OCCASIONAL PAPERS

of the

MUSEUM OF NATURAL HISTORY

The University of Kansas^ft
Lawrence, Kansas

UARVARO
NUMBER 60, PAGES 1-18 OCTOBER 20, 1976

SYSTEMATIC STUDIES OF DARTERS OF THE

SUBGENUS CATONOTUS (PERCIDAE),

WITH THE DESCRIPTION OF A NEW SPECIES

FROM THE LOWER CUMBERLAND AND

TENNESSEE RIVER SYSTEMS

By

Lawrence M. Page1 and Marvin E. Braasch2

In 1892 Etheostoma obeyense was described by P. H. Kirsch
from tributaries of the Cumberland River in Clinton County, Ken-
tucky, and since that time has been considered to be one of only
two species of barcheek darters in the Cumberland drainage, the
other being E. virgatiun. However, individuals considered until
now to be E. obeyense actually represent two distinct but morpho-
logically similar species. Etheostoma obeyense is distributed in
tributaries of the middle Cumberland River, and a second species,
described herein, is in tributaries of the lower Cumberland and
lower Tennessee rivers.

Methods

Characters. — The characters examined were numbers of lateral
scales, pored lateral line scales, scales above and below the lateral
line, scales around the caudal peduncle, transverse scales, infra-
orbital canal pores, supratemporal canal pores, preoperculomandib-
ular pores, dorsal fin spines, dorsal fin rays, branched caudal fin
rays, pectoral fin rays, anal fin spines, and anal fin rays; pigmenta-
tion patterns; squamation patterns {i.e., amount of squamation on
various areas of the body); head length/ standard length (HL/SL);
head width/SL (HW/SL); body depth/SL (BD/SL); caudal pe-
duncle depth/SL (CPD/SL); pectoral fin length/SL (P1L/SL);

1 Assistant Taxonomist, Illinois Natural History Survey, Urbana 61 SOI

2 Assistant Professor, Belleville Area College, Belleville, Illinois 62221

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

predorsal length/SL (PreDL/SL); second dorsal fin base length/
first dorsal fin base length (D2L/D1L); preorbital length/BD
(PreOL/BD); interorbital width/head width (IOW/HW); BD/
anal fin length (BD/AL); first dorsal fin height/DIL (D1H/D1L);
pelvic fin length/SL (P2L/SL); eye diameter/HL (ED/HL); eye
diameter/SL (ED/SL); gape width/preorbital length (GW/
PreOL); cheek bar width/PreOL ( CBW/PreOL ) ; and second dor-
sal fin length/distance from the anterior origin of the second dorsal
fin to the caudal base (D2FL/QL).

Counts and measurements were made as described by Hubbs
and Lagler (1964) with the following exceptions. The number of
transverse scales was counted in two ways — from the anal fin origin
to the first dorsal fin base, and from the anal fin origin to the second
dorsal fin base. Body depth was measured at the origin of the first
dorsal fin. First dorsal fin height is the length of the second spine.
Preorbital length is the distance from the tip of the snout to the
anterior margin of the orbit. Pectoral fin and pelvic fin lengths are
the lengths of the longest rays. Second dorsal fin length was meas-
ured from the origin of the fin to the tip of the most posterior-
reaching ray. Cheek bar width is its greatest transverse distance.
Eye diameter is the greatest transverse distance across the orbit.
Head canal pore counts were made on the left side only and in the
manner described by Hubbs and Cannon ( 1935) .

Analysis. — Counts and measurements (measurements are ex-
pressed as proportions of one another) were compared among
samples determined by sex (dimorphic), by size class (ontoge-
netic), by river system (geographic), and by longitude (geo-
graphic). For some comparisons (e.g. Table 3) adjustments were
made by limiting samples to reduce the influence of other potential
sources of variation (in Table 3, ontogenetic). One-way analysis of
variance tests were run to determine significances of differences in
means of samples determined by sex, size class, and geography.

Etheostoma smithi, new species
Slabrock Darter

Holotype. — Illinois Natural History Survey 75013, an adult male
51.7 mm SL (Fig. 1), collected in Ferguson Creek at the Route 70
bridge (37°6' N, 87°24' W) 3 km E Smithland, Livingston County,
Kentucky, on 26 April 1975 by L. M. Page, B. M. Burr, and J. A.
Boyd.

Paratopotijpes. — A total of 121 specimens deposited as follows:
13— Illinois Natural History Survey (INHS 75014, 25-51 mm SL, 26
Apr. 1971); 24 — Universitv of Michigan Museum of Zoology
(UMMZ 197498, 22-51 mm SL, 22 Apr. 1971); 24— U. S. National
Museum (USNM 214525, 26-49 mm SL, 20 May 1973); 14— The
University of Kansas (KU 16227, 15-41 mm SL, 29 May 1972);

SYSTEMATIC STUDIES OF DARTERS

1

Fig. 1. — Etheostoma smiihi holotype, male, 51.7 mm SL, Ferguson Creek,
Livingston Co., Kentucky, 26 April 1975.

13— Northeast Louisiana University (NLU 32515, 23-50 mm SL,
20 Feb. 1973); 13— Tulane University (TU 95964, 30-43 mm SL,
27 May 1971); 12— University of Alabama (UAIC 5021.01, 23-47
mm SL, IS Jan. 1972); 8— University of Tennessee (UT 91.1099,
30-50 mm SL, 3 May 1971).

