FAU Institutional Repository
http://purl.fcla.edu/fau/fauir
This paper was submitted by the faculty of FAU’s Harbor Branch Oceanographic Institute.
Notice: © 1998 Kluwer. This manuscript is an author version with the final publication available and may be cited as: Tucker, J. W., Jr. (1998). Marine fish culture. Boston: Kluwer Academic Publishers.
MARINE FISH CULTURE
by
John W. Tucker, Jr., Ph.D. Harbor Branch Oceanographic Institution and Florida Institute of Technology, Melbourne
KLUWER ACADEMIC PUBLISHERS Boston I Dordrecht I London Distributors for North, Central and South America: Kluwer Academic Publishers I 01 Philip Drive Assinippi Park Norwell, Massachusetts 02061 USA Telephone (781) 871-6600 Fax (781) 871-6528 E-Mail
Distributors for all other countries: Kluwer Academic Publishers Group Distribution Centre Post Office Box 322 3300 AH Dordrecht, THE NETHERLANDS Telephone 31 78 6392 392 Fax 31 78 6546 474 E-Mail
'' Electronic Services
Library of Congress Cataloging-in-Publication Data
Tucker, John W., 1948- Marine fish culture I by John W. Tucker, Jr. p. em. Includes bibliographical references (p. ) and index. ISBN 0-412-07151-7 (alk. paper) 1. Marine fishes. 2. Fish-culture. I. Title. SH163.T835 1998 639.3'2--dc21 98-42062 CIP
Copyright © 1998 by Kluwer Academic Publishers
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, mechanical, photo copying, recording, or otherwise, without the prior written permission of the publisher, Kluwer Academic Publishers, I 0 I Philip Drive, Assinippi Park, Norwell, Massachusetts 02061
Printed on acid-free paper.
Printed in the United States of America CONTENTS
Acknowledgments ...... ix
The Author ...... • ...... • ...... X 1 Introduction ...... Purposes ...... 1 Recent status ...... 1 Organization of the book ...... 12 Overview ...... 12 Summary ...... 16 2 Characteristics of Marine Fish ...... 17 Reproduction ...... 17 Development ...... 19 Critical periods ...... 32 Growth ...... 33 Wild vs hatchery fish ...... 33 Summary ...... 34 3 The Rearing Environment ...... 49 Temperature ...... 50 Salinity ...... 55 Solids ...... 59 Light...... 59 Color and texture of hatchery tanks and fittings ...... 66 Green-water culture ...... 67 Structured habitat ...... 67 Disturbances ...... 67 Microbial environment ...... 68 Water chemistry ...... 69 Waste products ...... 72 Gases ...... 81 Gas-related disorders ...... 90 Effects of seawater collection and usage ...... 96 Toxic metals ...... 97 Pesticides and other pollutants ...... 122 Toxic construction materials ...... 142 Toxicants accidentally introduced by the culturist or the building ..... 144 Therapeutic compounds ...... 145 Contaminant effects on beneficial bacteria ...... 145 Testing and electronic monitoring of water quality ...... 145 Predators ...... 145 Noxious microorganisms ...... 146 Environmental problems caused by aquacullure ...... 147 Summary ...... 148
4 Water Sources ...... 149 Criteria ...... 149 Major types ...... 150 Summary ...... 152
5 Water Treatment ...... 155 Requirements ...... 155 Materials ...... 155 Treatment options ...... 156 System design ...... 169 System monitoring and control ...... 172 Environmental considerations ...... 174 Summary ...... 174 6 Culture Units ...... 175 Considerations in choosing culture units ...... 175 Characteristics of culture units ...... 175 Applications of culture units ...... 191 Hatchery design ...... 208 Summary ...... 210 7 Obtaining Fish for Stocking ...... 211 Stock from the wild ...... 211 Stock from the hatchery ...... 211 Spermatogenesis (sperm formation) ...... 232 Oogenesis (egg formation) ...... 232 Oocyte maturation ...... 233 Endocrine control of oocyte maturation and ovulation ...... 237 Induced ovulation ...... 238 Timing and egg quality ...... 257 Artificial fertilization ...... 265 Care of eggs ...... 267 Storage of gametes ...... 269 Natural ovulation ...... 270 Care of broodfish ...... 289 Egg collection ...... 290 Induced vs natural ovulation ...... 290 Broodfish adaptability ...... 291 Examples ...... 291 Genetic considerations ...... 295 Hybridization ...... 296 Sex control ...... 296 Summary ...... 298
vi 8 Nutrition of Larval Fish ...... 299 Feeding criteria ...... 299 Choice and culture of foods ...... 307 General feeding practices ...... 336 Specific feeding practices ...... 352 General methods used in our hatchery ...... 372 Industrial-scale larval food processing in Italian hatcheries ...... 373 Summary ...... 374 9 Nutrition of Juvenile and Adult Fish ...... 375 Requirements and components ...... 375 Broodstock nutrition ...... 407 Nutritional disorders ...... 408 Environmental considerations ...... 411 Feed studies ...... 411 Suggested feed formu las ...... 460 Making and storing feeds ...... 461 Feeding methods ...... 464 Summary ...... 467
10 Energetics ...... 469 Energy budget components and influencing factors ...... 469 Specific examples ...... 472 Energy budgets for farms ...... 473 Summary ...... 473
11 Fish Health ...... 481 Disease prevention ...... 481 Therapeutic agents ...... 486 Biological disease agents and their control ...... 491 Neoplasias ...... 491 Viruses ...... _. _ ...... 491 Bacteria ...... _ ... _ ...... 495 Fungi ...... 500 Protozoans ...... 50 I Coelenterates ...... 504 Free-living flatworms ...... 504 Parasitic tlatworms ...... 504 Cestodes ...... 506 Nematodes ...... 506 A, anthocephalans ...... _ ..... _ ...... _ . .... 506 Crustaceans ...... _ ...... 506 Leeches ...... 507 Lampreys ...... 507 Other pathological conditions ...... 508 Genetic disorders ...... 508
vii Environmentally-induced disorders ...... 508 Overexposure to therapeutic chemicals ...... 510 Nutritional disorders ...... 511 Anaphylactic shock...... 513 Gas bladder disorders ...... 513 Susceptibility ...... 514 Environmental and human health considerations ...... 518 Summary ...... 520
12 Handling and Transporting Marine Fish ...... 521 Sensitivity of different stages ...... 521 Handling marine fish ...... 521 Grading marine fish ...... 525 Transporting marine fish ...... 526 . Summary ...... 532
13 Culture of Established and Potential Species-Food Fish . . . . . 533 Status ...... 533 Family accounts ...... 533 Environmental consideratious ...... 571 Summary ...... 571
14 Culture of Established and Potential Species-Bait Fish ...... 575 Family accounts ...... 575 Environmental considerations ...... 576 Summary ...... 576
15 Culture of Established and Potential Species-Ornamental Fish . 578 Family accounts ...... 578 Environmental considerations ...... 586 Summary ...... 586
16 The Future of Marine Fish Culture ...... 587 Obtaining fish for stocking ...... 587 Environment, water treatment, culture units ...... 587 Nutrition ...... 588 Health ...... 588
Literature Cited ...... 589
Glossary ...... 719 Appendix-Conversion Factors ...... 727
Species Index, by Family ...... 731
viii 1 INTRODUCTION
Why culture marine fish?
