UN

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

OCCASIONAL PAPERS

of the

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 119, PAGES 1-83 NOVEMBER 13, 1985

REVIEW OF THE SUBGENUS CATONOTUS (PERCIDAE)

WITH DESCRIPTIONS OF TWO NEW DARTERS OF

THE ETHEOSTOMA SQUAMICEPS SPECIES GROUP

By

Marvin E. Braasch1 and Richard L. Mayden2

INTRODUCTION

The Cumberland and Tennessee drainages together with tributaries of
the lower Ohio River possess one of the most diverse fish faunas in North
America. Many species native to these drainages are endemic. The
subgenus Catonotus is no exception to the pattern of diversity in this
region. With the exception of Etheostoma flabellare all of its members are
restricted to the Cumberland, Tennessee, and lower Ohio river drainages.

Prior to 1978, accounts of the spottail darter, Etheostoma squamiceps,
considered it to be a single variable species. Howell and Dingerkus (1978)
recognized the diversity in E. squamiceps when they described a highly
distinctive species, E. neopterum. Although their study was not inclusive
of the total range of E. squamiceps, they considered the possibility of
additional undescribed forms masquerading under the now more re-
strictive taxon, E. squamiceps. Braasch and Page (1979) additionally
recognized diversity in the spottail darter and described E. olivaceum, a
distinctive species known from the Caney Fork system since the 1960's.
Further, these authors had independently reached the same conclusion as
Howell and Dingerkus (1978), that the spottail darter represents a
polytypic taxon.

It is the purpose of this paper to examine morphological variation of E.
squamiceps and its relatives. Two new species of this group are described
herein. A brief review of other Catonotus is presented as well as
hypotheses of phylogenetic relationships and biogeography.

1 Belleville Area College, Belleville. Illinois 62221 USA

2 Museum of Natural History and Department of Systematics and Ecology. The
University of Kansas. Lawrence, Kansas 66045 USA, Present address: Department of
Biology. University of California, Los Angeles, California 90024

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

METHODS

Methods of data collection employed in this study follow Braasch and
Page (1979). Complete locality data of materials examined are available
from the authors. Museum acronyms are identified in acknowledgments,
except for the following: AMNH = American Museum of Natural History,
BMNH = British Museum of Natural History, MCZ = Museum of Com-
parative Zoology, and SU = Stanford University. The abbreviation MEB
refers to Marvin E. Braasch collections. The following abbreviations are
used throughout the text: SL = standard length, HL = head length,
BD = body depth, CPD = caudal peduncle depth, PreOL = preorbital
length, HL = head length, EW = eye diameter, DFB = spinous dorsal base
length, DFH = spinous dorsal height, P,L = pectoral fin length,
P2L = pelvic fin length. Analyses of phylogenetic relationships involved
methods of outgroup comparison and parsimony. Our use of this method
was aided through the computer program PAUP, Phylogenetic Analysis
Using Parsimony, provided by David Swofford of the Illinois Natural
History Survey, Champaign. Since the subgenus Catonotus has been
considered phylogenetically close to the subgenus Oligocephalus this
group was used as the primary outgroup. However, since Oligocephalus
has recently been split up taxonomically, we also made comparisons with
other species of Etheostoma, as well as Percina and Ammocrypta.

ACKNOWLEDGMENTS

We wish to express our thanks to the following individuals and their
institutions for providing information, supplying work space, and for the
loan of specimens during the course of this study: Herbert T. Boschung
and David L. Nieland, University of Alabama Ichthyological Collection
(UAIC), Royal D. Suttkus, Tulane University (TU), Lawrence M. Page
and Philip W. Smith, Illinois Natural History Survey (INHS), Reeve M.
Bailey, Robert R. Miller, and Douglas W. Nelson, University of Michigan
Museum of Zoology (UMMZ), Brooks M. Burr, Southern Illinois
University at Carbondale (SIUC), David A. Etnier and Bruce H. Bauer,
University of Tennessee (UT), Frank B. Cross and E. O. Wiley,
University of Kansas (KU), Neil H. Douglas, Northeast Louisiana
University (NLU), William Clay, University of Louisville (UL), Robert
Schoknecht, Cornell University (CU), and Richard Vari, Susan Jewett,
and Janet Gomon, United States National Museum (USNM). We are also
deepy indebted to many friends and students from Belleville Area College,
University of Kansas, Illinois Natural History Survey, and the University
of Tennessee at Knoxville and Martin for help in collecting species of the
subgenus Catonotus since the start of this study. We also wish to thank
David L. Swofford (INHS) for making his computer program PAUP
available to us, and Frank B. Cross, E. O. Wiley, David A. Etnier and
Robert E. Jenkins for reading and improving various parts of the
manuscript. Thomas M. Keevin (INHS) kindly supplied the photographs
of nesting adults. We thank Joseph T. Collins for helping proof the
manuscript and Sharon L. Dewey for suggesting the name crossopterum
for the fringed darter. We also thank Coletta Spencer and Jan Elder for

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 3

typing portions of the manuscript. Finally, we wish to thank the conserva-
tion departments of Kentucky, Tennessee, and Alabama for granting us
permits to collect and study these fishes in their states.

SUBGENERIC SYNOPSIS

Catonotus Agassiz, 1854, Amer. Journ. Sci. and Arts 17:305

Type-species Catonotus lineolatus Agassiz, 1854 by monotypy

Richia Coker, 1926, Bull. Bur. Fish. 42:106

Type-species Richia brevispina Coker, 1926

Invalid name— preoccupied by Richia Grote, 1887 (Lepidoptera)

Richiella Coker, 1927, Copeia 102:18

Type-species Richia brevispina Coker
Replacement for Richia Coker

Taxonomic History of the Subgenus

Rafinesque, in 1819, described Etheostoma flabellaris , the first species
of the group now known as Catonotus, and placed it along with
blennioides and Sciaena caprodes in the new genus Etheostoma. Rafin-
esque did not designate types for either the new genus or species (Collette
and Knappe, 1966). The genus Catonotus was erected by Agassiz in 1854,
when he described C. lineolatus and designated E. blennioides as type-
species for Rafinesque's Etheostoma. Putnam (1863) then added C.
kennicotti from southern Illinois to the genus. Jordan (1875) placed
flabellare in Catonotus and merged the genus Nothonotus Agassiz with it.
Jordan (1877a) then described E. squamiceps, placed it in this group, and
subtracted Nothonotus from Catonotus. Ignoring Agassiz's work, as well
as Jordan's earlier papers, Cope and Jordan (in Jordan, 1877b, p. 51)
designated flabellare as the type-species for Etheostoma, thus replacing
Catonotus with Etheostoma for the subgenus. Jordan and Evermann
(1896) merged many existing darter genera into the single genus Etheos-
toma Rafinesque, retaining the former genera as subgenera. The name
Catonotus was restored by Opinion 14, International Comm. Zool.
Nomen. 1910 (see Hemming, 1947 for discussion of Opinion 14) and has
been used as a subgenus by Bailey and Gosline (1955) and Page (1975a,
1981) in their systematic work on darters.

Relationships of the barcheek darters, E. virgatum Jordan and E.
obeyense Kirsch, to Catonotus were not early recognized. They were
placed in various subgenera (or genera) and were not formally listed as
Catonotus until Bailey and Gosline's (1955) review of the darter sub-
genera.

Collette (1965) divided Catonotus into two species groups— the E.
flabellare group {flabellare, kennicotti, and squamiceps) and the E.
virgatum group (virgatum and obeyense). Kuehne and Small (1971) added
E. barbouri as a third member of the virgatum group and presented a

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

detailed diagnosis for the subgenus. Page (1975a) further subdivided the
subgenus into three species groups— the flabellare group (flabellare and
kennicotti) , the squamiceps group (squamiceps) , and the virgatum group
(virgatum, obeyense, and barbouri) based on secondary sexual characteris-
tics and breeding behaviors.

Since 1976, four additional Catonotus have been described; smithi
Page and Braasch, 1976 and striatulum Page and Braasch, 1977 of the
virgatum group; and neopterum Howell and Dingerkus, 1978 and
olivaceum Braasch and Page, 1979 of the squamiceps group. The two
additional species of the squamiceps group described herein and an
undescribed species of the flabellare group (being studied by Dr. Robert
Jenkins) brings the total number of Catonotus species to twelve.

The Subgenus Catonotus Agassiz

Diagnosis — The following diagnosis is modified from Kuehne and
Small (1971) and Braasch and Page (1979): lateral line incomplete and
straight; preoperculomandibular canal pores 8-11; branchiostegal rays
usually 6; body fairly elongate; snout pointed; premaxillary frenum
present; vertebrae 32-37; first dorsal fin low; dorsal spines 7-11 and with
fleshy tips in breeding males; anal spines 2, the first thicker than the
second and more than half the length of the first ray; breeding tubercles
absent; anus not encircled by fleshy villi; genital papilla of females flat and
not bifurcated.

Key to Species Groups

1 . Nape and/or breast scaled, base of caudal fin with vertical row of 3

dark spots E. squamiceps group

Nape and breast naked, base of caudal fin without spots 2

2. Body and fins with red and blue coloration; cheeks with red and white
iridescent diagonal bar; fleshy knobs on dorsal fins small and opaque;
infraorbital canal pores 6 or fewer E. virgatum group

Body lacking red or blue colors; no brightly colored bar on cheeks;
large round fleshy golden knobs present on tips of dorsal spines of
males; infraorbital pores 7 or more E. flabellare group

ETHEOSTOMA SQUAMICEPS SPECIES GROUP

The E. squamiceps species group is a group of five closely related
usually allopatric species in the lower Ohio, Cumberland, and Tennessee
river drainages from southern Illinois and Indiana south to northern
Mississippi and Alabama. Species are moderately large to large Catonotus
(maximum body size = 85 mm SL) characterized by having three dark
spots at base of caudal fin, scales on nape, prepectoral, and breast (and
usually on cheeks and opercles), body and fin coloration not chromatic,
infraorbital canal complete or incomplete with 7 to 8 pores, and dorsal fin
spines with fleshy knobs in both males and females. The most important

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 5

characters distinguishing members of this group are coloration and shape
of dorsal fins of nuptial males. Since all species lack bright colors,
apparently fin coloration and shape are used for species recognition. These
characters are of taxonomic use only during the height of the breeding
season.

Etheostoma squamiceps Jordan
Spottail Darter
Figs. 1. 2, 3.
Etheostoma squamiceps. Jordan, 1877:11-12 (original description,
Russellville, KY)

Claricola squamiceps. Jordan, Evermann and Clark, 1930:292 (check-
list of fishes)

Catonotus squamiceps. Gerking, 1945:94 (status in Indiana)
Poecilichthys squamiceps. Moore and Cross, 1950:145-146 (separa-
tion of E. squamiceps and E. parvipinne)

Types— Etheostoma squamiceps was described from two specimens
from Russellville, Logan Co., Kentucky. The exact locality is unknown. It
could have been from either the Green River system of the Ohio drainage
or the Red River system (Cumberland drainage). Since the form E.
squamiceps (as used herein) is the only species of the E. squamiceps group
found in the Green or Red rivers, we restrict the types of Jordan to this
form. Lectotype designated by Collette and Knapp (1966): 59 mm SL male

4

j***«>

"Wffc.

V

Figure 1. Etheostoma squamiceps. Top. male 61.6 mm SL; Bottom, female 55.9 mm SL
(INHS 75747, Wabash R. System, Posey Co.. Indiana).

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Figure 2. First dorsal fin pigment patterns of breeding males of species in the Etheostoma
squamiceps species group. A) E. squamiceps, 68 mm SL (INHS 75747). B) E. neopterum, 64
mm SL (KU 20900). C) E. olivaceum, 61 mm SL (KU 14402). D) E. crossopterum, 70 mm
SL (KU 20896). E) E. nigripinne, 57 mm SL (KU 20000). Horizontal bar equals 1 mm.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

Figure 3. Second dorsal fin pigment patterns of breeding males of species in the Etheostoma
squamiceps species group. A) E. squamiceps, 68 mm SL (INHS 75747). B) E. neopterum, 56
mm SL (KU 20762). C) E. olivaceum, 61 mm SL (KU 14402). D) E. crossopterum, 70 mm
SL (KU 20896). E) E. nigripinne, 57 mm SL (KU 20000). Horizontal bar equals 1 mm.

8 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

USNM 1145 (erroneously given as USNM 1345 in original description).
The second specimen, a female (USNM 197968) became paralectotype.

Etymology — The name squamiceps is taken from the Latin "squama"
for scales and "ceps" for head in relation to the presence of scales on
opercles and cheeks. The common name refers to the three dark spots at
the base of the caudal fin.

Diagnosis — Dorsal fin rays with branches not adnate and possessing
moderately large, fleshy knobs in breeding males; anal fin rays usually 7;
pectoral fin rays usually 1 1; first dorsal fin without orange band.

Material examined— Lectotype: USNM 1145; Kentucky: Logan Co.,
Russellville. Paralectotype: USNM 197968; same locality data as lec-
totype.

Other specimens examined during the study are listed below in
geographic order by drainage and river system:

Ohio R. Drainage: Cache R. System: Illinois: Union Co.: INHS
17833 (19), 17838 (9), 17847 (7), 18110 (1), 18033 (2); KU 7267 (6).
Johnson Co.: INHS 2221 (1).

Tributaries Ohio R.: Illinois: Hardin Co.: INHS 985 (2), 994 (4),
999(1), 1027(6), 84986(1), 85003(1), 88504(16); UMMZ 111641 (1),
171805 (3), 171806 (9); TU 22377 (2). Pope Co.: INHS 1480 (3), 1484
(4), 1560 (1), 1576 (6), 1593 (21), 86187 (3), 88503 (2); UL 9176 (2),
9182 (3), 9186 (1), 12871 (8). Kentucky: Crittenden Co.: INHS 78340
(3), 78356 (1), 78358 (1), 78376 (26), 78379 (2), 78392 (2). Livingston
Co.: INHS 77707 (3), 78516 (7), 78534 (8).

Wabash R. System: Illinois: White Co.: UMMZ 102407 (4). Indiana:
Posey Co.: INHS 75747 (12), 73680 (1), 73681 (1). Gibson Co.: UMMZ
65040 (4).

Tradewater R. System: Kentucky: Caldwell Co.: INHS 86756 (33);
KU 14409(1); MEB 1 10 (28); SIUC uncat. (10); UL 51 14 (1). Crittenden
Co.: INHS 77716 (1), 77720 (2), 77730 (3). Christian Co.: SIUC uncat.

(1).

Lower Green R. System: Kentucky: Logan Co.: INHS 77734 (4).
Christian Co.: INHS 27501 (9); KU 16223 (3); MEB 111(1). Butler Co.:
KU 16432 (3).

Upper Green R. System: Kentucky: Grayson Co.: INHS 75755 (3);
UL 6280 (2), 7987 (4), 8677 (1). Adair Co.: KU 11565 (6), 14405 (4),
16428 (1). Larue Co.: KU 5816 (5); TU 19530 (3); UMMZ 154753 (1);
UT 91.2607 (3). Hardin Co.: UL 5031 (2), 5983 (3), 7023 (3), 8665 (1),
11896 (3), 12904 (3). Green Co.: UMMZ 86472 (1).

Barren R. System: Kentucky: Allen Co.: INHS 75991 (4); KU 1 1642
(1); UL 6878 (1). Tennessee: Macon Co.: UMMZ 165382 (1). Clay Co.:
INHS 82642 (7); UMMZ 165366 (4). Sumner Co.: MEB 107 (2), 108
(11).

Cumberland R. Drainage: Red R. System: Kentucky: Christian Co.:
INHS 75792 (1); UL 5286 (1). Logan Co.: INHS 78556 (3), 75800 (4);
UL 5833 (2), 6690(1). Todd Co.: INHS 77744 (1), 77696 (25), UL 5323

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 9

(1). Tennessee: Montgomery Co.: UMMZ 160995 (1). Robertson Co.:
MEB 112 (6), 184 (1).

Description — Etheostoma squamiceps is a large, robust Catonotus;
largest specimen a male 72.5 mm SL. Males generally larger than females,
largest female 58 mm SL. Infraorbital canal incomplete with narrow
interruption below eye, pores normally 4-3 or 4-4; preoperculomandibular
canal pores 10; supraorbital canal pores 4 or 5; supratemporal interrupted
medially, pores 2-2; lateral canal pores 5; frenum narrow to moderate;
branchiostegal rays 6 per side, membranes slightly to moderately joined;
preopercle entire; genital papilla broad and flat, not bifurcated.

Cheeks, opercles, breast and nape normally scaled. Lateral line scales
41-60, 12-51 pored, unpored scales 5-35; transverse scale rows below
lateral line 7-11 (usually 9), above lateral line 5-8 (6).

First dorsal fin with 7-11 (8-9) spines, fleshy knobs at tips present in
both sexes; second dorsal rays 12-15 (13); anal spines 2, anal rays 6-9
(7-8); branched caudal rays 14-17; pectoral jays 10-12 (11).

Body proportions: HL/SL 0.27-0.33 (X = 0.29); BD/SL 0.17-0.26
(0.21); CPD/SL 0.08-0.16 (0.01); P,L/SL 0.19-0.26 (0.23); P,L/SL
0.16-0.24 (0.20); PreOL/HL 0. 18-0.36 (0.25); EW/HL 0. 17-0.31 (6.25);
DFH/DFB 0.16-0.56 (0.31).

Coloration of freshly preserved specimens: General coloration of
females and non-breeding males consists of irregular mottling of brown on
a light tan background (Fig 1). Anterior lateral line pores unpigmented,
forming pale line against dark mottling. Dark humeral spot large. Base of
the caudal fin with three dark spots which may occasionally coalesce to
form a vertical bar. Dorsum normally paler than sides, especially in
specimens from the northern part of range where a sharply delineated wide
pale band extends posteriorly from occiput to caudal peduncle. Pale dorsal
area usually crossed by 8 (sometimes 9) dark bands. Venter normally
unmarked except for spot at base of each pectoral fin. Head typically with
pre-, post-, and suborbital dark bars. Cheeks usually mottled; may have a
dark diagonal line from below the eye posterodorsally connecting with
postorbital bar at the anterior edge of opercle.

Spinous dorsal fin colorless except for a few small melanophores
forming indistinct subdistal and basal bands; base of fin thickened;
membrane thickened at tip of spines, forming inconspicuous light colored
knobs. Membranes of soft dorsal fin with clusters of melanophores
forming 3-7 interrupted bands and rather indistinct basal band; base of fin
thickened. Caudal fin with melanophores clustered along rays and forming
6-8 narrow dark bands. Anal and pelvic fins unpigmented; pectoral fins
unpigmented to lightly barred with up to 9 narrow brown bands.

Breeding males dark. Sides uniformly dark brown or gray; caudal
spots enlarged to form vertical bar; dorsum uniformly dark; head much
darker, almost black, often obscuring regular markings; lower part of head
greatly swollen; venter dark gray (Fig. 1). During actual courtship and egg
care, strongly contrasting alternating dark and light bars develop on sides
(Page, 1975b, p. 10 for photograph). First dorsal fin black except for large

10

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

pale yellow or white knobs on spines and occasional small, unpigmented
vertical windows behind each spine; basal part of fin with narrow clear
band and membrane thickened (Fig. 2). Second dorsal fin black with a
broad distal white or pale yellow band and 4-6 rows of short, concave, pale
dashes across rays; basal portion of fin thickened; at peak of breeding
season tips of rays greatly elongated, terminating in round white or yellow
knobs (Fig. 3). Anal fin dark with pale distal margin; membrane thickened
around first spine. Pelvic fins dark except for paler membrane around
spine and pale distal margin. Caudal and pectoral fins unchanged from
non-breeding coloration. Genital papilla darkly pigmented.

Distribution— Etheostoma squamiceps is widespread, but not common,
in the Green drainage and Red river system of the Cumberland drainage in
Kentucky and Tennessee. It occurs more abundantly in Tradewater River
and nearby direct tributaries of Ohio River in Kentucky and southern
Illinois. Isolated populations occur in tributaries of Wabash River in
Gibson and Posey counties, Indiana, and White County, Illinois, and in
Cache River in southwestern Illinois (Fig. 4). Specimens from Kaskaskia
River, Shelby County, Illinois, reported in Forbes and Richardson
(1908:313) have been reidentified as Etheostoma asprigene (pers. obs.).

Variation — Etheostoma squamiceps shows little sexual dimorphism
except for size and pigmentation. Males have a deeper caudal peduncle,
smaller eye, and more pored lateral-line scales (a. = .001) than do females
(Table 1). Ontogenetic variation is more pronounced. Small specimens
( <40 mm SL) have a larger eye; longer pelvic fins; higher spinous dorsal
fins; fewer pored lateral-line scales; more unpored lateral-line scales

Figure 4. Distribution of Etheostoma squamiceps.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

11

(cx= .001); longer heads, and thinner caudal peduncles (a= .05) than do
larger individuals (Table 2).

To evaluate geographic variation, the range of E. squamiceps was
divided into eight operational units. The Green River system was divided
into three areas: Barren River, Upper Green (eastern headwaters plus
Nolin and Rough rivers), and lower Green (Mud and Pond rivers). Other
populations include Red River, Cache River, Wabash River, Tradewater
River, and small direct tributaries of Ohio River in southeastern Illinois
and northwestern Kentucky. Because of small sample size, the lower
Green and Barren populations were not used in analyzing proportions.

Geographic variation in pigmentation, squamation, meristics and
morphometries is extensive between populations of E. squamiceps.

Pigmentation: Dorsal pigmentation varies individually and geograph-
ically. Individuals from the Red river system have a narrow, light tan
dorsal band extending from nape area to caudal peduncle, which is crossed
by several dark cross bars. Populations in northern and western parts of
the range have reduced pigmentation and often have a wide pale dorsal
band which may even extend down onto the sides. Dorsal cross bars may
be reduced to a scalloping effect on the dorsal stripe.

Squamation: Of areas of the body exhibiting variation in squamation,
the peripheral populations (Cache, Ohio, Wabash, Barren rivers) show
reduced squamation on nape, opercles, and cheeks (Table 3; Figs. 5-7).

Meristics and morphometries: Dorsal spines, dorsal rays, anal rays,
pectoral rays, lateral-line scales, pored lateral-line scales, unpored lateral-

Table 1 . Morphological characters exhibiting sexual dimorphism in species of the Etheos-
toma squamiceps species group. Means and ranges of proportions in thousands of a

millimeter.

Species

N

MALES

N

FEMALES

&
Character

Range

X

SD

Range

X

SD

Etheostoma squamiceps

Pored Lateral Scales

48

29-51

38.5

5.46

34

24-47

35.8

5.78

EW/HL

49

171-318

243

0.03

34

192-308

268

0.03

CPD/SL

51

112-147

131

0.01

40

109-167

125

0.01

Etheostoma neopterum

P2L/SL

22

167-224

201

0.01

25

171-217

193

0.01

CPD/SL

22

105-158

128

0.02

25

111-138

120

0.01

Etheostoma crossopterum

Pored Lateral Scales

24

22-44

35.2

5.17

36

7-39

29.2

7.01

Unpored Lateral Scales

24

9-28

18.4

6.31

36

12-37

24'5

6.67

CPD/SL

23

111-142

128

0.01

36

92-133

188

0.01

EW/HL

23

186-292

239

0.03

36

211-318

268

0.03

PreOL/SL

23

222-294

261

0.02

36

167-304

246

0.03

P1L/SL

23

190-258

231

0.01

36

207-269

237

0.01

Etheostoma nigripinne

CPD/SL

46

116-159

137

0.01

24

96-154

129

0.14

EW/HL

46

188-308

253

0.03

24

222-300

268

0.02

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

line scales, transverse scale rows above and below lateral line, HL/SL,
BD/SL, P,L/SL, P2L/SL, PreOL/HL, EW/HL, and DFH/DFB all varied
significantly geographically (a = 0.001) (Tables 4-1 1).

