JNIVERSITY ©.
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
NATURAL HIST SURVE’

Life Histories of the Bandfin Darter, Etheostoma zonistium,
and the Firebelly Darter, Etheostoma pyrrhogaster,
in Western Kentucky

Douglas A. Carney
Brooks M. Burr

Illinois Natural History Survey Biological Notes 134
December 1989

Illinois Natural History Survey, Lorin I. Nevling, Chief
A Division of the Illinois Department of Energy and Natural Resources

Printed by authority of the State of Illinois
X02837-—2M-12-89
US ISSN 0073-490X

Cover photograph of a male Etheostoma zonistium was taken by W.N. Roston, Forsyth, Missouri.
Graphic Assistance: Molly Hardin Scott
Editors: Audrey S. Hodgins and Eva L. Steger

A catalog of the publications of the Illinois Natural History Survey is available without charge from the address below.
A price list and an order blank are included with the catalog.

Illinois Natural History Survey
Distribution Center

Natural Resources Building
607 East Peabody Drive
Champaign, Illinois 61820

Citation:
Carney, D.A., and B.M. Burr. 1989. Life histories of the bandfin darter, Etheostoma zonistium, and the firebelly darter,
Etheostoma pyrrhogaster, in western Kentucky. Illinois Natural History Survey Biological Notes 134. 16 p.

Life Histories of the Bandfin Darter, Etheostoma zonistium,
and the Firebelly Darter, Etheostoma pyrrhogaster,

in Western Kentucky

Bailey and Etnier (1988) described the bandfin darter,
Etheostoma zonistium, from Leath Creek, Chambers
Creek drainage, Hardin County, Tennessee, and the
firebelly darter, E. pyrrhogaster, from Clear Creek,
North Fork Obion River drainage, Henry County,
Tennessee. Both species are members of the subgenus
Nanostoma, which encompasses E. zonale and all species
previously referred to in the subgenus Ulocentra (Page
1981). Nanostoma includes thirteen taxonomically
described species, eight of which were described as
new species (Bouchard 1977; Page & Burr 1982; Bailey
& Etnier 1988; Etnier & Bailey 1989) or elevated from
synonomy (Etnier & Starnes 1986) in recent years. At
least five species of Nanostoma remain taxonomically
undescribed.

Substantial life history information is available for
four species of Nanostoma: Etheostoma zonale (Lachner
etal. 1950), E. coosae (O’Neil 1981), E. stmoterum (Page
& Mayden 1981), and E. baileyi (Clayton 1984). In
addition, the reproductive behavior of E. duryi (Page et
al. 1982) and E. rafinesquei and E. barrenense (Winn
1958a, b; Stiles 1974) has been described. Other
publications discuss fecundity and spawning substrate
(Weddle 1989), distributions (Bouchard 1972, 1973,
1974; Kuehne 1972; Fallo & Warren 1982; Burr &
Warren 1986), interspecific interactions (Greenburg
1984, 1985), food habits (Adamson & Wissing 1977;
Cordes & Page 1980), respiration and habitat selection
(Ultsch et al. 1978), and evolutionary trends in repro-
duction, morphology, and ecology (Page & Swofford
1984; Page 1985) of species of Nanostoma. Finally,
Kuehne and Barbour (1983) and Page (1983) provide
summaries of the natural history of members of
Nanostoma.

This paper was submitted in its original form by Douglas A.
Carney in partial fulfillment of the requirements for the
degree of Master of Arts in the Graduate School of Southern
Illinois University at Carbondale. Mr. Carney was formerly a

Technical Research Biologist with the Illinois Natural History

Survey and is now a Resource Planner with the Illinois
Department of Conservation. Dr. Brooks M. Burr is a
Professor in the Department of Zoology at Southern Illinois
University at Carbondale and an Affiliate of the Survey.

Douglas A. Carney and Brooks M. Burr

Etheostoma zonistium occurs in lower western tribu-
taries of the Tennessee River in western Kentucky and
Tennessee, barely extending into northern Mississippi
and Alabama, and in the Hatchie River drainage in
Tennessee and the Black Warrior River drainage in
Alabama (Fig. 1). Etheostoma pyrrhogaster occurs in
tributaries of the Mississippi River in extreme south-
western Kentucky and in western Tennessee (Fig. 1).
In Tennessee (Starnes & Etnier 1980) and in Kentucky
(Warren et al. 1986), E. pyrrhogaster is regarded as a
species of special concern because of its restricted
distribution and the loss of habitat resulting from the
channelization of streams.

Methods

Observations and minnow-seine (1.2 m x 3.1 m, 3.8-
mm mesh) collections were made at approximately
monthly intervals from 5 January to 13 November
1983. Additional observations and collections were
made during spawning periods in 1983 and 1984. Data
collected at West Fork Clarks River and at Terrapin
Creek on each sampling date included stream condi-
tions, darter habitat, and darter behavior. Water
velocity in darter habitat was measured during spawn-
ing periods with a Teledyne-Gurley Pygmy current
meter.

Fishes captured with the two darter species and the
habitats in which these captures occurred were quanti-
fied in a series of three monthly samples (May-July
1983). Unit-effort sampling consisted of five seining
efforts in each of four subjectively identified habitat
types:

Riffles, defined as shallow areas with swift current
and surface turbulence over gravel and sand substrate.

Raceways, defined as moderately shallow areas with
swift current and little to no surface turbulence over
gravel and sand substrate.

Stream margin zones, defined as areas of moderate
depth and current next to a bank (often undercut)
and primarily over sand (sometimes sand with large
stones or gravel).

Pools, defined as deep areas with sluggish current
over sand and detritus.

2 Illinois Natural History Survey Biological Notes

Kick-sets were used to capture fish in all habitat
types except pools, where seine hauls were made. All
captured fishes in each habitat type were identified
and enumerated.

Unit-effort sampling comprised 60 seining efforts
per study site (all habitat types combined). To facili-
tate discussion, an arbitrary rating of darter abundance
at each study site was established for quantified
sampling efforts:

Abundant, at least 60 individuals of a given species
captured.

Common, 12 to 59 individuals.

Uncommon, 6 to 11 individuals.

Rare, fewer than 6 individuals.

Specimens were preserved in 10 percent formalin
and stored in 70 percent ethanol. Totals of 336 E.
zonistium from West Fork Clarks River and 164 E.
pyrrhogaster from Terrapin Creek were preserved and
examined. Additional specimens from the ichthyologi-
cal collection at Southern Illinois University at Carbon-
dale were examined, and all darters collected during
the study were deposited at that facility.

All specimens were sexed, measured, and aged.
Sexing was by examination of the genital papillae or,
when necessary, by examination of the gonads. Meas-
urements were to the nearest 0.1 mm standard length
(SL). Age-to-year class was determined by counting
annuli on scales removed from the dorsum near the
junction of the first and second dorsal fins. Year-classes
were corroborated by analysis of length-frequency
distribution and number of annual rings on otoliths
(sagittae). Otoliths were removed by making an
incision from the breast to the sacculus and bending
the head back until the sagittae were exposed. Whole
sagittae were immersed in glycerine, and annuli were
counted with the aid of a fiber-optic light and dissect-
ing microscope. Age to month was determined by
using April, the month of greatest spawning activity, as
month zero.

Seasonal variation in the reproductive conditions
of males and females was determined by the gonadoso-
matic index (GSI). Gonad weight and adjusted body
weight (weight of the specimen minus gonads, stom-
ach, intestine, and liver) of alcohol-stored specimens
(blotted dry) were measured to the nearest 0.001 gm
on an analytical balance.

Ovaries of preserved specimens were dissected, and
oocytes were selected at random to create a subset of
total ova for diameter measurements. All fertilized
eggs recovered from aquaria were measured. Fertilized
eggs were incubated in an aerated malachite green
solution. The larval terminology used was that of
Snyder (1976). Diameters of ova and lengths of larvae
and genital papillae were measured to the nearest

No. 134

0.01 mm with an ocular micrometer. Ovum diameter
was calculated as the mean of three measurements.

Spawning behavior was observed in the laboratory
in 71-liter (30 cm high x 25 cm wide x 91 cm long)
aquaria. Adults in spawning condition were given a
choice of all known egg deposition sites used by other
species of darters (Page 1983:165): vertical rock,
vertical plant, raised horizontal rock, and a substrate of
mixed gravel and sand.

Regression coefficients were calculated by the
method of least squares using the Systems Regression
procedure of the Statistical Analysis System (SAS).
Correlations were Pearson product-moments. In
statistical tests, differences of P<0.05 were considered
significant.