PURPOSES Cultured marine fish are used mostly for food, stock enhancement, bait, or display in aquariums. They also are used in research to determine their requirements (e.g., physical, chemical, dietary needs; interactions with other organisms) to aid managers of wild or captive fish. Development of eggs and larvae is studied and documented to elucidate evolutionary and taxonomic relationships and to enable identification of captured early life stages. Species such as the sheepshead minnow are used in bioassays to determine the effects of pesticides and other potential pollutants.
RECENT STATUS In many areas of the world the demand for seafood exceeds the total potential catch from exploited and unexploited resources {e.g. the Caribbean region, Tucker and Jory, 1991). In 1993, 97.8% of the world production of Atlantic salmon came from farms, but only 7.2% of Pacific salmon were cultured (FAO, 1995). Fish culture has been successfi.d with many species in many countries (Tabies 1-1 to 1-5); however, the largest operations arc with fish such as carps, tilapias, and salmonids, which are relatively easy to produce in hatcheries. Although in the past marine fish were raised mainly from wild juveniles (e.g. eels, milkfish, yellowtail), hatche1y production is increasing. Some examples for which hatchery technology is well established are milkfish and Asian sea bass (barramundi) in southeast Asia, red sea bream in Japan, European sea bass and gilthead sea bream in the Mediterranean, and red drum in the United States. Because of greater complexity, not all aspects of technology are as advanced as for freshwater fish culture, but recent breakthroughs make marine fish culture a promising field. Total world aquaculture production of fish in 1993 was 11,284,725 mt (metric tonnes), including 85.1 o/o freshwater, 11.4% diadromous, and 3.5% marine species (FAO, 1998). In 1995, the proportions were 86.1 o/o, 10.2%, and 3.7% of 14,823,526 mt, and in 1996, 86.2%, 10.0%, and 3.8% of 16,664,491 mt (Table 1-1). Tables 1-2 through 1-4 give 1996 production data for all fish and countries surveyed by FAO (1998) , except freshwater fish raised in fresh water. Table 1-5 lists the major species or groups of farmed diadromous and marine fish with the countries producing the most in 1996. Tilapias were the main freshwater fish raised in brackish water (Table 1-2). Salmonoids, eels, milkfish, and ayu were the main diadromous fish raised in fresh water (Tables 1-3, 1- 5). Milkfish were by far the dominant diadromous fish in brackishwater culture, followed by salm onoids, barramundi, and eels; mainly salmon ids were raised in salt water. Of marine fish, mainly mullets and sea breams were raised in fresh water (Tables 1-4, 1-5). Mostly groupers, mullets, sea breams, Asian red snappers, and European sea bass were reared in brackish water. The main marine fish raised in salt water were yellowtail, red sea bream, hirame, puffers, sea basses (serranids), and other jacks in the Far East and gilthead sea bream, sea basses (Dicmtrarclms spp.), and turbot in Europe and the Mediterranean region. In 1995, about 95.9% of cultured freshwater fish were produced by 10 countries (Table 1-l): China (74.1%, nearly all carps), India (11.7%, nearly all cyprinids), Indonesia (2.4o/o, mostly carps and tilapias), Bangladesh (2.4%, a variety), United States (1.6%, nearly all channel catfish), Thailand ( 1.3% , mostly tilapias, Java barb, clariid catfishes), Viet Nam (0.9%, a variety), Philippines (0.6%, nearly all tilapias), Taiwan (0.5%, mostly tilapias), and Myanmar (0.5%, nearly all rohu). 2 Introduction
Table 1-l. Aquaculture production of freshwater, diadromous, and marine fish by country during 1996 (FAO, 1998, with permission from the Food and Agricultural Organization of the United Nations).
