Age and growth of blue catfish, Ictalurus furcatus (Lesueur, 1840), among three exploited and one unexploited waterbodies in Tennessee David R. Stewart, Jr. A thesis submitted to the graduate faculty in partial fulfillment of the requirements for the degree of Master of Science Middle Tennessee State University May 2009 ii Age and growth of blue catfish, Ictalurus furcatus (Lesueur, 1840), among three exploited and one unexploited waterbodies in Tennessee David Randall Stewart, Jr. Approved: George W. Benz, Ph.D., Major Professor Dennis M. Mullen, Ph.D., Committee Member Brian T. Miller, Ph.D., Committee Member George G. Murphy, Ph.D., Chair, Department of Biology Michael D. Allen, Ph.D., Dean, College of Graduate Studies iii Age and growth of blue catfish, Ictalurus furcatus (Lesueur, 1840), among three exploited and one unexploited waterbodies in Tennessee David R. Stewart Jr. Abstract The blue catfish, Ictalurus furcatus, is an important sport and commercial resource in Tennessee. Recently, the species has been designated a Tennessee sportfish, allowing the Tennessee Wildlife Resources Agency to apply for federal sportfishery enhancement funding to support its management. However, few studies have been published on blue catfish in Tennessee that contain biological data of interest to fishery managers. Furthermore, at least some of the results of the publications of interest are considered dubious, as more recent research has indicated that they are based on older and less reliable methods of analysis. With this in mind, the present study was conducted to primarily gather weight-length relationship data, relative weight estimates, and preliminary age and growth information for blue catfish in four Tennessee waterbodies: Lake Barkley, Kentucky Lake, the Mississippi River, and Fort Loudoun Reservoir. In this study, a total of 779 blue catfish were collected from fall 2006 to fall 2008 from fish wholesalers and commercial fishermen and biologists (Lake Barkley 169 fish sampled, Kentucky Lake 172 fish sampled, Mississippi River 248 fish sampled, Fort Loudoun Reservoir 190 fish sampled). Standard weight-length relationships for each waterbody, as well as a preliminary state-wide relationship based on pooled data provided good to excellent predictive power, with r2 values for the Kentucky Lake, Mississippi River, Fort iv Loudoun Reservoir, and the state-wide weight-length relationships all equaling 0.95 or higher and that of Lake Barkley equaling 0.74. Statistical differences in weight-length relationship slopes or intercepts were found between the following waterbodies: Lake Barkley vs. Kentucky Lake (Kentucky Lake fish adding body weight more rapidly with increasing body length than Lake Barkley fish), Lake Barkley vs. Fort Loudoun Reservoir (Fort Loudoun Reservoir fish adding body weight more rapidly with increasing body length than Lake Barkley fish), Kentucky Lake vs. the Mississippi River (Kentucky Lake fish adding weight more rapidly with increasing body length than Mississippi River fish), Mississippi River vs. Fort Loudon Reservoir (Fort Loudoun Reservoir fish adding weight more rapidly with increasing body length than Mississippi River fish), Lake Barkley vs. the Mississippi River (Lake Barkley fish weighing more overall than Mississippi River fish, but adding body weight at a similar rate per increase in body length), and Kentucky Lake vs. Fort Loudoun Reservoir (Kentucky Lake fish weighing more overall than Fort Loudoun Reservoir fish, but adding body weight at a similar rate per increase in body length). Calculations of relative weight revealed that blue catfish of all sizes in Lake Barkley and Kentucky Lake were above the species weight standard (by 21 and 12 percent, respectively); whereas, fish from the Mississippi River and Fort Loudoun Reservoir collections were below the standard (by 6 and 14 percent, respectively). Blue catfish aged in this study (using otoliths as the aging structure) ranged from 0 to 34 years old and 70 to 1,115 mm total length, with the 34-year-old individual representing the oldest blue catfish reported to date by any published study. Significant differences in growth were detected between some of the study waterbodies, with blue catfish in Kentucky Lake generally being longer per age than those in Fort v Loudoun Reservoir, in the Mississippi River generally being longer per age than those in Fort Loudoun Reservoir, in Lake Barkley generally being longer per age than those in the Mississippi River, in Lake Barkley generally exhibiting similar length at age as fish in Kentucky Lake, except for age 4 catfish, and in Lake Barkley generally being longer per age than those in Fort Loudoun Reservoir. This study was designed to detect and describe patterns, rather than reveal the processes underlying them. Consequently, the causes of the aforementioned differences and similarities in weight-length relationships, relative weight estimates, and age and growth patterns remain unknown. Nonetheless, the results provided by this study should prove valuable to fisheries managers in Tennessee as they consider how best to manage the state’s catfish resources. vi Acknowlegements I thank Cool Cats Fish Market, Harts Fish Market, B & F Fish Market, and Quillens Fish Market, and especially Danny Shelton, Tim Case, and Andy Eldridge (all Cool Cats Fish Market), Dennis Duncan and family (Harts Fish Market), and Jimmy Quillen (Quillens Fish Market), for their unmeasurable assistance and hospitality throughout the project. I thank George Scholten (Commercial and Reservoir Coordinator, Tennessee Wildlife Resources Agency, TWRA) for the opportunity to study catfish age and growth, guidance, and assistance regarding project and non-project matters; and Jackie Childress and Reggie Wiggins (both TWRA) for providing valuable assistance as we braved the commercial wholesalers in the midst of an all too hot summer to gather samples from Lake Barkley and Kentucky Lake. I thank Eric Ganus (TWRA) for assistance with sample collection and catfish aging, Pat Black (TWRA) for help with statistical software as well as asistance in the field, the TWRA big head carp round-up squandron (Bobby Wilson, George Scholten, Doug Peterson, Pat Black, Frank Fiss, Mike Bramlett, Mike Jolley, John Hammonds, Steve Henegar, Jim Negus, Rob Lindbom) and others I may have forgotten who assisted with sample collection in the Mississippi River; and Ed Scott (formerly with the Tennessee Valley Authority) for help with sample collection from Fort Loudoun Reservoir. A unquantifiable thanks is extended to Dr. George W. Benz (Middle Tennessee State University, MTSU) for valuable guidance and support during various stages of the project, and Drs. George W. Benz, Brian T. Miller, and Dennis M. Mullen (all MTSU) for constructive comments regarding this thesis. Special thanks go to Shirley and Randy Stewart for moral, loving, and financial support during my graduate program vii at MTSU. This project was partially funded by the Tennessee Wildlife Resources Agency, a George G. Murphy Research Scholarshp, a J. Gerald Parchment Biological Field Station Scholarship, and travel support from the MTSU Department of Biology, College of Basic and Applied Sciences, and the College of Graduate Studies, for which I am grateful. viii Table of Contents Abstract ..................................................................................................................... iii Acknowledgements ................................................................................................... vi Table of Contents ...................................................................................................... viii List of Tables ............................................................................................................ x List of Figures ........................................................................................................... xii Introduction ............................................................................................................... 1 Materials and Methods .............................................................................................. 5 Catfish Harvest Report .............................................................................. 5 Waterbodies Studied ................................................................................. 7 Sample Collection .................................................................................... 8 Otolith Removal and Processing .............................................................. 10 Data Anaylses ........................................................................................... 11 Results ....................................................................................................................... 14 Catfish Harvest Report ............................................................................. 14 Samples Collected .................................................................................... 16 Weight-length Relationships and Relative Weights ................................. 17 Age and Growth ....................................................................................... 19 Discussion ................................................................................................................. 22 Catfish Harvest Report ............................................................................. 23 Weight-length Relationships and Relative Weights ................................. 24 Age and Growth ....................................................................................... 26 ix Conclusions. ............................................................................................. 30 References ................................................................................................................. 34 Tables ........................................................................................................................ 39 Figures ....................................................................................................................... 56 x List of Tables Table 1. Sport fishery creel survey analysis results for catfish (blue catfish, Ictalurus furcatus; channel catfish, I. punctatus; flathead catfish, Pylodictis olivaris) and bullheads (black bullhead, Amerius melas; yellow bullhead, A. natalis) caught in three Tennessee waterbodies during 1998-2006. .............. 39 Table 2. Monthly 2005 commercial catfish harvest (kg) for 17 Tennessee waterbodies calculated from Tennessee Wildlife Resources Agency commercial fishing data. ......................................................................................................... 41 Table 3. Sport and commercial harvest of catfish (blue catfish, Ictalurus furcatus; channel catfish, I. puntatus; and flathead catfish, Pylodictis olivaris) for 22 waterbodies for the year 2006. . ........................................................................... 42 Table 4. Number of blue catfish, Ictalurus furcatus, sampled from four study waterbodies. ........................................................................................................ 44 Table 5. Parameters of log10 transformed weight-length linear regressions for four studied Tennessee waterbodies and log10 transformed weight-length linear regression for pooled data from the same four waterbodies used to represent Tennessee. ............................................................................................................ 47 Table 6. Overall and mean relative weights for five size classes of blue catfish, Ictalurus furcatus, sampled from four Tennessee waterbodies. ......................................... 48 Table 7. Mean total length at age (mm) for blue catfish, Ictalurus furcatus, in four Tennessee waterbodies. ....................................................................................... 