K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
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2013 SOUTHEASTERN NATURALIST 12(4):816–842
Conservation Assessment of the Yazoo Darter
(Etheostoma raneyi)
Ken A. Sterling1,2,*, Melvin L. Warren, Jr.3, and L. Gayle Henderson3
Abstract - We summarized all known historical and contemporary data on the geographic
distribution of Etheostoma raneyi (Yazoo Darter), a range-restricted endemic in the Little
Tallahatchie and Yocona rivers (upper Yazoo River basin), MS. We identified federal and
state land ownership in relation to the darter’s distribution and provided quantitative estimates
of abundance of the species. We also quantified sex ratio and mean size of males and
females, summarized abiotic and physical characteristics of streams supporting the species,
and characterized the fish assemblage most often associated with the Yazoo Darter. Yazoo
Darters are generally limited to headwater streams, have a female-skewed sex ratio, and
have larger males than females. Individuals in the Yocona River drainage are larger than
in the Little Tallahatchie River drainage. Abundance was highly variable among streams
within the two major drainages, but was similar within and between drainages. Yazoo Darter
habitat in the Little Tallahatchie River drainage has some protection because many streams
supporting this species are on land managed by federal or state agencies. Streams with Yazoo
Darters are far less common in the Yocona River drainage, have almost no protection,
and face growing pressure from urban expansion. For these reasons, management action is
urgently needed for Yocona River populations.
Introduction
Etheostoma raneyi Suttkus and Bart (Yazoo Darter) (Percidae: subgenus Adonia)
is a range-restricted fish endemic to the Yocona, Little Tallahatchie, and Tippah
river systems of the upper Yazoo River basin in north-central Mississippi (Fig. 1;
Johnston and Haag 1996, Suttkus et al. 1994, Thompson and Muncy 1986). The
species is classified as vulnerable by the Southeastern Fishes Council (Warren et
al. 2000) and American Fisheries Society (Jelks et al. 2008), as globally imperiled
by the Nature Conservancy (NatureServe 2013), and as sensitive by the USDA Forest
Service (USDA Forest Service 2013). The Mississippi Comprehensive Wildlife
Conservation Strategy lists the Yazoo Darter as a Tier 1 species of greatest conservation
need in the Upper East Gulf Coast Plain Ecoregion (Mississippi Natural
Heritage Program 2002).
Yazoo Darters are small (less than 65 mm SL), benthic insectivores living ≤3 years,
and most individuals do not survive their first year (Johnston and Haag 1996).
Recent phylogenetic analyses using mitochondrial DNA recovered two monophyletic
clades that are congruent with localities of Yazoo Darter specimens
from the Little Tallahatchie River and Yocona River drainages (Powers and
Warren 2009). Based on this genetic information, Powers and Warren (2009)
1Department of Biology, University of Mississippi, PO Box 1848, University, MS 38677.
2Current address - 385 East Center Street, Richfield, UT 84701. 3Center for Bottomland
Hardwoods Research, Southern Research Station, USDA, Forest Service, 1000 Front Street,
Oxford, MS 38655. *Corresponding author - kensterling39@gmail.com.
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2013 Southeastern Naturalist Vol. 12, No. 4
recommended these two allopatric populations of Yazoo Darters be treated as
separate management units.
Although not formally described until 1994 (Suttkus et al. 1994), the Yazoo
Darter was recognized as distinct in earlier surveys (Randolph and Kennedy 1974,
Thompson and Muncy 1986) within its range, which provided substantial historical
distributional information from the 1960s, 1970s, and 1980s. Suttkus et al. (1994)
indicated the first known collection of Yazoo Darters occurred in 1952 at Pumpkin
Creek (Lafayette County, MS) and provided other collection localities through the
early 1990s. S.T. Ross (2001; University of New Mexico, Albuquerque, NM, pers.
comm.) furnished records primarily from the 1980s through the mid-1990s. Under
the auspices of the USDA Forest Service (USFS), one of us (M.L.Warren, Jr.) conducted
an extensive set of surveys throughout the range of the species from 1999 to
2003 (Warren et al. 2002) and again from 2009 to 2011.
The goal of this study was to summarize known distributional, habitat, and
biological data for the species including new information from our recent work.
Specifically we had six objectives: 1) summarize all known historical and contemporary
data on geographic distribution of the species, 2) identify federal and
state land ownership in relation to the darter's distribution, 3) provide quantitative
estimates of the species’ abundance, 4) quantify sex ratio and mean size of male
and female fish, 5) summarize abiotic characteristics of streams supporting the species,
and 6) characterize the fish assemblage most often associated with the Yazoo
Darter. Our findings provide crucial information for the conservation of this species
and a basis for future research.
Field-site Description
The range of the Yazoo Darter lies within the Northern Hilly Gulf Coastal
Plain Ecoregion (Chapman et al. 2004) of north-central Mississippi (Fig. 1),
which consists of low rolling hills with elevations ranging from 80 to 180 m. The
region has experienced significant anthropogenic habitat alteration. Beginning in
the mid-19th century, forests were removed and land was converted to agricultural
use, leading to widespread and dramatic erosion, which filled stream valleys
with sediment and exacerbated flooding problems (Cooper and Knight 1991,
Shields et al. 1994). Localized efforts toward flood prevention and land reclamation
by straightening and channelizing streams met with little success (Shields et
al. 1994). The so-called Great Flood of 1927 affected seven states including Mississippi
and prompted the federal government to alter streams in an effort to prevent
catastrophic flooding. Within the range of the Yazoo Darter, large (>40,000
ha) flood-control impoundments were constructed on the Yocona and Tallahatchie
Rivers, extensive stream reaches were straightened and channelized, and
hundreds of headwater streams were impounded by small dams. These actions,
particularly stream channelization, altered stream gradients, which resulted in
stream incisement and headcutting in nearly all headwater streams (Shields et al.
1998). Channelized and incised streams tend to be shallow, sandy, homogeneous,
turbid, and unstable with flashy flows (Adams et al. 2004; Shields et al. 1994,
1998; Simon and Darby 1997).
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2013 Southeastern Naturalist Vol. 12, No. 4
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Methods
We used a hierarchical organization of sample locations based on watersheds for
comparative analyses detailed later in the Methods section (Fig. 1). Hereafter, we
use the phrases Yocona R. drainage and Tallahatchie R. drainage to refer to these
two river systems. We use the term unit to refer to groups of sample locations within
subdrainages of these two river systems. We grouped sample locations within the
Yocona R. drainage into two units: the Otoucalofa Creek Unit and the Yocona R.
Unit. We grouped locations in the Tallahatchie R. drainage into three units: the
Cypress Creek Unit, the Tippah River Unit, and the Tallahatchie R. Tributaries
Unit, which includes all locations within the Tallahatchie R. drainage except those
within the Tippah River and Cypress Creek units. We used the terms drainage and
unit to help distinguish these analytical groupings from more general references to
watersheds and tributaries, which are defined in the usual way .
Compilation of historical and current records
We compiled historical records (pre-1999) for Yazoo Darters from the following
sources: published literature (Johnston and Haag 1996, Randolph and Kennedy 1974,
Ross 2001, Suttkus et al. 1994, Thompson and Muncy 1986); unpublished data (Mississippi
Museum of Natural Sciences [T. Slack, Jackson MS, unpubl. data], Tulane
Museum of Natural History [H. Bart, Tulane University, Belle Chase LA, unpubl.
data]); and collection records from other USFS colleagues (W. Haag, USFS, Oxford,
MS, unpubl. data). We incorporated recent records (post-1998) from our own database
for the 1999–2003 USFS surveys, and from our own recent samples (2009–2011) into
the database (Appendix I). Here, we use the term location to refer to a physical site
within a stream that was sampled for fishes (i.e., the unique site IDs in Appendix 1).
Field methods
We predetermined reach lengths sampled for Yazoo Darters and other fishes in
order to make sampling effort proportional to stream size (Angermeier and Smogor
1995, Paller 1995). In 1999, we calculated reach lengths of streams 4–14 m wide as
20 times the average width. In 2000–2003, we sampled reaches that were 30 times the
average stream width. In 1999, we set a minimum reach length of 80 m for streams
less than 4 m average width. In later samples, minimum reach length was 120 m for streams
less than 4 m average width. We set a maximum reach length of 300 m for streams >15 m
average width. Changes in reach length were made to more thoroughly characterize
stream habitat and to increase the probability of detecting uncommon fishes.
