2009 SOUTHEASTERN NATURALIST 8(4):653–670 Distribution, Population Characteristics, and Physical Habitat Associations of Black Bass (Micropterus) in the Lower Eleven Point River, Arkansas Ronald L. Johnson1,*, Alan D. Christian1, Sam D. Henry2, and Sam W. Barkley2 Abstract - We report the longitudinal distribution and population characteristics of Micropterus dolomieu (Smallmouth Bass), M. salmoides (Largemouth Bass), and M. punctulatus (Spotted Bass) of the lower Eleven Point River of Arkansas. Smallmouth Bass were the most abundant species collected, followed by Largemouth Bass and Spotted Bass. Abundance of Smallmouth Bass was greatest upstream and declined significantly downstream; abundance of Spotted Bass was significantly greater downstream. Largemouth Bass were evenly distributed throughout the river. Associated with these species distribution trends were a downstream decline in stream slope and velocity, and an increase in the proportion of pools relative to riffl es and runs. Proportional stock structures were similar and relatively low for each species (range = 22.9–32.3). Growth rates of Smallmouth Bass and Spotted Bass were high relative to other study populations, with age-3 fish reaching ≈300 mm total length. Diets of adult Smallmouth Bass and Spotted Bass were similar. Diets of Smallmouth Bass varied by season (increased feeding on insects during the summer months) and length group (transition of feeding on insects to fishes to crayfishes). Relative weight was greater for Largemouth Bass (Wr = 96) and Spotted Bass (95) than for Smallmouth Bass (89). Based on habitat, abundance, diet and condition factors, the Arkansas portion of the Eleven Point River contains a viable and balanced population of black bass. Introduction The three black bass species common to the central US, Micropterus dolomieu Lacépède (Smallmouth Bass), M. punctulatus (Rafinesque) (Spotted Bass), and M. salmoides (Lacépède) (Largemouth Bass) are sympatric in many of the larger streams and reservoirs in Arkansas (c.f., Robison and Buchanan 1988). The construction of reservoirs in the southeastern US and the adaptive nature of these bass has enabled them to proliferate and greatly expanded the fisheries of these bass. Extensive research has gone into the adaptation of black bass into these reservoirs, particularly regarding their longitudinal distributions in association with hydrological characteristics (e.g., Applegate et al. 1967, Hubert 1975, Sammons and Bettoli 1999). Surprisingly, less research has been performed on the longitudinal distribution of all three black bass where they occur in sympatry within unregulated rivers (Balkenbush and Fisher 1999, Leonard and Jenkins 1952). Stream 1Arkansas State University, Department of Biological Sciences, State University, AR 72467. 2Arkansas Game and Fish Commission, 2920 McClellan Drive, Jonesboro, AR 72401. *Corresponding author - rlj@astate.edu. 654 Southeastern Naturalist Vol. 8, No. 4 population studies typically have focused on two black bass species, with most examining the Smallmouth Bass as it relates to another black bass (Scott and Angermeier 1998, Sowa and Rabeni 1995, Tillma et al. 1998). Stream habitat requirements of black bass differ among species. Smallmouth Bass prefer clear, cool streams having moderate current and a substrate of either gravel or rock (Coble 1975, Reynolds 1965). Smallmouth Bass prefer higher stream gradients than the other two bass species (Burton and Odum 1945, Trautman 1942). The Largemouth Bass is the most adaptive of these species to higher turbidity (Etnier and Starnes 1993) and low velocity (Fajen 1975a, Wheeler and Allen 2003). The Spotted Bass exhibits intermediate habitat requirements (Jenkins 1975). Our main objectives were to investigate the longitudinal distribution of these three black bass species in an unregulated river and to describe associations with hydrology and physical habitat. We also report length-at-age and diet of Smallmouth Bass and Spotted Bass from this system. Last, we discuss the potential population effects of new length-limit regulations for Smallmouth Bass. Study Site The Eleven Point River is a clear, predominantly spring-fed stream located in the Ozark Mountain region of southeast Missouri and northeast Arkansas (Fig. 