49 xi Table 8. Fishery mileposts for blue catfish, Ictalurus furcatus, in four Tennessee waterbodies. ....................................................................................................... 50 Table 9. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Lake Barkley (Tennessee and Kentucky). ................ 51 Table 10. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Kentucky Lake (Tennessee and Kentucky). .............. 52 Table 11. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Mississippi River (Tennessee and Louisianna). ........ 54 Table 12. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Fort Loudoun Reservoir (Tennessee), Lake Hugo (Oklahoma), Lake Ellsworth (Oklahoma), and Lake Eufaula (Oklahoma). ....... 55 xii List of Figures Figure 1. Major river systems of Tennessee. ........................................................... 56 Figure 2. Location and removal of sagittal otoliths from catfish. ............................ 57 Figure 3. Epoxy casts containing catfish otoliths. ................................................... 58 Figure 4. Weight-length relation for blue catfish, Ictalurus furcatus, collected from Lake Barkley (Tennessee portion of lake; formula for log10 transformed weight-length linear regression and r2 value provided at upper left). ........................................ 59 Figure 5. Weight-length relation for blue catfish, Ictalurus furcatus, collected from Kentucky Lake (Tennessee portion of lake; formula for log10 transformed weightlength linear regression and r2 value provided at upper left). .............................. 60 Figure 6. Weight-length relation for blue catfish, Ictalurus furcatus, collected from the Mississippi River (formula for log10 transformed weight-length linear regression and r2 value provided at upper left). ........................................................................... 61 Figure 7. Weight-length relation for blue catfish, Ictalurus furcatus, collected from Fort Loudoun Reservoir (formula for log10 transformed weight-length linear regression and r2 value provided at upper left). .................................................................... 62 Figure 8. Weight-length relation for blue catfish, Ictalurus furcatus, collected from four Tennessee waterbodies (Lake Barkley, Kentucky Lake, Mississippi River, Fort Loudoun Reservoir; formula for log10 transformed weight-length linear regression and r2 value provided at upper left). .................................................................... 63 Figure 9. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for four Tennessee waterbodies. ........................................................ 64 xiii Figure 10. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Lake Barkley. ............................................................................... 65 Figure 11. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Kentucky Lake. ............................................................................ 66 Figure 12. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Mississippi River. ........................................................................ 67 Figure 13. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Fort Loudoun Reservoir. .............................................................. 68 1 Introduction Ictalurids (Ictaluridae) are among the most frequently fished freshwater fishes in the United States. The National Survey of Fishing, Hunting, and Wildlife-Associated Recreation reported that, in 2006, seven million US anglers fished 98.2 million days for catfish (USFWS and USCB 2006). Catfish (blue catfish, Ictalurus furcatus (Lesueur 1840); channel catfish, I. punctatus (Rafinesque 1818); flathead catfish, Pylodictis olivaris (Rafinesque 1818); black bullhead, Ameiurus melas (Rafinesque 1820); and yellow bullhead, A. natalis (Lesueur 1819)), formed the third most sought after fish group, behind black bass (Micropterus spp.) and panfish (some non-bass members of Centrarchidae) (USFWS and USCB 2006). In Tennessee, blue catfish, channel catfish, and flathead catfish comprised the third most sought after group of fishes, behind black bass and crappie (black crappie, Pomoxis nigromaculatus (Lesueur 1829) and white crappie, P. annularis (Rafinesque 1818); (USFWS and USCB 2006; Pat Black, Tennessee Wildlife Resources Agency, unpublished report). The blue catfish is the largest ictalurid and can reach 165 cm total length (TL) and 68 kg in weight (Etnier and Starnes 1993). Although the species may sometimes be confused by anglers with other catfishes (unpublished observations of author), I. furcatus is easily differentiated from its congeners by the following characteristics: 30-36 anal fin rays, distinctive coloration (pale grey, white to blue dorsally; white ventrally), and a lower jaw that never extends beyond the upper jaw (Etnier and Starnes 1993). Blue catfish are native to 29 states within the Mississippi, Missouri, and Ohio river drainages, where they most commonly inhabit large bodies of water (Etnier and Starnes 1993). Blue 2 catfish generally mature at 35-66.2 cm TL and 4 or 5 years of age in the South and 38.148 cm TL and 6 or 7 years of age in the North (Graham 1999). Fourteen states, including Tennessee, support recreational and commercial fisheries for catfish (Graham 1999). Recreational angling for catfish has recently become more popular, as evidenced by increased media exposure, such as fishing-related television programming and sports magazines (Graham 1999). This increase in popularity has manifested itself in Tennessee as well, and in 2008 In-fisherman Magazine included Tennessee’s Cumberland River system as one of the ten best blue catfish fisheries in the nation (Hoffman 2008). Associated with this catfish angling invigoration, the International Game Fish Association has recognized four world-record blue catfish since 1996. The current all-tackle world record for the species, 56 kg and 147.3 cm TL (which broke the previous record by 1.4 kg) was caught in the Mississippi River near Alton, IL in 2005. Other large blue catfish have generated further attention, such as those landed at a 2008 Memphis, TN based Mississippi River catfish tournament, where for the first time, two catfish of any species weighing > 45 kg were brought to the scales (Brasher 2007). In Tennessee, there were 237 resident and 6 non-resident commercial fishing licenses issued in 2008 permitting holders to harvest catfish. Commercially harvested catfish (blue catfish, channel catfish, and flathead catfish) in Tennessee are sold to commercial fish wholesalers who distribute the flesh for human consumption. As of October 2008, the harvest price for catfish in Tennessee was US$0.45 per lb (Dennis Duncan, Hart’s Fish Market, personal communication). Blue catfish comprises the heaviest annual 3 commercial catch, followed by channel catfish, and flathead catfish (George Scholten, Tennessee Wildlife Resources Agency, personal communication). Regarding the two most-harvested catfishes, blue catfish are typically more desirable to commercial fishermen because consumers prefer their white flesh, which lacks the off-colored tint typically exhibited by flesh of channel catfish (Danny Shelton, Cool Cats Fish Market, personnel communication). Catfish (Ictalurus spp. and P. olivaris) were primarily managed as commercial species in Tennessee prior to 2003, and sportfishermen harvesting them were only required to hold a valid sportfishing license. The commercial harvest was regulated through license and gear restrictions, i.e., commercial fishermen were required to hold a valid general commercial fishing license and were restricted from fishing within 91 m (300 ft) of a river mouth or within 914 m (3,000 ft) downstream of any Tennessee Valley Authority or US Army Corps of Engineers dam (George Scholten, Tennessee Wildlife Resources Agency, personnel communication). In 2001, Arterburn et al. (2001) reported results of a comprehensive catfish angler survey within the greater Mississippi River basin that documented a widespread desire by anglers for increased management of catfish to enhance sportfishing. The Arterburn et al. (2001) survey results, along with the public’s growing interest in recreational catfish angling, prompted the Tennessee Wildlife Resources Agency (TWRA) to initiate a series of angler opinion surveys (2000, 2001, 2005) regarding catfish management (George Scholten, Tennessee Wildlife Resources Agency, unpublished data). Results of these surveys indicated that Tennessee’s catfish anglers supported the concept of TWRA managing catfish as sportfish by using a variety 4 of fisheries practices (George Scholten, Tennessee Wildlife Resources Agency, unpublished data). Those results, supported by attention that the catfishery received in association with the International Ictalurid Symposium (Irwin et al. 1999), prompted TWRA to re-evaluate catfish management within the state. Based on that process and subsequent deliberations, Tennessee’s catfish harvest regulations were modified as follows in March 2003: 1) catfish < 86.4 cm (< 34 in) TL could be harvested by commercial and sportfishermen, 2) sportfishermen could additionally harvest one catfish per day ≥ 86.4 cm TL, but only possess two catfish ≥ 86.4 cm TL at any one time, and 3) commercial fishermen could not fish within 91 m of a river mouth or within 914 m downstream of any Tennessee Valley Authority or US Army Corps of Engineers dam. After intense lobbying by commercial fishermen to establish parity with the sport fishery, commercial regulations were modified in 2006 such that commercial fishermen could also harvest one catfish per day ≥ 86.4 cm TL but only possess two catfish ≥ 86.4 cm TL at any one time. The Tennessee Wildlife Resources Commission reclassified blue catfish, channel catfish, and flathead catfish as sport and commercially fished species in fall 2007, a decision that allowed federal sportfish restoration funds to be utilized for catfish management. Some efforts by TWRA since that decision have been aimed at collecting basic information about catfish populations in Tennessee to aid in the assessment of the aforementioned management regulations. Age and growth data and weight-length relationships are used by biologists in a number of analytical assessments (e.g., estimation of growth rates, length and age at maturity, age at time of fishery recruitment, and duration fishes spend within a fishery) to 5 enhance fishery management and monitoring (Beverton and Holt 1957; Ricker 1975). Few age and growth data exist regarding catfish in Tennessee waters, with only two reports focusing on blue catfish. Conder and Hoffarth (1965) reported on the age and growth of blue catfish in Barkley Reservoir and Kentucky Lake and Hale and Timmons (1990) reported on age and growth of blue catfish in Kentucky Lake. Unfortunately, these studies assigned fish ages based on assessments of growth marks on pectoral fin spines, a methodology now considered unreliable because it results in the underestimation of the age of older fish (Buckmeier et al. 2002). Although age and growth data exist for blue catfish outside of Tennessee (but the majority of these based on fishes aged using fin spines) (Graham 1999), observed differences in age and growth of blue catfish amongst various waterbodies (Boxrucker and Kuklinski 2006) strengthens an argument in favor of obtaining waterbody specific data within Tennessee to support stock monitoring and management decisions. With the above in mind, the purpose of this study was to gather information useful to TWRA’s fishery managers regarding age and growth, weight-length relationships, and relative weight of blue catfish in three exploited and one unexploited Tennessee waterbodies. In addition, unpublished TWRA harvest data that was used to support the planning of the aforementioned study is also presented. Materials and Methods Catfish Harvest Report The catfish harvest component of this study was carried out to help identify waterbodies of