Analysis of Spawning Behavior, Habitat, and Season of the Federally Threatened Etheostoma scotti, Cherokee Darter (Osteichthyes: Percidae)
Casey M. Storey, Brady A. Porter, Mary C. Freeman, and Byron J. Freeman
Southeastern Naturalist, Volume 5, Number 3 (2006): 413–424
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2006 SOUTHEASTERN NATURALIST 5(3):413–424
Analysis of Spawning Behavior, Habitat, and Season
of the Federally Threatened Etheostoma scotti,
Cherokee Darter (Osteichthyes: Percidae)
Casey M. Storey1,5,*, Brady A. Porter2, Mary C. Freeman3,
and Byron J. Freeman4
Abstract - Etheostoma scotti (Cherokee darter) is a member of the subgenus
Ulocentra and a federally threatened endemic to the Etowah River system, GA. Field
observations of spawning behavior of the Cherokee darter were made at five stream
sites to identify spawning season and habitat over two field seasons. Cherokee
darters primarily spawn in pool habitats between mid-March and early June, at
temperatures between 11 and 18 ºC. Egg deposition was typically on large gravel
substrate, but ranged from gravel to bedrock in size and included woody debris.
Spawning occurred in a variety of depths (0.09–0.59 m) and velocities (0–0.68 m/s).
We examined spawning behavior and habitat of Etheostoma scotti Bauer,
Etnier, and Burkhead (Cherokee darter), a federally protected species endemic
to the Etowah River system in north Georgia. Described in 1995
(Bauer et al.1995) and listed as threatened in the same year (Federal Register
1994), the Cherokee darter typically inhabits small to medium-sized streams
in the Piedmont physiographic province.
Spawning behavior has been described for various members of the subgenus
Ulocentra (sensu Bouchard 1977, Bailey and Etnier 1988), of which
E. scotti is a member. This includes E. coosae (Fowler) (O’Neil 1981), E.
baileyi Page and Burr, E. etnieri Bouchard (Porterfield 1998), E. flavum
Etnier and Bailey (Keevin et al. 1998), E. pyrrhogaster Bailey and Etnier, E.
zonistium Bailey and Etnier (Carney and Burr 1989), and E. simoterum
(Cope) (Page and Mayden 1981). Typically, females attach single eggs to
rock substrate in a vertical position. The size of substrate most commonly
utilized by females in these descriptions is cobble or larger.
This study identified spawning period and environmental variables associated
with reproduction by the Cherokee darter, including substrate size for
egg deposition, general habitat types (i.e., pools, runs, riffles), current velocity,
and water depths.
1Institute of Ecology, University of Georgia, Athens, GA 30602-2202. 2Department
of Biological Sciences, Duquesne University, Pittsburgh, PA 15282. 3US Geological
Survey, Patuxent Wildlife Research Center, Athens, GA 30602. 4Georgia Museum of
Natural History, Natural History Building, University of Georgia, Athens, GA
30602-1882. 5CH2M HiLL, Inc., 115 Perimeter Place, NE Suite 700, Atlanta, GA
30346. *Corresponding author - casey.storey@CH2M.com.
414 Southeastern Naturalist Vol. 5, No. 3
Based on sampling conducted in winter 2002 across the Cherokee darter’s
range (Fig. 1), we chose the five sites where Cherokee darters were most
abundant. Sites selected were Shoal Creek in Dawson County, McCanless
Creek and Hickory Log Creek in Cherokee County, Butler Creek in Cobb
County, and Whitehead Creek in Paulding County. The lengths of the study
reaches varied from site to site depending on the densities of Cherokee darters,
property boundaries, access via roads, and habitat. The reach lengths ranged
from 40 m (Hickory Log Creek) to 140 m (Whitehead Creek).
Species of the subgenus Ulocentra are known to begin spawning in early
spring (Bauer et al. 1995, Boschung and Mayden 2004, Etnier and Starnes
1993, O’Neil 1981, Porterfield 1998, Suttkus and Etnier 1991). In an effort to
record conditions driving the onset of spawning, we began observations, via
snorkeling conducted by two observers, in March during base-flow conditions.
Observations were made over a 2-year period during March–June. The
observation period ended when spawning activity (including courting, spawning,
and chasing) was no longer observed over several visits to each site, or
until a majority of observed individuals no longer displayed courting behavior.
In 2002, observations began on 22 March and ended on 15 May, and in
2003, field observations began on 21 March and extended to 10 June. During
both years, an attempt was made to visit all five sites at regular intervals (once
every three to four weeks in 2002, and once every week or two in 2003). In the
Figure 1. Location of study sites in the Etowah River basin, GA.
