Western North American Naturalist

Volume 72 Number 1 Article 8

4-5-2012

Age and growth of spawning longnose ( osseus) in a north central Texas reservoir

Samuel W. Kelley U.S. Geological Survey, Wichita Falls, TX, [email protected]

Follow this and additional works at: https://scholarsarchive.byu.edu/wnan

Part of the Anatomy Commons, Botany Commons, Physiology Commons, and the Zoology Commons

Recommended Citation Kelley, Samuel W. (2012) "Age and growth of spawning longnose gar (Lepisosteus osseus) in a north central Texas reservoir," Western North American Naturalist: Vol. 72 : No. 1 , Article 8. Available at: https://scholarsarchive.byu.edu/wnan/vol72/iss1/8

This Article is brought to you for free and open access by the Western North American Naturalist Publications at BYU ScholarsArchive. It has been accepted for inclusion in Western North American Naturalist by an authorized editor of BYU ScholarsArchive. For more information, please contact [email protected], [email protected]. Western North American Naturalist 72(1), © 2012, pp. 69–77

AGE AND GROWTH OF SPAWNING LONGNOSE GAR (LEPISOSTEUS OSSEUS) IN A NORTH CENTRAL TEXAS RESERVOIR

Samuel W. Kelley1

ABSTRACT.—The longnose gar (Lepisosteus osseus) is a primitive predaceous fish common throughout much of the east central United States, but research on its age and growth in lacustrine systems is rare. To characterize gar age and growth, I used bowfishing to collect spawning longnose gar in spring 2010 from littoral zones at Lake Arrowhead, Clay County, Texas. Females were older than males but significantly exceeded males in total length and mass when age was controlled. Von Bertalanffy growth curves suggested that males had faster growth rates, smaller maximum lengths, and shorter life spans than did females. However, females were always longer than males at any given age. Bowfishing capture beyond dis- tances of 9 m was biased toward larger fish, but the method was viable for collecting spawning longnose gar at close range. This study will assist fisheries managers and aquaculturists by providing growth–age relationships for longnose gar in a southern lacustrine system.

RESUMEN.—El pejelagarto narigudo (Lepisosteus osseus) es un pez depredador primitivo que es común a lo largo de una gran parte del centro-este de los Estados Unidos; sin embargo, la investigación sobre su edad y crecimiento en sistemas lacustres es rara. En la primavera de 2010, utilicé pesca con arco y flecha para colectar pejelagartos narigudos en estado de desove de las zonas litorales del Lago Arrowhead, condado de Clay, Texas para caracterizar sus edades y crecimiento. Las hembras fueron mayores en edad que los machos, pero al corregir por la edad, éstas superaban considerablemente a los machos en el largo y la masa total. Las curvas de crecimiento de Von Bertalanffy indicaron que los machos tuvieron tasas de crecimiento más aceleradas, una longitud máxima menor y un promedio de vida más corto que las hembras. Sin embargo, las hembras a cualquier edad siempre superaron en longitud a los machos. La captura con arco y flecha de más allá de 9 metros se orientó hacia los peces más grandes, pero el método demostró ser viable para la captura del pejelagarto narigudo en estado de desove a distancias cercanas. Este estudio ayudará a los encargados de pesquerías y acuicultura porque describe las relaciones entre el crecimiento y la edad del pejelagarto narigudo en un sistema lacustre del sur.

