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2016 SOUTHEASTERN NATURALIST 15(1):102–114
Life History Traits of the Mirror Shiner, Notropis
spectrunculus, in Western North Carolina
Adric D. Olson1,* and Thomas H. Martin1
Abstract - We investigated the life history of Notropis spectrunculus (Mirror Shiner) at
4 locations in the Tennessee River drainage in western North Carolina that we sampled
monthly over 7 months. Specimens were collected by seining and examined to identify age,
growth, reproductive patterns, and feeding habits. Sexual maturity occurred at approximately
1 year of age. Spawning occurred from April to July with 13–331 mature oocytes (mean
= 115.53, SD = 75.36), and male breeding coloration was present in specimens collected in
May, June, and July. Gut contents consisted mainly of insect fragments, primarily Coleoptera
and Diptera. Fish were found to inhabit water 0.5–0.75 m deep with sandy substrate,
directly below flow-disrupting objects.
Introduction
Notropis spectrunculus (Cope) (Mirror Shiner) is a stream-dwelling fish of the
family Cyprinidae found in tributaries of the watershed of the Tennessee River
in deep pools just below riffles, rocky pools, and runs (Etnier and Starnes 1993).
Breeding coloration in males has been observed from mid-May to late-June, but
Etnier and Starnes (1993) state that the biology of the Mirror Shiner remains generally
unreported. Etnier and Starnes also noted that according to Mayden (1989), the
Mirror Shiner is most closely related to Notropis volucellus (Cope) (Mimic Shiner),
a theory confirmed since by Cashner et al. (201 1).
The Pigeon River, part of the Tennessee River watershed in North Carolina and
Tennessee, is a river system degraded by human use. A kraft paper and pulp mill
exists in the city of Canton, NC, which lies near the headwaters of the Pigeon River.
The mill diverted flow and polluted the river with effluent and artificially heated
water, destroying the natural ecosystem and tainting the color and smell of the Pigeon
River downstream into Tennessee. The mill underwent more than $300 million
in renovations and improvements between 1988 and 1994 to clean the water it was
releasing and decrease the amount of color and dioxins it releases and the volume
of water it uses (Bartlett 1995). Although the mill has significantly reduced its water
usage and the chemical pollutants it releases into the river, 2 low-head dams on
the Pigeon River located at the mill in Canton may prevent natural recolonization
below the mill from populations of fish from above the mill (LaVoie 2007), and issues
remain concerning the color and temperature of discharged water from the mill
(Hyatt 2010). Although the tributaries of the degraded section were not polluted by
the mill, recolonization of this reach of Pigeon River did not occur, potentially due
to a lack of population numbers in these tributaries or habitat characteristics that
1Department of Biology, Western Carolina University, Cullowhee, NC, 28723. *Corresponding
author - adric.olson@ttu.edu.
Manuscript Editor: Carol Johnston
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prevent migration of the fish. Furthermore, recolonization from below the mill is
also impossible due to a large downstream dam and reservoir.
The Pigeon River Recovery Project, jointly supported by the University of Tennessee,
the Tennessee Valley Authority, the Tennessee Department of Environment
and Conservation, the North Carolina Division of Water Quality, Blue Ridge Paper
Products, and the North Carolina Wildlife Resources Commission, has been trying to
reintroduce fish species thought to have been extirpated from the Pigeon River downstream
of Canton in an effort to restore naturally reproducing populations (University
of Tennessee Pigeon River Recovery Project 2004). The project has had success
establishing reproducing populations of some of the 24 target species, such as Etheostoma
zonale (Cope) (Banded Darter), Notropis leuciodus (Cope) (Tennessee Shiner),
Notropis rubricroceus (Cope) (Saffron Shiner), Hybopsis amblops (Rafinesque) (Bigeye
Chub), Percina evides (Jordan and Copeland) (Gilt Darter), Notropis telescopus
(Cope) (Telescope Shiner), Notropis photogenis (Cope) (Silver Shiner), and Notropis
micropteryx (Cope) (Highland Shiner). However, attempts to reintroduce other species,
including the Mirror Shiner, have been difficult here and elsewhere.
Between 2004 and 2012, more than 6000 Mirror Shiner individuals were relocated
to habitat below the mill in Canton, NC (J. Coombs, University of Tennessee
at Knoxville, Knoxville, TN, 2010 pers. comm.). However, unlike other fishes
reintroduced by the project, none of these individuals were recovered in surveys
conducted later in the year of the release. The lack of success in recapturing this
species, relative to the success in recapturing other reintroduced species, suggests
they may be experiencing severe mortality or dispersal out of the area. However,
little is known about the biology of the Mirror Shiner and so it is difficult to speculate
on possible reasons for the apparent failure of the reintroduction.
