North American Journal of Aquaculture 65:330–338, 2003 q Copyright by the American Fisheries Society 2003
Effects of Temperature and Salinity on Growth of Juvenile Black Sea Bass, with Implications for Aquaculture CHARLES F. COTTON* Department of Fisheries Science, Virginia Institute of Marine Science, College of William and Mary, Post Office Box 1346, Gloucester Point, Virginia 23062, USA
RANDAL L. WALKER University of Georgia Marine Extension Service, Shellfish Aquaculture Laboratory, 20 Ocean Science Circle, Savannah, Georgia 31411, USA
TODD C. RECICAR Skidaway Institute of Oceanography, 10 Ocean Science Circle, Savannah, Georgia 31411, USA Abstract.—The black sea bass Centropristis striata has recently gained popularity in the live seafood markets of the northeastern United States. Fish farmers need instruction on optimizing environmental parameters for the growth of black sea bass. In this study, optimal temperature and salinity were determined experimentally for the growth of juvenile black sea bass (initial mean weight, ;9.2 g). The temperature experiment compared growth at temperatures of 15, 20, 25, and 308C; the salinity experiment compared growth at 10, 20, and 30‰ salinities. Both of these experiments were performed in closed aquaria. At the end of the temperature experiment (6 weeks), Tukey’s Studentized range test (a 5 0.05) showed that fish reared at 258C were significantly larger than those reared at 208C and 308C. All of these temperatures produced significantly larger fish than did the 158C treatment. At the end of the salinity experiment (12 weeks), Tukey’s Studentized range test (a 5 0.05) showed that salinities of 20‰ and 30‰ did not produce significantly different weights in fish. However, both of these salinities produced significantly larger fish than did a salinity of 10‰. Given the results of these experiments, fish farmers can manipulate the environmental parameters of their aquacultural systems to optimize growth of juvenile black sea bass, thereby reducing the time required to produce a marketable product.
Black sea bass Centropristis striata is a commonly sought fish throughout its range from Florida to Massachusetts (Musick and Mercer 1977). From 1979 to 1990, recreational fishing accounted for 43% of black sea bass landings (by weight), while commercial landings accounted for 33% of the total harvest and headboat landings accounted for 24% (Vaughan et al. 1995). A developing market for sushi-grade black sea * Corresponding author:
[email protected] Received August 28, 2002; accepted April 25, 2003
bass is offering desirable prices for live fish. Currently in Georgia, fishers are offered about US$5.50 per kilogram for whole fish (C. Phillips, Phillips Seafood, Inc., personal communication). Live black sea bass, however, can be transported to markets in the northeastern United States and sold for $11.00–17.60 per kilogram (G. Kinard, Georgia Aquafarms, personal communication). Berlinsky et al. (2000) reported that the demand for black sea bass in the northeastern United States usually exceeds supply, thereby providing a lucrative market for live fish. Despite a sizable volume of literature pertaining to black sea bass aquaculture, commercial culture of this species has yet to fully develop. In Georgia and a few other states on the Atlantic coast, the live market has been supplied by pot-trapped fish taken from nearshore reefs. Fishers trap fish of minimum legal size and transport them to land-based tanks. There the fish are grown to a market size of about 900 g and are hauled alive to the markets of the Northeast. Vaughan et al. (1995) reported that catch per unit effort (CPUE) for black sea bass has declined in the South Atlantic Bight. New size regulations make the practice of trapping black sea bass more difficult as fishers find it harder to catch great numbers of minimum-legal-sized fish. Our research strives to satisfy demand and support an emerging aquacultural industry, while protecting wild stocks of black sea bass. These experiments are part of an ongoing effort to develop economically viable methods for mariculture of black sea bass using hatchery-reared fingerlings instead of wild-trapped adults and subadults. Our work was intended to build upon previous studies related to the aquaculture of juvenile
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FIGURE 1.—Plot of mean weight (6SE) versus time for juvenile black sea bass grown in closed tanks at temperatures of 15, 20, 25, and 308C.
