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Life History Traits of the Mirror Shiner, Notropis spectrunculus, in Western North Carolina
Adric D. Olson and Thomas H. Martin

Southeastern Naturalist, Volume 15, Issue 1 (2016): 102–114

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Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 102 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 Southeastern Naturalist 103 A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 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 Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 104 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. Southeastern Naturalist 105 A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 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. Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 106 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. Southeastern Naturalist 107 A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 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). Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 108 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. Southeastern Naturalist 109 A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 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. Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 110 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. Southeastern Naturalist 111 A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 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 Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 112 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. Literature Cited Aadland, L.P. 1993. Stream habitat types: Their fish assemblages and relationship to flow. North American Journal of Fisheries Management 13:790–806. Bartlett, R.A. 1995. Troubled Waters: Champion International and the Pigeon River Controversy. The University of Tennessee Press, Knoxville, TN. 376 pp. Black, J.D. 1945. Natural history of the Northern Mimic Shiner, Notropis volucellus volucellus Cope. Investigations of Indiana Lakes and Streams 2:449–466. Southeastern Naturalist 113 A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 Cashner, M.F., K.R. Piller, and H.L. Bart. 2011. Phylogenetic relationships of the North American cyprinid subgenus Hydrophlox. Molecular Phylogenetics and Evolution 59:725–735. Clayton, J.M. 2000. Life-history aspects of three minnow species of the subgenus Hydrophlox (Pisces: Cyprinidae): Notropis chiliticus, N. chlorocephalus, and N. lutipinnis. Ph.D. Dissertation. George Mason University, Fairfax, VA. Crim, L.W., and B.D. Glebe. 1990. Reproduction. Pp. 529–553, In C.B. Schreck and P.B. Moyle (Eds.). Methods for Fish Biology. American Fisheries Society, Bethesda, MD. Etnier, D.A., and W.C. Starnes. 1993. The Fishes of Tennessee. University of Tennessee Press, Knoxville, TN. 681 pp. Holder, D.S., and S.L. Powers. 2010. Life-history aspects of the Rainbow Shiner, Notropis chrosomus (Teleostei: Cyprinidae), in Northern Georgia. Southeastern Naturalist 9:347–358. Holm, S. 1979. A simple sequentially rejective multiple test procedure. Scandinavian Journal of Statistics 6:65–70. Hyatt, V. 2010, July 26. Environmentalists appeal mill permit. The Mountaineer, Canton, NC. Pp. 1A, 7A. LaVoie, M. 2007. Spatial and temporal patterns of drifting fish larvae in the upper Pigeon River, Haywood County, North Carolina. Master’s Thesis. Western Carolina University, Cullowhee, NC. Maceina, M.J., and P.W. Bettoli. 1998. Variation in Largemouth Bass recruitment in four mainstream impoundments of the Tennessee River. North American Journal of Fisheries Management 18:998–1003. Mayden, R.L. 1989. Phylogenetic studies of North American minnows, with emphasis on the genus Cyprinella (Teleostei: Cypriniformes). Miscellaneous Publications of the University of Kansas Museum of Natural History 80:1–189. Munz, J.T., and C.L. Higgins. 2013. The influence of discharge, photoperiod, and temperature on the reproductive ecology of cyprinids in the Paluxy River, Texas. Aquatic Ecology 47:67–74. Ogle, D.H. 2011. FSA: Fisheries Stock Analysis. R package version 0.2-7. Available online at https://fishr.wordpress.com/fsa/. Oliver, J.R. 1986. Comparative reproductive biology of the Cahaba Shiner, Notropis sp., and the Mimic Shiner, Notropis volucellus (Cope), from the Cahaba River drainage, Alabama. Master’s Thesis. Samford University, Birmingham, AL. Outten, L.M. 1958. Studies of the life history of the cyprinid fishes Notropis galacturus and rubricroceus. Journal of the Elisha Mitchell Scientific Society 74:123–134. R Core Team. 2013. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. Available online at http:// www.R-project.org/. Sokal, R.R., and F.J. Rohlf. 1981. Biometry: The Principles and Practice of Statistics in Biological Research. Freeman, New York, NY. 887 pp. United States Geological Survey (USGS). 2011. USGS Surface-water historical instantaneous data for the nation: Build time series. Available online at http://waterdata.usgs. gov/nwis/uv?referred_module=sw&search_criteria=state_cd&search_criteria=site_tp_ cd&submitted_form=introduction. Accessed 10 October 2011. University of Tennessee Pigeon River Restoration Project. 2004. Pigeon River Restoration Project. Available online at http://web.utk.edu/~mjwilson/index.php. Accessed 10 October 2011. Southeastern Naturalist A.D. Olson and T.H. Martin 2016 Vol. 15, No. 1 114 Wall, S.S., C.R. Berry Jr., C.M. Blausey, J.A. Jenks, and C.J. Kopplin. 2004. Fish-habitat modeling for gap analysis to conserve the endangered Topeka Shiner (Notropis topeka). Canadian Journal of Fisheries and Aquatic Sciences 61:954–973. Yanchis, T.L. 1993. Age, growth, and reproductive biology of Clinostomus funduloides raneyi in Caney Fork Creek, a Western North Carolina stream. Master’s Thesis. Western Carolina University, Cullowhee, NC.