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2010 SOUTHEASTERN NATURALIST 9(1):119–128
Life-history Aspects of Fundulus stellifer (Southern
Studfish) (Actinopterygii: Fundulidae) in Northern Georgia
Kerstin L. Edberg1 and Steven L. Powers2,*
Abstract - Fundulus stellifer (Southern Studfish) was studied using 12 monthly
collections from Moore’s Creek near Waleska, GA. Female specimens ranged from
24.72 mm to 94.29 mm SL and up to 15.90 g total weight. Male specimens ranged
from 27.81 mm to 96.07 mm SL and up to18.60 g total weight. Increases in size were
greatest in spring and summer. Maximum age appears to be just over 2 years for both
sexes, as the oldest specimens collected were estimated at 28 months of age. Spawning
likely occurred between late April and early June. Sexual maturity was reached
in the second full spawning season for both sexes, with females having a maximum
ova diameter of 2.75 mm. The primary diet of F. stellifer consisted of Chironomidae,
Branchiopoda, and unidentified insect parts, with other food items seasonally. The
amount of food consumed is greatest in the months preceding and during spawning
and lowest during fall and early winter.
Fundulus stellifer Jordan (Southern Studfish) is endemic to the Mobile
Basin (Boschung and Mayden 2004), a member of the subgenus Xenisma,
and most closely related to Fundulus catenatus Storer (Northern Studfish)
and Fundulus bifax Cashner et al. (Stippled Studfish) (Ghedotti et al. 2004).
Fundulus stellifer is relatively abundant within its range, which includes
small to medium streams in the Coosa and Alabama River systems of
northeast Georgia and central to south Alabama. It is easily recognized by
the superior placement of the mouth, random placement of brown to dark
red spots on the side of the body, and black pigmentation along the distal
margin of the caudal fin, a characteristic seen only in sexually mature males
(K.L. Edberg, pers. observ.; Thomerson 1969). There is little information
regarding the biology of F. stellifer except for a diet that may consist of
snails and other mollusks, as suggested by the structure of the pharyngeal
teeth of adults (Thomerson 1969). The spawning season is reported to occur
between April and May, as indicated by the capture of nuptial males during
these months (Boschung and Mayden 2004). The primary objective of this
study was to investigate aspects of the life history of F. stellifer.
Fundulus stellifer were collected from Moore Creek, just upstream
of its confl uence with Shoal Creek (34.3240°N, 84.5636°W), outside of
Waleska in Cherokee County, GA (Fig. 1). Moore Creek is characterized as a
1Department of Biology, Western Kentucky University, Bowling Green, KY 42101.
2Biology Department, Roanoke College, Salem, VA 24153. *Corresponding author -
120 Southeastern Naturalist Vol. 9, No. 1
second-order tributary to the Etowah River, with wetted widths ranging from
3.1 m to 6.4 m and depths <1.0 m at base-fl ow water levels. The substrate of
Moore Creek is primarily gravel to cobble with sporadic bedrock in riffl es,
gravel to sand in runs, and silt and sand in pools. Fundulus stellifer were
collected primarily from backwaters with heavy vegetation cover. Upstream
of the study area, the Moore Creek watershed is mostly forested with moderate
agricultural use and sparse residential development. Water temperatures
during the study period ranged from 5 °C in November 2006 to 25 °C in
June and August 2006. Species richness of fishes within the study area was
34 species from nine families collected between 2004 and 2007. A complete
list of species collected from Moore Creek near its Shoal Creek confl uence
can be found in O’Kelley and Powers (2007).