The longnose gar (Lepisosteus osseus) is a Morgan 1974), however, are lacking, particularly primitive predaceous fish that is abundant in the southern portion of the gar’s range. While throughout much of the east central United electrofishing is widely used to capture many States. It may exceed lengths of 1.8 m, it ap- fish species, including longnose gar (Sutton et proaches 23 kg in weight, and, among gar spe- al. 2009), it has produced poor results for lepi - cies in the United States, it is surpassed in size sosteids from Lake Arrowhead (Howell and only by the alligator gar ( spatula). Mauk 2004, 2008). The tendency of stunned Unlike the alligator gar, the longnose gar cur- gar to sink rather than float up for visible dip rently has a secure status; however, effective net captures (Burr 1931) may bias catch rates. management and conservation strategies require High conductivity, turbidity, and depths >1.2 m more biological data from lacustrine and riverine (frequent characteristics in southern reservoirs) systems across its range. Furthermore, lepiso - also reduce electrofishing efficiency. Addition- steids have been recognized as important envi- ally, most previous studies used passive capture ronmental indicators and biomarker species techniques (e.g., gill nets or trap boxes; Klaassen (Hartley et al. 1996, Huang et al. 1997, Huggett and Morgan 1974, Johnson and Noltie 1996, et al. 2001, Watanabe et al. 2003, Burger et Robertson et al. 2008) or a combination of al. 2004). passive and active capture techniques (e.g., Previous studies have provided insight on the gill nets, dip nets, trotlines, seining, and elec- growth and age of longnose gar in riverine sys- trofishing; Netsch and Witt 1962, Tyler et al. tems in the central United States. Compara- 1994, Johnson and Noltie 1997) to obtain tive data from lacustrine systems (Klaassen and specimens.

1U.S. Geological Survey, 3010 Buchanan St., Wichita Falls, TX 76308. E-mail: [email protected]

69 70 WESTERN NORTH AMERICAN NATURALIST [Volume 72

Fig. 1. Lake Arrowhead study site, Clay County, Texas. Black squares near the East Little Post Oak Creek and Deer Creek bridges indicate collection sites.