interest regarding the catfish age and growth studies that form the bulk of this report and to provide TWRA with some preliminary metrics regarding Tennessee’s catfish 6 harvest. Hence, no effort was made to measure harvest data variance or gather data in a manner that supported any statistical analyses. Tennessee Wildlife Resources Agency 1998-2006 creel survey data for blue catfish, channel catfish, and flathead catfish were collected and assessed, as available, for Lake Barkley, Kentucky Lake, and Fort Loudoun Reservoir. Specifically, annual creel data were reviewed and two annual species averages were calculated across the nine years for each waterbody: average mean total weight caught (kg) and average mean total weight harvested (kg). Annual TWRA creel data for blue catfish, channel catfish, flathead catfish, and the bullheads Amerius melas and A. natalis were combined to obtain average of the number of angler hours fished and the number of catfish caught per hour for each of the years 1998-2006. The 2005 TWRA commercial catfish harvest data (reported in lb for the following 17 waterbodies: Lake Barkley, Kentucky Lake, Mississippi River, Nolichucky River, Obion River, Perryville River, Reelfoot Lake, Chickamauga Lake, French Broad River, Hatchie River, Lake Nickajack, Old Hickory Reservoir, Watts Barr Reservoir, Cheatham Lake, Lake Douglas, Lake Guntersville, Hiwassee River) were converted to kg and tallied by month for four groups (blue catfish, channel catfish, flathead catfish, and catfish spp., the latter group referring to an unidentified mixture of individuals representing the three aforementioned species). The dollar value of the harvest was estimated by applying a US$0.45 per lb (US$ 0.992 per kg) multiplier (a value in line with information supplied by wholesale representatives, see above). Lastly, the total catfish harvest (kg; for blue catfish, channel catfish, and flathead catfish combined) for 22 Tennessee waterbodies (Chickamauga Lake, Lake Barkley, Kentucky Lake, Reelfoot Lake, Pickwick Lake, Watts Bar Reservoir, Old Hickory Lake, Lake Douglas, Percy Priest Reservoir, Lake Cherokee, 7 Normandy Lake, Woods Reservoir, Boone Lake, Tims Ford Lake, Norris Lake, Melton Hill Lake, Dale Hollow Lake, Lake Watauga, Center Hill Lake, South Holston Reservoir, Fort Loudoun Reservoir, Tellico Lake) was calculated by combining values for the sport harvest and commercial harvest for the year 2006 (George Scholten, Tennessee Wildlife Resources Agency, unpublished data). From this data, the contribution of the sport catfish and commercial catfish harvests were each be expressed as a percent of the total catfish harvest for each waterbody and for Tennessee as a whole, as was the harvest (kg) per ha for each waterbody. Waterbodies Studied Lake Barkley, Kentucky Lake, and the Mississippi River were selected for study as exploited waterbodies based on their high levels of catfish harvest in recent years (as determined from the catfish harvest analysis and discussion with TWRA staff). Fort Loudoun Reservoir was added to the study to represent an unexploited reservoir. Fort Loudoun Reservoir has been closed to commercial and recreational harvest since 1979 because of polychlorinated biphenyls (PCB) contamination (Bobby Wilson, Tennessee Wildlife Resources Agency, personnel communication). Lake Barkley is an impoundment of the Cumberland River possessing a 1,616-km shoreline (Tennessee portion of the lake has a surface area of 7,406 ha) located in southwestern Kentucky and north-central Tennessee (Figure 1). Kentucky Lake is the largest reservoir in Tennessee, with a surface area of 43,794 ha and over 3,322 km of shoreline extending from Stewart County to Hardin County, TN (Figure 1). The Mississippi River is the second longest river in North America and it borders Tennessee 8 for 269 km from Dyer County to Shelby County, TN (Figure 1). The Mississippi River is one of the most heavily fished rivers in North America. During 2005, Tennessee waters of the Mississippi River supported a commercial catfish harvest of 65,315 kg, with blue catfish comprising 78 percent (50,794 kg) of said harvest (Tennessee Wildlife Resources Agency, unpublished data). Fort Loudoun Reservoir is a 5,908 ha impoundment of the Tennessee River, with a 610-km shoreline in East Tennessee, West of Knoxville (Figure 1). Sample Collection Catfish samples from Lake Barkley and Kentucky Lake were collected by visiting commercial fish wholesalers (Hart’s Fish Market, Paris, TN; B&F Fish Market, Perryville, TN; Quillen’s Fish Market, Paris, TN; Cool Cats Fish Market, McKinnon, TN) during 2007 (May-October) and 2008 (May-October). Commercial fishermen selling to the wholesalers fished mainly with gill nets, but they occasionally used trotlines to harvest catfish. Gill net mesh sizes varied from 7.62-10.16 cm stretched mesh, with nets fished overnight. A minimum of ten fish per 25-mm length interval (intervals starting at 200 mm TL) was sought by unselectively gathering catfish from swim tanks at the commercial fish wholesalers or from the live wells of commercial fishing boats as catches were being offloaded at the wholesalers. In addition, six catfish from Kentucky Lake were obtained from commercial fisherman during the 2008 paddlefish (Polyodon spathula (Walbaum, 1792)) season; catfish comprise most of the paddlefish harvest bycatch. Because paddlefish are harvested using 15-cm stretched mesh gill nets, sampling that by-catch facilitated the collection of samples from catfish that were larger than those 9 normally seen at the commercial catfish wholesalers. All collected fish were measured (TL, nearest mm), weighed (nearest g), tagged in the head with a uniquely numbered tag (Floy® T-bar anchor tag, Floy Tag and Mfg., Inc., Seattle, WA), and decapitated. Processing procedures used by the commercial fish wholesalers precluded efforts to gather fish sex information. Sagittal otoliths were removed (see below) from each tagged head at the end of each sampling trip. Mississippi River sampling occurred during four days in November 2006 and 2007. Three sampling methods were used at each of the three collection locations along the Mississippi River (Caruthersville, MO; Ashport, TN; and Randolph Point, TN). Boat electrofishing using low-frequency pulsed-DC (15 pulses per s) was carried out in 10-min passes around wing dikes, rip-rap, and sand bars, with the electrofishing boat accompanied by two chase boats to increase netting efficiency. A 4.88-m knotless shrimp trawl (complete shrimp trawl, model number TRL16C, Memphis Net and Twine Co., Inc, Memphis, TN) was towed for 5 min around wing dikes, rip-rap, and sand bars. Two experimental gill nets (one a 46-m net with stretched mesh sizes ranging 2.54-10.16 cm and the second a 55-m net with stretched mesh sizes ranging 6.35-8.89 cm) were fished behind wing dikes for 4-8 hr during the day. Similar to the sampling methods used for Lake Barkley and Kentucky Lake, a minimum total of ten catfish per 25-mm length interval was sought as a combined sample from the three sample locations. All blue catfish were measured (TL, nearest mm), weighed (nearest g), and sexed (gonad inspection), prior to having their sagittal otoliths removed (see below). 10 Fort Loudoun Reservoir was sampled in January and February of 2007 using 38-91 m long trot-lines set at various locations within the reservoir. A total of nine lines were set at one time (total of 36 sets) and various baits (threadfin shad, Dorosoma petenense (Günther, 1867); common carp, Cyprinus carpio L., 1758; and flathead minnow, Pimephales promelas Rafinesque, 1820) were fished on 2/0 hooks hanging from 61-cm trotters. Trotters were spaced 152 cm apart and lines were fished overnight at depths ranging from 0.91-15.24 m. All blue catfish were measured (TL, nearest mm), weighed (nearest g), and sexed (gonad inspection), and sagittal otoliths were removed (see below) from each fish. Otolith Removal and Processing Sagittal otoliths (otoliths) were removed by transversly sectioning the supraoccipital bone 3-5 mm anterior to a plane bridging the locked pectoral spines (Figure 2; Buckmeier et al. 2002). Otoliths from each catfish were placed in an individual coin envelope bearing data regarding collection location, sample date, fish length, fish weight, fish sex, and tag number. Otoliths were allowed to dry in the envelopes for 2 weeks before soft tissue and debris were cleaned from them using sharp-tipped forceps. Clean otoliths were embedded in 2-part epoxy (EasyCast Epoxy resin and hardner, Fields Landing, CA) in a plastic mold (maroon 21 flat embedding mold, model number 2449M-AB, SPI Supplies, West Chester, PA) and allowed to harden for 3 days. Four steps were used to embed otoliths in the epoxy molds. First, the molds were slightly filled with 2-part epoxy and allowed to set for 3 days. Second, clean otoliths were aligned longitudinally in the epoxy mold (Figure 3). Each mold provided its own individual identification number. Lastly, 11 the epoxy molds were filled completely with the 2-part epoxy and allowed to set for 3 days. A LECO®VC-50 low-speed isomet saw was used to section otoliths just anterior to the anti-rostrum on the rostral side of the otolith (Figure 3). After sectioning, otoliths (still embedded in epoxy) were ground to the central portion of the sulcus toward the post-rostrum side of the otolith using 320 grit sand paper (Figure 3). Otoliths were examined under a stereo-microscope at 50× using fiber-optic lights to provide side illumination. Catfish ages were assigned using standard enumeration of growth rings as described by Buckmeier et al. (2002). An otolith “reading” training period during which the author read catfish otoliths with experienced TWRA biologists preceded the formal age assignment portion of the analysis. Data Analyses Throughout the data analysis, parametric statisical tests were used for data sets with normally distributed data. In instances when data were not normally distributed, data sets were either tranformed using standard methods for each analysis or nonparametric statistical tests were employed (as noted below). Association between catfish weight and length was determined using linear weightlength regressions generated by Statistical Analysis Software, (SAS; SAS Institute 2008), according to the following equation. linear equation: log10 (W) = -a + b log10 (TL) where: W = weight in (g) TL = total length in (mm) 12 a and b = the intercept and slope, respectively, of the weight-length regression Analysis of covariance (ANCOVA, using SAS) was used to test for significant differences in the weight-length regression slopes between pairs of study lakes (every possible pair being tested). In instances when a significant difference between slopes was not found, a subsequent ANCOVA using an equal slope (a dummy variable) for each waterbody was used to test for significant differences in weight-length regression intercepts according to methods fully explained in Pope and Kruse (2007). The relative weight index (Wr; see Wege and Anderson 1978) was calculated for catfish ≥ 160 mm TL from each of the four waterbodies according to the following equation. Wr = (W/Ws) × 100 where: W = observed weight (g) Ws = length specific standard weight (g) predicted from the species specific weight-length regression (see Anderson and Neumann 1996) Analysis of variance (ANOVA, using SAS) was used to test for significant differences in Wr between waterbodies that were sampled using similar capture methods (i.e., Kentucky Lake and Lake Barkley). A Kruskall-Wallis test (using SAS) was used to test for significant differences in Wr amonst the following five catfish length intervals for each waterbody: stock fish (fish ≥ 301 and ≤ 510 mm TL), quality fish (fish ≥ 511 and≤ 760 mm TL); preferred (fish ≥761 and ≤ 890 mm TL); memorable (fish ≥ 891 and ≤ 1,140 mm TL); and trophy (fish ≥ 1,140 mm TL). 