Study Areas

The site selected for study of Etheostoma zonistium was
West Fork Clarks River (Tennessee River drainage) at
the Kentucky Highway 121 bridge, 0.6 km east of
Coldwater, Calloway County, Kentucky (Fig. 2). At this
site, the stream is of moderate gradient and consists of
shallow riffles with gravel and sand substrate alternat-
ing with deep, sand-bottomed pools. Widths of 2 to 6
m and depths to 1.0 m are characteristic of the study
area during normal flow. Water is generally clear but
becomes turbid during flooding. Agricultural lands are
immediately adjacent to parts of the stream; however, a
narrow margin of deciduous trees lines most of the
study site. During 1983 monthly water temperature
ranged from 8.5°C in January to 25°C in July.

Nineteen fish species were collected from West
Fork Clarks River. Species taken, in addition to those
captured in the stream margins during unit-effort
sampling (Table 1), were Notropis lutrensis, N. umbra-
tilis, N. whipplei, Phenacobius mirabilis, Moxostoma

se

28 eae S g:
y

89° 87° 85° 83°

Figure 1. Ranges of Etheostoma zonistium (circles) and E.
pyrrhogaster (triangles). Life history study sites are enclosed by
open circles. Map modified from Bailey and Etnier (1988).

December 1989

erythrurum, Ameiurus natalis, Noturus nocturnus, Lepomis
cyanellus, and Percina maculata.

Etheostoma pyrrhogaster was studied in Terrapin
Creek (Obion River drainage) just north (upstream)
of the Tennessee Highway 69 bridge at the
Kentucky—Tennessee border, Graves County, Kentucky
(Fig. 3). Terrapin Creek has a low-to-moderate gradi-
ent and consists of sand- and gravel-bottomed riffles
alternating with sand-bottomed pools. Widths of 2 to 5
m and depths to 1.5 m are characteristic of Terrapin
Creek at normal flow. The water, as in West Fork
Clarks River, is turbid only during flooding. The study
area is immediately bordered by deciduous forest;
however, agricultural land is nearby. During 1983,
monthly water temperatures ranged from 8°C in
January to 20°C in June.

On 27 March and 18 April 1983, E. pyrrhogaster was
taken for aquarium spawning observation from a
backwater slough of Terrapin Creek, 0.8 km south of
Bell City, Graves County, Kentucky. The slough was

Figure 2. Study area for Etheostoma zonistium, West Fork
Clarks River, Calloway County, Kentucky, 18 April 1983.

Figure 3. Study area for Etheostoma pyrrhogaster, Terrapin
Creek, Graves County, Kentucky, 22 July 1983.

Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky 3

1 m wide with depths to 30 cm and had water tempera-
tures of 14°C and 17°C on the above sampling dates,
respectively. A silt substrate and abundant aquatic
vegetation were present in the slough, which was
surrounded by deciduous forest.

Fishes collected in Terrapin Creek and the back-
water slough during this study, in addition to those
taken during unit-effort sampling of stream margins
and pools (Table 2), include Umbra limi, Notropis
fumeus, Pimephales promelas, Moxostoma poecilurum,
Noturus hildebrandi, Fundulus olivaceus, Elassoma zo-
natum, Aphredoderus sayanus, Etheostoma chlorosomum, E.
gracile, and E. parvipinne. Other fishes reported from
the study area by Walsh and Burr (1981) but not taken
during this study were Esox americanus, Notemigonus

Table 1. Relative abundance of fishes captured with Etheo-
stoma zonistium in West Fork Clarks River, expressed as
percent of individuals (N= 167) captured by unit-effort
seining of stream margin habitat in May, June, and July 1983.

Species Percentage
Etheostoma zonistium 44.3
Campostoma oligolepis 22.2
Pimephales notatus 17.4
Semotilus atromaculatus 7.8
Hypentelium nigricans 2.4
Fundulus olivaceus 1.8
Erimyzon oblongus L.2
Etheostoma flabellare U2,
Etheostoma neopterum 1.2
Micropterus punctulatus 0.6

Table 2. Relative abundance of fishes captured with Etheo-
stoma pyrrhogaster in Terrapin Creek, expressed as percent of
individuals (N= 221) captured by unit-effort seining of
stream margin and pool habitat in May, June, and July 1983.

Species Percentage

Etheostoma nigrum S157
Semotilus atromaculatus Ded
Etheostoma lynceum 16.3
Notropis camurus 6.3
Etheostoma pyrrhogaster 6.3
La mpetra aepyplera 5.4
Phenacohius mirabilis 3.6
Percina sciera 3.2
Noturus phaeus 1.0
Gambusia affinis 1.0
Lepomis marginatus 1.0
Etheostoma swaini 1.0
Erimyzon oblongus 0.5
Minytrema melanops 0.5
Lepomis cyanellus 0.5
Lepomis macrochirus 0.5

4 Illinois Natural History Survey Biological Notes

crysoleucas, Notropis umbratilis, Ameiurus natalis, A. melas,
Lepomis gulosus, and L. megalotis. A total of 34 fish
species are known to occur in the Terrapin Creek
study area.

Habitat and Associated Species

In West Fork Clarks River, Etheostoma zonistium was
frequently collected over a substrate of sand or gravel
and sand near the stream margin at depths of 15 to 75
cm and was often found in emergent soft rush (Juncus
effusus). Unit-effort seining by habitat type confirmed
the observation that E. zonistiwm typically inhabited the
stream margin (Fig. 4). The species was abundant in
West Fork Clarks River as was E. flabellare; however, the
two were weakly associated due to the propensity of E.
flabellare for riffles (Fig. 4). Analysis of a 2 x 4 contin-
gency table of these species versus habitats by the log
likelihood ratio test (Sokal & Rohlf 1981) indicated a
significant association (G= 148.65, P<0.001) between
species and habitat. Etheostoma neopterum was rare at the
study site but was found with E. zonzstium in the stream
margin (Fig. 4), a habitat commonly noted for E.
neopterum (Howell & Dingerkus 1978; Braasch &
Mayden 1985). Fishes captured in greatest numbers
with E. zonistium were Campostoma oligolepis, Pimephales
notatus, and Semotilus atromaculatus (Table 1).

In Terrapin Creek, E. pyrrhogaster typically inhab-
ited the stream margin at depths of 30 cm to 1 m but
also occurred in deeper pools, often among exposed
tree roots in undercut banks over sand. Unit-effort
sampling indicated an affinity of individuals (N= 8) for
stream margin habitats; however, a number of indi-
viduals (N= 6) were also recovered from pools (Fig. 5).
Etheostoma pyrrhogaster was classified as uncommon in
Terrapin Creek with 18 individuals captured during
the unit-effort seining (Fig. 5). Although too few
individuals were captured to warrant statistical analysis
of the association between species and habitat, the
data represent a reliable estimate of the habitat affinity
of E. pyrrhogaster in Terrapin Creek as judged from
nonquantitative observations during the course of the
study. Fishes collected in greatest numbers with E.
pyrrhogaster were E. nigrum, Semotilus atromaculatus, and
E. lynceum (Table 2, Fig. 5).

Page and Mayden (1981) described characteristic
Nanostoma habitat as clean pools with moderate
current over bedrock, cobble, or gravel. The relatively
fine substrates over which E. zonisttum and E. pyrrhogas-
ter were taken, in comparison with other members of
Nanostoma, were indicative of their lowland stream
habitats.

Reproductive Cycle of the Male
Male Etheostoma zonistium and E. pyrrhogaster developed
brilliant nuptial coloration as described by Bailey and
Etnier (1988). Color intensification began as early as

TOTAL NUMBER OF DARTERS

TOTAL NUMBER OF DARTERS
>
3°

No. 134

October in individuals over 1 year of age. Two-year-old
males of both species collected in October were dark
red above the lateral line and orange-red ventrolater-
ally. At this time ventrolateral coloration extended
from the origin of the pelvic fin to the origin of the
caudal fin in EF. zonistium and from the origin of the
pelvic fin to midway through the caudal fin in E.
pyrrhogaster. By late March ventrolateral body colora-
tion was bright red midway through the caudal fin in
E. zonistium and throughout all but the margins of the
caudal fin in EF. pyrrhogaster. Commencement of
spawning activity in aquaria was accompanied by a
change in color of the chin and breast from blue-gray
to lavender in E. zonistiwm and from gray-green to
green in E. pyrrhogaster. By mid-July nuptial coloration
in field-collected individuals had faded considerably;
however, males of both species retained some red on
the body and dorsal fins throughout the year.

(J Etheostoma zonistium
Etheostoma flabellare
BB Etheostoma neopterum

STREAM
MARGIN

RIFFLE POOL

Figure 4. Number of darters captured in unit-effort seining
by habitat type, West Fork Clarks River, Calloway County,
Kentucky, May-July 1983.

(Etheostoma pyrrhogaster
Etheostoma lynceum

HBEtheostoma nigrum
EEJEtheostoma swaini

40

30

20

XM W000 q| NWA

'Y
Y Z
RACEWAY STREAM
MARGIN
Figure 5. Number of darters captured in unit-effort seining
by habitat type, Terrapin Creek, Graves County, Kentucky,

May-July 1983.