black sea bass (Harpster et al. 1977; Kim 1987; Berlinsky et al. 2000; Atwood et al. 2001) by experimentally determining the optimal temperature and salinity for this species. Our research should serve as a starting point for constructing an understanding of the salinity and temperature requirements across all life stages of the black sea bass, from fry to market size. Fish farmers are currently unaware of the effects of temperature and salinity on growth of juvenile black sea bass. The results of these experiments will guide fish farmers in adjusting environmental parameters to exploit optimal temperature and/or salinity for growth, thereby reducing the time to market for their product. Methods Broodstock were captured in pot traps off the coast of Virginia and later subjected to hormoneinduced spawning to produce the fingerlings used in these experiments. The fingerlings (mean weight 6 SE 5 1.29 6 0.02 g) were purchased
in September 2000 from Southland Fisheries Corporation in Edisto, South Carolina. Fish were kept for 3 months in 600-L, fiberglass holding tanks with a flow-through circulation of estuarine water from the Skidaway River, and supplemental aeration was provided by air stones. During the holding period, fish were fed Zeigler Salmon Starter (numbers 2 and 3 crumble) daily to apparent satiation. Temperature experiment.—Fish (9.0 6 0.16 g) were raised for 42 d in 76-L, closed glass aquaria in four climate-controlled rooms. Temperatures in each room (and thus water temperatures) were maintained at 15, 20, 25, and 308C, respectively. Fish were kept under constant light levels (30 lx) and provided with supplemental aeration by air stones. Tanks were equipped with an undergravel biofiltration system. Prior to the experiment, conditioning of the biofilter of each aquarium was performed (Moe 1992). Fish were fed a floating pellet (Rangen XTR 450, 2.4-mm pellet, 45% protein, 16% lipid) at a 3.0% daily ration (dry weight of food per wet weight of fish). Twelve tanks were stocked with 30 fish per tank. A random number table was used to assign one of four treatments to each tank, for a total of three replicate tanks per treatment. On a biweekly schedule, each fish from each tank was weighed, and tanks were cleaned. For the initial tank stocking and subsequent biweekly weighing, fish were individually weighed in a tared, 1-L beaker of seawater on an Acculab V-1200 balance. Fish were not anesthetized during weighing. After each weighing, new rations were calculated and adjusted for each tank. Treatment means (weight) and arcsine-transformed mortality data were analyzed by analysis of variance (ANOVA) and Tukey’s Studentized range test (SAS Institute 1989). Tests for normality and homoscedasticity were satisfied prior to employing the ANOVA and Tukey’s tests. Relative growth rate (RGR), specific growth rate (SGR), and daily
TABLE 1.—Initial and final weights, weight gain, relative growth rate (RGR), specific growth rate (SGR), daily weight gain (DWG), feed conversion ratio (FCR), and adjusted FCR (FCRadj ) for juvenile black sea bass grown in closed tanks at 15, 20, 25, and 308C. Initial and final weight values are presented as means 6 SEs. See text for definitions of variables. Temperature (8C) 15a 20 25 30 a
Initial weight (g) 8.94 8.76 8.96 9.34
6 6 6 6
0.28 0.34 0.29 0.38
Final weight (g)
Weight gain (g)
RGR (%)
SGR (%/d)
DWG (g/d)
FCR
FCRadj
6 6 6 6
20.23 4.60 7.87 5.23
22.6 52.5 87.8 56.0
20.06 1.00 1.50 1.06
20.005 0.110 0.187 0.125
25.62 2.86 1.57 14.84
236.09 2.02 1.32 2.23
8.71 13.36 16.83 14.57
0.30 0.59 0.70 0.68
Fish in the 158C treatment lost weight over the course of the 6-week experiment, thereby producing the negative FCR and growth rates.
