Molluscan Mariculture in theGreater Caribbean: An Overview
DARRYL E. JORY and EDWIN S. IVERSEN
Introduction
This paper reviews the history andcurrent status of both experimental andcommercial molluscan mariculture inthe greater Caribbean area (Fig. 1). Seafood is and has been a staple for Caribbean people since pre-Columbian times.
ABSIRACF-Marine mollusks suitable formariculture in the Caribbean area havereceived increased attention in recent yearsin an effort to produce more seafood for inhabitants of the area. However, molluscanmariculture in the Caribbean is still, witha few exceptions, in its infancy when compared with these activities elsewhere. Pilotand commercial culture operations forAmerican oysters, Crassostrea virginica, andmangrove oysters, C. rhyzophorae, exist inJamaica, Cuba, J.i'?nezuela, and Mexico, and
In many areas, however, seafood demand exceeds both the current catch andpotentially available resources. A recentreview of eastern Caribbean fisheries byOlsen et al. (1984) incorporated landarea, human population, shelf area,number of tourists, and fish landings byisland/nation. These figures, together
for rock mussels, Perna perna, in f.-enezuela.Extensive research has been carried out onthe mariculture potential ofthe queen conch,Strombus gigas, with experimental hatcheries in some countries (more are under construction or planned), and a commercialhatchery is already operating in Turks andCaicos. The culture potential of severalmollusks in the Caribbean, including nativeand exotic species, and several problems impeding increase in molluscan culture in theCaribbean are discussed.
with total seafood consumption rates(local residents and tourists), permitteda comparison between island-shelfpotential yield and demand for marineprotein in the Caribbean which revealedthat:
1) Only a few of the eastern island nations are currently supplying their ownseafood demand,
2) The current demand for seafood inthe area is about 775,000 metric tons (t),which greatly exceeds both currentlandings of about 87,000 t and the200,000 t potential yield, and
3) The shelf area of many islands is
Darryl E. lory and E. S. Iversen are with the Division of Biology and Living Resources, RosenstielSchool of Marine and Atmospheric Science,University of Miami, 4600 Rickenbacker Causeway, Miami, FL 33149.
VENEZUELA
+---------,I-----22·N
ATLANTIC OCEAN
PrOVJ(Jenctales I
85'W95'W
I
PACIFIC OCEAN
12·N----f------+--------'I.
Figure I.-Map of greater Caribbean region.
47(4), 1985 1
Figure 2.-Mangrove oyster culture system in coastal region.
generally too small to support additionalfishing effort, and although there aresome areas that can support additionalexploitation, increased seafood production must come from resources notalready exploited.
Although almost all island nationspresently have this serious seafood production deficit, some nations are inmore trouble than others. For example,the Dominican Republic, which currently lands about 6,435 t of finfish andshellfish, has a seafood demand of137,000 t and a potential yield from itsshelf of only 864 t.
Mollusk culture could increase seafood production in many Caribbeanisland nations and reduce the differencebetween present production and demand. Another reason for culturingmollusks in the Caribbean is that as export products (for food or the aquariumor ornamental shell trade) they generatehard currency to help alleviate serioustrade deficits that characterize many
2
Caribbean nations, as well as provideneeded employment. Finally, culturedmollusks, in particular cephalopods(Hanlon, In press) and the sea hare,Aplysia spp. (Fay, 1971), can and havebeen used extensively in neuroscienceand behavioral research.
However, despite these incentives,molluscan culture in the greater Caribbean is, with few exceptions, in its infancy when compared with the status ofthese activities in such countries asJapan, Australia, France, Spain, theNetherlands, and others. In addition,available information on the molluscanculture in the Caribbean is limited. Incontrast, the journal Aquaculture(Morse et al., 1984) devoted over 400pages to "Recent innovations in cultivation of Pacific mollusks." Possiblybecause of the vastness of the PacificOcean and variety of marine habitatsand molluscan species, the research effort is more extensive than in the Caribbean. Nonetheless, the importance of
increasing food production in the Caribbean is every bit as important as it is inthe Pacific, and perhaps even more soconsidering the large human populationand the relative lack of other foodsources in many Caribbean islandnations.
Present molluscan mariculture activities in the Caribbean can be divided into three categories: Semi-intensiveculture, extensive culture, and research.
Semi-intensive Culture
In the Caribbean, semi-intensive culture of mangrove oysters, Crassostrearhizophorae, is presently practiced inCuba and Jamaica, while Americanoysters, C. virginica, are raised in Mexico and Venezuela, and the South American rock mussel, Perna perna, iscultured only in Venezuela.
Cuba
Over a 12-year period (1963-74), biological and ecological studies and breed-
Marine Fisheries Review
Figure 3.-Mangrove oyster culture system in Cuba.
ing experiments supervised by UnitedNations Development Program/Foodand Agriculture Organization (UNDP/FAO) experts, were conducted on mangrove oysters in Cuba, and many locations were found suitable for culture. Amangrove oyster culture system wasdeveloped during the study period,based on known methods and the use ofinexpensive and readily available localmaterials (Nikolic et al., 1976).