Because spawning longnose gar often con- The lake has a surface area of 6036 ha, a maxi - gregate near the surface in littoral shallows, I mum depth of 14 m near the dam, and high tur- predicted that bowfishing would be a viable bidity. The lake elevation fluctuates from about collection method. Additionally, I desired to 1.2 to 1.8 m due to extensive municipal water characterize age and growth of longnose gar use by the city of Wichita Falls and neighboring from a southern reservoir system to provide a communities. comparison to previous studies from the central I collected longnose gar using archery (bow- United States. I hypothesized that female long- fishing) equipment and methods compliant nose gar would be generally larger and older with regulations of the Texas Parks and Wildlife than male longnose gar, as illustrated by previ - Department. Specifically, I took all available ous studies. Moreover, I expected that both shots at gar in range (distances ≤14 m), while I male and female longnose gar from a southern walked along the shoreline at 2 sites in the lower reservoir would grow faster than riverine con- region of the reservoir (near Deer Creek bridge geners farther north due to the longer growing and near East Little Post Oak Creek bridge; season and higher productivity of lacustrine Fig. 1). The collected specimens were assumed systems (Johnson and Noltie 1997). to be a random sample of the adult spawning population. METHODS Collected gar specimens were placed on ice and examined within 24 hours of capture. Total Longnose gar were collected 23–26 April length (TL, mm) and mass (g) were recorded and 29 May 2010 from Lake Arrowhead, a for each specimen. Sex was determined fol- reservoir impounded in 1966 by damming the lowing the methods of Ferrara and Irwin (2001), Little Wichita River in Clay County, Texas. al though sex was readily apparent from the 2012] AGE AND GROWTH OF SPAWNING LONGNOSE GAR 71 presence of eggs or milt during internal exam- normalize female mass. The length-to-mass ination. Age was determined by counting annuli relationship of both sexes combined was deter - on the largest left branchiostegal ray of each fish mined by linear regression using log10-trans- (Netsch and Witt 1962). The rays were removed, formed age and mass values. boiled briefly in water (Johnson and Noltie 1997), Initial results demonstrated that age, mass, cleaned in a dilute bleach solution (Klaassen TL at capture, and BCTLs of female gar were and Morgan 1974), and cleared in mineral oil. significantly greater than those for males. How - Annuli bands that extended completely across ever, age was suspected to exert a substantial the ray were counted using a dissecting micro- influence on these metrics; thus, statistical com - scope (Johnson and Noltie 1997, Love 2004). parisons were made between sexes using a Because of false annuli bands, a few gar speci- general linear model ANCOVA (Johnson and men ages were tentative. In those cases, a sec- Noltie 1997, Love 2004) with age and log10 age ond observer also counted annuli, and counts as covariates, TL and log10 BCTL as response were discussed until an agreement on age was variables, and sex as a factor variable. Before reached. A Wild Heerbrugg microscope with conducting this test, sex-by-age interactions an ocular micrometer was used to obtain all and assumptions of parallel slopes were tested branchiostegal ray measurements, including with preliminary analyses using multiple regres- those be tween each annulus for back-calculation sion and ANOVA (dependent variables [Y] = TL of TL by age. and log10 BCTL; numerical independent vari- Back-calculated TL (BCTL) was determined ables [X] = age and log10 age; categorical inde- following the methods and equations described pendent variable [X] = sex) to confirm that by Johnson and Noltie (1997 and references differences between sexes among mean TLs therein) and later slightly modified by Love and BCTLs were not dependent on age alone. (2004): Linear methods in fish growth calculations have fallen out of favor with many researchers; log TLb = a + (log TLc – a) (log BAa /log BLt), however, only Sutton et al. (2009) have pub- lished a growth curve for longnose gar using where TLb is the back-calculated TL at age b, nonlinear methods, although Ferrara (2001) and TLc is the TL of the fish at capture, BAa is the McGrath (2010) made noteworthy contributions branchiostegal ray length from the base to the using nonlinear methods as well. Thus, for com - median point on the annulus, and BLt is the total parative purposes, I derived growth estimate branchiostegal ray length from base to tip. The curves (by sex) from plotted BCTLs by age using constant (a) used in the above equation was ob- the von Bertalanffy (1938) growth model: tained from the regression (all specimens) of TL at capture on the total length of the bran- –K(t – t ) lt = L∞ (1 – e 0 ) , chiostegal ray (BL): where lt is length at age t, L∞ is the asymp- log TL = a + b log BL. totic length, K is the growth-rate parameter, and t0 is a theoretical age (usually a negative BCTLs yielded far greater numbers of age number) at which the total length is presumed estimates than did lengths at age of capture equal to zero. alone, providing total length estimates from All statistical tests were carried out using each year of life for every captured fish. Mean NCSS 2007 (NCSS, J. Hintze, Kaysville, UT). BCTLs