Material examined. — Drainages are listed in a west-to-east di-
rection. Numbers in parentheses are numbers of specimens exam-
ined. Complete collection locality data are available from the first
author. CUMBERLAND R. DR.: Ferguson Cr.: INHS 75013 (1),
75014 (5), 75016 (11), 75017 (10), UMMZ 197498 (11); Sandy
Cr.: INHS 75018 (2); Clay Lick Cr.: UMMZ 174983 (3); Goose
Cr.: UMMZ 174858 (2); South Cross Cr.: CU (Cornell University)
47400 (1); Elk Cr.: CU 47859 (17); Wells Cr.: INHS 75019 (1);
Richland Cr.: UMMZ 174482 (6), 177571 (5); Whites Cr.: USNM
188733 (5); Stones R.: INHS 75029 (21), TU 19470 (1), NLU
15779 (1); Drake Cr.: UMMZ 88070 (2); Bledsoe Cr.: UMMZ
174440 (1), UAIC 2974 (1); Spring Cr.: MEB (M. E. Braasch)
80 (7). TENNESSEE R. DR.: Whiteoak Cr.: INHS 75038 (3),
TU 89495 (1), UT 91.552 (2); Hurricane Cr.: INHS 75039 (4),
UT91. (2).

Etymology. — The new species is named for Dr. Philip W. Smith,
Head of the Section of Faunistic Surveys and Insect Identification
of the Illinois Natural History Survey, in recognition of his out-
standing contributions to vertebrate natural history. The common
name, slabrock darter, refers to the preference of the species for
slabrock habitats.

Diagnosis. — The subgenus Catonotus of Etheostoma was diag-
nosed by Kuehne and Small (1971). Etheostoma smithi is distin-
guished from all other members of the subgenus by the following
combination of characteristics: a bicolored bar on cheek; red and
blue pigments on fins and head; caudal fin without distinct black
vertical bands; infraorbital canal interrupted, with three pores an-
teriorly and one posteriorly (Fig. 2); 10 preoperculomandibular

4 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

pores; no conspicuous dark longitudinal stripes on sides; no (or
a weak) suborbital bar; adults (over 35 mm SL) with 13 or fewer
pored lateral line scales; usually with 14 dorsal fin rays.

Fig. 2. — Head canals of Etheostoma smithi (left) and E. obeijense (right).
The infraorbital canal of E. smithi has three pores anteriorly and one pos-
teriorly; of E. obeijense has four pores anteriorly and two posteriorly.

Comparisons. — Etheostoma smithi can be distinguished from
the other described barcheek darters as follows. Etheostoma obey-
ense, the closest relative, has four pores in the anterior segment of
the infraorbital canal, usually two pores in the posterior segment
of the infraorbital canal, 12 or more pored lateral line scales (in
adults), 13 dorsal fin rays, and a mean PreOL/BD of 0.44 (0.39 in
E. smithi) (Tables 1-3). Etheostoma harhouri has a strong tear-
drop, usually nine preoperculomandibular pores, and 13 dorsal fin
rays. Etheostoma virgatum has conspicuous dark longitudinal
stripes on the sides, usually four pores in the anterior segment of
the infraorbital canal, and two pores in the posterior segment of
the infraorbital canal.

Description. — A small Catonotus reaching 52 mm SL; elongated
with a terminal mouth, and compared to its closest relative, E.
obeijense, with a significantly (a = 0.005) shorter snout (preorbital
length). Infraorbital canal interrupted (Fig. 2) with three pores
anteriorly (99% of the specimens examined; N = 115) and one
pore posteriorly (99%; N = 115); supratemporal canal interrupted
medially with pores usually 2 — 2; usually 10, rarely nine, pre-
operculomandibular pores; snout with a rather broad frenum; six
branchiostegal rays on a side, with the branchiostegal membranes
separate or slightly fused; preopercle crenulate.

Head unsealed; body scaled except for the nape, breast, and
prepectoral area; nape unsealed to approximately the origin of the
first dorsal fin; 41 to 54 lateral scales; 2 to 13 (usually 10 or fewer)
pored lateral line scales; 14 to 17 transverse scales from anal fin
origin to first dorsal fin; 12 to 15 scales to second dorsal fin; 17 to
22 (usually 19 or 20) scales around caudal peduncle.

>

u

Q

oo

'X

CD
CM

to

CM

CM

CO

cm

CM
CM

i— I
CM

O
CM

T— 1

00
I— 1

t-

1— 1

CO

i— I

C/3

o

C/3

to

T— 1
1— 1

T3

O
Oh

co

CM

i— I

d

i—i

O

i—i

cx>

00

t>

CD

to

^r

co

cm

a!
c/TQ

■^ to ^_ CO CO C5 CM

TJH 6 00 00 00 rt rH

CM ^H CM CO CM CM CM

CM 00 CM CM -^ — I O

i-J to CD t^ to t> TP

CM O CM i— i CM — i i— (

i> WHTf qoqco

00 to it- -t C5 t^ CD

C5
CM

f — < t~- to -t t~-

^H

^H

CM

CM

CM

CO

CO

^

■— 1 i— 1

CD

CD

CM CM —I

^H
i-H

Tt<

CM i— i i— i

00

tJ<

CM i—l i—l CO

^H
■—I

co

CM

CM i— 1 i— 1 i-H i— 1

1—1
i-H

CM

CM

i—i i— ( CM

00

CM

CM

i-H i-H

CM

0>

Q .2

c

C

co

•c

U

c

o

PC U

6 c

e is

S J ^ »; c ~

M
C

Kl

-D ^ -2 ^ ^ ^ "

gQ

H

-

«

en

CD

u

1; S

r- wo

§ a

s.

