Recent Advances and New Species in Aquaculture

Recent Advances and New Species in Aquaculture Recent Advances and New Species in Aquaculture Edited by Ravi K. Foted...

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Recent Advances and New Species in Aquaculture

Recent Advances and New Species in Aquaculture Edited by

Ravi K. Fotedar Department of Environment and Agriculture School of Science Curtin University Perth, Western Australia

Bruce F. Phillips Department of Environment and Agriculture School of Science Curtin University Perth, Western Australia

A John Wiley & Sons, Ltd., Publication

This edition first published 2011 © 2011 by Blackwell Publishing Ltd Blackwell Publishing was acquired by John Wiley & Sons in February 2007. Blackwell’s publishing program has been merged with Wiley’s global Scientific, Technical and Medical business to form Wiley-Blackwell. Registered office: John Wiley & Sons, Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial offices: 9600 Garsington Road, Oxford, OX4 2DQ, UK The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK 2121 State Avenue, Ames, Iowa 50014-8300, USA For details of our global editorial offices, for customer services and for information about how to apply for permission to reuse the copyright material in this book please see our website at www.wiley.com/ wiley-blackwell. The right of the author to be identified as the author of this work has been asserted in accordance with the UK Copyright, Designs and Patents Act 1988. 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, electronic, mechanical, photocopying, recording or otherwise, except as permitted by the UK Copyright, Designs and Patents Act 1988, without the prior permission of the publisher. Designations used by companies to distinguish their products are often claimed as trademarks. All brand names and product names used in this book are trade names, service marks, trademarks or registered trademarks of their respective owners. The publisher is not associated with any product or vendor mentioned in this book. This publication is designed to provide accurate and authoritative information in regard to the subject matter covered. It is sold on the understanding that the publisher is not engaged in rendering professional services. If professional advice or other expert assistance is required, the services of a competent professional should be sought. Library of Congress Cataloging-in-Publication Data Recent advances and new species in aquaculture / edited by Ravi K. Fotedar, Bruce F. Phillips. p. ; cm. Includes bibliographical references and index. ISBN 978-1-4051-7664-4 (hardcover : alk. paper) 1. Aquaculture. I. Fotedar, Ravi K. II. Phillips, Bruce F. [DNLM: 1. Aquaculture–methods. 2. Fishes. 3. Palinuridae. SH 135] SH135.R428 2011 639.8–dc22 2010052329 A catalogue record for this book is available from the British Library. This book is published in the following electronic formats: ePDF 9781444341744; Wiley Online Library 9781444341775; ePub 9781444341751; Mobi 9781444341768 Set in 10/12 pt Times by Toppan Best-set Premedia Limited, Hong Kong

1

2011

Contents

Contributors Abbreviations and acronyms Preface Acknowledgements 1

Recent Developments Ravi Fotedar, Gopal Krishna, Uras Tantulo, Iain Mcgregor and Bruce Phillips 1.1 1.2 1.3 1.4 1.5 1.6 1.7

Introduction Disease resistance in aquaculture systems vis-à-vis breeding strategy Freshwater ornamental aquaculture – an industry view from Western Australia Use of immunostimulants as feed additives Alternative sites for aquaculture Future directions References

2 A Global Review of Spiny Lobster Aquaculture Bruce Phillips and Hirokazi Matsuda 2.1 2.2 2.3 2.4 2.5 2.6 3

Introduction Broodstock management Larval rearing Raising wild-caught pueruli and juveniles Future developments References

Slipper Lobsters Manambrakat Vijayakumaran and Edakkepravan V. Radhakrishnan 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 3.9

Introduction Biology Aquaculture potential Marketing Slipper lobster culture initiatives Hatchery production of seeds Factors influencing phyllosoma growth and survival Hatching and larval rearing in Thenus sp. Growth of juvenile slipper lobsters

ix xi xiv xv 1 1 1 5 10 11 16 16 22 22 28 30 46 65 68 85 85 87 88 89 90 90 95 99 101

vi

Contents

3.10 3.11 3.12 4

Culture of Thenus sp. Conclusions References

103 109 110

Mud Crab Aquaculture Brian D. Paterson and David L. Mann

115

4.1 4.2 4.3 4.4 4.5 4.6 5

Penaeid Prawns Ngo Van Hai, Ravi Fotedar and Nguyen Van Hao 5.1 5.2 5.3 5.4 5.5

6

Introduction Achievements Challenges Prospective/future outlook References

Cobia Culture Ravi Fotedar and Huynh Minh Sang 6.1 6.2 6.3 6.4 6.5 6.6 6.7 6.8 6.9 6.10 6.11

7

Introduction Portunid crab aquaculture Biology and life cycle Technology development Future developments References

Introduction Morphology Distribution Biological characteristics Nutritional requirement of cobia Hatchery Growout Disease and health management Post-harvest and marketing Challenges and opportunities References

Barramundi Aquaculture Suresh Job 7.1 7.2 7.3 7.4 7.5 7.6 7.7 7.8

Introduction Biology Hatchery production Hatchery culture Growout Nutrition and growth Health management Quality

115 116 119 122 125 131 136 136 137 157 159 161 179 179 179 181 181 183 186 189 191 194 195 196 199 199 200 202 207 211 213 216 221

Contents vii

7.9 7.10 7.11 7.12 8

Abalone Culture Mark Allsopp, Fabiola Lafarga-De la Cruz, Roberto Flores-Aguilar and Ellie Watts 8.1 8.2 8.3 8.4 8.5 8.6

9

11

Introduction The abalone market Abalone production technology Technological developments Future possibilities References

Seaweed Culture with Special Reference to Latin America Julieta Muñoz, Vivek Kumar and Ravi Fotedar 9.1 9.2 9.3 9.4 9.5 9.6 9.7 9.8

10

Sales and marketing Future directions Conclusions References

Introduction Seaweed utilisation Aquaculture Integrated aquaculture Post-harvest: agar extraction Cultivation in Latin America Conclusions References

221 222 224 224 231

231 231 233 245 249 249 252 252 252 254 257 259 261 266 268

Marine Ornamental Fish Culture Suresh Job

277

10.1 10.2 10.3 10.4 10.5 10.6 10.7 10.8

277 281 293 294 300 306 312 313

Introduction Broodstock and eggs Broodstock conditioning Larval culture Juveniles Commercial production Conclusions References

Tilapia Luan Dinh Tran, Trung Van Dinh, Thoa Phu Ngo and Ravi Fotedar

318

11.1 11.2 11.3 11.4 11.5 11.6

318 319 322 325 326 327

Introduction Seed production Culture practices Harvesting and value added products Genetic improvement of tilapia Environment and disease management

viii

Contents

11.7 11.8 11.9 12

13

Marketing of tilapia Conclusion References

327 330 331

Carp Polyculture in India Dilip Kumar

334

12.1 12.2 12.3 12.4 12.5 12.6 12.7 12.8 12.9 12.10

334 335 336 336 339 351 359 362 366 366

Introduction Freshwater aquaculture resources in India Development of aquaculture Commonly cultured species Aquaculture practices/systems Developments in culture practices Culture of pangasius (Pangasianodon hypophthalmus) Freshwater prawn farming Recent developments References

Future Directions Bruce Phillips, Ravi Fotedar, Jane Fewtrell and Simon Longbottom

368

13.1 13.2

368

13.3 13.4 13.5

Introduction Developments in managing the environmental impacts of aquaculture Ecolabelling The future References

Index Colour plate section facing page 48

368 378 381 381

387

Contributors

Mark Allsopp Level 1 Wakatu House 28 Montgomery Square PO Box 440, Nelson, New Zealand Trung Van Dinh Research Institute for Aquaculture No. 1 Dinh Bang, Tu Son, Bac Ninh, Vietnam Jane Fewtrell Department of Environment & Agriculture School of Science Faculty of Science and Engineering Curtin University GPO Box U1987, Perth, WA 6845, Australia Roberto Flores-Aguilar Centro de Investigación y Desarrollo de Recursos y Ambientes Costeros i-mar Universidad de Los Lagos Puerto Montt, Chile Ravi K. Fotedar Department of Environment & Agriculture School of Science Faculty of Science and Engineering Curtin University GPO Box U1987, Perth, WA 6845, Australia Ngo Van Hai Faculty of Agricultural Sciences and Food Technology Tien Giang University 119 Ap Bac St., Ward 5, My Tho City, Tien Giang Province, Vietnam Nguyen Van Hao Research Institute for Aquaculture No. 2 116 Nguyen Dinh Chieu Street, District 1 Ho Chi Minh City, Vietnam

Suresh Job Batavia Coast Maritime Institute Geraldton, WA 6530, Australia Gopal Krishna Central Institute of Fisheries Education Seven Bunglows, Versova Mumbai, 4000061, India Dilip Kumar Central Institute of Fisheries Education Seven Bunglows, Versova Mumbai, 400061, India Vivek Kumar Milne, 103-105 Welshpool Road Welshpool, WA 6106, Australia Fabiola Lafarga-De la Cruz Laboratorio de Genética y Biotecnología Acuícola Departamento de Oceanografía Facultad de Ciencias Naturales y Oceanográficas Centro de Biotecnología, Universidad de Concepción Casilla 160-C, Concepción, Chile Facultad de Ciencias Agronómicas, Universidad de Chile Santa Rosa 11315, La Pintana, Santiago, Chile Simon Longbottom Department of Environment & Agriculture School of Science Faculty of Science and Engineering Curtin University GPO Box U1987, Perth, WA 6845, Australia

x

Contributors

David L. Mann Bribie Island Research Centre Department of Employment, Economic Development and Innovation PO Box 2066, Bribie Island, Qld, Australia

Edakkepravan V. Radhakrishnan Central Marine Fisheries Research Institute Post Box No. 1603 Ernakulam North PO Kochi 682 018, India

Hirokazi Matsuda Mie Prefectural Science and Technology Promotion Center Fisheries Research Division Hamajima, Shima, Mie 517-0404 Japan

Huynh Minh Sang Institute of Oceanography 1 Cau Da Street Nha Trang City Khanh Hoa Province Vietnam

Iain Mcgregor Aquatico and Water Garden World Balcatta, Victoria Park and Armadale Western Australia Australia

Uras Tantulo Agriculture Faculty University of Palangka Raya Jl. Yos Sudarso, Palangka Raya Central Kalimantan, 73111A, Indonesia

Julieta Muñoz Department of Environment & Agriculture School of Science Faculty of Science and Engineering Curtin University GPO Box U1987, Perth, WA 6845, Australia

Luan Dinh Tran Research Institute for Aquaculture No. 1 Dinh Bang, Tu Son, Bac Ninh, Vietnam

Thoa Phu Ngo Research Institute for Aquaculture No. 1 Dinh Bang, Tu Son, Bac Ninh, Vietnam

Manambrakat Vijayakumaran National Institute of Ocean Technology NIOT Campus, Velachery – Tambaram Main Road Narayanapuram, Pallikaranai Chennai 600 100, Tamil Nadu, India

Brian D. Paterson Bribie Island Research Centre Department of Employment, Economic Development and Innovation PO Box 2066, Bribie Island, Qld, Australia

Ellie Watts Aquaculture Research Cawthron Institute 98, Halifax Street East Nelson, 7010, New Zealand

Bruce F. Phillips Department of Environment & Agriculture School of Science Faculty of Science and Engineering Curtin University GPO Box U1987, Perth, WA 6845, Australia

Abbreviations and Acronyms

AA amino acids AFLP amplified fragment length polymorphism AFR & DC Australian Fisheries Research and Development Corporation AR arachidonic acid AWG average weekly gain BLIS bacteriocin-like inhibitory substance BMP best management practices CFU colony forming unit CIFE Central Institute of Fisheries Education (India) CIFRI Central Inland Fisheries Research Institute (India) CL carapace length CMC carboxymethyl cellulose COP code of practices DAP di-ammonium phosphate DGC daily growth coefficient DGI daily growth increment DHA docosahexaenoic acid DHC differential haemocyte counts DO dissolved oxygen dph day(s) post-hatch ECP extracellular products EFA essential fatty acids EIA environmental impact assessment EMP environmental management plans EMS environmental management systems EPA eicosapentaenoic acid ERA ecological risk assessments EST expressed sequence tag FA fatty acid FCE feed conversion efficiency FCR feed conversion ratio FISH fluorescence in situ hybridisation FL fork length FOM final oocyte maturation FRP fibre reinforced plastic GAP good agricultural practice GIFT ‘Genetic Improvement of Farmed Tilapia’ GIS geographical information systems GM genetically modified or genetic modification

xii

Abbreviations and Acronyms

GnRHa GOC GSI HCG HUFA ICAR IHHNV IMC IMTA IP IRR L LC LHRH LM MAE MBV MDS ME mL MOS MoV MPEDA NGO(s) NNV NOEC NPU NPV OFBW ORP OTC OW PAP PCR PER PG PL POF proPO psu RAPD RAS RFLP RGR RT-PCR SDA SEM SGR(s)

gonadotropin-releasing hormone analog groundnut oil cake gonadosomatic index human chorionic gonadotropin high unsaturated fatty acids Indian Council of Agricultural Research infectious hypodermal hematopoietic necrosis virus Indian major carps integrated multi-trophic aquaculture intermoult period internal rate of return litre lipid class luteinising hormone releasing hormone light microscopy microwave-assisted extraction Monodon baculovirus moult-death syndrome metabolisable energy millilitre mannan oligosaccharide Mourilyan Virus Marine Products Export Development Authority (India) non-governmental organisation(s) nervous necrosis virus no-observable-effect-concentration net protein utilisation net present value ovary-free body weight oxidation–reduction potential oxytetracycline ocean water phagocytosis activating protein polymerase chain reaction protein efficiency ratio peptidoglycan post-larvae; phospholipid postovulatory follicles prophenoloxidase practical salinity unit random-amplified polymorphic DNA recirculating aquaculture system restriction fragment length polymorphisms relative growth rates reverse transcriptase-polymerase chain reaction specific dynamic action scanning electron microscopy specific growth rate(s)

Abbreviations and Acronyms

SNT ST TAG TAN TEM THC TL vBGF VER VNN WFC WMD

single nucleotide polymorphism sterol triacylglycerol total ammonia-nitrogen transmission electron microscopy total haemocyte count total length von Bertalanffy growth factor viral encephalopathy and retinopathy viral nervous necrosis World Fish Centre white muscle disease

xiii

Preface

There are many excellent books on aquaculture. However, the stimulus for this book was the absence of information on recent technological developments, new species, recent changes and some global trends, in a form suitable for academic level students. The introduction of new species into aquaculture is critical to the evolution of the global aquaculture industry, particularly as the species that are the basis of the current industry, such as salmon and black tiger prawns, reach maximum levels of production. The past decade has seen a remarkable growth in aquaculture production due to the surge in the development of new technologies and a better understanding in the production biology of new aquaculture species. The book is aimed at aquaculture students, the industry and interested members of the public, particularly in India, Australia, Vietnam and South America. It documents some of the important technological innovations of recent years used in the production technologies for the new species. In addition, the book highlights the increase in production of wellrecognised species such as carp and tilapia which has become possible because of the use of new technological and/or management tools. The book looks into the future by emphasising the need for the research that is required to make these new technologies sustainable. We had planned to include chapters on sea urchins, dolphin fish, composite fish farming in China, sea cucumbers, and some emerging species such as Fugu, mud skippers, cephalopods, southern blue fin tuna and reef-fish. However, we were unable to obtain contributions for these chapters; nevertheless we hope to include them in future editions of this volume. The chapters that are included all take different approaches and styles. This is partly because of the stage of development of aquaculture, but we also decided to keep to the original formats of individual authors as we felt this improved the presentation of the information for the reader.

Acknowledgements

Many people contributed to the development and production of this book. They are not acknowledged individually because of space availability, but all authors wish to thank the many colleagues who assisted them with their contributions. However, we wish to recognise the incredible amount of work carried out by Dr Seema Fotedar in polishing the manuscript for publication. Without Seema’s efforts this book might not have been published. We would also like to thank our postgraduate students who helped us in finding the latest references on the topics.

Fig. 1.1 Two metallic koi or ogons: one gold and one orange (with permission of David Prangell, from his thesis).

Fig. 1.2

Small butterﬂy koi. The larger ﬁsh shows the scale trait of ‘kin gin rin’.

(a)

(b) Fig. 1.3

(a) Long-ﬁnned kohaku. (b) Long-ﬁnned ‘Hi utsuri’.

Fig. 1.4

Long-ﬁnned golden ogon and hariwake showing pectoral ﬁn ray diversity.

Fig. 1.5 Koi with long mouth barbels and extended nostril phenotype.

Fig. 1.6

A typical inland saline water purpose-built pond in Wannamal, Western Australia.

Fig. 2.5

The vertically revolving system (VRR system) designed for Panulirus japonicus phyllosomas.

Aquaculture production 180 160

Indonesia India Taiwan Spain Singapore Philippines Japan Cuba Belize

140 Tonnage

120 100 80 60 40 20 0 1958

1963

1968

1973

1978

1983 1988 Years

1993 1998

2003

Fig. 2.9 Spiny lobster aquaculture production by country. Vietnam is not included because the scale of production is so large compared to the production of these other countries.

Phyllosoma

Nisto Gravid female

Adult Fig. 3.2

Life history of the slipper lobster Thenus orientalis.

(a)

(c)

(b)

(d)

Fig. 3.10 (a) Tail fan necrosis, (b) swollen vent, (c) regurgitated proventriculus and (d) oedematous pleopods in adult Thenus orientalis.

Fig. 6.1

A cobia ﬁngerling.

Fig. 7.5

Greenwater culture of larval barramundi.

Fig. 8.9 Wild H. laevigata, H. laevigata × H. scalaris (yybrid) and H. scalaris found in Western Australia.

Fig. 9.2 Mexico.

(a) Fig. 9.3

Eucheuma isiforme cultivated under experimental conditions at Dzilam de Bravo, Yucatan,

(b)

(c)

Kappaphycus alvarezii color strains cultivated in Mexico. (a) green; (b) red; (c) brown.

Fig. 10.2 substrate.

Yellow-stripe Premnas biaculeatus broodstock with a terracotta ﬂowerpot as a spawning

Fig. 10.3

Male Banggai cardinalﬁsh, Pterapogon kauderni, brooding eggs in its mouth.

Fig. 10.5 fridmani.

The most commonly cultured dottyback species, the Orchid dottyback, Pseudochromis

Fig. 10.6 Male Pseudochromis steenei using a PVC pipe as a spawning den.

Fig. 10.8

Captive-bred P. fridmani juveniles.

Fig. 10.9 Yellow-tail damselﬁsh, Chrysiptera parasema, juveniles shortly after settlement, showing the adult colouration.

Fig. 10.10 Juvenile Amphiprion ocellaris clownﬁsh.

Fig. 10.11 Newly released banggai cardinalﬁsh, Pterapogon kauderni, juveniles.

Fig. 10.12 Juvenile seahorse at the pelagic stage.

Fig. 10.13 Solomon Islands variant of the true black percula anemoneﬁsh, Amphiprion percula.

Fig. 10.14 Ocellaris clownﬁsh displaying partial broken banding on one side.

Fig. 13.1 Recirculating Aquaculture System at Curtin Aquatic Research Laboratory (CARL), Curtin University, Perth, Australia.

1 Recent Developments Ravi Fotedar, Gopal Krishna, Uras Tantulo, Iain Mcgregor and Bruce Phillips

1.1 INTRODUCTION The first decade of the twenty-first century saw a remarkable growth in aquaculture production due to the surge in the development of new technologies and better understanding of the production biology of new aquaculture species. A worldwide interest in the production biology of new candidate species for aquaculture and associated technology is not only deemed to be environmentally friendly, but could also lead to an increase in the productivity of aquaculture. This rise in interest in the subject has led to a gap in the published information, as only a few comprehensive textbooks are available to meet the demand. This chapter highlights the recent developments in biotechnology and the research attempts to extend aquaculture to non-traditional farming sites. The use of biotechnology during breeding strategy has been very impressive and has also been applied to deal with widespread disease issues through molecular genetics and through the use of specialised feed additives, which have a potential to enhance the immune-competence of the cultured species. Captive breeding is playing an increasingly important role, and has been commercialised while producing high-value freshwater ornamentals.

1.2

DISEASE RESISTANCE IN AQUACULTURE SYSTEMS VIS -À-VIS BREEDING STRATEGY

Disease outbreaks are major constraints in any intensive production system. Diseases that remain at a low level of incidence in natural populations may reach epidemic levels in intensive cultivation systems. Intensive management systems in livestock production encourage the unpredictable appearance of new diseases and changes in the characteristics of established diseases (Biggs 1985). If elimination of pathogens or control of culture conditions is difficult, selective breeding for host resistance to the pathogen may be an attractive option for disease control. Host

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

2

Recent Advances and New Species in Aquaculture

resistance should only be considered when (a) the disease causes severe damage, (b) there are no other existing simple, cost-effective control measures, (c) there is demonstrable genetic variation in resistance and (d) this is not coupled with an excessive level of negative associations with other desirable characteristics. The principles and concepts behind breeding programmes are based largely on experiences with plants and terrestrial animals as information from aquatic animals is very limited. With catastrophic diseases, such as white spot syndrome virus (WSSV), which cause mortalities of 98% or more, the frequency of resistance is low and it is suggested that for theoretical reasons single-gene, rather than polygenic, resistance is likely to develop. The low frequency of resistance genes in breeding populations may cause genetic bottlenecks, which will greatly reduce the genetic variation in the populations. In order to maintain the genetic variation the genes from the small numbers of survivors should be introgressed into populations with broader genetic variability. Genetic variation in resistance may be encountered either in the initial base populations or may arise spontaneously due to mutations. Once genetic variation has been detected, the most appropriate breeding methodology will depend on the nature of both the resistance and the disease(s) that are of interest to the producers. Most populations of farmed shrimp have only had a relatively short period to evolve and adapt to intensive cultivated production systems. In India modern intensive shrimp production systems provide almost ideal conditions for the propagation of diseases. The conditions favour epidemics and the appearance of apparently new diseases in intensive shrimp production systems. In Central and South America, Penaeus vannamei was widely devastated by Taura syndrome virus (TSV) in the early 1990s (Brock 1997). Later WSSV appeared in Asia and rapidly devastated the shrimp industry in many parts of the world. Both of these diseases were previously unreported. In Asia, epidemics of white spot and yellow head virus (YHV) have reduced production of various Penaeid shrimp species, including the native species P. monodon and the introduced species P. vannamei. As concepts behind disease control in aquatic animal species have been developed from warm-blooded terrestrial species, the major differences in their environments indicate that transfer of technology from one to the other should be carried out with caution. Warm-blooded terrestrial land animals maintain a relatively constant body temperature, whereas aquatic organisms are ecto-thermal and their body temperature fluctuates with that of the water in which they live. Similarly, the composition of the medium in which land animals live, the air, varies little, with such vital aspects as oxygen and carbon dioxide content relatively constant on a global basis. On the other hand, shrimps face tremendous variability in the environment in which they live, with dramatic changes often occurring abruptly. Stress, which is closely related to the manifestation of disease (Biggs 1985), is often induced by changes in such parameters as temperature, oxygen, salinity and ammonia. Vaccination is a common disease control measure in warm-blooded animals, protecting hundreds of millions of animals from disease and death (NOAH 2002). It is generally accepted that the crustaceans do not possess the capacity to acquire resistance and hence vaccination is not possible, although Witteveldt (2006) has questioned this assumption. In domesticated animal populations simple avoidance of diseases and pests has long been one of the most important means of disease control, with eradication of Newcastle disease in poultry and rinderpest and foot and mouth disease in cattle being well known cases (Biggs 1985). Disease avoidance or eradication is only possible in certain circumstances. Exclusion of diseases has been attempted with some success in shrimp cultivation

Recent Developments 3

(McIntosh 1999; Moss 1999), with various programmes emphasising the use of Specific Pathogen Free (SPF) stock in breeding programmes to minimise spread of diseases (Moss 1999; Moss et al. 2003; Lightner 2005a; Hennig et al. 2005). However, it is not easy to avoid or eradicate diseases in an open-air aquatic growout environment. With the exception of some diseases, such as yellow head virus (YHV) and monodon bacillus virus (MBV), most of the shrimp viruses have spread rapidly from the sites where they were first recognised (Lightner 1996, 2005b; Flegel et al. 2004). The recent epidemic of white spot syndrome indicates how rapidly an epidemic may spread in marine species. First detected in Taiwan in 1992 (Chou et al. 1995), WSSV spread rapidly to most Asian countries (Inouye et al. 1994; Wongteerasupaya et al. 1995; Flegel & Alday-Sanz 1998; Zhan et al. 1998) and by 1996 most shrimp-farming regions in Southeast Asia were affected (Flegel & Alday-Sanz 1998). In the western hemisphere the first outbreak of WSSV appeared in farmed P. vannamei and P. stylirostris in South Carolina (USA) in 1997 and it was associated with 95% of cumulative losses (Lightner 1999). By early 1999, WSSV had spread to farmed P. vannamei in Central America (Jory & Dixon 1999), reaching the Colombian Pacific coast in May of that year. The disease devastated most of the major shrimp-producing areas of the world. Attempts to eradicate or exclude it were mostly unsuccessful. WSSV appeared to have been successfully excluded from a few shrimp-producing regions, particularly the Atlantic coast of South America; however it has recently been reported in the cooler regions of southern Brazil (Anon 2005). It now appears that conditions on the Atlantic coast of South America were in general not conducive to the development of full-scale white spot due to the high water temperatures (Vidal et al. 2001). The absence of a white spot virus epidemic in this area appears to be related to the virus’s inability to replicate at the higher temperature rather than a temperature-mediated response by the shrimp (Reyes et al. 2007). Some areas in South and Southeast Asia may have escaped or have a low incidence of WSSV due to higher water temperatures. Shrimp farmers have in some cases reduced water exchange and appear to have achieved some level of control of WSSV with this practice, which probably both increases water temperatures and also reduces the chances of pathogens entering the ponds. In Thailand the use of specific pathogen free (SPF) stocks and biosecurity measures have reduced WSSV incidence dramatically. In many animals, disease resistance is both innate and acquired. Innate immunity is rapid, non-specific and acts as a first line of defence, while acquired resistance involves antigen-specific responses (Bishop et al. 2002). Shrimps possess an innate immune system that protects them from foreign organisms. Recently Witteveldt (2006) indicated that vaccination of shrimp against WSSV might be possible, which would open the way for the design of new strategies to control WSSV and other invertebrate pathogens. In addition, there may be possibilities to stimulate the immune system and a series of non-specific responses against invading organisms. Genetically controlled behavioural characteristics may also provide resistance to disease: for example genetically controlled hygienic behaviour in bees prevents chalk brood disease (Milne 1983). It is possible that genetic control of cannibalistic behaviour may be involved in providing a measure of resistance to infection (Gitterle et al. 2005). With diseases that are difficult to eradicate, control measures have been developed based on stimulation or enhancement of the natural defence mechanisms of the host organism, including selection for host resistance or tolerance to diseases and modification of the environment so that the disease is not favoured. Genetics-based host resistance is an attractive proposition from the point of view of the grower of improved stock. An advantage of host resistance is the minimal negative impact

4

Recent Advances and New Species in Aquaculture

on the environment. On the other hand, development of genetically based host resistance is often costly and may be impossible to achieve in the absence of useful levels of resistance. Selective breeding programmes inevitably lead to slower progress in other desirable characteristics in the breeding goal. Added to this, disease resistance may be negatively associated with other desirable characteristics. The genetic control of disease resistance in shrimps is not well understood and little research has been done in this area. Shrimps appear to have no acquired immune response, and in this sense they are perhaps somewhere between plants and mammals in their response. Inferences on various aspects of genetic control of disease resistance are mainly drawn from other species, particularly plants and mammals, of which there is a vast stock of knowledge. Selective breeding for disease resistance in plants has a longer history than in mammals. Vertical resistance provides effective immunity, normally through hypersensitivity, and is controlled by a single gene. Horizontal resistance does not provide total immunity but slows the spread of the disease and is controlled by many genes. In a selection programme, the selection protocol itself may affect the type of resistance that is encountered: selection procedures with a limited range of genetic variation and dosages or inoculum pressure that ensure more survivors are likely to lead to uncovering and selection for polygenic resistance, whereas natural selection in the field with larger genetic variation and extremely high mortalities (well over 99%) are likely to uncover single-gene resistance which will normally be dominant. In most of the animals studied, disease resistance is controlled quantitatively by multiple genes and breeding programmes are based on this assumption. However, breeders should not ignore the possibility of single-gene resistance, which has also been observed in animals and humans (Hills 2001). Fjalestad et al. (1993) suggest that in the fish farming environment, resistance to a given pathogen will normally develop slowly. However, resistance to serious pathogens may develop through natural selection in aquaculture populations where the animals have been exposed continuously to the pathogen for only a few generations, as in the case of TSV and with the QX disease in the Sydney rock oyster Saccostrea glomerata (Nell & Hand 2003). In shrimp, which has only recently been bred in captivity, most of the genes that control resistance will probably have come from the original native populations, although their frequency may have been radically altered as populations encountered vastly different conditions. Selection for disease resistance is directly related to its effect on growth and survival: the objective is not disease resistance per se but rather the impact that disease resistance will have on the desired performance characteristics of the selected stock. Diseases can directly affect both growth and survival. Diseases such as TSV and WSSV cause severe damage through mortality, although animals that survive may have reduced growth rates. Other diseases, for example Vibrio, may cause high mortality under some conditions, whilst in other conditions their main effect may be to reduce growth. Until now the main focus in selection for disease resistance in shrimps has been to improve survival in the face of epidemics of diseases such as TSV, which may cause mortalities of 70% or greater, and WSSV with mortalities close to 100%. The white spot case highlights the importance of having a broad genetic base so as to identify sources of resistance: the frequency of resistance genes appears to be very low and there may be sources of resistance that are not included in the initial populations. This case also highlights the difficulties encountered when there is a negative correlation between two or more desired traits. Selection procedures are needed to ensure selected stock will perform well commercially: this normally means having the ability to survive an epidemic.

Recent Developments 5

At present, selection for disease resistance in designed breeding schemes is normally carried out based on survival recorded in controlled challenge tests. In principle, the simplest programme of genetic stock improvement is to choose superior animals as breeders so that as generation succeeds generation the variation in the original population is translated into improved production. This straightforward approach can be guaranteed to work only if certain conditions are met:

• • • • •

the variation must be heritable so that the superior qualities of the parents are passed on to their offspring the qualities designated ‘superior ’ must be easy to recognise so that large numbers of animals can be classified quickly traits under selection must not be correlated in a way which is counterproductive it must be physically convenient to induce the selected individuals to mate and to keep the selected offspring separate from the rest of the population the progress of the selection programme must be carefully monitored to maintain the integrity of the experimental design over many generations.

1.3

FRESHWATER ORNAMENTAL AQUACULTURE – AN INDUSTRY VIEW FROM WESTERN AUSTRALIA

This section is based on a personal communication from Iain Mcgregor (2010), a leading freshwater ornamental aquaculturist in Western Australia. Some of the information also comes from leading magazines. The freshwater ornamental aquaculture industry in Western Australia has many complexities and provides unique challenges for the people working in it. This industry can be seen as typical of ornamental freshwater industries in the developed economies where captive breeding and other forms of technologies are employed. Most species are cultured in field conditions; some are kept in intensive situations such as recirculating systems or aquariums to match optimum requirements. To capitalise on time and space, complementary species are grown together. Aggressive or predatory species may present unique problems such as cannibalism. The greater the number of species in a polyculture situation the greater the complexities involved in successful production. Goldfish are probably the oldest species of ornamental fish and the shape, colour and physical mutations to choose from are mind-boggling, with new phenotypes appearing all the time. Other famous ornamental subjects such as discus fish and guppies have also been extensively developed, with many colour strains now available. These subjects have legions of avid admirers worldwide, to the extent that whole shows are now held for only one species. Japanese coloured carp or koi have a long history of culture for the dual purpose of table and ornamental qualities. Initially this fish was a protein source for people across Europe and Asia. As it spread to remote areas human culture started to mould the genetics of common carp (Cyprinis carpio). It was selected for high growth rates and reduced numbers of scales to make it easier to prepare for consumption. But in rural China and Japan another pressure was to change the destiny of this fish and launch it worldwide. The parts of the country where this happened suffered from extreme winter conditions and the locals were snowed in for extended periods. Fish for food were placed in a pond inside the homes and

6

Recent Advances and New Species in Aquaculture

used to get the snowbound inhabitants through this difficult season. This environment of isolation led to inbred genetics, leading to the mutation of colour. The unusual specimens must have attracted much attention, so that people began to take an interest in the fish beyond their value as a food source. As colours appeared and were mixed, new combinations arose and breeders began to select exclusively for superior specimens. Soon this diversity needed order and a basic grouping of 13 colour varieties was settled on as a rough guide and standard. One of the more amazing of these mutations is the trait that gives the fish a ‘metallic’ look (Fig. 1.1); so completely divergent is this from the brown/grey fish that shape is all they seem to have in common. The lustre of the skin on ‘Hikari muji mono’ and ‘Hikari moyo mono’ seems to glow, almost generating its own light. This intense colour trait has not been tried in all the traditional patterns but only needs a motivated breeder to achieve this. If the kaleidoscope of colour was not enough of a new factor, amazing reflecting scales looking like glitter appeared termed ‘kin gin rin’ (Fig. 1.2), and in a few years of breeding selected individuals, this trait could be expressed in five ways each illuminating the scale with a different facet. When combined with the metallic fish the the effect was very attractive. Long-fin or butterfly koi (Fig. 1.3a, 1.3b) seemed to have originated in Southeast Asia during the 1970s and distributed worldwide a short time after. The long-fin trait expresses itself with fins about twice the length of those in normal carp and some fish show great individuality of fin shape amongst themselves, such as with pectoral fins having each ray longer than the surrounding webbing and appearing ragged – in complete contrast to smooth entire-finned fish (Fig. 1.4).

Fig. 1.1 Two metallic koi or ogons: one gold and one orange (with permission of David Prangell, from his thesis). (Please see plate section for colour version of this figure.)

Recent Developments 7

Fig. 1.2 Small butterfly koi. The larger fish shows the scale trait of ‘kin gin rin’. (Please see plate section for colour version of this figure.)

In less than 150 years of culture, ornamental carp or ‘Nishikigoi’ have graced garden ponds and been collected and exhibited by enthusiasts across the globe. This appreciation of art placed on a fish reaches its peak at the Japan show, a place where fortunes are made or maintained by having the grand champion fish. Other genetic traits are the longer mouth barbels and larger nostrils that extend outside the head cavity (Fig. 1.5). They are also very robust individuals that grow strongly. As butterfly koi became a component of ornamental koi, the governing body was faced with a decision as to whether these fish had a place in the exhibition circuit. The Japanese group, the ZNA, made a decision against recognising the longer-finned fishes. This decision may have reduced their attractiveness to serious hobbyists. However, when a person who had no interest in this aspect was given a choice between short- and long-finned fish for their garden pond they usually chose the latter. These average pond keepers began creating an undeniable market for the illegitimate style. In recent years American koi shows have begun to give butterfly koi a class, and even in Japan, fish farms aim to service this part of the market. The butterfly koi looks set to become more popular than ever, despite traditional Japanese values and tastes in fish. Fish growers are responding to the demand and the genetics of this strain of koi are strengthening as each growing season passes. Higher quality stocks are being produced in greater numbers and each day novice koi keepers enter the market. What future can genetics hold for the appearance of butterfly koi? Soon they will be available in all colours with ‘kin gin rin’ scales. When this is achieved and all varieties are being supplied in large numbers, maybe the next horizon will be xanthic or albino fish with an underlying pattern of the major colours from traditional koi. Can the long-fin phenotype be exaggerated to be extra long, as has been bred into poultry or guinea pigs? Is body shape the next frontier, following down the same road as goldfish? Will some view short stumpy

8

Recent Advances and New Species in Aquaculture

(a)

(b) Fig. 1.3 (a) Long-finned kohaku. (b) Long-finned ‘Hi utsuri’. (Please see plate section for colour version of this figure.)

Recent Developments 9

Fig. 1.4 Long-finned golden ogon and hariwake showing pectoral fin ray diversity. (Please see plate section for colour version of this figure.)

Fig. 1.5 Koi with long mouth barbels and extended nostril phenotype. (Please see plate section for colour version of this figure.)

10

Recent Advances and New Species in Aquaculture

koi as beautiful? The technology for cloning carp currently exists and this procedure may provide a stimulus for new mutations. The possibilities that genetic engineering can offer are bound only by the imagination; gigantism may be the goal, or even salt tolerance! Indeed genetic manipulation may even be able to have a direct influence on where the colour on a fish is placed. As new markets unfold, new trends gain interest; these elegant fishes will continue to flourish. And for the breeders of butterfly koi each growing season holds new promise.

1.4

USE OF IMMUNOSTIMULANTS AS FEED ADDITIVES

Intensification of aquaculture has led to gradual and chronic environmental degradation, loss of biodiversity and eventually loss of productivity. In order to overcome the loss of productivity, the use of chemicals and antibiotics is an easy solution but not without drastic consequences on sustainability and the health of consumers of aquatic products. Recently, the use of environmentally friendly feed additives, namely probiotics and prebiotics, has become popular in the aquaculture industry (Vine et al. 2006; Soltanian et al. 2007). These specialised feed additives act as immunostimulants, which either enhance innate defence responses prior to exposure to a pathogen, or improve survival after the actual infection by pathogens (Bricknell & Dalmo 2005). Research on innate immune systems has revealed new insights into the management and control of diseases in aquaculture (Bachère 2003). It is believed that understanding the immune criteria as enhancement of non-specific defence responses against bacterial and viral injections is the most effective way for sustainable aquaculture production (Chang et al. 1999; Bachère 2003; Chang et al. 2003), but the benefit of immunostimulants is still doubtful in invertebrates (Marques et al. 2006). Immunostimulants fed to animals may not be effective against all diseases (Sakai 1999). However, Bricknell and Dalmo (2005) claimed that a great deal can be done to improve larval survival against bacterial and viral pathogens by the judicious use of immunostimulants. Recently, immunological techniques have identified several distinct types of collagen in P. japonicus (Mizuta et al. 1992), but the genetic control of the production of these techniques has not been investigated (Benzie 1998). Immunostimulants are obtained from various sources such as bacteria, brown and red algae and terrestrial fungi (Bricknell & Dalmo 2005), bacteria from aquatic habitats (Rengpipat et al. 1998) and marine yeast (Sajeevan et al. 2006). Immunostimulants can be divided into several groups, depending on their original sources such as bacteria, algaederived, animal-derived, nutritional factors and hormones or cytokines (Sakai 1999). Probiotics are defined as ‘live microorganisms which when administered in adequate amounts confer a health benefit to the host’ (FAO/WHO 2002) and prebiotics are defined as ‘nondigestible food ingredients that beneficially affect the growth and health of the host’ (Gibson & Roberfroid 1995). According to Kesarcodi-Watson et al. (2008), certain suggested immunostimulants (Itami et al. 1998; Smith et al. 2003) such as peptidoglycan (PG) and lipopolysaccharides can be considered as probiotics. In addition, a number of chemical agents, polysaccharides, plant extracts or some nutritional additives, act as immunostimulants (Sakai 1999; Gannam & Schrock 2001), are adjuncts to vaccination and provide a potential route to reduction of the widespread use of antibiotics (Burrells et al. 2001). Herbal immunostimulants, namely methnolics extracted from five different herbal medicinal plants, were shown to increase P. monodon resistance against viral pathogenesis caused

Recent Developments

11

by WSSV (Citarasu et al. 2006). The use of immunostimulants in penaeid prawn aquaculture is described in Chapter 5.

1.5

ALTERNATIVE SITES FOR AQUACULTURE

In terms of land availability, suitable sites for aquaculture are often not easily and readily available as most of them are expensive and have been occupied by other users (Allan et al. 2001). In addition, acquaculture activities may disturb other activities, or vice versa. In such conditions, aquaculture facilities often have to be built in areas that are essentially not suitable for aquaculture purposes as water and soil quality do not meet the aquaculture requirements. For example, in Asia, aquaculture ponds are built in mangrove areas, which have potential for acid sulphate released from the soils (Pillay 1993). This in turn can slow aquaculture production, as the cultured aquatic species are exposed to low water quality. In Thailand productive rice paddy fields have been used for inland saline prawn culture; the Thai authorities have now banned the practice as it has caused salinisation of agricultural land (Fegan 2001). Inland saline water (ISW) has resulted from anthropogenic activities. It has adversely influenced the agricultural outputs and environment in the USA, Australia, India, China and Israel (Allan et al. 2001). Several actions have been taken for remediation of these problems, including an attempt to use ISW for marine aquaculture (Fig. 1.6). This has been

Fig. 1.6 A typical inland saline water purpose-built pond in Wannamal, Western Australia. (Please see plate section for colour version of this figure.)

12

Recent Advances and New Species in Aquaculture

seen as a remedial approach to reduce the cost of groundwater pumping and create economic opportunities for the farmers in the affected areas (Doupé et al. 2003a,b). In addition, most ISW is suitable for aquaculture purposes as it is located in remote areas where land is cheap and disease-free, so there is potential for integration between aquaculture and agriculture (Gong et al. 2004).

1.5.1

Inland saline water aquaculture in different countries

To date, studies on the use of inland saline water (ISW) as an alternative source for marine culture have been attempted in countries like the USA (Forsberg et al. 1996; McMahon et al. 2001; Treece 2002; Saoud et al. 2003; Gong et al. 2004), India (Rahman et al. 2005; Jain et al. 2006) and Australia (Allan et al. 2001; Fielder et al. 2001; Partridge & Furey 2002; Prangnell & Fotedar 2005, 2006a,b; Partridge et al. 2006, 2008; Tantulo & Fotedar 2006, 2007). If the use of ISW as a medium for marine culture can be proven cost-effective, then it should be considered as an alternative for marine aquaculture. It has some comparative advantages over the coastal areas in terms of cheap land availability, better quarantine capability and freedom from conflict over the same resources (Allan et al. 2001). The use of ISW as a medium for culture of marine fish and prawns can only be successful if the K+ deficiency in ISW can be eliminated, e.g. by fortifying the K+ concentration in ISW from 50 to 100% of K+ concentration in ocean water (OW). This can be achieved by adding KCl in ISW, but high costs might be incurred when this method is used. Another method is by culturing hardy species like black tiger prawn and barramundi (Lates calcarifer) which have already been proven able to withstand the low salinities of ISW. 1.5.1.1 Australia In Australia, most studies have been conducted with the considerations that ISW has caused a negative impact to the physical, social and economic systems of the affected area (Allan et al. 2001). Attempts have been made to minimise these problems by pumping the ground ISW to lower the water table and then storing it in earthen ponds. This has been seen as an alternative source for marine aquaculture purposes (Allan et al. 2001). The main cause of increase in ISW and dry land salinity is the clearing of deep-rooted native plants and replacing them with shallow-rooted grain and pasture crops that have less capability to catch the excess surface rainwater (Walker et al. 1999; George & Coleman 2001) and keep the groundwater table at constant levels. This in turn has increased the saline groundwater table and brought it to the surface. In general, except for low K+ concentration, ionic composition and concentration of ISW is similar to the ocean water (Nulsen 1999). As with ocean water, sodium and chloride are the major ions that determine the salinity of the ISW (Rayment & Higginson 1992) and the osmolality of the haemolymph of the cultured species (Pequeux 1995). Low K+ concentration in ISW occurs due to adsorption of the K+ onto the clay (Allan et al. 2001). Although, K+ concentration only contributes a small part of the total ions making up the ISW, it plays an important role in the functioning of the physiological systems of the aquatic animals (Burton 1995; Shiau & Hsieh 2001). In crustaceans, K+ is very important to activate Na+/K+ ATPase (Skou 1957; Mantel & Farmer 1983), which is responsible for maintaining the ionic imbalance in the haemolymph. Therefore, alteration of Na+ and K+ ratio in the haemolymph may disturb entire physiological functions of the aquatic animals.

Recent Developments 13

As ISW is K+ deficient, most fish or prawns cannot survive in this type of water. Recent studies on prawn (Ingram et al. 2002; Saoud et al. 2003; Prangnell & Fotedar 2006a; Tantulo & Fotedar 2006) and fish culture (Fielder et al. 2001; Partridge & Creeper 2004) in this type of water have reported mortality. An approach to improve the survival rate of the cultured prawns is by fortifying K+ in ISW through the supplementation of KCl (Rahman et al. 2005; Prangnell & Fotedar 2005, 2006a,b; Tantulo & Fotedar 2006; Partridge & Lymbery 2008) and also by adding 5% KCl in fish diets (Gong et al. 2004). The physiological responses of the animals, including survival, growth, ionic and osmoregulation ability, have been analysed following rearing and culture in K+ deficient ISW, K+ fortified ISW and K+ supplemented diet (Gong et al. 2004; Prangnell & Fotedar 2006a; Tantulo & Fotedar 2006). However, as survival and growth rate might not give clear explanations of the effect of K+ deficient ISW on the cultured animals, detailed studies have been conducted on osmo- and iono-regulation of animals cultured in and exposed to ISW (Saoud et al. 2003; Prangnell & Fotedar 2006a; Tantulo & Fotedar 2007). The studies on the effect of K+ deficient ISW on osmo- and iono-regulation of prawns have been conducted on the basis that K+ is the principal intracellular cation in animals (Shiau & Hsieh 2001) and plays important role in creating differing electrical charges between inner and outer membranes (Burton 1995). K+ indirectly affects the haemolymph osmolality through the Na+/K+ ATPase activity. Na+/K+ ATPase activity is a mechanism that establishes the Na+ gradient in prawn haemolymph (Roer & Dillaman 1993). Tantulo and Fotedar (2007) revealed that a decrease in K+ concentration led to increased Na+ concentration in haemolymph of black tiger prawn (Penaeus monodon), which in turn increased the haemolymph osmolality and caused the death of the prawns. This is an indication that K+ is not isolated in its effect on the physiological system of the prawns or fish, but works in conjunction with Na+. Furthermore, the correct ratio of Na+ and K+ is very important for maintaining proper physiological functions of Litopenaeus vannammei (Zhu et al. 2004) and P. monodon (Tantulo 2007). The low K+ concentration in ISW can be increased either by supplementing ISW with KCl (Prangnell & Fotedar 2005, 2006a,b; Tantulo & Fotedar 2006, 2007) or spreading muriate of potash on the bottom of the earthen ponds (Collins et al. 2005; Partridge & Creeper 2004). Following addition of KCl, survival and growth rate of the prawns and finfish in ISW were similar to those cultured in OW (Collins et al. 2005; Prangnell & Fotedar 2005, 2006; Rahman et al. 2005; Tantulo & Fotedar 2006, 2007). In addition, K+ fortification in ISW also increased the osmo-regulation capacity of western king prawns and black tiger prawns and led to higher survival and growth rate of both prawns, similar to those cultured in OW (Prangnell & Fotedar 2005; Tantulo & Fotedar 2006, 2007). Exceptions to the negative effects of K+-deficient ISW have been observed in black tiger prawns (Tantulo & Fotedar 2007) and barramundi juveniles (Jain et al. 2006; Partridge et al. 2008) that were cultured in low salinity ISW of 5 and 15 ppt respectively. As the black tiger juveniles can strongly osmo-regulate their K+ concentratration, they exhibit similar survival and growth rates when cultured in ISW and OW of low salinities. Partridge et al. (2008) revealed that barramundi need more supplementation of the K+ concentration at higher salinity (45 ppt) than the fish reared in salinity close to the isosmotic line. On the other hand, fish or prawns reared at lower salinities may not need supplementation of K+. To date, most marine fish and prawn ISW culture attempts have been on an experimental scale (see Table 1.1 for Australia). It has been reported that some fish such as mulloway (Argyrosomus japonicus) can survive and grow well in K+-deficient ISW (Aquaculture SA 2003; Partridge & Lymbery 2009). However, barramundi died 10 days after stocking in

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Recent Advances and New Species in Aquaculture

Table 1.1 Aquatic species cultured in ISW either on an experimental (E) or commercial (C) scale in Australia. Species

State(s)

Scale

Reference(s)

Dunaliella salina

NT, SA, WA

E, C

Giant clam (Hippopus hippopus) Greenlip abalone (Haliotis laevigata) Pacific oysters (Crassostrea gigas) Sydney rock oysters (Saccostrea glomerata) Trochus (Trochus niloticus) Brine shrimp (Artemia salina)

NT

E

Paust (1999); George & Coleman (2001); McFarlane & Christie (2002); Collins et al. (2005) Lee (1999)

WA

E

Harris et al. (2005)

Vic

E

Ingram et al. (2002)

Vic

E

Ingram et al. (2002)

NT SA, Vic

E E, C

Banan prawns (Penaeus merguiensis) Kuruma prawns (Penaeus Japonicus) Tiger prawns (P. monodon)

Qld

E

Lee (1999) Hutchinson (1999); McFarlane & Christie (2002); Gooley et al. (1999); Gooley & Gavine (2003) Collins et al. (2005)

Vic

E

Ingram et al. (2002)

Western king prawns (Penaeus latisulcatus) Western rock lobster (Panulirus cygnus) Atlantic salmon (Salmo salar) Australian bass (Macquaria novemaculeata) Barramundi (Lates calcarifer)

WA

E

Ingram et al. (2002); Collins & Russell (2003); Doroudi et al. (2003); Collins et al. (2005); Rahman et al. (2005); Tantulo & Fotedar (2006) Prangnell (2006)

WA

E

Tantulo et al. (2005)

Vic Vic

E E

Ingram et al. (2002); Gooley & Gavine (2003) Ingram et al. (2002); Gooley & Gavine (2003)

NSW, WA, SA

E, C

Black bream (Acanthopagrus burcheri)

SA, Vic, WA

E, C

European carp (Cyprinus carpio) Greenback flounder (Rhombosolea tapirina) King George whiting (Sillaginodes punctatus) Mulloway (Argyrosomus hololepidotus)

Vic

E

Fielder & Allan (1997); Allan & Fielder (1999); Hutchinson (1999); Paust (1999); O’Sullivan (2003); Partridge & Creeper (2004); Partridge et al. (2006) Paust (1999); Walker et al. (1999); Ingram et al. (2002); Doupe et al. (2003a,b); Gooley & Gavine (2003) McKinnon et al. (1998)

SA, Vic

E

Hutchinson (1999); Ingram et al. (2002)

SA, WA

E

Hutchinson (1999); Partridge (2001)

NSW, Vic, WA

E, C

Rainbow trout (Onchorhynchus mykiss)

NSW, Vic, WA

E, C

Sand whiting (Sillago ciliate)

Vic

E

Doroudi et al. (2003, 2006); O’Sullivan (2003); Dutney (2004); Flowers & Hutchinson (2004); Partridge et al. (2006) Ingram et al. (2002); McFarlane & Christie (2002); Doupe et al. (2003a,b); Gooley & Gavine (2003); Partridge et al. (2006) Ingram et al. (2002)

NSW, NT, Qld, Vic, WA

Recent Developments 15 Table 1.1 (Continued ) Species

State(s)

Scale

Reference(s)

Silver perch (Bidyanus bidyanus)

NSW, SA, Vic

E, C

Snapper (Pagrus auratus)

NSW, SA, Vic, WA

E

Australian herring (Arripis Georgiana) Yellow-fin whiting (S. schomburgkii)

SA

E

Hutchinson (1999); Ingram et al. (2002); Doroudi et al. (2003); Gooley & Gavine (2003) Hutchinson (1999); Fielder et al. (2001); Ingram et al. (2002); Partridge & Furey (2002); O’Sullivan (2003) Hutchinson (1999)

SA

E

Hutchinson (1999)

NSW = New South Wales; SA = South Australia; Vic = Victoria; WA = Western Australia Source: Adapted from Prangnell (2006, unpublished thesis)

K+-deficient inland saline groundwater (Partridge & Creeper 2004; Partridge & Lymbery 2008). The fish survived and grew well when muriate of potash (KCl) was added to the ISW stocked in earthen ponds (Partridge et al. 2006, 2008), indicating that increasing K+ concentration in ISW has a positive outcome. Similar result was exhibited by snapper (Pagrus auratus) when cultured in ISW containing 5% K+ as in ocean water (Fielder et al. 2001). In this case, the mortality could be avoided by increasing the K+ concentration to 40% of K+ OW concentration (Fielder et al. 2001). 1.5.1.2 USA The use of inland saline water for prawn culture in the USA was first introduced in 1973 in West Texas where prawn production ranged from 3.36 to 5.04 t/ha (Treece 2002). Recently, an increased interest in using ISW for marine prawn and fish culture has become evident due to widespread outbreaks of disease, perception of a cheap under-utilised resource and less conflict with other users (Samocha et al. 2002). In the USA, most inland saline water (ISW) for marine prawn and fish culture is extracted from shallow saline groundwater. In Texas, inland saline groundwater (ISGW) has higher SO4−2 concentration when compared to ocean water at the same salinity (Forsberg et al. 1996). Boyd et al. (2009) did a study on the distribution of IGSW over the state of Alabama in order to map suitable areas of ISGW for aquaculture purposes. The suitability was assessed on the basis of chloride (Cl−) concentration above 126 mg/L. It has been reported that 238 out of 2,527 wells in Alabama had a concentration of Cl− above 126 mg/L. Although different in ion concentrations compared to those in OW, the ISGW has a potential for aquaculture purposes. The ISGW, when used for culturing, has been able to support the survival and growth of marine diatom (Phaeodacrylum tricomutum), brine shrimp (Artemia salina) (Brune et al. 1981); Pacific white shrimp (Penaeus vanname) (Smith & Lawrence 1990; Saoud et al. 2003) and red drum (Sciaenops ocellatus) (Forsberg et al. 1996; Forsberg & Neill 1997). Despite the great potential of ISGW for aquaculture purposes, some problems related to the osmo- and iono-regulatory capacity due to ionic imbalance of ISGW have been

16

Recent Advances and New Species in Aquaculture

notified by Gong et al. (2004). However, a modification of diet by supplementing additional dietary magnesium, potassium, phospholipids and cholesterol to a commercial shrimp feed has been proven effective to improve the osmo- and iono-regulatory capacity of prawns (Gong et al. 2004). 1.5.1.3 India In a similar manner to Australia, in the state of Haryana in India the extent of ISW is surging due to rising ground water tables, which bring saline water to the surface. The salinity of the ISW in the state ranges from 10–35 ppt with high Ca2+ and Mg2+, which has resulted in high water hardness (Jana et al. 2004). As a consequence, productive agricultural land has been destroyed. In India, research has focused on the use of ISW for culturing brackish water and freshwater species, such as black tiger prawns, milkfish (Chanos chanos), grey mullet (Mugil cephalus), barramundi and giant fresh prawn (Macrobrachium rosenbergii). Rahman et al. (2005) reported that black tiger prawns can experience high mortality when cultured in ISW, but survival and growth rates can be improved if the ISW is fortified with K+, Mg2+ and Ca2+. Grey mullet (Jana et al. 2004) and milkfish (Jana et al. 2006) can survive and grow well in ISW ponds. Similarly, Jain et al. (2006) reported that barramundi can survive and grow at lower salinity (15 ppt) compared to higher salinity (25 ppt) ISW. Research funded by the Australian Centre for International Agricultural Research (ACIAR) and the New South Wales Department of Primary Industries (DPI), in partnership with Murray Irrigation Ltd (Partners 2009), has focused on producing prawn larvae in ISW. Using ISW to produce giant freshwater prawn larvae has presented a problem, which is related to the ionic imbalance of the ISW. Jain et al. (2007) reported that prawn larvae only survived until 11 days and developed to stage IV unless the concentration of potassium and magnesium was increased, whilst concentration of calcium was decreased to a similar level to ocean water.

1.6

FUTURE DIRECTIONS

Research in the area of molecular genetics and immunostimulants to prevent disease outbreaks is still in progress. However, the success of the research needs to be quantified by transferring it into the production of disease-free aquaculture products, which is yet to be witnessed and documented. The use of population genetics to enhance the value of ornamental species by improving their colour schemes is restricted to a few freshwater species. Similarly, research into the inland saline water aquaculture has had mixed commercial outcomes.

1.7

REFERENCES

Allan, G.L. & Fielder, D.S. (1999) Inland saline aquaculture activities in NSW. In: Inland saline aquaculture (eds B. Smith & C. Barlow), pp. 14–15. Proceedings of a workshop held in Perth, Western Australia, 6–7 August 1997. ACIAR Proceedings No. 83, Australian Centre for International Agricultural Research, Canberra.

Recent Developments 17 Allan, G.L., Banens, B. & Fielder, D.S. (2001) Developing commercial inland saline aquaculture in Australia: Part 2. Resources inventory and assessment. FRDC Project 98/335, NSW Fisheries Final Report Series No. 31(ISSN 1440-3544) NSW Fisheries, Australia. Anonymous (2005) White Spot Disease, Brazil. http://www.aphis.usda.gov Aquaculture SA 2003. Mulloway aquaculture in South Australia. Primary Industries and Resources SA Fact Sheet FS35/03. Accessed from: www.pir.sa.gov.au/factsheets Axlerod, H.R. (1988) Koi Varieties: Japanese coloured carp-nishikigoi. TFH Publications, Neptune, New Jersey, USA. Bachère, E. (2003) Anti-infectious immune effectors in marine invertebrates: potential tools for disease control in larviculture. Aquaculture (3rd Fish and Shellfish Larviculture Symposium), 227, 427–438. Barrie, A. (1992) The Professional’s Book of Koi, pp. 528–525. TFH Publications, Neptune, New Jersey, USA. Benzie, J.A.H. (1998) Penaeid genetics and biotechnology. Aquaculture, 164, 23–47. Biggs, P.M. (1985) Infectious animal disease and its control. Philosophical Transactions of the Royal Society London B, 310, 259–274. Bishop, S., Chesnais, J. & Stear, M.J. (2002) Breeding for disease resistance: issues and opportunities. Proceedings of the 7th World Congress on Genetics Applied to Livestock Production. Communication 13–01, pp. 597–604. Montpellier, France. Boyd, C.A., Chaney, P.L., Boyd, C.E. & Rouse, D.B. (2009) Distribution of ground water suitable for use in saline-water aquaculture in Central and West-Central Alabama. Journal of Applied Aquaculture, 21, 228–240. Bricknell, I. & Dalmo, R.A. (2005) The use of immunostimulants in fish larval aquaculture. Fish & Shellfish Immunology, 19(5), 457–472. Brock, J.A. (1997) Taura syndrome, a disease important to shrimp farms in the Americas. Journal of World Aquaculture Society 13, 415–418. Brune, D.E., Reach, C. & O’Connor J.T. (1981) Inland saltwater as a medium for the production of biomass. Biotechnology and Bioengineering Symposium, 11, 79–93. Burrells, C., Williams, P.D. & Forno, P.F. (2001) Dietary nucleotides: a novel supplement in fish feeds: 1. Effects on resistance to disease in salmonids. Aquaculture, 199, 159–169. Burton, R.F. (1995) Cation balance in crustacean haemolymph: relationship to cell membrane potentials and membrane surface charge. Comparative Biochemistry and Physiology Part A: Physiology, 111, 125–131. Chang, C.F., Su, M.S., Chen, H.Y., Lo, C.F., Kou, G.-H. & Liao, I.C. (1999) Effect of dietary beta-1,3glucan on resistance to white spot syndrome virus (WSSV) in postlarval and juvenile Penaeus monodon. Diseases of Aquatic Organisms, 36, 163–168. Chang, C.-F., Su, M.-S., Chen, H.-Y. & Liao, I.-C. (2003) Dietary ß-1,3-glucan effectively improves immunity and survival of Penaeus monodon challenged with white spot syndrome virus. Fish & Shellfish Immunology, 15, 297–310. Chou, H.Y., Huang, C.Y., Wang, C.H., Chiang, H.C. & Lo, C.F. (1995) Pathogenicity of a baculovirus infection causing white spot syndrome in cultured penaeid shrimp in Taiwan. Diseases of Aquatic Organisms, 23, 165–173. Citarasu, T., Sivaram, V., Immanuel, G., Rout, N. & Murugan, V. (2006) Influence of selected Indian immunostimulant herbs against white spot syndrome virus (WSSV) infection in black tiger shrimp, Penaeus monodon with reference to haematological, biochemical and immunological changes. Fish & Shellfish Immunology, 21, 372–384. Collins, A. & Russell, B. (2003) Inland prawn farming trial in Australia. Global Aquaculture Advocate, 6(2), 84–85. Collins, A., Russell, B., Walls, A. & Hoang, T. (2005) Inland prawn farming. Department of Primary Industries and Fisheries, Queensland. Doroudi, M.S., Fielder, D.S., Allan, G.L. & Webster, G.K. (2003) Culture of marine and salt tolerant species using inland saline groundwater in Australia. Asia-Pacific Aquaculture, 2003, 73. Doroudi, M.S., Fielder, D.S., Allan, G.L. & Webster, G.K. (2006) Combined effects of salinity and potassium concentration on juvenile mulloway (Argyrosomus japonicus, Temminck and Schlegel) in inland saline groundwater. Aquaculture Research, 37, 1034–1039. Doupé, R.G., Lymbery, A.J., Sarre, G., Jenkins, G., Partridge, G. & George, R. (2003a) The national research and development plan for commercial inland saline aquaculture: A view from afar. Natural Resource Management, 6(1), 31–34.

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Doupé, R.G., Lymbery, A.J. & Starcevich, M.R. (2003b) Rethinking the Land: The development of inland saline aquaculture in Western Australia. International Journal of Agricultural Sustainability, 1, 30–37. Dutney, L. (2004) Commercial production of mulloway Argyrosomus japonicus using saline ground water. In: Australasian Aquaculture 2004, Sydney Convention Centre, Sydney. FAO/WHO (2002) Guidelines for the evaluation of probiotics in food. Report of a joint FAO/WHO working group on drafting guidelines for the evaluation of probiotics in food, London, Ontario, Canada, 30 April and 1 May 2002; http://www.who.int/foodsafety/fs_management/en/probiotic_guidelines.pdf. Fegan, D. (2001) Thailand’s Inland Farming Ban Continues. The Advocate, Dec., 63–64. Fielder, D.S. & Allan, G.L. (2001) Inland production of marine fish. The new rural industries: A handbook for farmers and investors. Retrieved 22 May 2006 from http://www.rirdc.gov.au/pub/handbook/ contents.html Fielder, D.S., Bardsley, W.J. & Allan, G.L. (2001) Survival and growth of Australian snapper, Pagrus auratus, in saline groundwater from inland New South Wales, Australia. Aquaculture, 201, 73–90. Fjalestad, K.T., Gjedrem, T. & Gjerde, B. (1993) Genetic improvement of disease resistance in fish: an overview. Aquaculture, 111, 65–74. Flegel, T.W. & Alday-Sanz, V. (1998) The crisis in Asian shrimp aquaculture: current status and future needs. Journal of Applied Ichthyology, 14, 269–273. Flegel, T.W., Nielsen, L., Thamavit, V., Kongtim, S. & Pasharawipas, T. (2004) Presence of multiple viruses in non-diseased, cultivated shrimp at harvest. Aquaculture 240, 55–68. Flowers, T.J. & Hutchinson, W.G. (2004) Preliminary studies towards the development of an aquaculture system to exploit saline groundwater from salt interception schemes in the Murray-Darling Basin. CNRM Final Report 2002/15, South Australian Research and Development Institute (Aquatic Sciences), Adelaide. Forsberg, J.A., Dorsett, P.W. & Neill, W.H. (1996) Survival and growth of red drum Sciaenops ocellatus in saline groundwaters of West Texas, USA. Journal of the World Aquaculture Society, 27, 462–474. Forsberg, J.A. & Neill, W.H. (1997) Saline groundwater as an aquaculture medium: physiological studies on the red drum, Sciaenops ocellatus. Environmental Biology of Fishes, 49, 119–128. Gannam, A.L. & Schrock, R.M. (2001) Immunostimulants in fish diets. In: Nutrition and Fish Health (eds. C. Lim & C.D. Webster), pp. 235–266. Food Products Press, New York. George, R. & Coleman, M. (2001) Hidden menace or opportunity – Groundwater hydrology, play as and commercial options for salinity in wheatbelt valleys. In: Proceeding of the wheatbelt valleys conference, Merredin, Western Australia. Gibson, G.R. & Roberfroid, M.B. (1995) Dietary modulation of the human colonic microbiota: introducing the concept of prebiotics. Journal of Nutrition, 125, 1401–1412. Gitterle, T., Salte, R., Gjerde, B., et al. (2005) Genetic (co)variation in resistance to white spot syndrome virus (WSSV) and harvest weight in Penaeus (Litopenaeus) vannamei. Aquaculture, 246, 139–149. Gong, H., Jiang, D.H., Lightner, D.V., Collins, C. & Brock, D. (2004) A dietary modification approach to improve the osmoregulatory capacity of Litopenaeus vannamei cultured in the Arizona desert. Aquaculture Nutrition, 10, 227–236. Gooley, G.J. & Gavine, F.M. (2003) Integrated agri-aquaculture systems – A resource handbook for Australian industry development. Rural Industries Research and Development Corporation, Canberra. Gooley, G.J., Ingram, B. & McKinnon, L. (1999) Inland saline aquaculture – a Victorian perspective. In: Inland saline aquaculture (eds B. Smith & C. Barlow), pp. 16–19. Proceedings of a workshop held in Perth, Western Australia, 6–7 August 1997. ACIAR Proceedings No. 83, Australian Centre for International Agricultural Research, Canberra. Harris, S., Savage, S. & Fotedar, R. (2005) Treated inland waters of Western Australia may be useable for greenlip abalone culture. Global Aquaculture Advocate, 8(6), 71–72. Hennig, O.L., Arce, S.M., Moss, S.M., Pantoja, C.R. & Lightner, D.V. (2005) Development of a specific pathogen free population of the Chinese fleshy prawn, Fenneropenaeus chinensis Part II. Secondary quarantine. Aquaculture, 250, 579–585. Hill, A.V.S. (2001) The genomics and genetics of human infectious disease susceptibility. Annual Review Genomics & Human Genetics, 2, 373–400. Hutchinson, W. (1999) Inland saline aquaculture in South Australia. In: Inland saline aquaculture (eds B. Smith & C. Barlow), pp. 20–23. Proceedings of a workshop held in Perth, Western Australia, 6–7 August 1997. ACIAR Proceedings No. 83, Australian Centre for International Agricultural Research, Canberra.

Recent Developments 19 Ingram, B.A., McKinnon, L.J. & Gooley, G.J. (2002) Growth and survival of selected aquatic animals in two saline groundwater evaporation basins: an Australian case study. Aquaculture Research, 33, 425–436. Inouye, K., Tamaki, M., Miwa, S., et al. (1994) Mass mortality of cultured kuruma shrimp Penaeus japonicus in Japan in 1993: electron microscopic evidence of the causative virus. Fish Pathology, 29, 149–158. Itami, T., Asano, M., Tokushige, K., et al. (1998) Enhancement of disease resistance of kuruma shrimp, Penaeus japonicus, after oral administration of peptidoglycan derived from Bifidobacterium thermophilum. Aquaculture, 164, 277–288. Jain, A.K., Kumar, G. & Mukherjee, S.C. (2006) Survival and growth of early juveniles of barramundi, Lates calcarifer (Bloch, 1790) in inland saline groundwater. Journal of Biological Research, 5, 93–97. Jain, A.K., Raju, K.D., Kumar G., Ojha, P.K. & Reddy, A.K. (2007) Strategic manipulation of inland saline groundwater to produce Macrobrachium rosenbergii (De Man) postlarvae. Journal of Biological Research, 8, 151–157. Jana, S.N., Garg, S.K. & Patra, B.C. (2004) Effect of periphyton on growth performance of grey mullet, Mugil cephalus (Linn.), in inland saline groundwater ponds. Journal of Applied Ichthyology, 20, 110–117. Jana, S.N., Garg, S.K., Thirunavukkarasu, A.R., Bhatnagar, A., Kalla, A. & Patra, B.C. (2006) Use of additional substrate to enhance growth performance of milkfish, Chanos chanos (Forsskal) in Inland Saline Groundwater ponds. Journal of Applied Aquaculture, 18, 1–20. Jory, D.E. & Dixon, H.M. (1999) Shrimp white spot syndrome virus in the western hemisphere. Aquaculture Magazine, 25, 83–91. Kesarcodi-Watson, A., Kaspar, H., Lategan, M.J. & Gibson, L. (2008) Probiotics in aquaculture: The need, principles and mechanisms of action and screening processes. Aquaculture, 274, 1–14. Lee, C.L. (1999) Potential of inland saline water for aquaculture of mollusc. In: Inland saline aquaculture (eds B. Smith & C. Barlow), pp. 37–39. Proceedings of a workshop held in Perth, Western Australia, 6–7 August 1997. ACIAR Proceedings No. 83, Australian Centre for International Agricultural Research, Canberra. Lightner, D.V. (1996) Epizootiology, distribution and the impact on international trade of two penaeid shrimp viruses in the Americas. Revue Scientifique et Technique, 15, 579–601. Lightner, D.V. (1999) The penaeid shrimp viruses TSV, IHHNV, WSSV, and YHV: current status in the Americas, available diagnostic methods and management strategies. Journal of Applied Aquaculture, 9, 27–52. Lightner, D.V. (2005a) Biosecurity in shrimp farming: pathogen exclusion through use of SPF stock and routine surveillance. Journal of World Aquaculture Society, 36, 229–248. Lightner, D.V. (2005b) The penaeid shrimp viral pandemics due to IHHNV, WSSV, TSV and YHV: History in the Americas and current status. In: Diseases in Asian Aquaculture. Crustacean Pathology and Diseases (eds P.J. Walker, R.G. Lester & M.G. Bondad-Reantaso), pp. 1–20. Fish Health Section, Asian Fisheries Society, Manila. Mantel, L.H. & Farmer L.L. (1983) Osmotic and ionic regulation. In: The Biology of Crustacea: Internal Anatomy and Physiological Regulation (eds D.E. Bliss & L.H. Mantel). Academic Press, New York. Marques, A., Dhont, J., Sorgeloos, P. & Bossier, P. (2006) Immunostimulatory nature of ß-glucans and baker ’s yeast in gnotobiotic Artemia challenge tests. Fish & Shellfish Immunology, 20, 682–692. McFarlane, J. & Christie, H. (2002) Inland saline aquaculture. In: PUR$L (Productive use and rehabilitation of saline lands) 8th national conference and workshop, National Dryland Salinity Program, Fremantle – Kojonup – Katanning, Western Australia. McIntosh, R.P. (1999) Changing paradigms in shrimp farming: 1. General description. Global Aquaculture Advocate, 2, 40–47. McKinnon, L., Ingram, B. & Gooley, G.J. (1998) Fish production from salt-affected land, profit potential from a persistent problem. Trees & Natural Resources, 40, 29–31. McMahon, D.Z., Baca, B., Samocha, T. & Jory, D.E. (2001) First commercial inland farm in Florida, USA uses zero discharge in low-salinity ponds. Global Aquaculture Advocate, 4, p.54. Milne, C.P.J. (1983) Honey bee (Hymenotpera: Apidae) hygienic behavior and resistance to chalkbrood. Annals of the Entomological Society of America, 76, 384–387. Mizuta, S., Yoshinaka, R., Sato, M., Suzuki, T. & Sakaguchi, M. (1992) Immunohistochemical localization of genetically distinct types of collagen in muscle of kuruma prawn Penaeus japonicus. Comparative Biochemistry & Physiology B, 103, 917–922.

