Magnolia ashei Weatherby, Rhodora 28: 35. 1926.

W. W. Ashe 3. United States of America. Florida: Okaloosa County, Rogue
Creek. April 26, 1925; June 6, 1925. Holotype: NCU?; isotypes: NO, OH.

MALVACEAE

Sida ampla M. E.Jones, Contr. W. Bot. 15: 146. 1929.

Jones 22858. Mexico. Nayarit: Acaponeta. February 25, 1927. Holotype: POM;
isotypes: NO, CAS, F, NY(2), WIS(2).

= Sida aggregata Presl; Fryxell, Syst. Bot. Monogr. 25: 382. 1988.

Sida wendtii Fryxell, Phytologia 46(6): 393. 1980.

Koch, Fryxell, ^ Wendt 79424. Mexico. Oaxaca: Santo Domingo Armenta,
Terraceria a Santo Domingo A., 1.5 km al S. de la carretera Pinotepa
Nacional-Acapulco, 29 km al O. de Pinotepa Nacional, sobre la carretera
a Acapulco, elevation 20 meters. November 8, 1979. Holotype: ENCB;
isotypes: NO, BM, BR, CAS, CHAPA, CTES, F, K, MARY, MICH, MO,
NY, TEX, WIS, XAL.

^Sida hrachystemon DC; Fryxell, Syst. Bot. Monogr. 25: 385. 1988.

MARCGRAVIACEAE

Marcgravia fosbergiana Ewan, Nat. Hist. Misc. 88: 2. 1951.

Ewan 16053. Colombia. Nariiio: Rio San Juan at pueblo of San Antonio,
elevation 1370 meters. September 3, 1944. Holotype: NA; isotypes: NO,
COL.

1993] Type Specimens of Vascular Plants 87

NYCTAGINACEAE

Mirabilis mac farlanei Constance 8c Rollins, Proc. Biol. Soc. Wash. 49: 148. 1936.

Constance, Rollins, Clements, & Dillon 1579. United States of America. Oregon:
Wallowa County, Lower Cottonwood Landing, between mouth of Somer's
Creek and Pittsburg Landing, T 27 N, R 1-2 W, Snake River Canyon. May
15, 1936. Isotypes: NO(2).

PIPERACEAE

Peperomia ewanii Trelease &: Yuncker, Piperac. N. South Amer. 2: 573, fig.
497. 1950.

Ewan 16574. Colombia. Narino: 6 km above San Jose, on trail to Las Mesas,
via Quebrada San Francisco, elevation 2700 meters. December 14, 1944.
Holotype: NA; isotypes: NO, ILL.

Peperomia glareosa Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part II: 50.
1936.

Ynes Mexia 6236. Peru. Loreto: above Pongo de Manseriche, bank of Rio
Santiago, elevation 200 meters. December 6, 1931. Holotype: UC; isotypes:
NO, OH.

Peperomia mansericheana Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part
II: 60. 1936.

Ynes Mexia 6196. Peru. Loreto: above Pongo de Manseriche, streamlet near
mouth of Rio Santiago, elevation 210 meters. December 1, 1931. Holotype:
UC; isotypes: NO, GH.

=Peperomia vulcanicola C. DC. var. mansericheana (Trelease) Trelease & Yunc-
ker, Piperac. N. South Amer. 2: 606. 1950.

Peperomia santiagoana Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part 11:
92, 1936.

Ynes Mexia 6146. Peru. Loreto: above Pongo de Manseriche on left bank of
Rio Santiago, elevation 200 meters. November 23, 1931. Holotype: UC;
isotypes: NO, GH.

Piper ewanii Yuncker, Piperac. N. South Amer. 1: 278. 1950.

Ewan 16752. Colombia. Putumayo: near San Diego de Colorado, vicinity of
Finca Pedrogoso, between Umbria and Puerto Asis. January 11, 1945.
Holotype: ILL; isotype: NO.

Piper indianonum Trelease, Field Mus. Nat. Hist., Bot. Ser 13, part II: 176.
1936.

Ynes Mexia 6396. Peru. Loreto: Distrito Iquitos, Fundo Indiana, elevation 1 10
meters. January 21, 1932. Holotype: UC; isotype: NO.

Piper paranapuranum Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part II: 204.
1936.

88 Tulane Studies in Zoology and Botany [Vol. 29

Ynes Mexia 6075. Peru. Loreto: near Yurimaguas, Rio Parana-pura, elevation
180 meters. October 24, 1931. Holotype: UC; isotypes: NO, GH.

Piper rio-nechianum Yuncker, Piperac. N. South Amer. 1:211 ,fig. 1 74. 1950.

Ewan 15827. Colombia. Antioquia: Dos Bocas, on Rio Nechi, near Pato. June
23-25, 1944. Holotype: ILL; isotypes: NO, NA.

Piper scabridulicaule Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part H: 229.
1936.

Ynes Mexia 6106 [ = 6100 ?]. Peru. Loreto: above Pongo de Manseriche at
mouth of Rio Santiago. November 14, 1931. Isotype: NO.

Piper tingens Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part H: 244. 1936.

Ynes Mexia 6072. Peru. Loreto: near Yurimaguas, Rio Parana-pura, elevation
180 meters. October 24, 1931. Holotype: UC; isotype: NO.

Piper tortuosivenosum Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part H:
244. 1936.

Ynes Mexia 6199. Peru. Loreto: above Pongo de Manseriche, near mouth of
Rio Santiago, elevation 210 meters. December 2, 193L Holotype: UC;
isotypes: NO, GH.

Piper tumidicondy Hi Trelease, Field Mus. Nat. Hist., Bot. Ser. 13, part H: 247.
1936.

Ynes Mexia 6431. Peru. Loreto: Distrito Iquitos, Rancho Indiana, elevation
110 meters. January 25, 1932. Holotype: UC; isotype: NO.

POLEMONIACEAE

Eriastrum brandegeae H. L. Mason, Madrono 8(3): 88. 1945.

Mason 12604. United States of America. California: Obsidian Ridge, southeast
of Borax Lake. June 28, 1945. Isotype: NO.

POLYGONACEAE

Chorizanthe procumbens Nutt. var. mexicana Goodman, Ann. Missouri Bot.
Card. 21: 87. 1934.

Fosberg 8281 . Mexico. Baja California: Tecate, elevation 600 meters. May 30,
1932. Holotype: MO; isotype: NO.

Eriogonum baileyi S. Wats. var. porphyneticum S. Stokes ex M. E.Jones, Contr.
W. Bot. 11: 17. 1903.

M. E. Jones s.n. United States of America. Nevada: Palisade. September 6,
1881. Isotype: NO.

= Eriogonum baileyi S. Wats. var. baileyi; ]ames L. Reveal [herbarium annotation,
1969].

Eriogonum campanulatum Nutt. subsp. leptothecum S. Stokes, Genus Eriog.
78. 1936.

1993] Type Specimens of Vascular Plants 89

S. G. Stokes 206. United States of America. Colorado: Boulder County, five
miles north of Boulder on road to Fort Collins. August 2, 1934. Holotype:
CAS 215300; isotype: NO.

Eriogonum eastwoodianum ]. T. Howell, Leafl. W. Bot. 2: 133. 1938.

Eastwood & Howell 5857 . United States of America. California: Fresno County,
in mountains south of Jacalitos Creek, 14 miles from Coalinga on road to
Parkfield. June 13, 1938. Holotype: CAS 257469; isotype: NO.

Eriogonum exitnum Tidestrom, Proc. Biol. Soc. Wash. 36: 181. 1923.

Heller 10649. United States of America. Nevada: Washoe County, mountains
west of Franktown, elevation 5800 feet. August 16, 1912. Holotype: US
509910; isotype: NO.

=Enogonum ovalifolium Nutt. var. eximium (Tidestrom) J. T. Howell, Mentzelia
1: 19. 1976 ["1975"].

Eriogonum heermannii Dur. &: Hilg. var. floccosum Munz, Manual S. Calif.
Bot. 597. 1935.

Munz 12915. United States of America. California: Mohave Desert, eastern
San Bernardino County, Clark Mountains, elevation 5200 feet. September
16, 1932. Holotype: POM 183921; isotype: NO.

Eriogonum nudicaule (Torrey) Small subsp. parleyense S. Stokes, Genus Eriog.
84. 1936.

Stokes 213. United States of America. Utah: Parley's Canyon, Wasatch
[Wahsatch] Mountains, elevation 6000 feet. August 18 [or 17?], 1934.
Holotype: CAS 215298; isotype: NO.

= Eriogonum brevicaule Nutt. var. brevicaule; S. Welsh et al. Utah Fl., Great
Basin Nat. Mem. 9: 476. 1987.

RANUNCULACEAE

Delphinium amabile Tidestrom subsp. clarianum Ewan, Univ. Colorado Stud.,
Ser. D, Phys. Sci. 2: 198. 1945.

M. Clary s.n. United States of America. California: Riverside-San Bernadino
County line, Lost Horse Canyon, Litde San Bernardino Mountains, Con-
chilla Range. May 12, 1932. Holotype: COLO; isotype: NO.

Delphinium bicornutum Hemsley subsp. oaxacanum Warnock, Madrono 31:
247. 1984.

Warnock 2536. Mexico. Oaxaca: about 1 1.5 mi. south of Miahuatlan on Puerto
Angel road (Pemex Route 175). October 19, 1981. Holotype: TEX; isotypes:
NO, ENCB, ILL, MEXU, MO, SHST.

Delphinium collinum Ewan, Bull. Torrey Bot. Club. 63: 338. 1936.

Epling ^ Robison s.n. United States of America. California: San Diego County,
ridge between Campbell's Ranch and Mason Valley. April 4, 1932.
Holotype: LA; isotype: NO.

90 Tulane Studies in Zoology and Botany [Vol. 29

Delphinium disjunctum Ewan, Nat. Hist. Misc. 61: 1-3. 1950.

Meyer i^ Rogers 3019. Mexico. Nuevo Leon: Cerro del Viejo, 15 mi. west of
Dulces Nombres, Municipality Zaragoza, elevation 2500 meters. August 19,
1948. Holotype: MO; isotype: NO.

Delphinium gypsophilum Ewan, Univ. Colorado Stud., Ser. D, Phys. Sci. 2: 189,
fig 54. 1945.

Ewan 10295. United States of America. California: West Fresno County, mouth
of Pinoche Creek, on east facing slope, elevation 500 feet. April 25, 1937.
Holotype: COLO; isotype: NO.

Delphinium hesperium A. Gray forma hirsutum Ewan, Univ. Colorado Stud.,
Ser. D, Phys. Sci. 2: 179. 1945.

Ewan 9562. United States of America. California: Sutter County, eastern side
of North Butte, elevation 800 feet. May 9, 1936. Holotype: COLO; isotype:
NO.

Delphinium hesperium A. Gray forma pai/escens Ewan, Univ. Colorado Stud.,
Ser. D, Phys. Sci. 2: 179. 1945.

Ewayi 8835. United States of America. California: Napa County, along Steel
Canyon Road, 0.5 mile from junction with Monticello-Winters Road. April
30, 1934. Holotype: COLO; isotype: NO.

Delphinium hutchinsoniae Ewan, Bull. Torrey Bot. Club 78: 379. 1951.

5. Hutchinson 7423. United States of America. California: Monterey County,
in sand on beach 2.5 miles south of Monterey, elevation 10 feet. May 28,
1938. Holotype: COLO; isotype: NO.

Delphinium menziesii DC. subsp. pyramidale Ewan, Univ. Colorado Stud., Ser.
D, Phys. Sci. 2(2): 107. 1945.

H. C. Gilbert II 33. United States of America. Oregon: Benton County, north-
west of Corvallis. May 6,1916. Holotype: OSC 4151; isotypes: NO, NDA.

Delphinium ramosum Rydberg forma sidalceoides Ewan, Bull. Torrey Bot.
Club 78: 380. 1951.

Weber 5798. United States of America. Colorado: Saguache County, along
road to Stone Cellar Ranger Station and Saguache Park, near junction of
main highway, 4 miles west of Cochetopa Pass, elevation 9700 feet. July
28, 1950. Holotype: COLO; isotype: NO.

Delphinium treleasei B. F. Bush, Minnesota Bot. Stud. 2: 444. 1900.

Bush 73. United States of America. Missouri: Eagle Rock. May 28, 1898.
Isotype: NO.

Ranunculus alveolatus Carter, Amer. J. Bot. 26(7): 555, fig. lefi. 1939.

A. Carter 1244. United States of America. California: Sacramento County,
between Fair Oaks and Folsom, 1.5 miles southeast of Orangevale, elevation
200 feet. April 25, 1937. Holotype: UC 604080; isotype: NO.

1993] Type Specimens of Vascular Plants 91

ROSACEAE

Horkelia caeruleomontana St. John, Fl. Southeastern Washington. 199. 1937.

St. John ^ Smith 8329. United States of America. Washington: Columbia
County, Wildcat Springs, R 40 E, T 7 N, elevation 5500 feet. July 5, 1927.
Holotype: State College of Washington; isotype: NO.

Malus coronaria (L.) Mill. var. puberula Rehder, Trees Shrubs 2: 229. 1913.

R. S. Cocks s.n. United States of America. Louisiana: Winnfield. April, 1913.
Holotype: A; isotype: NO.

Potentilla beanii Clokey, Bull. S. Calif. Acad. Sci. 38: 4. 1939.

Clokey 7974. United States of America. Nevada: Clark County, Charleston
Mountains, Ridge to Charleston Peak, elevation 3270 meters. July 22, 1938.
Holotype: Clokey herbarium; isotypes: NO, CI.

Potentilla cryptocauUs Clokey, Bull. S. Calif. Acad. Sci. 37: 4, pi. 1. 1938.

Clokey 7567. United States of America. Nevada: Clark County, Charleston
Mountains, Charleston Peak, elevation 3500 meters. July 18 [or 8?], 1937.
Holotype: Clokey herbarium; isotypes: NO, CI, GH.

=Ivesia cryptocauUs (Clokey) Keck, Lloydia 1: 130. 1938.

Rubus parviflorus Nutt. forma micradenius Fassett, Ann. Missouri Bot. Card.
28: 324. 1941.

Fassett 20193. United States of America. Colorado: Routt County, gorge below
Fish Creek Falls, Steamboat Springs. July 3, 1939. Holotype: WIS; isotype:
NO.

Rubus parviflorus Nutt. var. velutinus (Hook. 8c Arn.) Greene forma para-
trichophorus Fassett, Ann. Missouri Bot. Card. 28: 325. 1941.

M. E. Mathias 1389. United States of America. California: Monterey County,
about 12 miles south of Carmel, 0.5-1 miles from ocean, Palo Colorado
Canyon, Santa Lucia Mountains. August 7, 1938. Holotype: WIS; isotype:
NO.

Rubus parviflorus Nutt. var. velutinus (Hook. 8c Arn.) Greene forma
parahypomalacus Fassett, Ann. Missouri Bot. Card. 28: 325. 1941.

B. O. Schreiber 2538. United States of America. California: Marin County, 1.5
miles southeast of Abbot's Lagoon. August 6, 1938. Holotype: WIS; isotype:
NO.

Rubus parviflorus Nutt. var. ve/urin us (Hook. 8c Arn.) Greene iorma parbifarius

Fassett, Ann. Missouri Bot. Card. 28: 325. 1941.

L. Constance 2397. United States of America. California: Alameda County,
Berkeley, north slope of Strawberry Creek Canyon, 0.7 mile above its
mouth, elevation 1 100 feet. August 4, 1938. Holotype: WIS; isotype: NO.

92 Tulane Studies in Zoology and Botany [Vol. 29

RUBIACEAE

Galium munzii Hilend & Howell forma glabrum Ehrendorfer, Contr. Dudley
Herb. 5: 9. 1956.

Clokey 8134. United States of America. Nevada: Clark County, Charleston
Mountains, canyon above Wilson's Ranch, elevation 1230 meters. June 7,
1938. Holotype: OH; isotypes: NO, ARIZ, DZ, POM, RSA.

=Ga/mm waWA^tt'm A. Gray var.mag7i?/b/mm Dempster, Brittonia 10: 189. 1958.

Hedyotis acuminata (Cham. & Schlechtend.) Steud. forma brownii Fosberg,
Bishop Mus. Bull. 174: 79. 1943.

Degener et al. 11672. United States of America. Hawaii: Waimalu, Oahu, ele-
vation 1000 feet. November 3, 1935. Holotype: Degener herbarium;
isotype: NO.

= Hedyotis acuminata (Cham. 8c Schlechtend.) Steud.; W. L. Wagner, D. Herbst,
8c S. H. Sohmer, Manual Fl. PI. Hawaii, 1138. 1990.

Hedyotis halei Torrey 8c A. Gray, Fl. N. Amer. 2: 42. 1841.

J. Hale s.n. United States of America. Louisiana: near Alexandria [without
date]. Holotype: NY; isotype: NO.

= Pentodon pentandrus (Schum. &: Thonn.) Vatke; Correll & Johnston, Manual
Vase. PI. Texas, 1490. 1970.

Hedyotis thyrsoidea Fosberg, Bishop Mus. Bull. 174: 52. 1943.

Degener 11722. United States of America. Hawaii: near Laianui, Molokai. May
18, 1928. Holotype: Degener herbarium; isotype: NO.

= Hedyotis mannii Fosberg; W. L. Wagner, D. Herbst, & S. H. Sohmer, Manual
Fl. PI. Hawaii, 1148, 1151. 1990.

Perama mexiae Standley ex Steyermark, Mem. New York Bot. Gard. 10(5): 246,
fig. 81a-e. 1964.

Ynes Mexia 5837. Brazil. Minas Geraes: Diamantina, Olaria, elevation 1245
meters. May 14, 1931. Holotype: NY; isotype: NO.

l?udg-ea mexiae Standley, Field Mus. Nat. Hist., Bot. Ser. 13, part II: 165. 1936.

Ynes Mexia 6366. Peru. Loreto: above Pongo de Manseriche, right of mouth
of Rio Santiago, elevation 210 meters. January 1, 1932. Holotype: F
718409; isotype: NO.

SAXIFRAGACEAE

Heuchera duranii Bacigalupi, Contr. Dudley Herb. 1: 191. 1933.

Duran 2817. United States of America. California: Mono County, McAfee
Meadow, White Mountains, elevation 11,700 feet. July 26, 1930. Holotype:
DS 207811; isotype: NO.

1993] Type Specimens of Vascular Plants 93

SCROPHULARIACEAE

Castilleja clokeyi Pennell, Proc. Acad. Nat. Sci. Philadelphia 89: 420. 1938.

Clokey 7322. United States of America. Nevada: Clark County, Charleston
Mountains, on north slope of Kyle Canyon, elevation 2425 meters. July 8,
1936. Holotype: PH; isotype: NO.

Pedicularis semibarbata A Gray var. charlestonensis Pennell & Clokey, Bull. S.
Calif. Acad. Sci. 38: 6. 1939.

Clokey 7718. United States of America. Nevada: Clark County, Charleston
Mountains, Charleston Park, elevation 2300 meters. May 1 1, 1936 (flowers);
June 21, 1937 (fruit). Holotype: Clokey herbarium; isotypes: NO, PH.

= Pedicularis semibarbata subsp. charlestonensis (Pennell & Clokey) Clokey, Mad-
rono 8: 60. 1945.

Penstemon alluviorum Pennell in Small, Manual Southeastern Fl. 1203. 1933.

Pennell 11504. United States of America. Arkansas: Craighead County, Jones-
boro. June 7, 1923. Holotype: PH; isotype: NO.

= Penstemon digitalis Nutt. ex Sims; E. B. Smith, Atlas Vase. PI. Arkansas, ed.
2, 253. 1988.

Penstemon arkansanus PenneW, Proc. Acad. Nat. Sci. Philadelphia 73: 493. 1922.

Pennell 10640. United States of America. Arkansas: Pulaski County, Pulaski
Heights, Little Rock. May 31, 1920. Holotype: NY; isotype: NO.

=Penstemon pallidus Small subsp. arkansanus (Pennell) Bennett, Phytologia 9:
57. 1963.

Penstemone bicolor (Brandegee) Clokey & Keck subsp. roseus Clokey Sc Keck,
Bull. S. Cahf. Acad. Sci. 38: 12. 1939.

Clokey 5850. United States of America. Nevada: Clark County, Nelson, El
Dorado Canyon, elevation 600 meters. May 14, 1938. Holotype: Clokey
herbarium; isotypes: NO, CI.

Penstemon connatifolius A. Nelson, Amer. J. Bot. 18: 437. 1931.

Nelson 10314. United States of America. Arizona: Apache Trail, not many
miles from Roosevelt Dam. May 3, 1925. Holotype: RM; isotype: NO.

Penstemon keckii Clokey, Madrono 4: 128. 1937.

Clokey 7312. United States of America. Nevada: Clark County, Charleston
(Spring) Mountains, elevation 2670 meters. July 3, 1936. Holotype: Clokey
herbarium; isotypes: NO, and distributed in Plantae Exsiccatae Grayanae.

Penstemon multicaulis Pennell, Proc. Acad. Sci. Philadelphia 73: 494. 1922.

Pennell 10616. United States of America. Arkansas: Franklin County, Ozark.
May 29, 1920. Holotype: NY; isotypes: NO, PH.

= Penstemon arkansanus Pennell; E. B. Smith, Adas Vase. PI. Arkansas, ed. 2,
251. 1988.

94 Tulane Studies in Zoology and Botany [Vol. 29

Penstemon neotericus Keck, Univ. Calif. Publ. Bot. 16: 398,/?^. 13. 1932.

Keck 958. United States of America. California: Plumas County, halfway be-
tween Chester and Westwood, elevation 1465 m (4800 feet). July 24, 1930.
Holotype: DS 185575; isotypes: NO, CA, CI, GH, K, MISS, NY, PH, POM,
UC.

