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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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    1192 DAUGHERTY AND SUTTON

    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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