e

00

H-l

<

H
O
H

pcOtoto-HCDCDCMO

t^cor^to'co't^i-icD^'

i— I i— I i— IMHi- (i— I i— ii— I

C00500CMO0505CD1— I

^CD0qC5toCD0CtoO5

CM CMCM-tfCMCO^'cMCM

COtotoCOOOOCOl^OO

-tcDCDOico'i-HCDtoCD
i—i i — I i— I i— i i—l CM ^h ^h CM

00O)CD-tth-O05in
CM i— i i-h <—< i-H

<M

CO CM

CM

CM

CO

co

CO i—l
CM CM i—l
^H t- CM

i— I ~? i— I

i— I
CM

i— I CM
CM CM

CM HO)

co co co i-h

CO CM
i— I CM to CM

CM CM

CO

5*

05

to

O

ft

o

o

to

Z
H

=3

u

HO

«Ph

>>MH

C1^ >H >H >H >H ^

5 ^ m" h) I- [i, ^

1 « u w> fl co

2 3 » ,g « o

£ o > "5 c •£

: g 3 .-g £

u

CM

CD
1— I

to

co

CO
00
02
to

i-H

©

I-H

to

1— I

i—l
i-H

05
t>
to
CM

oo

H
O
H

6

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

S
to

to

-a
o

a

O
to

o
a

a
a
o

o

C
3
O

U

cm

w
j

a
<

<S

U

0)

g
w

C/3

o

OS

U

c

OJ

—

s

C/3
O

u

Q

C/3

'X

CM

>

u

Q

C/3

'X

-"

CO

in

a

C/3

4)

«

c

oooooooo
©do©©©©'©'

OOOOOOOOO
OCOOOOOOO

oooooooo

CObNOWOMM
CO i— I >— I CM i— I

oocoboinoNN i in

CO I— I i— I CM T— I 1— I

OOOOOCOOO

d> d> d> d> d> x^- <5 <5

CM © ©

MOO

©"©'©'©" ©' ©"©' ©'

© © © o ©
© o © © o

©©©©pi-nOO

co'cocococo'coco'co

00 t- t- © lO © b-

CO ^h — i CM

CM

00t~t-©lO©t^CM

CO i — I i— i i-^ i—i

6>c
c

o

o

U
c

a

■d

u

s '-

S OJ

U.Sgu
° ^ ,j C

z

■fHJ

Cy
60
03

C

Z

H

U

?0 S . -=5 "3 C •?

1- r£

Kl

oj .,_ s .a

=fciWK^

i-; g '■— CD

C/3 C

C

H

IT3

C/3

<

H
O
H

onini'OHffici
© oo i> co co oo co r^

(M r-i CO oi CO Ol m Ol

oo -r © cm >— i cc cm t—
coco^^riorfco-r

odd © © © © ©

©©cooo^t-©t-

t-cot-©co©c©

CM i— I i— I i— I t-H CM

OONWNnO!'* I©
CM i— I i— < I t—

i— iCOWCN©©!— iCD I CO
I CO

©TTOOt-O©©

©ioooo6o©io

OHOOf COM
© CM © © CO © © CM

©odd© ©do

0©_©©©©0©
•V CO Tj< tP co' •<* TP co'

t-coi>©©©©©

CM i— I i— I i— I r— I CM

t^CMt^©^©©© 100
CM i— I i— I i— I i— i i— I O

to

e
a

to

o

CM

z

H

M

i "*

00

<

H

O
H

5^ —

WW

>

u

Uffi

3
o
o

U

60 c ^

2 s D

tL- Ph J

Kl

SYSTEMATIC STUDIES OF DARTERS 7

Table 3. — Counts of dorsal fin rays in Etheostoma smithi and E. obeyense.

Species No. Fin Rays

&
Drainage 12 13 14 15 N X SD CV

Etheostoma smithi

Cumberland R. Drainage

Ferguson Cr., KY 10 23 5 38 13.9 0.62 4.5

tribs., Livingston, Crittenden,

Caldwell cos., KY 7 7 14.0 0.00 0.0

Elk Cr., TN 2 10 5 17 14.2 0.64 4.5

Richland Cr., TN 3 6 2 11 13.9 0.70 5.0

Whites Cr., TN 3 2 5 14.4 0.55 3.8

Stones R., TN 1 7 13 2 23 13.7 0.70 5.1

Spring Cr., TN 4 3 7 14.4 0.53 3.7

Tennessee R. Drainage 2 7 3 12 14.1 0.67 4.7

TOTALS: 1 24 73 22

Etheostoma obeyense

Obey R., TN 15 5 11 13.3 0.67 5.0

Wolf R. (ObeyR. trib.),

KY & TN

Marrowbone Cr., KY

Crocus Cr., KY

Reaver Cr., KY

Fishing Cr., KY

Pitman Cr., KY

Little S. Fk., KY

Rock Cr., TN

TOTALS: 27 94 40 3

Eight to 10 (usually 9) dorsal fin spines, without knobs; second
dorsal fin elongated with 13 to 15 (usually 14) rays; 11 to 16
(usually 14) branched caudal fin rays; 11 to 13 (usually 12) pec-
toral fin rays; usually 2 anal fin spines; 8 to 10 (usually 9) anal
fin rays; 1 pelvic fin spine; 5 pelvic fin rays.