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FIGURE 2.—Plots of (A) mean (6SE) dissolved oxygen concentration and (B) mean (6SE) total ammonia nitrogen concentration of each treatment for the duration of the temperature experiment, January 5–February 16, 2001 (1 ppm 5 1 mg/L). Measurements were taken at least weekly, and values of replicate tanks were averaged for each treatment.
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weight gain (DWG) were calculated with formulas (1), (2), and (3), respectively: RGR 5 100 3 (final mean weight 2 initial mean weight) 4 (initial mean weight)
(1)
SGR 5 100 3 [log e (final mean weight) 2 log e (initial mean weight)] 4 (time in days)}
(2)
DWG 5 (final mean weight 2 initial mean weight) 4 (time in days) (3) Feed conversion ratio (FCR) was calculated with formula (4). To account for weight loss attributed to mortality, an adjusted FCR was calculated with formula (5). The term ‘‘tank weight’’ refers to the total weight of all fish in the tank. FCR
5 (weight of feed consumed) 4 (final tank weight 2 initial tank weight)
(4)
FCRadj 5 (weight of feed consumed) 4 [(final tank weight) 2 (number of fish remaining at the end of the experiment) 3 (initial mean weight)] (5) Estuarine water withdrawn from the Skidaway River was sterilized with 12.5% sodium hypochlorite, and any excess chlorine was neutralized with sodium thiosulfate the following day. Water sterilized in this manner was used for initial filling of aquaria and water exchanges (about 80%) at least twice per week. Salinities ranged from 27‰ to 34‰ throughout the experiment and varied according to ambient salinity in the Skidaway River at the time of the water withdrawal. All tanks were given water of equal salinity during water changes, and salinity never changed by more than 3‰ after any single water change. Water temperature (8C), salinity (‰), dissolved oxygen (mg/L), total ammonia nitrogen (TAN; mg/L), and pH were measured at least weekly. Dissolved oxygen was measured with an Orion (Model 830) dissolved oxygen meter, and pH was measured with an Oakton (Model pH Testr2) pH meter. Salinity and temperature were measured with a refractometer and alcohol thermometer. Total ammonia nitrogen was measured with a LaMotte colorimeter (Model SMART). Salinity experiment.—Fish (9.4 6 0.30 g) were raised for 81 d in 114-L, closed glass aquaria, each assigned a salinity of 10, 20, or 30‰. Tanks were
FIGURE 3.—Plot of mean (6SE) weight versus time for juvenile black sea bass grown in closed tanks at 10, 20, and 30‰ salinities.
kept in a climate-controlled room, and room temperature was maintained at about 228C. Nine tanks were stocked with 30 fish per tank. A random number table was used to assign one of the three treatments to each batch of 30 fish prior to stocking, for a total of three replicates per treatment. Each batch of fish was incrementally acclimated to its assigned salinity treatment over the course of 4 d prior to the start of the experiment. Ambient salinity of the flow-through system in which the fish were held prior to stocking was 32‰. Salinities of three reservoirs (10, 20, and 30‰) were adjusted by diluting estuarine water from the Skidaway River. The diluted water was then sterilized as in the temperature experiment. The sterilized water was used for initial filling of aquaria and subsequent water exchanges (about 80%) at least twice per week. In all other respects, this experiment was conducted in the same manner as the temperature experiment. Computation and analysis of all growth, mortality, and feed conversion data were performed by the same methods used for the temperature experiment. Results Temperature In the temperature experiment (Figure 1), significant differences (P , 0.0001) in weight of fish were found after 6 weeks. Tukey’s Studentized range test (a 5 0.05) showed that fish reared at 258C were larger than those reared at 20 8C and 308C. All three temperatures produced significantly larger fish than did the 158C treatment. Fish
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TABLE 2.—Initial and final weights, weight gain, relative growth rate (RGR), specific growth rate (SGR), daily weight gain (DWG), feed conversion ratio (FCR), and adjusted FCR (FCRadj ) for juvenile black sea bass grown in closed tanks with 10, 20, and 30‰ salinities. Initial and final weight values are presented as means 6 SEs. See text for definitions of variables. Salinity (‰) 10 20 30
