UN\

HARVARD UNIVERSITY

Library of the

Museum of

Comparative Zoology

MUS. COMP. ZOOL
Y

OCCASIONAL PAPERS 3 ° 1982

of the

MUSEUM OF NATURAL HISTORY
The University of Kansas
Lawrence, Kansas

NUMBER 101, PAGES 1-20 JUNE IS, 19S2

THREE NEW SPECIES OF DARTERS (PERCIDAE,

ETHEOSTOMA) OF THE SUBGENUS NANOSTOMA

FROM KENTUCKY AND TENNESSEE

BY

Lawrence M. Page1 and Brooks M. Burr"

Among the freshwater fishes of North America are many recog-
nized but taxonomically nndescribed species. Some only recently
have been discovered but many have been known for decades. The
three species described herein have been recognized as distinct
species for at least 30 years. Because nndescribed species are known
only to a few specialists, they do not receive the scientific attention
given described species. For this reason, accentuated by the prepa-
ration of a book on Kentucky fishes (Burr, 19S0), we herein name
and describe Ethcostoma baileiji, E. rafincsc/uci, and E. barrcncn.se.
We hope, as did Kuehne and Small (1971) and Bouchard (1977),
that our action will promote descriptions of other well-known but
undescribed North American fishes.

In describing E. etnicri, Bouchard (1977) diagnosed Uloccntra
as a subgenus and discussed its close relationship to the subgenus
Etheostoma. Later, because of the conclusion that E. zonule was
more closely related to species of Uloccntra than to species of
Etheostoma s.s., Nanostoma was recognized as a subgenus and
Ulocentra was relegated to the synonymy of Nanostoma (Page,
1981). As diagnosed by Page (1981), Nanostoma contains five

1 Taxonomist, Illinois Natural History Survey, Champaign, Illinois 61820,
U.S.A. and Associate in Ichthyology, Museum of Natural History, The Univer-
sity of Kansas, Lawrence, Kansas 66045, U.S.A.

2 Associate Professor, Department of Zoology, Southern Illinois University
at Carbondale, Carbondale, Illinois 62901.

2 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

previously described species (E. zonale, E. simoterum* E. duryi,
E. coosae, and E. etnieri), the three species described herein, and
six or seven undescribed species.

With the exception of the wide-ranging E. zonale, all species of
Nanostoma are limited to southern tributaries of the Ohio River,
tributaries of the Mississippi River below the Ohio River, or Gulf
of Mexico drainages in Mississippi, Alabama, Georgia, and western
Florida. Most inhabit clear rocky pools of small to medium-sized
streams. Etheostoma zonale occupies an extensive area of the
Mississippi River system (Tsai and Raney, 1974) and has been
introduced into the Susquehanna River (Atlantic drainage) in
Pennsylvania (Denoncourt, et al., 1975).

METHODS

Counts and measurements follow llubbs and Lagler (1964)
except as follows. Transverse scales were counted from the1 anal
fin origin to the first dorsal fin. Body depth was measured at the
origin of the first dorsal fin. Pectoral and pelvic fin lengths were
lengths of the longest rays. Interorbital width was least fleshy
width. Number of lateral blotches was from opercle to caudal fin
base. Head canal pore counts follow Hubbs and Cannon (1935).

Sample means and modes were compared for geographic varia-
tion. When significant variation was absent, samples were suc-
cessively amalgamated by river system.

EtJwostoma baileiji Page and Burr, new species
Emerald Darter

Holotype. — Illinois Natural History Survey S74S4, an adult male
43.0 mmSL (Table 1) collected in Little Sexton Creek, tributary
of South Fork Kentucky River, 2 km E (town of) Sextons Creek
(37° 19'N, 83° 45'W), Clay County, Kentucky on 20 May 19S0
by W. L. Keller, L. E. Cordes, and L. M. Page.

Paratypes. — A total of 56 specimens from South Fork Kentucky
River system deposited as follows: 12 — Illinois Natural History
Survey (INIIS S6S71, 30-42 mm SL, same collection data as holo-
type); 9— The University of Kansas (KU 19109, 29-41 mm SL, same
locality as holotype, 22 March 1978) ; 9 — University of Tennessee
(UT 91.2360, 28-42 mm SL, same collection data as KU 19109);
16— Southern Illinois University at Carbondale (SIUC 1026, 27-42
mm SL, Red Bird River, Eriline, Clay County, Kentucky, 28 Octo-
ber 1980); 10— University of Florida, Florida State Museum (UF
32481, 26-39 mm SL, same collection data as SIUC 1026).

3 Etheostoma atripinne is at most only intraspecifically distinct from E.
simoterum (Etnier and Bouchard, in manuscript).

THREE NEW DARTERS 3

Material examined ( but not designated as types). — Museum ab-
breviations are identified in ACKNOWLEDGMENTS. Numbers
in parentheses are numbers of specimens on which counts or meas-
urements were made. Complete collection locality data arc- avail-
able from the authors. KENTUCKY R. SYSTEM: INIIS 78499 (1),
7S513 (3), 7S976 (1). 79029 (7), 7909S (1), 79197 (10), S6S71
( 10). KNPC KKolLEE (4), KKolLES (3). uncat. (demons Fork,
Breathitt Co, KY. 10 Mar. 1979) (5); SIUC 1027 (7); UL 6015
(3). 7133 (1), S051 (5). 11817 (4); UMMZ 177854 (10). CUM-
BERLAND R. SYSTEM: INIIS 83897 (3); KNPC KCOIBEL
(10), uncat. (Poor Fork. Harlan Co., KY, 22 June 1978) (3); SIUC
1028 (20). 1029 (1). 1030 (5), 1031 (3).

Etymology. — Named for Dr. Reeve M. Bailey, Curator Emeritus
of Fishes, University of Michigan Museum of Zoology, in recogni-
tion of his outstanding contributions to systematic ichthyology. The
common name, emerald darter, has been used for this species for
decades (e.g., Kuehne and Bailey, 1961) and refers to the domi-
nating color of the species.