POOL

RIFFLE

December 1989

The genital papilla of male E. zonistiwm was a white
to cream, somewhat flattened, broad-based structure
that terminated in a short tube (Fig. 6A, B). Mean
papilla length was 0.8 mm (range 0.5-1.1, sp = 0.15)
in 32 males that ranged in SL from 32 to 50 (x = 40.8)
mm. Genital papillae of male E. pyrrhogaster were

identical in structure and color to those of E. zonistium.

Mean papilla length was 0.7 mm (range 0.5-1.0, sp =
0.11) in 29 males that ranged in SL from 32.8 to 49.7
(x= 41.2) mm.

For both species, size and weight of testes gradually
increased as the spawning season approached. For
male E. zonistium, the relationship between weight of
the testes x 1000/adjusted body weight (Y) and the
month (X), with October = 1 and May = 8, was log Y=
0.152 + 0.323, with r= 0.932 (Fig. 7). In male E.
pyrrhogaster, the relationship, with October = 1 and
June = 9, was log Y= 0.391 + 0.275, with r= 0.852
(Fig. 8).

All 1-year-old males collected in April (N= 11 E.
zonistium, N= 9 E. pyrrhogaster) had enlarged testes and
were potential spawners. The proportionally largest
testes in E. zonistium equaled 1.6 percent of the ad-
justed body weight and were found in a 3l-mm SL
male collected on 18 April 1983. The proportionally

Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky

or

largest testes in E. pyrrhogaster equaled 1.4 percent of
the adjusted body weight and were taken from a 37-
mm SL male also collected on 18 April 1983.

Reproductive Cycle of the Female

Coloration of females showed no seasonal variation.
Bright colors in female Etheostoma zonistium included a
red ocellus in the first interradial membrane and a red
band in the distal one-half of the spinous dorsal fin.
Female E. pyrrhogaster also had a red ocellus; in some
individuals, a submedian red band was apparent in the
second dorsal fin.

The genital papillae of female E. zonistium (Fig. 6C,
D) and E. pyrrhogaster were thick, tubular structures
white to cream in color. In E. zonistium, mean papilla
length was 1.2 mm (range 0.8-1.7, sp = 0.20) in 47
females ranging in SL from 29.4 to 45.0 (x= 41.2) mm.
Mean papilla length of female E. pyrrhogaster was 1.3
mm (range 1.0—1.6, sp = 0.18) in 31 individuals
ranging in SL from 30.8 to 44.5 (x= 39.4) mm. As
females became distended with maturing ova, the
genital papillae shifted in position from lying against
the abdomen to an angle varying from acute to nearly
perpendicular to the abdominal wall. This change in
position was accompanied by a change in the distal

Figure 6. Genital papillae of Etheostoma zonistium from West Fork Clarks River, 18 April 1983: A) breeding male 43 mm SL,
lateral view; B) ventral view of A; C) breeding female 33 mm SL, lateral view; D) ventral view of C.

6 Illinois Natural History Survey Biological Notes

end of the papilla from somewhat rounded to flat-
tened, as the diameter of the gonopore increased.
Increased abdominal girth and change in position of
the genital papillae were first evident in late February
for females of both species.

Small white ova appeared in females of both
species in November. Intermediate-sized yellow ova
were present in each species in February, and by late
March mature ova were apparent. Mature ova were
translucent with a pitlike indentation. This indenta-
tion, also present in other darters (Page 1983:162), has
recently been shown to serve as a micropyle (J. M.
Grady, pers. comm.).

Mean diameter of white ova in November-collected
E. zonistium was 0.24 mm (range 0.17—0.34, sp = 0.062,
N= 20). January-collected E. pyrrhogaster had white ova
with a mean diameter of 0.26 mm (range 0.15—0.41, sp
= 0.068, N= 20). Mean diameters of yellow ova in
March-collected females were 0.78 mm (range
0.50—0.99, sp = 0.180, N= 20) in E. zonistium and 0.68
mm (range 0.54—0.86, sp = 0.098, N= 20) in E.
pyrrhogaster. Mean diameters of mature ova in March-
collected females were 1.08 mm (range 1.01-1.17, sp =
0.038, N= 20) in E. zonistium and 1.09 mm (range
1.02-1.18, sp = 0.048, N= 20) in E. pyrrhogaster.

5005 LOG Y=1.728+0.431X FEMALES r =0.974

¢

mes r=0.932

—
=
©
uu
=
>
(a)
Q 100
Q
=
w 50
=
sy 8
S °
o
°o . e
= ®

10
2 Sate
z fe}
e) 5 ° 8
© 20)
rs) 6 pers 6
—

4

5 8“ Log y=0.152+0.323x MALES 8
rr
=

MONTH

Figure 7. Monthly variation in gonad weight relative to
adjusted body weight in Etheostoma zonistium. Dots represent
individual females; circles represent individual males. Data
points for June and July were not included in the analysis but
are presented to illustrate the decline in relative gonad
weights.

1
OCT NOV DEC JAN FEB MAR APR MAY JUN JUL

No. 134

Counts of yellow and mature ova in 1-year-old E.
zonistium ranged from 41 to 137 (x= 77.0, sp = 25.17,
N=9) (Table 3). In contrast, females of 1-year-old E.
pyrrhogaster contained from 0 to 84 (x= 28.4, sp =
23.85, N= 9) yellow and mature ova (Table 4). Similar
counts of 2- to 3-year-old females ranged from 98 to
155 (x= 127.5, sp = 20.25, N= 4) in E. zonistium and
from 115 to 197 (x= 144.3, sp = 37.32, N= 3) in E.
pyrrhogaster.

Etheostoma simoterum females 12 months old con-
tained 110 to 240 mature (orange or translucent) ova
(Page & Mayden 1981). Total egg complements
ranged from 288 to 496 (x= 382.8) in E. coosae (O'Neil
1981) and 228 to 864 (x= 481.2) in E. bailey: (Clayton
1984). Winn (1958b) estimated that 1- and 2-year-old
female E. barrenense lay an average of 364 and 798 eggs
per spawning season, respectively. His estimates are
based on the premise that all ova present in the ovaries
were spawned in a season.

Some darters spawn more than one egg clutch per
year (Hubbs 1985), as demonstrated for E. (Boleosoma)
olmstedi (Gale & Deutsch 1985) and E. rafinesquez
(Weddle 1989). Both species were shown to spawn
over a period of several weeks at intervals ranging from

3 to 16 days (Gale & Deutsch 1985; Weddle 1989). One

500 LOG Y =1.995+0.355X FEMALES
r =0.919

—

ee

Oo

w

=

>

Qa

fe)

2 100

[@)

Ww

a

> 50 ‘
{a}

<

S H
Oo

en 8
x<

a 10

<x e
z

fe)

Oo 5

uw

fe)

is

) LOG Y =0.391+0.275X MALES 8
w

=

r=0.852

OCT NOV DEC JAN FEB MAR APR MAY JUN JUL
MONTH

Figure 8. Monthly variation in gonad weight relative to
adjusted body weight in Etheostoma pyrrhogaster. Dots repre-
sent individual females; circles represent individual males.
Data points for July were not included in the analysis but are
presented to illustrate the decline in relative gonad weight.

~I

December 1989 Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky

female E. rafinesquei spawned 13 clutches, a total of 780 1.728 + 0.431X, r= 0.974 (Fig. 7). The proportionally

eggs (Weddle 1989). These studies also suggest that largest ovaries (equaling 26.3% of the adjusted body
mature ova are recruited throughout the season and weight) were found in a 34.9-mm SL female collected
that darters collected early or late in the season would on 27 March 1983. In E. pyrrhogaster females the
contain lower numbers of mature ova than those relationship, with October = | and June = 9, was log Y
collected during the peak spawning period. If E. zonis- = 1.995 + 0.355X, r= 0.919 (Fig. 8). The proportionally
tiumand E. pyrrhogaster also produce multiple clutches, largest ovaries (equaling 19.4% of adjusted body
our counts of yellow and mature ova do not approxi- weight) were found in a 31.8-mm SL female collected
mate seasonal fecundity; however, they may represent on 18 April 1983.
two estimates of successive clutch sizes. We agree with All female E. zonistium and most female E.
Gale and Deutsch (1985) and Weddle (1989) that pyrrhogaster examined were sexually mature by the end
fecundity estimates based upon single-day ova counts of their first year of life. Individuals that failed to reach
may grossly underestimate seasonal fecundity. maturity within | year were female E. pyrrhogaster less
In E. zonistium the relationship between month than 27 mm SL (Table 4). Samples of E. pyrrhogaster
(X), with October = 1 and May = 8, and weight of taken on 27 March, 18 April, and 17 May 1983 yielded
ovaries X 1000/adjusted body weight (Y) was log Y= 29 first-year females, 7 (24%) of which were immature.

Table 3. Size, age, ovary weight, and number of ova from female Etheostoma zonistium collected on 27 March 1983.