SOLANACEAE

Solatium arborescens Clokey, Bull. S. Calif. Acad. Sci. 30: 60. 1931 ; non Solanum

arborescens Moench.

Clokey 5049 [5047 ?]. United States of America. California: Santa Cruz Island,
Pelican Bay, elevation 60 meters. May 23, 1930. Holotype: Clokey her-
barium; isotype: NO.

=Solanum clokeyi Munz, Bull. S. Calif. Acad. Sci. 31: 69. 1932.

=Solanum wallacei (A. Gray) Parish var. clokeyi (Munz) McMinn, Illustr. Manual
Calif. Shrubs, 491. 1939.

Solanum inopinum Ewan, Nat. Hist. Misc. 88: 8. 1951.

Ewan 16081. Colombia. Narino: vicinity of San Juan, on Rio San Juan. Sep-
tember 5, 1944. Holotype: NO(2).

Solanum siparunoides Ewan, Nat. Hist. Misc. 94: 1. 1952.

Ewan 15766. Colombia. Antioquia: Cordillera Central, Alto Capiro, above
Sonson-Abejorral camino. May 26-28, 1944. Holotype: NA; isotypes: NO,
COL.

TILIACEAE

Tilia floridana Small var. oblongifolia Sargent, Bot. Gaz. 66: 435. 1918.

Cocks 788. United States of America. Alabama: road to Hatcher's Bluff, near
Sardis. June 5, 1915; August 14, 1915 (fruit). Holotype: A; isotype: NO.

= Tilia caroliniana Mill.; J. 8c R. Kartesz, Synonymized Checklist Vase. Fl. U.
S. 462. 1980.

Tilia nuda Sargent var. glaucescens Sargent, Bot. Gaz. 66: 427. 1918.

Cocks 786. United States of America. Alabama: Dallas County, Cocks place,
Alabama River [G. . . 's Landing], Sardis. June 11, 1915; July 15, 1915
(fruit). Holotype: A; isotype: NO(2).

=Tilia leucocarpa Ashe var. glaucescens (Sargent) Sud worth. Checklist Forest
Trees U. S. 201. 1927.

VERBENACEAE

Aegiphila salticola Moldenke, Feddes Repert. Spec. Nov. Regni Veg. 37: 211.
1934.

1993] Type Specimens of Vascular Plants 95

Ynes Mexia 5922. Brazil. Para: Districto Acara, Thome Assu, Japanese Colony,
elevation 30 meters. July 16, 1931. Holotype: F; isotype: NO.

Lantana minasensis Moldenke, Phytologia 2: 138. 1946.

Ynes Mexia 4448a. Brazil. Minas Geraes: about km 2 along road from Vi^osa
to Barroso, elevation 690 meters. March 9, 1930. Holotype: NY; isotype:
NO.

Lantana rubella Moldenke, Phytologia 1: 464. 1940.

Ynes Mexia 5845. Brazil. Minas Geraes: Diamantina, upper slope of Serra do
Rio Grande, elevation 1290 meters. May 15, 1931. Holotype: NY; isotype:
NO.

Verbena hasslerana Briq. Bull. Herb. Boissier (II) 4: 1056. 1904.

Hassler 6464. Paraguay, [vicinity of Tobaty, September, 1900]. Syntypes: G-
DEL, NO [vide Moldenke, Phytologia 9: 177. 1963].

VOCHYSIACEAE

yoc/i/sjabraceiiniae Standley, Field Mus. Nat. Hist., Bot. Ser. 22(3): 149. 1940.

Ynes Mexia 6081. Peru. Loreto: near Yurimaguas on trail to San Ramon,
elevation 180 meters. October 25, 1931. Holotype: F; isotype: NO.

LIBRARY"

TULANE STUDIES IN , , , ,594

ZOOLOGY AND BOTANY' .^^^^

ri Art VAWD

UNIVERSITY
3lume 29, Number 2 May 1, 1994

THREE NEW SPECIES OF ETHEOSTOMA, SUBGENUS

ULOCENTRA, FROM THE GULF COASTAL PLAIN OF

SOUTHEASTERN UNITED STATES

ROYAL D. SUTTKUS

Museum of Natural History, Tulane University,
Belle Chasse, LA 70037

REEVE M. BAILEY

Museum of Zoology, The University of Michigan,
Ann Arbor, MI 48109-1079

AND

HENRY L. BART, JR.

Museum of Natural History, Tulane University,
Belle Chasse, LA 70037

FISHES OF THE BUFFALO RIVER SYSTEM,
WILKINSON COUNTY, SOUTHWESTERN MISSISSIPPI

MARK A. WARREN

Austin Community College, Cedar Park, TX 78613
ROBERT C. CASHNER

Department of Biological Sciences, University of New Orleans,
New Orleans, LA 70148

AND

ROYAL D. SUTTKUS

Museum of Natural History, Tulane University,
Belle Chasse, LA 70037

FISHES OF THE TANGIPAHOA RIVER SYSTEM,
MISSISSIPPI AND LOUISIANA

CHARLES L. KNIGHT

Mississippi Museum of Natural Science,
HI North Jefferson Street, Jackson, MS 39202

AND

ROBERT W. HASTINGS

Turtle Cove Environmental Research Station,
P.O. Box 585, Southeastern Louisiana University, Hammond, LA 70402

Tulane

University

New Orleans

TULANE STUDIES IN ZOOLOGY AND BOTANY

ISSN 0082-6782

Department of Ecology, Evolution, and Organismal Biology
Tulane University, New Orleans, Louisiana 70118

Steven P. Darwin, Editor

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THREE NEW SPECIES OF ETHEOSTOMA, SUBGENUS

ULOCENTRA, FROM THE GULF COASTAL PLAIN OF

SOUTHEASTERN UNITED STATES

Royal D. Suttkus

Museum of Natural History, Tulane University,
Belle Chasse, LA 70037

Reeve M. Bailey

Museum of Zoology, The University of Michigan,
Ann Arbor, MI 48109-1079

AND

Henry L. Bart, Jr.

Museum of Natural History, Tulane University,
Belle Chasse, LA 70037

Abstract

Etheostoma raneyi, E. lachneri, and E. ramseyi, three new darters of subgenus
Ulocentra occurring allopatrically in the Yazoo, Tombigbee, and Alabama River
systems, respectively, are described herein. The new species are distinguished
primarily on the basis of colors and color patterns of nuptial males. In E. raneyi,
the lateral stripe is composed of elongate blotches with even ventral margins
that barely extend below the lateral line. The lower side below the stripe is a
uniform red-orange. Etheostoma lachneri males have slightly oblique green bars
on the side of the body and caudal peduncle with bright orange in the in-
terspaces between bars. Etheostoma ramseyi males have ovoid to quadrate lateral
blotches that are connected with the dorsal saddle blotches. The orange ven-
trolateral coloration has dorsal extensions between the posterior lateral blotches.
Males and females of the three new species also exhibit slight differences in
body proportions. The three new Ulocentra lack a premaxillary frenum and
typically have one to several teeth on the head of the vomer, as is typical of
members of the E. duryi species group. The new forms plus E. colorosum and E.
tallapoosae are suggested to form a unique subgroup within the E. duryi group
in that they lack a red ocellus in the first interradial membrane and have incom-
plete bands of red and black blotches in the spinous dorsal fin. A principal
components analysis revealed differences in fin dimensions among species in
the five-member subgroup.

Introduction

The three species of darters described herein occur allopatrically in the
Yazoo, Tombigbee, and Alabama River systems. Until formal proposal of the
coastal darter {Etheostoma colorosum Suttkus and Bailey, 1993) one or both of the
easterly distributed forms were often thought to be conspecific with that species
(e.g., Kuehne and Barbour, 1983; Page and Burr, 1991). The three new species
share certain characteristics that suggest a close affinity with E. colorosum and
another nearby geographical representative, E. tallapoosae.

Tulane Studies in Zoology and Botany 29: 97-126. 1994.

97

98 Tulane Studies in Zoology and Botany [Vol. 29

Etheostoma colorosum occurs in coastal streams of southern Alabama and the
Panhandle of Florida from the Perdido through the Choctawhatchee rivers.
Populations of E. colorosum in western tributaries to the Escambia River system
are on the east side of a low drainage divide between the Escambia River and
lower eastern tributaries to the Alabama River. Thus, E. colorosum is isolated
from the new Alabama River system species.

Etheostoma tallapoosae Suttkus and Etnier (1991) occurs in the Tallapoosa
River system above the Fall Line and thus also is isolated from the new Alabama
River system species, which only occurs below the Fall Line in the Alabama River
system below the confluence of the Coosa and Tallapoosa rivers. However, the
new Alabama River species occurs below and above the Fall Line in the Cahaba
River system, a major tributary to the middle section of the Alabama River.

Identification of the three new species is based primarily on color and color
pattern of nuptial males. The subgenus, Ulocentra, as here used, follows
Bouchard (1977) and Bailey and Etnier (1988).

Materials and Methods

Specimens of the new species and comparative material used are primarily
from the collections of Academy of Natural Sciences of Philadelphia (ANSP);
Auburn University (AUM); Cornell University (CU); Mississippi State University
(MSU); Tulane University Museum of Natural History (TU); University of
Alabama (UAIC); University of Florida (UF); University of Michigan Museum
of Zoology (UMMZ); National Museum of Natural History (USNM); and Uni-
versity of Tennessee (UT).

Counts and measurements were made as described in Hubbs and Lagler
(1958) except as follows. Transverse body scales were counted from the origin
of the anal fm diagonally upward to the base of the spinous dorsal fm. Gill
rakers, counted on the anterior arch of either the left or the right side, include
both dorsal and ventral rudiments. Measurements were made with needle-point
dial calipers and recorded to the nearest 0.1 mm. Trans-pelvic width was meas-
ured between the outer bases of the pelvic spines. Our measurements of the
length of the spinous dorsal, soft dorsal, and anal fins are the total lengths of
these fins which is equal to the depressed lengths. Names used for associated
fish species follow Robins et al. (1991), except Etheostoma chlorosoma (Simons,
1992).

In the listing of type material, each catalog number is followed by the
number of specimens seen and the range of standard length (SL) in millimeters,
e.g. (10, 22 — 56). In addition to standard compass directions (with the following
"of deleted), the following abbreviations are used: mi. = mile(s), airmi. =
airmile(s), hwy. = highway, rd. = road, Cnty. = County, T = Township, R =
Range, Sec = Section. In lists of materials not designated as types, the catalog
number is followed by the number of specimens examined, enclosed in
parentheses. Highway numbers refer to state highways unless otherwise indi-
cated.

Etheostoma raneyi Suttkus and Bart, new species

Yazoo Darter
Figures 1, 6, 8, and 9

Etheostoma (Ulocentra) Bouchard, 1974: 41 (distribution). Randolph and Kennedy, 1974: 128-134
(Tippah River, Mississippi, records).

1 994] New Species of Etheostoma 99

Etheostoma sp. Clemmer et al., 1975: 8 (undescribed darter species, upper Yazoo River system,

Mississippi, rare). Jenkins, 1976: 644 (undescribed species, distribution).
Etheostoma {Ulocentra) sp. Deacon et al., 1979: 42 ("Yazoo darter," color illustration, male and female

from Puskus Creek, Lafayette Cnty., Mississippi). Kuehneand Barbour, 1983: 99-100 ("Yazoo

darter," briefly characterized, distribution).
Etheostoma sp. Page and Burr, 1991: 302, pi. 43, map 345 ("Yazoo darter," briefly characterized,

range).

Holotype: Adult male, TU 162795, 48.3 mm standard length. Hurricane
Creek, tributary to Tallahatchie River (Yazoo River system) at Mississippi High-
way 7, 2.5 miles north of Oxford (T7S, R3W, Sec 28), Lafayette Cnty., Missis-
sippi, 12 April 1992, R. D. Suttkus and H. L. Bart, Jr.

Paratopotypes: TU 162796 (14, 33 - 44), collected with holotype, distrib-
uted as follows: TU 162796 (12), UMMZ 222911 (2); TU 152358 (6, 27 - 31),
19 May 1988; UT 91.3542 (64, 22 - 47), 11 March 1989, distributed as follows:
UT 91.3542 (44), ANSP 170039 (2), CU 73780 (2), UMMZ 222912 (14), and
USNM 325768 (2); TU 158069 (8, 32-41), 18 May 1990; TU 163325 (13, 34
-44), 24 May 1992.

Other Paratypes: Tallahatchie River system, Mississippi. Lafayette Cnty.:
TU 152351 (18, 29 - 39), Graham Mill Creek, 0.7 mi. E Abbeville (T7S, R3W,
Sec 2), 19 May 1988, distributed as follows: TU 152351 (13), UMMZ 222910
(5); TU 158058 (3, 36 - 43), 18 May 1990; TU 162809 (4, 31 - 46), 12 April
1992; TU 66140 (17, 32 - 42), Puskus Creek, tributary to Little Tallahatchie
River at Hwy. 30, 7.9 mi. ENE Oxford (T8S, R2W, Sec 2), 8-9 October 1970,
distributed as follows: TU 66140 (13), UAIC 10681.01 (2), UP 100285 (2); TU
77310 (2, 35 - 47), 7 April 1972; TU 83595 (4, 31 - 48), 7 September 1973;
TU 84419 (1 1, 33 - 47), 13 October 1973, distributed as follows: TU 84419 (9),
CU 73781 (2); TU 87705 (2, 37 - 43), 7 May 1974; MSU 6038 (22, 32 - 44), 5
May 1976; AUM 20651 (4, 37 - 45), spring tributary to Puskus Creek at former
Minnows Inc. (T7S, R2W, Sec 34), 14 March 1980. Marshall Cnty.: TU 152338
(5, 34 - 40), Oak Chewalla Creek at Hwy. 310, 1.8 mi. W Malone (T6S, R3W,
Sec 9), 18 May 1988; TU 152346(5,38-44), 19 May 1988; TU 158054(1,29),
18 May 1990; TU 87058 (6, 30 - 41), Little Spring Creek at Hwy. 7, 0.5 mi. SW
Waterford (T5S, R3W, Sec 13), 27 October 1973; TU 87721 (1, 48), 7 May
1974; TU 152060 (1, 41), 10 May 1988; TU 84445 (1, 39), Little Spring Creek
at spillway out of Little Spring Lake, 1.6 mi. N Waterford (T5S, R3W, Sec 12),
13 October 1973; TU 152328 (4, 34 - 47), Big Spring Creek at US Hwy. 78, 3.8
mi. SE of junction US Hwy. 78 and Hwy. 7 (T4S, R2W, Sec 22), 18 May 1988;
UT 91.2176 (8, 31 - 47), 20 May 1981; UT 91.2167 (1, 27), Chewalla Creek,
tributary to Tippah River about 5 airmi. E Holly Springs (T4S, RIW, Sec 6), 20
May 1981.

Additional material examined, not designated as iypes: Vocona River system, Mis-
sissippi. Lafayette Cnty.: TU 152109 (10), tributary to Taylor Creek at Old Taylor Rd., 1.5 mi.
N Taylor (T9S, R4W, Sec 14), 11 May 1988; TU 87665 (3), Pumpkin Creek at Hwy. 334, 6.7 mi.
SE Oxford (T9S, R2W, Sec 19), 6 May 1974; TU 31 16 (1 1), Pumpkin Creek at US Hwy. 6, 6.5 mi.
E Oxford (T9S, R2W, Sec 3), 24 May 1952, TU 57136 (7), 17 April 1969, TU 152051 (22), 10 May
1988, UT 91.3496 (14), 22 October 1988, TU 155716 (4), 27 July 1989, and TU 158086 (3), 18
May 1990. Yalobusha Cnty.: TU 155210 (26), tributary to Otoucalofa Creek at Hwy. 32, 2.5 mi.
E Water Valley (T lis, R4W, Sec 15), 14 June 1989, distributed as follows: TU 155210 (21), UMMZ
222909 (5), TU 158100 (2), 18 May 1990, and TU 162832 (1), 12 April 1992; TU 155225 (3),
Gordon Branch, 17.2 mi. E Water Valley (TllS, R3W, Sec 7), 15 June 1989.

100 Tulane Studies in Zoology and Botany [Vol. 29

Diagnosis: Etheostoma raneyi is a member of subgenus Ulocentra as diagnosed
by Bouchard (1977) and Bailey and Etnier (1988). Further, E. raneyi lacks a
premaxillary frenum and the vomer usually bears a small cluster of slender
teeth and in these respects is typical of members of the E. duryi species group
of Bailey and Etnier (1988). Etheostoma raneyi lacks a red ocellus in the first
interradial membrane of the spinous dorsal fin, as do the other two new species
included in this study. The diagnostic features that distinguish the Yazoo darter
are the shape and position of the blotches in the lateral stripe and the ventral-lat-
eral coloration of the nuptial male. The more or less elongate brownish blotches
are aligned along the lateral line and have rather even ventral margins that
barely extend below the lateral line. The entire ventral-lateral area below the
lateral stripe of the nuptial male is a uniform reddish-orange. The red coloration
in the spinous dorsal fin is restricted to the posterior three to five interradial
membranes. There is an incomplete band of black blotches that extends from
the first to the seventh or eighth interradial membrane. These black blotches
are closer to the base of the fin than the aforementioned red blotches.

Description: Etheostoma raneyi reaches a maximum of 45 mm SL (females)
to 50 mm SL (males). Sexual dimorphism and apparent sexual maturity occur
at the end of the first year at a minimum of 28 mm SL (females) and 30 mm
SL (males).

Frequency distributions of scale and fin ray counts are presented in Tables
1-5. Number of vertebrae, branchiostegal rays, gill rakers, and preoperculoman-
dibular pores are given in Tables 6-7. The lateral line is usually complete with
(41 -) 43 - 49 (- 53) scales, occasionally the last scale is unpored. Transverse
scales 11 (2 specimens), 12 (36), 13 (57), 14 (51), 15 (16), or 16 (3). Caudal
peduncle scale rows number 15 (16), 16 (57), 17 (75), 18 (13) or 19 (4). Dorsal
fin has (9 -) 10 - 11 (- 12) spines and (9 -) 10-11 (- 12) soft rays. Anal fin
has 2 spines and (6 -) 7 - 8, modally 7, soft rays. Pectoral fin has (13 -) 14 - 15
rays. Vertebrae number 37 (8), 38 (41), or 39 (7); branchiostegal rays number
5 — 5 (149), 6 — 5 (4), 5 — 6 (6), or 6 — 6 (6); and gill rakers on first arch number
8 (6), 9 (19), or 10 (5). The sum of left and right preoperculomandibular pores
is nearly constant at 18 (in 26 of 30 specimens), of the remaining four, one has
16 pores and three have 17 pores. The opercle, cheek, and nape typically are

Table 1. Frequency distribution of transverse scale rows (anal to spinous dorsal) in three species
of Ulocentra.

11 12 13 14 15 16 17 N X S.D.

Etheostoma raneyi 2 36 57 51 16 3 165 13.3 1.0

Etheostoma lachneri 2 21 45 45 33 8 1 155 13.7 1.1

Etheostoma ramseyi 1 1 48 44 35 8 4 150 13.9 1.1

Table 2. Frequency distribution of number of scales around caudal peduncle in three species of

Ulocentra.

14 15 16 17 18 19 20 N x S.D.

Etheostoma raneyi 16 57 75 13 4 165 16.5 0.8

Etheostoma lachneri 1 23 35 97 37 18 2 213 17.0 1.1

Etheostoma ramseyi 2 20 45 85 45 12 3 212 16.9 1.1

1994]

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102 Tulane Studies in Zoology and Botany [Vol. 29

covered with exposed scales. The prepectoral area typically is covered with
scales. The breast is usually naked but occasionally has one or a few scales, and
the belly is completely covered with exposed scales.

In 10 specimens dissected for dentition, the vomer lacks teeth in one, the
head of the bone has two slender teeth in two, three in two, four in two, five in
two, six in one, mean 3.4 teeth. All 10 lack palatine teeth.

Proportional measurements appear in Tables 8-9. The 25 males were from
thirteen collections representing six different small stream systems. These collec-
tions were obtained during the months of April, May, June, September, and
October and thus represent nuptial and non-nuptial conditions. All proportional
measurements of the holotype are within the ranges given in Table 8. The 10
females were from six collections representing five different stream systems
obtained during April, May, and June and thus also represent nuptial and post-
nuptial conditions.

Colors of live and freshly preserved nuptial males, collected in March and
April are described from Hurricane Creek (Figure 1). Additional color descrip-
tions are provided for specimens collected in May from Hurricane, Graham
Mill, Pumpkin, Big Spring, and Taylor creeks. The entire lower side from the
midventral area up to the lower margin of the lateral stripe is orange to red-
orange color in large nuptial males. Small males, particularly in May collections,
have a varying amount of orange on the lower side, ranging from a narrow
stripe to a broad band that extends from axil of pectoral fin to the mid-area of
the caudal peduncle. The lateral stripe is composed of either a single or double
row of quadrate to oblong blotches. The single series exhibits a gradation from
red posteriorly to russet to brown anteriorly. This single row of blotches is
aligned along the upper margin of the lateral line in the posterior section but
more nearly straddles the lateral line anteriorly. Where a double row of blotches
occurs, the secondary series is composed of small brown blotches that border
the lower edge of the lateral line and typically occur in the interspaces between
the larger blotches. These two series of blotches are more or less suffused into
a single band at the height of nuptial coloration. This suffusion tends to dull
the brightness of the red-colored blotches in the posterior section of the primary
series. Posteriorly, some of the interspatial areas are overlaid with greenish-gray
pigment that extends ventrad from the dorsal-lateral area. The greenish-gray
pigment is usually in bar-shaped blotches on the caudal peduncle and base of
caudal fin and may extend slightly below the lower margin of the lateral stripe.
There is a scattering of irregularly shaped brown blotches along the dorsal-lat-
eral area. The mid-dorsal saddle blotches usually number eight and have a
greenish tinge in the nuptial male. The top of the head is dark brown. The
ventral surface of the head from the breast to the snout tip is pale and overlaid
with turquoise. The cheek, breast, gill membranes, lower part of opercle, and
side of snout are turquoise in color. There is a dark brown preorbital bar, a
dark brown suborbital bar, and several brown blotches along upper margin of
opercle.