Initial weight (g)
Final weight (g)
Weight gain (g)
RGR (%)
SGR (%/d)
DWG (g/d)
FCR
FCRadj
9.29 6 0.52 9.42 6 0.48 9.35 6 0.54
12.67 6 0.84 21.66 6 1.20 20.86 6 1.15
3.38 12.24 11.51
36.4 129.9 123.1
0.38 1.03 0.99
0.042 0.151 0.142
28.84 2.99 2.86
8.09 2.37 2.46
grown at 208C and 308C were not significantly different in weight. Weight gain, RGR, SGR, and DWG were highest in the 258C treatment (Table 1), and FCR was lowest in the 258C treatment. Mortality (mean 6 SE) for each treatment after 6 weeks was 12.2 6 2.9% at 158C, 13.3 6 1.9% at 208C, 12.2 6 1.1% at 258C, and 32.2 6 3.3% at 308C. Significant differences (P 5 0.0020) in mortality were found among treatments. Tukey’s Studentized range test (a 5 0.05) showed that mortality was significantly higher in the 308C treatment when compared to all other treatments. No significant differences in mortality were found among the 15, 20, and 258C treatments. Water quality data for the temperature experiment are presented in Figure 2. Dissolved oxygen ranged from 4.23 to 6.70 mg/L and varied by treatment due to the effect of temperature on oxygen saturation. Total ammonia nitrogen ranged from 0.00 to 1.17 mg/L throughout the experiment. The pH and salinity values were virtually constant (pH ø 7.2–8.0, salinity ø 27–34‰) throughout the experiment. Similarly, constant temperature was maintained in each treatment throughout the experiment. Salinity In the salinity experiment (Figure 3), significant differences (P , 0.0001) in weight of fish were found after 12 weeks. Tukey’s Studentized range test (a 5 0.05) showed that salinities of 20‰ and 30‰ produced significantly larger fish than did a salinity of 10‰. Fish reared in salinities of 20‰ and 30‰ were not significantly different in weight. Weight gain, RGR, SGR, and DWG were highest in the 20‰ treatment (Table 2). Of the positive values obtained for the feed conversion ratio, that for the 30‰ treatment was lowest, but FCRadj was lowest in the 20‰ treatment. A negative FCR was calculated for the 10‰ treatment, reflecting the loss of total weight in this treatment due to high mortality. Mortality for each treatment after 12 weeks was
41.1 6 2.9% at 10‰, 21.1 6 8.0% at 20‰, and 14.4 6 1.1% at 30‰. Significant differences (P 5 0.0209) in mortality were found among treatments. Tukey’s Studentized range test (a 5 0.05) showed that mortality was significantly higher for fish reared at 10‰ salinity than for fish reared at 30‰. Mortality in the 20‰ treatment was not significantly different from that of the other treatments. Water quality data for the salinity experiment are presented in Figure 4. Dissolved oxygen ranged from 5.20 to 6.43 mg/L and TAN ranged from 0.00 to 1.59 mg/L throughout the experiment. The pH and temperature values were virtually constant (pH ø 7.6–8.2, temperature ø 21–248C) throughout the experiment. Similarly, salinity was carefully maintained in each treatment during the experiment. Discussion Temperature The results of the temperature experiment indicate that juvenile black sea bass grow optimally at 258C. This is consistent with the results of another temperature experiment with black sea bass. Berlinsky et al. (2000) found that larval black sea bass grew significantly larger at 228C than at 188C. They also suggested that higher growth rates could be attained at higher temperatures. Similar temperatures have been reported for optimal growth of other subtropical and temperate fishes. McVey (1991) reported an optimal growth rate for European sea bass Dicentrarchus labrax at 228C. Woiwode and Adelman (1991) reported an optimal temperature range of 25.7–27.98C for growth of juvenile hybrid striped bass (female striped bass Morone saxatilis 3 male white bass M. chrysops), with a significant temperature–photoperiod interaction. We abandoned a previous temperature experiment designed identically to this experiment because of a bacterial infection with Mycobacteria marinum. Unpublished data from that experiment showed significantly higher growth rates at 308C
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FIGURE 4.—Plots of (A) mean (6SE) dissolved oxygen concentration and (B) mean (6SE) total ammonia nitrogen concentration of each treatment for the duration of the salinity experiment, January 15–April 6, 2001 (1 ppm 5 1 mg/L). Measurements were taken at least weekly, and values of replicate tanks were averaged for each treatment.
