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This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute.

Notice: ©2000 John Wiley & Sons, Inc. This manuscript is an author version with the final publication available at http://www.wiley.com/WileyCDA/ and may be cited as: Tucker, J. W., Jr., Venizelos, A., & Benetti, D. D. (2000). culture. In R. R. Stickney (ed.), Encyclopedia of Aquaculture. (pp. 418- 421). New York: John Wiley and Sons, Inc.

418 GROUPER CULlURE

BIBLIOGRAPHY REPROD UCTIO N

1. N. Stone, E. Park, L. Dorm an, and H. Thomforde, Baitfish. Most studied mature within 2 to 6 yrs (7). Many Culture in Arkansas: Golden Shiners, Goldfish and Fathead serranids are pro togynous hermaphrodites (8,9). Som e Minnows, Uni versity of Arkansas at Pin e Bluff, Pine Bluff, AK; species, as a rule, change from female to male with age, 1997. while others might change only if there is a shortage of 2. RG. Piper, LB. McElwain, L.E.Orm e, J .P. McCraren , L.G. males. In nature, Nassau groupers (Epinep helus striatus) Fowler, and J.R Leonard, Fish Hatchery Man agement, US spawn in large aggregations (100s to 1,000s of fish) Department of the In terior, Washington, DC, 1982. with a sex ratio near 1: 1. Gag groupers (Myc teroperca 3. N. Stone, E. Park, L. Dorman, and H. Thomforde, World microlepis ) spawn in harems, with a sex ratio often near Aquaculture, December 5- 13 (1997). 1 male : 10 females. For both species, individual spawning 4. J .T. Davis, in RR Stickn ey, ed., Cult ure of Nonsalmonid events usu ally involve small numbers of fish (e.g., 2 to 5). Freshwater Fishes, CRC Press, Boca Raton, FL, 1993, pp.307- 321. Small serranids often spawn in pairs without aggregating. A few species are simultaneou s hermaphrodites, but self­ fertilization seems to be rare. See also BAITFISH CULTURE . Voluntary spawning of captive groupers has occurred mostly with well-fed, uncrowded fish during the natural spawningseason under conditions of ambient temperature GROUPERCULT U RE and partial or total natural light (7,10). Day length seems to be a less important stimulus than temperature. At least J OHN W. TUCKER, J R. 27 serranid species have spawned voluntarily in captivity, Harbor Branch Oceanographic In stitution with groupers spawning in 1- to 21,200-m3 tanks or ponds Fort Pierce, Florida and 26- to 75-m3 cages. In Kuwait, 40 female and 9 male ARIETTAVENIZELOS orangespotted groupers ( coioides) held in a National Marine Fish eries Service , NOAA concrete tank spawned almost continuously for 50 days Virginia Key, Florida during April to June (11). In the Philippines, 1 female orangespotted grouper with 2 males held in a 48-m 3 DANIELD. BENETTI cage spawned 5 to 10 times a month for 4 months (12). Uni versity of Miami In Singapore, during December 1989 to October 1990 , Miami, Florid a 10 fem ale and 10 male brownmarbled groupers held in a 75-m3 cage spa wned 2 to 5 times during each of O UTLINE nine periods of 2 to 6 days , usu ally starting between the last qu arter moon and new moon (13). In Taiw an, Reprodu ction 8 female leopard coral trout held in a pond produced Larval Foods eggs 110 times during May to October (Che n et al., 1991a, cited in 7). In Florida, 3 or 4 female Nassau Compound Feeds groupers and 2 males held in a 37-m 3 raceway spawned Raising Groupers to Market Size near the full moon in March and April, with each Health female spawning as many as 9 times a day for 1 to 4 Annual Production days (14). General Comments Hormone-induced ovulation of rip e, wild , or captive Bibliography groupers also is reliable (7,10). At least 31 serranid species have been induced to ovulate . Typically, a female with fully Groupers are classified in 14 gene ra of the subfamily yolked oocytes will ovula te within 24 to 72 hours (usually Epinephelinae, which comprises at least hal f the approxi­ 36 to 50 hours) after the first of 1 to 3 injections of 500 to mately 449 species in the family . Throughout 1,000 IV human chorionic gonadotropin/kg body weigh t. most warm and temperate marine regions, serranids are Similar results have been obtained for several species highly value d for food, and both small and large sp ecies given 1 to 3 injections of 10 to 50 ug gonadotropin releasing are kept in aquariums. Maximum size ranges from 12 em hormone analog/kg body weight. GnRH-analogue implants (4.7 in .) total length (TL) for the Pacific creole-fish (Paran­ were effective for spa wning white groupers tEpinephelue thias colonus) to more than 4 m (13 ft) TL (440 kg, 968 lb) aeneus ). For six grouper species with egg diameters of 800 for the groper , or brindlebass, (Epinephelus lanceolatus ). to 1,000 urn, the minimum effective oocyte diameter before Several grouper species have been raised commercially inj ection was in th e range 41 to 61%. For Nas sau groupers, (mainly in Hong Kong, Taiwan, and the Southeast Asian the time from ovul ation to overripeness is only 1 to 2 hours region), usually by growing out captured wild juveniles. at 26 °C. Some species can grow from 15-20 g (0.5- 0.7 oz) to 1 kg Nassau gro upers at 6 kg can produ ce about 900 ,000 (2.2 lb) in about a year. Research has been conducted eggs per day by natural or hormone-induced ovul ation. At on spaw ning and rearing of dozens of serranid species the same size, hormone-treated brownmarbled groupers (Table 1). Th e main accomplishments are reviewed here. can produce 1.7 million eggs. Hormone-treated 1.5-kg Additional information can be found in various publica­ squaretail coraltrout can produce 400 ,000 eggs. AI-kg tions (1- 7). (2.2 Ib) redspotted grouper can produce more than GROUPER CULTURE 419