Body proportions: HL/SL, 0.26-0.36 (X = 0.30); HW/SL, 0.11-
0.16 (0.14); BD/SL, 0.15-0.21 (0.18); CPD/SL, 0.08-0.13 (0.10);
P1L/SL, 0.22-0.27 (0.25); PreDL/SL, 0.33-0.36 (0.34); PreOL/BD,
0.30-0.46 (0.39); D2L/D1L, 0.80-1.42 (1.19); IOW/HW, 0.29-0.52
(0.36); GW/PreOL, 0.82-1.32 (1.07); BD/AL— males, 0.49-0.61
(0.56); BD/AL— females, 0.57-0.72 (0.64); D2FL/QL— males, 0.83-
0.94 (0.90); D2FL/QL— females, 0.78-0.85 (0.82); ED/HL, 0.20-
0.33 (0.26); D1H/D1L, 0.33-0.51 (0.43); P2L/SL, 0.18-0.23 (0.20);
ED/SL, 0.06-0.11 (0.08); CBW/PreOL, 0.48-0.68 (0.57).

Coloration in life: Juveniles, females, and non-breeding males
with nearly identical pigmentation; large males sometimes darker,
retaining some breeding colors for variable lengths of time after
the breeding season; large, summer-collected males often have more
red in the fins than do females. In non-breeding individuals (Fig.

12

33

6

51

12.9

0.59

4.6

3

4

7

12.6

0.53

4.3

6

3

1

10

12.5

0.71

5.7

2

13

1

16

12.9

0.44

3.4

3

10

6

19

13.2

0.69

5.2

6

4

10

13.4

0.52

3.9

8

14

2 24

13.8

0.61

4.4

12

3

1 16

13.3

0.60

4.5

8

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

3), body straw-colored dorsally with 6 to 8 dark brown rectangular
saddles on the mid-dorsum — the first (sometimes absent) being
anterior to the origin of first dorsal fin and the last at the origin of
the caudal fin; dorsal saddles especially prominent on specimens
from Stones River. Series of 9 to 12 vertically elongated dark

D

Fig. 3. — A. Etheostoma smithi male, 44 mm SL, Ferguson Cr., Livingston
Co., Kentucky, 10 Dec. 1971; B. E. obeyense male, 54 mm, Smith Cr., Clinton
Co., Kentucky, 26 July 1975; C. E. smithi female, 36 mm, Ferguson Cr., Liv-
ingston Co., Kentucky, 24 Nov. 1972; D. E. obeyense female, 37 mm, Smith
Cr., Clinton Co., Kentucky, 2 July 1975.

SYSTEMATIC STUDIES OF DARTERS 9

brown blotches mid-lateraly extending only about two-thirds of
distance to midventer anteriorly, but encircling or nearly encircling
venter on caudal peduncle. Variously developed vermiculations of
same or slightly lighter color between lateral blotches and dorsal
saddles. Breast and belly white to straw color; breast usually
spotted with large melanophores. Humeral spot large and black.

Dorsum of head dark brown to black posteriorly and lighter
anteriorly with patches of melanophores; distinct preorbital bars
converging but not meeting on the upper lip; occasional faint sub-
orbital bar; cheeks heavily spotted with large melanophores. Cheek-
bar with black outline; lower half of cheek bar white to silver;
upper half red to orange; iridescence of lower half producing a
mirror effect. Opercle, underside of head, and branchiostegal mem-
branes heavily spotted with large melanophores.

First dorsal fin with large, anterior, basal, black blotch best
developed between second and fifth spines; rest of fin clear except
for scatterings of melanophores and a marginal or submarginal red
band; sometimes thin black band distally outlining red band. Sec-
ond dorsal and caudal fins with bands formed by concentrations of
red-brown and black pigments on fin membranes. Pectoral fin
similarly banded, with melanophores along rays, or clear. Pelvic
fin clear. Anal fin clear except for red-brown and black pigment
spots basally on membranes.

Breeding male (Fig. 1) spectacularly colored with bold, pre-
dominantly red dorsal, caudal, and anal fins; blotch in first dorsal
fin blacker and larger than in nonbreeding males and surrounded
by red; anal fin and lower half of caudal fin develop a blue-black
margin; breast and pelvic fins black; pectoral fins reddish medially
with wide, black margin, best developed ventrally; head dark and
swollen; cheek bar vivid; humeral spot especially prominent; verti-
cal bars on sides dark and more elongate than in nonbreeding males;
tubercles absent.

Variation. — Means of three of the 29 characteristics of Etheo-
stoma smithi tested show highly significant (a = 0.005) sexual
dimorphism. Males have more dorsal fin rays, a smaller BD/AL
ratio, and a larger D2FL/QL ratio ( Table 4 ) . Among these three
characters, only the D2FL/QL ratio is also sexually dimorphic in
both of the other species of Catonotus that have been examined —
E. kennicotti (Page and Smith 1976) and E. oheyense (Table 4).
A second dorsal fin longer in males than in females is characteristic
of all three species.