Cotmtry Freshwater fish Diadromous fish Marine fish (mt) (rnt) (mt)
Albania 73 e zo• Algeria 244 e n• 36. Argentina 10 1.242 Armenia 1,630. 1.430 . Australia 33 10,348 2,0\3 Austria 810 2.140 Azerbaijan 488c Bangladesh 340,828 Belarus 4,746 e 3e Belgium 300 546 Belize 287 Bhutan 30. Bolivia 80 300 Brazil 40,000 8oo• Brunei Darussalam 20 72 Bulgaria 1.900. 585 Burundi to• Cambodia 9,000 e Cameroon 55 e Canada 51 ,826 Central African Republic 3so• Chile 199,085 168 China (PRC) 10,651,754 147.316 240,592 Colombia 20.263 4,506 Congo Democratic Republic 7so• Congo Republic 5 Costa Rica 4,115 136 Cote d'lvoire 3so• Croatia 2,071 314 252 Cuba 27,987 Cyprus 105 348 Czech Republic 17,378 822 Denmark 41,428 Dominica zc Dominican Republic 343 300 20 Ecuador 330 35 Egypt 53,302 22.535 El Salvador 217 Estonia 61 211 Ethiopia 85 e Faeroe Islands 17,584 Fiji 122 Finland 57 17,605 France 10,143 54,405 2,728 French Polynesia 10 Gabon 3 Gambia 4 Germany 12,168 25,289 Ghana s5o • Greece 716 2.520 26,390 Guam 168 . 25• s• Guatemala 1,990 Guinea 4 Guyana t5o • Hondura~ 232 e Hong Kong 3.395 144 4,561 Hungary 8.076 4 Iceland 3,686 India 1,680,895 e Indonesia 343,000 160,120 11,300 Iran 27,914 ].900 Iraq 3,000 e Marine Fish Culture 3
Table 1-1. (Continued)
Country Freshwater fish Diadromous fi sh Marine fish (mt) (mt) (mt) Ireland 15,875 30 Israel 14,562 1,075 1,931 Italy 6,660 43,982 10,550 Jamaica 3,020 e Japan 15.170 65,142 247,822 Jordan 181 Kazakhstan 1,154 e Kenya 500 105 Kiribati 2" so• D.P. Republic of Korea 10,000 e 2.ooo• Republic of Korea 17,3 54 4,797 11 ,384 Kuwait 90 Kyrgyzstan 152. 9e Laos 14,400 e Latvia 379 I Lebanon 350 Lesoth o IO e 4e Libya 100 . Lithuania 1,537 Macedonia 341 570 Madagascar 2,650 Malawi 235. Malaysia 19,206 6,000 2,844 Mali 60. Malta 1.552 Martinique 13 10 Mauritius 71 16 Mayote I Me~i co 22,358 2,706 Moldova Republic 695. Morocco 750 153 e 1,225. Mozambique 37. Myanmar 68,123 e Namibia 5 • Nepal 10,649 Netherlands 1.200 3.050 New Zealand 3,500 Nicaragua 5 Niger 2 1. Nigeria 17,932 12 Norway 323,893 635 Pakistan 18,000 Panama 177 Papua New Guinea 10 . 8• Paraguay 350. Peru 88 1,148 Philippines 80,287 148,372 703 Poland 21.900 5,800 Portugal 2 1,412 867 Puerto Rico 68 Qatar Romania 13,520 e 380. Russian Federation 47,870 3,770 120 Rwanda so• Saudi Arabia 2.872 Senegal 53 Sierra Leone 30• Singapore 88 266 378 Slovakia 329 625 Slovenia 212 532 75 South Africa 110 790 Spain 160 25,975 6,944 Sri Lanka 3,500 e 4 Introduction
Table 1-l. (Continued)
Country Freshwater fish Diadromous fish Marine fish (mt) (mt) (nu)
St. Lucia l Sudan 1,000 Suriname I " Swaziland 88" Sweden 6.436 Switzerland 35 1.126 Syria 6,355 Taiwan (ROC) 69,343 92,014 13,505 Tajikistan 93 Tanzania 2oo• Thailand 188.3XO 3,750 800 Togo 21 Trinidad & Tobago 5' Tunisia 379 158 < 804 Turkey 780 I X,703 11.530 Turkmenistan 190 Uganda 210 Ukraine 32,195 107 157 United Kingdom 133 100,022 United States 230.853 41,855 Uruguay 6 12 Uzbekistan 5,006 Venezuela 2.400 235 VietNam 130.000' Yugoslavia 1.600. 5 Zambia 4,769 Zimbabwe 70. too•
Totals 14,365,816 1,67.\754 624.921
Total. all fish raised in all types of water in 1996 16,664.49 1
• indicates an estimate, in tables in this chapter. Table 1-2. World aquaculture production of freshwater fish in brack,;h ami salt water during 1996 (FAO, 1998, with permission from the Food and Agticultural Organization of the United Nations).