Table 1. Some Characteristics of Representative Groupers Raised Commercially (C) and Experimentally (E)

Egg Larval Market Maximum Type of Diam eter Duration Source of Size Maximum Age Species Culture" Locations (11 m) (d) Juveniles" (kg) Size" (yr)

Polka dot grouper C SEAsia 890 W 0.5 90 em Cromileptes alt ivelis Redspotted grouper C J apan 825 45-50 H,W 0.5 5 kg,60 em > 6 Epinephelus akaara Hong Kong Squaretail grouper C Hong Kong W 60 em Epinephelus areolatus Orangespotted grouper C SE Asia 807 35-40 H,W 2:95 em Epinephelus coioides Middle Eas t Brownmarbl ed grouper C SEAsia 840 35- 40 H,W 0.6 120 em Epinephelus fuscoguttatus C SE Asia 852 36-60 H,W 0.6 > 25 kg, 115 em Epin ephelu s malabaricus Sevenband grouper C J ap an 820 ~6 0 120 em Epinephelus septemfasciatus Nassau grouper E Carib bean 920 46-70 H 2 > 25 kg, 120 em 16 Epin ephelus striatus Greasy grouper C SE Asia 900 36-50+ H,W 0.6 75 em ~ 2 5 Epinephelus tau vina Leopard eora ltrout C SE Asia 875 ~5 5 H,W 2:20 kg, 80 em leopardus Chinese pereh C PR China ~2 0 00 H 0.45 > 5 kg S iniperca chuatsi

· C = commercial, E = exper imental. bH = from a hatchery, W = from wild stocks.

5 million eggs in a season, and a 6-kg Nassau grouper are not digested well until at least 2 to 4 weeks. Weaning can produce 3.3 million eggs in a 4-day period. can be comp leted just before or during transformation into With good timing and luck, groupers have been cau ght the juvenile stage, which occurs at 35 to 70 days after just before spawning and held in tanks or cages until they hatching, depending on species. ovulate naturally. Th e eggs are stripped , or rarely, the fish are left in the tank for voluntary or accidental fertilization to occur. COMPOUND FE EDS