2006 C.M. Storey, B.A. Porter, M.C. Freeman, and B.J. Freeman 415
2002 field season, a total of 27.5 hours of field observations were logged, with
13 visits made at all sites combined. Observation time in 2003 was greater,
with a total of 60.6 observation hours and 36 visits.
Prior to snorkel observations, water-quality parameters were measured
with a Hydrolab Datasonde® 4a and a Hach Turbidimeter® model 2100P.
Parameters measured included turbidity, conductivity, temperature, and pH.
In addition, ONSET Optic Stowaway Temp™ temperature loggers were
deployed at all sites during each year and set to record temperatures at 15-
Reaches were snorkeled in an upstream direction. Habitats observable
through a snorkel mask (depth > 0.2 m) were visually scanned for the presence
of Cherokee darters. Habitat too shallow to be observed through a snorkel
mask (depth < 0.2 m) was scanned from above the water surface for the
presence of Cherokee darters. Cherokee darters were counted, and individuals
were monitored for signs of spawning behavior. This behavior included the
mounting of females by males, chasing of females by males, or pairing, in
which a male and female were at rest immediately adjacent to one another on
the streambed. When courting or spawning behavior was observed, pairs were
followed until the activity ceased or the pair was lost. When spawning events
took place, the substrate upon which the egg was attached was flagged and left
in place until snorkeling of the reach was complete.
After the reach was completely snorkeled, measurements were taken at
all flagged spawning sites. Velocity (0.01 m/s) and depth (0.01 m) were
measured at five points adjacent to the spawn site (immediately upstream,
immediately downstream, adjacent right and left, and on top) with a Marsh-
McBirney Inc. FLO-MATE™ portable flow meter Model 2000 mounted on a
top-setting wading rod. All velocity measurements were taken at 60% depth.
Depth measurements were taken in tenths of feet and later converted to
meters. The dimensions of the substrate upon which the spawning took place
were measured to the nearest half centimeter and classified, based on the
median axis measurement, by Wentworth size category (Gordon et al. 1992).
The depth of the area immediately surrounding the spawn was measured to
the nearest 0.01 m.
In 2003, the locations of observed spawns were plotted within each reach
using temporary benchmarks. At sites where a minimum of ten spawns were
observed, we measured reach-scale spawning parameters using a Leica®
electronic total-station surveyor. After spawning observations were
concluded for the 2003 season, we measured longitudinal variation in bed
elevation and water-surface elevation along the length of the reach. Elevation
measurements were taken at 0.5–2.0-m (usually 1-m) intervals along the
thalweg during base-flow conditions, and were plotted to illustrate upstream
to downstream variation in channel form. The bed elevations of the previously
triangulated spawn localities were measured, and the results superimposed
upon the graphed bed and water-surface elevations to examine occurrence of
spawns relative to pool-run-riffle structure within the site.
416 Southeastern Naturalist Vol. 5, No. 3
Spawning acts were observed at all sites. During 2002, all observed
spawns occurred from 23 April–30 April, whereas in 2003, we observed
spawns from 21 March through early June (Table 1). Observations were
terminated after 10 June 2003, when we observed only 15 individuals and a
single spawning pair in the study reach at Hickory Log Creek (Table 1).
We observed a total of 63 spawning acts during the 2-year study. Spawning
occurred on a variety of substrates and at varying depths and velocities.
Females most frequently selected gravel and cobble-sized rocks upon which
Table 1. Recorded spawning activity of observed Etheostoma scotti at all sites (2002–2003) in
Georgia Streams. Behavior: M/M = male chasing or displaying to other male, M/F =male
chasing or displaying to female, PS = pseudo spawn, S = spawn, DS = dry spawn, C = courting,
NSA = no spawning related activity observed. NA = not available, ± = approximation of data.