Means of six of the 29 characteristics tested show highly sig-
nificant variation among three size classes, as follows: <35 mm
SL; 35.1-40.0 mm SL; and >40.0 mm SL. Only the values for the
smallest and largest classes are shown in Table 5. Larger E. smithi
have more pored lateral line scales. Relative to standard length,
larger individuals have a shorter head, body depth, pelvic fin

10 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

to

&
to
s>

to

O

a
a

c

to

1*3
a

5
*«

3
0)

W)

c
o

o

u

w

'X

v

a

as
•—

03

u

o o cm

CO CD CO

co i>

co m

-tf T CM

in o o

odd

■14.0

■0.667

•0.832

13.6-
0.609-
0.798-

co in
co —i

CO CD CO

co o d

i-H

cd as as

cq co ^

■^ cd co"

,59

0466

0307

o o o

■14.5

0.590

0.915

14.1-
0.530-

0.877-

O CD

cd as
co in co

-rod

r— 1

in as ©

03

K|

Q CM

to

S

to

qinn

co d in

C5 CO •— i

o m o

H(B*f

O O O

odd

CM —1 CD

t— CO i-H

co i> co

odd

d t^ 6

(OWN
CO CD t-

d d d

CO ^f CO

co as as

CO CD N

odd

CM CM CM

i— 1 i— 1 i— 1

co m t-

CM l> tJ<

CM r- i r— 1
CO CM CO

o o o

odd

l> o o
m co t

co in as

odd

as d ^
~r co as
co m co

odd

CO CO t>

in m r- 1
co in as

odd

1* f "^

c/3
Q

P Q CM

£ ffi Q

1*3

SYSTEMATIC STUDIES OF DARTERS

11

50

to
Si

to

-a
o

3
03

Co

O

to

o

—

as
o

-O

a

•c
>

13
o

X)

(II

c

o
g

O

in

u

c3

u

0)

N

CO

o

U

<y,

>y.

—< i> in in t> oo

CO

lO lO 00 CD CM

— i 05 o> co in

r-H CM O O ^H O

-r p p © © p
cm o © © © ©

00 © CM CO ©

O I- O CO CD

CD CO i— I CM CM O

00 O CD d o o

I I I I I I

00 00 b- 00 CO ©

CD

co co co o in

(NHrtMO

d d d d d

oo co in cm -v

© b- © CM CD
b- CM h i— i (M O

t^ © © © © ©

b- © © b- "^ ^
CM ■— I

HCIOWN00

oi in ^ d t^ co

CD 00 O tP CO
NCOrf«o]0O

O H O h (M O
00 O O O O O

' d d d d d

CM

00 CO l> 00 —I
CM O CM O O

oo co cm cm co ■— ;

do © do d

oo oo -H

^ CO —I

b- b- ©

Ot-00
CM CM O

o o o o o

oo t- b- cm in

i— i © i— < © ©
00 CO ^H CM CM O

in d d d d d

co co oo co oo co

CO ^H

cfl
CO

o

en

■« 5

a
o

<
o
to

Kl

2 K « 2 W W

CO Tf O i-J "tf -*

HCOOOOt^C)
CM CM i— ( t— I

CD CM CO CO CO
in T!< CM CO ©

co cm -h m i— i co

CO o o o o o

c6 d d d d d

^ ci N « f

© i— I CO I> CO ©
qq CO — ; CM O co

—t d d d d d
i i i i i i

^h co -* in in cm

^ © b- -# b- CD
J_, CM ^-J CM O CO

d d d d d

CO 00 ^H 05 00

ON©bt-

O CO i— J CM O CO

ooddddd

CD ^V CM CM CO CM

b- 'T ^ "* -* CM

to
to
S
to

to

o

a
g
o

CO

O

<»

CI

CD 0 p Tf i— i

1 —

co

"* !> O r-H CO
■ — 1 1 — 1 1 — 1

CI
CM

CS iH rH t^ O
i— 1 r-i CO I— I CO

p p p o o

-*'

d d d d d

CO CO CD CO 00
00 CO © CM O CO

^ co ih co — i in

p— I

p p p o o

CO

ci

—

co co d 06 co

CM CO © © CO

nrtcoot

d d d d d

CO CO CO CM CM

CM CO >— i O i— i

in co ^n co i— i in

co d d d d d

-r --i t— t- t- co

"^f CO CM CM CM CM

o

CD

c

13

&CO CflS73\

o'jJQQQh
Z Hi cq h w Q

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

length, and eye diameter; and relative to head length, a shorter
eye diameter.

Geographic variation among characteristics examined in Etheo-
stoma smithi is minor and no clinal or other consistent pattern of
variation is evident. Although a hiatus within the Cumberland
River distribution of E. smithi is as large as the hiatus between the
ranges of E. smithi and E. oheyense, no morphological difference
between the eastern and western populations of E. smithi was
found.

Ten of the 12 specimens examined from the Tennessee River
system have eight dorsal fin spines; a majority (56%) of the speci-
mens from the Cumberland River system have nine dorsal fin spines.
No other differences were found between the samples from these
two major river systems.

The sample from Stones River, Rutherford Co., Tennessee, is
the most unusual of the Etheostoma smithi examined. There is
more contrast between light and dark pigments on these individuals
than on individuals from other drainages, and the dorsal saddles
are especially prominent. The dorsal saddle anterior to the first
dorsal fin, which is usually weak or absent in E. smithi, is strongly
developed in the Stones River population. In addition, the Stones
River specimens have fewer than the usual number of dorsal fin
rays (Table 3) and a reduced number of pectoral rays (X = 11.5;
all other samples except that from Elk Creek, Stewart Co., Ten-
nessee, have means of 12.0 or more) .

The population in Richland Creek, Davidson Co., Tennessee,
is somewhat unusual, having especially low numbers of pored
lateral line scales (Table 1), of branched caudal fin rays (X =
12.9; all other samples have means over 14.1), and of lateral scales
(X = 45.1; other samples except that from Spring Creek, Wilson
Co., Tennessee, have means over 45.5) .