For fish samples (n = 93) collected in 1999–2003 (Warren et al. 2002), we
standardized effort for single-pass backpack electrofishing and seining to reduce
bias and ensure capture of a representative sample of all fishes. We predetermined
electrofishing effort by multiplying the reach length (see preceding paragraph) by 5
seconds (i.e., we electrofished 5 seconds/m) and we allocated time fished along the
entire reach and all available habitats. We indexed fish abundance as the number of
fish sampled per minute of electrofishing. We conducted 8 seine-hauls for streams
less than 5 m average width and 12 seine-hauls for streams >5 m average width. We defined
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2013 Southeastern Naturalist Vol. 12, No. 4
a haul as either a sustained pull within a stream habitat such as a pool or one setand-
kick in a riffle (Jenkins and Burkhead 1994). We allocated seine hauls along the
entire reach and attempted to sample all available habitats (riffles, runs, and pools).
For our 2009–2011 fish samples (n = 97), Yazoo Darters were specifically targeted
using only single-pass backpack electrofishing. We quantified abundance
of Yazoo Darters by electrofishing from 300–5734 seconds (mean = 1076.7, SE=
181.4) in a reach depending on the size of the stream. We sampled most streams
from March 2009 through July 2009. We recorded all fishes captured and measured,
and sexed all Yazoo Darters. Sex was determined by presence or absence of male
breeding colors, primarily the orange pigment present year round on mature males.
Immature fish (less than 30 mm) were not sexed but were used for all other analyses including
abundance estimates.
In a related study, we sampled three locations (sites 7180, 6821, and 6852;
Appendix 1) in separate streams periodically (June–July 2009, September–October
2009, January 2010, April–May 2010, September 2010, March 2011, June–
July 2011). A fourth location (site 7053; Appendix 1) was added to our periodic
sampling September–October 2009. At these locations, we used standardized
fish-sampling methods described previously for 2000–2003. All data collected
were used for all analyses, including abundance estimates.
We recorded habitat variables for our 1999–2003 samples after sampling for fishes.
Within each reach, we established 12 equally spaced transects (distance between
transects, 6.67–25 m) along the pre-determined fish-sampling reach. At each transect,
we measured wetted width and visually estimated stability (eroding or stable)
and height of each bank. However, because measures of right and left bank stability
and right and left bank height were highly correlated (data available on request), we
used data from the right bank only to reduce the number of variables used for analyses.
We measured water depth (cm) and water velocity (m/sec at 0.6 depth) at equally
spaced points along each transect . We also recorded presence or absence of detritus,
small wood (<10 cm diameter or <1.5 m in length), large wood (>10 cm diameter or
>1.5 m in length), and aquatic vegetation, and visually estimated percentage canopy
cover at each point as 0, 25, 50, 75, or 100%. We adjusted the number of points per
transect depending on stream width (transects >10 m in width = points at 2-m intervals;
5–10 m in width = points at 1-m intervals; less than 5 m in width = 5 sample points).
Because the number of points used to measure variables varied depending on stream
width, variables measured as present or absent are proportional.
Data analyses
We calculated abundance, sex ratios, and mean standard length (SL) of Yazoo
Darters using post-1998 data (Warren et al. 2002) and data from our recent surveys
(2009–2011). We estimated abundance at sampling locations as the number of Yazoo
Darters captured per minute of electrofishing (CPUE) ± 95% confidence intervals.
Yazoo Darters captured by seine are not included in the abundance estimates.
We calculated sex ratios, mean SL of male and female darters, and mean SL of
males and females combined for watershed units within the Tallahatchie R. drainage
and the Yocona R. drainage and for all sample locations within each drainage
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combined (Fig. 1). We tested if sex ratios deviated significantly from 1:1 in each
unit within a drainage (chi-square goodness of fit, nonparametric exact P-values;
StatXact version 8 [Cytel, Inc. 2007]) and for differences in SL between sexes
among units within drainages and between drainages (ANOVA, PopTools [Hood
Figure 1. Major drainages, units, counties, and cities within the range of the Yazoo Darter in
north-central Mississippi; red circles show location of all known Yazoo Darter collections.
Tallahatchie R. units are outlined and lettered as: A = Tippah River Unit, B = Tallahatchie
R. Tributaries Unit, and C = Cypress Creek Unit. Yocona R. watershed units are outlined
and lettered as: D = Yocona R. Unit and E = Otoucalofa Creek Unit.
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2013 Southeastern Naturalist Vol. 12, No. 4
2010]). We calculated 95% confidence intervals for sex ratios and mean SL for all
units and drainages.
Using data from the 1999–2003 locations at which Yazoo Darters were present,
(n = 37), we determined stream link (Osborne and Wiley 1992) and stream order
(Strahler 1957) from USGS 7.5-minute topographic maps by counting perennial
and intermittent streams. We counted perennial and intermittent streams because
stream-flow designations are unreliable in our region. As a consequence, our estimates
of these measures are inflated as compared to estimates obtained by counting
only perennial streams as described in the original papers. We also determined
watershed area (km2) for each of these locations using either USGS 7.5-minute
topographic maps or DeLorme TopoUSA version 7.1.0. We then calculated means
(± SD) for each of these variables and calculated mean values (± SD) for wetted
width, water depth, and water velocity from each site.
We tested possible relationships between abiotic habitat variables and Yazoo
Darter abundance and presence/absence data. First, we used principle components
analysis (PCA; PC-Ord ver. 5 [McCune and Mefford 1999]) to reduce 12 abiotic
variables (stream order, watershed area, wetted width, water depth, water velocity,
detritus, small wood, large wood, aquatic vegetation, canopy cover, bank height, and
bank stability) to a smaller number of synthetic variables that retained most of the
information from the original data. Mean values per location were calculated for all
variables except for proportional variables (detritus, small wood, large wood, aquatic
vegetation, and bank stability). Proportions were calculated from the presence or absence
of variables at sample points along transects (see Field methods section) except
for bank stability, for which we used the proportion of transects with stable banks. We
square-root transformed all data except proportional data, which we arc-sin squareroot
transformed. We determined the number of interpretable axes generated by PCA
using the broken-stick method (Jackson 1993). We then correlated (Pearson coefficient,
JMP 5.1 statistical software [SAS Institute 2002]) Yazoo Darter abundance at
each location with the site scores from each PCA axis. We also used logistic regression
to test for relationships between presence/absence data (likelihood ratio test; JMP 5.1
statistical software [SAS Institute 2002]) and site scores from each PCA axis.
We used indicator species analysis (ISA) as implemented in PC-ORD version
5.0 (McCune and Mefford 1999) to identify and test for significant fish species
associations with Yazoo Darters using Monte Carlo methods. The test statistic
is the maximum indicator value estimated for each species. Maximum indicator
values result from multiplying the proportional abundance of a species in a given
group relative to the abundance of that species in all groups and the proportional
frequency of a species in each group (Dufrene and Legendre 1997). Presence or
absence of Yazoo Darters per site was used as the grouping variable, and 10,000
permutations were used for Monte Carlo iterations. Species occurring at ≤5 (≈5%)
sites were dropped from the analysis. Lampreys were not identified to species in the
field because many individuals were larvae, but we believe that a high proportion
of lampreys sampled were Lampetra aepyptera Abbott (Least Brook Lamprey). For
this reason, we grouped all lampreys sampled.
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2013 Southeastern Naturalist Vol. 12, No. 4
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We omitted two locations sampled in the 1999–2003 surveys (sites 6814 and
6819; Appendix 1) from all of our analyses because they were statistical outliers
for watershed area, average wetted width, and average depth. We determined this
using an outlier box plot (JMP 5.1 statistical software; SAS Institute 2002). Each
location was from the mainstem Tallahatchie R. Canal, and each location had one
juvenile Yazoo Darter recorded. We address the implications of omitting these sites
in the Discussion section.
Results
Our data compilation of historical and contemporary sampling records yielded
224 records of Yazoo Darters, including multiple samples at the same location over
time, out of about 1200 total recorded samples for fishes within the known range
of the species (Figs. 2, 3; Appendix 1). At any given location of occurrence, Yazoo
Darters were detected from 1 to 23 times. A total of 2419 individual Yazoo Darters
were captured across all locations and samples. Of the 55 locations yielding Yazoo
Darters post-1998, 38 were new, previously unsampled locations. Sixteen locations
from the pre-1999 collections were resampled post-1998, with 13 yielding Yazoo
Darters. Two locations that yielded darters in 1999–2003 did not in 2009–2011
(sites 6820 and 6877).
Within its relatively narrow range, the Yazoo Darter is dispersed across numerous
tributaries in the middle Tallahatchie R. and middle Yocona R. drainages.