Diagnosis. — Member of subgenus Nanostoma as diagnosed by
Page (1981). As noted by Bouchard (1977) the most diagnostic
features of this group of darters are color characteristics. Etheo-
stoma baileyi is easily distinguished from all other darters by its
body and dorsal fin pigmentation. Side of body with 7-11 (usually
9-10; Table 2) small emerald-green squares (some appear as W's
when examined closely) subtending the lateral line; squares ex-
panded dorsally and ventrally as lime-green bars in breeding male.
Midline of dorsum with 7-10 (usually 8) emerald-green squares.
First dorsal fin green in male, clear in female, with red margin and
red spots on interradial membranes; red margin usually restricted
to distal one-fourth of fin except extends one-half way down height
of fin in first two membranes and occupies all of last membrane.
Without red or orange pigment except in first dorsal fin. Premaxil-
lary frennm narrow but always present. Modally five branchio-
stegal raws (Table 2) and 10 preoperculomandibular pores (Table

3).

Unlike E. baileyi, all other taxonomically described species of
Nanosto?na have red in second dorsal fin of adult male, and all
except E. zonule modally have nine preoperculomandibular pores.4
Etheostoma baileyi most closely resembles E. zonule, but E. zonule
differs by having only six dark squares on middorsum, dark vertical
bars rather than squares on side of body, and a more pointed snout.

Dcseription. — Smallest member of subgenus; maximum SL =
47 mm for males, 43 mm for females. Snont extremely blunt; mouth
small (Table 1) and subterminal. Body proportion valves given

4 Modal count for E. coosae is nine, not 10 as given by Page (1981:18).

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

_1

CO

H3

T3
(Ll

CJ

&3

o
c

3

o

cv>

W

co

Cu

c

0;

o

;-
3

P

B

c
o

<

CO

II
Z

-5'

3

►a

c

co

C
CO

Lo

■X

CO

CO

Q

co

'X

be

CO

'X

be

^h io io -r oi i—i oi co in cc cc cc i> ci oi rn pi [> 10 -r co cc
^odwjiCTfirJN^oO'tHin'^oNNoqHHOotJ

i— I i — I i—i> — I H CJi — I 1 — I i— ( I — I i — It— I

05

cncoNHCNCHOcw5t~cociri-x-*HC co
n-roTCC-r^xcccu-OTcoin-xacc

CM i— I CO i—l "—I Ol OI OI CM CM CI CM OI CI H H

IO

cc co co — t- a -r cr>
in in i~ — r io — i co — i oi oi io o>
ci ■— i in i^ cc in co oi •— i oi cm oi

CM-X C -"l-NCO
1- IO X 1- C J X h

ci ci ci ci n oi h h

cc ci

^-; co
05 ci
ci ol

in in — • oo ci ci —

co -r in co ci in co

cc i - co c ^ oi ci -r cc co
c m ci i — i i — i cm co co in ^
ci ci cm ci ci oi oi ci i— i

o

co c i- a in

-r -r -r -t co

Ol I— I

co-pinoimcccccocc i- in co
io r-i cj> •— < o oi cc co oi — o m

CO rt H rH C] oi oi oi CO i— I

co co Oi co — i t"- cc -f oi cc co co ^h t— i~ co oi t- co ci ■— i oi

lOco^occi-H-rcocoi— i co co -f i^ t> i— i co cc co ■— icocc

co cc co -t -r ~r -& *? oi co i— i co -r in ~? — i t~- o o co cri -r

CM

in t-

-r co
ci

'- -J t-

ro IO

in oi

c'l CO

-c!

cc
00 c

coincoinco-rcci->oo>ccc7ic^- ^ ^ -r —
co -r in co -^ o — i ci co co t^ co co cc oi in oi cc c75
' — I co ci oi cm oi "n

cc co -f -r oi
. o> in co> cc

Ol Ol CI CI OI OI

co

cc

Ol

cc t-coi-Hccci-r o t- co cc cc cc co

— i— — *■ co — n oi - v • — i cm t~- i — i h c? io fo cc -n o c

i— ! in co i- in co oi ^^ oi ci co m oi ci co ci co co oi

i- in w il 4 ci cc cc cc ^ o -i o -r in r- ici-t-ci

oi co -r in co t^ cc cc cc ci -r co -r — i cc co t^ co t^

i—l Ol i— I rt oi ci oi ci oi ci oi Ol i— l

—i •— i cc a — < cc co oi t- i — v a -v in co
^t'lncoininoii-HCCCioiccoicD a cc

•—( CC oi i— I '—i cl cl cl ol CI

i- in oi co co i- -r t- i- co i- co co co io co co in cc in i-h o

■-<-*ccccoi-oocccoccichicx-*xi-r-iin

-r co tr — r in i* t h o rr c

-r co c i—i t-

C5

CO

Ol

Ol

cc t-

ceo in in

~v oi i— i

■ i i

CD CO CO

ci °3 c^

CO

Cl

-*xoccc:rHOccNoixco^'tintccooi
-riocc-rco — cococooccoiococc'rot^i-cD

CO Ol Ol Ol Ol Ol Ol Ol CM Ol Ol i— i i—i

oi— -icdi-^io cc^iHCco^-*-r

t co in cc oi -r ^ on- ^ v- x io c x oi c o x

io i^ i- in co ci i— i ol oi co *t oi oi co ol co co h h

i i i i i i i i i i i i i i i i i i i

-rxoicccoicciocoTi-TCOco-rc
cotcc-tococojcccdcccioi — < co co cc cc cc -r

Ol i-H HC1 Cl Cl Ol Ol Ol Ol Ol i—i i—l

in

i- co c
-r
oi

-r -r f

in i—i
co co

in oi

CD O
CO Ol

CO CO

Ci -^

Ol

IO

oi

Cl

cc

Ol

bO

5)

tc

co

l) B

J3
M

CD
v.

*o o+

a co

C5 C35

Cl --1

-O O «0 Of

«oot «oot

bO U) 21 o?