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(not 258C, as in this experiment). One possible explanation for the apparent anomaly between the two experiments is that the former experiment used Georgia broodstock, whereas the latter experiment used Virginia broodstock. Mercer (1989) indicated that those fish came from two separate stocks. The northern stock would likely not experience temperatures as high as 308C, whereas the southern stock experiences this temperature annually and may be better adapted to grow under such conditions. For this reason, it is recommended that future research should report the origin of the broodstock for aquaculture studies of hatcheryreared juvenile black sea bass. Atwood et al. (2001) reported an upper thermal tolerance of 33.38C for black sea bass. Such a result explains the high mortality in the 308C treatment, since this temperature was near the upper thermal tolerance. The fish used in the Atwood et al. (2001) study were spawned from the northern stock, as were the fish in our experiment (H. L. Atwood, Clemson University, Department of Environmental Toxicology, personal communication). Black sea bass can tolerate a wide range of temperatures in the wild. Cupka et al. (1973) reported catching juvenile black sea bass in water temperatures ranging from 5.68C to 30.48C in South Carolina. The majority of these fish were caught in temperatures above 108C, and the authors claim that black sea bass exhibit no temperature preference above 108C. Similarly, Musick and Mercer (1977) reported catching fish in the Chesapeake Bight within a range of 6–268C, with the majority of fish being caught in temperatures above 98C. They also reported that juvenile black sea bass were caught in trawls in the York River only when bottom temperatures exceeded 9–108C. Mercer (1989) also describes black sea bass collections in a temperature range of 6–298C in North Carolina. Our results may also aid in understanding black sea bass population dynamics in the South Atlantic Bight. The southern population undergoes a major spawn in the spring and a minor spawn in the fall (Wenner et al. 1986). In Georgia, fish spawned in the spring will experience as much as 8 months of water temperatures of 208C or higher before winter arrives and retards their growth. Those fish spawned in the fall, however, will experience as little as 1 month of water temperatures of 208C or higher. These fish, although of the same year-class as the fish spawned in the spring, will presumably be smaller in size during the first year of growth. Fish farmers need to understand growth rates at
all relevant temperature ranges if they intend to use flow-through tanks or tanks with inadequate temperature regulation. The juveniles in our temperature experiment lost weight in the 15 8C treatment during the 6-week period. This is unacceptable for a commercial production facility; therefore, the fish farmer will need to utilize heaters during winter months, as water temperatures are typically lower than 158C in Georgia and will be much colder in more northerly latitudes. Fish farmers will also need to understand the degree of the differences in growth rates at these temperatures to determine how much heating will be required in the winter to balance optimal growth with the additional operating costs associated with increased water heating. Conversely, farmers will need to know how much chilling (if any) will be required in the summer to balance optimal growth with the increase in operating costs. Salinity The results of the salinity experiment indicate that juvenile black sea bass grow optimally at either 20‰ or 30‰ salinity. These results differ from another salinity experiment with black sea bass. Berlinsky et al. (2000) reported a significant increase in growth of juvenile black sea bass at 20‰ salinity compared to salinities of 32‰ or 10‰. The size of the fish was slightly different in the two experiments (3.7 g in the