Training course in fish stock assessment and fisheries management
Cat Fish Stock
Transcript of Cat Fish Stock
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North American Journal of Fisheries Management 25:11911201, 2005 [Article] Copyright by the American Fisheries Society 2005DOI: 10.1577/M03-251.1
Population Abundance and Stock Characteristics of Flathead
Catfish in the Lower St. Joseph River, Michigan
DANIEL J. DAUGHERTY AND TRENT M. SUTTON*
Department of Forestry and Natural Resources, Purdue University,
195 Marsteller Street, West Lafayette, Indiana 47907-1159, USA
Abstract.Little information exists regarding the biological attributes and stock dynamics of
flathead catfish Pylodictis olivaris in lotic systems throughout the northern United States. We
examined the population abundance, annual survival, growth, condition, size structure, and age
structure of flathead catfish in the lower St. Joseph River, Michigan, to direct future management
efforts in this system and increase our knowledge of northern flathead catfish stocks. Fish were
collected by means of electrofishing during June through September 2002 and 2003. Population
abundance was estimated at 5,452 individuals (range, 3,9857,277 fish), and annual survival was
estimated at 67%. Analysis of pectoral spine cross sections revealed that growth was greatest for
fish younger than age 6 (range, 8398 mm/year) and decreased among older age-classes (range,1870 mm/year). Relative weight was greatest for individuals less than 300 mm total length (TL;
115%) and declined with increasing fish length. The size structure and age structure of flathead
catfish were dominated by fish less than 400 mm TL and younger than age 4 (89% and 79%,
respectively), although individuals greater than 1,100 mm TL and up to age 17 were present in
the population. Relative stock density estimates indicated that 58% of fish greater than the minimum
stock size were of quality length, while flathead catfish of preferred, memorable, and trophy sizes
represented 28, 7, and 1% of fish collected, respectively. Despite their presence on the northern
fringe of the species geographic distribution, the flathead catfish in the lower St. Joseph River
exhibited biological characteristics and stock dynamics that were similar to those reported for
other lightly exploited stocks throughout the midwestern United States.
Flathead catfish Pylodictis olivaris are an im-
portant component of the North American catfish
fishery (Jackson 1999). This species is native to
the Mississippi River, Mobile River, and Rio
Grande River drainages, the Laurentian Great
Lakes basin, and northeastern Mexico (Hubbs and
Lagler 1947; Lee and Terrell 1987; Jackson 1999)
and supports recreational and commercial fisheries
throughout much of its geographic distribution
(Moss and Tucker 1989; Grussing et al. 2001). In
many states, flathead catfish offer anglers an op-
portunity to catch fish by means other than hook
and line (e.g., handgrabbing, bankpoling, etc.),while giving commercial fishers a high-quality
product preferred by many consumers (Quinn
1993; Grussing et al. 2001; Jackson 1999). Despite
the popularity of this species, knowledge regarding
the population characteristics of flathead catfish
stocks is limited. This lack of information is most
prevalent in moderate-size rivers, streams, and
creeks, where fisheries managers have found as-
sessment of catfish populations difficult due to the
solitary nature and low population abundances of
* Corresponding author: [email protected]
Received December 22, 2003; accepted January 12, 2005
Published online August 2, 2005
catfish (Stauffer et al. 1996; Vokoun and Rabeni
1999).Although a number of studies have examined
the population characteristics of flathead catfish in
lotic environments, most have focused on size
structure, age distribution, and growth in systems
throughout the southern United States (Purkett
1958; Minckley and Deacon 1959; Mayhew 1969;
Guier et al. 1984; Pisano et al. 1983; Young and
Marsh 1990; Grussing et al. 2001). Purkett (1958)
reported that flathead catfish grew an average of
76 mm/year in the Salt River, Missouri, and
reached a mean total length (TL) of 300 mm byage 4. Flathead catfish grew more slowly in the
Salt River system than in the Mississippi River
(IllinoisIowa and MissouriIowa), where fish
reached 381 and 432 mm TL, respectively, at the
age 4 (Barnickol and Starrett 1951). Similarly,
Minckley and Deacon (1959) reported that flathead
catfish reached 483 mm TL by age 4 in the Big
Blue River, Kansas. However, Guier et al. (1984)
reported faster growth in the Cape Fear River,
North Carolina, where individuals reached a mean
TL of 580 mm at the same age. The results of these
studies indicate that growth of flathead catfish issystem specific and that the length of the growing
season may not be a reliable predictor of growth
rate.