Farming facilities, generally locatedin estuarine areas, consist of stockadesof palm posts driven into muddy orsandy bottoms, arranged in line or in aquadrangle (Fig. 2). Posts extend about1-1.5 m above the surface of the waterand are placed 2.5 m apart, supporting6 m long, wooden traverse beams. Redmangrove terminal branches, suspendedfrom the traverse beams with tarredropes or monofIlament nylon thread, areused as spat collectors. The collectorsare checked at least once a month (Fig.3) to make certain that the binding ropesare securely fastened, that collectors arefavorably located in terms of tidal cycles,and to remove fouling organisms andpredators. The oysters are harvested 5-6months after placing the collectors. Subsequent harvests take place each monththereafter, when the largest oysters arecollected during monthly cleaning operations. (Nikolic et al., 1976).
The first commercial oyster farm inCuba, located on the northeastern shore,began operating in 1975. About 30 farmswere initially planned by Cuban officials, with an estimated potential production between 6,500 and 7,500 t.However, because of industrial pollutionin the culture areas, only about 20 percent of the area could be used. Presentlythere are 19 farms in operation, with anestimated potential production between900 and 1,100 t of unshucked oysters. Inthe last 3 years, wide fluctuations inseasonality and abundance of spat settlement have adversely affected growoutschemes. To optimize oyster growout,it is necessary to have an adequate, consistent source of spat. A hatchery underconstruction and supervision of FAO experts, combined with ongoing pilot scaleresearch on controlled reproduction andoyster larval maintenance, is expectedto produce sufficient spat consistently I.
47(4), 1985
Jamaica
In July 1977, a joint mangrove oysterculture project was set up by the International Development Research Center(IDRC) of Canada and the Governmentof Jamaica (through the Fisheries Division of the Ministry of Agriculture andthe University of the West Indies'Department of Zoology) to determine its
'Frias-Lepoureau, 1. A. Mariculture Section,Ministry of the Fishing Industry, Cuba. Personalcomrnun., I October 1984.
feasibility in arresting the loss in naturaloyster populations to land reclamation,particularly in the Kingston Harborarea, and to ensure a constant or increased supply of oysters. After 4 yearsthe culture system was considered viableso in 1980 an "Oyster Culture Unit" wasset up within the Ministry of Agricultureto operate pilot farms and provide extension services2•
2MooYoung, R. R. Inland Fisheries Project, Ministry of Agriculture, Jamaica. Personal comrnun.,8 November 1984.
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The system adapted known oyster raftculture methods to Jamaican conditions(Wade et aI., 1981). Spat collectors aremade of pieces of old car tires, cut into8 X 8 em squares and drilled in thecenter. These are strung together withmonofilament line, (10-12 collectors perstring), aged in seawater for 2 weeksbefore use, and then hung from bambooand mangrove racks in the intertidalzone to collect spat. When the preferreddensity of about 10 oysters per collector has settled, the collectors are restrung on long monofilament plasticlines, spaced with 10 em bamboo polesand tied to bamboo rafts. Flotation forthe rafts is provided by 44-gallon oildrums painted with antirust paint, andanchored by nylon ropes tied to80-pound concrete blocks. Market sizeof 7-8 em is reached in 6 months. Onlyabout 10 percent of production issuitable for marketing because of poaching, diseases, and fouling organisms, allmajor problems. In addition, somepotential growout sites located near urban centers may give rise to health problems caused by pollution; therefore, theJamaican government is consideringdepuration plants and strict marketingregulations (ADCP, 1983).
Oyster spat is presently collected atthe pilot farm of the Oyster Culture Unitin Bowden, Port Morant. Growout isonly carried out at this time in Port Antonio, where local fishermen built andhave maintained growout rafts sinceJanuary 1982. Growout was attemptedat Falmouth, but was discontinued dueto high coliform counts. Microbialstudies to institute preventive measuresare under way in a joint project betweenthe University of the West Indies and theUniversity of South Florida.
Belmont-Bluefields is a proposedgrowout site. There are plans to growthe oysters in baskets and trays, whichproduce single shells instead of clusters,and which have more appeal for tourists.There is a high demand for oysters fromhotels and restaurants, and all presentproduction is readily sold in Kingston.No reliable production figures are available but the quantities sold are reportedto be small. Three Jamaican workersfrom the Project have recently completed overseas training at DalhousieUniversity in Canada through IDRC
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scholarships. Finally, an experimentaloyster hatchery is being planned 3•
Mexico
In Mexico, commercial aquacultureactivities involving oysters (and sevenother groups of organisms which include shrimp and lobsters) are reservedexclusively for aquaculture cooperatives. Although the most successfuloyster farming cooperatives are locatedalong Mexico's Pacific coast (culturinga local species, Crassostrea corteziensis), experimental oyster culture in Mexico began in the lagoon of Tamiahua in1957 (Conrad, 1985). American oystersare also raised, and its most importantculture grounds are on Mexico's Gulfcoast, in the lagoons of Pueblo Viejo,Tamiahua, Tampamochoc, MachonaCarmen, Macoacan and others in theStates of Veracruz and Tabasco (Lizarraga, 1974).