Nuptial males have turquoise color in all fins except the pectorals. Nearly
the entire anal and pelvic fins are turquoise, with tips of the spines and soft rays
milky white. Two nuptial males from Hurricane Creek had a small red spot in
the posterior basal part of the anal fin. The margins of the caudal and first and
second dorsal fins are turquoise. The procurrent margins of the caudal fin are
more intense turquoise than the posterior margin. There are two orange to
red-orange blotches at the base of the caudal fin. The marginal band of tur-

1994] New Species of Etheostoma 103

quoise in the second dorsal fin is considerably broader than that in the spinous
dorsal fin. Proximally from the turquoise margin of the spinous dorsal fin there
are two distinct series of blotches. Posteriorly in the mid-portion of the fin there
is a series of red blotches; these number from three to six and are centered on
the interradial membranes. The posterior blotches are relatively large and di-
minish abruptly in size in the successive membranes anteriorly; there is some
pale cream to yellow color around the distal and proximal margins of the red
blotches. The other series of blotches is black; it extends from the first interra-
dial membrane to the sixth or seventh, and occurs in the basal half of each
interradial membrane (Figure 1). The posterior one to three black blotches are
proximal to the anteriormost red blotches described above.

Nuptial females have very little chromatic coloration. Occasionally a female
will have a few flecks of red in the posterior interradial membranes of the first
dorsal fin, three or four flecks of red in posterior interradial membranes of the
second dorsal fin, and more often will have upper and lower orange blotches at
the caudal fin base. Nearly all the pigmentation involves different shades of
brown. The blotches that compose the lateral stripe or band often are in one
series but occasionally form an interconnected double series. The latter pattern
usually has extensions above and below the lateral line. Females with a single
series of blotches have most of the pigment extending below the lateral line.
Regardless of whether the single or double pattern is present, the ventral margin
of the lateral stripe is more uneven in females than in nuptial males. Perhaps
the primary reason is that no suffusion or overlay of pigmentation occurs in the
females.

Distribution: Etheostoma raneyi is confined to tributaries to the Tal-
lahatchie and Yocona river systems in north-central Mississippi (Figures 6, 9).
Both rivers drain into the Yazoo River system. All of our material (five sites in
the Yocona River system and ten in the Tallahatchie River system) is from the
northern part of the North Central Plateau physiographic district (Lowe, 1925:
22-27). Our earliest record is that by R. D. Suttkus and two students on 24 May
1952, when eleven specimens were obtained from Pumpkin Creek, a tributary
to Yocona River. Additional collections were made periodically up to the present
dme. Randolph and Kennedy (1974), in a study of the Tippah River system,
reported samples from several sites in the upper part of that system. Their
easternmost locality occurs in the northern part of the Flatwoods physiographic
district (Figure 6). Kenneth W. Thompson and Robert J. Muncy were sponsored
by the U. S. Fish and Wildlife Service to compile a status report of the "Yazoo
River Darter," Etheostoma (Ulocentra) sp. Collections were obtained during the
period from 1980 to 1984. The report is undated but was submitted to the
contractor early in 1985. They found the "Yazoo River Darter" at three sites in
the Yocona River system and 14 sites in the Litde Tallahatchie River system.
They reported a total of 195 specimens, ranging from 20-61 mm SL. None
of the specimens collected by Randolph and Kennedy nor by Thompson and
Muncy was available to us. No other species of subgenus Ulocentra occurs within
the range of E. raneyi.

Habitat and biology: Hurricane Creek at Hwy. 7 (type locality) is a
muldple-channel stream that flows through a marsh area that varies from about
80 to more than 100 feet in width. There are several sets of beaver dams on the
upstream side of the highway. Much of the area is laden with a thick layer of
loose organic debris. There is an abundance of bur reed, Sparganium sp., along

104

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[Vol. 29

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106

Tulane Studies in Zoology and Botany

[Vol. 29

the main channel (10 - 15 feet in width) and along short stretches of lesser
channels. Also, there is an abundance of emergent aquatic plants along the
margins of the marshy stream system. Intermittently there are patches of sand
substrate along the main and lesser channels. Most of the Yazoo darters were
collected from near these patches of sand substrate.

1994]

New Species of Etheostoma

107

Figures 1-5. Etheostoma species from Mississippi, Alabama, and Florida. 1. E. raneyi, TU 162795,
holotype, adult male, 48 mm SL. Hurricane Creek, tributary to Tallahatchie River at Hwy. 7, 2.5
mi. N Oxford, Lafayette Cnty., Mississippi. Photo by Suttkus and Bart. 2. E. lachneri, TU 157555,
paratopotype, adult male, 45 mm SL. Wolf Creek, tributary to Little Souwilpa Creek at Hwy. 17,
1.0 mi. N Silas at US Hwy. 84, Choctaw Cnty., Alabama. Photo by Pierson. 3. E. ramseyi, TU 162574,
holotype, adult male, 49 mm SL. Beaver Creek at Wilcox Cnty. Rd. 27, 0.8 mi. W Sunny South,
Wilcox Cnty., Alabama. Photo by Suttkus. 4. E. tallapoosae, TU 157803, paratype, adult male, 55
mm SL. Gold Branch, tributary to Channahatchee Creek at Hwy. 63, 6.0 mi. N Eclectic, Flmore
Cnty., Alabama. Photo by Suttkus and Pierson. 5. E. colorosum, TU 162508, holotype, adult male,
50 mm SL. Pine Barren Creek, tributary to Escambia River, E.scambia Cnty., Florida. Photo by
Suttkus.

Water temperature in Hurricane Creek rises earlier in the spring than in
other nearby tributaries. On 11 March 1989 the water temperature was 16° C
and numerous Yazoo darters, nuptial males and females, were collected. On the
same day, Pumpkin (8° C) and Graham Mill (12° C) creeks were sampled. No
males and only a few females were collected. Other temperature records for
Hurricane Creek during which Yazoo darters were collected are as follows: 12
April 1992, 16° C; 18 May 1990, 21° C; 19 May 1988, 19° C; and 24 May 1992,
19° C. Water temperature records for 3 1 collections (including those cited above)
of Yazoo darters taken between 24 May 1952 and 24 May 1992 range from 16°

108

Tulane Studies in Zoology and Botany

[Vol. 29

91

90

89

34

33

34

33

91

90"

89°

Figure 6. Distribution oi Etheostoma raneyi in Mississippi (solid dots = material used in our study;
star in open circle = type locality; open circles = Randolph and Kennedy collection sites).

to 26° C. Two collections made at 25° C yielded four specimens each and one
collection at 26° C resulted in a single specimen. Observations on site, color
notes, and photos indicate that most spawning takes place during early spring
(March and April into May) with rising temperatures and tapers off or ceases
when water temperatures go above 20° or 21° C.

1994] New Species oi^ Etheostoma 109

Little and Big Spring creeks are similar to Hurricane Creek in that both have
an abundance of vegetation, primarily bur reed. Puskus and Oak Chewalla
creeks have primarily sand substrates with few or no aquatic plants. Graham
Mill and Taylor creeks have a mixture of sand and gravel substrate with man-de-
posited rubble at the bridge crossings and no aquatic vegetation. The tributary
to Otoucalofa Creek, 2.5 miles E Water Valley, has rubble at the bridge crossing
(where the Yazoo darters were obtained), sand and silted bottom upstream, and
sand and some hard clay bottom downstream with no aquatic vegetation. Most
of the streams inhabited by Yazoo darters are spring-fed as pointed out by
Thompson and Muncy in their status report. Many other sites within the Tal-
lahatchie and the Yocona river systems were sampled without obtaining Yazoo
darters. Invariably these sites were distant from spring sources, were disturbed
frequently by channelization, or had most or all the marginal vegetation re-
moved from banks. Individually and together these factors contribute to higher
water temperatures and unstable substrate, both conditions that appear to be
unsuitable for Yazoo darters.

The species associates of Etheostoma raneyi at the type locality based on four
collections are as follows: Lampetra aepyptera, Esox americanus, Erimyzon oblongus,
Cyprinella venusta, Lythrurus fumeus, Notemigonus crysoleucas, Noturus phaeus, Fun-
dulus olivaceus, Aphredoderus sayanus, Lepomis cyanellns, L. gulosus, L. macrochirns,
L. marginatus, L. punctatus, Micropterus salmoides, Etheostoma chlorosoma, E. lynceum,
E. nigrum, E. pannpinne, E. proeliare, E. swaini, and Percina sclera.

Etymology: We take pleasure in naming this darter, Etheostoma raneyi, for
the late Edward C. Raney in recognition of his numerous contributions to our
knowledge of the systematics and biology of darters. He had an intense interest
in darters and passed on his enthusiasm for them to a number of his students.
Dr. Raney authored or coauthored seven original descriptions of darters be-
tween the years 1939-1967.

Etheostoma lachneri Suttkus and Bailey, new species
Tombigbee Darter
Figures 2, 7, 8, and 9

Etheostoma (Ulocentra) sp. Smith- Vaniz, 1968: 136 (in part, reports from Black Warrior, upper and
lower Tombigbee rivers). Jenkins, 1976: 644 (in part, distribution). Knight and Cooper, 1987:
31, 33, 36, 38 (records in Otoucalofa Creek, Mississippi, habitat). Hubbard, et al. 1991: 3-8,
11 (records in Sucarnoochee River system). Boschung, 1992: 122 (in part, "orangestripe
darter," a complex of species).

Etheostoma sp. Mettee et al., 1983: 24 (in part), 26, 79, 82, 84, 86 (records from Citronelle and
Gilbertown oil fields, Alabama). O'Neil et al., 1984: 26 (distribution and abundance in Cit-
ronelle oil field, Alabama). Mettee et al., 1986: 19, 32 (records and abundance in Gilbertown
oil field, Alabama). Page, 1983: pi. 16G, 16H (in part, Minter Creek, Alabama, record).
Boschung, 1987: 187, 191 (distribution in upper Tombigbee River system, frequency of
occurrence, relative abundance). Page and Burr, 1991: 302, pi. 43, map 346 ("Coastal Plain
Darter," in part, briefly characterized, range).

Etheostoma sp. A. Mettee, 1978: 149 (habitat, distribution). Mettee et al., 1987: map (distribution,
species associates, habitat). Mettee et al., 1989a: 18, 19, 33-46, 188, map (Black Warrior River
system records, species associates, habitat). Mettee et al., 1989b: \52 (distribution).

Etheostoma sp. (colorosum) Hubbard, 1987: 24-26 (nomen nudum; misidentification; Noxubee River
system, Mississippi).

Holotype: Adult male, TU 162631, 41.6 mm standard length. Wolf Creek,
tributary to Little Souwilpa Creek at Alabama Highway 17, 1.0 mile north of
Silas at US Highway 84 (TION, R3W, Sec 31), Choctaw Cnty., Alabama, 1 April
1992, R. D. Suttkus.

110

Tulane Studies in Zoology and Botany

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Figure 7. Distribution o{ Etheostoma lachneri and E. ramseyi in the Mobile River Basin of Missis-
sippi, Alabama, Georgia, and Tennessee (star or triangle in open circle = type localities).

Paratopotypes: TU 162632 (30, 33 - 44), collected with holotype, distri-
buted as follows: TU 162632 (26), UMMZ 222913 (4); TU 56964 (43, 27 - 43),
22 March 1969, distributed as follows: TU 56964 (33), ANSP 170040 (2), CU
73782 (2), UAIC 10682.01 (2), UF 100286 (2), USNM 325769 (2); TU 151624
(50, 26 - 41), 9 April 1988; TU 152517 (75, 28 - 41), 26 May 1988, distributed
as follows: TU 152517 (65), UMMZ 222914 (10); TU 154427 (30, 32 - 46), 2
February 1989; UT 91.3567 (64, 33 - 45), 12 March 1989; TU 154488 (8, 35 -
46), 22 March 1989; TU 157555 (34, 33 - 45), 22 March 1990; TU 162344 (12,

1994] New Species of Etheostoma 1 1 1

35 - 48), 1 March 1992; TU 162558 (17, 35 - 49), 30 March 1992; TU 166059
(11, 30 - 46), 22 November 1992; TU 166419 (31, 32 - 46), 7 February 1993;
TU 166579 (57, 29 - 40), 25 March 1993; TU 166720 (31,31- 44), 16 April
1993.

Other Paratypes: TU 56990 (21, 28 - 42), Souwilpa Creek at Hwy. 17,
3.9 mi. S Gilbertown (TION, R3W, Sec 19), 22 March 1969; TU 76820 (1, 41),
Puss Cuss Creek, tributary to Okatuppa Creek at Hwy. 14, 4.2 mi. W Gilbertown
(TUN, R4W, Sec 28), 31 March 1972; TU 57007 (1, 32), Bogueloosa Creek at
Hwy. 17, 0.8 mi. N Toxey (TUN, R3W, Sec 17), 22 March 1969; TU 76831 (4,
31 - 39), tributary to Bogueloosa Creek at Hwy. 18, 0.6 mi. W Needham (T12N,
R4W, Sec 26), 31 March 1972.

Additional material examined, not designated as types: Tombigbee River system.
Alabama. Mobile Cnty.: TU 2598 (2), Sand Hill Creek, tributary to Chickasaw Creek at US Hwy.
45, 6.7 mi. S Citronelle (TIS, R3W, Sec 12), 4 June 1951, TU 32392 (51), 27 May 1964, and TU
162332 (3), 1 March 1992; TU 162324 (5), Cedar Creek at Hwy. 96, 4.8 mi. ENE Citronelle (T2N,
R2W, Sec 22), 1 March 1992. Washington Cnty.: TU 32409 (4), Taylors Creek, tributary to
Santa Bogue Creek at Hwy. 17, 10.2 mi. N Chatom (T7N, R3W, Sec 8), 27 May 1964; TU 162652
(7), Dunbar Creek at Hwy. 17, 1.7 mi. N Millry (T8N, R3W, Secl7), 1 April 1992. Clarke Cnty.:
TU 41434 (161), Salt Creek at Clarke Cnty. Rd. 15, 4.5 mi. S Jackson (T6N, R2E, Sec 34), 30 June
1966, TU 151775 (25), 16 April 1988, and TU 162395 (17), 2 March 1992; TU 32519 (20), tributary
to Salt Creek at Clarke Cnty. Rd. 15, 4.4 mi. S Jackson (T6N, R2E, Sec 34), 29 May 1964; TU 45503
(3), Rabbit Creek, tributary to Bassett Creek, 2.7 mi. NE Jackson (T7N, R2E, Sec 14), 16 April 1967;
TU 32466 (30), Bassett Creek, 2.0 mi. W Suggsville (T7N, R3E, Sec 12), 28 May 1964; TU 35314
(82), Bassett Creek at US Hwy. 84 at Allen, 8.0 mi. SE Grove Hill (T7N, R3E, Sec 12), 2 July 1964;
TU 152481 (19), Little Bassett Creek at Clarke Cnty. Rd. 30, 1.1 mi. E Dickinson (T9N, R4E, Sec
8), 26 May 1988; TU 59930 (29), Stave Creek at Hwy. 69, 5.4 mi. NW Jackson (T7N, RIE, Sec 25),
24 September 1969; TU 59959 (45), Jackson Creek at Hwy. 69, 9.7 mi. NW Jackson (T7N, RIE,
Sec 9), 25 September 1969, and MSU 1437 (13), 2 July 1970; TU 56939 (4), Tattilaba Creek at US
Hwy. 84, 7.4 mi. W Grove Hill (T9N, R2E, Sec 32), 19 March 1969; TU 43195 (2), Ulkinask Creek
at Clarke Cnty. Rd. 1, 2.0 mi. WNW Coffeeville (TION, RIW, Sec 33), 3 February 1967; TU 52128
(28), Pace Creek at Hwy. 69, 5.2 mi. N Coffeeville (TION, RIW, Sec 20), 23 May 1968; TU 52145
(34), Tallahatta Creek at Hwy. 69, 1 mi. SW Campbell (TUN, RIE, Sec 20), 23 May 1968. Choctaw
Cnty.: TU 52111 (5), Bladen Springs Run at Bladen Springs Park, 1 mi. NE town of Bladen
Springs (T9N, R2W, Sec 20), 23 May 1968, and TU 56951 (5), 19 March 1969; TU 76878 (8),
Wahalak Creek at Hwy. 17 at south edge of Butler (T13N, R2W, Sec 30), 31 March 1972. Maren(;o
Cnty.: TU 24539 (6), North and South Double Creek at Hwy. 69, 3 mi. N junction of Hwy. 69
and Hwy. 10 (T14N, RIE, Sec 34), 27 March 1961; TU 60931 (13), Beaver Creek at Hwy. 69, 1 1.9
mi. SW Linden (T15N, RIE, Sec 36), 18 December 1969; TU 163228 (20), Beaver Creek at US
Hwy. 43, 5.8 mi. N Dixons Mill (T14N, R3E, Sec 34), 1 1 May 1992, TU 166593 (3), 26 March 1993.
and TU 166748 (6), 17 April 1993. Sumter Cnty.: UMMZ 97797 (29), Sucarnoochee Creek, SE
Coatopa (T17N, RIW, Sec 6), 1 July 1931. Greene Cnty.: AUM 26795 (1), Brush Creek at Greene
Cnty. Rd. 133, SW Eutaw (T21N, RIE, Sec 3), 19 August 1990; TU 166606 (13), Minters Creek at
Greene Cnty. Rd. 213, 9.7 mi. N Eutaw (T23N, R2E, Sec 13), 26 March 1993, and TU 166762 (8),
17 April 1993. Hale Cnty.: TU 61373 (50), Elliotts Creek at Hwy. 69 at Moundville (T23N, R5E,
Sec 6), 20 December 1969, and AUM 26308 (7), 2 February 1990. Tuscaloosa Cnty.: UMMZ
177746 (17), tributary to Black Warrior River at Hwy. 69, 2.5 mi. N Tuscaloosa (T21S, R5E, Sec
10), 23 May 1956; TU 30194 (12), Carroll Creek at Hwy. 69, 5 mi. N Northport (r20S, RlOW, Sec
22), 10 April 1963; UMMZ 166363 (12), Carroll Creek at detour US Hwy. 43, 5 mi. N Tuscaloosa
(T20S, RlOW, Sec 20), 4 May 1952; UMMZ 166373 (8), Binnion Creek at US Hwy. 43, 5.5 mi. S
Samantha (T19S, RlOW, Sec 16), 4 May 1952. Pickens Cnty.: TU 154442 (1), Little Bear Creek
just off US Hwy. 82 at western edge of Gordo (T20S, R13W, Sec 9), 1 1 March 1989; TU 77044
(21), Bear Creek, tributary to Lubbub Creek, 2.0 mi. NNW Gordo (T20S, R13W, Sec 5), 3 April
1972; TU 76961 (1), Lindsey Branch, tributary to Tombigbee River, 6.6 mi. N Pickensville (T20S.
R17W, Sec 13), 2 April 1972. Lamar Cnty.: MSU 3882 (5), Luxapalila River at Hwy. 17, N Millport
(T17S, R15W, Sec 14), 17 November 1973; TU 163077 (3), Hells Creek at Lamar Cnty. Rd. 41
(T15S, R14W, Sec 23), 30 April 1992. Fayette Cnty.: TU 163098 (2), Hells Creek at Hwy. 18,
NW Fayette (T15S, R13W, Sec 17), 30 April 1992. Marion Cnty.: TU 30225 (10), New River at
US Hwy. 78 about 4 mi. W Elridge (TBS, RllW, Sec 10), 10 April 1963; TU 19080 (2), North
Fork Creek at US Hwy. 43, 10.2 mi. NNE Hamilton (T9S, R13W, Sec 33), 22 August 1958. Missis-

112 Tulane Studies in Zoology and Botany [Vol. 29

sippi. Lauderdale Cnty.: UMMZ 113865-6 (15), Ponta Creek, tributary to Sucarnoochee Creek
at US Hwy. 45, 12 mi. S Electric Mills (T8N, R17E, Sec 24), 3 June 1932. Kemper Cnty.: TU
127832 (4), Sucarnoochee Creek at Hwy. 16, 8.7 mi. W Scooba (TUN, R17E, Sec 19), 10 October
1982. Winston Cnty. MSU 999 (6), tributary to Hashuqua Creek at Hwy. 14, 2 mi. W Noxubee
Cnty. line (T14N, R14E, Sec 3), 24 May 1970. Choctaw Cnty.: MSU 722 (9), tributary to Noxubee
River about 2 mi. SE Choctaw Lake (T16N, RUE, Sec 2), 21 November 1969. Lowndes Cnty.:
TU 76945 (2), Ellis Creek, tributary to Tombigbee River about 6 mi. S Columbus (T19S, R17W,
Sec 17), 2 April 1972; TU 83698 (1), Tombigbee River at Buzzard Island (before impoundment)
(T19S, R18W, Sec 9), 8 September 1973; AUM 3349 (2), tributary to Buttahatchie River about 5
mi. N Caledonia (T15S, R17W, Sec 22), 2 October 1968.