Collecting more information about Mirror Shiners could be crucial to the reintroduction
efforts for the species. Therefore, this study describes some basic
life-history traits of the Mirror Shiner such as age, growth rate, diet, fecundity, and
habitat use.
Field-site Description
We identified focal sites with Mirror Shiners at 4 locations in Western North
Carolina: the Pigeon River upstream of Canton, Hominy Creek, and the Tuckasegee
River at East Laport and at Wilmot (Table 1, Fig. 1). We determined upstream land
Table 1. Field sites where Mirror Shiners were captured in western North Carolina showing locations
and characteristics.
Pigeon River Tuckasegee River Tuckasegee River Hominy Creek
Characteristic at Canton at East Laport at Wilmot at Candler
Latitude 35°31'31"N 35°17'49"N 35°24'14"N 35°32'07"N
Longitude 82°50'25"W 83°08'52"W 83°18'47"W 82°41'37"W
River width at field site ~30 m ~30 m ~40 m ~5 m
Average discharge in 2010 381.7 cfs 502.1 cfs 1007 cfs No gage
Upstream use Agriculture Agriculture, Agriculture, Sparse
residential residential, residential
paper mill
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use by both direct observation and through the use of aerial photos. Due to changing
river conditions, we did not sample the same location at each focal site each
trip, instead, we sampled eddies immediately below riffle areas. Although all of
our sampling locations within the 4 sites were in the same reach of each river, the
sampling location could be as far as 30 m from the previous sample location. Water
depth at the nearest USGS gaging station, obtained from www.usgs.gov, was noted
for each sampling event (USGS 2011).
Methods
We undertook field sampling once a month with a minimum of 2 weeks between
sampling events, following the approach in Yanchis (1993): we sampled sites with
a 5-m seine with 3-mm mesh size until 10 target-species fish had been caught or
an hour and a half of sampling was complete. Captured fish were euthanized with an
overdose of tricaine methanesulfonate, MS-222, and immediately placed on ice in
a cooler. In addition, we noted any observable breeding coloration. After returning
from the field, we stored fish in a freezer for future dissection .
During each sampling event, we noted the specific location in the stream where
specimens were captured, the substrate type at each collection location, and water
Figure 1. Map showing location of field sites in Western North Carolina in relation to major
rivers and cities. Sites are marked with triangles and correspond as follows: (1) Hominy
Creek at Candler, (2) Pigeon River at Canton, (3) Tuckasegee River at East Laport, and (4)
Tuckasegee River at Wilmot.
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depth at the nearest USGS gage. We measured water depth and water temperature
at the collection site and the distance from the sampling site to the nearest in-stream
obstacles such as riffles or boulders. We used these observations to compare river
characteristics of Mirror Shiner habitat.
Additionally, we sampled a 4-mile stretch of the Tuckasegee River at potential
Mirror Shiner habitat locations and at each noted substrate type, water depth, and
distance to flow-disrupting objects. We surveyed 14 sites with this method, which
aided with characterization of the microhabitats in which Mirror Shiners tend to be
found. We surveyed 1 site—the Tuckasegee River at Wilmot—using a surveyor’s
level to provide a detailed bathymetric map representative of the habitat occupied
by Mirror Shiners.
For dissection, specimens were slowly warmed to room temperature in a warmwater
bath. We took notes on coloration and measured standard length (SL) and
wet weight (to the nearest mm and nearest 0.01 g, respectively). We first made an
incision anteriorly from the anus to the pelvic girdle into each specimen, and then
another dorsally from each end of this first incision in order to allow access to the
internal organs of the fish for observation. We sexed fish by visual inspection of
gonads, except in small fish with poorly developed gonads where sex could not
be reliably determined. Finally, we removed the alimentary canal of each fish and
inspected the contents using a dissecting microscope. We identified to Order all
macroinvertebrates found inside and noted the presence of detritus and material
containing chloroplasts.