Seed is commonly collected on collars of oyster shell cultch, although rooftiles and wire-mesh or plastic bags arealso used. Spat collectors are placed instockades, hung from traverse beams,and after 2-3 months (when the seed is2-3 em) they are moved for growout.Two basic systems are used. In one thecollectors are placed on the bottom inareas consolidated with old oystershells, and in the other, a suspensionsystem is used with collectors strung ongalvanized wire strings, separated byplastic tubing, and hung in stockades.Growout time to commercial size (about8-10 em) is between 8 and 14 months,and the reported yield from these areasaverages 25 tons/ha (Haro et aI., 1983;Lizarraga, 1974).
The potential for increased oyster production in the Gulf of Mexico is considerable, with over 100,000 ha havingpotential for utilizing intensive culturetechniques (Haro et aI., 1983). Currentproduction is about 40,000 t/year, but adecline is predicted due to pollution inlagoons such as Tamiahua and others4,
where oil exploitation activities have
3Sessing,1. Jamaican aquaculturist, P.O. Box 642,Kingston 8, Jamaica. Personal commun., April1985.'Orbe, A. Centro de investigacion y EstudiosAvanzados lPN, Unidad Merida, Merida, Yucatan,Mexico. Personal commun., 6 November 1984.
resulted in destruction of natural beds(Conrad, 1985).
Venezuela
Experimental culture of the Americanoyster began in 1974 in the canals ofGuariquen, in the Gulf of Paria. TheCenter for Fisheries Research of Cumana, under the Ministry of Agriculture and Livestock, beginning in 1974,carried out several studies on the biology and culture potential of the American oyster (unpubl.) which stimulatedthe creation of two commercial venturesin 1980. For spat collection and growout these operations used old tires cutin strips and hung from floating woodenrafts. In November 1980, there were 120rafts: 100 rafts were 15 x 6 m with 420strips each and 20 rafts were 15 X 14m with 800 strips each. Commercialsize, 8-12 em, is reached in 1 year.
The marketed production in 1980,1981, and 1982 was 170, 176 and 132tons, respectively. The estimated production of a single operation in 1980 wasabout 1,000 t, most of which could notbe marketed because the marketingchannels were inadequate to handle production. Studies of the economic feasibility of smoking and canning oystermeats, possibly for export, have beenmade (Cervigon, 1983). However, in arecent visit to Venezuela, the seniorauthor was informed that marketing difficulties had apparently proved insurmountable and that both commercialventures would cease operations.
The mangrove oyster was also thesubject of experimental and commercialculture in Venezuela. Experimentalculture began in the early 1960's, andthere were two commercial attempts,both using wild spat and floating rafts:One, in 1969, in the Gulf of Cariaco andthe other in 1971 in la Restinga lagoon,Margarita Island. Both faced problemsof spat settling, competition, and shellbrittleness, and both failed due to marketing difficulties (Mandelli and Acuna,1975; Cervigon, 1983).
Culture of rock mussels, Perna perna,began in 1960 in the Gulf of Cariaco innortheastern Venezuela (Fig. 4) usingslightly modified Spanish raft-culturetechniques (Iversen, 1966). Presently,there are two private ventures with 20rafts and seven smaller ventures with a
Marine Fisheries Review
Figure 4.-Rock mussel rafts in Venezuela.
total of 45 rafts operated by fishermen'scooperatives. The latter have not produced any mussels in the last few yearsbecause of red tide problems in the areawhich, in August IfJ77, killed nine people who ate contaminated mussels.
Annual market production since IfJ72has ranged from 42 to 650 t. In 1980,production was estimated at 650 to 920t, but only 30 t were marketed due tored tide. In 1983, production was estimated at 182 t. This decrease is attributed to the replacing of wooden spat collectors by tire strips to which musselspat reportedly cannot attach securely.Besides red-tide-related problems, commercially successful mussel culture inVenezuela faces marketing problemssimilar to that of oysters (Salaya et al. ,
47(4), 1985
IfJ73; Mandelli and Acuna, lfJ75; Cervigon, 1983). Extensive research on therearing of mussel larvae has been recently carried out at the Instituto Oceanografico of Universidad de Oriente inCumana5. A project to set up a musseldepuration plant is being planned, andanother project for a large-scale musselfarming operation was recentlypresented to the National Council forScientific Research of Venezuela(CONICIT) and is presently beingevaluated6•
'Cervigon, F. Fundacion Cientifica Los Rogues,Caracas, Venezuela. Personal commun., May1985.6Robaina, G. Universidad de Oriente, Boca de Rio,Isla de Margarita, Venezuela. Personal commun.,December 1985.