13 Catfish growth was assessed using the von Bertalanffy growth model (see Ricker 1975) according to the following equation. Lt = L∞ {1 - e-k(t-to)} where: t = age Lt = mean total length at age L∞ = theoretical maximum total length k = Brody growth coefficient to = hypothetical age at length zero The model was fit for each waterbody using the software package, Fisheries Analyses and Simulation Tools (FAST) (Slipke and Maceina 2003), with the model parameter L∞ assigned as the length of the largest fish collected in each waterbody. FAST was also used to calculate mean length at age directly from the aged samples. Analysis of covariance (ANCOVA, using SAS) was used to compare the slopes of the log10 transformed mean TL at age regressions for Kentucky Lake, Mississippi River, and Fort Loudoun Reservoir, and a Tukey multiple comparison test (using SAS) was subsequently used to identify significant growth differences amongst these waterbodies. Because the convergence criteria for the Lake Barkley log10 transformed age regression failed (thus precluding the use of a SAS ANCOVA analysis), an unpaired Student’s t-test (using SAS) was used to evaluate mean total length at age for Lake Barkley vs. the remaining three waterbodies. 14 Results Catfish Harvest Report The creel survey analysis results indicated that the 1998-2006 mean total weight caught and mean total harvest for blue catfish, channel catfish, and flathead catfish were each highest (by far) in Kentucky Lake vs. Lake Barkley or Fort Loudoun, with the Kentucky Lake mean total weight caught equaling 92.9, 93.9, and 97.8 percent of the total mean total weight caught of blue catfish, channel catfish, and flathead catfish (respectively) for the three waterbodies combined (Table 1). Although a consideration of these means based on total fishing effort within the three waterbodies was not undertaken, the Kentucky Lake catch casually appeared disproportionately high when waterbody size (surface area in ha) was considered (Table 1). Mean total harvest accounted for 0-266 percent of the mean total weight caught for blue catfish, channel catfish, and flathead catfish within the three waterbodies. The 0 percent value stemmed from the lack of harvest in Fort Loudoun Reservoir associated with fishing advisory warnings, and the value of 266 percent (calculation double-checked for accuracy) for channel catfish in Lake Barkley obviously indicated an error or errors in the TWRA creel survey database. Although the analysis was not designed to statistically test any calculated values, it seems notable that the levels of retention (i.e., mean total harvest as a percent of mean total catch) for Lake Barkley and Kentucky Lake were within 10 percent for blue catfish, channel catfish, and flathead catfish (Table 1). In 2005, commercial fishermen harvested 892,827 kg of catfish from 17 Tennessee waterbodies at an estimated wholesale value of US$885,755, with blue catfish 15 comprising 70 percent (623,402 kg) and US$618,464, channel catfish comprising 25 percent (220,805 kg) and US$219,056, flathead catfish comprising 3 percent (29,322 kg) and US$29,089, and unidentified catfish spp. comprising 2 percent (19,298 kg) and US$19,146 of said take (Table 2). The greatest combined harvest of catfish occurred during the month of May (12 percent of the 2005 annual harvest; 108,681 kg), with most of the reported fish categories indicating the same (Table 2). Total catfish harvest increased 50.1 percent (36,270 kg) from April to May and decreased 22.2 percent (24,174 kg) May to June (Table 2). Blue catfish harvest exhibited similar changes during the same periods, i.e., a 67.7 percent (31,395 kg) increase and a 21.3 percent (16,604 kg) decrease, respectively (Table 2). During the fall, blue catfish harvest rose 27.9 percent (11,065 kg) from September to October before harvest decreased by 12.9 percent (6,548 kg) from October to November (Table 2). Channel catfish harvest exhibited an initial increase, 23.4 percent (4,308 kg), from February to March and remained high through May, before decreasing by 26.7 percent from May to June. Harvest then rose by 21.2 percent (3,418 kg) from June to August before plummeting 35.1 percent (6,873 kg) in September and oscillating through the fall and early winter (Table 2). Although overall harvest levels of flathead catfish were not as high as those of blue or channel catfish, flathead catfish harvest also exhibited a large increase from April to May, i.e., 93.5 percent (1,965 kg), and another large increase from September to October, i.e., 180.7 percent (3,432 kg). Although January and December 2005 exhibited low overall catfish harvests (i.e., 7.0 percent and 7.1 percent of the annual harvest, respectively), September exhibited the lowest harvest (6.1 percent of the annual harvest). Here it seems notable 16 that even during September, Tennessee’s commercial catfish harvest represented a US$53,967 wholesale enterprise. Amongst the 22 waterbodies for which annual harvest for the year 2006 was assessed, catfish harvest ranged from a high of 17.4 kg per ha in Chickamauga Lake to a low of 0 kg per ha in Tellico Lake (Table 3). The highest sport harvest per ha occurred in Chickamauga Lake (3.44 kg per ha) as did the highest commercial harvest (13.95 kg per ha) (Table 3). Regarding waterbodies reported on in this study, Lake Barkley was Tennessee’s second most heavily exploited reservoir (13.4 kg catfish per ha) during the post-2000 annual harvest assessment (Table 3). That take, totaling 99,390 kg, was comprised of 7.2 percent sport harvest (0.97 kg per ha) and 92.8 percent commercial harvest (12.44 kg per ha) (Table 3). During that same period, Kentucky Lake was ranked Tennessee’s third most heavily exploited reservoir (9.2 kg per ha) for catfish harvest, with said take (totaling 403,313 kg) consisting of 22.7 percent sport harvest (2.09 kg per ha) and 77.3 percent commercial harvest (7.11 kg per ha) (Table 3). Fort Loudoun Reservoir, open to fishing under a consumption advisory, naturally exhibited a negligible catfish harvest (Table 3). Samples Collected A total of 779 blue catfish were collected during this study from fall 2006 to fall 2008 from the four studied waterbodies. The Lake Barkley sample consisted of 169 fish ranging in length from 275-1,115 mm TL (Table 4). The Kentucky Lake sample consisted of 172 fish ranging in length from 266-1,191 mm TL (Table 4). The Mississippi River sample consisted of 248 fish ranging in length from 70-830 mm TL 17 (Table 4). The Fort Loudoun Reservoir sample consisted of 190 fish ranging in length from 255-1,105 mm TL (Table 4). Success in collecting 10 fish in each of the 25 mm study intervals was “good to excellent” in all waterbodies for intervals ranging from 451 to 650 mm, and spotty regarding size intervals consisted of fish < 451 or > 650 mm (Table 4). Of the fish collected, all except 12 fish from Kentucky Lake were aged (these unaged fish were, however, used in the weight-length and relative weight analyses). Additionally, the six large aged fish from Kentucky Lake (representing the entire sample collected by commercial paddlefishermen) were not used in any analysis because attempts to fit the von Bertalanffy growth model using these fish resulted in nonconvergence failure of model parameters, an indication that these larger fish were significantly dissimilar from the smaller individuals comprising the bulk of the Kentucky Lake sample. Weight-length Relationships and Relative Weights Linear regressions of log10 transformed weight-length data resulted in slopes ranging from 2.87 to 3.46 and intercepts ranging from -4.6 to -6.21 for the four waterbodies and a slope of 3.21 and intercept of -5.59 for pooled data considered to represent Tennessee (Table 5; Figures 4-8). More than 95 percent of the variation in log10 weight was described by the log10 length in the regressions of Kentucky Lake, the Mississippi River, Fort Loudoun Reservoir, and Tennessee overall; whereas, 74 percent of the same variation was described by log10 length in the regression representing Lake Barkley (Table 4). Analysis of covariance revealed weight-length regressions of the following pairs of waterbodies to be significantly different based on comparisons between their 18 slopes: Lake Barkley vs. Kentucky Lake (F = 18.40, df = 1, P < 0.0001; Kentucky Lake fish adding body weight more rapidly with increasing body length than Lake Barkley fish), Lake Barkley vs. Fort Loudoun Reservoir (F = 17.01, df = 1, P < 0.0001; Fort Loudoun Reservoir fish adding body weight more rapidly with increasing body length than Lake Barkley fish), Kentucky Lake vs. Mississippi River (F = 16.06, df = 1, P < 0.0001; Kentucky Lake fish adding body weight more rapidly with increasing body length than Mississippi River fish), Mississippi River vs. Fort Loudoun Reservoir (F = 15, df = 1, P < 0.0001; Fort Loudoun Reservoir fish adding body weight more rapidly with increasing body length than Mississippi River fish). Comparisons between weightlength regression slopes revealed insignificant differences for Lake Barkley vs. Mississippi River (F = 3.43, df = 1, P > 0.0648) and Kentucky Lake vs. Fort Loudoun Reservoir (F = 0.57, df = 1, P > 0.04510). Subsequent ANCOVA testing for significant differences between weight-length regression intercepts for the last two pairs of waterbodies revealed that the intercepts of Lake Barkley and the Mississippi River were significantly different (dummy slopes = 3.08, t = -5.36, df = 1, P < 0.0001; Lake Barkley fish weighing more overall than Mississippi River fish, but adding body weight at a similar rate per increase body length) as were the intercepts of Kentucky Lake and Fort Loudoun Reservoir (dummy slopes = 3.43, t = -12.85, df = 1, P < 0.0001; Kentucky Lake fish weighing more overall than Fort Loudoun Reservoir fish, but adding body weight at a similar rate per increase body length). In the latter two cases, the overall weight of catfish for Lake Barkley (intercept = -5.187) was larger than that of the Mississippi River (intercept = -5.279), and the overall catfish weight for Kentucky Lake (intercept = 6.112) was larger than that of Fort Loudoun Reservoir (intercept = -6.229). The weight- 19 length regression for Tennessee (i.e., a regression of pooled data from all four studied waterbodies) had a slope of 3.21 and an intercept of -5.59 and differed (i.e., different slope) significantly from the regressions of each of the four waterbodies (TN vs. Lake Barkley, F = 9.01, df = 1, P < 0.0028; TN vs. Kentucky Lake, F = 7.08, df =1, P < 0.0079; TN vs. Mississippi River, F = 7.02, df = 1, P < 0.0082; TN vs. Fort Loudoun Reservoir, F = 5.70, df = 1, P < 0.0171). Lake Barkley catfish had an overall Wr of 121 and Wr values among the stock, quality, preferred, and memorable length intervals (Table 6) did not differ significantly (χ2 = 5.70, df = 4, P > 0.2223). Kentucky Lake catfish had an overall Wr of 112 and a significant difference existed among Wr values of the stock, quality, preferred, and memorable length intervals (χ2 = 16.92, df = 4, P < 0.0020) (Table 6). Mississippi River catfish had an overall Wr of 94 and no significant difference existed among Wr values of the stock, quality, and preferred length intervals (χ2 = 4.13, df = 4, P > 0.2477) (Table 6). Fort Loudoun Reservoir catfish had an overall Wr of 86 and no significant difference existed among Wr values of the stock, quality, preferred, and memorable length intervals (χ2 = 9.06, df = 4, P > 0.0597) (Table 6). Kentucky Lake and Lake Barkley catfish overall Wr values were not significantly different from one another (F = 0.68, df = 1, P > 0.4087). Age and Growth The estimated age of blue catfish ranged from 0-34 years overall in the four study lakes, with age ranges for each waterbody as follows: Lake Barkley, 2-18 years; Kentucky Lake, 2-14 years; Mississippi River, 0-21 years; and Fort Loudoun Reservoir, 5-34 years 20 (Table 7; Figure 9). The 34-year old blue catfish from Fort Loudoun Reservoir (874 mm TL female) represents the oldest estimated age reported for this species (a 24-year-old blue catfish