Within the Tallahatchie R. drainage, the species is known from 11 individual tributaries
(18 locations) within the Tippah River Unit, 1 tributary (Puskus Creek: 15
locations) plus 2 locations in the mainstem within the Cypress Creek Unit, and 10
tributaries (31 locations) plus 2 locations in the mainstem within the Tallahatchie
R. Tributaries Unit. Within the Yocona R. drainage, the species is known from 4
tributaries (13 locations) in the Yocona R. Unit, and 10 tributaries (10 locations)
plus 2 locations in the mainstem within the Otoucalofa Creek Unit. All locations
with Yazoo Darters are within the boundaries of the Northern Hilly Gulf Coastal
Plain Ecoregion, with the possible exception of two locations (sites 6847 and 7175)
that are near the boundary with the Loess Plains Ecoregion (Chapman et al. 2004).
Of 93 locations of known occurrence of the Yazoo Darter, only 26% (24) are
on federally or state managed property. Twelve are on federal property managed
by the Holly Springs National Forest, 6 are on federal property managed by the
United States Army Corps of Engineers, and 6 are on state of Mississippi property
(University of Mississippi Field Station and Wall Doxey State Park) (Figs. 2, 3; Appendix
1). These sites represent 9 separate tributary streams. Another 40 locations
(43%) are ≤2 km from federal or state lands and represent 11 separate tributary
streams. Most of these locations (33) are in the Tallahatchie R. Tributaries, Tippah
River, and Cypress Creek units. The Yocona R. Unit has only 7 such locations,
confined to 2 tributaries, and the Otoucalofa Creek Unit has non e.
The Yazoo Darter is decidedly a species of small, flowing streams. At 37
locations yielding Yazoo Darters in the 1999–2003 survey, mean stream order,
stream link, watershed area, wetted width, water depth, and water velocity all are
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2013 Southeastern Naturalist Vol. 12, No. 4
indicative of small, shallow, flowing streams (Table 1). The narrow confidence
intervals on most of these variables suggest a high affinity for this range of habitat
conditions. Examination of survey results in large streams in the area lends further
support to the small-stream affinities of the species. A total of 91 samples in our
compiled database from mainstem reaches of the Tippah River, Tallahatchie R.,
Figure 2. Results of pre-1999 stream samples and landownership across the range of the
Yazoo Darter. Solid red circles represent locations that yielded Yazoo Darters, and open
circles represent locations that did not yield Yazoo Darters. The polygon encloses the proclamation
boundary of the Holly Springs National Forest; federal and state lands are color
coded (see legend).
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2013 Southeastern Naturalist Vol. 12, No. 4
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Cypress Creek, Yocona R., and Otoucalofa Creek did not yield Yazoo Darters.
However, one juvenile Yazoo Darter was captured from each of two locations in
the relatively large Tallahatchie R. Canal in August 1999. Two Yazoo Darters were
taken in the mainstem of Otoucalofa Creek (Ross 2001; S.T. Ross, unpubl. data)
at the confluence with Sarter Creek (site 4984) in May 1986. Two samples in July
Figure 3. Results of post-1998 stream samples and land ownership across the range of the
Yazoo Darter; solid red circles represent locations that have yielded Yazoo Darters, open
circles represent locations that have not yielded Yazoo Darters. The polygon encloses the
proclamation boundary of the Holly Springs National Forest; federal and state lands are
color-coded (see legend).
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2013 Southeastern Naturalist Vol. 12, No. 4
1985 (Ross 2001, S.T. Ross, unpubl. data) contained 26 Yazoo Darters near the
headwaters of Otoucalofa Creek in the mainstem of the stream (site 1129). Yazoo
Darters were also present in 7 samples from near the headwaters of Cypress Creek
in the mainstem (sites 6865 and 6867). We address these samples in the Discussion.
Mean abundance of Yazoo Darters at locations of occurrence varied among units
but within-unit variability was high (Fig. 4). Mean abundance across all units ranged
from 0.57 (Cypress Creek Unit) to 1.23 individuals/minute (Otoucalofa Creek Unit).
Notably the Yocona R. and the Cypress Creek Units had considerably lower mean
abundances (≈50% lower) than other units within their respective river drainages, but
confidence intervals showed broad overlap. Confidence intervals for most units were
wide, indicating a high level of among-site variation. No differences in mean abundance
were apparent between the Yocona R. and the Tallahatchie R. drainages.
Differences in standard length were apparent between sexes and between the Yocona
R. and Tallahatchie R. drainages. Males were significantly larger than females
in the Yocona R. drainage (df = 1, 95; F = 23.05; P ≤ 0.0001) and the Tallahatchie
R. drainage (df = 1, 309; F = 114.63; P ≤ 0.0001). Females (df = 1, 305; F = 22.23;
P ≤ 0.0001) and males (df = 1, 9; F = 4.11; P ≤ 0.045) were significantly larger in
Table 1. Means, standard deviation (± SD), and 95% confidence intervals (CI) for abiotic variables at
locations with Yazoo Darters sampled from 1999–2003 across all units and drainages (n = 37).
Order Link Area (km2) Width (m) Depth (cm) Velocity (m/sec)
Mean 3.24 28.30 20.86 4.25 14.77 0.22
± SD 0.98 41.33 23.96 2.00 11.51 0.15
Upper 95% CI 3.54 43.22 29.10 4.91 18.79 0.27
Lower 95% CI 2.92 16.89 13.79 3.65 11.49 0.18
Figure 4. Mean
abundance (fish/
minute of electrofishing;
± 95%
confidence intervals)
of Yazoo
Darters across
locations of occurrence
for each
unit and river
drainage. Otoucalofa
Creek Unit,
n = 8; Yocona R.
Unit, n = 13; Cypress
Creek Unit,
n = 21; Tippah
R. Unit, n = 22;
Tallahatchie R.
Tributaries Unit,
n =16.
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2013 Southeastern Naturalist Vol. 12, No. 4
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the Yocona R. drainage than in the Tallahatchie R. drainage (Table 2). Mean sizes
were similar among units in the Yocona R. drainage, but showed more variation
among units in the Tallahatchie R. drainage.
Sex ratios were skewed toward females and significantly deviated from expected
1:1 sex ratios in all units (Table 2). Sex ratios were similar among units in the Yocona
R. drainage. The percentage of males in those units was nearly identical and had
broadly overlapping confidence intervals. The Tallahatchie R. drainage units were
more variable. The percentage of males in the Tallahatchie R. Tributaries Unit were
lower than the percentage recorded in the Tippah River Unit and their confidence
intervals did not overlap. In the Cypress Creek Unit, the percentages of males were
intermediate between these two units. The percentage of males was similar in the Yocona
R. and Tallahatchie R. drainages, and the confidence intervals overlapped.
Among those locations where Yazoo Darters were present, ordination of abiotic
variables described a stream-size gradient, and an aquatic vegetation, stream flow,
and stream incision gradient. The first two PCA axes were regarded as interpretable,
with axis 1 and axis 2 explaining 34.0% and 18.4% of the dataset variance,
respectively. PCA axis 1 was positively correlated with forest canopy and bank
height and negatively correlated with watershed area, wetted width, and water
depth. PCA axis 2 was positively correlated with aquatic vegetation and streamcurrent
velocity and negatively correlated with bank height and forest canopy
Table 2. Mean ± SE standard length (SL, mm) of female and male Yazoo Darters by watershed unit
and drainage as well as of female and male Yazoo Darters combined by watershed unit and drainage,
percentage of male darters ± 95% confidence intervals (CI) in the sample, and male to female sex
ratios. Different superscripted letters indicate significant differences in length between Yazoo Darters
in the Yocona R. and Tallahatchie R. drainages.
Otoucalofa Tallahatchie R. Cypress Tippah
Creek Yocona R. Yocona R. Tributaries Creek River Tallahatchie R.
Unit Unit Drainage Unit Unit Unit Drainage
Female
mean SL 43.19 41.69 42.32A 38.45 42.87 40.15 39.79B
± SE 0.776 0.683 0.517 0.213 0.545 0.608 0.244
n 28 39 67 133 51 56 240
Male
mean SL 47.69 47.88 47.80A 43.26 47.24 46.44 45.48B
± SE 2.027 1.65 1.26 0.748 1.252 0.737 0.522
n 13 17 30 25 14 32 71
Male and female
mean SL 44.62 43.57 44.014A 39.21 43.81 42.44 41.09B
± SE 0.882 0.782 0.585 0.256 0.549 0.57 0.261
n 41 56 97 158 65 88 311
Percentage males 31.7 30.4 30.9 15.8 21.5 36.4 22.8
± 95% CI 14.24 12.04 9.2 5.69 9.99 10.05 4.67
Males:females 01:02.0 01:02.3 01:02.2 01:05.3 01:03.6 01:01.8 01:03.4
c² goodness of fit 5.48 8.64 73.82 21.06 6.55
Exact P 0.028 0.005 less than 0.0001 less than 0.0001 0.014
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2013 Southeastern Naturalist Vol. 12, No. 4
(Table 3). Yazoo Darter abundance was not correlated with site scores of PCA axis 1
(r = 0.048, P ≤ 0.21) or axis 2 ( r = 0.004, P ≤ 0.7).