In nature, juvenile and adult groupers eat mainly fish , LARVAL FOOD S crabs, shri mp, mantis shri mp, lobsters, and molluscs (16). Red groupers (Epinephelus mario) seem to prefer crabs With the notable exception of Siniperca spp., which have first, then shrimp. In Thai land and other areas , groupers large eggs and hatchlings ( ~5 mm) and are easy to feed have been fed mainly trash fish (with vitamins and and rear, grouper larvae usually are small and fragile and minerals) secondarily moist or semimoist pellets, and have relatively small mouths at first feeding. Yolk and oil rarely hi gh-protein dry pellets (17). A suitable starter tend to be exh austed quickly (7,15). Typically, the larval feed for gro upers would contain 50 to 60% high-qu ali ty period is long, and groupers tend to require live food longer protein, 12 to 16% fat, no more than 15% carbohydrate, than most marine fishe s that have been reared. less than 3% fiber, and less than 16% ash (7). Groupers Grouper larvae usually are raised in green water larger than 500 g (1.1 Ib) can be given a feed with (N annochloropsis, Tetraselmis, Chlorella spp.). At first approximately 45% protein, about 9% fat, and no more feeding, most species can eat small rotifers, bu t oyster or than 20% carbohydrate, 4% fiber, and 22% ash. Lower clam eggs and trochophore larvae sometimes are used as qu ality feeds likely would result in a higherfeed conversion a supplement. Growth and survival rates tend to increase ratio and possibly slower growth. if copepods or mixed zooplankton are included in th e diet, but care mu st be used to avoid introduction of pathogen s or predators. Enriched Artemia can be a staple food beginning RAISING GROUP ERS TO MARKET SIZE at 10 to 30 days, bu t their density should be controlled to minimize gorging. Microfeeds (artificial diets) have been In Indo-Pacific and Middl e Eastern regions, several species tried as a supplement during the first week, but probably of grouper are farmed in cages, ponds, and tanks, but 420 GROUPER CULTURE usually they are raised from wild juveniles and are larval tanks can allow the water to stratify and zooplank­ fed trash fish . They sometimes are fed small tilapia ton and fish to aggregate dangerously . With too much (Oreochrom is spp.) and occasionally are polycultured with turbulence, the fish are battered. Gorging on Artemia is them. Typical market size is 500 to 1,000 g (1.1- 2.2 lb), another source of mortality, and cannibalism among early which can be reached in 6 to 8 months of grow-out. The juveniles can be a problem. Larvae are fragile, and sur­ minimum size to begin grow -out, 75 to 100 mm, can be vival from eggs to juveniles often has been only 0 to 1%, obtained in nursery tanks, cages, or ponds. They are but juveniles and adults are among the hardiest of fish . stocked up to 60 fish/in" « 1 kg/m") in cages. In Taiwan, Grouper farming fluctuates because of variability in a pond farm typically stocks 60,000-80,000 groupers/ha the (mostly decreasing) supply of wild juveniles and and harvests 80% of them for a production of 30,000 to lack of sustained hatchery production for most species. 40,000 kg/ha; the groupers are fed mostly trash fish and Variability in quantity and quality of trash fish and grow from 46 mm to 600 g in 12 months and 2 kg (4.4lb) th e lack of economical compound feeds also has been a in 19 months (18). When fed pellets only, Nas sau groupers constraint in some areas. Nevertheless, the commercial can reach at least 450 g (L Ib) at 12 months and 2 kg feasibility of grouper culture has been proven in several (4.4 lb) at 24 months of age . countries, including Malaysia, Hong Kong, Taiwan, China, Philippines, Singapore, Japan, and Korea. HEALTH BIBLIOGRAPHY For groupers, snappers, and similar warmwater fish, gram-negative bacteria (Vibrio, Aeromonas, Pseudomonas, 1. Anonymous, Manual on Floating N etcage Fish Fa rm ing in Pasteurella spp.), Streptococcus, Mycobacterium, ectopar­ Singapore's Coastal Wat ers , Singapore Prim. Prod. Dept., asitic protozoans r.Amyloodinium ocellatus, sporozoans, 1986. Cryptocaryon irritans, Brooklynella spp.; Ichthyoph­ 2. J .J. Polovina and S. Ralston, eds., Trop ical Snappers and th irius sp. ), and monogeneans (Ne obenedenia m elleni, Group ers: Biology and Fisheries Management, Westview Diplectanum spp.) are among the most important Press, Boulder, 1987. pathogens (19- 23). In Singapore, sleepy grouper dis ease 3. S. Tookwinas, Actes Colloq. 9, 429-435 (1990). (lethal) probably was caused by a virus introduced with 4. M. Doi, M.B.H.M. Nawi , N.R.B.N . Lah, and Z.E. Talib, Arti­ wild juvenile groupers imported for cage farming (24). ficial Propagation of th e Grouper, Epinephelus suillus at th e Other viral pathogens and dis eases include golden eye dis­ Marine Finfish Ha tchery in Tanjong Demong, Terengganu, ease, red grouper reovirus, spinning grouper disease, and Malaysia , Dept. Fish., Kuala Lumpur, Mal aysia, 1991. viral nervous necrosis. Rancid dietary lipids are thought to 5. K Maruyama, K Nogami, Y Yoshida, and K. Fukunaga, cause nervous suffering dis ease of groupers, which could Th ird Asian Fisheries Forum, Asian Fish. Soc., Manila, 1994, pp. 446 -449. result in gill, blood, gas bladder, liver, heart, brain, and nerve damage (25). In J apan, pa steurellosis has been a 6. F. Arreguin-Sanchez, J .L. Munro, M.C. Balgos, and D. Pauly, eds., Biology, Fish eries and Culture ofTropical Groupers and major disease of young redspotted groupers (26). Snappers, ICLARM Conf. Proc. 48, Manila, 1994. 7. J .W. Tucker, Jr., Marine Fish Culture, Kluwer Acad emi c ANNUAL PRODUCTION Publishers, Boston, 1998. 8. D.Y Shapiro, in J .J. Polovina and S. Ralston, eds., Tropical In 1996, the Peoples' Republic of China produced Snappers and Groupers: Biology and Fisheries Management, 58,437 metric tons (mt) of Chinese perch (mandarin fish), Westview Press, Boulder, 1987, pp . 295-327. which has been considered a serranid, but has affinities 9. D.Y. Shapiro, J. Exp. Zool. 261 , 194-203 (1992) . with the centropomids (snooks), Malaysia produced 837 mt 10. J .W. Tucker, Jr., J. WorldAquacult. S oc. 25, 345-359 (1994 ). of greasy groupers and Hong Kong 360 mt. Hong Kong 11. N.A. Hu ssain and M. Higuchi, Aquaculture 19, 339-350 produced 750 mt of square tail groupers. Taiwan produced (1980). 1,883 mt of miscellaneous groupers, Thailand 600 mt, 12. J .D. Toledo, A. Nagai , and D. J avellana, Aquaculture 115, Philippines 595 mt, Singapore 93 mt, and Republic of 361 -367 (1993). Korea 9 mt. In 1995, Hong Kong produced 30 mt of 13. L.C. Lim , T.M. Chao, and L.T. Khoo, Singapore J . Prim. Ind. redspotted groupers (27). 18, 66-84 (1990). 14. J .W. Tucker, Jr., P.N. Woodward, and D.S. Sennett, J . World GENERAL COMMENTS Aquacult. So c. 27 , 373-383 (1996 ). 15. H. Kohn o, A. Ohno, and Y Taki, Third Asian Fish eries Commercial-scale hatcheries for redspotted grouper and Forum, Asian Fish. Soc., Manila, 1994, pp. 450 -453. kelp grouper (E. bruneus) in Japan and Malabar grouper 16. T. Brule, D.O. Avila, M.S. Crespo , and C. Deniel, Bull. Mar. in Taiwan and Thailand have raised large batches ofjuve­ Sc i. 55, 255-262 (1994). niles, with survival as high as 34% from hatchlings (7). The 17. N. Ruangpanit and R. Yashiro, in KL. Main and larval period is longer than for most cultured fishes . Some C. Rosenfeld, eds., Culture of High-value Marine Fishe s in groupers need small rotifers, trochophores, or copepods at Asia and th e United States, Oceanic Inst., Honolulu, 1995, first feeding. Proper ae ration is critical. Early grouper lar­ pp.167-183. vae, especially when stressed , sometimes exude an excess 18. Anonymous, Aqua Farm N ews 10(3), 9 (1992 ). of mucus, which can cause them to stick to each other, to 19. Y C. Chong and T.M. Chao, Commo n Diseases of Marine the surface film, or to solid objects. Too little turbulence in Foodfish, Sing apore Prim. Prod. Dept., 1986. GULF KILLIFISH CULTURE 421