1). The headwaters of the stream originate in Howell County, MO, and fl ows approximately 225 km south before joining the Spring River in Randolph County, AR. The upper 160 km of this stream occurs within the borders of Missouri, and the lower 65 km occurs in Arkansas. The Eleven Point River is a fourth-order stream within Arkansas. However, due to being predominantly spring fed, stream discharge is higher than would be expected, with 2002–2004 monthly discharge averages during the study period ranging from 12.5 to 56.4 m3s-1 (mean = 30.8 m3s-1 ± 3.33 SE; US Geological Survey gauge ). Stream input in the study area is limited to intermittent streams, and there are no National Point Source Discharge Elimination System permits in the study area. Stream gradient in the Arkansas portion of the Eleven Point River is approximately 0.57 m km-1 (0.057%). Dominant predatory sportfishes of the lower Eleven Point River within Arkansas include, in order of frequency: Smallmouth Bass, Ambloplites ariommus Viosca (Shadow Bass), Sander vitreus (Mitchill) (Walleye), Largemouth Bass, and Spotted Bass. Primary forage fishes include Nocomis biguttatus (Kirtland) (Horneyhead Chub), Pimephales notatus (Rafinesque) (Bluntnose Minnow), Hybopsis amblops (Rafinesque) (Bigeye Chub), Labidesthes sicculus (Cope) (Brook Silverside), Campostoma anomalum (Rafinesque) (Central Stoneroller), and several Notropis and Moxostoma species (Johnson and Beadles 1977). At the time of this study, the minimum size limit for Smallmouth Bass in the river was 305 mm, with a daily creel limit of four fish. Additionally, there was a statewide aggregate creel limit of 10 black bass. 2009 R.L. Johnson, A.D. Christian, S.D. Henry, and S.W. Barkley 655 Methods Population analysis Daytime electrofishing by boat occurred from within 1.5 km of the Missouri border to within 5 km of the river's confl uence with the Spring River (≈60 total km). The river was subdivided into five sections (Fig. 1; hereafter labeled as Sections A [upstream] through E [downstream]), primarily as a function of boat access and distance, with sampling performed from June to November in 2002 (9 sampling trips), January to October in 2003 (9 trips), and May to June in 2004 (7 trips). The entire 60-km stretch within Arkansas was sampled annually during these years. Total length (TL; mm) and mass (g) were collected from each black bass collected. Figure 1. Study sites (Sections A–E) for sampling black bass within the lower Eleven Point River, AR. 656 Southeastern Naturalist Vol. 8, No. 4 Population structure information (catch-per-effort [CPE], proportional stock structure [PSS], and relative stock structure [RSS]; Guy et al. 2006) was calculated for each species in each river section. Stock-sized individuals of each species were divided into three length groups for comparisons of condition and feeding. Stock size were 180 mm for Smallmouth Bass and Spotted Bass, and 200 mm for Largemouth Bass. Proportional stock structure (or, quality fish; Gabelhouse 1984) and RSS (or preferred fish) used the following parameters: Smallmouth and Spotted Bass, PSS 280 mm and RSS 350 mm; and for Largemouth Bass, PSS 300 mm and RSS 380 mm. Length groups were: stock–PSS (stock), PSS–RSS (RSSQP), and RSS+ (RSSP) (Guy et al. 2006). Relative weight (Wr) was calculated for stock-size individuals for each species using the parameters of Kolander et al. (1993), Wege and Anderson (1978), and Wiens et al. (1996) for Smallmouth Bass, Largemouth Bass, and Spotted Bass, respectively. Relative weights were compared among species and length groups using ANOVAs. ANOVAs demonstrating significance were followed with an a posteriori Tukey’s multiple comparison test to test for treatment and interaction effects. All significance levels were set at α = 0.05. Sagittal otoliths were extracted from 128 Smallmouth Bass and 76 Spotted Bass for age estimation. Thin sectioning of otoliths was performed longitudinally with a Buehler isomet low-speed saw. Sections were then mounted on petroglyph slides and polished for visualization of annuli, which were then observed and counted using light microscopy. Mean length and mass at age were then calculated. Stomach contents were extracted from Smallmouth Bass (n = 397) and Spotted Bass (n = 53) that were