Diagnosis: Etheostoma lachneri is a member of subgenus Ulocentra as diag-
nosed by Bouchard (1977) and Bailey and Etnier (1988). Etheostoma lachneri lacks
a premaxillary frenum and the vomer usually has a small cluster of two to five
or six teeth, occasionally lacking, as is typical of members of the E. duryi species
group of Bailey and Etnier (1988). Etheostoma lachneri lacks a red ocellus in the
first membrane of the spinous dorsal fin. The unique diagnostic features of E.
lachneri are the slighdy oblique, dark green bars on the side of body and caudal
peduncle, and the bright orange interspaces of the nuptial males. Etheostoma
lachneri is similar to the other two species described herein in that the red colora-
tion in the spinous dorsal fin is restricted to the posterior three to five interradial
membranes, and there is an incomplete band of black blotches that extends
from the first to the sixth or seventh interradial membrane. Both the spinous
and the soft dorsal fins oi E. lachneri are moderately elevated, in comparison to
other members of the colorosum subgroup.

Description: Etheostoma lachneri reaches a maximum of 47 mm SL
(females) to 52 mm SL (males). Frequency distributions of scale and fin-ray
counts are presented in Tables 1-5. Number of vertebrae, branchiostegal rays,
gill rakers, and preoperculomandibular pores are given in Tables 6-7. The lat-
eral line is usually complete with (39 -) 41 - 49 (- 51) scales, but occasionally
the last scale is unpored. Transverse scales number 1 1 (2 specimens), 12 (21),
13 (45), 14 (45), 15 (33), 16 (8), or 17 (1). Transverse scales from origin of
second dorsal fin to anal fin (an alternative count, see Bailey and Etnier 1988:
30) number 10 (1), 11 (31), 12 (19), 13 (7), mean of 58 counts, 11.5. Caudal
peduncle scale rows number 14 (1), 15 (23), 16 (35), 17 (97), 18 (37), 19 (18),
or 20 (2). Dorsal fin has (9 -) 10-11 (- 12) spines and (10 -) 11 - 12 (- 13)
soft rays. Anal fin has 2 spines and (6 -) 7 - 8 (- 9), modally 7, soft rays. Pectoral
fin has (13 -) 14 - 15 (- 16) rays. Vertebrae number 37 (11), 38 (66), 39 (13),
or 40 (2); branchiostegal rays number 5-^ (1), 5—5 (201), 6—5 (4), 5—6 (3),
or 6—6 (4); and gill rakers on first arch number 6 (7), 7 (32), 8 (90), 9 (44), 10
(8), or 11 (1). In general there is a weak clinal increase in the number of rakers
from the northern or headwater tributaries southward to the southern or lower
tributaries to the system. The sum of left and right preoperculomandibular
pores is nearly constant at 18 (in 187 of 198 specimens); of the remaining 11,
one has 16 pores, eight have 17 pores, and two have 19 pores. The opercle,
cheek, and nape are nearly or completely covered with exposed scales. The nape
is the most likely of the three areas to be incompletely covered with scales, and
the cheek is the least likely to be incompletely covered. The breast is consistently
scaleless.

In 10 specimens dissected, the vomer is toothless in three, has two slender
teeth in three, three teeth in three, and a cluster of five or six in one, mean 2.05
teeth; the palatines are uniformly toothless.

1994] New Species oi Etheostoma 1 13

Proportional measurements appear in Tables 8-9. Twelve of the 25 males
were collected at the type locality during March and May. The remainder of the
specimens came from six additional sites during March, April, and May. The
ten females came from seven different sites including the type locality. Three
specimens from the type locality were collected during March and the specimens
from the six other sites were collected during March, May, and August.

Colors of live and freshly preserved nuptial specimens, collected in March
and April, are described from Wolf Creek (type locality), tributary to Little
Souwilpa Creek of the Tombigbee River system (Figure 7). Nuptial males at
peak coloration have four to six, slightly oblique, dark green bars on the side of
body and caudal peduncle. The posterior bar occurs near the middle of the
caudal peduncle. Its broad base on the ventral part of the peduncle is usually
continuous posteriorly with the dark green or turquoise on lower procurrent
edge of caudal fm. Its dorsal extension on the side of peduncle narrows in width
to form a more or less triangular bar. Occasionally nuptial males have this
posterior bar connected to lower edge of a dorsal saddle blotch and rarely this
posterior, triangular bar is branched to form two dorsal projections. The penul-
timate bar has a narrower base than the ultimate bar and extends from the
anterior ventral part of caudal peduncle dorsad to the dorsal saddle blotch at
the posterior insertion of second dorsal fm. The next bar anteriorly lies between
the middle of the base of the anal fm and dorsal saddle blotch slightly anterior
to the middle of the second dorsal fm base. The next bar anteriorly extends
from the anterior base of anal fm dorsad to dorsal saddle blotch at posterior
base of spinous dorsal fm. Frequendy this bar does not extend to dorsal saddle
blotch but instead only about halfway between lateral line and dorsum, whereas
other nuptial males (Figure 2) have a partial bar that extends ventrad from
dorsal saddle blotch to slighdy below lateral line and not all the way to the base
of anal fm. Occasionally, large males have one or two more bars anterior to the
ones described above but these invariably are incomplete, extending from dorsal
saddle blotches ventrad to lateral line area or more or less centered vertically
across the lateral line. Prenupdal males from February and early March usually
have the triangular, uldmate bar well developed but the other bars are only
partially developed. The interspaces between the green bars of the nuptial males
are bright orange. This orange extends over entire ventral-lateral area anterior
to the anal fm insertion. The coloration of the dorsal-lateral area of body and
caudal peduncle varies considerably. Nuptial males have a row of orange to
red-orange blotches just above the lateral line although more frequently this
coloration is in a loose network pattern rather than discrete blotches. In addition
there are scattered flecks of orange or red-orange on the remainder of the
dorsal-lateral area. Other nuptial males have a reduced number of orange flecks
and blotches. The dorsal saddle blotches (typically eight) and the top of head
are dark brown with an overlay of green. There is a dark preorbital bar, a
suborbital bar, and several post-orbital dark brown blotches. The snout, lips, gill
membranes, isthmus, breast, as well as lower margins of cheek and opercle, are
green or turquoise.

The pectoral fms are clear in the upper two-thirds and light green or olive
on the lower third. The pelvics and all the median fms have some turquoise
color. The pelvic fms except for the tips of the spines and soft rays are deep
turquoise. The dps of rays are milky white. The anal fm is deep turquoise except
for a pale "window" in the posterior central area. The caudal fin typically has
bright turquoise procurrent marginal areas that are continuous with a broad

114 Tulane Studies in Zoology and Botany [Vol. 29

distal band of turquoise. The central basal area of the caudal fin is pale turquoise
or nearly clear. There are two orange blotches, one above and one below the
dark brown basi-caudal spot. The lower orange blotch is smaller and less distinct
than the upper. Both the spinous and soft dorsal fins have turquoise margins
and that on the soft dorsal fin is broader. Occasionally the turquoise will extend
over the distal half of the anterior part of the spinous dorsal fin. The complete
or nearly complete band of red blotches in the second dorsal fin and the incom-
plete, posteriorly located, red band in the first dorsal fin are similar to those
described for E. raneyi. The cream and yellow color around the red blotches in
both dorsal fins and the short band of black blotches in the first dorsal also are
similar in position, as described for E. raneyi.

Nuptial females have little or no chromatic coloration. Small nuptial females
lack chromatic coloration or have slight development of basi-caudal orange
blotches. Some large females have basi-caudal orange blotches plus greenish
lower procurrent margin of caudal fin and ventral posterior part of caudal
peduncle; a light green wash on chin, gill membranes, and breast; Hght green
along base of anal fin; and one or two small red blotches in the posterior mem-
branes of the spinous dorsal fin.

Distribution: Etheostoma lachneri is confined to the Tombigbee River sys-
tem, from slighdy above the Fall Line in several headwater tributaries to the
lower part of the system in Mobile Cnty., Alabama (Figures 7, 9; Mettee et al.,
1987: map, p. 173; Mettee et al., 1989a: map, p. 188; Boschung, 1989: map 107,
p. 93; Mettee et al., 1989b: map, p. 152). Most of the records in the upper
Tombigbee River system are in tributaries draining into the river from the east.
The reason for the abrupt demarkation of the western distribution is undoubt-
edly correlated with the fact that the Tombigbee River is the common border
for the western edge of the Fall Line Hills physiographic district and the eastern
border of the Black Belt physiographic district (Mettee et al., 1989a: 18, Figure
7). Moreover, each physiographic district has a distinct geological base.

Habitat and biology: Etheostoma lachneri occurs in a variety of habitats:
small streams with a mixture of sand and gravel substrate; rubble-strewn areas
at road crossings; and mixtures of sand, gravel, and hard clay or bedrock bot-
toms. The type locality. Wolf Creek, tributary to Litde Souwilpa Creek, is a small
stream 15 to 20 feet in width with sections of bedrock and sand bars upstream
of Hwy. 17. There are some deep potholes in the bedrock sections. Downstream
of Hwy. 17 the stream cascades down numerous ledges of bedrock. Wolf Creek
enters Little Souwilpa Creek at the lower end of an extensive cascade on the
Little Souwilpa, and has an overhanging canopy of shrubs and trees except at
the highway crossing.

Based on recent observations, prenuptial conditions were evident in Feb-
ruary and spawning occurs from early March to late April in the more southerly
areas and perhaps into late May in the more northerly areas. The sex ratio is
skewed toward more females in spawning aggregations. Based on nine collec-
tions of topotypes taken during February, March, and April, the 289 specimens
include 124 (42.9%) males and 165 (57.1%) females. The 124 males range from

27.0 to 48.6 mm in SL with a mean of 37.9 mm; the 165 females range from

26.1 to 43.7 mm, mean 35.6 mm.

The species associates oi Etheostoma lachneri at Wolf Creek (type locality), are
as follows: Erimyzon oblongus, Hypentelium etowanum, Moxostoma erythrurum, Moxos-
toma poecilurum, Campostoma sp., Cyprinella callistia, C. venusta, Luxilus

1994] New Species of f/Zi^oi/owa 115

chrysocephalus, Lythrurus bellus, Nocomis leptocephalus, Notropis ammophiius, N. baileyi,
N. buccatus, Pimephales notatus, Semotilus atromaculatus, Ameiurus natalis, Noturus
gyrinus, N. leptacanthvs, N. nocturnus, Fundulus olivaceus, Lepomis cyanellus, L. mac-
rochirus, L. megalotis, Etheostoma rupestre, E. stigmaeum, E. whipplei, and Percina
nigrofasciata.

Variation: The number of gill rakers, in general, varies from low in the
upstream tributaries to higher in the downstream tributaries to the Tombigbee
River. However, the lowermost tributary sampled showed a reverse trend.

Etymology: The name, Etheostoma lachneri, is in recognition of Ernest A.
Lachner, a few of whose early publications were concerned with detailed studies
of the biology of darters. Too, we recognize Dr. Lachner's exhaustive studies of
the systematics, zoogeography, and biology of the genus Nocomis, but perhaps
of equal importance to some of us (R. D. S.) was his seemingly untiring, unselfish
expenditure of time and energy to help young ichthyologists during their forma-
tive professional years while he was Curator of Fishes at the United States Na-
tional Museum.

Etheostoma ramseyi Suttkus and Bailey, new species

Alabama Darter
Figures 3, 7, 8, and 9

Etheostoma simoterum. Gilbert, 1891 : 155 (misidentification, "abundant in Alabama basin, and varying
more or less from the typical form").

Etheostoma (Ulocentra) sp. Smith-Vaniz, 1968: 136 (in part, reports from Cahaba and Alabama r.
systems). Jenkins, 1976: 644 (in part, distribution). Kuehne and Barbour, 1983: 97 ("Gulfcoast
Snubnose Darter," in part, characterized, range map, in part). Pierson et al., 1989: 1, 14, 15,
17, 26-50, 171 (habitat, map of localities in Cahaba River system, Alabama). Boschung, 1992:
122 (in part, "orangestripe darter," a complex of species).

Etheostoma sp. Suttkus and Boschung, 1990: 60 (Chilatchee Creek, Alabama record, species as-
sociates). Page and Burr, 1991: 302, map 346 (in part, characterized, range map in part).

Holotype: Adult male, TU 162574, 49.2 mm standard length, Beaver
Creek, tributary to Alabama River at Wilcox Cnty. Road 27, 0.8 mile west of
Sunny South (TUN, R4E, Sec 3), Wilcox Cnty., Alabama, 31 March 1992, R.
D. Suttkus.

Paratopotypes: TU 162575 (138, 29 - 46), collected with holotype, dis-
tributed as follows: TU 162575 (126), CU 73783 (2), UMMZ 222915 (4), USNM
325770 (2), UT 91.4345 (4); TU 158276 (48, 30 - 49), 26 May 1990, distributed
as follows: TU 158276 (42), ANSP 170041 (2), UAIC 10683.01 (2), UF 100287
(2); TU 166073 (20, 33 - 49), 22 November 1992; TU 166090 (15, 34 - 47), 23
November 1992; TU 166398 (33, 32 - 45), 7 February 1993; TU 166625 (19,
35 - 47), 27 March 1993; TU 166737 (20, 36 - 49), 17 April 1993; TU 167014
(29, 35-46), 17 May 1993.

Other Paratypes: Alabama. Clarke Cnty.: TU 158262 (7, 38-42), Beaver
Creek at dirt road, 0.8 mi. SE Atkinson (TUN, R4E, Sec 3). 26 May 1990.
Wilcox Cnty.: UAIC 2040 (20, 18 - 47) Beaver Creek, 2 mi. SE Pine Hill at
Cnty. Rd. (T12N, R5S, Sec 33), 30 May 1966; TU 44496 (26, 27 - 38), Turkey
Creek at Hwy. 5, 1.0 mi. S Kimbrough (T12N, R5E, Sec 10), 18 March 1967;
TU 154710 (12, 32 - 43), 9 May 1989. Marengo Cnty.: TU 32429 (142, 14 -
44), Dry Creek, tributary to Turkey Creek at Hwy. 10, 1.0 mi. E Vineland or

116 Tulane Studies in Zoology and Botany [Vol. 29

4.7 mi. W Hwy. 5 at Pine Hill (T12N, R4E, Sec 11), 28 May 1964; TU 151854
(33, 28 - 42), 17 April 1988; TU 157582 (35, 29 - 46), 22 March 1990; TU
157751 (29, 30 - 44), 11 April 1990, distributed as follows: TU 157751 (19),
UMMZ 222916 (10); TU 161920 (22, 30 - 45), 24 November 1991; TU 162361
(11,30-40), 2 March 1992; TU 162615 (15, 34 - 45), 31 March 1992; and TU
166409 (35, 34 - 49), 7 February 1993; TU 158313 (2, 40 - 46), Dry Creek at
Hwy. 25, 0.9 mi. S Vineland (T12N, R4E, Sec 10), 26 May 1990; TU 158293
(26, 32 - 46), Little Creek, tributary to Dry Creek at Hwy. 25, 0.3 mi. N Vineland
(T12N, R4E, Sec 3), 26 May 1990; TU 161862 (13, 30-41), 24 November 1991;
and TU 162377 (8, 33 - 39), 2 March 1992; TU 161881 (6, 33 - 38), Baptizing
Creek, tributary to Turkey Creek at Hwy. 25, 1.4 mi. N Vineland (T13N, R4E,
Sec 35), 24 November 1991; and TU 162601 (1, 34), 31 March 1992; TU 161904
(5, 32 - 40), Turkey Creek at Hwy. 25, 3.5 mi. N Vineland (T13N, R4E, Sec
23), 24 November 1991.

Additional material examined, not designated as types: Alabama River system.
Alabama. Baldwin-Monroe Cnty.: TU 32551 (1), Little River at Hwy. 59 at Chrysler, 13.5 mi.
W Uriah (T4N, R4E, Sec 19), 30 May 1964, TU 44399 (1), 17 March 1967, and TU 44419 (1), 18
March 1967. Escambia-Monroe Cnty.: TU 99950 (29), Little River at Escambia Cnty. Rd. I, 7.9
mi. NW McCullough (T3N, R5E, Sec 7), 20 October 1976; TU 153964 (10), Little River at Hwy.
21, 4.1 mi. S Uriah (T3N, R6E, Sec 5), 4 November 1988, TU 154479 (4), 21 March 1989, and TU
162548 (1), 30 March 1992. Clarke Cnty.: TU 34012 (4), Sand Hill Creek 1.1 mi. W Choctaw
Bluff (T5N, R3E, Sec 27), 1 July 1964, and TU 41447 (8), 30 June 1966; TU 32500 (6), Big Reedy
Creek at Clarke Cnty. Rd. 19, 1.5 mi. N Choctaw Bluff (T5N, R3E, Sec 22), 28 May 1964; TU
125380 (3), 14 May 1982, TU 163242 (13), 11 May 1992, and TU 165506 (3), 30 August 1992; TU
165527 (1), Sizemore Creek at Clarke Cnty. Rd. 2 (T5N, R4E, Sec 7), 30 August 1992; TU 137048
(12), Silver Creek, 5 mi. W Chance or 3.5 mi. E Scyrene (T9N, R5E, Sec 17), 8 August 1984, and
TU 152495 (8), 26 May 1988. Monroe Cnty.: TU 43254 (3), Wallers Creek, 5.7 mi. E Eureka
Landing on Alabama River (T5N, R4E, Sec 33), 5 February 1967; TU 153937 (1), Randons Creek
at Monroe Cnty. Rd. 23, 5 mi. NW Frisco City (T6N, R6E, Sec 15), 4 November 1988; UAIC 2357
(27), Randons Creek, approximately 2 mi. S US Hwy. 84, 25 September 1966. Wilcox Cnty.: TU
32454 (24), Bear Creek at Wilcox Cnty. Rd. 1, 3.1 mi. NW Lower Peach Tree (TUN, R5E, Sec 33),
28 May 1964, TU 60866 (32), 16 December 1969, TU 87039 (11), 21 March 1974, and TU 107599
(2), 26 March 1976; TU 47391 (1), Alabama River at Tait Bar, River Mile 122.4 (TUN, R6E, Sec
24), 7 August 1967; TU 3420 (2), Pursley Creek at Hwy. 11, 3.4 mi. SW Camden (TUN, R7E, Sec
2), 3 June 1951; TU 3067 (2), tributary to Pursley Creek at Hwy. 10, 1.8 mi. E Camden (T12N,
R8E, Sec 27), 3 June 1951; TU 47379 (1), Alabama River at Reeves Bar, River Mile 128.5 (TUN,
R7E, Sec 5), 7 August 1967; TU 47769 (2), Alabama River at Lower Canton Bar, River Mile 156.7
(T13N, R7E, Sec 23), 18 August 1967; TU 2575 (19), Turkey Creek, tributary to Pine Barren Creek
at Hwy. 10, 0.8 mi. W Awin (TION, RUE, Sec 16), 2 June 1951, and TU 32606 (66), 30 May 1964;
TU 60890 (154), Chilatchee Creek at Hwy. 5, 0.3 mi. N Alberta (T15N, R7E, Sec 30), 18 December
1969, TU 121394 (14), 15 May 1981, TU 125405 (44), 15 May 1982, TU 140941 (23), 24 May 1985,
TU 145528 (4), 20 May 1986, TU 151640 (21), 9 April 1988, TU 153093 (12), 2 August 1988, TU
153195 (26), 3 August 1988, TU 154403 (17), 3 February 1989, TU 154740 (33), 9 May 1989, TU
157470 (22), 1 March 1990, TU 157591 (13), 22 March 1990, TU 162585 (7), 31 March 1992, TU
162984 (24), 27 April 1992, TU 163218 (17), 11 May 1992, and TU 166792 (20), 18 April 1993.
Dallas Cnty.: TU 159006 (2), Tatum Creek, tributary to Bogue Chitto Creek at Dallas Cnty. Rd.
11 (T16N, R8E, Sec 28), 18 August 1990; TU 158962 (3), Dry Creek, tributary to Bogue Chitto
Creek at Dallas Cnty. Rd. 3 (T16N, R8E, Sec 17), 18 August 1990; TU 158982 (2), Bogue Chitto
Creek at Dallas Cnty. Rd. 182 (T16N, R7E, Sec 12), 18 August 1990; UAIC 2402 (14), Soapstone
Creek at US Hwy. 80, 7 mi. E Selma, 7 December 1966; TU 32648 (30), White Oak Creek at Hwy.
41, 13.3 mi. NE Camden (T13N, RIOE, Sec 8), 30 May 1964, TU 35376 (61), 29 June 1964, TU
157494 (9), 1 March 1990, TU 158887 (25), 17 August 1990, and TU 162970 (53), 26 April 1992;
TU 158926 (5), Musk Creek at Hwy. 41, 2.9 mi. S Sardis (T15N, RUE, Sec 29), 17 August 1990;
TU 35171 (6), Pine Flat Creek (Six Mile Creek) at Hwy. 41, 6 mi. S Selma (T16N, RUE, Sec 32),
27 June 1964, and TU 158945 (1), 17 August 1990. Autauga Cnty.: UAIC 2043 (13), Swift Creek,
Autaugaville on Hwy. 14 (T17N, R14E, Sec 22), 31 May 1966; UMMZ 146523 (1), Beaver Creek,
5 mi. W Autaugaville (T17N, R13E, Sec 14), 13 June 1942, and UMMZ 146547 (1), 20 December
1942; UMMZ 146539 (6), tributary to Beaver Creek, 5 mi. W Autaugaville (T17N, R13E), 20 March

1994] New Species of Etheostoma 117

1942; UMMZ 124134 (1), Autaugaville (T17N, R14E); UAIC 2311 (19). Autauga Creek between
Oak Grove and White City (T19N, R15E, Sec 16), 1 July 1966. Cuilion Cm v.: TU 15760') (13),
Bear Creek, tributary to Benson Creek at Chilton Cnty. Rd. 15, 4.8 mi. ENE Maplesville (12 IN,
R13E, Sec 16), 23 March 1990; TU 27347 (4), Mahan Creek, tributary to Little Clahaba River at
Chilton Cnty. Rd. 3213, 0.7 mi. NW Jemison (T23N, R13E, Sec 15). 20 December 1961. Perrv
Cnty. UMMZ 212385 (3), Silver Creek, 1 mi. E Suttle at US Hwy. 43 (T18N, R9E, Sec 21), 6
February 1965; TU 3271 1 (39), Ellards Creek, tributary to Taylor Creek, tributary to (.ahaba River
at Hwy. 5. 10.6 mi. SW Brent or 1.3 mi. S Bibb Cnty. line (T21N. R8E, Sec 10), 31 May 1964, lU
151661 (30), 9 April 1988, TU 151809 (57), 16 April 1988. TU 162997 (28). 27 April 1992, TU
166615 (1), 26 March 1993, and TU 166783 (8), 18 April 1993. Bibb Cn i y.: TU 15297 (1), Cahaba
River, 2.2 mi. N Centreville (T23N, R9E, Sec 14), 17 May 1957, and TU 35085 (1) 26 June 1964;
TU 151818 (68), Little Schultz Creek, 7.2 mi. N Centreville (T24N, R9E, Sec 24), 16 April 1988,
TU 163006 (14), 27 April 1992, and TU 166774 (11), 18 April 1993; TU 19414 (1), Cahaba River
at Hwy. 27, 8.5 mi. N Centreville (T24N, RIOE, Sec 33), 8 September 1958; FU 106753 (9), Coffee
Creek, tributary to Cahaba River at Bibb Cnty. Rd. 24 at Blocton (T22S, R6W. Sec 24). 14 April
1978; TU 25994 (14), Six Mile Creek, tributary to Little Cahaba River at Hwy. 25, about 10 mi. NE
Centreville (T23N, RUE, Sec 6), 17 April 1962; TU 152178 (5), Cooperas Creek, tributary to Six
Mile Creek (T23N, RUE, Sec 16), 12 May 1988; TU 152158 (24), Six Mile Creek (T23N, RllE,
Sec 15), 12 May 1988; TU 106776 (2), Little Cahaba River at Bulldog Bend at Bibb Cnty. Rd. 65
(T24N. RllE, Sec 19), 14 April 1978. Shelby Cnty.: TU 27376 (7), tributary to Shoal Creek,
tributary to Little Cahaba River, 2 mi. NNW Montevallo (T22S, R3W, Sec 19), 21 December 1961;
UMMZ 198756 (3), Big Cahawba River, Helena (T20S, R3W, Sec 15), 20 May 1889; AUM 9571 (1),
Little Cahaba River at Coal Branch, 2.0 airmi. SE Cahaba Heights (T18S, Rl W, Sec 19), 30 October
1971. Jefferson Cnty.: UMMZ 168646 (38), Little Cahaba River at US Hwy. 78 at Leeds (T17S,
RIE, Sec 21), 3 September 1954, and UMMZ 177738 (56), 22 May 1956. St. Clair Cnty.: TU
166768 (6), Little Cahaba River at Oliver Rd. in Leeds (T17S, RIE, Sec 15), 18 April 1993.