G c *r* ■—■

Of,

o+

cd a

CO Cl) w w
"i in cS (j

s ccccs

be bi

•r r:cccc-_-

r- E« C — —

1 — ™ r> i- trl ^ — 3 CC — -1 - -

-— CC r-

'■r. 's. — D

? "=

cc cc

y b 3

cfi I) (S
co m m

Cj „;; XT3 "O —

►q co y o (£;caOUciiii^<

CO CO Ph Uh CO CO

THREE NEW DARTERS

o
c

O

Kl

o

a

s.

co
o

C3
0)

c

C
C8

Cv
03

u

Ol

<

03

PC

z

0)

o

«

Z

>

O CD

•> CI

u

CO

d -r

Q

O CO

— h _ i
CO Ol

CO

z

CD

in

>

o i<

CO ■— 1

u

CO I>

co co

Lo

cd cd

-~ CO

d d

d d

'X

CD CD

CD t-

co co

CO CD

CD

CO

=8

CD O CD CD

in in »n in

oi co in m

3 as ^r cd

CD CO

Ol O CD Ol

CD CD CO CD

Ol

CO i— I ^ M

co o oi co

^r

-? ^ T CD

cd -r -r o
-r in . — i • — <

0 cd in oi

01 CO

CI" iH

S~. c; ^ T3

"o a o O j!

o

s

o

o

-D

O
U

2

u

CO

w
J

o
PL,

3

Dh

Q

z

>

u

Q
to

•X

CD

CO

>

U

Q

CO

'X

Z

CD
CO

CO

in i-j

CO CO

CO Ol

d d

CD CD

co co

CD CO

CD 3

— < in

CO CD

co t^

CD ^

in cd"

CO

cd q
t^ cd

-H CO

3 o

c-i a
t- tr-

lO o

co co
in in

Ol Ol

in in

d d

CO CD
CD d

Ol CD

-r in

oi

CD o

Ol TJ<

ci

— I CD
CD d

-r o
t in
d d

CO CD

co t-

i- -r

r- 1 oi

oi O

CO Ol

5. s

C3" Cj

Co c* -*»

I £ e

^ i; -^

? § B C

3 e a - ~*

s s ? ^

g s g 5 aj

S "g C; (J ^

Si) It) W

6 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

in Table 1 and shown in Fig. 1. Infraorbital canal with 7 or 8 pores
(Table 3), preoperculomandibular canal with 9-11 (usually 10)
pores (Table 3). Lateral line complete to end of hvpural plate
(56 specimens), or missing on 1 (15), 2 (21), 3 (10), 4 (9), 5 (6),
6 (1), 7 (1), or 10 (1) scales posteriorly. Genital papilla of breed-
ing female long and tubular, equal in length to about one-half
length of first anal spine; of male a short tube.

Breast unsealed; prepectoral area scaled. Belly fully scaled.
Nape usually fully scaled, rarely unsealed anteriorly. Cheek and
opercle one-half to fully scaled, scales often embedded. Lateral
scales (= lateral-line scales) 45-56 (Table 4); transverse scales
12-18 (Table 5); scales above lateral line 4 (4 specimens), 5 (95),
6 (20), or 7 (1); scales below lateral line 6 (5), 7 (69), 8 (35),
9 (8), or 10 (2); scales around caudal peduncle 15-21 (Table 6).
Dorsal fin spines 10-13 (Table 7); dorsal fin rays 10-12 (Table 7);
pectoral fin rays 13 (1), 14 (61), 15 (56), or 16 (2); anal fin spines
2; anal fin rays 6 (9), 7 (91), or 8 (20); branched caudal fin rays
11 (1), 12 (2), 13 (5), 14 (14), 15 (34), or 16 (1); total vertebrae
38 (7), 39 (39), 40 (13), or 41 (4).

Color in life: Juveniles and females olive above, yellow below,
with 7-11 small emerald-green squares along midside (subtending
lateral line) and 7-10 emerald-green squares along middle of back.
First middorsal square on anterior nape, second at origin of first
dorsal fin, third under first dorsal fin, fourth at posterior end of
first dorsal fin, fifth at origin of see md dorsal fin, sixth under second
dorsal fin, seventh on caudal peduncle, eighth at origin of caudal
fin. Upper side with wavy dark green lines; lower side, belly, and
breast without dark marks. Black prcorbital bar extends onto upper
lip. Black suborbital bar (teardrop) slants slightly posteriad Neu-
trally. Postorbital bar reduced to a black spot behind eye and
black dash on upper opercle. No medial black spot at base of
caudal fin (as is common in darters) but usually a small submedial
dark green blotch. First dorsal fin described above; second dorsal,
pectoral, and caudal fins with light green-brown bands; anal and
pelvic fins clear.

In breeding male (Fig. 1) lateral blotches expanded dorso-
ventrally into bright green bars; first dorsal fin green with bright
red margin and interradial spots; second dorsal and caudal fins
dusky yellow with green bases; cheek, opercle, and anal and pelvic
fins green.

Distribution (md Variation. — Etheostoma baileyi inhabits the
upper Cumberland and upper Kentucky River systems (Fig. 2).
In the Cumberland system below Cumberland Falls, E. baileyi is
restricted to the Rockcastle River and Big South Fork. Above
Cumberland Falls. E. baileyi is known only from Tellico Creek,
Clear Fork, and the Cumberland River. Although common in the

THREE NEW DARTERS

o
c

o

hq

o

ca

53

3
o
U

a
<

o
oo

6
2

U

Q
en

'X

2

in

in
io

no

io

CT>

XT

00

o
"5t<

CT

CO

00

co

=3

cj

O-
CO

ID

t>

CT rt;

oi

CO

co ~-f

CT

co

O tM
CT3 Cl

o

r— I

i-H oi

CD

CO

^H CT

CT
CO

in

CT O

i — I

Cl

1 — 1

in -t1

Cl

in

I— I ^H
i — I

-f CO
I — I

I- Cl

i-H

CT ^H
i-H

CO CO f-

O CO l>

I — I

in o co

i-H r— I

i— I i— I C]

Cl

-r —i

Cl

CO
Ol

CO
1—1

CO CO

in

co

CO

CT

8. e

u- ••- — ~C

a e e G

c a a j- -^

~ ~ ~ ^ u

8 8 5 "^ .5

a o i ,-t .

rt o 3 '"

bj W W

8

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

^M

B

Fig. 1. — (A) Etheostoma baileyi male, 43 mm SL, Little Sexton Creek,
Clay Co., KY., 20 May 1980; (B) E. rafinesquei male, 41 mm SL, Nolin
River, LaRue Co., KY., 19 May 1980; (C) E. barrenense male, 44 mm SL,
Trammel Creek, Allen Co., KY., 13 April 1978.