Peripheral populations from Cache, Tradewater, and Red rivers are
most extreme in variation. Anal rays generally 7 except in the Tradewater,
Barren, and Red rivers which typically have 8 rays (Table 7). Scales above
lateral line generally 8 except in populations from the Tradewater. lower
Green, and Barren rivers and to a lesser extent, the Red River which most
commonly have 9 rows (Table 10). Scales below lateral line generally 6
except in Wabash River populations which average more (Table 11).
Specimens from the Cache River have fewer dorsal rays and spines, and to
a lesser extent, populations from Red River, upper Green River, and small
tributaries of the Ohio River show some reduction in the number of dorsal
spines (Tables 5 and 6).

Habitat and Life History— The preferred habitat of E. squamiceps is
small, quiet streams with either large, flat rocks or with bedrock bottom
where they hide in crevices and under ledges. In this type of habitat E.

Table 2. Characters exhibiting ontogenetic variation in species of the Etheostoma squam-
iceps group. Means and ranges of proportions in thousands of a millimeter.

Species

N

<40 mm

SL

N

>40 mm SL

&
Character

Range

X

SD

Range

X

SD

Etheostoma squamiceps
Pored Lateral Scales

28

12-42

28.7

7.59

82

24-51

36.9

6.92

Unpored Lateral Scales

28

10-31

21.0

6.31

82

5-28

13.7

4.97

CPD/SL

27

98-161

123

0.02

91

109-167

128

0.01

EW/HL

28

235-333

282

0.03

82

171-318

253

0.03

DFH/DFB

27

278-538

388

0.03

83

206-550

324

0.04

Etheostoma neopterum
Pored Lateral Scales

59

14-41

34.1

4.81

47

9-42

36.6

3.56

Unpored Lateral Scales

59

9-31

14.8

4.16

47

7-23

13.1

3.45

HL/SL

59

282-356

313

0.01

47

237-330

301

0.02

CPD/SL

59

102-154

127

0.01

47

111-158

124

0.01

EW/HL

59

250-353

292

0.01

47

214-320

265

0.01

BD/SL

59

189-228

206

0.01

47

173-219

197

0.02

DFH/DFB

59

286-500

382

0.01

47

265-615

334

0.01

Etheostoma crossopterum
Pored Lateral Scales

55

7-43

23.9

8.28

60

7-44

31.6

6.98

Unpored Lateral Scales

55

15-42

29.6

7.67

60

9-37

22.1

7.17

HL/SL

57

284-350

306

0.01

59

267-321

297

0.01

EW/HL

57

227-333

289

0.02

59

186-318

257

0.03

DFH/DFB

57

222-538

356

0.05

59

200-588

306

0.07

P1L/SL

57

203-269

242

0.02

59

190-269

234

0.01

P2L/SL

57

176-242

209

0.01

59

165-248

199

0.02

Etheostoma niqripinne
HL/SL

30

278-328

297

0.01

70

264-318

291

0.01

EW/HL

30

250-333

280

0.02

70

188-308

258

0.03

DFH/DFB

30

211-500

339

0.06

70

174-381

280

0.05

P2L/SL

30

179-233

208

0.01

70

161-233

201

0.02

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

13

squamiceps may be abundant. Although they usually avoid riffles they may
occasionally be found in quiet riffles in late summer when pools are too
low to provide cover. They tend to avoid fast, gravelly streams and are
very rare over sand or mud substrate. Darter species most commonly
associated with E. squamiceps are Etheostoma barbouri in the eastern
portion of its range and E. kennicotti to the west. Other fishes commonly
found with E. squamiceps are Campostoma anomalum, Pimephales
notatus, Semotilus atromaculatus , Phoxinus erythrogaster, Fundulus
olivaceus, and Lepomis cyanellus.

Page (1974) gave detailed information on reproduction, growth, food,

Table 3. Squamation of nape, cheek, and opercular regions in species of the Etheostoma
squamiceps species group. N = naked, E = embedded and S = exposed.

Species
&

N

Nape

Cheek

Opercle

Population

N

E

S

N

E

S

N

E

S

Etheostoma squamiceps
Cache River

34

0

32

2

16

18

0

0

32

2

tribs.. Ohio River

35

0

35

0

0

35

0

0

35

0

Wabash River

14

0

13

1

0

12

2

0

13

1

Tradewater River

35

0

6

29

0

33

2

0

6

29

Lower Green River

14

0

10

4

2

12

0

0

10

4

Upper Green River

25

0

21

4

6

17

2

0

21

4

Barren River

14

0

13

1

0

14

0

0

13

1

Red River

42

0

36

6

0

39

3

0

36

6

TOTALS

213

0

166

47

24

180

9

0

166

47

Etheostoma neopterum
Clarks River

55

0

37

18

0

55

0

0

19

36

Big Sandy River

51

1

49

1

1

49

1

0

42

9

Duck River

23

0

23

0

0

23

0

0

23

0

Shoal Creek

26

0

16

10

5

15

6

0

14

12

TOTALS

155

1

125

29

6

142

7

0

98

57

Etheostoma crossopterum
Lower Cumberland River

46

0

46

0

20

26

0

0

43

3

Middle Cumberland River

58

0

57

1

20

38

0

0

55

3

tribs., Mississippi River
Duck River

11
48

3
0

8
43

0

5

11

22

0
26

0
0

3
0

8
45

0

3

Cypress Creek

42

0

31

11

0

42

0

0

17

25

TOTALS

205

3

185

17

73

132

0

3

168

34

Etheostoma nigripinne

western tribs.,

Tennessee River

37

0

32

5

0

37

0

0

32

5

eastern tribs.,

Tennessee River

24

0

24

0

0

24

0

0

21

3

southern tribs..

Tennessee River

64

0

53

11

0

63

1

0

53

11

Shoal Creek

31

0

27

4

2

29

0

0

25

6

Elk River

44

0

43

1

1

43

0

1

38

5

TOTALS

200

0

179

21

3

196

1

1

169

30

Etheostoma olivaceum

Caney Fork River

100

0

61

39

100

0

0

100

0

0

14

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Figure 5. Squamation of nape of species in the Etheostoma squamiceps species group. Each
circle represents a river system identifed in Table 3. In each circle stippled areas represent
embedded scales, solid black areas exposed scales, and white areas naked. Frequencies of
each condition are given in Table 3. Numbers refer to populations as outlined below. Area A
includes E. squamiceps, E. olivaceum, E. crossopterum, and E. nigripinne. Area B is E.
neopterum. Within area A, E. squamiceps = 1-8, E. crossopterum = 9-l3, E.
nigripinne = 14-18, and E. olivaceum = 19.

Figure 6. Squamation of opercles in the Etheostoma squamiceps species group. Area A
includes E. squamiceps, E. crossopterum, E. nigripinne, and E. olivaceum. Area B includes
E. neopterum. See Figure 5 and Table 3 for explanation of numbers and color codes.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

15

Figure 7. Squamation of cheek regions in the Etheostoma squamiceps species group. Area A
includes E. squamiceps, E. crossopterum, E. nigripinne and E. olivaceum. Area B includes
E. neopterum. See Figure 5 and Table 3 for explanation of numbers and color codes.

habitats, population composition, predation, and parasitism of this spe-
cies. This life history study was actually a composite of E. squamiceps
(Big Creek) and a heretofore undescribed species from the Cumberland
River system (Richland Creek). Life history data from this study,
however, are separable since data on these two species were presented
separately and ecological comparisons were made.

Etheostoma crossopterum new species

Fringed Darter

Figs. 2, 3, 8, 10.

Holotype— University of Kansas, KU 20896, breeding male 69.8 mm
SL (Fig. 8), collected from McKnight Branch, 8 km N Readyville,
Rutherford County, Tennessee, 2 April 1980, Marvin E. Braasch.

Paratypes — A total of 85 specimens deposited as follows: 5— Univer-
sity of Kansas (KU 20897, 46.6-75.6 mm SL, same data as holotype);
26— Illinois Natural History Survey (INHS 74918, 29.0-57.0 mm SL, East
Fork Stones River at Lofton, Rutherford Co., TN, 14 March 1976); 1 1 —
University of Michigan Museum of Zoology (UMMZ 174481, 31.6-53.4
mm SL, Richland Creek on N edge of Belle Meade, SSW Nashville,
Davidson Co., TN, 2 May 1954); 4— United States National Museum
(USNM 266733, 40.0-47.2 mm SL, same data as UMMZ 174481); 27-
UMMZ 177570 (15.0-59.5 mm SL, Richland Creek at Leake Ave., off US
70S, SW Nashville, Davidson Co., TN, 16 May 1956); 4— Southern
Illinois University at Carbondale (SIUC 9782, 46.6-59.4 mm SL, same
data as UMMZ 177570); 4— University of Tennessee (UT 91.838,

16

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

17

56.6-75.5 mm SL, Stones Creek at gravel road just off 1-40 (Mt. Juliet
exit), Wilson Co., TN, 12 May 1973); 3— UT 91.822 (48.8-73.0 mm SL,
same data as UT 91.838); 2— University of Alabama Ichthyological
Collection (UAIC 7713.01, 63.5-64.0 mm SL, same data as UT 91.838);
2— Tulane University (TU 139109, 59.4-64.7 mm SL, same data as UT
91.838).

Etymology — The name crossopterum is taken from the Greek
"crosso" meaning fringed or tassled and "pteron" for fin, in reference to
the morphology of the distal border of the second dorsal fin of breeding
males. The common name also refers to this character.

Diagnosis— Distal margin of second dorsal fin in breeding males with
broad light band, rays extending far beyond membrane margin, tipped in
black, and lacking fleshy knobs. Infraorbital canal incomplete. Anal fin
rays usually 7.

Table 5. Counts of dorsal fin spines in species of the Etheostoma squamiceps species group.

Species

&

Population

Number of Dorsal Spines

N

X

7

8

9

10 11

SD

Etheostoma squamiceps
Cache River

1

1

21

12

34

8.3

0.52

small tribs. , Ohio River

10

24

1

35

8.7

0.59

Wabash River

5

9

14

8.6

0.47

Tradewater River

5

21

9

35

9.1

0.62

Lower Green River

2

12

14

8.8

0.35

Upper Green River

11

12

2

25

8.6

0.62

Barren River

1

11

2

14

9.0

0.45

Red River

1

15

25

1

42

8.6

0.57

TOTALS

2

70

126

14 1

213

8.7

0.57

Etheostoma neopterum
Clarks River

1

51

3

55

9.0

0.26

Big Sandy River

13

34

4

51

8.8

0.55

Duck River

14

6

20

8.3

0.45

Shoal Creek

7

17

2

26

8.8

0.56

TOTALS

35

108

9

152

8.7

0.46

Etheostoma crossopterum
Lower Cumberland River

22

40

4

66

8.7

0.56

Middle Cumberland River

18

38

2

58

8.7

0.51

direct tribs.. Mississippi River

8

4

12

8.3

0.47

Duck River

25

25

50

8.5

0.50

Cypress Creek

27

6

33

9.1

0.38

TOTALS

73

134

12

219

8.7

0.56

Etheostoma nigripinne
western tribs., Tennessee River

1

25

10

1

37

8.3

0.56

eastern tribs., Tennessee River

1

18

11

30

8.3

0.53

southern tribs., Tennessee River

2

37

8

3

50

8.2

0.61

Shoal Creek

21

5

26

8.1

0.39

Elk River

3

27

18

2

50

8.3

0.66

TOTALS

7

128

52

6

193

8.3

0.59

Etheostoma olivaceum

Caney Fork River

7

43

7

57

9.2

0.45

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Material examined— Specimens examined during this study, but not
designated as types are arranged below by drainage and river system.

Mississippi R. Dr.: Tennessee, Obion Co.: INHS 27557 (3); KU
20925 (7), UT 91.189 (1), 91.166 (9), 91.1713 (7), 91.998 (10).

Tennessee R. Dr.: Duck R. System: Tennessee, Bedford Co.: INHS
77517 (5). 79364 (11), 76948 (14), 76137 (3); KU 20898 (1), 20946 (10),
20949 (28), 20959 (10); UMMZ 121038 (2), 121052 (6), 121094 (1),
121107(1), 121113(2), 121122(1), 121137(13), 121148(1), 121184(3),
121194 (18), 121252 (2), 121270 (11), 121260 (1), 121282 (6), 121294
(1), 121338 (1), 121351 (1): UT 91.746 (17), 91.768 (21), 91.774 (11),
91.1603 (4), 91.1478 (27); UAIC 2534 (1), 2535 (1), 2698 (2); MEB 114
(2), 115 (1), 116 (1), 117 (1), 118 (1). Marshall Co.: INHS 86781 (32);
KU 14193 (1), 16221 (31); UAIC 2858 (23); UMMZ 121389 (5). 121453
(2), 121479 (2); UT 91.1293 (2), 91.1294(1). Maury Co.: INHS 27582

Table 6. Counts of dorsal fin rays in species of the Etheostoma squamiceps species group.

Species

&

Population

Number of Dorsal F

in Ray

s

10 11

12

13

14

15

N

X

SD

Etheostoma squamiceps
Cache River

16

15

3

34

12.6

0.64

small tribs., Ohio River

7

26

2

35

12.8

0.48

Wabash River

5

7

2

14

12.8

0.67

Tradewater River

4

23

8

35

13.1

0.57

Lower Green River

3

7

4

14

13.0

0.70

Upper Green River

3

18

4

25

13.0

0.52

Barren River

4

5

4

1

14

13.1

0.91

Red River

6

20

16

42

13.2

0.68

TOTALS

48

121

43

1

213

12.9

0.64

Etheostoma neopterum
Clarks River

3

22

24

6

55

12.6

0.75

Big Sandy River

4 42

3

1

1

51

11.0

0.62

Duck River

2

17

1

20

11.9

0.38

Shoal Creek

1 12

13

26

11.5

0.58

TOTALS

5 59

55

26

7

152

11.8

0.58

Etheostoma crossopterum
Lower Cumberland River

2

25

37

2

66

12.5

0.60

Middle Cumberland River

2

31

24

1

58

12.4

0.58

direct tribs., Mississippi River

1

6

5

12

12.3

0.62

Duck River

8

23

19

50

13.2

0.70

Cypress Creek

6

23

4

33

13.9

0.54

TOTALS

5

70

95

45

4

219

12.8

0.62

Etheostoma nigripinne
western tribs., Tennessee River

1

4

21

11

37

13.1

0.70

eastern tribs., Tennessee River

1

14

14

1

30

13.5

0.61

southern tribs., Tennessee River

3

28

16

3

50

13.3

0.69

Shoal Creek

1

17

7

1

26

13.3

0.60

Elk River

1

13

26

10

50

12.9

0.72

TOTALS

2

22

106

58

5

193

13.2

0.69

Etheostoma olivaceum

Caney Fork River

4

60

35

1

100

12.3

0.57

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

19

it '»i"'tWji|hi<l<Hii7i

Figure 8. Etheostoma crossopterum, new species. Top, Holotype, male 69.8 mm SL (KU
20896); Bottom, Paratype, female, 49.8 mm SL (KU 20897).

(15), 79338 (1); KU 10674 (13), 10691 (1), 20929 (40); UMMZ 104685
(3), 104695(1), 104706(2), 104720(1), 104746(1), 104798(1), 105077
(5), 121514 (2), 121559 (5), 121598 (1), 121605 (1), 121608 (1); UT
91.745 (1), 91.764 (2); MEB 200 (12), 202 (2), 203 (1). Williamson Co.:
UMMZ 104776 (6), 121546 (4), 121566 (1), 121577 (5). Coffee Co.:
INHS 75761 (5); KU 20933 (1); UAIC 2418 (1); UMMZ 120841 (1),
120877 (7), 120889 (21), 120896 (7), 120906 (22), 120916 (1), 120926
(3), 120972 (9); UAIC 2418 (1); UT 91.627 (12), 91.714 (26), 91.715
(30), 91.1445 (8). Hickman Co.: INHS 77817 (4), 77636 (6), 77644 (4);
UMMZ 104757 (3), 104820 (1), 104974 (1), 105284 (5); MEB 97 (20).
Lawrence Co.: INHS 79312 (1); UMMZ 105123 (3), 105150 (4), 105169
(1); UT 91.99 (1); MEB 204 (2). Perry Co.: INHS 77661 (1): KU 10671
(5); UMMZ 105250 (2); TU 14694 (1); MEB 120 (2). Lewis Co.: KU
20852 (1); UMMZ 105109 (2); MEB 96 (3), 201 (5). Wayne Co.: INHS
86751 (14); KU 20899 (2); UMMZ 105180(3), 113945 (2); MEB 122(4),
123 (6), 205 (1), 206 (1). Humphreys Co.: INHS 77836 (10); UMMZ
104945 (2). Dickson Co.: UMMZ 104852 (1), 104881 (1).

Cypress Creek System: Tennessee: Wayne Co.: INHS 83082 (2);
MEB 100 (16). Alabama: Lauderdale Co.: INHS 76130 (17), 86778 (5);
UAIC 1589.11 (6), 1991.02 (2), 2509.05 (1), 3204.12 (23), 3205.05 (7),
3206.10 (16), 3207.10 (3), 3208.11 (10), 3238.08 (5), 3239.09 (14),
3240.12 (51), 3241.06 (1), 3242.09 (49), 3243.11 (40), 3520.09 (3).

Middle Cumberland R. Dr.: Tennessee: Rutherford Co.: INHS
83090(3), 74918(26), 86760(6). 86763(1); KU 11477(8); SIUC uncat.

20

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

(16); UMMZ 158764 (5). Cannon Co.: INHS 75873 (2), 86780 (3); KU
11681 (1), 20944(8); UT 91.1537 (1). Williamson Co.: INHS 75870 (1),
83122 (5), 75870 (1); MEB 124 (1); UMMZ 177581 (23). Dickson Co.
KU 14196 (2); TU 14663 (1). Davidson Co.: INHS 27463 (9), 75712 (11)
MEB 125 (1); UMMZ 96363 (53), 96371 (4), 174481 (15), 177570 (31)
UT 91.1297 (2), 91.1316 (4). Smith Co.: KU 12012 (5); MEB 126 (1)
Montgomery Co.: INHS 82996 (5), 83007 (3), 75766 (2); MEB 10 (1)
UMMZ 160995 (1). Sumner Co.: MEB 127 (11); UAIC 2974 (9); UMMZ
16402 (2). Cheatham Co.: INHS 82588 (1), 82596 (5), 88501 (1); UT
91.207 (4). Rutherford Co.: INHS 83085 (4). Trousdale Co.: INHS 83040
(1); MEB 129 (1), 130 (1); SIUC uncat. (5). Wilson Co.: KU 20938 (3);
MEB 132(17), 133(7); UT 91.165 (83), 91.822 (8), 91.838 (4), 91.1085
(4). Cannon Co.: UT 91.1537 (1).

Table 7. Counts of anal fin rays in species of the Etheostoma squamiceps species group.

Species

&

Population

Etheostoma squamiceps
Cache River
small tribs. , Ohio River
Wabash River
Tradewater River
Lower Green River
Upper Green River
Barren River
Red River

TOTALS

Etheostoma neopterum
Clarks River
Big Sandy River
Duck River
Shoal Creek

TOTALS

Etheostoma crossopterum
Lower Cumberland River
Middle Cumberland River
direct tribs., Mississippi River
Duck River
Cypress Creek
TOTALS

Etheostoma nigripinne
western tribs., Tennessee River
eastern tribs., Tennessee River
southern tribs., Tennessee River
Shoal Creek
Elk River

TOTALS 2

Etheostoma olivaceum
Caney Fork River

Number of Fin Rays

6

7

8

9

N

X

SD

1

27

5

1

34

7.2

0.15

4

15

15

1

35

7.4

0.72

1

12

1

14

7.0

0.63

1

11

21

2

35

7.7

0.63

2

5

7

14

7.4

0.72

3

14

7

1

25

7.2

0.71

5

6

3

14

7.9

0.74

1

12

26

3

42

7.7

0.62

13

101

88

11

213

7.5

0.65

1

15

29

10

55

7.9

0.72

2

26

22

1

51

7.5

0.60

19

1

20

8.1

0.22

1

21

4

26

7.1

0.43

4

62

74

12

152

7.7

0.49

3

35

27

1

66

7.4

0.60

6

38

14

58

7.1

0.57

2

7

2

1

12

7.2

0.79

1

15

33

1

50

7.7

0.55

19

13

1

33

7.5

0.55

12

114

89

4

219

7.4

0.59

1

14

19

3

37

7.6

0.66

8

22

30

7.7

0.44

21

28

1

50

7.6

0.53

12

14

26

7.5

0.50

1

31

18

50

7.3

0.51

86

40

101

15

193

55

7.6

7.3

0.55

0.45

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

21

Caney Fork R. System: Tennessee: Coffee Co.: UAIC 2417 (1), 2536
(1). Cannon Co.: INHS 82553 (1). Warren Co.: UT 91.646 (1).

Lower Cumberland R. Dr.: Kentucky: Lyon Co.: INHS 75877 (12);
KU 11581 (1); UMMZ 174973 (7). Trigg Co.: KU 10722 (1). Caldwell
Co.: INHS 75740 (3); UL 5080 (2), 6478 (2); UMMZ 174869 (7).
Livingston Co.: INHS 78519 (26), 75863 (14), 75874 (12), 75875 (23),
75876 (1), 79367 (7); MEB 134 (23). Christian Co.: INHS 77702 (9); UL
10780 (29); UMMZ 154770 (5). Tennessee: Houston Co.: INHS 75814
(1). Stewart Co.: INHS 83122 (2), 77739 (3); UMMZ 161005 (8), 168371
(1).

Description — Etheostoma crossopterum is a large, robust Catonotus;
largest specimen a male 83.5 mm SL. Males generally larger than females,
largest female 61 mm SL. Infraorbital canal incompete with narrow
interruption below eye, pores normally 4-3, occasionally 4-4; preoper-

Table 8. Counts of pectoral fin rays in species of the Etheostoma squamiceps species group.

Species

&

Population

Number

of Pecto

ral Fin

Rays

10

11

12

13

N

X

SD

Etheostoma squamiceps

Cache River

2

32

34

10.9

0.24

small tribs.. Ohio River

4

29

2

35

10.9

0.41

Wabash River

2

12

14

10.9

0.35

Tradewater River

2

27

6

35

11.1

0.46

Lower Green River

11

3

14

11.2

0.41

Upper Green River

1

23

1

25

11.0

0.28

Barren River

14

14

11.0

0.00

Red River

7

35

42

10.8

0.37

TOTALS

18

183

12

213

11.0

0.36

Etheostoma neopterum

Clarks River

16

37

2

55

11.7

0.51

Big Sandy River

4

43

4

51

12.0

0.40

Duck River

3

16

1

20

11.9

0.44

Shoal Creek

1

22

3

26

12.1

0.39

TOTALS

24

118

10

152

11.9

0.44

Etheostoma crossopterum

Lower Cumberland River

2

50

14

66

11.2

0.46

Middle Cumberland River

11

46

1

58

10.8

0.42

direct tribs., Mississippi River

2

8

2

12

11.0

0.58

Duck River

1

29

20

50

11.4

0.53

Cypress Creek

4

21

8

33

12 I

0.59

TOTALS

16

137

58

8

219

11.3

0.49

Etheostoma nigripinne

western tribs., Tennessee River

5

30

2

37

10.9

0.43

eastern tribs., Tennessee River

13

17

30

11.6

0.50

southern tribs., Tennessee River

19

30

1

50

11.6

0.52

Shoal Creek

23

3

26

III

0.32

Elk River

1

36

13

50

11.2

0.47

TOTALS

6

121

65

1

193

11.3

0.47

Etheostoma olivaceum

Caney Fork River

4

84

12

100

12.1

0.39

22

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TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

23

culomandibular canal pores 10; supraorbital canal pores 4 or 5; supratem-
poral canal interrupted medially, pores 2-2; lateral canal pores 5; frenum
narrow to moderate; branchiostegal rays six, membranes slightly to
moderately joined; preopercle entire. Genital papilla of females broad,
flat, and darkly pigmented.

Cheeks, opercles, breast and nape usually scaled. Lateral scale rows
38-64, with 7-44 pored, and 8-46 unpored; transverse scale rows below
lateral line 7-10 (usually 8-9), above lateral line 5-7 (6).