were plotted by year for both sexes Regression and von Bertalanffy (VB) plots were from age one to the oldest age of capture. Log- created using the regression wizard and dynamic transformed BCTLs were regressed separately fit wizard in SigmaPlot® 10.0 (Systat, San Jose, by sex on log-transformed age for comparison CA). Results were considered statistically sig- to previous studies. nificant at P < 0.05. Age, mass, and TL at capture of male and female gar were compared using 2-tailed equal- RESULTS variance t tests, following Shapiro–Wilk normal- ity tests and modified Levene tests to check for Twenty-four longnose gar (13ɉɉ, 11 ɊɊ) equal variance. Male and female mass were were collected via bowfishing (Fig. 1). Collec- log10-transformed prior to testing in order to tion efforts were hampered by highly turbid 72 WESTERN NORTH AMERICAN NATURALIST [Volume 72 waters from recent rainfall, short-lived spawn- 0.896; Fig. 2B), thus supporting use of the slope ing activities, and poor surface visibility due to value as a constant for deriving BCTLs. high winds. Visual observations suggested a Back-Calculated TL vs. Age male bias in the sex ratio of spawning long- nose gar, and it was not unusual to observe The assumption that the slopes of the BCTL- 2–6 males corralling a single large female to - versus-age relationship were equal between wards shoreline rocks in competitive efforts to males and females was not rejected (F = 1.938, fertilize her eggs. Additionally, females rarely df = 1, P = 0.165), and influence of age (covari- released all of their eggs at once but rather in ate) on BCTL was significant (F = 1180.4, df = 1, multiple locations near the shoreline, often P < 0.0001), validating the ANCOVA; however, allowing additional males opportunities for mean BCTLs between sexes differed signifi- fertilization. cantly when controlled for age (F = 567.46, Bowfishing proved to be most effective at df = 1, P < 0.0001), with females exhibiting shot distances of about 9 m or less. Distances significantly longer BCTLs than males (Fig. 3). >9 m were biased toward successful captures of larger specimens and against captures of Growth smaller specimens, as only a few females (i.e., Log-transformed regressions of growth pro- larger targets) were hit beyond 9 m. In this vided good fits for male and female longnose study, most shots were taken within 9 m of the gar from Lake Arrowhead (Fig. 3A). The VB collector because of the longnose ’ affinity growth curves were also a good fit for longnose for spawning along shoreline rocks. gar in this study; although, as in the regression analyses, they explained more of the variation Sexual Dimorphism in Age, Mass, and TL for males than for females. Mean male TLs did Mean age (+–SD) of captured female long- not exceed those of females at same-age compari- nose gar, as determined by branchiostegal ray sons. In addition, VB growth parameter values annuli counts, significantly exceeded that of were larger for males (K = 0.35) than for fe- captured males (ɊɊ = 16.2 +– 4.1 years, ɉɉ = males (K = 0.18), although females had greater + 9 – 2.9 years; t = 5.03, df = 22, P < 0.0001). asymptotic lengths (L∞ = 1307) than males Females also had significantly greater mean (L∞ = 923; Fig. 3B). (+–SD) mass (ɊɊ = 6878 +– 2760 g, ɉɉ = 2066 +– 584 g; t = 9.20, df = 22, P < 0.0001) DISCUSSION and TL (ɊɊ = 1253 +– 115 mm, ɉɉ = 887 +– 68 mm; t = 9.65, df = 22, P < 0.0001) than Bowfishing methods have seen little use in males; however, the influence of age on these scientific study but may provide insights into results was suspect, requiring further ANCOVA fish populations that are not easy to obtain analyses. Age was a significant covariate on using traditional methods, especially for large mass (F = 29.12, df = 1, P < 0.0001); never- fish in older age classes. Quinn (2010) lists theless, females significantly outweighed males 89%–100% of bowfishing captures in Arkansas when the model was controlled for age (F = as exceeding published size-at-maturity esti- 33.61, df = 1, P < 0.0001). The assumption mates for multiple species, including longnose that the slopes of TL at capture to age were gar, and this study supports his findings. Bow- equal between males and females was not fishing success depends largely upon location, rejected (F = 0.085, df = 1, P = 0.774), and season, and weather conditions (Quinn 2010), influence of age (covariate) on TL was signifi - as well as skill of the collector, but this is true cant (F = 31.02, df = 1, P < 0.0001); however, for all collection methods. captured females had significantly greater mean Longnose gar in this study were most simi- TLs than captured males when controlled for lar in age to those described by Netsch and age (F = 36.70, df = 1, P < 0.0001). Witt (1962) and were notably older than lacus- Despite dimorphism in mass, combined male trine gar from Kansas (Klaassen and Morgan and female TLs proved a good predictor of mass 1974). Mean BCTLs for male gar from Lake for both sexes, explaining the majority of varia- Arrowhead were slightly larger than those tion (r2 = 0.983) in a linear relationship (Fig. 2A). reported in previous studies for ages 2–10 but Additionally, branchiostegal ray length was a were similar to, or smaller than, mean BCTLs good predictor of TL for longnose gar (r2 = for riverine gar from Missouri at ages 1 and 2012] AGE AND GROWTH OF SPAWNING LONGNOSE GAR 73