Extensive Culture
The queen conch, Strombus gigas, isthe only marine mollusk considered tobe cultured extensively in the Caribbean. Increased demand for its meat,especially in u.s. markets, has resultedin overfishing and the decline of stocks,and has threatened its critical role as oneof the most important subsistence-levelfisheries of the area (Brownell andStevely, 1981).
Concern over the decline of stocks ledto extensive research on the mass-rearing of juveniles in hatcheries and usingthese juveniles to reestablish or replenish depleted natural populations. Suchresearch was carried on since about1980 in Bonaire, Puerto Rico, Los Ro-
5
ques Archipelago in Venezuela, Quintana Roo in Mexico, Turks and Caicos,Miami, and the Berry Islands in theBahamas (Iversen and Jory, 1985).
Preliminary results of many of theseresearch programs were presented in aConch Mariculture Session at the 35thAnnual Gulf and Caribbean FisheriesInstitute in Nassau, Bahamas, in November 1982. What has happened sincethen? The projects at the University ofMiami, Los Roques and the BerryIslands have ended due to lack of financial support. The project in Puerto Ricomight not receive further funding7.
However, there is also encouragingnews. The Bonaire hatchery reportedrearing and releasing 750,000 juvenilesoff Bonaire in 1984 in deep waterswhere they will not be easily accessibleto fishermen8• This may be an exampleof technological success, but does notimply economic feasibility. A hatcheryis being set up at the Hydrolab site atSalt River on St. Croix, U.S. VirginIslands, mainly to obtain animals forfurther research9. And construction ofa hatchery is about to begin on the islandof Martinique, French West Indies1o•
The senior author recently visited thePuerto Morelos hatchery in QuintanaRoo, Mexico, and was informed that itis preparing for its first field release ofhatchery-reared juveniles. Finally, in1984 Trade Wind Industries, Inc.ll, constructed and started operating the firstcommercial queen conch hatchery onthe island of Providenciales, Turks andCaicos. During the 1984 spawningseason, 20 larviculture tanks were inoperation, and the juvenile conchs produced were stocked in protective cagesnear the hatchery. The company reportedly has also acquired sea-bottom
'Appeldorn, R. Department of Marine Sciences,University of Puerto Rico, Puerto Rico. Personalcommun., October 1984."Hensen, R. Department of Agriculture and Fisheries, Bonaire. Personal commun., October 1984.·Coulston, M. L. Hydrolab Project West IndiesLaboratory, Fairleigh Dickinson University, SI.Croix. Personal commun., 22 October 1984.'OBazin, P. Association pour Ie Development deI'Aquaculture a la Martinique, Martinique. Personal commun., September 1984."Mention of trade names or commercial firmsdoes not imply endorsement by the NationalMarine Fisheries Service, NOAA.
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leases to grow conch to commercialsizel2.
The release size for hatchery-rearedconch discussed, and recommendedand/or used so far varies from 2 cm(Siddall, 1983), to 5 cm (Creswell,1984) to 12-15 cm (lory and Iversen,1983; Woon, 1983). The release size canhave important consequences on the rateof survival to market size. There is goodscientific evidence that natural mortalityis greater for smaller individuals, andhatchery-reared mollusks are no exception (Jory et al., 1984), including queenconch (Appeldoorn and Ballantine,1983; Jory and Iversen, 1983) as wellas other mollusks such as abalone13. Recent large-scale releases of small (2-5cm) hatchery-reared conchs in Bonaire,Mexico, Venezuela, and St. Croixshould provide an indication of the optimum release size.
Pilot Projects andResearch Activities
Bahamas
Molluscan mariculture has been attempted at three locations in the Bahamas. In the middle 1970's a private company conducted a pilot experiment withimported American oysters. In a ponddredged out for a dock project in Rudder Cut Cay, Exuma, the water was fertilized and imported oyster spat wasplaced in rafts. The project was promptly abandoned due to very slow growthrates.
The second study, growing oystersand clams in rafts by the Wallace GrovesAquaculture Foundation of Freeport,produced discouraging results. The thirdstudy is by Worldwide Protein BahamasLtd., using imported spat of Americanand European oysters and hard clams,Mercenaria mercenaria, grown in discharge canals from company shrimpponds on Long Island. Preliminaryresults indicate that fouling by algae andparticulate matter hinder production.Water temperatures were too high for
"Creswell, L. Center for Marine Biotechnology,Harbor Branch Institution, Inc., Florida. Personalcommun., November 1984.I3Haaker, P. California Department of Fish andGame. Personal commun. October 1983.
European oysters and the species maybe abandoned, but growout of Americanoysters and hard clams continues. Inaddition, the company recently obtainedpermission from Bahamian authoritiesto import Manila clams, Iapes japonicus, from the Philippines, for growouttrials l4.