Water Conductivity Turbidity
Males Females Behavior temp. (ºC) (S/cm) (NTU) pH
22-Mar-02 10 4 M/M 9.37 35.9 NA 7.95
23-Apr-02 NA NA M/F, M/M, S 18.09 52.0 NA 7.58
14-May-02 11 12 M/F, PS, C 16.76 51.0 NA 7.66
21-Mar-03 19 18 M/M, C 16.06 44.9 NA 7.74
29-Mar-03 9 5 PS, C 17.05 23.3 NA 7.69
1-Apr-03 20 21 M/F 9.31 24.0 3.39 7.22
13-Apr-03 19 13 S 16.09 49.0 2.78 NA
17-Apr-03 25 25 M/M, S, PS 15.01 52.0 2.55 NA
29-Apr-03 24 24 S 16.27 50.0 3.56 NA
13-May-03 19 16 M/F, S, DS, C 14.42 44.0 4.4 NA
27-May-03 13 13 NSA 17.61 47.0 NA NA
Hickory Log Creek
11-Apr-02 5 6 M/M 15.22 34.0 NA 6.95
23-Apr-02 NA NA M/F, S, 14.49 35.0 NA 7.14
14-May-02 2 10 NSA 14.70 36.0 NA 7.44
21-Mar-03 35 25 S, PS, 13.60 33.5 NA 7.27
29-Mar-03 27 57 M/M, C 15.25 8.1 NA 7.12
1-Apr-03 35 56 M/M, M/F 11.71 7.0 NA 7.15
9-Apr-03 14 26 NSA 11.85 36.0 11.5 NA
17-Apr-03 17 34 C 13.90 38.0 3.83 NA
28-Apr-03 17 15 S 14.62 37.0 5.84 7.09
13-May-03 10 15 M/F 16.93 9.43 6.12 7.46
27-May-03 9 8 S 16.70 34.0 NA 7.38
10-Jun-03 8 7 M/M, S 18.10 NA 6.11 NA
26-Mar-02 2 2 NSA 12.64 19.0 NA 6.8
11-Apr-02 2 2 M/F 15.22 20.1 NA 7.4
30-Apr-02 4 2 M/F, S, PS 16.50 20.0 NA 6.97
23-Mar-03 0 5 NSA 13.70 20.6 NA 7.27
1-Apr-03 8 9 NSA 14.32 NA NA 7.08
13-Apr-03 8 6 M/M, S 11.47 26.0 3.40 7.03
28-Apr-03 5 4 S 13.38 25.0 4.41 NA
13-May-03 3 3 C, M/F 13.02 19.0 5.13 7.11
27-May-03 2 4 NSA 14.63 20.0 NA 6.19
2006 C.M. Storey, B.A. Porter, M.C. Freeman, and B.J. Freeman 417
to deposit eggs. However, females used substrates varying from gravel to
bedrock, as well as woody debris (Fig. 2). Velocities at spawning sites
ranged from 0 to 0.68 m/s, with a mean of 0.24 m/s. Depth ranged from 0.09
to 0.59 m, with an average of 0.31 m (Fig. 3).
Water characteristics varied across sites and dates (Table 1). For the
dates when spawning was observed, temperatures ranged from 11.47 to
18.09 ºC, dissolved oxygen ranged from 8.84 to 11.32 mg/L, conductivity
ranged from 20.1 to 113 S/cm, turbidity from 2.55 to 6.11 NTU, and pH
ranged from 6.97 to 7.91. Continuous records of water temperature were
obtained for comparable periods in both years at three sites, and showed that
water temperatures were on average cooler during the 2003 spawning season
compared to 2002 (Table 2). Differences were most pronounced early in the
season. For example, at the Shoal Creek site (which had the most complete
Table 1, continued.
Water Conductivity Turbidity
Males Females Behavior temp. (ºC) (S/cm) (NTU) pH
4-Apr-02 1 3 NSA 14.56 78.0 NA 7.54
25-Apr-02 8 6 C, S, PS 16.90 89.0 NA 7.16
15-May-02 13 18 M/F, MM 14.23 100.0 NA 7.47
25-Mar-03 17 21 S, PS 13.52 109.0 NA 7.91
31-Mar-03 6± 8± C, PS, M/F 11.12 95.2 3.54 7.19
4-Apr-03 21 18 M/F, S, PS 16.80 106.0 4.5 7.45
15-Apr-03 23 25 C, MF 16.77 113.0 2.7 NA
12-May-03 11 18 S, PS, DS 16.45 113.0 NA 7.47
30-May-03 14 8 M/F 15.95 110.0 NA 7.22
4-Apr-02 3 2 M/M 12.94 34.0 NA NA
18-Apr-02 5 10 NSA 19.75 47.0 NA 7.36
25-Apr-02 5 6 M/M, C 18.92 10.0 NA 7.39
15-May-02 8 17 C, PS 16.51 47.0 NA 7.53
26-Mar-03 17 17 C, PS 17.08 41.1 NA 7.66
31-Mar-03 6 8 NSA 8.91 18.3 4.38 6.92
4-Apr-03 10 14 C, PS 14.59 45.0 3.4 7.37
15-Apr-03 10 14 C, S 14.05 46.0 3.01 NA
29-Apr-03 3± 8± NSA 18.32 47.0 NA 7.41
12-May-03 9 7 C 18.77 41.0 NA 7.3
30-May-03 7 2 NSA 17.10 41.0 NA 7.11
Table 2. Temperature data for three Etheostoma scotti study sites in Georgia streams, comparing
average daily temperature and total degree days (summed daily averages for dates included)
between 2002 and 2003.
