Distribution, Population Characteristics, and Physical
Habitat Associations of Black Bass (Micropterus) in the
Lower Eleven Point River, Arkansas
Ronald L. Johnson, Alan D. Christian, Sam D. Henry,
and Sam W. Barkley
Southeastern Naturalist, Volume 8, Number 4 (2009): 653–670
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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.
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 - email@example.com.
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
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
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.
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
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.
Fish Crayfish Insect Other Empty
Species/variable n % % % % %
Spring 65 57 43 2 7 35
Summer 292 39 48 32 8 37
Fall 38 37 58 11 0 50
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.
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. We
also thank R. Allen, C. Cato, and additional anonymous reviewers for improving the
quality of this manuscript.
Anderson, R.O., and A.S. Weithman. 1978. The concept of balance of coolwater
fish populations. Pp. 371–381, In R.L. Kendall (Ed.). Selected Coolwater
Fishes of North America. American Fisheries Society Special Publication 11.
Applegate, R.L., J.W. Mullan, and D.I. Morias. 1967. Food and growth of six
centrarchids from shoreline areas of Bull Shoals Reservoir. Proceedings of the
Annual Conference of the Southeastern Association of Game and Fish Commissioners
Austen, D.J., and D.J. Orth. 1985. Food utilization by riverine Smallmouth Bass in
relation to minimum length limits. Proceedings of the Southeastern Association
of Fish and Wildlife Agencies 39:97–107.
Austen, D.J., and D.J. Orth. 1988. Evaluation of a 305-mm minimum-length limit for
Smallmouth Bass in the New River, Virginia and West Virginia. North American
Journal of Fisheries Management 8:231–239.
Balkenbush, P.E., and W.L. Fisher. 1999. Population characteristics and management
of black bass in eastern Oklahoma streams. Proceedings of the Annual Conference
Southeastern Association of Fish and Wildlife Agencies 53:130–143.
Beamesderfer, R.C.P., and J.A. North. 1995. Growth, natural mortality, and predicted
response to fishing for Largemouth Bass and Smallmouth Bass populations in
North America. North American Journal of Fisheries Management 15:688–704.
Beamish, R.J., and G.A. McFarlane. 1987. Current trends in age determination methodology.
Pp. 15–42, In R.C. Summerfelt and G.E. Hall (Eds.). Age and Growth
of Fish. Iowa State University Press. Ames, IA.
Brown, D. 1932. The basses of Arkansas and some experiments in their propagation.
Transactions of the American Fisheries Society 61:83–85.
Burton, G.W., and E.P. Odum. 1945. The distribution of stream fish in the vicinity of
Mountain Lake, Virginia. Ecology 26:182–194.
Catchings, E.D., and R.O. Andress. 2005. Logan Martin reservoir management report.
Alabama Department of Conservation and Natural Resources, Division of
Wildlife and Freshwater Fisheries, Montgomery, AL.
Coble, D.W. 1975. Smallmouth Bass. Pp. 21–22, In R.H. Stroud and H. Clepper
(Eds.). Black Bass Biology and Management. Sport Fish Institute, Washington,
Dauwalter, D.C., and W.L. Fisher. 2008. Spatial and temporal patterns in stream
habitat and Smallmouth Bass populations in eastern Oklahoma. Transactions of
the American Fisheries Society 137:1072–1088.
DiCenzo, V.J., M.J. Maceina, and W.C. Reeves. 1995. Factors related to growth and
condition of the Alabama subspecies of Spotted Bass in reservoirs. North American
Journal of Fisheries Management 15:794–798.
Doan, K.H. 1940. Studies of the Smallmouth Bass. Journal of Wildlife Management
Echelle, A.A., and G.D. Schnell. 1976. Factor analysis of species associations among
fishes of the Kiamichi River, Oklahoma. Transactions of the American Fisheries
668 Southeastern Naturalist Vol. 8, No. 4
Etnier. D.A., and W.C. Starnes. 1993. The Fishes of Tennessee. The University of
Tennessee Press. Knoxville,TN.
Fajen, O.F. 1975a. Population dynamics of bass in rivers and streams. Pp. 195–203,
In R.H. Stroud and H. Clepper (Eds.). Black Bass Biology and Management.
Sport Fish Institute, Washington, DC.
Fajen, O.F. 1975b. The standing crop of Smallmouth Bass and associated species in
Courtois Creek. Pp. 240–249, In R.H. Stroud and H. Clepper (Eds.). Black Bass
Biology and Management. Sport Fish Institute, Washington, DC.