was reported by Holley (2006)). Parameters for the von Bertalanffy growth equation were as follows for the four study lakes: Lake Barkley, L∞ = 1,115 mm TL, k = 0.11, t0 = -0.693 (Figure 10); Kentucky Lake, L∞ = 940 mm TL, k = -0.126, t0 = -1.217 (Figure 11); Mississippi River, L∞ = 830 mm TL, k = -0.145, t0 = -1.019 (Figure 12); and Fort Loudoun Reservoir, L∞ = 1,105 mm TL, k = -0.044, t0 = -1.227 (Figure 13). The von Bertalanffy equation predicted that blue catfish spend roughly 11 years in the fishery in Lake Barkley, entering at age 1.5 and reaching the trophy size limit at age 12.5 (Figure 10). Blue catfish in Kentucky Lake recruited the fishery at about 1.5 years old and remain within the fishery until at least age 11 (i.e., the age of the oldest samples used in the analysis) (Figure 11). In the Mississippi River, blue catfish entered the fishery at about 1.5 years old and remain in the fishery until they were at least 21 years old (i.e., the age of the oldest samples used in the analysis) (Figure 12). In Fort Loudoun Reservoir blue catfish entered the fishery at age 3.5 and remain in the fishery until age 34 (Figure 13). Estimated ages of blue catfish at the memorable size (890 mm TL) were 15 and 34 years old in Lake Barkley and Fort Loudoun Reservoir, respectively (Figures 10, 13). The weight of catfish as they recruited the fishery ranged from 58 to 228 g in the four waterbodies, with entry-sized catfish in Kentucky Lake weighing the most and those in Fort Loudoun Reservoir weighing the least (Table 8; Figures 4-13). The weight of blue catfish at the midpoint age within the fishery and weight at the age when leaving the fishery could only be estimated from the populations sampled in Lake Barkley and Fort 21 Loudoun Reservoir (Table 8; Figures 4, 10; 7, 13). Lake Barkley blue catfish exhibited a fishery midpoint weight of 1,778 g and a fishery exit weight of 6,761 g; whereas, Fort Loudoun Reservoir catfish exhibited a fishery midpoint weight of 1,659 g and a fishery exit weight of 7,244 g. However, fishery midpoint blue catfish within Lake Barkley were estimated to be 6 years old, whereas, those in Fort Loudoun Reservoir were estimated to be 15 years old. Furthermore, fishery exit blue catfish within Lake Barkley were estimated to be 13 years old, whereas, those in Fort Loudoun Reservoir were estimated to be 34 years old. Analysis of covariance revealed a significant difference in the log10 transformed mean TL at age regressions among Kentucky Lake, Mississippi River, and Fort Loudoun Reservoir (F = 35.01, df = 2, P < 0.0001), and a subsequent Tukey multiple comparisons test indicated significant differences in catfish total length at age between Kentucky Lake and Fort Loudoun Reservoir (Tukey HSD 33.372, P < 0.05; with blue catfish in Kentucky Lake generally being longer at a given age than those in Fort Loudoun Reservoir) and Mississippi River vs. Fort Loudoun Reservoir (Tukey HSD 48.388, P < 0.05; with blue catfish in the Mississippi River generally being longer at a given age than those in Fort Loudoun Reservoir); whereas, no significant difference in catfish total length at age was discovered between Kentucky Lake and the Mississippi River (Tukey HSD 16.016, P > 0.05). Results of Student’s t-tests between Lake Barkley and Mississippi River blue catfish revealed significant differences for 2-year-old catfish (t = 2.36, df = 19P < 0.029), 4-year-old catfish (t = 2.41, df = 51, P < 0.020), and 5-year-old catfish (t = 5.06, df = 87, P < 0.0001; with blue catfish in Lake Barkley generally being 22 longer at a given age than those in the Mississippi River). Similar testing between Lake Barkley and Kentucky Lake blue catfish 2-10 years old revealed mean total length to only be different for age 4 catfish (t = 2.22, df = 36, P < 0.033; with age 4 blue catfish in Kentucky Lake being longer than those in Lake Barkley), while similar testing between Lake Barkley and Fort Loudoun Reservoir catfish 5-11 years old revealed mean total length to be different between waterbodies for all seven ages (age 5; t = 4.65, df = 18, P < 0.0002: age 6; t = 7.00, df = 27, P < 0.0001: age 7; t = 5.09, df = 55, P < 0.0001: age 8; t = 4.62, df = 22, P < 0.0001: age 9; t = 6.23, df = 12, P < 0.0001: age 10; t = 6.04, df = 16, P < 0.0001: age 11; t = 3.78, df = 6, P < 0.0092; with blue catfish in Lake Barkley being generally longer at a given age than those in Fort Loudoun Reservoir). Discussion The blue catfish is a highly sought sport and commercial species (Graham 1999). In Tennessee, the commercial blue catfish harvest is typically sold to commercial fish wholesalers or small restaurants (Hale and Timmons 1990). However, despite the sport and commercial interest in catfish in Tennessee, biological information on Ictalurus spp. in the state is limited, with only six such reports published in the primary literature (Schoffman 1954; Carroll and Hall 1964; Conder and Hoffarth 1965; Hargis 1966; and Hale and Timmons 1990). Of these, only two studies focused on blue catfish (Conder and Hoffarth 1965; and Hale and Timmons 1990). In addition, the grey literature supplies little more information on Ictalurus spp. in Tennessee other than occurrence records, with some of this information having been repackaged for publication in the peer-reviewed literature (e.g., Conder and Hoffarth 1965; Hale and Timmons 1990). 23 Commercial Harvest Report A comprehensive review by Graham (1999) revealed blue catfish to be recreationally important in 21 states and within 14 of said states (including Tennessee) they were considered to be commercially important as well. Graham (1999) reported commercial harvests of blue catfish (for an undesignated year or years) from nine states (Arkansas, Illinois, Indiana, Kentucky, Louisiana, Mississippi, Missouri, South Carolina, and Tennessee), with harvests ranging from < 22,222 kg (Indiana) to 4,888,889 kg (Louisiana). In Graham’s report, Tennessee’s commercial blue catfish harvest was 411,153 kg, a value that placed it fifth amongst the nine reporting states regarding total harvest. However, Graham (1999) reported Tennessee’s blue catfish sport harvest of 1,381,409 kg (i.e., 335 percent of the Tennessee commercial harvest) as the highest sport take of any of the five states for which sportfishing data were available. The Tennessee commercial catfish harvest in 2005 totaled 892,827 kg, with blue catfish comprising 70 percent (623,402 kg) of said take. And, regardless of the limitations of the harvest data reported herein, the typical annual blue catfish commercial harvest in Tennessee likely totals more than several hundred thousand kilograms, with a wholesale value of at least several hundred thousands of dollars. Nationally, Miranda (1999) reported the highest mean sportfishing harvest of Ictalurus spp. (i.e., channel and blue catfish) between 1946 and 1988 to be 26.7 kg per ha (waterbody not specified). That harvest rate pertained to channel and blue catfish, and thus is not directly comparable to data reported herein. However, it is notable that Tennessee’s waterbodies experienced a total annual blue catfish harvest as high as 17.4 kg per ha (Chickamauga Lake), with the highest commercial harvest within any Tennessee waterbody reported here equaling 13.95 kg per 24 ha (Chickamauga Lake) and with the highest sport harvest also coming from that same lake (3.44 kg per ha). In the Tennessee waterbodies fished commercially and for which data were gathered, the sport harvest per ha ranged from 3.99 percent (Reelfoot Lake) to 128.87 percent (Lake Douglas) of the commercial harvest per ha in the same waters. According to the monthly 2005 commercial catfish harvest reported herein, commercial catfishing in Tennessee is a 12-month per year activity, with even the smallest monthly harvest having a wholesale value of US$53,964. Fluctuations in monthly harvest noted above could be related to a variety of biological (e.g., fish migration, reproduction), physical (e.g., water temperature, flow regime), and socioeconomic (e.g., changes in fishing effort as fishermen are diverted to other higher value enterprises, such as paddlefish harvest, during certain seasons) factors. Weight-length Relationships and Relative Weights The weight of fishes and various biomass-related data are important information regarding fisheries management, and the standard weight-length relationship in particular is an important tool used in assessing harvest (Ricker 1975). Overall, the log10 transformed weight-length relationships for each of the four waterbodies provided a high level of explanation of catfish weight based on length, with r2 values for Kentucky Lake, Mississippi River, and Fort Loudoun Reservoir all above 0.95, and Lake Barkley equal to 0.74. Increased sampling within Lake Barkley (n = 168) probably would have resulted in a better weight-length fit, as several small fish in that sample displayed high weights (see Figure 4). Furthermore, when the data were pooled for the Tennessee-wide weightlength analysis (n = 773), an r2 value of 0.95 resulted. Two types of differences were 25 exhibited in the weight-length relationships of the four waterbodies. First, there were significant differences in the slopes of the weight-length regressions between; Lake Barkley and Kentucky Lake, Lake Barkley and Fort Loudoun Reservoir, Kentucky Lake and the Mississippi River, and the Mississippi River and Fort Loudoun Reservoir. Such differences denoted different rates of association between weight and length between waterbodies. Second, there were significant differences in overall weights (i.e., intercepts) despite the fact that the weight-length slopes were similar (this type of difference was exhibited between Lake Barkley and the Mississippi River, and Kentucky Lake and Fort Loudoun Reservoir). Possible biological explanations for the two types of difference are many (e.g., growth associated genetic differences between catfish populations, differences in forage bases between waters, etc.) and both types of difference could result from a similar cause. Although Lake Barkley and Kentucky Lake are only about 15-km apart and run roughly parallel to one another for 80 km (and thus might generally be considered to possess many environmental and biological similarities), blue catfish in Kentucky Lake gained weight at a faster rate than blue catfish in Lake Barkley. Although the pooled data analysis resulted in a Tennessee-wide weightlength relationship with a high fit, this relationship differed from each of the four waterbodies. Because of those differences, I believe that “lake specific” weight-length relationships (when available) should be used for management purposes (e.g., conversion of length data to biomass data or for yield modeling purposes). Relative weight indices are now commonly used in fisheries management to compare the weights of fishes from various waters to one another and to a species specific 26 standard (Anderson and Neumann 1996). In the present study, the relative weight of catfish in the Mississippi River and Fort Loudoun Reservoir exhibited values 6 percent and 14 percent below (respectively) the species standard while Lake Barkley and Kentucky Lake exhibited values 21 percent and 12 percent higher (respectively) than the species standard. An expansion of this analysis might point to waters such as Lake Barkley (especially) and Kentucky Lake as holding potential for designation as trophy blue catfish fisheries. Age and Growth Studies of fish age and growth arguably provide some of the most useful biological fisheries management data (e.g., see Beverton and Holt 1957; Ricker 1975). Blue catfish have been aged by interpreting annual growth marks on pectoral spines (primarily; e.g., Conder and Hoffarth 1965; Porter 1969; Freeze 1977; Hale and Timmons 1989, 1990) and otoliths (more recently; e.g., Holley 2006). However, to date neither of these two aging methods have been validated for blue catfish. Buckmeier et al. (2002) did validate the use of otoliths