First dorsal fin with 8-10 (8-9) spines; second dorsal rays 11-15
(12-14); anal spines 2, anal rays 6-9 (7-8); branched caudal rays 14-17;
pectoral rays 10-13 (1 1-12).

Body proportions: HL/SL 0.26-0.32 (X = 0.29); BD/SL 0.15-0.25
(0.21); CPD/SL 0.09-0.16 (0.13); P.L/SL 0.16-0.33 (0.23); P7L/SL

Table 10. Number of transverse scale rows above lateral line in species of the Etheostoma

squamiceps species group.

Species Number of

& Scales above lateral line

Population 5 6 8 N X SD

Etheostoma squamiceps

Cache River 26 8 34 6.2 0.42

small tribs., Ohio River 8 27 35 5.7 0.42

Wabash River 1 4 8 1 14 6.6 0.71

Tradewater River 2 16 17 35 6.4 0.59

Lower Green River 11 3 14 6.2 0.41

Upper Green River 8 17 25 5.6 0.46

Barren River 3 9 2 14 5.9 0.59

Red River 2 31 8 1 42 6.1 0.54

TOTALS 24 141 46 2 213 6.1 0.59

Etheostoma neopterum

Clarks River 38 17 55 5.3 0.46

Big Sandy River 1 1 39 1 51 5.8 0.44

Duck River 6 14 20 5.7 0.45

Shoal Creek 3 22 1 26 5.9 0.39

TOTALS 58 92 2 152 5.7 0.44

Etheostoma crossopterum

Lower Cumberland River 4 58 4 66 6.0 0.34

Middle Cumberland River 6 40 12 58 6.1 0.54

direct tribs., Mississippi River 2 9 1 12 5.9 0.49

Duck River 9 33 8 50 5.9 0.58

Cypress Creek 16 17 33 5.5 0.50

TOTALS 37 157 25 219 5.9 0.50

Etheostoma nigripinne

western tribs., Tennessee River 3 33 1 37 5.9 0.32

eastern tribs., Tennessee River 21 9 30 5.3 0.45

southern tribs., Tennessee River 8 40 2 50 5.8 0.43

Shoal Creek 2 24 26 5.9 0.26

Elk River 30 20 50 5.4 0.49

TOTALS 64 126 3 193 5.6 0.42

Etheostoma olivaceum

Caney Fork River 2 37 55 6 100 6.7 0.62

24

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

0.09-0.25 (0.19); PreOL/HL 0. 17-0.30 (0.24); EW/HL 0. 17-0.32 (0.25);
DFH/DFB 0.19-0.59 (0.29).

Coloration of freshly preserved specimens: Coloration similar to E.
squamiceps. Basic coloration of females and non-breeding males consists
of an irregular mottling of brown on light tan background. Dark humeral
spot large. Base of caudal fin with three well developed dark spots; middle
caudal spot often considerably darker than other two. Dorsum variable,
usually colored same as sides, occasionally paler and crossed by 8-9 short,
indistinct saddles. Venter usually unmarked. Markings on head typically
consist of pre-, post-, and suborbital dark bars. Cheeks often mottled and
may have a dark, diagonal line from below eye posterodorsally connecting
with postorbital bar at anterior edge of opercle.

First dorsal fin with faint, subdistal orange band formed by row of

Table 11. Number of transverse scale rows below lateral line in species of the Etheostoma

squamiceps species group.

Species

Number of

&

Scales below lateral line

N

X

Population

6 7

8

9

10

11 12

SD

Etheostoma squamiceps

Cache River

5

18

10

1

34

8.2

0.71

small tribs. , Ohio River

1

19

14

1

35

8.4

0.59

Wabash River

2

5

7

14

8.3

0.71

Tradewater River

4

14

13

4

35

9.4

0.84

Lower Green River

4

7

3

14

8.9

0.70

Upper Green River

1

15

9

25

8.3

0.54

Barren River

3

9

2

14

8.9

0.70

Red River

2

12

18

10

42

8.8

0.83

TOTALS

11

80

88

30

4

213

8.7

0.73

Etheostoma neopterum

Clarks River

16

33

6

55

8.8

0.60

Big Sandy River

11

37

3

51

8.8

0.50

Duck River

12

8

20

8.4

0.49

Shoal Creek

5 20

1

26

6.8

0.57

TOTALS

5 20

40

78

9

152

8.2

0.54

Etheostoma crossopterum

Lower Cumberland River

17

41

8

66

8.8

0.60

Middle Cumberland River

2

21

33

2

58

8.6

0.61

direct tribs.. Mississippi River

1

1

10

12

8.7

0.60

Duck River

9

31

10

50

8.2

0.96

Cypress Creek

6

18

8

1

33

8.1

0.75

TOTALS

18

88

102

11

219

8.4

0.73

Etheostoma nigripinne

western tribs., Tennessee River

2

26

9

37

8.1

0.51

eastern tribs., Tennessee River

6

21

3

30

7.9

0.53

southern tribs., Tennessee River

1 4

15

27

3

50

8.5

0.80

Shoal Creek

1

16

8

1

26

8.3

0.38

Elk River

14

35

1

50

7.7

0.48

TOTALS

1 27

113

48

4

193

8.1

0.62

Etheostoma olivaceum

Caney Fork River

2

16

42

32

7 1

100

9.3

0.92

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 25

short, vertical bars located between spines. Each bar consists of cluster of
dark melanophores overlaid by orange pigment which fades rapidly in
preservative. Proximal portion of membrane colorless except for scattered
melanophores, sometimes forming indistinct basal band. Base of fin
thickened; membrane slightly thickened at tip of spines, forming in-
conspicuous light colored knobs in both sexes. Soft dorsal fin with clusters
of melanophores forming 3-7 interrupted bands plus a rather indistinct
basal band; basal part of fin thickened. Caudal fin with melanophores
clustered along rays forming 6-8 narrow, dark, vertical bands. Anal and
pelvic fins unpigmented; pectorals unpigmented to lightly barred with up
to 9 narrow dark bands.

Breeding male coloration: Overall coloration dark (Fig. 8). Sides
uniformly dark brown or gray; caudal spots enlarged to form vertical bar;
dorsum uniformly dark; venter dark gray; head much darker, almost
black, often obscuring regular markings; lower part of head greatly
swollen. During courtship and egg-care, sides develop pattern of alternat-
ing dark and light bars (Fig. 8). First dorsal fin black except for large light
yellow or white knobs on spines and small, clear, round or tear-shaped
windows behind each spine; proximal part of fin with narrow clear band;
basal part of membrane thickened (Fig. 2). Second dorsal fin black with a
broad subdistal white band and 3;6 rows of short, pale dashes across rays,
dashes in some specimens appear to connect, forming wavy lines across
fin; tips of rays greatly elongated and extending far beyond membrane
margin, tips of rays black and lacking the large fleshy knobs of E.
squamiceps and E. neopterum (Fig. 3). Anal fin dark with pale distal
margin; membrane thickened around first spine. Pelvic fins dark except
for pale membrane around spine and light colored distal band. Caudal and
pectoral fins unchanged from non-breeding coloration.

Distribution— Etheostoma crossopterum occurs in the lower and mid-
dle Cumberland River drainage (except Red River) from mouth east to
Plunkett Creek in Smith County, Tennessee and throughout the Duck
River system (Tennessee Drainage) in central Tennessee. Isolated popula-
tions occur in the headwaters of Caney Fork River in central Tennessee,
direct tributaries of Mississippi River in western Tennessee and in Cypress
Creek in southern Tennessee and northern Alabama (Fig. 9).

Variation — Etheostoma crossopterum shows little sexual dimorphism
except for size and pigmentation. Males have more pored lateral-line
scales, fewer unpored lateral-line scales, deeper caudal peduncles, smaller
eyes (a=.001), and a longer preorbital length (a=.05) (Table 1).
Ontogenetic variation is more pronounced. Small specimens (<37.5 mm
SL) have fewer pored lateral-line scales, more unpored lateral-line scales,
larger heads, longer pectoral fins, higher first dorsal fins (a= .001), and
longer pelvic fins (a= .01) (Table 2).

To examine geographical variation, the range of E. crossopterum was
divided into five areas. The Cumberland River was divided at the mouth of
the Red River into an upstream middle Cumberland population and a
downstream lower Cumberland population. Additional populations in-

26

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

35-

37

_..3^- 35

89

86

Figure 9. Distribution of Etheostoma crossopterum.

elude Mississippi River tributaries in the Reelfoot Lake area of Tennessee,
the Duck River, and Cypress Creek.

Pigmentation: Although geographic variation in squamation and me-
ristics is extensive, there is little variation in pattern or coloration between
populations. Breeding males from the southernmost Cypress Creek
population are indistinguishable in body pigmentation or fin shape and
pattern from those of the centrally located Duck River or northern
Cumberland River. Within any one population, individual variation in
degree of breeding coloration attained is far greater than any variation
between populations.

Squamation: Several specimens from direct tributaries of the Mis-
sissippi River show reduced squamation on the nape, cheeks and opercles
(Figs. 5-7). Many specimens from throughout the range, except Cypress
Creek, lack scales on the cheek like the related E. olivaceum (Fig. 7)
which has a naked cheek. Variation also occurs in the degree of scale
exposure on the nape and operculum. The highest frequency of exposed
scales occurs in specimens from Cypress Creek (Figs. 5, 6).

Meristics and Morphometries: Anal fin rays, dorsal fin spines, dorsal
fin rays, pectoral fin rays, total lateral-line scales, pored lateral-line scales,
unpored lateral-line scales, transverse scale rows above lateral line,
transverse scale rows below lateral line, HL/SL, P,L/SL, P-.L/SL, CPD/
SL, EW/HL, BD/SL, and PreOL/HL all differed significantly between
populations (a= .05) (Tables 5-12).

Most characters appear to vary in a north-south cline. Although not

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

27

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28 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

included in the tables because of small sample size, specimens from
southern tributaries of Buffalo River are intermediate between those of
Cypress Creek and the Duck River population. Mississippi River speci-
mens vary independently from the main cline, probably indicating a long
isolation from the main gene pool.

Anal rays generally number 7 in all populations except Duck River
which averages slightly more (Table 7). Pectoral rays 1 1 in all populations
except Cypress Creek which modally has 12 (Table 8). Scales below
lateral line essentially 9 in populations from the Cumberland and Mis-
sissippi drainages and 8 in the Duck River and Cypress Creek systems
(Table 11). Scales above lateral line modally 6 except Cypress Creek
which is bimodal for 5 or 6 (Table 10). Dorsal spines modally 9 except in
populations from the Cumberland and Duck systems which have a
relatively high proportion of individuals with 8 spines (Table 5). Dorsal
rays highly variable between populations, ranging from a mode of 12 in
the Cumberland and Mississippi river drainages, to 13 in Duck River and
14 in Cypress Creek (Table 6).

Habitat and Life History— The preferred habitat of E. crossopterum is
small, quiet streams with large flat rocks or bedrock bottom. In streams of
this type in the Nashville Basin, E. crossopterum may be the most
abundant fish species. Although E. crossopterum has the same preferred
habitat as E. squamiceps, it shows less rigid habitat requirements. Small
populations may be found in small gravelly streams and in larger streams
along quiet margins and beneath undercut banks. Darter species most
commonly associated with E. crossopterum are E. simoterum atripinne
and E. smithi in the Cumberland River and E. luteovinctum and the golden
snubnose darter, E. (Nanostoma) sp. in the Duck River. Other species
commonly found with E. crossopterum are Campostoma anomalum,
Pimephales notatus, Semotilus atromaculatus, Fundulus catenatus, Lepo-
mis cyanellus, and Cottus carol inae.

Page (1974) gives detailed information on reproduction, growth, food
habits, population composition, predation, and parasitism on this species
from Richland Creek, Cumberland River drainage. As do other species of
Catonotus, E crossopterum spawns in cavities under rocks and attaches
the eggs to the undersides of stones (Fig. 10).

A natural hybrid E. crossopterum x E. luteovinctum was collected 21
July 1978 from a tributary of Wartrace Creek, Bedford County, Tennessee
by M. E. Braasch, P. W. Mayden, and M. Wright. This is the only known
E. crossopterum hybrid.

Etheostoma nigripinne new species
Blackfin Darter

Figs. 2, 3, 10, 11.

Holotype— University of Kansas, KU 20000, breeding male 56.5 mm
SL (Fig. 1 1), Pikes Peak Branch, tributary to the Tennessee River 1.6 km
W of TN Route 20 bridge, Decatur County, Tennessee, 5 April 1981,
Marvin E. Braasch.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

29

Paratopes— A total of 52 specimens deposited as follows: 2— Univer-
sity of Kansas (KU 20894, 50-59 mm SL, same data as holotype); 7— KU
20893 (36-52 mm SL, small tributary to Tennessee River, 0.5 km W
Tennessee River on TN Rt. 20, Decatur Co., TN, 14 May 1978); 2—
University of Alabama Ichthyological Collection (UAIC 7712.01, 43-49.5
mm SL, same data as KU 20893); 7— University of Michigan Museum of

\/

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Figure 10. Nest-guarding males and eggs off. crossopterum (Top, Bear Creek, Maury Co.,
TN) and E. nigripinne (Bottom, Whites Creek, Decatur Co., TN). Each male was guarding
the nest of eggs shown above before their removal. Photographs by Thomas M. Keevin.

30

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Zoology (UMMZ 212481, 28.5-51.7 mm SL, same collection data as KU
20893); 7-United States National Museum (USNM 268083, 35.6-53.3
mm SL, same data as KU 20893); 7— University of Tennessee (UT
91.2768, 35.9-61.5 mm SL, same data as KU 20893); 7— Illinois Natural
History Survey (INHS 68092, 33.8-62 mm SL, same data as KU 20893);
7— KU 20895 (36-46.2 mm SL, Whites Creek, 4 mi N Bath Springs,
Decatur Co., TN, 4 April 1978); 3— Southern Illinois University at
Carbondale (SIUC 9718, 39.8-60 mm SL, same data as KU 20895); 3—
Tulane University (TU 139110, 39.3-47.5 mm SL, same data as KU
20895).

Etymology— The specific name, nigripinne, comes from the Latin
"nigrum" black and "pinna" wing or fin and, along with the common
name, refers to the striking black dorsal, anal, and pelvic fins of breeding
males.

Diagnosis— Infraorbital canal incomplete; modally 8 transverse scale
rows below lateral line, modally 8 dorsal spines and 8 anal rays; cheeks
and opercles scaled; breeding males with second dorsal fin lacking pale
distal band and without knobs on rays; pelvic and anal fins black; breeding
males very dark.

Material examined— Specimens examined during this study, but not
designated as types are listed below by drainage and river system.

Western Tributaries Tennessee R.: Decatur Co.: INHS 27485 (19);
MEB 109 (37), 152 (3), 153 (15), 154 (1), 155 (7), 156 (1), 157 (12); UT
91.1047 (6), 91.1948 (3), 91.1172 (1).

■»»*(

Figure 11. Etheostoma nigripinne, new species. Top, Holotype, male 56.5 mm SL (KU
20000); Bottom, Paratype, female 43.8 mm SL (KU 20893).

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 31

Eastern Tributaries Tennessee R.: Perry Co.: MEB 158 (8), 170 (5);
UMMZ 177649 (5). Hardin Co.: MEB 103 (8), 104 (8), 105 (18), 163
(17), 164 (9), 165 (2), 166 (4), 167 (6), 168 (24), 169 (7), 171 (1); UT
91.254 (49), 91.1584 (2), 91.1594 (16). Wayne Co.: MEB 159 (7), 160
(7). 161 (1), 162 (7).

Northern and Southern Tributaries Tennessee R.: Tennessee:
Lincoln Co.: MEB 172 (4); UAIC 1972.05 (5), 1974.09 (2). Mississippi:
Tishomingo Co.: INHS 76082 (14), 76100 (16); UAIC 1350.04 (1),
2190.11 (5), 2299.12 (1). Alabama: Franklin Co.: KU 20920 (22); MEB
173 (6); UAIC 1883.08 (12), 1914.07 (10), 1921.07 (10), 2131.05 (1),
2132.09 (28), 2133.09 (13), 2135.07 (3), 2177.06 (1), 2178.13 (1),
2325.17 (5), 2522.10(1); UMMZ 1 13930 (1). Colbert Co.: UAIC 865.09
(5), 2136.10 (4), 2137.07 (2); UMMZ 114702 (7), 132424 (2), 132485
(1). Lauderdale Co.: INHS 27488 (16); MEB 174 (14); UAIC 2001.11
(1), 2002.01 (22). Lawrence Co.: MEB 175 (4); UAIC 1336.07 (1),
1337.06 (5), 1338.09 (2). Limestone Co.: MEB 176 (1); UMMZ 103760
(1), 114720(1), 115030(1), 115064(1), 159175 (1); UT 91.1423 (143).
Madison Co.: UAIC 1966.04 (3), 2543.08 (1). Jackson Co.: INHS 27511
(1); UMMZ 96468 (2). Morgan Co.: MEB 177 (1).

Shoal Creek System: Tennessee: Lawrence Co.: MEB 178 (20), 179
(6); UT 91.98 (5).

Elk R. System: Tennessee: Giles Co.: INHS 75398 (2); MEB 180
(18); UAIC 1978.01 (27), 2704.08 (5), 2794.09 (9), 2796.06 (1), 2869.09
(2), 2871.08 (1), 2872.13 (3), 2873.08 (3), 2955.10 (1), 2956.05 (4),
3201.13 (2), 3202.05 (1). Lincoln Co.: MEB 181 (16), 182 (10); UAIC
1970.04 (15), 1981.04 (3), 2944.03 (3), 2957.03 (3), 2965.16 (1),

2966.13 (21), 2967.12 (3), 2968.14 (21), 2971.06 (2), 2972.09 (12),
3203.17 (2), 3221.02 (3). Franklin Co.: UAIC 1978.01 (1), 2864.15 (8),

2865.14 (1), 2934.02 (2). Marshall Co.: KU 10687 (10), UAIC 2702.04
(1), 2969.03 (3). Coffee Co.: KU 12075 (1), UAIC 2831.07 (1), 2947.05
(14). Moore Co.: UAIC 2940.03 (1), 2948.05 (17). Alabama: Limestone
Co.: MEB 183 (14); UAIC 2497.09 (12), 2792.05 (2), 2961.07 (1).
Lauderdale Co.: UAIC 1987.05 (4), 3130.13 (2).

Duck R. System: Tennessee: Humphreys Co.: INHS 82932 (7).
Bedford Co.: UT 91.2056 (7). Coffee Co.: UT 91.627 (12).

Description— Etheostoma nigripinne is one of the smaller members of
the E. squamiceps group; largest specimen a male 63.5 mm SL. Males
generally larger than females, largest female 59.3 mm SL. Specimens of
either sex over 55 mm SL are rare. Infraorbital canal incomplete with
narrow interruption below eye, pores normally 3-4; preoperculomandibu-
lar canal pores 10; supraorbital canal pores 4 or 5; supratemporal canal
interrupted medially, pores 2-2; lateral canal pores 5; frenum moderate to
wide; branchiostegal rays 6, membranes slightly joined or free; preopercle
entire to slightly crenulate. Genital papilla of adults enlarged; breeding
females with a broad, flat and darkly pigmented papilla.

Cheeks, opercles, breast, and nape usually scaled. Lateral scale rows

32 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

42-58, with 10-45 pored, and 6-35 unpored; transverse scale rows below
lateral line 6-10 (usually 8), above lateral line 5-7 (5 or 6).

First dorsal fin with 7-10 spines (8); second dorsal rays 1 1-15 (13-14),
anal rays 6-9 (7-8); pectoral rays 10-13 (11-12).

Body proportions: HL/SL 0.27-0.33 (X-0.29); BD/SL 0.15-0.24
(0.21); CPD/SL 0.10-0.16 (0.13); P.L/SL 0.20-0.27 (0.23); P.L/SL
0.16-0.24 (0.20); PreOL/HL 0.17-0.33 (0.25); EW/HL 0.19-0.32 (6.26);
DFH/DFB 0.13-0.44 (0.27).

Coloration of freshly preserved specimens: Coloration similar to E.
squamiceps and E. crossopterum. Basic coloration of females and non-
breeding males consists of brown mottling on a tan background (Fig. 1 1).
In larger males, particularly from the southern part of range, mottling
tends to coalesce on sides to form a series of 12-13 small dark blotches
along lateral line. Small specimens have three dark caudal spots, but most
adults retain only the central one. Large dark humeral spot present. Dorsal
coloration variable. Specimens from southern populations have 8 dark
cross bars; third cross bar larger and darker than others and located
between dorsal fins. Cross bars much less prominent in specimens from
northern parts of range. Venter normally unmarked. Markings on head
typically consist of pre-, post-, and suborbital dark bars.

First dorsal fin with distinct subdistal orange band formed by a row of
vertical bars located between spines. Bars consist of clusters of
melanophores overlaid by orange pigment. Band much larger and brighter
in males than females. Most of membrane proximal to orange band dusted
with melanophores— thickly in large specimens, thinly in smaller. Basal
portion of membrane thickened; membrane slightly thickened at tips of
spines, forming inconspicuous light colored knobs. Soft dorsal fin with
clusters of melanophores along rays forming 3-8 interrupted bands plus an
indistinct basal band; basal part of membrane thickened. Caudal fin with
6-13 dark vertical bands. Anal and pelvic fins unpigmented; pectorals
unpigmented to lightly barred.

Breeding males very dark (Fig. 11); much darker than any other
species of the squamiceps group. Head, flanks, and dorsum black; venter
slightly paler; lower part of head swollen. During egg-care, sides may
develop a pattern of alternating vertical light and dark bars, similar to,
although more indistinct than, E. squamiceps, E. olivaceum or E.
crossopterum. Unlike these species, immediately upon preservation E.
nigripinne loses these bars and develops the jet black coloration. First
dorsal fin black except for large round, pale colored knobs on spines and
small round clear windows behind each spine; basal part of membrane
thickened and with narrow clear band above basal dark band (Fig. 2).
Second dorsal fin in highest males totally black; other males with 1-6 rows
of small pale dots or dashes across rays (Fig. 3). Rays elongated past
margin of fin in approximately 50% of the specimens. Basal part of fin
thickened and sometimes with clear band; distal half of membrane often
thickened and partially obscuring black color, giving appearance of gray
margin. Anal and pelvic fins black. Caudal fin with dark bands much

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 33

enlarged, but usually not touching. Pectoral fins unchanged from non-
breeding coloration.

There is variation in breeding color at any one location. Only a small
percentage of males will show strong breeding coloration. Whether this is
due to a very short breeding period per male or to individual variation is
unknown; however great care must be taken when identifying specimens
because non-breeding males resemble E. crossopterum males in having
pale borders to the second dorsal, anal, and pelvic fins.

Distribution— Etheostoma nigripinne occurs in tributaries of the Ten-
nessee River from the lower Duck River upstream to the Flint and Paint
Rock rivers (Fig. 12). At least two populations are also known from the
upper Duck River System in Coffee and Bedford counties, Tennessee, a
drainage typically occupied by E. crossopterum. Blackfin darters are
particularly abundant in the Elk River in southern Tennessee. The status of
E. nigripinne in Cypress Creek and southern tributaries of the Buffalo
River warrants further investigation. Although no specimens are known
from either place and both rivers contain populations of the closely related
E. crossopterum, small populations of E. nigripinne could occur in this
area. Etheostoma nigripinne occurs sympatrically with E. neopterum in
several creeks in Tennessee and Alabama and possibly with E. crossop-
terum in the Duck River in Tennessee. The occurrence of both E.
nigripinne and E. crossopterum in the Duck system is not unexpected since
other researchers have also found close relatives occurring together in this
system. Etnier and Jenkins (1980) reported two forms of Noturus elegans
in the Duck River System and E. duryi and "the golden snubnose" both
are known from the Duck (Etnier, pers. comm.)

Variation— Etheostoma nigripinne shows little sexual dimorphism
except for size and pigmentation. Males have deeper caudal peduncles and
smaller eyes (a =.05) than females (Table 1). Ontogenetic variation is
more pronounced. Small specimens (<40 mm SL) have longer heads,
longer pelvic fins, fewer pored lateral-line scales (a =.05), larger eyes,
and higher first dorsal fins (a = .001) than do larger specimens (Table 2).