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Whereas southern stocks of flathead catfish have
received attention from fisheries managers, little
information exists regarding the characteristics of
populations in the upper Midwest and Great Lakes
regions. Historically, northern populations haveexperienced lower exploitation rates and less an-
gling pressure than those in the south (Topp et al.
1994; Stauffer et al. 1996). These fishery char-
acteristics may result in differences in abundance,
growth, condition, age structure, and size structure
for flathead catfish stocks between northern and
southern systems. Stauffer et al. (1996) found that
growth rates of flathead catfish in the Minnesota
River, Minnesota, were 1352% lower than those
of fish in the southern United States. The authors
also noted that a high abundance of large fish was
present in the population: 20% of flathead catfishsampled were greater than 910 mm TL. These re-
sults suggest that differences in population char-
acteristics exist between northern and southern
flathead catfish stocks, supporting the need for ad-
ditional studies on fish in northern systems.
The popularity of recreational and commercial
fisheries for flathead catfish has increased sub-
stantially throughout their geographic range in re-
cent years (Summers 1986; Gilliland 1988; Quinn
1993; Cunningham 1995; Stauffer et al. 1996;
Jackson 1999). This trend is especially evident inthe northern United States, where flathead catfish
have historically attracted little angler interest
(Topp et al. 1994; Stauffer et al. 1996). Because
flathead catfish may be susceptible to overharvest
due to such factors as their low abundance and
aggressive behavior, knowledge of the life history
characteristics and population status of this species
is important for the implementation of appropriate
management strategies (Quinn 1993; Grussing et
al. 2001; Jackson 1999). Without this information,
management of flathead catfish populations would
be difficult, particularly as angling pressure in-
creases (Stauffer et al. 1996).
The flathead catfish population in the St. Joseph
River, a Lake Michigan tributary located in south-
western Michigan and northeastern Indiana, has
experienced an increase in angling pressure over
the past decade (J. Dexter, Michigan Department
of Natural Resources, personal communication).
Prior to 1996, an average of 14 flathead catfish
caught from the St. Joseph River each year was
entered into the annual state master angler award
program. Over the past 7 years, the average num-ber of entries submitted each year has decreased
by 50%. This decline in the catch of trophy fish
suggests an increase in the exploitation rate of flat-
head catfish in the St. Joseph River. However, the
current status of this fishery and the biological
attributes of the population remain unknown. The
objective of this study was to provide information
regarding the population characteristics and stockdynamics of flathead catfish in the St. Joseph River
system. Such information can be used to direct
future management efforts and i ncrease our knowl-
edge of flathead catfish populations in the northern
United States.
Study Site
The 338-km main-stem St. Joseph River is
joined by 2,641 km of tributaries, discharging an
average of 130 m3 /s from a watershed that drains
approximately 7,770 km2 in Michigan and 4,364
km2 in Indiana. The free-flowing (hereafter re-ferred to as lower) section of the St. Joseph River
is a 37.6-km reach between the Berrien Springs
Dam and the mouth of the river at Lake Michigan
(Figure 1). Because flathead catfish are not found
above the Berrien Springs Dam (J. Dexter, Mich-
igan Department of Natural Resources, personal
communication), the boundaries of this river reach
served as the upstream and downstream limits of
the sampling area. We divided the lower St. Joseph
River into four 9.4-km sampling sections to fa-
cilitate collection of flathead catfish (Figure 1).Sampling sections 1 and 2 are dominated by gravel
substrates and have a low to fair gradient (00.9
m/km), whereas sections 3 and 4 are dominated
by sand substrates and have a low gradient (0 0.5
m/km). Riparian land cover in the reach is dom-
inated by hardwood forests, although urban de-
velopment dominates section 4 as the river ap-
proaches Lake Michigan.