Scientific name Common name I 996 produclion (mt)
In brackish wain Oreoclzromis mossambicus Mozambiqul! tilapia 31,600 Oreoclzromis niloticus Nile tilapia 28,574 Oreochromis spp. (tilapias) 7.820 Cyprinus carpio Conunon carp 2,175 Micropterus sa/moides Largemouth ba~s 600 Chrysiclzthys nigrodigitatus (bagrid catfish) !55 Oreochromis spilurus 70 Oreochromis aureus Blue tilapia 40 93 Eleotridae (slccpas) 4 Barbodes gonionotus Java barb Osteichthyes (hony fishes) 45, 184
Total. freshwater fish raised in brackish water in 1996 11(>.182
In salt water Oreochromis spp. (tilapia.s) 3 Osteichthyes (bony fishes) 2
Total, freshwater fish raised in salt water in 1996
In this book. when listed in tables, general common nam~s arc in parenthcsc.< . In tables in this chapter, a superscript numher indkate.< that the amount is for a year hdorc 1996. Marine Fish Culture 5
Table 1-3. World aquaculture production of diadromous fish in fresh, brackish, and salt water during 1996 (FAO, 1998, with permission from the Food and Agricultural Organization of the United Nations). Scientific name Conunon name 1996 production (mt)
In fr~sh water Oncorh ynchus mykiss Rainbow trout 277,439 Anguilla japonica Japanese eel 177,520 Chanos chanos Milkfish 39.672 Anguilla spp. ("freshwater" eels) 29.065 Salmo spp. (salmons, trouts) 25,703 Plecoglossus a/r ive/is Ayu 10,007 Anguilla anguilla European eel 7,351 Sa/mo rnma Brown trout 7,057 Lares calcarifer Barramundi/Asian sea bass 4,763 Marone chrysops x M. scnari/is Hybrid striped bass 3,848 Coregonus spp. (whitefishes, ciscoes) 2,218 Acipenseridae (sturgeons) 903 Salvelimts f onrinalis Brook trout 568 Salmo solar Atlantic salmon 302 Oncorhynchus rshawyrscha Chinook salmon 200 Acipenser baerii Siberian sturgeon 172 Sa/velinus spp. (chars) 100 Coregonus lavarerus Po wan 93 Sa/velinus a/pinus Arctic char 90 Anguilla rosrrara American eel 38 95 Oncorhynchus kisurch Coho salmon 27 Lares niloricus Nile perch 12 Thymallus rhymallus Grayling 4 Acipenser rurhenrts Sterlet I Huso huso Beluga I
Total. diadromous fish raised in fresh water in 1996 587.116
In brackish water Chonos chanos Milkfish 324,828 Oncorh ynchus mykiss Rainbow trout 14.935 Lares calcarifer Barramundi/Asian sea bass 10,233 Salmo salar Atlantic salmon 7,647 Anguilla anguilla European eel 665 Salve litws a/pinus Arctic char 541 Anguilla spp. ("freshwater" eels) 350 Cenrropomus undecimalis Conm10n snook 300 Anguilla australis Shortfin eel 201 Anguilla japonica Japanese eel 93 Coregonus lavarellls Po wan 89 Macquaria ambigua Golden perch 8 Macculloche/la peelii Murray perch I Total, diadromous fish raised in brackish water in 1996 359,891
In salt water Salmo salar Atlantic salmon 547.694 Oncorh ynchus mykiss Rainbow trout 87,544 Oncorh ync hus kisutch Coho salmon 76,178 Oncorhynchus tshawytscha Chinook salmon 12.044 Salmo spp. (salmons, trouts) 1,330 Lates calcarifer Barramundi/A sian sea bass 888 Salmo tnma Brown trout 468 Anguilla anguilla European eel 401 Salvelimts spp. (chars) 200
Total , diadromous fish raised in salt water in 1996 726,747
Total. diadromous fish raised in all types of water in 1996 1,673,754 6 Introduction
Table 1-4. World aquaculture production of marine fish in fresh, brackish, and salt water during 1996 (FAO, 1998, with permission from the Food and Agricultural Organization of the United Nations).
Scientific name Common name 1996 production (mt) In fresh water Mugil cephalus Striped mullet 4,397 Sparidae (sea breams, porgies) 516 Dicentrarchus /abrax European sea bass 21 Mugilidae (mullets) 7 Osteichthyes (bony fishes) 1.150
Total, marine fish raised in fresh water in 1996 6,091
In brackish water Siniperca chuatsi Chinese perch 58.437 Mugil cephalus Striped mullet 24,660 Mugilidae (mullets) 11,676 Spams auratus Gilthead sea bream 8,895 Acanthopagrus schlegeli Black sea bream 3,037 Epinephelus spp. (groupers) 2,178 Lutjanus argentimaculatus Asian red snapper 2,007 Dicentrarchus labrax European sea bass 1,838 Sparidae (sea breams, porgies) 1,506 Epinephelus tauvina Greasy grouper 837 Serranidae (groupers, sea basses) 559 Pagrus major Red sea bream 559 Decapturus spp. (scads) 148 Lutjanidae (snappers) 121 Osteichthyes (bony fish c.•) 1,776
Total, marine fish raised in brackish water in 1996 118,234
In salt water Seriola quinqueradiata Yellowtail !45,889 Pagrus major Red sea bream 77,319 Sparus auratus Gilthead sea bream 23,832 Dicentrnrchus labrax European sea bass 19,231 Paralichthys o/ivaceus Hiramc 16,553 Tetraodontidae (puffers) 5,552 Dicentrarchus spp. (sea basses) 5,382 Trachurus japonicus Japanese horse mackerel 3,869 Scophthalmus maximus Turbot 2,588 Trachurus spp. (jack & horse mackerels) 2,343 Scorpaenidae (scorpionfishes) 2,036 Thunnus maccoyii Southem bluefin tuna 2,013 Epinephelus areola/us Squaretail grouper 750 Lutjanus argentimaculatus Asian red snapper 690 Epinephelus spp. (groupers) 407 Sparidae (sea breams, porgies) 368 Epinephe/us tauvina Greasy grouper 360 Trach inotus blochii Snubnosc pompano 325., Lutjarws russe/lii Russell's snapper 300 Lateolabrax japonicus Japanese sea bass 266 Rlwbdosargus sarba Silver sea bream 240 Mugilidae (mullets) 225 Pleuronectiformes (flatfishes) 218 Gadus morhua Atlantic cod 198 Cantherines spp. (filefishes) 148 94 Pagms pagrus Red porgy . 130 94 Diplodus sargus White sea bream 122 Acanthopagms berda River bream 90 Mugil cephalus Striped mullet 82 Lutjanus spp. (snappers) 80 Thmmus thyrmus Northern blucfin tuna 77 ------~------
Marine Fish Culture 7
Table 1-4. (Continued)
Scient i fie name Common name 1996 production (mt)