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Moss, S.M. (1999) Biosecure shrimp production: emerging technologies for a maturing industry. Global Aquaculture Advocate, 2, 50–52. Moss, S.M., Arce, S.M., Moss, D.R. & Otoshi, C.A. (2003) Disease prevention strategies for penaeid shrimp culture. In: Pathobiology and Aquaculture of Crustaceans: Proceedings of the 32nd US–Japan Natural Resources Aquaculture Panel, US–Japan Natural Resources Technical Report. Accessible from: http:// www.thefishsite.com/articles/123/disease-prevention-strategies-for-penaeid-shrimp-culture. Nell, J.A. & Hand, R.E. (2003) Evaluation of the progeny of second-generation Sydney rock oyster Saccostrea glomerata (Gould, 1850) breeding lines for resistance to QX disease Marteilia Sydney. Aquaculture, 228, 27–35. NOAH (2002) Vaccination of farm animals. Briefing Document No. 22. National Office of Animal Health. Middlesex, UK. http://www.noah.co.uk Nulsen, B. (1999) Inland Saline Waters in Australia. In: Inland saline aquaculture (eds B. Smith & C. Barlow), pp. 6–11. Proceedings of a workshop held in Perth, Western Australia, 6–7 August 1997. ACIAR Proceedings No. 83, Australian Centre for International Agricultural Research, Canberra. O’Sullivan, D. (2003) Inland saline research in WA. Austasia Aquaculture, August/September 2003, 36–44. Partridge, G.J. & Creeper, J. (2004) Skeletal myopathy in juvenile barramundi, Lates calcarifer (Bloch), cultured in potassium-deficient saline groundwater. Journal of Fish Diseases, 27(9), 523–530. Partridge, G.J. & Furey, A. (2002) Culturing snapper in Dumbleyung – A case study for determining the potential for inland saline groundwater to grow marine fish in Western Australia. In: PUR$L (Productive use and rehabilitation of saline lands) 8th national conference and workshop, National Dryland Salinity Program, Fremantle – Kojonup – Katanning, Western Australia. Partridge, G.J. & Lymbery, A.J. (2008) The effect of salinity on the requirement for potassium by barramundi (Lates calcarifer) in saline groundwater. Aquaculture, 278, 164–170. Partridge, G.J. & Lymbery, A.J. (2009) Effects of manganese on juvenile mulloway (Argyrosomus japonicus) cultured in water with varying salinity – Implications for inland mariculture. Aquaculture, 290, 311–316. Partridge, G.J., Sarre, G.A., Ginbey, B.M., Kay, G.D. & Jenkins, G.I. (2006) Finfish production in a static, inland saline water body using a Semi-Intensive Floating Tank System (SIFTS). Aquacultural Engineering, 35(2), 109–121. Partridge, G.J., Lymbery, A.J. & Bourke, D.K. (2008) Larval rearing of barramundi (Lates calcarifer) in saline groundwater. Aquaculture, 278, 171–174. Paust, G. (1999) Inland saline aquaculture in Western Australia. In: Inland saline aquaculture (eds B. Smith & C. Barlow), pp. 24–25. Proceedings of a workshop held in Perth, Western Australia, 6–7 August 1997. ACIAR Proceedings No. 83, Australian Centre for International Agricultural Research, Canberra. Pequeux, A. (1995) Osmotic regulation in crustaceans. Journal of Crustacean Biology, 15(1), 1–60. Pillay, T.V.R. (1993). Aquaculture: Principles and Practices. Fishing News Books, Oxford. Prangnell, D. (2006) Physiological responses of western king prawns, Penaeus latisulcatus, in inland saline water with different potassium concentrations. Doctoral thesis, Curtin University, Australia. Prangnell, D.I. & Fotedar, R. (2005) The effect of potassium concentration in inland saline water on the growth and survival of the western king shrimp, Penaeus latisulcatus, Kishinouye, 1896. Journal of Applied Aquaculture, 17, 19–33. Prangnell, D.I. & Fotedar, R. (2006a) Effect of sudden salinity change on Penaeus latisulcatus Kishinouye osmoregulation, ionoregulation and condition in inland saline water and potassium-fortified inland saline water. Comparative Biochemistry & Physiology – Part A: Molecular & Integrative Physiology, 145, 449–457. Prangnell, D.I. & Fotedar, R. (2006b) The growth and survival of western king prawns, Penaeus latisulcatus Kishinouye, in potassium-fortified inland saline water. Aquaculture, 259, 234–242. Rahman, S.U., Jain, A.K., Reddy, A.K., Kumar, G. & Raju, K.D. (2005) Ionic manipulation of inland saline groundwater for enhancing survival and growth of Penaeus monodon (Fabricius). Aquaculture Research, 36, 1149–1156. Rayment, G.E. & Higginson, F.R. (1992) Australian laboratory handbook of soil and water chemical methods. Inkata Press, Melbourne. Rengpipat, S., Phianphak, W., Piyatiratitivorakul, S. & Menasveta, P. (1998) Effects of a probiotic bacterium on black tiger shrimp Penaeus monodon survival and growth. Aquaculture, 167, 301–313. Reyes, A., Salazar, M. & Granja, C. (2007) Temperature modifies gene expression in subcuticular epithelial cells of white spot syndrome virus-infected Litopenaeus vannamei. Developmental & Comparative Immunology, 31, 23–29.

Recent Developments 21 Roer, R.D. & Dillaman, R.M. (1993) Molt-related change in integumental structure and function. In: The Crustacean Integument: Morphology and Biochemistry (eds M.N. Horst & J.A. Freeman), pp. 1–33. CRC Press, Boca Raton. Sajeevan, T.P., Philip, R. & Singh, I.S.B. (2006) Immunostimulatory effect of a marine yeast Candida sake S165 in Fenneropenaeus indicus. Aquaculture, 257, 150–155. Sakai, M. (1999) Current research status of fish immunostimulants. Aquaculture, 172, 63–92. Samocha, T.M., Hamper, L., Emberson, C.R., et al. (2002) Review of some recent developments in sustainable shrimp farming practices in Texas, Arizona, and Florida. Journal of Applied Aquaculture, 12(1), 1–42. Saoud, I.P., Davis, D.A. & Rouse, D.B. (2003) Suitability studies of inland well waters for Litopenaeus vannamei culture. Aquaculture, 217, 373–383. Shiau, S.Y. & Hsieh, J.F. (2001) Dietary potassium requirement of juvenile grass shrimp, Penaeus monodon. Fisheries Science, 67(4), 592–595. Skou, J.C. (1957) The influence of some cations on an adenosine triphosphatase from peripheral nerves. Biochimica et Biophysica Acta, 23, 394–401. Smith, L.L. & Lawrence, A.L. (1990) Feasibility of penaeid shrimp culture in inland saline groundwater-fed ponds. The Texas Journal of Science, 42, 3–12. Smith, V.J., Brown, J.H. & Hauton, C. (2003) Immunostimulation in crustaceans: does it really protect against infection? Fish & Shellfish Immunology, 15, 71–90. Soltanian, S., Francois, J.-M., Dhont, J., Arnouts, S., Sorgeloos, P. & Bossier, P. (2007) Enhanced disease resistance in Artemia by application of commercial [beta]-glucans sources and chitin in a gnotobiotic Artemia challenge test. Fish & Shellfish Immunology, 23, 1304–1314. Tantulo, U. (2007) Physiological responses of black tiger prawn (Penaeus monodon Fabricius, 1798) reared in inland saline water. PhD thesis, Curtin University, Australia. Tantulo, U. & Fotedar, R. (2006) Comparison of growth, osmoregulatory capacity, ionic regulation and organosomatic indices of black tiger prawn (Penaeus monodon Fabricius, 1798) juveniles reared in potassium fortified inland saline water and ocean water at different salinities. Aquaculture, 258, 594–605. Tantulo, U. & Fotedar, R. (2007) Osmo and ionic regulation of black tiger prawn (Penaeus monodon Fabricius 1798) juveniles exposed to K+ deficient inland saline water at different salinities. Comparative Biochemistry & Physiology – Part A: Molecular & Integrative Physiology, 146(2), 208–214. Treece, G. (2002) Inland shrimp farming in West Texas, USA. Global Aquaculture Advocate, 5(3), 46–47. Vidal, O.M., Granja, C.B., Aranguren, L.F., Brock, J.A. & Salazar, M. (2001) A profound effect of hyperthermia on survival of Litopenaeus vannamei juveniles infected with White Spot Syndrome Virus. Journal of World Aquaculture Society, 32, 364–372. Vine, N.G., Leukes, W.D. & Kaiser, H. (2006) Probiotics in marine larviculture. FEMS Microbiology Review, 30, 404–427. Walker, G., Gilfedder, M. & Williams, J. (1999) Effectiveness of current farming systems in the control of dryland salinity. Murray-Darling Basin Commission & CSIRO Land and Water, Canberra. Accessed Aug. 2010. Available from: http://www.clw.csiro.au/publications/Dryland.pdf Witteveldt, J. (2006) On the vaccination of shrimp against white spot syndrome virus. Summary Wageningen University dissertation, 3882. Wongteerasupaya, C., Vickers, J.E., Sriurairatana, S., et al. (1995) A non-occluded, systemic baculovirus that occurs in cells of ectodermal and mesodermal origin and causes high mortality in the black tiger prawn, Penaeus monodon. Diseases of Aquatic Organisms, 21, 69–77. Zhan, W.B., Wang, Y.H., Fryer, J.L., Yu, K.K. & Fukuda, H. (1998) White Spot Syndrome Virus infection of cultured shrimp in China. Journal of Aquatic Animal Health, 10, 405–410. Zhu, C., Dong, S., Wang, F. & Huang, G. (2004) Effects of Na/K ratio in seawater on growth and energy budget of juvenile Litopenaeus vannamei. Aquaculture, 234, 485–496.

2

A Global Review of Spiny Lobster Aquaculture

Bruce Phillips and Hirokazi Matsuda

2.1 INTRODUCTION There has been interest in spiny lobster aquaculture for over 100 years, but the first complete larval development was not achieved until fairly recently when Kittaka (1988) cultured Jasus lalandii, the South African rock (spiny) lobster, through its larval stages to the puerulus stage. Despite this success, when the prospects for spiny lobster aquaculture were reviewed by Kittaka and Booth (1994) they stated that ‘the greatest hurdle in the commercial culture of spiny lobster is the difficulty in growing species through their larval stages’. A great deal of research in this area has been conducted since that time and will be discussed in this chapter. Kittaka and Booth (2000) and Phillips and Liddy (2003) have reviewed spiny lobster aquaculture. Both provide excellent reviews of the published literature on the subject, but because of the commercial potential of spiny lobster aquaculture most of the recent developments are unpublished. In addition, it is a rapidly developing area and much of the research is ongoing or recently completed. This chapter examines the literature for both full culture of spiny lobsters and the growout of pueruli and juveniles, trying to bring together the latest research activities worldwide as well as assess future developments. Some earlier references can be found in Phillips and Melville-Smith (2006). In addition to reviewing the literature, we have sought to determine directly the level of interest in spiny lobster aquaculture worldwide. We have contacted colleagues, sought unpublished material, and made extensive use of the internet. A summary follows of the state of activity in the countries for which we were able to obtain information. We have ignored some papers describing attempts to raise larvae, where this appeared to be research conducted for scientific interest and opportunity rather than proposed applications to aquaculture. Likewise, we may have ignored some reports of raising juveniles for similar reasons.

2.1.1

Life history in the wild

When considering an animal for aquaculture it is critical to have a good understanding of its life history. The general life cycle of all the spiny lobster species is well

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

A Global Review of Spiny Lobster Aquaculture 23 Table 2.1 laboratory.

Species of spiny lobster where complete larval development has been achieved in the

Species

Number of instars

Duration of phyllosoma stages in the laboratory (months)

15

10

11

Kittaka (1988)

15–23

10.5–13.4

19

Sagmariasus verreauxi

16–17

6.1–11.6

25.5

Palinurus elephas Panulirus japonicus

6–9

2.0–4.2

11–15

Kittaka et al. (1988) Ritar & Smith (2005) Kittaka et al. (2005) Kittaka et al. (1997) Moss et al. (2000a) Ritar et al. (2006) Kittaka & Ikegami (1988)

Jasus lalandii Jasus edwardsii

Panulirus longipes bispinosus Panulirus penicillatus Panulirus homarus Panulirus argus Panulirus ornatus

20–31

7.5–12.6

Duration of puerulus stage in the laboratory (days)

9–26

Kittaka & Kimura (1989) Yamakawa et al. (1989) Sekine et al. (2000) Matsuda & Takenouchi (2005) Matsuda & Yamakawa (2000)

17

9.1–9.5

20

8.3–9.4

Matsuda et al. (2006)

5.5–8

Murakami, K. (2006)*

18–21

4.5–6.5

23–24

4–5

–

Author(s)

11–26

Goldstein et al. (2006) M.G. Kailis Pty Ltd (2006)** Northern Fisheries Centre 2007 (Jones C. pers. comm.) Australian Institute of Marine Science 2007 (Smith et al. 2009)

* Personal communication, not published ** Press release, Perth, 2 Aug 2006

known. However, some critical aspects such as the diet of the phyllosoma are poorly understood. The life cycle of spiny lobsters is complex and includes a long oceanic larval phase varying in length between species. In the Australian western rock lobster Panulirus cygnus, it is estimated to be 7 to 14 months (Rothlisberg 1988), but for the New Zealand red or Australian southern rock lobster Jasus edwardsii, it lasts at least 12–24 months (Booth 1994). Tropical species may have shorter oceanic cycles (Table 2.1), but there is little reliable data on these species. Spiny lobsters hatch as planktonic phyllosoma larvae (about 1–2 mm long) (Fig. 2.1) and develop through a series of moults, increasing in size. After developing in offshore waters, phyllosoma return towards the continental shelf where the final stage larvae metamorphose into the puerulus, a non-feeding stage (about 30 mm long), which then swims towards the coast.

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Recent Advances and New Species in Aquaculture

(a)

(b)

(c)

(d)

Fig. 2.1 Stages in early development of Panulirus cygnus: (a) newly hatched Stage I phyllosoma; (b) late-stage phyllosoma; (c) final phyllosoma (stage IX); (d) puerulus. (Photographs not to scale. Note: (b) and (c) are preserved specimens, and show sampling damage – ruptured cephalic discs and missing pereiopodal endopods.)

When the puerulus (Fig. 2.1) settles, it moults after a few days to weeks into a benthic juvenile stage. Small juveniles (often called post-pueruli up to a carapace size of about 20 mm) are usually found in shallow coastal reefs and larger juveniles and adults in deeper water offshore (the reverse in Palinurus elephas). It is in these shallow inshore or offshore depths that they reach maturity, mating takes place and the life cycle is completed.

2.1.2

Impetus for lobster farming and enhancement

Spiny lobsters are highly valued seafood, and wild stocks support some of the most valuable commercial fisheries in the world’s major oceans (Booth & Phillips 1994; Kittaka & Booth 2000). There is considerable interest in the acquaculture of spiny lobsters because of their consistently high demand and price, and because of the full exploitation of most natural stocks (Phillips et al. 2000). With most worldwide and Australian rock lobster fisheries now fully exploited, some form of aquaculture must be employed in order to sustain or increase production. Studies with juvenile and adult P. cygnus have shown they have relatively wide environmental tolerances and many behavioural, feeding and growth characteristics identifying them as suitable candidates for culture (Phillips 1985; Phillips et al. 2004). Juvenile and adult rock lobsters can be cultured in communal rearing systems because of their gregarious

A Global Review of Spiny Lobster Aquaculture 25

behaviour. They can be bred in captivity, adapt well to artificial conditions and feeding regimes of culture systems, and their growth rates can be increased by culturing at higher temperatures (Phillips 1985; Kittaka & Booth 2000). There is interest in rock lobster aquaculture in various forms, starting from different phases within the life cycle. These include the culture of larvae, on-growing of puerulus/ juveniles, and on-growing of adults. The on-growing of adults is not covered in this review, but the interested reader is referred to Hart and Van Barneveld (2000a,b) for further information.

2.1.3

History of larval rearing

2.1.3.1 Japan In Japan, capture of juvenile spiny lobsters is restricted by fishery regulations in each Prefectural government; hence wild juveniles have never been used for aquaculture. According to the regulations, the minimum size of Panulirus japonicus for legal capture ranges from 13 to 20 cm in body length (BL), which corresponds to two or three years after settlement. Since the minimum-sized lobsters have good economic value in themselves, there has been little attempt to develop aquaculture of captured lobsters. Instead, phyllosoma culture has long been studied in Japan to produce large numbers of juveniles. Research on phyllosoma culture in Japan has been taking place for more than 100 years. Matsuda and Takenouchi (2007) give a detailed description of the development of phyllosoma culture and research in Japan. The first trial of Panulirus japonicus phyllosoma culture was reported in 1899 (Hattori & Oishi 1899). Several trials followed but it took until 1957 to achieve the culture from hatch to subsequent instars by feeding of Artemia nauplii (Nonaka et al. 1958). After this success, research on phyllosoma culture became more active (e.g. Saisho 1966; Inoue 1981); phyllosoma were reared to more advanced stages as information on culturing conditions, such as environmental parameters and feeding, accumulated. Finally a first gilledstage larva of 30 mm BL was produced in 1978 by using a specially designed circular tank and the feeding of a combination of several diets, such as Artemia, natural zooplankton, and fish fry (Inoue 1981). Significant progress in culturing phyllosoma larvae of palinurid lobsters was made in 1987–88 (see Kittaka 2000). Kittaka (1988), using broodstock transferred from South Africa, cultured larvae of Jasus lalandii in closed-recirculation systems, and successfully produced a puerulus for the first time in 1987 by feeding mussel gonad and introducing the microalga Nannochloropsis oculata into the culture seawater. Then, by using methods similar to those with J. lalandii, pueruli of Jasus edwardsii and of Palinurus elephas were produced in the laboratory (Kittaka & Ikegami 1988; Kittaka et al. 1988). Kittaka et al. (1988) reported that they obtained pueruli of a hybrid between Jasus novaehollandiae and Jasus edwardsii, but J. novaehollandiae is now regarded as the same species as J. edwardsii (Ovenden et al. 1997). Kittaka and Kimura (1989) and Yamakawa et al. (1989) obtained pueruli and subsequent juveniles of P. japonicus in the same period. However, the survival rates from hatching to the juvenile stage were low, less than 1%, due in large part to the lack of information available on the optimal conditions for culturing middle- and late-stage larvae, as well as disease problems. As of 2009, research on the culture of P. japonicus larvae was underway at two laboratories in Japan: Mie Prefecture Fisheries Research Institute and the Minami-Izu Seafarming Centre of the Fisheries Research Agency. These laboratories have proposed optimal

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conditions for mass culture based on the biological, physiological and behavioural aspects of larvae (e.g. Matsuda et al. 2003; Murakami 2004; Matsuda & Takenouchi 2005), resulting in increasing survivorship, and the number of juveniles produced was up to about 300 in 2003 (Matsuda 2004). However, culture techniques for P. japonicus phyllosoma have not yet reached a sufficient level for practical use because it is difficult to prevent bacterial disease without antibiotics and to develop a large-scale system for culturing larvae. Much innovation will be needed for establishing mass culturing systems. Japan continues to show leadership in research in the area of phyllosoma rearing and held an International Workshop on Spiny Lobster Seed Production Technology in Mie, Japan in 2007 (Fisheries Research Agency 2007). 2.1.3.2 Australia There have been several reviews of the potential for aquaculture of rock lobsters in Australia including Chittleborough (1974a), Phillips (1985), Phillips and Evans (1997), Linton (1998) and Phillips et al. (2004). Both full aquaculture through the larval stages and raising of pueruli were suggested, but took a long time to gain acceptance. In 1998 Australia initiated an active research programme into spiny lobster aquaculture (Hart & Van Barneveld 2000a). The species being examined for aquaculture are Panulirus ornatus, J. edwardsii and Sagmariasus verreauxi, although some studies have been made on P. cygnus. The studies were originally coordinated through a subprogramme, the Rock Lobster Enhancement and Aquaculture of the Fisheries Research and Development Corporation. Research supported under this subprogramme includes: a review of the technical potential for rock lobster propagation in aquaculture systems published in reports edited by Hart and Van Barneveld (2000b) and Van Barneveld and Phillips (2002), and a review of the development of rock lobster propagation techniques for aquaculture edited by Crear and Hart (2000). Studies in Australia have included the condition of broodstock regarding the effects on larval quality (biochemical composition, survival, size); the determination of optimum environmental and system requirements for juvenile and adult rock lobster holding and growout; and feed development (Williams 2001, 2007). The morphology of the mouthparts and the digestive tract of phyllosoma larvae are also being investigated to help understand dietary requirements (Johnston 2000, 2003; Johnston et al. 2003; Johnston 2007). Another area of research is examining the manipulation of the moulting times using hormones (Hall et al. 2001). The studies on P. cygnus were conducted in Western Australia by Liddy (2004), seeking to determine the nutritional and feeding requirements of the larvae. These studies were published as Liddy and Phillips (2001) and Liddy et al. (2003, 2004a,b, 2005). The results showed longer initial starvation periods for Stage I larvae resulted in a significant decrease in survival, increase in total intermoult period, and decrease in size, suggesting that any delay in feeding should be avoided. However, larvae that moulted to Stage II and continued development to Stage III showed no significant differences in survival and growth, suggesting culture could persist with larvae if food availability was only limited for a short time (up to 2 days). Lipid was not a major component of early stage larvae and was not the major nutrient accumulated in fed larvae or used in starved larvae. However lipid became a greater component as larvae developed, suggesting that lipid assumes a greater importance as larvae progress through developmental stages.

A Global Review of Spiny Lobster Aquaculture 27

Experiments showed the lipid composition of dietary Artemia affect the lipid composition of early stage P. cygnus larvae. Addition of docosahexaenoic acid (DHA) and AA to enrichments resulted in increased levels of those fatty acids (FA) in the enrichments and Artemia, as well in phyllosoma. As with the accumulation/depletion experiment, the major lipid class (LC) in phyllosoma samples was phospholipid (PL) followed by sterol (ST), with the major LC in all enrichments and enriched Artemia being triacylglycerol (TAG) (Liddy et al. 2005). Not all of the research on J. edwardsii, P. ornatus or S. verreauxi has been published as it is considered ‘commercial in confidence’. However, some of the results have been released and are reviewed under appropriate headings below. The research is conducted and financed by two research organisations funded by the Commonwealth government of Australia, a university, a State-funded research agency and private companies, including Lobster Harvest Pty Ltd and Darden restaurants. Because of the complexity of the studies, the number of researchers, different organisations involved and the ‘commercial in confidence’ situation, it is not possible to give equal coverage to each species or topic, or in some cases to specify which species was the driving reason for the study conducted in the following topics. However, wherever possible this is indicated. 2.1.3.3

New Zealand

New Zealand researchers have carried out extensive studies on the food requirements of phyllosomas (mainly using Artemia). Studies have included the effects of Artemia densities and rations, Artemia sizes and size preferences (Tong et al. 1997; Moss et al. 1999; Moss & Tong 2000; Tong et al. 2000a,b,c). Studies have shown that the quality and size of the Artemia is important. Unlike researchers in Japan, they do not feed with nauplii; instead they feed with at least 1 mm Artemia and, as a preference, a size of 2–3 mm. The larger Artemia are readily captured by the phyllosomas, whereas nauplii (<0.8 mm) are less likely to be captured (Illingworth et al. 1997). In New Zealand an upwelling system was developed by Illingworth et al. (1997) and has proved useful for both J. edwardsii and S. verreauxi. The advantage of this system is that it allows regular cleaning of the culture tanks with little disturbance of the larvae. Using the upwelling system they have been able to produce Stage VIII larvae of J. edwardsii after about 90 days on a regular basis, which has enabled them to achieve larval survival rates of over 60% to Stage VIII, and they have been successful in rearing a few S. verreauxi phyllosoma larvae through to the puerulus stage (Moss et al. 2000a). 2.1.3.4 Brazil Studies are being conducted on phyllosoma culture, focusing on tropical species of spiny lobsters, Panulirus echinatus and Panulirus laevicauda, and also on the economic assessment of spiny lobster growout (Carvalho & Ogawa 2000). In September 2009 the State University of Paraiba (Universidade Estadual da Paraiba) hosted the first Brazilian International Spiny Lobster Culture workshop. The outcome of the workshop was a series of recommendations regarding future spiny lobster culture in the State of Paraiba in Brazil. These included the construction and deployment of puerulus collectors to monitor long-term settlement of puerulus on the nearby coastline, which showed great potential as a settlement site. Other longer-term goals included developing a

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broodstock programme for the three species of lobsters commonly found in this area (Panulirus argus, P. laevicauda and P. echinatus) and renovation of a local land-based shrimp farm for possible future phyllosoma culture. 2.1.3.5

People’s Republic of China

Chang-sheng et al. (2001) have studied feeding, growth and survival of P. stimpsoni phyllosomas. They studied the feeding, growth and survival in larval stages of Stage I–IV of Panulirus stimpsoni. Stage I larvae can fill their ‘stomach’ in 5–9 minutes, with digestion from full to empty taking 45–80 minutes. Results showed stage I phyllosoma are capable of catching different sized Artemia (1–3 days old). 2.1.3.6 India India has initiated a series of spiny lobster studies (Govind 1998). These include breeding programmes aimed towards induction of gonad maturation using hormonal and environmental manipulations, and also larval rearing experiments to shorten larval development and promote moulting through the administration of various moult-related hormones (Radhakrishnan et al. 1999). Radhakrishnan and Vijayakumaran (1993) have examined the early larval development of Panulirus homarus homarus. 2.1.3.7 Portugal Studies of the breeding and development of the early larval stages of Panulirus regius (Luis & Calado 2009) were sufficiently successful for these authors to recommend that this species could be a reasonable candidate for aquaculture in temperate zones.

2.2 2.2.1

BROODSTOCK MANAGEMENT Husbandry and manipulation of captive broodstock

In most cases lobsters for broodstock purposes have been captured from the wild in berried or near-berried condition, or caught in the closed season and held in the laboratory until mating and reproduction occurred. Chittleborough (1974b, 1976) had already demonstrated that P. cygnus, brought into the laboratory as pueruli or as mature lobsters, could be brought to sexual maturity using increasing temperature up to general tolerance limits. Management of P. ornatus broodstock in Australia has proven to be quite straightforward. The species is amenable to a captive environment, and will readily breed throughout summer. Environmental manipulation of breeding has been applied to achieve out-of-season breeding. Photoperiod is the primary cue, although temperature and social factors have a significant influence on breeding success (Knuckey & Cox 2004). Sachlikidis et al. (2005) conducted two experiments to assess the effect of photoperiod and temperature on spawning of P. ornatus. The results suggest photoperiod is the primary cue for the onset of gonad maturity and mating activity, with temperature playing a less important role. Physiological rest and possibly a moult may be required between breeding seasons before spawning can occur. Furthermore, temperature may be an important cue for pre-reproduction moulting.

A Global Review of Spiny Lobster Aquaculture 29

Similar results have occurred with other species. A gradual advancement in the breeding season over three years was achieved in J. edwardsii broodstock by manipulation of both temperature and photoperiod, resulting in mating, extrusion of eggs and hatching of viable larvae up to six months out of season compared to the natural pattern (Ritar & Smith 2008). For P. japonicus, photoperiod and temperature are important factors controlling ovarian development and spawning. Ovarian development progressed at 13–25 °C under long photoperiod ((light–dark cycle (LD) 14:10) and the development was accelerated at higher temperature. Under short photoperiod (LD 10:14), ovarian development depended on temperature: it progressed slowly at 13 °C, whereas it was prevented or considerably delayed at 19 °C and 25 °C (Matsuda et al. 2002).

2.2.2

Ensuring quality of the larvae

There have been three approaches to ensuring the quality of the larvae used in the studies to take the larval stages through their development and metamorphosis to the puerulus stage. 2.2.2.1 Selection of high quality females Smith et al. (2003a) examined photothermal manipulation of reproduction in broodstock and larval characteristics in newly hatched phyllosoma of J. edwardsii. Smith (2004) conducted an extensive study of the nutritional factors affecting larval competency in the spiny lobster, J. edwardsii. They studied the effect of embryo incubation temperature on indicators of larval viability in Stage I phyllosoma of the spiny lobster, and found a detrimental effect of warm incubation temperature during embryonic development of Stage I phyllosomas. The results of these studies were published as Smith et al. (2002b). Other studies on J. edwardsii include the influence of diet on broodstock lipid and fatty acid composition; larval competency and the effect of physical disturbance on reproductive performance; and tissue content, fecundity and quality of eggs and larvae after supplementing the diet of the broodstock with ascorbic acid-enriched Artemia biomass (Smith et al. 2004a,b,c; Smith et al. 2005; Smith et al. 2008 respectively). Smith and Ritar (2007) examined sexual maturation in captive spiny lobsters using J. edwardsii, looking at the relationship of fecundity and larval quality with maternal size. These results indicated that larger females produce larger, more viable larvae. 2.2.2.2 Improving the diet of phyllosomas The effect of diet on on-grown Artemia was studied by Nelson et al. (2002b). Smith et al. (2002a) and Ritar et al. (2003a,b, 2004) studied nutritional and bacterial profiles of juvenile Artemia fed different enrichments and during starvation. Smith et al. (2002c) examined the effects of starvation and feeding on the fatty acid profiles of Stage I phyllosoma of J. edwardsii. 2.2.2.3 Evaluation of larval competency A competency test for the Stage I larvae of several species has been developed (Smith et al. 2003b; Liddy 2004).

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In the case of J. edwardsii, larvae from captive broodstock incubated outside their normal temperature range and fed a restricted broodstock diet performed the poorest in the activity test in terms of large inactivity counts. By contrast, ambient incubated larvae from wild broodstock with free access to a wild diet demonstrated better performance, with low inactivity counts (Smith et al. 2003b). Elevated incubation temperature during embryonic development produced smaller larvae at hatch that performed worse in terms of survival during culture in both unfed and fed treatments. A positive correlation was obtained between survival in 14-day (Stage I) unfed larvae and animals cultured to 42 days (Stage IV), revealing that survival in short-term starvation trials in culture are a measure of the larval competence of congenerics in culture. Positive correlations were obtained between larval activity levels in 23 °C and 10‰ salinity and survival in both 14-day unfed larvae (Stage I) and larvae cultured for 42 days (Stage IV), indicating that J. edwardsii are amenable to the development of an activity test for the assessment of larval competency. However, further refinement of the temperature/salinity parameters for the activity test may be required to discern differences between ‘normal’ larvae obtained from broodstock held under ambient conditions during embryonic development.

2.3 2.3.1

LARVAL REARING Duration of larval development

Each species taken through its larval cycle in the laboratory has a different length of development, number of stages, etc. A list of the 10 species in which complete larval development has been achieved is given in Table 2.1. A complete review of the hatchery propagation of rock lobsters in Australia was made by Ritar et al. (2005) for a funding organisation, but has never been released. Panulirus ornatus Studies of the early development stages of P. ornatus were made by Talbot and McKinnon (2003), but this species was taken through its complete larval development from egg to juvenile by M.G. Kailis Pty Ltd (2006). However, since this is a private company few details are available. Lobster Harvest Pty Ltd was awarded a grant by the government of the Commonwealth of Australia to develop a complete industrial crustacean aquaculture process for commercial propagation from eggs of Moreton Bay bugs (Thenus spp.) and tropical rock lobster species including P. ornatus, to supply Australian, American, Asian and European seafood markets. The project aims to develop industrial aquaculture processes and protocols to deliver commercial quantities of full-sized bugs to market and hatchery-reared tropical rock lobster for growout by third parties. It will require research into fresh and formulated aquaculture diets, tank and water flow design, water treatment, stocking densities and disease management, to enable commercial production of juveniles at targeted survival rates at the greatest cost efficiency. The project also aims to complete lobster DNA research for use in a breeding programme to ensure sustainable competitive advantage. A larva of P. ornatus is reported to have achieved Stage IX out of XI stages in 211 days (Knuckey & Cox 2004). Jones (2007b) reported that the larvae of P. ornatus persist for 120 to 150 days, developing through 11 distinct stages and involving at least 20 moults. These statistics are based on successful rearing of P. ornatus through all stages to puerulus in the

A Global Review of Spiny Lobster Aquaculture 31

hatchery. This has now been achieved on a number of occasions, although in relatively small numbers and certainly not consistently. Because the larvae are physically very delicate, and are accustomed to ocean conditions, the key issues for captive rearing are to provide water that is as close as possible to oceanic in its cleanliness and chemistry, and provide food that delivers the equivalent nutrition of their wild diet. Jones (personal communication) has advised that subsequent to the 8th International Conference and Workshop on Lobster Biology & Management in Canada in 2007, his laboratory at the Northern Fisheries Centre in Cairns, Australia, was successful in producing in excess of 80 pueruli in a single batch. Smith et al. (2009) developed a description of the larval morphology of captive-reared P. ornatus spiny lobsters, benchmarked against wild-caught specimens. Observations were made on the plasticity of the larval duration in P. ornatus. Up to a total of 24 morphological increments were recorded in captive and wild P. ornatus phyllosoma. These were divided into 11 distinct stages by determining the commencement and completion of specific morphological traits. This descriptive morphological key provides a singular reference point for monitoring larval development in this species. The variable nature of the larval duration of P. ornatus suggests that the optimisation of husbandry and nutrition conditions may significantly reduce the length of the hatchery phase to as little as three months and enhance the possibility of providing seed stock for an aquaculture industry based on closed life cycle spiny lobster culture. Payne (2007) and Payne et al. (2006, 2007, 2008) studied the microbiology of larval rearing. The studies concentrated particularly on the microbial communities within the water column in addition to bacteria associated with cultured and wild P. ornatus phyllosomas using both culture-based and molecular microbiological techniques. An insight into the microbial dynamics of the larval rearing system is provided, which, combined with the microbial community overview, has allowed more effective microbial management regimes to be implemented, subsequently improving larval survival. The microbial population of the water column was dynamic and highly diverse, consisting largely of marine microorganisms affiliated with the classes a- and y-Proteobacteria and the Division Bacteroidetes. A small number of opportunistic pathogens were also retrieved, consisting mainly of Vibrio species; however, it is unclear whether these organisms within the water column are involved in the larval mortalities observed. Several Bacillus sp. strains were isolated from the water column and there is potential to use these as probiotics to benefit phyllosoma survival. Extensive colonisation of earlystage third moult phyllosomas by filamentous bacteria was observed following scanning electron microscopy analysis of phyllosomas and the majority of these bacteria were later confirmed as Thiothrix sp. following fluorescent in situ hybridisation analysis. This colonisation is predicted to have negative effects on the health of early stage phyllosomas by hindering their ability to moult and feed. As a result, the animals are likely to be susceptible to infection by pathogenic bacteria such as Vibrio spp. A diverse collection of Vibrio species featured prominently in culture-based and molecular analyses of cultured phyllosomas including potential pathogenic strains, such as Vibrio harveyi and Vibrio campbellii. As Thiothrix sp. is a sulphide-oxidising bacterium, the lowering or removal of sulphide compounds within the larval rearing system is a potential means of control. At present, ozonation of all incoming larval rearing water is serving as an effective Thiothrix sp. control mechanism. The microbial diversity associated with cultured and wild phyllosomas differed markedly using molecular-based clone library analysis. The cultured phyllosoma microbial

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community was dominated by y-Proteobacteria, principally comprising a large proportion of Vibrio and Pseudoalteromonas species, with many likely candidates responsible for compromising phyllosoma health and causing larval mortalities. In contrast, wild phyllosoma were largely dominated by, y-Protebacterial organisms, such as Sulfitobacter sp. The wild environment potentially promotes phyllosoma health through the presence of bacteria, such as Roseobacter sp., that possess favourable attributes for phyllosoma survival. By incorporating such potentially probiotic bacteria into the larval rearing of phyllosoma it may be possible to enhance phyllosoma health and subsequently improve larval survival. Whole phyllosoma survival assays were undertaken to identify both potential pathogenic and probiotic strains isolated during the course of this study. Bacteria were inoculated either directly to the water column or through an Artemia sp. feed vector and phyllosoma survival monitored to measure improved or reduced larval survival. No statistically significant increased mortality (pathogenic strains) or improved survival (probiotic strains) was identified in these animal assays. An absence of water exchanges, with progressively increasing organic load comprising phyllosoma health within all treatments was identified as one flaw that may have caused the negative results. However, candidate bacterial strains were identified for further study including a potentially probiotic Vibrio alginolyticus strain. It is also possible that the Vibrio sp. strains tested during the experiments were not pathogenic – an argument supported by the absence of haemolysin production in all isolates tested. The use of probiotic bacterial strains to improve survival in cultured P. ornatus phyllosoma has potential and is an area that currently requires further research. It is envisaged that an Australian aquaculture industry for P. ornatus will be viable in due time; however, at present further research is required into the monitoring and removal of pathogenic Vibrio spp., through the use of such techniques as real time polymerase chain reaction (PCR), bacteriophage therapy, quorum-sensing inhibition and probiotics. In addition, the development of improved nutritional sources to aid phyllosoma health and promote a robust immune defence for the animal are required if a successful commercial aquaculture industry for the species is to be established. Jasus edwardsii Kittaka et al. (1988) managed to achieve the complete larval development, from egg to juvenile, of this species. In New Zealand, Tong and his colleagues conducted similar work (Booth 1995; Kittaka et al. 2005; Ritar & Smith 2005). Studies of the larval development in Australia and New Zealand including growth and lipid composition, the influence of animal density and water turbulence on growth and survival, effect of predator/prey density and water dynamics on feed intake and growth, effects of light intensity, tolerance for ammonia in early stage phyllosoma, have been reported by Ritar (2001), Bermudes (2002), Ritar et al. (2002, 2003a,b), Nelson (2003), Nelson et al. (2003, 2004), Bermudes and Ritar (2004, 2005, 2008a,b), Smith and Ritar (2006, 2008), Smith et al. (2007), Bermudes et al. (2008). Smith et al. (2004c) examined the influence of diet on broodstock lipid and fatty acid composition and larval competency. Sagmariasus verreauxi This species has been grown through its complete larval development from egg to juvenile by Kittaka et al. (1997), Moss et al. (2000b), and Ritar et al. (2006). Kittaka et al. (1997) reported that the most successful results in the larval culture of any palinurid species so far have been achieved with S. verreauxi. Survival to the puerulus stage of this species was around 10%, a rate similar to that of several commercially cultured

A Global Review of Spiny Lobster Aquaculture 33

prawn species. Sagmariasus verreauxi may be considered a hardy species which is particularly suitable for larval culture. These results are the most promising for palinurid phyllosoma culture, but mortality is still high during the puerulus stage. Ritar et al. (2007) have expanded on the original information on this species. The phyllosoma larvae are hardy and develop rapidly to metamorphosis compared with other temperate rock lobster species and juveniles grow rapidly with anticipated harvest weight of 250 g in 12 months from metamorphosis. These characteristics make it comparable to the tropical rock lobster P. ornatus but with the added advantage of a market premium over other slower-growing temperate species. The study reported here outlines the results of the first year of larval culture and growout of S. verreauxi. Of the 139 phyllosoma that metamorphosed to the puerulus stage, the larval duration ranged from 7–10 months, considerably faster than the 12–18 months for J. edwardsii and other cultured temperate species. When cultured at 15, 18 or 21 °C, juveniles grew faster at the warmer water temperatures. The relatively high survival during larval rearing was attributed to the ozonation of culture seawater, which was found to be highly effective in water disinfection, minimising bacterial pathogens and reducing deaths, especially during the early stages when larvae appear more prone to disease. Studies at the University of Tasmania are currently embargoed by a contract with the Darden restaurants company. The following is a list of some theses currently under embargo in the university library: Crear 1998; Ward 1999, 2005; Westbury 1999; Calvert 2000; Tolomei 2000, 2006; Lyall 2001; Thomas 2001, Higgins 2002; Redd 2004; Patuawa 2005; Johns 2006; Louwen-Skovdam 2006; Jensen 2007; Gudekar 2009. Presumably these studies, plus any new theses that are added, will eventually be released from embargo. Panulirus japonicus This species was taken through its complete larval development from egg to juvenile by Kittaka and Kimura (1989), Yamakawa et al. (1989), Sekine et al. (2000) and Matsuda and Takenouchi (2005). There have been many studies in Japan on aspects of its food and feeding. Many such studies are in Japanese and therefore not readily accessible to outside researchers. However, relevant studies in English include those by Mikami et al. (1995) and Rodriguez-Souza et al. (1996, 1999, 2000). In Japan, tank design is seen as the most crucial element to be taken into account in constructing mass culture systems for phyllosomas. The shape of the tank greatly affects the strength and direction of water current, which affects larval behaviour and position in the tank and this, in turn, affects survival and growth. In general, phyllosomas show negative phototaxis except for a short period of time after hatch (Matsuda et al. 2006). This behaviour causes an aggregation on the tank floor. One option to minimise larval aggregation is to adjust the strength and direction of the water current. Circular tanks of 16–100 L were used in many studies on mass culture of phyllosomas, because the shape was considered to be the most effective way to create a circular current for dispersing and maintaining larvae in suspension (Inoue 1981; Kittaka 2000; Sekine et al. 2000) (Fig. 2.2). However, survival rates to the puerulus stage from hatch were low, less than 10%, and moreover the procedures to maintain circular tanks, such as removal of old food and exuviae, were complicated and probably not appropriate to scale up larval culture for commercial production of juvenile lobsters (Kittaka 2000). Recently, some newly designed tanks have been introduced to phyllosoma culture for the purpose of attaining high survival and efficiency of tank management. Fig. 2.3 shows a shallow elliptical tank of 40 L, with smoothly curved sides and a wide flat bottom. It was

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Recent Advances and New Species in Aquaculture

Fig. 2.2 A circular 50 L tank for phyllosoma culture of Panulirus japonicus used at the Minami-Izu Seafarming Centre, Japan.

designed to prevent excessive aggregation of larvae and allow them to be conveniently observed (Matsuda & Takenouchi 2005). By using this tank, survival rates of 37% to 54% from the middle phyllosoma stages (mean BL 11.5 mm) to the puerulus stage were obtained (Matsuda & Takenouchi 2005). Panulirus argus Several unpublished attempts at culturing larval P. argus were made in the late 1970s in the Florida Keys with only limited success at obtaining middle-stage larvae of 12 mm body length (Moe 1991). Goldstein et al. (2006, 2008) achieved the first complete larval development, from egg to juvenile. The rearing through all larval stages and pueruli to juvenile showed that P. argus has a relatively short larval period of 4.5–6.5 months and therefore has a high potential for aquaculture.

2.3.2

Culture systems for larval rearing

Different systems have been used for different species. A drum-shaped tank was examined for phyllosoma culture (Fig. 2.4). This tank, known as a ‘planktonkreisel’, was originally developed for culturing zooplankton by Greve (1975) and modified by Horita (2007). An advantage of this tank is that larvae are suspended in the water column with continuous

A Global Review of Spiny Lobster Aquaculture 35

Fig. 2.3 An elliptical tank for Panulirus japonicus phyllosomas used at the Mie Prefecture Fisheries Research Institute, Japan.

vertically rotating water current; a disadvantage is that it is difficult to transfer larvae between tanks and remove old food and exuviae. Several pueruli were produced with this tank in Toba Aquarium in Mie Prefecture (Horita 2007). Murakami et al. (2007) further improved the system and designed a vertically revolving tank (VRR System) with a capacity of 70 L (Fig. 2.5). The tank revolves by itself to provide a rotating current regulated accurately. The researchers obtained 34% survival to the puerulus stage from hatching. Moreover, another new culture system was proposed using larval reaction to light to manage the culture tank effectively, as mentioned in the section on environmental parameters below. Each culture system has some drawbacks and advantages, and there is currently no system to culture phyllosomas commercially. Further study on tank design will be required for improving large-scale culture systems.

2.3.3

Environmental parameters

Environmental parameters that affect survival and growth of phyllosoma cultured in the laboratory include food, temperature, salinity, photoperiod, light intensity, water flow rate, water current, and bacterial biota and number. This section explains some of the parameters and how they affect growth and survival of the larvae.

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Recent Advances and New Species in Aquaculture

Fig. 2.4

The planktonkreisel for Panulirus japonicus phyllosomas used at the Toba Aquarium, Japan.

2.3.3.1 Temperature Temperature is one of the most influential factors in phyllosoma culture. Each species has its own specific tolerance levels and growth and survival is highest at an optimal temperature level. Lower temperatures can cause larval mortality, because the intermoult period is prolonged and consequently the surface of larval body becomes soiled with foods given, which probably leads to failure at moulting. On the other hand, high temperatures induce deterioration of seawater, increased infection rates and an imbalance between energy uptake and consumption. Hence, it is crucial to determine the optimal temperature for larval culture. For P. japonicus phyllosomas, the effect of temperature on daily growth increment (DGI), calculated from the moult increment in BL divided by the intermoult period, was investigated throughout development (Matsuda & Yamakawa 1997). DGI is a good indicator to evaluate larval growth because moult increment and intermoult period can be considered simultaneously. When phyllosomas were cultured at four temperatures from 20 °C to 26 °C, DGI increased with increasing BL up to around 7 mm and then decreased gradually until reaching around 18 mm BL at all temperatures tested (Fig. 2.6). Subsequently, DGI increased again as larvae grew. Smaller larvae (c. 15 mm BL) showed the largest

A Global Review of Spiny Lobster Aquaculture 37

Fig. 2.5 The vertically revolving system (VRR system) designed for Panulirus japonicus phyllosomas. (Please see plate section for colour version of this figure.)

DGI at 24 °C (Fig. 2.6). Larval survival rates showed a trend similar to that of DGI: the survival rate at 24 °C was higher than those at 20, 22 and 26 °C for larger larvae. These results indicate that the optimal temperature is 26 °C for P. japonicus larvae smaller than around 15 mm BL and 24 °C for larger larvae. Jasus edwardsii inhabit relatively cold waters (Phillips et al. 2000). Accordingly, the optimal temperature for phyllosomas of this species is lower than for P. japonicus. When early and middle-stage larvae were cultured at 18, 21 and 24 °C, the intermoult period increased with decreasing temperature and the survival rates at 18 and 21 °C were higher than that at 24 °C (Tong et al. 2000b), suggesting that 21 °C was optimal for culturing these stages of this species. 2.3.3.2 Salinity Cultured larvae appear to show tolerance to a wide range of salinity. Salinity in the range from 21 to 36 psu had little effect on survival of early-stage larvae of P. japonicus (2.5 mm BL), but growth rate increased with increasing salinity (Matsuda 2005). For middle-stage larvae (11 mm BL), survival was little affected by salinity in the range from 21 to 33 psu; however, the survival was reduced at 36 psu (Matsuda unpubl.). The larval growth rate showed a trend similar to that of survival, indicating that the optimal range of salinity for middle-stage larvae of this species seems lower and narrower than that for early-stage larvae.

38 Recent Advances and New Species in Aquaculture 400 26°C

350

24°C Daily growth increment (mm)

300

22°C 20°C

250 200 150 100 50 0

0

5

10

15

20

25

30

Body length (mm) Fig. 2.6 Effect of temperature on daily growth increment, as calculated from moult increment in body length divided by intermoult period, throughout the entire phyllosoma phase of Panulirus japonicus.

A survey of wild late-stage larvae of P. japonicus found that they were distributed in the high-salinity water core (34.8 psu) of the ocean during the day (Yoshimura et al. 1999). This suggests that an optimal salinity for late-stage larvae may shift to a somewhat higher range than that for middle-stage larvae. 2.3.3.3

Light

Light conditions, such as photoperiod, light intensity and wavelength, are also important in culturing phyllosomas because they affect behaviour and physiological conditions. The effects of photoperiod and light intensity on larval growth and survival are discussed below, although it is difficult to determine their effects exactly because they are relatively small and experimental results are biased by larval conditions and different culturing systems. Photoperiod The effects of photoperiod vary depending on larval developmental stages. When earlystage larvae of P. japonicus were cultured at various light–dark (LD) cycles, the largest mean BL after four moults was found in continuous darkness, although there were no differences in survival (Matsuda et al. 1997). This was probably caused by lower metabolism rate in the dark than in the light. For late-stage larvae, on the other hand, longer photoperiod appeared to enhance survival and development. Survival of late-stage larvae to the puerulus was higher at LD 14:10 than at LD 10:14 and 12:12, as mentioned below in the section on

A Global Review of Spiny Lobster Aquaculture 39

metamorphosis. The results of the longer photoperiod experiment and increased numbers metamorphosing cannot be explained. Perhaps they were due to innate, selective behaviour (i.e. resulting from natural selection) in final phyllosomas of this species, which are physiologically nearing their metamorphic moult that puts them up nearer surface waters. A sudden change in photoperiod may have a negative effect because physiological conditions of larvae of P. japonicus are strongly regulated by an endogenous rhythm that is induced by an LD cycle (Mikami & Greenwood 1997; Matsuda et al. 2003). A stable LD cycle is recommended for culturing phyllosoma. Intensity Phyllosomas are sensitive to low light levels. Panulirus japonicus phyllosoma detect the light at 0.01 μmol s−1 m−2 (Matsuda et al. 2007) and J. edwardsii phyllosoma can sense even 0.001 μmol s−1 m−2 (Moss et al. 1999). Lower light intensities have little or no effect on the growth and survival of phyllosoma, while higher light intensities have a negative effect on the growth. Jasus edwardsii phyllosoma reared under lower intensities (0.001 μmol s−1 m−2 or completely dark) moulted to a larger size than those reared in higher intensities (0.1 or 10 μmol s−1 m−2) (Moss et al. 1999). The negative effect of higher intensity is probably caused by larval aggregating behaviour and by a difference in the distribution of the larvae and their food (Artemia) induced by a discrepancy between their responses to light. Newly hatched phyllosomas display a strongly positive reaction to light from fluorescent or incandescent lamps (Matsuda et al. 2006). This photopositive reaction soon or gradually disappears with development, and then larvae show a negative reaction to light although there are some individual differences in the response (Matsuda et al. 2006). These reactions to light lead to aggregation and entangling of the larvae, and also to reduction of food intake by causing a behavioural separation of larvae from Artemia (Mikami & Greenwood 1997; Moss et al. 1999), resulting in slow growth and low survival rates. Low light intensity, in general, is used for culturing phyllosoma because it helps maintain even distributions of larvae and Artemia in the tanks so that no behavioural separation between larvae and food occurs (Moss et al. 1999; Matsuda & Takenouchi 2005). In Japan, a clear glass tank utilising the larval response to light was designed to effectively manage culture (Matsuda, personal communication). The system consists of two circular tanks, which are connected but can be isolated from each other (Fig. 2.7). Phyllosomas are cultured in one tank and when it needs cleaning, light is directed at the side of the clean tank (in the case of early-stage larvae showing positive response) or at the side of the dirty tank (in the case of older larvae showing negative phototaxis) to transfer larvae between tanks. The response to light depends on moulting phase (pre-moult, intermoult or post-moult) and nutritional condition, as well as age, but around 80% of cultured larvae can be transferred passively using light, thereby reducing labour to clean the culture tanks. 2.3.3.4

Larval foods – live, inert and formulated options

Studies on the feeding and nutrition of the larvae were and are being conducted on several species and at several locations in Australia and New Zealand. Studies aim to characterise the role nutrition plays in phyllosoma culture and to identify critical nutrients required for successful larval rearing to puerulus. Which nutrients are critical? What nutrients can phyllosomas digest? Can an artificial diet for phyllosomas be developed to provide adequate nutrition and allow nutritional manipulation?

40

Recent Advances and New Species in Aquaculture

Fig. 2.7 The clear glass tank with rounded sides for phyllosoma culture of Panulirus japonicus (Matsuda unpublished).

The diet and feeding ecology of phyllosomas in the wild is unknown and this has led to a number of studies to improve our understanding of the diet of spiny lobster phyllosomas. Jeffs (2007) has reviewed the research undertaken in New Zealand in this critical area. Suzuki et al. (2007) reviewed the DNA approach being used in Japan to attempt to identify the diet in the wild. Takeuchi and Murakami (2007) review the studies in Japan to develop an artificial diet for P. japonicus. Linton et al. (2003) documented Artemia rearing techniques. A number of studies of the mouthparts, feeding behaviour and digestion of several species including J. edwardsii and S. verreauxi have been conducted to understand the feeding behaviour and assimilation processes (Johnston & Ritar 2001; Nelson et al. 2002a; Cox & Johnston 2003a,b, 2004; Johnston et al. 2005; Cox et al. 2008). Other studies have looked at digestive enzyme profiles (Johnston et al. 2004a,b). Konishi (2007) reported on the mouthparts of P. japonicus. Johnston (2007) examined feeding and digestion in P. ornatus. Johnston et al. (2008) described the changes in the structure and function of the mouthparts and foregut of early and late stage phyllosoma of P. ornatus, and Johnston (2007) the structure and function of the digestive gland and ontogenetic changes in enzyme activity of P. ornatus. Smith et al. (2009) examined the ability of P. ornatus phyllosoma to capture, feed and process diets of different texture and particle size during developmental stages I, III and V.

A Global Review of Spiny Lobster Aquaculture 41

Food was captured using the spines and dactyl on the second and third pereiopod and brought to the oral cavity assisted by the second and third maxillipeds. Optimal capture was obtained when foods were a gelatinous-muscular consistency, and hard particles were caught when embedded in a muscular carrier for the hard diet. A range of food textures, from gelatinous to hard, could be masticated in the oral cavity by stage V phyllosoma. Masticated particles were filtered by phyllosomas through spines and hairs in the proventriculus, during Stages I–II, and a filter press in Stages III–V. Maximum particle size passing into the midgut gland for digestion was 7, 3 and 0.5 μm in Stage I, III and V phyllosomas, respectively. B cells were prominent in the proximal ends of the caecum of Stage V phyllosomas and were visible in live animals under light microscopy. In Japan, various items have been examined for feeding to phyllosomas of palinurid lobsters (Kittaka 2000). However, in general, lobsters have exhibited narrow food preferences, limited to Artemia, mussel gonad, squid muscle, jellyfish and fish larvae. At present Artemia and mussel gonad represent the main food items in the culture of phyllosomas because they are available all year round and are easy to store and handle. When phyllosoma take food, they hold it with their maxillipeds and nibble small pieces with the mouthparts. Once they are satiated, they discard the food. This feeding behaviour makes it difficult to investigate how much of Artemia and mussel gonad they consume and assimilate. Matsuda et al. (2009) tried to estimate the relative assimilation of the two diets by cultured larvae of P. japonicus by using stable nitrogen isotope analysis. Calculations based on the measurements of the ratios of 15N to 14N in the bodies of phyllosoma (mean BL = 9.3 mm), Artemia and mussel gonad indicated that 66% of all nitrogen was derived from Artemia and 34% from mussel gonad, suggesting that Artemia is the more important food item at this larval size. Studies by Radford et al. (2008) of specific dynamic action (SDA) as an indictor of carbohydrate digestion in juvenile J. edwardsii have suggested that SDA would be a quick and effective way to determine food digestibility of components used in aquaculture. This technique may be useful in studying food digestibility in phyllosomas. 2.3.3.5

Ammonia

The water quality in aquaculture systems commonly deteriorates with the accumulation of nitrogenous wastes arising from intensive feeding regimes, especially during recirculation. Ammonia may be liberated in the aquatic environment from the decay of uneaten food and as the result of catabolism in cultured organisms. The tolerance of J. edwardsii phyllosomas to the effects of increasing levels of ammonia was assessed at Stages I to IV of larval development (Bermudes & Ritar 2008b). The median lethal concentration (96-hLC50) for total ammonia (and corresponding NH3-N) were 31.6 (0.97) mg 1−1, 45.7 (1.40) mg 1−1, 52.1 (1.59) mg 1−1and 35.5 (1.01) mg 1−1 at Stage I, II, III and IV, respectively. When Stage II larvae were cultured through to Stage III at total ammonia concentration of 0.5 (control), 1.4, 3.8, 6.3 and 9.5 mg 1−1, the intermoult period increased significantly at and above 6.3 mg 1−1 (from 11.2 days in controls to 13.2 days at 9.5 mg 1−1). Thus, the no-observable-effect-concentration (NOEC) of total ammonia (and corresponding NH3-N) at Stage II of Stage II 3.8 mg 1−1 (0.12 mg 1−1) was used as the divisor in the acute: chronic ratio (i.e., 96-hLC50 ÷ NOEC) to estimate the concentrations at Stages I, III, and IV, which were 2.7 (0.8) mg 1−1, 4.4 (0.14) mg 1−1, and 3.0 (0.09) mg 1−1, respectively. These results provide the basis for the design and management of systems for the culture of the larvae of J. edwardsii, and probably other species.

42

Recent Advances and New Species in Aquaculture

2.3.4

Diseases of larval culture

Diggles et al. (2000) reported a luminous vibriosis in J. edwardsii phyllosomas associated with infection by Vibrio harveyi and Diggles (2001) reported a mycosis of juvenile J. edwardsii caused by Haliphthoros sp. as well as possible methods of using chemical control of the latter. Handlinger et al. (2000) reported on a study of diseases in cultured phyllosomas of J. edwardsii. Flavobacterium sp., Vibrio anguillarum, V. alginolyticus and V. tubiashi were all identified. Tolomei et al. (2004) developed methods for the bacterial decontamination of ongrown Artemia, the introduction of which for food, introduces contamination of the culture water. Bourne et al. (2004, 2006) studied microbial community dynamics and biofilm development within a larval rearing tank for P. ornatus and Webster et al. (2006) studied the vibrionaceae infection of P. ornatus. Bourne et al. (2007) pointed out that significant challenges exist in producing commercial scale quantities of pueruli due to an extended larval phase, which acerbates a high rate of larval attrition caused by inadequate nutrition and a challenging microbial environment. In P. ornatus they investigated in a diverse and varied bacterial community in four compartments of the larval-rearing system: the water column, the biofilm, live feeds and the phyllosomas themselves. External fouling of phyllosomas by filamentous Thiothrix sp. was documented by scanning electron microscopy (SEM) and fluorescence in situ hybridisation (FISH). Internal proliferation of bacteria coinciding with mass mortality of phyllosomas was observed in histopathological analysis and identified as Vibrio sp. by specific labelling of sectioned hepatopancreas tissue using FISH. Of particular interest in relation to larval mortalities was a range of Vibrio species, isolated from the four rearing compartments, closely affiliated with V. alginolyticus, V. parahaemolyticus and V. harveyi. The presence of bacterial quorum-sensing signal molecules within the system was demonstrated in both biofilm and phyllosoma environments during a larval-rearing run. Interestingly, a large increase in quorum-sensing signal molecules corresponded with mass mortality in the phyllosomas. Vibrio harveyi is an economically important pathogen in the aquaculture systems for phyllosomas. It is therefore of interest to report a paper providing molecular identification, typing and tracking of Vibrio harveyi in aquaculture systems (Cano-Gomez et al. 2009).

2.3.5

Use of antibiotics

Use of antibiotics is generally discouraged and antibiotics are unlikely to be acceptable in commercial culture systems. However, use of antibiotics is standard practice in Japan in the research laboratory (Hamaski et al. 2007). Liddy (2004) conducted a trial using the antibiotic Trimidinein an attempt to control bacterial infection. The system receiving the antibiotic had much greater survival (estimated at 20%) to stage IV (about 33 days) than the system not receiving antibiotics (estimated at 5%). This indicates that bacteria, which were reduced in number in the antibiotic system (Table 2.1), were a major cause of mortality as all other factors between the two systems were the same. The system without antibiotics experienced high mortality at the first moult. In most cases the published results simply ‘fail to mention’ the use of antibiotics.

A Global Review of Spiny Lobster Aquaculture 43

2.3.6

Tanks and treatment of culture water

Ritar (2001) summarised the situation very well: The rearing of phyllosoma during their long larval development to settlement was accomplished in Japan with a ‘planktonkreisel’ (Kittaka 1994a,b) based on the system of Greve (1968) and Hughes et al. (1974), and in New Zealand with an upwelling tank (Illingworth et al. 1997). In both cases, the larvae and food were maintained in suspension through the upwelling effect of the upward jetting of water.

Static water systems have also been used to rear spiny lobster phyllosomas through all stages of development (Matsuda & Yamakawa 2000), but they are not suitable for culture on a medium to large scale. For all types of systems regular cleaning during the long larval phase is necessary to avoid hygiene problems such as microbial contamination and colonisation of the larval gut and exoskeleton which result in increased mortality. Thus, Illingworth et al. (1997) developed tanks that allowed the regular and frequent mass transfer of larvae to clean tanks with minimal mechanical handling compared with the tedious and frequent transfers of individual larvae required by Kittaka (1994b). However, the designs of culture vessels described by Kittaka (1994b) and Illingworth et al. (1997) are intricate and expensive and prohibit affordable replication. For this reason, an alternative inexpensive, medium-scale system of larval culture was sought to allow replicated comparisons of several experimental treatments reflecting growth and survival under conditions applicable to mass rearing. Ritar (2001) described a system used in the experimental rearing of phyllosomas of J. edwardsii. Seawater was filtered to 1 mm, heated to 18 °C and disinfected with ultraviolet light, then passed into circular 35 L vessels via a series of jets to achieve constant circular flow. Water exited through a screened drain fitted to the wall of the vessel and positioned to maintain a volume of 10 L with a turnover of 3–4 times h_1. Phyllosomas were cultured in this system from hatch to the final Stage XI when fed on-grown Artemia and pieces of mussel (Mytilus edulis). Liddy (2004) reported that three different systems were utilised during his project: two mass rearing systems and an individual holding system. The New Zealand upwelling systems (Illingworth et al. 1997) were used to rear large numbers of larvae; however, this system was limited in its use regarding experimental replication. This led to the usage of the systems designed in Tasmania (Ritar 2001). The Tasmanian tubs were successfully used for the experiments requiring replication and large numbers of larvae for biochemical analysis. Although this system was manually more time-consuming to operate, it was much more suitable for use in the project. This system also had the advantage of much lower setup costs due to its simplicity. The upwelling systems may be more useful for rearing large numbers of larvae for experiments taking place in different systems which provide the replication required for statistical analysis. The upwelling systems are designed more for rearing of larvae than for experimental purposes. The individual system, although timeconsuming, provided excellent survival of larvae in this system. The individual system used appears excellent for experiments requiring individual monitoring of larvae; however the number of larvae or replicates will be limited due to time and/or availability of staff. Ritar et al. (2006), in reporting on the successful rearing of J. edwardsii from egg to juvenile, discussed the use of ozonation to control water quality. For larvae reared from hatch to Stage III, survival was highest and bacterial contamination was lowest in seawater

44

Recent Advances and New Species in Aquaculture

ozonated at low and moderate levels (400 and 500 mV oxidation–reduction potential, ORP). By contrast, at high ozonation (600 mV), all larvae suffered deformities at the moult to Stage II and terminally starved, while in unozonated water (about 300 mV), all larvae died at Stage III probably as a consequence of Vibrio bacteria proliferation. In a second experiment between Stages VI to VIII, larval survival was highest in ozonated water that had been filtered through activated charcoal and coral sand, compared to ozonated water with no filtration or filtered only through activated charcoal. Ozonated water with the combined filtration was used subsequently but there were ongoing deformities, so the level was progressively reduced from 400 mV at Stage VIII to 330 mV at Stage X, at which time ozonation was discontinued. Larvae were then cultured in non-ozonated water to metamorphosis of eight pueruli at 377 to 437 days after hatch, of which two survived to juvenile. Ozonation was thus effective up to Stage IX in improving culture water to minimise bacterial disease without problems of larval deformities. Other studies using ozonation to oxidise and destroy pathogens and thereby improve survival of J. edwardsii phyllosoma were reported by Ritar and Smith (2005) and Ritar et al. (2006).

2.3.7

Metamorphosis

Metamorphosis is considered not to be triggered by external stimuli (McWilliam & Phillips 1997, 2007). In fact, phyllosomas cultured in the laboratory metamorphose without any apparent cues, under constant environmental and feeding conditions. However, as pointed out by McWilliam and Phillips (2007), while there is still no definite answer to the question ‘What triggers metamorphosis in phyllosomas?’, a high-energy diet, and nutritional levels in the final phyllosoma stage, and perhaps the penultimate stage also, are implicated in successful metamorphosis. Hence it is likely that the aquaculture design provides these feeding conditions. However, for larval P. japonicus longer photoperiod appears to improve the chance of successful metamorphosis. For phyllosomas at the penultimate stage (376 days old, mean BL = 23.5 mm) that had been cultured at an LD cycle of 12:12 and were then cultured under three LD cycles of 10:14, 12:12 or 14:10 at 25 °C for 90 days, the average time required to reach metamorphosis decreased with better survival as the length of the light period lengthened, because the larvae at 10:14 and 12:12 exhibited repeated moults in the final stage. It was found that BLs of phyllosoma at the time when metamorphosis occurred decreased significantly as the length of the light period increased (33.7 mm (n = 7) at 10:14, 31.5 mm (n = 7) at 12:12, 30.4 mm (n = 14) at 14:10). However, more research is necessary to determine the effect of photoperiod on metamorphosis, because there was a possibility that the extension of day-length from 12 to 14 hours may have been associated with some unknown endogenous rhythm to shorten intermoult periods and lead to precocious metamorphosis at smaller body lengths. Water currents, which are influenced by the design of the culture tank, the water inflow and the flow rate, affect the physical process of metamorphosis from the phyllosoma to the puerulus phase. A continuous vertically rotating current probably provides suitable conditions for a metamorphosing phyllosoma, but before we explain the reason(s) why this is the case, the physical process of metamorphosis will be described. Murakami et al. (2007) divided the process into five stages: in the first stage, the eyestalks gradually contract in length, bringing the eyes closer to the cephalon; in the second stage, the pereiopods begin

A Global Review of Spiny Lobster Aquaculture 45 100 Success rate (%)

(39) 80 60 (32) 40 20 0 100L planktonkreisel providing a rotating current

5L circular tank providing a slow current

Fig. 2.8 Success rate of metamorphosis of Panulirus japonicus. In the circular 5 L tank with a water flow rate of 0.5 L/min, the success rate was 38% (n = 32), while the success rate was 85% (n = 39) in the rotating current of the 100 L planktonkreisel with a water flow rate of 1.5 L/min.

to contract and become shorter, while still within the old shell; in the third stage the short pereiopods are completely extracted from the old pereiopod shell while still inside the thorax shell, but the abdomen remains unchanged within the shell; in the fourth stage; the abdomen rises inside the old shell of the thorax and then the whole body, except for the eyestalks, antennules and antennae, springs out of the old shell; in the fifth stage, the puerulus rapidly flexes its abdomen backwards by bending and stretching backwards while moulting the eyestalks, antennules and antennae. For pueruli, it is difficult to remove the old shell of antennae when they remain on the bottom of tank in still water, or in slow water current, because the old shell of the antennae is light and clings persistently, even though the pueruli rapidly flex backwards to complete exuviation. However, in the vertically rotating conditions, the old shell floats or sinks slowly in the current in the middle of the water column, while the newly metamorphosed puerulus, which is relatively heavy, sinks quickly, and these opposing movements assist in completing metamorphosis successfully. When phyllosomas metamorphosed in a circular 5 L tank with a water flow rate of 0.5 L/min, the success rate was 38% (n = 32), while the success rate was 85% (n = 39) in the rotating current of the 100 L planktonkreisel with a water flow rate of 1.5 L/min (Fig. 2.8). It was asserted by Yoshimura et al. (1999) that metamorphosis of P. japonicus takes place in the upper or middle layer of the ocean as metamorphosis in the upper water column, not on the bottom, may be necessary for larvae to succeed in moulting.

2.3.8

Economic feasibility of larval culture

Archer and Nichol (1996) reported that Palinurus elephas was a suitable candidate for aquaculture using an ‘expert choice’ model. Price and consumer acceptance were the most important economic factors considered in the model, whilst growth rate, seed availability and nutritional requirements rated highly as biological criteria. The major hurdle is associated with the larval development period. However, in P. elephas this is estimated to be only 4–7 months (in the wild) and Archer and Nichol believed that a dedicated research programme would solve this problem.

46 Recent Advances and New Species in Aquaculture

2.4

RAISING WILD -CAUGHT PUERULI AND JUVENILES

Due to the difficulties associated with culturing spiny lobster larvae from the egg through to the puerulus, particularly the length of time that this is likely to take, an easier alternative for aquaculture of rock lobsters is to begin the process with the puerulus stage or first juvenile captured in the wild (Hair et al. 2002). The reason for this idea was the successful development of collectors to catch the settling pueruli in good numbers. Many countries have an interest in spiny lobster aquaculture but only a few have produced any lobsters for the market (see Fig. 2.9). Spain, Japan, Belize, Cuba and Taiwan have all reported production in the past, but this has now ceased, or in the case of Taiwan, is minimal. Table 2.2 shows the recent aquaculture production of spiny lobsters by some countries worldwide. Not all countries report such production, or else it is included in their total lobster production. In India, we understand that following the 2004 tsunami, lobster production in cages on the southeast coast has not been re-established (Fig. 2.10a,b).

Aquaculture production 180 160

Indonesia India Taiwan Spain Singapore Philippines Japan Cuba Belize

140 Tonnage

120 100 80 60 40 20 0 1958

1963

1968

1973

1978

1983 1988 Years

1993 1998

2003

Fig. 2.9 Spiny lobster aquaculture production by country. Vietnam is not included because the scale of production is so large compared to the production of these other countries. (Please see plate section for colour version of this figure.)

Table 2.2 Total aquaculture production of spiny lobsters over the last 10 years by some countries (data from FAO). Country

1998

1999

2000

2001

2002

2003

2004

2005

2006

2007

Philippines Singapore Vietnam* Cuba Taiwan Indonesia India

10 12 1,000 2 12

19 11 500 1 18

27 6 600 22 6 5

17 10 1,000 13 1 10

10 6 1,127 11

18 11 1,200 3

19 14 1,120 1

19 5 1,200

23 8 1,900

64 2 1,400

10

10

20

50

100

100 3

* Not an exact match with FAO data

A Global Review of Spiny Lobster Aquaculture 47

(a)

(b) Fig. 2.10 (a) Floating cages used for growout of several Panulirus spp. in Tuticorin, India. Another floating cage can be seen in the background. (b) A view of the lobsters in the cage.

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Recent Advances and New Species in Aquaculture

2.4.1

Collection of seed

Collectors have been used successfully to catch the puerulus stage of spiny (rock) lobsters to provide animals for study in the laboratory, to investigate levels of puerulus settlement, and for mariculture purposes. Different types of collectors have been developed for different species and different areas. In Australia focus was mainly on the use of artificial seaweed collectors for P. cygnus; crevice-type collectors were developed for P. argus in the US and J. edwardsii in New Zealand. Workers in other countries, including Antigua, Bermuda, Cuba, Grenada, India, Jamaica, Japan, Mexico, and St Paul and Amsterdam Islands have also used collectors to catch the puerulus stage of Panulirus and Jasus spp. Collectors of various shape and size have been constructed from a range of materials. The biological response to collectors appears to vary among species, and the most appropriate collector has not yet been determined for any species. The type of mooring depends on the type of collector and the water depth. Most collectors have been set inshore (within 2 km of shore), where best catches have been made, either at the surface or near the sea floor at depths down to 12 m. Those near the surface require protection from the full force of waves and swell. Some collectors require conditioning before they are fully effective. The saturation level is unknown for any collector, but high catches on some suggest it is seldom approached. Lobsters remain on collectors for various times according to species and collector type (Phillips & Booth 1994). Pueruli can be consistently taken in relatively large numbers using artificial seaweed habitats near shore (Phillips 1985; Kittaka & Booth 1994), with several studies investigating cost-effective, large-scale harvesting of pueruli from the wild for growout purposes (Booth et al. 1999; Hart et al. 2000; Rossbach et al. 2001; van Barneveld 2001; Mills & Crear 2004; Gardner et al. 2006). Other major constraints to further development of the growout of juveniles appear to be the high cost and irregular abundance of wild seed (Phillips & Liddy 2003). Irregular abundance of wild seed has also been identified as a problem in New Zealand (de Zylva 2002) and a possible problem in Namibia (Keulder 2005). Jones (2007a) reviewed the growout system in South East Asia, particularly in Vietnam. Throughout Southeast Asia the focus of interest and development for lobster culture is now firmly on P. ornatus. The supply of seed focuses on the swimming puerulus, and methods were developed to catch these as they swam in towards the coast. Specific locations were identified where pueruli were somewhat aggregated, typically in bays or channels where north-flowing currents develop. Eighty percent of the seed catch is now by this method using surface nets deployed between two stationary boats, at night. A light is positioned at the net centre (to attract fish) on the surface and the net is set at about 9 pm, retrieved at 1 am and again at 5 am. Each net may return up to 15 pueruli in one night, although the average is usually less than 10. Juvenile lobsters that have settled to the bottom are also fished using alternative methods, which represent habitat enhancement. Timber poles, and coral rocks are drilled with 10 and 15 mm diameter holes to provide refuge for the small lobsters. Bundles of netting are also used. These are placed in shallow water between 1 and 5 m depth, generally 100 to 500 m off the beach. Divers swim down and retrieve the small lobsters. These are serviced every 2–3 days through the season. The catch of these more advanced juveniles represents 15 to 20% of the total seed catch, which varies from 1–2 million annually. Jones (2010a,b) reviewed the current situation and discussed future developments in Vietnam, Indonesia and Australia, and Phillips and Fotedar (2010) also review the

A Global Review of Spiny Lobster Aquaculture 49

situation in Vietnam and compared its aquaculture production of spiny lobsters to other countries.