Diagnosis: Etheostoma ramseyi is a member of subgenus Ulocentra as diag-
nosed by Bouchard (1977) and Bailey and Etnier (1988). Etheostoma ramseyi lacks
a premaxillary frenum and the head of the vomer usually has one or a few
slender teeth and therefore is typical of members of the Etheostoma duryi species
group of Bailey and Etnier (1988). Etheostoma ramseyi, like E. lachneri and E.
raneyi, lacks a red ocellus in the first interradial membrane of the spinous dorsal
fin. The unique features of nuptial male E. ramseyi are the size and disposition
of the dark lateral blotches along the lateral line, the presence of a series of
orange to russet blotches dorsad of the lateral blotches, the interconnection of
lateral blotches with respective dorsal saddle blotches, and the continuous
orange coloration of the ventral lateral part of the body and caudal peduncle
with dorsal extensions between posterior lateral blotches on caudal peduncle.
The lateral blotches are of moderate size and are ovoid to rounded quadrate in
shape.

Description: Etheostoma ramseyi reaches a maximum of 45 mm SL (females)
to 51 mm SL (males). Sexual dimorphism and apparent sexual maturity occur
at the end of the first year at a minimum of 29 mm SL for both females and
males. Frequency distributions of scale and fin-ray counts are presented in Ta-
bles 1-5. Number of vertebrae, branchiostegal rays, gill rakers, and preoper-
culomandibular pores are given in Tables 6-7. The lateral line is usually com-
plete with (40 -) 42 - 49 (- 53) scales, occasionally the last one or two scales are
unpored. Transverse scales number 12 (11 specimens), 13 (48), 14 (44), 15 (35),
16 (8), or 17 (4). Transverse scales from origin of second dorsal fin to anal fin
(an alternative count, see Bailey and Etnier 1988: 30) number 1 1 (10), 12 (33),

13 (17), 14 (2), mean of 62 counts, 12.2. Caudal peduncle scale rows number

14 (2), 15 (20), 16 (45), 17 (85), 18 (45), 19 (12), or 20 (3). Dorsal fin has (8 -)
10 - 1 1 (- 12) spines and (10 -) 1 1 - 12 (- 13) sofi rays. Anal fin has 2 spmes

118 Tulane Studies in Zoology and Botany [Vol. 29

and 6 - 7 (- 8), modally 7, soft rays. Pectoral fin has 13-15 rays. Vertebrae
number 37 (23), 38 (88), or 39 (19); branchiostegal rays number 4 — 4 (1), 5—3
(1), 5^4 (3), 5_5 (181), 5—6 (2), or 6—6 (1), and gill rakers on first arch
number 6 (6), 7 (20), 8 (43), 9 (78), 10 (36), or 11 (3). In general there is a
decrease in the number of gill rakers from the northern or headwater tributaries
southward to the southern or lower tributaries to the Alabama River system.
The sum of left and right preoperculomandibular pores is nearly constant at 18
(171 of 179 specimens), of the remainder, one has 16 pores, seven have 17
pores. The opercle, cheek, nape, and prepectoral area typically are covered with
exposed scales. Occasionally the opercle, cheek, or nape has slightly reduced
scale coverage and in a few individuals some of the cheek scales are embedded.
The belly is consistently covered with exposed scales whereas the breast is con-
sistently scaleless.

In 10 specimens examined for dentition, the vomer lacks teeth in one, the
head of the vomer bears a single slender tooth in three, there are two teeth in
two, three teeth in three, and four teeth in one, mean 2.0; all 10 lack palatine
teeth.

Proportional measurements appear in Tables 8-9. The 25 males represent
specimens from five different months: March (9 specimens). May (10), June (1),
July (2), and December (3). The ten females represent specimens from two
months: March (6) and May (4).

Colors of live and freshly preserved specimens, collected in February, March,
and May, are described from Beaver Creek, the type locality. We preface our
descriptive remarks by saying that perhaps E. ramseyi is the most variable of the
three new species in its nuptial coloration, especially when considering different
sub-populations. Nuptial males have orange coloration on nearly the entire
lower side of body and caudal peduncle. Except for extreme anterior part of
body the orange extends dorsad to the ventral margin of lateral brown blotches
(typically eight) and up to lateral line between lateral blotches. The orange from
the venter fuses or merges with orange blotches above the lateral line in the last
two interspaces on the caudal peduncle. The orange on the last interspace is
continuous with two elongate orange blotches at base of caudal fin. The penul-
timate, lateral, brown blotch, plus the next four anteriorly, have an overlay of
green that extends diagonally down from a respective dorsal saddle blotch (Fig-
ure 3). Occasionally the green overlay extends downward over the penultimate
lateral brown blotch to the midventral line. A similar extension of the next
anterior lateral blotch is rare. The lateral brown blotches are moderate in size
and are centered along the lateral line. There is a series of distinct oblong
blotches on the dorsal lateral area just above the lateral line. These occur be-
tween the diagonal green bars that extend from dorsal saddle blotches to the
lateral brown blotches along the lateral line. This series of blotches above the
lateral line grades from bright red or red-orange posteriorly to russet anteriorly.
As mentioned above, the last two red or red-orange blotches merge with the
orange of the ventral lateral area. The red or red-orange blotches are brighter
in large prenuptial males and in small nuptial males than in large nuptial males.
Moreover the posterior two blotches on the caudal peduncle in prenuptial males
are separate from the ventral lateral orange on the caudal peduncle. Thus the
ventral lateral coloration becomes more extensive and the blotches in the series
above the lateral line become darker with the approach of the height of nuptial
condition. Nuptial males have a variable amount of red or orange pigment high
on the dorsal lateral area on the otherwise pale areas between the dorsal saddle

1994] Nevj Species of Etheostoma 119

blotches. Additional information regarding coloration is given in the section on
variation. The dorsal saddle blotches (typically eight) are dark brown with an
overlay of green. The breast, gill membranes, cheek, lower part of opercle, and
side and anterior tip of snout are light to dark turquoise. There are dark brown
preorbital and suborbital bars, as well as several brown blotches along upper
margin of the opercle. The turquoise, red, black, yellow, and cream pigmenta-
tion on both the spinous and soft dorsal fins of the nuptial male E. ramseyi is
similar to that described for E. raneyi and E. lachneri. The caudal fin of E. ramseyi
nuptial male has turquoise edging on procurrent margins and light turquoise
on outer part of both upper and lower lobes. The two elongate orange blotches
at base of the caudal fin are prominent and are continuous with orange on
caudal peduncle. The anal fin is a uniform turquoise and seldom has a pale
"window" as is common in nuptial male E. lachneri. The pelvic fins are turquoise
with milky white tips on the anterior rays. The pectoral fins of nuptial males
from the type locality lack turquoise but do have some yellowish pigment along
the lower rays.

Nuptial females have some bright coloration although greatly reduced in
coverage compared to that of the nuptial male. Large females have brighter
coloration than small females and the smallest sexually mature females may
completely lack bright colors. A sample taken from White Oak Creek, Dallas
Cnty., Alabama on 26 April 1992 contained 39 females. These specimens were
examined in detail for presence of red or orange coloration, and were tallied
under six categories. Thirty eight of the 39 specimens had several blotches of
orange in the lateral series just above lateral line; 37 of 39 had the two basi-
caudal orange blotches; 34 had one or more red spots in spinous dorsal fin; 24
had some orange on lower side of body; 22 had some red spots in second dorsal
fin; and only six of the 39 females had some flecks or small blotches of red or
orange high on the upper side between the dorsal saddle blotches.

Distribution: Etheostoma ramseyi is distributed mosdy below the Fall Line
in the Alabama River system but is relatively common in headwater tributaries
to the Cahaba River above the Fall Line (Figures 7, 9). No other species of
subgenus Ulocentra occurs within the range of E. ramseyi. We do not believe the
absence of E. ramseyi in the north-westerly flowing tributaries to the upper
Alabama River and the lower Tallapoosa River below the Fall Line is an artifact
due to lack of collecting effort. The reason for the absence in this area is not
fully understood, but perhaps it is because this area represents the eastern exten-
sion of the Black Belt district and there is limited suitable habitat (Harper, 1943).

Habitat and biology: Beaver Creek at the type locality has a diversity
of habitats. In general there is an alternation of pools and riffles, raceways or
cascades. Pools vary from one to three feet in depth during low flow and have
sand or bedrock substrates. The substrate elsewhere is gravel, sand, rock, clay
rubble, or bedrock. There are patches o( Justicia on some of the small gravel
bars. The stream is 15 to 20 feet in width at low flow and is bordered by steep
banks, especially at the road crossing. The banks are wooded and thus the
stream is shaded during the summer months.

Based on recent observations, spawning occurs from early March to late
April or early May at the type locality, however the spawning period may extend
earlier into late February at some more southern localities or later into May
farther north. Air temperature at any particular time has little correlation with
reproductive activities. However, water temperature is direcdy related to nuptial

120 Tulane Studies in Zoology and Botany [Vol. 29

development and spawning activities. Although water temperatures are related
to air temperature, the proximity of spring flow, nature of stream substrate, and
the vegetation coverage in the watershed, particularly in the riparian strip, en-
hance or decrease the impact on stream temperature. Within the same 24 hour
period (17-18 April 1993) water temperature of Litde Cahaba River at Leeds,
St. Clair Cnty. (above the Fall Line) was 15° C, Little Schultz Creek, 7.2 mi. N
Centreville, Bibb Cnty. was 17° C, Ellards Creek, 10.6 mi. S Centreville, Perry
Cnty., was 16° C, Chilatchee Creek, 0.3 mi. N Alberta, Wilcox Cnty. was 20° C,
and at Beaver Creek, the type and most southern locality, was 14° C. Most
spawning is completed before the water temperature reaches 20° C. Short
periods of 20° C may not terminate spawning activities but when temperatures
of 20° C or higher are sustained then termination of spawning is almost sure to
occur.

The sex ratio is skewed toward more females in spawning aggregations.
Based on five collections of topotypes taken during February to May, the 259
specimens include 91 (35.1%) males and 168 (64.9%) females. The males range
from 29.1 to 49.2 mm SL, mean 39.8 mm; the females from 28.9 to 44.2 mm
SL, mean 37.4 mm.

The species associates of E. ramseyi at the type locality are as follows:
Ichthyomyzon gagei, Moxostoma erythrurum, M. poecilurum, Campostoma sp., Hybog-
nathus nuchalis, Luxilus chrysocephalus, Lythrurus bellus, Nocomis leptocephalus, Not-
ropis ammophilus, N. baileyi, N. buccatus, Pimephales notatus, Semotilus atromaculatus,
Ictalurus punctatus, Noturus funebris, N. leptacanthus, Fundulus olivaceus, Gambusia
affinis, Lepomis cyanellus, L. macrochirus, L. megalotis, Etheostoma nigrum, E. whipplei,
Percina maculata, and P. nigrofasciata.

Variation: Number of gill rakers is the only meristic character that shows
some geographic variation. We did not detect any geographical trends in the
morphometries that are presented in Tables 8 and 9. Although not fully inves-
tigated by us, we believe that nuptial males in the upper part of the Cahaba
River system display more and larger blotches of red on the dorsal lateral area
of body, between the dark brown dorsal saddle blotches, than the lower Cahaba
and Alabama river populations. Females display a similar trend. Also we ob-
served that, in some samples, both males and females were darker than in other
samples, however, we believe this to be a direct correlation with substrate color
and perhaps water clarity.

Etymology: We take pleasure in naming this darter for John S. Ramsey,
who contributed to our knowledge of the systematics, zoogeography, and biol-
ogy of the darters and other fishes in the southeastern United States. Dr. Ramsey
generously contributed his field notes and preliminary analyses.

Comparisons

The typical nuptial coloration of males of Etheostoma raneyi, E. lachneri, and
E. ramseyi is illustrated in Figures 1-3 respectively. The lateral stripe of E. raneyi
is composed of two sets of blotches. One set of oblong or elongate blotches
aligned along the lateral line is more or less interposed by a second set of oval
or quadrate blotches. Although blotches in both of these series are variable in
shape and density of pigmentation, their ventral borders form a continuous
rather smooth margin. The entire lower side below this margin is orange to
red-orange (Figure 1). Etheostoma ramseyi has a single series of blotches along the

1994] New Species of £^/i^o5/or«a 121

lateral line and the orange or red-orange of the lower side extends dorsad
between the blotches, that is, covering the interspaces and thus producing a
scalloped pattern along the lateral blotches. The second set of blotches occurs
well above the lateral line. Occasionally the penultimate blotch along the lateral
line is continuous with green or turquoise coloration on the ventral part of the
caudal peduncle (Figure 3). Etheostoma lachneri is unique in the presence of two
to five complete, slightly oblique green bars on the posterior half of the body.
Those males with fewer than four complete bars usually have one or more
incomplete bars. The interspaces between the bars and the anterior ventral
lateral area of body are deep orange. The orange coloration above the lateral
line is not in the form of compact blotches but instead is an irregular network
of color (Figure 2).

The illustration of the male E. raney'i (Yazoo darter) in Deacon et al. (1979:
42) does not show the typical bright nuptial coloration. The color illustration of
the male E. raneyi (Yazoo darter) in Page and Burr (1991 : plate 43) is an accurate
depiction of the lateral blotches and the ventral lateral coloradon, however, the
spinous dorsal fin does not have a complete band of red blotches across the
entire fm as shown in their illustration. Also there is an incomplete band of black
blotches (not shown in their illustration) along the proximal part of anterior
membranes of the spinous dorsal fm.

We are somewhat perplexed with the illustration of the "Coastal Plain darter"
(Page, 1983: plate 16G). The locality data given are Minter Creek, Greene Cnty.,
Alabama and date of collection 17 April 1977. Our first collection of 13 speci-
mens (4 males) from Minter(s) Creek, Greene Cnty., Alabama was obtained on
26 March 1993. All four males had the green-barred pattern as shown in Figure
2. Our second collection (8 specimens) from the same site was taken on 17 April
1993 and the three males also had the barred pattern, although none was as
developed as illustrated in Figure 2. The pale "window" in the anal fin, the
noticeably elevated spinous dorsal fin, and the network of orange coloration
above the lateral line lead us to confirm the Page (1983: plate 16G) illustration
as Etheostoma lachneri. We suspect that the photo was taken quite a while after
capture and as a result much of the green and turquoise color had already
faded. We are unaware of any published illustradons of Etheostoma ramseyi.

A review of the eight scale, fin-ray, and vertebral counts (Tables 1-6) reveals
that Etheostoma raneyi, E. lachneri, and E. ramseyi are quite similar in these meris-
tics. The number of gill rakers on first arch in E. raneyi ranged from 8 to 10,
mean, 9.0; the number in both E. lachneri and E. ramseyi ranged from 6 to 11.
means, 8.1 and 8.7 respectively (Table 7).

Thirteen of nineteen proportional measurements on 25 adult males and 10
adult females of each of the three species (Tables 8-9) show differences that we
interpret as independent species variadon and not a clinal geographic pattern
of a single species. For males, the westernmost species, E. raneyi, is intermediate
between E. lachneri and E. ramseyi in eight of thirteen proportions. Etheostoma
raneyi has the longest head, anal soft ray, pelvic fins, and greatest trans-pelvic
width but the shortest soft dorsal fin. Etheostoma lachneri, which is intermediate
geographically, has the longest or greatest measurement for eight of the propor-
tions (longest dorsal spine, soft dorsal fin, dorsal soft ray, anal fin, first anal
spine, caudal fin, pectoral fin, and greatest body depth); is intermediate in
length of head and length of longest anal soft ray; is equal to E. ramseyi in length
of pelvic fins; and has the shortest caudal peduncle, and least trans-pelvic width.
Etheostoma ramseyi has the shortest or least average measurement for nine of the

122 Tulane Studies in Zoology and Botany [Vol. 29

thirteen proportions (longest dorsal spine, dorsal soft ray, anal fin, first anal
spine, anal soft ray, caudal fin, pectoral fin, head length, and the least body
depth measurement); intermediate in length of soft dorsal fin and trans-pelvic
width, equal to E. lachneri in length of pelvic fins; and has the longest caudal
peduncle.

For females (Table 9), Etheostoma raneyi has the longest or greatest measure-
ment for 1 1 of the thirteen proportions (longest head, dorsal spine, dorsal soft
ray, caudal peduncle, anal fin, first anal spine, and soft ray, caudal fin, pelvic
fins, greatest body depth, and trans-pelvic width) but intermediate pectoral fin
length. Etheostoma lachneri (intermediate geographically), has the shortest or least
measurement for 12 of the 13 proportions (recall that E. lachneri males have the
longest or greatest measurements of the three species for eight of the 13 propor-
tions). Females of all three species have nearly identical soft dorsal fin length
measurements. Etheostoma ramseyi females have the longest pectoral fins.

Comparison of species in a five-member subgroup: We here propose that
Etheostoma raneyi, E. lachneri, E. ramseyi, E. colorosum, and E. tallapoosae (Figures
1-5) form a subgroup of the E. duryi species group of Bailey and Etnier (1988).
The combination of the absence of the red ocellus in the first membrane of the
spinous dorsal fin, the incomplete band of red blotches in the spinous dorsal fin
(primarily on posterior membranes only) and the incomplete band of black
blotches in the spinous dorsal fin (on all anterior membranes but lacking on
posterior few) unites these five species but at the same time distinguishes them
from all other described species of the subgenus (Ulocentra). Separate principal
components analyses (PCA) were performed on morphometric data for 25 males
and 10 females of the three new species plus E. tallapoosae and E. colorosum to
characterize overall shape relationships. The analyses were performed with
Statistical Analysis System (SAS) software. Log-transformed data for 19 mor-
phometric variables were first regressed on standard length and the residuals
retained for PCA. This partial regression procedure effectively removed a
strong size-related trend in the data evidenced by highly inter-correlated vari-
ables and a high degree of variability explained by the first principal component
axis (PCI). After partialling out the effect of standard length, variables were
largely uncorrected and PCI accounted for only 34% of the variability in the
data for males and 24% of the variability for females.

Results of the PCAs are presented in Figure 8 as polygons representing
projections of data for males and females of each of the five species in the space
of the first two principal components. In both sets of plots, species are separated
mainly along PCI which relates primarily to fin dimensions (positive values
representing long and elevated fins). There is considerable overlap among
species, but overlap is greater among males than among females. The greatest
separation among species is between E. tallapoosae and E. colorosum, with the
latter tending to have the longer and more elevated fins. Also of interest is that
E. colorosum is completely separate (both males and females) from E. tallapoosae
and E. ramseyi in shape characteristics (Figure 8) although the latter two species
are close geographical associates in an adjacent drainage system. The new
species exhibit considerable overlap in shape characteristics with E. ramseyi being
more similar to E. tallapoosae, and E. lachneri and E. raneyi being intermediate
between the two extremes.