Berlinsky et al. experiment versus 9.4 g in our experiment) and could account for the conflicting results. Atwood et al. (2001) showed that the lower threshold of salinity tolerance for black sea bass is 4–6‰. Berlinsky et al. (2000) also anecdotally mentioned 9‰ as the lowest observable tolerance of salinity for black sea bass. This explains the high mortality we observed in the 10‰ treatment, as this salinity was near the lower threshold of salinity tolerance for black sea bass. Black sea bass can tolerate a wide range of salinities in the wild. Cupka et al. (1973) reported catching juvenile fish in salinities from 8.8‰ to 37.8‰, but most were caught in salinities above 30‰, with an overall increasing CPUE as salinity increased. Mercer (1989) also reported catching fish in salinities ranging from 1‰ to 36‰. Fish caught at the lower ranges of these reported salinities were presumably just passing through these hyposaline areas or perhaps were captured below the pycnocline, since Atwood et al. (2001) reported a lower lethal threshold of between 4‰ and 6‰. Duplication of this experiment for subadult and
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adult black sea bass will be important. Cupka et al. (1973) and Musick and Mercer (1977) reported that juvenile black sea bass are known to inhabit estuarine areas as well as offshore reefs. Additionally, Waltz et al. (1979) showed a positive correlation between black sea bass age and distance offshore. This suggests that older black sea bass might not be able to tolerate a salinity range as low as that experienced by the juveniles in our experiment. Therefore, older black sea bass may prove to exhibit significantly lower growth rates at 20‰ than at 30‰. Although growth was significantly reduced and mortality was significantly higher in the 10‰ salinity treatment, the fish exhibited positive growth. For aquaculturists intending to grow black sea bass in facilities located some distance inland, the salinity of the available water source may be about 10‰. In addition, one farmer intends to make artificial seawater by use of well water and Instant Ocean salt mix. Knowledge of the degree of difference in growth rates at 10, 20, and 30‰ will be important in determining acceptable losses in growth in terms of the cost of increasing the salinity levels with artificial salt. Given the high mortality, low growth rates, and high FCR exhibited in this experiment, raising juvenile black sea bass in water with 10‰ for extended periods of time would not be advisable. Ephemeral exposure to lower salinities may eventually prove to be a useful tool for managing parasites, such as dinoflagellates Amyloodinium spp., which have been problematic in culturing red drum Sciaenops ocellatus. Water quality was never problematic during either experiment (Figures 2, 4). Mean TAN concentrations were occasionally recorded at levels above 1 mg/L in both experiments, and as high as 3.5 mg/L in one tank of the salinity experiment, but these levels never seemed to adversely affect the fish. Additionally, Hoff (1970) noted that the fish in his experiments experienced dissolved oxygen levels as low as 3 mg/L for periods of 1–3 d ‘‘with no apparent effect’’ on the fish. Black sea bass have exhibited hardiness, excellent market potential, fast growth, and tolerance of deleterious water quality (Hoff 1970; Roberts et al. 1976; Harpster et al. 1977; Tucker 1984; Kim 1987; Berlinsky et al. 2000; Kupper et al. 2000; Walker and Moroney 2000). Black sea bass also quickly adapt to a tank environment and readily accept artificial food (Hoff 1970; Kim 1987; Berlinsky et al. 2000). With its high resilience, adapt-