2.4.2

Activities in individual countries

2.4.2.1 Australia 2.4.2.1.1

Shore-based operations

Panulirus ornatus Jones (2000) and Jones et al. (2001) investigated the effect of density on growout of juvenile (3.24 g) P. ornatus, in a flow-through holding system. Results showed survival and size were unaffected by the densities tested (14, 28 and 43/m2). Mortality was consistent through time and was almost entirely attributable to cannibalism of post-moult individuals. The cannibalism may have been due to inappropriate shelter and feeding strategy. Despite higher mortality than anticipated, growth was rapid, representing a specific growth rate of 1.56% per day, sufficient to permit growth to in excess of 1 kg in 18 months. Johnston (2007) and Johnston et al. (2007a) evaluated partial replacement of live and fresh feeds with formulated diets in P. ornatus including effects of diet form, particle size, feeding stimulants and protein. Dry-pelleted diets for P. ornatus have demonstrated not only diet acceptance but also high survival rates (59%) (Williams et al. 2000). A dietary protein specification of at least 43% appears to be the minimum requirement for acceptable growth and survival rates and this is likely to be dependent on the energy concentration of the diet. Jones and Shanks (2008b) have reviewed the requirements for the aquaculture of P. ornatus in Australia. Panulirus cygnus Growout of pueruli or juveniles is possible in the laboratory. Australian experience with P. cygnus pueruli was that they can be raised in the laboratory, with up to 95% survival (Phillips 1985). At a constant temperature of 25 °C, P. cygnus juveniles can be raised under laboratory conditions from the puerulus stage (of carapace length (CL) about 8 mm) to commercial size (CL 76 mm) in about 2.1 years. This is approximately half the time required in the wild, where juveniles take on average 4 years to reach the legal minimum size (Chittleborough 1974a; Phillips 1985). The protein and lipid requirements of P. cygnus post-puerulus were studied using diets with varying protein levels (30, 35, 40, 45, 50 and 55%) at each of two lipid levels (6 and 10%) (Glencross et al. 2001). Growth was better in animals fed diets with protein levels greater than 50%, and significantly better in those fed the lower lipid diets, with survival in all treatments equal to or greater than 75%. Results indicate a very high dietary protein requirement and a low capacity to use dietary lipid. The results of this study support that P. cygnus post-puerulus have a very high dietary protein requirement and a poor capacity to utilise dietary lipid. It should be noted, though, that the relative growth of the P. cygnus post-puerulus fed the pelleted diets was considerably lower than of those fed the reference diet of fresh mussels. Additional studies by Johnston et al. (2006, 2007b, 2008) have elucidated stocking densities, types of shelter, survival and growth when fed natural and formulated diets and temperatures and feeding frequencies for P. cygnus growout. The implications for P. cygnus culture are that temperatures should be maintained close to 23 °C during the entire growout

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period, with due care taken to minimise mortalities through adequate provision of food and shelter. Feeding P. cygnus once daily to excess just prior to dusk to coincide with nocturnal feeding behaviour is recommended. Moyle et al. (2009) continued these studies of P. cygnus and studied the effect of stocking densities on growth and survival. The results of the study support the results of previous studies that indicate that P. cygnus postpueruli should be stocked at densities less than 100/ m2 in order to minimise the negative effect of density on growth and survival. In addition to studies of the growout of pueruli and juveniles, a number of studies in Australia, particularly on P. cygnus, have investigated catching, handling, transport, holding and shipping live lobsters to international markets. These have included studies by Paterson and Spanoghe (1997), Jussila et al. (2001), Tsvetnenko et al. (2001), Evans et al. (2002), Paterson et al. (2005), Fotedar et al. (2001, 2006), Fotedar (2006), and Fotedar and Evans (2011). An understanding of the influence of transport conditions on the health and condition of crustaceans might be crucial to sustained success in the live export market. Measurement of immune system parameters can provide an indication of the health of the animals, as can the measurement of physiological stress parameters. If the response to stress associated with live transport can be quantified, then it may be possible to develop alternative practices to minimise levels of these stressors. The results, summarised from Fotedar and Evans (2011), show various haematological and immunological parameters such as total haemocyte counts (THC), differential haemocyte counts (DHC), clotting time, phenoloxidase activity, phagocytic index, reactive oxygen intermediates and antibacterial activity have all been considered as potential health or disease markers in crustaceans and can be used to assess stress responses to live transport. However, influences of stress on immune responses are diverse and usually depend on the nature of the stressor, the immune parameter studied and the physiological status of the animal (Lacoste et al. 2002). Thus it is unlikely that one parameter alone would predict health status and future mortalities. In studies on immune responses to live transport of the spiny lobster P. cygnus, measurement of a suite of immune parameters in harvested lobsters assessed by factory grading staff as either healthy or unhealthy, provided evidence of decreased immunocompetence in the unhealthy lobsters. Since it is unlikely that an unhealthy lobster would have entered a lobster pot, it can be assumed that this decrease in immunocompetence, and, therefore, increased susceptibility to disease, was induced through exposure to environmental stressors during the harvesting and storage procedures (Evans et al. 2002). Choice of a combination of parameters is important in assessment of stress response or health status. In animal research, some individual variability is always found due to factors such as moult stage, age and size of animals. Crustaceans may also vary in their normal response and capacity to respond to different factors due to natural genetic variation. This individual variability can result in high variation between animals and mask the trends within the group. Likewise, a trend visible in a group may have limited predictive ability when applied to an individual. Hence, it is important to understand the purpose of the test. In addition, some parameters may have a relatively wide normal range in unstressed animals while others are easily altered by sampling procedures (Fotedar 2006). Studies of shell colour are important because of consumer response to specific colours in the marketplace. Coloration in crustaceans is affected by a combination of factors: diet, environment and genetics. A study by Wade et al. (2009) has shown that a protein called crustacyanin forms a specific interaction with a carotenoid, astaxanthin. This presents a unique avenue to understand and optimise colour production in farmed spiny lobsters and enhance the value of these seafood products.

A Global Review of Spiny Lobster Aquaculture 51

Jasus edwardsii Juveniles are usually fed a diet of fresh mussels. However, dry and moist manufactured feeds have been examined as cheap alternatives to fresh mussels for J. edwardsii (Hart & Van Barneveld 2000b). Lobsters fed cultured mussels can reach a market size of 250–300 g within 3–4 years (Holland & Jeffs 2000). Crear et al. (2000) reported that growth of juvenile J. edwardsii is influenced by diet and temperature, whilst survival is influenced by diet and tank environment. Geddes et al. (2001) reported on a study of the optimum environmental and system requirements for juvenile and adult rock lobster holding and growout of J. edwardsii in South Australia. Simon and James (2007) studied the effect of different holding systems and diets on the performance of juvenile J. edwardsii and found that sea-cage culture of juvenile spiny lobsters can provide significant growth advantages over tank systems, possibly due to supplementary nutrition from fouling organisms. However, seacage structural design, feeding and diet type are important factors influencing the performance of J. edwardsii juveniles in sea-cages. Studies in Tasmania have defined the optimum growout temperature for J. edwardsii to be 19–21 °C (Thomas et al. 2000). In New Zealand, James and Tong (2000) examined the effects of stocking density and shelter on growth and survival of early juvenile J. edwardsii. They found growth (length and weight) was significantly slower when densities increased from 50 through 100 and 150 to 200 animals/m2. Survival rates were high, with little change at densities tested. However, they found that shelter was essential to maintain high survival rates but had no effect on growth. They recommended commercial operation densities of 50–100 rock lobsters/m2, with adequate shelter to maximise growth and survival. Crear et al. (2002) examined commercial shrimp growout pellets as diets for juvenile southern rock lobsters J. edwardsii and the influence on growth, survival, colour and biochemical composition. Johnston et al. (2003) studied dietary carbohydrate: lipid ratios and nutritional condition. Measuring food intake, food availability and the relationship to survival, growth, dominance and agonistic behaviour in J. edwardsii were studied by Thomas et al. (2002, 2003). Tolomei et al. (2003) examined the effects of diet immersion time on growth, survival and feeding behaviour, while Ward et al. (2003) determined the optimal dietary protein level, at two lipid levels. The studies were wide ranging and included oxygen consumption (Crear & Forteath 2000, 2001a,b); production of ammonia (Crear & Forteath 2002); effect of photoperiod on growth and survival (Crear et al. 2003), comparison of wild and cultured adults for growth, sensory analysis and oil composition (Nelson et al. 2005). Oliver & MacDiarmid (2000) examined blood refractive index and weight/carapace length ratios as indices of condition in juveniles. McLeod et al. (2004) studied the changes in the body composition of adult male southern rock lobster, J. edwardsii, during starvation. Johnston (2003) investigated the ontogenetic changes in digestive enzymology of J. edwardsii. The study suggested that carbohydrate and lipid are utilised first, followed by protein. Consistently high levels of lipase in the puerulus digestive gland confirmed the importance of lipid as a major energy substrate. Radford et al. (2008) studied theeffects of dietary carbohydrates on growth of J. edwardsii. The results indicated that glycogen from mussels and agar could be used as carbohydrate source in the future development of a practical diet for spiny lobsters. Simon (2009c) studied digestive enzyme response to natural and formulated diets in cultured J. edwardsii. Simon (2009c) examined enzyme response to natural and formulated diets. Rearing of juvenile lobsters on the formulated diet for six months resulted in a marked decrease in the

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digestive capacity (i.e., total and specific enzyme activity of the foregut and digestive gland) and nutritional condition of lobsters. Overall, these results suggest that difficulty in the digestive processing of formulated feeds may help to explain the bottlenecks encountered in developing more effective formulated diets for juvenile spiny lobster culture. Improvements in the dissolution of dietary ingredients upon entering the foregut, and in the digestibility of dietary carbohydrate sources, may assist in further improving the performance of formulated diets for lobsters. Simon (2009d) examined the effect of carbohydrate source, inclusion level of gelatinised starch, feed binder and fishmeal particle size on the apparent digestibility of formulated diets for spiny lobster juveniles of Jasus edwardsii. The results indicated that using digestible carbohydrate sources (dextrin, carboxymethyl cellulose (CMC), and native wheat starch) for energy, reducing the inclusion level of gelatinised starch, and using CMC or gelatine as binders improve the apparent digestibility of formulated diets. Furthermore, the results also indicate that the use of more soluble and pre-hydrolysed protein sources in diets for J. edwardsii may greatly improve digestibility which is critical in these spiny lobsters where overall food intake is limited. Ritar and Smith (2008) reviewed the whole question of reproduction and growth in marine lobsters. Handlinger et al. (2000, 2001, 2006) reported on a two-year study of diseases in juveniles of J. edwardsii. Several fouling organisms were found, including Leucothrix-like bacteria, stalked peritrick ciliates, Chilodonella-like flagellates, amoebae and fungi. The overall level of disease was low. 2.4.2.1.2

Sea-cage operations

Panulirus ornatus Jones and Shanks (2008a) reported the growout of P. ornatus in floating cages within shrimp ponds in Cairns, Australia. Early indications from the trial were encouraging and indicated that the production of P. ornatus on a formulated diet, in shrimp farm ponds may be possible. 2.4.2.2 New Zealand 2.4.2.2.1 Shore-based operations James (2007) has summarised the situation in New Zealand. The National Institute of Water and Atmospheric Research (NIWA) undertook an extensive programme of research over a 10-year period to investigate the optimal conditions for on-growing wild-caught pueruli of J. edwardsii. The initial focus was on land-based holding systems and measured the efficacy of natural diets. The research showed that the optimal stocking density for captive lobsters was 50 lobsters/m2/internal surface area of the container (James et al. 2002). Other studies have looked at the growth and survival rates with and without shelters (James et al. 2002); salinity (Moss et al. 2000b); holding tank design (James et al. 2003). However, after the negative conclusions of the economic study by Jeffs and Hooker (2000) and several attempts at commercial on-growing of pueruli in land-based facilities, the focus on lobster aquaculture shifted from land- to sea-based on-growing. 2.4.2.2.2 Sea-cage operations Following up on initial studies by Jeffs and James (2001), James (2007) and James and Simon (2008) reported that J. edwardsii do well in sea-cages in New Zealand. Commercial

A Global Review of Spiny Lobster Aquaculture 53

scale sea-cage on-growing trials reared J. edwardsii pueruli to a marketable size of 200– 250 g in 2–3 years. NIWA has continued sea-cage research and also the development of an artificial diet to improve the economic viability of on growing in sea-cages. These growout studies in New Zealand were on the level of commercial-scale operations and were conducted with commercial partners. Studies of J. edwardsii continue in New Zealand. Radford et al. (2004) reported on temporal variation in the specific dynamic action of juveniles; while Radford et al. (2005) looked at haemolymph glucose concentrations of juvenile feeding on different carbohydrate diets and Radford et al. (2007) examined effects of dietary carbohydrate on growth of juveniles. These studies have been so successful that NIWA in New Zealand now offers lobster aquaculture services to the public and one of its pamphlets states ‘Lobster Aquaculture is here’! 2.4.2.3 India 2.4.2.3.1 Shore-based operations In India it has been shown that juveniles of Panulirus homarus homarus can be cultured to 200 g with good survival in 5–6 months (Vijayakumaran & Radhakrishnan 1984). In India, another type of culture system used is ‘pit culture’. Pits are dug into hardened substrate above the low tide mark, with water exchange occurring at high tide. Shelters are supplied in the pits, and the entire area is covered with nets fixed along the sides, preventing escape of the lobsters. There is some supplementary feeding with trash fish, but feeding mostly relies on natural production within the pits (Philipose 1994). Studies in India have also indicated the feasibility of commercial culture. Both P. homarus homarus and P. ornatus juveniles can be economically cultured from wild caught juveniles, with growth from 50–100 g to 250 g in 5–6 months (Kaleemur Rahman & Srikrishnadhas 1994; Kaleemur Rahman et al. 1997). India has initiated a series of spiny lobster studies (Govind 1998), feed trials to enhance growth of juveniles (Raj et al. 1998) and also sea-ranching to study survival and growth of juveniles are being investigated (Achary et al. 1998). Another area of interest is in water quality and disease management in lobster culture. Vijayakumaran et al. (2009) reported that pueruli and post-pueruli, early juveniles and sub-adults of the spiny lobster, P. homarus, and juveniles of P. ornatus were grown in different floating sea-cages along the southeast coast of India from May 2003 to May 2007. This study indicated that post-pueruli of P. homarus can be grown to over 200 g in 12 months and up to 350 g in 16 to 17 months in-sea cages. Juveniles (average weight 76.35 ± 34.50 g) of P. ornatus, reared with P. homarus at a stocking density of 80 individuals/ m2, showed a weight gain of 139 g in 155 days at a rate of 0.89 ± 0.32 g per day with an SGR of 0.67. Live marine clams, Donax spp., were the main feed, supplemented with the gastropod Xancus pyrum, the green mussel Perna viridis, marine crab (Charibdis sp.), squid (Loligo sp.), and fish such as clupeids and Leognathus sp. Pueruli and post-pueruli settled in large numbers (up to 35 individuals/month in one cage) both inside and outside the cages. Kizhakudan et al. (2007) studied the high-density growout techniques for P. homarus in the laboratory. An average weight gain of 30–50 g was obtained in 220 days, with a survival rate of 50%. Repeated antennal ablation was found to greatly reduce the territorial defence and aggressive behaviour among the lobsters, and led to reduced cannibalism, improved growth rates and better performance under high-density stocking (3–5 kg/m2).

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Pellet diets ranging in diameter from 1.8–2 mm were found to be better for smaller lobsters in terms of wastage and consumption while pellets of 2 mm and 2.2 mm size were better for bigger lobsters. 2.4.2.3.2 Sea-cage operations India has initiated a series of studies on the aquaculture of spiny lobsters (Anon 1998). These include breeding programmes aimed at induction of gonad maturation using hormonal and environmental manipulations, and also larval rearing experiments to shorten larval development and promote moulting through the administration of various moultrelated hormones (Radhakrishnan et al. 1999). Feed trials are being investigated to enhance growth of juveniles (Raj et al. 1998), and sea ranching trials are also being held to study survival and growth of juveniles (Achary et al. 1998). In 2010 an International Conference on Recent Advances in Lobster Biology, Aquaculture and Management was held in Chennai, India. This provided an opportunity to identify a range of research that has been conducted in India over the last few years on various aspects of spiny lobster aquaculture. Some of these studies were presented at the conference, but some were published. These have been incorporated into the References to this chapter and can be categorised as studies principally concerned with Transportation of spiny lobsters (Immanuel et al. 2006; Verghese et al. 2008; Jayagopal & Vijayakumaran 2010); Growout conditions (Vijayakumaran et al. 2005; Radhakrishnan et al. 2007, 2009; Anbarasu et al. 2010; Balaji et al. 2010; Ganesh et al. 2010; Jha et al. 2010; Leema et al. 2010; Magesh et al. 2010; Muthu Rathinam et al. 2010; Remany et al. 2010; Vijayakumaran et al. 2010); Health and disease (Verghese et al. 2007; Leema et al. 2009; Vijayan et al. 2010a,b); Review of prospects for spiny lobster aquaculture (Senthil Murugan et al. 2005; Radhakrishnan 2006; Fotedar & Phillips 2010; Radhakrishnan et al. 2010); Growout trials (Thiruvengadam & Srikrishnadhas 2007; Vijayakumaran et al. 2007; Kizhakudan et al. 2010; Gulshad et al. 2010; Kizhakudan 2010; Rao et al. 2010; Thampi et al. 2010; Vijayakumaran et al. 2010). 2.4.2.4 Vietnam Marine cage culture of spiny lobsters began in the 1980s. Vietnam is already producing, in 40,000 sea-cages, more than 2,000 tons (2008) of cage-raised P. ornatus, P. homarus homarus, Panulirus stimpsoni and Panuliris longipipes longipes annually (worth about $US100 million) which are exported to China, Japan, Hong Kong and Thailand (Tuan et al. 2000; Tuan & Mao 2004; K. Williams, personal communication). These lobsters can be grown to 1 kg in 18–24 months from puerulus (Fig. 2.11a,b). Jones (2007a) reviewed the growout system in Southeast Asia, particularly Vietnam, where the focus of interest and development for lobster culture is on P. ornatus. The growout phase that follows extends over some 18–20 months to enable growth to 1 kg. Cages are of various dimensions but generally between 3 × 3 m and 6 × 6 m, and 3 to 5 m deep. In the early days, the cages were tied to timber posts fixed into the bottom in relatively shallow water, but most farms are now further offshore in much deeper water, and the cages are tethered to floating frames. The industry exclusively uses fishery bycatch for the food, which is supplied fresh to the farm each day and is fed 1 to 2 times per day. There are currently about 40,000 growout cages dedicated to lobster, the bulk of which are of the floating cage variety. Annual production for 2007 was 1,400 tons valued at around $US50 million, but it is understood that the catch in 2008 was approximately 800 tons more. Each

A Global Review of Spiny Lobster Aquaculture 55

(a)

(b) Fig. 2.11 (a) Growout facilities in Vietnam. (b) Typical examples of the Panulirus ornatus grown in the floating cages.

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cage will have on average 60 to 80 lobsters at harvest. Mortality through the nursery phase is high, but during growout is less than 10%. During 2008, the cost of seed was particularly high due to reduced catch. Prices as high as $US17 per puerulus were paid, although the average across the season was around $10 to $12. Juveniles fetch a higher price than pueruli. The prices are driven by supply and demand, and although they seem high the industry is profitable, and has had a significant beneficial impact on poverty in many coastal areas. On 9–10 December 2008 an international symposium was held in Nha Trang, Vietnam, on ‘Spiny lobster aquaculture in the Asia–Pacific region’. Twenty papers, 16 reporting Australian Centre for International Agricultural Research project research, were presented in four sessions with the following themes: sustainable lobster aquaculture; improving lobster nursery culture; lobster growout culture systems; and lobster growout feeds and feeding practices (Williams 2009). 2.4.2.5 Cuba Perera (2008) reported that experiments have been conducted studying the growout of P. argus juveniles. These included studies of lobster diets that included gastropods, pelecypoda and crustaceans to provide efficient use of dietary proteins (Diaz-Inglesias et al. 2002), and pellets as an alternative to fresh foods (Perera & Diaz-Inglesias 2004). A study to evaluate practical diets was conducted by Perera et al. (2005), who reported that squid and fishmeal diets for lobsters increased the nutritional value of the diet for P. argus, but growout trials are needed to evaluated whether these protein sources in formulated diets improve growth sufficiently enough to warrant using them. Another group is studying the immune system of P. argus (Perdomo-Morales et al. 2007, 2008), and other studies of its main defence mechanisms are being conducted. Perera et al. (2008a,b) studied changes in the digestive enzymes through developmental and moult stages in P. argus. 2.4.2.6 Singapore Cage farming of Panulirus polyphagus is being conducted (Chou & Lee 1997). 2.4.2.7 South Africa Hazell et al. (2000) studied the effect of temperature on growth of juvenile J. lalandii. Juveniles at 15 °C had growth rates twice as fast as those at 10 or 19 °C. Temperature influenced intermoult period, but food had no effect on moult increment. Aquaculture of J. lalandii has been considered but is hindered by a lack of data on suitable growout conditions. Dubber et al. (2004) studied the effects of temperature and diet on its survival, and also the quantity of food uptake. An optimal temperature of 18 °C for survival was determined for J. lalandii. Esterhuizen (2004) developed a protocol for rearing juvenile J. lalandii. The objectives of the project were to investigate the nutritional requirements, as well as the effect of stocking density and tank design on growth and survival of J. lalandii. The results indicate that density had a significant effect on growth and survival of post-pueruli. An initial stocking density of 75 post-pueruli/m2 was regarded as optimal both in terms of the growth rate and biomass production per tank.

A Global Review of Spiny Lobster Aquaculture 57

Paulet et al. (2007) and Paulet (2008) reported that they were working towards a sustainable, pelleted diet for Panulirus homarus rubellus. Progress to date includes the development and testing of preliminary diets and reformulation of diets based on those results. In addition, behavioural trials have been conducted in order to determine acceptability and preference of the various formulated diets. Reasonable growth and good survival have been attained, and are expected to improve as diet formulations become more specific. This research will include: a protocol for shore-based growout of lobsters, a description of the nutritional requirements for lobster culture, a contribution to knowledge of lobster behaviour and physiological requirements and an economic model for lobster culture. Kemp et al. (2009) examined ammonia excretion dynamics in the east coast rock lobster Panulirus homarus rubellus. 2.4.2.8

Taiwan

Pond raising of wild-caught juveniles of P. japonicus and P. polyphagus is undertaken in Taiwan in ponds designed for shrimp (Wickens & Lee 2002). Chen (1990) reported that one of the largest farms on Taiwan has 1.3 ha of water with the capacity to produce about 150,000 market-size lobster a year. In 1987 there were about 20 lobster farms in Taiwan, which produced about 400,000 lobsters. This quantity could hardly meet the demand of consumers, and because lobster culture is limited by seed supply there was mass importation of lobsters from different parts of the world. Some were imported from Singapore and Hainan Island (PR China) but the largest supply of live lobsters was from mainland (PR China), whose fishermen trade live lobsters for money or goods with Taiwanese fishermen at sea. Things have clearly changed. No production has been reported from Taiwan since 2001 by FAO and P. homarus spp. is stated to be one of the developing aquaculture species (Cin Liao & Chao 2008). 2.4.2.9

Belize & Bahamas

Brown et al. (1995) reported evaluations in the Bahamas of a water-stable feed for the culture of P. argus. Dalhgren and Stains (2007) reported on experimental growout aquaculture of P. argus in Belize and the Bahamas. Growth and survival rates from the Bahamas suggested that if an adequate supply of pueruli is available, large-scale growout may be possible, as it was possible to raise the pueruli to legal size in less than18 months. There were problems with disease in the Belize trials. 2.4.2.10 Malaysia The culture of spiny lobsters in Malaysia is still in its infancy, with activities based in the State of Sabah – mainly concentrated in Darvel Bay area. Spiny lobster culture involved both growout of juveniles in sea-cages and holding of adults in cement tanks and sea-cages. Seed are obtained from local waters as well as from neighbouring countries. In Darvel Bay, spiny lobsters are reared in sea-cages at depths of 10–20 m. Food consists of chopped trash fish, given 1–2 times daily (Biusing & Fui Lin 2004).

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2.4.2.11 Philippines Growout industries exist in the Philippines for several tropical species and are supported by research programmes by ACIAR in Australia. Other than Vietnam, the Philippines was the largest producer as of 2009.

2.4.2.12 USA Panulirus argus Lellis (1991) at the Harbor Branch Oceanographic Institution (Florida) suggested growout of P. argus could be a useful approach in the Caribbean, and conducted some growout trials which indicated optimal growth at 29–30 °C. Pardee and Foster (1999) conducted some growout experiments of P. argus using four diets, at two densities, to examine how the costs of feeding and facilities might affect economic viability of an aquaculture enterprise. Dahlgren and Stains (2007) reported on experimental growout aquaculture of P. argus in Belize in 2003 and 2004. A major problem with a virus infection, DNA virus (PaV1) (identified by Shields & Behringer 2004), also found in wild populations of P. argus, together with other unknown disease problems, essentially wiped out the experimental population of lobsters. Research in Puerto Rico was funded by the National and Atmospheric Administration’s Small Business Innovation Research programme in 2004 and a further grant from NOAA is being used to evaluate cage-culture of P. argus by a firm called ‘Snapperfarm Inc’. This has now expanded considerably. Davis et al. reported that beginning in February 2005, Harbor Branch Oceanographic Institution, Snapperfarm, Inc. and several collaborators began a NOAA SBIR Phase II project in Culebra, Puerto Rico, to: a) develop optimal methods to collect pueruli on Witham collectors in the vicinity of fish sea-cages, and b) develop methods to grow lobsters from pueruli to market size in submerged sea-cages. Results for the first seven months show that the optimal depth of collection of pueruli in the Vieques Passage, Puerto Rico is between approximately 15–26 m with minimal pueruli collection at approximately 3 m depth. Peak recruitment pulses occurred during the summer months, particularly in June with 93 pueruli collected off one Witham collector located at approximately 26 m. The highest total numbers of pueruli were collected in July; 474 pueruli were collected from 19 collectors located in water depths of approximately 15– 26 m. Submerged sea-cage prototypes for juvenile growout were also due to be tested. These results demonstrate that it is possible to collect a large number of spiny lobster pueruli to culture to marketable size as an additional source of income for fish sea-cage culture in Puerto Rico. The Harbor Branch Oceanographic Institution is already offering training in spiny lobster culture methods, so the development of the industry is accelerating. Poseidon Science is another company in the USA conducting research on growout of spiny lobsters and offering its services on the internet. Perera et al. (2007) studied the metabolic rate of P. argus. Their results suggested that the inclusion of squid and high-quality fishmeal in local fishmeal diets increases the nutritional value and that squid enhances digestive protease activities in the hepatopancreas. However, growout trials are needed to fully demonstrate the growth-enhancing effect of these protein sources in formulated diets for juvenile P. argus and to decide whether the growth rate increase is sufficient to warrant using these protein sources.

A Global Review of Spiny Lobster Aquaculture 59

The effect of feeding frequency on growth was studied by Cox and Davis (2006), and Cox and Davis (2009) evaluated seven potential diets for the culture of post-pueruli of P. argus. The results demonstrated that a seafood-based juvenile formulation produced the fastest growth rate over a 28-day period. The composition of the seafood-based diet is given in Table 4 in an article by Cox and Davis (2009). This is a good start, but a lot more testing needs to be undertaken.

2.4.2.13 Indonesia Lobster culture in eastern Indonesia was first established in 2000 as a by-product of seaweed and grouper culture, which had been in operation since the 1990s. Puerulus and were often observed settling on the floats, cages and other materials associated with seaweed and grouper culture. They were captured by hand and retained in separate cages, in which they grew well, and thus was born lobster aquaculture (Jones 2007a; Priyambodo 2009; Priyambodo & Jaya 2010). Production of P. homarus spp. at Lombok is about 100 tons per year. P. ornatus and P. polyphagus are both being examined for growout at Lombok but this is in the developmental stage. This research is linked to research in Australia and Vietnam through funding from the Australian Centre for International Agricultural Research, Canberra, Australia. About 1,000 small-scale growout industries already exist in Indonesia.

2.4.2.14 British Virgin Islands Power et al. (2005) made a preliminary examination of the feasibility of commercial aquaculture of P. argus based on pueruli collections. They also tested methods of collection of pueruli, sources of local food for juvenile lobsters and holding systems, and determined the rearing period to reach minimum market size.

2.4.2.15 Namibia Namibia is interested in the possibility of the growout of J. lalandii and investigations to assist this were conducted as part of PhD studies (Keulder 2005). Studies of the level of pueruli settlement were conducted. These studies indicated that there was considerable interannual variation in the levels of pueruli settlement in the area studied, and this would limit production unless other sources of pueruli could be located. Pilot culture of rock lobster J. lalandii began in Lüderitz (Namibia) in 2007 and has been flourishing since then. Lobster culture in Lüderitz has been represented by an experimental station, which employs a total of 12 people on a permanent basis and 16 casual employees (New Era 2009).

2.4.2.16 Turks & Caicos Newspaper reports on 16 February 2010 indicated that scientists from the Darden Restaurant Company would spend the next five years, from 2010, determining if the Turks and Caicos is a suitable environment for the cultivation of P. argus. Darden has already spent 18 months conducting preliminary research in the South Caicos.

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2.4.2.17 Mexico Mexico has a stabled seed collection and commercial growout facility for P. argus on Isla Mujeres, Quintana Roo. Preliminary studies in 2008 and 2009 were conducted on the possibility of the aquaculture of Panulirus interruptus on the Pacific coast. Methods to capture puerulus were tested and the captured animals were grown out near El Rosario. Juveniles reached 220 g in 12 months and 350 g in 18 months with good survival (Diaz-Inglesias E., CICESE, Ensenada, Mexico, personal communication). 2.4.2.18 Other countries Other countries that are believed to have some involvement or interest in spiny lobster aquaculture include Ireland, the People’s Republic of China, the United Kingdom, Spain, the Seychelles, Sri Lanka, Iran, Saudi Arabia, Thailand, Burma, Croatia, Tahiti, Mexico and Norway.

2.4.3

Economic feasibility for puerulus and juvenile growout

The costs of larval culture or capture of pueruli from the wild, as well as growout tanks, food for the post-puerulus and juvenile lobsters, the impact of diseases and mortality of the lobsters during the holding period, and the quality of the lobsters at the time of submission to the market, are all matters to be investigated before a viable industry can be established. Food may make up to 50% of production costs in aquaculture; therefore development of a suitable pelleted diet is crucial for commercially viable spiny lobster aquaculture. The food must have high nutritional value and acceptance, be available year round, easy to store and handle, and be reasonably priced (Booth & Kittaka 2000). Jasus edwardsii Jeffs and Hooker (2000), using results from experimental growout of J. edwardsii from the previous 20 years in New Zealand, assessed the economic feasibility of commercial spiny lobster aquaculture in temperate waters. The assessment indicated that while the culture of J. edwardsii in land-based tank systems is biologically feasible, it would only be commercially viable if it were possible to reduce infrastructure and operating costs (feed, freight and packaging, labour, electricity and puerulus collection costs). One possibility to reduce infrastructure costs is the use of sea-cage culture or sea-ranching. The assessment also highlighted the need for a cost-effective artificial feed to improve production costs. Feed costs have the potential to be reduced substantially, with cheaper foods than farmed mussels providing improved food conversion. Low feed consumption has been suggested as a major impediment for development of effective formulated diets. Simon and Jeffs (2008) studied feeding and gut evacuation rates in J. edwardsii and reported that a slow appetite revival and difficulties in processing and digestion of formulated diets appear as major issues to be resolved to improve the performance of formulated diets for spiny lobsters. Simon (2009a) studied feed intake and its relation to foregut capacity in juvenile J. edwardsii. The results indicated that J. edwardsii juveniles have a small foregut capacity (2.5–3% BW) that limits food intake when diets are fed every 48 hours. There appears to be no advantage in dry matter intake by providing the nutrient-dense dry formulated diet of this study compared with mussel flesh. Formulated

A Global Review of Spiny Lobster Aquaculture 61

diets would need to be fed more frequently and be highly digestible if they are to deliver sufficient nutrition to maximise growth for commercial aquaculture. Ward and Carter (2009) made a study of the nutritional value of alternative lipid sources to juvenile J. edwardsii. Jasus lalandii An economic assessment of the viability of raising the pueruli was conducted in South Africa by Esterhuizen (2004). The study examined the economic feasibility by modelling a hypothetical shore-based rock lobster farm. A projection of production costs and revenues was based on the typical costs of a shore-based abalone farm and the current market prices for wild harvested J. lalandii. The economic viability was evaluated using benefit–cost ratios, payback period, internal rate of return and breakeven analysis. Sensitivity analyses revealed that the projected lobster growth and survival rates were the main biological factors influencing the economic feasibility of the hypothetical rock lobster farm. An assumed four-year growout period at a low stocking density yielded more lucrative internal rate of return (IRR), benefit–cost ratio, payback period and net present values (NPV) than a five-year growout period at a high density. The four-year growout scenario proved to be more robust to the fluctuating SA rand/US$ exchange rate and could accommodate a lower lobster survival rate. Results presented in this study indicate that rock lobster farming is a marginal commercial prospect based on current production performance and costs. Panulirus ornatus Linton (1998) reviewed the data on the potential for tropical rock lobster aquaculture in Queensland (Australia) and suggested that P. ornatus was a likely candidate. Jones et al. (2001) reported that P. ornatus post-pueruli could be raised from 3 g to 1 kg within 18 months and that this species has an excellent potential for commercial aquaculture. Jones (2009a) reported on the temperature and salinity tolerances of this species. Jeffs and Davis (2003) reported that P. ornatus in Southeast Asia and P. argus in the Caribbean have the highest potential for aquaculture due to their high growth rates. In a comparative diet study with green-lipped mussels, Smith et al. (2005) reported that P. ornatus grew well when fed high protein, high lipid, pelleted feeds. Development of an artificial diet for growout of pueruli and juveniles of all species is a very high priority. Considerable research has focused on J. edwardsii and P. ornatus. Smith et al. (2003) and Williams (2007) provided substantial reviews of the nutritional requirements and feeds development for juvenile spiny lobsters of both species. Irvin and Williams (2007) examined the apparent digestibility of selected marine and terrestrial feed ingredients for tropical spiny lobster Panulirus ornatus. Simon (2009b) identified digestible carbohydrate sources for inclusion in formulated diets for juvenile J. edwardsii. Huynh and Fotedar (2010) studied the effects of dietary mannan oligosaccharide (MOS) (Bio-Mos®, Alltech, USA) on the growth, survival, physiology, bacteria and morphology of the gut and immune response to bacterial infection of tropical rock lobsters P. ornatus juveniles. Dietary inclusion level of MOS at 0.4% was tested against the control diet (trash fish) without MOS inclusion. At the end of 56 days of rearing period, a challenged test was also conducted to evaluate the bacterial infection resistant ability of the lobsters fed the two diets. Lobster juvenile fed MOS diet attained 2.86 ± 0.07 g of total weigh and 66.67 ± 4.76% survival rate which were higher (P < 0.05) than the lobsters fed control diet (2.35 ± 0.14 g total weight and 54.76 ± 2.38% survival rate, respectively) thus providing the higher (P < 0.05) specific

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growth rate (SGR) and average weekly gain (AWG) of lobsters fed MOS diet. Physiological condition indicators such as wet tail muscle index (Tw/B), wet hepatosomatic index (Hiw) and dry tail muscle index (Td/B) of the lobsters fed MOS-supplemented diet were higher (P < 0.05) than that of the lobsters fed the control diet. Bacteria in the gut (both total aerobic and Vibrio spp.) and gut’s absorption surface indicated by the internal perimeter/external perimeter ratio were also higher (P < 0.05) when the lobsters were fed MOS diet. Lobsters fed MOS diet were in better immune condition, shown by higher THC and GC, and lower bacteraemia. Survival, THC and GC were not different among the lobsters fed either MOS or control diet after 3 days of bacterial infection, while bacteraemia was lower in the lobsters fed MOS diet. After 7 days of bacterial infection the lobsters fed MOS diet showed higher survival, THC, GC and lower bacteraemia than the lobsters fed the control diet. The experimental trial demonstrated the ability of MOS to improve the growth performance, survival, physiological condition, gut health and immune responses of tropical spiny lobsters juveniles. Panulirus homarus homarus and P. ornatus Rahman and Srikrishnadhas (1994) reported that both P. homarus homarus and P. ornatus could be economically cultured in India from wild-caught juveniles. Techniques to achieve this are being investigated, including propagation of the larvae to produce pueruli. Panulirus argus Fraga et al. (2000) evaluated the protein level requirements in the growth of juvenile (90 g) P. argus using isocaloric and isolipidic diets. Best growth rates over the 90-day trial were obtained with 30 and 35% protein diets, having the highest number of moults and length increment per moult. Survival was high for all treatments (80–100%). Carvalho and Ogawa (2000) reported on the economic assessment of spiny lobster growout in marine cages in Brazil. Depreciation of the cages was the major cost item, followed by cost of storing harvested lobsters. Despite this, results indicated the economic feasibility of the activity in this country. Although Jeffs and Davis (2003) reported that P. ornatus in Southeast Asia (together with P. argus in the Caribbean) have the highest potential for aquaculture due to their high growth rate, they also pointed out that feed is likely to make up more than 25% of the production costs. The latter is probably the single most important obstacle to large-scale commercial aquaculture development of P. argus in the USA. Cruz et al. (2006) examined large-scale collection of pueruli of P. argus from the wild for growout. The results indicated a cost of about US$1.50 per puerulus and US$4.14 per juvenile, which was expensive. In Cuba juvenile mortality is not high (42% after settlement) and based on these data they estimated that juvenile removals from the wild would have a negative impact on pre-recruits and subsequent commercial catches. Jeffs et al. (2007) reviewed the available post-larval survival data for P. argus species in the wild and found that there was substantive evidence that the harvesting of seed lobsters for aquaculture could increase overall lobster production, whilst maintaining sufficient seed lobsters to maintain wild fisheries. However, a review of the available information on the patterns of seed lobster settlement suggests that there is a lack of information on the physical processes, as well as the behaviour and ecology of P. argus post-larvae that would be useful for directing the efficient harvesting of seed lobsters for aquaculture. Jeffs and Davis (2008) made a comprehensive report on the potential for harvesting seed of P. argus for aquaculture.

A Global Review of Spiny Lobster Aquaculture 63

Panulirus cygnus Melville-Smith et al. (2009) determined that western rock lobsters have many biological attributes that are consistent with their suitability for aquaculture. Most significantly, postpueruli, year 1 and year 2 post-settlement juveniles can be stocked at very high densities (up to 100 m−2 for post-pueruli) without adverse effects on growth. This species tolerates low flow rates and high ammonia concentrations and has significantly faster growth rates at 23 °C compared with ambient temperatures, without impacting the survival of postpueruli and only minimally impacting the survival of older juveniles. Male pueruli held at 23 °C can potentially reach legal size (76 mm CL) within 2.3 years, whereas females can reach legal size within 2.5 years. These are substantially faster growth rates than wild lobsters, which reach legal size in approximately 4 years. Lobsters of this species consumed formulated pellet diets reasonably well, but superior growth was achieved when supplemented with fresh mussels. A novel rigid plastic mesh shelter design significantly improved survival compared with the brick shelters that have been used in many other growout studies. A number of aspects require further work before P. cygnus culture would be a commercial proposition. Development of a more palatable, nutritionally complete diet specific for P. cygnus is essential for the economic viability of commercial operations. Commercial scale trials need to be conducted to determine optimum tank specifications for commercial culture. An economic analysis needs to be undertaken to identify those parameters that are most sensitive to the profitability of a western rock lobster growout venture, so that it can be used to focus future research on the economically critical issues. Previous studies have shown that large numbers, potentially of commercial quantities, of pueruli and post-pueruli can be harvested from Western Australian waters. At this stage, this appears to be unique in Australian waters. A commercial attempt to catch pueruli of P. cygnus for growout and to establish a pilot project for lobster aquaculture at Jurien in Western Australia was established in 2006 but went out of business in 2008.

2.4.4

Conflicts with fishers and ecosystem considerations

From studies on P. argus and P. cygnus there is known to be a high mortality of pueruli and post-pueruli, particularly in the first year after the settlement of the puerulus (Herrnkind & Butler 1994; Phillips 2003). However, the possible impacts on the wild fisheries of removing pueruli and post-pueruli from the wild for the purposes of aquaculture need to be assessed to avoid conflicts with the wild-caught lobster fisheries, particularly in years of low puerulus settlement (Kittaka & Booth 1994, 2000). In Australia, a major study to assess this question was undertaken for P. cygnus and the results were published by Phillips et al. (2003). These authors concluded that the impact of P. cygnus pueruli removals for aquaculture would be slight, mainly because of the high natural mortality in the wild: 80–98% during the first year after settlement of the pueruli. In 1996 the New Zealand government approved the collection of young settled J. edwardsii for growout on marine aquaculture farms. For every ton of quota withdrawn from the commercial fishery for J. edwardsii, 40,000 pueruli and post-pueruli can be taken for aquaculture. Monthly catches in the thousands have been achieved (Booth et al. 1999). However, as of August 2000 only two farms were still in operation. De Zylva (2002) reported on the commercial farm at Napier that was just marketing its first crop of lobsters. More recently, there is mention of commercial partners in the new cage-growout studies,

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but no further information is supplied (James & Simon 2008). Similar approvals for collection and raising of J. edwardsii juveniles were given in Tasmania, Australia and a call for applications for seven rock lobster aquaculture trial permits was made in 2001 (Anonymous 2001; Treloggen 2001). To satisfy concerns from the fishers of the wild fishery, it was proposed by the government that after a year of culture (or if they have reached 35 mm long), a number of juveniles equal to 5% of the number of pueruli originally taken must be released, plus another 20% of the surviving cultured lobsters numbers (Hart & Van Barneveld 2000b; Mills et al. 2000). The success of this method is dependant on the successful survival of the released growout juveniles. Six applicants were licensed to capture and grow out pueruli and juveniles of J. edwardsii. However, these licences were eventually abandoned because the licence holders regarded the conditions imposed as unfeasible. Acoustic tracking showed that movement, behaviour and habitat choice were similar for wild-caught and on-grown J. edwardsii juveniles released into the wild. Modelling from released tagged juveniles showed high survival for re-seeded juveniles, with no detectable difference in survival over one month between wild or on-grown juveniles (Mills et al. 2000). In Belize, it is understood that an initial approval for growout of P. argus pueruli and juveniles was withdrawn after complaints from the wild-catching lobster fishing industry.

2.4.5

Reseeding of wild stocks

Conan (1986) comprehensively reviewed enhancement of lobster stocks. It involves all types of protection of the early life history stages to increase yields from the fisheries. As Conan pointed out, ‘References on recruitment enhancement are extremely scarce’. Japan has pursued a plan for stock enhancement of the Japanese wild stocks of P. japonicus and, in addition to encouraging culture of the larvae, has a programme of research into the best methods for safe release of pueruli and young juveniles into the wild. Studies in Tasmania, Australia, are being conducted on on-growing J. edwardsii pueruli to enhance rock lobster stocks while providing animals for commercial culture (Hart & Van Barneveld 2000a; Mills et al. 2000). Joint research between New Zealand and Australia has been conducted to assess if lobsters released for reseeding of wild stocks will survive and behave in a similar manner to wild lobsters. Several studies have been conducted but, according to studies by Mills et al. (2000), Mills et al. (2005), Oliver et al. (2005), Mills et al. (2006), survival rates of J. edwardsii released from aquaculture to the wild were equivalent to the survival of wild rock lobsters released in the same manner. Vaitheeswaran and Srikrishnadhas (2007) described efforts to enhance the stock of spiny lobsters in the wild. To conserve lobsters and to enhance lobster stock in the Gulf of Mannar in southern India, sea-ranching of spiny lobsters was undertaken. The spiny lobsters such as P. homarus homarus, P. ornatus and P. versicolor are commonly available in the Gulf of Mannar. Berried animals of these lobsters were collected from commercial catches and reared in the wet laboratory, hatched out phyllosoma larvae were sea-ranched in 08° 52′N 78° 15′E at Tuticorin on the coast of Gulf of Mannar. Thirty-five adult females P. homarus, two P. ornatus and seven P. versicolor were reared in the cages and all hatched out phyllosoma larvae. To enhance wild stock, 60, 16, 026, 3.25 and 6.87 millions of larvae, respectively, of P. homarus homarus, P. ornatus and P. versicolor were sea-ranched on the southeast coast of India.

A Global Review of Spiny Lobster Aquaculture 65

2.5

FUTURE DEVELOPMENTS

Propagation of larvae through to the puerulus stage, rather than capture and raising of wild pueruli, is likely to be the best short-term option. Complete aquaculture from egg to adult would allow for genetic or selective breeding. Growout studies of pueruli are useful for research purposes but the number of pueruli available for growout may be limited. Concern over the number that should be collected, particularly in years of low pueruli settlement, will always make this option difficult. Therefore, to ensure future expansion and sustainability of the rock lobster aquaculture industry, phyllosomas must be cultured through the full larval cycle to the post-puerulus stage. Hamaski et al. (2007) have identified the direction for mass culture of phyllosomas in Japan: To improve the survival rate and expand the scale of phyllosoma larvae, firstly, behavioural characteristics such as phototaxis and geotaxis, and sinking speed: then, control measures such as using an agitator similar to that which improved the larval survival rate in the snow cabs, should be investigated. Further, it is important to study bacterial diseases and their control measures.

Jones (2007b,c,d) reviewed the technical progress towards propagation of spiny (rock) lobsters in Australia. Interest in the development of rock lobster aquaculture has risen sharply in the past decade, and Australia has been at the forefront of research to develop propagation technology. A nationally coordinated research programme targeting rock lobster aquaculture was developed in 1999 involving multiple agencies. The research is market driven, and thus has focused on the four species that are currently fished in Australian waters and have strongly developed markets. Ideally, rock lobster aquaculture will involve closed-cycle culture, which is effectively independent of wild populations. This will involve managed production of the various life history phases including breeding, larval rearing, nursery and growout. Larval rearing has proven to be the most challenging. Like all rock lobsters, the Australian species have prolonged larval life histories that are significantly longer than any commercially successful aquaculture species in the world. Three of the species of interest (J. edwardsii, P. ornatus and S. verreauxi) have now been successfully cultured from egg through to puerulus, as a direct result of the research programme. Nevertheless, the magnitude of the technical challenge to develop commercially applicable methods that enable consistent, economically viable, larval production is not to be underestimated. Phyllosomas are extremely delicate and accustomed to the pristine environmental conditions of the open ocean. Simulating such conditions in the hatchery is clearly achievable, but maintaining suitable water quality over long periods is very difficult. Similarly, provision of appropriate nutrition is equally problematic and often in conflict with maintenance of water quality. On the basis of progress to date with P. ornatus in Australia, the R&D effort is now moving to a commercialisation phase. Commercial partnerships have been established with aquaculture companies to take the research forward. We believe another 3 to 5 years of research will be necessary to provide commercially robust technology, and even if that is successful there is still the issue of economics. Can lobsters be propagated for a reasonable and cost-effective price? Once we can produce some consistency in rearing larvae through to the puerulus stage, it will be possible to consolidate the economic data to answer that question.

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India, Thailand, Burma, Taiwan, Singapore, the Philippines and Vietnam are all involved in commercial spiny lobster aquaculture operations to date. All of these operations are based on catching pueruli from the wild and raising or on-growing pueruli to a commercial-sized lobster. The costs of infrastructure development, collection and raising of pueruli and small juveniles has been discussed by a few authors including Linton (1998) and Jeffs and Hooker (2000), but a lot of the current information is ‘commercial in confidence’. Therefore it is difficult to assess the real costs of spiny lobster aquaculture at this time. In Australia, J edwardsii and P. ornatus remain the main target species for complete aquaculture. The following is a public statement available on the website of the Marine Research Laboratories, University of Tasmania, which was selected by the Fisheries Research and Development Corporation to lead the national project on propagation. The statement was made in 2006. Considerable progress has been made towards developing an efficient method of capturing pueruli from the wild for on-growing. Commercial puerulus collectors were developed that were lightweight and easily deployed. Coastal areas were surveyed to identify where pueruli settle. Juvenile rock lobsters can now be cultured to a market size of 250 g in two years. Artificial diets were developed, although a diet of fresh mussels produced the best growth rates. Adjusting the level of carotenoids in the diet was found to control the colour of the cultured lobsters. Lobsters grow slowly during winter at <12 °C suggesting the use of heated water will be necessary, at least for part of the year. Indeed, they can be grown at temperatures of up to 22 °C without adverse effects on growth, survival or feeding efficiency. To better design and manage culture systems it is necessary to understand the specific behaviours of cultured lobsters. These have been identified, characterised and quantified under a range of stocking densities, lobster sizes and shelter availability. The research conducted over the past five years has shown that juvenile rock lobsters can be farmed to market size in sea cages or in land-based tanks. However, the supply of the puerulus or juvenile stages has been unreliable. Therefore, to achieve a sustainable lobster farming industry it is necessary to close the life cycle and culture animals from eggs. Lobsters have a long larval phase and for southern rock lobsters it can take up to 2 years in the wild for the phyllosomas to develop through the 11 morphological stages from egg hatch until metamorphosis to the puerulus. Lobster broodstock hatch their eggs over a brief period in the spring, limiting the time during which research on larval rearing is conducted. Nowadays, the breeding season can be extended, by holding animals in altered water temperature and light to mate and hatch at other times of the year. Larval quality is being assessed from different groups using an activity test developed to measure larval viability at hatch. No matter what the broodstock background, application of the rapid test ensures that only the best larvae are used for extended hatchery rearing. In 2005, a major breakthrough in the project occurred with the culture of larvae through all the phyllosoma stages to the pueruli and then to the juvenile of J. edwardsii in a little over one year. This is the first time this has been achieved in Australia and the first time it has been achieved on any spiny lobsters species anywhere in the world without the use of antibiotics. The current research in Tasmania focuses on minimising the disease-causing microorganisms, which typically afflict larvae in high intensity culture, mostly as a result of feeding live Artemia. Various disinfection treatments, including ozonation and probiotics, have been found to be effective in reducing bacterial contamination in the culture water, Artemia and the larvae and pueruli. The optimum size and density of Artemia have been determined

A Global Review of Spiny Lobster Aquaculture 67

and their nutritional quality has been improved with a range of enrichments. However, in the longer-term, the goal will be to develop cleaner alternative formulated diets. This research is being conducted in collaboration with researchers in Japan, New Zealand and Queensland.

The above information has been updated by Buxton et al. (2007). The potential for spiny rock lobster aquaculture (temperate and tropical) species is of significant international interest. The rapid expansion of a new sector based on the capture and on-growing of P. ornatus seed stock (pueruli and juveniles) in Vietnam, with live product destined for the north Asian market, is an indication of the worldwide potential for rock lobster farming. In Australia, several species are of interest because of their strong market demand, although suitable aquaculture technology is yet to be developed. The possibility of collection of wild seed for ranching has been examined, but is generally unpopular because of concerns about sustainability and possible impingement on associated wild fisheries. In New Zealand, commercial collection and on-growing at several locations was initially successful but then faltered because of poor settlement on collectors. The alternative, hatchery propagation of lobsters, has been pioneered by Japan, but is now the focus of several laboratories around the world, and particularly in Australia. Should hatchery production of lobsters be successful, the nature of the subsequent growout has yet to be determined. Buxton et al. (2007) indicated that the most important areas for research in Australia are:

• • •

To understand disinfection and water treatments to further reduce microbial problems – maximise the antibacterial activity while suppressing toxic residues To further develop systems to allow scaling up for semi-commercial production To examine formulated diets as substitutes or supplements to live Artemia and seafood.

Jones (2007a) reviewed the growout of P. ornatus pueruli and juveniles in Southeast Asia and suggested that the industry faces a number of significant problems. Environmental degradation from the farms is increasing and this is leading to increased incidence of disease. The seed catch has fallen for the first time and may not be sustainable. Mortality of the early stages is severe and must be curbed. There has been some increased interest in alternative species, but none of these has the value of P. ornatus. To address these problems, and to develop growout technology that suits Australian circumstances, a collaborative project was developed between Australia and Vietnam within the Australian Centre for International Agricultural Research, which is part of Australia’s foreign aid programme. This project is now well advanced in generating results, with benefits flowing to both Vietnam and Australia. Growout of pueruli is now well established in many countries, especially Vietnam, the Philippines and Indonesia, and these will be joined by other countries in due course. However, in spite of the large amount of research conducted and the significant advances in recent years, commercial aquaculture of spiny lobsters through propagation of the larvae is not yet taking place. However, with the recent developments becoming more widespread, and species such as P. ornatus likely to have a larval period under culture of only three months, spiny lobster aquaculture offers such a high financial reward for success that efforts will continue and expand.

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The most recent reviews of the subject are those of Jones (2009b), Jeffs (2010) and Rogers et al. (2010). Rogers, the Chairman of Lobster Harvest Pty Ltd, expects to commence commercial hatchery production of Panulirus ornatus within 2 to 3 years. The Australian Institute of Marine Science in Australia was awarded an $A55 million grant by the Australian Government in 2009, most of which will be used to assist the Institute’s efforts to develop spiny lobster aquaculture (Thyer 2009).

2.6

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A Global Review of Spiny Lobster Aquaculture 83 Thomas, C., Carter, C. & Crear, B.J. (2002) Potential use of radiography in measuring feed intake of southern rock lobster (Jasus edwardsii). Journal of Experimental Marine Biology & Ecology, 273, 189–198. Thomas, C., Carter, C. & Crear, B.J. (2003) Feed availability and its relationship to survival, growth, dominance and agonistic behaviour of the southern rock lobsters, Jasus edwardsii in captivity. Aquaculture, 215, 45–65. Thyer, R. (2009) Farmed lobsters face a delicate larval journey. Fish, September 2009, 21–22. Tolomei, A. (2000) Chemoattractants and their role in the feeding behaviour, growth and survival of Jasus edwardsii. Honours thesis, School of Aquaculture, University of Tasmania. Tolomei, A.J. (2006) Microbial control in southern rock lobster Jasus edwardsii and Artemia. PhD thesis. School of Aquaculture, University of Tasmania. Tolomei, A., Burke, C., Crear, B. & Carson, J. (2004) Bacterial decontamination of on-grown Artemia. Aquaculture, 232, 357–371. Tolomei, A., Crear, B. & Johnston, D. (2003) Diet immersion time: effects on growth, survival and feeding behavior of juvenile southern rock lobster, Jasus edwardsii. Aquaculture, 219, 303–316. Tong, L.J., Moss, G.A., Paewai, M.P. & Pickering, T.D. (1997) Effect of brine-shrimp numbers on growth and survival of early stage phyllosoma larvae of the rock lobster Jasus edwardsii. Marine & Freshwater Research, 48, 935–940. Tong, L.J., Moss, G.A. & Paewai, M.P. (2000a) Effect of brine shrimp size on the consumption rate, growth, and survival of early stage phyllosoma larvae of the rock lobster Jasus edwardsii. New Zealand Journal of Marine & Freshwater Research, 34(3), 469–474. Tong, L.J., Moss, G.A., Paewai, M.P. & Pickering, T.D. (2000b) Effect of temperature and feeding rate on the growth and survival of early and mid-stage phyllosomas of the spiny lobster Jasus edwardsii. Marine & Freshwater Research, 51, 235–241. Tong, L.J., Moss, G.A., Pickering, T.D. & Paewai, M.P. (2000c) Temperature effects on embryo and early larval development of the spiny lobster Jasus edwardsii, and description of a method to predict larval hatch times. Marine & Freshwater Research, 51(3), 243–248. Treloggen, R. (2001) TRLFA news. Fishing Today, 14(1) February/March, 27–28. Tsvetnenko, E., Fotedar, S. & Evans, L. (2001) Antibacterial activity in the hemolymph of western rock lobster, Panulirus cygnus. Marine & Freshwater Research, 52(8), 1407–1412. Tuan, L.A., Nho, N.T. & Hambrey, J. (2000) Status of cage mariculture in Vietnam. In: Liao, I.C. & Lin, C.K. (eds.) Cage Aquaculture in Asia: Proceedings of the First International Symposium on Cage Aquaculture in Asia, Asian Fisheries Society, Manila, and World Aquaculture Society – Southeast Asian Chapter, Bangkok. Tuan, L.A. & Mao, N.D. (2004) Present Status of Lobster Cage Culture in Vietnam. In: Spiny lobster ecology and exploitation in the South China Sea region (ed. K.C. Williams). Nha Trang, Vietnam, ACIAR. Vaitheeswaran, T. & Srikrishnadhas, B. (2007) Sea ranching of spiny lobster for stock enhancement in the southeastern coast of India. 8th International Conference & Workshop on Lobster Biology & Management, Charlottetown, Canada, p. 38. Van Barneveld, R. (2001) Rock lobster enhancement and aquaculture in Australia. In: Australian Aquaculture Yearbook (ed. P. Shelley). National Aquaculture Council. Executive Media Pty Ltd., Melbourne. Van Barneveld, R. & Phillips, B. (eds) (2002) Developments in rock lobster enhancement, aquaculture and post harvest practices. Proceedings of the Fourth Annual Rock Lobster Post-Harvest Subprogram/ Rock Lobster Enhancement and Aquaculture Workshop, Cairns, Australia. FRDC, Canberra. Verghese, B., Radhakrishnan, E.V. & Padhi, A. (2007) Effects of environmental parameters on immune response of the Indian spiny lobster Panulirus homarus (Linnaeus, 1758). Fish & Shellfish Immunology, 23(5), 928–936. Verghese, B., Radhakrishnan, E.V. & Padhi, A. (2008) Effect of moulting, eyestalk ablation, starvation and transportation on the immune response of the Indian spiny lobster Panulirus homarus. Aquaculture Research, 79(9), 1004–1013. Vijayan K.K., Sharma, K. & Kizhakkudan, J. et al. (2010a) Gaffkemia (red tail disease) – an emerging disease problem in lobster holding facilities in southern India. International Conference on Recent Advances in Lobster Biology, Aquaculture and Management, 4–8 January 2010, Chennai. Abstract, pp. 61–62. Vijayan K.K., Sanil, N.K. & Krupesha S. (2010b) Health management concepts in lobster mariculture. International Conference on Recent Advances in Lobster Biology, Aquaculture and Management, 4–8 January 2010, Chennai. Lobster Research in India, 41–40.

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Vijayakumaran, M. & Radhakrishnan, E.V. (1984) Effect of eyestalk ablation in the spiny lobster Panulirus homarus (Linnaeus): 2. On food intake and conversion. Indian Journal of Fisheries, 31, 148–55. Vijayakumaran, M., Senthil Murugan, T., Remany, M.C., et al. (2005) Captive breeding of the spiny lobster, Panulirus homarus. New Zealand Journal of Marine & Freshwater Research, 39, 325–334. Vijayakumaran, M., Venkatesan, R., Senthil Murugan T., et al. (2007a) Farming of spiny lobsters in sea cages. 8th International Conference & Workshop on Lobster Biology & Management, Charlottetown, Canada. Abstract, p. 38. Vijayakumaran, M., Senthil Murugan, B.T., Remany, M.C. et al. (2007b) Growth of the deep-sea lobster, Palinustus waguensis in captivity. 8th International Conference & Workshop on Lobster Biology & Management, Charlottetown, Canada. Abstract, p. 121. Vijayakumaran, M.R., Venkatesan, R., Senthil Murugan, T., et al (2009) Farming of spiny lobsters in sea cages in India. New Zealand Journal of Marine & Freshwater Research, 43, 623–634. Vijayakumaran M., Anbarasu, M. & Kumar T.S. (2010) What helps tropical spiny lobsters to grow better in communal rearing – physical or chemical interactions? International Conference on Recent Advances in Lobster Biology, Aquaculture and Management, 4–8 January 2010, Chennai. Abstract, p. 41. Wade, N.M., Tollenaere, A., Hall, M.R. & Degnan, B.M. (2009) Evolution of a novel carotenoids– binding protein responsible for crustacean shell color. Molecular Biology & Evolution, 26, 1851–1864. Ward, L.R. (1999) Protein and protein: energy ratio requirements of the southern rock lobster, Jasus edwardsii. BSc (Hons) thesis, School of Aquaculture, University of Tasmania. Ward, L.R. (2005) Protein and lipid nutrition of southern rock lobster, Jasus edwardsii. PhD thesis, School of Aquaculture, University of Tasmania. Ward, L.R. & Carter, C.G. (2009) An evaluation of the nutritional value of alternative lipid sources to juvenile southern rock lobster, Jasus edwardsii. Aquaculture, 296, 292–298. Ward, L.R., Carter, C.G., Crear, B.J. & Smith, D.M. (2003) Optimal dietary protein level for juvenile southern rock lobster, Jasus edwardsii, at two lipid levels. Aquaculture, 217, 483–500. Webster, N.S., Bourne, D.G. & Hall, M.R. (2006) Vibrionaceae infection in phyllosomas of the tropical rock lobster Panulirus ornatus as detected by fluorescence in situ hydridisation. Aquaculture, 255, 173–178. Westbury, H.G.J. (1999) Agonistic behaviour and its relationship to stocking density, size and feeding regime in cultured juvenile southern rock lobster Jasus edwardsii. School of Aquaculture, University of Tasmania. Wickens, J.F. & Lee, D.O’C. (2002) Crustacean Farming: Ranching and Culture. Blackwell Science, Oxford. Williams, K. (2001) Rock lobster enhancement &aquaculture Subprogram Project 3: Feed development for rock lobster aquaculture. Final report of Project 98/303 to Fisheries Research and Development Corporation. (Australia). Williams, K. (2007) Nutritional requirements and feed development for post-larval spiny lobster: A review. Aquaculture, 263, 1–14. Williams K.C. (2009) Spiny lobster aquaculture in the Asia–Pacific region. Proceedings of an international symposium held at Nha Trang, Vietnam, 9–10 December 2008. ACIAR Proceedings No. 132. Australian Centre for International Agricultural Research: Canberra. Williams, K.C., Smith, D.M. & Irvin, S. (2000) Development of a dry pelleted diet for the tropical spiny lobster Panulirus ornatus. Abstract, Third World Fisheries Congress, Beijing. Yamakawa, T., Nishimura, M., Matsuda, H., Tsujigado A. & Kamiya, N. (1989) Complete larval rearing of the Japanese spiny lobster Panulirus japonicus. Nippon Suisan Gakkaishi, 55(4), 745. Yoshimura, T., Yamakawa, H. & Kozasa, E. (1999) Distribution of final stage phyllosoma larvae and freeswimming pueruli of Panulirus japonicus around the Kuroshio Current off southern Kyusyu, Japan. Marine Biology, 133, 293–306.

3

Slipper Lobsters

Manambrakat Vijayakumaran and Edakkepravan V. Radhakrishnan

3.1 INTRODUCTION Slipper lobsters belonging to four subfamilies, Ibacinae, Arctidinae, Scyllarinae and Theninae (Holthuis 1991, 2002) and are widespread in shallow, temperate and tropical seas (Baisre 1994). Among approximately 80 species of scyllarids (Holthuis 2002), only about 30 larger slipper lobster species are of commercial interest (Holthuis 1991). Species belonging to the genera Scyllarides, Thenus and Ibacus are used as food, although other species are used as baits (Spanier & Lavalli 2007). Some of the smaller species of scyllarids, such as Arctides regalis, Scyllarus americanus and Petrarctus rugosus, have commercial prospects in aquarium trade owing to their attractive forms and colours (see Spanier & Lavalli 2007; Sharp et al. 2007; Kumar et al. 2009). Slipper lobsters contribute less than 2% of the total lobster landings of about 227,000 tons (FAO statistics reported in Spanier & Lavalli 2007). Thenus sp. is the most economically significant species as it contributes towards many commercial trawl fisheries along the tropical coasts of the Indian Ocean and the Western Pacific region (Jones 2007). Catches of other genera such as Ibacus, Scyllarides, Paribacus and Evibacus throughout the Indian and Pacific oceans, and of Scyllarides in the Mediterranean, are generally negligible compared to Thenus landings (Holthuis 1991; Jones 2007). In the Western and Central Pacific, however, slipper lobsters contribute about 25 to 50% of total lobster catch. About 75 to 94% of the slipper lobster catch belongs to Thenus sp., comprising mainly a single species, Thenus orientalis (Fig. 3.1a,b). Modern molecular studies have revealed that there may be five or six species of Thenus (Burten & Davie 2007). Exploitation of Thenus sp. in Australia and Southeast Asia (Jones 2007) is as bycatch by trawlers. More than 90% of Thenus sp. landed in Australia, where it is popularly called Moreton Bay bugs or Bay lobsters, is from Queensland waters with a production ranging from 430 to 750 tons (DPI Brisbane 2002). Thenus orientalis is the only slipper lobster of commercial significance among the rich diversity of scyllarid lobsters recorded from the Indian coast (Radhakrishnan et al. 2007). They appear as by-catch in trawl fisheries and although catch rates are low they constitute the most important component of the lobster fishery on the northwest, southwest and

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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(a)

(b)

Fig. 3.1 Thenus orientalis, the most important commercially exploited slipper lobster for which aquaculture technology is available; (a) dorsal and (b) ventral view.

southeast coasts of India. In the northwest, along the Mumbai coast, the Thenus fishery collapsed in 1995 and has yet to recover, causing concern about the sustainability of the slipper lobster fishery (Deshmukh 2001). The other slipper lobsters of commercial importance in Australia are the Balmian bugs or Ibacus sp. About 200 tons of Ibacus sp. belonging to three species, Ibacus chacei, I. peroni and I. brucei (95% I. chacei and I. peroni) are caught in Australia (Haddy et al. 2007). The Galapagos slipper lobster, Scyllarides astori, contributes a minor fishery of 12–14 tons (see Spanier & Lavalli 2007), whereas Scyllarides latus are caught in very small quantities in the Mediterranean and Israel (Holthuis 1991). There is little or no information on other slipper lobster fisheries (Spanier & Lavalli 2007). Consistent with its insignificant fishery, the biology of slipper lobsters has attracted less attention compared to the commercially important nephropid and spiny/rock lobsters. Holthuis (1991) compiled all the taxonomical and biological features of slipper lobsters, reviewing the classification of the whole group. Recently, Lavalli and Spanier made a great contribution to present all available information on slipper lobsters in one place. The Biology and Fisheries of the Slipper Lobster, edited by these scientists and published by the CRC Press in 2007, is a compendium on the ecology, distribution, fishery and biology of slipper lobsters, and provides an insight into the future areas of research to be undertaken on slipper lobsters. Mikami and Greenwood (1997a) completed the life history of two species of Thenus: T. orientalis and T. indicus, and produced the post-larva nisto with 80% survival from egg in about 25–30 days. This feat has revived interest in slipper lobsters as

Slipper Lobsters 87

Thenus sp. is considered to be one of the most promising species for aquaculture. This chapter describes the prospects of slipper lobster aquaculture.

3.2 3.2.1

BIOLOGY Morphology and anatomy

Slipper lobster has the typical segmented body structure of a decapod crustacean with three distinct body regions, the cephalothorax (14 segments fused together), the abdomen (6 segments) and the telson (single segment) (Holthuis 1991). Each body segment bears specialised biramous appendages composed of an exopod and an endopod, except in few scyllarids that lack exopods in one or more maxillipeds. The cephalothorax contains the antennule (first antennae), the antenna, specialised mouthparts (mandible, maxillae (2) and maxillipeds (3)) and five pereiopods or walking legs. The abdomen bears five pairs of pleopods that are specialised for forward swimming in nistos and reproduction in adults and the last abdominal segment bears the uropods, which are modified pleopods (Lavalli & Spanier 2007). The bifurcated flagellum of the antennule of scyllarids is attached to a three-segmented peduncle, while the flagellum of the antenna, whip-like in nephropids and palinurids, is a single plate-like segment and is attached to five additional and highly flattened antennal segments (Holthuis 1991), a clear distinguishing feature for scyllarid lobsters. The internal body plan of scyllarids is not well described but is believed to be similar to that of other lobsters (Lavalli & Spanier 2007).

3.2.2

Life cycle

The females brood the eggs in their pleopods and release the fully developed phyllosoma larvae (Fig. 3.2), while a few scyllarids such as Thenus sp. release a short-lived naupliosoma that moults into phyllosoma within hours of release (Mikami & Greenwood 1997a). The naupliosoma has well developed appendages except for first maxillipeds and fourth and fifth pereiopods (Sekiguchi et al. 2007). As the phyllosoma develops, the appendages enlarge and new appendages develop at successive moults, but in all instars the abdomen is extremely small, relative to the cephalic shield and thorax. Pleopods are absent in phyllosomas (Johnston 2007; Lavalli et al. 2007). The branchial cavity and gill buds appear only in the last stage of phyllosoma, which is believed to respire through the entire body surface and osmo-regulate via the ventral surface of the cephalic shield (Haond et al. 2001). The gilled phyllosoma undergoes a metamorphic moult into the post-larval nisto, which looks more like an adult with varying sizes among species (3–20 mm carapace length (Sekiguchi et al. 2007). The nisto is an important phase in the life history of scyllarids and it settles into the benthic habitat. In some species, such as Thenus sp., the nisto settles into the typical adult grounds, while others settle in the juvenile ground and later travel to the adult habitat (Sekiguchi et al. 2007; Lavalli et al. 2007). While the phyllosoma swims with the help of its setose exopodites, the nistos use pleopods for forward swimming and use backward swimming or tail flipping as an escape mechanism. However, in a few adult slipper lobsters, pleopods are used for long-duration swimming (Lavalli et al. 2007; Jones 2007).

88 Recent Advances and New Species in Aquaculture

Phyllosoma

Nisto Gravid female

Adult Fig. 3.2 Life history of the slipper lobster Thenus orientalis. (Please see plate section for colour version of this figure.)

The juveniles and sub-adults of slipper lobsters are sparingly reported. In a few species larger juveniles and sub-adults are caught along with adults, while juveniles of T. orientalis are caught in shrimp trawls and gill nets (Kizhakudan et al. 2004a). The growth in the wild population is faster in some species such as Thenus sp. which grow to adults in two to three years, while other scyllarids grow more slowly, taking up to six years to mature (Hearn et al. 2007; Radhakrishnan et al. 2007; Haddy et al. 2007).

3.3

AQUACULTURE POTENTIAL

Since the beginning of the 21st century aquaculture has been diversifying, with a large number of new species and groups (e.g. sea urchins and other echinoderms) registering the largest growth (FAO 2006). Lobsters – clawed, spiny or slipper – are one of the most sought-after groups for mariculture owing to their high price, especially the live ones, which are valued twice as much as frozen in Southeast Asian markets. Breeding and hatchery production of seed, fast growth rate, feed and food conversion efficiency, hardiness of the animals in high-density culture, water quality, market value and production technologies are the main factors that make aquaculture a successful commercial venture. Notwithstanding the short larval life and the highly successful production of seeds

Slipper Lobsters 89

in hatchery, high cannibalism requiring individual compartments for growout has discouraged aquaculture of the clawed lobster (Homarus americanus) (Aiken & Waddy 1980). Spiny lobsters are hardy animals with good growth rates for juveniles, but the long larval life extending over several months, with limited success in production of seeds, has discouraged its large-scale aquaculture (Kittaka & Booth 2000). The alternative strategy adopted for spiny lobster culture was large-scale collection of its post-larva (puerulus) and early juveniles without affecting the fishery and to grow them in sea-cages, as is being carried out in Vietnam, India and the Philippines (Tan 1997; Tuan & Mao 2004; Vijayakumaran et al. 2009) and being trialled in Australia and New Zealand (Sheppard et al. 2002). With a shorter larval life, high growth rates for juveniles, hardiness, and a good market value, the slipper lobster is fast emerging as a new species of aquaculture interest. In Arctidinae, the adult females are large (carapace length (CL) > 100 mm) and produce smaller eggs with a larval development period of at least eight months. The adult females of Ibacinae and Theninae are also relatively large (CL > 70 mm) and produce larger eggs that complete developments in about a month (Theninae) or in 2–4 months (Ibacinae) (see Sekiguchi et al. 2007). Scyllarinae are the smallest of the scyllarids (CL < 33 mm) with more than half the number of species in Scyllaridae (Holthuis 2002). Ibacus, Thenus and Scyllarus larvae hatch at a more advanced stage and hence have shorter larval stages. This shorter larval life and hardy nature of the larvae have aided in development of large-scale aquaculture of Thenus sp. in recent years (Mikami & Kuballa 2007). Maintenance of the fragile phyllosomas larvae in controlled conditions for a period ranging up to 12 months for production of puerulus was the major impediment in developing culture techniques for lobsters. Mass culture of phyllosomas did not look feasible, until recent studies reported mass production of post-larva (nisto) of Thenus sp. in 25 to 30 days with high survival rates of over 80% (Mikami & Greenwood 1997a). Outside Australia, successful completion of larval development of T. orientalis was achieved in India with 22% survival (Kizhakudan et al. 2004b). The early juveniles (5–10 g) of this species were also grown up to 150 to 175 g in 10 to 11 months in static aquaria (Anon. 2007). Among the other larger slipper lobsters, the Ibacinae appears to be a candidate for large-scale aquaculture as the larvae were grown to nisto in about 54–70 days (Stewart & Kennelly 2000). But the slow growth rate of this species, along with the low survival in rearing trials of larvae, suggest that further trials are needed to evaluate its aquaculture potential.

3.4

MARKETING

Slipper lobsters, generally caught in shrimp and fish trawls as by-catch, are almost dead by the time they reach the deck and are marketed as frozen product. The unusual shape and its unfamiliarity to the consumer makes slipper lobster a less sought-after commodity compared with rock lobsters (Mosing & Fallu 2006). However, with the global production as low as 2,000 to 4,000 tonnes (FAO 2006), the slipper lobster has an assured market demand at present and there is tremendous potential for improving it, both in quantity and value, by appropriate marketing. The Southeast Asian customer considers live crustacean superior to the frozen or chilled ones and is prepared to pay almost double the price for live ones. Aquaculture of such high-value products would enable the farmer to be highly successful if he can achieve

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consistent supply of live animals, and assure quality and standard size (Anon. 2008a). Successful aquaculturists can balance the risk against the odds and the costs, and monitor the production trends to predict the results. However, it is important to know precisely when to sell his produce (Mosing & Fallu 2006; Anon. 2008a).

3.5

SLIPPER LOBSTER CULTURE INITIATIVES

Based on the success of mass-scale production of Thenus sp., the Australian Fresh Research and Development Corporation (AFR & DC) initiated pilot-scale production of Thenus sp. at the department of Primary Industries & Fisheries, Bribie Island Aquaculture research Centre (BIARC) (Mikami 2007). The Thenus sp. were grown from egg to 250 g size in about 400 days with >80% survival rate. The production technique was fine-tuned for over 11 years with each of the batches typically comprising 40,000 animals (Anon. 2008a). The first commercial-scale culture of Bay lobsters (Moreton Bay bugs, Thenus sp.) was initiated by the Australian Fresh Corporation Pty Ltd. at Cudgeon, west of Kingscliff in northern New South Wales (Mikami 2007; Anon. 2008a). A second commercial growout of Bay lobsters is underway in Australia with the Western Australian Ministry for Innovation, Industry, Science and Research announcing a $2,082,237 Commercial Ready grant to Lobster Harvest Pty Ltd, Fremantle, to develop a process to propagate two species of lobsters. The project aims to develop industrial aquaculture processes and protocols to deliver commercial quantities of full-sized Thenus sp. to market and hatchery-reared tropical rock lobster to growout by third parties (Anon. 2008b). In India, the juveniles of T. orientalis weighing approximately 5 g each were grown to over 150 g in 250 days at a stocking density of 30–35 individuals/m2 with live marine wedge clam, Donax cuneatus as feed, in a static semi-enclosed intensive system (Anon. 2007).