Fundulus stellifer were collected using a Smith-Root model 24 backpack
electrofisher and a 3.3-m x 1.3-m seine with 9.5 mm mesh during twelve
mid-monthly surveys from February 2006 to January 2007. A total of 240
specimens was collected, preserved in 10% formalin in the field, rinsed with
water after two weeks, and transferred into 70% ethyl alcohol (EtOH) for
long-term storage in the University of Alabama Ichthyological Collection
(UAIC 15028–15039). Standard length (SL) of each specimen was measured
using digital calipers and recorded to the nearest 0.01 mm. Specimens were
blotted dry, and their total weight (TW), eviscerated weight (EW) and gonad
weight (GW) were measured using a digital analytical scale and recorded
to the nearest 0.001 g. Sexual size dimorphism was not detected using a
Figure 1. Map of
study area in Moore
Waleska in Cherokee
2010 K.L. Edberg and S.L. Powers 121
two-sample t-test of SL (t = 1.3, P = 0.18); therefore, statistical analyses
were performed without regard to sex, except for those involving GSI, in
which sexes were treated separately. All statistical analyses were carried out
using Data Desk 6.0 (Data Description, Inc., Ithaca, NY) with alpha values
for all tests set at 0.05. Standard deviation is abbreviated as SD.
To illustrate age and growth patterns, SL was plotted against month.
Gaps of >10 mm SL in specimens from a given month were considered
indicative of age groups (e.g., for October all specimens were 29.35–44.79,
64.56–75.58, and 90.7–93.52 mm SL, with each cluster lacking gaps <10
mm) (Fig. 2). Due to the high GSI values seen in April and May, spawning
is assumed to occur in May for estimating the age of individuals. Specimens
<12 months of age were counted as age 0+, specimens between 12 and 23
months were counted as age 1+, specimens >24 months were counted as age
2+. Regressions by least sum of squares were performed for SL and EW to
describe length-weight relationships of F. stellifer.
The anterior one-third of the gastrointestinal track was opened and the
contents were removed, weighed to the nearest 0.001 g on a digital analytical
scale, and placed in 70% EtOH for storage. Empty guts were recorded with
a weight of zero. One-way analysis of variance (ANOVA) was performed on
the weight of gut contents among different months to test for differences in
amount of food consumed throughout the year. Taxa richness of gut contents
was the total number of different food items in each specimen. An ANOVA
was performed on the taxa richness of gut contents per month to test for
dietary changes throughout the year. To test for infl uence of body size on
the amount of food consumed, gut content weight was regressed with SL by
least sum of squares. Taxa richness of gut contents was also regressed with
SL to test for an infl uence of size on food items. Food items were enumerated
and identified to the lowest possible taxonomic category following Thorp
and Covich (1991) and Merritt and Cummins (1996). Due to mastication by
the pharyngeal teeth, food items were often unidentifiable past the level of
family, order, or class. Presence/absence of sand, detritus, and unidentifiable
insect parts were recorded. Niche breadth (NB) was calculated following
Levins (1969) to compare diet of F. stellifer to that of Hypentelium etowanum
Jordan (Alabama Hog Sucker) and Notropis xaenocephalus (Jordan)
(Coosa Shiner), two syntopic species for which diet has been documented
within the study area. Values approaching 1.0 are considered generalists,
while smaller values indicate a more specialist diet.
Reproductive condition was investigated by calculating gonadosomatic
index (GSI = GW/EW). A one-way ANOVA was performed to test mean
differences in GSI among months for both males and females. In gravid
females, greatly enlarged (≈2 mm in diameter), fully yolked, mature oocytes
were counted, and five representatives were measured using digital calipers
to provide an approximation of oocyte size and number (Heins and Baker
1988). Smaller oocytes (<0.5 mm in diameter) were also counted and measured
to provide a size comparison to mature oocytes. Oocyte counts include
all oocytes (small and large) present in the ovary of the female. Oocyte identification procedures can be found in Heins and Baker (1988). Regressions
122 Southeastern Naturalist Vol. 9, No. 1
were performed to test for an infl uence of SL on the number of mature and
immature oocytes present in a mature female.