To evaluate geographic variation, the range of E. nigripinne was
divided into five areas. Direct tributaries of Tennessee River in Decatur,
Benton and Perry counties were designated as western tributaries; Hardin,
Indian and Horse creeks in Wayne and Hardin counties, TN were
designated as eastern tributaries; all tributaries eastward from Tishomingo
County, MS to Paint Rock River in northeast AL with the exception of
Shoal Creek and Elk River were designated as southern tributaries; Shoal
Creek and Elk River were treated separately.

There is extensive variation in pigmentation, meristics, and propor-
tions between populations of E. nigripinne.

Pigmentation: Intensity of pigmentation both in breeding and non-
breeding coloration varies clinally from north to south. Individuals from
Shoal Creek and Elk River have darker dorsal cross-bars, a darker orange
band in the first dorsal fin, and a row of lateral blotches along the lateral

34

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Figure 12. Distribution of Etheostoma nigripinne.

line. Breeding males from the southern tributaries are usually the darkest,
with many individuals being jet black.

Squamation: Little variation occurs with respect to squamation of the
head. Most populations have scales on the cheeks, opercles, and nape
except in Elk River where a small percentage of the population lacks scales
on the cheeks and opercles (Figs. 6, 7). When scales are present they are
most commonly embedded although a considerable number of individuals
have exposed scales on the nape and operculum (Figs. 5, 6).

Meristics and Morphometries: Dorsal fin rays, pectoral fin rays, anal
fin rays, total lateral scale rows, pored lateral-line scales, unpored lateral-
line scales, transverse scale rows above lateral line, transverse scale rows
below lateral line, HL/SL, BD/SL, DFH/DFB, CPD/SL. EW/HL, P,L/
SL, and PreOL/HL all differed significantly (a = 0.05) between popula-
tions off. nigripinne (Tables 6-11, 13).

Most characters varied semi-clinally from upstream to downstream
(southeast to northwest). Further, most values were lowest in Elk River
and highest in western (i.e., CPD/SL, PreOL/HL, EW/HL) or eastern
tributaries (i.e., P,L/SL, anal fin rays). A few characters are reversed and
have high values at the northern end of the cline (i.e., dorsal fin rays,
pectoral rays, pored lateral-line scales).

Habitat and Life History— The preferred habitat of E. nigripinne is
small bedrock bottom creeks, often less than 1 m wide. A small creek
seems to be more important than substrate; in northern Alabama small

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

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populations exist in mud bottom creeks where they hide under debris. The
darter species most commonly associated with E. nigripinne is E. duryi,
with which it is almost always found. Other species of fishes commonly
occurring with the blackfin darter are Hemitremia flammea (eastern part of
range), Phoxinus erythrogaster (western part of range), Clinostomus
funduloides , Campostoma anomalum, Semotilus atromaculatus and Cottus
carolinae.

Although males begin to acquire breeding coloration in late March and
early April, they do not attain full coloration of their dorsal fins until
spawning time and begin to lose the coloration after the eggs hatch. Males
guarding eggs were found in a tributary of Cane Creek, Lincoln Co., TN
on 30 April 1978 and in a tributary of Hardin Creek, Wayne Co., TN on
15 May 1978. Eggs were fastened to the undersides of flat rocks (Fig. 10)
as in E. crossopterum, E. olivaceum, and E. squamiceps (Page, 1974;
1980) and some E. neopterum (Page and Mayden, 1979). No other life
history data are available.

Etheostoma neopterum Howell and Dingerkus
Lollypop Darter
Figs. 2, 3, 13.
Etheostoma neopterum. Howell and Dingerkus, 1978.

Types — Etheostoma neopterum was described from Little Butler
Creek, Wayne Co., Tennessee. Holotype USNM 217529. Paratypes
deposited at USNM, CU, AMNH, UMMZ, TU. UAIC, and UT.

Diagnosis— Etheostoma neopterum may be separated from all other
Catonotus in having branches of dorsal fin rays adnate, modally 11-12
dorsal rays, and 12 pectoral fin rays. Rays of second dorsal fin of breeding
males elongate and with expanded knobs distally. Second dorsal fin
membranes of males with round clear areas.

Material examined— Specimens examined during this study are listed
below by drainage and system.

Mississippi R. Dr.: Bayou Du Chien System: Kentucky: Hickman
Co.: UL 5992 (1). Graves Co.: KU 20900 (1).

Tennessee R. Dr.: Clarks R. System and Tributaries Lower
Tennessee R.: Kentucky: Marshall Co.: INHS 75759 (1), 75784 (3),
77680 (11); KU 12744 (2), 20902 (8); MEB 85 (41); UL 12689(7), 12867
(18). Calloway Co.: INHS 78268 (1), 75540 (2), 75616 (25), 77602 (2),
77773 (23); KU 20901 (3); MEB 86 (15). Graves Co.: SIUC uncat. (7);
UL 12693 (3). Livingston Co.: SIUC uncat. (77).

Big Sandy R. System: Tennessee: Benton Co.: INHS 79348 (2).
79351 (8); KU 16271 (57), 20762 (1); MEB 140 (5), 141 (2), 142 (50);
TU 14713 (3); UT 91.194 (37). Henderson Co.: KU 9809 (2). Carroll
Co.: KU 9917 (1). Humphreys Co.: INHS 77624 (3), 77810 (23); MEB
143 (3), 144 (6), 145 (9); UT 91.78 (1). Decatur Co.: MEB 91 (2), 146
(8). Henry Co.: INHS 77780 (12). Houston Co.: MEB 147 (3).

Duck R. System: Tennessee: Hickman Co.: INHS 77593 (3), 77786

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

37

(2), 77817 (4); KU 14399 (13), 16216 (22); MEB 97 (20). Wayne Co.:
MEB 136(2). 137(1), 138(1). Maury Co.: MEB 139(1). Perry Co.: KU
17684 (8), 20903 (1).

Eastern Tributaries Tennessee R.: Tennessee: Perry Co.: MEB 148
(2), 151 (4). Hardin Co.: MEB 149 (1), 150 (2).

Shoal Creek System: Tennessee: Lawrence Co.: MEB 135 (2), 186
(4). Wayne Co.: CU 64511 (21), 64513 (9). Alabama: Lauderdale Co.:
CU 64515 (7).

Description— The following description is based primarily on material
at CU, KU, and INHS. Data in parentheses include extreme values
reported in Howell and Dingerkus (1978).

Etheostoma neopterum (Fig. 13) is a medium-sized Catonotus, with
males (up to 63 mm SL) generally larger than females (maximum 61 mm
SL). Infraorbital canal complete with 8 pores (northern portion of range)
or incomplete, pores 4-3 or 4-4 (southern portion of range); preoperculo-
mandibular canal pores 10; supraorbital canal pores 4 or 5; supratemporal
canal interrupted, pores 2-2; lateral canal pores 5; frenum moderate;
branchiostegal rays 6 per side, membranes free to moderately joined;
preopercle entire. Genital papilla enlarged; breeding females with broad
and flat papilla.

Cheeks, opercles, breast, and nape usually scaled. Total lateral scale
rows 43-55 (usually 41-58), with 14-45 pored and 7-33 unpored; trans-
verse scale rows above lateral line 5-7 (usually 5-6), below lateral line
8-10 (usually 8-9).

VWM*;.*.

<?*$£

Figure 13. Etheostoma neopterum. Top, male 61.9 mm SL (KU 20900, Bayou de Chien,
Graves Co., KY); Bottom, female 55.5 mm SL (KU 9809, Big Sandy River, Henderson Co.,

TN).

38 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Spinous dorsal with 8-10 spines (usually 8-9); second dorsal rays
10(9)-14 (usually 11-13); anal spines 2, anal rays 6-9 (usually 7-8);
pectoral rays 11(10)-13 (usually 12).

Body proportions: HL/SL 0.27-0.36 (X = 0.30), BD/SL 0.16-0.22
(0.20); CPD/SL 0.11-0.15 (0.13); P,L/SL 0.19-0.27 (0.23); P.L/SL
0.17-0.23 (0.20); PreOL/SL 0.19-0.30 (0.24); EW/HL 0.22-0.35 (6.27);
DFH/DFB 0.27-0.61 (0.35).

Coloration of freshly preserved specimens: Females and non-breeding
males irregularly mottled with brown on tan background, similar to other
species of squamiceps group (Fig. 13). Large dark humeral spot present,
as well as 1-3 dark caudal spots. Head with pre-, post-, and suborbital dark
bars. First dorsal fin clear except for orange distal band formed of clusters
of pigment; other fins lightly barred to clear.

Breeding males darker, sides grey to dark brown (Fig. 13). First dorsal
fin black except for clear basal band, small round clear windows behind
spines, and large yellow or white, round, fleshy knobs at tip of each spine
(Fig. 2). Second dorsal black except for clear basal band and 2-4
horizontal rows of clear round spots, which may be only anterior to a ray
or on both anterior and posterior sides of a ray, occasionally connected by
a narrow crescent or semi-horizontal pale line. Tips of greatly elongated
rays with large yellow or white fleshy knobs (Fig. 3). Anal and pelvic fins
dark, caudal and pectoral fins similar to non-breeding males. More
detailed descriptions, including photographs, are given by Howell and
Dingerkus (1978).

Distribution — Etheostoma neopterum occurs in tributaries of Ten-
nessee River from the mouth south to Shoal Creek in northern Alabama
and in tributaries of Mississippi River in western Kentucky (Fig. 14). The
lollypop darter is very abundant in the Clarks River area of western
Kentucky and the Big Sandy River area of Tennessee and may be the most
common species of fish in some small creeks in western Kentucky. In the
southern half of its range it is quite sporadic, most collections contain only
a few individuals. It occurs sympatrically with E. nigripinne in direct
tributaries of Tennessee River from Decatur Co., TN south to Shoal
Creek, AL and with E. crossopterum in Duck River. Because of
difficulties in separating small E. neopterum from E. crossopterum and E.
nigripinne, E. neopterum may be more common in the southern portion of
its range than these records indicate.

Variation— Since Howell and Dingerkus did not use specimens from
the northern or eastern portions of the range in their description and
analysis of variation, data on specimens from the Clarks, Duck, and Big
Sandy rivers and Shoal Creek are compared.

Geographic variation is extensive. Anal rays, pectoral rays, dorsal
rays, dorsal spines, total lateral-line scales, pored lateral-line scales,
unpored lateral-line scales, transverse scale rows above and below lateral
line, HL/SL, P2L/SL, BD/SL, DFH/DFB, CPD/SL, and EW/HL (Tables
6-11, and 14) varied geographically.

Variation of most characters is commonly in a north-south cline. Duck

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

39

Table 14. Geographically variable body proportions in Etheostoma neopterum. Measure-
ments and means in hundreths of a millimeter.

HL/SL

EW/HL

BD/SL

Population

N

Range

X

SD

Range

X

SD

Range

X

SD

Clarks River

26

24-

-31

29

0.01

22

-29

26

0.02

16-

-22

19

0.01

Big Sandy River

21

29-

-33

31

0.01

23-

-32

27

0.01

18-

-22

20

0.01

Duck River

16

29-

-33

31

0.01

24-

-33

28

0.02

19-

-22

20

0.01

Shoal Creek

26

27-

-31

29

0.01

18

-26

21

0.02

20-

-23

21

0.01

TOTALS

89

24-

-31

30

0.01

18-

-33

26

0.02

16-

-23

20

0.01

N

P2L/SL

DFH/DFB

CPD/SL

Range

X

SD

Range

X

SD

Range

X

SD

Clarks River

26

17-

-21

19

0.01

28-

-62

34

0.02

11-

-16

12

0.01

Big Sandy River

21

18-

-22

20

0.01

27-

-38

32

0.01

11-

-14

12

0.01

Duck River

16

19-

-22

21

0.01

33-

-45

38

0.02

12-

-15

13

0.01

Shoal Creek

26

18-

-23

20

0.01

21-

-32

29

0.03

10-

-14

12

0.01

TOTALS

89

17-

-23

20

0.01

21-

-62

33

0.02

10-

-16

12

0.01

River specimens often vary from the Clarks River-Shoal Creek cline. Anal
rays usually 7 or 8 in Clarks and Big Sandy rivers, 8 in the Duck River,
and 9 in Shoal Creek (Table 7). Pectoral fin rays modally number 12
except in Clarks River where a fairly high percentage of individuals (30%)
have 1 1 (Table 8). Scale rows below lateral line modally 9 in Clarks and
Big Sandy rivers, 8 in Duck River, and 7 in Shoal Creek (Table 1 1). Scale
rows above lateral line typically 6, except in a high percentage of Clarks
River specimens which have 5 rows (Table 10). Dorsal spines modally 9
except in Duck River where 8 spines are typical (Table 5). Populations
from Clarks River modally have 12-13 dorsal rays, those from Big Sandy
River and Shoal Creek typically have 1 1 rays and Duck River specimens
have 12 (Table 6).

Other characters which show significant variation include the infraor-
bital canal and squamation of the head and nape. Specimens from northern
populations have the infraorbital canal complete with 8 pores while most
specimens from southern populations have the canal incomplete with pores
4-3 or 4-4. Embedded or exposed scales normally present on cheeks,
opercles, and nape. Clarks River and Shoal Creek specimens have the
highest percentage of exposed scales while Duck River specimens lack
exposed scales (Table 3, Figs. 5-7).

Habitat and Life History— Throughout most of its range, E. neopterum
occurs in small to medium size sandy bottom streams where it hides in and
around debris. In Duck River and Shoal Creek it occurs in gravel bottom
pools. Like all species of the squamiceps group, E. neopterum is often
found beneath undercut banks and exposed tree roots at margins of pools.
Darter species most commonly associated with E. neopterum are E.
(Nanostoma) sp. (red snubnose darter), E. gracile, and E. chlorosomum in
the northern portion of the range, and E. caeruleum, E. simoterum, E.
spectabile, E. flabellare, and E. blennius in southern and eastern areas.

40 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Other associated species include Notropis umbratilis, Pimephales notatus,
Semotilus atromaculatus , Erimyzon oblongus, Fundulus olivaceus, and
Lepomis cyanellus in the northern portion of the range, and Campostoma
anomalum, Clinostomus funduloides , Notropis boops, N. chrysocephalus,
N. telescopus, Semotilus atromaculatus, Fundulus catenatus, and Cottus
carolinae in the southern and eastern portions.

Spawning occurs in late April or early May. In suitable habitat, eggs
are fastened to the undersides of flat rocks (Page and Mayden, 1979).
However, they may fasten eggs to sticks or leaves (Howell and Dingerkus,
1978) as do E. gracile and E. chlorosomum. A higher percentage of E.
neopterum males show breeding coloration and dorsal fin morphology
than in other species of the group. The reason for this is unknown. Known
predators of E. neopterum include Cottus carolinae and the salamander
Necturus maculosus (Howell and Dingerkus, 1978).

Etheostoma olivaceum Braasch and Page

Dirty Darter

Figs. 2, 3, 15.

Etheostoma olivaceum. Braasch and Page, 1979

Types — Etheostoma olivaceum was described from Rock Springs
Branch at Rock Springs Church, 2 km N Buffalo Valley, Putnam Co, TN.
Holotype INHS 75734. Paratypes deposited at INHS, USNM, UMMZ,
UT, TU, UAIC, NLU, and KU.

Diagnosis— Scales present on nape, prepectoral area, and belly, but
absent from cheeks and opercles; transverse scale rows above lateral line
usually 7; dorsal fin rays usually 12; infraorbital canal complete; branches
of second dorsal fin not adnate; suborbital bar absent.

Material examined— Materials examined during this study include:
Tennessee: Dekalb Co.: KU 16214 (7). Putnam Co.: KU 13043 (11).
Smith Co.: KU 11442 (33), 14402 (14), 16207 (6). Wilson Co.: KU
11584 (1).

Description — Etheostoma olivaceum (Fig. 15) is a moderate sized
Catonotus; largest specimen a male 67 mm SL, although specimens larger
than 55 mm SL are rare. Males generally larger than females, largest
female 55.5 mm SL. Infraorbital canal complete with 8 or 9 pores. Total
lateral scale rows 44-58, pored lateral-line scales 13-46, unpored 10-37;
transverse scale rows above lateral line 5-8 (6-7) and below lateral line
7-12 (8-10); dorsal spines 8-10 (9), with fleshy knobs in both sexes; dorsal
rays 11-14 (12-13); anal spines 2, anal rays 6-9 (7-8); pectoral rays 11-13
(12); branched caudal fin rays 14-16.

Coloration of freshly preserved specimens: Coloration of non-breed-
ing males and females variable. Pale individuals have indistinct mottling or
vertical bars on a yellow or tan background. Darker individuals may be
uniformly dark grey or olive-brown. Head markings consist of pre- and
postorbital dark bars. No suborbital bar present. Dark humeral spot
present; caudal spots, if present, never darker than other markings. First

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

41

dorsal fin with orange subdistal band (fading to brown in alcohol); rest of
fin colorless except for scattered melanophores; basal portion of fin
thickened; light colored fleshy knobs present on spines. Second dorsal,
caudal, and pectoral fins with dark bands; anal and pelvic fins colorless.

Breeding males very dark (Fig. 15). Sides and head uniform brown or
black; dorsum and venter somewhat paler. Flesh of head swollen. First
dorsal fin black with elongate white knobs on spines and small round clear
windows behind each spine; membranes with small black spot distally
behind each spine (Fig. 2). Second dorsal fin black with 3-5 narrow gold
bands composed of U-shaped markings along each ray. Distal margin of
fin pale; rays produced beyond membranes and with black tips (Fig. 3).
Caudal fin with broad pale margin and 5-10 narrow gold bands alternating
with black bands. Anal fin dark with pale margin. Pelvic fins dark with
white knobs on spines. Breeding tubercles absent. For detailed descrip-
tion, variation, and photographs see Braasch and Page (1979).

Distribution — Etheostoma olivaceum is restricted to small tributaries
of lower Caney Fork River and a few nearby direct tributaries of
Cumberland River in central Tennessee (Fig. 14).

Habitat and Life History— Etheostoma olivaceum occurs in small
bedrock bottom streams where it hides under flat rocks. Occasionally
males may be found in quiet riffles. Darter species commonly associated
with E. olivaceum include E. simoterum, E. flabellare, E. caeruleum, and

37-

35-

-37

35

89

86

Figure 14. Distribution of Etheostoma olivaceum (triangles) and Etheostoma neopterum
(circles).

42

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Figure 15. Etheostoma olivaceum. Top, male 59.3 mm SL; Bottom, female 49.6 mm SL
(KU 11442, Dry Creek, Smith Co., TN).

E. spectabile. Other associated species include Campostoma anomalum,
Notropis ardens, N. boops, N. chrysocephalus , N. galacturus, Semotilus
atromaculatus , Rhinichthys atratulus, Fundulus catenatus, and Cottus
carolinae.

Page (1980) presents a thorough life history of the dirty darter.
Maturity occurs at one year for both sexes. Spawning occurs in late March
to early May. Mature females produce up to 112 mature oocytes. Eggs are
attached to the undersides of flat rocks as in other Catonotus and may
number as many as 1500. Males are highly territorial and singly attend to
nests. Small crustaceans and immature insects comprise most of the diet of
young and adults.

SPECIES COMPARISONS IN THE
ETHEOSTOMA SQUAMICEPS GROUP

Although nuptial males may be easily identified, females and non-
breeding males are often difficult to separate. Most species may be
separated by range. The only known areas of sympatry are Duck River
(crossopterum, neopterum, nigripinne) and direct tributaries of Tennessee
River from Shoal Creek north to Decatur and Perry counties, TN
(neopterum, nigripinne).

Counts and Proportions — All adult E. neopterum (specimens larger
than 40 mm SL) may be identified by adnate branches of dorsal rays.
Etheostoma olivaceum may be separated from all species except some E.
neopterum by its complete infraorbital canal (neopterum may be complete
or incomplete). While no single meristic character is diagnostic, taken as a

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 43

group they help to verify identification. Etheostoma squamiceps differs
from the other species by having shorter pelvic fins, larger eye, lower first
dorsal fin, and (along with neopterum) fewer unpored lateral-line scales.
Etheostoma nigripinne differs from others in generally having 8 dorsal
spines, 8 anal rays, and 8 lateral scale rows below the lateral line.
Etheostoma crossopterum can be separated from the other species meris-
tically by having 7 anal rays and 9 dorsal spines. Etheostoma neopterum
has more anal rays, fewer dorsal rays, and (along with squamiceps) fewer
unpored lateral-line scales. Etheostoma olivaceum has a greater number of
unpored lateral-line scales.

Coloration of Breeding Males— Although the basic pattern is similar
for all five species, major differences occur in dorsal fin color and
morphology. As stated previously, dorsal fin pattern is the last part of the
overall breeding coloration acquired and the first lost. The most likely
errors would be to confuse sub-breeding E. nigripinne or E. squamiceps
with E. crossopterum.

Color of First Dorsal Fin: The basic pattern is to have the membrane
black with a small clear spot or window posterior to each spine and a clear
basal band. Each spine has a light yellow or white fleshy knob. Males off.
squamiceps have small, elongate knobs and the clear windows of the
membrane, when present, are narrow vertical slits. Etheostoma crossop-
terum similarly has elongated small knobs, slightly larger than those off.
squamiceps , and round or teardrop-shaped windows. Etheostoma neop-
terum has large round knobs on spines and round windows located about 2/?
of the way up the fin. Etheostoma nigripinne has large round knobs and
round windows located just below the fleshy knobs. The membranes are
much darker in E. nigripinne than in the other species and the broad, clear
basal band is absent or restricted to only a narrow clear line above a dark
basal band. E. olivaceum has small elongated knobs, round windows and a
clear basal band.

Color and Morphology of Second Dorsal Fin: The basic pattern is a
black membrane with several pale cross bars and a clear basal band. Males
of E. squamiceps have moderately large yellow fleshy knobs distally on
branches of each ray; on each ray several U-shaped pale dashes form
discontinuous horizontal wavy bands (Fig. 3). Males of E. crossopterum
lack fleshy knobs and have elongated, black-tipped rays with a sub-
marginal pale band; rays crossed by pale semi-horizontal to U-shaped
dashes with rows almost touching, forming horizontal pale bands. In E.
neopterum, large fleshy knobs similar to those of E. squamiceps develop
on rays but differs in that branches of rays are adnate with one knob per ray
(squamiceps has two per ray). Fins of northern populations (Bayou du
Chien, Clarks, Big Sandy, Duck rivers) of E. neopterum have rather large
round pale spots anterior to rays, rather than elongated dashes, forming
very disconnected horizontal bands (Fig. 3). Lollypop darters from the
middle Tennessee tend to develop small clear spots anterior and posterior
to rays which may be connected by light concave or horizontal lines. In E.
nigripinne males with maximum breeding coloration, the fin is totally

44 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

black; in males with less than maximum coloration a few horizontal pale
dashes are present. Males of E. olivaceum have elongate rays with sub-
marginal pale band and black tips. Their rays are crossed by several U-
shaped light dashes which connect to form wavy horizontal bands.

Coloration of Body: All species for which life history information is
available have dark heads and sides with pale horizontal bands. Etheos-
toma nigripinne differs from others in that after preservation the body
becomes jet black, while in other species the pale bands remain prominent
with the head much darker than the sides.

Coloration of Females and Non-Breeding Males— Coloration of all
species is very similar. Etheostoma olivaceum differs from all other
species except E. neopterum from the Duck River in lacking a suborbital
bar and by having reduced caudal spots. Most E. olivaceum can also be
separated by their uniform gray or brown coloration as compared to the
irregular mottling of others. Although Howell and Dingerkus (1978)
suggested using nape coloration as a character in separating species of this
group, it has very 'limited usefulness because of extreme intra-specific
variation.

The presence or absence of an orange or gold stripe in the first dorsal
fin is of some diagnostic value. Etheostoma squamiceps lacks a band while
it is very prominent in E. nigripinne. Etheostoma olivaceum, E. neop-
terum, and E. crossopterum are intermediate. This band is useful for
separating sub-breeding male and female E. nigripinne from E. crossop-
terum. The bright orange band clearly shows through the dark pigmenta-
tion of E. nigripinne, while the duller band of E. crossopterum is quickly
hidden by melanophores.