sacrificed for age estimation and from live fish using gastric evacuation. All stomach contents were identified in the field to broad taxa (fish, crayfish, insects, other). More precise identifications were hindered due to the fl ushing of evacuation disrupting the structure of invertebrates, in addition to the partial digestion of prey in fish stomachs. We used chi square analysis to compare among Smallmouth Bass by season (spring, summer, and fall) and by length group, and between Smallmouth Bass and Spotted Bass. Hydrology and physical habitat analysis Length of the five sections in an upstream-to-downstream order was: Section A (5.40 km), Section B (8.74 km), Section C (15.33 km), Section D (9.80 km), and Section E (17.29 km). Stream gradient for each section was determined using USGS topographic maps, comparing elevation decline over river distance for each section. Habitat type, location (GPS, NAD83 datum), and wetted width were recorded for the entire study area. The length of each habitat (riffl e, run, pool) was measured in the laboratory with ArcMap (ESRI, Inc.), based on the starting and ending coordinates and tracing tools. We standardized riffl e, run, and pool area to area per km by dividing the area of each habitat by the length of each section. Regression analysis compared habitat composition among sections. 2009 R.L. Johnson, A.D. Christian, S.D. Henry, and S.W. Barkley 657 Water clarity and velocity analyses were performed during low fl ow in July. Water clarity (m) was measured at the tail of nine riffl es within each section and averaged using a Secchi tube and disk. A Secchi tube (7.9 cm diameter, 125 cm length) has a disk attached to the end of the tube, with water clarity measured by filling the tube with water and slowly releasing the water, via a notch on the side, until the disk becomes visible. The height of the water at the point the disk became visible is the Secchi depth; if the disk is visible while the tube is filled with water, then the water clarity is greater than the height of the tube (>1.25 m). Riffl e velocity (m s-1) was measured at three locations across the wetted width (i.e., 0.25, 0.50, and 0.75 width) for the nine riffl es per section using a Marsh-McBirney Flowmeter at 60 percent of the riffl e depth. Comparison of physical data with species distributions We performed principal components analysis (PCA) on the data matrix of CPE, PSS, and RSS for each species of the five sampling reaches using the software program PCOrd (Version 4) (McCune and Mefford 1999). The resulting multivariate matrix was analyzed using PCA to reduce the multivariate data into three meaningful axes and resulting loading scores. A secondary matrix of five physical variables of water clarity, riffl es, runs, and pool frequency per km, and riffl e, run, and pool area per km was used to correlate with the main matrix loading scores. Results Population analysis Smallmouth Bass (n = 1032) were the most abundant gamefish species collected, followed by Shadow Bass (631), Walleye (301), Largemouth Bass (197), and Spotted Bass (166). There were significantly more Smallmouth Bass caught per effort than both Largemouth Bass and Spotted Bass (F2,46 = 51.155, P < 0.0001; Table 1). Significant differences in Smallmouth Bass CPE were also identified longitudinally among sections (F4,19 = 3.866, P = 0.018), with densities declining downstream. The highest mean catch rate for Smallmouth Bass occurred upstream at Section A (29.8 fish/h), and catch rates were significantly lower at Sections D and E (P < 0.01). No Smallmouth Bass were collected in the lower 10 km of Section E. Largemouth Bass had high variation in catch rates, and there were no significant differences among sections (range = 2.4 to 11.6 fish/h; F4,19 = 0.365, P = 0.831; Table 1). However, significant differences in CPE among sections occurred for Spotted Bass (F4,19 = 6.617, P = 0.002), with greater numbers collected downstream. Section E had the highest CPE for Spotted Bass (9.8 fish/h), which was significantly higher than CPE for Sections A and B (P < 0.01). Most individuals collected for each species were stock size or greater (82% for Smallmouth Bass, 90% for Spotted Bass, and 85% for Largemouth Bass), which may be due to the size biases of boat electrofishing 