Fig. 2. Attribute relationships for combined adult male (filled triangles; n = 13) and female (open circles; n = 11) spawning longnose gar (Lepisosteus osseus) collected from Lake Arrowhead, Clay County, Texas, April–May 2010: A, mass (M) versus total length (TL); B, total length (TL) versus branchiostegal ray length (BL).

11–14 (Fig. 4A). Mean BCTLs for female long- Growth of young (<1-year-old) longnose gar nose gar from Lake Arrowhead exceeded those is very rapid and does not conform well to the of riverine gar from Missouri (Netsch and Witt VB growth curve, as evidenced by poorer fit 1962, Johnson and Noltie 1997) and reservoir results with trial inputs provided by Echelle gar from Kansas (Klaassen and Morgan 1974) and Riggs (1972) of TLs equaling 10 mm at at all ages, though some overlap was apparent hatch and 20 mm at 10 days. The VB growth (Fig. 4B). curve for Lake Arrowhead female longnose gar 74 WESTERN NORTH AMERICAN NATURALIST [Volume 72

Log10 (Age, years) Mean Back-Calculated Total Length (mm) Total Mean Back-Calculated

Age (years)

Fig. 3. Length versus age relationships based on branchiostegal ray annuli counts (n = 117) for male (filled triangles; n = 13) and counts (n =178) for female (open circles; n = 11) spawning longnose gar (Lepisosteus osseus) collected from Lake Arrowhead, Clay County, Texas, April–May 2010: A, log10-transformed back-calculated total length versus log10- transformed age; B, mean (+–SD) back-calculated total length versus age plots plus von Bertalanffy growth curves (dotted lines = 95% confidence intervals). Data points lacking +– SD error bars indicate singular specimens at those age classes. appeared to slightly overestimate TL at age 1, TLs and diverge from the standard asymptotic and underestimate TL at ages >19 years (Fig. VB growth curve. Larger gar tend to prey upon 3B). Although increased sample size might larger forage species (Tyler et al. 1994) and may ameliorate some of these disparities, results exhibit a bimodal growth pattern as a result of from this study and from Netsch and Witt (1962) diet shifts to larger prey. Overall, VB growth suggest that large, old female gar have variable parameter values suggested that males in Lake 2012] AGE AND GROWTH OF SPAWNING LONGNOSE GAR 75

Age (years)

Age (years)

Fig. 4. Mean (+– SD) back-calculated total length by age comparison plots based on branchiostegal ray annuli counts for spawning longnose gar (Lepisosteus osseus) from Lake Arrowhead (filled circles, solid lines), Clay County, Texas, April–May 2010: A, males (n = 13), annuli counts (n = 117); B, females (n = 11), annuli counts (n = 178). Data points from Lake Arrowhead lacking +– SD error bars indicate singular specimens at those age classes. Comparative means are derived from Netsch and Witt (1962; open triangles, dash-dot-dot line), Klaassen and Morgan (1974; filled squares, long-dash line), and Johnson and Noltie (1997; open circles, dotted line; inverted filled triangles, short-dash line). Means from Johnson and Noltie (1997) are derived from graphical estimates with an estimated projection error of +– 10 mm.