20. W.-Y. Tseng and S.K Ho, Grouper Culture-A Practical of Mexico coast to eastern Mexico. The fish is also found in Manual, Chien Chieng Publisher, Kaohsiung, Taiwan, 1988 . Cuba (1). 21. D. Gallet de Saint Aurin, J .C. Raymond, and V. Vianas, Actes Male Gulf killifish are a uniform greenish-silver in Colloq. 9, 143-160 (1990). color, while the females are darker and have prominent 22. W.G. Dyer, E.H. Williams, Jr., and L. Bunkley-Williams, J. spots on their bodies and fins . Eggs are laid on suitable Parasitol. 73, 399-401 (1992 ). substrates, such as submerged or emergent vegetation 23. K.-K Lee, Microbial Pathogenesis 19, 39-48 (1995 ). in nature. Spawning can occur over a period of several 24. F.H.C . Chua, M.L. Ng, KL. Ng, J .J. Loo, and J .Y. Wee, J. months, from early spring to early fall. Reproduction has Fish Dis. 17,417-427 (1994 ). been known to occur in the salinity range of 3-20 parts 25. D.-K. Hua, M.-L. Cai , and Z.-Y. Zhang, Third Asian Fisheries per thousand (ppt ), Forum, Asian Fish. Soc., Manila, 1994, pp. 357-360. Eggs hatch in 10-21 days; the time depends on 26. H. Sako , in K.L. Main and C. Rosenfeld , eds., Aquaculture temperature and salinity. Higher temperatures reduce Health Management Strategies for Marine Fishes , Oceanic incubation time, as do lower salinities. Egg hatching has Inst., Honolulu, 1996, pp. 81-90. been reported over a salinity range from 0 to 40 ppt 27. FAD, Aquaculture Production Statistics 1987-1996. FAD with subsequent good fish growth; survival has been Fish. Circ. No. 815 , Rev. 10, 1998. reported over a salinity range from 5 to 40 ppt. Fry survival and growth are reduced in freshwater or at salinities in excess of 60 ppt, yet juveniles arid adults GULF KILLIFISH CULTURE can tolerate very low salinities « 1 ppt) and near-zero dissolved oxygen concentrations for at least briefperiods of