ability, and desirable market value, this fish is highly recommended for mariculture. Acknowledgments The authors would like to thank Georgia Sea Grant College Program for its partial funding of this research. In addition, the invaluable assistance of Mary Sweeney-Reeves and Dodie Thompson made this work possible. The editorial reviews offered by Cecil Jennings and Richard Lee were greatly appreciated. The authors would also like to thank Georgia Aquafarms for loan of equipment, boat time, and hours of insightful advice on growing black sea bass. The authors do not endorse any of the products mentioned in this report. References Atwood, H. L., S. P. Young, J. R. Tomasso, and T. I. J. Smith. 2001. Salinity and temperature tolerances of black sea bass juveniles. North American Journal of Aquaculture 63:285–288. Berlinsky, D., M. Watson, G. Nardi, and T. M. Bradley. 2000. Investigations of selected parameters for growth of larval and juvenile black sea bass, Centropristis striata L. Journal of the World Aquaculture Society 31:426–435. Harpster, B. V., D. E. Roberts, Jr., and G. E. Bruger. 1977. Growth and feed conversion in juvenile southern sea bass, Centropristis melana (Ginsburg), fed commercial and semi-natural diets. Proceedings of the World Mariculture Society 8(12):795–809. Hoff, F. H., Jr. 1970. Artificial spawning of black sea bass, Centropristes striatus melanus Ginsburg, aided by chorionic gonadotropic hormones. Florida Department of Natural Resources Marine Research Laboratory Special Scientific Report 25. Kim, J. W. 1987. Growth potential of young black sea bass Centropristis striata, in artificial environments. Doctoral dissertation. Old Dominion University, Norfolk, Virginia. Kupper, R. W., D. H. Hurley, and R. L. Walker. 2000. A comparison of six diets on the growth of black sea bass, Centropristis striata, in an aquacultural environment. University of Georgia, Marine Extension Bulletin 21, Savannah. McVey, J. P. 1991. CRC handbook of mariculture, volume II. Finfish aquaculture. CRC Press, Ann Arbor, Michigan. Mercer, L. P. 1989. Species profiles: Life histories and environmental requirements of coastal fishes and invertebrates (South Atlantic), black sea bass. U.S. Fish and Wildlife Service Biological Report 82. Moe, M. A. 1992. The marine aquarium handbook: beginner to breeder. Green Turtle Publications, Plantation, Florida. Musick, J. A., and L. P. Mercer. 1977. Seasonal distribution of black sea bass, Centropristis striata, in the Mid-Atlantic Bight with comments on the ecology and fisheries of the species. Transactions of the American Fisheries Society 106:12–25.
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Roberts, D. E., Jr., B. V. Harpster, W. K. Havens, and K. R. Halscott. 1976. Facilities and methodology for the culture of the southern sea bass (Centropristis melana). Proceedings of the World Mariculture Society 7(1):163–198. SAS Institute. 1989. SAS/STAT user’s guide, version 6, 4th edition, volume 1. SAS Institute, Cary, North Carolina. Tucker, J. W. 1984. Hormone induced ovulation of black sea bass and rearing of larvae. Progressive FishCulturist 46:201–204. Vaughan, D. S., M. R. Collins, and D. J. Schmidt. 1995. Population characteristics of the black sea bass Centropristis striata from the southeastern U.S. Bulletin of Marine Science 56:250–267. Walker, R. L., and D. A. Moroney. 2000. Growth of juvenile black sea bass, Centropristis striata, fed
either a commercial salmon or trout diet. University of Georgia, Marine Extension Bulletin 22, Savannah. Waltz, W., W. A. Roumillat, and P. K. Ashe. 1979. Distribution, age structure, and sex composition of the black sea bass, Centropristis striata, sampled along the southeastern coast of the United States. South Carolina Marine Resources Center Technical Report 43. Wenner, C. A., W. A. Roumillat, and C. W. Waltz. 1986. Contributions to the life history of black sea bass, Centropristis striata, off the southeastern United States. U.S. National Marine Fisheries Service Fishery Bulletin 84:723–741. Woiwode, J. G., and I. R. Adelman. 1991. Effects of temperature, photoperiod, and ration size on growth of hybrid striped bass x white bass. Transactions of the American Fisheries Society 120:217–229.