Turks and Caicos
The Smithsonian Institution's MarineSystems Laboratory has recently soughtto determine the culture potential of thetopshell or magpie shell, Cittariumpica. Its life cycle has been closed (Heslinga and Hillmann, 1981), and preliminary results indicate that juvenilesplaced in floating cages can reachmarket size in 12-18 months. Similarresearch has also been carried out in theDominican Republic, Antigua, and St.Vincent15. The Mariculture Team of theMarine Systems Laboratory visited theisland of St. Lucia where they demonstrated topshell culture methods; theseappeared to be easily adapted to localconditions, and the feasibility of establishing a pilot project was indicated 16.
Topshells are widely consumed in theCaribbean; the species has been overfished in many areas and several nationsare contemplating plans to regulate itsfisheries. Mariculture may be a viableoption to increase production since topshells appear to fulfill many of thecriteria for commercial culture.
St. Lucia
In 1983 a joint "Experimental OysterProject" was started on St. Lucia between a local yacht charter company andthe Fisheries Management Unit of theMinistry of Agriculture, Lands, Fisheries, and Cooperatives. ImportedJapanese oyster, Crassostrea gigas, spatwere placed in bamboo and plastic meshrafts at two sites. Unfortunately, the raftsat one site were destroyed but there are
I4Higgs, C. Ministry of Agriculture, Fisheries andLocal Government, Nassau, Bahamas. Personalcommun., October 1984."Bernard, W. L. Marine Systems Laboratory,Smithsonian Institution, Washington, D.C. Personal commun., October 1985.'6Walters, H. D. Ministry of Agriculture, Lands,Fisheries, and Cooperatives, Castries, SI. Lucia.Personal commun., 5 October 1984.
Marine Fisheries Review
plans for a second attempt. In addition,several local private concerns have contacted the Fisheries Management Unitfor advice regarding oyster culturel6.
Panama
Between January and December 1979,two pilot studies were conducted todetermine culture potential of the mangrove oyster. Two experimental farmswere located in the Archipelago deBocas del Toro, using the Cuban culturesystem described earlier in this paper.Preliminary results were encouragingbut the project was not scaled-up to acommercial level 17. University of Panama and University of Delaware scientists are jointly experimenting with commercially important bivalves withencouraging results at the University ofPanama's Centro de Ciencias del Mary Limnologia laboratory on Panama'sPacific coast18.
Colombia
Studies on mangrove oyster biology,preparatory to culture, have been carriedout at the Cienaga Grande de SantaMarta and in the Gulf of Uraba on theCaribbean coast of Colombia. Researchhas concentrated on basic biologicalaspects of the oysters and testing for thebest methods of spat collection andgrowout. Results are very encouraging;it may be possible to raise oysters inclosed suspended baskets to commercialsize (6-8 cm) in about 7 months. Bottom culturing methods are not suitablein the Gulf of Uraba due to the heavysedimentation coming from the RioAtrato (Wedler, 1980; Aguilera, 1984).No plans for commercial scale-up werementioned.
St. Kitts and Nevis
In 1981 the Government of St. Kittsand Nevis and the IDRC tried unsuccessfully to establish a pilot project toculture mussels. The project failed because no suitable local species of mussel
I7Arosemena, D. H. Departamento de Direccionde Recursos Marinos, Panama. Personal commun., 28 September 1984.I8D'Croz, L., and 1. R. Villalaz. Centro de Ciencias del Mar y Limnologia, Facultad de CienciasNaturales y Farmacia, Universidad de Panama.Personal commun., 29 October 1984.
47(4), 1985
could be found. In 1983 the project wasredesigned into a Conch ManagementProgram with a "red algae, Graci/ariaspp., Research Project" as a subcomponent, which continues19•
Nicaragua
In 1976, an oyster culture pilot studywas carried out with assistance fromJapanese scientists, using strung scallopshells separated by PVC spacers andhung from mangrove structures. Spatwas successfully collected and grownduring the dry season (May-December)but failed during the rainy season dueto depressed salinities and high sedimentation in the estuarine site. Oystersgrew well up to 81 mm, but mortalitieswere more than 80 percent. Becausemost of the Nicaraguan coastline whereoysters can be raised has similar estuarine conditions, government officialsare not inclined to conduct further oysterculture experiments2o.
Puerto Rico
Experiments to determine the commercial feasibility of mangrove oysterculture started in December 1972, withthe joint support of the Puerto RicoDepartment of Agriculture and the U.S.National Marine Fisheries Service.These experiments consisted of monitoring growth of collected spat in bags,with frames made of concrete-coatedplywood and plastic sheeting suspendedfrom 3 x 4 m rafts. Oysters reportedlyreached market size in 2 months (Watters and Prinslow, 1975). No commercial oyster culture presently exists inPuerto Rico. The Commonwealth Government encourages such nondestructive uses of its coastal lagoons which arecoming under increasing developmentpressure21.