Shoal Creek Whitehead Creek Hickory Log Creek
27 March–26 May 5 April–29 May 11 April–26 May
Year 2002 2003 2002 2003 2002 2003
Average daily temp. (oC) 15.1 14.7 16.7 15.7 16.5 15.9
Total degree days 924 827 918 861 757 730
418 Southeastern Naturalist Vol. 5, No. 3
Figure 2. Spawning substrate utilization by Etheostoma scotti across all localities in
the Etowah River basin, GA, observed through seasons 2002 and 2003. Substrate
types follow Wentworth size scale (Gordon et al. 1992). The y-axis shows the
substrate selection frequency.
Figure 3. Plot of depths and velocities at observed Etheostoma scotti spawns for all
sites in the Etowah River basin, GA, 2002 and 2003. Large squares = McCanless
Creek, small squares = Hickory Log Creek, open circles = Whitehead Creek, closed
circles = Shoal Creek, triangles = Butler Creek.
2006 C.M. Storey, B.A. Porter, M.C. Freeman, and B.J. Freeman 419
temperature record), daily temperatures during April averaged 15.3 in 2002
compared to 13.7 in 2003.
During spawning observation, male Cherokee darters were often observed
displaying perpendicular to the long axis of females, with dorsal fins erect and
nuptial colors at high intensity. Males often pursued receptive females,
mounting them and periodically pecking at their nape to instigate spawning
behavior. Females moved about scanning substrate in what appeared to be an
attempt to select a suitable egg-deposition point. Often, females would peck
substrate, possibly attempting to signify to the male her readiness to spawn.
This pecking often elicited quivering by the male as he mounted or positioned
himself parallel to the female. Then the female continued moving about in
search of further appropriate egg-deposition points, with the male matching
her movements. Once suitable substrate was found, the female would again
peck at a spot, usually on the vertical surface of the substrate, and the male
would begin to quiver and release milt. Very soon after pecking, and while the
male was still quivering, the female would quiver briefly and quickly move
forward with her ovipositor touching the exact point that she pecked. After
this series of events, a slightly opaque and colorless egg could be seen
attached within small dimples or fissures upon the substrate selected.
We observed multiple spawning acts on a single rock only once. Females
appeared to select substrate free of algal growth or sediment coatings. In
many situations, females would remove fine debris or algae by pecking the
substrate, frequently several times. This was most often noted in streams or
reaches with dense algal growth. In many instances, when frequent pecks
were made, the female would abandon the site and continue the search for
Occasionally, a female would select/peck a substrate and the male
would begin to quiver; but, instead of placing her ovipositor over the
cleaned crevice, the female would abruptly change positions or move off
the substrate and continue to search. We termed this behavior pseudospawning.
The observation of a pseudo-spawn indicated that spawning
was taking place on the date of the observation (Table 1). This was useful
in determining the duration of spawning and in signifying spawning pairs
that should be observed. Towards the end of the spawning season we
observed “dry spawning.” In this situation, the entire courting and spawning
ritual would be carried out by a pair, but after positioning of the
ovipositor by the female, an egg would not be observed in the spot. This
behavior was noted at McCanless Creek (13 May, 2003) and Butler Creek
(12 May, 2003) and we treated it as a true spawn, taking measurements as
outlined in the methods.
At the conclusion of the snorkel observations, McCanless Creek and
Butler Creek were selected for longitudinal mapping, because 10 or more
spawns were observed in these creeks. Observed spawns plotted against the
longitudinal variation in bed and water-surface elevation indicated that
420 Southeastern Naturalist Vol. 5, No. 3
Figure 4. Total-station analysis of water-surface elevation (upper line), bed elevation
(lower line), plotted downstream (0 m) to upstream for the Butler Creek study site in
the Etowah River basin, GA, shown with habitat types. Etheostoma scotti spawning
locations and elevations are indicated by shaded circles; locations shown above or
below the bed are lateral to the thalweg.
Figure 5. Total-station analysis of water-surface elevation (upper line) and bed
elevation (lower line), plotted downstream (0 m) to upstream for the McCanless
Creek study site in the Etowah River basin, GA, shown with habitat types.
Etheostoma scotti spawning locations and elevations are indicated by shaded circles;
locations shown above or below the bed are lateral to the thalweg.