Rockcastle River and elsewhere locally, E. baileyi is sporadically
distributed and generally uncommon in the Cumberland system.

Etheostoma baileyi is more widespread and common in the Ken-
tucky River system and is present in the Red River and the South,
Middle, and North forks of the Kentucky River. It does not occur
in the Dix River (contra Branson and Batch, 1974).

Etheostoma baileyi and another darter, E. sagitta, are the only
fishes restricted in distribution to the Cumberland and Kentucky
River systems (Jenkins, et al., 1972). Kuehne and Bailey (1961)

THREE NEW DARTERS 9

concluded that E. sagitta and E. baileyi entered the Kentucky sys-
tem from the Cumberland as a result of stream capture. Jenkins,
et al. (1972) postulated that E. baileyi entered the Cumberland
from the Kentucky system. Ericymba buccata, common in the Ken-
tucky system, present in the upper but not the lower Cumberland
system, and absent in the Tennessee system, must have entered
the Cumberland from the Kentucky system. Variation in E. nigrum
suggests that it also entered the upper Cumberland from the Ken-
tucky system (or the Big Sandy system) and differentiated into the
endemic subspecies E, n. susanae, then re-invaded the upper
Kentucky system which today contains E. n. nigrum x susanae inter-
grades (Staines and Starnes, 1979). Variation in Nocomis micro-
pogon also suggests transfers between upper Kentucky and upper
Cumberland fishes (Lachner and Jenkins, 1971).

Whether the original transfer of E. baileyi and other darters was
from the Kentucky to the Cumberland or vice versa is impossible
to determine. However, E. sagitta and the blackfin darter, Percina
(Odontopholis) species, both occur in the upper Kentucky system
and in one other river (the Cumberland and Green rivers, respec-
tively) in Kentucky and have as their only close relatives species
restricted to the Missouri River system in Missouri. The Kentucky
River was formerly a tributary of the upper Mississippi River (via
preglacial Teays River), as was the Missouri River (Pflieger, 1971).
This suggests that the darters first were in the Kentucky River and
later invaded other streams. If so, E. baileyi and E. nigrum prob-
ably were included in the transfer which carried E. sagitta from
the Kentucky system to the Cumberland system.

The only notable variation among characteristics examined in
E. baileyi is between the Kentucky River and the Cumberland
River populations. Kentucky River individuals have higher modal
numbers of lateral blotches (Table 2) and infraorbital pores (Table
3), and more lateral scales (Table 4) and transverse scales (Table
5). In E. sagitta the reverse was found; almost all meristic counts
were higher in the Cumberland River population (E. s. sagitta)
than in the Kentucky River population (E. s. spilotum) (Kuehne
and Bailey, 1961).

Habitat and Ecology. — Etheostoma baileyi occupies rock-bot-
tomed, current-swept pools of small to large streams. Where popu-
lations are large some individuals are found in rock riffles, especially
along the riffle margins. Individuals in breeding condition have
been collected in late March (IXIIS 79197), April (INHS 78976),
May (INHS 86871), and early June (UL 11817).

Etheostoma rafinesquei Burr and Page, new species
Kentucky Snubnose Darter
Holotype. — Illinois Natural History Survey 87488, an adult male

10

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Fig. 2. — Distribution of localities from which Etheostoma baileyi, E. rafines-

(jiiei, and E. banencn.se have been collected.

47.5 mm SL (Table 1) collected in Barren Run, tributary of North
Fork Nolin River, 2 km NW (town of) Barren Run (37° 3WN,
85° 4S'YV), LaRue County, Kentucky on 29 May 19S1 by B. M.
Burr and L. M. Page.

Paratypes. — A total of 54 specimens collected with the holotype
deposited as follows: 11 — Illinois Natural History Survey (INHS
87482, 31-48 mm); 11— The University of Kansas (KU 19111, 31-48
mm SL); 11— University of Tennessee ( UT 91.2359, 31-46 mm SL);
11— Southern Illinois University at Carbondale (SIUC 1025, 32-44
mm SL); and 10 — University of Florida, Florida State Museum
(UF 32482, 31-48 mm SL, same locality as holotvpe, 27 September
1980).

Material examined (but not designated as types). — GREEN R.
SYSTEM: INHS 74849 (3), 75395 (7), 76034 (3), 79138 (1),
84058 (2), 86874 (5), 86878 (10); SIUC 1015 (1), 1016 (1), 1017
(10); UL 5262 (9), 5379 (2), 5712 (2), 5729 (3), 5801 (13), 5817
(2), 5981 (22), 10250 (1); UMMZ 177559 (10). BARREN R.
SYSTEM: SIUC 1018 (7).

Etymology. — Named for Constantino S. Rafinesque, a pioneer
naturalist and former Professor of Botany and Natural History,
Transylvania University, Kentucky, who described 36 of Kentucky's

THREE NEW DARTERS

11

o

o
c

o

"So
O

tq

o

CO

53

c
U

in

w
j

B
<

o

GO

U

'X

3C

£

IO

CC

Ol

'3

C/5

^h in

O OT>

CO -^

CD CD*

03 05

cd in

f i-l
00 O

d d

d .-i

in p

q i>

■H CO

i—l i— 1

•—1 ■— I

■—I i— i

in in

CO

O}

C5 O
CM

00
1—1

in Tf<

C] CO

C5 -r

in o

CD Ol

i—i

CD
CO

2. =

a « « T! ^

~ ~ 2 U '->

C ~ ~ ~ 5

■*-.*-_ — — —

y >■ > _ ~

- p ~ ,r -

ia ~ - ^/ -e

tt) fcj K|

12 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

presently recognized species of fishes. The common name, Kentucky
snubnose darter, refers to the geographic domain of the species
and its inclusion in the subgenus Nanostoma, generally referred to
as snubnose darters. Etheostoma rafinesc/uei is the only fish re-
stricted to Kentucky.