U.S. Virgin Islands
Since May 1972, the St. Croix Arti-
"Wilkins, R. Department of Agriculture, St. Kittsand Nevis. Personal commun., 2S September1984.2°Martinez Casco, S. Centro de InvestigacionesPesqueras, Instituto Nicaraguense de la Pesca. Personal commun., 4 October 1984.2ITorres, F. Corporation for the Development andAdministration of the Marine Resources, PuertoRico. Personal commun., 7 November 1984.
ficial Upwelling Project on the northshore of the island produced phytoplankton by pumping nutrient-rich seawater from 870 m depth into 100 m2
ponds. This was used to feed oysters,clams, and scallops which grew adequately. A pilot-scale operation wascarried out between October 1976 andOctober 1978 in which Manila clamsproduced in the Project Hatchery weresuccessfully grown to determine yieldsand production costs. Very encouragingresults were also obtained culturingbrine shrimp (Roels et al., 1979). TheProject apparently developed into TheMaritek Corporation in 1980 (SeafoodBusiness Report, 1984), and is presentlyengaged in penaeid shrimp culture in theBahamas22.
Other Candidates
Caribbean Species
Milk Conch
The milk conch, Strombus costatus,although considerably smaller than thequeen conch, is nevertheless of commercial value. Its biology has beenstudied and it has been reared for possible use in the marine aquarium tradeand for extensive mariculture. Hatcherytechniques, predation problems, andoutlook are very similar to those for thequeen conch (Appeldoorn and Ballantine, 1983).
Great White Lucineand Gaudy Asaphis
In many Caribbean areas, low primary productivity generally precludesculturing suspension-feeding mollusks.Recent research has shown two clamspecies, the great white lucine, Codakiaorbicularis, and the gaudy asaphis, Asaphis deflorata, to have possible chemoautotrophic capabilities through a symbiotic relationship with sulphur-fixingbacteria within their gill tissues; hence,they have been suggested as viable mariculture candidates. Both species havebeen reared in the laboratory, andchemical analyses have shown that the
22Higgs, C. Ministry of Agriculture, Fisheries andLocal Government, Nassau, Bahamas. Personalcommun., July 1984.
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great white lucine is relatively high inprotein, carbohydrates, and calories butlow in cholesterol compared with otherclam species (Berg and Alatalo, 1982).It has also been speculated that industrial sulfide wastes may be adaptedto a mariculture system involving theseclams (Berg and Alatalo, 1984). Nopilot or commercial projects to rearthese clams are planned 23.
Cephalopods
Many cephalopod species are important research subjects in neuroscience,environmental toxicology, learning behavior, and other areas. The highlydeveloped giant axon of squids, for example, is used in numerous models ofvisual experimentation (Hanlon andForsythe, In press). In addition, fisheries biologists have recently begunusing cultured cephalopods for life-cycleanalyses (Hanlon, In press). The advantages of laboratory-cultured cephalopodsto the researcher include the consistentavailability of experimental animals ofknown species, age, sex, and environmental background (Hanlon and Forsythe, In press). In the Western Atlantic, cephalopods are presently beingcultured on an experimental scale at theMarine Biomedical Institute of the University of Texas Medical Branch,Galveston, Tex.24, and at the Centro deInvestigaciones Cientificas de la Universidad de Oriente, on Margarita Islandoff northeastern Venezuela (Robaina,1983).
Roger Hanlon of the Texas MarineBiomedical Institute reports24 that thelife cycles of several Octopus and Loligospecies have been closed, and extensiveinformation pertinent to their potentialcommercial mariculture has been accumulated. He further reports receivingrequests from people interested in culturing cephalopods commercially in thesouthern Caribbean.
The only known commercial cultureoperation of cephalopods is in Japan,where Octopus vulgaris is reared; production in recent years was about 50 tannually (Boletzky and Hanlon, 1983).However, "the outlook for future com-
23Berg, C. 1. Woods Hole Oceanographic Institution. Personal commun., 7 November 1984.24R. Hanlon, Marine Biomedical Institute, Texas.Personal commun., 5 November 1984.
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mercial culture of cephalopods is unpredictable, because it is predominantly an economic consideration. Whenand if the capture fishery cannot meetthe market demand, culture will receiveemphasis" (Hanlon, In press). Finally,cephalopods may have much potentialin the aquarium trade, because of thespectacular color changes of manyspecies and because they can be easilymaintained in aquariums (Hanlon, Inpress).
ScallopsScallops of the family Pectinidae are
the basis of several important commercial fisheries around the world. Commercial scallop farming in Japan hasbeen very successful, and scientists elsewhere are trying to adapt Japanese techniques to their own countries (Wood,1978). Pilot research continues worldwide, and in Latin America, Peru(Wolff, 1984), and Mexico (Kimbrough,1983) have recently reported commercial culture operations. Those, however,are on the Pacific coast; no pilot- orcommercial-scale scallop culture projectis known in the Caribbean. Research oninduced reproduction and larval rearingis being carried on in Venezuela25. Berg(1984) recently suggested scallop cultureas having potential in Bermuda.