We used a chi-square goodness-of-fit test to test for departure from a 1:1 sex
ratio. We removed the gonads and weighed them to the nearest 0.01 g, and then
computed the gonadosomatic indices (GSI) according to the formula (Crim and
Glebe 1990):
GSI = gonad weight (g) / body weight (g) x 100
Eggs were removed from the body cavity and counted. We used an ANOVA followed
by a Tukey’s HSD test to compare GSI between months. We also utilized an
ANOVA, excluding outlying values, to test for a significant relationship between
egg count and size. Differences in the number of fish observed with a particular diet
item between sites were tested with a G-test of homogeneity (as in Sokal and Rohlf
1981). To determine if a sex-dependent growth rate existed, we tested the statistical
significance of the interaction term in an ANOVA relating standard length to age
and sex. We used pair-wise analyses to test for differences in growth rates among
sites, and Holm’s sequentially rejective method to control experiment-wise error
rates (Holm 1979).
We determined age by inspection of monthly length–weight scatterplots for all
fish caught. We interpreted distinct clusters of points to represent different ages.
Age, determined in this way, was added to month caught after birth month to determine
age of specimens (Holder and Powers 2010). We determined birth month
by analyzing GSI, breeding coloration, and the first presence of young-of-year in
the samples.
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Length at age was fit to a von Bertalanf fy growth model:
E[L|t] = L∞(1 - e-K(t - t0)),
where E[L|t] is the expected or average length at age t, L∞ is the asymptotic average
length, K is the Brody growth rate coefficient (units are year-1), and t0 is a modeling
artifact that represents the age when the average length was zero (the model was
fit using the FSA package for R, Ogle 2011). We computed all statistical analyses
using R (version 2.14.1; R Core Team 2013) or Microsoft Excel (version 2010),
with alpha = 0.05.
Results
We collected and dissected 238 Mirror Shiners. All fish were captured over
sandy substrate, in pools 0.5 to 1.0 m in depth where the beginning of the pool was
1.0 m or less in distance to a water-flow altering obstacle in the river. We sexed 81
individuals: 47 male and 34 female. This sex ratio was not significantly different
from a 1:1 ratio (P = 0.15). All individuals able to be sexed were at least 1 year old.
The largest individual was a female of 77 mm SL and 2.96 g total weight. The largest
male was 71 mm SL and 2.39 g total weight. However, the largest individuals of
each sex were not the oldest. The oldest individual was a 46-month-old male. The
oldest female was 37 months old. Weight increased with standard length according
to the equation weight (g) = 0.000002 * length (mm) ^ 3.3048 (r2 = 0.91).
The maximum age of fish observed was nearly 4 years old. At least one individual
of each sex was found in their third year; however, only 5 individuals were aged
to 3 years. Mortality rate was consistent for age classes 0 through 2. Survivorship
fit a weighted catch-curve estimate of mortality (Fig. 2), as described by Maceina
and Bettoli (1998). Catch data from year 3 was excluded from the estimate due to
low catch numbers. Using this method, we computed a mortality rate of 36.7%.
However, mortality rates calculated in this fashion assume a stable age distribution,
which may be unlikely for these sampled populations.
No differences were found between male and female growth (P = 0.40, F =
0.70, df = 1). Statistical significance was found among growth rates at the different
sites in 3 of the 6 possible comparisons. The growth rate at the Hominy Creek
site was significantly higher than that at Canton (P < 0.001, F = 164.6, df = 3, 113)
and Wilmot (P < 0.001, F = 130.9, df = 3, 109). The growth rate at East Laport
was also significantly higher than that at Canton (P < 0.001, F = 75.8, df = 3, 120).
The growth rate at Hominy Creek appeared to be the highest, although it was not
significantly different from that at East Laport (P = 0.126, F = 165.4, df = 3, 97).
The growth rate at Canton appeared to be lower than that at Wilmot, although not
significantly (P = 0.135, F = 46.3, df = 3, 132). The growth rate from all sites combined
was 1.05 mm/month.
The fit of length at age to a Von Bertalanffy growth model (Fig. 3) suggests an
asymptotic average standard length of 91 mm and a Brody growth rate of 0.0246
year-1. We fit the growth model using data from all sites combined to produce a
more general growth curve for the species rather than for a particular location.
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Figure 3. Fitted
line plot for the a
Von Bertalanffy
growth model for
Mirror Shiners
with approximate
9 5 % b o o t s t r a p
confidence intervals
shown as interior
dashed lines
and 95% bootstrap
prediction bounds
shown as exterior
dashed lines.
Confidence intervals
and prediction
bounds based on
1000 bootstraps.
For this model, L∞
= 91.0, K = 0.0246,
and t0 = -15.4. All
of these parameters
were statistically
significant.