Diagnosis. — Member of subgenus Nanostoma as diagnosed by
Page (1981). Distinguished from other taxonomically described
species of Nanostoma, except E. zonale, by its low lateral scale count
of 37-42 (usually 38-41; Table 4) and low transverse scale count
of 10-12 (Table 5); E. zonale has 36-63 lateral scales and 9-19 trans-
verse scales, other described species of Nanostoma have 42-61 lateral
scales and usually 13 or more transverse scales although E. duryi
has as few as 11 and E. simoterum as few as 12 (Tsai and Raney,
1974; Bouchard, 1977). Distinguished from E. zonale and all other
darters by its body and median fin pigmentation. Side of body
with 7-10 (usually 8-9; Table 2) dark bars 5-6 scale rows in height.
Dorsum with 7-9 dark blotches; first blotch largest, covering 90%
of nape, always extending posteriorly to origin of dorsal fin. Cross-
hatched appearance on side of body created by dark (red-orange
in breeding male) spots on posterior edges of scales on lower half
of body and on 1-2 scale rows above lateral line. First dorsal fin
with red spot in first interradial membrane. Snout blue-green.
Breeding male with blue-green lips, opercle, anal fin, and procurrent
caudal fin membranes; red-orange breast; red distally and basally
on interradial membranes of first dorsal fin; blue-green medially
(and occasionally basally) on first 4-6 interradial membranes of
first dorsal fin; second dorsal fin red-purple medially. Premaxillary
frenum pit-sent (narrow) in over 90% of specimens. Modally five
branchiostegal rays (Table 2) and 9 preoperculomandibular pores
(Table 3). Breast usually (69% of specimens) scaled posteriorly.

Other taxonomically described species of Nanostoma have
greatly contrasting color patterns (Bouchard, 1977). Etheostoma
rafinesquei most closely resembles E. barrenense, but E. barrenense
has 43-50 lateral scales, shorter and wider lateral blotches that are
confluent with a dark midlateral stripe, the first dorsal dark blotch
confined to anterior half of nape, and an unsealed breast (95% of
specimens).

Description. — Moderate-sized member of subgenus, maximum
SL = 54 mm for males, 49 mm for females. Snout extremely blunt;
month small (Table 1) and snbterminal. Body proportion values
given in Table 1 and shown in Fig. 1. Infraorbital canal with 7-9
(usually 8) pores (Table 3); preoperculomandibular canal with
8-10 (usually 9) pores (Table 3). Lateral line complete to end of
hypural plate (112 specimens), or missing on 1(2) scale posteriorly.
Genital papillae as in E. bailei/i.

Breast unsealed in 27 specimens, variously scaled (to 50%) in

THREE NEW DARTERS 13

61 specimens; prepectoral area scaled. Belly, nape, and opercle
fully scaled. Cheek unsealed (26) or fully scaled (64); scales
often embedded. Lateral scales 37-42 (Table 4); transverse scales
10-12 (Table 5); seales above lateral line 3 (27) or 4 (68); scales
below lateral line 5 (3), 6 (24), or 7 (67); scales around caudal
peduncle 14-16 (Table 6). Dorsal fin spines 10-12 (Table 7); dor-
sal fin rays 9-13 (Table 7); pectoral fin rays 14 (68) or 15 (45);
anal fin spines 1 (2) or 2 (97); anal fin rays 6 (8), 7 (74), or 8
(17); branched caudal fin rays 12 (3), 13 (32), 14 (33), or 15 (30);
total vertebrae 38 (14), 39 (25), or 40 (5).

Color in life: Juveniles and females yellowish with 7-10 verti-
cally elongated dark blotches along side, occasionally partly con-
fluent with, but always extending well above and below a black
lateral stripe. Dorsum with 7-9 square, vague to dark blotches.
First blotch largest, covering 90% of nape; second and third under
middle of first dorsal fin, fourth between dorsal fins, fifth and sixth
under second dorsal fin, seventh on caudal peduncle, eighth at
origin of caudal fin. Posterior edges of scales on side of body dark.
Preorbital bar darker than snout, extending to upper lip. Black
suborbital bar. Dark dashes behind eye on cheek and opercle.
Pelvic and anal fins clear. Pectoral, second dorsal, and caudal fins
with dusky bands on rays. First interradial membrane of first dor-
sal fin with red spot; remaining interradial membranes with dusky,
sometimes reddish, dashes. First dorsal fin usually with a distal
or subdistal dark band.

Breeding male (Fig. 1) brightly colored as described in Diag-
nosis with lateral blotches distinct and greatly expanded dorso-
ventrally.

Distribution. — Etheostoma rafincsquci occurs in the upper Green
and lower Barren River systems (Fig. 2). In the Upper Green River
system it is most common in tributaries of the Nolin River and
fairly common in the Little Barren River and Russell, Brush, Pitman,
and Goose creeks. In tributaries of the Gasper River, a tributary
of the Barren River, E. rafinesquei seems to be most common in
headwaters, particularly Wiggington Creek, Logan County. Al-
though E. rafinesquei and its sister species, E. barrenense, both
inhabit the Barren system, they maintain allopatric ranges. Except
for one upstream locality, E. rafinesquei is not known from the main
channel of the Green River.

The Green River system supports 151 species of fishes, five of
which are introduced and five of which are endemic. Two of the
endemics, E. barrcnense and Moxostoma atripinne, are restricted
to the upper Barren system; two, E. helium and E. barbouri, to the
upper Green and upper Barren systems; and one, E. rafinesquei, to
the upper Green and lower Barren systems. Two crayfishes, Barbi-

14

OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

o

Co

o
c

S

Co

6
££

o
v

a

0)
CD

3

CD
a

"as

o

T3

o

o
U

CO

w

►J

<

ca

o

c
Z

U

>y.