Ordination of all sites sampled described a stream-size gradient, and a depth,
woody debris, and stream-incisement gradient. Again, the first 2 PCA axes were
regarded as interpretable, with axis 1 and axis 2 explaining 25.1% and 21.5% of the
dataset variance, respectively. Axis 1 was positively correlated with watershed area,
wetted width, and stream order, and negatively correlated with canopy, detritus, and
small woody debris. Axis 2 was positively correlated with depth, large and small
woody debris, and detritus, and negatively associated with bank height, stream
order, and canopy. Presence/absence data for Yazoo Darters were not significantly
associated with PCA axis 1 (c2 = 0.14, P ≤ 0.71) or axis two ( c2 = 2.05, P ≤ 0.15).
Results from indicator species analysis show that 6 species were significantly associated
with Yazoo Darters: Noturus phaeus Taylor (Brown Madtom) (P < 0.001),
lamprey (P < 0.001), Etheostoma lynceum Hay (Brighteye Darter) (P < 0.002),
Etheostoma swaini Jordan (Gulf Darter) (P < 0.002), Percina sciera Swain (Dusky
Darter) (P < 0.002), and Hypentelium nigricans Lesueur (Northern Hog Sucker)
(P < 0.027) (Table 4). Of the 71 fish species we recorded in our study, 60 of them
occurred at least once at locations with Yazoo Darters.
Table 4. Fish species significantly associated with Yazoo Darters (indicator species analysis) showing
the number of locations (total locations, n = 93) where a species was sampled (n), the percent of sites
yielding Yazoo Darters where a species was sampled (%), the maximum indicator value (MI value),
and P-value.
Species n % MI value P-value
Lamprey spp. 35 59 45.2 0.0002
Brown Madtom 50 74 51.8 0.0003
Brighteye Darter 29 50 36.7 0.0017
Dusky Darter 47 68 45.9 0.0019
Gulf Darter 14 29 24.5 0.0022
Northern Hog Sucker 19 32 22.8 0.0272
Table 3. Loadings from principal components analysis (PCA) of abiotic variables for locations yielding
Yazoo Darters (abundance data; PCA 1) and for locations yielding and not yielding Yazoo Darters
(presence/absence data; PCA 2) sampled from 1999–2003.
Abiotic variable PCA 1 axis 1 PCA 1 axis 2 PCA 2 axis 1 PCA 2 axis 2
Stream order -0.7048 -0.4407 0.5767 -0.2967
Area -0.8588 -0.2841 0.7186 0.4104
Width -0.8628 -0.2585 0.8398 0.3432
Depth -0.8087 0.3571 0.5367 0.6487
Velocity -0.4718 0.5036 0.5054 -0.1754
Detritus -0.4064 -0.222 -0.4718 0.5694
Small wood -0.5273 0.1426 -0.3803 0.7082
Bank height 0.3442 -0.4881 0.4034 -0.4513
Bank stability -0.3488 -0.2493 0.0622 0.1063
Large wood -0.5142 -0.344 -0.1574 0.7885
Aquatic vegetation -0.1455 0.8894 0.0266 0.149
Canopy 0.5134 -0.4532 -0.5957 -0.2721
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
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Discussion
Additional sampling efforts will likely result in the discovery of new localities
with Yazoo Darters, but our results and records from the combined database suggest
that few additional tributary populations are likely to be discovered. Our sampling
effort was mainly in the central and western portions of the Yocona and Tallahatchie
R. drainages within the known range of the Yazoo Darter. Sampling along the eastern
portions of the known range of Yazoo Darters has been less intense and appears
to have the highest potential to yield new tributary records for the species.
Land ownership patterns in relation to Yazoo Darter distribution paint a mixed
picture in terms of long-term persistence of the species. Many locations harboring
Yazoo Darters in the Tallahatchie R. drainage are in watersheds offering some
measure of protection due to state or federal management for timber, recreation, or
research. As such, these locations should be at substantially less risk of degradation
than streams traversing private land. In contrast, Yazoo Darter locations in the
Yocona R. and Otoucalofa Creek Units are on private lands and lack the protection
afforded by public or conservation ownership. In particular, Yazoo Darters apparently
occur in only 4 small tributaries of the Yocona R. Unit, and all 4 of these
tributaries are likely to be affected by continued urban expansion from the city of
Oxford, MS. The uppermost headwaters of 2 of these tributaries, Pumpkin and Yellow
Leaf Creeks, are on National Forest land. The other 2 tributaries, Taylor and
Morris Creeks, flow completely through privately owned lands and have been impacted
by development (K. Sterling, pers. observ.), and are subject to deforestation
and urban development. Locations within the Otoucalofa Creek Unit face pressure
from agricultural activities and from urbanization near the city of Water Valley, MS.
Our quantitative habitat analyses clearly indicated that Yazoo Darters consistently
occupy small, shallow, headwater streams, an observation also made
by others (Johnston and Haag 1996, Suttkus et al. 1994, Thompson and Muncy
1986). However, single young-of-the-year juvenile Yazoo Darters were captured
at two locations (sites 6814 and 6819) in the Tallahatchie R. Canal in late summer.
These two individuals may have been waifs from tributaries that were displaced
downstream during a high-flow event, or were moving out of headwater streams
to avoid adverse low-flow conditions of late summer. Alternatively, these fish may
evidence a generalized movement of juvenile Yazoo Darters from headwaters to
larger streams. If juvenile Yazoo Darters commonly disperse across drainages at
around 6 months of age, then we would expect the numerous other fish samples
from mainstem reaches of the Tippah R. (and large channelized tributaries like
Potts Creek), Tallahatchie R., Yocona R., and Otoucalofa Creek to have also contained
Yazoo Darters. We doubt that the Tallahatchie R. Canal provides quality
Yazoo Darter habitat, and we do not believe that the degraded habitat present in the
Canal could support reproducing, permanent populations. Two Yazoo Darters were
sampled (Ross 2001) from the mainstem of Otoucalofa Creek (site 4984) at the
confluence with Sarter Creek. Because the sample was taken in May, we doubt that
these individuals could have been juveniles. The watershed area above this location
is only about 110 km2, and channelized portions of the stream appear on maps
829
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
to be well downstream. This location may be considered normal habitat for Yazoo
Darters. Alternatively, these individuals may have come from Sarter Creek. In any
case, it is likely that Yazoo Darters occasionally venture into uncharacteristically
large streams as evidenced by these unusual occurrences. Another location (site
1129) farther upstream in the mainstem Otoucalofa is certainly typical Yazoo Darter
habitat because that section of Otoucalofa Creek is a second-order stream with a
watershed area of about 9 km2. Yazoo Darters were also present in seven samples
from near the headwaters of Cypress Creek in the mainstem (sites 6865 and 6867).
At these locations, Cypress Creek is a second-order stream with a watershed area
of about 15.5 km2 and it also appears to be suitable Yazoo Darter habitat.
Our measures of abundance did not yield any clear patterns within or among
watershed units. Because variation was relatively high, it seems likely that repeated
sampling over time would be needed to precisely estimate relative abundances
among watersheds.
Male Yazoo Darters were significantly larger than females, a pattern consistent
with other snubnose darter species (Boschung et al. 1992, Powers and Mayden 2003,
Suttkus and Etnier 1991). However, mean size of male and female Yazoo Darters
from the Yocona R. drainage was greater than mean size in the Tallahatchie R.
drainage. This finding may reflect genetic differences between populations in the
respective rivers as revealed by MtDNA analysis (Powers and Warren 2009) but may
also indicate disparity between the two drainages in factors such as food availability
or survivorship. However, we are unaware of differences in the two drainages (e.g.,
productivity, predation) that would affect growth or survivorship. The size disparity
between populations in the two drainages deserves further investigation.