Distribution and Habitat. — Etheostoma smithi occurs in tribu-
taries of the lower Cumberland River from near the mouth of the
Cumberland River in western Kentucky to near the Caney Fork
system in north central Tennessee and in tributaries of the lower
Tennessee River (Fig. 4). Based on presently available locality
data on museum specimens, there is a large hiatus in the Cumber-
land River distribution of the species in Tennessee; the hiatus
extends from approximately the abrupt bend in the Cumberland
River at Cumberland City to near Nashville. A collecting trip by
the authors in November, 1975, to this area failed to produce any
specimens of E. smithi. At present E. smithi is known in the Ten-
nessee River system from only two tributaries: Whiteoak Creek,
a direct tributary of the lower Tennessee River in Humphreys and
Houston counties, Tennessee; and Hurricane Creek, a tributary of
the lower Duck River in Humphreys County, Tennessee.

SYSTEMATIC STUDIES OF DARTERS

13

-37

-35

Fig. 4. — Distribution of localities from which Eiheostoma smiihi (triangles)
and E. obeyense (circles) have been collected.

With the exception of the approximately 900 specimens of E.
smithi collected from the type-locality, most collections of the
species are quite small. The species is usually present in small
populations, and difficulty in collecting is further exacerbated by
the difficulty of collecting in slabrock pools.

The habitat of E. smithi is shallow, slabrock pools in streams.
A life history study similar to those recently published on other
Catonotus (Page 1974, 1975a) is completed for E. smithi in Fer-
guson Creek, Livingston Co., Kentucky, and is awaiting publication.

Etheostoma obeyense Kirsch 1892
Barcheek Darter

Etheostoma obeyense Kirsch, 1892:292 (original description).
Poecilichthys obeyensis. — Shoup, Peyton, and Gentry, 1941:70
(list of fishes of the Obey River).

Types. — Etheostoma obeyense was described in 1892 on the
basis of 210 specimens from four Cumberland River tributaries in
Clinton County, Kentucky. Of the 210, 21 were from Indian Creek,
36 from Spring Creek, 23 from Smith Creek, and 130 from Albany
Branch. The original description included a figure of the new
species, later reproduced by Jordan and Evermann (1896) who
also redescribed the species and selected a lectotype (USNM

14 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

45565). Collette and Knapp (1966) erred in stating that the figure
of E. obeyense in Jordan and Evermann (1896) was of the lecto-
type.

Collette and Knapp (1966) located the following types of E.
obeyense: lectotype, USNM 45565 (male, trib., Cumberland R.,
52 mm SL); paralectotypes, SU (Stanford University) 5116 (41
specimens, Albany Br., 28-51 mm SL), SU 1918 (38, Spring Cr.,
32-55 mm SL), and BMNH (British Museum) 1892.12.30.136-139
(4, Albany Br., 35-41 mm SL). In addition UMMZ has a series of
43 paralectotypes (UMMZ 61558, 30-44 mm SL) from Albany Br.,
collected in 1891 by P. H. Kirsch. The collection was formerly IU
(Indiana University) 4318.

Material examined. — Drainages are listed in a west-to-east di-
rection. Numbers in parentheses are numbers of specimens exam-
ined. Complete collection locality data are available from the
first author. CUMBERLAND R. DR.: Obey R.: KU 11512 (11);
Wolf R. (Obey R. Dr.:): UMMZ 125463 (10), 125669 (2), 125678
(1), KU 11533 (25), USNM 204345 (10), UT 91.183 (2), TU
32849 (1); Kettle Cr.: UMMZ 125608 (1); Marrowbone Cr.:
UMMZ 154638 (7); Crocus Cr.: UMMZ 177990 (10); Beaver Cr.:
UMMZ 169479 (16); Fishing Cr.: INHS 75015 (6), UMMZ
169381 (3), 171603 (10); Pitman Cr.: UMMZ 168017 (10); Little
S. Fk.: UMMZ 169489 (4), UT 91.452 (10), 91.665 (10); Rock
Cr.: UTuncat. (16).

Diagnosis. — Etheostoma obeyense is distinguished from all other
members of the subgenus by the following combination of char-
acteristics: a bicolored bar on cheek; red and blue pigments on
fins and head; caudal fin without distinct black vertical bands; in-
fraorbital canal interrupted, with four pores anteriorly and usually
two pores posteriorly (Fig. 2); 10 preoperculomandibular pores;
no conspicuous dark longitudinal stripes on sides; no (or a weak)
suborbital bar; adults (over 35 mm SL) with 12 or more pored
lateral line scales; usually with 13 dorsal fin rays.

Comparisons. — Etheostoma smithi was contrasted earlier with
E. obeyense. Etheostona barbouri has a strong suborbital bar, 12
or fewer pored lateral line scales, usually nine preoperculomandib-
ular pores, three pores in the anterior segment of the infraorbital
canal, and one pore in the posterior segment of the infraorbital
canal. Etheostoma virgatum has conspicuous dark longitudinal
stripes on the sides.

Description. — A moderate-sized Catonotus reaching 70 mm SL
(about 33% larger than E. smithi); elongated with a terminal
mouth, and a significantly longer snout (preorbital length) than
that of E. smithi. Infraorbital canal interrupted (Fig. 2) with four
pores anteriorly (96% of the specimens examined; N = 112) and
usually two pores posteriorly (71%; N = 112); supratemporal canal
interrupted medially with pores usually 2—2; usually 10, rarely

SYSTEMATIC STUDIES OF DARTERS 15

nine, preoperculomandibular pores; snout with a rather broad
frenum; six branchiostegal rays on a side with the branchiostegal
membranes separate or slightly fused; preopercle crenulate.