CM

CD

DO

in

PC
08

CD

a

CO

q cd

-t t>

■<* io

l- CO

i-H CM

cc oc

CO o
CO CM

o d

i— ( i—l

CM in

O CO

in cd

i— i i— i

co i>

^H i—l

i— i i—i

in in

CM I-

i— 1 CM

i-H CM

CD

co ©

i— 1 CM

■—I

in

OT CD
i-H i-H

t~- CO

co co

co in

co —

CO i-H

CO CM

"S *

=: C

Cr SO

Co ~ .—

SO <v> Si

£ -~ 5j

« s; ^ c

« a s Jr, m

~ ~ zz cj o

£ S — ^ — ,

O O O ~= C

"+- H-i -^ C r-

c^ c^ ^ « 7*

o o o r R 3

50 50 SO U -^

W uq tq

8

S

O
■4«a

cu

CD

c
U

i-

w

<

PC

2

CO

c
Z

>

U

Q
s.

X

CO

—

CM

CD

CO
i—i

CM

CD

CO

'o

CU

a
co

t>

in

cc co

ln

in

in ■*

CO

co

CO

co

-* i—i
co in

d

d

d d

i-H

in

O CO

!— 1

i

i—i

>-i d

i—i i—i

I-

co

CM

i — i

in in

cm in

i-H "*T

CD

^H

cm

CD

co co
i—i

cd

co

CI

^r in

CM

co co

CD CO

>

i— i

^H

CO i-H

u

^r

"tf

co in

Q

CO

l> CD

co in

CO

d

d

d d

CD

CM

q q

'X

d

i—l

i— I ^H

Z

t-

CM

1 — 1

in in

CM CO

CD

CO

in

in

rr ^h

ci = S> ;

~ g> g

g G G C

a = a r -*

~ ~ ; 1; y

: s~ a

O 3 S

Co ^D

o o

1*1 y y

THREE NEW DARTERS 15

cambarus cornutus and Orconectes barrenense, are endemic to the
upper Green and Barren systems.

Habitat and Ecology. — Etheostoma rafinesquei generally occu-
pies second-, third-, and fourth-order upland streams where it is
most common in rock- or gravel-bottomed pools. During fall and
spring it is sometimes common in riffles or their margins, especially
around emergent vegetation.

Winn (1958a, b) collected E. rafinesquei in breeding condition
from South Fork Nolin River on S April 1953, transferred them to
an aquarium, and observed spawning behavior. Stiles (1974) ob-
served spawning in streams in the springs of 1971, 1972, and 1973.
Eggs are laid singly on the sides of stones, usually with the spawn-
ing pair in a vertical position. Etheostoma rafinesquei in breeding
condition has been collected as early as mid-March (INHS 76034)
and as late as 25 May (SIUC 1020); most ripe individuals are in
April collections.

Etheostoma harrcneme Burr and Page, new species
Splendid darter

Holotype. — Illinois Natural History Survey 87491, an adult male
48.8 mm SL (Table 1) collected in Trammel Creek, tributary of
Drakes Creek, 3 km SW Halfway (36° 48'N, 86°18'W) Allen
County, Kentucky on 25 May 1981 by L. M. Page and B. M. Burr.

Paratypes. — A total of 44 specimens deposited as follows: 12 —
Illinois Natural History Survey ( INHS 87492, 32-47 mm SL, same
collection data as holotype ) ; 7 — The University of Kansas ( KU
19110, 33-43 mm SL, Trammel Creek, 6 km SSW Scottsville, Allen
County, Kentucky, 13 April 1978); 6 — University of Tennessee (UT
91.235S, 37-42 mm SL, same collection data as KU 19110); 9—
Southern Illinois University at Carbondale (SIUC 1009, 35-45 mm
SL, Indian Creek, 4.8 km E Fountain Run, Monroe Co., Kentucky,
18 March 1980 ) ; 10 — University of Florida, Florida State Museum
(UF 32483, 32-44 mm SL, same collection data as SIUC 1009).

Material examined (but not designed as types). — BARREN R.
SYSTEM: INHS 74879 (1), 75064^(10), 75468 (10), 75997 (7),
76871 (8), 77070 (2), 77514 (2), 79290 (10), 82653 (2), 84050
(3), 84057 (2); SIUC 1009 (10), 1011 (2), 1013 (2), 1010 (5);
UL 5187 (7), 5364 (10), 564S (1); UMMZ 177622 (10).

Etymology. — The name barrenense is an adjective referring to
the Barren River to which the new species is endemic. The common
name, splendid darter, alludes to the bright colors of the breeding
male and has been used by ichthyologists for this species for many
years.

Diagnosis. — Member of subgenus Nanostoma as diagnosed by
Page ( 1981 ) . Distinguished from other described members of sub-

16 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

genus by body and median fin pigmentation. Side of body with
7-10 (usually 8-9; Table 2) round to square black blotches always
confluent with a midlateral dark stripe; blotches not elongated dor-
soventrally on breeding male. Dorsum with 7-9 (usually 8) dark
blotches, the first confined to anterior half of nape. First dorsal
fin with red spot in first interradial membrane. Snout green. Breed-
ing male with bright blue anal and pelvic fins and procurrent
caudal fin membranes; red-orange breast; red on lower side, and
in a thin stripe above midlateral stripe; red distally and medially
on interradial membranes of first dorsal fin; black or green basally
on first eight membranes of first dorsal fin; second dorsal fin red-
purple medially and, on posterior half, basally. Premaxillary frenum
narrow but always present. Modally five branchiostegal rays (Table
2) and nine preoperculomandibular pores (Table 3). Breast usu-
ally unsealed (95% of specimens).

Most similar to E. rafinesquci (see Diagnosis for E. rafinesquci).
Etlieostoma duryi has broad black stripe along side but lacks a
premaxillary frenum, and the breeding male lacks red pigment
above and below black lateral stripe and bright blue in anal, caudal,
and pelvic fins.

Description. — Moderate-sized member of subgenus, maximum
SL = 53 mm for males, 47 mm for females. Snout extremely blunt;
mouth small (Table 1) and subterminal. Body proportion values
given in Table 1 and shown in Fig. 1. Infraorbital canal with 7-9
(usually S) pores (Table 3); preoperculomandibular canal with
8-10 (usually 9) pores (Table 3). Lateral line complete to end of
hypural plate (104 specimens) or missing on 1 (5), 2 (2), or 4 (1)
scales posteriorly. Genital papillae as in E. baileyi.