as an accurate aging structure in channel catfish in an aging study of known age, 1-4 year-old fish. In that study, the use of pectoral spines as an aging structure (two methods: cut spines and basal–articulatory process) was also assessed. Overall, the use of pectoral spines was considered to be less accurate than the use of otoliths, with age estimates stemming from the cut spine method generally agreeing with otolith estimates for 1-3 year-old fish, but tending to underestimate the age of catfish over 3 years of age. The use of the basal-articulatory process of pectoral spines was shown to result in erratic and relatively inaccurate age estimates. Based on their results, 27 Buckmeier et al. (2002) recommended that otoliths be used as the standard aging structure to estimate the age of channel catfish. More recently, Sakaris and Irwin (2008) validated the use of otoliths for the study of daily growth increments for young-of-theyear channel catfish as old as 119 days post-hatch, with considerable accuracy in fishes up to 60 days old. Ideally, age and growth studies of fishes are conducted with multiple “readers” (age interpreters) so that individual biases can be identified and an assessment of aging precision can be conducted (e.g., Isermann et al. 2003). However, given the exigencies facing fishery management agencies, some aging efforts rely on single readers, as was the case in the present study. While this presents an increased level of uncertainty regarding the assigned ages, the bias injected into an aging study by any individual reader should be consistent such that general comparisons between populations of the same species are not invalidated. Although a single reader was used to assign ages in this study, recently, a second reader (Eric Ganus, Tennessee Wildlife Resources Agency) independently estimated the ages of a subset (n = 40) of the blue catfish otoliths used in this study from Kentucky Lake. All 40 of those estimates agreed with the ages assigned to the same fish by this study. Thus, conclusions presented here regarding the major differences in blue catfish growth (length at age) amongst the four studied waterbodies are likely robust. Notably, blue catfish in Fort Loudoun Reservoir grew at a considerably slower rate than conspecifics in Lake Barkley, Kentucky Lake, and the Mississippi River (Figure 9). To date, no growth assessment of various year classes (through the use of age and growth back-calculation methods; see Ricker 1975) of blue catfish has been reported for 28 any waterbody. The method of assessing growth used in the present study (i.e., a nonback calculation method) assigned length at age to individual catfish based on the length and age at the time of capture. Although such a method facilitates the rapid accumulation of length at age data and requires less expensive aging equipment (such that the method is preferred by some agency biologists for some studies), this method injects two sources of potential bias into the growth analysis. For one, if fishes from different waterbodies are captured at significantly different growth periods during the year (e.g., spring vs. fall of the same year), differences in growth since last annulus formation (year mark) on the aging structure can confound the analysis by increasing the variance associated with the length at age model. In the present study this bias was likely spread evenly amongst the studied waterbodies, as for the most part, sample collection from the study lakes proceeded similarly throughout the overall sampling period. A second bias that gets interjected by a non-back calculation method of aging is that each length at age value is specific to a particular year class of fish. Hence, if significant differences in growth amongst year classes within a waterbody exist, the overall variance associated with the growth model becomes inflated. The ability to determine just how egregious this type of bias is can only be determined in light of a concurrent assessment of the growth of each year class within the pooled sample. Thus, results stemming from a non-back calculation study of growth do not directly correlate to mean length at age data stemming from the averaging of length at age data of independent year classes, and the results of comparisons between the two must be considered tentative. 29 In light of the above, direct comparisons of blue catfish growth estimates reported by the present study with those reported by others must be interpreted with caution. Furthermore, based on the results of Buckmeier et al. (2002) one might expect studies that employed pectoral spines as the aging structure to report somewhat inflated values for length at age of catfish older than age 3. Blue catfish growth reported by this study within the Tennessee portion of Lake Barkley was faster for age 1-5 fish compared to blue catfish growth reported by Freeze (1977) for the Kentucky portion of the lake (Table 9). Age 6 and age 7 catfish in the Kentucky portion of the lake were estimated to be longer (Freeze 1977) than those reported by this study for the Tennessee portion of the lake. The difference in the general rate of growth between these two studies may have resulted from age underestimation of catfish deemed age 5 and age 6 by Freeze (1977), age overestimation of catfish deemed age 5 and age 6 by the present study, differences in growth between different regions of the lake, differences in growth over the 30 years separating these two studies, differences in sample size between studies, or combinations of these variables. Blue catfish growth reported by this study for Kentucky Lake was faster than any of the previous five studies of blue catfish growth in this lake in either Tennessee or Kentucky waters (Table 10). An inspection of growth increments (see Table 10) indicates some possible aging errors in the previous studies (i.e., what appear as large departures from the expected lessening of growth increments with age). However, a combination of year class and sample size effects can sometimes produce such appearances. Also, Hale and Timmons (1990) documented differences in growth 30 between blue catfish within the lacustrine and riverine portions of this lake in Tennessee waters, with the lacustrine catfish exhibiting faster growth (Table 10). Hale and Timmons (1990) proposed that a higher mortality rate (associated with high fishing mortality) within the lacustrine portion of the lake was the cause of this difference. Some evidence also exists that blue catfish in Kentucky waters of Kentucky Lake grew slower than those in Tennessee waters of the lake (cf. growth data of Hale and Timmons 1989 and 1990, as reported in Table 10). Blue catfish growth reported by this study for the Mississippi River exhibited slower growth than that reported by Kelley and Carver (1966) in their study of catfish (sample size of only 57 fish) in the Mississippi River Delta (Table 11). In addition to the dissimilarity in sample size and aging method between these two studies, the notable difference in growth could have been attributed to the lower Mississippi River possessing a longer growing season, higher prey density, or many other factors. Blue catfish in Fort Loudoun Reservoir (the slowest growing catfish in the present study) exhibited slower growth than blue catfish in three Oklahoma lakes purported to exhibit slow growth due to stunting associated with overpopulation (Boxrucker and Kuklinski 2006; Table 12), while blue catfish growth rates for Lake Barkley, Kentucky Lake, and the Mississippi River were all faster than the slow-growth Oklahoma lakes (cf. length at age data in Tables 9-12). Conclusions The present study focused on three exploited (Lake Barkley, Kentucky Lake, Mississippi River) and one recently unexploited (Fort Loudoun Reservoir) waterbodies. Unexploited 31 waters are often studied in conjunction with exploited waters in fisheries studies in hope of using them to define natural mortality and to otherwise consider the effects of fishing harvest on fish populations (e.g., see Beckwith et al. 1986). However, in the present case, although differences in the weight-length relationship, relative weight, and length at age between Fort Loudoun Reservoir and at least some of the exploited waters were documented by this study, the differences are not necessarily related to the unexploited status of Fort Loudoun Reservoir. For example, differences between blue catfish in Fort Loudoun Reservoir and the exploited waterbodies could be related to “lake specific” nonfishery-related factors such as lake productivity or forage base. To robustly consider the effects of exploitation, multiple unexploited waters are typically studied along with multiple exploited waters; and, it is further desirable to assess exploited waters in light of their relative levels of exploitation (e.g., see Beckwith et al. 1986). Because this study was designed to detect and describe patterns rather than reveal the biological processes underlying them, the causes of the differences and similarities in weight-length relationships, relative weight estimates, and age and growth patterns remain unknown. That said, the results of this study, although preliminary, should prove immediately valuable as baseline data for fisheries managers in Tennessee as they consider how best to manage the state’s catfish resources. With that task in mind, following are some recommendations regarding future studies in support of catfish management in Tennessee. One could argue that two primary research agendas should exist regarding catfish management in Tennessee: defining the fishery and gathering biological information necessary for sound management. 32 The state would benefit from continued assessments aimed at defining the monthly and annual sport and commercial harvest in waters throughout the state. Such assessments should include harvest as well as some metric of effort. Potentially important ancillary data (e.g., water temperature, wholesale price of catfish, type of gear fished by sport and commercial fishermen, etc.) should also be routinely gathered. As Tennessee is generally known as a state of opportunity regarding a quality catfishing experience (Hoffman 2008), the state might also consider greater promotion of catfishing to increase tourism and stimulate other economic benefits. Studies aimed at defining the economic impact of catfishing relative to other types of fishing within the state might also help to properly prioritize catfish management in relation to other fisheries exigencies. Several studies would be beneficial to gathering biological information required to manage catfish in Tennessee. For example, to date no report exists that validates otolith aging methods in blue catfish. Given the life history of Ictalurus spp., such a validation study should be designed to investigate possible differences between male and female catfishes, as mature males may develop false annuli associated with the periods during which they guard the developing eggs. Such a study would be relatively easy to design and would lend confidence to studies requiring age assessments. Similarly, differences in growth amongst year classes of fishes can be pronounced (for results of a recent study focusing on flathead catfish see Jones and Noltie (2007)) and yet no data regarding this phenomenon exists for blue catfish. With this in mind, a study in one to several of Tennessee’s most important reservoirs should investigate possible variations in blue catfish growth associated with sex and year class. Valuable too would be investigations 33 of possible association between sex and weight-length relationships and relative weight, fecundity at age, age and length at maturity, and studies aimed at estimating numbers and biomass of blue catfish in waters throughout the state, especially as these numbers relate to variable year class strength and other metrics affecting recruitment to the fishery. The ultimate goal of these studies would be to set the stage for a more ambitious statewide study that focuses on yield modeling (Beverton and Holt 1957; Ricker 1975; Beckwith et al. 1986) to assess changes in catfish biomass associated with various real or hypothetical management changes (e.g., alterations of harvest size limits). Certainly studies such as these will generate a better understanding of blue catfish as an important renewable natural resource within Tennessee. 