3.6 3.6.1

HATCHERY PRODUCTION OF SEEDS Broodstock management

Thenus sp. are the only slipper lobsters whose aquaculture potential has been demonstrated. Berried females of Thenus sp. are available in Australian waters throughout the year and the females usually spawn twice a year (Mikami & Kuballa 2007). In India, ovigerous females of T. orientalis are caught almost throughout the year with two annual spawning periods during June to August and February to March in the southeast (Subramaniam 2004; Radhakrishnan et al. 2007) and from August to April, with peak spawning during October to January, along the northwest coasts (Kabli & Kagwade 1996; Deshmukh 2001; Radhakrishnan et al. 2007) (Fig. 3.3). Vijayakumaran (unpubl. data) and Kizhakudan et al. (2004b) raised captive broodstock of T. orientalis from juveniles collected from bottom-set gill nets, while Mikami and Greenwood (1997a) produced broodstock of Thenus sp. from hatching in 8–10 months (Mikami 2007; Anon. 2008a). The captive breeders of T. orientalis mated and spawned from January to May and no spawning was recorded during May to December (M. Vijayakumaran, unpubl. data). The commercial-scale culture of Bay lobsters (Thenus sp.) by the Australian Fresh Corporation Pty Ltd requires two breeders per day for

Slipper Lobsters 91 20 18

Frequency (%)

16 14

Spawners

12 10

Recruits

8 6 4 2 0 A

M

J

J

A

S

O

N

D

J

F

M

Months Fig. 3.3

Annual breeding cycle and recruitment of Thenus orientalis in India.

production of 3 tonnes of Thenus sp. per day (1,000 tonnes per year) in their first phase of production (Mikami 2007). This is possible only by induced spawning by environmental manipulation throughout the year in Thenus sp. Broodstocks were reared in static aquaria with 100% exchange of water daily (Mikami 2007) or in static aquaria with in situ recirculation biological filters and 50% daily water exchange (Kizhakudan et al. 2004b). Holding temperature for berried females were between 24 and 28 °C (Mikami 2007) in Australia, whereas Vijayakumaran and Radhakrishnan (pers. obs) reared the broodstock at the ambient temperature of 26–32 °C. The broodstock was fed ad libitum on clams (D. cuneatus and Meritrix casta) and green mussel (Perna viridis) by the Indian investigators while bivalves and squids were used in Australia (Mikami 2007). Mating in Thenus appears to be a brief encounter (Jones 1998). In palinurids the spermatophoric mass hardens into a tough matrix on the sternum of the female, which has well-developed claws on the fifth pereiopods to pinch and scrape it during ovulation (Aiken & Waddy 1980). As no spermatophoric mass has been seen in wild breeders and the chelae of pereiopods are weak and fixed, it was thought that in Thenus it would be soft and shortlived, smilar to that of Jasus (MacDiarmid & Kittaka 2000). This was confirmed by Kizhakudan et al. (2004b), who observed the spermatophoric mass adhering to the postventral sternite and anterior abdominal region of the female in the form of a longitudinal white, jelly-like mass in T. orientalis and P. rugosus. Ritz (see Jones 2007) observed fertilisation and oviposition within 6 hours of mating in Thenus sp., whereas (Kizhakudan et al. (2004b) reported that the egg extrusion in T. orientalis and P. rugosus started within 5–7 hours of mating and was completed within 6–8 hours and the spermatophoric mass was lost in about 12 hours after mating. The egg development period in Thenus varied from 20–23 days at 26–32 °C. Few breeders mated again and produced a second batch of eggs within 1–2 months of the first larval

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release (Senthil et al. 2004). Mikami and Kuballa (2007) also reported that Thenus sp. breed twice in a year. In T. orientalis, the authors observed that the total number of phyllosoma released by captive broodstock (2,820–10,600) was much less compared with wild breeders (7,913–43,275). Jones (1998) reported that the mean fecundity of T. indicus is 12,455 eggs (range 3,686–25,314) and that of T. orientalis is 32,230 (range 5,579–54,746). The relationship of fecundity and size for these two species has been described by the following functions: T. indicus: fecundity = 658.7 × CL (mm) − 26,329 and T. orientalis: fecundity = 1,273.2 × CL (mm) − 6,7049 (Jones 2007). Kagwade and Kabli (1996) reported a linear relationship between total length, the fecundity in T. orientalis, expressed as: Fecundity (in thousand eggs) = −46.66 + 0.4164 × TL.

3.6.2

Larval development in scyllarids

A flat and transparent leaf-like body, a long developmental period and shorter post-larval phase are common features for both palinurid and scyllarid lobsters. The planktonic developmental period in scyllarids ranges from a few weeks to more than 9 months (Booth et al., 2005; Mikami & Kuballa 2007; Sekiguchi et al. 2007). The phyllosoma go through a number of moults or instars as they develop, leading to a new stage in development. Each stage may be associated with a single or multiple instars. In scyllarids, the number of instars varies from four (each stage with a single instar) in T. orientalis to more than 16 (eight stages) for Scyllarides haani (Table 3.1). The nisto settles in its natural habitat soon after completion of larval development as in Theninae, or after many days as in many scyllarids. The nisto provides the link between the planktonic and benthic life history phases in scyllarids (Booth et al. 2005; Mikami & Kuballa 2007; Sekiguchi et al. 2007). A description of larval morphology of many scyllarids from plankton collections is given by Holthuis (2002). There is relatively little work on the early life histories of scyllarids, as adults of only few scyllarids are big enough to have any commercial importance (Sekiguchi et al. 2007). Attempts have been made to study the larval developments of 22 species of scyllarids (see Mikami & Kuballa 2007; Kumar et al. 2009), but a complete description of the larval development is available for only a few species (Table 3.1) such as Ibacus ciliatus, I. novemdentatus (Takahashi & Saisho 1978), I. peroni (Marinovic et al. 1994), Chelarctus cultrifer (Matsuda & Mikami reported in Mikami & Kuballa 2007), Crenarctus bicuspidatus (Matsuda reported in Mikami & Kuballa 2007), Scyllarus americanus (Robertson 1968), S. arctus (Pessani et al. 1999), Petrarctus (formerly Scyllarus) demani (Ito & Lucas 1990), P. rugosus (Kumar et al. 2009), and T. orientalis and Thenus sp. (Mikami & Greenwood 1997a). Phyllosomas of most scyllarids are primarily predators and use their pereiopods to fix and hold food items. The mouth and foregut structures suggest consumption of soft fleshy food (Mikami & Greenwood 1997a; Mikami & Takashima 2000). Many scyllarid phyllosoma were reported in close association with medusae (Hernkind et al. 1976; Barnet et al. 1986). Sims and Brown (1968) suggested a feeding association of scyllarid phyllosoma with medusae as nematocysts were found in the faeces of a giant phyllosoma (possibly of Paribaccus sp.) and of other (unnamed) phyllosomas. Foods such as Artemia, fish larvae, ctenophores, Sagitta, bivalve gonads and jellyfish are used in phyllosoma culture (Mikami & Kuballa 2007). Mikami and Takashima (2000) first described the morphological development of the proventriculus in the phyllosomal and nisto stages of the scyllarids. Filtering food particles

Theninae Thenus orientalis (Lund, 1793) Thenus indicus (Leach, 1816)

Scyllarinae Chelarctus cultrifer (Ortmann, 1897) Crenarctus bicuspidatus Scyllarus americanus (S.I. Smith, 1869) Scyllarus arctus (Linnaeus, 1758) Petractus demani (Holthuis, 1946) Petractus rugosus

Ibacus novemdentatus Gibbes, 1850 Ibacus peronii Leach, 1815

Ibacinae Ibacus ciliatus

Species

4 + nisto + juvenile

8 + nisto + juvenile instar 1) 8 + nisto

28

27

24.9 to 28.1

25.5

Fresh flesh of Donax brazieri

Artemia salina for ∼80 days then whisked fish and beef Artemia nauplii plus chopped Gafrarium sp. after the 5th instar Artimea nauplii plus flesh of mussel/clam/ fish, and frozen cyclop-eeze after the 4th instar

20 ± 1

192 at 20 ± 1 46 to nisto 51

25

32–40

16 postlarva

Artemia nauplii

24

Artemia nauplii plus chopped gonads of the mussel Mytilus galloprovincialis after 30 days Artimia nauplii with chopped Mytulis edulis

51–62

24.3

20.5 and 23.5

Stage 11 + nisto + juvenile 6–7 Postlarva

79

6 + nisto + juveniles

23–25

Artemia nauplii plus the meat of the shortnecked clam Tapes philippinarum in later stages Artemia nauplii plus the meat of the shortnecked clam Tapes philippinarum in later stages Artemia nauplii plus ovaries of mussel Mytilus edulis after 3rd stage

Feed

159

65

7 + nisto

25

Temperature (°C)

Stage 10 + nisto

54–76

Days of culture

7–8 + nisto

No. of stages attained

Table. 3.1 Completed larval rearing of Scyllarid lobsters.

Mikami & Greenwood (1997a)

Kumar et al. 2009

Pessani et al. (1999) Ito & Lucas (1990)

Matsuda & Mikami (unpublished) Matsuda (unpublished) Robertson (1968)

Marinovic et al. (1994)

Takahashi & Saisho (1978)

Takahashi & Saisho (1978)

Author

Slipper Lobsters 93

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previously masticated by the mouthparts is apparently the main function of the phyllosomal proventriculus. The nisto is a week feeder as it lacks phyllosoma-like filtering and the development of juvenile/adult masticating functions in the proventriculus (see Johnston 2007 for a full description of the gastric system of scyllarids). In T. orientalis the nisto is much closer to the adult than the nisto of other scyllarid sp. (Barnet et al. 1986) and its functional role may not be identical in all species. The nisto of T. orientalis has poorly developed pleopods and is a poor swimmer (Barnet et al. 1984) and may not be morphologically equipped for active dispersal or site selection. Almost all nistos described to date are at first virtually transparent, possibly evolved to avoid predation as they move inshore (Mikami & Kuballa 2007; Kizhakudan et al. 2004b). The nistos are mostly backward swimming (Barnet et al. 1986; Jacklyn & Ritz 1986) and, like adults, they can also sink passively and swim forward by vigorous beating of the setose pleopods (Robertson 1968). Sand burying during day may also be quite widespread in the nisto stage.

3.6.3

Growth of phyllosoma

Moulting or periodic shedding of the exoskeleton and expansion of the soft new exoskeleton before it hardens again, to allow for growth of tissue within, regulate growth in crustaceans. A range of environmental cues affect moulting, which is regulated by hormonal signals (Skinner 1985). The endocrine control of the moulting cycle appears similar in larval and adult nephropid lobsters and the demonstration of the sinus gland – X organ complex in the stage 1 phyllosoma of P. japonicus (see Mikami & Kuballa 2007) indicates that the hormones regulating moult cycle are functional even at the earliest stages of larval development. Manipulation of endocrine pathways might help in reducing instar duration or number of stages required for metamorphosis in scyllarid species that have a particularly long larval cycle, such as Scyllarides and Parribacus, enhancing their culture possibilities (Mikami & Kuballa 2007).

3.6.4

Larval nutrition

Nutrition is vital in phyllosoma rearing. Survival, intermoult period and growth of phyllosoma are influenced by starvation and feeding duration. Dietary deficiencies introduced extra moults, atypical morphological development, and poor survival in phyllosomas of Thenus sp. (Mikami 1995) and P. homarus (Vijayakumaran & Radhakrishnan 1986). Excessive moults, incomplete metamorphosis, and morphological anomalies owing to poor nutrition are also reported in nephropid larvae (Charmantier & Aiken 1987). Delay in first feeding beyond the period when 50% of the phyllosoma would have moulted to the second instar (1.7 days for T. indicus) affects moulting, growth and survival (Mikami 1995) leading to the conclusion that Thenus phyllosomas first assimilate an energy reserve for moulting and use additional reserves for growth. Establishing nutritional requirements of phyllosoma through different larval stage is essential as different types and sizes of food are required by different stages. Morphology of the maxillipeds (mouthparts) and thoracic appendages could give valuable clues to the feeding habits across species (Phillips & Sastry 1980). Scyllarid phyllosomas have welldeveloped mouthparts from stage 1, with sharp incisor processes on the mandibles and well-developed setation on the first maxillae and second maxillipeds (Johnston 2007). The

Slipper Lobsters 95

incisor and molar mandibular processes assist in mastication of food. In the absence of a gastric mill, the scyllarid phyllosoma does not grind food and the foregut sorts and filters food particles previously masticated by the mandibles, suggesting it prefers a gelatinous diet that requires little additional mastication (Johnston 2007). The stomach appears to squeeze and filter the food particles previously cut and chewed within the mouthparts (Mikami et al. 1994). The midgut gland of the phyllosoma occupies most of the inner carapace and has a single layered midgut lumen (Mikami et al. 1994). Successful completion of most phyllosoma rearing was possible by supplementing chopped flesh of bivalves along with the main feed, the Artemia nauplii (Takahashi & Saisho 1978; Ito & Lucas 1990; Marinovic et al. 1994; Mikami & Greenwood 1997a; Kumar et al. 2009). Whipped fish and beef were used by Pessani et al. (1999) to feed later stages of Scyllarid arctus during the successful completion of larval rearing of this species to nisto. Zooplankton such as Daphnia sp., Sagitta sp., fish larvae and hydromedusae have been fed to phyllosoma, but none of the studies was able to complete the larval life cycle (Saisho 1966; Ritz & Thomas 1973; Radhakrishnan & Vijayakumaran 1995). Takahashi and Saisho (1978) could rear I. ciliatus and I. novemdentatus phyllosoma using the shortnecked clam, Tapes phillipinarum, while Ito and Lucas (1990) used Venus clam, Geranium sp., to complete larval rearing of Petrarctus demani. Successful rearing of several palinurid species and the scyllarid, I. peroni were conducted feeding on the flesh of the blue mussel, Mytilus edulis (Marinovic et al. 1994; Kittaka & Booth 2000) whereas the green mussel (P. viridis) and clam (D. cuneatus and M. casta) were used along with Artemia nauplii for completion of larval rearing in T. orientalis (Kizhakudan et al. 2004b). The introduction of molluscan flesh seems to be a key factor contributing to the complete rearing of phyllosomas. Mikami and Greenwood (1997a) found that fresh bivalve flesh contributed to high survival of T. orientalis phyllosomas with larger moult increments and shorter intermoult periods. But this was not possible with frozen mussels as its nutritional quality reduced during the freezing process.

3.6.5

Larval diseases

Disease outbreaks are being increasingly recognised as a significant constraint in the hatchery production of lobster larvae. The low level of nutrients in the open ocean, the natural habitat of phyllosoma, ensures an apparently pathogen-free environment (Phillips & Sastry 1980). Phyllosoma is very sensitive to the microbial load in the water column. Vibriosis (mainly caused by gram negative Vibrios) remains the most important disease problem associated with larval rearing of lobsters. Vijayakumaran and Radhakrishnan (pers. obs.) have observed that the phyllosoma larvae and the live feed, Artemia nauplii, have high Vibrio load resulting in mass mortality. Fouling with filamentous bacteria (Leucothrix sp.) and protozoans such as Zoothamnium sp., Vorticella sp. and Acinata sp. was another serious problem in larval rearing. The live feed, Artemia nauplii, could introduce bacteria almost continuously to the culture system.

3.7

FACTORS INFLUENCING PHYLLOSOMA GROWTH AND SURVIVAL

Phyllosoma require good quality water for normal development and the provision of optimum conditions is the key to the success of its rearing. Optimum conditions for rearing

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phyllosoma may vary among species and it is often very difficult to maintain hygienic conditions in a large-scale hatchery operation during a lengthy larval phase (Mikami 2007; Mikami & Kuballa 2007).

3.7.1

Rearing tank design

The delicate body of phyllosoma could be damaged by abrasion caused by water circulation, aeration and the structure of the rearing tank. The rearing tank design should have controlled flow of water to avoid physical damage and keep the larvae in suspension (Mikami 2007). Sufficient provision of quality feed is another important criterion for successful larval rearing. Many authors have reared phyllosomas in static water in plastic or glass containers ranging from 50 ml capacity to several litres. Individual rearing in small containers gave better survival and it was easier to follow the development through many moult cycles. Kittaka (1994) used a vertical upwelling system, which was subsequently adopted by many other researchers. Strong aeration or water movement, as in the upwelling system, tends to damage body parts of the fragile phyllosomas. Attempts to prevent this have led to the development of horizontal doughnut-shaped tanks (Matsuda & Takenouchi 2005), rectangular shallow plastic trays (Kumar et al. 2009) and raceways (Fig. 3.4) (Mikami & Kuballa 2007) for phyllosoma rearing. Optimal tank design with a consideration of hydrodynamics for water circulation is crucial for the maintenance of the fragile phyllosomas (Mikami & Kuballa 2007).

Fig. 3.4 Protoype 300 mm raceways for larval rearing of Thenus sp. (reproduced from Mikami & Kuballa 2007; used with permission).

Slipper Lobsters 97

Matsuda and Takenouchi (2005) obtained higher survival for P. japonicus phyllosoma in doughnut-shaped tanks with horizontal water movements. The doughnut-shaped tanks and raceways with horizontal slow movement of water were easy to maintain. Large numbers of Thenus sp. nistos were produced in the raceway tanks and this model has been adopted for seed production in commercial Thenus culture ventures in Australia (Mikami 2007). In raceways, the floor space requirements are greatly increased, but the accessibility to larvae also increases along with space available for rearing large numbers of larvae, making this the only system currently able to produce large numbers of juveniles (Mikami & Kuballa 2004). Passive water movement in the tank facilitating swimming and feeding behaviour of phyllosoma, efficient food dispersal along with accessibility to larvae were responsible for the high success rate of phyllosomas rearing in raceways (Mikami & Kuballa 2004).

3.7.2

Environmental factors

3.7.2.1 Temperature Temperature is probably the most influential factor in phyllosoma growth and survival. An increase in temperature increases growth rate, but beyond optimum range the moult increment may be reduced, affecting survival (Mikami 1995). The upper limit of temperature differs for each species and in different ecological zones (Table 3.2). Mortality increases at high temperature, probably due to poor nutritional conditions and high bacterial growth in the system (Mikami 1995). Mikami and Greenwood (1997a) suggested 27 °C as the optimum temperature for rearing T. orientalis phyllosoma, whereas Vijayakumaran and Radhakrishnan (pers. obs.) found 28–29 °C to be optimum for the same species in India and above 30 °C the bacterial load of both phyllosoma and the rearing system increased, causing mortality. If all other rearing conditions are kept at optimal level it may be possible to increase the temperature, but the larvae may be exposed to higher stress levels leading to mortality if any one of the other rearing parameters is altered.

Table 3.2 Effect of temperature on phyllosoma rearing in Scyllarid lobsters. Species

Tested temperature range (°C)

Optimum temperature (°C)

Reference

Scyllarus americanus Scyllarus aequinoctialis Scyllarus arctus

10.0–30.0

Shortest duration at 25.0 Highest survival to Nisto at 20.0 Maximum survival at 24

Robertson (1968) Robertson (1979)

Ibacus peronii

20.7–23.3

Thenus orientalis Thenus sp.

26.0–32.0

Full development to Nisto in 192 days at 20.0–21.0 Higher survival at 20.7 but low moult frequency Higher survival and growth between 28.0 and 29.0. Higher mortality above 30.0 High survival and growth at 27 and reduced moult increments and survival above 27

Pessani et al. (1999) Marinovic et al. (1994) M. Vijayakumaran (pers. obs) Mikami & Greenwood (1997a)

10.0–30.0

27.0 ± 0.5 25.5–27.0 for nisto

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3.7.2.2 Salinity Salinity level of oceanic seawater (∼35 psu) is ideal for phyllosoma rearing, even though the phyllosma can tolerate a wide range of salinities (Robertson 1968; Mikami 1995). Suboptimal salinity resulted in low growth in T. orientalis (Mikami 1995) or in extra moult stage in Scyllarus americanus (Robertson 1968). When grown under different salinity regimes of 25–35 psu, the survival and growth of phyllosomas of the palinurid P. homarus and the scyllarid, T. orientalis were best at 32–33 psu (M. Vijayakumaran, pers. obs.). Increased osmo-regulatory demands may be the cause for reduced growth under low salinities as crustacean larvae maintain their haemolymph hyper-osmotic to seawater by a weak hyper-regulation (Mantel & Farmer 1983). Under low salinity levels, the mechanism of hyper-regulation may require more energy to maintain body fluids within the phyllosomas at the required osmolality, and this energy drain may result in slower growth (Mikami & Kuballa 2007).

3.7.2.3 Dissolved oxygen Dissolved oxygen (DO) level of rearing water is an important factor in phyllosoma growth and survival and has to be maintained above 5 mg/L. High mortality of stage II phyllosomas of P. homarus (15 larvae/litre) occurred when the DO level fell to <3 mg/L in the vertical upwelling rearing tanks. The default was traced to a faulty air pump in the recirculating system. Similarly, the total ammonia in the rearing tank has to be kept below 1 ppm for better growth and survival of phyllosomas (M. Vijayakumaran, pers. obs.).

3.7.2.4

Light

The duration and timing of moult cycle in phyllosomas is affected by photoperiod and temperature. In groups exposed to natural daylight, 24 hour light and 24 hour dark respectively, synchronised moulting was observed only in those exposed to natural light (T. orientalis: Mikami & Greenwood 1997a). In T. orientalis, a rise in temperature up to 27 °C induced a shorter moulting period and survival decreased dramatically when larvae were exposed to temperature above 30 °C or below 16 °C (Mikami 1995). Early-stage phyllosomas are strongly positively phototropic (Radhakrishnan & Vijayakumaran 1986; Mikami 1995) and this reaction might affect feeding opportunities of phyllosomas in culture systems. As the most common larval feed, live Artemia nauplii are also positively phototropic; the encounter between the predator and the prey is enhanced, resulting in more efficient feeding. However, under conditions of total darkness there was no effect on growth of phyllosomas of P. homarus (Radhakrishnan & Vijayakumaran 1986) or T. orientalis phyllosomas and nistos, suggesting that light may modify their behaviour, but does not necessarily affect their metabolic rate (Mikami & Greenwood 1997a). The growth and survival of phyllosoma of T. orientalis (Mikami & Greenwood 1997b) was not affected when exposed to natural light and 24 hour darkness, but were lower for those larvae exposed to 24 hour light. Moulting was synchronised in natural light and occurred around dawn while under 24 hour darkness and light it was erratic and occurred irregularly (Mikami & Greenwood 1997b). Light may modify the behaviour of phyllosoma, but may not modify their metabolism (Mikami & Greenwood 1997b).

Slipper Lobsters 99

3.7.3

Biological factors

Stocking density is an important factor in the success of phyllosoma rearing and it will vary according to the rearing system. Solitary rearing usually gives maximum growth rates, while high density affects growth of scyllarid phyllosoma (Mikami 1995). Mortalities under high densities in phyllosoma instars of T. orientalis was related to high levels of cannibalism only in cases where insufficient food was provided (Mikami 1995). This suggests that phyllosoma can be reared at high densities (>10 phyllosomas per litre) if adequate quality and quantity of food is consistently available. Bacterial/viral control is a key issue for the rearing of long-lived oceanic phyllosomas under controlled environments. Micro algae that can control nutrients and bacterial growth were used in the early rearing trials of phyllosoma (Kittaka & Booth 2000). Fouling by epibionts such as filamentous bacteria and protozoa is another problem encountered in rearing phyllosomas. Malachite green treatment was effective in removing the epibionts such as Zoothamnium and Leucothrix (Vijayakumaran & Radhakrishnan 2003), but it is a banned chemical for hatchery use owing to its carcinogenic implications. Among other chemicals tested for controlling epibionts, methylene blue proved to be equally effective. However, the use of disinfectants and antimicrobial drugs has limited success in the prevention or cure of lobster larval disease and they are not considered as suitable long-term solutions for commercial-scale rearing. Moreover, the abuse of antimicrobial drugs could produce resistant bacteria, which can transfer their resistance genes to other bacteria that have never been exposed to the antibiotic (Kittaka & Booth 2000; Mikami 2007). UV light and ozone treatment are now being used extensively for microbial control in hatcheries. Use of UV treated and ozone-sterilised water have improved survival of phyllosoma to a great extent (Mikami & Kuballa 2004; Ritar et al. 2006). These treatments have now become important components in recirculation systems to remove microbes from incoming water to the tanks. Ozone can also remove microbes from the body of phyllosoma and live feeds in the rearing system. Ozone is, however, toxic to all living organisms and can cause abnormalities in phyllosoma (Ritar et al. 2006), and therefore should be maintained at low levels (<0.05 ppm) (Mikami & Kuballa 2007).

3.8 3.8.1

HATCHING AND LARVAL REARING IN THENUS SP. Hatching and phyllosoma rearing

Hatching in Thenus occurs just before sunrise (Mikami & Kuballa 2007). Phyllosoma of Thenus sp. are positively phototactic, which helps in harvesting healthy larvae from the hatching tank with the help of light (Mikami & Kuballa 2007). Thenus orientalis and T. indica have a short-lived naupliosoma, four phyllosoma stages (Table 3.3) and a nisto that Table 3.3 Duration of phyllosoma instars of the slipper lobster Thenus orientalis. Phyllosomal instar

Duration (days) of phyllosomas instar Mikami & Greenwood (1977a)

I II III IV

6–9 6–9 6–13 7–16

Kizhakkudan et al. (2004) 6 5 7 7

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settles into their benthic habitat and moults into a juvenile (Mikami & Greenwood 1997a). The average duration to attain post larval nisto stage is 25 to 30 days at 26–27 °C with a minimum of 21 days, the shortest duration of phyllosoma stage in any lobster (Mikami & Greenwood 1997a). The protocol for rearing phyllosomas of T. orientalis is given in Table 3.4.

3.8.2

Nursery rearing

Scyllarid post-larvae are more like their adults. Nistos of offshore species such as Thenus sp., Ibacus sp. and Evibacus sp. just settle by burying into the sandy bottom, their natural habitat. The nistos of oceamic species such as Scyllarides and Parribacus have to swim long distances to settle, suggesting that the swimming capability of lobster post-larvae correlates with the habitat of the adult stage (see Sekiguchi et al. 2007; Mikami & Kuballa

Table 3.4 The protocol for rearing Thenus orientalis seeds in hatchery. Stage of lobster/larvae

Environmental/physical/biological parameters

Broodstock

Rearing system: Static tank or raceways with UV/ozone-treated seawater Salinity: 30–35 psu Temperature: 26–29 °C DO*: >5 mg/L Total Ammonia:
Phyllosoma

Horizontal rearing system – Doughnut-shaped tank/raceway with UV/ ozone-treated seawater, ozone <0.05 ppm Stocking density: up to 10individual/L Salinity: 32–35 psu Temperature: 26–28 °C DO: >5 mg/L Total Ammonia: <1 ppm; Feed: Artemia; fresh bivalves/squid/fish

Nisto

Static tank or raceways with good water movement UV/Ozone treated sea water Stocking density: High; vary with rearing system Salinity: 32–35 psu Temperature: 26–29 °C DO: >5 mg/L Total Ammonia: <1 ppm Feed: No feed required

Juvenile (nursery rearing)

Static tank or raceways UV/ozone-treated seawater Stocking density: Varies with rearing system Salinity: 32–35 psu Temperature: 26–29 °C DO: >5 mg/L Total Ammonia: <1 ppm Feed: bivalve/squid/fish

* DO = dissolved oxygen

Slipper Lobsters 101

2007). In the hatchery, nistos of Thenus sp. settle immediately after metamorphosis and bury themselves in the sand (Mikami 1995; Kizhakudan et al. 2004b). Good water movement is essential for nisto in hatchery as static water tends to cause mortality (Mikami 1995). As the nistos of Thenus sp. are non-feeding, Mikami and Greenwood (1997a) did not feed them throughout their entire seven-day period. The non-feeding status of these postlarvae is reflected in the structure of their mouthparts and foregut (see Johnston 2007 for details), which is less developed compared with the phyllosomas or juveniles and is ill equipped to both ingest and digest food (see Mikami & Kuballa 2007). The nisto has to survive solely on energy reserves accumulated throughout the phyllosoma stages till their settlement and transformation to juveniles. Understanding of the basic biology of nisto and juvenile stages is necessary to grow nisto to juveniles and further to market-size adults. Mikami and Greenwood (1997a) observed that metamorphosis of Thenus sp. phyllosoma into nisto stage occurs just after sunset. The non-feeding nistos of scyllarid lobsters require the least maintenance, enabling high-density stocking as they settle by burrowing into sand immediately after metamorphosis. This facilitates reduced rearing space and husbandry, important factors in the success of commercial rearing (Mikami & Kuballa 2007).

3.9

GROWTH OF JUVENILE SLIPPER LOBSTERS

Growth process has been studied only in few scyllarids such as Scytlarides, mainly of the Mediterranean species (see Bianchini & Ragonese 2007) and in Thenus sp. (Mikami 2007; see Radhakrishnan et al. 2007). The parameters of the seasonalised vBGF (von Bertalanffy growth factor) obtained in the laboratory for Scyllarides latus (Latreille, 1802) are CLoo = 127.2 mm, k = 0.20, C = 1.0, ts = 0.83, which are in agreement with data from long-term recaptures in the wild, and not far from the values for the Galapagos species S. astori Holthuis, 1960 (Hearn 2006). The vBGF suggests that Scyllarides species are long lived, with a yearly moult in the adult phase, and incremental increases in carapace length per moult of around 6% (Bianchini & Ragonese 2007). Kabli and Kagwade (1996) estimated the vBGF for T. orientalis, showing that its growth pattern was retrogressive and geometric throughout life. Growth in the genus Ibacus also has been studied, both in captivity and in the wild. Stewart and Kennelly (1997, 2000) found that while males and females of I. peronii (Leach 1815) differ in growth rates, no sex-related differences exist in I. chacei (Brown & Holthuis 1998). Rudloe (1983) examined the aquaculture potential of Scyllarides nodifer (Stimpson 1866), hypothesising that growth from a post-larva to a 300 g animal should require 9 to 10 moults and approximately 18 months. Bianchini et al. (2001), using medium- and long-term recaptures, combined with laboratory data, derived the parameters of the vBGF, and compared observed and expected measurements for S. latus (Bianchini & Ragonese 2007). Growth rates tend to rise with temperature, but mortality also increases at the highest viable temperature (Hartnoll 1982). In artificial conditions, the water quality is crucial in maintaining high growth rates. Dissolved oxygen levels, nitrogenous waste levels, crowding or isolation, and shelter may play a role in moulting and growth of animals in captivity. The growth rate of crustaceans is also influenced by feed, and reduces when feeding is inadequate (Hartnoll 1982), through extended intermoult periods and smaller moult increments (Chittleborough 1975).

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Little or no information is available on the nutritional requirements of the slipper lobsters making it more difficult to formulate diets, where even the feed presentation plays a role in the lobster nutrition (Tolomei et al. 2003). Weight increment per moult in slipper lobsters varies with size, with the maximum recorded in smallest individuals. In the Mediterranean slipper lobster, S. latus, the mean increments range from 24% for the smallest individuals to 6% for animals over 500 g of weight. In general the exuvia, the shed exoskeleton, represents 26% of the premoult body weight, with an almost linear relationship with the total premoult weight. Thus, when considering the shed exoskeleton, the average animal encompasses an actual weight increase/recovery of almost 40%. Such data for S. latus are similar to those for Ibacus sp., where growth increments are higher in juveniles than mature adults (20–35% vs. 11–15%, respectively) (Haddy et al. 2005). In T. orientalis, the average weight increase per moult in 10 g juvenile is above 90% of premoult weight and it reduces gradually to around 1511% in lobsters weighing 280 g. The shed exuvia weight represents 25–30% depending on size, making it more than 100% actual weight increase per moult in smaller lobsters (M. Vijayakumaran, unpubl. data). Thenus orientalis (Lund) is found in the coastal waters of India where it breeds and moults throughout the year, with alternating peaks for moulting and reproduction. Females mature at a carapace width (CW) of 7.3 ± 0.1 cm (see Radhakrishnan et al. 2007).

3.9.1

Juvenile growout

The inability to produce large numbers of juveniles in hatchery has traditionally been the major constraint in commercialisation of slipper lobster farming. Recent success in producing thousands of juveniles of Thenus sp. at a very high survival rate (>80%) (Mikami 2007; Mikami & Kuballa 2007) has generated interest in farming of this species and two big projects have already been initiated in Australia. It is possible to grow Thenus sp. in static indoor tanks, with daily partial exchange of water; in recirculation system with a series of tanks and with partial exchange of water, or in raceways connected to the recirculation system. As Thenus sp. occupy sandy or muddy habitat at the sea bottom, it may also be possible to culture them in earthen shrimp ponds that use seawater or creek water for farming. The studies of Vijayakumaran and Radhakrishnan have shown that T. orientals can be grown in indoor tanks without a sandy substratum to burrow, indicating that high-density culture in sea-cages is possible as practised for palinurids. Feed is an important component of successful growout of juvenile scyllarids. Bivalve molluscs are the preferred diet of scyllarids. Unlike palinurids, which break the shell of bivalve molluscs such as clam and mussels along its outer thin margins, the scyllarids cannot break the shell of bivalves. Instead, they wedge open the shell using sharp pereiopod dactylus (Johnston 2007). The mouthparts of adult Thenus sp. are not heavily calcified, and the flesh is masticated by the shearing action of the asymmetrical mandibles. The stomach has well-developed teeth in the gastric mill with large incisor processes and chitinous plates, which masticate food by a cutting action by the lateral teeth, with only a minor role for the medial tooth. The digestive gland, which is similar in structure and function to palinurids and nephropids, has high concentrations of proteolytic enzymes consistent with a carnivorous diet (Johnston 2007). To date no commercial artificial feed is developed for

Slipper Lobsters 103

any lobster species and the juvenile growout is entirely dependent on natural feed such as mussels, clams, cuttlefish waste and trash fish.

3.10 3.10.1

CULTURE OF THENUS SP. Culture in indoor tanks

Encouraging growth and survival of juvenile T. orientalis in large cement tanks, at the Kovalam Field Centre of Central Marine Fisheries Research Institute, Chennai, India (Anon. 2007), has generated interest in indoor culture of this species. Wild juvenile lobsters (average 20 mm CL and 5 g weight) were grown in cement tanks of 12.5 m2 floor space and sand bed, which served as a substrate as well as a biological filter. Juveniles were stocked at a density of 30–35 individuals/m2 with water column of 0.5 m height with 30% water was exchanged per day. The environmental parameters in the rearing system were salinity 36–38 psu, pH 7.8–8.2 and temperature 27–29 °C. Lobsters fed ad libitum with live marine wedge clam, D. cuneatus attained 140–175 g (0.60 to 0.70 g per day) in 250 days with an overall survival of 90%. A similar growth rate was obtained with wild juveniles (10 g) grown (7 individuals/m2) in one-ton capacity circular fibre reinforced plastic (FRP) tanks, with intertidal sand bed covering 50% of the bottom, at the NIOT laboratory, also in Chennai, India. The juveniles, fed with the marine clam D. cuneatus and the green mussel P. viridis at a rate of 5% body weight per day, grew to over 150 g in 350 days with 6 moults. The water quality parameters were: salinity – 30–33 psu, pH 7.8–8.2, temperature 26–32 °C; dissolved oxygen >4 mg/L and total ammonia <1 ppm. The growth was faster up to 100 g and then reduced, partly due to erratic feeding at later stages of the trial in addition to attainment of sexual maturity. On average, the juveniles moulted 6 times with the intermoult period varying from 25–30 days in early stage to 111–114 days towards the end of the trial. In another trial, juveniles of T. orientalis with average weight of 5 g stocked at a density of 20 individuals/m2 in small 200 L FRP tank attained a weight of 95–100 g in 6 months. Three different feeds, mussel, clam and oyster were fed at 10% body weight and all the three proved to be equally effective (Vijayanand et al. 2004). These three trials give ample proof that T. orientalis can be successfully reared in static indoor tanks at high density. Frequent occurrence of diseases, seasonal variations in water quality parameters such as salinity, the need to maintain higher temperature of rearing water in temperate countries, and the difficulty in maintaining discharge concentrations of farm effluents have led to excellent models of recirculation systems for broodstock maintenance, larval rearing, nursery rearing and for indoor growout in tanks and raceways. A typical recirculatory system (Fig. 3.5) would have a settling tank, a bead filter to remove large particle, an ozoniser, connected to an ozone generator (plus an UV treatment unit if necessary), a protein skimmer to remove dissolved organic matter and finally a biofilter (beads, balls, biofilm or seaweed) from which recirculated water flows into the rearing tanks. The recirculatory system is successfully used in hatcheries for rearing fish larvae, in nurseries for fry and fingerlings and for growout in intensive culture of fishes such as eels (Heinsbroek & Kamstra 1990), sea bream (Ellner et al. 1996), and turbot (Rodrigo & Jorge 2003). The growth and survival of T. orientalis in static indoor tanks suggests that a series of such tanks can be connected to efficient recirculatory systems to intensively grow this species to market size.

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Tank

Tank

Tank

Tank

Tank

Tank

Tank

Tank

Automatic flow changers Air Drain

Bead Recirculation Filter Filters Pump

Automatic Bead filter washing cycle

Fig. 3.5

3.10.2

UPS

Settler Timer

Recirculatory system for indoor culture of Thenus lobsters.

Culture in raceways

The Australian Fresh Research and Development Corporation (AFR & DC) conducted pilot-scale production trials of Thenus spp. in shallow raceways with 40,000 animals in each batch (Anon. 2008a). In 12 months, the lobsters were grown from egg up to 250 g adults with an 80% plus survival rate. Juveniles moulted about 19 times to reach 250 g (Anon. 2008a). Mortalities in preliminary trials were traced to bacterial disease of larvae accentuated by poor nutrition. The bacteria infecting phyllosoma were present in seawater and opportunistically attacked phyllosomas’ weakened sub-optimal environmental and nutritional conditions. Modification to larval production system and optimisation of nutritional requirements reduced the incidence of bacteria-related mortality (Mikami 2007). The nisto and juveniles were more robust and no disease problem was encountered in their growth. Dosing of ozone to ensure sterilisation of seawater minimised the level of bacteriarelated mortality (Mikami 2007). Based on the pilot production system, the Australian Fresh Corporation Pty has initiated commercial production of Thenus sp. in northern New South Wales. Recirculation aquaculture technology would be used to grow animals in shallow raceways (Fig. 3.6). The project is designed to produce two products; live hard-shell animals of an average 218 g and soft-shell animals (frozen or fresh chilled) of an average 45 g (Mikami 2007). The

Slipper Lobsters 105

Fig. 3.6 Thenus growout raceway tank (reproduced from Mikami & Kuballa 2007; used with permission).

production target is 3 tonnes per day in an annual production of 1,000 tons in stage 1, which would require a standing stock of approximately 400 tonnes of animals (Fig. 3.7) (Mikami & Kuballa 2007). The stage 1 would occupy an area of approximately 15 ha and include hatchery and growout facilities, car parking plus processing building, workshop facilities, administration building, seawater and freshwater storage tanks, as well as seawater and wastewater pipelines and pump house. Stage 2 and stage 3 would replicate stage 1 with an expected total annual production of 3,000 tons. The production process will be housed in a large facility with provision to control temperature and continuous monitoring of water quality and high standards of hygiene. The facility will have back-up power supplies and filtration systems and adequate inputs such as feed, labour, etc., to cover an increased degree of variability at full-scale production process problems not experienced in pilot trials (Anon. 2008a). Production of soft-shell slipper lobster would be an important innovation for realising high value for the product in aquaculture. Controlling the moulting process is critical to success in the production of soft-shell animals. Moult stages can be easily identified by the presence of exoskeleton ecdysial sutures (crack lines) on the gill chambers. Thenus sp. show synchronised moulting around sunset when animals are under a natural day–night condition. The timing and synchrony of moulting can be manipulated by altering the day– night cycle, whereas the length of the actual moult stage can be shortened or prolonged by

106

Recent Advances and New Species in Aquaculture Fresh seawater 600m3 per day (8 days storage)

555kg pipis or equivalent molluscs per day

2 berried females per day

Seawater intake & storage Broodstock (100% per day)

Food preparation

Larval rearing area

7032kg squid per day

(75% per day) (270kg)

(5000m3)

Nursery area (100% per day)

Growout I (6762kg)

Freshwater

Growout II

Softshell 450kg per day Hardshell 2.5t per day

(5% per day)

Waste water treatment

Discharge seawater 600m3 per day Fig. 3.7 Diagram of the proposed Thenus aquaculture facility in northern New South Wales, Australia. Courtesy of The Lobster Newsletter, 21(2), 18–21.

manipulating temperature. During the intermoult stage for juveniles (average weight 79 g), body weight increases only by 9%, whereas most of the weight gain (an additional 44%) occurs within the short period just before and after the actual moult stage. These findings have application for the development of ‘softshell’ products, which can be harvested at around the actual moult stage (Mikami 2005).

3.10.3

Culture in earthen ponds

Earthen ponds are traditionally used for production of shrimps along the coastal zone in many countries. In India, many small and large shrimp farms have temporarily stopped farming of shrimp (Penaeus monodon) due to frequent occurrence of the viral disease white spot disease syndrome (WSSV), and a fall in the selling price of shrimps on the international market which has reduced the viability of shrimp farming. Such shrimp farms sourcing water from the sea with a salinity range of 30–35 psu could be used for stocking Thenus sp. as the muddy bottom would be an ideal habitat for the lobster. The mud spiny lobster, P. polyphagus, with similar habitat to Thenus sp., could survive and moult in earthen ponds with muddy bottom (Vijayakumaran & Radhakrishnan unpubl. data). Similarly, it may be possible to grow Thenus sp. in earthen shrimp ponds with suitable environmental parameters. Successful rearing of sub-adults of P. ornatus in fixed cages in shrimp ponds in Australia in field trials by Jones and Shanks (2008) (Fig. 3.8) is an encouraging sign for extending lobster farming to coastal earthen ponds.

Slipper Lobsters 107

Fig. 3.8 Lobster cages in the intake channel adjacent to shrimp pond in Australia. From ‘Spiny lobster ecology and exploitation in the South China Sea region’, (ed. K.C. Williams). ACIAR Proceedings No. 120, 21–25. Australian Centre for International Agricultural Research: Canberra.

3.10.4

Culture in sea-cages

That spiny lobsters can be grown successfully in sea-cages has been demonstrated in Vietnam, the Philippines, India, Australia and New Zealand. In laboratory rearing, Thenus sp. would bury in the sand bed provided at the bottom of the tank during the day and come out for feeding at night. In a few instances it was possible to rear T. orientalis without sand bed in FRP tanks (M. Vijayakumaran unpubl. data). Hence, it would be worthwhile to try to grow Thenus sp. in floating sea-cages with hard bottom and side walls like the ones used for rearing spiny lobsters in India or in net cages as used in Vietnam (Fig. 3.9a,b,c). In India, spiny lobsters are stocked in sea-cages at a density of up to 100 individuals/m2 (Vijayakumaran et al. 2009) and it may also be possible to stock slipper lobsters at high densities in sea-cages.

3.10.5

Disease in growout

No serious health problems were reported in nisto, juveniles and adult Thenus sp. when grown in high densities in indoor cement tanks in India (Anon. 2007) and in shallow raceways in the pilot-scale operation in Australia. In both these instances the temperature of rearing water was 27–28 °C. However, adult Thenus grown in two-tonne FRP tanks with sand bed at the NIOT laboratories in India developed shell lesion. The necrotic lesion first

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(a)

(b)

(c) Fig. 3.9

Floating sea-cages used for spiny lobster culture. (a, b) India; (c) Vietnam.

Slipper Lobsters

(a)

(c)

109

(b)

(d)

Fig. 3.10 (a) Tail fan necrosis, (b) swollen vent, (c) regurgitated proventriculus and (d) oedematous pleopods in adult Thenus orientalis. (Please see plate section for colour version of this figure.)

appeared in the tail fan and later spread to the pleopods (Fig. 3.10a,b,c,d). When the lesion progressed to the dermis, it increased further after each moult and turned into a melanised, often ulcerative, lesion. Green colonies of Vibrios, generally considered as pathogenic, were predominant in the ulcerative area (up to 12 × 104 colonies/g tissue), but the haemolymph was almost free of vibrios. Vibrio alginolyticus and Vibrio parahemolyticus were the two species isolated from the diseased lobsters. While V. alginolyticus formed yellow colony, generally termed as non-pathogenic and beneficial, V. parahemolyticus formed green colonies in TCBS agar. Sudden death of few lobsters with oedematous pleopods and regurgitated and bulged proventriculous was also observed. The rearing temperature was between 26 and 32 °C and most of the death occurred during summer when the temperature was above 30 °C. Vibrios that are ubiquitous are opportunistic pathogens and could turn pathogenic when the lobsters become weaker if the culture conditions are not favourable to lobsters.

3.11 CONCLUSIONS Successful aquaculture of slipper lobsters relies heavily on improving our understanding of their larval life cycle, moulting and nutritional needs. The potential of large-scale

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culture of scyllarids has been demonstrated by the high survival obtained recently in larval rearing and good growth rate and survival obtained in juvenile growout (Mikami 2007). To successfully develop the aquaculture of scyllarid species, biological, technical and economic issues should be addressed. Identification of factors affecting phyllosomas growth and the ways in which larval and juvenile nutrition can be optimised are important areas to be addressed. The development of productive, large-scale rearing systems with appropriate hydrodynamics and control of microbial conditions within the system are the technical challenges facing large-scale farming of slipper lobsters (Mikami & Kuballa 2007). The development of large-scale aquaculture for any species is dependent on economic feasibility based on market demand, price and evaluation of costs for optimal operations.

ACKNOWLEDGEMENTS We are thankful to Prof. T. Subramoniam, Kruba Ratnam, S. Muthukumar and T.S. Kumar, National Institute of Ocean Technology, Chennai, India for help rendered in preparation of the manuscript.

3.12

REFERENCES

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Kittaka, J. (1988) Culture of palinurid Jasus lalandi from egg stage to puerulus. Nippon Suisan Gakkaishi, 54, 87–93. Kittaka, J. (1994) Larval rearing. In: Spiny Lobster Management (eds B.F. Phillips, J.S. Cobb & J. Kittaka), pp. 402–423. Blackwell Science, Oxford. Kittaka, J. & Booth, J.D. (2000) Prospects for aquaculture. In: Spiny Lobsters: Fisheries and Culture (eds B.F. Phillips & J. Kittaka), 2nd edn, pp. 465–473. Blackwell Science, Oxford. Kizhakudan, J., Thirumilu, K., Rajapackiam. P., et al. (2004a) Captive breeding and seed production of scyllarid lobsters – opening new vistas in crustacean aquaculture. Marine Fisheries Information Service Technical & Extension Series, 181, 1–4. Kizhakudan, J., Thirumilu, K., Rajapackiam. P., et al. (2004b) Fishery of the sand lobster Thenus orientalis (Lund) by bottom-set gillnets along Tamil Nadu coast. Marine Fisheries Information Service Technical & Extension Series, 181, 6–7. Kumar, T.S., Vijayakumaran, M., Senthil, M.T., et al. (2009) Captive breeding and larval development of the scyllarine lobster Petrarctus rugosus. New Zealand Journal of Marine & Freshwater Research, 43, 101–112. Lavalli, K.L. & Spanier, E. (2007) Introduction to the biology and fishery of slipper lobsters. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 3–24. CRC Press, Boca Raton, London, New York. Lavalli, K.L., Spanier, E. & Grasso, F. (2007) Behaviour and sensory biology of slipper lobsters. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 133–182. CRC Press, Boca Raton, London, New York. MacDiarmid, A.B. & Kittaka, J. (2000) Breeding. In: Spiny Lobsters: Fisheries and Culture (eds B.F. Phillips & J. Kittaka), 2nd edn, pp. 485–507. Blackwell Science, Oxford. Mantel, L.H. & Farmer, L.L. (1983) Osmotic and ionic regulation. In: The Biology of Crustacea: Internal Anatomy and Physiological Regulation Vol. 5 (ed. L.H. Mantel), pp. 53–161. Academic Press, New York. Marinovic, B., Lemmens, J.W.T.J. & Knott, B. (1994) Larval development of Ibacus peroni Leach (Decapoda: Scyllaridae) under laboratory conditions. Journal of Crustacean Biology, 14, 80–96. Matsuda, H. & Takenouchi, T. (2005) New tank design for larval culture of Japanese spiny lobster, Panulirus japonicus. New Zealand Journal of Marine & Freshwater Research, 39, 279–285. Mikami, S. (1995) Larviculture of Thenus (Decapoda, Scyllaridae), the Moreton Bay bugs. PhD thesis, University of Queensland. Mikami, S. (2005) Moulting behaviour responses of Bay lobster, Thenus orientalis, to environmental manipulation. New Zealand Journal of Marine & Freshwater Research, 39, 287–292. Mikami, S. (2007) Prospects of aquaculture of bay lobsters (Thenus spp.). Bulletin of Fisheries Research Agency, 20, 45–50. Mikami, S. & Greenwood, J.G. (1997a) The complete development and comparative morphology of larval Thenus sp. Nov. (Decapoda: Scyllaridae) reared in the laboratory. Journal of Crustacean Biology, 17(2), 289–308. Mikami, S. & Greenwood, J.G. (1997b) Influence of light regimes on phyllosomal growth and timing of moulting in Thenus orientalis (Lund) (Decapoda: Scyllaridae). Marine & Freshwater Research, 48, 777–782. Mikami, S. & Kuballa, A.V. (2004) Overview of lobster aquaculture research. Proceedings of the Second hatchery and feed technology Workshop (eds. S. Kolkovski, J. Heine & S. Clarke), pp. 132–135. Western Australian Department of Fisheries – Research Division, Sydney. Mikami, S. & Kuballa, A.V. (2007) Factors important in larval and post larval molting, growth and rearing. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 91–110. CRC Press, Boca Raton, London, New York. Mikami, S. & Takashima, F. (2000) Functional morphology of the digestive system. In: Spiny Lobsters: Fisheries and Culture (eds B.F. Phillips & J. Kittaka), 2nd edn, pp. 601–610. Blackwell Science, Oxford. Mikami, S., Greenwood, J. & Takashima, F. (1994) Functional morphology and cytology of the phyllosomal digestive system of Ibacus cliatus and Panulirus japonicas (decapoda, Scyllaridae and Palinuridae). Crustaceana, 67, 212–225. Mosing, J. & Fallu, R. (2006) Australian Fish Farmer. A Practical Guide to Aquaculture, p. 444. Landlinks Press, Collingwood, Victoria. Pessani, D., Pisa, G. & Gatelli, R. (1999) The complete larval development of Scyllarus arctus (Decapoda: Scyllaridae) in the laboratory, pp. 143–144. Abstract of the 7th Colloquium Crustacea Decapoda Mediterranea, Lisbon.

Slipper Lobsters 113 Phillips, B.F. & Sastry, A.N. (1980) Larval ecology. In: The Biology and Management of Lobsters, Vol. 2: Ecology and management (eds J.S. Cobb & B.F. Phillips), pp. 11–57. Academic Press, New York. Radhakrishnan, E.V. & Vijayakumaran, M. (1986) Observations on the feeding and moulting of laboratory reared phyllosoma larvae of the spiny lobster, Panulirus homarus (Linnaeus) under different light regimes. Proceedings of the Symposium on Coastal Aquaculture (Marine Biological Association India), 4, 1261–1266. Radhakrishnan, E.V. & Vijayakumaran, M. (1995) Early larval development of the spiny lobster, Panulirus homarus (Linnaeus 1758). Crustaceana, 68(2), 151–159. Radhakrishnan, E.V., Manisseri, M.K. & Deshmukh, D. (2007) Biology and fishery of the slipper lobster Thenus orientalis, in India. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 309–324. CRC Press, Boca Raton, London, New York. Ritar, A.J., Greg, G.S. & Thomas, C.W. (2006) Ozonation of seawater improves the survival of larval southern rock lobster, Jasus edwardsii, in culture from egg to juvenile. Aquaculture, 261, 1014–1025. Ritz, D.A. & Thomas, L.R. (1973) The larval and post larval stages of Ibacus peronii Leach (Decapoda, Reptantia, Scyllaridae). Crustaceana, 24, 5–16. Robertson, P.B. (1968) The complete larval development of the sand lobster, Scyllarus americanus (Smith) (Decapoda, Scyllaridae) in the laboratory with notes on on larvae from plankton. Bulletin of Marine Science, 18(2), 294–342. Robertson, P.B. (1979) Larval development of the scyllarid lobster Scyllarus planorbis Holthuis reared in the laboratory. Bulletin of Marine Science, 29(3), 320–328. Rodrigo, L.A. & Jorge, O.F. (2003) Culture of turbot (Scophthalmus maximus) juveniles using shallow raceways tanks and recirculation. Aquacultural Engineering, 32, 113–127. Rudloe, A. (1983) Preliminary studies of the mariculture potential of the slipper lobster Scyllarides nodifer. Aquaculture, 34, 165–169. Saisho, T. (1966) Studies on the phyllosoma larvae with reference to the oceanographical conditions. Memoirs of the Faculty of Fisheries, Kagoshima University, 9, 84–90. Sekiguchi H., Booth, J.D. & Webber, W.R. (2007) Early life histories of slipper lobsters. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 69–90. CRC Press, Boca Raton, London, New York. Senthil, M.T., Vijayakumaran, M., Remany, M.C., et al. (2004) Early phyllosoma larval stages of the sand lobster, Thenus orientalis (Lund, 1793). In: Proceedings of MBR 2004 National Seminar on New Frontiers in Marine Bioscience Research (eds S.A.H. Abidi, M. Ravindran, R. Venkatesan & M. Vijayakumaran), pp. 161–168. Allied Publishers, New Delhi. Sharp, W.C., Hunt, J.H. & Teehanhern, W.H. (2007) Observations on the ecology of Scyllarides aequinoctialis, S. nodifer, and Parribacus antarcticus and a description of the Florida scyllarid lobster fishery. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 231–242. CRC Press, Boca Raton, London, New York. Sheppard, J.K., Bruce, M.P. & Jeffs, A.G. (2002) Optimal feed pellet size for culturing juvenile spiny lobster Jasus edwarsii (Hutton 1875) in New Zealand. Aquaculture Research, 33, 913–916. Sims, H.W. & Brown, C.L. (1968) A giant scyllarid phyllosoma larva taken north of Bermuda (Palinuridae). Crustaceana, (Suppl. 2), 80–82. Skinner, D.M. (1985) Moulting and Regeneration. In: The Biology of Crustacea: Integument, Pigments and Hormonal Processes, Vol. 9 (ed. D.E. Bliss), pp. 44–128. Academic Press, New York. Spanier, E. & Lavalli, K.A. (2007) Slipper lobster fisheries – present status and future perspectives. In: The Biology and Fisheries of the Slipper Lobster (eds K.A. Lavalli & E. Spanier), pp. 377–391. CRC Press, Boca Raton, London, New York. Stewart, J. & Kennelly, S.J. (1997) Fecundity and egg-size of the Balmian bug Ibacus peroni (Leach 1815) (Decapoda, Scyllaridae) of the east coast of Australia. Crustaceana, 70, 191–197. Stewart, J. & Kennelly, S.J. (2000) Growth of the scyllarid lobsters Ibacus peroni and I. chacei. Marine Biology, 136, 921–930. Subramaniam, T. (2004) Fishery of sand lobster Thenus orientalis (Lund) along Chennai coast. Indian Journal of Fisheries, 51, 111–115. Takahashi, M. & Saisho, T. (1978) The complete larval development of the scyllarid lobster, Ibacus ciliates (Von Siebold) and Ibacus novemdentatus Gibbes in the laboratory. Memoirs of the Faculty of Fisheries, Kagoshima University, 27, 305–353. Tan, C.L. (1997) A livelihood option for coastal communities: Lobster culture in Guimaras (Philippines). Aquaculture Asia, 35–37.

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Thomas, C.W., Carter, C.G. & Crear, B.J. (2003) Feed availability and its relationship to survival, growth, dominance and antagonistic behaviour of the southern rock lobster, Jasus edwarsii in captivity. Aquaculture, 215, 45–65. Tolomei, A., Crear, B. & Johnston D. (2003) Diet immersion time: effects on growth, survival and feeding behaviour of juvenile southern rock lobster, Jasus edwarsii. Aquaculture, 219, 303–316. Tuan, L.A. & Mao, N.D. (2004) Present Status of Lobster Cage Culture in Vietnam. In: Spiny lobster ecology and exploitation in the South China Sea region. Proceedings of a workshop held at the Institute of Oceanography, Nha Trang, Vietnam, July 2004 (ed. K.C. Williams), pp. 21–25. CSIRO, Australian Centre for International Agricultural Research, Canberra. Vijayakumaran, M. & Radhakrishnan, E.V. (1986) Effect of food density on feeding and moulting of phyllosoma larvae of the spiny lobster Panulirus homarus (Linnaeus). Proceedings of the Symposium on Coastal aquaculture (Marine Biological Association of India), 4, 1281–1285. Vijayakumaran, M. & Radhakrishnan, E.V. (2003) Control of epibionts with chemical disinfectants in the phyllosoma larvae of the spiny lobster Panulirus homarus (Linnaeus). In: Aquaculture Medicine, Centre for Fish Disease Diagnosis and Management (eds I.S.B. Singh, S.S. Pai, R. Philip & A. Mohandas), pp. 69–72. CUSAT, Kochi, India. Vijayakumaran, M., Venkatesan, R., Senthil, M.T., et al. (2009) Farming of lobsters in sea cages in India. New Zealand Journal of Marine & Freshwater Research, 43, 623–634. Vijayanand, P., Murugan, A., Saravanakumar, K., et al. (2004) Abstract. Experimental fattening of sand lobster Thenus orientalis (Lund). In: Ocean Life Food & medicine Expo 2004. International Conference & Exposition on Marine Living Resources of India for Food and Medicine. Aquaculture Foundation of India, Chennai, p. 52.

4

Mud Crab Aquaculture

Brian D. Paterson and David L. Mann

4.1 INTRODUCTION Portunid crabs emerged as a new aquaculture species because of their high value and the ready availability (at the time) of wild-caught ‘seed’ crabs. They also fit into existing coastal mariculture infrastructure – an important feature in areas where shrimp farming has been hit with viruses. However, the different marketing options available and the biological quirks of crabs make them a different prospect for farming from shrimp. Portunid crabs, particularly mud crabs, have a heavy, calcified exoskeleton and robust, strong claws. They are built to dismember and crush other crabs and molluscs – habits that are obviously not a good fit with high-density mariculture! The biggest crab tends to win all the fights – so it is no surprise that portunid crabs are amongst the fastest-growing crustaceans on earth. In the fishery, portunids take two years to reach commercial size (300–800 g depending on species), but in aquaculture this can be reduced to within one year (Trino et al. 1999; Christensen et al. 2004; Ut et al. 2007b). Juvenile portunid crabs typically double their weight each time they moult, but this weight change is tempered as maturation sets in by redirecting energy into sexual development (Hill 1976). While the weight change is massive upon moulting, the crabs are soft and helpless for about an hour. Even when their cuticle has hardened up on the outside, days and weeks may elapse before they have filled their impressive carapace with meat. For significant periods after moulting the crabs can be ‘empty’. Scylla and other portunids are usually marketed live. Mud crabs are well suited to live marketing, which is an advantage for countries with limited infrastructure for seafood processing and distribution. Because of their heavy shell and lack of abdomen, edible recovery in crabs is lower than it is for tailed decapods like shrimp or lobster – but in mud crabs, the gonads of mature females provide an added premium (Chiou & Huang 2003). Portunid crabs are ‘farmed’ in a variety of ways. The simplest forms of this are arguably not aquaculture at all. Firstly, soft empty post-moult crabs are harvested from the fishery,

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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individually contained and fed with trash fish to ‘fatten’ them for market. In one of Nature’s ironies, the opposite practice, soft-shell crab production, is increasingly becoming favoured; catching hard-shell crabs, feeding them individually until they moult and double their weight and then marketing them while they are soft. Competition for a limited resource of wild crabs has inevitably seen these live-holding practices evolve a longer-term focus. Here, farmers purchase small juveniles or ‘seed’ crabs from the wild and rear them extensively in earthen ponds for on-growing to commercial size on a diet of trash fish. Marketing the crop as hard-shell is still the most common practice, but regional improvements in transport and processing infrastructure mean that production of soft-shell product for export is increasing. Inevitably, demand for crab ‘seed’ soon became so intense that coastal mud crab stocks diminished and supply of juveniles to crab farming became a significant bottleneck to industry growth (Shelley 2008). In response to this, mud crab hatchery methods have now been developed, following an international collaborative effort, putting crab farming on a more sustainable footing. This chapter describes the current status of mud crab farming in Asia; discusses the suitability of the mud crab biology and life cycle to aquaculture; gives an update on recent developments in research and examines a number of significant areas requiring further development

4.2

PORTUNID CRAB AQUACULTURE

Crab farming is largely centred on Asia, and marine farming of portunid crabs (primarily mud crabs Scylla spp.) accounts for only a third of global farmed crab production (about 660,000 tons). The remainder involves freshwater farming of the Chinese mitten crab Eriocheir japonica sinensis (FAO 2008). The high value of the latter means that portunid crabs account for only one-fifth of the total value of farmed crab production, which is US$2.8 billion (FAO 2008). Aquaculture of mud crabs Scylla spp. and more recently swimming crabs Portunus spp. was originally based on collection and extensive on growing of juvenile or ‘seed’ crabs in earthen ponds. Farming occurs in coastal areas of southern Asia and Southeast Asia and in southern China. While all crabs are generally harvested for the hard-shell live market, production of soft-shell crabs is becoming more common. Mud crab females with mature ovaries also attract a price premium, presenting an opportunity for mono-sex culture (Trino et al. 1999; Khatun et al. 2009). The relatively high prices and ready live transportability and marketability of Scylla spp., added to the fact that they did not initially fall victim to significant pathogens such as WSSV (Lavilla-Pitogo et al. 2007), also meant that crab farming was for many years an attractive alternative in areas where viral outbreaks prevented penaeid shrimp farming. However, this explosion of interest quickly threatened the sustainability of coastal crab populations. The subsequent recruitment difficulties were at best a tight brake upon industry expansion and at worst stopped crab farming altogether in some regions. Wild juveniles also presented some practical problems. As demand forced harvesting at smaller and smaller sizes it became more and more difficult to identify them accurately to species, meaning that in some areas farmers may inadvertently buy less desirable varieties (such as S. olivacea) in the mix of juveniles on offer (Christensen et al. 2004; Walton et al. 2006a).

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Fortunately, the recent development of hatchery and nursery techniques for mud crabs and other species promises to set the industry on a more sustainable footing, and pave the way for an expansion of production (Christensen et al. 2004; Walton et al. 2006a; Mann et al. 2007; Rodriguez et al. 2007).

4.2.1

Growout

Scylla spp. are apparently the fastest-growing of all commercially farmed crustaceans. Average weekly gain for mud crabs Scylla spp. grown to harvest size in ponds is calculated at around 10 g/wk compared to values of only about 2 g/wk for both the so-called ‘giant’ tiger shrimp Penaeus monodon and the freshwater prawn Macrobrachium rosenbergii (Wickens & Lee 2002). Apparent growth rates for Scylla serrata and S. paramamosain calculated from production figures (Trino et al. 1999; Christensen et al. 2004) give incredible average weekly gains of 14 to 28 g. It is easy to understand why evolution would favour mud crabs that grow fast – giving them protection from other crabs and predators. However, it is also possible that at least some of the commercial growth is illusory. For example, if large crabs kill off smaller crabs, or if farmers don’t harvest any residue of unmarketable small individuals in the pond, then the observed average size can easily become inflated. The downside for crab farmers is that the production from mud crab ponds is about one-tenth of what is possible from a well-managed intensive penaeid shrimp pond (up to 2 tons/ha compared to up to 20 tons for shrimp (Wickens & Lee 2002). The prices for mud crabs would need to be high indeed to make up for that low productivity. Yields for mud crabs in ponds are in fact similar to those for other commercial decapods with claws; freshwater prawns and crayfish (Wickens & Lee 2002). Like the latter, Scylla species and other swimming crabs have to be stocked at low densities (<1 crab/m2) to boost survival. Mud crabs are grown in earthen ponds, with perimeter fences to prevent the crabs from walking out (Fig. 4.1). They are known to burrow in the wild, but this behaviour does not seem to become commonly expressed when crabs are reared in ponds. Soon after stocking, the ‘natural’ levels of food in the pond environment are sufficient to satisfy the early-stage crablets. The crab’s diet is eventually supplemented with low-cost local fishery products and wastes (Christensen et al. 2004). The crop is initially trapped out for sale or finally gathered by drain harvesting. Obviously, the crabs must be tied (restraining their claws) immediately to allow for efficient bulk shipment. Polyculture is possible with mud crabs and other portunids under certain circumstances. For example, it is feasible to grow mud crabs as a ‘house-keeping’ crop underneath floating pens or cages holding fish. Generally however, free polyculture of fish and crabs together is not recommended. Depending upon the timing of the introduction of each crop and the relative size and growth rate of the fry, the fish could eat the juvenile crabs – or vice versa (particularly when rearing a swimming portunid like P. pelagicus).

4.2.2

Soft- shell crabs

Large mud crab juveniles up to 60 g in size are induced to moult using multiple limb autotomy (removing all claws and legs, leaving only the paddles) and stocked at high density and fed in small earthen ponds (Dat 1999). After about one and half weeks the pond

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Fig. 4.1 pond.

Fences serve an important purpose in this crab farm in Vietnam; they keep the crabs in the

is drain harvested and the pre-moult crabs with well developed limb-buds are transferred in small groups into floating baskets so that they can be regularly checked and harvested as soon as they moult. Confining crabs in this manner allows them to be stored and checked efficiently at much higher densities than is typical for crab culture. For this reason, greater care is required in feed management and control of pond water quality to avoid stressing the crabs, particularly as they undergo moulting (Dat 1999).

4.2.3

Crab fattening

A broad category that spans several short-term culture practices, ‘fattening’ is primarily a value-adding process, like soft-shell crab production. For example, low-value ‘lean’ marketsized mud crab may be taken and fed for a short period to harden the shell and restore the flesh content (Dat 1999). But taken more loosely, ‘fattening’ also encompasses other practices such as maturation of females for sale, as well as a kind of proto-aquaculture – the growth of sub-commercial sized mud crabs through a single moult to improve their marketability (Trino & Rodriguez 2001). As such, fattening practices can vary in their intensity, from labour-intensive housing and maintenance of crabs in baskets or enclosures through to pond practices similar to longer-term pond culture (Shelley 2008; Khatun et al. 2009). Crabs are housed and fed intensively in bamboo cages with up to 25 kg crabs/m2, so man-

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agement of fresh feed to the ponds is crucial to avoid deterioration in water quality (Dat 1999).

4.3

BIOLOGY AND LIFE CYCLE

Mud crabs possess almost all the key characteristics required of a viable candidate for aquaculture (high market price; rapid growth; simple feed and abiotic requirements; ready availability/production of juveniles and no disease hurdles) (Wickens & Lee 2002) but they fail in one crucial area. Their aggressive appearance and behaviour, probably a major reason for their premium market value, is a profound biological obstacle to intensive communal culture.

4.3.1

Habits

Mud crabs (Scylla spp.) are amongst the largest of the Portunidae, the swimming crabs, a group that includes a number of other commercially important fishery species in the IndoPacific (Portunus spp.) and in the Americas (Callinectes spp.). Four species of Scylla are recognised, of which the largest, Scylla serrata, is the most widespread, ranging from the coast of southern Africa around the coast of southern Asia through to the oceanic Western Pacific ( Keenan et al. 1998; Keenan 1999; Macintosh et al. 2002). While all species are to varying degrees estuarine in habit (they are all quite literally ‘mangrove crabs’), spawning Scylla serrata migrate the furthest from the mangal, and spawn in the open sea. The other Scylla species are largely restricted to Southeast Asia, and their life histories are more closely tied to estuaries. They were only distinguished relatively recently from amongst the existing diversity of local Scylla serrata stocks. Despite the large size of these crabs as adults (300 g or more), the fifth leg is a typical swimming paddle. While not pelagic by preference, the crabs are able to swim, even as adults. Typically, they have a cryptic benthic habit, and are readily found on intertidal flats and mangrove fringes (Hill et al. 1982; Le Vay et al. 2001; Walton et al. 2006b). To survive in this habitat, Scylla species possess a thick exoskeleton, robust claws and a capacity for prolonged survival out of water (Varley & Greenaway 1992). As adults, they are nocturnal predators of molluscs and other crabs (Hill 1976, 1979). While foraging, their singular morphology and sheer size probably protect them from all but the largest and most determined of predators. While they dig burrows in mangroves as adults, as juveniles they hide during daylight under rocks, amongst marine plants such as sea grasses or by burrowing into the sediment. As they grow, their habitat preferences change; for example juvenile S. serrata is readily found at high densities on complex intertidal mudflats while the adults forage sub-tidally at much lower densities (Hill et al. 1982; Walton et al. 2006b). Growing best under tropical conditions of temperature, their estuarine lifestyle means that juvenile and adult mud crabs tolerate a wide range of salinities (Davenport & Wong 1987; Chen & Chia 1997; Nurdiani & Zeng 2007).

4.3.2

Growth

The widely occurring Scylla serrata is the fastest- and largest-growing of the four scylla species, exceeding 1 kg as an adult (Williams & Primavera 2001). In contrast, tropical

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Fig. 4.2 Juvenile mud crabs double their weight when they moult. In this example, the post-moult crab has also substantially regenerated a claw (missing from the exuvia on the right).

Scylla paramamosain, the species the most commonly farmed, matures at a slightly smaller size, and is marketed at only 200–250 g. S. serrata moults 15–17 times after metamorphosis, over 2 or 3 years under natural conditions (Williams & Primavera 2001). Juvenile and sub-adult mud crabs, like other portunids, double their weight at each moult by inflating a greatly expanded new cuticle through imbibing large amounts of seawater (Neufeld & Cameron 1994) (Fig. 4.2). As expected, the moult increments of portunids decrease below 100% of pre-moult weight with the onset of sexual maturation (Josileen & Menon 2005; Paterson et al. 2007). Immediately after moulting, the crabs are soft and vulnerable to predation. The large moult increment also means that recently moulted crabs extracted from the fishery (when they harden and first resume feeding) are more or less ‘empty’ of edible meat. The crabs take some time to ‘grow into’ their newly expanded shell.

4.3.3

Reproduction and life history

Scylla, like other portunids, display mate-guarding behaviour. Hard-shelled males compete for mature pre-moult females, and the successful male shelters the female with his body, protecting the female during ecdysis and mating immediately afterwards (Jivoff & Hines 1998). This strategy is expressed through a familiar form of sexual growth dimorphism. The males ultimately grow to a larger total weight and invest proportionally more of their body mass into their claws. In contrast, the cost of gonadal development and spawning

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Fig. 4.3 Female mud-crabs (in this case Scylla paramamosain) are readily distinguished from males by their proportionally smaller claws and wider abdominal flap.

reduces total somatic growth amongst females, and they show no exaggeration of claw size (Heasman 1980) (Fig. 4.3). After mating and hardening, the female retains the spermatophore internally and uses it later to fertilise the extruded eggs she attaches to the ovigerous pleopods within her abdominal flap. Mature S. serrata females can extrude egg sponges totalling around 3 million eggs. Mud crabs spawn all year, especially in the tropics, and can fertilise several batches of eggs using sperm from just one mating. The spermatophore serum contains potent antimicrobial compounds to ensure longevity of the spermatozoa (Jayasankar & Subramoniam 1999). The ovigerous females then migrate so that the eggs hatch offshore. There the newly hatched larvae feed on zooplankton, growing through five zoeal moults before metamorphosing into the ‘lobster-like’ megalops stage. This recruits into the estuary and it settles amongst the tidal flats or mangrove fringe areas and undergoes a further metamorphic moult to the first crab stage or ‘crablet’.

4.3.4

Mud crabs as candidates for aquaculture

Decapods with large chelae are often difficult to farm because of aggression and territorial behaviour (Wickens & Lee 2002) and, locked in an ‘arms race’ of their own, Scylla species probably take the concept of ‘difficult’ to an entirely new level. It is likely that in terms of evolution, the need to win fights above all other factors drives the phenomenal growth rate

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achieved by mud crabs and other portunids. The biology of mud crabs is a story of rapid somatic growth rate, extraordinary levels of fecundity and dispersal. As explained in the previous section, the high fecundity meant that an industry could develop rapidly, based upon a ready supply of wild juveniles. However, their prolific life strategy did not always support the huge fishing pressure that resulted. Thus the primary focus of technology development (section 4.4) has been to develop sustainable methods of larval rearing.

4.4

TECHNOLOGY DEVELOPMENT

Practical hatchery methods are no longer a bottleneck preventing sustainable farm production of mud crabs (Marichamy & Rajapackiam 2001; Quinitio et al. 2001; Wang et al. 2005). Variations on these methods applied to mud crabs and other portunid species has enabled routine production of crablets for fishery enhancement/ restocking purposes (Zmora et al. 2005; Obata et al. 2006; Lebata et al. 2009) as well as evaluation of other species as candidates for aquaculture (Marshall et al. 2005; Parado-Estepa et al. 2007; Nicholson et al. 2008).