The smallest specimen collected during this study was taken in July,
measuring 16.3 mm in length and weighing 0.076 g TW. The gender of this
specimen could not be determined. The largest male and female specimens
were 96.07 mm and 18.6 g and 94.29 mm and 15.9 g, respectively. The earliest
young-of-year capture occurred in June, with specimens ranging from 17.66
to 19.08 mm (mean = 18.38, SD = 1.0). For all collections used in this study,
female to male ratio was 0.78:1, but no sexual size dimorphism was detected
in regards to SL (P = 0.1813) and EW (P = 0.0742). Standard length increased
with age in months (R2 = 0.92, P < 0.001), and preliminary examination of
the data suggested a curvilinear relationship existed between SL and EW.
We therefore log-transformed EW before regressing it with SL. This analysis
produced a significant linear relationship (R2 = 0.95, P < 0.001). Increases in
SL were greatest among months in the spring (Fig. 2). Of the 240 specimens
collected, 52% were age 0+, 39% were age 1+, and 9% were age 2+ age. The
maximum ages of males and females were 28 and 26 months, respectively.
The diet of F. stellifer consisted mostly of Chironomidae larvae (68% of
total diet by number) and Branchiopoda (12% of total diet) (Table 1). Levins’
NB value for F. stellifer was 0.0996. Data for Hypentelium etowanum
and Notropis xaenocephalus from recently published life-history studies
(Jolly and Powers 2008; O’Kelley and Powers 2007) produced NB values of
0.0791 and 0.275, respectively. Unidentified insect parts were in 22% of the
individuals examined, and sand particles were in 27%, peaking in June and
Figure 2. Standard length (SL) in mm by month of collection (1 = January, 2 = February,
etc.) for Fundulus stellifer collected from Moore Creek, Cherokee County, GA
between February 2006 and January 2007.
2010 K.L. Edberg and S.L. Powers 123
September (present in 11 individuals each month). Feeding was not uniform
throughout the year (F = 5.8, P < 0.001), with February and May having the
greatest weight of gut contents. Much of the variation in the weight of gut
contents can by explained by SL (R2 = 0.37, P < 0.001). There was also a significant relationship between SL and taxa richness of food items (P = 0.027),
but the relationship had low explanatory power (R2 = 0.02). Taxa richness
of food items did not differ statistically (F = 1.79, P = 0.06) throughout
the year. Collembola, Hirudinae, and Physa sp. made up moderate to large
components of the diet in winter, spring, and summer respectively, but were
largely absent from other seasons. The greatest proportion of empty guts occurred
in November, with a total of 41% of individuals lacking food items
that month (Table 1).
Female GSI peaked in May (mean GSI = 0.0974, maximum GSI = 0.211;
Fig. 3). Gravid females were collected in March, April, and May, with
oocytes ranging in size from 0.34 mm to 2.75 mm. The number of oocytes
present in a female was significantly related to SL (R2 = 0.58, P = 0.010),
with a mean of 308 mature oocytes found in gravid females (SD = 139.5).
The youngest age at sexual maturation for females was 22 months. The highest
mean GSI value for males (0.0025) was in February, and maximum male
GSI (0.0045) was in April (Fig. 4). Elevated GSI values were found in males
collected from February through July.
Our study suggests that specimens <1 year of age are generally <40 mm
SL, those 1+ are generally 45–80 mm SL, and those 2+ are 85–95 mm SL.
Maximum age appears to be just over 2 years. Growth appears to increase in
Figure 3. Gonadosomatic Index (GSI) by month of collection (1 = January, 2 =
February, etc.) for female Fundulus stellifer collected from Moore Creek, Cherokee
County, GA between February 2006 and January 2007.
124 Southeastern Naturalist Vol. 9, No. 1
Table 1. Stomach contents of Fundulus stellifer from Moore Creek, Cherokee County, GA by month. Numbers for each food item indicate total number of
individuals found. Detritus, sand, and unidentified insect parts are noted by occurrence due to difficulty quantifying them (e.g., the occurrence of sand in three
specimens is denoted as “3”).