34 References Anderson, R. O., and R. M. Neumann. 1996. Length, weight, and associated structural indicies. Pages 447-482 in B. R. Murphy, and D. W. Willis, editors. Fisheries techniques, second edition. American Fisheries Society, Bethesda, Maryland. Arterburn, J. E., D. J. Kirby, and C. R. Berry, Jr. 2001. Biologist opinions and angler attitudes concerning catfish and their management in the Mississippi River basin. Mississippi Interstate Cooperative Resources Association, Final Report, Des Moines, Iowa. Beckwith, Jr., E. E., G. W. Benz, and R. P. Jacobs. 1986. Stock assessment of largemouth bass (Micropterus salmoides) in selected lakes with special reference to the 305 mm minimum size limit. Final Federal Report 1980-1986. Dingle-Johnson Project No. F-57-R. Connecticut Department of Environmental Protection, Hartford, Connecticut. Beverton, R. J. H., and S. J. Holt. 1957. On the dynamics of exploited fish populations. Chapman and Hall, London, U.K. Boxrucker, J., and K. Kuklinski. 2006. Abundance, growth, and mortality of selected Oklahoma blue catfish populations: implications for management of trophy fisheries. Proceedings of the Annual Conference of the Annual Associates of the Fish and Wildlife Agencies 60:152-156. 35 Brasher, B. 2007. Monster catfish highlights victory. Memphis commercial appeal. Available: http://www.commercialappeal.com/news/2007/nov/05/monster-catfishhighlights-victory/. Buckmeier, D. L., E. R. Irwin, R. K. Betsill, and J. A. Prentice. 2002. Validity of otoliths and pectoral spines for estimating ages of channel catfish. North American Journal of Fisheries Management 22:934-942. Caroll, B. B., and G. E. Hall. 1964. Growth of catfishes in Norris Reservoir, Tennessee. Journal of Tennessee Academy of Science 39:86-91. Conder, J. R., and R. Hoffarth. 1965. Growth of channel catfish, Ictalurus punctatus, and blue catfish, Ictalurus furcatus, in the Kentucky Lake portion of the Tennessee River in Tennessee. Proceedings of the Annual Conference of the Southeastern Association of Game and Fish Commissioners 16:348-354. Etnier, D. A., and W. C. Starnes. 1993. The fishes of Tennessee. University of Tennessee Press, Knoxville, Tennessee. Freeze, T. M. 1977. Age and growth, condition, and length-weight relationships of Ictalurus furcatus and Ictalurus punctatus from Lake Barkley and Kentucky Lakes, Kentucky. Master’s thesis. Murray State University, Murray, Kentucky. Graham, K. 1999. A review of the biology and management of blue catfish. Pages 3749 in E. R. Irwin, W. A. Hubert, C. F. Rabeni, H. L. Schramm, Jr., and T. Coon, editors. Catfish 2000: proceedings of the international ictalurid symposium. American Fisheries Society, Symposium 24, Bethesda, Maryland. 36 Hale, R. S., and T. J. Timmons. 1989. Comparative age and growth of blue catfish in the Kentuck portion of Kentucky Lake between 1967 and 1985. Transactions of the Kentucky Academy of Science 50:22-26. Hale, R. S., and T. J. Timmons. 1990. Growth of blue catfish in the lacustrine and riverine areas of the Tennessee portion of Kentucky Lake. Journal of the Tennessee Academy of Science 65:86-90. Hargis, H. L. 1966. Haul seine and trap net development and improvement based on related biological fishery research in Tennessee reservoirs. Job Completion Report 45-R-1, Tennessee Game and Fish Commision, Nashville, Tennessee. Hoffman, S. 2008. Top blue cat waters. In-Fisherman. Available: http://www.infisherman.com/magazine/exclusives/if2807_cats/. Holley, M. P. 2006. An evaluation of the catfish fishery in Wilson Reservoir, Alabama. Masters thesis. Auburn University, Auburn, Alabama. Irwin, E. R., W. A. Hubert, C. F. Rabeni, H. L. Schramm, Jr., and T. Coon, editors. 1999. Catfish 2000: proceedings of the international ictalurid symposium. American Fisheries Society, Symposium 24, Bethesda, Maryland. Isermann, D. A., J. R. Meerbeek, G. D. Scholten, and D. W. Willis. 2003. Evaluation of three different structures used for walleye age estimation with emphasis on removal and processing times. North American Journal of Fisheries Management 23:625-631. 37 Jones, B. D., and D. B. Noltie. 2007. Flooded flatheads: evidence of increased growth in Mississippi River Pylodictis olivaris (Pisces: Ictaluridae) following the great midwest flood of 1993. Hydrobiologia 592:183-209. Kelley, Jr., J. R., and D. C. Carver. 1966. Age and growth of blue catfish, Ictalurus furcatus (LeSuer 1840) in the recent delta of the Mississippi River. Proceedings of the Annual Conference Southeastern Association of Game and Fish Commissioners 19:296-299. Miranda, L. E. 1999. Recreational harvest in reservoirs in the USA. Fisheries Management and Ecology 6:499-513. Pope, K. L., and C. G. Kruse. 2007. Condition. Pages 423-471 in C. S. Guy, and M. L. Brown, editors. Analysis and interpretation of freshwater fisheries data. American Fisheries Society, Bethesda, Maryland. Porter, M. E. 1969. Comparative age, growth, and condition studies on the blue catfish, Ictalurus furcatus (LeSuer), and the channel catfish, Ictalurus punctatus (Rafinesque), within the Kentucky waters of Kentucky Lake. Master’s thesis. Murray State University, Murray, Kentucky. Ricker, W. E. 1975. Computation and interpretation of biological statistics of fish populations. Bulletin of the Fisheries Research Board of Canada 191:1-382. Sakaris, P. C., and E. R. Irwin. 2008. Validation of daily ring deposition in the otoliths of age-0 channel catfish. North American Journal of Fisheries Management 28:212218. 38 SAS Institute. 2008. User’s guide, version 9.1.3. SAS Institute, Cary, North Carolina. Schoffman, R. J. 1954. Age and growth of the channel catfish in Reelfoot Lake, Tennessee. Journal of the Tennessee Academy of Science 39:2-8. Slipke, J. W., and M. J. Maceina. 2003. Fishery analysis and simulation tools (FAST 3.0). Department of Fisheries and Allied Aquacultures, Auburn University, Auburn, Alabama. USFWS (Fish and Wildlife Service), and USCB (Census Bureau). 2006. National Survey of Fishing, Hunting, and Wildlife-Associated Recreation. U.S. Department of the Interior and U.S. Department of Commerce, Washington, D.C. Wege, G. J., and R. O. Anderson. 1978. Relative weight (Wr): a new index of condition for largemouth bass. Pages 79-91 in G. D. Novinger, and J. G. Dillard, editors. New approaches to the management of small impoundments. American Fisheries Society, North Central Division, Special Publication 5, Bethesda, Maryland. 39 Table 1. Sport fishery creel survey analysis resultsa for catfish (blue catfish, Ictalurus furcatus; channel catfish, I. punctatus; flathead catfish, Pylodictis olivaris) and bullheads (black bullhead, Amerius melas; yellow bullhead, A. natalis) caught in three Tennessee waterbodies during 1998-2006. Values in parentheses represent percent of corresponding grand total. Reservoir Species Mean total weight caught (kg) Mean total harvest (kg) Mean total harvest as percent of mean total weight caught Lake Barkley Angler hours fished for catfish and bullheads Number of catfish and bullheads caught hr-1 17,567 1.52 Mean total weight caught ha-1 (kg ha-1) 7,359 (6.3%) 6,071 82% 0.99 Channel catfish 2,411b (6.1%) 6,409b 266% 0.32 Flathead catfish 114 (2.2%) 98 86% 0.015 Kentucky Lake 399,254 1.05 Blue catfish 107,917 (92.9%) 99,224 92% 2.46 Channel catfish 36,999 (93.9%) 32,779 89% 0.84 Flathead catfish 5,179 (97.8%) 4,228 82% 0.19 Table 1. Continued on Subsequent Page Blue catfish 39 40 Table 1. Continued. Reservoir Species Mean total weight caught (kg) Mean total harvest (kg) Mean total harvest as percent of mean total weight caught Fort Loudoun Reservoirc Blue catfish 931 (0.8%) 0d Grand Total Angler hours fished for catfish and bullheads Number of catfish and bullheads caught hr-1 9,806 0.22 0% 0.99 426,627 Blue catfish 116,207 105,295 Channel catfish 39,410 39,188 Flathead catfish 5,293 4,326 Mean total weight caught ha-1 (kg ha-1) 0.32 0.015 2.46 a Data analyzed from Tennessee Wildlife Resources Agency creel surveys, 1998-2006. A reported harvest > the corresponding catch (both calculated values double checked for accuracy) indicates a reporting error in the TWRA creel survey. c Blue catfish were the only species of catfish reported, other catfishes are not generally considered abundant in this reservoir. d Lack of harvest corresponds to a period when the reservoir was under a fishery advisory warning regarding fish consumption. b 40 41 Table 2. Monthly 2005 commercial catfish harvest (kg) for 17 Tennessee waterbodies calculated from Tennessee Wildlife Resources Agency commercial fishing data.a 2005 Monthly Commercial Harvest (kg) Harvested group Jan. Feb. Mar. Apr. May Jun. Jul. Aug. Sept. Oct. Nov. Dec. Blue catfish 623,402 44,809 51,245 45,292 46,351 77,746 61,142 52,149 65,061 39,719 50,784 44,236 44,868 Channel catfish 220,805 15,890 18,390 22,698 21,958 22,029 16,153 16,205 19,571 12,698 17,489 21,703 16,021 Flathead catfish 29,322 652 819 1,575 2,100 4,065 3,351 2,907 3,499 1,899 5,331 2,041 1,083 Catfish spp.b 19,298 722 1,075 1,829 2,002 4,841 3,861 1,201 732 82 18 1,775 1,160 Totals 892,827 62,073 71,529 71,394 72,411 108,681 84,507 72,462 88,863 54,398 73,622 69,755 63,132 a Waterbodies: Lake Barkley, Kentucky Lake, Mississippi River, Nolichucky River, Obion River, Perryville River, Reelfoot Lake, Chickamauga Lake, French Broad River, Hatchie River, Lake Nickajack, Old Hickory Reservoir, Watts Barr Reservoir, Cheatham Lake, Lake Douglas, Lake Guntersville, Hiwassee River b Catfish spp. refers to a mixture of blue catfish, channel catfish, and flathead catfish not identified by species when reported. 41 42 Table 3. Sport and commercial harvest of catfish (blue catfish, Ictalurus furcatus; channel catfish, I. punctatus; and flathead catfish, Pylodictis olivaris) for 22 Tennessee waterbodies for the year 2006.a Sport harvest (kg) Commercial harvest (kg) Total harvest (kg) Sport harvest (% of total) Commercial harvest (% of total) Sport harvest ha-1 (kg) Commercial harvest ha-1 (kg) Harvest ha-1 (kg) Chickamauga Lake 13,962 47,991 194,780 242,771 19.8 80.2 3.44 13.95 17.4 Lake Barkleyb 7,406 7,147 92,243 99,390 7.2 92.8 0.97 12.46 13.4 Kentucky Lake b 43,794 91,727 311,586 403,313 22.7 77.3 2.09 7.11 9.2 Reelfoot Lake 4,220 1,028 25,348 26,376 3.9 96.1 0.24 6.01 6.3 Pickwick Lake 2,492 2,590 11,879 14,469 17.9 82.1 1.04 4.77 5.8 Watts Bar Reservoir 15,783 23,477 38,861 62,338 37.7 62.3 1.49 2.46 3.9 Old Hickory Lake 9,105 5,104 21,976 27,080 18.8 81.2 0.56 2.41 3.0 Lake Douglas 12,383 15,440 11,975 27,415 56.3 43.7 1.25 0.97 2.2 Percy Priest Reservoir 5,747 4,869 0 4,869 100 0 0.85 0 0.85 Lake Cherokee 12,262 4,914 0 4,914 100 0 0.40 0 0.4 Normandy Lake 1,233 441 0 441 100 0 0.36 0 0.36 Woods Reservoir 1,481 282 0 282 100 0 0.19 0 0.19 Boone Lake 1,829 289 0 289 100 0 0.16 0 0.16 Table 3. Continued on Subsequent Page Surface area (ha) Waterbody 42 43 Table 3. Continued. Surface area (ha) Sport harvest (kg) Commercial harvest (kg) Total harvest (kg) Sport harvest (% of total) Commercial harvest (% of total) Sport harvest ha-1 (kg) Commercial harvest ha-1 (kg) Harvest ha-1 (kg) Tims Ford Lake 4,290 312 0 312 100 0 0.07 0 0.07 Norris Lake 13,840 823 0 823 100 0 0.06 0 0.06 Melton Hill Lake 1,898 108 0 108 100 0 0.06 0 0.06 Dale Hallow Lake 7,069 525 0 525 100 0 0.07 0 0.07 Lake Watauga 2,602 82 0 82 100 0 0.03 0 0.03 Center Hill Lake 7,373 175 0 175 100 0 0.02 0 0.02 South Holston Reservoir 2,564 26 0 26 100 0 0.01 0 0.01 Fort Loudoun Reservoirc 5,908 27 0 27 100 0 0.005 0 0.005 Tellico Lake 6,498 0 0 0 100 0 0 0 0 207,377 708,648 916,025 22.6% 77.4% 0.61±0.18 2.28±0.88 2.93±1.03 