Figure 2. Catchcurve
regression
showing mortality
of Mirror Shiners
in western North
C a r o l i n a . Va r i -
able A is yearly
mortality rate and
variable Z is instantaneous
mortality
rate. Graph
obtained using the
FSA package for R
(Ogle 2011).
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Ovigerous females had 13–331 oocytes. Ovigerous females in their 23rd month
had an average of 133 oocytes. Females in their 24th month had an average of 95
eggs. Females in their 25th month had an average of 70 eggs. The lone 37-month-old
female with oocytes had 331. Oocyte count was not size dependent (P = 0.63, F =
0.23, df = 1).
Individual GSIs ranged from 2.26 to 19.23. Average GSI was significantly higher
in May than in April (difference of 10.7 ± 6.6; P < 0.001) and June (difference
of 6.2 ± 5.5; P = 0.02) but not July (difference of 5.2 ± 6.1; P = 0.11) (Fig. 4). No
other comparisons were significant.
One-hundred twenty-seven individuals (53% of those examined) contained food,
including the following (and number of individuals in which an item was found):
detritus (80), Diptera (22), Coleoptera (20), Ephemeroptera (14), Hemiptera (4), Hymenoptera
(9), Lepidoptera (1), Megaloptera (7), Odonata (11), Trichoptera (10), and
material containing chloroplasts (16) (Fig. 5). No significant difference in frequency
Figure 4. Boxplots of GSI of female Mirror Shiner by month with outliers (those farther
than 1.5x the interquartile range) shown as circles. Only May was significantly different
from the other months.
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of occurrence of diet items was found between any of the sites by using a pairwise
G-test. For the G-test, we combined all observations that were not part of Coleoptera,
Diptera, or Detritus into an “other” category.
We captured all specimens in water ranging from 0.5 to 0.75 m in depth.
As water level changed, Mirror Shiners moved to maintain this microhabitat.
Furthermore, the locations where we captured specimens were all immediately
downstream (less than 1.0 m) from a riffle or rock in the stream that disrupted
streamflow. All pools in which we collected specimens had a sandy substrate.
Sites that lacked a sandy substrate, a pool beneath a flow-disrupting object, or
water 0.5 to 0.75 m in depth also lacked Mirror Shiners (Table 2). Water temperature
was consistently lower at the Hominy Creek and East Laport sites than
at Canton and Wilmot (Fig. 6).
Figure 5. Percent occurrence of gut contents in Mirror Shiner specimens by site. Macroinvertebrates
are identified to order.
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Discussion
The Mirror Shiner seems to have a similar life history to those of other Notropis
species, especially the Mimic Shiner. They have comparable diet, fecundity,
Table 2. Stream conditions at sites where Mirror Shiners were found or were not found while sampling
the Tuckasegee River. Each row represents a different sampling location.
Substrate Distance to
type Water depth (cm) flow-disrupting object Were Mirror Shiners present?
Sand 27 <1 m No
Sand 53 <1 m Yes
Sand 63 <1 m Yes
Sand 17 >1 m No
Cobble 12 <1 m No
Cobble 44 >1 m No
Cobble 37 >1 m No
Cobble 40 >1 m No
Cobble 43 >1 m No
Cobble 56 >1 m No
Cobble 39 >1 m No
Boulder 134 >1 m No
Boulder 75 >1 m No
Boulder 83 >1 m No
Figure 6. Line
graph showing
water temperature
at each of
the field sites
over the course
of the study.
T u c k a s e g e e
River at Wilmot
is indicated
with a plus
sign, Tuckasegee
River at
Laport is indicated
with a
square, Pigeon
River at Canton
is indicated
with a triangle,
and Hominy
Creek at Candler
is indicated
with a circle.
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total size and growth rate, and GSI. They also exhibit habitat specificity, and this
may be the most important reason that reintroduction efforts for the species have
been difficult.
A small survivorship to maximum age is not uncommon for shiners (Holder and
Powers 2010), which could be due to individuals recruiting into a size class that is
heavily preyed upon by larger fish or due to an energetic investment in reproduction
that proves fatal after the breeding season. Many Notropis spp. have a maximum
lifespan of 3 years, although the Tennessee Shiner and the Saffron Shiner have been
reported to live to a maximum of 5 years (Clayton 2000, Outten 1958).
The field site at Hominy Creek was the location with the least amount of disturbance
upstream, based on both observations in the field and of aerial photographs,
and the coldest water. East Laport had the second coldest water temperature during
the study and is also the second least-disturbed location. As Table 1 shows, there is
significant agricultural use upstream of the Canton site and a paper mill upstream
of the Wilmot site.