Filipek. S., et al. 1995. Arkansas Smallmouth management plan. Project F-42. Arkansas
Game and Fish Commission, Little Rock, AR.
Fiss, F.C., T.A. Cleveland, B.D. Carter, R.D. Bivens, and J.M. Swearengin. 2001.
Population characteristics of riverine Smallmouth Bass in Tennessee, simulated
effects of length limits, and management recommendations. Tennessee Wildlife
Resources Agency, Fisheries Report 01-19. Nashville,TN.
Fleener, G.G. 1975. Harvest of Smallmouth Bass and associated species in Courtois
Creek. Pp. 250–256, In R.H. Stroud and H. Clepper (Eds.). Black Bass Biology
and Management. Sport Fish Institute, Washington, DC.
Gabelhouse, D.W., Jr. 1984. A length-categorization system to assess fish stocks.
North American Journal of Fisheries Management 4:273–285.
Greene, J.C., D.L. Abernethy, and R.A. McVay. 2006. Coosa River management
report. Alabama Department of Conservation and Natural Resources, Division of
Wildlife and Freshwater Fisheries, Montgomery, AL.
Guy, C.S., R.M. Neumann, and D.W. Willis. 2006. New terminology for proportional
stock density (PSD) and relative stock density (RSD): Proportional size structure
(PSS). Fisheries 31:86–87.
Howland, J.W. 1931. Studies on the Kentucky Black Bass (Micropterus pseudaplites
Hubbs). Transactions of the American Fisheries Society 61:89–94.
Hubert, W.A. 1975. Age and growth of three black bass species in Pickwick Reservoir.
Proceedings of the Annual Conference of the Southeastern Association of
Game and Fish Commissioners 29:126–134.
Hubert, W.A. 1977. Comparative food habits of Smallmouth and Largemouth Basses
in Pickwick Reservoir. Journal of Alabama Academy of Science 48:167–178.
Jenkins, R.M. 1975. Black bass crops and species associations in reservoirs. Pp.
114–124, In R.H. Stroud and H. Clepper (Eds.). Black Bass Biology and Management.
Sport Fish Institute, Washington, DC.
Johnson, B.M., and J.K. Beadles. 1977. Fishes of the Eleven Point River within Arkansas.
Proceedings of the Arkansas Academy of Sciences 31:58–61.
Kolander, T.D., D.W. Willis, and B.R. Murphy. 1993. Proposed revision of the
standard weight (Ws) equation for Smallmouth Bass. North American Journal of
Fisheries Management 13:398–400.
Leonard, E.M., and R.M. Jenkins. 1952. Growth of the basses of the Illinois River,
Oklahoma. Proceedings of the Oklahoma Academy of Sciences 33:21–29.
Lewis, W.H., and D.R. Helms. 1964. Vulnerability of forage organisms to Largemouth
Bass. American Fisheries Society, Transactions 93:315–318.
Lobb III, M.D., and D.J. Orth. 1991. Habitat use by an assemblage of fish in a large
warmwater stream. Transactions of the American Fisheries Society 120:65–78.
Long, J.M., and W.L. Fisher. 2001. Precision and bias of Largemouth, Smallmouth,
and Spotted Bass ages estimated from scales, whole otoliths, and sectioned otoliths.
North American Journal of Fisheries Management 21:636–645.
2009 R.L. Johnson, A.D. Christian, S.D. Henry, and S.W. Barkley 669
McClendon, D.D., and C.F. Rabeni. 1987. Physical and biological variables useful
for predicting population characteristics of Smallmouth Bass and Rock Bass in an
Ozark stream. North American Journal of Fisheries Management 7:46–56.
McCune, B., and M.J. Mefford. 1999. PC-ORD. Multivariate Analysis of Ecological
Data, Version 4 Edition. MjM Software Design, Gleneden Beach, OR.
Mettee, M.F., P.E. O’Neil, and J.M. Pierson. 1996. Fishes of Alabama and the Mobile
Basin. Oxmoor House, Inc, Birmingham, AL.