Sex ratios were skewed toward females in all watershed units analyzed. This
finding is consistent with more spatially and sample-limited work for Yazoo Darters
(Johnston and Haag 1996). The pattern is typical of most other snubnose darters for
which sex ratios have been reported (Carney and Burr 1989, Khudamrongsawat and
Kuhajda 2007, Page and Mayden 1981, Suttkus and Bailey 1993, but see Clayton
1984 on Etheostoma baileyi Page and Burr [Emerald Darter]). We did not examine
sex ratios by age class, but in one population, sex ratios of Yazoo Darters at hatching
were close to 1:1 (Johnston and Haag 1996), as in some other snubnose darters
(Barton and Powers 2010, Carney and Burr 1989), and then, presumably, malebiased
mortality in the first year skewed sex ratios. Because skewed sex ratios can
dramatically affect effective population sizes (Allendorf and Luikart 2007), further
investigation of the driving mechanisms behind differential survival in the Yazoo
Darter is warranted.
The relatively low variation in our measures of stream order, watershed area,
and current velocity indicate that Yazoo Darters are generally constrained to smaller
headwater streams, a conclusion supported by nearly all known records of Yazoo
Darter samples as discussed previously. Thus, headwater habitat preservation and
restoration will be essential to help ensure persistence of the species. Investigation
of the mode and timing of dispersal between headwater streams is needed, as is
identification of potential barriers to dispersal.
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
830
We did not find any relationships between Yazoo Darter abundance and measured
habitat variables. It is possible that we failed to measure some variable of
overriding importance, such as water temperature or dissolved oxygen, which
would explain the insignificant results of the logistic regression. Another, not
mutually exclusive, possibility is that the habitat requirements of Yazoo Darters
represent thresholds. In this case, once the habitat requirements of the species
are met, relative abundance is not influenced by variation in habitat. This theory
may explain why we did not find a strong correlation between abundance and indicators
of stream incision (bank height and bank stability). Other factors such as
the influence of groundwater and springs may be important (Suttkus et al. 1994).
Some of the densest populations we sampled were in streams receiving spring flow
(e.g., Chewalla Creek tributary, site 6851; Big Spring tributary, site 6852; and Bay
Springs Branch, site 7171; see Appendix 1). Our attempt to quantify habitat may
have been at too large a spatial scale (120–300 m) because Yazoo Darters were not
evenly distributed throughout a stream reach. As a result, we may have been measuring
variables in unsuitable habitat as well as suitable habitat within our study
reaches. Johnston and Haag (1996) concluded that Yazoo Darters were habitat
generalists, but their study focused on a single population and, given their sample
numbers, the habitat was likely of relatively high quality and not limiting. Based
on the patchy nature of the Yazoo Darter’s spatial distribution within and among
watersheds, and our field observations of streams and mesohabitat in which it does
and does not occur, we feel the species is likely habitat-limited at landscape and
even meso- or microhabitat scales.
Across their ranges, species associates of Yazoo Darters occupy a range of
stream sizes from the smallest headwater streams (Brown Madtom, lamprey) to
medium-sized streams and small rivers (Brighteye Darter, Dusky Darter, Northern
Hog Sucker) (Etnier and Starnes 1993, Ross 2001). Our study was not designed to
detect or describe fine-scale ecological interactions or even ecological similarities
among these species. Even so, all of them co-occurred with the Yazoo Darter in
small stream habitats more often than expected by chance, and some shared ecological
traits among the associates are apparent. Similar to the Yazoo Darter, most
of the associates are strongly rheophilic, benthic, and small bodied. Even for the
largest associate, the Northern Hog Sucker, our catch was composed almost entirely
of juveniles (M.L. Warren, pers. observ.). Interestingly, within the Yazoo R. basin,
the brook lampreys encountered in our study streams (i.e., predominantly ammocoetes
of Least Brook Lamprey), show a distribution nearly identical to that of the
Yazoo Darter, and they are confined to portions of the Little Tallahatchie, Tippah,
and Yocona rivers in the Northern Hilly Gulf Coastal Plain Ecoregion (Ross 2001).
Northern Hog Suckers are similarly distributed in the area, being absent from
most channelized main channels (Ross 2001). The Brown Madtom and Brighteye
Darter are more widespread in the Yazoo R. basin than the Yazoo Darter, but most
records are along a north–south band describing the Northern Hilly Gulf Coastal
Plain Ecoregion (Ross 2001). At the level of meso-habitat, the Brown Madtom often
inhabits tiny streams and is strongly associated with stream flow and complex
831
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
habitat provided by woody structure that is missing from channelized and highly
incised habitats in the region (Chan and Parsons 2000, Monzyk et al. 1997). Although
many populations were likely eliminated by channelization within its range,
the Brighteye Darter appears to be most common in better-quality streams that are
least affected by channelization (Etnier and Starnes 1993). Similarly, Gulf Darters
are associated with flow, woody debris, and Sparganium spp. (bur-reeds), which
we and others noted is often present and abundant at sites with high Yazoo Darter
densities (Suttkus et al. 1994). Overall, we believe this group of frequent associates
is an indicator of high-quality habitats associated with the Yazoo Darter, and their
confinement to particular sites is a result of stream degradation over much of the
stream system in the region.
Our data show that populations of the Yazoo Darter in the Yocona R. drainage
are far less numerous relative to the Tallahatchie R. drainage, and these
populations have no protection from continued urban development or habitat
modification (i.e., impoundments and stream alteration on private lands). Genetic
work indicates that Yazoo Darters in the Yocona R. drainage have lower allelic
richness, observed heterozygosity, and gene diversity relative to Yazoo Darters
in the Tallahatchie R. drainage, and that they are isolated within tributary streams
(Sterling et al. 2012). Personal observations (K. Sterling) suggest that suitable
Yazoo Darter habitat within these highly modified tributary streams is uncommon
due to habitat homogenization. For these reasons, and because populations of
Yazoo Darters in the Yocona R. drainage are genetically distinct from those in the
Tallahatchie R. drainage (Powers and Warren 2009), management action should
be focused on Yocona R. drainage populations. Standardized, quantitative habitat
surveys should be conducted throughout each Yocona R. drainage watershed
that harbors Yazoo Darters in an effort to provide baseline data for monitoring
efforts. This should also include quantification of watershed-scale land-use and
land-cover variables to track changes due to urbanization. Our own first efforts
at modeling Yazoo Darter and habitat associations should be improved upon. If
satisfactory models can be produced, results could be coupled with results from
stream habitat surveys, results from this study, and the existing literature to build
a stream-habitat-restoration strategy.
Within the Tallahatchie R. drainage, sampling records indicate Yazoo Darter
populations have not been extirpated. Even so, because sampling records only
extend back several decades for most populations, this finding should be regarded
with caution. Risk of extirpation within the entire drainage in the near term is
somewhat minimized due to the fact that many populations are located on or near
state- or federally managed lands. However, because Yazoo Darters are genetically
differentiated among headwater tributaries within drainages, and the mainstem Tallahatchie,
Tippah, and Yocona rivers are apparently barriers to dispersal (Sterling
et al. 2012), continued monitoring of populations is warranted. Extirpation of any
headwater population would result in loss of important genetic diversity and would
preclude future efforts via human-assisted migration to increase genetic diversity
and adaptive potential in the face of a changing climate.
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
832
The Yazoo Darter is a charismatic, endemic species that greatly enhances the
natural heritage of northern Mississippi where few vertebrate endemics occur. Preservation
of this fish in the short-term is an achievable goal that should be a priority
for federal and state agencies and the public.
Acknowledgments
We thank the many people who generously contributed to this work by assisting in the
field and laboratory, sharing information and ideas, providing logistical support, and offering
numerous other professional courtesies: S. Adams, H. Bart, M. Bland, A. Clingenpeel, A.
Commens-Carson, D. Drennen, T. Fletcher, W. Haag, H. Halverson, C. Harwell, C. Jenkins,
C. Kilcrease, S. Krieger, D. Martinovic, F. McEwen, G. McWhirter, A. Pabst, S. Powers, R.
Reekstin, M. Roberts, S. Ross, T. Slack, and L. Staton. Two anonymous reviewers contributed
greatly toward improving this manuscript and deserve thanks for their efforts. We are
also grateful to numerous private landowners who graciously granted permission to survey
streams on their property. The study was supported by a USDA Forest Service Chief’s grant,
and funds from National Forests of Mississippi, Southern Region, USDA Forest Service;
the Center for Bottomland Hardwoods Research, Southern Research Station, USDA Forest
Service; the US Fish and Wildlife Service, Mississippi Ecological Services Office; and a state
wildlife grant from the Mississippi Museum of Natural Science, Jackson, MS.
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Appendix 1. Records for all known samples yielding Yazoo Darters (1952–2011) listed by watershed unit (Fig. 1) and site. T = total number
of Yazoo Darters sampled; CPUE = fish per hour; Unit = watershed unit; UT = Unnamed tributary. OC = Otoucalofa Creek; YR = Yocona
River; CC = Cypress Creek; TRT = Tallahatchie River Tributaries; TR= Tippah River. USFS = USDA Forest Service; UMFS = University
of Mississippi Field Station; USACE = US Army Corps of Engineers; WDSP = Wall Doxey State Park.