4.4.1

Hatchery

Where spawning occurs year-round in the tropics, hatchery production can rely upon collection of egg-bearing females. Because female Scylla store viable spermatophores for months after mating (Jayasankar & Subramoniam 1999), mature female mud crabs can also be brought into maturation tanks and allowed to spawn naturally, though spawning can be accelerated by eyestalk ablation (Millamena & Quinitio 2000; Zeng 2007). During maturation, broodstock can be fed either on fresh diets or fresh diets supplemented with formulated ingredients (Millamena & Quinitio 2000; Alava et al. 2007a). While HUFA lipids are recognised to be an important component of the maturation diets, further research in this area is required (Alava et al. 2007b; Parado-Estepa et al. 2007). Larval rearing methods for mud crabs have been readily adopted by shrimp hatcheries, where perhaps the only departure from conventional shrimp practices involves the feeding of live rotifers to the first larval stage (Ruscoe et al. 2004; Davis et al. 2005); a practice more common with finfish larval rearing. Where Artemia of appropriate nutritional quality are available, then feeding early ‘teardrop’ or ‘umbrella’ stage Artemia to the Z1s is also possible (Nghia et al. 2007); the important thing being to not overwhelm the small crab larvae with large prey items. An example of a rotifer/Artemia feeding table for Scylla serrata is presented in Table 4.1. Batch tank culture is typically practised, with periodic exchanges to control microbial and chemical quality of the water (Seneriches-Abiera et al. 2007). Temperatures are kept as high as practical to facilitate rapid development (28–30 °C). Because larval development of Scylla species proceeds in or near estuaries, salinities close to full marine salinity are not obligatory even for Scylla serrata (Nurdiani & Zeng 2007). It is also true that the salinity in estuaries can fall dangerously low during heavy rain. For this reason, it is always prudent for hatcheries in or near estuaries to have ample seawater storage capacity. Larval rearing of Scylla species can involve relatively high mortality. Of course, the easiest way to increase percentage survival is to dramatically reduce larval densities. But in the end, high survival in this case would either reduce the hatchery output or would

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Table 4.1 An example of a feeding schedule for Scylla serrata larvae culture. zoea1 Rotifers, L strain Artemia nauplii

zoea2

zoea4

zoea5

megalop

crab 1

10 to 15 per mL

1.0 to 1.5 per mL 0.2 to 0.3 per mL

Artemia juveniles

0.1 per mL

Artemia adults Particulate diet

zoea3

1.0 to 2.0 g per ton

require construction of a huge inefficient hatchery! In fact, the large egg masses produced by Scylla spp. mean that there is usually a gross excess of zoea. At a typical growout density of say 0.5 crab/m2, a single spawner hatching 3 million eggs could theoretically stock 600 hectares of ponds if all larvae survived. This is enough to satisfy the requirements of an entire country! So what does it matter if survival through the hatchery is only 5 to 10%? Crab zoeas are predators and rely on their feeding appendages to fragment and render their food (Lumasag et al. 2007). A recent study recommended that tanks have darker backgrounds to aid feeding efficiency and reduce stress (Rabbani & Zeng 2005), though feeding efficiency will also be influenced by prey density. Rotifers and Artemia are of course used for convenience in aquaculture (Holme et al. 2006a), and rotifers are likely to be considerably easier to catch than the larvae’s natural food (Lumasag et al. 2007). Nutritional problems may cause some reported difficulties in survival and growth. Cannibalism can be a problem, ironically, when survival is high at high densities, when zoea moult through to the next stage and prey upon smaller larvae remaining in the previous instar. This can become particularly acute when development in larval populations becomes very asynchronous – larvae begin moulting through to the clawed, lobster-like megalops while there is still a large residue of Z4s and Z5s in the tank. ‘Moult-death syndrome’ (MDS) is also sometimes reported. This occurs when larvae ‘hang-up’ or are unable to free themselves from their old exoskeleton, and die when moulting. The exact cause of this syndrome is not fully understood, but inadequate nutrition is often suspected (Holme et al. 2006b, 2007a). Larval nutrition studies have benefited from the development of particulate larval diets, but more research is needed (Holme et al. 2009). Live feeds are not nutritionally complete and are routinely fortified either with algae or commercial booster formulations (Genodepa et al. 2007). Fully manufactured larval diets are attractive because of the expense and labour involved in growing live feed and algae cultures (Holme et al. 2006a). Initial trials using micro-bound diets are promising, though there is still some distance to go before they can be used commercially (Holme et al. 2006a). Current test diets are well accepted by the zoea, which is already an achievement given that the larvae must manipulate the diet particles within them disintegrating and fouling the water (Genodepa et al. 2007; Lumasag et al. 2007). Studies already show that dietary HUFA levels can improve survival (Nghia

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et al. 2007) but practical demonstration of this can be complicated by other uncontrolled factors impinging upon hatchery outcomes. Bacterial problems are largely blamed for the ‘crashes’ in hatchery tanks, leading in some cases to near-total mortality. For this reason, there has been too much reliance upon antibiotics such as oxytetracycline (OTC) in hatcheries to manage the risks of batch failure (Nghia et al. 2007). Fungi have also caused issues in crab hatcheries, requiring a different solution (de Pedro et al. 2007), but widespread use of antibiotics in this manner is not recommended (and it is often illegal in some countries). The biggest problem here is not the tiny residues of antibiotics that might persist in the larvae beyond the hatchery phase and any uncertainty over food safety risk to humans. The greatest threat from unregulated use of antibiotics is the build-up of resistance to those antibiotics amongst environmental microbes, which can not only recoil back on the hatchery operators but also lead to the transfer to resistance to other animal (and human) pathogens (Akinbowale et al. 2006, 2007).

4.4.2

Nursery

High-density nursery production has emerged as an important intermediate practice between hatchery production of megalops and first crablets (Crab 1) on the one hand and the routine growout to harvest size at low density on the other. The megalops is readily transported in water (Quinitio 2000), but on growing a couple of instars post-settlement has a practical benefit in that the more robust crablets can be shipped to farms in damp packing material, without water. Nursery conditioning is also beneficial in cases where juveniles are reared for restocking into the wild (Davis et al. 2005; Lebata et al. 2009). Farmers are understandably reluctant to take on the increased risk of on-growing small hatchery-reared juveniles, at least until they are confident in the consistency and quality of the product. While on the face of it these high-density nursery methods make more efficient use of space during early growth stages, the work in this area is also providing us with our first insights into the growth and survival of mud crabs and other portunids at high densities. Mortalities of 50% or more are still reported during the nursery phase of S. serata and S. paramamosain (Mann et al. 2007; Rodriguez et al. 2007; Ut et al. 2007a). Cannibalism is believed to be a major cause of this mortality, but inter-batch variation is also reported, implying differences in the ‘condition’ of megalopa leaving the hatchery (Mann et al. 2007). Reducing stocking density reduces but does not eliminate the percentage mortality (Ut et al. 2007a). As crablets grow, one crab occupies more area and the acceptable stocking density declines. Cannibalism and aggression amongst crablets leaves its mark. Within weeks of stocking a pond with post-settlement P. pelagicus, the stable isotopic profile of the largest juveniles in a pond indicated a mixed diet of pellets and other crabs (Møller et al. 2008). The survivors from even a brief period of nursery growout already show a relatively wide divergence in size (as many as four or five instars may be present at harvest!), and a high proportion of crablets are missing one or both chelae, presumably because of agonistic encounters with other individuals (Mann et al. 2007). Repeated loss and regeneration of claws by some individuals may explain the wide divergence in size (Paterson et al. 2007). Mortality is reduced by including shelter or artificial habitats in nursery systems (Mann et al. 2007; Ut et al. 2007a). Increased environmental complexity, even through the addition of simple objects such as as paving stones, apparently confers a benefit by expanding the

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available surface area or reducing the encounters between individuals (Marshall et al. 2005; Mann et al. 2007; Ut et al. 2007a).

4.4.3

Growout and feeding

Recent mud crab growout research has examined practical issues such as stocking density and diet supplementation in extensive pond or mangrove-enclosure systems. To reduce the costs of aggression and cannibalism on survival during growout, crabs must be stocked at 0.5 crab/m2 – anything more than that and mortality becomes unacceptable (Trino et al. 1999). Stocking density is so low that in the first few months there is enough biota in the pond environment to sustain the crablets. ‘Supplemental’ feeding is only required once the crabs outgrow the natural productivity (Christensen et al. 2004). The crabs are fed using fresh feeds, such as fish and molluscs and fishery by-products, since Scylla species are understood to be predators, at least as adults (Hill 1976). Development of formulated diets is underway (Trino et al. 2001), and for experimental work, mud crabs and Portunus spp. can be grown on high-quality fishmeal-based penaeid shrimp diets (Mann et al. 2007; Paterson et al. 2007) though these are too expensive for commercial adoption. Amongst mud crabs, males grow significantly heavier than females, while ‘roed-up’ females achieve a market premium (Trino & Rodriguez 2001). Monosex culture also apparently alleviates cannibalism when fattening ‘lean’ male and female S. serrata (Cholik & Hanafi 1992; Trino & Rodriguez 2001). This finding deserves closer attention as it apparently contradicts the outcome expected from normal male ‘mate guarding’ behaviour; crucially the survival of each sex in the mixed-sex treatment was not reported. Trials of monosex male and female production of S. serrata show that males and females show similar levels of survival when not mixed together, but there are different economic outcomes depending upon the respective growth rates of males or females (Trino et al. 1999, 2001). Interestingly, monosex culture did not improve survival of S. olivacea during fattening in bamboo pens, though this is consistent with the presence of natural mangrove habitat and with the sexes being of similar average weight (Khatun et al. 2009).

4.5

FUTURE DEVELOPMENTS

Improvements in productivity that address cannibalism loom large as areas of development for mud crab farming. This involves two broad areas, i.e. improving survival in the ponds and investigating the feasibility of containerised production. The latter research area emerges from experience gained from individual rearing of crabs for feed research. Development of alternatives to ‘trash fish’ is also becoming a priority because low-cost feed is emerging as the next bottleneck to industry expansion. Of course, while sustainable hatchery production is now common, some effort is being made to minimise industry reliance upon antibiotics.

4.5.1

Questions about growth

Growout feed research has been expedited by the ready supply of hatchery-reared juveniles. Of course, the cannibalistic nature of the crabs has also meant that these experiments

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involve distribution of feeds to large numbers of isolated crablets – using individuals as true replicates. Unlike growth trials involving fish or shrimp, where growth is continuous or relatively so, growth trials with rapidly growing juvenile portunids involve large amounts of individual variation. The juveniles basically double their weight each time they moult. Added to this, at any given time treatments might contain individuals of three different instars, and so differ up to fourfold in wet weight. Fortunately, using initial weight as a covariate accounts for much of the variation in size in any given instar – an interesting point for future selective breeding programmes. Compared in terms of specific growth rates (SGR), wild and hatchery-reared seed crabs achieve similar growth rates and survival in ponds (Ut et al. 2007b). However, SGR is unlikely to be a suitable growth model for crabs – for example, the graphs in one recent study show that growth is not exponential and only departs slightly from linear (Christensen et al. 2004). Daily Growth Coefficient (DGC) may be more widely applicable, but as it calculates a change in a single dimension from a change of weight or volume the efficacy of this ‘width’ needs to be judged against simply measuring the growth of carapace width. Issues of measuring growth aside, published growth rates for mud crabs in nursery and growout ponds should be treated with caution because of the bias inherent in sampling and harvesting. Calculating growth to harvest as average weekly gain (AWG) (grams per week (g/wk)) shows that crops of S. serrata, S. paramamosain and S. olivacea all achieve impressive AWGs of 13–27 g/wk after 120–160 days (Trino et al.1999; Trino & Rodriguez 2002; Christensen et al. 2004). However, crabs measured/harvested following indoor feed trials or nursery trials, show much lower AWG values (0.05 to 5 g/wk) (Sheen & Wu 1999; Sheen 2000; Catacutan 2002; Ut et al. 2007b), as do crabs reared in nursery ponds (AWG of 0.7 to 3.6 g/wk) (Rodriguez et al. 2007). The highest of these values (2–5 g/wk) may be consistent with the slower weekly gains expected at early instars. But the reports of growth rates for extended periods as slow as or even slower than that typical of slower-growing aquaculture species such as the giant tiger shrimp Penaeus monodon ought to ring alarm bells. The recorded growth of less than 0.1 g/wk for early instar juveniles for several weeks (Sheen & Wu 1999; Sheen 2000) is a concern since this is around half of the typical farm growout period. Scylla would normally be close to 100 g in weight after 60–80 days. Animals will grow poorly in sub-optimal conditions. But what is ‘sub-optimal’ for mud crabs? Factors possibly explaining slower growth by crabs from laboratory feeding trials include lower ambient temperatures, disturbance during regular cleaning/maintenance, inadequate nutrition, or even social isolation. But this does not explain why crablets confined in net enclosures would grow more slowly but survive better than crabs at the same density free in the pond (Rodriguez et al. 2007). Even considering that it would be impossible to find absolutely all of the smallest pond individuals hidden in the mud and algal slime (Rodriguez et al. 2007), this does not seem to account for the wide differences in average weight observed (4–6 g, rather than 10–16 g). While certain physico-chemical parameters may also differ between enclosures (e.g. fouling by epiphytes) and open ponds, a ‘social’ dimension to growth should also be considered. By analogy with freshwater crayfish, perhaps individuals in an enclosure can become stuck in a social hierarchy if they continually encounter the same individuals (Hemsworth et al. 2006).

4.5.2

Feed development

The digestive enzyme profiles of S. serrata and P. pelagicus are typical of carnivores (Figueiredo & Anderson 2009). Reports of digestive cellulases in predatory S. serrata may

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seem to be a paradox. However, it may simply indicate that juvenile Scylla has a more omnivorous diet than adults (Pavasovic et al. 2004), or that the enzymes are produced by endosymbionts (Figueiredo & Anderson 2009). Whatever the origin of the cellulases, the net result is that S. serrata can readily digest a wide range of plant meals (Catacutan et al. 2003; Tuan et al. 2006; Truong et al. 2008) and these observations are being extended to S. paramamosain, with some evidence emerging of differences between the Scylla species (Truong et al. 2009). The crucial question is the extent to which fishmeal can be replaced with these plant meals in manufactured diets without compromising growth (Truong et al. 2009). Scylla serrata is reported to require a diet of 32–40% crude protein and 6–12% crude lipid, along with certain nutrients such as cholesterol (Sheen 2000; Catacutan 2002). Interestingly, the usual caution about elevated dietary lipid for crustaceans does not appear to hold for Scylla serrata, where levels as high as 13.8% lipid did not impair growth or lead to abnormal accumulation of lipid (Sheen & Wu 1999). Even the megalops stage of S. serrata tolerates relatively high dietary lipid levels (Holme et al. 2007a), an outcome raising the prospect that protein in mud crab diets can be spared using lipid (Holme et al. 2009).

4.5.3

Addressing aggression and cannibalism

Cannibalism remains a major obstacle to improving the productivity of portunid crab growout. The factors contributing to cannibalism are poorly understood except at the most superficial level. However if these factors could be addressed on-farm, then this would improve the profitability of pond production (Fig. 4.4). Research using nursery-sized animals has shown that mortality from cannibalism is reduced by lowering stocking density and adding artificial habitat, both strategies that reduce encounters between individuals (Mann et al. 2007; Ut et al. 2007a). The cannibals are the larger individuals present (Marshall et al. 2005; Møller et al. 2008) and the victims tend to be animals that have not reached post-moult (Marshall et al. 2005). Artificial habitat, which apparently works by spreading the population out and ‘reducing’ the local population density, is a strategy also relevant to farming other clawed decapods (Baird et al. 2006). A distinction should probably be drawn in future studies between ‘habitat’ and ‘shelter ’, in that ‘habitat’ refers to enhancing the general landscape within the pond, while a ‘shelter ’, like a pot, is a resource, and potentially a focus of aggression between individuals. Complicating the pond floor terrain with ‘habitat’, however, raises practical issues for pond management. Grading appears to be a promising strategy to adopt, if practical issues can be overcome (Marshall et al. 2005). Left alone, even a graded sample of crablets rapidly spreads across several instars (Mann et al. 2007). Regular harvesting and grading of crabs is out of the question. Partial harvesting is a simple ‘high-end’ approach to grading, targeting ‘shooters’ as they moult through to harvest size (Trino & Rodriguez 2001). This regular ‘fishing mortality’ has the double advantage of removing dominant individuals and steadily reducing the population density. Another passive approach from the low end could be drawn by analogy with ecological studies of crab nurseries in the wild (Hovel & Lipcius 2002; Hovel & Fonseca 2005). This is to distribute a nursery ‘habitat’ in areas of the pond; for example a bundled net with a mesh size only suited to small individuals. While this ‘nursery’ would initially protect them, the crabs would eventually have to abandon that nursery as they grow.

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Fig. 4.4

Injury and size variation are significant issues for nursery production of mud crab juveniles.

Understanding the social behaviour of crabs in ponds is probably the key to mitigating cannibalism. What we know about the role of body size, sex or moult stage (Marshall et al. 2005; Trino & Rodriguez 2001), must be weighed against the recognition that observations in clear water laboratory conditions are unusual – vision can play no role in the real night-time encounters between crabs in the depths of a turbid pond. There is as yet no evidence that distance chemoreception plays a role in targeting of moulting individuals, but this cannot be ruled out due to the sophisticated chemical awareness of crabs (Wall et al. 2009). That study, however, demonstrated that size determined a communally reared crab’s response to the odour of crushed conspecific – an outcome consistent with observations that larger individuals are cannibals. We need to establish how this behaviour is acquired and reinforced. It remains to be demonstrated whether active cannibals become larger or whether large crabs are simply opportunistic cannibals. Ordinarily, the odour of ‘blood in the water ’ or the presence of carcass in a trap is repellent to crabs (Moore & Howarth 1996; Diaz et al. 2003).

4.5.4

Containerised production

Individual rearing of crabs is an obvious solution to the mortality and injury that occurs when crabs are reared ‘socially’. Experience shows that crabs show almost no mortality when raised in a properly maintained system, albeit with a slower growth rate than the breakneck rates of growth achieved in ponds.

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Fig. 4.5 High production densities can be achieved at an experimental scale by growing crabs in containers (such as for this stable isotope feeding trial), but what are the consequences for growth rate and the economics of a scaled-up commercial system?

However, indoor containerised rearing of crustaceans is not yet common, presumably because of the extra start-up costs and the higher risks (Fig. 4.5). Containerised growth of crabs and other crustaceans adds an extra dimension to the already technically intense setup and running costs of a recirculation aquaculture system (RAS) – fish won’t climb out of the tank! Judging by accounts in the literature (Nicosia & Lavalli 1999; Barki et al. 2006), the technology for homarid lobsters and freshwater crayfish is further advanced than that for mud crabs. However, the longstanding but largely unrealised interest in containerised production of homarid lobsters is a sobering reminder that technological solutions in aquaculture are not always cost-effective (Kazmierczak & Caffey 1996). While a million mud crabs under a large pavilion sounds like a lot, biologically it is not large-scale at all for a species that produces millions of eggs per spawner. And a RAS crab farmer would potentially operate all year round, moving small cohorts (perhaps tens of thousands) through each stage of the farm. In this context, a mud crab hatchery supporting a RAS system and operating a modest genetic selection programme would find itself tipping most of its larval production down the drain. One such hatchery could supply many farms! Distributed to farms, one million seed crabs would require 200 hectares of water at 0.5 crabs/m2. But safe in 25 × 25 cm containers they take up just 6.25 hectares of water. The implied efficiency of land use is compelling – but can the numbers stack up? It is worth considering briefly what issues to consider when running an indoor crab farm as a RAS system.

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RAS economics are improved by reducing capital and operating costs and/ or increasing revenue (eg. sale price) (De Ionno et al. 2006). It is generally recommended that RAS systems concentrate upon exotic and/or high-value species because they generally cannot compete against well-established competition from pond aquaculture or wild fisheries (Funge-Smith & Phillips 2001; Zohar et al. 2005). It could be argued that pond production of mud crabs is already so well established in Southeast Asia as to push RAS mud crab production out of contention on the international stage. Certainly, any proponents would have to weigh up the market carefully before proceeding. Production of higher value ‘softshell’ crabs looks to be a better candidate for RAS production, but even this market may not prove resilient to cheaper pond-reared competition. Still, it is hard to believe that money cannot be made selling something that doubles its weight before harvest simply by drinking seawater! RAS systems ideally spread their fixed costs by operating at a large economy of scale (Dunning et al. 1998). For fish, large-scale generally means bigger tanks. For crabs, it means deciding how small the compartments can be. A corollary of having a large economy of scale is that each part of the facility must be stocked as fully as possible (Dunning et al. 1998), hence the importance of working out the minimum practical size for compartments (Nicholson et al. 2008). RAS systems also have some practical advantages for complete closure of the species’ life cycle (Zohar et al. 2005), but intensification needs to be practised all the way through the cycle. For crabs, juveniles could be grown intensively in a succession of appropriately sized nursery containers and then distributed from there into the growout containers (Nicholson et al. 2008). Even with finfish, regular stock handling comes at a cost (Dunning et al. 1998), so handling a million individual crabs would be particularly onerous. In welldesigned infrastructure, the containers could be handled robotically, but of course this adds further to the set-up and running costs of the enterprise. Operating the RAS system at maximum efficiency means there is little margin for error (Zohar et al. 2005). To begin with, the recirculation and remediation systems in place must be rated realistically to the designed output of the plant (Dunning et al. 1998). Much also hinges upon the skills and expertise of RAS farm operators and the decisions they make (Kazmierczak & Caffey 1996) – even before one takes into account the ‘newness’ of operating a containerised system. On the face of it, a large RAS crab farm requires huge numbers of individual containers, which need to be cleaned and maintained, plus a more expensive and complicated water reticulation system. While hygienically challenging, a containerised system can be automatically fed and uneaten feed recorded (crabs will change their feed intake during the moult cycle). To avoid disturbance, crabs and feed can be photographed using infrared cameras in the otherwise shadowy confines of their containers (Fig. 4.6).

4.5.5

Improving hatchery efficiency

Finding alternatives to the use of antibiotics in hatcheries is a major priority. While acceptable levels of survival to metamorphosis occur in hatcheries, bolstering these figures using antibiotics to control the bacterial flora is not acceptable. More work needs to be done, to encourage adoption of alternatives such as ozonation or use of specific probiotics (Nghia et al. 2007). Development of artificial larval diets for mud crabs, a worthy aim in its own right in terms of refining hatchery management practices, also promises to remove a major vector of microbial contamination – via the growout and enrichment of live feeds (Holme et al. 2009).

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Fig. 4.6 Containers may suit cryptic species like mud crabs – and yet individuals can be examined with minimal disturbance using infra-red LEDs and cameras to reveal the animal within.

4.6

REFERENCES

Akinbowale, O.L., Peng, H. & Barton, M.D. (2006) Antimicrobial resistance in bacteria isolated from aquaculture sources in Australia. Journal of Applied Microbiology, 100, 1103–1113. Akinbowale, O.L., Peng, H. & Barton, M.D. (2007) Diversity of tetracycline resistance genes in bacteria from aquaculture sources in Australia. Journal of Applied Microbiology, 103, 2016–2025. Alava, V.R., Quinitio, E.T., de Pedro, J.B., Orosco, Z.G.A. & Wille, M. (2007a) Reproductive performance, lipids and fatty acids of mud crab Scylla serrata (Forsskal) fed dietary lipid levels. Aquaculture Research, 38, 1442–1451. Alava, V.R., Quinitio, E.T., de Pedro, J.B., Priolo, F.M.P., Orozco, Z.G.A. & Wille, M. (2007b) Lipids and fatty acids in wild and pond-reared mud crab Scylla serrata (Forsskal) during ovarian maturation and spawning. Aquaculture Research, 38, 1468–1477. Baird, H.P., Patullo, B.W. & Macmillan, D.L. (2006) Reducing aggression between freshwater crayfish (Cherax destructor Clark: Decapoda, Parastacidae) by increasing habitat complexity. Aquaculture Research, 37, 1419–1428. Barki, A., Karplus, I., Manor, R., Parnes, S., Aflalo, E.D. & Sagi, A. (2006) Growth of redclaw crayfish (Cherax quadricarinatus) in a three-dimensional compartments system: Does a neighbour matter? Aquaculture, 252, 348–355. Catacutan, M.R. (2002) Growth and body composition of juvenile mud crab, Scylla serrata, fed different dietary protein and lipid levels and protein to energy ratios. Aquaculture, 208, 113–123. Catacutan, M.R., Eusebio, P.S. & Teshima, S. (2003) Apparent digestibility of selected feedstuffs by mud crab, Scylla serrata. Aquaculture, 216, 253–261. Chen, J.C. & Chia, P.G. (1997) Osmotic and ionic concentrations of Scylla serrata (Forskaal) subjected to different salinity levels. Comparative Biochemistry and Physiology, 117A, 239–244.

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Chiou, T.-K. & Huang, J.-P. (2003) Chemical constituents in the abdominal muscle of cultured mud crab Scylla serrata in relation to seasonal variation and maturation. Fisheries Science, 69, 597–604. Cholik, F. & Hanafi, A. (1992) A review of the status of the mud crab (Scylla sp) fishery and culture in Indonesia. In: The Mud Crab: Report of the Seminar of Mud Crab Culture and Trade (ed. C.A. Angell), pp. 13–28. Bay of Bengal Programme, Madras, India. Christensen, S.M., Macintosh, D.J. & Phuong, N.T. (2004) Pond production of the mud crabs Scylla paramamosain (Estampador) and S. olivacea (Herbst) in the Mekong Delta, Vietnam, using two different supplementary diets. Aquaculture Research, 35, 1013–1024. Dat, H.D. (1999) Description of mud crab (Scylla spp.) culture methods in Vietnam. In: Mud Crab Aquaculture and Biology (eds C.P. Keenan & A. Blackshaw), pp. 67–71. Proceedings of an international scientific forum held in Darwin, Australia, 21–24 April 1997. Australian Centre for International Agricultural Research. Davenport, J. & Wong, T.M. (1987) Responses of adult mud crabs (Scylla serrata) (Forskal) to salinity and low oxygen tension. Comparative Biochemistry & Physiology, 86A, 43–47. Davis, J.A., Wille, M., Hecht, T. & Sorgeloos, P. (2005) Optimal first feed organism for South African mud crab Scylla serrata (Forskal) larvae. Aquaculture International, 13, 187–201. De Ionno, P.N., Wines, G.L., Jones, P.L. & Collins, R.O. (2006) A bioeconomic evaluation of a commercial scale recirculating finfish growout system – An Australian perspective. Aquaculture, 259, 315–327. de Pedro, J.B., Quinitio, E.T. & Parado-Estepa, F.D. (2007) Formalin as an alternative to trifluralin as prophylaxis against fungal infection in mud crab Scylla serrata (Forsskal) larvae. Aquaculture Research, 38, 1554–1562. Diaz, H., Orihuela, B., Forward Jr, R.B. & Rittschof, D. (2003) Orientation of juvenile blue crabs Callinectes sapidus Rathbun, to currents, chemicals and visual cues. Journal of Crustacean Biology, 23, 15–22. Dunning, R.D., Losordo, T.M. & Hobbs, A.O. (1998) The Economics of Recirculating Tank Systems: A Spreadsheet for Individual Analysis, SRAC Publication No. 456. Southern Regional Aquaculture Center. FAO (2008) The State of World Fisheries and Aquaculture. Food and Agriculture Organisation of the United Nations, Rome. Figueiredo, M.S.R.B. & Anderson, A.J. (2009) Digestive enzyme spectra in crustacean decapods (Paleomonidae, Portunidae and Penaeidae) feeding in the natural habitat. Aquaculture Research, 40, 282–291. Funge-Smith, S. & Phillips, M.J. (2001) Aquaculture systems and species. In: Aquaculture in the Third Millennium (eds R.P. Subasinghe, P. Bueno, M.J. Phillips, C. Hough, S.E. McGladdery & J.R. Arthur), pp. 129–135. Technical Proceedings of the Conference on Aquaculture in the Third Millennium, 20–25 February 2000. NACA, Bangkok and FAO, Rome. Genodepa, J., Zeng, C. & Southgate, P.C. (2007) Influence of binder type on leaching rate and ingestion of microbound diets by mud crab, Scylla serrata (Forsskal), larvae. Aquaculture Research, 38, 1486–1494. Heasman, M.P. (1980) Aspects of the general biology and fishery of the mud crab Scylla serrata (Forsskal) in Moreton Bay, Queensland. PhD thesis, Department of Zoology, University of Queensland. Hemsworth, R., Villareal, W., Patullo, B.W. & Macmillan, D.L. (2006) Crustacean social behaviour changes in response to isolation. Biological Bulletin, 213, 187–195. Hill, B.J. (1976) Natural food, foregut clearance-rate and activity of the crab Scylla serrata. Marine Biology, 34, 1432–1793. Hill, B.J. (1979) Aspects of the feeding strategy of the predatory crab Scylla serrata. Marine Biology, 55, 209–214. Hill, B.J., Williams, M.J. & Dutton, P. (1982) Distribution of juvenile, subadult and adult Scylla serrata (Crustacea: Portunidae) on tidal flats in Australia. Marine Biology, 69, 117–120. Holme, M.H., Zeng, C. & Southgate, P.C. (2006a) Use of microbound diets for larval culture of the mud crab, Scylla serrata. Aquaculture, 257, 482–490. Holme, M.H., Zeng, C. & Southgate, P.C. (2006b) The effects of supplemental dietary cholesterol on growth, development and survival of mud crab, Scylla serrata, megalopa fed semi-purified diets. Aquaculture, 261, 1328–1334. Holme, M.H., Southgate, P.C. & Zeng, C. (2007a) Survival, development and growth response of mud crab, Scylla serrata, megalopae fed semi-purified diets containing various fish oil: corn oil ratios. Aquaculture, 269, 427–435.

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Holme, M.H., Southgate, P.C. & Zeng, C. (2007b) Assessment of dietary lecithin and cholesterol requirements of mud crab, Scylla serrata, megalopa using semi-purified microbound diets. Aquaculture Nutrition, 13, 413–423. Holme, M.H., Zeng, C.S. & Southgate, P.C. (2009) A review of recent progress toward development of a formulated microbound diet for mud crab, Scylla serrata, larvae and their nutritional requirements. Aquaculture, 286, 164–175. Hovel, K.A. & Lipcius, R.N. (2002) Effects of seagrass habitat fragmentation on juvenile blue crab survival and abundance. Journal of Experimental Marine Biology & Ecology, 271, 75–98. Hovel, K.A. & Fonseca, M.S. (2005) Influence of seagrass landscape structure on the juvenile blue crab habitat-survival function. Marine Ecology Progress Series, 300, 179–191. Jayasankar, V. & Subramoniam, T. (1999) Antibacterial activity of seminal plasma of the mud crab Scylla serrata (Forsskal). Journal of Experimental Marine Biology & Ecology, 236, 253–259. Jivoff, P. & Hines, A.H. (1998) Female behaviour, sexual competition and mate guarding in the blue crab, Callinectes sapidus. Animal Behaviour, 55, 589–603. Josileen, J. & Menon, N.G. (2005) Growth of the blue swimmer crab, Portunus pelagicus (Linnaeus, 1758) (Decapoda, Brachyura) in captivity. Crustaceana, 78, 1–18. Kazmierczak, R.F. & Caffey, R.H. (1996) The Bioeconomics of Recirculating Aquaculture Systems. Louisiana State University Agricultural Centre Bulletin 854. Keenan, C.P. (1999) The fourth species of Scylla. In: Mud Crab Aquaculture and Biology (eds C.P. Keenan & A. Blackshaw), pp. 4–58. Proceedings of an International Scientific Forum held in Darwin, Australia, 21–24 April 1997. ACIAR Proceedings No. 78. Australian Centre for International Agricultural Research, Canberra. Keenan, C.P., Davie, P.J.F. & Mann, D.L. (1998) A revision of the genus Scylla de Haan, 1833 (Crustacea: Decapoda: Brachyura: Portunidae) (1998). Raffles Bulletin of Zoology, 46, 217–245. Khatun, M., Kamal, D. & Yi, Y. (2009) Comparisons of growth and economic performance among monosex and mixed-sex culture of red mud crab (Scylla olivacea Herbst, 1796) in bamboo pens in the tidal flats of mangrove forests, Bangladesh. Aquaculture Research, 40, 473–485. Lavilla-Pitogo, C.R., de la Pena, L.D. & Catedral, D.D. (2007) Enhancement of white spot syndrome virus load in hatchery-reared mud crab Scylla serrata (Forsskal, 1775) juveniles at a low temperature. Aquaculture Research, 38, 1600–1603. Le Vay, L., Ut, V.N. & Jones, D.A. (2001) Seasonal abundance and recruitment in an estuarine population of mud crabs, Scylla paramamosain, in the Mekong Delta, Vietnam. Hydrobiologia, 449, 231–240. Lebata, M., Le Vay, L., Walton, M.E., et al. (2009) Evaluation of hatchery-based enhancement of the mud crab, Scylla spp., fisheries in mangroves: comparison of species and release strategies. Marine & Freshwater Research, 60, 58–69. Lumasag, G.J., Quinitio, E.T., Aguilar, R.O., Baldevarona, R.B. & Saclauso, C.A. (2007) Ontogeny of feeding apparatus and foregut of mud crab Scylla serrata Forsskal larvae. Aquaculture Research, 38, 1500–1511. Macintosh, D.J., Overton, J.L.& Thu, H.V.T. (2002) Confirmation of two common mud crab species (genus Scylla) in the mangrove ecosystem of the Mekong Delta, Vietnam. Journal of Shellfish Research, 21, 259–265. Mann, D.L., Asakawa, T., Kelly, B., Lindsay, T. & Paterson, B. (2007) Stocking density and artificial habitat influence stock structure and yield from intensive nursery systems for mud crabs Scylla serrata (Forsskal 1775). Aquaculture Research, 38, 1580–1587. Marichamy, R. & Rajapackiam, S. (2001) The aquaculture of Scylla species in India. Asian Fisheries Science, 14, 231–238. Marshall, S., Warburton, K., Paterson, B. & Mann, D. (2005) Cannibalism in juvenile blue-swimmer crabs Portunus pelagicus (Linnaeus, 1766): effects of body size, moult stage and refuge availability. Applied Animal Behaviour Science, 90, 65–82. Millamena, O.M. & Quinitio, E. (2000) The effects of diets on reproductive performance of eyestalk ablated and intact mud crab Scylla serrata. Aquaculture 181, 1–2. Møller, H., Lee, S.Y., Paterson, B. & Mann, D. (2008) Cannibalism contributes significantly to the diet of cultured sand crabs, Portunus pelagicus (L.): a dual stable isotope study. Journal of Experimental Marine Biology & Ecology, 361, 75–82. Moore, P.G. & Howarth, J. (1996) Foraging by marine scavengers: Effects of relatedness, bait damage and hunger. Journal of Sea Research, 36, 267–273.

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Neufeld, D.S. & Cameron, J.N. (1994) Mechanism of the net uptake of water in moulting blue crabs (Callinectes sapidus) acclimated to high and low salinities. Journal of Experimental Biology, 188, 11–23. Nghia, T.T., Wille, M., Binh, T.C., Thanh, H.P., Van Danh, N. & Sorgeloos, P. (2007) Improved techniques for rearing mud crab Scylla paramamosain (Estampador 1949) larvae. Aquaculture Research, 38, 1539–1553. Nicholson, S., Mann, D., Fotedar, R. & Paterson, B. (2008) The effects of holding space on growth and survival of individually reared three-spot crab (Portunus sanguinolentus). Aquacultural Engineering, 39, 30–36. Nicosia, F. & Lavalli, K. (1999) Homarid lobster hatcheries: their history and role in research, management, and aquaculture. Marine Fisheries Research, 61(2), 1–57. Nurdiani, R. & Zeng, C. (2007) Effects of temperature and salinity on the survival and development of mud crab, Scylla serrata (Forsskal), larvae. Aquaculture Research, 38, 1529–1538. Obata, Y., Imai, H., Kitakado, T., Hamasaki, K. & Kitada, S. (2006) The contribution of stocked mud crabs Scylla paramamosain to commercial catches in Japan, estimated using a genetic stock identification technique. Fisheries Research, 80, 113–121. Parado-Estepa, F.D., Quinitio, E.T. & Rodriguez, E. (2007) Seed production of Charybdis feriatus (Linnaeus). Aquaculture Research, 38, 1452–1458. Paterson, B., Mann, D., Kelly, B. & Barchiesi, M. (2007) Limb-loss in pond-reared blue swimmer crabs Portunus pelagicus (L.): effect on growth in an indoor shedding system. Aquaculture Research, 38, 1569–1579. Pavasovic, M., Richardson, N.A., Anderson, A.J., Mann, D. & Mather, P.B. (2004) Effect of pH, temperature and diet on digestive enzyme profiles in the mud crab, Scylla serrata. Aquaculture, 242, 641–654. Quinitio, E.T. (2000) Transport of Scylla serrata megalopae at various densities and durations. Aquaculture, 185, 1–2. Quinitio, E.T., Parado-Estepa, F.D., Millamena, O.M. & Rodriguez, E.M. (2001) Seed production of mud crab Scylla serrata juveniles. Asian Fisheries Science, 14, 161–174. Rabbani, A.G. & Zeng, C. (2005) Effects of tank colour on larval survival and development of mud crab Scylla serrata (Forskal). Aquaculture Research, 36, 1112–1119. Rodriguez, E.M., Parado-Estepa, F.D. & Quinitio, E.T. (2007) Extension of nursery culture of Scylla serrata (Forsskal) juveniles in net cages and ponds. Aquaculture Research, 38, 1588–1592. Ruscoe, I.M., Williams, G.R. & Shelley, C.C. (2004) Limiting the use of rotifers to the first zoeal stage in mud crab (Scylla serrata Forskal) larval rearing. Aquaculture, 231, 517–527. Seneriches-Abiera, M.L., Parado-Estepa, F. & Gonzales, G.A. (2007) Acute toxicity of nitrite to mud crab Scylla serrata (Forsskal) larvae. Aquaculture Research, 38, 1495–1499. Sheen, S.S. (2000) Dietary cholesterol requirement of juvenile mud crab Scylla serrata. Aquaculture, 189, 3–4. Sheen, S.S. & Wu, S.W. (1999) The effects of dietary lipid levels on the growth response of juvenile mud crab Scylla serrata. Aquaculture, 175, 143–153. Shelley, C. (2008) Capture-based aquaculture of mud-crabs (Scylla spp.). In: Capture-based Aquaculture. Global Overview (eds A. Lovatelli & P.F. Holthuis), pp. 255–269. FAO Fisheries Technical Paper No. 508. FAO, Rome. Trino, A.T. & Rodriguez, E.M. (2001) Mud crab fattening in ponds. Asian Fisheries Science, 14, 211–216. Trino, A.T. & Rodriguez, E.M. (2002) Pen culture of mud crab Scylla serrata in tidal flats reforested with mangrove trees. Aquaculture, 211, 125–134. Trino, A.T., Millamena, O.M. & Keenan, C. (1999) Commercial evaluation of monosex pond culture of the mud crab Scylla species at three stocking densities in the Philippines. Aquaculture, 174, 1–2. Trino, A.T., Millamena, O.M. & Keenan, C.P. (2001) Pond culture of mud crab Scylla serrata (Forskal) fed formulated diet with or without vitamin and mineral supplements. Asian Fisheries Science, 14, 191–200. Truong, P.H., Anderson, A.J., Mather, P.B., Paterson, B.D. & Richardson, N.A. (2008) Effect of selected feed meals and starches on diet digestibility in the mud crab, Scylla serrata. Aquaculture Research 39, 1778–1786. Truong, P.H., Anderson, A.J., Mather, P.B., Paterson, B.D. & Richardson, N.A. (2009) Apparent digestibility of selected feed ingredients in diets formulated for the sub-adult mud crab, Scylla paramamosain, in Vietnam. Aquaculture Research, 40, 322–328.

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5 Penaeid Prawns Ngo Van Hai, Ravi Fotedar and Nguyen Van Hao

5.1 INTRODUCTION Although prawn prices have shown a steady decrease, prawn farming has continued to be attractive to local farmers, investors and local governments because of its high profitability and the market’s demand for its products (Cao 2007). Aquaculture acts as an engine for economic growth in rural areas (Ahmed et al. 2007). Prawn aquaculture has been creating jobs and bringing sizeable incomes to farmers living in remote and coastal areas. Among 110 species of 12 genera belonging to the Family Penaeidae (Flegel 2007), Penaeus, Litopenaeus, Marsupenaeus, Metapenaeus and Fenneropenaeus are commonly cultured worldwide. Black tiger prawn, Penaeus monodon (Fig. 5.1), remains the most widely farmed species (Ahmed et al. 2007), making a major contribution to global prawn production (Deachamag et al. 2006). Its popularity is due to its rapid growth, low-cost food requirements and ability to reach a marketable size more rapidly compared to the other commonly farmed species (Kenway et al. 2006). In recent decades, although total marine fish, crustacean and mollusc production has increased continuously (FAO 2009), global production of P. monodon (Fig. 5.1) has declined due in part to farmers taking up production of white prawns, Litopenaeus vannamei, based on the availability of specific pathogen free (SPF) stocks (Wyban et al. 1993). These two species are the most important cultured prawn species in the world (Pongtippatee et al. 2007), making a significant contribution to the economic development of tropical, subtropical and temperate areas alike (Bachère 2003). Prawn production has been declining because of a reliance on wild broodstock, introduction of disease and an inability to conduct a selective breeding programme (Macbeth et al. 2007). Hatchery owners prefer wild spawners rather than pond-reared ones as wild prawns are larger, more fecund and yield more viable nauplii (Sugama et al. 2002). Captive-reared broodstock has poor reproductive performance (Macbeth et al. 2007) with unreliability of spawning, poor egg production, low hatch rate and consequent low naupliar production (Menasveta et al. 1993). However, it is anticipated that substantial benefits could arise from a selective improvement programme (Kenway et al. 2006).

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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Fig. 5.1

Black tiger prawn (Penaeus monodon).

It was falsely assumed that the practical problems would be overcome by focusing on simpler approaches to husbandry and nutrition (Benzie 1998). Research on prawns has been focusing not only on techniques and chemicals to improve disease resistance or production, but also on the domestication of some specific species. The stages of development of the species targeted for domestication need to be understood. Improvements in reproductive performance, through husbandry practices (Kian et al. 2004) and selection for reproductive performance, are essential for achieving domestication of specific species (Macbeth et al. 2007). Biological and biotechnological research needs to focus on investigating the species’ genome and physiology (Benzie 1998). The impacts of genetic and physiological factors on aquatic organisms have been mentioned more frequently in international journals in recent years. These works have opened a collaborative network to complementary research areas related to prawn physiology, genetics, pathogens and environment. This chapter highlights the achievements, challenges faced, and future perspectives for the penaeid aquaculture industry.

5.2 5.2.1

ACHIEVEMENTS Biological and environmental aspects

5.2.1.1 Integration between biological, ecological and environmental aspects Understanding of the habitat/environment requirements, reproduction ecology, nutritional requirements/trophodynamics and reseeding/stock management leads to better species selection, site evaluation and culture practices (Rothlisberg 1998). Environmental factors

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play a vital role and are significantly associated with disease outbreaks (Prayitno & Latchford 1995). Penaeid prawns exhibit various physiological responses to environmental stressors (Fingerman et al. 1996; Munro & Owens 2007). Increased susceptibility of farmed prawns to diseases occurs as a consequence of the impairment of the immune defence mechanisms resulting from physical and environmental abuse. Characteristics of aquaculture systems, including salinity and temperature, influence the physiological and immunological responses of penaeid prawns (Alabi et al. 2000). The interaction between salinity and temperature influences the growth and survival of prawns (O’Brien 1994). The critical thermal minima and lower incipient lethal temperature of Penaeus semisulcatus were not affected by salinity (Kir & Kumlu 2008). In penaeid prawns, survival is not determined solely by temperature, as salinity may exert an influence on the cold tolerance (Kumlu & Kir 2005). Variations in salinity cause a change in the immune response of Penaeus spp., which are more susceptible to white spot syndrome virus (WSSV) under salinity stress (Yu et al. 2003; Joseph & Philip 2007). Most outbreaks of luminous bacterial diseases caused by Vibrio harveyi occur during rainy seasons when salinity is influenced by rainfall (Prayitno & Latchford 1995). Although P. japonicus (Fig. 5.2) can adapt to a wide salinity range of 3–32 ppt, they are more susceptible to ammonia toxicity and spend more energy to compensate for the cost of osmo-regulation at low salinity (Li et al. 2007). Therefore, in low

Fig. 5.2

Kuruma or Japanese prawn (Penaeus japonicus).

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salinity farms one needs to be alert to ammonia nitrogen levels in the rearing water, to avoid unnecessary crop losses. Temperature affects the development of WSSV in P. japonicus (Guan et al. 2003) and L. vannamei (Vidal et al. 2002). Daily temperature fluctuation has both negative and positive effects on disease resistance and mortality of L. vannamei infected with WSSV (Rahman et al. 2007a,b). High water temperature prevented the onset of disease and reduced mortality of WSSV-inoculated L. vannamei regardless of the route of inoculation or virus titer used (Rahman et al. 2006). On the other hand, changes of temperature were reported to decrease resistance of L. vannamei (Cheng et al. 2005) and P. monodon (Wang & Chen 2006) to V. alginolyticus and Photobacterium damselae, respectively. After dissolved oxygen, ammonia-nitrogen, built up from the metabolism of feed, is a second limiting factor to an increase in production (Ebeling et al. 2006). While water quality variables, i.e. pH, salinity, chlorine, temperature, dissolved oxygen, nitrite and nitrate can be excluded as minor influences, ammonium toxicity is a causal factor for mortality and reduction of growth of P. monodon post-larvae (PL) (Nga et al. 2005). An increase in ammonia from 3 to 6 ppm when water pH increases from 7 to 8.5 caused low survivals of P. monodon larvae, but no interactive effect of pH and ammonia was detected (Noor-Hamid et al. 1994). L. vannamei was safe at nitrite-nitrogen up to 25.7 mg/L in salinity of 35 ppt (Lin & Chen 2003). Little has been published on the effects of the physical environment and mechanisms on inducement of spawning in P. monodon (Benzie 1997). Prawns benefit from a wide range of organisms within greenwater systems, including bacteria, phytoplankton, protists, rotifers and nematodes (Decamp et al. 2003; Schuur 2003). Knowledge of the community structure of suspended organic matter called floc, consisting of phytoplankton/algae and bacteria, helps to develop cost-effective prawn feed formulas (Forster & Dominy 2005). These flocs confer benefits to prawn culture such as an enhancement of animal growth from nutrients and micronutrients (Epp et al. 2002), an improvement of product quality in terms of colour, sensory properties and flavour materials (Breithaupt 2004), an improvement in water quality (Tacon et al. 2002), an acceleration of animal health and a decrease in mortality (Linan-Cabello et al. 2002; Burford et al. 2004). Bio-floc production depends on the quality of the added substrates such as C:N ratio, bioavailability and other factors (Avnimelech 2007). Flocs from prawn culture water are rich in protein, vitamins and minerals (Tacon et al. 2002). Uptake of flocs by aquatic animals depends on the animal species/size, feeding traits, floc size, floc density, and biodegradation of flocs. It is also dependent on the microbial community associated with flocs such as bacteria, protozoa and others (Avnimelech 2007). Microbial growth fundamentals are used to characterise production of volatile and total suspended solids for autotrophic and heterotrophic systems (Ebeling et al. 2006). The taxonomic markers (carotenoid fucoxanthin, lutein, zeaxanthin, peridinin and alloxanthin) and chlorophyll-a, as independent and dependent variables, respectively, are used as simple, rapid and reliable methods for determining and monitoring the microbial community change in aquatic environment (Ju et al. 2008). Diatoms are a source of food and nutrients for most aquatic animals, while chlorophytes are less valuable as these are not effectively consumed by zooplankton and prawns (Boyd 1989). Cyanobacteria and dinoflagellates are known to impart unpleasant flavours to animals and to degrade water quality by producing toxic compounds (Pearl & Tucker 1995). As toxic algal blooms cause economic losses worldwide, biological impacts of cyanoprokaryotic toxins, such as cylindrospermopsin, microcystins (Reinikainen et al. 2001;

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Fig. 5.3

Western king prawn (Penaeus latisulcatus).

Meriluoto & Codd 2005; Ramos et al. 2005; Zurawell et al. 2005) and nodularins, on aquatic animals have been reviewed (Landsberg 2002; Zimba et al. 2006). Environmental effects play important roles in determining maturation and larvae quality of penaeid prawns (Chow & Sandifer 1991; Benzie 1997). The influence of interactions between genotypes and temperature on the survival and growth of P. japonicus suggests that these factors can affect the efficiency of selection of prawn-breeding programmes (Coman et al. 2002). A feature of captive-reared penaeids is low reproductive success (Benzie 1998), not only because of genetic effects of inbreeding depression (Sbordoni et al. 1987), but also poor water quality and lack of key nutrient elements (Primavera 1985). Prawn farms need to meet the required conditions for specific species to obtain desired results. L. vannamei PL can grow well in fresh water supplemented with major iron, potassium, magnesium, sulphate to a final salinity of 0.7 ppt (Green 2008). P. monodon (Tantulo & Fotedar 2007) and P. latisulcatus (Fig. 5.3) (Prangnell & Fotedar 2005) can also grow in inland saline water dosed with potassium. The bioremediation capability using probiotics has been increasing rapidly with the demand for environmentally friendly and sustainable aquaculture (Wang & Han 2007). Probiotics can improve water quality (Thompson et al. 1999; Verschuere et al. 2000; Hai et al. 2009) as probiotics are usually Gram-positive bacteria, which convert organic matter back to CO2 more effectively than Gram-negative. These studies have explained the expansion of aquaculture areas and increased production of penaeid prawns worldwide. 5.2.1.2 Culture systems Effective exploration of available water bodies has led to rapid developments in prawn aquaculture. Consequently, there is little distinction between the various farming systems: intensive, semi-intensive, improved extensive, or extensive. In intensive cultivation systems, higher stocking densities may be used to improve final production, but may easily cause microbial problems, resulting in poor growth and mass mortalities (Skjermo & Vadstein 1999).

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The impact of crowding (50–100 P. monodon PL/L) on growth and survival is due to some excreted chemical compounds and water quality variables (Nga et al. 2005). In intensive culture, successful growth and survival of juvenile L. vannamei was achieved at densities up to 4,514 prawn/m3 (Samocha 2001; Samocha et al. 2001), juvenile brown tiger prawn Penaeus esculentus at 3,636 prawn/m3 (Crocos et al. 2003) and L. setiferus at 284.1 prawn/m3 (Williams et al. 1996), but 510 prawn/m3 resulted in poor growth in P. monodon (AQUACOP 1984). The production of juvenile P. esculentus was less costeffective at the high stocking density of 5,720–11,430 prawn/m3 due to slow growth and survival (Arnold et al. 2006b). Various applications for enhancing production designed to improve water quality, nutrient absorption and disease resistance, have been gaining increased attention worldwide. Final biomass was increased by the addition of artificial substrates, which enhanced growth and survival rates of P. monodon (Arnold et al. 2006a). The artificial substrates, AquaMat®, reduced competition for space by providing more surface area for prawns to inhabit, and reduced competition for food by accelerating colonisation of epiphytic biota as a natural food source for P. esculentus prawns (Arnold et al. 2005). Concerns about the environmental impacts and nitrogen budget of prawn farming, and the incidence of disease, have led to a progressive reduction in water exchange up to 0% (Bratvold & Browdy 2001; Tacon et al. 2002; Burford et al. 2003; Perez-Velazquez et al. 2008). In addition, the greenwater culture system has been applied widely in penaeid aquaculture. An understanding of microflora structure with dominance of filamentous fungi Penicillium and Aspergillus species and yeast Rhodotorula and Saccharomyces in greenwater culture systems for P. monodon shows that it plays a fundamental role in the control of luminous Vibrio (Leano et al. 2005). The application of new techniques to improve production has produced diversified culture systems, in which environmental and biological technologies have been explored. Fertilisers and liming materials are commonly used but can cause eutrophication. On the other hand, oxidants, disinfectants, osmo-regulators, algicides, coagulants, herbicides and probiotics are being used less frequently in aquaculture. However, these compounds or biological products degrade or precipitate quickly, are not bioaccumulative and do not cause environmental perturbations in natural waters (Boyd & Massaut 1999). Numerous studies on the role of nutrition, growth, maturation, and egg and larval quantity and quality, are easy to access from relevant international journals. Protein is a major factor in nutrition. Protein requirements vary due to biotic (species, size, sex) and abiotic factors (temperature, salinity) (Perez-Velazquez et al. 2008) and are defined as the required amount per animal biomass per day (Kureshy & Davis 2002). The availability of fishmeal as the main protein source is decreasing while demand for it is rising within the aquaculture industry because of the expansion of the industry. Consequently there is a need to find a viable alternative to fishmeal (Smith et al. 2007). Nutritionists have tried to replace fishmeal protein, at least partially, with plant proteins (Cuzon et al. 2004) as these protein sources are low in cost (Amaya et al. 2007). Replacement of fishmeal protein with plant sources (solvent-extracted soybean meal, corn gluten meal and corn fermented soluble) in prawn diets at 9, 6, 3 and 0% did not alter growth and production (Amaya et al. 2007). Other ingredients investigated include soybean meal, field peas and canola. Replacement of soybean meal by lupin meal in diets for P. monodon was also examined (Sudaryono et al. 1999b). Lupin (Lupinus angustifolius) kernel meal can partially replace the fishmeal protein in diets for P. monodon (Sudaryono et al. 1999a,c; Smith et al. 2007).

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Generally, the use of all-plant protein feed is limited because of a deficiency or imbalance of essential amino acids, reduced levels of minerals, limited levels of high unsaturated fatty acids (HUFA), the presence of anti-nutritional factors or toxins and decreased palatability (Davis et al. 2004). Plant protein can replace animal protein sources without adverse effects in prawn performance or production economics (Amaya et al. 2007). Fishmeal and marine oil sources can be removed from prawn feeds if suitable alternative sources of proteins and lipids are provided to meet the essential amino acid and fatty acid requirements of prawns (Davis et al. 2004). Moreover, the complete substitution of fishmeal and animal by-product meals with other protein sources can only be achieved when basic conditions for specific species such as adequate lipids, phosphorus and amino acids are provided (Amaya et al. 2007). Live feeds are an essential part of any marine hatchery. Studies on live food are still emerging in the larviculture industry, especially in enrichment. Live feeds lack some vital ingredients in the nutritional requirements of aquatic animals. A system designed for simple enrichment has been established for various uses such as in the comparison/evaluation of commercial and experimental enrichment products, and to monitor bacteria (Kolkovski et al. 2004). Enrichment of live feed such as Artemia, with essential amino acid, highly unsaturated fatty acid (HUFA) (Robin 1998; Czesny et al. 1999; Han et al. 2000), docosahexaenoic acid (DHA) (Bransden et al. 2005), eicosapentaenoic acid (EPA) (Harel et al. 2002), essential fatty acids (EFA) (Monroig et al. 2006a,b), free amino acid (Tonheim et al. 2000), bacterial probiotics (Gatesoupe 2002; Hai et al. unpublished), yeast and microalgae (Marques et al. 2004) or vitamin A or C (Monroig et al. 2007a) have been trialled, for use in the cultivation of aquatic animals. Enriching Artemia with commercial preparations such as Super Selco®, DHA Selco, DHA protein Selco®, DC-DHA Selco, Algamac-3050, and marine oils that are high in HUFAs, enhances their nutrition value for larval development (McEvoy et al. 1998; Payne & Rippingale 2000; Evjemo et al. 2001; Gatesoupe 2002; Woods 2003; Ritar et al. 2004; Hanaee et al. 2005; Monroig et al. 2007b; Hamre & Harboe 2008). Changes in the enrichment diets offer possibilities to cover the needs of various species, and help reduce problems related to diseases, stress resistance, malnutrition and pigmentation in numerous aquatic animal species (Sorgeloos et al. 2001). In addition, microparticulate microbound diet has characteristics that can be used effectively in the culture of carnivorous species/stages of crustaceans that are currently replicated on live Artemia nauplii (D’Abramo et al. 2006). Although most tissues can synthesise nucleotides de novo, some such as liver cells, immune cells and the cells in intestines are lacking this capacity and are dependent on preformed nucleotides (Quan 1992). Exogenous nucleotides present essential physiological and biochemical functions such as encoding and deciphering genetic information, mediating energy metabolism and cell signalling or components of coenzymes, allosteric effectors and cellular agonists (Cosgrove 1998). These nucleotides with various effects on digestion, absorption, metabolism and influences on different physiological responses (immunogenes and modulation of immunoglobulin production) show promise as dietary supplements to enhance disease resistance in aquaculture (Burrells et al. 2001; Li & Gatlin III 2006). These findings relate to age/size-related responses and appropriate doses and timing of administration (Li & Gatlin III 2006). Supplementation of diets with high levels of selenium is harmful for L. vannamei and increases toxicity of nitrite-nitrogen as an imbalance between prooxidant forces and antioxidant defences is one of the mechanisms of nitrite-nitrogen in this prawn (Wang et al. 2006a).

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To use the feed effectively, attention must be paid to feeding performances of penaeid larvae with various amount of food. Depending on the stage of development of the animals, different quality and quantities of food are essential for growth and development. A more spaced food offering stimulates L. vannamei’s search for and ingestion of feed, and feeding at three times per day assures lower labour costs (Pontes et al. 2008). In addition, proper feed manufacture and feedstuff processing have been found to improve the overall nutritional quality (Davis & Arnold 2000; Hernandez et al. 2004; Samocha et al. 2004). Extrusion has the advantage of deactivating and/or destroying some of the heat-sensitive anti-nutritional factors in plant protein sources (Carver et al. 1989).

5.2.2

Physiological aspects

Research on prawn physiology has been focused on hormones or endocrines as primary targets for further studies on domesticated programmes. For example, human growth hormone supplemented with diets has an influence on the growth of L. vannamei larvae and on their resistance to salinity stress (Laufer & Sagi 1991). In individual growth model for penaeids, stimulation of physiological processes such as ingestion, assimilation, faeces production, respiration and female reproduction, is integrated into population dynamic models in terms of biomass and density through the stages of the life cycles. These models can be applied for aquaculture and management of penaeid stocks (Franco et al. 2006). The integration by neuro-receptors between the effects of environmental influences and internal processes on reproduction is used to determine the production of hormones from key organs, maturation and reproduction (Benzie 1997). The sinus gland in the eyestalk affects a range of physiology such as moulting, glucose metabolism, pigment dispersion, various aspects of carbohydrate and protein metabolism, ion regulation, respiration and reproduction (Benzie 1998). In recent decades, reliable techniques for inducing ovarian maturation and spawning as an alternative to the conventional and destructive unilateral eyestalk ablation have been experimented with (Alfaro et al. 2004). Various alternatives to ablation have been evaluated such as environmental control, crustacean endocrinology or implantation (Vaca & Alfaro 2000). According to Wongprasert et al. (2006), serotonin (5-hydroxytryptamine, 5HT) can stimulate the release of several crustacean hormones such as hyperglycemic (Keller et al. 1985), red pigmentdispersing (Rao & Fingerman 1975), neurodepressing (Arechiga et al. 1985), moultinhibiting (Mattson & Spaziani 1985) and gonad-stimulating hormone (Kullkarni et al. 1992). The 5HT has been reported to induce ovarian maturation and spawning in crayfish (Procambarus clarkia) (Sarojini et al. 1995), L. vannamei (Vaca & Alfaro 2000) and P. monodon (Wongprasert et al. 2006). The combination of 5HT and dopamine antagonist, spiperone, stimulated ovarian maturation, spawning and the release of maturation-promoting pheromones of L. stylirostris and L. vannamei (Alfaro et al. 2004). The 5HT-injected P. monodon had ovarian maturation and spawning rate at a level comparable to that of unilateral eyestalk-ablated ones (Wongprasert et al. 2006). The combined injection of 5HT and spiperone not only induces maturation and spawning of L. stylirostris and L. vannamei, but also stimulates the release of maturation-promoting pheromones into the water (Alfaro et al. 2004). The hatching rate and the amount of nauplii produced per 5HT-injected spawner were higher than those per eyestalk-ablated one (Wongprasert et al. 2006). In addition, ovarian development and oocyte maturation in crustaceans are regulated by steroid hormones similar to teleost fish and amphibia (Fairs et al. 1990). The mandibular

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organ of crustaceans synthesises and secretes steroids and terpenoids, which stimulate oocyte growth (Tsukimura & Kamemoto 1991). Terpenoid hormone, methylfarnesoate, stimulates and enhances ovarian maturation of crayfish, and acts as a gonadatropin in crustaceans (Laufer et al. 1998). The isolation and characteristics of key molecules involved in the control of reproduction and the formation of yolk in penaeids have progressed (Benzie 1997). Vitellin and vitellogenin synthesis in the ovary or in the hepathopancreas of penaeids, and its transport from the hepathopancreas to the ovary via the haemolymph, have been studied (Browdy et al. 1990; Fainzilber et al. 1992; Shariff et al. 1992) in order to further investigate the isolates and characterise penaeid hormones, vitellin and vitellogenin.

5.2.3

Diseases

Major losses in penaeid aquaculture come about because of problems with water quality control and vibrios. Vibrios are opportunistic bacteria and are normally present in the culture facilities as well as in the intestinal tracts and live feed. These bacteria usually cause diseases under sub-optimal culture conditions (Decamp et al. 2008). As aquatic environment contains diverse microbial communities of pathogenic, innocuous and beneficial bacteria, the key is to maintain the balance between pathogenic and potentially beneficial bacterial strains in the culture environment (Schulze et al. 2006). Disease problems have devastated the aquaculture industry in the past and continue to threaten it; thus, the prevention and control of diseases is the priority for prawn aquaculture (Bachère 2000). Tools need to be developed for the rapid diagnosis and control of pathogens (Benzie 1998). Diagnosis of disease and prawn viruses has been conducted using histopathology, polymerasase chain reaction (PCR), electron microscopy and serology methods (Munro & Owens 2007). Invertebrates lack immunoglobulins (Igs), T-cell receptors and major histocompatibility complex (Mhc) high diversity molecules (Arala-Chaves & Sequeira 2000), which are an adaptive secondary memory immune response in vertebrates (Klein 1989). But some adaptive immunity does exist in invertebrates, particularly in crustaceans, and this is the basis for the evolution of research on immunology and the development of vaccination strategies (Arala-Chaves & Sequeira, 2000). Invertebrates have also developed theimmune memory of adhesion molecules such as catherins, Ig-like proteins, integrins, collagens and laminins (Johansson 1999; Arala-Chaves & Sequeira 2000). Although invertebrates lack a true adaptive immune response, several empirical prawn microbial vaccinations have been attempted. P. japonicus exposed to WSSV became resistant to subsequent challenge with the virus (Sánchez-Martínez et al. 2007) due to a substantial reduction of the virus from a humoral neutralising factor in the prawn immune system (Wu & Muroga 2004). P. monodon fed ß-1,3-glucan (Kenkyu 1994) and killed vibrios (Teunissen et al. 1998) had enhanced resistance to vibrios. Therefore, these results pave the way for producing vaccines against WSSV. Crustacean haemocytes with three main cell types of hyaline, semigranular and granular (Hose et al. 1990), play an important role in phagocytosis, melanisation, cytotoxicity and cell–cell communication. Haemocytes are responsible for clotting, exoskeleton hardening and elimination of foreign materials (Song & Hsieh 1994). Semigranular and granular cells store and release prophenoloxidase (proPO) (Johansson et al. 2000). Phenoloxidase, one of the most important enzymes involved in the innate immune system of invertebrates (Cerenius & Söderhäll 2004), is synthesised as proPO, and is activated by the serine

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proteinase cascade, which is triggered by microbial carbohydrates such as ß-1,3-glucan (Vargas-Albores & Yepiz-Plascencia 2000) and lipopolysacchride (Lee et al. 2000). Several proteins, viz. peroxinectin, transglutaminase and clotting protein, are components of or associated with the activation system and are involved in cell adhesion, de-granulation, encapsulation enhancement, phagocytosis and cytotoxic reactions (Cerenius & Söderhäll 2004). In invertebrates, proPO mRNA expression is only present in haemocytes, and studies on proPO activity have led to a thorough understanding of their roles in the immune systems of prawns (Wang et al. 2006b). Activation of the proPO system was independent of bacterial virulence (Alabi et al. 2000). Moulting, development, reproductive status, nutrition conditions and diseases influence haemocyte abundance (Le Moullac et al. 1997; Cheng & Chen 2001). Decreases in number of haemocytes correlate well with the reduction of resistance to pathogens (Le Moullac et al. 1998). Total haemocyte count (THC) is a useful indicator of prawn health (Sarathi et al. 2007) as the probiotic-fed prawns were shown to be healthy with an increase in THC (Hai et al. 2009). Prolongation of clotting is a consequence of haemocyte disappearances and high levels of WSSV protein in haemolymph of WSSV-infected F. indicus (Sarathi et al. 2007). A study on in vivo and in vitro effects of bacteria, extracellular products (ECP) and cysteine protease of P. monodon coagulogen/haemostasis led to the investigation of pathophysiological changes caused by V. harveyi injection, particularly a relationship among haemolymph coagulation, diseases and bacteremia/sepsis (Lee et al. 1999). Lysozyme is one of the earliest known antibacterial peptides secreted by prawns, is ubiquitous in many eukaryotes and prokaryotes, and has been recognised as the molecule involved in non-specific innate immune systems (de-la-Re-Vega et al. 2006; Tyagi et al. 2007). In invertebrates, lysozyme is an expression that refers to the response to bacterial challenges (Somboonwiwat et al. 2006; Burge et al. 2007). There are six types of lysozyme such as chicken, goose, plant, bacterial, T4 phage and invertebrate types (Hikima et al. 2003). Chicken types are active against both Gram-positive and Gram-negative bacteria (de-la-Re-Vega et al. 2006; Tyagi et al. 2007). P. japonicus lysozyme shows lytic activity to several infectious pathogens belonging to Vibrio spp. (Hikima et al. 2003). The chickentype lysozyme from L. vannamei showed activity against V. alginolyticus, V. parahaemolyticus and V. cholerae (de-la-Re-Vega et al. 2006). Lysozyme activity was not detected in the plasma of vaccinated P. vannamei (Alabi et al. 2000). Live feeds used for larviculture operations are a source of microbial contamination. For example, Artemia nauplii are heavily contaminated with bacteria, mostly Vibrio spp., which are potentially pathogenic and may cause stress in prawn PL up to the point where they become more susceptible to viral infections (Lavens & Sorgeloos 2000). Brachionus and Artemia serve as vectors of WSSV to inject prawns (Sahul Hameed et al. 2002; Zhang et al. 2006). Rotifers serve as vectors in WSSV transmission to prawns when ingested (Yan et al. 2004; Zhang et al. 2006; Yan et al. 2007). The WSSV has a wide host range including copepods, prawns, crayfish, crab, lobster, freshwater crab and prawn (Wang et al. 1998; Chen et al. 2000; Sahul Hameed et al. 2001, 2002, 2003a), while rotifers are used as live feed for larvae of these organisms (Zhang et al. 2006). It is noted that WSSV cannot be detected in prawns before PL10 (Yoganandhan et al. 2003). There are approximately 18 viruses that are reported in penaeid prawns (Munro & Owens 2007). These viruses may interact with each other once they have infected prawns. There are no adequate treatments available against the most dangerous virus for penaeid prawn aquaculture, WSSV (Witteveldt et al. 2004b; Rahman et al. 2006), which causes up to 100% mortality within 7–10 days in commercial prawn farms (Lightner 1996). Once

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WSSV is introduced into the culture system, it spreads rapidly and is uncontrollable (Yi et al. 2003). As WSSV and yellow head-like virus are well known to be transmitted vertically and horizontally in penaeid prawns and other cultured and wild crustacean species and aquatic organisms, the control strategies have focused on the exclusion of viruses from culture system by such means as chemotherapy, vaccines and other methods related to environmental aspects (Munro & Owens 2007; Sánchez-Martínez et al. 2007). The yellow head-like virus disease resistance is ‘active accommodation’ (a tolerance mechanism) involving the binding of viral antigens to cellular receptors during the early life stages of prawns (Flegel & Pasharawipas 1998). Viral diseases are often accompanied by bacterial infections (Tyagi et al. 2007). It is advantageous to determine the likelihood of the occurrence of disease and the mixture of disease expression in the dual infection of monodon baculovirus (MBV), infectious hypodermal hematopoietic necrosis virus (IHHNV) and Mourilyan Virus (MoV) in the same prawns (Munro & Owens 2007). The process of disease causation is not fully understood, as not all infected prawns with a specific virus exhibit disease symptoms or suffer mortalities (Munro & Owens 2007). Vibriosis is the most prevalent bacterial disease, which causes mass mortalities in both hatcheries and growout ponds (Fig. 5.4a,b) in penaeid prawn aquaculture (Saulnier et al. 2000). The common vibrios encountered in prawns are V. harveyi, V. parahaemolyticus, V. alginolyticus and V. anguilaram (Tyagi et al. 2007). A method for detection of V. harveyi contamination or infection could facilitate disease prevention in the prawn aquaculture industry due to the close phylogenetic relationship of this species to other Vibrio species, i.e. V. parahaemolyticus, V. alginolyticus, V. campbellii and V. carchariae (Kita-Tsukamoto et al. 1993; Pedersen et al. 1998). V. harveyi is one of the most disastrous pathogens, and causes high mortality in the prawn culture industry worldwide (Liu et al. 1996). V. harveyi was isolated as dominant luminous species with 94.05% in all penaeid prawn hatchery components except eggs and UV-treated water, and 97.3% in prawn intestines (Abraham & Palaniappan 2004). Broodstock maturation and spawning facilities can be the main sources of V. harveyi (Chrisolite et al. 2008). Effects of V. harveyi and V. splendidus on P. monodon larvae related to the age of prawns (Baticados et al. 1990; Prayitno & Latchford 1995). It confirms that older prawns develop an increasing resistance to those pathogens (Prayitno & Latchford 1995). Besides V. harveyi, other species such as V. splendidus (Lavilla-Pitogo et al. 1990), P. phosphoreum (Prayitno & Latchford 1995), P. damselae, V. parahaemolyticus, V. alginolyticus, V. fluvialis and V. vulnificus (Lightner 1993; Mohney et al. 1994) are luminous bacterial species. The application of antibiotics is expensive and detrimental and can result in prevalence of drug residues in reared animals (Moriarty 1999). The use of antibiotics should be rejected not only for prophylactic but also for therapeutical treatment (Lavens & Sorgeloos 2000). The application of immunostimulants in prawn culture alters the use of antibiotics and chemotherapeutics (Chythanya et al. 2002; Chotigeat et al. 2004; Supamattaya et al. 2005; Sánchez-Martínez et al. 2007; Decamp et al. 2008). Immunostimulants produce biochemical compounds such as bacteriocins, lysozymes, proteases and hydroperoxide (Balcázar et al. 2007) that activate the immune systems of animals and render them more resistant to infections by viruses, bacteria, fungi and parasites (Raa 1996). These substances also inhibit the bacterial pathogens in aquaculture systems (Rengpipat et al. 1998; Gram et al. 1999). These effects mean they are a better and more effective alternative than administering antibiotics to manage prawn health (Moriarty 1997; Verschuere et al. 2000. Comparison was made between chloramphenicol and Arthrobacter XE-7, regarded as probiotics, on protection of P. chinensis larvae from pathogenic vibrios (Li et al. 2006).

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(a)

(b) Fig. 5.4

A typical semi-intensive prawn farm with (a) aerator and (b) feeding tray.