Waterbody Total Mean (± SE) a Data primarily from unpublished records of the Tennessee Wildlife Resources Agency (see text for details). Heavily exploited reservoir included in study. c Lightly exploited reservoir included in study. b 43 44 Table 4. Number of blue catfish, Ictalurus furcatus, sampled from four study waterbodies. Number of Fish per Total Length Interval ≤ 225 ≥ 226 ≤ 250 ≥ 251 ≤ 275 ≥ 276 ≤ 300 ≥ 301 ≤ 325 ≥ 326 ≤ 350 ≥ 351 ≤ 375 ≥ 376 ≤ 400 ≥ 401 ≤ 425 ≥ 426 ≤ 450 ≥ 451 ≤ 475 ≥ 476 ≤ 500 Lake Barkley 169 0 0 1 0 1 2 4 6 6 5 12 12 Kentucky Lake 172 0 0 3 2 1 2 2 7 3 7 13 12 Mississippi River 248 43 5 6 5 2 3 16 15 21 16 17 19 Fort Loudoun Reservoir 190 0 0 3 3 1 4 4 5 0 10 14 6 Total 779 43 5 13 10 5 11 26 33 30 38 56 49 Table 4. Continued on Subsequent Page Waterbody Total number of fish sampled 44 45 Table 4. Continued. Number of Fish per Total Length Interval ≥ 501 ≤ 525 ≥ 526 ≤ 550 ≥ 551 ≤ 575 ≥ 576 ≤ 600 ≥ 601 ≤ 625 ≥ 626 ≤ 650 ≥ 651 ≤ 675 ≥ 676 ≤ 700 ≥ 701 ≤ 725 ≥ 726 ≤ 750 ≥ 751 ≤ 775 ≥ 776 ≤ 800 Lake Barkley 17 14 20 7 11 9 14 4 2 5 10 2 Kentucky Lake 11 8 17 13 14 9 6 5 7 7 5 0 Mississippi River 23 16 4 10 7 4 6 3 1 2 0 1 Fort Loudoun Reservoir 13 7 22 14 12 11 5 9 7 3 3 6 Totals 64 45 63 44 44 33 31 21 17 17 18 9 Table 4. Continued on Subsequent Page Waterbody 45 46 Table 4. Continued. Number of Fish per Total Length Interval Waterbody ≥ 801 ≤ 825 ≥ 826 ≤ 850 ≥ 851 ≤ 875 ≥ 876 ≤ 900 ≥ 901 ≤ 925 ≥ 926 ≤ 950 ≥ 951 ≤ 975 ≥ 976 ≤ 1,000 ≥ 1,001 Lake Barkley 0 0 0 1 1 0 0 0 3 Kentucky Lake 1 2 0 2 0 3 0 0 10 Mississippi River 2 1 0 0 0 0 0 0 0 Fort Loudoun Reservoir 2 8 6 4 1 1 1 2 3 Totals 5 11 6 7 2 4 1 2 16 46 47 Table 5. Parameters of log10 transformed weight-length linear regressions for four studied Tennessee waterbodies and log10 transformed weight-length linear regression for pooled data from the same four waterbodies used to represent Tennessee. Weight-Length Parameters Waterbody Slope ± 95% CI y-intercept ± 95% CI r2 Lake Barkley 2.87 ± 2.61-3.13 -4.60 ± -5.32--3.88 0.74 Kentucky Lake 3.46 ± 3.35-3.57 -6.20 ± -6.50--5.90 0.96 Mississippi River 3.10 ± 2.5-3.17 -5.33 ± -5.51--5.15 0.97 Fort Loudoun Reservoir 3.41 ± 3.23-3.59 -6.16± -6.45--5.87 0.96 Tennessee 3.21 ± 3.17-3.26 -5.59 ± -5.72--5.46 0.95 48 Table 6. Overall and mean relative weights for five size classes of blue catfish, Ictalurus furcatus, sampled from four Tennessee waterbodies.a Mean Relative Weights (Wr) Overall Wr Stock (≥ 300 to ≤ 509 mm TL) Quality (≥ 510 to ≤ 759 mm TL) Preferred (≥ 760 to ≤ 889 mm TL) Memorable (≥ 890 to ≤ 1,139 mm TL) 121 156.45 98.06 102.81 105.85 Kentucky Lake 112 106.18 114.53 121.17 103.62 Mississippi River 94 95.85 89.27 105.42 Fort Loudoun Reservoir 86 86.83 86.1 87.09 Waterbody Lake Barkley a 78.5 Lack of data prevented calculation of trophy size interval values. 48 49 Table 7. Mean total length at age (mm) for blue catfish, Ictalurus furcatus, in four Tennessee waterbodies. Age 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 32 33 34c Number of fish a Lake Barkleya Kentucky Lakea 318 405 479 559 547 575 631 681 709 693 787 271 535 525 550 560 595 670 704 740 1079 1092 1144 108 236 271 376 428 471 522 558 602 664 679 650 688 812 1115 1035 830 169 172 Heavily exploited reservoir included in study. Lightly exploited reservoir included in study. c Oldest blue catfish “aged” to date. b Mississippi Rivera 248 Fort Loudoun Reservoirb 329 281 317 382 445 446 441 472 474 508 527 579 592 602 636 625 718 710 729 667 759 826 812 839 837 900 874 190 50 Table 8. Fishery mileposts for blue catfish, Ictalurus furcatus, in four Tennessee waterbodies.a Note that values for some mileposts could not be estimated for some waters because of a lack of data. Waterbody Lake Barkley Number of years within fishery Age (yr) leaving fisheryc Weight (g) entering fishery Weight (g) at midpoint within fishery Weight (g) leaving fishery 1.5 11 12.5 158 1,778 6,761 1,659 7,244 Kentucky Lake 1 228 Mississippi River 1 138 Fort Loudoun Reservoir a Age (yr) entering fisheryb 3.5 30.5 34 58 Age estimations calculated using waterbody specific von Bertalanffy model parameters (see Figures 1013) as described by Slipke and Maceiana (2006). Weight estimations calculated using waterbody specific weight-length associations (see Figures 4-7). b Age entering fishery calculated as corresponding to a total length of 205 mm. c Age leaving fishery calculated as corresponding to a total length of 864 mm. 51 52 Table 9. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Lake Barkley (Tennessee and Kentucky). Length at Age (mm) (Growth Increment from Previous Age; mm) Waterbody (State) Study Aging Structurea 1 2 3 4 5 6 7 (TN) Present studyb O 190 290 (100) 372 (82) 450 (78) 519 (69) 581 (62) 637 (56) (KY) Freeze (1977)c P 76 188 (112) 302 (114) 376 (74) 455 (79) 584 (129) 658 (74) Lake Barkley a O = otoliths, P = pectoral spines. Length at age and growth increment values as predicted by the waterbody specific von Bertalanffy growth model (see Figure 10). c Length at age and incremental growth values derived from reported mean length at age values. b 51 53 Table 10. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Kentucky Lake (Tennessee and Kentucky). Length at Age (mm) (Growth Increment from Previous Age; mm) Waterbody (State) Study Aging Structurea 1 2 3 4 5 6 7 8 9 10 11 (TN) Present Studyb O 229 313 (84) 388 (75) 453 (65) 511 (58) 561 (50) 606 (45) 646 (40) 681 (35) 711 (30) 873 (27) (TN) Hale and Timmons (1989)c P 132 221 (89) 274 (53) 318 (44) 363 (45) 424 (61) 485 (61) 549 (64) 584 (35) 607 (23) 693 (86) (TN) Hale and Timmons (1990)c P 142 229 (87) 287 (58) 343 (56) 401 (58) 447 (46) 500 (53) 423 (-77) 551 (128) 587 (36) (TN) Hale and Timmons (1990)c P 145 239 (94) 295 (56) 356 (61) 427 (71) 483 (56) 551 (68) 627 (76) 671 (44) (KY) Conder and Hoffarth (1965)c P 135 198 (63) 252 (54) 297 (45) 356 (59) 429 (73) 513 (84) 582 (69) 699 (117) Kentucky Lake 846 (147) 52 54 Table 10. Continued. Length at Age (mm) (Growth Increment from Previous Age; mm) Waterbody (State) Study Aging Structurea 1 2 3 4 5 6 7 8 9 10 11 (TN) Present Studyb O 229 313 (84) 388 (75) 453 (65) 511 (58) 561 (50) 606 (45) 646 (40) 681 (35) 711 (30) 873 (27) (KY) Porter (1969)c P 117 213 (96) 310 (97) 391 (81) 480 (89) 559 (79) 627 (68) (KY) Freeze (1977)c P 76 165 (89) 239 (74) 302 (63) 311 (9) 432 (121) 483 (51) 564 (81) 666 (102) Kentucky Lake a O = otoliths, P = pectoral spines. Length at age and growth increment values as predicted by the waterbody specific von Bertalanffy growth model (see Figure 11). c Length at age and incremental growth values derived from reported mean length at age values. b 53 55 Table 11. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Mississippi River (Tennessee and Louisiana). Length at Age (mm) (Growth Increment from Previous Age; mm) Waterbody (State) Study Aging Structurea 1 2 3 4 5 6 (TN) Present Studyb O 211 294 (83) 367 (73) 429 (62) 483 (54) 530 (47) (LA) Kelley and Carver (1966)c P 191 386 (195) 508 (122) 638 (130) 749 (111) 848 (99) Mississippi River a O = otoliths, P = pectoral spines. Length at age and growth increment values as predicted by the waterbody specific von Bertalanffy growth model (see Figure 12). c Length at age and incremental growth values derived from reported mean length at age values. b 54 56 Table 12. Comparison of blue catfish, Ictalurus furcatus, length age data stemming from studies in Fort Loudoun Reservoir (Tennessee), Lake Hugo (Oklahoma), Lake Ellsworth (Oklahoma), and Lake Eufaula (Oklahoma). Length at Age (mm) (Growth Increment from Previous Age; mm) Waterbody (State) Study Aging Structurea 1 2 3 4 5 6 7 8 9 10 11 12 13 Fort Loudoun Reservoir (TN) Present Studyb O 103 146 188 227 265 301 336 369 400 431 460 488 514 (43) (42) (39) (38) (36) (35) (33) (31) (31) (29) (28) (26) Lake Hugo (OK) Boxrucker and Kuklinski (2006)c 223 273 320 331 371 412 450 474 487 (55) (50) (47) (11) (40) (41) (38) (24) (13) Lake Ellsworth (OK) Boxrucker and Kuklinski (2006)c 186 222 234 253 274 298 321 339 384 394 414 382 (20) (36) (12) (19) (21) (24) (23) (18) (45) (10) (20) (-32) Lake Eufaula (OK) Boxrucker and Kuklinski (2006)c 203 256 295 273 351 357 375 415 427 473 479 491 (47) (53) (39) (-22) (78) (6) (18) (40) (12) (46) (6) (12) P P P 168 166 156 a O = otoliths, P = pectoral spines. Length at age and growth increment values as predicted by the waterbody specific von Bertalanffy growth model (see Figure 13). c Length at age and incremental growth values derived from reported mean length at age values. b 55 57 1 3 2 4 Figure 1. Major river systems of Tennessee. Blue catfish (Ictalurus furcatus) samples were collected from the following four waterbodies as part of this study: Lake Barkley (1), Kentucky Lake (2), Mississippi River (3), and Fort Loudoun Reservoir (4). 56 58 B A C D Figure 2. Location and removal of sattigal otoliths from catfish. A. Otoliths are located within a plane (dashed line) lying just anterior to (ca. 3-5 mm) a transverse plane (solid line) bridging the proximal ends of the locked pectoral spines. B. Sawblade lying along plane of section used to expose otoliths. C. Catfish head being sectioned to expose otoliths. D. Right sagittal otolith being removed with a forceps. Figure 2A and 2B modified from Buckmeier et al. (2002). 59 r ar ss Figure 3. Epoxy casts containing catfish otoliths. For aging purposes, the sagittal otolith was transversely sectioned just anterior to the antirostrum (ar) on the rostrum side (r) using a low-speed isomet saw. Sand paper (200-500 grit) was then used to grind the otolith to the ventral sulcus (s). 60 log10 W = -4.60 + 2.87log10TL r2 = 0.74 n = 168 Figure 4. Weight-length relationship for blue catfish, Ictalurus furcatus, collected from Lake Barkley (Tennessee portion of lake; formula for log10 transformed weightlength linear regression and r2 value provided at upper left). 61 log10 W = -6.21 + 3.46log10TL r2 = 0.96 n = 166 Figure 5. Weight-length relationship for blue catfish, Ictalurus furcatus, collected from Kentucky Lake (Tennessee portion of lake; formula for log10 transformed weight-length linear regression and r2 value provided at upper left). 62 log10 W = -5.33 + 3.10log10TL r2 = 0.97 n = 205 Figure 6. Weight-length relationship for blue catfish, Ictalurus furcatus, collected from the Mississippi River (formula for log10 transformed weight-length linear regression and r2 value provided at upper left). 63 log10 W = -6.16 + 3.41log10TL r2 = 0.96 n = 190 Figure 7. Weight-length relationship for blue catfish, Ictalurus furcatus, collected from Fort Loudoun Reservoir (formula for log10 transformed weight-length linear regression and r2 provided value at upper left). 64 log10 W = -5.59 + 3.21log10TL r2 = 0.95 n = 773 Figure 8. Weight-length relationship for blue catfish, Ictalurus furcatus, collected from four Tennessee waterbodies (Lake Barkley, Kentucky Lake, Mississippi River, Fort Loudoun Reservoir; formula for log10 transformed weight-length linear regression and r2 value provided at upper left). 65 864 mm TL (Trophy size limit) A C D B A) Lake Barkley B) Kentucky Lake C) Mississippi River D) Fort Loudoun Reservoir Figure 9. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curves for four Tennessee waterbodies. 66 1,115 mm TL (L∞) 864 mm TL (TN trophy size limit) Lt = 1,115 {1 – e -0.11(t +0.693)} r2 = 0.83 n = 169 Figure 10. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Lake Barkley. 67 940 mm TL (L∞) 864 mm TL (TN trophy size limit) Lt = 940 {1 – e -0.126(t +1.217)} r2 = 0.92 n = 155 Figure 11. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Kentucky Lake. 68 864 mm TL (TN trophy size limit) 830 mm TL (L∞) Lt = 830 {1 – e -0.145(t +1.019)} r2 = 0.97 n = 207 Figure 12. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Mississippi River. 69 1,105 mm (L∞) 864 mm TL (TN trophy size limit) Lt = 1,105 {1 – e -0.044(t +1.227)} r2 = 0.96 n = 190 Figure 13. Blue catfish, Ictalurus furcatus, von Bertalanffy growth curve and model parameters for Fort Loudoun Reservoir.