Key to Species of the Etheostoma squamiceps Group

1 . Breeding males with distal fleshy knobs on dorsal fin rays (Fig. 3A-B)
2

Breeding males without fleshy knobs on dorsal fin rays (rays may be
elongated) (Fig. 3C-E) 3

2. Dorsal fin rays with 3 branches, anterior branch reaching margin
separate from posterior two (Fig. 3A); spinous dorsal fin without
orange subdistal band E. squamiceps

Dorsal fin rays with all branches adnate (appearing unbranched) (Fig.

3B); spinous dorsal fin with dull orange subdistal band

E. neopterum

3. Infraorbital canal complete with 8 pores, opercle unsealed

E. olivaceum

Infraorbital canal incomplete (pores 4-3 or 4-4), opercle scaled. . 4

4. Second dorsal in breeding males with broad white sub-marginal band,
rays elongated with black tips (Fig. 3D); first dorsal with dull orange

subdistal band E. crossopterum n. sp.

Second dorsal fin in breeding males without white submarginal band,

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 45

rays elongated or distal part of ray unicolor (Fig. 3E); first dorsal fin
with bright, distinct subdistal orange band, not hidden by melano-
phores E. nigripinne n. sp.

ETHEOSTOMA VIRGATUM SPECIES GROUP

The E. virgatum species group, or barcheek darters, is a clade of five
closely related species. Members occur in the Green, Cumberland, and
Tennessee river drainages of Kentucky and Tennessee. Barcheek darters
are small to moderately large Catonotus characterized by lacking scales on
nape, cheeks, opercles, and prepectoral areas; nuptial coloration of males
with reds and blues (at least in fins) and by the presence of an iridescent
red and white diagonal bar on their cheeks.

Etheostoma virgatum (Jordan)

Striped Darter

Fig. 16.

Poecilichthys virgatus. Jordan, 1880

Types— Poecilichthys virgatus was described from the Rockcastle
River at Livingston, Rockcastle County, Kentucky. Lectotype USNM
23456; paralectotype UMMZ 187511.

New Material Examined: Localities in addition to those listed in Page
and Braasch (1977) include: Tennessee: Canon Co. creek: KU 20945 (16).
Putnam Co., Calfkiller River: KU 11598 (2). Rutherford Co., Overall
Creek: KU 20908 (4), MEB (10). Williamson Co., Mayes Creek: KU

20909 (23); MEB (28).

Diagnosis: Fins with reds and blues, face with iridescent red and white
diagonal bar; infraorbital canal incomplete, pores 4-2; sides with continu-
ous dark longitudinal lines.

Description— Etheostoma virgatum (Fig. 16) is a moderately large
Catonotus reaching approximately 65 mm SL. Infraorbital canal in-
complete, pores 4-2; preoperculomandibular canal pores 10; frenum
moderate. Prepectoral area, nape, cheeks, opercles, and breast naked.
Total lateral-line scales 38-58, 0-25 pored. Dorsal fin spines 7-10 and with
small fleshy knobs; dorsal rays 11-17; anal spines 2, anal rays 7-11;
pectoral rays 9-14.

Coloration of freshly preserved specimens: Females and non-breeding
males light brown dorsally with six dark brown rectangular saddles. Sides
with distinct dark longitudinal stripes formed by a longitudinal, medial
row of melanophores through each row of scales. Stripes most distinct on
dorsal two-thirds of sides. Sides with a series of nine to 11 rectangular
dark blotches. Head dark brown dorsally; well developed preorbital black
bar present, suborbital bar weak or absent. Cheek bar with upper portions
red or orange, lower portion silvery white and whole bar outlined by gold
(fading quickly in preservative). First dorsal fin with black, basal blotch
extending from first or second to fifth or sixth spine; remainder of
membrane often red. Second dorsal, caudal, and pectoral fins with dark

46

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

B

.+J&P&-

- ^^

D

Figure 16. Adult males of species of the Etheostoma virgatum species group. A) E. obeyense
(42.1 mm SL, Cumberland Co., KY. 2 April 1966, KU 11552). B) E. virgatum (45.7 mm
SL, Cannon Co., TN, KU 20945). C) E. striatulum (38.5 mm SL, Bedford Co., TN. 5 April
1969, KU 16218). D) E. smithi (33 mm SL, Livingston Co., KY, 29 May 1972, KU 16227).
E) E. barbouri (50.1 mm SL, Adair Co., KY, 8 April 1969, KU 16226).

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

47

bands, membranes sometimes with red; anal fin orange, red, or yellow
basally.

Breeding males much brighter. Dorsal, caudal, pectoral, and anal fins
mostly red; anal, lower caudal, and lower pectoral (and sometimes first
and second dorsal) with blue-black margin. Black blotch in first dorsal fin
intense; head dark and swollen; cheek bar vivid, lateral stripes dark and
distinct.

For a more detailed description, analysis of variation and photographs
of adults, see Page and Braasch (1977).

Distribution— Etheostoma virgatum may be found in three separate
areas in the Cumberland drainage (Fig. 17). These include the Rockcastle
River area in KY, upper Caney Fork River in central TN, and tributaries
of the middle Cumberland from the Stones to Red River.

Habitat and Life History — Etheostoma virgatum occupies a wider
range of habitats than do most barcheek darters. They may be quite
common in small bedrock creeks, such as those of the Nashville Basin,
where they hide beneath flat rocks. They also occupy gravel bottom pools
and gentle riffles of larger streams such as in the Rockcastle and Harpeth
rivers. As an example of this habitat diversity, in October 1978 two
sizeable collections of E. virgatum were made in the Harpeth River
system, TN. One was from a large stream with moderate current where E.
virgatum were in large gravel bottom pools approximately 30 m wide and
0.5-1 m deep. The second collection was from a small, almost inter-
mittent bedrock bottom creek approximately 3 m wide and 1.5 m deep. In
larger streams, E. virgatum is often associated with emergent vegetation,
such as Dianthera, or it may be found under tree roots or undercut banks;
it is also found in slower riffles and gravel bottom pools with no cover.

Figure 17. Distributions of species of the Etheostoma virgatum species group.

48 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Little is known of the biology of this species. Adults are known to
fasten their eggs on the undersides of rocks in a manner similar to the other
Catonotus (Page, 1983). Presumably males guard their eggs until hatching
occurs. Kuehne and Barbour (1983) found this species to be one of the first
darters to successfully reinvade silted stripmine streams of the Rockcastle
System. They also noted several adults in steady current entering small
tributaries in April, presumably to spawn.

Etheostoma obeyense Kirsch

Barcheek Darter

Fig. 16.

Etheostoma obeyense. Kirsch, 1892

Types— Etheostoma obeyense was described from several tributaries of
Cumberland River, Clinton County, Kentucky. Lectotype USNM 45565,
other types in SU, BMNH, and UMMZ.

Material Examined— Localities in addition to Page and Braasch (1976)
include: Kentucky: Lincoln Co., Green River: KU 7091 (3).

Diagnosis— Fins with reds and blues, face with an iridescent red and
white diagonal bar; infraorbital canal incomplete, pores 4-2; preoperculo-
mandibular pores 10; no dark longitudinal stripes on sides; suborbital bar
weak or absent.

Description— Ethesotoma obeyense (Fig. 16) is a large Catonotus,
reaching 70 mm SL. Infraorbital canal incomplete, pores 4-2; preoper-
culomandibular canal pores usually 10; frenum rather broad;
branchiostegal membranes free or slightly connected. Nape, cheeks,
opercles, and breast naked; total lateral scale rows 39-56, 10-26 pored;
transverse scale rows above and below lateral line 5-9. Dorsal fin spines
8-10 (9) and with fleshy knobs on tips; dorsal rays 12-15 (13); anal spines
2, anal rays 7-10 (9); pectoral rays 11-14 (12).

Coloration: Females and non-breeding males straw-colored dorsally
with 6 to 8 dark saddles. Flanks with a series of 9 to 12 vertically
elongated dark blotches. Fins identical to those of E. virgatum. First
dorsal with black basal blotch and some red in membranes; other fins
lightly barred and often with some red or orange. Head darker dorsally
and with red and white diagonal bar on cheek. Dark preorbital bar present,
suborbital bar absent or very weak. Breeding males similar, but with more
intense colors, head darker and swollen.

For a more detailed description, analysis of variation, and photographs
of adults see Page and Braasch (1976).

Distribution— Etheostoma obeyense is found in tributaries of Cumber-
land River from the Big South Fork west to Obey River in Kentucky and
Tennessee (Fig. 17).

Habitat and Life History— The preferred habitat of E. obeyense is
small bedrock streams where they may be very abundant. In these streams
they particularly tend to hide in crevices in the bedrock. Although we have
collected obeyense from gravel bottom pools in larger streams, they are
very rare outside their preferred habitat.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 49

Page et al. (1981) reported on nesting habits of E. obeyense from
Smith Creek, Clinton Co., Kentucky. Eggs were found attached to the
underside of a stone like other Catonotus. The 100 eggs were approxi-
mately 2 mm in diameter and at various stages of development when found
in late May. Nothing is known about spawning habits of this species,
although they are presumably similar to those of other barcheeks, where
known. Page's (1975a) observations of "E. obeyense" spawning habits
were later determined to be of E. smithi (Page and Braasch, 1976).

Etheostoma smithi Page and Braasch

Slabrock Darter

Fig. 16.

Etheostoma smithi. Page and Braasch, 1976

Types — Etheostoma smithi was described from Ferguson Creek,
Livingston Co., Kentucky. Holotype INHS 75013. Paratypes deposited at
INHS, KU, UMMZ, USNM, NLU, TU, UAIC, and UT.

New Material Examined — Localities in addition to those listed in Page
and Braasch (1976) include: Tennessee: Trousdale Co., Rocky Creek: KU
20904 (4). Wilson Co., Spring Creek: KU 20905 (5), creek: 20939 (4).
Humphreys Co. INHS 77627 (3). Sumner Co., INHS 77497 (7); MEB
(18). Caldwell Co., INHS 79368 (3). Rutherford Co., MEB (6).

Diagnosis — Fins with reds and blues, face with an iridescent red and
white diagonal bar; infraorbital canal incomplete, pores 3-1; no dark
longitudinal stripes or markings on sides; modally 14 dorsal fin rays.

Description— Etheostoma smithi (Fig. 16) is a small Catonotus, largest
specimen 52 mm SL. Infraorbital canal incomplete, pores 3-1; anal rays
8-10 (usually 9), anal spines 2; dorsal spines 8-10 (9); dorsal rays 13-15
(14); pectoral rays 11-13 (12). Prepectoral area, cheeks, opercles, nape,
and breast naked; lateral scale rows 41-54, 2-13 pored; transverse scale
rows below lateral line 5-9 (6-8), scale rows above 5-9 (6-8). Tips of
dorsal fin spines of breeding males with small sub-conical knobs.

Coloration: Coloration in both non-breeding males, females, and
breeding males almost identical to those of E. obeyense. Differences
include a tendency for the lateral blotches to show more vertical elongation
in E. smithi and the background coloration of E. smithi to be darker than
E. obeyense.

For a more detailed description, analysis of variation, and photographs
of adults see Page and Braasch (1976).

Distribution — Etheostoma smithi occurs in small tributaries of Cum-
berland River from the mouth upstream to Wilson and Trousdale counties
in central Tennessee and in lower Duck River and nearby Tennessee River
(Fig. 17). There are no records for the Red or Harpeth rivers in Tennessee
nor the lower Tennessee River in Kentucky.

Habitat and Eife History— In the Nashville Basin, the frequented
habitat of E. smithi is small creeks with a bottom of bedrock. However, in
the Duck River they occur in gravel bottom pools and in the lower
Cumberland River their habitat is variable, usually pools with either sand,

50 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

gravel, or bedrock substrate. They seem to be most common in creeks
5-10 m wide. Occasionally E. smithi is found in large rivers. It has
recently been collected several times in Cumberland River (Bruce Bauer,
pers. comm.). The large river records presumably represent strays and not
breeding populations.

Page and Burr (1976) detailed the life history of this species,
describing reproduction, growth, food, and population composition in
Ferguson Creek, Livingston Co., Kentucky. Males and females attain
maturity at one year and live up to 2 years. Young and adults feed
primarily on small crustaceans and immature aquatic insects. Spawning
occurs between late April and mid-June. Up to 600 eggs are fastened to the
undersides of flat rocks where they are guarded by a single male.

Etheostoma striatulum Page and Braasch

Striated Darter

Fig. 16.

Etheostoma striatulum. Page and Braasch, 1977

Types— Etheostoma striatulum was described from Wartrace Creek,
Bedford County, Tennessee. Holotype INHS 75035. Paratypes deposited
at INHS, UT, TU, NLU, USNM, KU, UAIC.

New Material Examined— Localities in addition to those listed in Page
and Braasch (1977) include: Tennessee: Bedford Co., Hurricane Creek:
KU 20950 (4); MEB (1). Marshall Co., INHS 86776 (1); MEB (1), Maury
Co., MEB (15).

Diagnosis— Fins with reds and blues, cheek with an iridescent red and
white diagonal bar; infraorbital canal incomplete, pores 3-1; dorsal fin
spines usually 8; pectoral rays usually 11; sides with dark, longitudinal
striations formed by concentrations of melanophores on median and
posterior edge of scales.

Description— Etheostoma striatulum (Fig. 16) is a small Catonotus,
reaching 49 mm SL. Infraorbital canal incomplete, pores 3-1; preoper-
culomandibular canal pores 10. Prepectoral area, nape, cheeks, opercles,
breast, and nape naked. Total lateral scale rows 38-50, 2-18 pored,
transverse scale rows above lateral line 4-6 (5), 6-8 below lateral line.
Anal rays 7-10 (usually 9), spines 2; dorsal spines 8-9 (8), dorsal rays
12-14 (13); pectoral rays 10-12 (11). Tips of dorsal spines of breeding
males with small round knobs.

Coloration: Color very similar to E. virgatum, differing primarily in
lateral pattern. Lateral stripes in E. virgatum are darker and more
continuous than in E. striatulum. Stripes in striatulum appear discon-
tinuous; because of posterior clusters of melanophores on scales, stripes
appear diagonal as well as horizontal. E. virgatum usually have 10 or more
stripes, whereas E. striatulum has 8 or less.

For a complete description, review of variation, and photographs of
adults see Page and Braasch (1977).

Distribution— Etheostoma striatulum is restricted to small tributaries

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 51

of Duck River in Bedford, Marshall and Maury counties, Tennessee (Fig.
17).

Habitat and Life History— The preferred habitat of E. striatulum is
small bedrock streams characteristic of the Nashville Basin. They tend to
congregate near margins of quiet pools where they hide under flat rocks, in
vegetation, or beneath under-cut banks or tree roots. At least one
collection (University of Tennessee) is known from Duck River proper.
Although this probably represents a stray, it indicates that E. striatulum is
not rigidly restricted to the small creeks.

Page (1980) conducted an intensive life history study on this species.
Both sexes reach maturity at one year and live up to 17 months. Adult
females produce up to 108 mature oocytes and are significantly smaller
than males. Spawning occurs between late March and early May. Eggs are
attached to the undersides of flat rocks in the typical Catonotus manner and
are guarded by a single male. Up to 327 round, translucent eggs have been
found in a single nest. As in other barcheeks, the diet of E. striatulum
varies with age, but consists primarily of small crustaceans and immature
aquatic insects.

Etheostoma barbouri Kuehne and Small

Teardrop Darter

Fig. 16.

Etheostoma barbouri. Kuehne and Small, 1971

Types— Etheostoma barbouri was described from Brush Creek, 2.7 mi
N Liberty, Casey County, Kentucky. Holotype USNM 204339. Paratypes
were deposited at USNM and UMMZ.

New Material Examined— Localities in addition to those cited in
Kuehne and Small (1971) include: Kentucky: Adair Co., Petty' s Creek:
KU 11564 (175), 14406 (22); creek 7080 (6). Lincoln Co., Green River:
KU 5809 (7). Tennessee: Sumner Co., Little Trammel Fork: KU 14397
(12).

Diagnosis— Fins with reds and blues, cheek region with red and white
iridescent diagonal bar; suborbital bar dark and well developed. Infraorbi-
tal canal incomplete, pores 3-1; preoperculomandibular canal pores 9.

Description— The following description is based primarily on speci-
mens at KU and INHS. Etheostoma barbouri (Fig. 16) is a small
Catonotus, maximum size 50 mm SL. Infraorbital canal incomplete, pores
3-1; preoperculomandibular canal pores modally 9. Prepectoral area,
cheeks, opercles, nape, and breast unsealed. Total lateral scale rows
40-51, pored 0-16, unpored 29-49; transverse scale rows below lateral line
6-10, scale rows above 2-6 (3-4). Anal spines 2, anal rays 7-11 (9); dorsal
spines 7-10 (9), dorsal rays 11-15 (13); pectoral rays 11-13 (12). Tips of
dorsal spines of breeding males with small sub-conical knobs.

Coloration: Coloration very similar to E. virgatum, differing in the
presence of a dark suborbital bar (or teardrop), lateral longitudinal stripes
less intense and discontinuous, and lateral blotches larger, more intense,
and vertically elongated, sometimes approaching vertical bands. Kuehne

52 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

and Small (1971) present a color photograph and a more detailed
description and analysis of variation.

Distribution— Etheostoma barbouri is restricted to the Green River
system where it occurs in Barren River and tributaries of upper and middle
Green River (Fig. 17).

Habitat and Life History— The preferred habitat of E. barbouri in the
Green River system is small to medium size bedrock creeks. In the Barren
River system, it is found in gravel or sand bottom pools. Flynn and Hoyt
(1979) detailed the life history of E. barbouri in Alexander and Little
Muddy creeks, Edmonson and Butler counties, Kentucky. Both sexes
attain maturity at one year and live up to 2 + years. Males are typically
larger than females. Spawning occurs between March and May. Eggs are
attached to the undersides of slab rocks and guarded by a single male (Page
et al. 1982). In two nests the embryos numbered 42 and 70 and averaged
2. 1 mm in diameter. Females produce 17 to 48 mature oocytes depending
on age. Oocytes averaged 1.6 mm in diameter. Adults and juveniles feed
heaviest during spring and winter months; throughout the year they feed
primarily on small crustaceans and immature aquatic insects. Some
cannibalism was observed and many teardrop darters were parasitized by
Crassiphiala bulboglossa and Contracaecum sp.

SPECIES COMPARISONS IN THE
ETHEOSTOMA VIRGATUM GROUP

The E. virgatum group consists of two moderate sized (obeyense and
virgatum) and three small species (smithi, striatulum, barbouri). Although
differing in color pattern (Fig. 16), there are few differences in meristics
between species.

Color Pattern — Three species (virgatum, striatulum, barbouri) have
dark longitudinal stripes along sides. These stripes are continuous in E.
virgatum and broken into dots or striations in E. striatulum and E.
barbouri (Fig. 16). Etheostoma barbouri may be separated from E.
striatulum by the presence of a well developed dark suborbital bar. In
addition, E. striatulum has posterior markings on each scale giving the
appearance of having oblique rows of striations as well as horizontal ones.

Etheostoma obeyense and E. smithi lack horizontal stripes and are
almost identical to each other in pattern. Etheostoma smithi has a darker
ground color and more elongated lateral blotches than does E. obeyense.

Meristics and Morphometries— Counts and proportions are very simi-
lar in all species. Major differences exist in the number of infraorbital
canal pores and pored lateral-line scales. Infraorbital pores are 4-2 in E.
obeyense and E. virgatum and 3-1 in E. smithi, E. striatulum and E.
barbouri. Etheostoma obeyense and E. virgatum also have modally more
pored lateral-line scales than the others, although differences between all
five barcheeks exist (obeyense 17.5, virgatum 13.0, striatulum 9.6, smithi
7.5 and barbouri 5.2) (Page, 1981). Etheostoma barbouri has modally
fewer preoperculomandibular canal pores (9 vs. 10 in others); E.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 53

striatulum has fewer pectoral fin rays and fewer total lateral-line scales; E.
obeyense and E. smithi have shorter pelvic fins, and E. obeyense and E.
barbouri share a longer preorbital length.

Key to species of the Etheostoma virgatum group

1. Scales on side with medial row of dark melanophores laterally,
forming continuous horizontal stripes (Fig. 16B) . . . . E. virgatum
Melanophores not forming continuous horizontal stripes 2

2. Scales on sides, at least posteriorly, with central dark pigment spot

formed by group of large melanophores 3

Scales with melanopores evenly, or randomly, distributed and thus
not forming discrete pigment spots 4

3. Preoperculomandibular canal pores modally 10, restricted to Duck
River system (Fig. 16C) E. striatulum

Preoperculomandibular canal pores modally 9, restricted to the Green
River system of Kentucky and Tennessee (Fig. 16E) . . E. barbouri

4. Infraorbital canal pores 4 (3 + 1), pored lateral line scales 0-10 (Fig.
16D) E. smithi

Infraorbital canal pores 6 (4 + 2), pored lateral line scales 1 1-30 (Fig.
16 A) E. obeyense

ETHEOSTOMA FLABELLARE SPECIES GROUP

The E. flabellare species group consists of two described and one or
more undescribed species. The described species include E. flabellare and
E. kennicotti. The widespread E. flabellare is outside the scope of this
paper and is only briefly considered here for comparative purposes. This
species is presently being studied by Lawrence McGeehan, Ohio State
University. Beside the nominal form, two subspecies of E. flabellare are
currently recognized, E. f. lineolatum and E. f. brevispina. The earlier
recognized subspecies of E. kennicotti, cumberlandicum was considered
invalid by Page and Smith (1976).

Etheostoma flabellare may be separated from E. kennicotti by its
broadly connected branchiostegal membranes and different color pattern.

Etheostoma kennicotti (Putnam)

Stripetail Darter

Fig. 18.

Catonotus kennicotti. Putnam, 1863

Etheostoma cumberlandicum. Jordan and Swain, 1883

Types — Catonotus kennicotti was described from a rocky brook, Union
County, Illinois. Smith and Page (1975) designated MCZ 24665 as
lectotype.

Diagnosis— No red or blue in fins or on body; no red or white diagonal

54

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

bar on face; bicolored spinous dorsal, dark distally and clear basally;
prepectoral area, cheeks, opercles, and nape unsealed; branchiostegal
membrane slightly to moderately connected; anal fin rays usually 7; dorsal
rays usually 12.

Material Examined-INHS 972, 983, 1495, 1515, 1575, 1592, 2218,
77899, 88502; KU 14401, 15145, 20911, 20942.

Description — The following description is based on original data and
Page and Smith (1976). Etheostoma kennieotti (Fig. 18) is a moderately
large Catonotus. Largest specimens approach 70 mm SL, but most
specimens are much smaller. Infraorbital canal incomplete, pores 4-1;
cheeks, opercles, nape, prepectoral area, and breast naked. Total lateral
scale rows 38-55, with 6-37 pored and 14-40 unpored. Transverse scale
rows above lateral line 4-7 (usually 5), below lateral line 6-10 (8). Anal
rays 6-9 (7); first dorsal with 6-9 (7) spines, fleshy knobs present in both
sexes; dorsal fin rays 10-13 (11-12); pectoral fin rays 11-14 (12-13).

Coloration: Sides yellow to dark brown with 9-1 1 dark blotches; head
darker, particularly dorsally; pre- and postorbital dark stripes present.
First dorsal fin colorless except for dark subdistal band and spines
sometimes with yellow or orange fleshy knobs (color occasionally extends
down onto membrane). Second dorsal, caudal, and pectoral fins with dark
bars; anal fin usually colorless. Breeding males with heads darker; flanks
flesh-colored to orange with vertically elongated lateral blotches; dorsal
spines with conspicuous orange knobs. For a more detailed description

.

- * * I I I c

Figure 18. Top, Etheostoma kennieotti, male 53.8 mm SL (KU 14401, Hatfield Creek.
Cambell Co., TN). Bottom, Etheostoma flabellare , male 53.3 mm SL (KU 11670, Goose
Creek. Trousdale Co., TN).