658 Southeastern Naturalist Vol. 8, No. 4 (Reynolds 1996). There were no significant differences among species for PSS (P = 0.173; Table 1) or RSS (P = 0.140). No site effect was identified for any species for PSS or RSS (range of P values of 0.191 to 0.964). However, the middle three river sections (B, C, and D) exhibited Smallmouth Bass PSSs > 30, and RSS for Smallmouth and Largemouth Bass generally increased downstream. Length at age of Smallmouth Bass and Spotted Bass were similar, exhibiting linear growth to age 6 (Table 2). It takes 3 years for Smallmouth Bass to reach legal harvest size (305 mm), and 4–5 years for Smallmouth or Spotted Bass to reach preferred size (> 350 mm). Longevity was estimated to be 6 and 11 years for Smallmouth Bass and Spotted Bass, respectively. Maximum mass for an individual Largemouth Bass was 2008 g, for Smallmouth Bass was 1199 g, and for Spotted Bass was 686 g. Table 1. Mean catch per effort (CPE), proportional stock structure (PSS), and relative stock structure (RSS) for black bass of the lower Eleven Point River, AR. Standard errors are in parentheses below means. River Smallmouth Bass Spotted Bass Largemouth Bass section n CPE PSS RSS CPE PSS RSS CPE PSS RSS Section A 3 29.8 28.0 6.8 1.0 16.7 0.0 10.0 15.5 2.9 (5.46) (3.44) (1.95) (0.33) (33.33) (0.00) (3.68) (12.12) (1.43) Section B 4 27.8 36.4 9.0 1.1 40.0 20.0 6.2 30.4 7.8 (6.33) (5.75) (2.73) (0.31) (12.50) (12.50) (1.59) (6.25) (4.72) Section C 9 24.2 32.0 10.8 3.5 10.2 2.5 11.6 32.5 5.6 (2.38) (4.39) (1.70) (0.60) (2.95) (0.92) (2.05) (11.21) (7.41) Section D 4 17.0 37.9 11.2 4.0 25.5 3.9 8.6 40.2 14.8 (3.39) (4.15) (2.02) (0.77) (6.03) (3.23) (1.93) (18.28) (9.14) Section E 4 11.9 20.5 16.0 9.8 28.0 8.6 2.4 37.0 5.9 (3.12) (12.53) (10.51) (2.22) (11.66) (2.61) (0.49) (18.86) (5.00) Totals 24 23.9 32.3 10.3 6.4 22.8 6.1 8.9 30.2 7.7 (2.16) (2.86) (1.92) (0.65) (5.33) (2.24) (0.98) (6.31) (3.49) Table 2. Mean TL (mm) and mass (g) at age for Smallmouth and Spotted Bass in the lower Eleven Point River, AR. Standard errors are in parentheses after means. Smallmouth Bass Spotted Bass Age n TL Mass n TL Mass 1 29 148.7 (5.25) 46.4 (4.50) 21 164.5 (4.23) 61.9 (6.03) 2 76 219.2 (5.68) 147.9 (12.93) 18 214.1 (5.82) 121.9 (6.06) 3 38 291.8 (5.54) 340.9 (16.91) 24 307.0 (6.83) 204.1 (10.09) 4 6 346.0 (10.26) 532.0 (47.41) 8 286.0 (11.76) 338.0 (64.67) 5 21 383.6 (5.63) 781.6 (30.80) 1 396.0 (N/A) N/A (N/A) 6 8 422.0 (12.38) 870.6 (80.25) 1 365.0 (N/A) 683.0 (N/A) 7 0 - - 2 356.0 (15.00) 475.0 (N/A) 11 0 - - 1 380.0 (N/A) 623.0 (N/A) 2009 R.L. Johnson, A.D. Christian, S.D. Henry, and S.W. Barkley 659 Each of the three black bass species were characterized by high Wr (Table 3). Spotted Bass (Wr = 93) and Largemouth Bass (96) had signifi- cantly greater Wr than Smallmouth Bass (89) (F2,893 = 215.076, P < 0.001), and Largemouth Bass had greater Wr than Spotted Bass (P < 0.01). Wr for Smallmouth Bass were similar for all three length groups (F2,681 = 1.365, P = 0.498). Wr was more variable among Spotted Bass length groups, with longer fish having significantly lower relative weights (F2,128 = 17.222, P < 0.001; stock size > RSSQP and RSSP, P < 0.01; PSSQP > RSSP, P < 0.05). Conversely, relative weights were greater for the longest length group of Largemouth Bass (F2,78 = 12.085; P < 0.001; RSSP > stock, P < 0.01). Of the 397 Smallmouth Bass analyzed for stomach contents, 153 (39%) had empty stomachs, whereas 28 of 53 (53%) of Spotted Bass sampled had empty stomachs (Table 4). Crayfish (found in 48% of Smallmouth Bass sampled), fish (42%), and insects (25%) were consumed most frequently by Smallmouth Bass (Table 4). Diets of Spotted Bass were equally distributed among fishes and crayfish, with few insects (4%) occurring in stomachs. Diets were significantly different among species (χ2 = 271.43, P < 0.001). Few of the fish consumed as prey by either Smallmouth Bass or Spotted Bass were centrarchids (Micropterus, Lepomis, Ambloplites); most individuals Table 3. Wr of black bass as related to length group in the lower Eleven Point River, AR. Standard errors are in parentheses after means. Smallmouth Bass Spotted Bass Largemouth Bass Size n Mean Wr n Mean Wr n Mean Wr Stock Size 433 88.4 (1.36) 97 95.3 (1.31) 57 94.1 (1.33) PSS 174 89.1 (1.61) 