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Proportion
Sand 3 8 2 4 11 8 3 11 3 2 6 61
Detritus 6 8 6 7 16 13 8 8 12 3 3 5 95
Unidentified parts 6 2 2 4 8 7 3 5 4 3 1 4 49
Nematoda 8 8 0.003
Trematoda 1 1 0.000
Ancylidae 1 1 0.000
Physa 2 5 4 12 16 2 41 0.015
Oligochaeta 1 1 0.000
Hirudinae 18 16 1 35 0.013
Araneae 1 1 2 0.001
Acari 2 1 3 0.001
Branchiopoda 5 21 45 68 14 10 19 16 73 21 34 2 328 0.122
2010 K.L. Edberg and S.L. Powers 125
Table 1, continued.
Jan Feb Mar Apr May Jun Jul Aug Sep Oct Nov Dec Total Proportion
Collembola 20 10 3 1 50 84 0.031
Hemiptera 1 1 0.000
Megaloptera 5 5 0.002
Ephemeroptera 3 4 1 3 18 2 4 1 11 3 50 0.019
Plecoptera 1 1 1 3 8 14 0.005
Coleoptera 1 2 1 4 0.001
Colepotera Larvae 1 2 3 0.001
Hymenoptera 5 2 7 0.003
Trichoptera 1 5 2 3 1 1 13 0.005
Diptera 1 1 0.000
Chironomidae larvae 6 168 62 545 221 356 126 85 109 29 12 103 1822 0.679
Unidentified pupae 1 4 5 15 9 1 5 1 2 43 0.016
Unidentified adults 3 1 2 2 1 2 1 12 0.004
# of specimens 21 20 16 20 19 21 14 19 16 21 17 17 221
Empty stomachs 6 1 1 1 0 0 1 0 1 8 7 3 29
% Empty 28.57 5.00 6.25 5.00 0.00 0.00 7.14 0.00 6.25 38.10 41.18 17.65 13.12
Total items 57 235 134 663 301 428 180 150 216 78 61 181 2684
Mean items/stomach 5.2 16.8 16.8 66.3 25.1 35.7 20.0 10.7 21.6 8.7 6.1 20.1 107.4
126 Southeastern Naturalist Vol. 9, No. 1
spring and level off in fall. This pattern coincides with the feeding patterns
of F. stellifer addressed below. The low proportion of age 2+ individuals
suggests that few individuals survive to reproductive maturity. This high
mortality prior to sexual maturity has been observed in other syntopic
species such as Hypentelium etowanum (O’Kelley and Powers 2007) and
Notropis xaenocephalus (Jolly and Powers 2008), and is common among
stream fishes (Boschung and Mayden 2004). The low proportion of specimens
<25 mm SL collected could be explained by these small individuals
passing through the 9.5-mm mesh of the seine. Sexual dimorphism appears
to be restricted to male specimens developing a black band on the distal
caudal margin as they approach sexual maturity.
Increased feeding during spring and summer months appears to coincide
with increased energetic requirements associated with gamete production,
growth, and spawning. The high numeric proportion of food items as Chironomidae
and Branchiopoda (67.9% and 12.2%, respectively) may indicate
selective feeding, as these data contrast with the diets of the syntopic Hypentelium
etowanum and Notropis xaenocephalus in which Chironomidae made
up 88% and 19.5%, and Branchiopoda 3.59% and 0% of their food items, respectively
(Jolly and Powers 2008, O’Kelley and Powers 2007). Contrasting
NB values (especially between F. stellifer and N. xaenocephalus) provide
further evidence of this selective feeding. The high number of Chironomidae
in the diet of F. stellifer may also be explained by densities often >50,000
Chironimidae/m2 in aquatic environments (Coffman and Ferrington 1996),
making them the most abundant food resource available and the bulk of the
diet of most stream fishes (Alford and Beckett 2007, Matthews et al. 1982).