The lower St. Joseph River supports a resident
warmwater fish community; the most abundant
species are flathead catfish, channel catfish Ictal-
urus punctatus, freshwater drum Aplodinotus grun-
niens, common carp Cyprinus carpio, and gizzard
shad Dorosoma cepedianum (Wesley and Duffy
1999). Mean channel width in the lower St. Joseph
River is 130 m (range, 50250 m), and mean water
depth is 1.6 m (range, 0.38.2 m), although lateral-
scour pools with water depths in excess of 7 m
occur throughout the reach. Midsummer (June
August) water temperature can be as high as 28C;
mean dissolved oxygen is 9.6 mg/L (range, 6.9
14.1 mg/L) and turbidity is 16.5 nephelometric
turbidity units (NTU; range, 1127 NTU). In-stream habitat is composed primarily of woody
debris and rip-rap; few aquatic macrophytes exist
in the system.
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1193ST. JOSEPH RIVER FLATHEAD CATFISH
FIGURE 1.Map of the lower, free-flowing reach of the St. Joseph River, Michigan. Lines bisecting the river
indicate the upper and lower boundaries of the four sections used for the sampling of flathead catfish.
Methods
Flathead catfish were collected during June
through September 2002 and 2003. We used the
modified predator approach defined by Vokoun and
Rabeni (1999). Fish were sampled once per week
from each section of the study area by use of 24
38-V AC produced by a three-bar magnetic motor,
as described by Morris and Novak (1968). Al-
though sampling efforts were conducted in all hab-
itat types (i.e., main-channel pools, riffles, and
runs with and without structure), efforts were con-
centrated in structural habitats typically selected
by flathead catfish (e.g., large woody debris jams,
timbered channels, undercut banks, rip-rap, etc.;
Cunningham 2000). At each sampling location,two 18-gauge insulated wires (each 6.1 m in
length) with the distal ends connected to
aluminum-bar electrodes (30.5 cm long and 2.5
cm wide) were deployed approximately 45 m up-
stream and downstream of the boat. Electrical cur-
rent was applied continuously for 90 s at each
sampling location. During August 2003, flatheadcatfish were also sampled from section 4 by means
of low-frequency (20% pulse width), low-pulse
(7.5 pulse/s) DC boat electrofishing, as described
by Stauffer and Koenen (1999). A chase boat was
used during all sampling periods in 2003 to ca pture
flathead catfish that surfaced downstream of the
boat carrying the electrofishing unit.
Population characteristics.All captured flat-
head catfish were measured for TL (nearest 1 mm)
and wet weight (nearest 1 g). The spine of the left
pectoral fin was removed and dried for subsequentage and growth analyses in the laboratory. Ab-
solute population abundance of flathead catfish
was calculated by use of the Schnabel estimator
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1194 DAUGHERTY AND SUTTON
for closed populations (Schnabel 1938). Because
assessments of flathead catfish movement in the
lower St. Joseph River indicated that individuals
remained within the study reach during the entire
study period (Daugherty and Sutton 2005), we as-sumed the population to be geographically closed.
Ninety-five percent confidence intervals (CIs)
were calculated with the Poisson distribution by
substituting the confidence limits for the number
of marked individuals at large in the population
(R) into the Schnabel estimator (Van Den Avyle
1993). Flathead catfish density (i.e., the number of
individuals per river kilometer [rkm]) was calcu-
lated by dividing the estimate of absolute abun-
dance by the length of the study reach.
In the laboratory, pectoral spines were sectioned
by use of a rotary tool fitted with a diamond-coated
cutting disk. To avoid potential fish age underes-
timation due to erosion of the central lumen (Turn-
er 1982; Nash and Irwin 1999), multiple cross sec-
tions were cut between the distal end of the basal
recess and the proximal end of the spine dentations
(Layher 1981; Crumpton et al. 1987). Spine sec-
tions were then visually examined and discarded
until enlargement of the central lumen was no lon-
ger obvious (Munger et al. 1994). Cross sections
of each spine were mounted on microscope slides
and viewed with transmitted light at 36135magnification under a zoom stereomicroscope fit-
ted with a digital camera. Images of each spine
were analyzed with IPLab 3.6 image analysis soft-
ware (Scanalytics, Fairfax, Virginia), which was
used to determine the number of annuli present,
the length of the spine radius, and the radial dis-
tance to each annulus for back-calculation of
length at age. All measurements were recorded to
the nearest 0.01 mm. Each spine image was ana-
lyzed independently by two readers, and disagree-
ments between readers were reconciled with a sub-sequent concert read.