27 89.0 (2.61) 17 98.5 (1.61) RSS 77 88.3 (1.31) 7 84.0 (2.02) 7 102.6 (2.74) Totals 684 88.5 (1.00) 131 93.4 (1.34) 81 95.7 (1.04) Table 4. Percents of prey items in Smallmouth Bass and Spotted Bass of the Eleven Point River, AR, with Smallmouth Bass sorted by season and length. Prey item Fish Crayfish Insect Other Empty Species/variable n % % % % % Smallmouth Bass Season Spring 65 57 43 2 7 35 Summer 292 39 48 32 8 37 Fall 38 37 58 11 0 50 TL 100; Slipke et al. 1998) were greater than we report for the Eleven Point River population. Both Spotted and Largemouth Bass relative weights were greater than for Smallmouth Bass, yet values for Spotted Bass (93.4) are greater than a stream in Alabama (Wr of 90 in Alabama; Greene et al. 2006), and yet lower than those reported for other streams (e.g., Wr of 103 in Kansas [Tillma et al. 1998] and 101 in Virginia [Scott and Angermeier 1998]). 666 Southeastern Naturalist Vol. 8, No. 4 Both Smallmouth and Spotted Bass are considered to be opportunistic in their prey selections (Austen and Orth 1985, Lewis and Helms 1964). Primary differences in diets of Smallmouth Bass and Spotted Bass were in the greater consumption of insects during the summer for Smallmouth Bass. Otherwise, diets of both species were quite similar; the reliance by both on crayfish and fish is consistent with many other studies (e.g., for Smallmouth Bass: Doan 1940, Watt 1959; for Spotted Bass: Mettee et al. 1996, Smith and Page 1969). Insects are often a greater component of Spotted Bass diets in other streams (e.g., Ryan et al. 1970, Scott and Angermeier 1998, Smith and Page 1969). Interspecific competition from feeding on similar prey may be reduced by differing habitat requirements (Scott and Angermeier 1998). The transition of the dietary focus of Smallmouth Bass in the Eleven Point River from insects to crayfish and fish with increasing TL is also consistent with previous studies (Robertson and Winemiller 2001, Roell and Orth 1993, Scott and Angermeier 1998). Our study has demonstrated that Smallmouth Bass are abundant throughout the lower Eleven Point River in Arkansas. Growth rates and size structure are high relative to other Ozark streams and comparable to most southeastern streams. Study of black bass populations and hydrology upstream in the Missouri section of the Eleven Point River would provide further insight into this system. In 2006, the length and creel limits in the Eleven Point River of Arkansas were altered from a 305 mm minimum TL and a daily creel limit of four fish to a 350 mm minimum TL and two fish per day, respectively. At the time of the study, Smallmouth Bass required a little more than 3 years to reach the legal limit of 305 mm. The extension of the length limit to 350 mm would allow for an additional year of growth prior to harvest if present growth dynamics are not negatively altered by an increase in standing stock. In addition to enhancing recruitment by allowing Smallmouth Bass to reproduce an additional year prior to harvest, the addition of trophy restrictions on a stream system resulted in increases of catch and release practices by anglers and increases in both the size structure and numbers of Smallmouth Bass in a Tennessee river (Slipke et al. 1988). Conversely, adverse effects on size structure of Smallmouth Bass populations have been noted following implementation of 300-mm creel limits (Austen and Orth 1988, Paragamian 1984b). The effects of the recent changes in harvest limits on the Smallmouth Bass population in the Eleven Point River remain to be seen. Both Spotted Bass and Largemouth Bass were common in the river, particularly Spotted Bass downstream. However, neither species was abundant enough to warrant special management considerations. Acknowledgments This research was funded by the Arkansas Game and Fish Commission, the Federal Aid to Sport Fish Restoration under Project F-39-R, and the RISE REU program at Arkansas State University (NSF Grant to R. Hannigan and C.B. Dowling [DBI 0552608]). We thank Allison Asher, Esther Daniells, Holly Martin, Andrew Peck, 2009 R.L. Johnson, A.D. Christian, S.D. Henry, and S.W. Barkley 667 and J.P. Fairhead for assistance with the collection and data entry of habitat data. 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