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5.2.4

Immunological aspects

Although antibiotics and diverse types of feed additives are known to be used for the control of bacteriosis, reduction of stress and improvement in the health of cultured prawns, the regular or indiscriminate use of antibiotics or chemotherapeutic agents has led to problems of drug resistance (Karunasagar et al. 1994; Weston 1996; Esiobu et al. 2002; Balcázar et al. 2006b). The use of preventive and environment-friendly approaches such as antibacterial peptides, probiotics and prebiotics is becoming increasingly important in aquaculture (Sakai 1999; Bachère 2003; Vine et al. 2006; Soltanian et al. 2007). Immunostimulants including probiotics and prebiotics have become important for heightening the activity of non-specific defence mechanisms and conferring protection against diseases (Jeney & Anderson 1993). Immunostimulants are obtained from various sources such as bacteria, brown and red algae and terrestrial fungi (Bricknell & Dalmo 2005), bacteria from aquatic habitats (Rengpipat et al. 1998) and marine yeast (Sajeevan et al. 2006). Immunostimulants can be divided into several groups, depending on their original sources such as bacteria, algaederived, animal-derived, nutritional factors and hormones or cytokines (Sakai 1999). Probiotics are defined as ‘live microorganisms which when administered in adequate amounts confer a health benefit to the host’ (FAO/WHO 2002) and prebiotics are defined as ‘nondigestible food ingredients that beneficially affect the growth and health of the host’ (Gibson & Roberfroid 1995). According to Kesarcodi-Watson et al. (2008), certain suggested immunostimulants (Itami et al. 1998; Smith et al. 2003) such as peptidoglycan (PG) and lipopolysaccharides can be considered as probiotics. In addition, a number of chemical agents, polysaccharides, plant extracts or some nutritional additives, act as immunostimulants (Sakai 1999; Gannam & Schrock 2001), are adjuncts to vaccination and provide a potential route to reduction of the widespread use of antibiotics (Burrells et al. 2001). Herbal immunostimulants, namely methnolics extracted from five different herbal medicinal plants, were shown to increase P. monodon resistance against viral pathogenesis caused by WSSV (Citarasu et al. 2006). Probiotics isolated from aquatic environments include vibrionaceae, pseudomonads, lactic acid bacteria, Bacillus spp. and yeast (Gatesoupe 1999). Some probiotic bacteria and vibrios are the most common genera associated with crustaceans (Moriarty 1997) and common inhabitants of aquatic environments including prawn culture ponds (Otta et al. 1999a). For example, live Bacillus sp. and V. alginolyticus are probiotic (Gullian et al. 2004). Probiotic Pseudomonas spp. in the marine environment produce a wide range of secondary metabolites (Raaijmakers et al. 1997; Vijayan et al. 2006). Pseudomonas PS-102 isolated from brackish water are used as probiotics for management and control of bacterial infections in P. monodon (Vijayan et al. 2006). Four probiotic strains isolated from the gastrointestinal tract of L. vannamei showed antagonism towards prawn-pathogenic bacterium, V. parahaemolyticus (Balcázar et al. 2007). Probiotics should be tested for their inhibitory activity on different species of Vibrio and Aeromonas (Vijayan et al. 2006; Hai et al. 2007) that have been considered as major pathogens in aquaculture system (Singermann 1990). Currently, four methods: double layer, well-diffusion, cross-streak and disc-diffusion, are commonly employed to screen for inhibitory substances in vitro (Kesarcodi-Watson et al. 2008). For instance, Hai et al. (2007) have used bacteriocin-like inhibitory substance (BLIS), modified BLIS, disc-diffusion, well-diffusion and co-culture methods to select the most suitable probiotics for use in the cultivation of P. latisulcatus. Probiotics have to be evaluated for safety to the hosts

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(Verschuere et al. 2000). Initial in vitro screening is followed by small-scale tests either for pathogenicity to the host (Chythanya et al. 2002) or host protection when challenged with pathogens (Rengpipat et al. 1998; Gram et al. 2001; Irianto & Austin 2002b; Vaseeharan et al. 2004). Probiotics Pseudomonas spp. at 107 CFU/mL did not cause any harmful effects to P. monodon larvae (Vijayan et al. 2006) and P. latisulcatus juvenile (Hai et al. 2009). In agreement with these studies, immersion challenges indicated effectiveness at reducing disease caused by V. parahaemolyticus in probiotic-fed L vannamei (Balcázar et al. 2007). There are two major limitations to the screening approach. Firstly, other modes of probiotic activities do not display on agar plates and secondly, positive results in vitro fail to determine the real in vivo effect (Kesarcodi-Watson et al. 2008). Certain properties that aid in the correct establishment of new, effective and safe probiotic products include i) no harm to the host; ii) accepted by the host through ingestion and colonisation and proliferation within the host; iii) reach target organs where they can have an impact; iv) work in both in vivo and in vitro conditions; v) do not contain virulence resistance genes or antibacterial resistance genes (Verschuere et al. 2000; KesarcodiWatson et al. 2008). Probiotics can be applied directly into the rearing water or supplemented with feed or live feeds (Skjermo & Vadstein 1999; Hai et al. 2009; Hai & Fotedar, 2009). Appropriate probiotic density is at 105 CFU/mL (Moriarty 1998; Hai et al. 2009). However, overdosages or prolonged administrations of probiotics can induce immunosuppression of continuous responses (Sakai 1999). Probiotics can be used singly or a combination (Gatesoupe 2002; Kesarcodi-Watson et al. 2008) or even mixture between probiotics and prebiotics. Probiotics based on a single strain are less effective than those based on mixed strains (Hai et al. 2009; Hai & Fotedar 2009). Multistrain and multispecies probiotics have positively provided synergistic bacteria with complementary modes of action to enhance protection (Timmermans et al. 2004). It has been argued that in aquaculture the microbial habitat changes continuously. Therefore, it is unlikely that a single bacterial species will be able to remain dominant in a continuously changing environment. A beneficial bacterium will probably dominate the associated microbiota when several bacteria are administered rather than when only one probiotic strain is involved (Verschuere et al. 2000). Current probiotic applications and scientific data on mechanisms of action indicate that the non-viable microbial components act in a beneficial manner and this benefit is not limited just to the intestinal tracts (Salmien et al. 1999). The application of probiotics improves the survival, growth rates and feed conversion ratio (FCR) of prawns (Balcázar et al. 2007; Hai et al. 2009). Bacterial strain PM-4 promoted the growth of P. monodon nauplii (Maeda & Liao, 1992). Photosynthetic bacteria and Bacillus spp. improved the growth performance of L. vannamei with high levels of lipase and cellulase activity (Wang 2007). F. indicus grew better due to increases in specific activities of amylase, total protease and lipase when they were exposed to probiotic Bacillus (Ziaei-Nejad et al. 2006). The survival and wet weight of L. vannamei-fed probiotics were higher than those receiving oxytetracycline and the control (Garriques & Arevalo 1995). Probiotics participate in the digestive processes by producing extracellular enzymes such as proteases, carbohydrolases and lipases and providing growth factors (Arlleano & Olmos 2002; Ochoa & Olmos 2006). Probiotics reduced the FCR of L. vannamei (Balcázar et al. 2007). PG-fed P. japonicus showed higher survival than the control (Itami et al. 1998). Inhibitory compounds excreted by probiotics, which produce a hostile environment for pathogens, include bacteriocines, lysozymes, proteases and hydroperoxide (Balcázar et al.

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2007). Probiotics can produce extracellular enzymes such as proteases, carbohydrolases and lipases, and provide growth factors (Arellano & Olmos 2002; Ochoa & Olmos 2006). Competition for adhesion receptors with pathogens may be the first probiotic effect (Montes & Pugh 1993). Microbial populations release chemical substances, which have bactericidal and bacteriostatic effects on other microbial populations. Probiotics promote the defence of the gut flora against pathogens (Skjermo & Vadstein 1999). Therefore, it can alter interpopulation relationships by influencing the outcome of competition for chemicals or available energy (Pybus et al. 1994). Prebiotics, namely ß-glucan and mannan oligosaccharides (Bio-Mos®), enhance the immune responses of prawns (Chang et al. 2003; Fritts & Waldroup 2003; Hai & Fotedar 2009). According to Reid (2008), prebiotics is seen as a means of influencing the gut microbiota and risk of allergy, and probiotics influence newborn health and the health function of the liver and pancreas of organisms. Generally, regular consumption of probiotics and prebiotics enhances immune function, improves colonic integrity, decreases the incidence and duration of intestinal injections, down-regulates allergic response and improves digestion (Douglas & Sanders 2008). ß-glucan enhanced the non-specific immune system to resist bacterial injections (Burrells et al. 2001), improved the survival of WSSV-infected P. monodon (Chang et al. 2003) and increased the survival of brooder P. monodon reared in both indoor and outdoor conditions (Chang et al. 2000), but an increase in dosages of ß-1,3-glucan in diets for L. vannamei from 0.02 to 0.1% did not affect the expressions of the genes related to immune proteins in haemocytes and hepatopancreas (Wang et al. 2008). Supplement of ß-1,3-glucan with feed enhanced haemocyte phagocytic activity, cell adhesion, superoxide anion production in P. monodon (Song & Hsieh 1994; Chang et al. 2000) and synthesis of cells and ProPO in L. vannamei (López et al. 2003). P. japonicus treated with peptidoglycan enhanced the phagocytic activity of granulocytes and increased prawn resistance to V. penaeicida (Itami et al. 1998). ß-1,3-glucan increases the resistance against vibriosis in P. japonicus (Itami et al. 1998) and against vibriosis and WSSV in P. monodon (Sung et al. 1998). ß-glucan and Vibrio bacterins do not change the haemolymph protein profile of P. monodon, but haemagglutination activity partly accounts for the immunomodulatory activity of these immunostimulants (Pais et al. 2008). The brooder P. monodon proved healthier with their haemocyte phagocytic activity, cell adhesion and superoxide anion production when they were fed ß-1,3-glucan at 0.02% diets (Chang et al. 2000). L vannamei fed ß-1,3-glucan or vitamin C showed a significantly greater growth rate than those without any immunostimulants (López et al. 2003). Dead bacteria and probiotics are claimed to induce and build up protection against a wide range of diseases in invertebrates (Keith et al. 1992; Alabi et al. 1999; Vici et al. 2000; Marques et al. 2006a). The use of immunostimulants is a new preventative approach or strategy for microbial management to reduce mortalities caused by infectious diseases (Skjermo & Bergh 2004; Bricknell & Dalmo 2005), and to maintain the good health of cultured organisms (Marques et al. 2006b). Immunostimulants help prawns overcome stress conditions, including handling, grading, vaccination, net changing, salt water transfer, anti-parasite bath treatments (Burrells et al. 2001; Bricknell & Dalmo 2005), and unilateral eyestalk ablation (Maggioni et al. 2004). Immunostimulants stimulate various components of the cellular and humoral immune systems, and act as adjuvants to increase vaccine effectiveness (Sakai 1999). Bacillus S11 provided disease protection by activating cellular and humoral immune defences (Rengpipat et al. 2000). Immunostimulants have an influence on the non-specific immune elements such as phagocytic cell activity, natural killer cell activity, lysozyme levels, complement levels and total immunoglobulin levels, in which immunostimulants

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mainly facilitate the function of phagocytic cells and increase their bactericidal activities (Sakai 1999). The modes of action of immunostimulants are i) to improve water quality, ii) to produce inhibitory compounds or antibiotics, iii) to compete for chemicals or energy/nutrient and for adhesion sites, iv) to enhance the immune responses, v) to interact with phytoplankton, vi) to supply sources of macro- and micronutrients and enzymatic contribution to digestion, vii) to modulate interactions with the environment and viii) to develop beneficial immune responses (Gatesoupe 1999; Verschuere et al. 2000; Irianto & Austin 2002a; Abidi 2003; Balcázar et al. 2006b). To use immunostimulants effectively, the timing, dosages, methods of administration and physiological conditions need to be considered properly (Sakai 1999). Immunostimulants should be administered into the culture systems before disease outbreaks occur, to reduce disease-related losses (Anderson 1992). Short-duration feeding of immunostimulants, followed by a period of control diet feeding, produces optimal effects on immune response and disease resistance (Bagni et al. 2000; Couso et al. 2003; Bridle et al. 2005). There are various ways to introduce immunostimulants such as supplement with feed, immersion, submersion, oral administration and intramuscular or intra-arterial/intraperitoneal injections, of which the supplement with feed or in-feed route is considered more productive and practical than others (Azad et al. 2005; Hai et al. 2009). In P. japonicus, feeding 0.2 mg peptidoglycan (PG)/kg BW/day was enough to increase the phagocytic activity of granulocytes (Itami et al. 1998). In contrast, oral administration is considered the most practical method for prawn immunostimulants (Huang et al. 2006). Oral administration has advantages for prawns regardless of their size (Sakai 1999), as prawns can be treated at any stage of culture period. Oral administration of ß-1,3-glucan (Itami et al. 1994) or PG (Itami et al. 1998) to prawns enhanced disease resistance. Specific immune responses and protection can be induced in P. monodon by oral vaccination (Witteveldt et al. 2004b). The immersion method is useful although it is limited to the juvenile stage or to the stage before releasing to growout ponds (Itami et al. 1998). P. monodon (Sung et al. 1994) and P. latisulcatus (Hai et al. 2009) immersed in a suspension of ß-glucan or probiotics showed higher disease resistance than those in the control, respectively. P. monodon grew faster with glucan immersion at 0.5, 1 and 2 mg/mL than at 0.25 mg/mL (Sung et al. 1994). In P. japonicus, injected or immersed with the formalin-killed Vibrio bacterin, mortalities were reduced when challenged with Vibrio injection (Itami et al. 1989). Intramuscular immunisation of the WSSV enveloped protein VP19 and VP28 increased the P. monodon survival, and this result offers new strategies to control viral diseases in prawns and other crustaceans (Witteveldt et al. 2004a). An intramuscular injection of inactivated WSSV vaccines, followed by an intramuscular challenge with WSSV resulted in mixed protection of P. japonicus, and recombinant rVP28 induced resistance, while heatinactivated WSSV did not induce resistance in the prawns. These results suggested that it is possible to vaccinate prawns with recombinant proteins against WSSV (Namikoshi et al. 2004).

5.2.5

Genetic aspect

Various molecular techniques have been developed as the basic tools to improve both quantity and quality of prawn production. For instance, the development of cell lines for prawns and the development of molecular probes provide a basis for description of the

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prawn genome, for understanding disease resistance, for developing control of reproduction and for developing specific disease-free or disease-resistant domesticated lines (Benzie 1998). Light microscopy (LM), transmission electron microscopy (TEM) and scanning electron microscopy (SEM) have been used to examine the morphological events of P. monodon acrosome reaction during an in vitro incubation and during spawning (Pongtippatee et al. 2007). Other techniques have also been established and widely applied such as polymerase chain reaction (PCR) and molecular markers. For example, nine immune-related genes in L. vannamei were detected using conventional reverse transcription polymerase chain reaction (RT)-PCR, quantitative RT-PCR and in situ hybridisation. These provide knowledge on physiological repression to activate and modulate the immune response, in L. vannamei, and guidelines for analysing gene expression for future immunological studies (Wang et al. 2007). Research on genetic aspects contributes to the development of appropriate technologies to achieve reliable production of penaeid prawns. Genetic factors influence larval quality of P. monodon (Benzie et al. 1997), in which 10% control of growth rate of post-larvae (PL) by genetic factors is substantial enough to expect a good response to selective breeding (Benzie 1997). Massive selection in prawns has improved production performances, but is quickly counterbalanced by a rapid growth in the inbreeding coefficient, which affects survival, reproduction and growth rate (de Lima et al. 2008). Loss of genetic variability caused by inadequate breeding strategies, for instance, in P. stylirostris (Bierne et al. 2000), P. monodon (Xu et al. 2001) and L. vannamei (Garcia et al. 1994; Wolfus et al. 1997) has been intensified by improper genetic management of cultured populations, resulting in inbreeding depression and negative impact on production (De Donato et al. 2005). This means that the loss of genetic diversity is detrimental to populations and affects their sustainability (Dunham 2004). Genetic improvement may help to solve the problems of diseases and small size for markets (Fu et al. 2004). Some antiviral genes from P. monodon can aid in prawn viral disease control (Luo et al. 2003). The peptides derived from these genes can bind to WSSV and block virus infection, and act as a potential for an antiviral peptide drug (Yi et al. 2003). Specific silencing RNAs of WSSV genes can suppress WSSV efficiently in L. vannamei and provide a potential approach to the therapy of prawn viral diseases (Wu et al. 2007). Successful genetic improvement of Australian strains of P. japonicus has prompted the prevention of unauthorised breeding from the elite genotypes (Coman et al. 2008). Molecular markers can be used for solving the problem of unauthorised reproduction of prawn strains (Valerio-García & Grijalva-Chon 2008). The populations of species were clustered into geographical groups, consistent with regions that have been in separate sea basins, isolated sea arms or at past low sea level stands. Some species showed no variation over thousands of kilometres, while others revealed differentiation over hundreds of kilometres (Sugama et al. 2002). Popular genetic markers used in aquaculture include allozymes, mitochondrial DNA, restriction fragment length polymorphisms (RFLP), random-amplified polymorphic DNA (RAPD), amplified fragment length polymorphism (AFLP), microsatellites, single nucleotide polymorphism (SNT) and expressed sequence tag (EST). Penaeid prawns show little allozyme polymorphism (Tam & Chu 1993). Allozyme data showed that P. monodon populations from Western Australia differed in allozyme frequencies from populations on the northern and eastern coasts (Benzie et al. 1992). The principles, potential, requirements, advantages and disadvantages of the applications of these genetic markers have been discussed by Liu and Cordes (2004). RAPD provides

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the markers for prawn breeding programmes as well as for other species (Garcia & Benzie 1995). AFLP markers have been developed as an alternative to microsatellites for linkage mapping studies, and are robust with some polymorphisms conserved across families (Moore et al. 1999). DNA-base markers show higher levels of variation and have greater sensitivity in detecting genetic variability in inbred cultured stocks than allozyme variants (Benzie, 1998). To date, many types of molecular markers have been used to analyse the genetic parameters of wild and farmed populations, in which DNA microsatellites are becoming a standard for highly desirable characteristics of reproducibility, polymorphism and codominance (Schlötterer 2000). The development of molecular markers such as microsatellites needs to be pursued to provide the tools for marker-assisted selection and to further investigate wild stock structures (Benzie 1997). Microsatellites are polymorphic and can be used as genetic markers (Divu et al. 2008). Microsatellite markers have been isolated from L. vannamei (Garcia et al. 1996), P. monodon (Xu et al. 2001), P. setiferus (Ball et al. 1998) and P. japonicus (Moore et al. 1999) for genetic studies. Microsatellite markers are used to investigate the diversity levels in populations for implementing breeding programmes (Wolfus et al. 1997). The microsatellite technique opens a new perspective for study of the structure of closely related populations (Estoup et al. 1998). As the development of polymorphic genetic markers is an essential tool for identifying parentage relationships, providing pedigree information in aquaculture selection programmes (O’Reilly & Wright 1995; Ferguson & Danzmann 1998), the application of DNA markers allows for the investigation of the parentage assignments, genetic variability and inbreeding, species/strain identification and the construction of high-resolution linkage maps for aquaculture species (Liu & Cordes 2004). The application of microsatellite markers reveals considerable progress in verifying pedigree in lines and populations. Microsatellite markers are also used for tracking pedigrees in breeding programmes and marker-assisted selection (Moore et al. 1999), and also provide such data for stock management, selective breeding programmes and sustainable use of wild resources (Liu & Cordes 2004). Polyploid penaeids have been successfully produced using chemicals and temperature shock (Xiang et al. 1992; AQUACOP et al. 1993). Studies on triploid induction techniques have been developed for penaeid prawn species such as P. chinensis (Zhang et al. 2003, 2004; Xiang et al. 2006), L. vannamei (Dumas & Ramos 1999), P. japonicus (Norris et al. 2005; Sellars et al. 2006a; Coman et al. 2008). The difference in growth performances including the specific growth rate (SGR), feeding rate, feed conversion efficacy (FCE) and intermoult period (IP) between triploid and diploid of Fenneropenaeus chinensis are related to salinity concentrations of 20 and 20–30 g/L, respectively (Zhang et al. 2008). Although cryopreservation of spermatozoa has been developed as a tool for the improvement of broodstock management and for genetic selection, this technique as applied to aquatic stocks has not been widely implemented (Christensen & Tiersch 1997). Cryopreservation of spermatophores from crustaceans has received less attention (Gwo 2000), but cryopreservation of sperm has been achieved in penaeids (Clark & Griffin 1993) such as in P. monodon (Bart et al. 2006; Vuthiphandchai et al. 2007) and in L. vannamei (Lezcano et al. 2004). This technique is useful in breeding programmes of specific domesticated species such as P. monodon in Australia and Thailand (Withayachumnarnkul et al. 2001; Coman et al. 2006) and is necessary to ensure that sperm is available for breeding programmes. Cryopreserved P. monodon spermatophores offer an alternative and reliable approach for access to male gametes for use in the artificial insemination of female

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broodstock in breeding programmes (Bart et al. 2006; Vuthiphandchai et al. 2007). Cryopreservation of P. monodon spermatophores resulted in a decline in the numbers of bacteria, especially eliminated pathogenic bacteria (Vibrio spp. and P. aeruginosa), during long-term storage of 90–120 days in liquid nitrogen (Nimrat et al. 2008). The genetic basis for sex determination has not been studied in decapods, while experimental sex reversal has been used to identify the genetic sex determination mechanism of a number of crustacean species (Legrand et al. 1987). The growth-related genes in the southwestern Atlantic pink shrimp Farfantepenaeus paulensis have been detected under DDRT-PCR protocol (Kamimura et al. 2008). The result opens an initial approach towards exploration of genetic regulation of growth in prawn culture. No reports on transgenic prawns have been published (Benzie 1998). The introduction of DNA into prawn eggs by microinjection has been successfully achieved (Cadoret et al. 1991; Preston & Atkinson 1995). ß-glucuronidase reporter genes have been expressed in prawn eggs after injection of foreign DNA (Rothlisberg 1998). The innate immune system is the only defence weapon of invertebrates (Magnadóttir 2006). Innate immune defence relies on the production of more than 400 antimicrobial peptides, which can be found in both prokaryotic and eukaryotic organisms (Bachère 2003). Antimicrobial peptides are produced in the blood cells of crustaceans (Destoumieux et al. 2000). These peptides have a huge range of antimicrobial functions against Gram-positive and Gram-negative bacteria, and against filamentous fungi (Bachère 2003). Using the same molecular approach of expressed sequence tag analysis, a putative homologue of the 11.5kDa polypeptide has been characterised from the haemocyte of L. vannamei and L. setiferus (Gross et al. 2001). Homologies to the limulidae anti-lipopolysaccharide factor have been found in P. monodon (Tassanakajon et al. 2000). A family of antimicrobial peptides, called penaeidins, has been fully characterised and cloned from the haemocyte of L. vannamei (Destoumieux et al. 1997). These peptides are ubiquitous in crustaceans (Bachère 2003). The penaeidins are original 5.5–6.5-kDa peptides and display the dual function of antimicrobial activities (antifungal and anti-Gram positive bacterial activities) (Destoumieux et al. 1999) and a chitin-binding property (Bachère et al. 2000). Alpha-2-macroglobulin, protein that interacts with phagocytosis activating protein (PAP), may facilitate the entry of glutathione-S-transferase-PAP into phagocytic cells, and increases the survival of WSSVinfected P. monodon (Chotigeat et al. 2007).

5.2.6

Domesticated programmes

Some wild stocks are now being threatened or rendered extinct so the development of domesticated strains is urgently needed (Laubier & Laubier 1993) to alleviate problems of broodstock shortages (Coman et al. 2007). Most farmed prawns are currently produced from ocean- and wild-caught broodstock (Coman et al. 2002), which are often infected with prawn viral diseases (Flegel & Alday-Sanz 1998; Otta et al. 1999b). Specific pathogen free or disease-resistant domesticated stocks need to be developed (Wyban et al. 1993; Carr et al. 1996; Benzie 1998; Hennig et al. 2005; Pantoja et al. 2005). Research has been focused on the deleterious consequences of uncontrolled inbreeding, and engendered attempts to maintain larger population sizes in breeding programmes have been mentioned (Malecha & Hedgecock 1989). Intense global efforts are being made to develop captive breeding programmes in biosecure settings for most commercially farmed prawn species (Browdy 1998; Paibulkichakul

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et al. 2008). Recently, attempts to domesticate high-value prawn species have been focusing on P. monodon (Kenway et al. 2006; Wongprasert et al. 2006; Macbeth et al. 2007; Kenway et al. 2008) L. vannamei (Alfaro et al. 2004), F. chinensis (Hennig et al. 2005; Pantoja et al. 2005) and P. japonicus (Sellars et al. 2005a,b, 2006b, 2007). To date, P. monodon have been difficult to breed in captive conditions (Menasveta et al. 1993; Benzie 1998) and have not been possible to domesticate on a commercial scale (Macbeth et al. 2007). However, other species have been successfully domesticated on a commercial scale, including L. vannamei (Argue et al. 2002; Pascual et al. 2004) and P. stylirostris (Goyard et al. 2002). The minimum requirements for spawning have been established for a number of penaeid species (McVey 1993). These requirements include nutrient metabolism, genetics and environmental factors (Primavera 1985; McVey 1993; Benzie et al. 1997). Dietary supplementation with 8% fish oil (12% total lipid) and 280 mg/kg astaxanthin improved P. monodon maturation and spawning success (Paibulkichakul et al. 2008). Considerable efforts have been made to domesticate prawns using both green and clear water systems (Coman et al. 2005). Greenwater rearing systems are more problematic in maintaining biosecurity because of difficulties in eradicating and preventing the introduction of pathogens into the systems (Sellars et al. 2006b), while clear water systems such as raceways or tanks allow greater control over influents and are more suited to biosecure broodstock production (Coman et al. 2005). Determining the optimum husbandry protocols and diets for prawns may lead to a maximisation of the reproductive outputs of the domesticated stocks (Coman et al. 2007). Reproduction capacity of P. japonicus was negatively impacted by ionising radiation at 20 Gray for females and 10 and 20 Gray for males (Sellars et al. 2005b). Morphometric data and reliable characteristics are used to assess broodstock quality and assist in the choice of individuals for inclusion in breeding programmes (Goswami et al. 1990). A selective breeding programme of P. monodon reared in captivity in tanks over three generations gave full pedigree information and concurrently improved both growth and survival (Kenway et al. 2006). Rearing density is a key factor influencing the growth and survival rates of P. japonicus (Coman et al. 2004). The decrease in densities from 3 to 1 prawn/m2 of second-generation domesticated female P. monodon resulted in an improvement in growth and reproductive performance such as per spawning or per female (Coman et al. 2007). P. japonicus broodstock reared in indoor tanks are less at risk of Mourilyan virus (MoV) infection and have a greater capacity to tolerate infection compared to those reared in farm ponds (Sellars et al. 2006b). Evaluation of the reproductive performances of prawn stocks is based on a comparison with wild-caught broodstock (Palacios et al. 1999; Peixoto et al. 2003), while the value of domestication programmes is assessed not only through this direct comparison but also by comparisons of the overall economic benefits to the industry (Coman et al. 2006). These beneficial achievements include percentages of female spawning (Hansford & Marsden 1995), time from ablation to first spawning, number of spawning per female (Menasveta et al. 1993, 1994) and hatching rate per spawning (Hansford & Marsden 1995). Captive-reared broodstock P. monodon have exhibited poor reproductive performance (Benzie 1997) with limitation in attempts to domesticate, when estimates of genetic correlations between reproductive strains and weight or growth were done (Macbeth et al. 2007). Captive-reared P. monodon have been reported to mature and spawn less frequently than wild-caught ones (Menasveta et al. 1993). In captive breeding programmes of P. japonicus from egg to maturity, broodstock reared in indoor tanks showed a higher survival than

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sibling broodstock reared in farm ponds (Sellars et al. 2006b). In contrast, wild females outperformed domesticated females in terms of maturation, spawning and total egg production in P. monodon (Menasveta et al. 1993; Hall et al. 2003) and P. stylirostris (Mendoza 1997). The survival of PL20 from hybridisation between some Penaeus spp. was lower than 1% (Lin et al. 1988; Bray et al. 1990a, b; Benzie et al. 1995). Most of these hybrids were not reared for long enough to mature, and, if they were, no maturation of female was recorded and no matings were observed among them (Bray et al. 1990a,b). The goal of the reproduction programmes of tank-reared P. monodon stocks is to improve the quality of female broodstock (Coman et al. 2006).

5.2.7

Food security

Food has to be safe for use; however, the absolute safety of any human endeavour is impossible to achieve (Hammes & Hertel 2002). Although micronutrients are widely used in aquaculture, their effectiveness is not well known. For example, boron, as a fertiliser in agriculture and as an ingredient of pesticides (Li et al. 2008), is an essential micronutrient for the normal development and growth of organisms (Moseman 1994). L. vannamei at low salinity of 3 ppt is more sensitive to the ambient boron toxicity than at a higher salinity of 20 ppt (Li et al. 2008). Although some proposed methods for the control of disease in aquaculture such as the use of probiotics, immunostimulants and zoo-techniques have made some progress (Robert & Gérard 1999; Bachère 2003), antibiotics and chemicals have been intensively used as preventive and curative means (Barg & Lavilla-Pitogo 1996; Bachère 2003). The question is, what are the consequences of probiotics in nutrition and on the well-being of the consumers (Hammes & Hertel 2002)? Besides an appearance of antibiotic-resistant bacteria caused by regular use or misuse of antibiotics (Karunasagar et al. 1994; Molina-Aja et al. 2002; Chelossi et al. 2003; Sahul Hameed et al. 2003b), residue from the antibiotics and chemicals has degraded the environment (Kautsky et al. 2000; Bachère 2003) and could pose a risk to humans (Schwarz et al. 2001; Kesarcodi-Watson et al. 2008). The transferral process of resistant genes between bacteria (Schwarz et al. 2001) means that antibiotic-resistant bacteria originating from prawn farms could potentially transfer plasmids to bacteria involved in human health problems (Kesarcodi-Watson et al. 2008). Fortunately, there is insufficient data to show a linkage to resistant gene transferral to humans (Kesarcodi-Watson et al. 2008). Governments and organisations have introduced much tighter restrictions for antibiotic use in animal production. For example, the European Union initially put a ban on the use of avoparcin in 1997 and in 1999 also banned virginiamycin, spiramcin, tylosin and bacitracin as well as growth promoters and all nontherapeutic antimicrobials (Turnidge 2004; Delsol et al. 2005). Many Asian countries have increased antibiotic controls because of foreign restrictions on antibiotic-contaminated products for export markets. Nevertheless, trace levels of chloramphenicol have been detected in prawn products from Thailand, Myanmar, India, Pakistan and Vietnam (Heckman 2004; Kesarcodi-Watson et al. 2008). The GLOBALGAP standard is designed to minimise detrimental environmental impacts of farming operations, to reduce the use of chemical inputs and to ensure a responsible approach to the health and welfare of producers and consumers. This standard serves as a practical manual for good agricultural practice (GAP) for all interested parties, i.e. producers, suppliers, retailers, journalists and consumers. Therefore, country prawn producers try to establish efficient certification standards and

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procedures for their market competion and sustainable bussiness. Moreover, producers need to obtain other certifications such as BMP, GAP or EuroGAP to meet the requirements of strict overseas country/region markets.

5.3

CHALLENGES

The potential challenges to penaeid aquaculture need to be recognised and identified as the constraints must be addressed and overcome during the development process of prawn farming. The first challenge is to expand production to meet the increasing demand for aquaculture products (Zhang 2007), which can no longer be supplied by captured fisheries because of dwindling aquatic stocks and overfishing (Ahmed et al. 2007). Development strategies and policies are clearly defined for specific areas or regions. Sustainable development of penaeid prawns will depend upon feasible solutions for environmental problems, proper development planning and policy enforcement (Cao 2007), as the development of prawn aquaculture is associated with disease and environmental problems. For example, there is still no effective solution for suspended solid and sediment in prawn ponds. The coastal zone serves as the basic ecosystem for prawns and is easily influenced by any small changes of environmental conditions under prawn culture activities. Its effectiveness takes a long time to recover. Diversification of culture environments and species, development of hatcheries for high commercial value species and/or high demand by consumers, development of domestic feed industry, and provision of insurance and other related services are still unexplored economically in penaeid aquaculture (Hishamunda 2007). The diversity of cultured species has suffered from the introduction of exotic species into aquaculture because of problems such as environmental degradation and the transfer of new diseases. For example, the introduction of L. vannamei has been causing negative effects to environment and biodiversity in Thailand, Malaysia and Indonesia. The prawn farmers in these countries have complained that L. vannamei can grow and survive normally, but have abdominal segment deformity diseases caused by a new virus (Sakaew et al. 2008). This problem may also occur in Vietnam or other Asian prawn producer countries. L. vannamei served as a Taura syndrome virus carrier, when they were introduced from the Pacific coast of the Americas to Asia in the late 1990s (Sakaew et al. 2008). In addition, the world tends to consume less large, expensive prawns and more small, cheap prawns. Therefore, prawn farmers are rushing to farm L. vannamei instead of P. monodon due to the shorter period required to realise more profit more quickly. However, it is predicted that consumers’ demand for L. vannamei is just temporary, and P. monodon will remain the top choice for the long term. This may challenge the prawn farmers in operating their farms. Practical problems relating to growth and survival rate, disease prevention, maturation and larval quality or even chemicals/antibiotic residues in aquaculture products, are becoming more common under traditional/conventional culture. Water pollution, which has been reaching warning alert levels in some countries, affects aquaculture production directly or indirectly at various levels. Intensification of aquaculture activities has increased the occurrence of diseases (Shariff et al. 2001). Massive loss caused by diseases has closed some commercial farming or even aquaculture regions. Defective rearing conditions such as high density, excess food or poor food quality are associated with mortalities caused by either heterotrophic or pathogenic bacterial strains such as Pseudomonas, Aeromonas and Vibrio,

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in which Vibrio species are responsible for disease outbreaks in all stages of development of prawns such as V. damsela (Song et al. 1993), V. alginolyticus, V. parahaemolyticus (Lightner 1992), V. penaeicida (Costa et al. 1998) or V. harveyi (Karunasagar et al. 1994; Robertson et al. 1998). Viral diseases have emerged as the most serious enzootic pathogens. Moreover, difficulties in controlling diseases come from the various susceptibilities of aquatic animals to the diversity of pathogens (Bachère 2003). Although immunostimulants have been recommended for disease control, outcomes are dissimilar. These results are attributed to a lack of an understanding of the close relationship between the hosts, pathogens and environmental stressors (Sánchez-Martínez et al. 2007). Doubts about the efficacy and safety of probiotics on the market arise from the use of ineffective bacterial species/strains, unrealistic claims, lack of scientific evidence, poor quality control during product processing, and inappropriate delivery methods leading to contamination or reduced performance (Temmerman et al. 2003). On the other hand, vibriosis can be controlled by the use of antibiotics, vaccine and immunostimulants such as probiotics and prebiotics, but viral injection in prawns has no effective therapeutic or prophylactic countermeasures (Itami et al., 1998). The question raised here is the value of biochemical compounds produced from immunostimulants for cost-effective control of disease injections in aquaculture, especially for long-lasting protection in juveniles and adults of prawns (Smith et al. 2003). The health and zootechnical performances can be improved by the prophylactic use of probiotics in cultured crustacean species. These uses should be considered to be a kind of risk insurance in that they do not provide notable benefits when the culture is performing under optimal conditions and in the absence of opportunistic pathogens, but are helpful if infectious diseases break out (Verschuere et al. 2000). Once it has been decided to apply probiotics on a large scale, production of large amounts of bacterial biomass/probiotics requires appropriate quality control to avoid contamination by other bacteria (Verschuere et al. 2000). Therefore, the development of appropriate probiotics/prebiotics for speciesspecific situations is not a simple task and requires empirical and fundamental research, full-scale trials, appropriate product monitoring and control measures (Decamp et al. 2008). A question emerges as to how many opportunities are left for pathogens to grow and become a threat, when appropriate probiotics dominate other microbial communities, and one useful approach is to carefully monitor shifts in the overall microbial community (Verschuere et al. 2000). Few papers on effective ways of carrying out sustainable invertebrate aquaculture can be found because of a lack of fundamental data on the biology and genetic characterisation of prawns (Bachère 2003). The progress in penaeid prawn genetic and biotechnology research has been slow due to the gap in knowledge of the fundamental aspects of specific species biology (Benzie 1998). Loss of genetic variability is maximised in L. vannamei closed broodstock-rearing systems, where post-larvae are produced from crosses of breeders collected from an associated growout farm after mass selection (de Lima et al. 2008). The difficulty of providing controlled conditions and producing large numbers of spawners leads to a slowdown in efforts to estimate the heritability of quantitative traits of farmed species (Chow & Sandifer 1991). Fully artificial maturation diets as the key goals to reduce production costs and to increase spawner yield are not available (Benzie 1997). Environmental and endocrine replacements for eyestalk ablation have not fully taken the vital role in breeding programme (Benzie 1997). Large-scale production of high-quality larvae from pond-reared stock depends on an understanding of the role of genetics and

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environment on maturation and larval quality (Benzie 1997). No data exist for synthetic models of endocrine control of either male or female crustaceans (Benzie 1997). Heavy exploitation of populations may put the wild populations at risk, with potentially deleterious long-term consequences for conservation, fisheries and future aquaculture development (Sugama et al. 2002). Moreover, the introduction of large numbers of spawners of P. monodon from the Andaman Sea to the Gulf of Thailand has changed the genetic structure of the wild stocks in the area (Sodsuk et al. 1996). Attention needs to be paid to the distribution of cultured stocks as they may endanger wild genetic resources (Benzie 1996). One problem is male infertility in the domesticated P. monodon due to low sperm count in the spermatophore (Pratoomchat et al. 1993) or low capacity of individual sperm to fertilise the egg, or both (Pongtippatee et al. 2007). Understanding morphological events of sperm of P. monodon before and at the time of fertilisation is the key for further research on sperm infertility in this species (Pongtippatee et al. 2007). As can be seen, while prawn production has been in a decline due to environmental degradation and diseases, the global demand for prawn products has been continuously increasing. Meanwhile, experts assert that the requirement on the quality of prawn products in the global markets should be increasingly stricter, and prawn prices have been declining worldwide. Therefore, these significant effects may influence the prawn producers’ income markedly. The establishment or implementation of prawn farming now needs to be considered carefully in all aspects of not only technology but also quality and market aspects in terms of sustainable development.

5.4

PROSPECTIVE/FUTURE OUTLOOK

Food requirements for prawns, such as proteins, lipids, carbohydrates, coppers, vitamins, cholesterol, essential fatty acids and minerals, are now well known, hence feed can be more accurately formulated for specific species (Cuzon et al. 2004). Replacement of expensive proteins with readily available and cheap protein sources is possible in terms of economics and environmental considerations because, although gut physiologies, digestive enzymes and habitats are similar, feeding habits can change (Cuzon et al. 2004). In addition, the world demand for aquatic products mainly comes from aquaculture as the production from fisheries is limited. Among aquatic products, prawns still remain the top choice. Fundamental work on physiology, genetics and immunology at a molecular level has been progressing to create new tools to assist producers in adequate health management and prevention of diseases (Bachère 2003). Important progress has been made in the prevention and control of diseases, with reliable diagnosis methods based on molecular probedetection techniques (Walker & Subasinghe 2000). Genetic molecular markers and the criteria of survival to injections are used for establishing a marker-assisted selection approach in the cultured species (Bachère 2003). Studies on the properties and function of penaeidins led to the understanding of physiology and their capacity to respond to pathological injuries (Bachère 2003). Molecular and biotechnological tools are available for investigating details of the genome and biology of penaeid prawns (Benzie 1998). Information concerning the amount and distribution of genetic variation is vital for the design and implementation of adequate management strategies for the species (Sugama et al. 2002). Therefore, an integrated approach to the control of diseases is required with

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consideration of the combination of ecological, environmental and zootechnical aspects of production (Bachère 2000). Fortunately, the farmers’ experiences on prawn culture have been accumulated and improved by time. In addition, biodiversity of prawn species and various culture systems has been determined in terms of effectiveness and sustainability. Nowadays, organic farming is emerging to meet the strict requirements of global markets and sustainable development in prawn aquaculture. Prawn producers need to preserve genetic diversity for further development of prawn aquaculture, and should make sure that the introduction of non-native stocks to increase production does not have adverse effects on the genetic integrity of local populations (Chareontawee et al. 2007). Interestingly, the strict requirements to rear the product without using antibiotics and banned chemicals, while also minimising effects on the culture environment or trademark, has led to the development of greater competition and sustainable development in prawn aquaculture. Although unplanned expansion of penaeid prawn aquaculture in coastal areas in recent decades has resulted in an increase in production, this activity has been the result of its production ecosystems. The history of this process needs to be explored to avoid the mistakes experienced elsewhere (Hishamunda 2007). The development of appropriate management practices, the control of environmental variables, genetic improvement, understanding of virus physiology, modulation of the prawn immune system; all have contributed to the effective control of WSSV in prawn aquaculture so far. Therefore, successful farms are the ones using specific-pathogen-free prawns, correctly monitoring samples and undertaking evaluation using PCR, histopathology, filing and using serological methods on newly introduced broostock, eggs, post-larvae and juvenile prawns (Sánchez-Martínez et al. 2007). Generally, progress in immunology and physiology of the cultured species provides potential approaches for disease control at various ontogentic stages of production of penaeids (Bachère 2003). Alternative prophylactic methods including the use of probiotics and prebiotics are proposed for the control of diseases in aquaculture, as enhancing resistance against pathogens of prawns by improving prawns’ immune systems is considered one of the effective strategies for the control of prawn diseases (Huang et al. 2006). These methods help protect aquatic animals against infections (Verschuere et al. 2000; Vici et al. 2000; Bachère 2003) as probiotics are an effective addition to disease control strategies in aquaculture (Irianto & Austin 2002a; Balcázar et al. 2006a). It is possible to vaccinate P. japonicus with recombinant proteins against WSSV (Namikoshi et al. 2004). Interestingly, probiotics as an adjuvant for detoxification protocols (Brudnak 2002) may emerge as an idea for aquaculture. There are doubts about the use of probiotics, especially regarding the limitations and inside effects of pharmaceutical agents, and consumer demand for natural products is high. The establishment of a consensus on probiotic product regulations is required to assist in the enforcement of guidelines and standards, and appropriate clinical studies need to be carried out on the mechanisms of actions of strains (Reid 2006). Knowledge of the dynamics of bacterial activity, differences in efficacies of vaccines, different responses of various aspects of the immune systems, and limited duration of antibacterial activity increases manipulative advantages in prawn culture systems (Alabi et al. 2000). There is an increasing need to control, prevent or minimise the devasting effects of disease in prawn culture without using toxic chemicals or antibiotics (Smith et al. 2003). The development of domesticated programmes to relieve pressure on wild stock, and to improve industrial production through selective breeding programmes, will require the inclusion of appropriate genetic variation in the broodstock population (Taniguchi et al.

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1994). Rearing large numbers of families from defined matings has been successful in P. vannamei (Wyban et al. 1993), P. monodon (Benzie 1997) and P. stylirostris (Bedier et al. 1996). Sperm quality of pond-reared P. monodon males was not different from that of wild males (Pratoomchat et al. 1993) so that husbandry methods can provide adequate conditions to achieve normal male fertility levels. Domesticated programmes in biosecure tank systems help alleviate high pressure on wild-caught broodstock, enable control over pathogens and generate farmed stocks. These programmes, with improvements in broodstock diets and husbandry, will meet the reproductive outputs from tank-reared prawns (Coman et al. 2006). Through breeding programmes using massive selection and inbreeding effects, a L. vannamei strain produced animals free of the three most dangerous diseases of TSV, WSSV and YHV, so this is a great resource for the prawn culture industry (De Donato et al. 2005). Browdy (1998) believed that if broodstock diet, health and other conditions in a maturation system are well maintained, the quality of spawns of captive matured females may be as good as that of wild females. The long-term sustainability of the prawn industry depends on the equilibrium among environmental quality, prevention of diseases by adequate diagnosis and epidemiological surveys of the pathogens, and the prawn health status, as well as selection programmes to obtain disease-resistant prawns (Bachère, 2000). Research and its application have been developing to meet the increasing demand on global aquatic products, and bringing perspective to penaeid prawn aquaculture. Strategic advances for penaeid prawn aquaculture have anticipated the combination of endocrine control of reproduction, its effects on egg quality, molecular markers, maturation diets and practical husbandry methods (Benzie, 1997). Finally, macro-government monitoring should be established to prevent disease spread and to increase production for sustainable development of penaeid prawn aquaculture.

5.5

REFERENCES

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Penaeid Prawns 177 Verschuere, L., Rombaut, G., Sorgeloos, P. & Verstraete, W. (2000) Probiotic bacteria as biological control agents in aquaculture. Microbiology & Molecular Biology Reviews. American Society for Microbiology, 64, 655–671. Vici, V., Bright Singh, I.S. & Bhat, S.G. (2000) Application of bacterins and yeast Acremonium dysporii to protect the larvae of Macrobrachium rosenbergii from vibriosis. Fish & Shellfish Immunology, 10, 559–563. Vidal, O.M., Granja, C.B., Aranguren, F., Brock, J.A. & Salazar, M. (2002) A profound effect of hyperthermia on survival of Litopenaeus vannamei juveniles infected with while spot synderome virus. Journal of World Aquaculture Society, 32, 364–372. Vijayan, K.K., Bright Singh, I.S., Jayaprakash, N.S., et al. (2006) A brackishwater isolate of Pseudomonas PS-102, a potential antagonistic bacterium against pathogenic vibrios in penaeid and non-penaeid rearing systems. Aquaculture, 251, 192–200. Vine, N.G., Leukes, W.D. & Kaiser, H. (2006) Probiotics in marine larviculture. FEMS Microbiology Reviews, 30, 404–427. Vuthiphandchai, V., Nimrat, S., Kotchalat, S. & Bart, A.N. (2007) Developement of a cryoprevervation protocol for long-term storage of black tiger shrimp (Penaeus monodon) spermatophores. Theriogenology, 68, 1192–1199. Walker, P., Subasinghe, R. (2000) DNA-based molecular diagnosis techniques. FAO Fisheries Technical Paper 395, 93. FAO, Rome. Wang, Y.-B. (2007) Effect of probiotics on growth performance and digestive enzyme activity of the shrimp Penaeus vannamei. Aquaculture, 269, 259–264. Wang, F.-I. & Chen, J.-C. (2006) The immune response of tiger shrimp Penaeus monodon and its susceptibility to Photobacterium damselae subsp. damselae under temperature stress. Aquaculture, 258, 34–41. Wang, Y.-B. & Han, J.-Z. (2007) The role of probiotic cell wall hydrophobicity in bioremediation of aquaculture. Aquaculture, 269, 349–354. Wang, Y.-C., Lo, C.-F., Chang, P.-S. & Kou, G.-H. (1998) Experimental infection of white spot baculovirus in some cultured and wild decapods in Taiwan. Aquaculture, 164, 221–231. Wang, W.-N., Wang, A.-L. & Zhang, Y.-J. (2006a) Effect of dietary higher level of selenium and nitrite concentration on the cellular defense response of Penaeus vannamei. Aquaculture, 256, 558–563. Wang, Y.-C., Chang, P.-S. & Chen, H.-Y. (2006b) Tissue distribution of prophenoloxidase transcript in the Pacific white shrimp Litopenaeus vannamei. Fish & Shellfish Immunology, 20, 414–418. Wang, Y.-C., Chang, P.-S. & Chen, H.-Y. (2007) Tissue expressions of nine genes important to immune defence of the Pacific white shrimp Litopenaeus vannamei. Fish & Shellfish Immunology, 23, 1161–1177. Wang, Y.-C., Chang, P.-S. & Chen, H.-Y. (2008) Differential time-series expression of immune-related genes of Pacific white shrimp Litopenaeus vannamei in response to dietary inclusion of ß-1,3-glucan. Fish & Shellfish Immunology, 24, 113–121. Welker, T.L., Lim, C., Yildirim-Aksoy, M., Shelby, R. & Klesius, P.H. (2007) Immune response and resistance to stress and Edwardsiella ictaluri challenge in channel catfish, Ictalurus punctatus, fed diets containing commercial whole-cell yeast or yeast subcomponents. Journal of World Aquaculture Society, 38, 24–35. Weston, D.P. (1996) Environmental considerations in the use of antibacterial drugs in aquaculture. In: Baird, D., Beveridge, M.V.M., Kelly, L.A. & Muir, J.F. (eds.), Aquaculture and Water Resource Management, pp. 140–165. Blackwell, Oxford. Williams, A.S., Davis, D.A. & Arnold, C.R. (1996) Density-dependent growth and survival of Penaeus setiferus and Penaeus vannamei in a semi-closed recirculating system. Journal of World Aquaculture Society, 27, 107–112. Withayachumnarnkul, B., Plodpai, P., Nash, G. & Fegan, D. (2001) Growth rate and reproductive performance of F4 domesticated Penaeus monodon broodstock. The 3rd national symposium of marine shrimp, 8–9 November 2001, Queen Sirikit National Convention Center, Bangkok, Thailand, pp. 33–40. Witteveldt, J., Vlak, J.M. & van Hulten, M.C. (2004a) Protection of Penaeus monodon against white spot syndrom virus using a WSSV subunit vaccine. Fish & Shellfish Immunology, 16, 571–579. Witteveldt, J., Cifuentes, C.C., Vlak, J.M. & van Hulten, M.C. (2004b) Protection of Penaeus monodon against white spot syndrom virus by oral vaccination. Journal of Virology, 78, 2057–2061. Wolfus, G.M., Garcia, D.K. & Alcivar-Warren, A. (1997) Application of the microsatellite technique for analyzing genetic diversity in shrimp breeding programs. Aquaculture, 152, 35–47.

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Wongprasert, K., Asuvapongpatana, S., Poltana, P., Tiensuwan, M. & Withyachumnarnkul, B. (2006) Serotonin stimulates ovarian maturation and spawning in the black tiger shrimp Penaeus monodon. Aquaculture, 261, 1447–1454. Woods, C.M.C. (2003) Effects of varying Artemia enrichment on growth and survival of juvenile seahorses, Hippocampus abdominalis. Aquaculture, 220, 537–548. Wu, J.L. & Muroga, K. (2004) Apoptosis does not play an important role in the resistance of ‘immune’ Penaeus japonicus against white spot syndrom virus. Journal of Fish Diseases, 27, 15–21. Wu, Y., Lu, L., Yang, L.-S., Weng, S.-P., Chan, S.-M. & He, J.-G. (2007) Inhibition of white spot syndrome virus in Litopenaeus vannamei shrimp by sequence-specific siRNA. Aquaculture, 271, 21–30. Wyban, J.A., Swingle, J.A., Sweeney, J.N. & Pruder, G.D. (1993) Specific pathogen-free Penaeus vannamei. World Aquaculture, 24, 39–45. Xiang, J., Li, F., Zhang, C., et al. (2006) Evaluation of induced triploid shrimp Penaeus (Fenneropenaeus) chinensis cultured under laboratory conditions. Aquaculture, 259, 108–115. Xiang, J.H., Zhou, L.H., Liu, R.Y., et al. (1992) Introduction of the tetraploids of the chinese shrimp, Penaeus chinensis, Proceedings of the Asia Pacific Conference on Agricultural Biotechnology Beijing, 20–24 Aug. 1992, pp. 841–846. China Science and Technology Press, Beijing. Xu, Z., Primavera, J.H., de la Pena, L.D., Pettit, P., Belak, J. & Alcivar-Warren, A. (2001) Genetic diversity of wild and cultured black tiger shrimp (Penaeus monodon) in the Philippines using microsatellites. Aquaculture, 199, 13–40. Yan, D.C., Dong, S.L., Huang, J., Yu, X.M., Feng, M.Y. & Liu, X.Y. (2004) White spot syndrome virus (WSSV) detected by PCR in retifers and retofer resting eggs from shrimp pond sediments. Diseases of Aquatic Organisms, 59, 69–73. Yan, D.-C., Feng, S.-Y., Huang, J. & Dong, S.-L. (2007) Rotifer cellular membranes bind to white spot syndrome virus (WSSV). Aquaculture, 273, 423–426. Yeh, S.-T. & Chen, J.-C. (2008) Immunomodulation by carrageenans in the white shrimp Litopenaeus vannamei and its resistance against Vibrio alginolyticus. Aquaculture, 276, 22–28. Yi, G., Qian, J., Wang, Z. & Qi, Y. (2003) A phage-displayed peptide can inhibit infection by white spot syndrom virus of shrimp. The Journal of General Virology, 84, 2545–2553. Yoganandhan, K., Narayanan, R.B. & Sahul Hameed, A.S. (2003) Larvae and early post-larvae of Penaeus monodon (Fabricius) experimentally injected with white spot syndrome virus (WSSV) show no significant mortality. Journal of Fish Diseases, 26, 385–391. Yu, Z.M., Li, C.W. & Guan, Y.Q. (2003) Effect of salinity on the immune reponses and outbreak of white spot syndrom in the shrimp Marsupenaeus japonicus. Ophelia, 57, 99–106. Zhang, C., Li, F., Yu, K. & Xiang, J. (2008) Comparative growth performances of diploid and triploid Chinese shrimp Fenneropenaeus chinensis (Osbeck, 1765) under different salinities. Aquaculture Research, 39, 962–969. Zhang, C., Li, F., Yu, K., Wu, C., Guo, Z. & Xiang, J. (2004) Haematological changes of sibling dipoid and tripoid Fenneropenaeus chinensis after being challenged with pathogens. Journal of Fisheries of China/Shuichan Xuebao, 28, 535–540. Zhang, J.-S., Dong, S.-L., Tian, X.-L., Dong, Y.-W., Liu, X.-Y. & Yan, D.-C. (2006) Studies on the rotifer (Brachionus urceus Linnaeus, 1758) as a vector in white spot syndrome virus (WSSV) transmission. Aquaculture, 261, 1181–1185. Zhang, X. (2007). Aquaculture in China. In: Species and system selection for sustainable aquaculture (eds P. Leung, C.-S. Lee & P.J. O’Bryen), pp. 131–143. Blackwell Publishing, Ames, Iowa. Zhang, X., Li, F. & Xiang, J. (2003) Chromosome behaviour of heat shock induced trilpoid in Fenneropenaeus chinensis. Chinese Journal of Oceanology & Limnology, 21, 222–228. Ziaei-Nejad, S., Rezaei, M.H., Takami, G.A., Lovett, D.L., Mirvaghefi, A.-R. & Shakouri, M. (2006) The effect of Bacillus spp. bacteria used as probiotics on digestive enzyme activity, survival and growth in the Indian white shrimp Fenneropenaeus indicus. Aquaculture, 252, 516–524. Zimba, P.V., Camus, A., Allen, E.H. & Burkholder, J.M. (2006) Co-occurrence of white shrimp, Litopenaeus vannamei, mortalities and microcystin toxin in a southeastern USA shrimp facility. Aquaculture, 261, 1048–1055. Zurawell, R.W., Chen, H., Burke, J.M. & Prepas, E.E. (2005) Hepatotoxic cyanobacteria: a review of the biological importance of microcystins in freshwater environments. Journal of Toxicology & Environmental Health, 8, 1–37.

6

Cobia Culture

Ravi Fotedar and Huynh Minh Sang

6.1 INTRODUCTION Culture of cobia began in the late 1990s after the technology of mass fry production was developed in 1997. In recent years, cobia has emerged as potentially the most important species in the aquaculture sector. Cobia is one of the species that fulfil the conditions and criteria set by aquaculturists and has been regarded as having the greatest potential among all candidate species for offshore cage culture in tropical waters. Cobia can be grown from fingerling (Fig. 6.1) to 4–6 kg in 1 year (Franks et al. 1999) and there is usually a very high survival rate of around 90%. Besides exhibiting a rapid growth rate, cobia also has a good flesh quality, which is the most important characteristic in marketing (Kaiser & Holt 2005). Due to its suitability for culture along with its high value in international markets, the technology for cobia culture has developed rapidly in the past few years. Efforts are now being focused on the areas of mass propagation through natural spawning of broodstocks, advanced larval rearing techniques, nursery production in tanks, ponds and nearshore cages, and culture in offshore cages. Since wild-caught cobia does not represent a major fishery and the farming of cobia is in its infancy, details regarding the market and trade of this species are notably lacking. As a result of its current limited availability, many seafood consumers have probably never tasted cobia. Increasing supplies from aquaculture (Fig. 6.2), combined with effective marketing of this firm-textured, white-fleshed fish will be critically important for future market expansion.

6.2

MORPHOLOGY

Cobia, Rachycentron canadum (Linnaeus 1766), is the sole representative of the family Rachycentridae. The fish has an elongate body that is strongly rounded with a broad, flat head. The mouth is terminal in position with a projecting lower jaw. The first dorsal fin comprises seven to nine short, stout isolated spines and the second dorsal fin is long with

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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Fig. 6.1

A cobia fingerling. (Please see plate section for colour version of this figure.)

Fig. 6.2

Table-sized cobia from a cage-farming operation in Vietnam.

anterior rays forming a raised lobe in adults. The anal fin is similar to second dorsal and both are covered in thick skin. The caudal fin is lunate and crescent shaped in adults with the upper lobe longer than the lower lobe. The caudal fin in the very young is paddle shaped. Scales are small and embedded in the skin. Coloration of the cobia is highly variable. Adults are uniform chocolate brown to bronze on their back and sides with a white to cream col-

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oured belly. Two lateral stripes running the full length of their body may be present. The stripes can range from stark white to bronze in colour. Stripes in juveniles tend to be brighter in colour ranging from stark white to chartreuse, possibly with a touch of rose on the caudal fin (Hammond 2001).

6.3

DISTRIBUTION

Cobia is distributed worldwide, occurring in tropical and subtropical seas except for the eastern Pacific (Briggs 1960; Shaffer & Nakamura 1989). Cobia occurs in the western Atlantic Ocean from Massachusetts and Bermuda to Argentina (Briggs 1960). Shaffer and Nakamura (1989) reported cobia to be most abundant along the south Atlantic coast of the USA and the northern Gulf of Mexico. The Marine Recreational Fisheries Statistical Survey conducted by the National Marine Fisheries Service recorded recreational cobia landings from Texas to New York (Hammond 2001). Tagging studies have documented the movement of fish in both directions between the Gulf and the Carolinas, suggesting gene exchange between Atlantic and Gulf stocks. Cobia inhabits both coastal and continental shelf waters, and adult fish have been found in bays and estuaries as well as at depths of 1,200 m). Temperature appears to be the primary factor in determining their range; although specimens have been collected in waters from 16 to 32 °C, they appear to prefer temperatures above 20 °C. Capture data indicate that, during cooler months of the year throughout their range, cobia either migrate to warmer water in a north–south pattern or move further offshore to deeper water. Cobias tolerate a wide range of salinities. Specimens have been collected in waters with 22 to 44 ppt salinity and reared in culture systems down to 5 ppt (Kaiser & Holt 2005).

6.4 6.4.1

BIOLOGICAL CHARACTERISTICS Growth

Male cobia collected in North Carolina mature at age 2, when they are 60 to 65 cm in length. Females also matured at age 2, but averaged 80 cm (Smith 1995). The oldest fish sampled in the Gulf were found to be an 11-year-old female and a 9-year-old male (Franks et al. 1999). A female cobia caught from the Gulf of Mexico measured over 160 cm in fork length (FL) (measured from tip of upper jaw to fork in tail) and weighed 62.2 kg (Franks et al. 1999). Female cobia from the northeastern Gulf of Mexico have a mean fork length of 1,050 mm and are significantly larger than males, which have a mean FL of 952 mm. Male cobia (525–1,330 mm FL) ranged from age 0 to 9, and females (493–1,651 mm FL) ranged from age 0 to 11. The relationship of observed fork length and age was described by the von Bertalanffy growth equation for males FLt = 1171 × (1-exp [−0.432(t + 1.150)]) and for females FLt = 1555 × (1-exp [−0.272(t + 1.254)]). Growth in length for both sexes was relatively fast through age 2, after which growth slowed gradually. Estimates of the von Bertalanffy growth equation parameters L∞ and K were significantly different for males and females, whereas differences in estimates for to were not significant (Franks et al. 1999). The instantaneous rate of total mortality (Z) estimated by catch curve analysis for fully recruited ages 4–8 is 0.75 (Franks et al. 1999).

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6.4.2

Reproduction

Reproductive biology of cobia differs among regions. In the Gulf of Mexico in the southern United States, spawning occurred from April through September. Some female cobia (17–32%) had spent or regressed ovaries by July. Total male to female sex ratio is 2.7:1. Gonadosomatic index peaked between May and July throughout the region. Ovaries of females contained both postovulatory follicles (POF) and oocytes in final oocyte maturation (FOM) during all months of the reproductive season. Mean batch fecundity ranged from 377,000 ± 64,500 to 1,980,500 ± 1,598,500 eggs. Batch fecundity showed a positive relationship with fork length and ovary-free body weight (OFBW). Relative batch fecundity was not significantly different among months during the spawning season and averaged 53.1 ± 9.4 eggs/g OFBW (Brown-Peterson et al. 2001). Cobia from the southeastern United States and north-central Gulf of Mexico were estimated to spawn once every 5 days, whereas cobia from the western Gulf of Mexico were estimated to spawn once every 9 to 12 days (Lotz et al. 1996; Brown-Peterson et al. 2001). The reproductive biology of cobia in Hormozgan province (Iran) is different from the Gulf of Mexico (Valinassab et al. 2008). Spawning occurrs from July to the beginning of September. The total male to female sex ratio was 1.3:1.0, which was significantly different from the normal ratio, 1:1 (P < 0.05). The highest sex ratio difference was seen in April. The average absolute fecundity was 1,684,954 eggs. Maximum ova diameter was 0.575 mm belonging to the stage 4 and the minimum was estimated at 0.250 mm belonging to the stage 2. Ova diameter average increased from April onwards and its peak was in July. In Iran, cobia has partial synchronism in oocytes and is a total spawner species (Valinassab et al. 2008). By contrast, the sex ratio of males to females of cobia from northeastern Australian waters is 0.46:1 (van der Velde et al. 2010). Cobia develops hydrated oocytes during a protracted spawning season between September and June. Gonadosomatic index peaks from October to December, coinciding with the monsoon or ‘wet’ season. Estimated length at first maturity for female cobia is 671 mm FL. Length at 50% maturity (L50) for females is estimated at 784 mm FL (1–2 years of age). Batch fecundity ranges from 577,468 to 7,372,283 eggs with a mean of 2,877,669 (± SD 1,603,760) eggs. Relative batch fecundity is 249 eggs per gram, and there is no relationship between relative fecundity and fork length but there is a significant positive relationship between the total numbers of eggs produced and fork length. Spawning frequency is 7.6 days (van der Velde et al. 2010). A positive correlation was found between the proportion of floating eggs and hatch rate for both spawning seasons (Faulk & Holt 2008).

6.4.3

Feeding

During the larvae stage, cobia feed on zooplankton, especially on copepoda. The juvenile and adult cobia is a carnivorous species and feeds primarily on benthic/epibenthic crustaceans and fishes (Shaffer & Nakamura 1989). The stomach of cobia (373–1,530 mm fork length), from the Gulf of Mexico, contained 77.6% of crustacean among the total number of identifiable prey. The second most important prey category was fish, which was dominated by hardhead catfish, Arius felis, and eels. Fish occurred in 58.5% of the total sampling stomachs but only accounted for 20.3% of the total number of prey. The importance of fish as prey increased with increasing size (length) of cobia, with the largest size class of cobia

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(1,150–1,530 mm fork length) showing the highest percentage frequency occurrence of fish prey (84.4%) (Meyer & Franks 1996). The feeding of cobia was dependant upon prey availability rather than upon a few specific food organisms (Meyer & Franks 1996).

6.5 6.5.1

NUTRITIONAL REQUIREMENT OF COBIA Protein and amino acids

Correctly formulating diets (Fig. 6.3) with respect to protein is essential for maximising production efficiency and profits. For instance, an over-formulation of protein increases the cost of the feed unnecessarily and may lead to increased protein catabolism and nitrogen excretion (Fraser & Davies 2009). As a carnivorous fish, cobia requires relatively high concentration of protein in the diet for optimum growth. According to Chou et al. (2001), the optimum concentration of crude protein for maximum growth and lowest FCR is 44.5%. The information on specific amino acid requirements of cobia is relatively limited (Fraser & Davies 2009). The requirement of methionine in juvenile cobia, in terms of maximum growth and lowest FCR, was found to be 1.19% of dry diet in the presence of 0.67% cysteine corresponding to 2.64% of dietary protein dry weight (Zhou et al. 2006). These values for methionine and lysine are in accordance with those of commonly cultured fish, 1.8–3.2% and 3.2–6.2% dietary protein respectively (Wilson 2002; Zhou et al. 2006, 2007).

Fig. 6.3

Formulated diet used for growing cobia in cages.

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6.5.2

Lipid and essential fatty acids

Biochemical composition of eggs and yolksac larvae reflect the basic nutritional requirements of first feeding larvae (Sargent et al. 1999). Thus, cobia larvae and juveniles require high amounts of the highly unsaturated fatty acids (HUFAs), docosahexaenoic acid (DHA 22:6n-3), eicosapentaenoic acid (EPA 20:5n-3) and arachidonic acid (ARA 20:4n-6) as these account for approximately 80% of the PUFAs in cobia eggs and yolksac (Holt et al. 2007). The cobia larvae maintain the high lipid and PUFA content with docosahexaenoic acid (DHA, 22:6n-3), eicosapentaenoic acid (EPA, 20:5n-3) and arachidonic acid (ARA, 20:4n-6) comprising the majority of the PUFAs. In cobia eggs and 1 day post-hatch (dph) larvae the DHA: EPA is 3.3 and the EPA:ARA is 2.2 due to the high ARA content (Faulk & Holt 2003). The initial optimum dietary lipid requirement for juvenile cobia was determined at 5.76% dry matter (Chou et al. 2001). High dietary lipid levels above 15% produced little practical benefit because of higher fat accretion in cobia. Juvenile cobia (7.71 g) displayed no significant difference in weight gain in fish fed diets containing 5% and 15% dietary lipid, though both of these displayed significantly increased growth over the fish on the 25% lipid diet. Additionally, the 25% lipid diet significantly reduced daily feed intake, compared with 5% and 15% dietary lipid levels, suggesting lipid levels above 15% reduce growth due to a reduction in feeding (Wang et al. 2005). Craig et al. (2006) also observed significantly reduced weight gain in juvenile cobia (49.3 g) fed a diet with the highest lipid level tested in an experiment using a factorial design containing two levels of crude protein (40% and 50%) and three levels of lipid (6%, 12% and 18%). In contrast, in an identical experiment with smaller cobia (7.4 g) there was no significant effect on weight gain caused by the different dietary lipid levels, suggesting varying lipid requirements during different life stages of cobia. Apparently cobia exhibits a lower capacity for protein sparing by dietary lipid, preferring mainly protein as the primary dietary energy source to lipid. Excess lipid levels would appear to reduce growth and fish health and, as fish oil is an expensive product, increase production costs (Craig et al. 2006).

6.5.3

Vitamins and trace elements

Both vitamins and minerals play important roles in normal life processes in fish, and deficiencies in these essential micronutrients lead to chronic pathologies and eventually mortalities. However, with cobia there is very limited information on the roles of micronutrients. Only the role of choline on survival and growth of cobia was recently reported by Mai et al. (2009). It is suggested that dietary choline significantly influenced survival, feeding rate, weight gain, feed efficiency ratio and hepatosomatic index, as well as the choline concentrations in the liver and muscle of cobia. Based on weight gain, liver and muscle choline concentrations, choline requirements were found to be 696, 877 and 950 mg choline kg−1 diet in the form of choline chloride, respectively (Mai et al. 2009). Except for Selenium (Se) requirement, there is no information available on the requirement of trace elements in the diets of cobia. The Se requirements of juvenile cobia were between 0.788 and 0.811 mg Se kg−1 diet in the form of seleno-dl-methionine (Liu et al. 2010). The dietary Se level can influence the survival, growth rate, feed efficiency and the Se concentrations in the whole body and vertebra of cobia juveniles (Liu et al. 2010).

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185

Energy requirements

Temperature plays an important role in energy budget and growth of cobia. Growth and feed conversion efficiency (FCE) were maximised at an intermediate temperature range of 27–29 °C in juvenile cobia. At higher temperatures metabolism impacted on growth as feeding was still active. In contrast, at lower temperatures growth was reduced due to a decrease in feed consumption (Sun et al. 2006). For the larger cobia (330 g), greatest weight gain occurred in fish maintained at 29 °C, compared with 23 °C and 18 °C, during a 70-day feeding trial. Feed conversion ratio values of cobia showed no difference between the fish held at 23 and 29 °C. Fish previously maintained at sub-optimal temperatures (18 °C for 70 days) before temperatures were elevated close to the optimum (1 degree per day to maximum of 29 °C over 54 days), displayed significant increases in specific growth rates (SGRs) compared with fish maintained at 29 °C throughout (1.23 and 1.04 respectively) (Schwarz et al. 2007). It is recommended that the optimal dietary protein to energy ratio for juvenile cobia should be 33.8 mg protein kj-1 in diets containing large amounts of carbohydrates (Webb et al. 2010).

6.5.6

Feed ingredients

The factors to be considered when formulating a feed are: (1) the formulation needs to provide adequate nutrient and energy levels; (2) the final product needs to be palatable; (3) ingredients should not impact negatively on final product quality; and (4) feed needs to be easily manufactured and be cost-efficient (Fraser & Davies 2009). Apparent digestibility is one of the important criteria to evaluate the nutrient value and feed ingredients. Zhou et al. (2004) reported the digestibility of selected feed ingredients for juvenile cobia. Apparent digestibility coefficients (dry matter, crude protein, crude lipid and gross energy) were highest for fishmeal and corn gluten meal. Carbohydrate-rich products produced the lowest digestibility levels and were not recommended for use as a fishmeal replacement in cobia. Poultry and corn meal had higher digestion coefficients than plant meals, indicating a greater potential for these ingredients to become dietary replacements for fishmeal in cobia. Cobia feed contains large quantities of fishmeal and oil as protein and lipid sources. This increases the demand for fishmeal, though juvenile cobia are able to utilise relatively high levels of dextrin (Webb et al. 2010). Thus, recent work has focused on finding alternative protein sources. In cobia, a number of alternative sources have shown potential for use in feeds, including soybean (Chou et al. 2004) and organically certifiable yeast-based protein (Lunger et al. 2006). Chou et al. (2004) conducted a feeding trial to determine the amount of soybean meal that could replace fishmeal in formulated diets without reducing growth. Juvenile cobia (initial mean weight, 32 g) were fed 48% crude protein diets in which dietary protein was supplied by brown fishmeal or a mixture of hexane-extracted soybean meal and fishmeal, resulting in 10%, 20%, 30%, 40%, 50% and 60% of fishmeal protein being replaced by soybean protein. The fish readily accepted all seven experimental diets and no fish died during the trial. Detrimental effects on growth performance were obvious when half of the fishmeal protein was replaced by soybean protein. Significant differences in fish weight gain, feed conversion ratio (FCR), protein efficiency ratio (PER) and net protein utilisation (NPU) were observed when the replacement level for fishmeal protein was increased from 40% to 50%, indicating that up to 40% of fishmeal protein can be replaced by soybean

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meal protein without causing a reduction in growth and protein utilisation. On the other hand, quadratic regression analysis showed a growth optimum at 16.9% replacement of fishmeal protein by soybean meal protein (Chou et al. 2004). These findings were supported by the study of Zhou et al. (2005) to determine the potential use of defatted soybean meal (roasted and solvent-extracted) as a partial replacement for fishmeal in the diet for juvenile cobia. The results indicated that up to 40% of fishmeal protein could be replaced by defatted soybean meal without causing significant reduction in growth. The optimum level of fishmeal protein replacement with defatted soybean meal, determined by quadratic regression analysis, was 18.92% on the basis of maximum weight gain (Zhou et al. 2005). Lunger et al. (2006) reported that cobia fed a diet containing 25% of dietary protein from the yeast-based protein source had equal weight gain and feed conversion ratio values to fish fed the control diet composed of 100% fishmeal. Biological indices including hepatosomatic index, visceral somatic index and muscle ratio were all similarly affected by inclusion of the yeast-based protein source, with significant impacts when inclusion levels rose above 25% of dietary protein. This study suggested a minimum 25% of dietary protein can be provided by yeast-based protein in diets for cobia (Lunger et al. 2006). Further studies by Lunger et al. (2007) suggested that taurine supplementation has a significant impact on growth and feed efficiency of juvenile cobia when they are fed diets containing high levels of plant-based proteins as replacements for fishmeal. Additionally, alternative proteins, especially those of plant and yeast-based origin, can be incorporated at very high levels in diets for cobia with proper amino acid supplementation (Lunger et al. 2007). Cobia juveniles are also capable of digesting chitin from crustacean processing wastes through strong endogenous chitinolytic enzymes (Fines & Holt 2010), increasing the sustainability of cobia culture by reducing inclusion rates of fishmeal and other expensive feed ingredients. The research of Salze et al. (2010) has shown the importance of alternative ingredients such as soy protein concentrates for the replacement of fishmeal and fish oil in formulated diets for cobia. In their two feeding trials, Salze et al. (2010) demonstrated that juvenile cobia can show excellent performance when the fishmeal component of their feed was replaced by up to 75%. Dietary inclusion of mannan oligosaccharides (MOS) did not produce any significant effects; though there was a beneficial trend (Salze et al. 2010). Sun and Chen (2009) described all the quadric relationships between food conversion efficiency and temperature in cobia culture. Energy budgets of juvenile cobia at satiation ration were: 100 C = 7.0 F + 7.7 U + 69.0 R + 16.4 G (or 100 A = 81 R + 19 G) at 33°C 100 C = 6.8 F + 7.9 U + 68.0 R + 17.3 G (or 100 A = 80 R + 20 G) at 27°C and 100 C = 6.3 F + 8.4 U + 77.2 R + 8.2 G (or 100 A = 90 R + 10 G) at 21°C C, where C is food energy, A is assimilated energy, F is faeces energy, U is excretion energy, R is metabolism energy and G is growth energy (Sun & Chen 2009).

6.6 6.6.1

HATCHERY Egg production

The rapid development of cobia aquaculture has focused research on spawning and improved larval rearing techniques (Holt et al. 2007). Researchers in the US have used hormones to

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induce adult cobia caught during their natural spawning season to produce eggs. Both HCG (human chorionic gonadotropin) injected at 275 IU/kg and a slow-release pellet containing salmon GnRHa (gonadotropin-releasing hormone analog) implanted in fish have resulted in spawns (Franks et al. 2001). Arnold et al. (2002)) reported the technique for induced spawning of cobia by ambient seasonal cycles in tanks. The fish were held in a 35,000 L recirculating semi-oval broodstock tank measuring 7.5 m in length, 4.5 m wide, and 1.1 m deep. The broodstock tank was located in an outdoor heated greenhouse constructed of clear fibreglass siding and exposed to ambient photoperiod conditions. Three separate pieces of grey vinyl tarp 0.3 m apart were located 1 m over the tank to reduce the intense direct sunlight during warmer months of the year. The light intensity measured at the water ’s surface on a clear, sunny day ranged from 1,000–2,000 lux, depending on where the reading was taken in the broodstock tank. The sides of the tank were open and a 0.6 m-high plastic mesh netting fence encircled the entire tank on its top edge. Water temperature was maintained during the winter months using submerged heaters in the filter box, and two natural gas heaters were utilised to warm the air in the greenhouse. Two 10.16 cm diameter PVC airlift pipes located in the tank along both of the 7.5 m sides created a clockwise current that the fish were usually observed swimming against. The tank was lined with a 2 mm-thick black plastic liner and water was circulated via airlift at a rate of 70 L/min. At this rate the total volume of the system passed through the external biological filter containing 1 m3 of 5.84 cm Lanpac media (Lantec Products, Inc., Agoura Hills, California, USA) approximately 3 times every 24 hours. Water exchange or addition was <5% of the tank volume per week, temperature ranged from 19–28.5 °C, salinity from 27–34 ppt, NH3 <0.3 mg/L, and the pH was maintained at 7.3–7.8 with the periodic addition of soda ash. The fish were fed once daily at a rate of 2–3% body weight a diet predominantly composed of shrimp by-catch, 95% of which was made up of three fish species: spot Leiostomus xunthurus, pinfish Lugodon rhomboids, and Atlantic croaker Micropogonius undulutus. In addition, smaller portions of squid and shrimp supplemented the fish diet two times a week and made up approximately 20% of the total diet. The spawning of cobia was observed during April and May. Faulk and Holt (2008) showed that amino acid profile was independent of the season. No relationship between egg quality and amino acid content was noted, with the most prominent amino acids being glutamate, leucine, alanine, proline, lysine and aspartate; nor were any differences detected between spawning seasons (Faulk & Holt 2008). In Taiwan, cobia intended for broodstock are obtained from hatcheries and reared in open cages until they attain sexual maturity (about 1.5–2 years when fish weighs about 10 kg). Maturing brooders are selected from sea cages and transferred to land-based spawning ponds (400–600 m2 area; 1.5 m depth) with flow-through seawater, at a density of 100 fish per pond and a sex ratio of about 1:1 (male/female). Fish are fed to satiation with raw fish (e.g. sardines, mackerels, squids) once or twice a day. Brooders spawn spontaneously all year round, with a peak in spring and autumn when water temperature is maintained at 23–27 °C. The fertilised eggs are collected using a seine net installed against the current created by paddlewheels. The eggs are then transferred to outdoor larval rearing ponds (earthen ponds; <5,000 m2 area; 1–1.2 m water depth) with well maintained greenwater (Chlorella sp.) and an abundant number of copepods. Water exchange was minimal or unnecessary in the early stage as long as the greenwater was maintained (Liao et al. 2004). Cobia can also be induced to spawn at the desired time by putting them through simulated seasons at regular intervals of photo-thermal condition. In the typical conditioning cycle, photoperiod ranges from 10 (winter) to 14 (summer) hours of daylight and water

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temperature ranges from 20 to 26 °C). Tank spawning generally begins at 13 to 14 hours of daylight and 25.5 to 27 °C). It will continue for several months if these conditions are maintained (Kaiser & Holt 2005).