Standard Adjusted body Age Ovary weight Number of Number of Number of yellow
length weight (g)* (months) (g) yellow ova mature ova and mature ova
26.5 0.200 11 0.028 38 20 58
26.6 0.181 ll 0.019 4) 0 41
29.0 0.264 11 0.047 74 0 74
30.9 0.312 11 0.062 72 55 137
see 0.302 11 0.048 60 0 60
32.4 0.409 11 0.067 77 0 77
34.9 0.514 11 0.135 0 78 78
35.1 0.482 11 0.061 79 0) 79
36.8 0.541 11 0.093 89 0 89
42.1 0.841 23 0.175 20 135 155
42.3 0.932 23 0.146 14 117 131
42.8 0.995 23 0.134 98 0 98
45.0 1.072 35 0.138 126 0 126
% 35.) 0.543 16 0.089 61 31 93

“Weight of the specimen minus gonads, stomach, intestine, and liver.

Table 4. Size, age, ovary weight, and number of ova from female Etheostoma pyrrhogaster collected on 27 March 1983.

Standard Adjusted body Age Ovary weight Number of Number of Number of yellow

length weight (g)* (months) (g) yellow ova mature ova and mature ova
24.5 0.137 11 0.001 0 0 0
25.6 0.175 1] 0.003 0 0 0
26.1 0.168 11 0.002 0 0 0
27.2 0.199 1] 0.018 27 0 27
28.3 0.243 11 0.011 9 0 9
29.0 0.247 1] 0.012 23 0 23
29.8 0.312 11 0.042 37 24 61
30.7 0.342 11 0.044 52 0 52
31.5 0.275 11 0.055 63 21 84
42.6 0.901 23 0.154 115 0 115
43.7 1.000 23 0.153 95 102 197
44.5 1.125 35 0.150 68 53 121
x 29.5 0.402 14 0.050 38 15 53

“Weight of the specimen with gonads, stomach, intestine, and liver removed.

8 Illinois Natural History Survey Biological Notes

Spawning
Observations suggested that both species began
spawning in March and continued to May or June
(Tables 3 & 4; Figs. 7 & 8). Spawning appeared to peak
in March and April, with declines in the gonadoso-
matic index thereafter (Figs. 7 & 8). Low values for
female Etheostoma zonistium collected in June (range
18-48, x= 30.5, N= 6) and males (range 1-9, x= 4.2,
N= 6) (Fig. 7) suggested little spawning potential
during that month. Female E. pyrrhogaster collected in
June maintained relatively higher values (Fig. 8) and
large yellow ova, but the four specimens captured
lacked mature ova.

Water temperatures in both West Fork Clarks River
and Terrapin Creek ranged from 11°C on 27 March to
17°C on 17 May 1983. On 18 June 1983, water tem-
perature was 21°C in West Fork Clarks River and 20°C
in Terrapin Creek.

Spawning occurs primarily in April among other
species of Nanostoma with distributions limited to the
southeastern United States; for example, E. barrenense
and E. rafinesquei (Stiles 1974; Page & Burr 1982), E.
coosae (O’Neil 1981), and E. simoterum (Page & Mayden
1981). May was reported to be the most active month
of spawning for E. baileyi (Clayton 1984). The spawning
period for E. zonale is May and June in Pennsylvania
(Lachner et al. 1950) and April and May in Missouri
(Pflieger 1975:312).

Etheostoma zonistium was removed from the study
site on 27 March 1983 (N= 6) and 12 April 1984 (N=
8) for observation of spawning in aquaria. Stream
temperatures were 11°C and 14°C on those dates, re-
spectively. Darters in breeding condition were col-
lected near the stream margin at depths of 38 to 46 cm
over sand and gravel. Current speed ranged from 0.26
to 0.32 m/sec at sites of capture. Ripe individuals were
most common among emergent vegetation, tree roots,
and brush piles. Attempts to observe spawning in the
stream were unsuccessful.

Etheostoma pyrrhogaster was removed from the back-
water slough on 18 April 1983 (N= 5) and from
Terrapin Creek proper on 12 April 1984 (N= 8) for
observation of spawning in aquaria. Stream tempera-
ture was 14°C on both dates. Darter collection sites in
Terrapin Creek on 12 April 1984 had moderate flow
(0.31 to 0.40 m/sec) with depths of 27 to 38 cm over
sand and stones. Etheostoma pyrrhogaster was captured
primarily near the bank among tree roots and/or large
stones (10 to 20 cm in diameter). The presence of ripe
adults in the backwater slough (see Study Areas for
habitat characteristics) suggests that this area may have
been used for spawning.

Spawning E. zonistiwm were observed in aquaria
from 28 March to 1 April 1983 and from 14 to 17 April

No. 134

1984; water temperature ranged from 17 to 22°C.
Individuals of E. pyrrhogaster were injected with human
chorionic gonadotropin (approximately 50 IU/fish)
on 20 April 1983 after it appeared that an increase in
water temperature was not sufficient to induce spawn-
ing in captivity. Subsequent to hormone injections,
these darters spawned in aquaria on 21 April 1983 ata
water temperature of 22°C. Individuals collected in
April 1984 did not spawn in the aquarium before or
after injection.

The spawning behaviors of E. zonistiwm and E.
pyrrhogaster were virtually identical. A male approached
a female and followed her, crossing over her dorsum
in a zigzag pattern. The male often rubbed his pectoral
fin, and occasionally his chin, on the female’s nape.
The male then followed the female to an egg deposi-
tion site. Once the site was selected, the male assumed
an S-shaped posture over the female with his caudal
peduncle placed adjacent to hers. While in this
position, they vibrated rapidly and a single egg was
deposited and fertilized. During egg deposition, the
female had her mouth open. Open-mouth behavior
was previously noted in spawning Etheostoma proeliare
(Burr & Page 1978) and E. asprigene (Cummings et al.
1984). A period of inactivity (approximately 30 sec)
followed each spawning act. Eggs were occasionally
eaten by the parents. Spawning darters were promiscu-
ous; for example, a male spawned with several females,
and a female spawned with several males. Agonistic
encounters between rival males occurred during and
just prior to spawning. An area of approximately 15 cm
in diameter was defended by males from intrusion by
rivals. The largest males were generally dominant and
spawned more frequently. Females congregated in a
loosely knit group when not engaged in spawning.
Occasionally a female would leave the group and swim
in front of a male that was following another female.
The male then followed the intruding female and
spawned with her.

Frequency of spawning acts by males of each
species was recorded during peak spawning activity. In
observations of two rival males of E. zonistium, a 47-mm
SL male spawned an average of once every 3 min and a
48.7-mm SL male spawned an average of once every
2.1 min over a 30-min period. During this time 11
aggressive encounters occurred between the two
males, one encounter every 2.7 min. Similarly, a male
E. pyrrhogaster (49.7 mm SL) was noted to spawn about
every 2.5 min during 30 min of observation.

Etheostoma zonistium spawned vertically on aquarium
corners and walls and on vertical rocks and plants
(Ceratophyllum demersum). Eggs deposited on rocks were
generally placed in small crevices. The species spawned
horizontally on sand and gravel substrate (Fig. 9) and

December 1989

on the top and the underside of raised horizontal
rocks. The two most frequent spawning/egg deposi-
tion sites of EF. zonistium were vertical corners of the
aquarium and sand and gravel substrate.

Etheostoma pyrrhogaster spawned vertically on aquar-
ium corners and on rocks. The most frequent spawn-
ing site observed in the aquarium was sand and gravel
substrate at the aquarium wall interface. They spawned
in a horizontal position at this site.

On 14 April 1984, an attempt was made to induce
interspecific spawning between E. zonistiwm and E.
pyrrhogaster. Male E. pyrrhogaster (N= 2) were placed
with female E. zonistium (N= 5) in an aquarium.
Conversely, male E. zonistium (N= 2) and female E.
pyrrhogaster (N= 2) were combined. No spawning
attempts were observed between male E. pyrrhogaster
and female E. zonistium. One spawning attempt was
observed within 1 hour of introduction between a male
E. zonistium and a female E. pyrrhogaster; however, egg
deposition either did not occur or the egg was quickly
ingested by one of the parents. Behavior during this
interspecific spawning act did not deviate from the
general sequences described earlier. Agonistic behay-
ior between males and the courting of females contin-
ued for 4 hours, but females were unreceptive. All
darters in the interspecific spawning experiment were
subsequently injected with human chorionic gona-
dotropin, but no further breeding behavior was
observed.

Winn (1958a) observed spawning between a male
E. barrenense and a female E. rafinesquei that resulted in
fertilized eggs. Males of these two species of Nanostoma
display dissimilar dorsal fin banding patterns (Page &
Burr 1982), and these patterns are believed to be a
factor in sex and/or species recognition (Page 1983:
180). Winn (1958b) concluded that characteristic
movements are important in sex recognition by darters
during spawning. Dorsal fin patterns of E. zonistium
and E. pyrrhogaster are dissimilar yet interspecific

Figure 9. Spawning Etheostoma zonistium collected 27 March
1983 from West Fork Clarks River: horizontal position,

Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky 9

spawning was attempted, presumably because of the
similar behavior patterns of these species.