No difference existed between male and female growth rate. Pooled growth rate
was similar to that of Notropis chrosomus (Jordan) (Rainbow Shiner) (1.35 mm/
month) (Holder and Powers 2010). Growth rate was higher at the colder and less
disturbed sites, which could affect the reintroduction efforts of the Pigeon River
Recovery Project because the reach of the Pigeon River in which Mirror Shiners
are being relocated is downstream of a paper mill and therefore unnaturally warm
and disturbed.
Since GSI in May was statistically higher than in April and June and the most
intense breeding coloration was observed in males in May, peak breeding season
for the Mirror Shiner likely occurs in May. Breeding activity probably starts in
April, runs through the middle of the summer, and ends in July. Specimens collected
late in the year were beginning to mature, but lacked developed eggs, which
indicates a single yearly breeding season. We observed no spawning behavior, but
Mimic Shiners likely spawn nocturnally in open water (Black 1945). GSI values
were comparable to those of Mimic Shiners (Munz and Higgins 2013). Breeding
timing of the Mirror Shiner is comparable to that of the Tennessee Shiner, with a
single breeding season peaking in spring to early summer; although different from
that of the Mimic Shiner, which may spawn throughout the spring, summer, and fall
(Etnier and Starnes 1993).
Egg count for individuals in year 2 decreased as the year progressed, although
not significantly. The egg count for the female collected in year 3 was higher than
the counts for those collected in year 2, but only 1 ovigerous year-3 individual
was collected. No ovigerous year-1 individuals were collected. Egg count was
similar to what was found by Oliver (1986) in the Mimic Shiner, with up to 386
eggs per individual.
The predominant food items were Diptera, Coleoptera, and Ephemeroptera. A
large number of individuals were found containing detritus (33.8% of individuals
containing food), but it is unlikely that this large amount of detritus is an indication
that the Mirror Shiner is primarily a detritivore. This finding likely represents
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significantly digested, and thus unidentifiable, food in the guts of specimens. The
observation that many different prey items were found supports the hypothesis
that the Mirror Shiner is an opportunistic drift-feeder. The lack of a significant difference
in diet among sites further corroborates this hypothesis. Diet was similar
to that of the Mimic Shiner—primarily aquatic and terrestrial insects (Etnier and
Starnes 1993).
A high survivorship into year 2 appears especially important for the Mirror
Shiner because no year-1 individuals were collected with eggs. Although not
closely related to the Mirror Shiner (Cashner et al. 2011), Notropis spp. about
which more life history is known, such as the Rainbow Shiner and Notropis nubilus
(Forbes) (Ozark Minnow), appear to reach sexual maturity at 1 year of age. Other
related species, including Notropis lutipinnis (Jordan and Brayton) (Yellowfin
Shiner) and the Saffron Shiner, do not reach maturity until 2 years of age (Holder
and Powers 2000).
Crucially, Mirror Shiners were only captured in microhabitats fulfilling a specific
set of stream conditions. The collection location for each sampling event varied primarily
with water depth. Furthermore, Mirror Shiners were present only over sandy
substrates in pools just below an in-stream flow disturbing object. Therefore, it is apparent
that Mirror Shiners are habitat specific. Other Notropis spp. have been shown
to exhibit habitat-specific tendencies as well (Aadland 1993, Wall et al. 2004).
In conclusion, the habitat specificity of the Mirror Shiner suggests that the
fish may be patchily distributed. Therefore, reintroduction projects should take
care to reintroduce Mirror Shiners into river stretches that contain proper habitat
and to sample the appropriate microhabitat to observe surviving transplants. The
damming of the Pigeon River at Canton may have altered the particle sizes of
transported sediment, reducing the sandy habitat in pools that this species needs to
survive. In order to reintroduce Mirror Shiners successfully, stretches of rivers with
several sandy-bottom pools immediately downstream of in-stream flow-disrupting
objects should be used. Also, the stretch of river designated for reintroduction of
Mirror Shiners should have several sandy bottom pools of different depths to ensure
proper habitat for the fish throughout river height changes .
Acknowledgments
Thanks are due to A.D. Olson’s committee: Dr. Thomas Martin, Dr. Joseph Pechmann,
Dr. Seàn O’Connell, and Dr. Greg Adkison. The authors would also like to thank Dan Dawson
for help generating the map presented herein.
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