Head, nape to the origin of the first dorsal fin, breast, and pre-
pectoral area unsealed; 39 to 56 lateral scales; 10 to 26 (usually
12 or more) pored lateral line scales in adults; 12 to 18 transverse
scales from anal fin origin to first dorsal fin; 12 to 15 scales to the
second dorsal fin; 16 to 22 (usually 19) scales around the caudal
peduncle.

Eight to 10 (usually 9) dorsal fin spines, without knobs; second
dorsal fin elongated with 12 to 15 (usually 13) rays; 13 to 16
(usually 14) branched caudal fin rays; 11 to 14 (usually 12) pec-
toral fin rays; usually 2 anal fin spines; 7 to 10 (usually 9) anal
fin rays; 1 pelvic fin spine; 5 pelvic fin rays.

Body proportions: HL/SL, 0.20-0.35 (X = 0.32); HW/SL,
0.11-0.15 (0.14); BD/SL, 0.15-0.22 (0.18); CPD/SL, 0.08-0.14
(0.11); P1L/SL, 0.20-0.28 (0.25); PreDL/SL— males, 0.34-0.37
(0.35); PreDL/SL— females, 0.35-0.38 (0.37); PreOL/BD, 0.35-
0.71 (0.44); D2L/D1L, 0.81-1.13 (0.95); IOW/HW, 0.28-0.44
(0.37); GW/PreOL, 0.67-1.17 (0.93); BD/AL— males, 0.49-0.63
(0.56); BD/AL— females, 0.58-0.79 (0.69); D2FL/QL— males, 0.85-
0.99 (0.92); D2FL/QL— females, 0.73-0.86 (0.79); ED/HL, 0.20-
0.50 (0.28); D1H/D1L, 0.31-0.73 (0.45); P2L/SL, 0.16-0.27 (0.21);
ED/SL, 0.06-0.13 (0.09); CBW/PreOL, 0.43-0.66 (0.54).

Coloration in life: The coloration of Etheostoma obeyense, in-
cluding that of the breeding male, is nearly identical to that of
E. smithi. A subtle difference in E. obeyense, both in non-breeding
individuals and breeding males, is a tendency for the mid-lateral
pigment to be more concentrated into a series of squarish, but
diffuse blotches, as opposed to the more vertically elongated
blotches of E. smithi ( Fig. 3).

Kuehne and Small ( 1971 ) included a color photo of a breeding
male E. obeyense collected on 2 April 1969 and photographed one
day later. Although the blue-black margins on the lower caudal,
anal and lower pectoral fins are not shown, these dark margins are
as characteristic of E. obeyense breeding males as of E. smithi.
Several other collections of preserved breeding E. obeyense (in-
cluding UMMZ 154638 and UT 91.665) retain the dark fin margins
as described for E. smithi. Tubercles do not develop.

Variation. — Means of three of the 29 characteristics tested show
highly significant (a = 0.005) sexual dimorphism. Males have
smaller PreDL/SL and BD/AL ratios, and a larger D2FL/QL ratio
than do females (Table 4). The sexual dimorphism in the PreDL/
SL ratio is not expressed in E. smithi, and the dimorphism in the
number of dorsal fin rays in E. smithi is not found in E. obeyense
(although there is a significantly higher number in males than in
females of E. obeyense at a = 0.05).

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Means of six of the 29 characteristics tested showed highly
significant variation among three different size classes ( <35 mm
SL, 35.1-40.0 mm SL, and >40 mm SL). Data for only the smallest
and largest classes are shown in Table 5. Five of the six charac-
teristics show the same pattern of variation as in E. smithi; larger
E. obeyense have more pored lateral line scales and, relative to
standard length, a shorter head, body depth, and eye diameter.
In addition, larger E. obeyense have a shorter first dorsal fin height
in relation to fin length. The ontogenetic variation in pelvic fin
length expressed in E. smithi is not found in E. obeyense.

The two characters consistently showing ontogenetic variation
in the three Catonotus species examined, E. smithi, E. obeyense,
and E. kennicotti (Page and Smith 1976), are the number of pored
lateral line scales and relative head length.

Geographic variation among characteristics examined is greater
in E. obeyense than in E. smithi but also shows no clinal or other
consistent pattern. The samples examined from Fishing Creek,
Pulaski Co., Kentucky, and Rock Creek, Pickett Co., Tennessee,
have especially high numbers of pored lateral line scales (Table 1)
and pectoral fin rays (X = 12.9 and 13.0, respectively; all other
samples have means under 12.5). The samples from Marrowbone
Creek, Cumberland Co., and Beaver Creek, Wayne Co., Kentucky,
have especially low counts of pores in the posterior segment of the
infraorbital canal (Table 2). The sample from Crocus Creek, Cum-
berland Co., Kentucky, has a high number of lateral scales (X =
51.4; all other samples have means under 49.8).

Distribution and Habitat. — Etheostoma obeyense occurs in trib-
utaries of the middle Cumberland River from the Obey River sys-
tem east to tributaries of the Big South Fork (Fig. 4). The species
generally is common in this rather restricted area of northcentral
Tennessee and southcentral Kentucky, as first noted by Kirsch
( 1892 ) in the original description of the species.

Little ecological information is available on E. obeyense other
than observations by Kuehne and Small ( 1971 ) that the species
occurs in pools of 2nd, 3rd, and 4th order streams, and that of
Comisky and Etnier ( 1972 ) that the species is common over sand
in shallow water. We found E. obeyense to be most common in
bedrock pools ( in which crevices are occupied as hiding places by
the darters) and in slabrock pools in small streams.