Breast unsealed (97 specimens), with 2 scales (2), 10% scaled
(2), or 40% scaled (1); prepectoral area scaled. Belly, nape, and
opercle fully scaled. Cheek unsealed (6) or fully scaled (96); scales
often embedded. Lateral scales 42-49 (Table 4); transverse scales
12-14 (Table 5); scales above lateral line 3 (2), 4 (99), or 5 (10);
scales below lateral line 6 (1), 7 (26), 8 (79), or 9 (5); scales
around caudal peduncle 15-18 (Table 6). Dorsal fin spines 10-12
(Table 7); dorsal fin rays 10-13 (Table 7); pectoral fin rays 13
(15), 14 (87), or 15 (10); anal fin spines 2; anal fin rays 6 (9),
7 (59), or 8 (44); branched caudal fin rays 12 (1), 13' (22), 14
(46), or 15 (42); total vertebrae 38 (11), 39 (32), or 40 (5).

Color in life: Juveniles and females yellowish with 7-10 round
to square black blotches along side always confluent with a lateral
black stripe. Blotches extend only slightly below, rarely above,
lateral stripe. Dorsum with 7-9 (usually 8) vague to dark, round
to square blotches; first blotch confined to anterior half of nape,
second under origin of first dorsal fin, third and fourth under first
dorsal fin, fifth and sixth under second dorsal fin, seventh on caudal

THREE NEW DARTERS 17

peduncle, and eighth at origin of caudal fin. Centers and edges
of body scales above lateral stripe dark. Preorbital bar usually
poorly developed, but extending to upper lip. Suborbital bar re-
duced to a dusky dash, occasionally absent. Dark dashes behind
eye on cheek and opercle. Pectoral, pelvic, and anal fins clear.
Second dorsal and caudal fins with dusky bands on rays. First
dorsal fin with red spot in first interradial membrane and occasion-
ally with red spots on margin. Snout green. Upper half of cheek
and opercle olive-green. Posterior half of body with red specks
above and below lateral stripe.

Breeding male (Fig. 1) brightly colored as described in Diag-
nosis.

Distribution. — Etheostoma barrenense is restricted to the upper
Barren River system in Kentucky and Tennessee (Fig. 2). It is
common, especially in Drakes Creek, Trammel Creek, Beaver Creek,
Peter Creek, Long Fork, Fallen Timber Creek, Bays Fork, and East
Fork Barren River. The only other fish species endemic to the
Barren River is the highly distinctive Moxostoma atripinne.

Habitat and Ecology. — Etheostoma barrenense occurs commonly
in pools, and occasionally along the margins of riffles and raceways
over limestone bedrock, clean gravel, or around large rocks.
It occupies second-, third-, and fourth-order streams.

Winn (1958a, b) collected E. barrenense in breeding condition,
made' field observations on territoriality and spawning in April 1953
and April 1954, and observed spawning in an aquarium. Stiles
(1974) observed courtship and spawning in streams in the springs
of 1971, 1972, and 1973. Spawning occurs in a vertical position;
eggs are laid on the sides of rocks. Breeding specimens are present
in collections made as early as 18 March (SIUC 1009) and as late
as 17 June (INIIS 75064) but are most prevalent in collections
made in April and May.

RELATIONSHIPS

Relationships among species of Nanostoma are difficult to de-
cipher because published morphological data on the remaining
undescribed species are scarce to nonexistent, differentiating char-
acteristics among species are predominantly color features which
are highly variable, and speciation appears to have been recent and
characteristics clearly recognizable as synapomorphies are rare.

The one clearly synapomorphous character state among Najio-
stoma is a mode of only five branchiostegal rays; a mode of six
rays in E. coosae (and in most other Etheostoma) separates it as a
sister group to other species of Nanostoma. Unfortunately a few
populations of E. zonale modally have six branchiostegal rays, con-
fusing the phylogenetic relationships of E. zonale to other species
of Nanostoma. Other character states recognized as derived within

18 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Nanostoma but also shared with species outstide the subgenus
Nanostoma are a mode of nine preoperculomandibular pores (10
in E. zonale and E. baileyi), mode of eight dorsal dark blotches
(E. zonale has six), absence of prevomerine teeth (present in E.
coosae and E. zonale), and loss of scales on the breast (variable
in E. zonale, present posteriorly in E. etnieri and E. rafinesquei) .

The loss of the premaxillary frenum is a derived state but too
much intraspecific variation occurs for the characteristic to impart
much phylogenetic information. Likewise the presence in some
species of a red spot anteriorly in the first dorsal fin is difficult to
interpret phylogenetically; in some species it is a distinct spot (E.
simoterum), but in others is expanded dorsoventrally (E. baileyi and
E. cluryi) or is part of a red band (E. eoosae and E. rafinesquei).
Other color characteristics are equally or more difficult to relate to
one another, although phylogenetic patterns may become more clear
when the remaining six or so recognized but undescribed species
are described. For now. all that is apparent is that E. eoosae, E.
zonale, and E. baileyi are early offshoots from the main line of
evolution in Nanostoma, and that all other species probably arc
more closely related to one another than to these three species.

ACKNOWLEDGMENTS

Field assistance was provided by L. E. Cordes, \Y. L. Keller,
R. L. Mayden, and S. J. Walsh, to whom we are most grateful.
R. A. Brandon, M. A. Morris, and M. E. Retzer read and improved
the manuscript. Loans of specimens were provided by M. L. War-
ren, Jr., Kentucky Nature Preserves Commission (KNPC), W. D.
Pearson, University of Louisville (UL), and R. M. Bailey, Univer-
sity of Michigan Museum of Zoology (UMMZ). Karen Schmitt of
the Scientific Photography and Illustration Facility of the Southern
Illinois University at Carbondale (SIUC) assisted in the prepara-
tion of the map. Other technical assistance was provided by B. P.
Sweeney, L. Le Mere, and L. Woodrum. all of the Illinois Natural
History Survey (INTIS).