6.6.2

Larvae and juvenile rearing

Cobia eggs are 1.20 to 1.40 mm in diameter, heavily pigmented, and hatch in 24 hours at 27–29 °C (Kaiser & Holt 2005). Liao et al. (2004) reported the ‘greenwater ’ method of rearing the cobia larvae. Cobia larvae are transferred to greenwater nursery ponds where they are reared on copepod nauplii and rotifers until day 20 (survival of larvae to day 20 is reported to be 5 to 10%). Then they enter a three-stage nursery pond system. During the first stage, days 20 to 45, the cobia are weaned onto pelleted floating food, size-graded every 4 to 7 days to reduce cannibalism, and grown to 2–5 g. The second stage (days 45 to 75) uses larger ponds (more than 0.07 acres or 300 m2), where the fish are fed to satiation five to six times daily and reared to 30 g. In the final nursery stage (from day 75 to day 150 to 180), the cobia are grown to 600–1,000 g either in large ponds or nearshore cages, depending on the operation (Liao et al. 2004). Production in these extensive systems requires large land areas with little control over the output, but the growth rates are rapid (Holt et al. 2007). In the United States, cobia larval rearing is undertaken in a 300 L rearing tank. The typical larval rearing system is described by Holt et al. (2007). Eggs are usually stocked into rearing tanks at a density of 5 to 10 per litre. Larvae in tanks are fed enriched rotifers (3 to 5 rotifers/ ml) beginning on the third day post-hatch and continuing for a minimum of 4 days. Enriched Artemia preparations are fed from that point until weaning (generally day 25 to 30), after which the cobia are given only dry feed (Kaiser & Holt 2005) (Table 6.1). Other protocols on larviculture were developed at Virginia Seafood Agricultural Research and Extension Center (Virginia, USA) (Holt et al. 2007). The first feeding began at 2 dph, described as follows: L-type rotifers were fed from 2 through 8 dph (coinciding with algal additions) and were enriched for 12 hours with Protein Selco Plus (INVE, Salt Lake City, Utah, USA). AF speciality Artemia (INVE) were fed from 6 through 10 dph, unenriched EG Artemia (INVE) were fed from 8 through 12 dph, and EG enriched Artemia (INVE) were fed from 10 through 25 dph. EG Artemia were enriched for 24 hours with DC DHA Selco (INVE). Cofeeding of larvae with Otohime Marine Larvae Weaning Diets began on 18 dph, weaning procedures began on 22 dph, and all fish were fully weaned and Artemia were discontinued by 25 dph. On 28 dph all surviving animals were weighed and counted. From 28 dph onwards no further mortalities were recorded. The average fingerling weight at 35 dph was 1.77 g (Holt et al. 2007). Table 6.1 Summary of larval cobia feeding regime used at the University of Texas Fisheries and Mariculture Laboratory. Days post-hatch (dph)

Feeding regime

3–9 6–26 16 onwards

Rotifers as first feed Artemia as second feed Weaning to dry feed

Note: Both rotifers and Artemia are enriched with a commercially available preparation. Adapted from: Kaiser & Holt (2005)

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In intensive systems growth and in some cases survival of larvae, can be affected by density-dependent processes. Cobia growth and survival were negatively correlated with increasing density (1–20 larvae/L) in 300 L tanks with temperature and photoperiod control (Hitzfelder et al. 2006). However, survival and growth performance of cobia juvenile (6.7 ± 0.2 g) were not affected by the stocking density in the range of 0.04 to 0.44 g of fish per litre (Webb et al. 2007). Culture of cobia juveniles may be practical in salinities as low as 5 ppt (Resley et al. 2006). Benetti et al. (2008) developed protocols for larviculture of cobia by incorporating the use of probiotics and prophylaxis, minimising microalgae use, and including commercially available ingredients for live feed enrichment. Their trials generated 125,328 fingerlings in four tanks in just two months; levels of production that could sustain a commercial operation and indicate that cobia aquaculture could be viable in the Americas.

6.6.3

Nutrient and diets for larvae rearing

Cobia larvae require high level of lipid and PUFA content in the diets. This can be satisfied by enriching rotifers with a product high in ARA as well as in DHA and EPA (Holt et al. 2007). Rotifers enriched with Aquagrow Advantage + Aquagrow arachidonic acid (Advanced BioNutrition Corporation, Columbia, Maryland, USA) have such a profile. The addition of an ARA supplement to the Algamac 2000 (Aquafauna Bio-Marine, Hawthorne, California, USA) enrichment increased the level of ARA in live prey from 2.4 to 3.9% for rotifers and 2.4 to 3.7% Artemia but did not increase the amount of ARA in the whole body tissues of day 7 and 16 cobia larvae, nor were any differences in growth and survival detected (Faulk & Holt 2005). Enriching Artemia with Aquagrow or Algamac products increased the levels of ARA and DHA in Artemia. The incorporation of DHA from enriched Artemia into larval tissues was positively correlated with levels measured in the dietary lipids, indicating preferential retention of this fatty acid. As the amount of DHA increased in the live prey, a corresponding increase was evident in the larval tissue (Holt et al. 2007). Rotifers and Artemia enriched with Isochrysis galbana or commercial products (Algamac 2000 and Aquagrow) in conjunction with greenwater culture provided the best growth and survival of cobia larvae in recirculating aquaculture systems. Survival of cobia larvae was significantly improved from 12 to 25% in 16 dph larvae by the addition of live algae (I. galbana or Nannochloris oculata) to the rearing tanks (Faulk & Holt 2005). The possibility of using umbrella-stage Artemia also presents an opportunity to simplify the rearing protocol and to reduce the production costs of cobia larviculture (Nhu et al. 2009). While using the smaller-sized Vietnam Artemia franciscana (AF) strain instead of the Great Salt Lake A. franciscana strain, Nhu et al. (2009) showed that the rotifer-feeding period could be shortened by 3 days, resulting in significant improvements in larval survival and growth.

6.7

GROWOUT

Benetti et al. (2010) for the first time demonstrated that cobia culture is technically feasible in submerged open ocean cages (Fig.6.4a and b). Cobia grew to a mean weight of 6.035 kg 2.10%/day of specific growth rate at the Puerto Rico site and 3.545 kg in 346 days, equivalent to 2.04%/day of specific growth rate in the Bahamas (Benetti et al. 2010). The

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(a)

(b) Fig. 6.4

(a) and (b) Cobia cage culture in the ocean.

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length–weight relationships at these two sites demonstrated that the cultured cobia can exhibit greater condition factors than their wild counterparts. The results have also shown that the growth rates of cobia were inversely related to temperature and increasing stocking densities.

6.8

DISEASE AND HEALTH MANAGEMENT

Parasites and other pathogens in cobia culture have been reviewed by McLean et al. (2008). The main disease-causing organisms in cobia cultures are classified as crustacean parasites, metazoan parasites, mysosporidia parasites, protozoan parasites, bacterial pathogen and viruses. An increase in the number of cage farmers, due to successful production of cobia, has resulted in a rise in the incidence of disease. For example, in 2002 total cobia production in Taiwan fell to 1,000 tons from 3,000 tons in 2001 because of numerous disease outbreaks (Liao et al. 2004). Therefore, managing disease and parasite issues has been identified as one of the major challenges in cobia culture (FAO 2007). Chen et al. (2001) have identified some diseases and parasites outbreaks during different rearing stages of cobia.

6.8.1

Crustacean parasites

Epizootics of parasitic crustaceans can result in serious economic losses during cobia culture. Ten crustacean parasitic species are reported to infect cobia (McLean et al. 2008). The most common of them are sea lice Caligus lalandei (Chang & Wang 2000), C. epidemicus (Ho et al. 2004) and Parapetalus occidentalis (Ho & Lin 2001). In Puerto Rico, wild cobia were reported to be infected by Tuxophorus caligodes, Euryphorus nordmanni, L. longiventris, L. hemiramphi, and C. haemulonis (Bunkley-Williams & Williams 2006). Parasitic crustaceans often possess attachment organs but others may move freely on the host’s surface, causing widespread necrosis and disruption of the protective mucus covering of the skin at anchor points. The size and age of the host, host health status, the species of parasite, and developmental stages present, affect the severity of the disease. Severe wounding can result in cobia death due to osmotic imbalance or by providing entry points for secondary pathogens. Economic losses incurred other than direct mortality of farmed cobia are generally due to reduced growth of infected fish, negative impacts on edible tissues and costs associated with treatments (McLean et al. 2008).

6.8.2

Metazoan parasites

Digenea parasite infections on wild adult cobia caught from the Gulf of Tonkin and South China Sea, Vietnam, are reported by Arthur and Te (2006). These parasites are Aponurus carangis, Bucephalus varicus, Derogenes varicus, Dinurus selari, Lepidapedon megalaspi, Neometanematobothrioides rachycentri, Paracryptogonimus morosovi, Phyllodistomum parukhini, Stephanostomum imparispine, Tormopsolus filiformis and Tubulovesicula angusticauda. Mclean et al. (2008) compiled a list of other digenea parasites on cobia, which include Tormopsolus spatulum, Pseudolepidapedon pudens, Lecithochirium monticellii, Stephanostomum dentatum, S. cloacum S. pseudoditrematis, S. microsomum, S. rachycentronis, Mabiarama prevesiculata, Plerurus digitatus, Sclerodistomum rachycentri and S. cobia.

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Monogenetic trematodes, more usually referred to as flukes or flatworms, parasitise the skin, gills and fins of cobia. Some species may be found in the gut, body cavity and even blood vascular system. They attach to the fish via a variety of suckers and hooks located at their posterior end (opisthaptor) while using the anterior end or prohaptor for feeding and to assist movement (McLean et al. 2008). In wild cobia, Dionchus rachycentris and D. agassizi which infect the gills of fishes have been described (Bullard et al. 2000). Neobenedenia girellae causes extensive necrosis of the dorsal head region which, together with Streptococcus infection, ultimately causes blindness in infected animals (Chiau et al. 2004). Liao et al. (2004) reported the infection of Neobenedenia melleni on cultured cobia in Taiwan. Cobia have also been reported with infections of tapeworm including Nybelinia bisulcata, Callitetrarhynchus gracilis, Rhinebothrium flexile, Rhynchobothrium longispine and Trypanorhyncha sp. Tapeworms can reduce growth rates and have a negative impact on feed efficiencies in captive cobia (McLean et al. 2008). In a similar way to tapeworms, nematodes (roundworms) can reduce reproductive performance and have a negative impact on feed conversion efficiencies, leading to reduced growth and overall performance of cobia. Economic loss can also occur due to the burrowing of worms into various tissues including musculature (McLean et al. 2008). BunkleyWilliams and Williams (2006) reported two nematodes, Mabiarama prevesiculata, from the stomach of cobia in Brazil and Goezia pelagia, which was isolated from fish in the Gulf of Mexico. Wild cobia from Vietnam has also been reported to be infected by Anisakis sp. and Philometroides sp. (Arthur & Te 2006).

6.8.3

Myxosporidia

Myxosporidian parasites are one of the major causes for concern during cobia aquaculture (McLean et al. 2008). Cumulative mortality of 90% in cultured cobia was reported within 1 month of the infection. Cobia of 45 g exhibited anaemia and ascites, and mottled red and grey, extremely enlarged kidney with cream-coloured patches or spherical nodules. Extrasporogonic or sporogonic stages of a myxosporean appeared in the blood, glomerulus, renal tubules and renal interstitium. Many sporogonic stages are attached to the brush border of the epithelium of the renal tubules (Chen et al. 2001).

6.8.4

Protozoan parasites

Ciliophoran parasites were generally reported during the hatchery phase, while Trichodina have been recorded during the nursery phase of cobia (FAO 2007). The parasite, isolated from the skin and gills of infected fish, was associated with lethargy and inappetance, but the low level of mortality was coupled with secondary infections. Cobia also succumb to the prostomatean Cryptocaryon irritans which causes marine white spot (McLean et al. 2008). Cryptocaryon and Brooklynella hostilis was reported to cause the massive death of 30,000 stressed juvenile cobia in Puerto Rico (Bunkley-Williams & Williams 2006). Brooklynellosis, more commonly referred to as slime-blotch or clownfish disease, is characterised by body discolouration, lethargy, inappetance, increased mucus production and coughing where the gills are infected. Brooklynella reproduce by binary fission so that infections can rapidly build and spread in enclosed environments such as recirculating systems. Often, Brooklynellosis is associated with other opportunistic diseases (McLean et al. 2008).

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The parasitic flagellate Ichthyobodo infests gills and skin of its host and has become a serious issue during mariculture operations, especially when undertaken in tanks (McLean et al. 2008). This parasite was isolated from cobia in a commercial aquaculture operation in Puerto Rico (Bunkley-Williams & Williams 2006). The stalked colonial ciliate Epistylis has also been reported to grow on larval cobia (McLean et al. 2008). This parasite appears as a fluffy grey/white or brown/red-coloured growth on the skin and fins of infected cobia and can be isolated from the mouth and gills of the fish. Severe infestations with Epistylis are often associated with secondary infections and are indicative of poor water quality and high organic loads. Several methods are available to combat fish infected with external protozoan parasites and these usually involve some form of bath (formalin- or copper-based) (FAO 2007). Experimental vaccines employing attenuated strains of parasitic protozoa or recombinant antigenic surface proteins have been developed and may prove useful for cobia aquaculture in the future (McLean et al. 2008). Kaiser and Holt (2005) reported the infection of cobia by dinoflagellates such as Amyloodinium ocellatum. This parasite causes hyperplasia, inflammation and necrosis at infected sites and disrupts gas exchange in the gills of juvenile cobia. Skin infestations cause fish to flash and chafe, whereas gill clogging triggers a coughing response. Severe infestations of Amyloodinium may trigger rapid mortality.

6.8.5

Bacterial pathogens

Bacterial disease caused by Photobacterium sp. has been considered as a major problem for cobia throughout the production cycle. Photobacteriosis in caged cobia has resulted in 80% mortalities at some sites (McLean et al. 2008). Liao et al. (2004) encountered outbreaks of vibriosis, mycobacteriosis, furunculosis and streptococcosis in cobia culture in Taiwan. Cobia from all stages of the production cycle may succumb to vibriosis. Several species of Vibrio have been isolated from moribund farmed cobia including V. alginolyticus (Rajan et al. 2001), V. harveyi (Liu et al. 2004), V. parahaemolyticus and V. vulnificus (Lopez et al. 2002) and vibriosis has accounted for 45% mortalities in cage-stocked juvenile cobia (McLean et al. 2008). Multiple pathogen including Aeromonas hydrophila, Citrobacter sp. and Mycobacterium marinum were also reported to infect cultured cobia (Lowry & Smith 2006). The clinical signs of vibriosis disease in cobia widely vary from no external indications through to a darkening of the skin, lethargy, inappetance, exophthalmia, swollen abdomen, pale gill colour, erosion and hemorrhage in the fins and skin lesioning. Internally, ascites may be present in the peritoneal cavity and the host liver and kidney may pale in colour while the spleen may have white tubercules present. For photobacteriosis, the signals include skin ulceration and a buildup of whitish granulomatous tissue on the kidney, liver and spleen (McLean et al. 2008). Restriction or ban of the antibiotics used as additives in fish feed has prompted interest in developing alternative strategies for health promotion and disease control. Thus, in recent years, there has been considerable research on dietary supplementation in which various health-promoting compounds have been studied (Gatlin et al. 2006). In cobia culture, dietary inclusion of some immunistimulants to improve the resistance to bacterial infection has been reported. Incorporation of beta 1,3–1,6-glucan at 0.5% of the diet was found to enhance resistance of cobia to P. damselae and Streptococcus iniae challenge (Chang et al. 2006). Leaño et al. (2003) concluded that levamisole inclusion at 500–1,000 mg kg−1 diet

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reduced the virulence of P. damselae, apparently by enhancing the leukocyte response of 12–25 g juveniles. It is likely that the intestine represents a major portal of entry for this pathogen. Therefore, it is possible that benefits may accrue from using dietary mannan oligosaccharides since these complex carbohydrates appear to assist in maintaining and enhancing the barrier integrity of the cobia gut (Salze et al. 2008).

6.8.6

Viral disease

There is very limited information regarding virus infection in cobia. Chi et al. (2003) reported the presence of beta-nodavirus NNV (nervous necrosis virus) causing death of cobia.

6.9

POST-HARVEST AND MARKETING

It is a common practice to starve fish one day before harvesting (FAO 2007). The fish are killed, bled and chilled and subsequently packed in layers of ice in insulated boxes as a whole or fillets. Most of the fresh cobia should reach market within 24 to 48 hours of harvest. The flesh of cobia is high in nutritional value; particularly rich in vitamin E and DHA. The nutritional profile of cobia is shown in Fig. 6.5. Cobia enjoys a relatively high market value compared to other finfish in Taiwan Province of China. Larger cobia (8–10 kg) are sold whole there, while Japan is the primary destination for smaller (6–8 kg) fish which are sold both whole and headless (some for sashimi), with fillet product typically exported to other markets. The prices for cobia vary according to size; the market value in Taiwan for whole fish of 7.7 kg and larger was approximately US$5.50/kg in 2004 and reported to be less than that for other smaller fish. Cobia exported to Japan for sashimi markets are whole/gutted or headless (Liao et al. 2004). The Taiwanese market accepts a diverse product range of fresh and frozen cobia, which includes whole round cobia, gilled and gutted cobia, skin-on cobia fillets, fresh or frozen skinless cobia fillets, cobia heads, cobia fish liver and cut cobia fish bone (Council of Agriculture Taiwan n.d.). 400

363

350 mg/100 g

300 250 186

200 150 100 50

70

54 2.58

0.21

3.2

Vitamin E

Niacin

0 Cholesterol Vitamin B12

Natrium

Potassium

Phosphate

Fig. 6.5 Nutritional profile of cobia (Adapted from: http://www.agexporter.com.tw/eng_web/flower_ view.jsp?cid=70032).

Cobia Culture

6.10

195

CHALLENGES AND OPPORTUNITIES

The expansion of cobia culture is still hindered by the lack of a regular supply of seed, as larval nutrition and the impact of formulated diets on broodstock condition and subsequent larval health are still not well understood. Further, the spawning behaviour of cobia and environmental clues which trigger the spawning of cobia are very much site-specific and need further research in order to ensure a constant supply of viable cobia fingerlings from the hatchery. The survival of 20-day-old larvae is below commercially acceptable levels due to the lack of understanding of broodstock management and nutrition requirements of larvae. Stress during transport from nursery tanks/inshore cages to growout cages and diseases during the nursery phase are other areas that need further research. As pressure for the intensification of cobia culture increases there will be a need for further research on the use of commercially viable dietary immunostimulants to improve immunocompetence under stressful, intense farming conditions. Depending on pond management, culture density, feed rate and water exchange rates, the aquaculture production of cobia in ponds may result in water quality issues and excess nutrient loading in the effluent. Large-scale production of cobia at various life stages could potentially impact many coastal areas where such activities are located and production discharge would require monitoring. Cobia growout thus far has been reported in nearshore and offshore cage systems, which will also have some form of environmental impact. These operations carry with them some inherent risks including, but not limited to, escapees (genetic pollution), disease transmission, and nutrient loading in and around farm sites. Since 1998, when significant aquaculture production of cobia was first reported, it has been noted in Taiwan Province of China that while cobia production is successful and expanding there, diseases affecting the fish in culture are a major issue and best management practices must be implemented wherever possible. Because of the annual threat of typhoons, some farmers there have located their growout systems in more enclosed, lower energy areas, at the expense of higher water flow and flushing rates. This intensification of nearshore production has been reported to result in increased disease outbreaks and, in some cases, lower quality flesh from cobia raised in cage systems from more polluted areas. Expansion of cobia cage culture in areas such as the Caribbean and Central America where cage sites may be close to sensitive coral reef would also require producers to monitor impacts and the nutrient loading of the nearby ecosystem. Cage culture environmental impacts are site-specific and, while water exchange rates and dilution factors in many areas are sufficient to prevent excess nutrient loading, every site has a carrying capacity that needs to be considered. Another environmental consideration is that since cobia are higher trophic level carnivores, it is necessary to use feed containing fairly high crude protein levels (the specific nutritional requirements are currently being investigated) with a certain portion of this protein obtained from fishmeal. For aquaculture to continue supplying the increasing demand for seafood worldwide, additional research into supplemental or alternative protein sources for use in feeds for species such as cobia is imperative. To that end, research on the specific nutritional requirements of this species in terms of protein and lipid levels and fishmeal substitution in feed has been conducted and is ongoing. Improvements during the nursery phase of cobia production are critical if culturists are going to be able to supply the massive numbers of juveniles that would be required to stock large, commercial cage operations. Towards that goal, US and Taiwanese researchers are

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working on the development of intensive and super-intensive recirculating systems for juvenile production, utilising temperature control, UV sterilisation, drum filters, protein skimmers and oxygenation units. So far, a Taiwanese high-density juvenile production system has been quite successful, with optimum production and survival rates for cobia grown from 7 to 76 g in four weeks at initial stocking densities of 370 fish/m3 and final harvest values reported to be 28 kg/m3. As cobia aquaculture expands in the future, the efficient production of a high quality product with minimal environmental impact during culture should be the desired goal. The future global supply, market, and pricing of cobia will certainly be affected by the future production (or lack of) cobia in China and Taiwan as well as in other countries in Southeast Asia.

6.11 REFERENCES Arnold, C.R., Kaiser, J.B. & Holt, G.J. (2002) Spawning of cobia Rachycentron canadurn in captivity. Journal of World Aquaculture Society, 33, 205–208. Arthur, J.R., & Te, B.Q. (2006) Checklist of the parasites of fishes of Viet Nam. FAO Fisheries Technical Paper No. 369/2. Rome. Benetti, D.D., Sardenberg, B., Welch, A., Hoenig, R., Orhun, M.R. & Zink, I. (2008) Intensive larval husbandry and fingerling production of cobia Rachycentron canadum. Aquaculture, 281, 22–27. Benetti, D.D., O’Hanlon, B., Rivera, J.A., Welch, A.W., Maxey, C. & Orhun, M.R. (2010) Growth rates of cobia (Rachycentron canadum) cultured in open ocean submerged cages in the Caribbean. Aquaculture, 302, 195–201. Briggs, J.C. (1960) Fishes of worldwide (circumtropical) distribution. Copeia, 3, 171– 180. Brown-Peterson, N.J., Overstreet, R.M., Lotz, J.M., Franks, J.S. & Burns, K.M. (2001) Reproductive biology of cobia, Rachycentron canadum, from coastal waters of the southern United States. Fisheries Bulletin, 99, 15–28. Bullard, S.A., Benz, G.W. & Braswell, J.S. (2000) Dionchus postoncomiracidia (Monogenea: Dionchidae) from the skin of blacktip sharks, Carcharhinus limbatus (Carcharhinidae). Journal of Parasitology, 86, 245–250. Bunkley-Williams, L. & Williams, E.H.J. (2006) New records of parasites for culture cobia, Rachycentron canadum (Perciformes: Rachycentridae) in Puerto Rico. Revista de Biologia Tropical, 54 (Suppl. 3), 1–7. Chang, C.F., Yang, J.H. & Chang, S.L. (2006) Application of dietary beta-1,3–1, 6–glucan in enhancing resistance of cobia (Rachycentron canadum) against Photbacterium damselae subsp. Piscicida and Streptococcus iniae infections. Journal of Taiwan Fisheries Research, 14, 75–87. Chang, P., & Wang, Y. (2000). Studies on the Caligusiasis and Benedeniasis of marine cage cultured fish in Pingtung area of Taiwan. Proceedings of the First International Symposium on Cage Aquaculture in Asia, Asian Fisheries Society, Manila, Philippines. Chen, S.C., Kou, R.J., Wu, C.T., Wang, P.C. & Su, F.Z. (2001) Mass mortality associated with a Sphaerosporalike myxosporidean infestation in juvenile cobia, Rachycentron canadum (L.), marine cage cultured in Taiwan. Journal of Fish Diseases, 24, 189–195. Chi, S.C., Shieh, J.R. & Lin, S. (2003) Genetic and antigenic analysis of betanodaviruses isolated from aquatic organisms in Taiwan. Diseases of Aquatic Organisms, 55, 221–228. Chiau, W.Y., Chou, C.L. & Shih, Y.C. (2004) Marine aquaculture in Chinese Taipei: Status, institutions and challenges. APEC Bulletin on Marine Resource Conservation and Fisheries, 6, 15–20. Chou, R.L., Su, M.S. & Chen, H.Y. (2001) Optimal dietary protein and lipid levels for juvenile cobia (Rachycentron canadum). Aquaculture, 193, 81–89. Chou, R.L., Her, B.Y., Su, M.S., Wang, G.H., Wu, Y.H. & Chen, H.Y. (2004) Substituting fish meal with soybean meal in diets of juvenile cobia Rachycentron canadum. Aquaculture, 229, 325–333. Craig, S.R., Schwarz, M.H. & Mclean, E. (2006) Juvenile cobia (Rachycentron canadum) can utilize a wide range of protein and lipid levels without impacts on production characteristics. Aquaculture, 261, 384–391.

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FAO (2007) Cultured aquatic species information program: Rachycentron canadum. http: //www. fao. org/ fi/website/FI Faulk, C.K. & Holt, G.J. (2003) Lipid nutrition and feeding of cobia (Rachycentron canadum) larvae. Journal of World Aquaculture Society, 34, 368–378. Faulk, C.K. & Holt, G.J. (2005) Advances in rearing cobia Rachycentron canadum larvae in recirculating aquaculture systems: live prey enrichment and greenwater culture. Aquaculture, 249, 231–243. Faulk, C.K. & Holt, G.J. (2008) Biochemical composition and quality of captive-spawned cobia Rachycentron canadum eggs. Aquaculture, 279, 70–76. Fines, B.C. & Holt, G.J. (2010) Chitinase and apparent digestibility of chitin in the digestive tract of juvenile cobia, Rachycentron canadum. Aquaculture, 303, 34–39. Franks, J.S., Warren, J.R. & Buchanan, M.V. (1999) Age and growth of cobia, Rachycentron canadum, from the northeastern Gulf of Mexico. Fisheries Bulletin, 97, 459–471. Franks, J.S., Ogle, J.T., Lotz, J.M., Nicholson, L.C., Barnes, D.N. & Larson, K.M. (2001) Spontaneous spawning of cobia, Rachycentron canadum, induced by human chorionic gonadotropin (HCG), with comments on fertilization, hatching, and larval development. Proceedings Carribbean Fisheries International, 52, 598–609. Fraser, T.W.K. & Davies, S.J. (2009) Nutritional requirements of cobia, Rachycentron canadum (Linnaeus): a review. Aquaculture Research, 40, 1219–1234. Gatlin, D.M., Li, P., Wang, X., Burr, J.S., Castille, F. & Lawrence, A.L. (2006) Potential application of prebiotics in aquaculture. Symposium Internacional de Nutricion Acucola. Mexico. Hammond, D.L. (2001) Status of the South Carolina fisheries for cobia. Science Marine Research, 83, 12. Hitzfelder, G.M., Holt, G.J., Fox, J.M. & Mckee, D.A. (2006) The effect of rearing density on growth and survival of cobia, Rachycentron canadum, larvae in a closed recirculating aquaculture system. Journal of World Aquaculture Society, 37, 204–209. Ho, J.S. & Lin, C.L. (2001) Parapetalus occidentalis Wilson (Copepoda, Caligidae) parasitic on both wild and farmed cobia (Rachycentron canadum) in Taiwan. Journal of the Fisheries Society of Taiwan, 28, 305–316. Ho, J., Kim, I., Cruz-Lacierda, E.R. & Nagasawa, K. (2004) Sea lice (Copepoda, Caligidae) parasitic on marine cultured and wild fishes of the Philippines. Journal of the Fisheries Society of Taiwan, 31, 235–249. Holt, G.J., Faulk, C.K. & Schwarz, M.H. (2007) A review of the larviculture of cobia Rachycentron canadum, a warm water marine fish. Aquaculture, 268, 181–187. Kaiser, J.B. & Holt, G.J. (2005) Species profile Cobia. SRAC Publication 7202. Leaño, E.M., Chang, S.L., Guo, J.J., Chang, S.L. & Liao, I.C. (2003) Levamisole enhances non-specific immune response of cobia, Rachycentron canadum, fingerlings. Journal of the Fisheries Society of Taiwan, 30, 321–330. Liao, I.C., Huang, T.-S., Tsai, W.-S., Hsueh, C.-M., Chang, S.-L. & Leaño, E.M. (2004) Cobia culture in Taiwan: current status and problems. Aquaculture, 237, 155–165. Liu, K., Wang, X.J., Ai, Q., Mai, K. & Zhang, W. (2010) Dietary selenium requirement for juvenile cobia, Rachycentron canadum L. Aquaculture Research, doi: 10.1111/j.1365–2109.2010.02562.x Liu, P., Lin, J., Chuang, W. & Lee, K. (2004) Isolation and characterization of pathogenic Vibrio harveyi (V. carchariae) from the farmed marine cobia fish Rachycentron canadum L. with gastroenteritis syndrome. World Journal of Microbiology & Biotechnology, 20, 495–499. Lopez, C., Rajan, P.R., Lin, J.H., Kuo, T. & Yang, H. (2002) Disease outbreak in seafarmed cobia (Rachycentron canadum) associated with Vibrio spp., Photobacterium damselae ssp. piscicida, mongenean and myxosporean parasites. Bulletin of the European Association of Fish Pathologists, 22, 206–211. Lowry, T. & Smith, S.A. (2006) Mycobacterium sp. infection in cultured cobia (Rachycentron canadum). Bulletin of the European Association of Fish Pathologists, 26, 87–92. Lotz, J.M., Overstreet, R.M. & Franks, J.S. (1996) Gonadal maturation in the cobia, Rachycentron canadum, from the Northcentral Gulf of Mexico. Gulf Research Reports, 9, 147–159. Lunger, A.N., Craig, S.R. & Mclean, E. (2006) Replacement of fish meal in cobia (Rachycentron canadum) diets using an organically certified protein. Aquaculture, 257, 393–399. Lunger, A.N., Mclean, E., Gaylord, T.G., Huhn, D. & Craig, S.R. (2007) Taurine supplementation to alternative dietary proteins used in fish meal replacement enhances growth of juvenile cobia (Rachycentron canadum). Aquaculture, 271, 401–410. Mai, K., Xiao, L., Ai, Q., et al. (2009) Dietary choline requirement for juvenile cobia, Rachycentron canadum. Aquaculture, 289, 124–128.

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McLean, E., Salze, G. & Craig, S.R. (2008) Parasites, diseases and deformitties of cobia. Ribarstvo, 66, 1–16. Meyer, G.H. & Franks, J.S. (1996) Food of cobia, Rachycentron canadum from the Northcentral Gulf of Mexico. Fisheries Bulletin, 9, 161–167. Nhu, V.C., Dierckens, K., Nguyen, T.H., Tran, M.T. & Sorgeloos, P. (2009) Can umbrella-stage Artemia franciscana substitute enriched rotifers for Cobia (Rachycentron canadum) fish larvae? Aquaculture, 289, 64–69. Rajan, P.R., Lopez, C., Lin, J.H. & Yang, H. (2001) Vibrio alginolyticus infection in cobia (Rachycentron canadum) cultured in Taiwan. Bulletin of the European Association of Fish Pathologists, 21, 228–234. Resley, M.J., Webb Jr., K.A. & Holt, G.J. (2006) Growth and survival of juvenile cobia, Rachycentron canadum, at different salinities in a recirculating aquaculture system. Aquaculture, 253, 398–407. Salze, G., McLean, E., Schwarz, M.H. & Craig, S.R. (2008) Dietary mannan oligosaccharide enhances salinity tolerance and gut development of larval cobia. Aquaculture, 274 (1), 148–52. Salze, G., Mclean, E., Battle, P.R., Schwarz, M.H. & Craig, S.R. (2010) Use of soy protein concentrate and novel ingredients in the total elimination of fish meal and fish oil in diets for juvenile cobia, Rachycentron canadum. Aquaculture, 298, 294–299. Sargent, J., McEvoy, L., Estevez, A. et al. (1999) Lipid nutrition of marine fish during early development: current status and future directions. Aquaculture, 179, 217–229. Schwarz, M.H., Mowry, D., Mclean, E. & Craig, S.R. (2007) Performance of advanced juvenile cobia, Rachycentron canadum, reared under different thermal regimes: evidence for compensatory growth and a method for cold banking. Journal of Applied Aquaculture, 19, 71–84. Shaffer, R.V. & Nakamura, E.L. (1989) Synopsis of biological data on the cobia, Rachycentron canadum. (Pisces: Rachycentridae). FAO Fisheries Synopsis 153 (NMFS/S 153). US Dep. Commer. NOAA Tech. Rep., 82, 21. Smith, J.W. (1995) Life history of cobia, Rachycentron canadum, (Osteichthyes: Rachycentridae), in North Carolina waters. Brimleyana, 23, 1–23. Sun, L. & Chen, H. (2009) Effects of ration and temperature on growth, fecal production, nitrogenous excretion and energy budget of juvenile cobia (Rachycentron canadum). Aquaculture, 292, 197–206. Sun, L.H., Chen, H.R. & Huang, L.M. (2006) Effect of temperature on growth and energy budget of juvenile cobia (Rachycentron canadum). Aquaculture, 261, 872–878. Valinassab, T., Ashtari, S., Sedghi, N. & Daghoghi, B. (2008) Reproductive biology of Rachycentron canadum in the Persian Gulf (Hormozgan Province waters). Iranian Scientific Fisheries Journal, 17, 143–152. Van der Velde, T.D., Griffiths, S.P. & Fry, G.C. (2010) Reproductive biology of the commercially and recreationally important cobia Rachycentron canadum in northeastern Australia. Fisheries Science, 76, 33–43. Wang, J.T., Liu, J.L., Tian, L.X. et al. (2005) Effect of dietary lipid level on growth performance, lipid deposition, hepatic lipogenesis in juvenile cobia (Rachycentron canadum). Aquaculture, 249, 439–447. Webb Jr., K.A., Hitzfelder, G.M., Faulk, C.K. & Holt, G.J. (2007) Growth of juvenile cobia, Rachycentron canadum, at three different densities in a recirculating aquaculture system. Aquaculture, 264, 223–227. Webb Jr., K.A., Rawlinson, L.T. & Holt, G.J. (2010) Effects of dietary starches and the protein to energy ratio on growth and feed efficiency of juvenile cobia, Rachycentron canadum. Aquaculture Nutrition, 16, 447–456. Wilson, R.P. (2002) Amino acids and proteins. In: Fish Nutrition (eds J. Halver & R. Hardy). Academic Press, San Diego. Zhou, Q.C., Tan, B.P., Mai, K.S. & Liu, Y.H. (2004) Apparent digestibility of selected feed ingredients for juvenile cobia Rachycentron canadum. Aquaculture, 241, 441–451. Zhou, Q.C., Mai, K.S., Tan, B.P. & Liu, Y.J. (2005) Partial replacement of fishmeal by soybean meal in diets for juvenile cobia (Rachycentron canadum). Aquaculture Nutrition, 11, 175–182. Zhou, Q.C., Wu, Z.H., Tan, B.P., Chi, S.Y. & Yang, Q.H. (2006) Optimal dietary methionine requirement for juvenile cobia (Rachycentron canadum). Aquaculture, 258, 551–557. Zhou, Q.-C., Wu, Z.H., Chi, S.Y. & Yang, Q.H. (2007) Dietary lysine requirement of juvenile cobia (Rachycentron canadum). Aquaculture, 273, 634–640.

7

Barramundi Aquaculture

Suresh Job

7.1 INTRODUCTION Barramundi (Lates calcarifer) are widely distributed throughout the Indo-West Pacific region from the Arabian Gulf to southern China, the Philippines, Indonesia, Papua New Guinea and northern Australia (Nelson 1994). They are more commonly known as Asian seabass or giant seaperch in countries outside of Australia. Barramundi were originally classified as belonging to the family Centropomidae, subfamily Latinae, but have since been re-classified as belonging to the family Latidae (Nelson 1994). Aquaculture production of barramundi began in Thailand in the 1970s (Yue et al. 2009), and has since spread to other countries in Southeast Asia and elsewhere. They are now an important aquaculture species in much of Southeast Asia and Australia (Tucker et al. 2002). Global aquaculture production of barramundi from 1998 to 2008 (FAO 2010) is shown in Table 7.1. Barramundi are farmed in a range of countries including Indonesia, Malaysia, Singapore, Philippines, Brunei, Hong Kong, PR China, Thailand, Taiwan, Saudi Arabia, Australia and the USA (Boonyaratpalin 1997; Chou & Lee 1997; Frost et al. 2006; Zhu et al. 2006, Yue et al. 2009). Barramundi are socially and economically important in recreational and commercial fisheries, and in aquaculture. In Australia, for example, they are the highest-ranked sport fishing species in northern Australia (McDougall 2004), and an estimated 300 tons were harvested by the recreational fishing sector in 1999 in Queensland alone (Williams 2002). Barramundi aquaculture is growing rapidly in countries such as Australia and the USA. In Australia, for example, barramundi aquaculture began in the mid-1980s (Palmer et al. 2007). Barramundi aquaculture production has since increased from 980 tons in 2000/2001 to 2,075 tons in 2005/2006 and 2,996 tons in 2007/2008 and the value of production increased from $9.2 million to about $34 million over that time period (ABARE 2009). Barramundi are an ideal tropical finfish species for aquaculture in many ways. They are hardy, tolerate crowded conditions and handling, and thrive in a wide range of physiological and environmental conditions including high turbidity, and varying salinities and

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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Recent Advances and New Species in Aquaculture Table 7.1 Global barramundi production from 1998–2008. (Based on FAO (2010) data for giant seaperch (Lates calcarifer)). Year 1998 1999 2000 2001 2002 2003 2004 2005 2006 2007 2008

Value (US$000) 78,000 81,000 72,000 87,000 63,000 98,000 78,000 82,000 88,000 118,000 158,000

Quantity (tons) 21,000 23,000 21,000 27,000 26,000 28,000 30,000 32,000 33,000 34,000 45,000

temperatures (Boonyaratpalin 1997; Rajaguru 2002; Yue et al. 2009). In addition, they are fast growing, with a growth rate of approximately 1 kg/year, can reach a marketable size (350 g–5 kg) in 6–24 months (depending on conditions), have a desirable white flesh, and command a relatively high market price (Boonyaratpalin 1997; Yue et al. 2009). They are also highly fecund, with relatively well established hatchery production and nursery protocols.

7.2 7.2.1

BIOLOGY Life history strategy

Barramundi are tropical euryhaline fish that can tolerate a wide range of salinity levels from fresh to seawater, and inhabit a wide range of habitats, from freshwater to brackish and marine systems. Barramundi are found in a number of habitats including coastal regions, estuaries, landlocked freshwater billabongs and rivers (Davis 1984a,b; Guiguen et al. 1994; Balston 2009). They display a catadromous life history strategy, where the adults breed in brackish or salt water (salinities between 22 and 40 ‰). As a rule, breeding takes place in river mouths and bays near areas of suitable nursery habitat (Guiguen et al. 1994). Larval barramundi develop in coastal waters and remain there until they reach approximately 5 mm in length (Moore & Reynolds 1982; Almendras et al. 1988). They then move into coastal swamps or similar habitats until they reach approximately 20 cm in length and 6 months of age (Almendras et al. 1988). Areas such as mangrove swamps and low-lying land that becomes flooded during spring tides and monsoonal rains provide ideal habitat for juvenile barramundi (Schipp 1996). Where rivers are not readily accessible, they tend to remain in the estuaries. They then transition into the freshwater sections of rivers and creeks where these are accessible, to continue their growth and development. In general, juveniles spend the beginning of their lives in the freshwater reaches of rivers, migrating to a brackish or marine environment to breed at 3–4 years of age (Davis 1984a; Almendras et al. 1988; Guiguen et al. 1994).

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Barramundi are hermaphroditic and begin their lives as males, changing sex to females at a weight of around 5 kg (protandrous hermaphrodites). Barramundi generally reach sexual maturity as males at 3–4 years of age (60–70 cm TL), although some fish may mature a little earlier. At this stage, males that are in rivers and creeks migrate to the sea to spawn. In the wild, most males participate in one or more spawning seasons before undergoing a sexual change (protandry), becoming functional females by the next breeding season (Schipp 1996). In general, barramundi males change into females at 6–8 years of age (85–100 cm TL). Fish less than 80 cm length are usually males and those longer than 100 cm are females. However, this varies in many areas, and some sexually precocious populations change sex at younger ages (4–5 years) and sizes (30–50 cm TL) (Davis 1984a). Broodstock held under captive conditions have been found to change sex to females at a smaller size than wild fish. This may be the result of the captive environment or hormone treatments used during the spawning season (Schipp 1996).

7.2.2

Spawning

In tropical areas, barramundi spawn during the summer (Rimmer & Russell 1998). Water temperature is the primary environmental determinant of spawning seasonality, with rainfall patterns (the monsoon or wet season) and photoperiod being secondary drivers (Rimmer & Russell 1998; Balston 2009). In Australia (Southern Hemisphere), barramundi generally spawn between October and March, depending on latitude (Makaira 1999). In the Philippines (Northern Hemisphere), the spawning season for barramundi is between May and September (Harvey et al. 1985), with July and August being the peak period (Garcia 1990a). In areas close to the equator, spawning may occur throughout the year (Sampath-Kumar et al. 1995) or may be more influenced by the onset of the monsoon (wet) season. Barramundi are pelagic spawners, where the eggs and sperm are broadcast into the water column for external fertilisation. In the wild, spawning takes place at night, generally at or after dusk, and appears to be related to the lunar cycle. Barramundi generally spawn after the full and new moons during the spawning season (Schipp 1996), although this may be modified so that spawning occurs on an incoming tide (which may help to transport eggs and larvae into estuarine areas). Spawning generally occurs over 2–3 nights. In some regions, barramundi return to freshwater areas after spawning. In other regions, mature fish tend to remain in coastal and estuarine areas. Movements between river/ estuarine systems tends to be limited in most cases. This has resulted in the development of different strains of barramundi in different locations. In Australia, for example, there are at least 16 different recognised genetic strains (Makaira 1999). Barramundi are highly fecund, and a large female (>120 cm TL) can produce several million eggs per spawning season (Schipp et al. 2007). The eggs hatch within 18 hours and the fry are ready to feed two days later, beginning the cycle once more. Survival of barramundi eggs and larvae is higher in high salinity environments (Moore 1982; Schipp 1996; Balston 2009). Barramundi spawning and the survival of barramundi larvae and juveniles appear to be enhanced by warm sea surface temperatures, high rainfall, increased freshwater flow and low evaporation rates (Balston 2009).

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7.2.3

Physiology

Barramundi can reach up to 1.8 m in length and weigh up to 60 kg, and can live for over 25–30 years (Staunton-Smith et al. 2004). They are carnivorous, and feed on live prey such as fish and prawns. Barramundi adults are crepuscular visual feeders in the wild, and show a peak in feeding at dusk (Barlow et al. 1995). They have been reported to accumulate heavy metals such as mercury (Currey et al. 1992) and organochlorine pesticides (Jabber et al. 2001). Barramundi are able to tolerate temperatures up to 35 °C for prolonged periods, and have a critical thermal maximum of 44.5 °C (Rajaguru 2002).

7.3 7.3.1

HATCHERY PRODUCTION Broodstock

Broodstock are generally maintained in either floating cages (Garcia 1990a; SampathKumar et al. 1995) or tanks (Harvey et al. 1985; Almendras et al. 1988; Schipp et al. 2007). Barramundi hatcheries generally hold between 25 and 70 brood fish, ranging in size from 3 to 20 kg. The availability of excess broodstock helps to ensure egg supply, but increases the cost of holding broodstock. The actual number required depends on the reproductive performance of the fish and the need to maintain adequate genetic diversity in the cultured fish. Broodstock tanks tend to have low stocking densities (from 0.5–1 fish/m3 depending on the size of the broodstock). Broodstock tanks vary in size, but are generally at least 20 m3 in volume, with a depth of at least 1.5–2 m. The sex ratio of females to males in the tanks generally ranges from 1:1 to 1:2 or 2:1 (Almendras et al. 1988; Garcia 1990a,b). Almendras et al. (1988) recommend a female to male ratio of 1:2, and report that egg fertility and hatching rates are higher with this ratio compared to a 1:1 ratio. The environmental conditions in the broodstock tanks are generally set to broadly reflect those experienced by the fish in the wild during the summer spawning period. In general, water quality parameters are maintained at salinity 28–36 ppt, temperature 28–30 °C, pH 7.8–8.4, photoperiod 14L:10D, dissolved oxygen >5.5 mg/L. This helps to condition the fish and to ensure that they can be spawned either naturally or through the use of hormones. Broodstock nutrition is based on the use of a varied diet of trash or bait fish, prawns and squid, usually with the addition of a vitamin supplement (Harvey et al. 1985; Boonyaratpalin 1997). Broodstock are fed an average of 1–2% of their body weight per day. In many cases, broodstock are only fed 2–3 times per week, but with the total summing to the equivalent of 1–2% of their body weight per day.

7.3.2

Spawning

Barramundi will only spawn in salt water (28–36 ppt), and barramundi raised in fresh water must be placed in salt water prior to the breeding season. Spawning occurs at dusk and in the early part of the night. Barramundi appear capable of multiple spawnings during the spawning season, both in the wild (Davis 1984a) and in captivity (Garcia 1989a). Female barramundi generally produce 6–8 million eggs depending on the size of the female (Schipp et al. 2007).

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A number of methods have been used successfully to breed barramundi for aquaculture production. The simplest is perhaps allowing them to spawn naturally in net cages (SampathKumar et al. 1995) or tanks (Boonyaratpalin 1997). Natural spawning results in the highest quality and largest number of eggs, but is dependent on suitable environmental conditions being available for spawning (Boonyaratpalin 1997). Induced spawning is used where greater control of the supply of eggs is required, especially for off-season production of eggs, or where natural production fails to take place (Harvey et al. 1985; Garcia 1990a,b; Boonyaratpalin 1997; Mylonas et al. 2010). Induced spawning is usually done using a combination of environmental manipulation and hormone injection (Garcia 1990a,b). Environmental manipulation is achieved by increasing the water temperature and photoperiod. Broodstock system temperatures are generally raised to 28–30 °C, and the photoperiod increased to approximately 14L:10D. Environmental manipulation allows gonadal maturation to occur. Gonadal development is confirmed by ovarian biopsy through cannulation (Fig. 7.1) (Garcia 1989a,b). Final gonadal maturation and spawning is induced through the use of exogenous hormones, once the environmental manipulations have progressed gonadal development to a suitable stage. Females with an oocyte diameter of at least 400 microns are suitable for hormonal induction (Garcia 1989b). Males should have dense sperm (milt) when cannulated, or display expressible milt with gentle abdominal pressure (Garcia 1989b, 1990a,b). The spermatozoa should display a high degree of sperm activation with sea water (Almendras et al. 1988). Egg quality and size, as well as sperm quality and viability, are confirmed through microscopic examination (Garcia 1989a). A range of hormones, at various dose rates, have been successfully used with barramundi, including HCG, GnRHa and LHRHa (Harvey et al. 1985; Lim et al. 1986; Almendras

Fig. 7.1

Ascertaining the status of gonadal maturation through broodstock cannulation.

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et al. 1988; Garcia 1989a,c, 1990a,b). Luteinising hormone-releasing hormone analogues (LHRHa) is currently perhaps the most commonly used. Hormones are injected either intramuscularly (at the base of the pectoral fin) or intraperitoneally. LHRHa dosages generally vary from 5–25 micrograms/kg body weight, but can be as high as 50–100 micrograms/ kg body weight (Garcia 1990a; Schipp et al. 2007). A dosage rate of approximately 25 micrograms/kg body weight generally yields good results. The spawning rate (number of spawnings of each fish over four consecutive days) increases with the dosage used within the dose limits that have been tested (Garcia 1990a). A dose of 5 micrograms/kg body weight generally produces a single spawning event, while doses of over 10 μg/kg body weight cause spawnings over 2–4 nights (Garcia 1990a). Males do not generally have to be injected in order to spawn (Garrett & O’Brien 1994), but a few males are usually injected at approximately half the dosage used for the females (Harvey et al. 1985; Garcia 1990a; Schipp et al. 2007). Hormone treatment does not appear to influence the number, fertilisation and hatching rates of spawned eggs (Garcia 1990a). Fish are generally injected with hormones early in the day. Spawning occurs at night 34–40 hours after hormone injection (Garcia 1989c). In most cases (depending on hormone dose used), spawning occurs over 2–4 consecutive nights. Egg production, fertilisation rate and hatching rate are usually highest on the first night, and decline over the following nights (Garcia 1989c). In most cases, only eggs from the first night or first two nights are used. Egg production on the first night is generally approximately 300,000 eggs per kg body weight of fish (Garcia 1989c, 1990a). In general, each female produces 3 to 6 million eggs in a spawning, depending on size and condition. The eggs are immediately fertilised by the males (Garrett & O’Brien 1994). An alternative method that has been used to induce spawning in barramundi is the use of slow-release hormone pellets (Harvey et al. 1985; Garcia 1989a, 1990b). The hormone pellets usually consist of LHRHa in a 95% cholesterol binder. Slow-release hormone pellets may help to enable early spawning of barramundi in the absence of environmental manipulation, but may require multiple treatments with the pellets before spawning occurs (Garcia 1990b). Higher dose rates of LHRHa result in an increase in spawning rate (e.g. spawnings over 2–4 nights), but low to moderate doses of LHRHa in the pellets results in better fertilisation rates and hatching success than high doses (Garcia 1989a). A dose of 38–75 μg LHRHa/kg body weight of fish has been recommended for best results (Garcia 1989a). Artificial fertilisation has also been used to obtain barramundi eggs for aquaculture production (Boonyaratpalin 1997). Spawning adults are collected and strip-spawned to obtain the eggs and milt. The eggs are then fertilised manually. Work has also been done on the cryopreservation of barramundi spermatozoa, and indicates that DMSO and glycerol both can be used successfully to preserve the sperm (Leung 1987; Palmer et al. 1993). In general, DMSO gives better results, but may be toxic at very high doses (Leung 1987; Palmer et al. 1993). Cryopreserved sperm can be activated after being held in liquid nitrogen for 90 days, and used to successfully fertilise eggs (Palmer et al. 1993). The larvae produced appear to show no obvious abnormalities.

7.3.3

Hatching

In natural or induced spawning, eggs and sperm are released into the water column and fertilisation occurs externally. Egg size is generally 0.74–0.85 mm in diameter, with a single

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Fig. 7.2

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Positively buoyant barramundi eggs in the hatching tank.

oil droplet (0.23–0.26 mm diameter) (Schipp et al. 2007). Fertilised, viable eggs are positively or neutrally buoyant (Fig. 7.2) (Schipp et al. 2007). In tank-based broodstock systems, eggs are collected in collection baskets through which the effluent water passes. Eggs are then removed early in the morning of the next day, and placed into hatching tanks. Eggs may be treated with ozone (0.2 ppm for 2.5 minutes) or other chemicals to kill off any potential pathogens that may be on the eggs. Eggs are generally stocked in the hatching tanks (Fig. 7.3) at a density of approximately 1,000 eggs/L, but densities of up to 2,000 eggs/L can be used (Schipp et al. 2007). Fertilised eggs develop rapidly, and hatch within 12–17 hours after fertilisation at 27– 30 °C. Hatching rates can vary substantially from less than 20% to over 70%. Following hatching, any dead eggs and/or larvae are usually siphoned off from the bottom of the hatching tanks. The newly hatched larvae are transferred into the larvae rearing tanks or ponds.

7.3.4

Larvae

Newly hatched larvae (Fig. 7.4) have a large yolk sack that is generally absorbed within 48 hours of hatching. The mouth and gut of the larvae develop the day after hatching. Larvae commence feeding approximately 48 hours after hatching. Larval survival is highest when larvae are fed no later than 48 hours after hatching, and declines relatively steeply with delayed feeding (Kailasam et al. 2007).

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Fig. 7.3

Egg hatching tank.

Fig. 7.4

Newly hatched barramundi larvae in the hatching tank.

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7.4

207

HATCHERY CULTURE

The two main culture systems for barramundi are extensive larval culture and intensive larval culture. Each of these two culture system has a number of variations (e.g. clearwater vs. greenwater culture systems in intensive larval culture). Extensive culture is based on the rearing of the larvae at relatively low densities in fertilised marine or brackish water ponds (Rimmer & Russell 1998). Intensive culture is based on the rearing of larvae in tanks at relatively high densities (Schipp et al. 2007). Each system has its advantages and disadvantages, depending on site characteristics. The major advantage of extensive larval rearing is that it requires less labour and facilities compared to intensive culture systems, and therefore results in cheaper production of fingerlings. Larvae also tend to grow faster and are generally more robust (Rimmer & Rutledge 1991). The major advantage of intensive culture systems is that larval survival rates are far more reliable, thus resulting in more stable production of fingerlings. Also, growth rates are more consistent and controllable. Overall, there has been a marked shift towards intensive culture systems in barramundi production over the past decade due to the greater reliability of survival rates and fingerling supply.

7.4.1

Extensive culture

In extensive culture, the larval rearing ponds are filled and fertilised with a combination of organic and inorganic fertilisers 8 to 10 days before the larvae are stocked. Fertiliser regimes for extensive pond culture vary across sites depending on the natural productivity of the seawater and pond soil at each site. Schipp (1996) details rates for the Northern Territory, Australia. The ponds develop natural phytoplankton and zooplankton blooms that the larvae then feed on. Larvae are stocked before first feeding, which is up to two days after hatching, and prey on naturally occurring pond zooplankton. Water turbidity is monitored regularly, and inorganic fertiliser is added when necessary to maintain pond productivity. Copepods are usually the major component of zooplankton in a brackish water pond, and copepod nauplii are ideal food organisms for newly hatched fish larvae. Zooplankton populations need to be monitored at least twice weekly, and rotifers may also be introduced to ponds if necessary (Rimmer & Rutledge 1991). Water quality in the ponds is routinely monitored for stratification as well as temperature, salinity, dissolved oxygen, pH and turbidity. Barramundi are usually stocked at a density of up to 900,000 larvae per hectare. Larval growth is rapid in ponds provided adequate food is available, and the larvae can reach 25 to 35 mm total length (TL) in about three weeks (Rimmer & Rutledge 1991). The fingerlings are generally harvested at 25–35 mm TL, and transferred into weaning tanks at densities of 5,000–10,000 m-3. Weaning is usually complete within a few days (Rimmer & Rutledge 1991). Extensive culture systems result in far more variable survival rates as compared to intensive culture systems (Rimmer & Russell 1998). However, growth rates are, on average, higher in extensive culture systems. The costs of extensive culture systems are about half that of intensive systems (Rimmer & Russell 1998). A number of farms in northern Australia use extensive larval rearing procedures due primarily to the lower costs involved (Barlow et al. 1996; Lobegeiger et al. 1998).

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7.4.2

Intensive culture

In intensive culture, larvae can be cultured in hatcheries using either clear water or greenwater techniques (Palmer et al. 2007). Clear water techniques rely on very strict controls over the entire rearing environment and process. Optimal water quality is maintained in these systems through high rates of water exchange to remove waste products. These systems can be either flow-through or recirculation-based. Recirculating aquaculture systems (RAS) in larval culture usually include water purification components such as UV sterilisers to minimise pathogen levels. Greenwater culture systems are based on the addition of microalgae (e.g. Nannochloropsis oculata) to the rearing tanks (Palmer et al. 2007). Algal densities usually range from 1 × 104 cells/mL to 1 × 106 cells/mL. Palmer et al. (2007) reviewed the benefits of using greenwater culture. In brief, microalgae may aid in maintaining water quality and in reducing the levels of pathogenic bacteria. The algae may also help maintain the nutritional value of rotifers added as food for the larvae, and improve the visibility of live food organisms. Water changes are restricted in greenwater culture systems (Palmer et al. 2007). In the early larval stages, water changes are generally limited to the regular addition of new algae to maintain algal densities in the larval tank. Algae is usually added to the larval tanks throughout the rotifer feeding stage, and the early part of the Artemia feeding stage, before being discontinued (Moretti et al. 1999; Curnow et al. 2006). There has been a marked shift towards greenwater culture over the past decade, and most barramundi hatcheries operate using greenwater techniques (Fig. 7.5). Regardless of systems used, the approximate water quality conditions maintained in intensive larval

Fig. 7.5 figure.)

Greenwater culture of larval barramundi. (Please see plate section for colour version of this

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culture are: temperature: 26–31 °C; salinity: 28 ppt–36 ppt; pH: 8.0–8.4; DO: >90% saturation; Ammonia: 0–0.1 ppm; Nitrite: 0–0.2 ppm; Nitrate: 0–1 ppm.

7.4.3

Larval feeding

Larvae are generally stocked in the larval rearing tanks at a density of approximately 20 larvae per litre. Larvae begin first feeding 48 hours post-hatch and yolk absorption is completed by 96 hours post-hatch (Kailasam et al. 2007). Rotifers are the first food for barramundi larvae, and are added to the rearing tanks from the second day after hatching. Delayed initial feeding has a marked negative impact on larval growth and survival, with a delay until 120 hours post-hatch resulting in complete mortality (Kailasam et al. 2007). Barramundi larvae are visual feeders, and require light to feed (Barlow et al. 1993, 1995). Larval barramundi also display a peak in feeding at dusk, but are unable to feed in total darkness (Barlow et al. 1993, 1995). In contrast, juvenile fish in tanks are apparently able to feed even in darkness (Harpaz et al. 2005a). Larvae are usually fed with rotifers until 12–15 days post-hatch (dph) (Curnow et al. 2006). Rotifer densities range from 5–20 rotifers/mL. Rotifers are added to the rearing tanks 2–4 times a day to ensure that adequate densities are constantly maintained. Rotifers are enriched with microalgae, specific enrichment products high in ω3 HUFAs, or a combination of microalgae and specific enrichment products. The use of copepods as a live feed for barramundi larvae has been trialled recently, and results in better growth and survival compared to rotifers and Artemia (Rajkumar & Kumaraguru Vasagam 2006). Brine shrimp (Artemia sp.) are added to the rearing tanks from the 8th day onwards. There is a substantial period of overlap where both rotifers and Artemia are added simultaneously (weaning period). Artemia densities range from 0.5–10 per mL depending on the age/size of the larvae, where higher densities are maintained as the larvae grow. Artemia are generally enriched with commercial HUFA enrichment products (Curnow et al. 2006). In general, rotifers are discontinued from approximately 13–15 dph, and Artemia are discontinued from approximately 20 dph (Curnow et al. 2006). From Day 15 onwards, the larvae are weaned gradually onto artificial diets (Fig. 7.6). The first artificial diet is generally a pellet of similar size to Artemia (e.g. Gemma 300, Proton 3/5, etc.) (Curnow et al. 2006). The size of the pellet used is gradually increased as the larvae grow. Recent studies have shown that direct weaning from rotifers to artificial feeds is possible, but results in lower survival rates than with a standard feeding protocol that includes Artemia (Curnow et al. 2006). Artificial feeds on their own were unsuitable and led to very high mortality rates (Curnow et al. 2006). One possible reason for this is that fish larvae rely on the enzymes present in live feeds to help digest the food effectively. Thus, younger larvae with their incompletely developed digestive systems are unable to adequately digest and utilise artificial diets (reviewed in Curnow et al. 2006). The acceptability and performance of different artificial pelleted microdiets varies depending on their lipid content and level of leaching of feed attractants such as amino acids (Curnow et al. 2006). High-lipid microdiets with higher levels of leaching of feed attractants are better accepted by fish larvae (Curnow et al. 2006). In some Asian countries, barramundi larvae and young juveniles may be weaned from Artemia onto minced trash fish (bait fish) instead of artificial diets (Boonyaratpalin 1997). This tends to result in poorer growth and survival rates than with standard hatchery protocols.

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Fig. 7.6 Juvenile barramundi at the pelleted feed stage. An auto-feeder is visible at the back end of the tank.

7.4.4

Nursery phase

Barramundi larvae metamorphose into juveniles at approximately 21 days old and 20– 35 mm in total length (TL). At a size of 3–5 cm, the juveniles (Fig. 7.7) are transferred from the larval systems into nursery tanks, ponds or floating cages (Rimmer 1995; Schipp et al. 2007). At this stage, they can be slowly adapted to fresh water or brackish water if required. The optimum salinity range for barramundi fingerlings is 0–36 ppt (Schipp 1996), but barramundi fingerlings can survive in waters with a salinity of over 50 ppt. The optimum temperature for growth is between 28 and 32 °C, but barramundi fingerlings can survive temperatures as low as 16 °C and as high as 35 °C. Juveniles at this stage are generally fed to satiation with either artificial (pelleted) diets or minced bait fish, etc. They are maintained in the nursery systems until they have reached 5–10 cm in length (TL). Stocking densities vary depending on the specific nursery system and size of fish, and range from 20–300 fish/m3. Barramundi adapt easily to high stocking densities, provided that adequate water quality is maintained. Dissolved oxygen levels should be greater than 4 ppm, and ammonia levels should not exceed 1 mg/L. Barramundi will tolerate exposure to levels outside of these ranges, but usually display slower growth rates and increased stress. Prolonged exposure to sub-optimal conditions will result in increased incidence of disease and death. Barramundi are highly cannibalistic and will eat siblings that are less than two-thirds of their own size. Cannibalism is perhaps the biggest cause of mortality in the late stages of larval culture and in the nursery stage. From approximately day 17 onwards, larger fish tend to attack their smaller siblings. Even in cases where the smaller fish survives, the

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Fig. 7.7

211

Juvenile barramundi ready to move into the nursery phase.

injuries from the attack tend to cause mortality a short time later. In some cases, the larger fish also dies from choking on the smaller fish. Cannibalism is managed by grading the fish regularly and separating them into different size classes (Shelley 1993; Rimmer 1995; Schipp et al. 2007). Fingerlings less than 50 mm TL require grading every 3–4 days. Once they reach 50 mm TL, grading is only required once a week until they reach 80–100 mm TL. Fish may be graded again at 200 mm TL and finally at 300 mm TL (Schipp 1996). There are a number of grading systems in use, and most facilities have their own modified systems. Regardless of the system, they all work on the same principle. In essence, a screen is used that enables smaller fish to pass through, but retains the larger fish. The larger sized fish are then transferred into separate tanks. This also helps to reduce size variation between groups, while reducing mortality rates.

7.5

GROWOUT

Barramundi reach the growout stage at a size of 75–100 mm (Tucker 1998; Schipp et al. 2007). Barramundi are grown out in marine, brackish water or freshwater environments. Net cages, ponds and tanks are the three main systems used for barramundi growout. Most barramundi growout systems in tropical areas use net cages or ponds, while indoor recirculation-based tank growout systems are used primarily in non-tropical areas. The optimum temperature for barramundi culture in terms of growth efficiency is between 27 and 31 °C, with good growth rates between 26 and 33 °C (Williams & Barlow

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1999; Katersky & Carter 2005, 2007a,b; Williams et al. 2006; Glencross 2008). Barramundi are able to survive at temperatures below this range, but show decreased metabolism and growth and generally stop feeding at temperatures below 20 °C. Growth rates are highly variable depending on environmental factors, feed type, stocking density and growout system used. Fish generally grow from fingerlings to 300–800 g within 6–12 months, and to 2–3 kg in approximately 2 years. There are no significant differences in growth rates between fresh water and salt water. Barramundi are relatively hardy fish, and survival rates in growout systems tend to be high in the absence of severe disease outbreaks or other major issues.

7.5.1

Cage culture

Net cages range in size from 3 × 3 m up to 10 × 10 m, and 2–4 m deep. Cages are also frequently placed within ponds or even in recirculating systems. Cage culture in estuarine or marine waters has advantages over other systems, particularly where large-scale production is envisaged. Cage-based systems require regular maintenance, particularly to ensure that the cage mesh does not clog up due to biofouling (Rimmer & Russell 1998). Also, holes due to predators, etc need to be repaired rapidly to prevent fish escapes, particularly in open water areas (Schipp 1996). Predator nets are generally used around the cages to minimise the occurrence of holes (Barlow 1998). Stocking densities for cage culture are 40–50 fish/m3 for the first 2–3 months, reducing to 10–30 fish/m3 after that (Rimmer & Russell 1998). Depending on cage size and environmental conditions, stocking biomass in cage systems range from 15 kg/m3 to as high as 60 kg/m3.

7.5.2

Recirculating systems

The second method of on-growing barramundi is intensive production in an indoor, controlled environment. Recirculating tank-based growout systems generally use underground (bore) fresh water or brackish water and a high level of recirculation through filters (Rimmer & Russell 1998). The major advantage of these systems is that they can be located close to markets, as long as water is easily available. The high degree of control in these systems, particularly of temperature and feeding, allows year-long production and rapid growth. However, capital and operating costs are generally higher than for outdoor cage systems (Barlow 1998). In recirculating systems, stocking densities are usually maintained at between 15 and 40 kg/m3, depending on the technologies and methodologies used. At optimum temperatures, barramundi can reach market size (500 g) in 6–12 months. It also avoids the environmental concerns associated with release of nutrients to open waterways from pond or cage culture operations (Schipp 1996).

7.5.3

Pond culture

Earthen ponds (without cages) are a common growout system in tropical areas of Australia and Asia, and are known as free-ranging (Rimmer & Russell 1998; Russell & Rimmer 2004). In addition, cage culture of fish less than 120 to 150 mm TL and free-ranging of

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larger fish are sometimes combined (Schipp 1996). Ponds are usually aerated, with a water exchange rate of 5–10% of pond volume per day. Growth rates are generally faster than with cages or tanks. Fish are more difficult to harvest, and this is usually done through seine netting. Stocking densities are generally between 0.25 and 2 fish/m2.

7.6

NUTRITION AND GROWTH

Barramundi are fed with commercially available pellets in Australia, but still tend to be fed with trash/bait fish in many parts of Asia (Rimmer & Russell 1998; Schipp et al. 2007). A substantial amount of work has been done on the nutritional requirements of barramundi (Boonyaratpalin 1997; Williams & Barlow 1999; Glencross 2006). However, much of the work has focused on smaller fish (<300 g initial size), and less is known about the requirements of larger sized fish. In general, the nutritional requirements of barramundi are broadly similar to those of other tropical marine carnivorous species (Boonyaratpalin 1997; Glencross 2006). Much of the focus in barramundi nutrition has been on improving feed quality. There is also an increasing focus on shifting away from the use of trash/bait fish and more into the use of formulated dry feeds derived from sustainable source materials such as plantbased meals (Catacutan & Coloso 1997; Tantikitti et al. 2005). Furthermore, the nutritional quality and food conversion efficiency of formulated feeds is generally superior to that of trash/bait fish (Catacutan & Coloso 1997). As a result, improved growth rates and survival, as well as, improved feed conversion ratios (FCR) and reduced waste production are obtained with formulated diets in barramundi (Tantikitti et al. 2005). Feeding and growth in fish are affected by internal and external factors (Tucker 1998; Volkoff et al. 2009). Intrinsic factors include age, size, growth rate, reproductive status and genetic variability. External factors include abiotic factors such as temperature, photoperiod and salinity as well as biotic factors such as stocking densities and feed composition and quantity (Volkoff et al. 2009). In addition, fish may display diurnal or circannual rhythms (Tucker 1998; Volkoff et al. 2009). The extent of endocrine control of feeding in fish is still uncertain (Volkoff et al. 2009). Optimisation of feeding and nutrition in barramundi is generally undertaken by manipulating diet components such as feed composition, feed ration, and frequency and time of feeding. The nutritional requirements of barramundi change with the age and/or size of fish (Boonyaratpalin 1997; Nankervis et al. 2000; Williams et al. 2003a; Glencross 2008; Glencross et al. 2008). Smaller barramundi have a higher requirement for protein, but reduced requirement for lipid and gross energy compared to larger barramundi. Thus, diets formulated for smaller barramundi generally contain higher protein, lower lipid and lower energy levels than diets formulated for larger fish (Glencross 2006). Smaller fish require a higher percentage of their body weight in feed per day, but the overall amount required increases with fish size (Boonyaratpalin 1997; Williams & Barlow 1998a; Glencross 2006, 2008). Feed intake also increases with increasing water temperature, and optimum growth occurs at around 26–31 °C (Williams & Barlow 1999; Katersky & Carter 2005, 2007a; Williams et al. 2006; Glencross 2008). Williams and Barlow (1999) suggested that small barramundi should be fed twice per day and large barramundi should be fed once per day. There is no advantage to feeding barramundi more frequently (Williams & Barlow 1999). The amount of weight gained per day in absolute terms increases with

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fish size (Glencross 2006). Under Australian conditions, weight gain is estimated to increase from 2.24 g/day in 50 g fish to 10.70 g/day in 2,000 g fish cultured at 30 °C (Glencross 2008). Feed tables have been developed for barramundi grown under different conditions and at different sizes (Boonyaratpalin 1997; Glencross 2008). The most comprehensive feed table to date is that developed by Glencross (2008) for Australian conditions, which accounts for a range of different temperatures and different energy density diets, and covers the fish size range from 10 g to 3,000 g. With dry feeds, Harpaz et al. (2005a) suggested that juvenile barramundi (approximately 20 g starting weight) should be fed at approximately 4% of biomass per day. Glencross (2008) suggested that juvenile barramundi of 10 g should be fed 4.20–5.04% of their body weight per day at 26 °C, depending on the energy density of the diet (15 MJ per kg digestible energy vs 18 MJ per kg digestible energy). In comparison, 3000 g adult fish should be fed 0.51–0.61% of their body weight per day at 26 °C, again depending on the energy density of the diet (Glencross 2008). While feeding at higher levels had no benefit, feeding at 2% of biomass per day resulted in reduced growth, (Harpaz et al. 2005a). Restricted feeding can also lead to lower survival rates as a result of increased injuries from aggression (Harpaz et al. 2005a). With wet feeds (trash/bait fish), Boonyaratpalin (1997) suggested that fish under 5 g should be fed 7.18% of their body weight per day. This decreases with fish size, and fish between 28 and 45 g should be fed 3.5% of their body weight per day (Boonyaratpalin 1997). Barramundi perform best with diets that contain approximately 45–55% protein (Catacutan & Coloso 1995, 1997; Williams & Barlow 1999; Williams et al. 2003a). Smaller fish would generally perform better on diets at the high end of the range, while larger barramundi would perform well on diets at the lower end of the range. The specific amino acid requirements of barramundi are still poorly understood, but are thought to be similar to those of other carnivorous fish (Glencross 2006). Higher protein levels result in better growth rates, provided that the energy density of the diet is also high (Williams et al. 2003a). However, growth in larger fish is more closely correlated with energy density of the feed than with the protein content, although there is still a positive correlation between growth and protein intake (Glencross et al. 2008). The optimum lipid level for barramundi diets is between 15 and 22% lipid (Catacutan & Coloso 1995, 1997; Williams & Barlow 1999; Williams et al. 2003a). The minimum lipid level for adequate growth appears to be approximately 10%, and lipid levels of 5% caused fatty acid deficiency symptoms irrespective of protein levels (Catacucan & Coloso 1995). In general, barramundi grow better when fed with diets containing higher lipid levels, provided that protein levels in the diet are also high (Williams et al. 2003a). However, very high lipid levels can lead to excessive fat deposition in the fish carcass (Catacucan & Coloso 1997; Glencross et al. 2008). As with other carnivorous marine fish species, barramundi have a relatively high requirement for ω3 HUFAs. The ω3 HUFA requirement is approximately 1.5–1.7% of the total diet, with 1.7% recommended (Boonyaratpalin 1997). While barramundi do not have a specific requirement for carbohydrates, carbohydrates appear to be effective in producing protein-sparing and lipid-sparing effects in juvenile barramundi (Catacucan & Coloso 1997; Nankervis et al. 2000). Barramundi juveniles are able to effectively utilise gelatinised starch as a source of carbohydrate up to an inclusion rate of 17–20% of the diet (Catacucan & Coloso 1997; Nankervis et al. 2000). Catacucan and Coloso (1997) suggested a carbohydrate level of 10–17% for barramundi diets.