The five species of the subgroup are allopatric and no one of them is known
to be sympatric with any other species of subgenus Ulocentra (Figure 9). Two (E.
raneyi and E. colorosum) inhabit discrete drainage areas. The other three live in
separate areas of the Mobile Basin; of these E. lachneri lives in the Coastal Plain

1994]

New Species of Etheostoma

123

0 —

MALES

tallapoosae

colorosum

raneyi

FEMALES

tallapoosae ramseyi

Figure 8. PCA polygons representing projections of morphometric data for Etheostoma raneyi, E.
lachneri, E. ramseyi, E. tallapoosae, and E. colorosum.

to the Fall Line in the Tombigbee River system, E. ramseyi occurs in the Alabama
River system largely below the Fall Line, but crosses that line in the Cahaba
River system. Etheostoma tallapoosae inhabits the Tallapoosa River system above
the Fall Line. Six additional species of subgenus Ulocentra (one yet unnamed)
occur in the Mobile River system, all above the Fall Line.

124

Tulane Studies in Zoology and Botany

[Vol. 29

33"

31"

G U

O F

SCALE OF MILES

M t: X Y

c o

91

89

87

85

Figure 9. Distribution of five darters of duryi species group oi Etheostoma subgenus Ulocentra: E.
raneyi, E. lachneri, E. ramseyi, E. colorosum, and E. tallapoosae in Mississippi, Alabama, and parts of
adjacent states.

Acknowledgments

Study materials and distributional records of the three new snubnose darters
have been accumulating over the years, due to the efforts of many individuals
including past students and faculty of Auburn University, Cornell University,
Mississippi State University, Tulane University, University of Alabama, Univer-
sity of Michigan, University of New Orleans, and University of Tennessee. We
extend our thanks to Glenn H. Clemmer (formerly at Mississippi State Univer-
sity) and John S. Ramsey (formerly at Auburn University) for the loan of mate-
rials under their care.

1994] New Species of Etheostoma 125

We are indeed grateful to Maurice F. Mettee, Geological Survey of Alabama,
and J. Malcolm Pierson, Alabama Power Company, for their participation in
collecting and photographing specimens and the donation of color slides. We
are indebted to David A. Etnier for his generosity in allowing us to utilize and
distribute specimens formerly housed at the University of Tennessee, and for
the loan of additional series. We thank James D. Williams for bringing to our
attention the various U. S. Fish and Wildlife reports on fishes in northern Mis-
sissippi. We extend our thanks to two reviewers, Herbert T. Boschung and
David A. Etnier, for their many helpful suggestions.

Literature Cited

Bailey, R. M. and D. A. Etnier. 1988. Comments on the subgenera of darters (Percidae) with

descriptions of two new species of Etheostoma (Ulocentra) from southeastern United States. Misc.

Publ. Mus. Zool. Univ. Michigan 175: 1-48.
Boschung, H. T. 1987. Physical factors and the distribution and abundance of fishes in the upper

Tombigbee River system of Alabama and Mississippi, with emphasis on the Tennessee- Tombig-

bee Waterway. In W. J. Matthews and D. C. Heins, eds. Community and Evolutionary Ecology

of North American Stream Fishes (pp. 184-192). Univ. Oklahoma Press, Norman.
Boschung, H. T. 1989. Atlas of fishes of the upper Tombigbee River drainage, Alabama-Missis-
sippi. Proc. Southeast. Fishes Council 19: 1-104.
Boschung, H. T. 1992. Catalog of freshwater and marine fishes of Alabama. Bull. Alabama Mus.

Nat. Hist. No. 14 (266 pp.).
Bouchard, R. W. 1974. The subgenus Ulocentra (Percidae, Etheostomatini) in western Kentucky,

western Tennessee and northern Mississippi. Assoc. Southeast. Biologists Bull. 21:41 (abstract).
Bouchard, R. W. 1977. Etheostoma etnieri, a new percid fish from the Caney Fork (Cumberland)

River system, Tennessee, with a redescription of the subgenus Ulocentra. Tulane Stud. Zool. Bot.

19: 105-130.
Clemmer, G. H., R. D. Suttkus, and J. S. Ramsey. 1975. A preliminary checklist of endangered

and rare fishes of Mississippi. In Mississippi Game and Fish Comm. Preliminary list of rare and

threatened vertebrates in Mississippi (pp. 6-1 1). Jackson, Mississippi.
Deacon, J. E., G. Kobetich, J. D. Williams, and S. Contreras. 1979. Fishes of North America

endangered, threatened, or of special concern: 1979. Fisheries 4(2): 29-44.
Gilbert, C. H. 1891. Report on explorations made in Alabama during 1889 with notes on the

fishes of the Tennessee, Alabama, and Escambia rivers. Bull. U. S. Fish Commission (1889) 9:

143-159.
Harper, R. M. 1943. Forests of Alabama. Geological Survey of Alabama Monograph 10(230 pp.).
Hubbard, W. D. 1987. Distribution and diversity of fishes in the Noxubee River drainage, Missis-
sippi. J. Mississippi Acad. Sci. 32: 19-29.
Hubbard, W. D., C. E. Tucker, and H. Boschung. 1991. Fishes of the Sucarnoochee River

system, Alabama and Mississippi. Proc. Southeast. Fishes Council 24: 1-11.
HuBBS, C. L. AND K. F. Lagler. 1958. Fishes of the Great Lakes region. Cranbrook Inst. Sci. Bull.

26 (213 pp.).
Jenkins, R. E. 1976. A list of the undescribed freshwater fish species of continental United States

and Canada, with additions to the 1970 checklist. Copeia 1976: 642-644.
Knight, S. S. and C. M. Cooper. 1987. Fishes of Otoucalofa Creek, Missis.sippi prior to major

channel modifications. J. Mississippi Acad. Sci. 32: 31-38.
Kuehne, R. a. and R. W. Barbour. 1983. The American Darters. Univ. Kentucky Press,

Lexington (201 pp.).
Lowe, E. N. 1925. Geology and mineral resources of Mississippi. Mississippi State Geol. Survey

Bull. 20 (140 pp.).
Mettee, M. F. 1978. The fishes of the Birmingham-Jefferson County region of Alabama with

ecologic and taxonomic notes. Geol. Survey Alabama Bull. 115 (182 pp.).
Mettee, M. F., S. C. Harris, and P. E. O'Neil. 1986. Biological diversity and water quahty of

streams draining the Gilbertown oil field area. Geol. Survey Alabama Circular 120 (32 pp.).
Mettee, M. F., P. E. O'Neil, and S. C. Harris. 1983. A biological inventory of streams draining

the Citronelle, Pollard, and Gilbertown oil fields in Alabama. Geol. Survey Alabama Circular 108

(101 pp.).
Mettee, M. F., P. E. O'Neil, J. M. Pierson, and R. D. Suttkus. 1989a. Fishes of the Black

Warrior River system in Alabama. Geol. Survey Alabama Bull. 133 (201 pp.).

126 Tulane Studies in Zoology and Botany

Mettee, M. F., p. E. O'Neil, J. M. Pierson, and R. D. Suttkus. 1989b. Fishes of the western

Mobile River basin in Alabama and Mississippi. Geol. Survey Alabama Atlas 24 (170 pp.).
Mettee, M. F., P. E. O'Neil, R. D. Suttkus, and J. M. Pierson. 1987. Fishes of the lower

Tombigbee River system in Alabama and Mississippi. Geol. Survey Alabama Bull. 107 (186 pp.).
O'Neil, P. E., S. C. Harris, and M. F. Mettee. 1984. The distribution and abundance of aquatic

organisms inhabiting streams draining the Citronelle oil field, Alabama. Geol. Survey Alabama

Circular 99 (26 pp.).
Page, L. M. 1983. Handbook of Darters. T. F. H. Publications, Inc. Neptune City, New Jersey (271

pp.).
Page, L. M. and B. M. Burr. 1991. A Field Guide to Freshwater Fishes, North America, North

of Mexico. Houghton Mifflin Co., Boston, Mass. (432 pp.).
Pierson, J. M., W. M. Howell, R. A. Stiles, M. F. Mettee, P. E. O'Neil, R. D. Suttkus, and

J. S. Ramsey. 1989. Fishes of the Cahaba River system in Alabama. Geol. Survey Alabama Bull.

134 (183 pp.).
Randolph, K. N. and M. L. Kennedy. 1974. The fishes of the Tippah River system Mississippi,

with notes on habitats and distribution. J. Mississippi Acad. Sci. 19: 128-134.
Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W.

B. Scott. 1991. Common and scientific names of fishes from the United States and Canada

(5th ed.). Amer. Fish. Soc. Spec. Publ. 20: 1-183.
Simons, A. M. 1992. Phylogenetic relationships of the Boleosoma species group (Percidae: Etheos-

toma). In R. L. Mayden (ed.), Systematics, Historical Ecology, and North American Freshwater

Fishes (pp. 268-292). Stanford Univ. Press.
Smith-Vaniz, W. F. 1968. Freshwater Fishes of Alabama. Auburn Univ. Agric. Experiment Sta.

(211 pp.).
Suttkus, R. D. and R. M. Bailey. 1993. Etheostoma colorosum and E. bellator, two new darters,

subgenus Ulocentra, from southeastern United States. Tulane Stud. Zool. Bot. 29: 1-28.
Suttkus, R. D. and H. T. Boschung. 1990. Notropis ammophihis, a new cyprinid fish from south-
eastern United States. Tulane Stud. Zool. Bot. 27: 49-63.
Suttkus, R. D. and D. A. Etnier. 1991. Etheostoma tallapoosae and E. brevirostrum, two new darters,

subgenus Ulocentra, from the Alabama River drainage. Tulane Stud. Zool. Bot. 28: 1-24.

FISHES OF THE BUFFALO RIVER SYSTEM,
WILKINSON COUNTY, SOUTHWESTERN MISSISSIPPI

Mark A. Warren

AiLUiti Community College, Cedar Park, TX 78613

Robert C. Cashner

Department of Biological Sciences, University of New Orleans,
New Orleans, LA 70148'

AND

Royal D. Suttkus

Museum of Natural History, Tulane University,
Belle Chasse, LA 70037

Abstract

The occurrence and distribution of the fishes of the Buffalo River in south-
western Mississippi were determined from two major surveys conducted from
1968-1971 and 1986-1991, as well as museum and literature records. In all, 95
species, from 20 families and 50 genera, were documented from the Buffalo
River system. Cyprinids, centrarchids, poeciliids, fundulids, and clupeids were
numerically dominant in both surveys. Longitudinal distribution of fishes in
Buffalo River showed progressive downstream addition, with a large-scale re-
placement in the lower reaches. Monthly collections were made at six permanent
stations from September 1986 to October 1987. Community similarity indices
revealed relatively low faunal similarity between upper and lower sections of
Buffalo River, and a strong pattern of seasonal variation in the lower section of
the river.

Introduction

The importance of the Mississippi River in fish biogeography has been dis-
cussed in detail (Guillory, 1978, 1982; Grady et al., 1983; Conner and Suttkus,
1986). Fishes have been surveyed in several tributaries to the eastern part of the
Mississippi River in Mississippi and Louisiana (Guillory, 1981, 1982; Grady et
al., 1983; Cashner et al., 1979; Grady and Cashner, 1988; Ebert et al., 1985;
Matthews, 1978), but only a few of the detailed results of these surveys have
been published.

Buffalo River (or Buffalo Bayou) is the lowermost eastern major tributary
to the Mississippi River. Fish collections were made sporadically in the Buffalo
River by R. D. Suttkus and students at Tulane University in the mid-1950's and
early 1960's. The first comprehensive survey of the drainage was from
November 1968 to July 1971, in which 68 collections were made at 17 sites
located primarily in the upper and middle reaches of the river (Cashner et al.,
1976). The present study included a second major survey conducted from Sep-
tember 1986 to November 1991. During this period, 140 collections were made

Tulane Studies in Zoology and Botany 29: 127-139. 1994.

127

128

Tulane Studies in Zoology and Botany

[Vol. 29

at 60 different localities; 82 collections were made during monthly samples at 6
stations from September 1986 to October 1987. The two surveys represent a
combined total of 208 collections at 60 localities. The purpose of this paper is
to present a list of species for the Buffalo River drainage, to document the
pattern of distribution of these species, and to assess seasonal variation in the
fish assemblages at the six stations sampled monthly from 1986-1987. A paper
currently in preparation will compare the results of the two major surveys and
assess the changes in fish assemblages over a time span of nearly two decades.

Study Area

Except for extreme headwaters in Amite County, Buffalo River is within
Wilkinson County, Mississippi (Figure 1). The spring-fed headwaters of the
river originate in a forested area, 3.5 miles north of Centreville, Mississippi. The
river flows in a westerly direction, then north, increasing in size to a 4th-order
stream (based on Strahler, 1957) as it receives runoff from small creeks and its
first major tributary. Little Buffalo River. Buffalo River continues to flow north-
ward, gradually increasing in width and depth. Near the vicinity of the Missis-
sippi Highway 563 bridge, the river starts a northwesterly course and continues
in this direction until it divides into several channels and forms a series of bald
cypress dominated swamps near Lessley, Mississippi. Then the river flows south-
westerly to Lake Mary, an ox-bow lake formed from the Mississippi River, which
is also connected to the Homochitto River to the north. After Lake Mary, Buf-
falo River flows nearly due south, entering the Mississippi River near River Mile

Figure 1. Map of Buffalo River, Wilkinson County, Mississippi, with collecting localities indicated.

1994] Buffalo River Fishes 129

313, approximately 1.5 miles southwest of Fort Adams, Mississippi. The eleva-
tion of the Buffalo River is 380 feet at the headwaters and 25 feet at the conflu-
ence with the Mississippi River. The average gradient is 5.6 feet/mile and the
watershed area is approximately 344 square miles.

The river crosses several geological formations along its course. Headwaters
are characterized by deep cut banks, a narrow channel, relatively clear water,
and alternating pool, raceway, and riffle-type habitats. Here the stream exposes
the alluvial sands and gravels of the Citronelle formation (Brown and Guyton,
1943; Fisk, 1951; Sibley, 1972). Sand is the dominant substrate throughout the
upper headwaters, however, gravel is relatively abundant in riffle areas, and
hard packed clays are found in some raceways. Farther downstream, the river
traverses broad depositional Pleistocene terraces formed by Mississippi River
sediments (Fisk, 1951; Sibley, 1972). The terraces, as well as the Citronelle for-
mation to the east, are covered by loess soils (Snowden and Priddy, 1968). In
this section, the river becomes larger, wider, more turbid, and mixed gravel
substrates are replaced by shifting sand and mud. The lower reaches of Buffalo
River cut through the recent alluvial floodplain of the Mississippi River (Fisk,
1944, 1951; Sibley, 1972). In the lower section, water levels fluctuate dramatic-
ally and are directly influenced by the Mississippi River. Slow, turbid water
flowing over soft mud substrate, rich in organic detritus, is characteristic of this
reach.

Terrestrial vegetation providing stream cover is abundant in the middle to
upper regions of Buffalo River. A mixed coniferous-hardwood forest makes up
most of the stream canopy. Aquatic vegetation is occasionally present in the
upper portions of the river, with blooms of floating pond-scum {Spirogyra sp.)
common during the spring and summer. Upland pine forest is gradually re-
placed by cypress-hardwood in the lower reaches of Buffalo River. Black willow
{Salix nigra), eastern cotton wood {Populus deltoides) and bald cypress {Taxodium
distichum) dominate. Aquatic vegetation is also abundant, with duck-weed {Lemna
sp.), smart-weed (Polygonum sp.), duck-potato (Sagittaria latifolia), coon-tail
(Ceratophyllum demersum) and American lotus {Nelumbo luted) most prevalent
(Caplenor et al., 1968; Fassett, 1957).

Materials and Methods

At each locality, during both surveys, a 650-1300 foot longitudinal reach of
the stream was seined for approximately 1.0-1.5 hours or until all of the major
macrohabitats were sampled. Collections were made using seines 10-50 feet long
of 3/16 inch or 1/4 inch mesh. During the latter survey, in 1987, monofilament
nylon gillnets, 150 feet x 6 feet with a 2 inch square mesh, were set at stations
5 and 6 on two different occasions (Figure 1). Monthly collections, from Sep-
tember 1986 to October 1987, were made at six stations established from the
upper reaches of Buffalo River to near its confluence with the Mississippi River
(Figure 1).

Specimens were fixed in the field with 10% formalin and stored for one
week, then washed and transferred to 50% isopropyl alcohol for permanent
storage. Nomenclature used in this paper follows Robins et al. (1991), except
for the recognition of the families Fundulidae and Elassomatidae. All specimens
collected during this study were deposited in the Vertebrate Collection at the
University of New Orleans (UNOVC) or the Tulane Museum of Natural History
(TU). Detailed locations of all 60 collecting sites are given in the Appendix.

130 Tulane Studies in Zoology and Botany [Vol. 29

Analyses of Data

Morisita's Index (!„,) and percent similarity index (PSI) were used to assess
faunal similarity between stations 1-6. All species were included in the analysis
following Matthews (1986a). Indices of similarity have been widely used in com-
paring sites within and between streams. Although the ranges of values consid-
ered by different authors to be high vary somewhat, we follow Matthews et al.
(1988) in using values greater than 0.7 for PSI and 1^ as indicative of relatively
high degree of similarity between stations. Diversity indices (H') were calculated
(to the log base 2) for stations 1-6 (Shannon and Weaver, 1962).

Results

A total of 71,559 specimens of 74 species in 17 families was recorded from
the Buffalo River drainage in the 1968-1971 survey, and in the 1986-1991 sur-
vey, 60,830 specimens of 88 species in 17 families were collected. Based on the
two surveys, plus museum and literature records, the known ichthyofauna for
the Buffalo River presendy stands at 95 species in 20 families (Table 1).

Cyprinidae was the dominant family, both in number of species represented
(26% of total) and individuals (74% of total) collected during the two surveys.
The three numerically dominant species, Cyprinella camura, Notropis longirostris,
and Cyprinella venusta, made up 54% of the total specimens. The cyprinids were
followed by Fundulidae and Poeciliidae (5% each) and Centrarchidae (6%) in
number of individuals. The Percidae, with 17%, and the Centrarchidae, with
14%, ranked second and third, respectively, behind the cyprinids in number of
species represented.

During the survey of 6 permanent stations in 1986-1987, 82 collections were
made yielding 79 species in 15 families (Table 2). There was an obvious change
in the numerical dominance of species between the upper, middle, and lower
reaches of the Buffalo River (Table 2). At stations 1 and 2, Cyprinella camura and
Notropis longirostris were the most abundant and accounted for 54% of the speci-
mens captured. At stations 3 and 4, the numerical abundance of Cyprinella cam-
ura and Notropis longirostris decreased, and Cyprinella venusta and Hybognathus
nuchalis increased. These two species comprised 52% of the total specimens
collected at stations 3 and 4, compared with 26% for Cyprinella camura and
Notropis longirostris. In the lower portions of the river, at stations 5 and 6,
Dorosoma petenense, Gambusia affinis, and Lepomis macrochirus were the most abun-
dant forms taken.

There was virtually no similarity between the upper and middle stations 1-4,
(Figure 1) and the lower ones (5 and 6, Table 3). I^^ values among the upper
four sites ranged from 0.34-0.94. In all but a few cases the PSI estimates were
lower than I^, as predicted by Linton et al. (1981). Only stations 1 and 2 and 3
and 4 had both I^ and PSI values suggesting relatively high faunal similarity.
Based on the I^, alone, stations 1 and 2 and 1 and 3 could be regarded as highly
similar to each other. Stations 5 and 6 were the only other paired stations that
had Ij„ values near 0.70.

The most dissimilar adjacent stations were 4 and 5 (I^^ =0.20; PSI =0.17),
which had only 40% of their species in common. Cyprinella venusta and Hybog-
nathus nuchalis accounted for 56% of the specimens taken at Station 4, but only
5% at Station 5. Notropis texanus (11%) and Lythrurus fumeus (7%) were the most
common cyprinids at Station 5, and slow-water forms, such as Gambusia affinis,
Labidesthes sicculus, and Lepomis macrochirus, made up nearly 50% of the catch
(compared to about 7% at Station 4).

1994] Buffalo River Fishes 131

The number of species per site, for the six permanent stations, increased
gradually from 30 to 37 downstream from Station 1 to Station 4, which was
located near the end of the lower middle reaches of Buffalo River (Table 4).
The number of species increased to 47 at Station 5 and was virtually the same
at Station 6. Species diversity was highest at Station 5 (H' = 2.56); values at the
other 5 stations varied from 2.02 to 2.15.

Seasonal Variatioyi at Six Stations.

Similarity across seasons in the fish assemblages at the six stations was
examined for species collected monthly by seines from October 1986 to Sep-
tember 1987. The year was divided into four seasons, autumn (September-
November), winter (December-February), spring (March-May), and summer
(June-August) (Table 5). Streamwide, the species richness and abundance were
highest in the summer and autumn, with 61 species and 12,352 specimens, and
63 species and 8190 specimens, respectively. Fifty-eight species and 7398 speci-
mens were taken in spring, and the winter season yielded 3188 specimens, rep-
resenting 45 species.