The significant relationship between weight of gut contents and SL suggests
Figure 4. Gonadosomatic Index (GSI) by month of collection (1 = January, 2 = February,
etc.) for male Fundulus stellifer collected from Moore Creek, Cherokee County,
GA between February 2006 and January 2007.
2010 K.L. Edberg and S.L. Powers 127
that the amount of food consumed increases with size. The significant relationship
between taxa richness of food items and SL also suggests that diet
may become more varied in larger specimens, but the low explanatory power
value (R2 = 0.02) suggests these changes are slight. Taxa richness of food
items does not appear to change throughout the year, but certain taxa appear
to be seasonally important. Weight of gut contents differs throughout the
year, with elevated levels from late winter through summer. This variation
coincides with increased energetic requirements during periods of spawning
and increased growth.
The near restriction of Collembola, Hirudinae, and Physa sp. as food
items to specific seasons may be evidence of selective feeding, but may also
be explained by several relatively simple phenomena making them widely
available as prey items during specific times of year. As Collembola are
mostly soil dwelling or semi-aquatic (Merritt and Cummins 1996), the rising
waters of winter may fl ood previously dry riparian areas, making them easy
prey during winter months. Hirudinae are largely annual with eggs hatching
in the spring (Thorp and Covich 1991), which may provide an abundance
of small leaches as prey items at that time of year. Physa sp. may be largely
absent from Moore’s Creek except for summer months because pulmonate
snails are known to burrow into the substrate during cold periods (Thorp
and Covich 1991). Therefore, these data do not provide clear evidence of F.
stellifer selecting these specific items during particular seasons, but rather
may identify F. stellifer as opportunistic in feeding on whatever food items
are present in their habitat.
The GSI of both males and females was highest in spring, but males
displayed high GSI values earlier and later in the year compared to females.
Elevated female GSI in April and May, followed by a precipitous drop
in female GSI in June indicates spawning occurs in April and May, with
water temperatures near 16 °C, and is finished by mid-June. One specimen
collected in July appeared to be spent, as there were no oocytes and only a
membrane remaining. Mature oocytes were not observed in females younger
than 22 months of age. Subsequently, spawning does not appear to occur
until the second spring of life. We found that maximum female lifespan was
approximately 26 months. Thus, it appears that F. stellifer will go through
no more than one spawning cycle during their lifespan. No spawning activity
was observed, but during the spring months, specimens were collected from
backwater pools with heavy submerged and emergent macrophyte vegetation,
suggesting spawning habitat is likely similar to that inhabited during
the rest of the year. Young-of-year were also collected primarily from these
same areas of heavy vegetation or leaf cover.
Although F. stellifer and F. catenatus are closely related, they appear to
have several differences in life-history traits. Fundulus catenatus has a longer
lifespan (5+ years) and larger maximum size (nearly 115 mm SL) (Fisher
1981), compared to F. stellifer (just over 2 years and maximum SL of less
than 95 mm). Fundulus catenatus also has a longer spawning period which
lasts from April to July (Fisher 1981), whereas the F. stellifer spawning period
occurs from April to early June. Fundulus stellifer appears to be more
128 Southeastern Naturalist Vol. 9, No. 1
fecund with a mean of 308 mature oocytes in gravid females and a maximum
ova diameter of 2.75 mm, while F. catenatus have 28–245 mature oocytes
and larger ova up to 3.5 mm (Fisher 1981). Diptera larvae also appear to
make up the bulk of the diet of both species. Seasonal shifts in food items
appear common between the two species, but differences in pharyngeal teeth
morphology (Thomerson 1969) is likely adaptive, as the more molariform
teeth of F. stellifer likely crush mollusks such as Physa.
We thank S.D. Barton, D.S. Holder, and C.K. Ray for assistance with field and
lab work. Fishes were collected under Georgia Scientific Collecting Permit number
16494 issued to S.L. Powers. We thank Reinhardt College for the use of equipment
essential to this research. This study was conducted in part as an undergraduate independent
research project at Reinhardt College by K.L. Edberg.
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