2006 C.M. Storey, B.A. Porter, M.C. Freeman, and B.J. Freeman 421
spawning was most concentrated in topographic valleys (Figs. 4 and 5).
Despite the availability of riffles throughout the studied reaches, the selected
spawning habitat was consistently located at the margins, heads, or tails of
pools, as well as in run habitat.
Field observations correspond with laboratory observations of Cherokee
darters spawning (Bauer et al. 1995); for example, females were observed
“visually scrutinizing” their surroundings, presumably in search of appropriate
spawning substrata. Additionally, in the laboratory and in the field, male
Cherokee darters tend to maintain a “roving territory” around the female
with which they are courting. As the female may travel several meters or
more through a variety of habitats, the male maintains a territorial area.
When other males enter this loosely defined territory, the male will often
give chase. In this process, he may lose track of the female or have another
male take his place.
Cherokee darters appear to have a potentially prolonged spawning
period, extending from at least the middle of March to early June, corresponding
to water temperatures of 11.5 to 18.1 ºC. The extended spawning
season observed in 2003 (in contrast to 2002) may have been related to
cooler temperatures. Indeed, temperature has been cited as a major factor
contributing to the length of spawning season in percids (Hubbs 1985). In
addition, we hypothesize that cooler temperatures in 2003 may have been
responsible for the observed “dry spawn” behavior noted near the end of the
spawning period, if appropriate water temperatures stimulate spawning behavior
despite the inability of females to deposit or produce additional eggs.
Spawning sites characteristically occurred in run and pool habitats, with
moderate depths and velocities (i.e., usually < 0.5 m and < 0.4 m/s, respectively),
and most often, on gravel-sized sediment. The use of gravel for egg
attachment contrasts with the use of cobble and boulder substrates
frequently reported for other Ulocentra species (Porterfield 1998), but
Cherokee darters also utilized a variety of bed sediments, from medium
gravel to bedrock. Woody debris was also used.
Other Ulocentra also peck substrate prior to egg placement (Porterfield
1998). After observing numerous spawning Cherokee darters, we propose
that this behavior probably serves two purposes. The first is to stimulate the
male and to signal the onset of egg deposition. Secondly, this behavior may
clear each site for strong adherence of the individual egg. Thus, availability
of bed sediments that are relatively free of fine sediment and algal growth
may affect habitat suitability for spawning.
Future research into a relationship between the pecking behavior and
periphyton density could be helpful in determining to what extent nutrient
loading and algal growth may limit spawning habitat. Such an analysis could
422 Southeastern Naturalist Vol. 5, No. 3
offer an explanation for pecking behavior beyond its use in courtship rituals.
If this behavior varies among Ulocentra species, then this variation could
predict differences in population responses to stream eutrophication.
Our observations on habitat use and spawning period provide information
that should be useful for avoiding development-related impacts on
Cherokee darters. Urbanization frequently leads to a loss of stream fish
species (Walters et al., in press; Wang et al. 2001). Habitat alteration
relative to requirements for reproduction may contribute to these losses.
From our field observations, it is clear that a reduction in the availability of
gravel and cobble substrate from sedimentation could reduce the spawning
success of the Cherokee darter. Excessive sediment inputs can additionally
cause infilling of pools (the primary sites of spawning) and alteration of
depth and velocity regimes (Berkman and Rabeni 1987, Waters 1995).
Lower Cherokee darter abundances have been associated with increasing
levels of fine sediments and higher bed mobility (Roy 2004), possibly
reflecting lower reproductive success and demonstrating the importance of
erosion and sedimentation controls. Roy (2004) also related lower Cherokee
darter abundances to increased magnitude and flashiness of
stormflows, which is in turn related to increased urban and impervious land
cover. Increasing runoff from impervious surfaces and loss of riparian
buffers may also increase stream temperatures (Barton et al. 1985, Osborne
and Kovacic 1993, Poole and Berman 2001), adversely affecting Cherokee
darter reproduction. We observed spawning to occur across a temperature
range of 11.0–18.1 ºC. While this range is relatively large, the protection of
forested riparian buffers is suggested throughout the range of this species
in order to prevent the early elevation of temperatures that could shorten
the spawning season. Finally, culvert installation and other types of urban
and suburban development that cause instream disturbance could be timed
to avoid disturbing spawning and incubation of eggs. Conservatively, restricting
instream disturbance during a period spanning March to early
June should benefit Cherokee darter reproduction.
We thank Megan Hagler, Jane Rogers, Ryan Creehan, and Jesslyn Shields for
their assistance in various aspects of this project. We especially thank property
owners that granted access to our snorkel sites through both field seasons. This
research was funded by the US Department of the Interior through Cooperative
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