The greatest stability in fish assemblages for the 12-month period occurred
at Station 2 (Table 5). The !„, values among all seasons were high for this station
(0.80-0.93). For Station 1, autumn-winter and winter-spring I„, values averaged
0.90, but the summer assemblage was much more variable, with I^^ for spring-
summer of 0.60 and 0.67 for summer-autumn. The faunal similarity at the
middle sites, stations 3 and 4, was also highest between autumn, winter, and
spring samples (x = 0.75) and lowest between the summer and the other seasons
(x = 0.48). The lower stations, 5 and 6, were the most variable between seasons.
Only autumn and winter samples at Station 5 had high similarity (0.92). The
comparisons of faunal assemblages for the other seasonal pairings at these two
stations ranged from 0.04 — 0.61 (x = 0.32). The greatest seasonal variability
was at Station 6, where \^ values for the four seasons averaged just 0.25.

Longitudinal Distribution of Fishes.

Species richness in Buffalo River increases from the headwaters to the
mouth. Thirty species were recorded at Station 1, and 51 by Station 4, where
the river channel becomes noticeably wider and deeper (Table 4). A rather
striking change in the fish fauna occurs between stations 4 and 5, where the
river takes on a more distinctive lowland character. Stations 5 and 6 had the
highest species number, with a total of 47 and 45 species, respectively. At these
two lower stations, 28 species not collected in the upper and middle portions
were added, bringing the total for the six permanent stations to 79.

Only six species appeared to be restricted to the upper reaches, and 14
others were more or less confined to the upper and middle portions, stations
1-4, of the stream course (Table 2). A group of 35 species was found primarily
in the lower reaches, at stations 5 and 6, and 13 species were distributed
throughout all of the six stations or were absent from only one site (Table 2).

Discussion

The general pattern of species addition has been documented in many
studies for rivers and streams in the eastern and central United States (e.g.,
Larimore and Smith, 1963; Sheldon, 1968; Jenkins and Freeman, 1972). The
Buffalo River exhibits a pattern of increased species richness downstream ac-

132

Tulane Studies in Zoology and Botany

[Vol. 29

Table 1. Abundance of fish species recorded from Buffalo River, Mississippi, in 1968-1971 and
1986-1991 surveys. Asterisk indicates species based on catch by commercial fishermen or
museum record.

Family and Species

Number of Specimens
1968-1971 1986-1991

Total

Petromyzontidae (lampreys)

Ichthyomyzon gagei

Lampetra aepyptera
Polyodontidae (paddlefishes)

Polyodon spathula*
Lepisosteidae (gars)

Lepisosteus oculatus

Lepisosteus osseus

Lepisosteus platostomus
Amiidae (bowfins)

Amia calva
Anguillidae (freshwater eels)

Anguilla rostrata
Clupeidae (herrings)

Alosa chrysochloris

Dorosoma cepedianum

Dorosoma petenense
Cyprinidae (carps & minnows)

Campostoma anomalum*

Cyprinella camura

Cyprinella lutrensis

Cyprinella venusta

Cyprinus carpio

Hybognathus hayi

Hybognathus nuchalis

Luxilus chrysocephalus

Lythrurus fumeus

Lythrurus umbratilis

Macrhybopsis storeriana

Nocomis leptocephalus

Notemigonus crysoleucas

Notropis atherinoides

Notropis blennius

Notropis buchanani

Notropis longirostris

Notropis shumardi

Notropis texanus

Notropis volucellus

Notropis winchelli

Opsopoeodus emiliae

Pimephales notatus

Pimephales vigilax

Semotilus atromaculatus
Catostomidae (suckers)

Carpiodes carpio

Erimyzon oblongus

Hypentelium nigricans

Ictiobus bubalus

Minytrema melanops

Moxostoma poecilurum
Ictaluridae (catfishes)

Ameiurus melas

Ameiurus natalis

Ictalurus furcatus

Ictalurus punctatus

Noturus gyrinus

Noturus miurus

15

9

24

12

2

14

0

5

5

2

36

38

0

2

2

0

3

3

1

6

7

14

115

129

366

3695

4061

0

1

1

22427

5328

27755

25

871

896

11304

7846

19150

0

3

3

2

22

24

340

6781

7121

2372

2843

5215

53

1051

1104

85

330

415

0

3

3

1569

1085

2654

7

358

365

15

57

72

0

182

182

1

0

1

18910

5987

24897

19

631

650

27

1715

1742

405

280

685

114

332

446

55

316

371

538

390

928

2061

1337

3398

57

267

324

21

11

32

62

20

82

79

18

97

9

156

165

2

44

46

32

17

49

0

6

6

26

108

134

1

1

2

9

126

135

0

10

10

1585

113

1698

1994]

Buffalo River Fishes

133

Table 1. (Continued)

Family and Species

Number of Specimens
1968-1971 1986-1991

Total

Notiuus phaeus

Pylodiclis olivatis
Esocidae (pikes & pickerels)

Esox amei'icaniL'i*
Aphredoderidae (pirate perches)

Aphredoderm sayaiius
Fundulidae (killifishes)

Fii ndiilits catenatus

Fundulus chrysotus

Funduliis dispar

Fundulus nolalus

Fundulus uUvaceus
Poeciliidae (livebearers)

GamhiLsia affinis
Atherinidae (silversides)

Labidesthes sicculus

Menidia bejyllina
Syngnathidae (pipefishes)

Syngjiathus scuvelU
Percichthyidae (temperate basses)

Morone chrysops

Morone mississippiensis

Morone saxatilis
Centrarchidae (sunfishes)

AmblopUtes ariommus

Lepomis cyanellus

Lepomis gulostis

Lepomis Immilis

Lepomis macrochirus

Lepomis megalotis

Lepomis microlophus

Lepomis punclatus

Lepomis symmetricus

Micropterus punctulatus

Micropterus salmoides

Pomoxis annularis

Pomoxis nigromaculatus
Elassomatidae (pygmy sunfishes)

Elassoma zonalum
Percidae (perches & darters)

Ammocrypta beani

Etheostoma chlorosomum

Etheostoma fusiforme

Etheostoma granle

Etheostoma histrio

Etheostoma lynceum

Etheostoma pan'ipinne

Etheostoma proeliaire

Etheostoma stigmaeum

Etheostoma swaini

Etheostoma whipplei

Percina caprodes

Percina sciera

Percina shumardi

Percina vigil

Stizostedion canadense
Sciaenidae (drums)

Aplodinotus grunniens

277

84

361

1

3

4

0

0

I

26

94

120

2652

1145

3797

0

533

533

0

372

372

13

171

184

1457

541

1998

639

78
46

6530

1301
415

22

7169

1379
461

26

2

61

63

0

5

5

0

7

7

2

0

2

107

84

191

25

62

87

15

30

45

461

4648

5109

975

508

1483

21

192

213

0

233

233

0

72

72

364

74

438

15

223

248

1

185

186

2

112

114

352

48

400

2

52

54

0

5

5

2

3

5

0

2

2

495

74

569

14

3

17

10

26

36

138

71

209

72

76

148

87

64

151

0

1

1

571

99

670

0

1

1

0

50

50

1

0

1

23

26

Total

132,396

134

Tulane Studies in Zoology and Botany

[Vol. 29

Table 2. Species and number of specimens collected during the 1986-1987 survey at six permanent
stations in Buffalo River, Mississippi.

Station
3 4

6 Total

Ichthyomyzon gagei
Lampetra aepyptera
Lepisosteus oculalus
L. osseus
L. platostomus
Alosa chrysochloris
Dorosoma cepedianum
D. petenense
Cyprinella camura
C. lutrensis
C. venusta
Cyprinus carpio
Hybognathus hayi
H. nuchalis
Luxilus chrysocephalus
Lythrurus furneus
L. umbratilis
Nocomis leptocephalus
Notemigonus crysoleucas
Notropis atherinoides
N. blennius
N. longirostris
N. shumardi
N. texanus
N. volucellus
N. winchelli
Opsopoeodus emiliae
Pimephales notatus
P. vigilax

Semotilus atromaculatus
Carpiodes carpio
Erimyzon oblongus
Hypentelium nigricans
Ictiobus bubalus
Minytrema melanops
Moxostoma poecilurum
Ameiurus natalis
Ictalums punctatus
Noturus gyrinus
N. miurus
N. phaeus

Aphredoderus sayanus
Fundulus catenatus
F. chrysotus
F. dispar
F. notatus
F. olivaceus
Gambitsia affinis
Labidesthes sicculus
Menidia beryllina
Syngnathus scovelli
Morons chrysops
M. mississippiensis
M. saxatilis
'Lepomis cyanellus
L. gulosus
L. humilis

4

4

1

2

1
1

18

7

23

1

6

78

2750

1

25

1

2

6

96

2757

1208

964

1094

945

4211

2

96

47

145

282

94

1515
2

2682

103

496

5172
2

1

4

4

9

538

843

2035

334

26

3776

211

635

91

54

991

6

14

590

58

668

12

274

8

294

144

438

5

587

3

7

4
11

187

28

182

201
39

182

1086

1364

742

807

631

3999
631

2

156

928

189

1275

1

14

232

7

254

24

2

71

152

256

38

249
294

38

63

43

81

3

228

1

145

416

117

80

759

10

2

12

1

1

9

11

3

1

2

6

3

11

2

41

151

16

151

41

2

6

6

14

5

1

6

1

2

16

1

2
4

9

30
5

19

67

86

3

67

70

1

1

26

28

310

363

30

17

429

273

12

720
441
273

2

3

141

15

161

107

94

50

69

2

322

12

8

260

459

1733

2587

5059

45

803

105

20

81

184

1

58
1
I

929

289

21

58

1
1

21

23

10

5

59

4

18

24

1
1

23
25

1994]

Buffalo River Fishes

135

Table 2. (Continued)

L. macrochinis

L. megalulis

L. microlophus

L. punctatiLs

L. symmetric us

Micropterus punctulatus

M. salmoides

Pomoxis annularis

P. nigromaculatus

Ammocjypta beani

Etheosloma chlorosomum

E. gracile

E. lynceum

E. pannpinne

E. proeliare

E. stigmaeum

E. swaiyii

E. whipplei

Percina caprodes

P. sciera

P. vigil

Aplodinotiis grunniens

Station
'^ 4

fi Total

21

45

32

56

1637

2505

4296

80

17

35

72

126
139

23

8

353

147

1

222
50

1

224
50

1

11

7

26

4

49

1

7

1

67
6

125

179
58

201

179

64

2

2

9

22
1

9

1
1

35

10

2

19

33
2

1
1

1

2

19

1

55
1

20
3

6

2

4

4

5

21

4

7

9

2

1

1

23

1

32

12

14

13

71

30

3

2

5

1

10

40

11

Totals

4234 4346 5328 8412 8388 10864 41572

Table 3. Community similarity indices for the six permanent stations for Buffalo River, Mississippi,
1986-1987. First value I„„ PSI in parentheses.

Station 12 3 4 5 6

1

—

2

.89(.72)

—

3

.77(.61)

.55(.43)

—

4

.60(.46)

.34(.28)

.94(.79)

...

5

.05(.08)

.02(.03)

.15(.13)

.20(.17)

6

.03(.06)

.02(.04)

.15(.12)

.16(.15)

.67(.50)

Table 4. Species richness and Shannon Diversity (H') for collections made during the 1986-1987
survey at the six permanent stations, Buffalo River, Mississippi.

Station

Species
Site

Total Species
for Drainage

\

11'

1
2
3
4
5
6

30

30

4234

2.114

33

38

4346

2.026

34

42

5328

2.077

37

51

8412

2.068

47

71

8388

2.563

45

79

10864

2.148

136 Tulane Studies in Zoology and Botany [Vol. 29

Table 5. Seasonal similarity (I,„) for the six permanent stations, Buffalo River, Mississippi, during
1986-1987.

Station

Aut-Win

Win-Spr

Spr-Sum

Sum-Aut

Mean

1

.91

.89

.60

.67

.77

2

.92

.80

.93

.90

.89

3

.75

.92

.43

.68

.70

4

.71

.61

.36

.44

.53

5

.92

.22

.27

.61

.51

6

.40

.37

.04

.18

.25

companied by an abrupt change in faunal components in the lowermost reaches.
The major replacement of species between stations 4 and 5 is associated with
the river becoming more lowland in character. A cypress swamp located at
Station 5 probably acts as a natural barrier to fishes adapted to higher gradient
conditions (Guillory, 1978; Matthews and Styron, 1981). Ichthyofaunal surveys
conducted in drainages adjacent to the Buffalo River have shown similar pat-
terns of species distribution (e.g., Guillory, 1982; Grady et al., 1983).

A downstream increase in species diversity occurs in most lotic systems
(Hynes, 1970; Hawkes, 1975; Horwitz, 1978), and this pattern is reflected in
samples at the six permanent stations on Buffalo River. Various authors have
suggested that such patterns are due to an increase in available habitat or trophic
diversity (Kuehne, 1962; Sheldon, 1968; Horwitz, 1978). Increasing stream size
and habitat heterogeneity, along with the less variable physicochemical environ-
ment found in the lower mainstream, produces a more temporally stable envi-
ronment, as compared to the headwaters, and could partially explain the high
species richness and diversity values seen for stations 5 and 6 (Sheldon, 1968;
Evans and Noble, 1979; Matthews and Styron, 1981; Schlosser, 1987). Some
authors have suggested that characteristics of local fish assemblages are strongly
related to stream order (Kuehne, 1962; Harrel et al., 1967; Whiteside and
McNatt, 1972), and that stream order represents a discrete biological unit for
fishes (Lotrich, 1973). We found no evidence of changes in fish assemblages
correlated with changes in stream order using methods described in Matthews
(1986b). The striking differences in faunal composition between stations 4 and
5 are partially enhanced by the distance between the sampling sites (21 miles)
rather than just a sudden change in environmental conditions. Later collections,
made between stations 4 and 5, revealed a more gradual change in the fish
assemblages.

Stations 5 and 6 exhibited the greatest faunal change when compared to the
other four permanent stations. Guillory (1982) concluded that transients from
the faunally rich Mississippi River and adjacent floodplain were largely respon-
sible for the high diversity at sites along Thompson Creek similar in habitat
complexity to stations 5 and 6.

The diversity and species recorded for Buffalo River (95) is comparable to
the richest faunas reported for tributaries to the lower eastern Mississippi River,
e.g., Thompson Creek (89 species; Guillory, 1981), Bayou Sara (80 species;
Grady et al., 1983), Homochitto River (105 species; Ebert et al., 1985), and
Bayou Pierre (95 species; Matthews, 1978).

Acknowledgments

We would like to thank R. Ary, G. Bueche, J. Craven, P. Denette, M. Farabee,
S. Herbert, J. Knight, F. McCormick, D. Roberts, Marianne Warren, and A.

1994] Buffalo River Fishes 137

Whitehurst for field assistance. R. Mayden and B. Kuhajda (UAIC), and J.
Caruso (TU) made specimens available for examination and confirmed species
identifications. S. Ross (USM) made available species records tor Buffalo River.
J. Dale (Mississippi CooperaUve Extension Office), C. Hollins (U. S. Soil and
Conservation Office), M. Plunkett (U. S. Geological Survey), and T. Reynolds
(U. S. Army Corps of Engineers) extended numerous courtesies. M. Stevenson,
P. Denette, and especially W.J. Matthews made helpful suggestions during the
course of the study and on the manuscript. Funding for this study was provided
in part by a grant from the UNO Research Council and NSF BSR 87707760 to
RCC.

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Appendix. Locations of collection sites in Mississippi. Stations 1-6 (1-6 in Figure 1) were sampled
monthly during 1986-1987, and at least one collection was made at the other 54 sites identified as
7-60 in Figure 1.

Station 1: Little Buffalo River at Macedonia Road, 2.8 miles west of Centreville, and 6.4 miles north
of Highway 24 (T2N, RIE, Sec 5). Station 2: Buffalo River at Hiram-McGraw Road, 2.8 miles west
of Centreville, 1.4 miles north of Highway 24 (T2N, RIE, Sec 34). Station 3: Buffalo River at
Highway 563, 0.5 miles south of Wilkinson (T3N, RIW, Sec 17). Station 4: Buffalo River at US
Highway 61, 8.5 miles north of Woodville (T3N, R2W, Sec 21). Station 5: Buffalo River at Highway
24, 5.1 miles west of Lessley (T2N, R4W, Sec 10). Station 6: Buffalo River at Jackson Point Road,
1.9 miles west of Fort Adams (T2N, R5W, Sec 18). Station 7: Little Buffalo River at Bluff Springs
Road, 3.4 miles west of Gloster (T3N, RIE, Sec 23). Station 8: Little Buffalo River at Lower Gloster
Road, 3.5 miles west of Gloster (T3N, RIE, Sec 26). Station 9: Unnamed tributary to Little Buffalo
River at Mt. Carmel Road, 3.3 miles west of Gloster (T3N, RIE, Sec 26). Station 10: Buffalo River
at Dr. Anderson Road, 3.9 miles north of Centreville (T2N, RIE, Sec 11). Station 11: Buffalo River
at Macedonia Road, 2.3 miles northwest of Centreville (T2N, RIE, Sec 24). Station 12: Smith Creek

1994] Buffalo River Fishes 139

at Macedonia Road, 1.2 miles northwest of Centreville (T2N, RIE, Sec 29). Station 13: liiifTalo River
at Willie Neyland Road, 1.5 miles north of Highway 24 (T2N, RIW, Sec 25). Station 14: |a(kson
Creek at White School Road, 1.0 miles north of Highway 24 (T2N, RIW, Sec 26). Station 15: Biili.ilo
River at Jones- White Road, ?>.{) miles north of Highway 24 ( r2N. RIW, Sec 24). Sution 16: Browns
Creek at White School Road, 3.5 miles north of Highway 24 ( r2N, RIW, Sec 15). Station 17: Buffalo
River at Little Buffalo River confluence, at White School Road, 5.0 miles north of Highway 24 ( r2N,
RIW, Sec 14). Station 18: Dry Fork Creek at Highway 563, 3.2 miles north of Woodville or 0.6 mile
north of Highway 61 at Highway 563 bridge (T2N, R2W, Sec 12). Station 19: Dixon Creek at Henley
Road, 9.0 miles north of Highway 24 (T3N, RIE, Sec 32). Station 20: Caledonia Creek at William
Anderson Road, 6.5 miles southwest of Wilkinson (T3N, RIW, Sec 40). Station 21: Caledonia Creek
at William Anderson Road, 5.0 miles southwest of Wilkinson ( I 3N, RIW, Sec 39). Station 22: Willis
Creek at Walter Anderson Road, 5 miles southwest of Wilkinson (T3N, RIW, Sec 30). Station 23:
Piney Creek at Highway 563, 3.0 miles northwest of Wilkinson (T3N, RIW, Sec 14). Station 24: Dry
Creek at Walter Anderson Road, 3.0 miles southeast of Wilkinson (T3N, RIW, Sec 25). Station 25:
Buffalo River at Dry Creek confluence, at Walter Anderson Road, 3.6 miles southeast of Wilkinson
(T3N, RIW, Sec 31). Station 26: Buffalo River at Otis Jackson Road, 2.0 miles southeast of Wilkinson
(T3N, RIW, Sec 17). Station 27: Silver Creek at Highway 563, 1.0 mile east of Wilkinson (T3N,
RIW, Sec 16). Station 28: Fords Creek at Fords Creek Road, 3.6 miles north of Woodville (T2N,
R2W, Sec 3). Station 29: Fords Creek at US Highway 61, 6.6 miles north of Woodville (T3N, R2W,
Sec 41). Station 30: Buffalo River at Mill Creek confluence, 1.4 miles west of US Highway 61 ( r3N,
R2W, Sec 22). Station 31: Phipps Creek at Doloroso Road, 4.0 miles east of Doloroso (T4N, R2W,
Sec 29). Station 32: Hazlit Creek at Doloroso Road, 0.7 mile east of Doloroso (T4N, R2W, Sec 26).
Station 33: Big Piney Creek at US Highway 61, 4.0 miles southeast of Doloroso (T3N, R2W, Sec 5).
Station 34: Buffalo River at Sanders Fork Road, 3.5 miles northeast of Lanehart (T3N, R2W, Sec
11). Station 35: Buffalo River 3.5 miles north of Lanehart (T3N, R2W, Sec 10). Station 36: Steels
Creek at Woodlawn Road, 7.5 miles northeast of Lanehart (T3N, R3W, Sec 12). Station 37: Beaver
Creek at Beaver Creek Road, 1.5 miles southwest of Lanehart (T3N, R3W, Sec 43). Station 38:
Buffalo River at Beaver Creek confluence, 2.0 miles north of Beaver Creek Road (T3N, R3W, Sec
28). Station 39: Buffalo River at Southland Road, 5.0 miles north of Lessey (T3N, R3W, Sec 25).
Station 40: Unnamed tributary to Buffalo River at Pleasant Valley Road, 3.0 miles north of Highway 24
(T3N, R4W, Sec 42). Station 41: Buffalo River at Pleasant Valley Road, 2.5 miles northwest of
Highway 24 (T3N, R4W, Sec 27). Station 42: Lake Mary at Percy Creek-Sand Road, bridge at public
boat ramp, 3.0 miles northwest of Highway 24 (T3N, R4W, Sec 36). Station 43: Unnamed cypress
swamp lake, 3.7 miles north of Percy Creek-Sand Road (T2N, R4W, Sec 6). Station 44: Percy Creek
0.1 mile south of Highway 24 at Lessley (T2N, R3W, Sec 17). Station 45: Percy Creek at Bullet
Branch confluence at Highway 24, 2.5 miles west of Lessley (T2N, R3W, Sec 12). Station 46: Hayes
Creek at Highway 24, 3.0 miles west of Lessley (T2N, R4W, Sec 17). Station 47: Percy Creek at
Percy Creek-Sand Road, 1.5 miles north of Highway 24 at Ford (T2N, R4W, Sec 6). Station 48:
Smith Creek at Highway 24, 5.0 miles northwest of Fort Adams (T2N, R4W, Sec 20). Station 49:
Buffalo River 0.6 mile south of- Percy Creek-Sand Road bridge (T2N, R4W, Sec 10). Station 50:
Bloomer Creek at Highway 24 bridge, 3.1 miles northeast of Fort Adams (T2N, R4W, Sec 21).
Station 51: Belmont Lake at unmarked road, 1.3 miles northeast of Fort Adams (T2N, R4W, Sec
41). Station 52: Belmont Slough at unmarked road, 3.1 miles northeast of Fort Adams (T2N, R4W,
Sec 27). Station 53: Buffalo River at confluence with Lake Mary Canal at unmarked road, 2.3 miles
west of Highway 24, 4.9 miles northeast of Fort Adams (T2N, R4W, Sec 1 1). Station 54: Buffalo
River at unmarked road, 1.5 miles south of confluence with Lake Mary Canal, 3.2 miles west of
Highway 24 (T2N, R4W, Sec 24). Station 55: Unnamed tributary to Buffalo River at Pond Road,
0.7 mile east of Highway 24 (TIN, R4W, Sec 11). Station 56: Buffalo River, 1.5 miles north of
Jackson Point Road, 1.9 miles west of Fort Adams (T2N, R4W, Sec 42). Station 57: Buffalo River.
1.0 miles north of Jackson Point Road, 1.9 miles west of Fort Adams (T2N, R5W, Sec 15). Station
58: Buffalo River, 0.5 mile north of Jackson Point Road, 1.9 miles west of Fort Adams (T2N, R5W,
Sec 19). Station 59: Buffalo River, 0.5 mile south of Jackson Point Road, 1.9 miles west of Fort
Adams (T2N, R4W, Sec 43). Sution 60: Buffalo River at Mississippi River confluence, 1.5 miles
southeast of Jackson Point Road, 1.9 miles west of Fort Adams (TIN, R4W, Sec 10).