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 55

and analysis of variation see Page (1975a) and Page and Smith (1976).

Distribution — Etheostoma kennicotti occurs in the Tennessee, upper
Cumberland, Green, and lower Ohio rivers. Although common in the
Shawnee Hills and upper Cumberland River, it is sporadic or absent
throughout much of the range (Page and Smith, 1976, fig. 2). Wherever it
occurs, however, it is often abundant.

Habitat and Life History— Page (1975b), Page and Smith (1976), and
Page and Schemske (1978) give the preferred habitat of E. kennicotti as
slab-rock pools. Throughout most of the range we have found them in
gravel or sand-bottom pools or gentle gravel riffles of small spring-fed
creeks to mid-sized rivers. Often E. kennicotti hides among emergent
vegetation at margins of pools or riffles.

Complete life history information of E. kennicotti is provided by Page
(1975b). Males and females mature in one year and live up to 2 + years.
Spawning occurs in April and May when eggs are attached to the
undersides of flat rocks. Eggs in nests range from 50 to 400 and are
guarded by highly territorial males. Females are generally smaller than
males and are known to produce 24 to 130 mature oocytes. Adults and
juveniles feed primarily on small crustaceans and immature aquatic
insects.

Etheostoma flabellare (Rafinesque)
Fantail Darter
Fig. 18.
Etheostoma flabellaris. Rafinesque, 1819
Etheostoma fontinalis. Rafinesque, 1820
Etheostoma linsleyi. Storer, 1851
Catonotus lineolatus. Agassiz, 1854
Oligocephalus humeralis. Girard, 1859
Catonotus fasciatus. Girard, 1859
Richia brevispina. Coker, 1926

Types — Etheostoma flabellaris was described from specimens col-
lected in the Ohio River. No types are available as Rafinesque traditionally
did not preserve any type material. Based on the drawing illustrated in his
notebook, however, there is no problem with the identity of the animal
drawn (Collette and Knapp, 1966).

Diagnoses — No red or blue coloration; no scales on cheeks or
opercles; branchiostegal membranes broadly connected; transverse scale
rows above lateral line usually 7 and below lateral line usually 9.

Description — The following brief description of E. flabellare (Fig. 18)
is based on Trautman (1957), Cross (1967), and Pflieger (1975) plus
specimens from the upper Tennessee and lower Cumberland rivers. This
highly variable species probably represents a complex of forms.

Infraorbital canal incomplete, pores 4-2. Nape, cheeks, and opercles
unsealed; total lateral scale rows 40-57, 8-37 pored. Transverse scale rows
above lateral line 7-9, below lateral line 8-10.

56 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Dorsal spines 7-10, dorsal rays 13-14; anal rays 7-9; pectoral rays
12-13.

Coloration variable. Ground color yellow to black with 10-15 dark
crossbars extending to below lateral line. Sides often with longitudinal
stripes. Humeral spot dark. Head darker than body, with pre- and
postorbital bars. Spinous dorsal with distal and basal orange or brown
bands. Spines with fleshy, orange knobs. Soft dorsal, caudal, and pectoral
with dark bands; anal and pelvic fins colorless. Breeding males similar,
but head darker and with flanks flesh or orange. Orange dorsal fin knobs
enlarged.

Distribution— Etheostomaflabellare occurs from the east coast west to
the eastern Great Plains and south in the Mississippi valley to Mississippi.
In the Ohio River system, E. flabellare is widespread and common. In the
upper Cumberland and the Tennessee River drainages in northeast
Alabama and east Tennessee it is common in the Blue Ridge Province
along North Carolina border, but is virtually absent in the Ridge and
Valley Province (Page, 1983).

Habitat and Life History— Etheostoma flabellare occupies quiet to
moderate riffles of gravel or rocks. Stream size is of little importance as
they can be found in small brooks and large rivers. Considerable work has
been done on the growth, reproduction, population dynamics, and diet of
this species. This information is summarized by Page (1983). Basically,
the biology of E. flabellare is similar to other Catonotus. Both sexes
mature in one year and live up to 4 years. Males live longer than females
and are generally larger. Spawning occurs between April and July, but
geographic variation has been observed. Eggs are adhesive and are
attached to the underside of a flat rock. Spawning occurs beneath the flat
rock in cavity excavated by the male. Females produce up to 562 eggs
which average 2.2-2.7 mm. As do other darters, E. flabellare feeds
primarily on small crustaceans and immature aquatic insects.

RELATIONSHIPS

Previous discussions of relationships in Catonotus have varied from
comments of sister species relationships based on morphology to an
electrophoretic examination of six species. Wolfe et al. (1979) examined
E. squamiceps, E. flabellare, E. kennicotti, and three barcheek species for
three protein systems. They interpreted their results from an unrooted
Wagner Network analysis of patrastic distances as supporting a close
similarity of the E. virgatum group, which they equated with this group
being natural. Further, they related E. kennicotti to E. squamiceps and
these two species to the E. virgatum group rather than either to E.
flabellare.

Results from morphological studies of the evolution of Catonotus are
varied. Collette (1966) viewed the subgenus as two major groups, one
comprised of E. squamiceps, E. kennicotti, and E. flabellare, all sharing
well developed fleshy knobs on dorsal spines. The other group being the E.
virgatum species group which never or only occasionally develop small

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 57

incipient fleshy knobs (Collette, 1966). Page (1975b) evaluated these
hypothesized relationships and disagreed with Collette after examining
other morphological and breeding characteristics. Page (1975b) consid-
ered the E. virgatum group to be most closely related to a group including
E. kennicotti and E. flabellare (flabellare species group), and he consid-
ered E. squamiceps to be the most primitive lineage of the subgenus.
Kuehne and Small (1971) viewed their new species, E. barbouri, as a close
relative of E. virgatum and E. obeyense. Page and Braasch (1976)
considered E. smithi and E. obeyense to be closest relatives, closely
related to the other barcheeks, E. virgatum and E. barbouri. They further
noted a close relationship of the clade E. kennicotti-E. flabellare to all
barcheeks. Prior to the description of E. striatulum, it had long been
considered a close relative off. virgatum, but Page and Braasch (1977)
concluded that there is little evidence to consider E. striatulum more
closely related to E. virgatum than to any other barcheek.

Etheostoma squamiceps had long been considered a single species until
the descriptions of E. neopterum (Howell and Dingerkus, 1978) and E.
olivaceum (Braasch and Page, 1979). Howell and Dingerkus (1978)
placed E. neopterum in the E. squamiceps group, but considered it more
primitive than E. squamiceps. Braasch and Page (1979) also considered
their new species E. olivaceum to be a member of the E. squamiceps group
and suggested that it was more closely related to the polytypic E.
squamiceps than to E. neopterum.

Our analysis of morphological and behavioral characters presented in
the preceding sections of this review and summarized in Appendix 1 and 2
supports a phylogenetic history of Catonotus illustrated in Figure 19. We
consider the subgenus to be monophyletic with three major subclades — (1)
the squamiceps species group, (2) the flabellare species group, and (3) the
virgatum species group (Fig. 19). As discussed above, this arrangement of
species groups is consistent with previous studies, with the exception of
Collette (1966) and Wolfe et al. (1979). Species relationships within the
virgatum group and squamiceps group, however, vary somewhat from
previous hypotheses. We believe these inconsistencies stem from grouping
species with primitive characters as compared to consistent grouping using
derived characters.

Monophyly of Catonotus. — In this analysis the monophyly (i.e. natu-
ralness) of Catonotus is supported by 8 characters (Fig. 19). These include
(1) an interrupted infraorbital canal, (2) flattened genital papillae in
females, (3) spawning occurring on the undersides of stones, (4) presence
of fleshy knobs on the tips of dorsal fin spines of males, (5) an incomplete
lateral line, (6) a low first dorsal fin, (7) swollen heads in breeding males,
and (8) a habitat preference for slab rock pools (See Appendix 1 and 2 for
implied transformation series for these and other characters used in this
analysis). Character argumentation is presented below.

Most species of Catonotus have an interrupted infraorbital canal.
Having an incomplete infraorbital canal series is considered derived
(Page, 1977; 1981). Etheostoma olivaceum and some populations of E.

58

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

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TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 59

neopterum have a complete canal, a condition we consider a reversal.

Females of all Catonotus have flattened genital papillae, a morphology
unique to this subgenus (Page, 1983). Females of E. sellare of the
subgenus Etheostoma, species of the subgenus Boleosoma, and species of
the E. maculatum species group of the subgenus Nothonotus also have
flattened papillae, but their structures are different and are not considered
homologous with the type found in Catonotus (Page, 1983).

All species of this subgenus spawn under slab stones (Page et al.,
1982). This egg clustering behavior is considered derived and unique to
this group of darters. The primitive spawning behavior in darters involves
burying eggs in the substrate. Some other darters are known to attach their
eggs to the undersides of objects, but certain aspects of their behaviors are
quite different from Catonotus (i.e. multiple layers of eggs) (Page et al.,
1982).

Breeding males of all species of Catonotus develop round to elongate
fleshy knobs at the tips of their dorsal fin spines (Mayden, 1985). These
structures have been reported on several occasions for the flabellare and
squamiceps species groups, but were mentioned only once for the
virgatum group by Collette (1965). Knobs were reported as missing from
the virgatum group by Page and Braasch (1976, 1977), Page (1981, 1983),
and Page and Burr (1976). Although somewhat smaller than in the other
two clades, fleshy knobs are present in the virgatum group and are
undoubtedly homologous with those of the other species groups, support-
ing the rnonophyly of Catonotus (Mayden, 1985). Analogous fleshy
structures, similar to the knobs of species of Catonotus, are also found on
spinous dorsal fins of breeding male Boleosoma.

All species of Catonotus have an incomplete lateral line. We consider
the reduced lateral line derived since having a complete lateral line is
primitive in darters (Page, 1977). Among the more highly derived
barcheeks the lateral line becomes increasingly more reduced.

In Catonotus the shape of the first dorsal fin is distinctive. Adults of
species of the squamiceps and flabellare groups have a low first dorsal fin
(Figs. 1,8, 11, 13, 15, 18). Young of these species, however, have higher
dorsals relative to the fin base (Table 2), more like adults of other darters.
Adult barcheeks, however, have a slightly more elevated first dorsal fin,
more like other darters (Fig. 16).

Breeding male Catonotus develop greatly swollen heads during court-
ship, spawning, and egg guarding. We consider this a derived condition in
Etheostoma since such extreme development is presumably restricted to
this subgenus. The development of a swollen head is believed to have a
secretory function while guarding eggs (Cross, 1967; Page, 1983).

The habitat preferred by most species of Catonotus is distinctive
among darters. All species, with the exception of E. flabellare, E.
kennicotti, and E. neopterum prefer slabrock pools (Table 15, also see
previous discussion of habitat and competition in Catonotus, and Page and
Schemske, 1978). Although varied, not many other darters can be
characterized as preferring this particular type of habitat. We therefore

60 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

suggest that this behavioral characteristic is derived for Catonotus and has
been modified in E. flabellare, E. kennicotti, and E. neopterum (Table
15).

Although the above derived characters of Catonotus support the
monophyly of the group, the independence of some characters remains to
be tested. Five characters 1) flattened papilla, 2) spawning on undersides
of stones, 3) fleshy knobs on spines, 4) low first dorsal fin, and 5) swollen
heads of males may be correlated and realistically represent a single
character complex relating to mode of reproduction. We say this because
in other darters and some other fishes when the mode of reproduction
involves attaching eggs to the underside of an object some of the above
listed characters are sometimes present. Examples include the Etheostoma
maculatum species group in which some species are known to attach their
eggs to the undersides of rocks in fast water and also have a flattened
papilla (Page et al., 1982). In the subgenus Boleosoma of Etheostoma
females have flat, bifurcate papillae and males have swollen knobs on
dorsal spines. Species of this group are also known to deposit eggs on the
undersides of objects (Page, 1983). Further, in the cyprinid genus
Pimephales, some species attach their eggs to the undersides of debris or
stones and males have swollen heads. Thus, the above list of characters
may be correlated and represent a single character. However, since in no
species outside Catonotus can we find complete duplication of all the
morphological characters with a similar (but not homologous) reproduc-
tive mode (e.g., species of the E. maculatum group lack swollen knobs and
heads and reduced dorsal fins; Boleosoma lack reduced fins and swollen
heads) we hesitate to combine all of these characters into one single
character complex.

Monophyly of the flabellare-virgatum group. — The sister group rela-
tionship of these two clades is supported by 10 shared derived characters
(Fig. 19). These include (1) further reduction of the pored lateral line to
less than 40%, (2) a naked nape, (3) naked prepectoral area, (4) naked
opercles, (5) naked cheeks, (6) a weakly developed suborbital bar, (7) a
further reduction of the infraorbital canal, (8) unique nuptial structures on

Table 15. Habitat preference of Catonotus. 1 = preferred habitat, 2 = less common habitat,
3 = least preferred habitat, x = habitat occupied in restricted portion of range or seasonally.

Sand-Gravel

Slabrock

Gravel Riffle

Slab or Rock

Species

Pool

Pool

Riffle

E. flabellare
E. kennicotti

X

1

X

2

1
3

1

E. squamiceps

1

X

E. neopterum

1

X

E. crossopterum

2

E. nigripinne
E. olivaceum

2
2

2

E. virgatum
E. obevense

2
2

2

E. smithi

X

E. harbour i

2

E. striatulum

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 61

the ventrolateral body scales of breeding males, (9) a change in scale rows
above the lateral line, and ( 10) a change in the number of scale rows below
the lateral line. Changes in the number of scale rows above and below the
lateral line are discussed at the end of this section.

All species of the flabellare and virgatum groups have an increased
reduction in the number of pored lateral line scales to 40% or less from the
60-75% pores scales of the squamiceps group. As discussed above, having
a complete lateral line is primitive for darters. The higher percentage of
pored scales in the squamiceps group is therefore considered primitive
within Catonotus.

All species of this group lack scales on the nape, prepectoral areas,
opercles and cheeks. These conditions are considered derived in
Catonotus, since most darters and all other Catonotus have scales on these
regions. Since E. olivaceum can be related to the squamiceps group with
several characters (Fig. 19) and not with the flabellare-virgatum group we
believe the loss of cheek and opercle scales in E. olivaceum was
independent. Further, the occurrence of reduced scales in only two of
these four regions in E. olivaceum argues for independence of the four
regions to scale loss. If these areas were linked in any way such that a loss
of scales in one region resulted in a loss of scales in the other regions, then
one would expect the nape of E. olivaceum to be naked, when in fact it is
always scaled. In the flabellare-virgatum clade then, the loss of scales in
all four regions represents four different independent characters and not a
single character— reduced head squamation.

A strongly developed "teardrop" or suborbital bar is common among
darters and in the squamiceps group. The loss or reduced development of
the bar in the flabellare-virgatum group is then interpreted here as a shared
derived character of this group. Etheostoma barbouri is unique among
species of this clade in having a well developed bar (Kuehne and Small,
1977).

Species of the flabellare-virgatum clade share the loss of the anterior-
most pore of the posterior segment of the infraorbital canal. As discussed
above, having a complete infraorbital canal is primitive in darters and the
interrupted canal of Catonotus is derived. Species of the squamiceps group
usually have four pores anteriorly and 3-4 (usually 3) pores posteriorly.
Members of the flabellare-virgatum clade have further reduced their head
canal system with a maximum of 2 pores in the posterior canal segment.
We consider this morphology derived since this is a further step in
reduction from a primitive complete system and the squamiceps group has
a more complete system of 3 pores posteriorly.

The last derived character shared by the flabellare-virgatum group is
the occurrence of nuptial ridges on the ventrolateral body scales of
breeding males. In males with maximum breeding coloration, scales below
the lateral line from 6-7 scale rows posterior to the pectoral fin insertion to
the caudal fin have centrally located swollen fleshy ridges radiating more-
or-less from the center of each scale (Mayden, 1985). Species in the
squamiceps group and all other darters lack similar structures.

62 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Monophyly of the Etheostoma virgatum group. — The naturalness of the
virgatum species group is supported by their common possession of at least
six derived characters (Fig. 19). These include (1) the presence of an
iridescent red and white cheek bar, (2) reduced knobs on the tips of dorsal
spines, (3) a further reduction in the number of pored lateral-line scales,
(4) a distinctive first dorsal fin pigmentation pattern, (5) a preopercle with
a crenulate edge, and (6) 9 anal rays.

All species of the barcheek group have a bright red and white colored
bar on both sides of their head covering parts of their cheeks and
preopercles. We consider this feature to be derived for this group since it is
unique among darters.

Breeding males of all species of Catonotus have knobs on the spines of
the first dorsal fin (Mayden, 1985). In the squamiceps and flabellare
species groups the knobs are large, but in the virgatum group they are
small. We believe the most parsimonious interpretation of the distribution
of this character is to consider the reduced knob morphology of barcheeks
a derived state. Further, if comparisons are made to other taxa with knobs
(Boleosoma), the knobs present in the flabellare and squamiceps groups
are more like those of these other taxa (in size).

All species of the virgatum group share a further reduction of pored
lateral-line scales to less than 40% of the total. This is a greater reduction
than present in the squamiceps and flabellare groups. We interpret this
further reduction as a derived condition for this group since having a
complete lateral line is primitive in darters (Page, 1977).

The dorsal fin pigmentation pattern found in this group is unique
among Catonotus, as well as other Etheostoma. The fin of breeding males
is predominantly red with a large intense black basal blotch extending
from the first to the sixth spine. Species of the squamiceps and flabellare
groups have an orange band in a darkly pigmented first dorsal fin. We
consider the virgatum color pattern derived because it is not known to
occur in any other darters and the orange band present in the squamiceps
and flabellare groups is comonly shared outside of the subgenus.

The bright coloration of other unpaired, as well as paired, fins in
species of this group may also be interpreted as synapomorphic characters.
The second dorsal and anal fins of breeding males are predominantly
bright red; the caudal fin is red with a blue ventral margin; pectoral fins
are reddish medially and blue ventrally; pelvic fins are blue-black. Fins of
other Catonotus are not brightly colored.

All members of the virgatum group share a crenulate preopercle, while
other Catonotus have a smooth preopercle. Since most darters have a
smooth opercle, we consider the crenulate preopercle of the virgatum
group to be the derived condition. E. okaloosae of the subgenus Belophlox
and some species in the subgenus Boleichthys have a serrated preopercle
which is quite different from the condition found in the virgatum group.

Within Catonotus the modal number of anal fin rays varies between
species from 7 to 9. The particular distribution of anal fin rays, however,
suggests that the ancestral condition in this group was a modal number of 8

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 63

rays (Appendix 2). Species of the virgatwn group uniquely share one of the
two derived conditions present in the subgenus, 9 anal rays. The other
derived condition, having modally 7 anal rays, has occurred independently
in three more primitive members of the subgenus and will be discussed
below.

Within the virgatwn group we consider E. obeyense to be the sister
taxon to all other presently recognized barcheeks (Fig. 19). This conclu-
sion is based on the observation that other barcheeks share four derived
characters, (1) fewer than 35% of the lateral line scales with pores, (2)
longitudinal stripes along their flanks, (3) changes in the number of scale
rows above, and (4) below the lateral line. We consider these conditions
derived since having a complete lateral line is primitive and the longitudi-
nal stripes are nearly unique to this group of Catonotus. Longitudinal
striations also occur in some parts of the range of E. flabellare and have
apparently been lost in E. barbouri. Changes in the numbers of scale rows
above and below the lateral line are discussed below.

Etheostoma virgatwn is hypothesized to be the sister group of the three
remaining species, E. striatulum, E. barbouri and E. smithi (Fig. 19). The
latter three species share three derived characters not shared with E.
virgatwn, (1) reduction of the anterior segment of the infraorbital canal to
3 pores, (2) reduction of the posterior infraorbital canal to one pore, and
(3) reduction to approximately 25 % or less of the lateral body scales with
pores. The reduction in the number of infraorbital pores is considered
derived since having a complete infraorbital canal is primitive in darters
and having 4 anterior and 2 posterior pores is primitively shared with other
Catonotus as well as other Etheostoma. The pored lateral line reduction is
considered derived because it represents a further modification from the
primitive complete condition. The presence of only three pores in the
anterior infraorbital canal segment is unique to this group of Catonotus,
but the reduction to one pore in the posterior segment is also known to
occur in E. kennicotti (Fig. 19).

Among the remaining three barcheek species, E barbouri and E. smithi
are hypothesized to be sister species, most closely related to E. striatulum.
Etheostoma barbouri and E. smithi share a more derived condition of the
lateral line in having a reduction to fewer than 20% of the scales being
pored.

Monophyly of the E. flabellare group.— At least four shared derived
characters corroborate the monophyly of this species group (Fig. 19)
These include (1) a modal number of 7-8 dorsal fin spines, (2) very large
knobs on dorsal spines, (3) knobs on dorsal spines being golden, and (4)
the presence of these knobs on both males and females.

The modal number of dorsal spines varies in Catonotus from 7 to 9.
The distribution of the modal number of spines indicates that primitively
Catonotus had 9 dorsal spines (Appendix 2). Changes from this condition
to 8 and 7 have occurred three times, once to 8 and 7 in the flabellare
group and twice to 8 in E. nigripinne and E. striatulum (Appendix 2). We

64 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

therefore consider the reduction to 8 and 7 dorsal spines in E. flabellare
and E. kennicotti to represent evidence of their monophyletic origin. Since
E. flabellare is a wide-ranging species with variation in the number of
dorsal spines this species may actually represent a polytypic member of the
subgenus with some geographic forms (those with a mode of 7 dorsal
spines) more closely related to E. kennicotti (which modally has the
further derived state of 7 spines) than to other E. flabellare with modally 8
spines.

Further corroborating the monophyly of this species pair are the well-
developed knobs on dorsal spines, their presence in both males and
females, and their bright golden color. In members of the squamiceps and
virgatum groups knobs are opaque to light yellow. In both of these groups
females do not develop these structures and the size of knobs in breeding
males is reduced compared to males of E. flabellare or E. kennicotti.
Thus, because of the shared nature of these characters between the non-
sister groups {squamiceps and virgatum groups) we interpret the condi-
tions shared in the flabellare group derived.

Monophyly of the Etheostoma squamiceps group. — The monophyly of this
species group is supported by their sharing at least six advanced character-
istics (Fig. 19). These include (1) three distinct dark spots at the base of
the caudal fin, (2) rays of the second dorsal fin of breeding males
extending well beyond the fin margin, (3) having clear windows in the
heavily pigmented membranes of the first dorsal fins, (4) having a wide
subdistal white band in the second dorsal fin, (5) spawning adults only
briefly inverting to spawning surface, and (6) females spawning more than
one egg at a time.

Having rays of the second dorsal fin extending beyond the fin margin is
apparently unique to this species group of darters. In other Catonotus, as
well as other darters, the fin edge may be rough with high points
corresponding to tips of fin rays and lower areas in between represented by
a slightly lower membrane, but never do the rays extend out beyond the
membrane as in members of the squamiceps species group (Fig. 3).

The first dorsal fin pigment pattern found in males of this group (Fig.
2) is, again, presumably unique. Other Catonotus species lack any
development of clear windows in the fin membranes and we are unaware
of any similar development of this pattern in other darters.

Breeding behavior may also serve to support the monophyly of this
species group. Page (1975b) compared the spawning behaviors of the
different species groups of Catonotus. He examined variation in the
inversion behavior of males and females to the spawning surface and the
number of eggs spawned at one time. Where known, males of the
flabellare-virgatum group lay 1 egg at a time on the breeding surface and
remain inverted for several hours (Page 1975a, b; 1983). Spawning
behaviors are known for 3 of the 5 species of the squamiceps group, E.
squamiceps, E. crossopterum (Page, 1974, 1975b) and E. olivaceum
(Page, 1980). No data are available for E. neopterum and E. nigripinne,
although we would predict, as did Page and Mayden (1979), their

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 65

behaviors to be very similar. In the squamiceps group the male and female
only briefly invert to deposit more than one egg at a time (Page, 1974).
Therefore, there are presently two modes of spawning in Catonotus,
representing two characters, duration of female inversion and the number
of eggs spawned with each inversion. Either a female is inverted for an
extended period of time and spawns a single egg at a time, or they invert
only briefly and spawn up to three eggs at a time.