Relationships

No evidence of intergradation or of an east-to-west cline linking
the two populations now regarded as E. smithi and E. obeyense
was found. Such evidence would have suggested incomplete dif-
ferentiation of the two populations.

Etheostoma smithi and E. obeyense are morphologically similar

SYSTEMATIC STUDIES OF DARTERS 17

species, allopatric but occupying the same river system ( Fig. 4 ) ,
and presumably have differentiated in relatively recent time. They
have until now been considered to be one species. Recent examples
of this misinterpretation are in a publication by Kuehne and Small
(1971) in which their "Lower Cumberland E. obeyense" was ac-
tually E. smithi, and in a publication by Page (1975b) in which
the "E. obeyense" on which spawning observations were made
actually was E. smithi.

Reduced meristics, reductions in the lateralis system, and a
smaller maximum body size are derived characteristics of darters.
Thus, smaller numbers of pored lateral line scales and infraorbital
canal pores, and the smaller maximum size of E. smithi, suggest
that E. smithi is a derivative of E. obeyense. The only character
found at variance with this evidence is a higher modal number of
dorsal fin rays in E. smithi than in E. obeyense.

Other close relatives of E. smithi and E. obeyense are the other
barcheek darters, E. virgatum and E. barbouri, and secondarily,
the nonbarcheek species of Catonotns, E. kennicotti and E. fla-
bellare (Page 1975b).

Acknowledgments

We are indebted to P. W. Smith, B. M. Burr, and M. A. Morris,
all of the Illinois Natural History Survey, for criticism on our early
manuscript; to The University of Kansas Museum of Natural History
and its personnel for support and guidance during the early stages
of study on Catonotns; to D. A. Etnier, University of Tennessee,
for information on the distribution of Tennessee Catonotns; and to
the following ichthyologists and their institutions for the loans of
specimens: R. M. Bailey, University of Michigan Museum of Zool-
ogy; G. C. Beckham, Cornell University; H. T. Boschung, Univer-
sity of Alabama; B. B. Collette, United States National Museum of
Natural History; F. B. Cross, University of Kansas; N. H. Douglas,
Northeast Louisiana University; D. A. Etnier, University of Ten-
nessee; and R. D. Suttkus, Tulane University. Technical assistance
was provided by B. P. Sweeney, S. D. Cowan, C. W. Ronto, L.
LeMere, and L. Farlow, all of the Illinois Natural History Survey.

Summary

Three species of barcheek darters (Etheostoma virgatum, E.
obeyense, and E. smithi) occur in the Cumberland River system.
Etheostoma obeyense Kirsch and E. smithi, herein described, are
closely related but can be distinguished as follows: E. obeyense
has four pores in the anterior segment of the infraorbital canal,
usually two pores in the posterior segment of the infraorbital canal,
12 or more pored lateral line scales (in adults), modally 13 dorsal
fin rays, and a mean PreOL/BD of 0.44; E. smithi has three pores

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

in the anterior segment of the infraorbital canal, one pore in the
posterior segment of the infraorbital canal, 13 or fewer pored
lateral line scales, modally 14 dorsal fin rays, and a mean PreOL/
BD of 0.39. Eiheostoma obeyense occurs in the middle Cumberland
River system. Eiheostoma smithi occurs in the lower Cumberland
and lower Tennessee river systems.

Literature Cited

Collette, B. B., Knapp, L. W. 1966. Catalog of type specimens of the

darters (Pisces, Pericidae, Etheostomatini). Proc. U. S. Nat. Mus. 119

(3550): 1-88.
Comiskey, C. E., Etnier, D. A. 1972. Fishes of the Big South Fork of the

Cumberland River. J. Tennessee Acad. Sci. 47(4) : 140-145.
Hubbs, C. L., Cannon, M. D. 1935. The darters of the genera Hololepis

and Villora. Univ. Michigan Mus. Zool. Misc. Publ. 30. 93 p.
Hubbs, C. L., Lagler, K. F. 1964. Fishes of the Great Lakes region. Univ.

Michigan Press, Ann Arbor. 213 p.
Jordan, D. S., Evermann, B. W. 1896. The fishes of North and Middle

America: a descriptive catalogue of the species of fish-like vetrebrates

found in the waters of North America, north of the Isthmus of Panama.

U. S. Nat. Mus. Bull. 47. 4 pts.
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, 10:289-292.
Kuehxe, R. A., Small, Jr., J. W. 1971. Etheostoma barbouri, a new darter

( Percidae, Etheostomatini ) from the Green River with notes on the

subgenus Catonotus. Copeia 1971 (l):18-26.
Page, L. M. 1974. The life history of the spottail darter, Etheostoma squami-

ceps, in Big Creek, Illinois, and Ferguson Creek, Kentucky. Illinois

Nat. Hist. Surv. Biol. Notes 89. 20 p.
Page, L. M. 1975a. The life history of the stripetail darter, Etheosoma kenni-

cotti in Big Creek, Illinois. Illinois Nat. Hist. Surv. Biol. Notes 93. 15 p.
Page, L. M. 1975b. Relations among the darters of the subgenus Catonotus

of Etheostoma. Copeia 1975 (4):782-784.
Page, L. M., Smith, P. W. 1976. Variation and systematics of the stripetail

darter, Etheostoma kennicotti. Copeia 1976 (3):532-541.