SUMMARY

Three new species of the subgenus Nanostoma arc described.
Etheostoma baileyi inhabits the upper Cumberland and upper Ken-
tucky River systems, Kentucky and Tennessee, and is distinguished
from other darters by its pigmentation, especially the row of small
emerald green squares along the side and the red margin and
interradial red spots on the first dorsal fin. Etheostoma rafinesquei,
in the upper Green and lower Barren River systems. Kentucky, is
distinguished from other species of Nanostoma except E. zonale
by its low lateral scale count, and from all darters by the combi-

THREE NEW DARTERS 19

nation of a series of dark bars along; the side of the body, a cross-
hatched appearance created by dark (red-orange in breeding male)
spots on the posterior edges of lateral scales, and modally five
branchiostegal rays and nine preoperculomandibular pores. Etlwo-
stoma barrenense, in the upper Barren River system, Kentucky and
Tennessee, is distinguished by having modally five branchiostegal
rays and nine preoperculomandibular pores, a series of dark blotches
along the side strong])' confluent with a midlateral stripe, and, in
the breeding male, red on the lower side and in a thin stripe above
the midlateral stripe, and bright blue anal and pelvic fins.

LITERATURE CITED

Rouchard, R. W. 1977. EtJieostoma etnieri, a new percid fish from the Caney
Fork (Cumberland) River system, Tennessee, with a redescription of
the subgenus Ulocentra. Tulane Stud. Zool. Rot. 19:105-130.

Braxsox, R. A., and Batch, D. L. 1974. Fishes of the Red River drainage,
eastern Kentueky. Univ. Press of Kentueky, Lexington. 67 p.

Burr, R. M. 1980. A distributional checklist of the fishes of Kentueky. Brim-
leyana 3:53-84.

Dexoxcourt, R. F., IIocutt, C. II., and Stauffer, J. R. Jr. 1975. Exten-
sions of the known ranges of Ericymba buccata Cope and Etheostoma
zonule (Cope) in die Susquehanna River drainage. Proc. Pa. Acad. Sci.
49:45-46.

Hubrs, C. L., and Cannon, M. D. 1935. The darters of the genera Hololepis
and Villora. Misc. Publ. Univ. Mich. Mus. Zoo]. 30. 93 pp.

Hubbs, C. L., and Lagler, K. F. 1964. Fishes of the Great Lakes Region.
Univ. Mich. Press, Ann Arbor. 213 pp.

Jexkixs, R. E., Lachxer, E. A., and Schwartz, F. J. 1972. Fishes of the
central Appalachian drainages: their distribution and dispersal. In:
The distributional history of the biota of the southern Appalachians,
Part III: Vertebrates, by P. C. Holt, R. A. Paterson, and J. P. Hubbard.
Va. Polytech. Inst. State Univ. Res. Div. Monogr. 4:43-117.

Kuehxe, R. A., and Bailey, R. M. 1961. Stream capture and the distribution
of the percid fish Etheostoma sagitta, with geologic and taxonomic con-
siderations. Copeia 1961 ( 1 ) : 1-8.

Kuehxe, R. A., and Small, J. W., Jr. 1971. Etheostoma barbouri, a new
darter ( Percidae, Etheostomatini ) from the Green River with notes on
the subgenus Catonotus. Copeia 1971 ( 1 ): 18-26.

Lachxer, E. A., and Jexkixs, R. E. 1971. Systematics, distribution, and
evolution of the chub genus Nocomis Girard (Pisces, Cyprinidae) of
eastern United States, with descriptions of new species. Smithson. Con-
trib. 85. 97 p.

Pace, L. M. 1981. The genera and subgenera of darters (Percidae, Etheosto-
matini). Occ. Pap. Mus. Nat. Hist. Univ. Kan. (90):l-69.

Pfliecer, W. L. 1971. A distributional study of Missouri fishes. Publ. Mus.

Nat. Hist. Univ. Kan. 20:225-570.
Starxes, W. C, and Starxes, L. R. 1979. Taxonomic status of the percid

fish Etheostoma nigrum susanae. Copeia 1979(3) : 426-430.
Stiles, R. A. 1974. The reproductive behavior of the Green and Rarren

River Ulocentra {Ostcichthijes; Percidae: Etheostoma) . ASR Bull.

21:86-87.

20 OCCASIONAL PAPERS MUSEUM OF NATURAL HISTORY

Tsai, C, and Raney, E. C. 1974. Systematics of the banded darter, Ethco-
stoma zonule (Pisces: Percidae). Copeia 1974 ( 1 ) : 1-24.

Winn, H. E. 1958a. Observations on the reproductive habits of darters (Pisces-
Percidae). Am. Midi. Nat. 59:190-212.

Winn, H. E. 1958b. Comparative reproductive behavior and ecology of four-
teen species of darters ( Pisces- Percidae ) . Ecol. Mong. 28:155-191.

University of Kansas Publications

MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural
History, beginning with volume 1 in 1946, was discontinued with
volume 20 in 1971. Shorter research papers formerly published in the
above series are now published as Occasional Papers, Museum of
Natural History. The Miscellaneous Publications, Museum of Natural
History, began with number 1 in 1946. Longer research papers are
published in that series. Monographs of the Museum of Natural
History were initiated in 1970. All manuscripts are subject to critical
review by intra- and extramural specialists; final acceptance is at the
discretion of the publications committee.

Institutional libraries interested in exchanging publications may
obtain the Occasional Papers and Miscellaneous Publications by
addressing the Exchange Librarian, The University of Kansas Li-
brary, Lawrence, Kansas 66045. Individuals may purchase separate
numbers of all series. Prices may be obtained upon request addressed
to Publications Secretary, Museum of Natural History, The Univer-
sity of Kansas, Lawrence, Kansas 66045.

Editor: E. O. Wiley
Managing Editor: Joseph T. Collins

PRINTED BY

UNIVERSITY OF KANSAS PRINTING SERVICE

LAWRENCE, KANSAS

170 IHt

3 2044 093 361 699

DATE DUE

ARfefc-3-2004

DEMCO, INC. 38-2931