Site ID Date Source T CPUE Unit Stream Ownership Lat Long
1090 10/10/1985 Ross et al. 2001 1 OC Dickey Creek Private 34.168 89.438
1090 6/17/2009 2009–2011 data 1 8.49 OC Dickey Creek Private 34.168 89.438
640 6/19/2009 2009–2011 data 7 54.90 OC Johnson Creek Private 34.123 89.641
749 6/14/1989 Suttkus et al. 1994 26 OC UT Otoucalofa Creek Private 34.141 89.589
749 5/18/1990 Suttkus et al. 1994 2 OC UT Otoucalofa Creek Private 34.141 89.589
749 4/12/1992 Suttkus et al. 1994 1 OC UT Otoucalofa Creek Private 34.141 89.589
7177 6/30/2009 2009–2011 data 2 21.88 OC Spring Creek Private 34.153 89.529
7178 6/18/2009 2009–2011 data 2 8.91 OC Moore Creek Private 34.156 89.548
7179 6/18/2009 2009–2011 data 16 116.10 OC Mill Creek Private 34.166 89.520
7186 6/19/2009 2009–2011 data 13 144.00 OC UT Otoucalofa Creek Private 34.125 89.610
841 6/15/1989 Suttkus et al. 1994 3 OC Gordon Branch Private 34.140 89.549
841 6/30/2009 2009–2011 data 15 169.80 OC Gordon Branch Private 34.140 89.549
990 7/23/1985 Ross et al. 2001 4 OC Smith Creek Private 34.138 89.474
1129 7/8/1985 Ross et al. 2001 14 OC Otoucalofa Creek Private 34.133 89.412
1129 7/8/1985 Ross et al. 2001 12 OC Otoucalofa Creek Private 34.133 89.412
4984 5/14/1986 Ross et al. 2001 2 OC Otoucalofa Creek Private 34.162 89.512
5034 7/10/1985 Ross et al. 2001 1 OC Shippy Creek Private 34.153 89.433
6858 6/11/1999 1999–2003 data 2 8.87 YR Pumpkin Creek Private 34.327 89.397
6859 6/11/1999 1999–2003 data 6 52.68 YR Pumpkin Creek Private 34.339 89.384
5028 5/6/1974 Suttkus et al. 1994 3 YR Pumpkin Creek Private 34.286 89.445
1164 5/24/1952 Suttkus et al. 1994 11 YR Pumpkin Creek Private 34.327 89.397
1164 4/17/1969 Suttkus et al. 1994 7 YR Pumpkin Creek Private 34.327 89.397
1164 5/10/1988 Suttkus et al. 1994 22 YR Pumpkin Creek Private 34.327 89.397
1164 10/22/1988 Suttkus et al. 1994 14 YR Pumpkin Creek Private 34.327 89.397
1164 7/27/1989 Suttkus et al. 1994 4 YR Pumpkin Creek Private 34.327 89.397
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2013 Southeastern Naturalist Vol. 12, No. 4
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Site ID Date Source T CPUE Unit Stream Ownership Lat Long
1164 6/16/2009 2009–2011 data 1 5.19 YR Pumpkin Creek Private 34.327 89.397
1164 5/18/1990 Suttkus et al. 1994 3 YR Pumpkin Creek Private 34.327 89.397
7181 6/30/2009 2009–2011 data 1 8.28 YR UT of Pumpkin Creek Private 34.291 89.440
6860 9/1/2009 1999–2003 data 2 11.98 YR Yellow Leaf Creek Private 34.368 89.428
6861 6/9/1999 1999–2003 data 5 34.04 YR Yellow Leaf Creek Private 34.375 89.421
6862 6/9/1999 1999–2003 data 3 18.00 YR Yellow Leaf Creek Private 34.374 89.421
6863 6/9/1999 1999–2003 data 7 81.55 YR Yellow Leaf Creek Private 34.379 89.413
765 5/11/1988 Suttkus et al. 1994 10 YR UT of Taylor Creek Private 34.123 89.641
765 6/26/2009 2009–2011 data 12 64.96 YR UT of Taylor Creek Private 34.123 89.641
768 8/20/1991 Ross et al. 2001 1 YR Taylor Creek Private 34.271 89.580
7176 3/24/1993 Johnston and Haag 1996 24 YR Morris Creek Private 34.300 89.549
7176 4/26/1993 Johnston and Haag 1996 9 YR Morris Creek Private 34.300 89.549
7176 3/11/1994 Johnston and Haag 1996 40 YR Morris Creek Private 34.300 89.549
7176 4/7/1994 Johnston and Haag 1996 27 YR Morris Creek Private 34.300 89.549
7176 5/2/1994 Johnston and Haag 1996 11 YR Morris Creek Private 34.300 89.549
7176 5/17/1994 Johnston and Haag 1996 8 YR Morris Creek Private 34.300 89.549
7180 9/24/1993 Johnston and Haag 1996 21 YR Morris Creek Private 34.283 89.544
7180 10/21/1993 Johnston and Haag 1996 18 YR Morris Creek Private 34.283 89.544
7180 11/19/1993 Johnston and Haag 1996 19 YR Morris Creek Private 34.283 89.544
7180 12/14/1993 Johnston and Haag 1996 10 YR Morris Creek Private 34.283 89.544
7180 1/12/1994 Johnston and Haag 1996 21 YR Morris Creek Private 34.283 89.544
7180 2/23/1994 Johnston and Haag 1996 11 YR Morris Creek Private 34.283 89.544
7180 5/20/1993 Johnston and Haag 1996 14 YR Morris Creek Private 34.283 89.544
7180 3/23/1994 Johnston and Haag 1996 10 YR Morris Creek Private 34.283 89.544
7180 6/25/1993 Johnston and Haag 1996 17 YR Morris Creek Private 34.283 89.544
7180 4/20/1994 Johnston and Haag 1996 11 YR Morris Creek Private 34.283 89.544
7180 7/26/1993 Johnston and Haag 1996 22 YR Morris Creek Private 34.283 89.544
7180 8/27/1993 Johnston and Haag 1996 30 YR Morris Creek Private 34.283 89.544
7180 3/24/1993 Johnston and Haag 1996 19 YR Morris Creek Private 34.283 89.544
7180 4/26/1993 Johnston and Haag 1996 5 YR Morris Creek Private 34.283 89.544
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2013 Southeastern Naturalist Vol. 12, No. 4
Site ID Date Source T CPUE Unit Stream Ownership Lat Long
7180 5/2/1994 Johnston and Haag 1996 4 YR Morris Creek Private 34.283 89.544
7180 5/17/1994 Johnston and Haag 1996 8 YR Morris Creek Private 34.283 89.544
7180 6/2/2009 2009–2011 data 11 50.83 YR Morris Creek Private 34.282 89.543
7180 9/10/2009 2009–2011 data 13 67.05 YR Morris Creek Private 34.282 89.543
7180 1/14/2010 2009–2011 data 3 10.56 YR Morris Creek Private 34.282 89.543
7180 4/23/2010 2009–2011 data 12 YR Morris Creek Private 34.282 89.543
7180 9/10/2010 2009–2011 data 8 YR Morris Creek Private 34.282 89.543
7180 3/2/2011 2009–2011 data 4 YR Morris Creek Private 34.282 89.543
7180 7/1/2011 2009–2011 data 25 YR Morris Creek Private 34.282 89.543
6865 3/9/1982 Thompson and Muncy 1986 1 CC Cypress Creek Private 34.393 89.286
6865 6/1/1999 1999–2003 data 4 30.13 CC Cypress Creek Private 34.393 89.286
6865 4/1/2009 2009–2011 data 11 23.02 CC Cypress Creek Private 34.393 89.286
6867 6/4/1999 1999–2003 data 9 62.53 CC Cypress Creek Private 34.382 89.298
6867 7/23/2009 1999–2003 data 4 23.41 CC Cypress Creek Private 34.382 89.298
6867 3/30/2009 2009–2011 data 2 12.83 CC Cypress Creek Private 34.382 89.298
6867 4/7/2009 2009–2011 data 1 6.50 CC Cypress Creek Private 34.382 89.298
6875 10/17/1980 Thompson and Muncy 1986 9 CC Puskus Creek Private 34.396 89.372
6875 7/27/1981 Thompson and Muncy 1986 13 CC Puskus Creek Private 34.396 89.372
6875 9/29/1983 Thompson and Muncy 1986 12 CC Puskus Creek Private 34.396 89.372
6875 6/2/1999 1999–2003 data 6 44.08 CC Puskus Creek Private 34.396 89.372
6875 3/25/2009 2009–2011 data 10 CC Puskus Creek Private 34.396 89.372