In aquaria, E. zonistium and E. pyrrhogaster attached
eggs to a site and left them unattended; that is, they
were egg attachers as defined by Page (1983:165, 167;
1985). All other members of Nanostoma for which
reproductive behaviors have been reported are also
egg attachers: EF. zonale (Pflieger 1975:312; Trautman
1957:663), E. simoterum (Page & Mayden 1981), E.
rafinesquei and FE. barrenense (Winn 1958a, b), E. coosae
(O’Neil 1981), and E. duryi (Page et al. 1982). Hori-
zontal spawning on the substrate was observed in E.
pyrrhogaster and to a lesser extent in E. zonistium. When
spawning on the substrate, these species placed eggs
on the surface of the gravel, a behavior not associated
with typical egg-buriers. Members of Nanostoma have
most frequently been observed spawning in a vertical
postion on the sides of rocks; for example, E. simoterum
(Page & Mayden 1981), FE. rafinesquei, E. barrenense
(Winn 1958a, b; Page & Burr 1982), and E. duryi (Page
et al. 1982). A female E. simoterwm occasionally buried
eggs in gravel but not as deeply as more typical egg-
burying species (Page & Mayden 1981). Etheostoma
coosae spawns at angles from horizontal to vertical and
in an inverted position on most inclined, creviced
surfaces (O’Neil 1981). The high incidence of horizon-
tal substrate spawning observed in E. zonistium and E.
pyrrhogaster may be related to the constraints of aquar-
ium confinement (Winn 1958b) or to their lowland
stream habitat, which offers little or no bedrock and
few large rock surfaces.

Aggressive behavior among male E. zonistium and E.
pyrrhogaster was not considered a strict expression of
territoriality because males occasionally changed
locations in the aquaria and actively defended these
“new” spaces. This behavior accorded with the defini-
tion of Stiles (1972), who regarded the aggressive
defense of space surrounding an individual fish, no
matter where that fish was located, as maintenance of
individual space. Winn (1958b) referred to “moving
territories,” a term that apparently described the same
phenomenon.

Based on field and aquaria observations, Winn
(1958b) described territories of breeding male EF.
barrenense and E. rafinesquei as 50 to over 100 cmin
diameter, centered around a large rock and apparently
stationary. Field observations of these two species by
Stiles (1974) indicated that breeding males were highly
aggressive toward other males that invaded their
individual space but were not territorial in the strict
sense. Page and Mayden (1981) stated that breeding E.
simoterum males held in aquaria did not defend station-
ary territories but were combative toward one another
when in close proximity.

10 Illinois Natural History Survey Biological Notes

Development and Growth

Eggs were fertilized and larvae were reared in aquaria.
Fertilized eggs of Etheostoma zonistium and E. pyrrhogas-
ter were spherical, translucent, demersal, and adhesive.
A yellow, acentric oil droplet was present in eggs of
each species.

Fertilized eggs of E. zonistium averaged 1.7 mm in
diameter (range 1.5-2.1, sp = 0.13, N= 26). Eggs
incubated at 20 + 2°C hatched in 163 to 170 hr (6.8 to
7.1 days). Only three embryos survived to hatching.
Total lengths (TL) of two measurable E. zonistium
protolarvae were 4.33 mm and 4.25 mm. Both speci-
mens had 15 preanal myomeres and 21 postanal
myomeres.

Fertilized ova of E. pyrrhogaster averaged 1.6 mm in
diameter (range 1.4-1.7, sp = 0.12, N= 13). Eggs
incubated at 25 + 3°C hatched in 146 to 190 hrs (6.1 to
7.9 days). Protolarvae averaged 4.08 mm TL (range
3.39—4.55, sp = 0.243, N= 7). Myomere counts were as
follows: 15 preanal, 21 postanal (N= 5), 15 preanal, 20
postanal (N= 1), and 16 preanal, 25 postanal (N= 1).

Page (1983:168-169) presented a regression of egg
incubation period on temperature for all darters on
which this information was available. Egg incubation in
E. zonistium appeared to conform to Page’s (1983:169)
regression line; however, eggs of E. pyrrhogaster re-
quired an unusually long incubation time in relation
to temperature.

Myomere counts were similar for E. zonistium (4.33
and 4.25 mm TL), and E. pyrrhogaster (3.39 to 4.55 mm
TL): modally 15 preanal and 21 postanal. Among
other members of Nanostoma, counts are available only

Figure 10. Protolarva of Etheostoma zonistium, the product of
laboratory spawning by individuals collected in West Fork
Clarks River: A) protolarva, 4.3 mm TL, age 170 hr, lateral
view; B) ventral view of A; C) dorsal view of A.

No. 134

for E. zonale, which had 14 to 15 preanal and 22
postanal myomeres at 5 to 7 mm TL (Auer 1982).

Protolarvae of E. zonistiwm and E. pyrrhogaster (Figs.
10 & 11) had the following characteristics in common:
mouth subterminal, well developed, and overlapped by
subocular tissue (lateral view); snout rounded; eyes
darkly pigmented (iris, black; pupil, olive); head
sparsely pigmented dorsally; antero-ventral yolk sac
with large oil droplet; single row of melanophores on
each side of ventral midline (along length of anal fin);
single row of melanophores ventrally along anal tube;
and yolk sac heavily pigmented ventrally and lightly
pigmented laterally (lower half only).

Protolarvae of E. zonisttwm (N= 2) (Fig. 10) lacked
two to five melanophores on the lateral body region
that were present in protolarvae of E. pyrrhogaster (N=
7) (Fig. 11). The mouth terminated on a horizontal
plane even with the bottom of the pupil in E. pyrrhogas-
ter and below the pupil in E. zonistium.

Growth analyses are based on specimens collected
in the field throughout the study. Males of both
species appeared to be larger than females of the same
age, but growth curves were not calculated by sex due
to the paucity of 2- and 3-year-old males. The relation-
ship between standard length (Y) and age in months
(X) for E. zonistium (N= 336) was Y= 4.43 + 11.75 log
X, with r= 0.848 (Fig. 12). At 12 months, males aver-
aged 33.8 mm SL and were significantly (t= 3.359,

P< 0.01, df= 29) longer than females, which averaged
31.3 mm SL. The largest specimens collected from
West Fork Clarks River were a 50.4-mm SL male and a
46.7-mm SL female, both taken on 5 January 1983. The
ichthyological collection at Southern Illinois University

Figure 11. Protolarva of Etheostoma pyrrhogaster, the product
of laboratory spawning by individuals collected in Terrapin
Creek: A) protolarva, 4.0 mm TL, age 190 hr, lateral view; B)
ventral view of A; C) dorsal view of A.

STANDARD LENGTH (mm)

STANDARD LENGTH (mm)

December 1989

at Carbondale contains two larger males (51.6 and 53.0
mm SL) taken from Panther Creek, Graves County,
Kentucky, on 16 June 1979. The oldest EF. zonistiwm
taken from the study area were two males and two
females 35 months of age, captured on 27 March 1983.
The males measured 48.0 and 49.7 mm SL and the
females were 44.8 and 45.0 mm SL.

For E. pyrrhogaster (N= 164), the relationship
between standard length (Y ) and age in months (X)
was Y= 0.114 + 13.01 log X, with r= 0.861 (Fig. 13). At
12 months, males averaged 35.7 mm SL and were
significantly (t= 3.394, P< 0.01, df= 16) longer than
females, which averaged 32.2 mm SL. The largest and
oldest male, collected from a Terrapin Creek back-
water slough on 18 April 1983, was 36 months old and
measured 49.7 mm SL. The two largest females (both
44.5 mm SL) were taken from Terrapin Creek on 5
January and 27 March 1983. The female collected in
March was also the oldest at 35 months of age.

One-year-old males of each species were signifi-
cantly longer than females at the end of their first year.
Growth curves indicated that E. zonistiwm (Fig. 12) was
slightly longer at 6 months of age than E. pyrrhogaster
(Fig. 13) but had a comparatively slower growth rate
thereafter.

wn
on

Y=4.43+11.75 LOG X 2
r=0.848

a
oO

a
v1)

ny
wn

Zea 6) (OG 02. 14.16) 18) 2072224926 128) (30) 132) 134° 36

MONTHS OF AGE

Figure 12. Size distribution by age of Etheostoma zonistium
collected in West Fork Clarks River, Calloway County,
Kentucky, between 5 January and 13 November 1983. Dots
represent sample means for males; circles represent sample
means for females. Regression line is for both sexes. A total
of 336 specimens is represented.