Evynnis japonica Crimson sea bream 52 94 Serranidae (groupers, sea basses) 36 Epinephelus akaara Redspotted grouper 30 95 Solea solea Conm10n sole 29 Decapterus spp. (scads) 20 Pleurogrammus azot!IIS Atka mackerel 19 Acanthopagrus schlegeli Black sea bream 18 Diplod•ts spp. (sea breams) 15 Rachycentron canadum Cobia 13 Sciaenops oce/latus Red drum 10 Stephanolepis cirrhifer Thread-sailfin filefish 7 Sciaenidae (drums) 2 Siganus spp. (rabbitfishes) 2 Seriola dumerili Greater amberjack I Dentex dentex ConmJOn dentex 1 Siganus callaliCIIintus P.:arly spinefoot I Osteichthyes (bony fishes) 189,342 Total, marine fish raised in salt wat.:r in 1996 500,596
Total, marine fish raised in all types of water in 19% 624,921
About 95.6% of cultured diadromous fish were produced by 20 countries (Table 1-l): Norway (19.4%, mostly Atlantic salmon, some rainbow trout), Chile (11.9%, salmons, rainbow trout), Indonesia (9.6%, nearly all milkfish), Philippines (8.9%, nearly all milkfish), China (8.8%, eels and others), United Kingdom (6.0%, mostly Adantic salmon, some rainbow trout), Taiwan (5. 5%, milkfish, eels, barramundi, others), Japan (3.9%, mostly eels, coho salmon, rainbow trout, ayu), France (3.2%, mostly rainbow trout), Canada (3 .1 %, salmons, rainbow trout), Italy (2.6%, rainbow trout, eels, sturgeon), Denmark (2.5%, mainly rainbow trout and eels), United States (2.5%, mostly rainbow trout, Adantic salmon, hybrid striped bass), Spain (1.6%, mostly rainbow trout, some Atlantic salmon and eels), Germany ( 1.5%, mostly rainbow trout, some brown trout), Finland (1.0%, nearly all rainbow trout), Faeroe Islands (1.0%, mostly Atlantic salmon, some rainbow trout), Turkey (I. I%, salmon ids), Ireland (0.9%, Atlantic salmon and rainbow trout), and Australia (0.6%, Atlantic salmon, rainbow trout, barramundi, eels), with most of the remainder raised in other Asian countries. About 97.4% of cultured marine fish were produced by 12 countries (Table 1-1): Japan (39.6%, mostly yellowtail, red sea bream, hirame, puffers), China (38.5%, Chinese perch and others), Greece (4.2%, mostly gilthead sea bream, European sea bass, striped mullet), Egypt (3.6%, striped mullet, gilthead sea bream, European sea bass), Taiwan (2.2%, mostly barramundi, black sea bream, striped mullet, groupers, red sea bream, other sea breams), Turkey (1.8%, gilthead sea bream , sea basses), Republic of Korea (1.8%, mostly hirame, scorpion fishes , Japanese sea bass), Indonesia ( 1.8%, mullets), Italy ( 1.7%, European sea bass, gilthead sea bream, striped mullet), Spain ( 1.1 o/o, mostly gilthead sea bream, turbot, European sea bass), Hong Kong (0.7%, mostly striped mullet, groupers, snappers, silver sea bream), and Malaysia (0.4%, mostly eels, barramundi, Asian red snappers, greasy groupers). More regional information can be obtained from publications such as: Liao ( 1988), Joseph (1990), and Proceedings of the First, Second, Third, and later Asian Fisheries Forums (Maclean ct al., 1986; Hirano and Hanyu, 1990; Chou et al., 1994), Guerrero and Garcia (1991), and Lacanilao et al. (1994) for Asia; Davy (1990) and Yoshimura et al. (1996a) for Japan; Guerrero ( 1996) and Ri ce and DeVera (I 998) for the Philippines; AI-Thobaity and James (1994) for Saudi Arabia; Kissil (1996) for Israel; Barnabe (1990a) for Europe; Flos et al. (1990) and Larrazabal (1992) for Spain; Aiken (1990) and Needham (1990) for 8
Table 1-5. Aquaculture production of major diadromous and marine fish by the highest-producing countries during 1996 (FAO, 1998, with permission from the Food and Agricultural Organization of the United Nations). Numbers in parentheses are metric tonnes for the year.
Scientific name Common name First Second Third Founh
Diadromous Acipenseridae-sturgeons Acipenser baeri Siberian sturgeon France (160) Uruguay (12) Acipe11Ser ruthenus Sterlet Ukraine (1) Huso huso Beluga Ukraine (l) Acipenseridae (sturgeons) Russian Federation (600) Italy (280) Estonia (17) Ukraine (5) Anguillidae-"freshwater" eels Anguilla anguilla European eel Italy (3000) Netherlands (2800) Denmark (900) Greece (584) Anguilla australis Shonfin eel Australia (201) Anguilla japonica Japanese eel China (147 ,316) Taiwan (25,063) Malaysia (3635) Republic of Korea (I 599) Anguilla rostrata American eel Dominican Republic (38)~5 Anguilla spp. ("freshwater" eels) Japan (28,595) Indonesia (820) Chanidae-milklish C/umos chanos Milkfish Indonesia (157.600) Philippines (148.372) Taiwan (58,453 ) Kiribati (50)e Osmeridae--smelts Plecoglossus a/rive/is Ayu Japan (977 5) Taiwan (232) Salmonidae-salmons, trouts Coregonus lavaretus Po wan Finland (92) Czech Republic (90) Coregonus spp. (whitefishes. ciscoes) Armenia (1150)" Russian Federation (970) Macedonia (70) Switzerland (26) Oncorhynchus kisurch Coho salmon Chile (66,988) Japan (840 1) Canada (756) United States (33) Oncorhynchus mykiss Rainbow trout Chile (54.429) France (50,625) Italy (40.702) Denmark (40,278) Oncorh ynchus lshawyrscha Chinook salmon Canada (8403) New Zealand (3500) Chile (341) Salmo salar Atlantic salmon Norway (301.426) United Kingdom (83.344) Chile (77.327) Canada (36,352) Salmo tnata Brown trout Germany (2529) France (2 118) Russian Federation (2200) United Kingdom (345) Salmo spp. (salmons, trouts) Turkey (18,51 0) Japan (4788) Greece (1927) Canada (I 000) Salvelinus a/pinus Arctic char Iceland (541) France (90) Salvelinus fontinalis Brook trout France ( 450) Czech Republic (82) Slovakia (25) Slovenia & U.K. (5) Salvelinus spp. (chars) Norway (200) Sweden (100) 17zymallus tlzymal/us Grayling France (2) Slovenia (2) Centropomidae-snooks Cemropomus undecimalis Common snook Dominican Republic (300) L.ates calcarifer Barramundi Taiwan (6981) Thailand (3750) Malaysia (2365) Indonesia (] 700) Lates niloticus Nile perch Kigeria (12) Percichthyidae-temperate basses i'v!OI·one chrysops x M. saxatilis Hybrid striped bass United States (3561) Israel (287) i'r!accul/ochella peelii Murray perch Australia (I) Macquaria ambigua Golden perch Australia (8)