Orth, D.J., D.D. Oakey, and O.E. Maughan, 1983. Population characteristics of
Smallmouth Bass in Glover Creek. Southeast Oklahoma. Proceedings of the
Oklahoma Academy of Science 63:37–41
Paragamian. V.L. 1984a. Population characteristics of Smallmouth Bass in five Iowa
streams and management recommendations. North American Journal of Fisheries
Paragamian, V.L. 1984b. Evaluation of a 12.0-inch minimum-length limit on Smallmouth
Bass in the Maquoketa River, Iowa. North American Journal of Fisheries
Quinn, J. 2003. Characteristics of fish populations in the rivers and streams of
Arkansas. Arkansas Game and Fish Commission, Report STP2003-07, Little
Reed, M.S., and C.F. Rabeni. 1989. Characteristics of an unexploited Smallmouth
Bass population in a Missouri Ozark stream. North American Journal of Fisheries
Reynolds, J.B. 1965. Life history of Smallmouth Bass, Micropterus dolomieui
Lacepede, in the Des Moines River, Boone County, Iowa. Iowa State Journal of
Reynolds, J.B. 1996. Electrofishing. Pp. 221–253, In B.R. Murphy and D.W.
Willis (Eds.). Fisheries Techniques, 2nd Edition. American Fisheries Society.
Robertson, M.S., and K.O. Winemiller. 2001. Diet and growth of Smallmouth Bass
in the Devils River, Texas. Southwestern Naturalist 46:216–221.
Robison, H.W., and T.M. Buchanan. 1988. Fishes of Arkansas. The University of
Arkansas Press, Fayetteville, AR.
Roell, M.J., and D.J. Orth. 1993. Trophic basis of production of stream-dwelling
Smallmouth Bass, Rock Bass, and Flathead Catfish in relation to invertebrate bait
harvest. Transactions of the American Fisheries Society 122:46–62.
Ryan, P.W., J.W. Avault, Jr., and R.O. Smitherman. 1970. Food habits and spawning
of the Spotted Bass in Tchefuncte River, southeastern Louisiana. Progressive
Sammons, S.M., and P.W. Bettoli. 1999. Spatial and temporal variation in electrofishing catch rates of three species of black bass (Micropterus spp.) from Normandy
Reservoir, Tennessee. North American Journal of Fisheries Management
Scott, M.C., and P.L. Angermeier. 1998. Resource use by two sympatric black basses
in impounded and riverine sections of the New River, Virginia. North American
Journal of Fisheries Management 18:221–235.
Slipke, J.W., M.J. Maceina, V.H. Travnichek, and K.C. Weathers. 1998. Effects of a
356-mm minimum-length limit on the population characteristics and sport fishery
of Smallmouth Bass in the Shoals Reach of the Tennessee River, Alabama. North
American Journal of Fisheries Management 18:76–84.
670 Southeastern Naturalist Vol. 8, No. 4
Smith, P.W., and L.M. Page. 1969. The food of Spotted Bass in streams of the Wabash
River drainage. Transactions of the American Fisheries Society 98:647–651.
Sowa, S.P., and C.E. Rabeni. 1995. Regional evaluation of the relation of habitat to
distribution and abundance of Smallmouth Bass and Largemouth Bass in Missouri
streams. Transactions of the American Fisheries Society 124:240–251.
Stewig, J.D., and D.R. DeVries. 2004. The fish community of a fl ow-impacted river
reach in Alabama, USA, with emphasis on Largemouth Bass and Spotted Bass.
Journal of Freshwater Ecology 19:387–400.
Tillma, J.S., C.S. Guy, and C.S. Mammoliti. 1998. Relations among habitat and
population characteristics of Spotted Bass in Kansas streams. North American
Journal of Fisheries Management 18:886–893.
Trautman, M.B. 1942. Fish distribution and abundance correlated with stream gradients
as a consideration in stocking programs. Transactions of the North American
Wildlife Conference 7:211–224.
Watt, K.E.F. 1959. Studies on population productivity II. Factors governing productivity
in a population of Smallmouth Bass. Ecological Monographs 29:367–392.
Wege, G.J., and R.O. Anderson. 1978. Relative weight (Wr), a new index of condition
for Largemouth Bass. Pp. 79–91, In G.D. Novinger and J.G. Dillard (Eds.). New
Approaches to the Management of Small Impoundments. American Fisheries
Society, North Central Division, Special Publication 5, Bethesda, MD.
Wheeler, A.P., and M.S. Allen. 2003. Habitat and diet partitioning between Shoal
Bass and Largemouth Bass in the Chippola River, Florida. Transactions of the
American Fisheries Society 132:438–449.
Wiens, J.R., C.S. Guy, and M.L. Brown. 1996. A revised standard weight (Ws)
equation for Spotted Bass. North American Journal of Fisheries Management
Zweifel, R.D., R.S. Hayward, and C.F. Rabeni. 1999. Bioenergetics insight into
black bass distribution shifts in Ozark border region streams. North American
Journal of Fisheries Management 19:192–197.