Length at age for each flathead catfish was back-
calculated by means of a modification of the Fras-
erLee method (DeVries and Frie 1996). Back-
calculated length-at-age estimates were used to de-
termine the age at which flathead catfish were re-
cruited to the harvestable fishery (minimum length
limit, 381 mm TL) and to calculate age-specific
annual growth rates (mm/year). A von Bertalanffy
growth model was constructed for the population
in FAST 2.0 (Auburn University, Auburn, Ala-bama). Mean TL and age-at-capture data were used
to estimate the theoretical maximum fish length
(L
), Brody growth coefficient (K), and time period
when fish body length equals 0 mm (to) for the
von Bertalanffy growth model.
Fish condition was calculated as relative weight
(Wr). Length-specific standard weights (Ws) for
flathead catfish were derived from the standard-weight equation, log10(Ws) 5.542 3.23log10(TL)
(Bister et al. 2000). Relative weight was not cal-
culated for flathead catfish less than 130 mm in
length because the standard-weight equation ex-
cluded smaller individuals due to variance-to-
mean errors greater than 0.02 (Bister et al. 2000).
Relative stock density (RSD) indices (e.g., quality
[Q], preferred, memorable, and trophy) were cal-
culated based on length categories developed by
Bister et al. (2000). Minimum TLs for each cat-
egory were as follows: 350 mm for stock length,
510 mm for quality length, 710 mm for preferredlength, 860 mm for memorable length, and 1,020
mm for trophy length.
Due to variable capture rates of flathead catfish
ages 512, annual survival was estimated based
on the Robson and Chapman (1961) method.
Ninety-five percent CIs for these estimates were
calculated by use of the methods described by
Ricker (1975). The estimate of annual survival ob-
tained from the Robson and Chapman (1961)
method was used to calculate the instantaneous
total mortality rate Z. Confidence intervals for Zwere calculated by substituting the confidence lim-
its of the annual survival rate into the calculation
of Z. Age-classes younger than the first age-class
fully recruited to the sampling gear, as well as age-
classes with limited sample sizes (N 5), were
omitted from the estimation of annual survival
(Ricker 1975; Van Den Avyle 1993).
Data analysis.Piecewise linear regression was
used to determine the relationship between mean
annual growth increment and fish age. Simple lin-
ear regression was employed to determine the re-
lationship between relative condition and fish TL.
Mean TL of flathead catfish in the lower St. Joseph
River was regressed as a function of fish age, and
analysis of covariance (ANCOVA) was used to
compare this regression with growth relationships
for nine other flathead catfish populations. Signif-
icant differences among regression coefficients
were determined by the Tukeys honestly signifi-
cant difference test. All statistical analyses em-
ployed a significance level of 0.05.
Results
A total of 653 flathead catfish were collected
from 759 sites on 32 sampling occasions during
June through September 2002 and 2003. One-
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1195ST. JOSEPH RIVER FLATHEAD CATFISH
FIGURE 2.Length-frequency distribution of flathead
catfish collected from the lower St. Joseph River, Mich-
igan, during 2002 and 2003.
FIGURE 3.Age-frequency distribution of flathead
catfish collected from the lower St. Joseph River, Mich-
igan, during 2002 and 2003.
hundred forty-six flathead catfish were collected
by low-frequency electrofishing during 15 d of
sampling in 2002. In 2003, 447 individuals were
collected during 17 sampling occasions. An ad-
ditional 60 fish were collected during August 2003
by means of DC boat electrofishing.
Six percent (N 40) of the flathead catfish
tagged during the study period were recaptured
during subsequent sampling occasions. The ab-
solute abundance of flathead catfish in the lowerSt. Joseph River was estimated at 5,452 individ-
uals (95% CI 3,9857,277). The density of flat-
head catfish was estimated at 145 fish/rkm. Annual
survival of flathead catfish in the lower St. Joseph
River was 0.672 (95% CI 0.6700.673). The Z
derived from this approach was 0.397 (95% CI
0.3960.400).
Flathead catfish captured during the study
ranged from 87 to 1,132 mm TL (Figure 2). In-
dividuals less than 100 mm TL represented 1% of
fish sampled, whereas flathead catfish ranging
from 100 to 400 mm TL made up the greatest
proportion of fish collected (89%). Capture rates
of flathead catfish greater than 400 mm TL varied
among length-classes (Figure 2). Fish ranging
from 500 to 600 mm TL composed 3% of the catch,
whereas fish that were 700800 mm TL repre-
sented 7% of fish sampled. Flathead catfish 900
mm TL and larger made up 5% of the population.