Marine
Gadidae-cods Gadus morhua Atlantic cod 1\orway (198) Scorpaenidae--scorpionfishes Scorpaenidae (scorpionfishes) Republic of Korea (2036) Hexagrammidae-greenlings Pleurogrammus azonus Atka mackerel Republic of Korea (19) Percichthyidae-temperate basses Dice11trarchus labrax European sea bass Greece (11,662) Italy (3800) France ( 1997) Egypt (977) Dicentrarchus spp. (sea basses) Turkey (5210) Croatia (172) Lateolabrax japonicus Japanese sea bass Republic of Korea (266) Serranidae-groupers, sea basses Epinephelus akaara Redspotted grouper Hong Kong (30)95 Epinephelus areolatus Squaretail grouper Hong Kong (750) Epinephelus tauvina Greasy grouper Malaysia (837) Hong Kong (360) Epinephelus spp. (groupers) Taiwan (1883) Thailand (600) Singapore (93) Republic of Korea (9) Siniperca chuatsi Chinese perch China (58,437) Serranidae (groupers, sea basses) Philippines (595)
Pomatomidae-bluefishes Pomatomus sa/latrix Bluefish Tunisia (29)92 10
Table 1-5. (Continued)
Scientific name Common name First Second Third Fourth
Rachycentridae-cobia Rachycentron canadum Cobia Taiwan (13) Carangidae-jacks Decapturus spp. (scads) Taiwan (168) Seriola dumeri/i Greater amberjack Spain (1) Seriola quinqueradiata Yellowtail Japan (145,773) Republic of Korea (116) Trachinotus blochii Snubnose pompano Hong Kong (325)95 Trachurus japonicus Japanese horse mackerel Japan (3869) Trachurus spp. (jack & horse mackerels) Japan (2343) Lutjanidae-snappers Lmjanus argentimaculatus Asian red snapper Malaysia (2007) Hong Kong (690) Lu tjanus spp. (snappers) Singapore (80) Lutjanidae (snappers) Taiwan (121) Lurjanus russellii Russell's snapper Hong Kong (300) Sparidae-sea breams, porgies Acanrhopagrus berda River bream Kuwait (90) Acanthopagrus schlegeli Black sea bream Taiwan (3055) Japan (80)94 Dent ex dent ex Common dentex Spain (1) Diplodus sargus White sea bream Greece (122) Diplodus spp. (sea breams) Cyprus (15) Evytmis japonica Crimson sea bream Japan (52) 94 Pagrus major Red sea bream Japan (77,092) Taiwan (759) Republic of Korea (27) Pagrus pagrus Red porgy Greece (130) 94 Rhabdosargus sarba Si Iver sea bream Hong Kong (240) Sparus auratus Gilthead sea bream Greece (13.799) Turkey (6320) Spain (3818) Italy (3650) Sparidae (sea breams, porgies) Taiwan (2363) Republic of Korea (16) Cyprus (11 ) Sciaenidae-drums Sciaenops ocellatus Red drum Martinique (1 0) Sciaenidae (drums) Taiwan (2) Mugilidae-muUets liza ramada Thinlip mullet Tunisia (20)92 Mugil cephalus Striped mullet Egypt (20,101) Italy (3100) Taiwan (2321) Hong Kong (1561) Mugilidae (mullets) Indonesia (11 ,300) Thailand (200) Ukraine (157) Spain (125) Siganidae-rabbittishes Siganus canaliculatus Pearly spinefoot Qatar (1) Siganus spp. (rabbitfishes) Philippines (2) Scombridae-mackerels Thunnus rnaccoyii Southern bluefin tuna Australia (2013) Thunnus th ynnus Northern bluefin tuna Spain (77) Bothidae-lefteye flounders Paralichthys olivaceus Hirame Republic of Korea (8861) Japan (7692) Scophthalmus maximus Turbot Spain (2189) France (225) Portugal (144) Ireland (30) Soleidae-soles Solea solea Common sole Spain (23) Portugal (6) Pleuronectiformes-Oattishes Pleuronectiformes (flatfishes) Chile (168) Russian Federation (50) Balistidae-leatherjackets Cantherines spp. (filefishes) Japan (148)94 Stephanolepis cirrhifer Thread-sailfin filefish Republic of Korea (7) Tetraodontidae-puffers Tetraodontidae (puffers) Japan (5552) MisceUaneous bony fishes Osteichthyes (bony fishes) China (182,155) Japan (5501) Taiwan (2820) Hong Kong (660) e estimate. A superscript number indicates the year of the last reJX>rted value, which is given. 12 Introduction
Canada; Conte (1990) for California; NRC (1992) for the United States; Hargreaves and Alston (1991) and Tucker and Jory (1991) for the Caribbean; Benetti et al. (1994, 1995a) for Ecuador; O'Sullivan (1990, 1992) for Australia; and Hecht et al. (I 992) for South Africa. Current news is provided by Aquaculture Europe (magazine), Aquaculture Magazine, Fish Farming International (newspaper), f-ish Farming News (newspaper), World Aquaculture (magazine), and others. Scientific information is published mainly in aquaculture journals such as Aquaculture, Aquaculture International, Aquaculture Nutrition, Aquaculture Research, Aquacultural Engineering, International Journal of Aquaculture and Fisheries Technology, Journal of Applied Aquaculture, Journal of Aquaculture in the Tropics, Journal of the World Aquaculture Society, Northern Aquaculture, Progressive Fish Culturist, Suisanzoshoku (Aquaculture), but also in regional aquaculture journals, fish biology journals (e.g., Asian Fisheries Science, Bulletin of the Japane~e Society of Scientific Fisheries--now Fisheries Science), and symposium proceedings. Development of marine fish culture, at least in the United States, has been slow. In the U.S., only a few freshwater food fish (trout, salmon, channel catfish, and hybrid striped bass), and ornamental fish (e.g., cichlids, goldfish, guppies, platys) are raised in large numbers. Red drum have been raised in small numbers for food, and clown fish, gobies, and dottybacks support a small ornamental marine fish industry. Research has been done on several marine species that have good potential, but actual farming of these seems more likely in other countries. This book describes important aspects of existing technology and identifies sources of additional information.