FISHES OF THE TANGIPAHOA RIVER SYSTEM,
MISSISSIPPI AND LOUISIANA

Charles L. Knight

Mississippi Museum of Natural Science,
111 North Jefferson Street, Jackson, MS 39202

AND

Robert W. Hastings

Turtle Cove Environmental Research Station,
P.O. Box 585, Southeastern Louisiana University, Hammond, LA 70402

Abstract

An ichthyofaunal survey of the Tangipahoa River system in the Lake
Pontchartrain drainage was conducted over an 18 month period in order to
compile a comprehensive list of fish inhabitants, and to compare these data with
lists of species reported from other streams in the Lake Pontchartrain, lower
Mississippi River, and Pearl River drainages. Eighty-five collections from 59
sites yielded 11,081 specimens including 71 species and 21 families. Over 50%
of the total catch was represented by nine species of Cyprinidae followed by
Centrarchidae (17%) with 14 species. Cyprinella venusta, Lythrurus roseipinnis, and
Gambusia affinis were most abundant at 22.6%, 10.3%, and 9.6% of the total
catch, respectively. Lepomis macrochirus (69.4%), Lepomis megalotis (64.7%), and
Gambusia affinis (60.0%) displayed the greatest frequency of occurrence. Three
species collected, Alosa chrysochloris, Heterandria formosa, and Etheostoma par-
vipinne, were considered new records for the Tangipahoa River system. Fishes
identified during our study combined with records from other sources bring
the total species reported to inhabit the Tangipahoa River system to 92. Within
the Lake Pontchartrain drainage, coefficient of similarity calculations indicated
that the Tickfaw, Amite, and Tchefuncte rivers supported fish assemblages most
similar to the Tangipahoa, while comparisions with streams from neighboring
drainage systems suggested intermediate species similarity.

Introduction

The Mississippi River is an apparent barrier for east and west dispersal of
many species of freshwater fishes in the United States (Lee et al., 1980). Douglas
(1974) reported 25 species of fishes occurring in Louisiana exclusively east of
the Mississippi River. Two additional fishes which may be added to this list are
the northern studfish {Fundulus catenatus) (Laiche, 1978); and the broadstripe
topminnow {Fundulus euryzonus), a species that is endemic to the Tangipahoa
and Amite rivers of the Lake Pontchartrain drainage (Suttkus and Cashner,
1981).

Tulane Studies in Zoology and Botany 29: 141-150. 1994.

141

142

Tulane Studies in Zoology and Botany

[Vol. 29

Most streams in the Lake Pontchartrain drainage have had their
ichthyofauna surveyed within the past 20 years (e.g., Tickfaw River, Saul, 1974;
Bayou Lacombe, Sobczak, 1976, and Farabee, 1992; Amite River, Laiche, 1978;
Blind River, Watson et al., 1981) (Figure 1). Prior to our study, however, no
comprehensive survey had been conducted on the Tangipahoa River system. In
addition to the studies conducted in the Lake Pontchartrain drainage, ichthyo-
faunal surveys have been made in most tributaries of the neighboring lower
Mississippi River (e.g., Homochitto River, Ebert et al., 1985; Buffalo River,
Cashner et al., 1976; Bayou Sara, Grady et al., 1983; Thompson Creek, Guillory,
1974, 1981, 1982) and Pearl River (e.g., Bogue Chitto River, Dosher, 1976)
drainages.

Figure 1. The Lake Pontchartrain, lower Mississippi River, and Pearl River drainage in southwest
Mississippi and southeast Louisiana. (1. lower Mississippi River drainage, 2. Lake Pontchartrain
drainage, 3. Pearl River drainage; A. Homochitto River, B. Buffalo Bayou, C. Bayou Sara, D.
Thompson Creek, E. Comite River, F. Amite River, G. Tickfaw River, H. Tangipahoa River, L
Tchefuncte River, J. Bogue Chitto River, K. Bayou Lacombe, L. Blind River).

1994] Tangipahoa River Fishes 143

This study was conducted to provide a comprehensive list of fish species and
their relative abundances in the Tangipahoa River system, and to compare these
data with those from other streams in the Lake Pontchartrain, lower Mississippi
River, and Pearl River drainages in order to contribute to a greater understand-
ing of the zoogeography of this unique area.

Study Area

The Tangipahoa River originates in Lincoln County, Mississippi, and
empties into the Lake Pontchartrain estuary in Tangipahoa Parish, Louisiana.
The entire watershed drains approximately 1997 sq. km (Sloss, 1977).

Elevations range from approximately 97.5 m above sea level in the upper
reaches of the Tangipahoa River to sea level at the confluence with Lake
Pontchartrain. Mean annual precipitation is 152.4 cm, and mean temperature
is 19.4° C (Newton, 1972).

Historically, the upper basin was dominated by longleaf pine {Pinus palvstris)
(Brown, 1945). Recently, however, most of the longleaf pine forests have been
replaced by loblolly pine {Pinus taeda), a tree planted extensively by the logging
industry (Louisiana Department of Natural Resources, 1974). The lower 15 km
of the river is deep and sluggish as it enters bald cypress {Taxodium distichum)
and tupelo gum {Nyssa aquatica) swamps bordering Lake Pontchartrain.

The stream bed is characterized by areas of sand and sand mixed with gravel
in the upper to mid reaches, and primarily sandy clay and silt downstream.
Common aquatic macrophytes in this stream include eelgrass {Vallisneria
americana), various water lilies {Nymphaea spp.), water milfoil {Myriophylum
spicatum), and bushy- pond weed (Najas guadalupensis).

The Tangipahoa River is used for swimming, fishing, and canoeing. Lake
Tangipahoa, a 283 ha man-made lake in Percy Quin State Park, interrupts the
river near its headwaters in Pike County, Mississippi. The Louisiana Department
of Wildlife and Fisheries (LDWF) classifies the Tangipahoa as a "natural and
scenic" river (LDWF, 1981).

M'aterials and Methods

Most collections were taken by a combination of a Coffelt BP-IC backpack
electro-shocker and 0.65 cm mesh minnow seine. In areas too deep for these
methods to be effective, gill nets of varying mesh sizes were used.

Upon capture, specimens were fixed in 10% formalin for one week, soaked
in water for one day, and then transferred to 40% isopropyl alcohol for identifi-
cation and storage in the Southeastern Louisiana University Museum. Identifi-
cation was based on Cook (1959), Douglas (1974), Parker et al. (1972), and
Pflieger (1975). Nomenclature followed Robins et al. (1991). Collection records
made prior to this study, from the Mississippi Museum of Natural Science,
Southeastern Louisiana University Museum, and University of New Orleans
Museum, were examined for additional species.

Fifty-nine sites were sampled between 1 July 1985 and 15 December 1986,
and included 85 collections. Most sites were selected for accessibility and diver-
sity of habitat and were usually at bridge crossings of highways or backroads.
Care was taken to include the mainstream, all major tributaries, and all stream
orders. Six stream orders were recognized using 15' United States Geological

144 Tulane Studies in Zoology and Botany [Vol. 29

Survey topographical maps, according to the method described by Horton
(1945), and modified by Strahler (1954, 1957). Jaccard Coefficient of Similarity
(South wood, 1966) was calculated as a correlation of species similarity between
the Tangipahoa River and other rivers in the Lake Pontchartrain, lower Missis-
sippi River, and Pearl River drainages.

Results and Discussion

A total of 11,081 fishes was collected during this study and included 71
species and 21 families. Blacktail shiners {Cyprinella venusta) occurred in the
greatest numbers at 251 1 individuals, and 22.7% of the total catch. The cherry-
fin shiner {Lythrurus roseipinnis) and mosquitofish (Gambusia affinis) followed at
1136 (10.3%) and 1062 (9.6%), respectively. Bluegill (Lepomis macroc hints), most
frequently encountered in different collections, were observed 59 times at 69.4%
frequency of occurrence. Longear sunfish (Lepomis megalotis) were found in 55
collections (64.7%) while mosquitofish (G. affinis) were caught on 51 occasions
(60.0%).

Families represented by the greatest number of individuals were Cyprinidae
at 53.4% and Centrarchidae at 17.1% of the total catch. Families from which
the most species were collected were Centrarchidae with 14, and Cyprinidae and
Percidae each with 9 species. Approximately 94% of the total catch consisted of
six families (Cyprinidae, Centrarchidae, Poeciliidae, Percidae, Cyprinodontidae,
and Catostomidae).

Species number generally increased in stream orders 1 through 5, with 25,
39, 48, 48, and 54 species collected, respectively. In the sixth order reach where
sampling efforts proved less effective, 42 species were collected. Large numbers
of banded pygmy sunfish {Elassoma zonatum) were almost entirely restricted to
first order streams. Other fishes that tended to be more common in lower stream
orders were the creek chubsucker (Erimyzon oblongus) and green sunfish {Lepomis
cyanellus). Fishes showing a preference for the higher stream orders included
the bullhead minnow {Pimephales vigilax), inland silverside {Labidesthes sicculus),
mimic shiner {Notropis volucellus), threadfin shad (Dorosoma petenense), and clear
chub {Notropis winchelli).

Three fishes collected in this study and seven species represented by speci-
mens in the Southeastern Louisiana University museum collections had not been
previously reported from the Tangipahoa River. The skipjack herring {Alosa
chrysochloris), was captured at two locations in the main river channel in
Louisiana, least killifish {Heterandria formosa), in a small pool near Hammond,
Louisiana, and goldstripe darter {Etheostoma parvipinne) near the headwaters of
Terry's Creek in Mississippi. None of these were new records for the Lake
Pontchartrain drainage, although the goldstripe darter appears to be rare in this
region.

Fishes not previously documented from this system but located in museum
collections were the blue catfish {Ictalurus furcatus), lake chubsucker {Erimyzon
sucetta), gulf killifish {Fundulus grandis), rainwater killifish {Lucania parva) , inland
silverside {Menidia beryllina), swamp darter {Etheostoma fusiforme), and saddleback
darter {Percina vigil). None of these is apparently common in the river.

Twenty-one additional species of fishes not captured during this study have
been reported from the Tangipahoa River by Douglas (1974) and Davis et al.
(1970). In addition, six species were reported to be distributed statewide in

1994] Tangipahoa River Fishes 145

Louisiana by Douglas (1974), although specific references to these fishes occur-
ring in the Tangipahoa River were not indicated. Therefore, we have exchided
these six species from this discussion. Two species, the speckled chub (Macrhyhop-
sis aestivalis) and an undescribed species o^ Percina (Thompson, 1980), reported
by Douglas (1974) and also found in the Southeastern Louisiana University
Museum, inhabited the Tangipahoa River at one time. However, current evi-
dence of the occurrence of these fishes could not be found during this study.
They have apparently been reduced in numbers in recent times and may even
be extirpated from the river. Discrepancies in the distribution of the cypress
minnow {Hybognathus hayi), as presented by Douglas (1974), were noted by Saul
(1974). Evidence of this fish occurring in the Tangipahoa River was not substan-
tiated.

Davis et al. (1970) noted the occurrence of several fishes in the lower Tan-
gipahoa River, which could not be documented by other sources. Three of these
(Atlantic sturgeon, Acipenser oxyrhynchus; paddlefish, Polyodon spathula; and
southern flounder, Paralichthys lethostigrna) occur in Lake Pontchartrain (pers.
obs.) and seem likely inhabitants of this river. Paddlefish are commonly seen by
boaters on the river (pers. comm.). Three additional species (crystal darter,
Ammocrypta asprella; redfin darter, Etheostoma ivhipplei; and freckled darter, Per-
cina lenticula) occur in adjacent drainages (Pearl River and/or lower Mississippi),
but have not been reported from the Lake Pontchartrain drainage by other
investigators and their existence in the Tangipahoa River seems unlikely.

One specimen of striped bass {Morone saxatilis) was captured from the lower
channel. This species had not been reported from the river since Bean (1884)
mentioned collecting specimens and seeing large schools of these fish near
Osyka, Mississippi (Mcllwain, 1967). The number of lateral line scales (154)
indicated that our specimen was not a naturally occurring fish, but instead, a
member of the Atlantic coast race introduced as part of restocking efforts of
the LDWF which began in 1967 (Mr. Bennie Fontenot, LDWF, pers. comm.,
1986).

The Florida subspecies of the largemouth bass {Micropterus salmoides
floridanus) is another fish which has been stocked in the Tangipahoa River. A
total of 86,300 Florida largemouth bass were stocked in the river between 1981
and 1986 (Harris, 1987). All specimens of largemouth bass captured (40) were
northern largemouth bass {Micropterus salmoides salmoides).

Changes in geological and other environmental factors have resulted in com-
plex distribution patterns of fishes inhabiting southwestern Mississippi and
southeastern Louisiana. Aspects of these patterns were discussed by Guillory
and Conner (1973), Bart and Cashner (1980), and others. Certain fishes of this
region such as the frecklebelly madtom (Noturus munitus) and the silverjaw min-
now (Notropis buccata) reach their western distribution limits at the Pearl River.
Others, such as the bluehead chub (Nocomis leptocephalus), however, exhibit dis-
junct distribution patterns. The bluehead chub is quite abundant in the Bogue
Chitto River, a tributary of the Pearl River, and is also found in some lower
tributaries of the Mississippi River, but does not occur in the intervening Lake
Pontchartrain drainage (Douglas, 1974). Suttkus and Clemmer (1977) proposed
Bayou Pierre as a historical pathway for the dispersal of fishes such as the
northern studfish and bluntface shiner {Cyprinella camura) between the lower
Mississippi River and the Pearl River systems. Similarly, varying patterns were
observed within the Lake Pontchartrain drainage. For example, the emerald
shiner {Notropis atherinoides), found throughout Mississippi and Louisiana includ-

146

Tulane Studies in Zoology and Botany

[Vol. 29

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1994] Tangipahoa River Fishes 147

ing most of the Lake Pontchartrain drainage (Douglas, 1974), has not been
reported from the Tangipahoa River.

Fish species collected in the Tangipahoa River during this study and previous
studies were compared with those from Bayous Lacombe and Sara, the
Tchefuncte, Tickfaw, Amite, Blind, Homochitto, Bogue Chitto rivers, and
Thompson Creek (Table 1). Eighteen species, 8 of which were centrarchids,
occurred in all of these stream systems. Thirty species were found in all six
streams of the Lake Pontchartrain drainage. Within the Lake Pontchartrain
drainage, the Tickfaw, Amite, and Tchefuncte rivers tended to support ichthyo-
faunal assemblages most similar to that of the Tangipahoa River. These four
rivers are ecologically and physically similar (Laiche, 1978) with suitable habitats
for a variety of cyprinid, centrarchid, and percid species, many of which are
common to all of these rivers. Bayou Lacombe and the Blind River, however,
were quite different from the Tangipahoa River. These systems are in areas of
low relief with less diversity of habitat, and consequently had significantly fewer
species, especially among the minnows (Cyprinidae), madtom catfishes (Ic-
taluridae), and darters (Percidae). They also tended to have more marine
species. Comparisons with the Homochitto River, Bayou Sara, Thompson
Creek, and the Bogue Chitto River suggested that these streams were inter-
mediate in similarity to the Tangipahoa.

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1994] Tangipahoa River Fishes 149

Appendix. A checklist of fishes of the Tangipahoa River, southwest Mississippi, and southeast
Louisiana. (Source codes: Do = Douglas, 1974; Da = Davis et al., 1970; SLU = Southeastern
Louisiana University Museum collection. Sources only provided for those species not collected dur-
ing this study).

Family and Species Source

Petromyzontidae (lampreys)

Icthyomyzon castaneus, chestnut lamprey

/. gagei, southern brook lamprey
Acipenseridae (sturgeons)

Acipenser oxyrhynchiis, Atlantic sturgeon Da

Polyodontidae (paddlefishes)

Polyodori spathula, paddlefish Da

Lepisosteidae (gars)

Lepisosteus ociilaliis, spotted gar

L. osseus, longnose gar Da, SLU

L. platostomus, shortnose gar Da, Do

L. spatula, alligator gar
Amiidae (bowfins)

Amia calva, bowfin
Anguillidae (freshwater eels)

Anguilla rostrata, American eel Do

Clupeidae (herrings)

Alosa alabamae, Alabama shad

A. chrysochloris, skipjack herring

Dorosoma cepedianum, gizzard shad

D. petenense, threadfin shad
Engraulidae (anchovies)

Anchoa mitchilli, bay anchovy
Cyprinidae (carps and minnows)

Cyprinella venusta, blacktail shiner

Hybognathus hayi, cypress minnow Do

Luxilus chrysocephalus, striped shiner

Lythrurus fumeus, ribbon shiner Do

L. roseipinnis, cherryfin shiner

Macrhybopsis aestivalis, speckled chub Do, SLU

Notemigonus crysoleucas, golden shiner

Notropis longirostris, longnose shiner

A', texanus, weed shiner

N. volucellus, mimic shiner

A', winchelli, clear chub

Opsopoeodus emiliae, pugnose minnow

Pimephales vigilax, bullhead minnow
Catostomidae (suckers)

Erimyzon oblongus, creek chubsucker

E. sucetta, lake chubsucker Do. SLU
E. tenuis, sharpfin chubsucker

Hypentelium nigricans, northern hog sucker

Ictiobus bubalus, smallmouth buffalo Do, Da

Minytrema melanops, spotted sucker
Moxostoma poecilurum, blacktail redhorse
Ictaluridae (bullhead catfishes)
Ameiurus melas, black bullhead
A. natalis, yellow bullhead

Ictalurusfurcatus, blue catfish SLU

/. punclattis, channel catfish
Noturus gyrinus, tadpole madtom
A^. leptacanthus, speckled madtom
A^. miurus, brindled madtom
A', noctumus, freckled madtom
Pylodictis olivaris, flathead catfish

150 Tulane Studies in Zoology and Botany

Appendix. (Continued)

Family and Species Source

Esocidae (pikes)

Esox americanus, grass pickerel

E. niger, chain pickerel
Aphredoderidae (pirate perches)

Aphredoderus sayanus, pirate perch
Cyprinodontidae (killifishes)

Fundulus chrysotus, golden topminnow

F. euryzonus, broadstripe topminnow

F. grandis, gulf killifish SLU

F. notti, bayou topminnow

F. olivaceus, blackspotted topminnow

Lucania parva, rainwater killifish SLU

Poeciliidae (livebearers)

Gambusia affinis, mosquitofish

Heterandriaformosa, least killifish
Atherinidae (silversides)

Labidesthes sicculus, brook silverside

Menidia beryllina, inland silverside SLU

Percichthyidae (temperate basses)

Morone mississippiensis, yellow bass

M. saxatilis, striped bass
Centrarchidae (sunfishes)

Ambloplites ariommus, shadow bass

Centrarchus macropterus, flier

Elassoma zonatum, banded pigmy sunfish

Lepomis cyanellus, green sunfish

L. gulosus, warmouth

L. macrochirus, bluegill

L. marginatus, dollar sunfish

L. megalotis, longear sunfish

L. microlophiis, redear sunfish

L. punctatus, spotted sunfish

Micropterus punctulatus, spotted bass

M. salmoides, largemouth bass

Pomoxis annularis, white crappie

P. nigromaculatus, black crappie
Percidae (perches)

Ammocrypta asprella, crystal darter Da

A. beani, naked sand darter

Etheostoma chlorosomum, bluntnose darter

E.fusiforme, swamp darter SLU

E. parvipinne, goldstripe darter

E. proeliare, cypress darter

E. stigmaeum, speckled darter

E. swaini, gulf darter

E. lynceum, brighteye darter

E. whipplei, redfin darter Da

Percina caprodes, logperch Do, SLU

P. lenticula, freckled darter Da

P nigrofasciata, blackbanded darter

P. sclera, dusky darter

P. vigil, saddleback darter SLU

Sciaenidae (drums)

Aplodinotus grunniens, freshwater drum
Mugilidae (mullets)

Mugil cephalus, striped mullet
Bothidae (flounders)

Paralichthyes lethostigma, southern flounder Da

Soleidae (soles)

Trinectes maculatus, hogchoker

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