Pectoral spine cross sections were examined for
94% (N 612) of the collected fish. Age and
growth analyses were omitted for 41 individuals
either because of missing or broken pec toral spinesor because of pectoral spine size. Pectoral spines
collected from individuals less than 130 mm TL
were too small to accurately cross-section. The age
of flathead catfish ranged from 1 to 17 years (Fig-
ure 3). Age-1 flathead catfish represented 22% of
the fish sampled, age-2 fish represented 36%, age-
3 fish represented 15%, and age-4 fish represented
6%; these data indicate that the fish did not become
fully vulnerable to the sampling gear until age 2.
Capture rates of age-512 flathead catfish varied
among age-classes (Figure 3). Age-9 flathead cat-
fish composed the greatest proportion (4%) of fish
captured within this age range. Fish older than age12 made up 2% of the population (Figure 3).
Length-at-age estimates for flathead catfish in
the lower St. Joseph River indicated that flathead
catfish reached a mean TL of 102 mm by age 1
and were recruited to the harvestable fishery by
age 4, when the mean TL was 401 mm (Figure 4).
The mean length at age of fish in the oldest age-
class (age 17) was 1,016 mm TL. Based on the
von Bertalanffy growth equation, the predicted age
at which flathead catfish were recruited to the har-
vestable fishery was 4 years, while the maximum
attainable TL of flathead catfish in the St. Joseph
River was estimated at 1,174 mm (Figure 4).
Mean annual growth increments of flathead cat-
fish were greatest among fish younger than age 6
(range, 8398 mm/year). The slope of the linear
regression relationship over this age range indi-
cated that annual growth declined by less than 5
mm/year during this period (Figure 5). The great-
est declines in the annual growth rate of flathead
catfish in the lower St. Joseph River were observed
between ages 6 and 8 (range, 4270 mm/year).
Piecewise regression of these age-classes indicatedthat annual growth declined by approximately 14
mm/year. Annual growth rates declined little
among fish older than age 8; however, fish growth
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1196 DAUGHERTY AND SUTTON
FIGURE 4.Mean length at age and fitted von Ber-
talanffy growth equation (t age in years; lt length
at time t) for flathead catfish collected from the lower
St. Joseph River, Michigan, during 2002 and 2003. Errorbars represent 95% confidence intervals about the mean
length at age. Confidence intervals were not calculated
for age 16 because only one fish was collected.
FIGURE 6.Relationship between mean relative
weight (Wr) and total length (TL) for flathead catfish
collected from the lower St. Joseph River, Michigan,
during 2002 and 2003. Individuals were pooled into 100-
mm length groups. Error bars represent 95% confidence
intervals. Confidence intervals were not calculated for
the 1,100-mm size-class because only one fish was col-
lected.
FIGURE 5.Relationship between age (X) and calcu-
lated mean annual growth increment (Y) for each age-
class (i) of flathead catfish collected from the lower St.
Joseph River, Michigan, during 2002 and 2003. Error
bars represent 95% confidence intervals.
TABLE 1.Comparative relative stock density (RSD)
indices for flathead catfish in the lower St. Joseph River
(SJ), Michigan, the Flint River (FL), Georgia, and the
Cape Fear River (CF), North Carolina.
Indexcategory
MinimumTL (mm)
SJfrequency
(%)a
FLfrequency
(%)b
CFfrequency
(%)c
Quality 510 58 57 49
Preferred 710 28 28 21
Memorable 860 7 14 9
Trophy 1,020 1 5 1
a This study.b Quinn (1988).c Ashley and Buff (1986).
was less than 30 mm/year for these older age-
classes.
An inverse relationship was observed between
Wr and TL of flathead catfish in the lower St. Jo-
seph River (r2 0.75; P 0.001; Figure 6). Fish
condition was similar for individuals less than 300
mm TL (Wr
115%), whereas condition variedamong 300700-mm fish. Mean Wr of 300399-
mm fish was 98%, while the mean Wr for 400
700-mm fish was 113%. Relative weight was low-
est for flathead catfish greater than 700 mm TL
(mean, 88%).