6874 6/2/1999 1999–2003 data 2 CC Puskus Creek USFS 34.394 89.371
6878 8/3/1999 1999–2003 data 1 4.89 CC Puskus Creek USFS 34.445 89.336
6878 5/25/2000 1999–2003 data 1 3.99 CC Puskus Creek USFS 34.445 89.336
6878 7/24/2009 1999–2003 data 2 3.27 CC Puskus Creek USFS 34.445 89.336
7183 3/25/2009 2009–2011 data 5 27.48 CC Puskus Creek USFS 34.445 89.336
7183 3/30/2009 2009–2011 data 7 20.47 CC Puskus Creek USFS 34.445 89.336
1267 10/9/1970 Suttkus et al. 1994 17 CC Puskus Creek Private 34.415 89.372
1267 4/7/1972 Suttkus et al. 1994 2 CC Puskus Creek Private 34.415 89.372
1267 9/7/1973 Suttkus et al. 1994 4 CC Puskus Creek Private 34.415 89.372
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
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Site ID Date Source T CPUE Unit Stream Ownership Lat Long
1267 10/13/1973 Suttkus et al. 1994 11 CC Puskus Creek Private 34.415 89.372
1246 5/7/1974 Suttkus et al. 1994 2 CC Puskus Creek Private 34.411 89.377
1267 5/5/1976 Suttkus et al. 1994 22 CC Puskus Creek Private 34.415 89.372
1267 3/24/1982 Ross et al. 2001 2 CC Puskus Creek Private 34.415 89.372
1261 8/20/1991 Ross et al. 2001 6 CC Puskus Creek Private 34.411 89.374
1358 8/8/1984 Thompson and Muncy 1986 8 CC Puskus Creek Private 34.446 89.330
1358 7/18/1977 Ross et al. 2001 10 CC Puskus Creek Private 34.446 89.330
7170 8/8/1984 Thompson and Muncy 1986 4 CC UT Puskus Creek Private 34.439 89.385
7170 4/1/2009 2009–2011 data 3 6.07 CC UT Puskus Creek Private 34.439 89.385
6877 7/28/1999 1999–2003 data 2 9.07 CC UT Puskus Creek USFS 34.450 89.349
7182 4/14/2009 2009–2011 data 10 38.42 CC UT Puskus Creek Private 34.431 89.375
6879 9/14/1999 1999–2003 data 4 26.60 CC Bay Springs Branch UMFS 34.428 89.396
7171 2/21/1981 Thompson and Muncy 1986 1 CC Bay Springs Branch UMFS 34.428 89.395
7171 3/11/1982 Thompson and Muncy 1986 5 CC Bay Springs Branch UMFS 34.428 89.395
7171 3/14/1984 Thompson and Muncy 1986 2 CC Bay Springs Branch UMFS 34.428 89.395
7171 4/22/1984 Thompson and Muncy 1986 7 CC Bay Springs Branch UMFS 34.428 89.395
7171 10/29/1984 Thompson and Muncy 1986 11 CC Bay Springs Branch UMFS 34.428 89.395
1268 3/14/1980 Suttkus et al. 1994 4 CC Bay Springs Branch UMFS 34.425 89.386
1268 3/14/1980 Suttkus et al. 1994 3 CC Bay Springs Branch UMFS 34.425 89.386
7171 4/14/2009 2009–2011 data 10 52.17 CC Bay Springs Branch UMFS 34.428 89.395
7171 2/25/2009 2009–2011 data 156 120.50 CC Bay Springs Branch UMFS 34.428 89.395
7171 9/30/2009 2009–2011 data 74 49.79 CC Bay Springs Branch UMFS 34.428 89.395
7171 2/10/2010 2009–2011 data 190 117.40 CC Bay Springs Branch UMFS 34.428 89.395
1187 8/21/1991 Ross et al. 2001 5 CC UT Bay Springs Branch UMFS 34.421 89.388
1187 8/21/1991 Ross et al. 2001 4 CC UT Bay Springs Branch UMFS 34.421 89.388
7187 7/26/1984 Thompson and Muncy 1986 1 TRT Graham Mill Creek USACE 34.511 89.494
7187 11/6/1984 Thompson and Muncy 1986 2 TRT Graham Mill Creek USACE 34.511 89.494
5000 10/5/1995 Ross et al. 2001 1 TRT Lee Creek USACE 34.513 89.490
5000 5/20/1996 Ross et al. 2001 1 TRT Lee Creek USACE 34.513 89.490
5000 8/5/1996 Ross et al. 2001 1 TRT Lee Creek USACE 34.513 89.490
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2013 Southeastern Naturalist Vol. 12, No. 4
Site ID Date Source T CPUE Unit Stream Ownership Lat Long
971 5/19/1988 Suttkus et al. 1994 18 TRT Graham Mill Creek Private 34.503 89.490
971 5/18/1990 Suttkus et al. 1994 3 TRT Graham Mill Creek Private 34.503 89.490
971 4/12/1992 Suttkus et al. 1994 4 TRT Graham Mill Creek Private 34.503 89.490
5038 10/2/1995 Ross et al. 2001 8 TRT Bagley Creek USFS 34.495 89.413
5038 7/30/1996 Ross et al. 2001 1 TRT Bagley Creek USFS 34.495 89.413
5041 5/17/1996 Ross et al. 2001 2 TRT Bagley Creek USFS 34.482 89.409
5041 7/30/1996 Ross et al. 2001 8 TRT Bagley Creek USFS 34.482 89.409
6881 6/7/1999 1999–2003 data 1 8.35 TRT Bagley Creek USFS 34.481 89.405
6814 8/10/1999 1999–2003 data 1 2.36 TRT Tallahatchie R. Canal USACE 34.528 89.366
6819 8/9/1999 1999–2003 data 1 TRT Tallahatchie R. Canal Private 34.482 89.225
5188 10/4/1995 Ross et al. 2001 4 TRT Mitchell Creek Private 34.519 89.203
5188 5/21/1996 Ross et al. 2001 3 TRT Mitchell Creek Private 34.519 89.203
5188 8/6/1996 Ross et al. 2001 1 TRT Mitchell Creek Private 34.519 89.203
6820 8/2/1999 1999–2003 data 4 36.00 TRT Mitchell Creek Private 34.521 89.203
6853 7/29/1999 1999–2003 data 2 8.05 TRT Big Spring Creek Private 34.634 89.397
7175 4/7/1984 Thompson and Muncy 1986 6 TRT Big Spring Creek Private 34.721 89.406
1201 5/18/1988 Suttkus et al. 1994 4 TRT Big Spring Creek Private 34.711 89.394
1215 5/20/1981 Suttkus et al. 1994 8 TRT Big Spring Creek Private 34.711 89.391
1215 4/13/1984 Ross et al. 2001 9 TRT Big Spring Creek Private 34.711 89.391
1201 3/16/1983 Ross et al. 2001 4 TRT Big Spring Creek Private 34.711 89.394
1201 4/29/1982 Ross et al. 2001 2 TRT Big Spring Creek Private 34.711 89.394
1215 5/12/1982 Ross et al. 2001 2 TRT Big Spring Creek Private 34.711 89.394
6852 7/30/1999 1999–2003 data 32 163.90 TRT UT Big Spring Creek Private 34.663 89.412
6852 6/23/2009 2009–2011 data 73 411.30 TRT UT Big Spring Creek Private 34.663 89.412
6852 9/11/2009 2009–2011 data 24 131.50 TRT UT Big Spring Creek Private 34.663 89.412
6852 10/19/2009 2009–2011 data 52 TRT UT Big Spring Creek Private 34.663 89.412
6852 1/26/2010 2009–2011 data 62 185.80 TRT UT Big Spring Creek Private 34.663 89.412
6852 5/24/2010 2009–2011 data 51 TRT UT Big Spring Creek Private 34.663 89.412
6852 9/2/2010 2009–2011 data 54 TRT UT Big Spring Creek Private 34.663 89.412
6852 3/3/2011 2009–2011 data 45 TRT UT Big Spring Creek Private 34.663 89.412
K.A. Sterling, M.L. Warren, Jr., and L.G. Henderson
2013 Southeastern Naturalist Vol. 12, No. 4
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Site ID Date Source T CPUE Unit Stream Ownership Lat Long
6852 6/29/2011 2009–2011 data 11 TRT UT Big Spring Creek Private 34.663 89.412
6882 8/5/1999 1999–2003 data 5 43.27 TRT Lee Creek Private 34.473 89.446
6883 5/24/2000 1999–2003 data 5 22.75 TRT Lee Creek USFS 34.497 89.456
6883 8/5/1999 1999–2003 data 1 7.79 TRT Lee Creek USFS 34.497 89.456
4993 5/20/1996 Ross et al. 2001 3 TRT Lee Creek USFS 34.498 89.457