ORGANIZATION OF THE BOOK This book is written from the viewpoint of an ex-larval fish taxonomist/physiologist who over the last 27 yr has intensively reared some of the more difficult marine fish (Table 1-6) on research and pilot scales. During hundreds of rearing studies, procedures for spawning, larviculture, and grow-out have been developed. It is not intended to be a manual for marine fish culture, but a combination reference and textbook. Most of the chapters are literature reviews combined with original data and information gained by experience. Extent of the reviews depends on both the importance and the volume of information available. Fairly extensive reviews arc included on reproduction and nutrition, with more lim ired ones on characteristics of marine fish, water quality, and fish health. Factors limiting the development of commcrci;~l marine fish farming are in five main categories: water quality (chapters 3, 4, and 5), reproduction (chapter 7), nutrition (chapters 8, 9, and 1 0), health management (chapter I I), and economics (briefly discussed in this chapter). Water quality is mainly an engineering problem and depends on a good source and an appropriate treatment system . Especially during the last 15 yr, much progress has been made in the areas of reproduction and nutrition of marine fish. f-ish health depends mostly on water quality and nutrition. Economics is based on supply and demand balances. Chapter 2 describes characteristics of fish relative to rearing them. Chapter 6 describes containers used for rearing fish. Chapter 12 describes handling and shipping methods for live fish. Chapters 13, 14 , and 15 give a synopsis for representative species. Chapter 16 briefly speculates on the future of marine fish culture. For the sake of completeness and to facilitate comparisons, euryhalinc tilapias and diadromous fish such as sturgeons, eels, and salmons are included. Because much already has been written on their culture, their tre;~tment here is brief.
OVERVIEW Before getting into the details, here arc some general comments about biological, engineering, siting, economic, social, environmental, and genetic bcrors relevant to culture of saltwater fish. Marine Fish Culture 13
Table 1-6. Fish reared by the author.
Scientific name Common name Stages First year
Family Anguillidae-"freshwater" eels Anguilla rostrata American eel Larva to early juvenile 1975
Family Clupeidae-herrings Brevoorlia patronus Gulf menhaden Egg to adult 1984
Family Engraulidae-anchovies Anchoa mitchil/i Bay anchovy Egg to mid-stage larva 1978
Family Cyprinodontidae-killilishes Cyprinodon variegatus Sheepshcad minnow Egg to adult to egg 1971
Family l'oeciliidac-livcbcarers Gambusia affinis Western mosqu.itofish Larva to adult to larva 1971 Poecilia reticulata Guppy Larva to adult to larva 1964 Xiphoplwrus varia/us Variable platyfish Larva to adult to larva 1984
Family Centropomidae-snooks Centropomus undecimalis Common snook Egg to adult 1983 Cenlropomus pecti!wtus Tarpon snook Larva to adult 1983 Laces calcarifer Barramund.i, Asian sea bass Egg to late juvenile 1985
Family Percichthyidae-tempcrdte basses Morone clzrysops x M. saxatilis White bass x striped bass Early juvenile to adult 1988
Family Scrranidac-groupcrs, sea basses Centropristis striata Black sea bass Egg to early juvenile 1979 Epinephelus fulvus x E. guttatus Coney x red h.ind Egg to mid-stage larva 1994 Epinephelus fulvus x P. furcifer Coney x creole-fish Egg to mid-stage larva 1994 Epinephelus fuscogulfatus Brown marbled grouper Egg to mid-stage larva 1991 Epinephelus gurta/us Red hind Egg to late juvenile 1994 Epinephelus morio x C. striata Red grouper x black sea bass Egg to early larva 1994 Epinephelus slriatus Nassau grouper Egg to adult to egg 1987 Paranthias furcifer Creole-fish Egg to mid-stage larva 1994 Plectropomus areola/us Squaretail corahrout Egg to mid-stage larva 1991 Serranus subligarius Belted sandfish Egg to adult to egg 1991
Family Coryphaenidae-dolphins Coryphaena hippurus Dolphin Egg to juvenile 1994
Family Lutjanidae-snappers Ocyun;s chrysurus Yellowtail snapper Egg to mid-stage larva 1988
Family Sparidae-sea breams, porgies Archosargus probatocephalus Sheepshead Egg to adult 1984
Family Sciaenidae-drums Cynoscion nebulosus Spotted sea trout Egg to adult 1985 Leiostomus xantlzurus Spot Egg to adult 1984 Memicirrhus americanus Southern k.ingfish Egg to early juvenile 1979 Pogonias cromis Black drum Early juvenile to adult 1994 Sciaenops ocel/alus Red drum Early juvenile to adult 1991
Family Cichlidae-cichlids Oreoc/zromis mossambicus Mor.1mbique tilapia Egg to adult to egg 1985 14 Introduction
Table 1-6. (Continued)
Scientific name Common name Stages First year
Family Pomacentridae--damseUishes Abudefduf saxatilis Sergeant major Egg to adult 1994 Stegastes leucosticws Beaugregory Egg to mid-stage larva 1993
Family Mugilidae-mullets Mugil cephalus Striped mullet Egg to adult 1995 Mugil curema White mullet Egg to late juvenile 1972
Family Labridae-wrasses Halichoeres bivittatus Slippery dick Egg to early larva 1993 l.aclmolaimus ma.ximus Hagfish Egg to early larva 1986 Thalassoma bifasciatum Bluehead wrasse Egg to early larva 1993
Family Ephippidae-spadefishes Chaetodipterus faber Atlantic spadefish Egg to mid-stage juvenile 1980
Family Bothidae-lefteye lloundcrs Scophthalmus aquosus Windowpane Egg to adult 1972
Balistidae-leatherjackets Batistes vetula Queen triggerfish Egg to late larva 1992