The RSD estimates indicated that 58% of fish
greater than the minimum stock size (350 mm TL)
were quality length (510 mm TL; Table 1). Flat-
head catfish in the preferred-size range (710 mm
TL) composed 28% of the stock, while memorable-
size individuals (860 mm TL) made up 7% of
fish collected. Trophy fish (1,020 mm TL) rep-resented 1% of fish captured during the study pe-
riod.
Discussion
Unexploited fish stocks are characterized by
high population abundance, a low rate of annual
mortality, a broad range of fish age-classes and
length-classes, and decreased annual growth (Cla-
dy et al. 1975; Goedde and Coble 1981). In ad-
dition, the presence of large fish with reduced con-
dition further suggests little or no exploitation due
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1197ST. JOSEPH RIVER FLATHEAD CATFISH
to increased competition for suitable prey resourc-
es and the increased occurrence of senescence
(Van Den Avyle 1993). Previous studies have char-
acterized the flathead catfish as a long-lived spe-
cies (
15 years) that exhibits relatively fastgrowth (approximately 100 mm/year; Barnickol
and Starrett 1951; Layher and Boles 1979; Guier
et al. 1984; Jackson 1999; Nash and Irwin 1999).
In addition, flathead catfish are known to reach
TLs in excess of 900 mm in systems with low rates
of exploitation (Hesse 1994; Stauffer et al. 1996).
Our study results suggest that the population char-
acteristics and stock dynamics of flathead catfish
in the lower St. Joseph River are similar to those
of other midwestern stocks that receive low ex-
ploitation rates.
Absolute population abundance estimates offlathead catfish in lotic systems have not been re-
ported; therefore, we could not make direct com-
parisons with other studies. However, the esti-
mated density (145 fish/rkm) of flathead catfish in
the lower St. Joseph River was comparable to den-
sities reported for other systems. Marsh et al.
(1988) estimated that the density of flathead catfish
ranged from 155 to 259 fish/km in the lower Col-
orado River, Arizona. Similarly, flathead catfish
density estimates ranged from 159 to 249 fish/km
in the Missouri River, Nebraska (Tondreau 1988).Therefore, our estimate of flathead catfish density
demonstrates that the lower St. Joseph River sup-
ports fish numbers similar to those seen in other
riverine systems.
Survival rates for flathead catfish have not been
quantified in lotic systems because of the limited
data availability and an overreliance on length- and
age-frequency distributions to qualitatively de-
scribe fish survival. For example, Stauffer et al.
(1996) attempted to estimate survival and mortal-
ity rates for flathead catfish and found that the
number of captured fish from each age-class did
not decline with increasing age. As a result, these
authors concluded that flathead catfish survival
was high. Although estimates of flathead catfish
survival are not available, the annual survival of
fish in the lower St. Joseph River (67%) was in
agreement with estimates for lightly exploited
populations of channel catfish. For example, Ku-
beny (1992) determined that the annual survival
of channel catfish was 74% in the James River,
South Dakota, whereas annual survival rates as
high as 87% have been reported for some Iowastreams (Paragamian 1990). Similarly, Gerhardt
and Hubert (1991) estimated the annual survival
of channel catfish to be 77% in the Powder River,
Wyoming; high survival was attributed to a low
rate of exploitation. The similarity in annual sur-
vival rate between the lower St. Joseph River and
these previously studied systems further suggests
relatively low exploitation of flathead catfish inthe St. Joseph River.
Because flathead catfish are long-lived and have
relatively low fecundity (approximately 6,900
11,300 eggs/female), the size structure and age
structure of populations of this species may be
altered by excessive harvest (Jenkins and Burk-
head 1994; Stauffer et al. 1996; Jackson 1999).
Overexploited populations of flathead catfish have
been characterized by the presence of large num-
bers of small fish (400 mm TL) and the absence
of larger individuals. In contrast, populations that
receive little harvest pressure possess a more uni-form length-frequency distribution over a greater
range of fish length-classes. Hesse (1994) reported
that the percentage of flathead catfish greater than
the legal length limit (457 mm TL) in the highly
exploited Missouri River was less than 5% be-
tween 1974 and 1993. In contrast, greater than
25% of flathead catfish collected in the lightly ex-
ploited Flint River, Georgia, were greater than this
length (Quinn 1989). Flathead catfish greater than
500 mm TL represented 47% of fish sampled in
the Minnesota River (Stauffer et al. 1996), whereexploitation of flathead catfish has been estimated
at less than 1% (Leitch and Baltezore 1987). Al-
though a large proportion (89%) of the flathead
catfish collected in the lower St. Joseph River were
less than 400 mm, the relative frequency of fish
greater than this length and the presence of fish as
old as age 17 suggest high survival and a low rate
of exploitation in this system. However, the sharp
decline in abundance between ages 3 and 4, which
corresponds to the age at which most fish were
recruited to the harvestable fishery (381 mm
TL), suggests that flathead catfish do experience
some harvest pressure in the lower St. Joseph Riv-
er.