Pearl Oysters
Pearl oysters have been successfullycultured for many years in Japan,Republic of Korea, China, Australia, Indonesia, the Philippines, and othercountries, and the techniques are wellknown. Berg (1984), in reviewing theculture potential of Bermudian bivalves,mentioned that there appears to be noreason why these techniques could notbe successfully applied to Atlantic pearloysters, Pinctada spp. On Mexico'sPacific coast the pearl oyster, P mazatlanica, has been experimentally cultured26. No information is available onpresent or recent research on pearloyster culture in the Caribbean, although in Venezuela aspects of itsbiology have been studied to some ex-
"Padron, M. Universidad de Oriente. Boca deRio, Isla de Margarita, Venezuela. Personal commun., 5 December 1985.26Diaz, G. 1. J. 1969. Cultivo experimental demadreperla, Pinc/aOO maza/tanica, en la Bahia deLa Paz, Baja Calif., Mex., (mimeogr.) 12 p.
tent (Martinez, 1971), and a researchproposal is currently being evaluated27•
Pen Shells
The culture potential of pen shells,Pinna carnea and Atrina rigida, wasalso reviewed by Berg (1984), who concluded that they seem to have poorpotential because of the possibly longplanktonic development which wouldmake larval rearing difficult. He alsomentioned that they may be suitable asan additional species in a polyculturesystem, since they seem to invest littleenergy developing viscera and shell andtherefore might grow very fast. Penshells are very valuable in several Caribbean countries, and in Mexico, wherethey command higher prices thanshrimp and as high as abalone (aboutU.S.$IO.OO/kg as of early 1985)28. Ashellfish hatchery to produce larvae ofcommercially important bivalves, including several species of oysters,clams, scallops, and pen shells, recently started operating in Bahia Kino, in theGulf of California (O'Sullivan, 1984).No commercial or research projects areknown in the Caribbean.
Exotic SpeciesGreen Mussels
A species from the Indo-Pacific, thegreen mussel, Perna viridis, is considered to be a good candidate for introduction into the Caribbean. Presentlythere are over 5,000 ha under its culturein Thailand and the Philippines. Research at the Harbor Branch Institutionin Florida has shown considerablepotential for this species' culture inlocalized Caribbean areas where primary productivity is sufficient to support filter-feeding bivalves29.
Giant Clams
Species of giant clams, Tridacnaspp., are being intensively studied atvarious institutions in the Philippines,Australia, Micronesia, and Californiafor possible commercial culture. These
"Robaina, G. Universidad de Oriente, Boca deRio, Isla de Margarita, Venezuela. Personal commun., 4 December 1985.28Reyes, C. Instituto Tecnologico y de EstudiosSuperiores de Monterrey, Guaymas, Sonora, Mex.Personal commun., March 1985.2·Creswell, L. Center for Marine Biotechnology,Harbor Branch Institution, R. Personal commun.,November 1984.
Marine Fisheries Review
clams have a thin layer of tissue in theirmantle where zooxanthellae live andprovide food (Munro and Heslinga,1983). This aspect of their biology isunique and actually is a strong plus forthe species' introduction to the Caribbean, particularly in areas of low productivity that cannot support filterfeeding bivalves such as oysters ormussels. Cultured tridacnids from Palaumay soon be introduced to the island ofGuadaloupe, pending approval of importpermits30. However, as with any otherexotic species, the possible ecologicalconsequences of introducing any newspecies into a new environment mustfirst be carefully considered, as isstrongly urged by Munro and Heslinga(1983).
Discussion and Conclusions
Molluscan mariculture is nonexistentin several Caribbean countries; in a fewit is at an experimental or incipient stageof development while only a handful ofcommercial molluscan culture operations presently exist anywhere in thearea. Three species of bivalves (American and mangrove oysters and rockmussels) are raised in semi-intensivefacilities; one gastropod, the queenconch, is raised in extensive culture(mass-reared in hatcheries and releasedto augment natural populations).
Techniques for oyster and mussel culture are very similar, involving wild spatcollection on different substrates (i.e.,from mangrove branches to syntheticropes) and growout with the collectorssuspended from floating rafts or woodenstockades. These techniques haveproven successful in some Caribbeancountries and have potential in severalothers. Queen conch culture involvesmass-rearing juveniles in hatcheries andfield growout to commercial size; thefirst has had limited success so far whilethe second remains unproven.
Introduction of exotic molluscanspecies into the Caribbean must be carefully considered. Several native species,particularly filter-feeding bivalves, appear suitable and should be exhaustively studied before considering any introduction. An exception may be that of
30Heslinga, G. Micronesian Mariculture Demonstration Center, Palau. Personal commun.,February 1985.