55

50] ¥ =0.114+13.01 LOG x
45 r=0.861

2 4 6 8 10 12 14 16 18 20 22 24 26 28 30 32 34 36

MONTHS OF AGE

Figure 13. Size distribution by age of Etheostoma pyrrhogaster
collected in Terrapin Creek, Graves County, Kentucky,
between 5 January and 13 November 1983. Dots represent
sample means for males; circles represent sample means for
females. Regression line is for both sexes. A total of 164
specimens is represented.

Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky 1]

Composition and Survival

Of 336 Etheostoma zonistium collected in West Fork
Clarks River between 5 January and 13 November
1983, 58.6 percent were up to 1| year of age, 34.8
percent were over | and up to 2 years, and 6.6 percent
were over 2 years (Table 5). The overall sex ratio was
1.7 females to 1 male (y2= 23.04, P<0.001). Darters 1
year old or less (N= 197) were present at a female-to-
male ratio of 1.4:1 (y?= 4.88, P<0.05). Those 1+ years
of age (N= 117) exhibited a 2.44:1 ratio of females to
males (x? = 20.52, P<0.001). At 2+ years (N= 22), E.
zonistium was collected at a ratio of 2.14:1, but the ratio
was insignificant, presumably a result of small sample
size.

Of 164 E. pyrrhogaster collected in the Terrapin
Creek drainage between 5 January and 13 November
1983, 59.8 percent were up to | year of age, 26.8
percent were over | and up to 2 years, and 13.4
percent were over 2 years (Table 5). The overall sex
ratio was 1.3 females to 1 male, a nonsignificant
deviation from 1:1. The only age class to show signifi-
cantly different proportions between the sexes were 1+
darters (N= 44), which yielded 3.9 females to 1 male
(y? = 15.36, P< 0.001).

The overall sex ratio of EF. zonistium deviated
significantly from 1:1 but that of EF. pyrrhogaster did
not. Significant deviations did occur in 1+ year classes
of both species, and in each case of significantly
skewed sex ratios, females outnumbered males. In
general, however, darter populations do not deviate
from a 1:1 sex ratio (Page 1983:169). Exceptions to
this generality occur among highly territorial members
of the subgenus Catonotus, for example, Etheostoma
flabellare (Lake 1936), E. squamiceps (Page 1974), E.
kennicotti (Page 1975), E. smithi (Page & Burr 1976),
and E. olivaceum (Page 1980). Selection pressure for a
sex ratio favoring females in E. squamiceps and E.
kennicotti was presumably a result of the dominance of
a proportionately few large males during spawning

Table 5. Distribution of sexes and age groups in samples of
Etheostoma zonistium and E. pyrrhogaster collected between
5 January and 13 November 1983.

Number by age group

Sex a | 1+ 2+ Total
E. zonistium

Male 83 34 7 124
Female 114 83 5 212
Total 197 117 22 336

E. pyrrhogaster
Male 52 9 12 73
Female 46 35 10 9]
Total 98 44 22 164

12 Illinois Natural History Survey Biological Notes

(Page 1974, 1975). Females significantly outnumbered
males in the 1+ year class of E. coosae (O’Neil 1981)
and in E. simoterum overall (Page & Mayden 1981). An
E. baileyi population did not deviate significantly from
a 1:1 sex ratio except for the 3+ year class, in which
males outnumbered females (Clayton 1984). Males of
E. zonale outnumbered females in their second and
third years in western Pennsylvania (Lachner et al.
1950). Four of the six species of Nanostoma for which
life histories have been completed indicate tendencies
for sex ratios favoring females, particularly in the 1+
year class. If we assume no sampling bias, males may
experience higher mortality in their first year. Both
sexes may be more susceptible to large predators than
shallow riffle-inhabiting darters, but predation on
small, brightly colored males may be greater than
predation on females and such differential predation
may result in sex ratios skewed towards females.
Relative survival values for the 124 male and 212
female EF. zonistium collected in West Fork Clarks River
showed that 40.5 percent of the first-year males and
72.8 percent of the first-year females survived into their
second year. Only 20.6 percent of second-year males
and 18.1 percent of second-year females survived into
their third year. Of the total sample of 336 E. zonistium,
59.4 percent of first-year individuals survived into their
second year and 18.8 percent into their third year.

No. 134

Relative survival values for the 73 male and 91
female E. pyrrhogaster from the Terrapin Creek drain-
age showed that 17.3 percent of the first-year males
and 76.1 percent of the first-year females survived to
their second year. More third-year males (N= 12) than
second-year males (N= 9) were taken during the study;
however, data suggested that only 28.6 percent of the
second-year females survived until their third year. Of
the total sample of 164 E. pyrrhogaster, 44.9 percent of
first-year individuals survived into their second year
and 50 percent into their third year.

Diet
Stomach contents of 89 Etheostoma zonistium and 81
E. pyrrhogaster were tabulated by month of collection
(Tables 6 & 7). The diets of these species, similar to
those of most darters (Page 1983:171), were composed
primarily of microcrustaceans and aquatic insects.
Dipteran larvae were reported to be a predominant
food of the closely related darters E. zonale (Forbes &
Richardson 1920; Adamson & Wissing 1977; Cordes &
Page 1980; Wynes & Wissing 1982), E. coosae (O'Neil
1981), and E. simoterum (Page & Mayden 1981).
Predominant food items of both E. zonistiwm and
E. pyrrhogaster were larval chironomids, and these were
consumed in greatest abundance in May, June, and
July. The diet of E. pyrrhogaster included ephemerop-

Table 6. Stomach contents of Etheostoma zonistium from West Fork Clarks River by month of collection. Mean number of food
organisms per stomach is followed parenthetically by percentage of stomachs in which food organisms occurred.

March
N= 10

Jan Feb

Food organism N=10 N=11

Gastropoda — = = =

Arachnida
Acarina — = —

Crustacea
Cladocera — — — -—

Ostracoda _ a — 0.3 (27)

Copepoda — — 0.2 (20) —=
Eubranchiopoda

Conchostraca — — _ —

Insecta
Plecoptera 0.1 (10) — — —
Ephemeroptera = 0.2 (18)
Trichoptera — — = =

Diptera
Tipulidae _— — — =
Simuliidae —_ 0.2 (18) —_ —
Chironomidae 8.5 (50) — 23.8 (73)
Ceratopogonidae = — —
Empididae — _
Unknown pupae =

Teleostomi eggs —_ _— —

Empty —

April
N=11

0.1 (09)

11.5 (73)

May June July Oct Nov
N=9 N=10 N=9 N=8 N=11
— _ a 0.4 (13) —
_ 0.1 (10) _- 1.9 (50) 0.1 (9)
0.1 (11) 0.2 (20) — == =
0.7 (33) 0.6 (33) 0.3 (25) _
_— — _ 0.2 (13) —

— — —- 0.1(13) 0.4 (18)
— 0.1 (10) 0.1 (11) — —
= 0.2 (10) _— = on
Ose) 2.1 (20) 3.4 (11) — 0.1 (9)
31.3 (89) 109.1 (100) 31.1 (100) 9.1 (88) 2.6 (45)
= 0.6 (30) 0.3 (11) — _
— —_ 0.2 (11) — —
0.3 (22) 1.0 (50) 1.7 (56) 0.9 (38) --
1.2 (33) 0.3 (20) _ _ —
(9) = = = (59)

December 1989

teran nymphs and trichopteran larvae throughout the
year; however, E. zonistiwm fed on these larval insects

less consistently. Empty stomachs were encountered in
individuals of both species between January and April.

Predation
Potential predators captured in West Fork Clarks River
were examined for darter remains in the stomach and
gut: three Micropterus punctulatus, two Lepomis cyanellus,

one L. marginatus, three L. megalotis, and two L. gulosus.

At Terrapin Creek one Aphredoderus sayanus, three
Lepomis cyanellus, one L. macrochirus, and two L. margi-
natus were examined. No evidence of fish predation
was found.

Parasitism

None of the 89 Etheostoma zonistium stomachs exam-
ined contained parasites. One of 81 E. pyrrhogaster
stomachs contained Leptorhynchoides thecatus (Acanth-
ocephala) (April). Of 336 E. zonistium examined for
external parasites, 7 harbored cysts thought to be

trematode metacercaria (2 in January and 5 in March).

The only external parasite on the 164 E. pyrrhogaster
examined was a leech from a specimen collected in

January.

Summary

Major aspects of the life history of Etheostoma zonistium
in West Fork Clarks River and E. pyrrhogaster in Terra-
pin Creek are summarized in Table 8.

Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky 13

Acknowledgments

For assistance in the field we are grateful to Alan L.
Balliett, James M. Grady, Bernard R. Kuhajda, Richard
W. Liedtke, Curt Means, John Mellen, Richard J.
Packauskas, Thomas E. Shepard, Melvin L. Warren, Jr.,
and Lisa A. Whiteside. J.M. Grady and B.R. Kuhajda
kindly provided assistance and advice on various
aspects of this study. Margaret Krzyzanowska of South-
ern Illinois University at Carbondale (SIUC), Depart-
ment of Educational Psychology, assisted with statistical
analyses. Genital papillae drawings were prepared by
J-M. Grady and larval drawings by Kevin S. Cummings.
Victor Santucci assisted in the counting of myomeres.
David F. Oetinger (Kentucky Wesleyan College)
identified parasites. Karen A. Schmitt and Karen L.
Fiorino of the Scientific Photography and Illustration
Facility, SIUC, assisted in the preparation of figures.
Dr. William N. Roston graciously provided the slide
used on the cover. Drafts of the manuscript were typed
by Betsy Lewis of SIUC and Mary Lou Williamson of
the Illinois Natural History Survey (INHS). Roy C.
Heidinger, John E. McPherson, Jr., Lawrence M. Page
(INHS), Melvin L. Warren, Jr. (SIUC), Gordon K.
Weddle (Campbellsville College), and two anonymous
reviewers provided helpful comments on the manu-
script. To all of these people we are sincerely grateful.

Table 7. Stomach contents of Etheostoma pyrrhogaster from Terrapin Creek drainage by month of collection. Mean number of
food organisms per stomach is followed parenthetically by percentage of stomachs in which food organisms occurred.

Jan Feb
N=14 N=11

March

Food organism N= 10

Turbellaria _— —_ —
Gastropoda — = i a=

Arachnida
Acarina

Collembola

Crustacea
Cladocera — ~~ — —
Ostracoda — a a =
Copepoda _ _ 0.7 (40)

Insecta
Plecoptera —
Ephemeroptera 0.1 (7)
Trichoptera 0.1 (7)

0.2 (18)
0.1 (9)
0.2 (18)

0.1 (10) —

0.1 (10) —_
Diptera

Simuliidae — — —
Chironomidae 7.9(71) 20.5 (100) 15.6 (90)
Ceratopogonidae — — _ —
Unknown pupae — 0.2 (9) 0.4 (30) —

Empty (21) — (10) (13)

April
N=8

0.3 (13)

0.5 (50)

0.3 (30) 0.1 (13)

13.0 (88) 31.1 (100)

May June July Oct Nov
N=7 N=4 N=9 N=8 N=10
0.7 (43) — — 0.4 (30) —
0.1 (14) _ ~ a 0.3 (30)
_ 0.3 (25) = _ 0.1 (10)
0.7 (43) — 0.1 (11) — 1.3 (60)

0.1 (14) — ws =, i2
0.1 (14) = 0.9 (56) 0.9 (25) 0.2 (10)
0.3 (29) 0.5 (25) 1.0 (44) 0.1 (13) 6.6 (100)

0.2 (11) = —
89.1 (100) 15.3 (100) 11.4 (90)
O.1 (11) _— _—
2.1 (78) 1.9 (25) 0.2 (20)

88.8 (100)

0.1 (14) 0.8 (50)

14

Illinois Natural History Survey Biological Notes

No. 134

Table 8. Summary of life history information on Etheostoma zonistium in West Fork Clarks River and E. pyrrhogasterin Terrapin

Creek.

Characteristic

Principal habitat

Age at sexual maturity
Size at sexual maturity

Sexual dimorphism

Breeding tubercles
Description of genital papillae

Number of mature ova in
preserved females

Description of mature ova
Description of fertilized eggs
Spawning period

Spawning habitat
Egg deposition sites in aquaria

Spawning position

Egg guarding
Incubation period
Size at hatching

Influence of sex on growth rate

Sex ratio
Longevity
Maximum size
Migrations
Territoriality

Principal diet

Literature Cited

Adamson, S.W., and T.E. Wissing. 1977. Food habits and
feeding periodicity of the rainbow, fantail, and banded
darters in Four Mile Creek. Ohio Journal of Science

77:164-169.

Etheostoma zonisttum
Stream margins with sand, gravel, and
emergent vegetation
1 year

Females average 31 mm; males average
34mm

Males are larger than females and brightly
colored

Absent

Long and tubular in females, short and
flattened in males

20-135

1.1 mm in diameter, translucent with a
pitlike indentation

1.7 mm in diameter, spherical, translucent,
demersal, and adhesive

From March into May; water temperature
11-17°C
Unknown in nature

Aquarium walls, sand and gravel substrate,
rocks, and plants

Male and female in vertical or horizontal
position

None observed

163-170 hrs at 20 + 2°C

4.3 mm TL

At | year, males significantly larger
than females

1.7 females :1 male
35 months

50 mm SL

None observed
None observed

Immature aquatic insects and
microcrustaceans

Etheostoma pyrrhogaster
Stream margins with undercut banks and
exposed tree roots over sand
1 or 2 years

Females average 32 mm; males average
36 mm

Males are larger than females and brightly
colored

Absent

Long and tubular in females, short and
flattened in males

21-102

1.1 mm in diameter, translucent with a
pitlike indentation

1.6 mm in diameter, spherical, translucent,
demersal, and adhesive

From March into June; water temperature
11-20°C

Unknown in nature

Sand and gravel substrate, aquarium
corners, and rocks

Male and female in vertical or horizontal
position

None observed

146-190 hrs at 25 + 3°C

3.4-4.6 mm TL

At 1 year, males significantly larger
than females

1.2 females :1 male
36 months

50 mm SL

None observed
None observed

Immature aquatic insects and
microcrustaceans

Bouchard, R.W. 1972. A new darter of the subgenus Ulocentra
from the Caney Fork River system in Tennessee (Osteich-
thyes, Perciformes, Percidae). Association of Southeast-
ern Biologists Bulletin 18:56.

Bouchard, R.W. 1973. A preliminary survey of the distribu-

tion of fishes in the Caney Fork River system, Tennessee.

Auer, N.A. 1982. Family Percidae, perches. Pages 581-648 in
N.A. Auer, ed. Identification of larval fishes of the Great
Lakes basin with emphasis on the Lake Michigan
drainage. Great Lakes Fishery Commission, Ann Arbor,
Michigan. Special Publication 82-3. 744 p.

Bailey, R.M., and D.A. Etnier. 1988. Comments on the
subgenera of darters (Percidae) with descriptions of two
new species of Etheostoma (Ulocentra) from southeastern
United States. University of Michigan Museum of
Zoology Miscellaneous Publications 175. 48 p.

Association of Southeastern Biologists Bulletin 20:40.

Bouchard, R.W. 1974. The subgenus Ulocentra (Percidae,
Etheostomatini) in western Kentucky, western Tennes-
see, and northern Mississippi. Association of Southeast-
ern Biologists Bulletin 21:41.

Bouchard, R.W. 1977. Etheostoma etnieri, a new percid fish
from the Caney Fork (Cumberland) River system,
Tennessee, with a redescription of the subgenus Ulocen-
tra. Tulane Studies in Zoology and Botany 19:105-130.

December 1989

Braasch, M.E., and R.L. Mayden. 1985. Review of the
subgenus Catonotus (Percidae) with descriptions of two
new darters of the Etheostoma squamiceps species group.
University of Kansas Museum of Natural History Occa-
sional Papers 119. 83 p.

Burr, B.M., and L.M. Page. 1978. The life history of the
cypress darter, Etheostoma proeliare, in Max Creek, Illinois.
Illinois Natural History Survey Biological Notes 106. 15 p.

Burr, B.M., and M. L. Warren, Jr. 1986. A distributional atlas
of Kentucky fishes. Kentucky Nature Preserves Commis-
sion Scientific and Technical Series 4. 398 p.

Clayton, J.M. 1984. Population differences and life history of
the emerald darter, Etheostoma baileyi (Pisces: Percidae).
MSS. Thesis, University of Kentucky, Lexington. 95 p.

Cordes, L.E., and L.M. Page. 1980. Feeding chronology and
diet composition of two darters (Percidae) in the
Iroquois River system, Illinois. American Midland
Naturalist 104:202-206.

Cummings, K.S., J.M. Grady, and B.M. Burr. 1984. The life
history of the mud darter, Etheostoma asprigene, in Lake
Creek, Illinois. Illinois Natural History Survey Biological
Notes 122. 16 p.

Etnier, D.A., and R.M. Bailey. 1989. Etheostoma (Ulocentra)
flavum, a new darter from Tennessee and Cumberland
river drainages. University of Michigan Museum of
Zoology Occasional Papers 717. 24 p.

Etnier, D.A., and W.C. Starnes. 1986. Etheostoma lynceum
removed from the synonomy of E. zonale (Pisces, Per-
cidae). Copeia 1986:832-836.

Fallo, G.J., and M.L. Warren, Jr. 1982. Distribution and
habitat of Etheostoma atripinne in Kentucky. Kentucky
Academy of Science Transactions 43:131-136.

Forbes, S.A., and R.E. Richardson. 1920. The fishes of
Illinois. 2nd ed. Illinois Natural History Survey. cxxxvi +
357 p.