The RSD estimates of flathead catfish in the low-
er St. Joseph River were within the range reported
for other populations of this species (Table 1).
Quinn (1989) reported the RSD-Q of flathead cat-
fish in the Flint River fishery to be 57%, whereas
fish in the Cape Fear River had a RSD-Q of 49%
(Ashley and Buff 1986). The proportions of pre-
ferred, memorable, and trophy sizes of fish in the
lower St. Joseph River were also similar to pro-portions reported for these stocks. In other lotic
systems receiving low rates of exploitation, trophy
fish have been found to make up less than 5% of
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1198 DAUGHERTY AND SUTTON
TABLE 2.Comparative regression relationships based on mean length-at-age data, mean annual regional surface air
temperature (MAST), and geographic location for flathead catfish populations in 10 U.S. rivers. The MAST data were
based on 19711999 averages published by the National Oceanic and Atmospheric Administration. Regression equations
with common letters were not significantly different (ANCOVA: F 3.59; df 9, 69; P 0.001). Fish older than
age 12 were not included in the analysis because of small sample sizes (N 5 fish) or limited availability among
populations.
River CoordinatesMAST
(C) Regression r2 N P
Minnesota River (Minnesota)a 44N, 94W 7.5 TL 111.7 69.4 (Age) z 0.96 12 0.0001
St. Joseph River (Michigan)b 42N, 86W 7.5 TL 69.2 73 (Age) z 0.96 12 0.0001
Mississippi River (IowaIllinois)c 41N, 90W 10 TL 98.9 71.7 (Age) z 0.99 11 0.0001
Salt River (Missouri)d 40N, 92W 12.5 TL 34.6 61.7 (Age) z 0.99 9 0.0001
Missouri River (Nebraska)e 40N, 95W 10 TL 43.4 80.5 (Age) z 0.93 7 0.0004
Blue River (Kansas)f 39N, 96W 12.5 TL 46.0 108.4 (Age) y 0.99 7 0.0001
Flint River (Georgia)g 33N, 83W 17.5 TL 147.6 105.3 (Age) y 0.97 7 0.0001
Cape Fear River (North Carolina)h 35N, 79W 15 TL 117.7 101.5 (Age) y 0.97 8 0.0001
Colorado River (ArizonaCalifornia)i 32N, 114W 22.5 TL 6.6 112.9 (Age) y 0.99 9 0.0001
Rio Grande River (Texas)j 30N, 101W 20 TL 72.3 107.9 (Age) y 0.99 7 0.0001
a Stauffer et al. (1996).b This study.c Barnickol and Starrett (1951).d Purkett (1958).e Holz (1969).f Minckley and Deacon (1959).g Quinn (1988).h Guier et al. (1981).i Young and Marsh (1990).j Pate (1980).
flathead catfish populations (Hesse et al. 1978;
Pugibet and Jackson 1991; Insaurralde 1992).
These studies corroborate our results, indicatingthat the size structure of flathead catfish in the
lower St. Joseph River provides further evidence
of low rates of exploitation.
Length-at-age data for flathead catfish in the
lower St. Joseph River were within the range of
growth estimates reported for other midwestern
populations of this species. However, fish growth
was slower than that reported for populations
throughout the southern United States (Table 2).
The growth of fish in our system most closely re-
sembled the growth of flathead catfish in the Mis-
sissippi River, IowaIllinois, and the Minnesota
River, whereas the average growth rate of flathead
catfish in southern systems was 35% greater than
that of St. Joseph River fish (Table 2). Similar
latitudinal trends in growth have been reported for
other fish species (Power and McKinley 1997;
Braaten and Guy 2002). Our results suggest that
the thermal gradient and length of the growing
season associated with geographic location are re-
liable indicators of flathead catfish growth rates.