Biological Considerations The best candidates for aquaculture will grow fast and efficently and will survive well, despite the stresses of captivity. Sufficient seed will be readily available. From a biological point of view, there are four main areas in which available information and knowledge presently limit the potential of marine fish culture: obtaining eggs, maintaining a suitable environment, ensuring good nutrition, and controlling disease. The most common bottleneck is lack of sufficient survival of larvae to the juvenile stage, which would result from good environment, nutrition, and health. Juvenile survival usually is not a problem. Many marine species can be reared in fresh water after their osmoregulatory capabilities have become adequate. Snook, barramundi, shecpshead, red drum, spotted seatrout, southern flounder, striped mullet, and others can survive and grow well in fresh water at some point before they become juveniles (within 3-5 weeks). But they have to be in salt water for those first few weeks, and salt water adds many more problems for the culturist. Engineering Considerations Weather can be especially hazardous to a coastal facility (e.g., hurricanes, northeasters, erosion). Salt brings other problems, notably corrosion of most metals used in construction. Some water quality problems (e.g., higher ammonia toxicity) can be more important in salt water. Maintenance of high water quality often is the most critical factor, but just as often the most difficult and frequently the least understood and most underestimated. Siting Considerations Problems can occur in site selection, broodstock
Environmental Considerations Noxious exotic species can be introduced by aquaculture, unless strict precautions arc taken. In many cases, the fish eventually do escape, for example, because of an accident or ponds overflowing during a flood. Some introduced fish have eaten or displaced more desirable indigenous species or othe1wise become a nuisance, for example, Nile perch introduced in certain African lakes and grass carp in the U.S. Tilapia are well established in several states of the U.S. (Courtenay et al., 1991). Exotic species should be raised only in closed, recirculating systems or in such a way that the effluent cannot carry exotic fish or exotic pathogens into local habitats where they could survive. When fish are farmed, space and resources sometimes are taken away from natural populations~.g., natural water bodies are filled, fenced, or diked; and cages also take up space. In coastal environments, where productivity typically is high, humans cannot compete well with nature. Monoculrure practices often result in more of a particular species than would grow nawrally in a given area, but, if left alone, overall productivity in natllre usually is higher (Turner, 1977). Alternative approaches include strong offshore cages or land-based grow-out using recirculating water systems wheneve r possible to minimize effluent disposal problems. Addition and enhanced cycling of nutrients can increase plant biomass and productivity, sometimes resulting in problems such as phytoplankton blooms or ponds clogged with weeds. Physical and chemical effects of solid or dissolved wastes can reduce habitat quality near culture facilities. Excess sediment and nutrients have to be removed from tank or pond wastewater before it is discharged, and lagoon treatment seems the most practical way (Chen et al., 1997). Introduction of exotic diseases and parasites (e.g., Thompson, 1990), proliferation of rare pre-existing pathogens, and development of resistant or new strains (Aiken, 1991; Edwards, 1991, and others in the same issue) can occur. Crowded aquaculture conditions often enhance growth of viruses, bacteria, and parasites. Changes in composition, abundance, and antibiotic resistance of bacteria have been documented at fish farms. Further discussion of the environmental impact of aquaculture can be found in publications such as: Herke (1976, 1977, 1978), Gowen and Bradbury (1987), Smith and Pestano-Smith (1985), Stickney (1988), Anonymous ( 1989) , Lam (1990), De Pauw and Joyce (1992), and Pillay (1992). Genetic Considerations In fish farming, artificial selection is practiced: because survival is higher than in nature, the ratio of offspring to parents is much higher. In addition, relatively few broodfish are employed. If large numbers of reared fish are released, genetic diversity in the natural population could be reduced (Bartley ct al., 1992; Garcia-Vazquez et al., 1995).
SUMMARY A species good for commercial aquaculture is likely to have the following characteristics: high value, easy to get eggs, larvae, or juveniles of, easy to rear, easy and cheap to feed, can be crowded, grows fast and efficiently. Some ideas that might contribute to success in rearing marine fish are: Although the basic needs of some species are very well known, marine fish culture is still very much an art. Manuals might help, but there is no substitute for experience and good judgement. Consider well the needs of the fish in choosing the site, in designing the facility, and in operating the facility. Be sure the water quality is suitable before making a major committment. Plan for all possible contingencies. Look cl osely at rearing conditions and at how the fish develop and behave, especially in response to the conditions or changes in them . Use the past experience of others and yourself at all stages. Evaluate, reevaluate, and revise rearing strategies whenever possible. Keep what is good, but make sure it is repeatable. Eventually, the art will become somewhat more of a science.