47(4), 1985
exotic species with phototrophic capabilities, such as giant clams, which maybe suitable for introduction in thoseareas where the naturally low primaryproductivity precludes the culture offilter-feeding species. However, muchcare should be exercised if these or anyother exotic species is introduced intothe Caribbean, and guidelines such asthose recommended by the InternationalCouncil for the Exploration of the Seashould be followed (ICES, 1972).
Several problems still hamper Caribbean molluscan culture. These includeinsufficient biological information onpotential candidate species, dependencyon wild spat, lack of economic information, few trained technicians, inadequatemarketing channels, low primary productivity in many areas, and pollutionand public health considerations.
The FAa Species Identification Sheetsfor Fishery Purposes of the WesternCentral Atlantic (Fischer, 1978) list 37bivalves and 24 gastropods which arelarge, edible, and common enough toserve as human food. Several of thesespecies of oysters, clams, scallops, arks,mussels, and pen shells, as well asothers not included in the FAa sheets,may meet the criteria for culture candidates proposed by Bardach et al. (1972)and Webber and Riordan (1975).
However, adequate biological information on which to judge the feasibility of culture projects for many speciesis lacking, particularly concerningreproductive aspects (spawning season,larval stage requirements) and growthrates and factors which affect it. Research in progress at several Caribbeaninstitutions will help alleviate this. However, economic support for researchsometimes may end prematurely, as inthe case of the queen conch, whosebiology and culture potential (includinghatchery techniques) was intensively investigated at several institutions for afew years, until funding was discontinued. UNDP/FAO has advocated establishment of a Caribbean RegionalAquaculture Center, which has alsobeen strongly supported by countries inthe region and may be approved and implemented in the near future3l.
"Choudhury, P. C. Fishery Resources and Environment Division, FAO, Rome. Personal commun., 28 September 1984.
To depend on natural spat settlementas a source of seed for commercialgrowout is generally not advisable because of the wide fluctuations in spatabundance and settlement due to bioticand abiotic factors. Cuba, for example,has reported such problems for theiroyster culture operations, and is investigating controlled reproduction andoyster larval maintenance as well as construction of a hatchery. Other countriessuch as Mexico, Jamaica, Panama, andVenezuela are operating or planninghatcheries and/or are involved in activeresearch for this purpose.
Economic problems also hamperdevelopment of Caribbean molluscanculture (as well as culture of most othermarine species), particularly the distribution channels and availability of theproducts. Acceptability of seafood is nota problem: Most Caribbean islands andcountries bordering the Caribbean havehigh rates of seafood consumption (>20g/person per day), and in some of thesecountries fish constitutes almost 20 percent of the total protein source (Olsenet aI., 1984). Adequate distributionchannels and product availability problems must be viewed in terms of theadded technological and economic burden involved in processing and marketing highly perishable products in placeswhere refrigeration may be unavailableor inadequate (May, 1978). That inadequate marketing channels are a problemtoday is exemplified by the Venezuelanoyster culture operations discussedpreviously.
Another problem is pollution and related public health considerations. Sincethe Caribbean is generally a developingregion with relatively modest industrialization and urbanization, water pollutionfrom land-produced wastes has notreached the alarming levels of more industrialized regions. However, there arelocalized areas where marine pollutionfrom industrial, domestic, and agricultural wastes, and from oil productionand transport, is a problem (Rodriguez,1981), and this is likely to become aproblem in other areas as industrialization and urbanization occur.
Estuarine areas are particularly affected by pollution and urban development. Besides being highly productivenursery grounds for many commerciallyimportant animals, clean estuaries are
9
also needed for molluscan culture.Public health aspects of pollution mustbe especially considered because the occurrence of human viruses in mollusksfrom waters lightly to moderately polluted is well documented (Vaughn andLandry, 1984) and because informationon the extent of pollution in the Caribbean (i.e., from sewage) is very limited(Rodriguez, 1981).
Finally, the harvesting of culturedmollusks may often have to be restrictedowing either to periodic red tide outbreaks, as in the case of the Venezuelanmussel culture operations discussed, orto ciguatera poisoning, which may occur in populations of topshells of certain areas (Olsen et al., 1984).
In conclusion, molluscan mariculturein the Caribbean has a long way to goto partially augment catches from traditional capture fisheries. It is doubtfulthat it can soon achieve the productionper unit area obtained in other parts ofthe world because it is still in its infancyand many of the problems remain unresolved. However, current and plannedresearch are encouraging.
Acknowledgments
This report is based on a search of theavailable literature and on personal communications with researchers, fisheriesofficers, and administrators involved inor knowledgeable about aquacultureactivities throughout the Caribbean,whose contributions we thankfully acknowledge. A shortened version of thismanuscript was presented at the November 1984 MEXUS-GULF IV and Gulfand Caribbean Fisheries Institute 37thAnnual Meeting in Cancun, Mexico.
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