Since depositing eggs on the undersides of stones is a rare behavior in
darters, it is difficult to determine the polarity of these behaviors.
However, if comparisons are made with other Etheostoma with this type of
reproduction then behaviors found in the flabellare-virgatum group can be
considered primitive. Species with similar reproductive activities include
the subgenus Boleosoma (Page et al., 1981) and some members of the
subgenus Nothonotus (Page et al., 1982). At present spawning behavior is
known only for species of Boleosoma, and in this group females remain
inverted beneath the spawning surface for several hours and deposit eggs
one at a time (for summary see Page, 1983), like the flabellare-virgatum
group.

Within the squamiceps clade E. squamiceps and E. neopterum are
considered sister species (Fig. 19). These two species together form the
sister group to the remaining species E. olivaceum, E. crossopterum, and
E. nigripinne (Fig. 19). The sister group relationship between E. squam-
iceps and E. neopterum is evidenced by their sharing the unique mor-
phology of having the tips of their second dorsal fin rays being expanded
into knobs (Figs. 1, 3, 13) and not pointed or flat as in other Catonotus
(Fig. 3) as well as other darters. We consider this morphology derived
since it is unique among darters.

The monophyly of E. olivaceum, E. crossopterum, and E. nigripinne is
supported by their sharing an advanced color pattern of the second dorsal
fin. Breeding males of all three species have black tips on rays of the
second dorsal fin. We consider this color pattern derived since second
dorsal fin ray tips of other Catonotus, including E. squamiceps and E.
neopterum, are pale during all seasons of the year. Males of E. nigripinne
differ from the other two species in that the black tips of rays, dorsal to a
broad white subdistal band, are obscured in maximally colored spawning
males by a complete blackening of the fin (Figs. 3, 11). However, males of
E. nigripinne prior to spawning have the black tips and a white subdistal
band like E. crossopterum and E. olivaceum.

Within this group of three species relationships are close, but we
consider E. nigripinne and E. crossopterum to be more closely related to
each other than either is to E. olivaceum (Fig. 19). This relationship is
supported by their sharing modally 1 1 pectoral rays and white horizontal
to semi-horizontal dashes in the second dorsal fin. The primitive modal
number of pectoral rays in the subgenus Catonotus and in the squamiceps
species group is 12. Changes from this condition have occurred three times
in the evolution of Catonotus, once in E. striatulum and twice in the
squamiceps group (once in E. squamiceps and once in the common

66 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

ancestor of E. nigripinne and E. crossopterum). Etheostoma olivaceum
and E. neopterum have retained the primitive condition of having modally
12 pectoral fin rays. One may wish to view the distribution of this
character in the squamiceps group as an early change from 12 to 11 in the
ancestor of the squamiceps group and later independent reversals to the
primitive condition in E. neopterum and E. olivaceum. This is congruent
with our hypothesized phylogeny, but is not the most parsimonious
explanation. It requires additional steps to explain the evolution of this
character. An equally parsimonious scenario as the one we have chosen to
adopt here would be the evolution of 1 1 rays in a common ancestor of the
squamiceps group and a reversal to 12 rays in a common ancestor of E.
neopterum and E. olivaceum. This hypothesis, however, is in conflict with
other characters and is not adopted here.

When present, depigmented dashes in the second dorsal fin in
Catonotus vary from being U-shaped to horizontal dashes. Etheostoma
squamiceps, E. olivaceum, E. flabellare, and E. kennicotti all have several
U-shaped pale dashes crossing rays of the second dorsal fin (Figs. 1,3, 15,
18). In E. neopterum the pale areas are generally large dots (Figs. 3, 13)
and in the barcheeks pale areas are absent. In breeding males of E.
nigripinne and most males of E. crossopterum the pale areas are in the
shape of horizontal dashes. This character appears to be fixed in E.
nigripinne, but is more variable in E. crossopterum as males with a semi-
horizontal pattern exist. Aside from having the fixed horizontal pattern, E.
nigripinne differs from E. crossopterum (and other species) in having only
a few rows of very small horizontal pale lines in the second dorsal fin and
having a completely black fin in males with maximum breeding coloration.

Additional characters.— Because of the inherent variability of some
meristic data between outgroups and within species it is difficult to
determine some polarities a priori in Catonotus. However, once less
variable characters are logically polarized, data from these meristic
characters may then be examined for their probable evolution in
Catonotus, trying in each case to minimize the number of steps necessary
to explain the evolution of the characters in the subgenus. Characters
included here are numbers of dorsal rays, dorsal spines, anal rays, and
transverse scales above and below the lateral line (Fig. 19, Appendix 1 and
2).

Interpretation of the number of anal fin rays was briefly discussed
above under the monophyly of the virgatum and flabellare groups. Eight
anal rays appear to be the condition most likely present in the common
ancestor of Catonotus. This condition is present in E. kennicotti and all
species of the squamiceps species group except E. squamiceps and E.
crossopterum. The occurrence of seven rays in these latter two species is
considered convergent because of conflict with other more conservative
characteristics suggestive of alternate relationships (Fig. 19). As discussed
above, the occurrence of 9 anal rays in all species of the virgatum group is
interpreted as an independent development from the ancestral mode of 8.

The evolution of dorsal spines is discussed briefly under the mono-

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 67

phyly of \h& flabellare group. All Catonotus have 9 dorsal spines except E.
nigripinne, E. striatulum, and most E. flabellare, which typically have 8
and E. kennicotti which modally has 7. With reference to relationships in
Catonotus (Fig. 19) we therefore consider a mode of 9 dorsal spines to be
primitive in Catonotus and 8 spines to be derived independently in E.
nigripinne, E. striatulum, and the ancestor of the flabe Hare group. In the
flabellare group further reduction has occurred in some E. flabellare and
E. kennicotti, with 7 spines.

Interpretation of the evolution of dorsal fin rays is somewhat less
confusing. Modal numbers of dorsal rays in Catonotus varies from 1 1 to
14. The most parsimonious explanation of the evolution of this character
would be that primitively in this subgenus the modal number of dorsal rays
was 13, since primitive members of all clades have this number. Changes
have occurred twice in the squamiceps group, once in E. neopterum with
1 1 rays and once in E. olivaceum with 12 rays. Among the remaining
Catonotus, E. kennicotti shifted to a mode of 12 and E. smithi typically has
14 rays.

Transverse scale rows above and below the lateral line are more
difficult to interpret. With reference to the phylogeny (Fig. 19), however,
one may hypothesize that the ancestral Catonotus species had 6 scale rows
above the lateral line. A change occurred only once in the squamiceps
group, in E. olivaceum which modally has 7 scale rows. A change from 6
to 5 scale rows in the ancestral species of the flabellare-virgatum group is
congruent with the phylogeny presented above. This was modified with a
change to 7 rows in E. flabellare and a reversal to 6 rows in the ancestral
species of the virgatum-striatulum-barbouri-smithi clade. Within this latter
group, changes to 5 and 4 scale rows occurred in striatulum and barbouri,
respectively. This interpretation of the evolution of the modal number of
scale rows above the lateral line is one of many, but is most congruent with
the evolution of other characters.

The evolution of scale rows below the lateral line in Catonotus is
equally perplexing. The modal number of rows varies between 7 and 9.
Nine rows appears to represent the primitive condition in Catonotus and is
present in all the squamiceps group species (except E. nigripinne) and E.
flabellare. Etheostoma kennicotti and E. obeyense either independently
evolved 8 scale rows from the primitive 9 or the ancestor of the flabellare-
virgatum group evolved 8 and E. flabellare reversed back to modally
having 9. The ancestral species to the virgatum-striatulum-barbouri-smithi
group evolved 7 scale rows from the ancestor of the virgatum group,
which presumably had 8 rows. Etheostoma striatulum is at variance with
its closest relatives in having reversed its number of scale rows below the
lateral line to 8.

These are but just a few of the many possible evolutionary interpreta-
tions of meristic variation in Catonotus. We have chosen to prefer the
above scenario over alternative scenarios because of minimal conflict with
other more stable characteristics.

68 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

ZOOGEOGRAPHY

Although all 12 described species of Catonotus occur in the same
general area, regions now drained by the Cumberland, Tennessee, and
lower Ohio river drainages, we know of no case in which more than four
species can be found at a particular locale. As discussed briefly under each
species group and by many biogeographers, the distribution of a species is
controlled by geological or ecological barriers and dispersal routes.
Within the range of a species, its presence or absence in a particular stream
is dependent on the availability of suitable habitat, and given that suitable
habitat exists, competition between species may make suitable habitat
unavailable. Below we discuss the biogeography of the subgenus
Catonotus with respect to possible origins of the various species and
attempt to correlate distributional patterns with available geological and
drainage histories of the region. In addition, the maintenance of these
species distributions are discussed with reference to habitat availability and
interspecific competition.

The complex distributional patterns of species in Catonotus together
with the embryonic state of our knowledge of the subgeneric relationships
of darters and our limited understanding of the early geological history of
the region makes it difficult to determine the antiquity of this group and its
geographic origin. The large degree of sympatry between and within
groups and the high diversity of Catonotus in the Cumberland River
drainage implies that the subgenus may be quite old (probably mid-
Tertiary) and had its origin in the Cumberland drainage.

Etheostoma squamiceps group. — The earliest division of this group was
the separation of the squamiceps-neopterum ancestral species from the
ancestor of E. crossopterum, E. olivaceum and E. nigripinne (Fig. 19).
Presumably, the ancestor of the squamiceps-neopterum pair gained access
to the Ohio River from the Cumberland drainage and both differentiated
from their ancestral form. The exact means by which these two species
pairs became isolated is problematical. One scenario is that access to the
Ohio drainage was gained at some date when the Ohio and Cumberland
rivers formed a single system and the event(s) correlated with the initial
speciation event was the separation of these two river drainages (Fig.
20A). Thus, there would have been a widespread "squamiceps " ancestral
species in the Ohio and Cumberland river drainages which gained access
to both of these drainages prior to our existing preglacial drainage
patterns, over a landscape unknown to us today. The vicariance event
leading to the separation of the two ancestral species, one in the Ohio and
one in the Cumberland, involved drainage alterations which culminated in
our existing pre-Pleistocene patterns of the Ohio and Cumberland rivers
where both flowed independently into the Coastal Plain (Fig. 20A).

An alternative scenario is that the ancestral "squamiceps" species
gained access to the Ohio River drainage through stream capture between
the two drainages. The most likely area involved would be capture
between a tributary of the Cumberland River by a tributary of the Barren

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

69

Figure 20. Hypothesized biogeographic history of the squamiceps species group. (A)
Preglacial drainage patterns and distributions of ancestral species of the squamiceps-
neopterum pair (horizontal) and olivaceum-crossopterum-nigripinne group (vertical).
Drainage patterns from Thornbury ( 1965). Pflieger ( 1970), and Starnes and Etnier (in press).
(B) Late Tertiary-early Pleistocene drainage patterns with E. squamiceps in the Ohio
drainage, E. neopterum (stippled) in the lower Cumberland-Duck drainage, E. olivaceum on
eastern Highland Rim and ancestor off. crossopterum and E. nigripinne in the Cumberland
and upper Duck drainages. (C) Late Pleistocene capture of middle and upper Tennessee
system by Duck system and movement of E. neopterum and E. nigripinne (form) into
Tennessee drainage, with differentiation of the latter.

70 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

River. Tributaries of these two rivers interdigitate in northern Tennessee
and the divide is easily breached. Having gained entry into the Green
River, it could have easily spread throughout the upper Ohio drainage
(which was small at the time) (Fig. 20A) and attained a range now
occupied by E. squamiceps (Fig. 20A).

Separation of the Ohio River form into E. squamiceps and E.
neopterum probably occurred when a population became isolated in the
lower Cumberland River drainage, which at that time basically paralleled
the Ohio River (Fig. 20A). The most likely area of connection is a break in
the divide near Karnak, Pulaski County, Illinois, which today is occupied
by a drainage canal. Once in the lower Cumberland drainage it diverged
from E. squamiceps and became distributed in sandy bottom tributaries of
the upper Mississippi Embayment from the Obion River in western
Kentucky to the mouth of the Duck River. It would have been absent in the
middle Tennessee since at this time the middle and upper Tennessee River
were not connected to the present lower Tennessee, but had a separate
entry into the Coastal Plain and the Gulf of Mexico through eastern
Tennessee or northern Mississippi (Starnes and Etnier, in press) until mid-
to late Tertiary (Fig. 20B). Thus, no Catonotus was in this portion of the
present-day Tennessee River system. This is further supported by the
absence of any representative of the subgenus in gulf drainages. Move-
ment of E. neopterum into the middle Tennessee could not have occurred
until late Tertiary when a temporary drainage advantage in the middle and
upper Tennessee captured the Duck River (Starnes and Etnier, in press).

The Cumberland River form attained a distribution in the Preglacial
Cumberland drainage as far downstream as what is now northwest
Tennessee (Fig. 20A and B). In the Mississippi Embayment this form was
probably restricted to small streams along bluffs immediately adjacent to
the river and was probably never common or widespread. The absence of
appropriate habitat in the lower portions of the Duck River where it passed
through the lowland Mississippi Embayment precluded its entry into the
system. Access to the Duck system did not occur until the Pleistocene
when the center of Highland Rim peneplain, formed in the Miocene
(Thornbury, 1965), hollowed out to form the Nashville Basin. Develop-
ment of this low relief basin may have allowed the Cumberland form to
gain access to the Duck through stream capture. Correlated with the
development of the Nashville Basin was the isolation of a population in the
Caney Fork system on the eastern remnant of the Highland peneplain (Fig.
20B). This eastern population differentiated into E. olivaceum.

Later in the Pleistocene, the middle and upper portions of the present-
day Tennessee River were captured by the Cumberland drainage (Fig.
20C), thus creating drainage patterns of today (Starnes and Etnier. in
press). This event was instrumental in allowing the Nashville Basin and
lower Cumberland form (ancestor of crossopterum and nigripinne) to gain
access to the middle Tennessee River drainage and later differentiate into
E. nigripinne (Fig. 20C). Factors responsible for separating these two
forms and promoting their differentiation would most likely have been the

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 71

marginal habitat of the western Highland Rim. Alternatively, one may
hypothesize that the ancestor of E. nigripinne may have been introduced
into tributaries of the middle Tennessee River through stream capture
across the Duck-Tennessee divide, thus isolating it and favoring differ-
entiation. Marginal habitat on the western Highland Rim would have been
instrumental in this scenario in preventing contact of these two forms at the
confluence of the Duck and Tennessee rivers.

Since the origins of these species, there have been several minor
modifications of their ranges. Adjustments in the range of E. squamiceps
include the elimination of the northern part of its range by glaciation and
its invasion of the Red River system (Cumberland Dr.) presumably via
stream capture.

Distributional changes in E. neopterum include dispersal south through
tributaries of the Tennessee River into northern Alabama (with some
differentiation) and east into the lower Duck River after the connection of
the Tennessee-Duck drainages in the Pleistocene. Many of these streams
have marginal habitat and populations are low. Mississippi Embayment
populations have been eliminated except for those in the Bayou de Chien of
western Kentucky (Fig. 14).

Modifications of the range of E. crossopterum include invasion of the
upper Caney Fork via stream capture from the Duck River (also known for
several other taxa), invasion of Cypress Creek (middle Tennessee
drainage) via stream capture from the Duck River, movement west onto
the western Highland Rim, where habitats are marginal and populations
are low, and elimination of most of the populations in the Mississippi
Embayment. After the Ohio River was diverted south into the stream bed
of the Cumberland and the Mississippi River was diverted east to join the
new Ohio River near Cairo, Illinois, the changed size and nature of the
river and its floodplain would have adversely affected populations of E.
crossopterum living in the small creeks immediately adjacent to the
original river. The only population remaining is in a few small creeks near
Reelfoot Lake in northwestern Tennessee (Fig. 9).

Etheostoma nigripinne has spread east throughout most of the Elk
River and tributaries of the Tennessee as far as the lower Paint Rock River
in northwestern Alabama, and north into tributaries of the Tennessee River
as far as the mouth of the Duck River. Populations present in the upper
Duck River (Bedford and Coffee Counties) may represent introductions
via stream capture within the eastern Highland Rim between the upper Elk
and Duck rivers. In this region headwaters of these two drainages
interdigitate considerably and the divide is minor. Alternatively, these
populations may indicate previous dispersal throughout the Duck system.
Whatever the case this phenomenon is not unique to E. crossopterum and
E. nigripinne since sympatric occurrences of two forms of Noturus
elegans (Etnier and Jenkins, 1980) and E. duryi and the "golden snubnose
darter" (Etnier, pers. comm.) are known from the Duck system.

Etheostoma olivaceum is the only species which has been unable to
expand its range significantly. It is blocked on the east by the Cumberland

72 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Plateau escarpment and on the west by possible competition with E.
crossopterum.

Etheostoma virgatum group.— The distribution patterns of this group are
complex (Fig. 17) and as noted earlier the geological history of this region
is poorly known. Thus, our interpretation of the biogeography of this
group with reference to its evolutionary history (Fig. 19) is difficult. The
origin of the ancestral barcheek species was presumably in the ancestral
Cumberland drainage since most primitive members of the subgenus
(obeyense and virgatum) both occur in this drainage and the two species
occurring outside the Cumberland are closely related to E. smithi in the
Cumberland drainage. The easiest geological event to correlate with the
evolutionary history of this group is the formation of the Nashville Basin.
This event was probably closely tied to the origin of the ancestral species
of E. smithi, E. striatulum and E. barbouri. Prior to the formation of the
Nashville Basin the ancestral species of this group plus E. virgatum was
widespread in the middle Cumberland River on the Highland peneplain.
Once the basin hollowed out E. virgatum became isolated from the basin
form, presumably being left on some part of the existing Highland Rim.
Later in the Pleistocene, stream capture within the basin introduced a
population into the Duck system which later differentiated into E.
striatulum. It is possible that this capture was also involved in introducing
the Cumberland ancestor of E. crossopterum and E. nigripinne into the
Duck system.

The origin of E. barbouri in the Green River system could have been
correlated with a possible stream capture across the middle Cumberland-
Barren divide, introducing the ancestor of E. barbouri into the Green
system. Alternatively, the origin of species of this species pair (smithi and
barbouri) could have been correlated with glacial processes fracturing the
range of a once more widespread ancestral species inhabiting the lower
Cumberland, Green River system, and tributaries of the Ohio River in
intervening areas. Glacial outwash and lakes in the area of the present
lower Ohio and Green rivers could have eliminated populations of the
ancestor of these species in the lower Green and Ohio tributaries, thus
promoting their differentiation.

Events correlated with the early origins of E. obeyense and E.
virgatum are difficult to determine. Etheostoma obeyense could have been
an isolate on the Cumberland Plateau when the ancestor of the remaining
barcheeks (Fig. 19) moved down over the 100 meter escarpment onto the
Highland Rim peneplain. Because populations of this species are also
known from the eastern Highland Rim, this scenario necessitates its later
dispersal downstream. Alternatively, E. obeyense may have evolved on the
eastern Highland Rim and later dispersed up onto the Cumberland Plateau.

Page and Schemske (1978) suggested that the distribution of E.
virgatum in three segments of the Cumberland River drainage may
indicate that this species was once more widespread and these populations
represent relicts. If E. virgatum is a single species and was once more
widespread we cannot address events leading to its initial differentiation.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 73

Alternatively, it is possible that E. virgatum evolved in one of the main
tributaries of the Cumberland River (in the Nashville Basin, eastern
Highland Rim or Cumberland Plateau) and dispersed to other drainages.
Because E. virgatum occupies a wider range of habitat types than most
other barcheeks, it seems possible that it could have originated on the
eastern Highland Rim in possibly the Caney Fork system when the
Nashville Basin was formed and has subsequently spread down into the
basin and up onto the Cumberland Plateau.

In summary, the early ancestors of the barcheek group may have been
distributed on the Cumberland Plateau. Movement down onto the High-
land Rim peneplain in the late Tertiary isolated E. obeyense from the
ancestor of the remaining species. Etheostoma virgatum may have differ-
entiated from the ancestral species of the remaining three taxa when the
Nashville Dome hollowed out to form the Nashville Basin, leaving the
former species on the eastern rim and the latter ancestor in the basin. The
origin of E. virgatum may then have been correlated with the origin of E.
olivaceum. Etheostoma striatulum was next temporally and became
isolated in the Duck system via stream capture between the Cumberland
and Duck in the basin. This event may have been correlated with the entry
of the crossopterum-nigripinne ancestor into the Duck. The origins of E.
smithi and E. barbouri may have resulted when glaciation fractured the
range of a more widespread species into lower Cumberland and upper
Green River populations, respectively. Alternatively, E. barbouri may
have evolved in the upper Green River after a population of the ancestor of
these two species was captured from the ancestral Cumberland drainage.

Adjustments of the distributions of E. smithi since its origin include
possible dispersal into tributaries of the lower Cumberland River (given
that the latter of the above two scenarios for the origin of this species and
E. barbouri is accepted) and the invasion of some tributaries of the lower
Tennessee via stream capture or dispersal as hypothesized by Starnes and
Etnier (in press) (Fig. 17). A likely location for stream capture between
the Cumberland and Tennessee rivers would be White Oak Creek
(Tennessee drainage) and Yellow Creek, a tributary of the Cumberland.
Etheostoma smithi, a large river and reservoir tolerant species, could have
dispersed into the lower Cumberland drainage.

Alterations of the range of E. virgatum might have included down-
stream invasion of the Stones, Harpeth, and Red rivers and upstream
dispersal into the Rockcastle River. Given that the origin of E. barbouri
included a stream capture, it has experienced considerable dispersal since
its origin. Modifications of the range of E. striatulum have apparently only
included reduction of its range. Page (1980) and Kuehne and Barbour
(1983) noted that populations of this species have been reduced due to
agricultural practices.

Etheostoma flabellare group.— Until work on the undescribed species of
this group and an analysis of the variation off. flabellare is completed it is
impossible to discuss much about the zoogeography of this group.
However, their ancestral species must have been widespread and consider-

74 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

able dispersal, largely on the part of the widely tolerant E. flabellare, has
occurred. One possible explanation for the origin of these two species is
that E. flabellare evolved in the Teays-Mississippi system and E. ken-
nicotti in the preglacial Ohio-Cumberland drainage. The hypothesis of a
widespread E. flabellare was also considered by Pflieger (1975). Neither
species was present in the middle and upper Tennessee prior to the
connection of this drainage with the Duck-Cumberland drainage in the
Pleistocene since no relatives occur in the Mobile Bay drainage and would
likely be represented there given that either form was present at the time of
the Tennessee R. -Coosa R. connection. Once the Ohio River was diverted
south into its present location during the Pleistocene, E. flabellare gained
access to the Cumberland and Duck drainages. Later in the Pleistocene
when the Tennessee united, access to the middle and upper portions were
available for both species.

Comparative Biogeography.— historical events involved in the evolution
of Catonotus are not unique to this group. Several other darters,
particularly the subgenus Nanostoma (sensu Page, 1981), appear to have
experienced a similar history. Relationships of species in these groups,
however, are not as well known as in Catonotus. Within Nanostoma, E.
barrenense and E. rafinesquei (Page and Burr, 1982) together are
congruent with the distribution of E. barbouri and are partially congruent
with E. squamiceps. Other species endemic to the Green River system
include the fishes Moxostoma (Tfioburnia) atripinne and E. bellum, and the
crayfish Barbieambarus cornutus and Orconectes barrenense (Page and
Burr, 1982). Based on results from the present study we would predict that
these species would have their closest relatives in the Cumberland, lower
Ohio, and perhaps Tennessee rivers. Within the subgenus Tfwbumia,
however, there are only two other species, both found in Atlantic Slope
drainages. Results from a phylogenetic study of this group (Buth, 1979)
indicate that the two east coast species are most closely related, and M.
atripinne forms the sister group to these. Thus, at least within this group
some extinction may have occurred. Relationships of E. bellum are not as
thoroughly resolved but Zorach (1968) considered this species close to E.
ruffilineatum of the Cumberland and Tennessee rivers and later (Zorach,
1972) placed it in the species group with E. camurum of the Ohio,
Cumberland, and Tennessee rivers and E. chlorobranchium of the upper
Tennessee.