4993 7/28/1995 Ross et al. 2001 2 TRT Lee Creek USFS 34.498 89.457
4993 8/5/1996 Ross et al. 2001 1 TRT Lee Creek USFS 34.498 89.457
4997 5/7/1993 Ross et al. 2001 1 TRT Lee Creek USACE 34.513 89.491
5015 5/20/1993 Ross et al. 2001 2 TRT Lee Creek Private 34.499 89.465
7049 7/3/2002 1999–2003 data 4 TRT Mill Creek Private 34.546 89.227
1053 10/27/1973 Suttkus et al. 1994 6 TRT Little Spring Creek Private 34.642 89.464
1053 5/7/1974 Suttkus et al. 1994 1 TRT Little Spring Creek Private 34.642 89.464
1053 5/10/1988 Suttkus et al. 1994 1 TRT Little Spring Creek Private 34.642 89.464
7083 5/30/2003 1999–2003 data 1 2.58 TRT Little Spring Creek Private 34.642 89.464
7174 7/13/1984 Thompson and Muncy 1986 9 TRT Little Spring Creek WDSP 34.667 89.467
7174 8/17/1984 Thompson and Muncy 1986 3 TRT Little Spring Creek WDSP 34.667 89.467
7174 3/8/1985 Thompson and Muncy 1986 16 TRT Little Spring Creek WDSP 34.667 89.467
7184 4/2/2010 2009–2011 data 1 TRT Little Spring Creek USACE 34.576 89.475
1039 10/13/1973 Suttkus et al. 1994 1 TRT Little Spring Creek WDSP 34.66 89.467
7089 10/16/1981 Thompson and Muncy 1986 2 TRT Oak Chewalla Creek USACE 34.582 89.511
7089 9/14/1983 Thompson and Muncy 1986 1 TRT Oak Chewalla Creek USACE 34.582 89.511
7089 5/22/2003 1999–2003 data 1 3.12 TRT Oak Chewalla Creek USACE 34.582 89.511
942 5/18/1988 Suttkus et al. 1994 5 TRT Oak Chewalla Creek USACE 34.582 89.509
942 5/19/1988 Suttkus et al. 1994 5 TRT Oak Chewalla Creek USACE 34.582 89.509
942 5/18/1990 Suttkus et al. 1994 1 TRT Oak Chewalla Creek USACE 34.582 89.509
7090 5/22/2003 1999–2003 data 3 15.84 TRT Oak Chewalla Private 34.613 89.518
7091 5/23/2003 1999–2003 data 2 11.76 TRT Fice Creek Private 34.421 89.246
7173 8/26/1982 Thompson and Muncy 1986 10 TRT Blackwater Creek USACE 34.569 89.609
7173 8/11/1984 Thompson and Muncy 1986 5 TRT Blackwater Creek USACE 34.569 89.609
935 4/12/1992 Suttkus et al. 1994 1 TRT Hurricane Creek Private 34.446 89.509
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2013 Southeastern Naturalist Vol. 12, No. 4
Site ID Date Source T CPUE Unit Stream Ownership Lat Long
935 4/12/1992 Suttkus et al. 1994 14 TRT Hurricane Creek Private 34.446 89.509
935 5/19/1988 Suttkus et al. 1994 6 TRT Hurricane Creek Private 34.446 89.509
935 3/11/1989 Suttkus et al. 1994 64 TRT Hurricane Creek Private 34.446 89.509
935 5/18/1990 Suttkus et al. 1994 8 TRT Hurricane Creek Private 34.446 89.509
935 5/24/1992 Suttkus et al. 1994 13 TRT Hurricane Creek Private 34.446 89.509
4994 4/7/1984 Thompson and Muncy 1986 18 TRT Hurricane Creek Private 34.425 89.496
4994 4/18/1984 Thompson and Muncy 1986 4 TRT Hurricane Creek Private 34.425 89.496
4994 4/22/1984 Thompson and Muncy 1986 3 TRT Hurricane Creek Private 34.425 89.496
4994 11/10/1984 Thompson and Muncy 1986 4 TRT Hurricane Creek Private 34.425 89.496
7172 8/2/1984 Thompson and Muncy 1986 1 TRT Hurricane Creek Private 34.457 89.545
6821 – Randolph and Kennedy 1974 0 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 7/21/1999 1999–2003 data 6 15.35 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 6/25/2009 2009–2011 data 13 91.59 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 7/2/2009 2009–2011 data 25 91.93 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 10/21/2009 2009–2011 data 12 57.37 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 1/27/2010 2009–2011 data 13 24.55 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 5/26/2010 2009–2011 data 12 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 9/7/2010 2009–2011 data 27 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 3/15/2011 2009–2011 data 9 TR Yellow Rabbit Creek USFS 34.819 89.105
6821 6/30/2011 2009–2011 data 12 TR Yellow Rabbit Creek USFS 34.819 89.105
6822 – Randolph and Kennedy 1974 0 TR Yellow Rabbit Creek Private 34.773 89.144
6822 7/22/1999 1999–2003 data 4 15.50 TR Yellow Rabbit Creek Private 34.773 89.144
6827 7/12/1999 1999–2003 data 15 103.80 TR Wagner Creek Private 34.768 89.229
6829 6/17/1999 1999–2003 data 20 159.60 TR UT Tippah River USFS 34.708 89.255
6829 6/25/2009 2009–2011 data 11 110.90 TR UT Tippah River USFS 34.708 89.255
6830 6/17/1999 1999–2003 data 2 17.60 TR UT Tippah River Private 34.681 89.281
6832 7/9/1999 1999–2003 data 6 53.33 TR UT Tippah River USFS 34.660 89.287
6847 6/16/1999 1999–2003 data 13 112.80 TR Chewalla Creek Private 34.814 89.368
6849 6/21/1999 1999–2003 data 1 6.41 TR Chewalla Creek USFS 34.697 89.331
1325 5/20/1981 Suttkus et al. 1994 1 TR Chewalla Creek Private 34.767 89.349
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2013 Southeastern Naturalist Vol. 12, No. 4
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Site ID Date Source T CPUE Unit Stream Ownership Lat Long
6851 7/8/1999 1999–2003 data 28 209.60 TR UT Chewalla Creek USFS 34.733 89.303
7085 7/12/1984 Thompson and Muncy 1986 3 TR UT Chewalla Creek USFS 34.76 89.332
7085 7/2/2003 1999–2003 data 13 53.46 TR UT Chewalla Creek USFS 34.76 89.332
7085 6/24/2009 2009–2011 data 4 19.20 TR UT Chewalla Creek USFS 34.76 89.332
7085 6/25/2009 2009–2011 data 5 TR UT Chewalla Creek USFS 34.76 89.332
7185 6/24/2009 2009–2011 data 7 68.29 TR UT Chewalla Creek Private 34.725 89.305
1348 9/7/1968 Randolph and Kennedy 1974 1 TR UT Chewalla Creek Private 34.764 89.343
1550 2/24/1984 Thompson and Muncy 1986 2 TR Big Snow Creek Private 34.815 89.240
1687 9/11/1968 Randolph and Kennedy 1974 1 TR Rhoden Creek Private 34.757 89.169
1878 10/5/1968 Randolph and Kennedy 1974 3 TR Sorghum Creek Private 34.707 89.071
7053 5/24/2002 1999–2003 data 1 TR South Fork Chilli Creek Private 34.682 89.172
7053 9/1/2009 1999–2003 data 22 119.80 TR South Fork Chilli Creek Private 34.682 89.172
7053 10/20/2009 2009–2011 data 3 14.50 TR South Fork Chilli Creek Private 34.682 89.172
7053 1/15/2010 2009–2011 data 11 37.43 TR South Fork Chilli Creek Private 34.682 89.172
7053 5/27/2010 2009–2011 data 1 TR South Fork Chilli Creek Private 34.682 89.172
7053 9/8/2010 2009–2011 data 14 TR South Fork Chilli Creek Private 34.682 89.172
7053 3/4/2011 2009–2011 data 3 TR South Fork Chilli Creek Private 34.682 89.172
7053 6/29/2011 2009–2011 data 3 TR South Fork Chilli Creek Private 34.682 89.172
7080 7/2/2003 1999–2003 data 2 5.82 TR Shelby Creek Private 34.843 89.039