R OBERT R. S TI CKNEY time. Anecdotal reports of high mortalities associated with Texas Sea Grant College Program high pH (9.0- 9.5) have occurred (J ack Booth, personal Bryan, Texas communication), but the mortalities may nave been related to high percentages of un-ionized ammonia (NH3 ). Un-ionized ammonia is the more toxic form. In solution, OUTLINE total ammonia is a combination of ionized ( N~ + ) and un­ ionized ammonia. The percentage of un-ionized ammonia Introduction in the mixture increases with increased pH . In any case, Life History one desirable feature of Gulf killifish is their ability to Culture Techniques survive under the stressful conditions that exist in bait Bibliography buckets.

INTRODUCTION CULTURE TECHNIQUES

The Gulf killifish (Fund ulus grandis ), also commonly The recommended stocking rate for Gulf killifish of known in Texas as the mudfish or mudminnow and in 5-7.5 em (2- 3 in. ) is 30,000lha (12,000/ac) in a ratio oftwo Alabama as the bull minnow, is a widely used baitfish females for each male in advance of the spawning season. within the marine recreational fishing community along Commercial minnow feeds have been used successfully, as the Florida and northern Gulf of Mexico coasts of the have floating catfish feeds and agricultural by-products. United States. Historically, the species has been captured In ponds, natural vegetation can provide suitable from nature and sold through retail bait outlets; however, spawning substrate, but the standard practice is to employ supplies from that source have waned in recent years, and spawning mats that consist of about 1 kg (2.2 lb) of both interest and activity associated with the culture of Spanish moss sandwiched at a thickness of 5-7.5 em Gulfkillifish have developed in areas to which that fish is (2- 3 in. ) between pieces of plastic-coated wire mesh. The native. pieces of wire mesh are typically 0.7 x 1 m (2 x 3 ft) in Gulf killifish are found in coastal marshland areas. size, though any convenient size is suitable. Materials They are common in grassy bays and canals, and even in other than Spanish moss have been used, but Spanish adjacent freshwater areas, because they have a very wide moss continues to be commonly employed and is locally tolerance for salinity. While their maximum length (1) is available along much of the Gulfcoast. reportedly 180 mm (7.1 in .), fish in excess of 15 ern (6 in .) Spawning mats should be evenly distributed along the are not commonly seen. Bait dealers prefer fish of 7.5 ern edges of each brood pond at a rate of no less than 125lha (3 in.) or slightly less. Bait-sized fish can be produced (50/a). In windy areas, the mats should be placed along the from eggs in a few months. A brief guide to their culture, upwind side of the ponds. Placement in that less turbid by Strawn et aI., in 1986, continues to be a highly useful pond region helps avoid silting in of the mats, which would publication, which, unless otherwise noted, forms the basis result in smothering of the eggs. of the following discussion (2). The mats should be fully submerged and should be su spended a few em (1 em = 2.5 in. ) above the bottom, LIFE HISTORY as the fish will deposit their adhesive eggs on all mat surfaces . If the mats lie on the bottom, access to their Gulfkillifish can be found along both the Atlantic and Gulf undersides is denied and the mud can work up into the coasts of Florida, in the Florida Keys, and along the Gulf mats and cause eggs to die.