Arrowhead grew faster and had smaller maxi- differences are due to disparate latitudes, habitat mum lengths and shorter life spans than did (riverine vs. lacustrine system), sex ratios, or females (Fig. 3B). methodology is unclear. Longnose gar are sexu- Comparisons of VB growth curves for long- ally dimorphic (Netsch and Witt 1962, Tyler et nose gar in this study to those of Sutton et al. al. 1994, Johnson and Noltie 1997, McGrath (2009) reveal dissimilarities, but whether those 2010), and previous studies have also indicated 76 WESTERN NORTH AMERICAN NATURALIST [Volume 72 a male-biased sex ratio in longnose gar (Tyler et gorge on forage fish that congregate in the al. 1994, Johnson and Noltie 1996, 1997, Ferrara highly oxygenated waters. If prey abundance 2001, McGrath 2010; but see Klaassen and Mor - is a primary driving factor of growth potential, gan 1974); however, Sutton et al. (2009) did not then riverine gar proximal to dam structures sex captured longnose gar but generated a single may have the capacity to match growth of reser- growth curve for both sexes combined using a voir gar, as illustrated by the longstanding (1954) VB model of mean TL to age at capture. Their world record longnose gar (22.82 kg, 1842 mm) combined sex growth curve for riverine long- from the Trinity River in Houston County, Texas nose gar from Indiana and Illinois yield pa- (Meyer 2008). rameter estimates (L∞ = 1009, K = 0.21, t0 = This study contributes to the biological –1.62) that lie between those of males and knowledge of the longnose gar and may assist females in this study (Fig. 3B). conservationists and aquaculturists by provid- Comparisons of VB growth curves for long- ing pertinent growth and age data from a nose gar in this study are similar to those of southern lacustrine system. Whereas fisheries McGrath (2010) (females: L∞ = 1132, K = 0.18, managers once embraced eradication of gar t0 = –1.55; males: L∞ = 875, K = 0.23, t0 = species from public and private waters, future –2.11). Like males in McGrath (2010), males in initiatives may reverse these policies and per- this study had greater growth-rate parameter haps support stocking efforts of longnose gar values than females, suggesting more rapid as an apex predator in select lacustrine waters. male growth. The male growth-rate parameter There is also a growing body of research on value from this study exceeded that of male the use of lepisosteids, including longnose gar, riverine gar in Virginia, yet female growth as aquaculture candidates for food and fisheries parameters were equal. However, asymptotic restocking (e.g., Alfaro et al. 2008, Jaroszewska lengths of male and female longnose gar from et al. 2010). Although some results of this Virginia were smaller than those presented study corroborate previous research, much of herein (Fig. 3B). the data herein are unique. Specifically, this Ferrara (2001) reported a VB growth curve report provides the first published record of for riverine female gar in Alabama with L∞ (1) a scientific collection of longnose gar ac - (1306) and K (0.17) values similar to those I quired solely using bowfishing methods, (2) obtained (Fig. 3B). These similarities may be due sexually dimorphic VB growth curves for spawn- to similar growing season lengths and higher ing longnose gar based on BCTLs, and (3) log- mean temperatures. These conditions likely al - transformed regression and VB growth curves low for longer, more efficient foraging periods. for longnose gar from any reservoir system. Moreover, Ferrara (2001) collected gar immedi- ately below a dam in a tailrace that supported an ACKNOWLEDGMENTS abundance of shad ( spp.), which may have contributed to greater growth potential. I thank Allyse Ferrara for freely providing Lake Arrowhead has an abundance of gizzard both raw data and results from her Ph.D. shad (Dorosoma cepedianum), with Howell and research regarding longnose gar in Alabama Mauk (2008) reporting catch rates of 576 per for comparison to this study. Gary Burke and hour. Moreover, collected gizzard shad displayed an anony mous reviewer provided helpful com- a high index (88.5%) of vulnerability (DiCenzo ments on earlier drafts of the manuscript. et al. 1996) based on catch sizes ≤203 mm. Klaassen and Morgan (1974) suggested that long- LITERATURE CITED nose gar may grow faster and larger in reservoirs than in riverine systems because of increased ALFARO, R.M., C.A. GONZÁLEZ, AND A.M. FERRARA. 2008. Gar biology and culture: status and prospects. Aqua- abundance of gizzard shad prey. Results of this culture Research 39:748–763. study seem to support this idea, and prey abun- BURGER, J., E.F. ORLANDO, M. GOCHFELD, G.A. BINCZIK, dance may be a factor contributing to the large AND L.J. GUILLETTE JR. 2004. Metal levels in tissues sizes reported herein. Such generalizations may of Florida gar (Lepisosteus platyrhincus) from Lake be skewed, however, because personal obser- Okeechobee. Environmental Monitoring and Assess- ment 90:187–201. vations and previous research indicate that river- BURR, J.G. 1931. Electricity as a means of garfish and carp ine gar migrate upstream (Johnson and Noltie control. Transactions of the American Fisheries Society 1996), often congregating in dam tailraces to 61:174–182. 2012] AGE AND GROWTH OF SPAWNING LONGNOSE GAR 77