Gale, W.F., and W.G. Deutsch. 1985. Fecundity and spawning
frequency of captive tessellated darters—fractional
spawners. American Fisheries Society Transactions
114:220-229.

Greenburg, L.A. 1984. Habitat utilization by two species of
stream fishes. American Society of Zoologists 24:128A.

Greenburg, L.A. 1985. Distributional ecology of two species
of darters. Ecological Society of America Bulletin 66:182.

Howell, W.M., and G. Dingerkus. 1978. Etheostoma neopterum,
a new percid fish from the Tennessee River system in
Alabama and Tennessee. Alabama Museum of Natural
History Bulletin 3:13-26.

Hubbs, C. 1985. Darter reproductive seasons. Copeia
1985:56-68.

Kuehne, R.A. 1972. Appearance and distribution of the
named and unnamed snubnose darters (Ulocentra),
Etheostoma, Percidae, Osteichthyes). Association of
Southeastern Biologists Bulletin 19:80.

Kuehne, R.A., and R.W. Barbour. 1983. The American

darters. The University Press of Kentucky [Lexington].
173 p.

Carney and Burr: Bandfin and Firebelly Darters in Western Kentucky 15

Lachner, E.A., E.F. Westlake, and P.S. Handwerk. 1950.
Studies on the biology of some percid fishes from
western Pennsylvania. American Midland Naturalist
43:92-111.

Lake, C.T. 1936. The life history of the fan-tailed darter
Catonotus flabellanis flabellaris (Rafinesque). American
Midland Naturalist 17:816-830.

O'Neil, P.E. 1981. Life history of Etheostoma coosae (Pisces:
Percidae) in Barbaree Creek, Alabama. Tulane Studies in
Zoology and Botany 23:75-83.

Page, L.M. 1974. The life history of the spottail darter,
Etheostoma squamiceps, in Big Creek, Illinois, and Ferguson
Creek, Kentucky. Illinois Natural History Survey Biologi-
cal Notes 89. 20 p.

Page, L.M. 1975. The life history of the stripetail darter,
Etheostoma kennicotti, in Big Creek, Illinois. Illinois Natural
History Survey Biological Notes 93. 15 p.

Page, L.M. 1980. The life histories of Etheostoma olivaceum and
Etheostoma striatulum, two species of darters in central
Tennessee. Illinois Natural History Survey Biological
Notes 113. 14 p.

Page, L.M. 1981. The genera and subgenera of darters
(Percidae, Etheostomatini). University of Kansas
Museum of Natural History Occasional Papers 90. 69 p.

Page, L.M. 1983. Handbook of darters. T.F.H. Publications,
Inc., Neptune City, New Jersey. 271 p.

Page, L.M. 1985. Evolution of reproductive behaviors in
percid fishes. Ilinois Natural History Survey Bulletin
33:275-295.

Page, L.M., and B.M. Burr. 1976. The life history of the
slabrock darter, Etheostoma smith, in Ferguson Creek,
Kentucky. Illinois Natural History Survey Biological
Notes 99. 12 p.

Page, L.M., and B.M. Burr. 1982. Three new species of
darters (Percidae, Etheostoma) of the subgenus Nanostoma
from Kentucky and Tennessee. University of Kansas
Museum of Natural History Occasional Papers 101. 20 p.

Page, L.M., and R. L. Mayden. 1981. The life history of the
Tennessee snubnose darter, Etheostoma simoterum, in
Brush Creek, Tennessee. Illinois Natural History Survey
Biological Notes 117. 11 p.

Page, L.M., M.E. Retzer, and R.A. Stiles. 1982. Spawning
behavior in seven species of darters (Pisces: Percidae).
Brimleyana 8:135-143.

Page, L.M., and D.L. Swofford. 1984. Morphological corre-
lates of ecological specialization in darters. Environ-
mental Biology of Fishes 11:139-159.

Pflieger, W.L. 1975. The fishes of Missouri. Missouri Depart-
ment of Conservation [Jefferson City]. 343 p.

Snyder, D.E. 1976. Terminologies for intervals of larval fish
development. Pages 41-58 in J. Boreman, ed. Great
Lakes fish egg and larvae identification. U.S. Fish and
Wildlife Service National Power Plant Team, Ann Arbor,
Michigan. 220 p.

Sokal, R.R., and F. J.Rohlf. 1981. Biometry. 2nd ed. W.H.
Freeman and Company, San Francisco. 859 p.

16 Illinois Natural History Survey Biological Notes

Starnes, W.C., and D.A. Etnier. 1980. Pages B-1—-B-134 in D.C.
Eager and R.M. Hatcher, eds. Tennessee’s rare wildlife.
Vol. 1, The vertebrates. Tennessee Department of Con-
servation and Tennessee Wildlife Resources Agency.

Stiles, R.A. 1972. The comparative ecology of three species of
Nothonotus (Percidae - Etheostoma) in Tennessee’s Little
River. Ph.D. Dissertation, University of Tennessee,
Knoxville. 97 p.

Stiles, R.A. 1974. The reproductive behavior of the Green
and Barren River Ulocentra (Osteichthyes: Percidae:
Etheostoma). Association of Southeastern Biologists
Bulletin 21:86-87.

Trautman, M.B. 1957. The fishes of Ohio. Ohio State
University Press [Columbus]. 683 p.

Ultsch, G.R., H. Boschung, and M.J. Ross. 1978. Metabolism,
critical oxygen tension, and habitat selection in darters
(Etheostoma). Ecology 59:99-107.

Walsh, S.J., and B.M. Burr. 1981. Distribution, morphology
and life history of the least brook lamprey, Lampetra
aepyptera (Pisces: Petromyzontidae), in Kentucky.
Brimleyana 6:83-100.

No. 134

Warren, M.L., Jr., W.H. Davis, R.R. Hannan, M. Evans, D.L.
Batch, B.D. Anderson, B. Palmer-Ball, Jr., J.R. Mac-
Gregor, R.R. Cicerello, R. Athey, B.A. Branson, GJ. Fallo,
B.M. Burr, M.E. Medley, and J.M. Baskin. 1986. Endan-
gered, threatened, and rare plants and animals of
Kentucky. Kentucky Academy of Science Transactions
47:83-98.

Weddle, G.K. 1989. Fecundity and spawning substrate
preferences of captive Kentucky snubnose darters
(Etheostoma rafinesquet) , multiple spawners. Kentucky
Academy of Science Transactions 50:127.

Winn, H.E. 1958a. Observations on the reproductive habits
of darters (Pisces: Percidae). American Midland Natural-
ist 59:190-212.

Winn, H.E.. 1958b. Comparative reproductive behavior and
ecology of fourteen species of darters (Pisces: Percidae).
Ecological Monographs 28:155-191.

Wynes, D.L., and T.E. Wissing. 1982. Resource sharing
among darters in an Ohio stream. American Midland
Naturalist 107:294-304.

Manuscripts of high quality dealing with any aspect of natural history will be considered for publication in one of the Illinois
Natural History Survey series: Bulletin, Biological Notes, Circular, and Special Publications. Authors who are not employees of
the Survey are required to pay printing costs. Manuscripts should follow the recommendations of the third edition of the Council
of Biology Editors Style Manual except that journal names in literature cited are to be spelled in full. The Survey expects to
publish only one or two manuscripts by non-Survey authors yearly. Send three copies of manuscripts to be considered for publi-
cation to Office of the Chief, Illinois Natural History Survey, 607 East Peabody Drive, Champaign, Illinois 61820. Before a
manuscript is accepted for publication in the Bulletin or in Biological Notes, two or more outside referees must recommend it.

(hires Pine. a) paginas net . i

AT” fost Vs. Gaye Ory i 4 J

aye lewdAisie aAyd 7
: i

i!

| 7 > .
———— ee ————EEEEeEeeEeEEEeEeEeeeerereoorr

a

Tarehadeda
edad
rt

scptetet

jee
tne

Ditheats
ete
jieted tse

ee
aed
i

Peiiehie tube t ete
Braet s

ayiet

phabwsbet ic htecaniam
Tee ttt
i

Penta
‘eto br este

etl ieethseaeiae
eu he 4

i
pene
atari
Hvicetch

PAT cemhiage
Pym tyemae i?

iaea bl tes
riya izkl

wabare

1 Wigedt

En,
Hiatal

sberatebeiead
the

SE eekret

iv 28
rf reer ewL
Hace ytrey ney

re

f

Falylateages
ve
Peevey

fet

lhl)

era

ue

Bitte
Ae ey

¢
fy

it

*
6

ve
te

“a i siya ti ad tivanery fee
Miter 3

r Anny at
mete tet ee eh
id wie

t o
ay

bay
they . nl iinet:

Spite

iis

Tepematen woe
etree

Te ore

Ue

‘ 1
SONU ty ete
OMT Seat
Erith)
oe

velrytee
Wey

ge

on
ME Cr ln ne
: ra

Sy epepees

os

“Le ot urbe hrePLrLp

a Y