The declines in annual growth rates observed
for flathead catfish in the lower St. Joseph Riverhave been observed for other flathead catfish
stocks throughout the species range. Purkett
(1958) found that the mean annual growth incre-
ment of age-14 flathead catfish was 68 mm/year,
while the growth of fish older than age 4 decreased
by 26%. Stauffer et al. (1996) estimated that thegrowth rate of age-15 flathead catfish in the Min-
nesota River averaged 120 mm/year. In contrast,
the mean annual growth increment for fish aged
610 in this system decreased to 50 mm/year,
whereas the growth of fish older than age 10 av-
eraged 34 mm/year. Mayhew (1969) reported that
annual growth increments were greater than 120
mm/year for fish younger than age 3, whereas the
growth of older age-classes declined to less than
40 mm/year by age 7. Previous studies have de-
termined that flathead catfish become reproduc-
tively mature between ages 3 and 5 (Barnickol and
Starrett 1951; Minckley and Deacon 1959; Turner
and Summerfelt 1971). Although the onset of re-
productive maturity in flathead catfish may con-
tribute to declines in the annual growth rate, the
variation among populations suggests that addi-
tional factors (e.g., population density, prey avail-
ability, etc.) may also affect annual growth on a
system-specific basis.
The relationship between Wr and TL of flathead
catfish in the lower St. Joseph River contrasted
with results reported for other populations of flat-head catfish. Guier et al. (1984) and Lemmons
(1995) found that the condition of flathead catfish
increased with body length. Although food habits
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1199ST. JOSEPH RIVER FLATHEAD CATFISH
and relative abundance of prey types for flathead
catfish were not examined in our study, the decline
in Wr with increasing length may indicate a lack
of adequate prey sizes for larger fish. Adult flat-
head catfish are known to be highly piscivorous(Minckley and Deacon 1959; Turner and Sum-
merfelt 1971; Jackson 1999); forage fish make up
as much as 95% of the diet of fish greater than
600 mm TL (Weller and Robbins 1999). Studies
by Brown and Dendy (1961) and Haas et al. (2001)
found a positive relationship between prey size and
the TL of flathead catfish, suggesting that larger
fish selected larger forage. The high relative abun-
dance of large flathead catfish in the lower St. Jo-
seph River, which results from low exploitation
and high survival, may limit the availability of
prey items selected by these fish. Future studiesshould assess flathead catfish diet and the popu-
lation structure and proportional stock densities of
forage fishes to provide a better understanding of
predatorprey dynamics in this system.
The results of our study indicate that flathead
catfish in the lower St. Joseph River are compa-
rable in biological attributes and stock dynamics
to populations throughout the midwestern United
States. The survival rate, size structure, and age
distribution of flathead catfish in the lower St. Jo-
seph River were comparable to those of stocksknown to receive low exploitation rates, suggest-
ing that harvest pressure is relatively low in this
system. However, little information exists regard-
ing system-specific biotic and abiotic factors that
influence the stock characteristics and population
dynamics of flathead catfish. For example, future
studies should attempt to determine factors that
influence the abundance, growth, and condition of
flathead catfish. As angler attitudes and interests
change regarding this species, efforts must be un-
dertaken to quantify exploitation rates and sub-
sequent impacts on flathead catfish survival and
population structure. Studies that examine flathead
catfish recruitment would increase our understand-
ing of stock structure and describe annual vari-
ability in year-class strength and recruitment dy-
namics. These data would provide fisheries man-
agers with the information necessary to develop
appropriate management strategies for flathead
catfish populations throughout the species geo-
graphic range.
Acknowledgments
We would like to thank S. Donabauer, L. Zurita,
R. Wyld, A. Gima, and S. Reed for their assistance
with field data collections. J. Wesley and J. Dexter,
Michigan Department of Natural Resources, Fish-
eries Division, provided assistance with project
development and sampling equipment. Construc-
tive comments on earlier drafts of this manuscript
by A. Benson, M. Hansen, P. Hrodey, R. Swihart,H. Weeks, and two anonymous reviewers im-
proved this manuscript. Funding for this project
was provided by the Great Lakes Fishery Trust and
Purdue University Department of Forestry and
Natural Resources. This research was approved for
publication as manuscript number 17405 by the
Purdue University Agricultural Research Pro-
grams.
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