Other darter species have striking distributional similarities with
species of Catonotus. Etheostoma etnieri and E. olivaceum in the Caney
Fork River; the "golden snubnose darter" and the pair E. s. simoterum
and E. s. atripinne with the species pair E. crossopterum-E. nigripinne,
except that E. nigripinne does not ascend the Tennessee River as far as E.
simoterum; E. duryi, E. jessiae, and E. nigripinne, except again the latter
does not extend as far upstream; the "red snubnose darter" and E.
neopterum; and E. microlepidum of the Cumberland, E. aquali of the
Duck and E. sp. of the Elk (Williams and Etnier, 1978) with E.
crossopterum and E. nigripinne. Although relationships of these species

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 75

are not fully resolved and distributions of some are not exactly the same,
primarily due to habitat differences, it will be interesting to see how much
congruence with the subgenus Catonotus is observed.

HABITAT AND COMPETITION

In addition to geological and drainage histories of a region determining
species ranges in Catonotus, habitat association and competition with
congenerics may have had a profound effect on the distributions of these
species. This aspect of Catonotus biogeography has also been discussed by
Page and Schemske (1978). Each species group in Catonotus can be
characterized as having a preferred habitat, although there is variation
within groups and even within a species. This is somewhat at variance with
ideas presented by Page and Schemske (1978); our interpretations are
stated below. The flabellare group prefers sand or gravel pools and gravel
riffles while the squamiceps and virgatum groups prefer slab or bedrock
pools. Table 15 lists the habitat preferences of the different Catonotus
species based on collections from over 400 localities (MEB). We suggest,
therefore, that these designations represent the normal habitats for each
species. In fall and winter, however, strays may be found in any habitat.
Since breeding locations of some species and populations are unknown, it
is also possible that they migrate to different habitats for spawning. For
example, whether populations of E. kennicotti and E. neopterum in sandy
creeks have modified their spawning behavior or whether they migrate to
an area with flat rocks is not known.

Throughout most of the Green, middle and lower Cumberland, and
middle and lower Tennessee rivers, members of each species-group are
generally allopatric but often are sympatric with a representative of
another Catonotus species group. Although E. kennicotti and E. flabellare
commonly occur together, there are no records of any two members of the
virgatum group occurring together, and instances of sympatry
between members of the squamiceps group are rare.

Etheostoma flabellare group. — Etheostoma flabellare is the only true riffle
inhabiting Catonotus and is the most widespread in distribution. Although
sympatric with all other Catonotus (Table 16) it is not truly syntopic with
any and therefore would presumably not be in direct competition.

Page and Schemske (1978) hypothesized that the absence of E.
kennicotti in the lower Cumberland River drainage is a result of its failure
to compete successfully with other pool inhabiting Catonotus. We believe
this to be only part of the explanation. Since E. kennicotti is known to
occur with E. flabellare, E. barbouri, E. squamiceps, E. nigripinne, and
E. neopterum, competition with other Catonotus must not be the most
important factor in determining its distribution. In fact, E. kennicotti
rarely occurs without at least one other Catonotus being present. The total
number of Catonotus species inhabiting the region does not seem to limit
the occurrence of E. kennicotti. Although seven species of Catonotus
inhabit the middle and lower Cumberland River no more than three are

76

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Table 16. Sympatric populations of Catonotus species. X = sympatric population,
? = probable sympatric distribution based on literature reference or ranges.

Species

&

u

3

3

&

c
c

.§.

-a

s

■3 es

£5 -c

■5 c

•S TOTALS

flabellare

X

X

X

X

X

X

X

X

X

X

X

11

kennicotti

X

X

X

9

X

7

X

5(7)

squamiceps

neopterum

olivaceum

X
X
X

X
X

X

X

X

3

4
1

crossopterum
nigripinne

obevense

X
X
X

9

X

7

X
X

X

X

X

5(6)

3

1(2)

virgatum
striatulum

X
X

X
X

2
2

smithi

X

X

2

barbouri

X

X

X

3

found in any one stream and E. kennicotti occurs with at least that many
species in the Green River System (squamiceps, flabellare, barbouri). If
competition with other Catonotus is not sufficient to explain the distribu-
tion of E. kennicotti perhaps the answer is competitive interactions
between E. kennicotti and other darters. Throughout most of its range, E.
kennicotti prefers sand and gravel pools (Table 15), but in the upper
Cumberland, upper Tennessee and Shawnee Hills it also inhabits slabrock
pools and sometimes gravel or rock riffles. In these areas, there are few
species of darters and competition could be minimal. For example, in the
Tradewater River (Shawnee Hills), the only common darters are E.
kennicotti, E. squamiceps, and E. nigrum; thus E. kennicotti and E.
squamiceps are ubiquitous in these streams, inhabiting most pools and
riffles. Etheostoma kennicotti inhabits the upper Green River, but is absent
from the Barren River— a system rich in darter species. Likewise E.
kennicotti occurs in some areas of the Buffalo, but not the rest of the Duck
River System. Probably E. kennicotti (or the ancestral species) was
primitively a slab rock species, but switched to a sandy or gravel pool
habitat (at least for most of the year), except at localities lacking
competitors where it reverts to its ancestral habitat.

Etheostoma squamiceps group. — Distributions of members of the squam-
iceps group are apparently not affected by competition with members of
other species groups. Etheostoma squamiceps species occur sympatrically
with all other Catonotus except E. obeyense and occur syntopically with all
of these species except E. flabellare. Within their own group, however,
competition may be the most important factor limiting species distribu-
tions. Except for E. neopterum, members of this group have the same
preferred habitat, shallow pools, usually with a bedrock substrate,
containing flat slab rocks. Etheostoma crossopterum and E. nigripinne

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 11

have less rigid requirements than E. squamiceps and E. olivaceum and are
sometimes found in small numbers in gravel or sandy pools. Apparently,
however, no two slab rock species can exist syntopically, as is similarily
observed in the slab pool inhabiting species of the virgatum group.
Although E. crossopterum, E. squamiceps, and E. olivaceum inhabit the
middle and lower Cumberland River drainage and E. crossopterum and E.
nigripinne inhabit the Duck River, no mixed collections are known.
Etheostoma neopterum is the only exception to this generalization. The
preferred habitat of E. neopterum is sandy streams, a habitat where no
syntopic occurrences with other members of the squamiceps group species
are known. Mixed collections off. neopterum and E. crossopterum or E.
neopterum and E. nigripinne occur in gravel bottom streams— a habitat
marginal for both species. Apparently, neither species can attain high
enough population levels in these marginal areas to exclude the other
species. It is noteworthy that E. neopterum has the most distinctive
breeding characteristics of any species of the squamiceps group and thus
could probably co-exist with less chance of any interspecific mating.

Etheostoma virgatum group.— The distributional pattern of barcheek
darters is unusually compact but complex (Fig. 17). Three of the five
species are nearly sympatric in the middle and lower Cumberland River
drainage. The remaining two species, E. barbouri and E. striatulum,
occur only in the Green and Duck river systems, respectively. Etheostoma
obeyense and E. smithi occupy the same microhabitat— small, permanent,
quiet, bedrock bottom creeks where they hide under flat rocks, among
emergent vegetation or in cracks in the bedrock. Etheostoma obeyense
inhabits streams on the eastern Highland Rim and E. smithi in the
Nashville Basin and the western Highland Rim. Etheostoma virgatum, if a
single species, is more versatile, occupying habitats ranging from small
intermittent creeks to fairly large rivers. It may also inhabit slab rock
pools, gravel bottomed pools or quiet gravel riffles. Apparently E. smithi
may exclude E. virgatum from some areas, but specialized habitat
requirements limit its distribution to small direct tributaries of the
Cumberland River. The most important limiting factor seems to be riffles.
Larger streams and those creeks on the eastern slope of the Nashville
Dome have long gravel riffles, some of which are quite rapid. Because of
its large size and wide habitat tolerance, E. virgatum can bypass these
riffles and invade areas not occupied by E. smithi. Thus, E. virgatum
occupies large tributaries (Red, Harpeth, Stones, Caney Fork rivers) while
E. smithi is found in the smaller, quiet tributaries. Page and Schemske
(1978) describe the ranges of E. smithi and E. virgatum as relictual—
remnants of formerly wide ranges. It is also possible that their distribu-
tions represent dynamic equilibrium between two competing relatives and
is the result of recent expansion of ranges rather than contractions. One
interesting situation is the distribution of E. smithi and E. virgatum in the
Stones River. Etheostoma virgatum is common throughout the lower river
and West Fork while E. smithi is found in only two tributaries of the East
Fork. This represents the only population of E. smithi in a large tributary

78 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

of the Cumberland River. In 1967, extensive collecting in Bradleys Creek,
one of the known E. smithi localities, failed to reveal any barcheeks. In
1978 additional collections in this region indicated that E. smithi was
common at the same localities. Between 1967 and 1978, the Percy Priest
Reservoir was constructed on the lower Stones River and the lake extends
almost to the E. smithi localities. Perhaps habitats created by the lake gave
E. smithi a chance to bypass the expansive riffles which formerly acted as
barriers. It will be interesting to see if this species spreads throughout the
rest of the Stones River system.

The distribution of E. obeyense is much harder to interpret. Appar-
ently, it can exclude E. virgatum just as E. smithi can, and because of its
larger size, can inhabit larger streams. Thus, E. obeyense may have kept
E. virgatum from the South Fork and Obey rivers, they only sizeable
tributaries of the middle and lower Cumberland drainage where E.
virgatum is lacking. The Cumberland Plateau escarpment acts as a barrier
on the east for E. obeyense but the barrier against westward expansion is
unclear. Competition with E. smithi is a possible explanation although
there is a hiatus of several kilometers between their two ranges.

Factors regulating the distribution of E. barbouri and E. striatulum
apparently are historical rather than biological. Both species are the only
virgatum group members in their respective areas, although both occur
syntopically with other pool inhabiting Catonotus and competition with
other barcheeks is not a factor in explaining their present distributions.
Etheostoma barbouri occurs throughout the Green and Barren rivers
wherever suitable habitat is present. Etheostoma striatulum inhabits every
permanent tributary of the Duck River in the Nashville Basin, but has been
unable to move out of the Basin onto the Highland Rim even though some
streams appear to have suitable habitat.

In reviewing ecological preferences, distributions, and competition
between species of Catonotus, the following pattern emerges:

1. The ancestral Catonotus species inhabited slab rock pools.

2. Interspecific competition importantly affects distributions within
species groups, but members of different species groups often occur
syntopically.

3. Within species groups, sympatry is most likely between species
with a long evolutionary separation, and with some habitat and
secondary sexual divergence.

4. The species having the largest ranges iflabellare, kennicotti, vir-
gatum) owe their "success' ' to modified habitat preferences that
minimize competition with other Catonotus. Etheostoma flabellare has
shifted from a pool to a riffle habitat and has been most successful in
expanding its range. Etheostoma kennicotti and E. virgatum are less
dependent on a specific habitat and flexible enough to use any areas not
already occupied by other Catonotus. This flexibility was accompanied
by a loss of competitive advantage within any one habitat and both

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 79

species are strongly influenced by competition by other Catonotus
species as well as other species.

LITERATURE CITED

Agassiz, L. 1854. Notice of a collection of fishes from the southern bend of the Tennessee

River, Alabama. Amer. Journ. Sci. and Arts, ser. 2, Vol. 17, pp. 297-308,

353-369.
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.

Mich. No. 93. 44 p.
Braasch, M. E.. and L. M. Page. 1979. Systematic studies of darters of the subgenus

Catonotus (Percidae), with the description of a new species from Caney Fork,

Tennessee. Occ. Pap. Mus. Nat. Hist. Univ. Kansas 78:1-10.
Buth, D. G. 1979. Genetic relationships among the torrent suckers, genus Thoburnia.

Biochem. Syst. and Ecol. 7:311-316.
Coker, R. E. 1926. New genus of darter from western North Carolina. Bull. Bur. Fish.

42:105-108.
Coker, R. E. 1927. Richiella to replace Richia as name for genus of darter. Copeia

162:17-18.
Collette, B. B. 1965. Systematic significance of breeding tubercles in fishes of the family

Percidae. Proc. U.S. Nat. Mus. 117(3518):567-614.
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Percidae, Etheostomatini). Proc. U.S. Nat. Mus. 1 19(3550): 1-88.
Cross. F. B. 1967. Handbook of fishes of Kansas. Misc. Pub. Mus. Nat. Hist. Univ. Kansas

45. 357 p.
Etnier, D. A. 1980. Etheostoma atripinne and Etheostoma simoterum. pp. 625 and 693. In

D. S. Lee, et al. Atlas of North American Freshwater Fishes. N.C. State Museum

Natural History, Raleigh, i-x + 854 p.
Etnier, D. A., and R. E. Jenkins. 1980. Noturus stanauli, a new madtom catfish

(Ictaluridae) from the Clinch and Duck Rivers, Tennessee. Bull. Alabama Mus.

Nat. Hist. 5:17-22.
Flynn, R. B.. and R. D. Hoyt. 1979. The life history of the teardrop darter, Etheostoma

barbouri Kuehne and Small. Am. Midi. Nat. 101:127-141.
Forbes, S. A., and R. E. Richardson. 1908. The fishes of Illinois. 111. Nat. Hist. Surv.,

Urbana. 357 p.
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Streams, 3:1-137.
Girard, C. 1859. Ichthyological notices, 5-27. Proc. Acad. Nat. Sci. Philadelphia, vol.

11:56-68.
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Percomorphi). Bull. International Comm. Zool. Nomen. l(23):235-244.
Howell, W. M., and G. Dingerkus. 1978. Etheostoma neopterum, a new percid fish from

the Tennessee River system in Alabama and Tennessee. Bull. Alabama Mus. Nat.

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Geol. Surv. Ind. 1874 (1875): 197-228.
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collections of the Untied States National Museum. II. A.— Notes on Cottidae,

Etheostomatidae, Percidae, Centrarchidae, Aphododeridae, Dorysomatidae, and

Cyprinidae, with revisions of the genera and descriptions of new or little known

species. Bull. U.S. Nat. Mus. 10:1-68.
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1877:42-82.
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80 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Cumberland, Whitley County, Kentucky, with descriptions of three new species.

Proc. U.S. Nat. Mus. 6:248-251.
Jordan. D. S., and B. W. Evermann. 1896. The fishes of North and Middle America: a

descriptive catalogue of the species of fishlike vertebrates found in the waters of

North America, north of the Isthmus of Panama. Bull. U.S. Nat. Mus. 47, pt.

1:1-1240.
Jordan, D. S., B. W. Evermann, and H. W. Clark. 1930. Checklist of the fishes and

fishlike vertebrates of North and Middle America north of the northern boundary of

Venezuela and Columbia. Rept. U.S. Comm. Fish., App. 10, 670 p.
Kirsch, P. H. 1892. Notes on the streams and fishes of Clinton County, Kentucky, with a

description of a new darter. Bull. U.S. Fish. Comm., 1890, vol. 10:289-292.
Kuehne, R. A., and R. W. Barbour. 1983. The American darters. The University Press of

Kentucky, Lexington. 177 p.
Kuehne. R. A., and J. W. Small. Jr. 1971. Etheostoma barbouri, a new darter (Percidae,

Etheostomatini) from the Green River with notes on the subgenus Catonotus.

Copeia 1971:18-26.
Mayden, R. L. 1985. Nuptial structures in the subgenus Catonotus, genus Etheostoma

(Percidae). Copeia 1985:580-583.
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Poecilichthys parvipinnis (Gilbert and Swain). Copeia 1950:139-148.
Page. L. M. 1974. Life history of the spottail darter, Etheostoma squamiceps, in Big Creek,

Illinois, and Ferguson Creek, Kentucky. 111. Nat. Hist. Surv. Biol. Notes, 89:1-20.
Page, L. M. 1975a. Relations among the darters of the subgenus Catonotus of Etheostoma.

Copeia 1975:782-784.
Page, L. M. 1975b. The life history of the stripetail darter. Etheostoma kennicotti, in Big

Creek, 111. 111. Nat. Hist. Surv. Biol. Notes 93:1-15.
Page, L. M. 1977. The lateralis system of darters (Etheostomatini). Copeia 1977:472-475.
Page, L. M. 1980. The life histories of Etheostoma olivaceum and Etheostoma striatulum,

two species of darters in central Tennessee. 111. Nat. Hist. Surv. Biol. Notes

113:1-14.
Page, L. M. 1981. The genera and subgenera of darters (Percidae, Etheostomatini). Occ.

Pap. Mus. Nat. Hist. Univ. Kansas 90:69 p.
Page, L. M. 1983. Handbook of darters. T. F. H. Publications, Neptune City, N.J. 271 pp.
Page, L. M., and M. E. Braasch. 1976. Systematic studies of darters of the subgenus

Catonotus (Percidae). with descriptions of a new species from the lower Cumber-
land and Tennessee River system. Occ. Pap. Mus. Nat. Hist. Univ. Kansas 60: 1-18.
Page, L. M., and M. E. Braasch. 1977. Systematic studies of darters of the subgenus

Catonotus (Percidae), with the description of a new species from the Duck River

system. Occ. Pap. Mus. Nat. Hist. Univ. Kansas 63:1-18.
Page, L. M.. and B. M. Burr. 1976. The life history of the slabrock darter, Etheostoma

smithi, in Ferguson Creek, Kentucky. 111. Nat. Hist. Surv. Biol. Notes 99:1-12.
Page, L. M., and B. M. Burr. 1982. Three new species of darters (Percidae, Etheostoma)

of the subgenus Nanostoma from Kentucky and Tennessee. Occ. Pap. Mus. Nat.

Hist. Univ. Kansas. 101:1-20.
Page, L. M. and R. L. Mayden. 1979. Nesting site of the lollypop darter, Etheostoma

neopterum. Trans. Ky. Acad. Sci. 40(l-2):56-57.
Page, L. M., and D. W. Schemske. 1978. the effect of interspecific competition on the

distribution and size of darters of the subgenus Catonotus (Percidae: Etheostoma).

Copeia 1978:406-412.
Page, L. M., and P. W. Smith. 1976. Variation and systematics of the stripetail darter,

Etheostoma kennicotti. Copeia 1976:532-541.
Page, L. M., W. L. Keller, and L. E. Cordes. 1981. Etheostoma (Beleosoma)

longimanum and E. (Catonotus) obeyense, two more darters confired as egg- .

clusters. Trans. Ky. Acad. Sci. 42(l-2):35-36.
Page, L. M., M.E. Retzer, and R. A. Stiles. 1982. Spawning behavior in seven species of

darters (Pisces: Percidae). Brimleyana 8:135-143.
Pflieger, W. L. 1971. A distributional study of Missouri fishes. Univ. Kansas Publ. Mus.

Nat. Hist. 20(3):226-570.

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS 81

longimanum and E. (Catonotus) obeyense, two more darters confirmed as egg-

clusterers. Trans. KY. Acad. Sci. 42(l-2):35-36.
Page, L. M., M. E. Retzer. and R. A. Stiles. 1982. Spawning behavior in seven species of

darters (Pisces: Percidae). Brimleyana 8:135-143.
Pflieger. W. L. 1975. The fishes of Missouri. Missouri Dept. of Conservation, Jefferson

City. 343 p.
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exchange for other specimens, with annotations. Bull. Mus. Comp. Zool. Harvard

1(1):2-16.
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l'inte'rieur des Etats - Unis d'Amerique, durant l'annee 1818. Journ. Phys., Chim.,

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Catonotus kennicotti Putnam. Copeia 1975:777-778.
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the Tennessee and Cumberland rivers drainage realm. In: Zoogeography of North

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six species of darters in the subgenus Catonotus. Biochem. Syst. and Ecol. 7:81-85.
Zorach, T. 1968. Etheostoma bellum, a new darter of the subgenus Nothonotus from the

Green River System, Kentucky and Tennessee. Copeia 1968:474-482.
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robranchium New Species, with a discussion of the subgenus Nothonotus. Copeia

1972:427-447.

82 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

APPENDIX 1
Characters and implied transformations used in the phylogenetic
analysis of species of Catonotus. The condition listed first and generally
coded as 0 is considered primitive in Catonotus. See text for explanations
of implied transformations.

1. Genital papillae of females: tubular, conical = 0, flat and not bifurcate = 1.

2. Spawning location: in gravel = 0, on undersides of slab rocks = 1.

3. Dorsal fin shape: high and rounded edge=0, low and straight edge= 1.

4. Knobs on first dorsal fin: absent = 0, large = 1, small = 2.

5. Head of breeding males: not swollen = 0, swollen = 1.

6. Infraorbital pore 4: present = 0, absent = 1.

7. Lateral line: completely pored = 0. <75% pored=l, <45% pored = 2, <40%
pored = 3, <35% pored = 4, <30% pored = 5, <20% pored = 6.

8. Habitat preference: gravel riffle, raceway = 0, slabrock pool= 1, sand-gravel pool = 2.

9. Nape squamation: scaled = 0, naked = 1.

10. Prepectoral squamation: scaled = 0, naked=l.

1 1. Opercle squamation: scaled = 0, naked = 1.

12. Cheek squamation: scaled = 0, naked = 1.

13. Suborbital bar: strongly developed = 1, absent = 0, weakly developed = 2.

14. First dorsal fin pigmentation: orange band present = I, no orange band, but similar in
other respects = 2, reds and black basoanterior spot = 0.

15. Infraorbital pore 3: present = 0, absent = 1.

16. Iridescent red and white cheek bar: absent = 0, present = 1.

17. Preopercle edge: entire = 0, crenulate= 1.

18. Longitudinal stripes on flanks: absent = 0. present = 1. strongly developed = 2.

19. Infraorbital pore 2: present = 0, absent = 1.

20. Infraorbital pore 5: present = 0, absent = 1.

21. Caudal spots: absent = 0, present=l.

22. Rays of second dorsal fin: extending only to fin margin = 0, extending well beyond fin
margin = 1 .

23. Windows in first dorsal fin membranes: absent = 0, present = 1.

24. Number of eggs spawned by a female at a time: one = 0, greater than one (usually 3) = 1 .

25. Female spawning behavior: inverted for extended period of time = 0, inverted

briefly = 1.

26. Tips of rays of second dorsal fin: pointed = 0, expanded into large knobs = 1.

27. Branches of rays of second dorsal: separate = 0. adnate (appearing unbranched) = 1 .

28. Tips of second dorsal fin rays: white = 0, black = 1.

29. Modal number of pectoral fin rays: 12 = 0, 11 = 1.

30. Knobs on first dorsal fin: normal = 0, greatly enlarged = 1.

31. Color of knobs on first dorsal fin: opaque or white = 0, golden = 1.

32. Sexual distribution of first dorsal fin knobs: typically males only = 0, males and
females = 1 .

33. Modal number of dorsal fin spines: 9 = 0, 8=1, 7 = 2.

34. White lines in second dorsal fin: concave = 0, tending to form horizontal dashes =1.

35. Modal number of anal fin rays: 8=1, 7 = 0, 9 = 2.

36. Modal number of dorsal fin rays: 13 = 0, 12= 1.

37. Modal number of dorsal fin rays: 13 = 0, 11 = 1.

38. Modal number of dorsal fin rays: 13 = 0, 14= 1.

39. Second dorsal fin color pattern: submarginal colored = 0, submarginal region with broad,
white band= 1. submarginal region black = 2.

40. Preoperculomandibular pores: 10 = 0, 9=1.

41. Clear basal band in first dorsal fin: present = 0. absent = 1.

42. Knobs on first dorsal fin: elongate = 0, round = 1.

43. Windows in membranes of first dorsal fin: round = 1, vertical slits = 0, ovals = 2.

44. Scales above lateral line: 4= 1, 5 = 2, 6 = 3, 7 = 4.

45. Scales below lateral line: 7=1, 8 = 2, 9 = 3.

46. Modified fleshy ridges on ventrolateral scales of breeding males: absent = 0, present = 1 .

TWO NEW DARTERS OF THE SUBGENUS CATONOTUS

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