DICENZO, V.J., M.J. MACEINA, AND M.R. STIMERT. 1996. KLAASSEN, H.E., AND K.L. MORGAN. 1974. Age and growth Relations between reservoir trophic state and gizzard of longnose gar in Tuttle Creek Reservoir, Kansas. shad population characteristics in Alabama reservoirs. Transactions of the American Fisheries Society 103: North American Journal of Fisheries Management 402–405. 16:888–895. LOVE, J.W. 2004. Age, growth, and reproduction of spotted ECHELLE, A.A., AND C.D. RIGGS. 1972. Aspects of the gar (Lepisosteus oculatus), from the Lake Pontchar- early life history of gars (Lepisosteus) in Lake Texoma. train estuary, Louisiana. Southwestern Naturalist 49: Transactions of the American Fisheries Society 101: 18–23. 106–112. MCGRATH, P.E. 2010. The life history of longnose gar, FERRARA, A.M. 2001. Life-history strategy of Lepisostei- Lepisosteus osseus, an apex predator in the tidal waters dae: implications for the conservation and management of Virginia. Doctoral dissertation, College of William of alligator gar. Doctoral dissertation, Auburn Univer- and Mary, Williamsburg, VA. sity, Auburn, AL. MEYER, W.P. 2008. Gar hunter, writer, stockbroker—Town - FERRARA, A.M., AND E.R. IRWIN. 2001. A standardized pro- send Miller’s 1954 longnose gar record still stands cedure for internal sex identification in Lepisosteidae. today [online]. [Cited 9 February 2011]. Texas Parks and North American Journal of Fisheries Management Wildlife Magazine: November. Available from: http:// 21:956–961. www.tpwmagazine.com/archive/2008/nov/legend/. HARTLEY, W.R., A. THIYAGARAJAH, AND A.M. TREINIES. NETSCH, N.F., AND A. WITT JR. 1962. Contributions to the 1996. Liver lesions in the gar fish (Lepisosteidae) as life history of the longnose gar, (Lepisosteus osseus) in biomarkers of exposure. Marine Environmental Re- Missouri. Transactions of the American Fisheries search 42:217–221. Society 91:251–262. HOWELL, M., AND R. MAUK. 2004. Statewide freshwater QUINN, J.W. 2010. A survey of bowfishing tournaments in fisheries monitoring and management program survey Arkansas. North American Journal of Fisheries Man- report for Arrowhead Reservoir, 2003. Texas Parks agement 30:1376–1384. and Wildlife Department, Federal Aid Report F-30- ROBERTSON, C.R., S.C. ZEUG, AND K.O. WINEMILLER. 2008. R-29, Austin, TX. Associations between hydrological connectivity and ______. 2008. Statewide freshwater fisheries monitoring resource partitioning among sympatric gar species and management program survey report for Arrow- (Lepisosteidae) in a Texas river and associated oxbows. head Reservoir, 2007. Texas Parks and Wildlife Depart - Ecology of Freshwater Fish 17:119–129. ment, Federal Aid Report F-30-R-33, Austin, TX. SUTTON, T.M., A.C. GRIER, AND L.D. FRANKLAND. 2009. HUANG, T.L., P.O. OBIH, R. JAISWAL, W.R. HARTLEY, AND Stock structure and dynamics of longnose gar and A. THIYAGARAJAH. 1997. Evaluation of liver and brain in the Wabash River, Indiana–Illinois. esterases in the spotted gar fish (Lepisosteus oculatus) Journal of Freshwater Ecology 24:657–666. as biomarkers of effect in the lower Mississippi TYLER, J.D., J.R. WEBB, T.R. WRIGHT, J.D. HARGETT, River basin. Bulletin of Environmental Contamina- K.J. MASK, AND D.R. SCHUCKER. 1994. Food habits, tion and Toxicology 58:688–695. sex ratios, and size of longnose gar in southwestern HUGGETT, D.B., J.A. STEEVENS, J.C. ALLGOOD, C.B. LUTKEN, Oklahoma. Proceedings of the Oklahoma Academy C.A. GRACE, AND W. H . B ENSON. 2001. Mercury in of Science 74:41–42. sediment and fish from north Mississippi lakes. VON BERTALANFFY, L. 1938. A quantitative theory of organic Chemosphere 42:923–929. growth. Human Biology 10:181–213. JAROSZEWSKA, M., K. DABROWSKI, AND G. RODRIGUEZ. WATANABE, K.H., F.W. DESIMONE, A. THIYAGARAJAH, 2010. Development of testis and digestive tract in W. R . H ARTLEY, AND A.E. HINDRICHS. 2003. Fish tissue longnose gar (Lepisosteus osseus) at the onset of exoge- quality in the lower Mississippi River and health nous feeding of larvae and in juveniles. Aquaculture risks from fish consumption. Science of the Total Research 41:1486–1497. Environment 302:109–126. JOHNSON, B.L., AND D.B. NOLTIE. 1996. Migratory dynamics of stream-spawning longnose gar (Lepisosteus osseus). Received 9 June 2011 Ecology of Freshwater Fish 5:97–107. Accepted 11 October 2011 ______. 1997. Demography, growth, and reproductive allo- cation in stream spawning longnose gar. Transactions of the American Fisheries Society 126:438–466.