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The vitamin and mineral requirements of barramundi are still poorly understood. Quantitative minimum requirements have been established for ascorbic acid (30 mg/kg of diet as ascobyl-2-monophosphate-magnesium), pyridoxine (5–10 mg/kg of diet) and pantothenic acid (15 mg/kg of diet as calcium D-pantothenate) (Boonyaratpalin 1997; Phromkunthong et al. 1997; Glencross 2006). The phosphorous requirements of barramundi are estimated to be 5.5–6.5 g/kg of diet (Boonyaratpalin 1997; Glencross 2006). Harpaz et al. (2005b) suggested that adding salt to the diet of barramundi grown in fresh water, up to the 4% inclusion level, improves feed utilisation. While deficiency symptoms have been observed for a number of vitamins when using purified diets in laboratory trials, the main vitamins that are likely to cause deficiency symptoms with the use of practical diets are thiamine, riboflavin, panthothenic acid and Vitamin C (Boonyaratpalin 1997). Barramundi perform well on diets with a gross energy density of approximately 18– 23 MJ/kg (Catacutan & Coloso 1995, 1997; Williams & Barlow 1999; Nankervis et al. 2000; Williams et al. 2003a; Glencross et al. 2008). Increases in dietary energy result in higher growth rates and better feed conversion ratios, and also lead to increased protein and lipid gain (Nankervis et al. 2000). Body lipid content is positively correlated with lipid content of the diet, and increases with dietary energy (Catacucan & Coloso 1995; Nankervis et al. 2000; Williams et al. 2003a). The quantity eaten decreases with increasing energy density of the diet (Nankervis et al. 2000; Glencross 2006). Larger fish require higher energy density diets than smaller fish (Williams et al 2003a, Glencross 2008). A metabolisable energy (ME) density of 24 MJ/kg has been recommended for large barramundi (Glencross et al. 2008). Barramundi perform well with a crude protein to gross energy ratio of approximately 25–30 mg/kJ (Catacutan & Coloso 1995, 1997; Williams et al. 2003a; Glencross 2006; Glencross et al. 2008). The optimum metabolisable protein to metabolisable energy ratio for large barramundi (approximately 1.6 kg) is 18.4 g/MJ (Glencross et al. 2008). A range of alternative protein sources have been trialled in barramundi as partial protein replacements for fishmeal (Boonyaratpalin et al. 1998; Williams et al. 2003b; Glencross 2006; Katersky & Carter 2009). Meat meal successfully maintained growth rates and protein growth efficiency at inclusion levels of 40–50% (Williams 1998; Williams et al. 2003b). The ability of barramundi to utilise soybean-based proteins depends on the type of soybean product used (Boonyaratpalin et al. 1998; Tantikitti et al. 2005). Solvent-extracted soybean meal supports good growth rates (Boonyaratpalin et al. 1998; Williams 1998). The maximum inclusion level with solvent-extracted soybean is approximately 30% (Williams 1998). Tantikitti et al. (2005) suggest that 10% of fishmeal protein can be replaced with defatted soybean meal without significant adverse impacts on growth performance. Excessively high replacement levels impair growth performance (Boonyaratpalin et al. 1998; Tantikitti et al. 2005; Glencross 2006). Partial replacement of fish oil with soybean oil also has no significant effects on growth or food intake (Raso & Anderson 2003). Good results have been obtained with lupin-based ingredients (Williams 1998; Katersky & Carter 2009). Lupin-based ingredients have been successfully used to replace fishmeal at an inclusion level of up to 45% without negatively affecting growth performance or protein synthesis (Katersky & Carter 2009). A range of FCR values have been obtained under laboratory conditions, depending on diet composition, age of fish, etc. These can be as low as 0.8–0.9:1 for young fish fed on high-protein and high-lipid formulated diets (Williams et al. 2003a). In general, formulated feeds produce FCRs of between 1.0–1.6: 1 under laboratory conditions depending on moisture content and feed composition (Catacutan & Coloso 1995, 1997; Williams et al.

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2003a; Glencross 2006; Glencross et al. 2008). FCRs under practical farm conditions generally range from 1.2 to 2.0 (Schipp et al. 2007). FCRs with trash/bait fish are substantially higher, and range from 3.8 to 7.4 (Boonyaratpalin 1997). Growth rate and FCR both improve with increasing feed ration up until just before the satiety feeding level, and then begin to decline (Glencross 2008). Barramundi display compensatory growth in response to short periods of food deprivation or food rationing (Tian & Qin 2003, 2004). Fish starved for one week were able to compensate for this over a three-week re-feeding period (Tian & Qin 2003). Growth lost during starvation for 2 or 3 weeks, however, could not be fully compensated (Tian & Qin 2003). Similarly, fish fed at 50% or 75% of satiation for 2 weeks were able to display a compensatory growth response over a 5-week re-feeding period (Tian & Qin 2004). However, fish fed at 0% or 25% of satiation could not recover the lost growth.

7.7

HEALTH MANAGEMENT

The intensification and rapid expansion of barramundi farming over the past few decades has led to an increase in the incidence of various diseases (Anderson & Norton 1991; Seng 1997; Kumar et al. 2007, 2008; Parameswaran et al. 2008). Barramundi are susceptible to a wide range of diseases including parasites (Herbert et al. 1995; Seng 1997; Ryan 2010), bacteria (Soltani et al. 1996; Azad et al. 2004; Delamare-Deboutteville et al. 2006; Kumar et al. 2007, 2008; Bromage & Owens 2009) and viruses (Munday et al. 1992, 2002; Azad et al. 2006; Parameswaran et al. 2008). These diseases cause significant economic losses to the barramundi industry each year. Two of the main diseases that are particularly problematic in the barramundi aquaculture industry are viral nervous necrosis (VNN, also known as viral encephalopathy and retinopathy or VER) and Streptococcus iniae.

7.7.1

Viral nervous necrosis

Viral nervous necrosis (VNN) is caused by nervous necrosis virus. The causative agent is an RNA virus with a diameter of 20–34 nm (Wu & Chi 2006; Chia et al. 2010). It is classified as a betanodavirus (piscine nodavirus) of the Nodaviridae (Chia et al. 2010). The disease occurs in both wild and farmed fish (Gomez et al. 2004), and has been reported in over 40 species of fish from different locations (Azad et al. 2006). The disease can affect fish of all sizes and ages, and causes high mortality episodes in the hatchery and nursery stages (Glazebrook et al. 1990; Munday et al. 1992; Le Moullac et al. 2003; Azad et al. 2005; Parameswaran et al. 2008), as well as in net cages during growout (Munday et al. 1992; Fukuda et al. 1996; Le Breton et al. 1997). The disease is fast acting, and can cause very high mortalities within a short period of time (Munday & Nakai 1997; Parameswaran et al. 2008). 7.7.1.1 Diagnosis The disease affects the nervous system. On a histopathological level, it causes necrosis and degeneration of the nervous system, including vacuolation of the brain and retina (Glazebrook et al. 1990; Munday et al. 1992; Wu & Chi 2006; Parameswaran et al. 2008).

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Clinical symptoms include anorexia, abnormal swimming behaviour (usually whirling or spiralling swimming behaviour) and darkened body colouration (Munday et al. 1992; Munday & Nakai 1997; Le Moullac et al. 2003; Wu & Chi 2006; Parameswaran et al. 2008; Chia et al. 2010). Infected fish show rapid progression of the disease (Munday et al. 1992). The incubation period for the virus is approximately 4–6 days (Glazebrook et al. 1990; Azad et al. 2006). In hatcheries, the first overt signs may start to manifest as early as 9 days post-hatch, and mass infection becomes obvious at 15–18 days post-hatch (Munday et al. 1992). The presence of the virus in barramundi larvae can generally be detected using immunohistochemistry-based testing (immunofluorescence/immunoperoxidase) from as early as 4–6 days post-hatch (Azad et al. 2006). This is similar to what has been observed in other species (Breuil et al. 2002). Standard histopathological symptoms such as vacuolation of nervous tissue become obvious from 15 days post-hatch (Azad et al. 2006). The clinical (behavioural) symptoms usually start to manifest approximately 24–36 hours after the standard histopathological symptoms (Munday et al. 1992). Mortality rates of greater than 50% occur within the first month (Munday et al. 1992). Fingerlings that survive the initial disease do not survive long during the nursery and growout stage, and usually succumb to other pathogens (Munday et al. 1992). Larvae that have been directly exposed to the virus through immersion develop clinical symptoms 2 days post-infection, and can result in 100% mortality within 4 days of infection (Parameswaran et al. 2008). Juveniles that have been subjected to intramuscular injection of the virus develop clinical symptoms 10 days post-infection, and can also result in 100% mortality by 15 days post-infection (Parameswaran et al. 2008). 7.7.1.2

Transmission

Transmission of the virus appears to occur through multiple pathways. Vertical transmission from the mother to the eggs may occur in some fish species including barramundi (Grotmol et al. 1999; Grotmol & Totland 2000; Breuil et al. 2002; Azad et al. 2006). Nodavirus has been detected in the connective tissue around the oocytes, and may contaminate the eggs during spawning and hatching (Grotmol et al. 1999; Grotmol & Totland 2000; Breuil et al. 2002; Azad et al. 2006). Contamination of the larvae could occur either through direct contact with the egg casing during hatching or through the hatching water (Breuil et al. 2002). The virus then enters the larvae through the musculature before progressing into the nerve cells (Azad et al. 2006). Another mode of transmission is horizontal transmission via the surrounding water (Munday et al. 1992). Eggs, larvae, juveniles and adults may become infected by nodavirus present in the surrounding water. Barramundi larvae have been shown to be susceptible to nodavirus present in the surrounding water, and juveniles can become infected through the musculature (Parameswaran et al. 2008). Similarly, eggs of other species have been shown to be susceptible to nodavirus present in the surrounding water (Breuil et al. 2002). The virus is also easily transferred from infected fish to uninfected fish through shedding into the water column (Schipp et al. 2007). 7.7.1.3 Prevention and treatment The management of nodavirus issues in barramundi aquaculture requires action on a number of levels. Improved hygiene has been shown to be effective in minimising

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Fig. 7.8

Regular health checks are essential for managing disease issues.

nodavirus infections in hatcheries (Munday et al. 1992). Furthermore, removing juveniles from the larval culture area and transferring them elsewhere, followed by a thorough sterilisation process of the larval culture area, helps to reduce the risk of nodavirus infections. Stress appears to be a major trigger of the disease manifesting in fish, and reducing stress levels helps to minimise the occurrence of outbreaks. Stress factors that could potentially be involved include poor water quality, overcrowding, and poor nutrition, amongst others. Given the possibility of vertical transmission of the virus from parent fish, increased testing of broodstock and oocytes could enable the selection of disease-free fish for broodstock (Le Moullac et al. 2003). A key challenge is to be certain that broodstock are indeed disease-free as the disease may be present in low numbers and/or be latent, and thus not easily detected (Schipp et al. 2007). Thus, multiple and regular testing (Fig. 7.8) may be required to ensure that broodstock are indeed disease-free. Also, strict hygiene protocols are required to prevent infection of broodstock through other fish or the water, and to ensure that they remain disease-free. Surface sterilisation of the spawned eggs using ozone has been used in an attempt to minimise the risk of transmission of nodavirus to the eggs during spawning (Grotmol & Totland 2000; Breuil et al. 2002; Buchan et al. 2006). A common method is to expose the eggs to short duration baths in ozonated water shortly after hatching. The effectiveness of ozone-based egg sterilisation as a means of reducing the risk of nodavirus transmission has been demonstrated in a range of species (Grotmol & Totland 2000; Breuil et al. 2002;

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Buchan et al. 2005). Caution needs to be exercised in ensuring that the dosage and duration of the bath treatment are adequate for sterilisation, but do not overly impact on egg hatching rates (Grotmol et al. 2003). The specific dosage required varies between fish species (Grotmol et al. 2003) and a dose of 0.5 mg/L for 2 minutes has been recommended for barramundi (Schipp et al. 2007). Aside from its potential benefits in treating nodavirus, sterilisation may also help to reduce the occurrence of fungal and bacterial infections in the eggs. There are currently no effective treatments for nodavirus, and strict testing is done in many places to minimise the spread of the disease. In Australia, for example, barramundi fingerlings from hatcheries are tested at 2–3 weeks of age, and require disease-free certification before they can be transported off-site. Infected fish are usually sacrificed. One of the challenges is that larvae and juveniles may not show any obvious clinical symptoms in some cases, but still be positive for nodavirus under histopathological, PCR, ELISA or immunohistochemistry testing. Furthermore, there may occasionally be a mismatch between histopathological diagnoses and diagnosis based on more sensitive tests such as PCR, especially where there are no overt clinical symptoms. Thus, PCR-based tests may indicate that fish are nodavirus-positive, while standard histology-based tests may indicate that they are not. As a precautionary measure, some places routinely undertake testing using both standard histology and PCR, and utilise the PCR results where there is a mismatch between the two tests. Tests based on ELISA or immunohistochemistry techniques have also been trailed in an attempt to develop a more sensitive and accurate test for nodavirus. New vaccines are currently being developed to improve resistance to nodavirus infections in fish (Tanaka et al. 2001; Thiery et al. 2006; Pakingking et al. 2009, 2010). These vaccines induce the production of nodavirus-neutralising antibodies, and could provide substantial protection even in the longer term (Pakingking et al. 2009, 2010). The effectiveness of these vaccines in commercial barramundi aquaculture is currently unknown. There is also the risk that the virus may be able to mutate, and that resistant forms may develop with widespread use. Regardless, the further development of vaccines may help to reduce the impact of nodavirus infections in the future as the industry develops.

7.7.2

Streptococcus iniae

Streptococcus iniae are a gram-positive coccoid bacterium (Bromage et al. 1999). It is one of the groups of bacteria that causes the disease Streptococcosis (Bromage et al. 1999). The disease occurs in both freshwater and saltwater fish, and can result in severe infections and high mortalities in barramundi (Bromage et al. 1999; Bromage & Owens 2002, 2009; Schipp et al. 2007; Nawawi et al. 2008). It is rapidly becoming one of the most serious bacterial diseases impacting on aquacultured barramundi. Streptococcus iniae appears to be widespread in warm water aquaculture, and infections have been reported in a number of countries (Bromage et al. 1999; Agnew & Barnes 2007). While they can occur in barramundi of all sizes and ages, they appear to be most commonly reported from fish in the growout stage of culture, particularly younger fish. Their impact on the barramundi aquaculture industry can be quite high, and approximately 90% of cumulative fish losses in at least one site in Australia have been attributed to S. iniae (Bromage & Owens 2009).

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7.7.2.1 Diagnosis Streptococcosis causes damage to the nervous system, meningoencephalitis and septicemia. The behavioural symptoms tend to vary between species and between cases. Symptoms in barramundi include lethargy, loss of orientation, erratic swimming behaviour, corneal opacity, ascites, exopthalmia, increased ventilation rate and internal and/or external haemorrhaging (Bromage et al. 1999; Bromage & Owens 2002; Schipp et al. 2007). In acute cases, fish may be found moribund or dead with few visible symptoms aside from slight corneal opacity (Bromage et al. 1999; Bromage & Owens 2002; Schipp et al. 2007). Preliminary identification of S. iniae is usually done using standard bacterial testing methods such as bacterial plating. Definitive diagnosis of S. iniae as the causative agent is generally done using biochemical and physiological testing (Bromage et al. 1999). However, in some cases definitive diagnosis may require molecular genetics methods, such as DNA sequencing and DNA-DNA hybridisation. S. iniae is highly pathogenic and lethal, and mortality rates of greater than 40% within 48 hours have been observed under experimental conditions (Bromage et al. 1999). Fish that survive an infection may carry the disease asymptomatically in the nervous tissue. The modes of transmission of S. iniae are still unclear. Bromage and Owens (2009) suggest that transmission from the host fish to the surrounding water (and/or vice versa) occurs. The bacteria appear to be absent from the surrounding water when there are no sick fish and vice versa. Transmission to humans and other mammals have also been reported, and involves direct contact with infected fish (Agnew & Barnes 2007). Thus, there may be multiple modes of transmission of the pathogen. It has been demonstrated that barramundi can become infected by oral administration of the bacterium, as well as by immersion in water containing the bacterium (Bromage & Owens 2002). The results suggest that sub-acute infection is more likely to occur via oral transmission. Acute infections that result in mass mortalities are more likely to occur as a result of the increased presence of the bacterium in the environment (Bromage & Owens 2002).

7.7.2.2

Prevention and treatment

Streptococcus iniae can be treated successfully in fish using a range of antibiotics, including enrofloxacin, amoxicillin, erythromycin and oxytetracycline (Agnew & Barnes 2007). The results have been good with enrofloxacin, although there are concerns with the development of resistant strains of the pathogen (Stoffregen et al. 1996). The results have been mixed with oxytetracycline (Agnew & Barnes 2007). Vaccines have been developed for S. Iniae (Delamare-Deboutteville et al. 2006; Russo & Yanong 2009). Vaccines for barramundi are generally developed from autogenous strains of the bacterium (Schipp et al. 2007). The vaccine can be administered either by intraperitoneal injection or via immersion bath (Delamare-Deboutteville et al. 2006). While both methods of administering the vaccine can be successful, intraperitoneal injection generally results is a stronger and longer lasting response (Delamare-Deboutteville et al. 2006). The vaccine is usually administered when the fish are approximately 7–10 g body weight (Schipp et al. 2007). At present the most effective way of dealing with the disease on farms is through management interventions. Outbreaks of the disease are strongly correlated to temperature. The optimum temperature for the disease outbreaks is between 25 and 28 °C (Bromage & Owens 2009). Temperatures on either side of this range have a far lower risk of resulting in a

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disease outbreak either in the laboratory or in net cages on farms (Bromage & Owens 2009). Thus, management interventions such as reducing stocking density, feeding rates and handling during the time period when the temperatures are close to the optimum range for the disease have been demonstrated to be effective in reducing the risk of outbreaks (Bromage & Owens 2009).

7.8 7.8.1

QUALITY Post-harvest handling

While much of the focus in the industry to date has been on the larval culture and growout of the barramundi, and on sales and marketing, new research suggests that handling of stock during and after harvest also plays a key role (Wilkinson et al. 2008). Fish that have been harvested using rested harvest techniques are easier to process and have better quality flesh compared to fish harvested using standard techniques (Wilkinson et al. 2008). The rested harvest technique causes less stress and minimises exercise in the fish during harvest, thus improving flesh quality. However, the cost-effectiveness and practicality of this technique has yet to be tested on large-scale commercial farms.

7.8.2

Sensory characteristics

A key challenge with barramundi aquaculture in freshwater ponds or net cages has been the presence of a muddy or earthy taint to the flavour of the fish flesh (Percival et al. 2008). This is caused by the presence of the compounds geosmin (GSM) and/or 2-methylisoborneol (MIB) in the fresh water (Percival et al. 2008). These compounds are metabolites produced by algae and cyanobacteria in the fresh water (Howgate 2004). This muddy taste is absent in saltwater-farmed fish (Percival et al. 2008). The muddy taste varies between different parts of the fish, and is strongest in areas with higher lipid content (Percival et al. 2008). As larger fish have higher lipid content than smaller fish, larger fish also tend to have a noticeably stronger muddy taste overall (Percival et al. 2008). Purging the fish in either brackish or salt water or in fresh water that is free of GSM and MIB for approximately 1 week effectively removes the muddy taste from the flesh of the fish (Percival et al. 2008). There is no preference for wild-caught versus either saltwater-farmed or purged freshwaterfarmed barramundi (Percival et al. 2008) in sensory tests. The effect of different diets and culture methods on the sensory characteristics of barramundi has also been evaluated. In general, the differences between different diets are relatively small (Glencross et al. 2008). There is a correlation between lipid levels in the diet and the dryness/oiliness of the flesh of the fish. Also, bait fish-fed barramundi tended to taste more fishy that pellet-fed fish (Glencross et al. 2008). While there were differences in the other sensory characteristics, these were relatively minor.

7.9

SALES AND MARKETING

Barramundi are sold at a range of sizes, and in a variety of forms (Rimmer & Russell 1998). Live fish are sold at a premium price. Plate size fish (350–500 g) are popular in Asia, and

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are sold either live or fresh/chilled. Most recirculating system production is based close to urban centres, and the fish tend to be sold at plate size. The plate sized fish market, however, is fairly saturated, and the focus has now shifted to the production of larger sized fish. Most pond and cage produced fish outside of Asia are sold at larger sizes of 2 kg or more. These can be sold as whole chilled fish, gilled and gutted or as fillets and cutlets (Rimmer & Russell 1998). Gilled and gutted fish range in size from 500 g–2 kg, while whole chilled fish and fillets and cutlets tend to be over 2 kg (Rimmer & Russell 1998). The economics of production favour larger fish, but the risks also increase and returns are delayed. The price/kg of barramundi varies with the form it is sold in. Prices also vary over time depending on supply and demand within specific market segments. Average prices have declined over the past decade for some barramundi products, primarily due to increased production, as well as increased competition from substitute products. A number of marketing strategies have been explored to differentiate barramundi from other fish, and to help position it as a premium product. These include promoting saltwaterfarmed fish as being of premium quality as these do not suffer from the muddy taste issues associated with some freshwater production. Also, Australian and US production is marketed as barramundi to help differentiate it from Asian production (which is usually called Asian seabass or giant seaperch.

7.10 7.10.1

FUTURE DIRECTIONS Selective breeding

Selective breeding of barramundi broodstock is the next key step for the barramundi aquaculture industry, and will be required to maintain the rapid growth of the industry in the Asia-Pacific region and elsewhere (Zhu et al. 2006; Wang et al. 2007; Yue et al. 2009). There are significant ecological and genetic differences between wild barramundi populations, and a better understanding of these differences will help the development of targeted selective breeding programmes for aquaculture (Zhu et al. 2006; Yue et al. 2009). This genetic variation is currently being documented in a number of countries including Australia (Salini & Shaklee 1988; Shaklee et al. 1993; Keenan 1994; Chenoweth et al. 1998a,b; Doupe & Lymbery 1999; Frost et al. 2006), Singapore (Zhu et al. 2006; Wang et al. 2007; Yue et al. 2002, 2009), Malaysia (Norfatimah et al. 2009), and elsewhere. There are at least 16 genetically distinct barramundi populations across northern Australia (Keenan 1994), and at least 7 distinct populations in Queensland alone (Shaklee et al. 1993). There are also marked genetic differences within Asia (Zhu et al. 2006; Norfatimah et al. 2009; Yue et al. 2009), and between Asian and Australian barramundi populations (Yue et al. 2009). The results to date indicate that wild populations of barramundi display high levels of genetic variability, which can be harnessed for aquaculture (Yue et al. 2009). Comparisons between wild and aquacultured stock, however, suggest that this genetic diversity is being lost. Wild populations display higher levels of allelic diversity, allele richness and heterozygosity (gene diversity) than cultured stock (Yue et al. 2009). Cultured stock from both Asia and Australia appear to have experienced a recent bottleneck, and may have been established from a small number of founder individuals, or a relatively small effective population size due to differential contribution of broodstock to surviving larvae and juve-

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niles (Frost et al. 2006; Yue et al. 2009). This loss of genetic diversity in aquaculture tends to be exacerbated over multiple generations of captive breeding (Frost et al. 2006; Yue et al. 2009). One of the key challenges with barramundi aquaculture is that most existing breeding programmes are not designed with the intention of maximising genetic potential or selectively breeding for specific traits (Le Moullac et al. 2003). Barramundi hatcheries use mass spawning with varying numbers of males and females, and there is generally little monitoring and control of the relative contributions of the different broodstock to the next generation of fish (F1 generation). As a result, the effective population size may be half or less of the observed population size due to the design of the breeding program (Frost et al. 2006). With the increase in genetic information on barramundi populations, the industry is moving rapidly towards implementing better designed breeding programmes. With the development of these new programmes there should be a marked improvement in the quality of barramundi production.

7.10.2

Inland saline aquaculture

Increasing groundwater salinity, due to secondary salinity, is an issue in over 20 countries worldwide, and affects over 380 million hectares of land (Ghassemi et al. 1995; Partridge & Lymbery 2008). In Australia, for example, approximately 5.8 million hectares of agricultural land is affected by secondary salinity. There has been a significant amount of interest worldwide in utilising these areas more productively, and aquaculture has been identified as one potential use (Allan et al. 2001a,b). The advantages of utilising inland saline areas for aquaculture include their lower cost, reduced environmental sensitivity, reduced risk of conflict with other potential uses, and reduced risk of transfer of pathogens (Partridge et al. 2008). A number of fish and other species have been evaluated for aquaculture potential in inland saline areas, including barramundi (Partridge & Lymbery 2008; Partridge et al. 2008). The results to date have been mixed. A key challenge with inland saline aquaculture is variability in the ionic composition of saline groundwater between locations. While the ionic composition of saline groundwater is generally broadly similar to that of seawater, it is also influenced by the nature and timing of recharge and the nature of the ground material (Partridge & Lymbery 2008). A deficiency in potassium in saline groundwater is common in most areas. This generally makes these areas unsuitable for aquaculture without ongoing potassium supplementation. The potential to culture barramundi in inland saline areas has been explored in recent years (Partridge & Creeper 2004; Partridge & Lymbery 2008; Partridge et al. 2006, 2008). Barramundi can be successfully cultured in inland saline areas, but as with other species, requires adequate potassium ion concentrations in the water. However, the level of potassium supplementation required for barramundi juveniles may be lower than for many other fish species considered for inland saline aquaculture, provided that the salinity is reduced to approximately 15 ppt (Partridge & Lymbery 2008). With reduced salinity and potassium ion concentrations of at least 20% K-equivalence (Jain et al. 2006) or 25% K-equivalence (Partridge & Lymbery 2008), the survival and growth of barramundi juveniles is similar to that achieved in normal seawater at 15 ppt salinity (Partridge & Lymbery 2008). Barramundi larvae also survive and grow in saline groundwater at lower salinity levels, but have a higher requirement for potassium than juveniles (Jain et al. 2006; Partridge

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et al. 2008). Larvae were unable to survive in unsupplemented saline groundwater with 38% K-equivalence, even at 15 ppt salinity (Partridge et al. 2008). In contrast, their survival in 100% K-equivalence saline groundwater was broadly similar to that observed in normal seawater (Partridge et al. 2008). Overall, the potential for barramundi aquaculture in saline groundwater appears relatively promising compared to other fish species that are being considered. However, it appears that barramundi growout may be more feasible in inland saline areas rather than hatchery production.

7.11

CONCLUSIONS

The barramundi aquaculture industry is growing at a rapid rate, and there is great potential for the continued development of the industry. A number of key advances have been made over the past few decades, and there have been significant gains in knowledge as well as production efficiency. Progress in barramundi nutrition, disease management, and larval culture in particular has been rapid over the past decade or so. New directions for the industry as it starts to mature include work on selective breeding and aquaculture in nontraditional areas such as inland saline areas, as well as greater attention to product quality.

7.12

REFERENCES

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Fukuda Y., Nguyen, H.D., Furuhashi, M. & Nakai, T. (1996) Mass mortality of cultured sevenband grouper, Epinephelus septemfasciatus, associated with viral nervous necrosis. Fish Pathology, 31, 165–170. Garcia, L.M.B. (1989a) Dose-dependent spawning response of mature female sea bass, Lates calcarifer (Bloch), to pelleted luteinizing hormone-releasing hormone analogue (LHRHa). Aquaculture, 77, 85–96. Garcia, L.M.B. (1989b) Development of an ovarian biopsy technique in the sea bass, Lates calcarifer (Bloch). Aquaculture, 77, 97–102. Garcia, L.M.B. (1989c) Spawning response of mature female sea bass, Lates calcarifer (Bloch), to a single injection of luteinizing hormone-releasing hormone analogue: effect of dose and initial oocyte size. Journal of Applied Ichthyology, 5, 177–184. Garcia, L.M.B. (1990a) Spawning response latency and egg production capacity of LHRHa injected mature female sea bass, Lates calcarifer Bloch. Journal of Applied Ichthyology, 6, 167–172. Garcia, L.M.B. (1990b) Advancement of sexual maturation and spawning of sea bass, Lates calcarifer (Bloch), using pelleted luteinizing hormone-releasing hormone analogue and 17a-methyltestosterone. Aquaculture, 86, 333–345. Garrett, R.N. & O’Brien, J.J. (1994) All-year-around spawnings of hatchery barramundi in Australia. Austasia Aquaculture, 8(2), 40–42. Ghassemi, F., Jakeman, A. & Nix, H. (1995) Salinisation of Land and Water Resources: Humans Causes, Extent, Management and Case Studies. University of New South Wales Press Ltd, Sydney. Glazebrook, J.S., Heasman, M.P. & de Beer, S.W. (1990) Picorna-like viral particles associated with mass mortalities in larval barramundi, Lates calcarifer Bloch. Journal of Fish Diseases, 13(3), 245–249. Glencross, B. (2006) The nutritional management of barramundi, Lates calcarifer – a review. Aquaculture Nutrition, 12, 291–309. Glencross, B. (2008) A factorial growth and feed utilization model for barramundi, Lates calcarifer based on Australian production conditions. Aquaculture Nutrition, 14, 360–373. Glencross, B., Michael, R., Austen, K. & Hauler, R. (2008) Productivity, carcass composition, waste output and sensory characteristics of large barramundi Lates calcarifer fed high-nutrient density diets. Aquaculture, 284, 167–173. Gomez, D.K., Sato, J., Mushiake, K., Isshiki, T., Okinaka, Y. & Nakai, T. (2004) PCR-based detection of betanodaviruses from cultured and wild marine fish with no clinical signs. Journal of Fish Diseases, 27, 603–608. Grotmol, S. & Totland, G.K. (2000) Surface disinfection of Atlantic halibut Hippoglossus hippoglossus eggs with ozonated sea-water inactivates nodavirus and increases survival of the larvae. Diseases of Aquatic Organisms, 39(2), 89–96. Grotmol, S., Bergh, O. & Totland, G.K. (1999) Transmission of viral encephalopathy and retinopathy (VER) to yolk-sac larvae of the Atlantic halibut Hippoglossus hippoglossus: occurrence of nodavirus in various organs and a possible route of infection. Diseases of Aquatic Organisms, 36, 95–106. Grotmol, S., Dahl-Paulsen, E. & Totland, G.K. (2003) Hatchability of eggs from Atlantic cod, turbot and Atlantic halibut after disinfection with ozonated seawater. Aquaculture, 221, 245–254. Guiguen, Y., Cauty, C., Fostier, A., Fuchs, J. & Jalabert, B. (1994) Reproductive cycle and sex inversion of the seabass, Lates calcarifer, reared in sea cages in French Polynesia: histological and morphometric description. Environmental Biology of Fishes, 39, 1573–5133. Harpaz, S., Hakim, Y., Barki, A., Karplus, I., Slosman, T. & Eroldogan, O.T. (2005a) Effects of different feeding levels during day and/or night on growth and brush-border enzyme activity in juvenile Lates calcarifer reared in freshwater re-circulating tanks. Aquaculture, 248, 325–335. Harpaz, S., Hakim, Y., Slosman, T. & Eroldogan, O.T. (2005b) Effects of adding salt to the diet of Asian sea bass Lates calcarifer reared in fresh or saltwater recirculating tanks, on growth and brush border activity. Aquaculture, 248, 315–324. Harvey, B., Nacario, J., Crim, L.W., Juario, J.V. & Marte, C.L. (1985) Induced spawning of sea bass, Lates calcarifer, and rabbitfish, Siganus guttatus, after implantation of pelleted LHRH analogue. Aquaculture, 47, 53–59. Herbert, B.W., Shaharom, F.M. & Anderson, I.G. (1995) Histopathology of cultured sea bass (Lates calcarifer) (Centropomidae) infected with Cruoricola lates (Trematoda: Sanguinicolidae) from Pulau Ketam, Malaysia. International Journal of Parasitology, 25, 3–13. Howgate, P. (2004) Tainting of farmed fish by geosmin and 2-methyl-isoborneol: a review of sensory aspects and of uptake/depuration. Aquaculture, 234, 155–181.

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8

Abalone Culture

Mark Allsopp, Fabiola Lafarga-De la Cruz, Roberto Flores-Aguilar and Ellie Watts

8.1 INTRODUCTION Abalone is a prized seafood delicacy worldwide. Abalone are marine gastropod molluscs of the family Haliotidae, also called sea snails, ear-shells or sea ears. They possess a single shell, which is a low open spiral structure, and a large muscular foot that is used to attach to hard surfaces. The family Haliotidae contains one genus, Haliotis, and about 100 species are recognised worldwide (Jia & Chen 2001). More information on the biology of abalone can be found in Jia and Chen (2001). Aquaculture activities have grown considerably in the past decade, increasing their contribution to the global market as fisheries continue to decline worldwide. Abalone aquaculture industry has rapidly developed from about 3,000 tons in 2000 to over 40,000 tons in 2008 (FAO 2010). The principal countries producing cultured abalone are China, Korea and Taiwan. Several other countries including Australia, Chile, Mexico, New Zealand, South Africa, Thailand and the United States are also developing abalone aquaculture industries. With the maturity of production lines from farms worldwide the industry has established markets in mainland China through Hong Kong, Japan and Singapore. The demand for cocktail-size abalone has driven the expansion and development of the industry throughout the producer countries. A variety of abalone species are cultivated around the world (see Table 8.1).

8.2

THE ABALONE MARKET

The two largest consumers of wild and cultivated abalone are China and Japan. Generally the Chinese prefer a lighter coloured ‘foot’ and the Japanese a darker one; a characteristic

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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Table 8.1 Abalone species cultivated around the world. Country

Commonly cultivated species

Australia New Zealand China and Taiwan Taiwan Korea

Haliotis laevigata and Haliotis rubra Haliotis iris and Haliotis australis Haliotis discus hannai Haliotis diversicolor Haliotis discus, Haliotis discus hannai, Haliotis diversicolor, Haliotis diversicolor supertexta Haliotis discus hannai Haliotis asinina Haliotis midae Haliotis discus hannai, Haliotis rufescens Haliotis rufescens, Haliotis fulgens, Haliotis corrugata Haliotis rufescens, Haliotis fulgens, Haliotis corrugata Haliotis kamtschatkana Haliotis tuberculata

Japan Thailand South Africa and Namibia Chile Mexico USA Canada Ireland

Production 2008* (tons) 504 8 33,010 348 5,146

NA 30 1,040 515 60 175 NA NA

Note: NA = No data available * Production data obtained from FAO Fisheries and Aquaculture Information and Statistics Service website

that varies between species. The preferred size category is between 200 and 300 g per abalone, but most cultivated abalone are sold at between 50 and 150 g.

8.2.1

Japan

Japan is the largest world consumer of live, fresh and frozen abalone. These product forms are generally identified as having the highest premium on the world market. Because Japan is the largest consumer of premium-quality abalone, Japanese consumer preferences are important in understanding the premium abalone markets. The Japanese native fishery is historically significant and highly valued as a cultural resource. It has given rise to cultural traditions and consumer tastes that make the appearance of an abalone as important as taste and texture when determining the value of the product (Oakes & Ponte 1996).

8.2.2

Mainland China

Mainland China is the largest consumer of abalone, a fact that often remains unrecognised because consumption of abalone in China is almost entirely in the canned form. In regions such as Japan and the USA, canned abalone is generally not considered a premium product. Canned abalone has a traditional place in Chinese society as an item of prestige, often presented as a show of affluence or a demonstration of respect. Considered customary in banquets and traditional feasts, a single can of abalone is often given as a token of respect. The strong traditions surrounding abalone consumption in China have created a stratified market, based on perceived quality differences between popular brand names and countries of origin. The major distribution point for canned abalone destined for mainland China is through Hong Kong (Oakes & Ponte 1996).

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USA

In the USA there is a traditional market for abalone, which is mainly in California, where there was a flourishing fishery until the early 1970s. In the California market tradition, abalone are removed from the shell and sliced into steaks, which are tenderised and then fried. At one time in California, abalone was an abundant, low-cost regional delicacy, but as the fishery dwindled due to over-harvest, constricting supplies caused the market price to increase to a level that has severely restricted demand for the product. The traditional US market now consists primarily of expensive, white tablecloth restaurants in California. The emergence of Asian communities as a significant abalone market in major US metropolitan areas has spurred the demand for specialty food products. This has kindled a demand for Asian-style abalone products in the US market. The market niche is mainly for fresh abalone meat used in Japanese sushi, but a brisk market for live cultured abalone has developed in recent years (Oakes & Ponte 1996).

8.2.4

Southeast Asia

Lucrative markets exist for live abalone in Hong Kong, Taiwan, Singapore, Thailand and other Asian metropolitan centres. The Hong Kong market is the largest and best established of the Asian markets. As well as acting as the gateway to China, Hong Kong offers a direct market for premium abalone products in many forms. Product demand throughout Southeast Asia is based on established markets, which are similar to those in Hong Kong. As Asian affluence increases, these market areas will become a more important market factor. The combined influence of China and Southeast Asia will be significant in determining the location and product concepts best suited for future production sites (Oakes & Ponte 1996).

8.2.5

Europe

Although Europe is not a major market area for cultured or fishery-caught abalone, there is a regional demand arising from the traditional fishery for H. tuberculuta. This market is concentrated in France, but there is some demand throughout the UK and the rest of Europe. This demand is generally under-supplied and could be developed if supplies were available. The European abalone species are small and traditional product presentations are well suited to the smaller (100 g) abalone produced by culturists. Therefore, this region is of great interest for future market expansion.

8.3

ABALONE PRODUCTION TECHNOLOGY

Though culture of abalone has developed in several countries, this section focuses on developments in South America, Australia and New Zealand.

8.3.1

Chile

In Chile, aquaculture is an important income source for the economy. It produced nearly 853,000 tons with a value of US$5.3 billion by 2007, positioning Chile among the top ten

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world aquaculture producers (FAO 2009). Among the aquaculture resources exploited in Chile, abalone was introduced in the late 1970s as a means of diversification, taking into consideration its high commercial value and an unsatisfied demand worldwide (FloresAguilar et al. 2007). Currently, the abalone industry is supported by two foreign species: Haliotis rufescens, red abalone from California and Japanese or ezo abalone, Haliotis discus hannai from Japan. Red abalone was first introduced in 1977, for experimentation in closed systems by Fundación Chile and Universidad Católica del Norte (UCN) (Godoy et al. 1992). Later, in 1982, UCN introduced and adapted the culture technology of the Japanese abalone in collaboration with the Japan International Cooperation Agency (JICA). However, abalone culture technology transfer began only in 1992, when red abalone culture was authorised, in the sea off Chiloé Island in southern Chile, for a subsidiary company of Fundación Chile. Japanese abalone commercial culture started in 1996 in northern Chile where several companies adapted the technology for land-based culture, as abalone seed were provided by the UCN’s Center of Abalone Production. The first official red (1 ton) and Japanese abalone (8 tons) production were registered by the Undersecretary of Fisheries of Chile, in 1998 and 2003 respectively. Exports began in 1999 with 36 tons of red abalone. Currently, there are 19 farms in operation (11 in the north and eight in the south). Seed production is mainly in the northern region, while most growout systems are land-based in the north and in-water in the south of Chile. Since 2002, Chilean legislation has allowed both species to be cultured in land-based semi-closed systems, while red abalone may also be cultured in water-suspended systems between Seno del Reloncavi and Skyring Peninsula in southern Chile (Resolution 30 September 2002). On the other hand, since 2004 the culture of both species has been permitted in the sea but only in two of the three actual culture regions located in northern Chile, and the stock has to be single-sex individuals and sited over a soft substrate area (Subpesca 2006). Currently, Chile is positioned as the fifth abalone producer worldwide with a production volume of 479 tons and an estimated value of US$11.5 million. Red abalone production accounts for 97.5% of total production, as this species has been well adapted to full-cycle culture in northern and southern Chile (Enríquez & Villagrán 2008). On the other hand, Japanese abalone has not adapted well because of its minor resistance to the Chilean culture conditions (i.e. water temperature and type of macroalgae availability) and is actually considered as an emergent species (less than 5 tons a year). Unfortunately, no other abalone species have been introduced, because Chilean legislation allows only these two species to be imported and cultured, and efforts to introduce new exotic species (not endemic) had been laborious, time consuming and unsuccessful. However, research on hybridisation between red and Japanese abalone has proved to be potentially important to diversify and to improve the Chilean abalone industry. Moreover, abalone farming can be considered a young industry, with 70% of the farms just starting the phase of commercialisation and exportation. 8.3.1.1 Conditioning and spawning induction Abalone culture technology in Chile is fully integrated in the northern region, where hatchery, nursery and growout operations are undertaken by most of the farms. Only two of all the southern farms possess all culture phases, and the rest are only in-water growout facilities that are provided with red abalone seeds in the range of sizes of 15–25 mm by northern

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farms. Hatchery facilities are composed of a broodstock area, a spawning area and a larval rearing system. Adult abalone are maintained in a specially designed unit, separated by sexes at stocking biomasses of 25 g/L, in continuously running water at ambient temperature and normal photoperiod 12D:12L. Water is usually filtered up to 25 μm, but some farms use 50 μm. Acceptable water quality parameters are: water temperature 12–20 °C, pH 7.4–8.5, dissolved oxygen 7–10 mg/L, alkalinity 120–180 ppm, salinity 34–36 psu, ammonium 0.0–0.02 ppm, nitrite 0.0–0.2 ppm and nitrate 0.0–2.0 ppm. Feeding rates are around 10–20% of body weight per day, supplied with a fresh mixed macroalgae diet made up mostly of Macrocystis sp. (90%), Lessonia sp., red algae Gracilaria sp. and green algae Ulva sp. Spawning adults normally used are 2 to 6 years old, with a visual gonad index of 2+ to 3+. If hybridisation is desired red abalones of 2–4 years old should be used to improve fertilisation and hatching rates (unpublished data). Spawning induction is usually undergone by chemical stimulation using doses of TRIS-H2O2reactive (Morse et al. 1976) in UV-irradiated water filtered at 1 μm after 1 hour of desiccation at ambient temperature. But temperature and UV induction are also applied in some facilities, normally by raising temperature gradually up to 5 °C at rate of 1 °C /hour. Normally, females are induced 15–30 minutes before males, but if hybridisation is undertaken males should be induced at least 15 minutes before females, to assure sperm availability as the fertilisation window for successful hybrid crosses is less than 20 minutes. At increasing egg age fertilisation rates drop sharply (Lafarga-De la Cruz et al. 2010). Female gametes are collected, and fertilisation is done in 20 L containers with sperm concentrations in the order of 106 sperms/mL for homospecific crosses, and 107 sperms/mL for heterospecific crosses (Lafarga-De la Cruz et al. 2010), and contact times around 2–6 minutes. Fertilised eggs are rinsed several times by decantation with UV-irradiated water. Finally, fertilised eggs are placed forming a monolayer at the bottom of the hatching tanks (50–100 L) and left static overnight, in a controlled-temperature room (17 °C). After 16–18 hours, trocophore larvae hatch out and they are collected and selected (>150 μm) in upwelling tanks for its larval culture period of 5 to 7 days, depending on water temperature. Antibacterial treatments are recommended daily during this period, as well as maintenance activities. Normal larval development is followed daily by microscopic observations. Both closed systems and flow-through systems are used for larval culture, and when the abalone larvae are competent (observation of the third tubule in cephalic tentacles, and characteristic larvae’s foot movements) they are transferred to the post-larval and juvenile tanks. 8.3.1.2

Nursery technology

The nursery facility for rearing abalone from post-larval to juvenile seed size (17–28 mm) is based on the Japanese plastic plate system for larval settlement (Fig. 8.1). Preconditioning of plates is normally with naturally occurring diatoms, but some farms also use cultured microalgae (mainly Ulvella sp., Cocconeis spp. and Navicula spp.). Abalones remain in nursery between 3 and 10 months, depending on the type of culture system used. Land-based systems use abalones of 10–15 mm in shell length into the production system, where they are maintained for 24–48 months. On the other hand, in-water (non-land based) systems use slightly bigger animals (20–25 mm) for grow-out.

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Fig. 8.1

Japanese plate system for post-larval and juvenile culture.

8.3.1.3 Growout technology Abalone farms in the north are characterised by land-based growout operations employing a substantial infrastructure; with many raceway tanks (Fig. 8.2) and integration of all phases of production. The growout tanks are 10 × 1.5 × 0.7 m, made of fibreglass, with a total volume around 11,000 L, with compartments having a conic-shaped bottom to facilitate the cleaning process. They have 6 baskets inside (1.5 × 1.5 m and 0.6 m) (Fig. 8.3), made of plastic mesh (6 mm hole diameter) and plates where the abalone is attached. This makes a total surface of 100 square metres available for the abalone, and the plates hold the organisms off the bottom where the waste debris from the abalone and algae is accumulated. A 1 mm thick HDPE plastic plate covers each basket and weights are added on to keep the shelters inside the water. Ambient temperature seawater is used in a flow-through system. The seawater exchange rate is 9 tons per hour, and filtered seawater to 90 μm is used. Air is pumped to each tank constantly. The main food for growing out abalone is brown algae. Three brown algal species are normally used: Lessonia trabeculata, Lessonia nigrescens and Macrocystis integrifolia, with L. trabeculata being the most abundant. In the north, there is a regulation that only registered companies may harvest kelp and they have to comply with scientific management regulations in order to maintain the sustainability of the resource. Most of this kelp is

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Fig. 8.2

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Raceways growout culture system (P. Camanchaca Company, Caldera Chile).

harvested at low tide and is cut with a knife by fishermen holding contracts with these companies. The capacity of abalone per basket and shelter is constant and the number depends on the abalone size. Two times a week the abalone are fed ad libitum, and the cleaning of the tank depends on the time of the year and the algal feed but is normally once a week. The tank is emptied and the tank surfaces are scrubbed with a brush and then refilled with fresh seawater. Most of the land-based abalone aquaculture farms in Chile monitor water quality as temperature and oxygen once a day, and salinity and phytoplankton and bacteria at least once a month. In compliance with the regulations of the federal agencies (Decreto Supremo No. 90/1996. Ministry of Economy), levels of suspended solids, oxygen, ammonia and temperature amongst other factors have to be continuously monitored. The exact monitoring requirements and their frequency varies, however, according to the size and location of the farm and type of feed used. In the south of the country, sea-based growout systems are widely used. The small farmers use the barrel culture system (Fig. 8.4), and only the larger production companies with inventories over 2 million abalone use cage-based growout systems. These are either plastic moulded cages or iron-galvanized structures covered with netting. The cage size is normally 2 × 1 × 1 m, but the most advanced cages are 3 × 1 × 1 m. These cages (Fig. 8.5) have vertical plates as surface for the abalone to attach. In these plates a maximum capacity of 65% of the total surface area of abalone is allowed.

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Fig. 8.3

Baskets with shelter plates for juvenile culture.

Fig. 8.4

Barrel abalone growout system.

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Fig. 8.5

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Abalone growout cage with its HPDE plastic plates.

These containers are suspended in a typical long line, and the kelp Macrocystis pyrífera is the most widely used algal species to feed abalone. To operate the culture containers the bigger farms use barges with a hoist to lift the cages. The Macrocystis feed is harvested from small boats and cut with a knife. On the small farms staff collect the seaweed manually, while the bigger companies pay local fishermen to supply the algae. Macrocystis is abundant in summer, but almost disappears in winter, forcing farmers to purchase cultured red algae, Gracilaria chilensis. While there is constant supply of cultivated Gracilaria in the south, the growers claim that Macrocystis produces much better abalone growth rates. No kelp harvest permits are required in the southern regions. Artificial feeds are used in some phases of the abalone growth, especially in land-based farms in the north, in both nursery and growout operations. At some land-based farms, abalone of all sizes receive a combination of artificial and kelp diet. No manufactured diets are used on the in-water farms because environmental regulations restrict aquaculture operations using formulated feeds. As a result of the large-scale salmon culture in the region, aquaculture operations using pelleted feeds are deemed ‘intensive’ farms and sea concessions will only be granted if they are a minimum of 2.8 km from neighbouring concessions. This makes it very difficult to find suitable areas for abalone culture that comply with this regulation. As a result all farms in the south use seaweeds as feed. One company in the north has experimented with a recirculation system for more than five years and it is proving very successful.

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8.3.2

Australia

In 2008, abalone aquaculture emerged as one of the fastest-growing agribusiness sectors in Australia. With 850 tons produced in 2006/7, worth AU$42.5 million to the Australian economy, this is estimated to grow to 1,500 tons over the next five to ten years, worth $75 million (Fleming 2008). 8.3.2.1 Conditioning and spawning technology Until recently it had been assumed that Australian abalone farmers have found it far more reliable to collect conditioned animals from the sea rather than condition them in tanks (Fleming & Roberts 2001). However, high-quality gametes can now be obtained in winter from H. laevigata held in a flow-through broodstock system developed by researchers from the Western Australian Fisheries Department and industry (Freeman et al. 2006). Water temperature is considered the main exogenous factor that regulates the reproductive cycle of abalone (Landau 1991; Hone et al. 1999; Fleming & Roberts 2001; Maguire 2001; Plant et al. 2003). Fleming and Roberts (2001) indicated that for H. rubra temperatures between 15 and 17 °C are optimal. In a study undertaken at Ocean Wave Seafoods farm at Lara, Victoria, Plant et al. (2003) determined that H. rubra can be brought into spawning condition when kept at constant temperature (18 °C). The best spawning results were achieved after 120 days, when about 2 million eggs were spawned per female in 60% of those tested, with a 75% fertilisation rate. The results clearly showed that conditioning at a constant, increasing temperature delivers an increase in spawning success. The researchers believe the process should be applicable to other species. Fleming and Roberts (2001) found that temperatures between 17 and 19 °C were optimal for conditioning H. laevigata. In Western Australia, Freeman (2001) has found that H. laevigata can be spawned all year round when water temperatures in the conditioning room ranged from 14.2–19.23 °C. The most successful spawning events with highest egg productions occurred out of the ‘natural spawning season’ in most groups. Spawning events during this period can be highly beneficial to farmers as they can take advantage of the enhanced growth of juveniles during the early summer months. Animals can be weaned off the plates before the highest summer temperatures occur. High water temperature during the weaning process can cause high mortalities in some regions of South Australia (Maguire 2001). Abalone can be stimulated with a single stimulus, or a combination of stimuli including temperature changes, treating seawater with UV, ozone or hydrogen peroxide, handling animals, or exposing them to air, depending on the species of interest (Maguire 2001). Hone et al. (1999) outlined the procedure for spawning H. laevigata as follows: place the abalone in clear aquaria and reduce the light and noise levels. After a few hours of acclimation in ambient temperature water, activate the UV filter. If after 5 hours there is no activity, further stimulate the abalone by placing an immersion heater in the tanks and rapidly raise the temperature by 3–5 °C. 8.3.2.2

Hatching

Fertilised eggs from H. laevigata hatch after approximately 16 hours at 18 °C and are called trochophore larvae. Newly hatched trochophore larvae swim to the surface and then can be easily separated from the unhatched eggs and discarded egg cases by decanting off the

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top water layer into a clean tank for subsequent larval rearing (Maguire 2001). Abalone Farms Australia (AFA) in Tasmania use a system where the larvae hatch from the negatively buoyant eggs on the bottom of a tank and swim to the surface of the tank where there is a small weir on the sides that leads directly to the larval rearing tanks. This system requires minimal labour (Cropp, pers. comm. 2002). The non-feeding larvae develop over about five days at 17 °C to 18 °C and about four days at 20 °C. Densities are kept at less than 25/ ml to ensure the highest water quality, and to reduce the chances of bacteria growing on the tank surfaces (Maguire 2001). Survival rates of over 80% are common if proper care is taken during larval rearing (Fleming & Roberts 2001). 8.3.2.3 Nursery phase Towards the end of larval development, the larvae sink to the bottom of the container and begin exploring for a suitable surface for settlement (Maguire 2001). Benthic biofilm consisting of bacteria and mixed species of diatoms growing on PVC settlement plates have traditionally been used as a settlement substrate in abalone nurseries worldwide. This process is unpredictable and larval settlement rates can be low (1–10% of larvae) (Daume 2003). Enhanced settlement up to 80% has been obtained in small-scale experiments through the use of the non-geniculate coralline red alga, Sporolithron durum (Daume 2003). Currently naturally developing diatom films on plastic plates are predominantly used as the settlement cue in abalone nurseries in Australia. Some diatom species produce better settlement than others; for example Daume (2003) found that H. laevigata settled particularly well on the diatom Navicula ramosissima. Isolating particular diatom species and growing them in monoculture before inoculating settlement tanks in the nursery provides greater control. This practice has not, however, been embraced by the industry because it is believed that the gain does not justify the extra costs involved in the scale-up diatom culture site. Roberts and Lapworth (2001) explain that some diatom species are not good for settlement, and strains that are excessively mobile or form 3-dimensional colonies can prevent successful settlement. Therefore con-specific substrate films may play a significant role in increasing settlement rates. Hatcheries in Japan culture the microalgae Ulvella lens to improve settlement of the Japanese abalone Haliotis discus hannai (Takahashi & Koganezawa 1988). Takahashi and Koganezawa (1988) reported settlement rates of 67% on U. lens, which was not previously grazed on by juvenile abalone. Pre-grazed U. lens yielded a settlement rate of 93–100% (Takahashi & Koganezawa 1988; Seki 1997). The first investigation of the settlement of Australian abalone species on U. lens was conducted by Daume (2003), who found that settlement for H. rubra was higher on U. lens than on diatom films. This study suggested that settlement plates seeded with U. lens could induce high and consistent settlement of H. rubra. In addition, H. laevigata settled well on U. lens compared to some diatom films. But more experiments are needed to explore the abalone growth and settlement rates achieved on different diatom species and cell densities (Daume 2003). Settlement and metamorphosis occur typically within one to three days after the larvae are introduced to the settlement tank (Maguire 2001). The transition from a free-swimming larva to a juvenile, living permanently on a hard surface, is a critical phase in the life of an abalone and mortality can be very high (∼90%) although the technology is improving (Daume 2003).

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8.3.2.4 Growout phase The weight of the meat compared to the length of the shell is referred to as the meat weight: shell length ratio. This ratio is important for farmers, because though the product is normally sold as net meat weight, shell length is normally used to assess the size of abalone. Therefore, the aim of growout phase for farmers is to produce abalone with greater meat content. Freeman (2001) found that abalone reared at high densities have a higher meat to shell length ratio than those cultured at lower densities. In Australia, abalone is usually cultured on land using tanks, troughs or raceway systems. Additionally, abalone can be reared in barrels or sea-cages hanging from buoys or rafts (Maguire 2001). There is currently much greater emphasis on land-based growout systems than sea-based systems in Australia. A wide range of system designs are currently implemented in the country, ranging from deep to shallow and round to rectangular tanks. An early land-based system in South Australia for H. laevigata culture used a ‘surfboard’ design, which was shallow concrete tanks/raceways, usually with a length of approximately 2 m. These small ‘surfboard’ raceway tank systems are usually sited indoors or in shade cloth enclosures. The ideal slope for a raceway is 1:100, which helps to prevent mass mortalities in the event of a pump failure by allowing the tanks to drain but also allowing the bottom surface to remain wet. Shelters (hides) are not used since they disrupt the water flow (Freeman 2001). There is also a culture system based on much larger concrete tanks using the same principles of shallow water depth and high water exchange. It employs a shaded outdoor system similar to the ‘surfboard’ system. This is also currently being used in South Australia. Some growers have successfully adapted the typical ‘Taiwanese’ culture method of deep tanks with strong aeration and numerous shelters for refuge (Forster 1996). These tanks are designed so they can be drained regularly to remove solid wastes. They seem particularly well suited to H. rubra, but are also used for H. laevigata (Maguire 2001). A farm in Tasmania currently operates this system successfully to cultivate H. laevigata. Annual survival rates can be 80–95%, with growth rates of 20–30 mm a year in landbased systems for H. laevigata in the southern regions of Australia (Fleming 1995). These rates are not always achieved, however, as growth rates vary significantly with the seasons (Fleming & Roberts 2001). Barrel or cage culture for abalone offers a low capital cost alternative, but can have high maintenance costs (Forster 1996). The barrels or cages can be hung from longlines supported by buoys or attached to rafts and large cages that can be placed on the ocean floor, provided that precautions are taken to prevent predation by crabs and starfish. Supplying feed to submerged cages has been simplified by the development of a surface feeding system that pumps feed from a surface vessel into the rafts or cages on the ocean floor (O’Brien 1996a,b). The highest growth rates achieved in a trial involving a range of locations, culture systems (land- and sea-based) and species in Western Australia was with H. laevigata grown in barrels at Albany (Fleming & Roberts 2001). The water flow rate is critical because it must be sufficient to encourage feeding behaviour, maintain dissolved oxygen levels and remove wastes, but not so fast as to wash away the feed before it can be consumed. Stocking density and feed type are important factors when using this culture system. In general, higher stocking densities in land-based systems,

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with most of the floor area covered by abalone, encourage more uniform distribution of H. laevigata. Throughout the growout phase, abalone densities are regularly reduced and size grading is carried out (Edwards et al. 1999). Biofouling can greatly increase the maintenance costs of production systems and can directly smother the abalone by covering the respiratory pores. New Australian technology aimed to reduce biofouling on molluscs, and plastic mesh based growout systems, may be critical for the success of sea-based abalone farming in Australia (Maguire 2001).

8.3.3

New Zealand

Paua (Māori name) is the common name for abalone in New Zealand. Three species of abalone occur naturally in New Zealand; black foot paua (Haliotis iris) (Fig. 8.6) yellow foot paua (Haliotis australis) and white foot paua (Haliotis virginea). Black foot paua (Fig. 8.8) is the largest abalone species in New Zealand and is most commonly found in shallow waters at depths less than 6 m all around mainland New Zealand, Stewart Island and the Chatham Islands. They often form large clusters in the sub-littoral zone on open, exposed coasts where drift seaweed accumulates and there is good water movement. Black foot paua grow to about 180 mm in shell length (the legal size for wild harvesting is 125 mm, measured as the longest shell length).

Fig. 8.6

Paua abalone.

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Fig. 8.7

Trays with black foot paua abalone.

The New Zealand paua farming industry had its beginning in the mid-1980s when hatchery techniques were developed for the New Zealand species. Paua farms in New Zealand have traditionally been land-based and configured to operate on flow-through water supply where the water is pumped from the sea, over the paua and then allowed to return to the sea. Recirculation technology has gained favour in recent years and offers several advantages over flow-through by improving performance, raising efficiencies, reducing costs and reducing risk. These advantages include maintaining consistent rearing temperatures, protecting farms from fluctuating environmental conditions and improving overall biosecurity. Although the trend is towards recirculation, at least two farms currently operating use sea-based barrel culture technology; however, the focus for these is on the production of paua pearls, which employs a more extensive approach using freshly harvested seaweeds. Early efforts on paua commercial production used seaweeds for feed. However, now most land-based farms rely on manufactured pellet feeds because of their convenience, consistency of food quality, difficulties in obtaining sufficient quantities of seaweed and the additional benefit of reducing the black pigment in the foot. Only black foot paua are currently farmed in New Zealand. This is mainly because of the larger size of the black foot compared with the other two New Zealand species, and their iridescent shell coloration, utilised for the production of cultured paua pearls. Paua

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aquaculture has excellent potential for New Zealand due to strong overseas demand and high prices attained for live meat (more than NZ$60 per kg). Currently, the industry in New Zealand is based around 11 farms, mostly onshore systems that on-grow hatchery-reared juveniles to market size for their meat. The largest of these is OceaNZ Blue Ltd (OBL) in Northland, which produces the bulk of New Zealand’s farmed abalone. OBL is on track to become the first 100 ton production facility in New Zealand and operates using a semi-recirculating system. Two paua farms are focusing primarily on the production of juvenile paua for the purpose of reseeding to enhance coastal areas depleted of wild stocks. This area of production is expected to gain momentum as enhancement efforts increase and the potential to identify reseeds from wild stock are realised through the development of molecular markers.

8.4 8.4.1

TECHNOLOGICAL DEVELOPMENTS Polyploid induction

Chromosome set manipulation in molluscs has received wide attention in the past two decades. Research has primarily focused on the induction and evaluation of triploidy in bivalve species of commercial importance (Beaumont & Fairbrother 1991; Nell 2002). The principal value of triploids for aquaculture arises from their sterility, presumably because of failure in synapse of the three sets of chromosomes during meiosis. Sterility may lead to faster growth of (adult) triploids owing to energy reallocation from reproduction to somatic growth. Sterility may also result in better meat quality of triploids in association with their reduced spawning activities (Beaumont & Fairbrother 1991). Triploid animals are produced, by inhibiting the release of the first or second polar body, through the application of a chemical or environmental stress soon after fertilisation. The retention of an extra set of chromosomes within the developing embryo results in an animal containing three sets of chromosomes per cell (triploid) instead of the usual two sets (diploid). Triploidy can be induced using multiple techniques. 8.4.1.1 6- dimethylaminopurine exposure Fertilised eggs can be exposed to 100 μM solution of 6-DMAP (6-dimethylaminopurine) between 15 and 20 minutes post-fertilisation. Following a 15-minute exposure to 6-DMAP, the eggs are rinsed and placed in hatching trays. 8.4.1.2 Cytochalasin B (CB) exposure Induction of triploidy has been achieved applying CB at a concentration of 0.70 mg/L after 27 minutes for a duration of 10 minutes. Eggs are then rinsed and placed in hatching trays. 8.4.1.3 Temperature shock Exposure of fertilised eggs to 3 °C at 15 minutes post-fertilisation for duration of 10 minutes and/or exposure to 26 °C water 15 minutes post-fertilisation has induced triploidy in various abalone species.

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8.4.2

Hybridisation

Intraspecific (crossbreeding) and interspecific hybridisation within the family Haliotidae can be used as a strategy to improve the profitability of abalone farming. An effort by farmers to reduce production costs rather than increase the price of their product may become a more realistic method of increasing profitability. It would be expected that hybrids that display positive characteristics of heterosis would be an integral part of such an effort. However, as pointed out by some authors, if hybrids are to be used for aquaculture, it is fundamental that they are made sterile or that efficient methods to prevent escape are developed, because of the abalone’s established ability to produce offspring among them (F2 progeny) and backcrossing with parental species (gene introgression risk). Escape could lead to a possible genetic impact of hybridisation on natural abalone populations. Hybrids often have superior characteristics than their parents, such as greater in size, resistance to disease, number of offspring produced. Heterosis, or hybrid vigour, is a term used to describe the phenomenon in which the performance of an F1 hybrid, generated by the crossing of two genetically different individuals, is superior to that of the better parent (heterobeltiosis) or average between both parents (middle parent heterosis). Heterosis has been extensively used in breeding programmes of aquaculture resources as a way for genetic improvement of cultured species (Hulata 2001). There are two major hypotheses that have been promulgated to explain heterosis: the dominance hypothesis and the overdominance hypothesis. The dominance hypothesis suggests that heterosis is due to cancelling of deleterious recessives contributed by one parent, by dominant harmless alleles contributed by the other parent in the heterozygous F1 (Jones 1917). On the other hand, over-dominance hypothesis assumes that the heterozygous combination of the alleles at a single locus is superior to either of the homozygous combinations of the alleles at that locus (Shull 1908). As already pointed out in some research, intra-specific cross breeding has produced positive heterotic hybrid abalones. With regard to the analysis of the genetic change in parent abalone and their hybrids, Wan et al. (2001) obtained inter- and intra-population similarity indexes and genetic distances by Random Amplified Polymorphic DNA (RAPD) analysis of H. discus hannai and H. discus discus, and their reciprocal hybrids. From the 113 bands of RAPD analysis some were special (diagnostic) for each kind of abalone and some others were common between two and three of the kinds. Results of inter-population similarity indexes showed greater variation in hybrids, whereas genetic distances between the hybrids and the two parental species differed, being more similar to H. discus discus for both hybrid crosses. Later on, Wan et al. (2004) continued their comparative studies on genetic diversities among these two parental species and their hybrids, in order to understand better the process of heterosis, using AFLP as molecular markers. Results showed significant differences between parental populations in 88 out of 552 loci found. In hybrids, more loci with lower frequency were amplified, whereas those with 0 and 100% frequency were less amplified when compared with parents. Lower similarity indexes and higher heterozygocity in hybrids resulted in an increased genetic diversity of hybrids. And as noted before with RAPD analysis (Wan et al. 2001), the genetic distances between reciprocal F1 generations and H. discus discus were both smaller than those between hybrids and H. discus hannai. Recently, Liu and Zhang (2007) conducted a genetic study using AFLP markers to analyse the F1 intraspecific hybrid family of H. discus hannai from two distinct geographical populations. Results showed fragments exclusively for females, males and certain others common for both, some segregating in agreement with Mendelian 1:1 ratio and other with 3:1 ratio,

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showing ratios of segregation distortion of 27.4, 25.9 and 33.9%, respectively. The authors hypothesised that segregation distortion may be associated with the incompatibility of genes between the two populations of H. discus hannai used in this study (Liu & Zhang 2007). Aquaculture has produced hybrids to improve growth and survival in the past. The yabby industry found that the crossing of female Cherax rotundus and male Cherax albidus produces only male progeny that grow faster than mixed sex groups of C. albidus. The results of trials under model pond conditions found the hybrid to have a final harvest value 4.6 times greater than that of C. albidus yabbies (Lawrence 2004). This hybrid offers considerable potential to increase the profitability of yabby farming. Hybrid striped bass are currently cultured in the United States and other countries, including Taiwan, Israel and Italy. Hybrid striped bass are a cross between striped bass, Morone saxatilis and white bass, M. chrysops. When the female parent is a striped bass the hybrid is called a palmetto bass; when the female is a white bass the hybrid is called a sunshine bass. Ludwig (2004) found that at 5 days of age the palmetto bass are 6 to 9 mm long, while those of sunshine bass are only 3 to 5 mm in length. Landau (1991) in an experiment using a hybrid from male blue catfish and female channel catfish, grown for 220 days in earthen ponds, found that the harvest of the hybrids was 13.5% greater than that of channel catfish. Additionally, the hybrids were more easily captured by seining, were more uniform in size and had a greater dress-out percentage (weight after the head and skin were removed and it is eviscerated, divided by the live weight of the fish). Heterotic effects in the hybridisation of abalone have been reported as increased growth and survival (Leighton & Lewis 1982; Koike et al. 1988) compared to those of the parent species. Hoshikawa et al. (1998) studied the heterotic effect of hybridisation on growth. It was only observed at high water temperature (18 °C) in the form of superior growth rates and not at the lower water temperature (8 °C) in the cross between pinto abalone, Haliotis kamtschatkana and ezo abalone, H. discus. The daily shell growth was significantly higher in the hybrid at 18 °C with 33.4 to 55.6 μm/day compared to 6.5 to 10.9 μm/day for pinto abalone and 31.2 μm/day for ezo abalone (Hoshikawa et al. 1998). Heterotic effect in growth and survival can also be observed at ambient temperature as demonstrated by Wang and Jiachun (1999), who found the growth and survival rates in the F1 offspring from the cross breeding of red abalone H. rufescens and pacific abalone H. discus hannai were higher than those of H. discus hannai. Juveniles of the hybrid between Haliotis rufescens and Haliotis fulgens, and also between H. rufescens and Haliotis sorenseni displayed superior growth at ambient temperature than the parent species (Hoshikawa et al. 1998). Hybrid abalone have been produced in Australia by crossing female Halitosis rubra and male H. laevigata abalone (Freeman 2001). These individuals have been produced in an attempt to find an abalone that has the best characteristics in terms of growth rate, meat to shell ratio, meat texture and market appeal. The ‘tiger ’ abalone which is a cross between a female H. laevigata and a male H. rubra, is commercially cultured in Tasmania by Tas Tiger Pty Ltd. Hone et al. (1999) indicated the tiger abalone has significant market value. A hybrid between H. laevigata and H. scalaris has recently been produced at SAM abalone in Port Lincoln, South Australia. It has shown a faster growth rate during the months between December to March than that of both parent species. Fig. 8.8 is a photograph of H. scalaris, the hybrid, and H. laevigata and clearly demonstrates the size difference. This particular hybrid cross has also been found to occur in the wild. The Western Australian Fisheries Department has specimens that have been found in the south west of Australia (Fig. 8.9).

Fig. 8.8

Left to right: H. scalaris, H. laevigata × H. scalaris (hybrid) and H. laevigata.

Fig. 8.9 Wild H. laevigata, H. laevigata × H. scalaris (yybrid) and H. scalaris found in Western Australia. (Please see plate section for colour version of this figure.)

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249

FUTURE POSSIBILITIES

In South America, the biggest constraint for the industry is considered to be the supply of abalone feed. Some farmers are searching for a better supply of algae, including culturing Macrocystis pyrifera as the solution while some others are experimenting with manufactured diets of their own production or from domestic and foreign feed-producing mills. In New Zealand, prohibitive harvesting costs and limits on harvesting have resulted in abalone farmers using artificial feed pellets. Since abalone needs to actively search for static artificial food, studies are being undertaken to improve the attractiveness of such feeds by the addition of chemical attractants, feeding stimulants or dried seaweed fragments (Allen et al. 2006). On the other hand, artificial diets have been considered expensive and unsuitable for ocean-based longline culture of abalone and suitability of a variety of seaweeds is being investigated (Qi et al. 2010). Abalone production systems need to optimise culture configurations to improve productivity. In Chile, efficiencies of basket and lantern nets suspended in seawater tanks were compared for growth and survival of juvenile abalone (Pereira & Rasse 2007). Lantern systems were found to have a larger carrying capacity while occupying less water column space. The lantern nets provided better growth, and were more economical and easier to handle (Pereira & Rasse 2007). Recirculating aquaculture systems (RAS) are considered to have reduced environmental impacts. Installation of baffles within the RAS culture tanks has been trialled to allow high-density culture (Park et al. 2008). Recently, Integrated multi-trophic aquaculture (IMTA) involving integration of fed species and extractive species been developed and is gaining recognition as a sustainable approach to aquaculture (Nobre et al. 2010). In South Africa, IMTA has been implemented using seaweed Ulva lactuca and abalone Haliotis midae, where the algae take up the dissolved inorganic nutrients from the system and the produced algal biomass provides renewable feed for the other cultivated species, abalone (Nobre et al. 2010). Another major challenge for the industry is to diversify the market. Most abalone were sold in Japan in the past but now Chinese, United States and European companies are making big efforts to develop the market and specifically the presentation of canned abalone.

8.6

REFERENCES

Allen, V.J., Marsden, I.D., Ragg, N.L.C. & Gieseg, S. (2006) the effects of tactile stimulants on feeding, growth, behaviour and meat quality of cultured blackfoot abalone, Haliotis iris. Aquaculture, 257, 294–308. Beaumont A.R. & Fairbrother J.E. (1991) Ploidy manipulation in molluscan shellfish: a review. Journal of Shellfish Research, 10, 1–18. Cropp, M. (2002) Abalone Farms Australia Pty. Ltd. Personal Communication, Bicheno Tasmania, (Owner/ Manager). Daume, S. (2003) Early life history of abalone (Haliotis rubra and H. laevigata): Settlement, survival and early growth, Fisheries Research Contract Report. Department of Fisheries, Perth, Western Australia. Edwards, S., Ralph, J. & Geraghty, D. (1999) Movement of gut contents in blacklip abalone (Haliotis rubra) is not mediated by peristalsis. In: Proceedings of the 6th Annual Abalone Aquaculture Workshop, April, 1999, NSW (ed. P. Hone), pp. 102–111. Fisheries Research and Development Corporation, Sydney.

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Enríquez, R. & Villagrán, R. (2008) La experiencia del desarrollo del cultivo de abalón (Haliotis spp.) en Chile: Oportunidades y desafios. Revue Scientifique et Technique (International Office of Epixootics), 27, 103–112. Fleming, A. (2008) Global sustainability targets of Australian abalone farmers. Accessed July 2010. Available from http://www.thefishsite.com/articles/567/global-sustainability-targets-of-australianabalone-farmers). Fleming, A.E. (1995) Growth, intake, feed conversion efficiency and chemosensory preference of the Australian abalone, Haliotis rubra. Aquaculture, 132(3–4), 297–311. Fleming, A.E. & Roberts, R. (eds) (2001) Proceedings of the 7th Annual Abalone Aquaculture Workshop (Australasian abalone aquaculture Conference, Dunedin, New Zealand). FRDC Abalone Aquaculture Subprogram, Williamstown Victoria. Flores-Aguilar, R., Gutiérrez, A., Ellwanger, A. & Searcy-Bernal, R. (2007) Development and current status of abalone aquaculture in Chile. Journal of Shellfish Research, 26, 705–711. Food and Agriculture Organisation of the United Nations (FAO) (2009) World aquaculture production. FAO Fisheries and Aquaculture Department, online database. Available from http://www.fao.org/ fishery/statistics/global-aquaculture-production/en. Food and Agriculture Organisation of the United Nations (FAO) (2010) FAO Fisheries and Aquaculture Information and Statistics Service. Available from http://www.fao.org/figs/servelet. Forster, A. (ed.) (1996) Proceedings of the Abalone Aquaculture Workshop, December (1995), Albany, Western Australia. Aquaculture Development Council and Fisheries Department of Western Australia. Freeman, K.A. (2001) Aquaculture and related biological attributes of abalone species in Australia – a Review. Fisheries Research Report, 128, Fisheries Western Australia, Perth. Freeman, K.A., Daume, S., Rowe, M., Parsons, S., Lambert, R. & Maguire, G.B. (2006) Effects of season, temperature control, broodstock conditioning period and handling on incidence of controlled and uncontrolled spawning of greenlip abalone (Haliotis laevigata Donovan) in Western Australia. Journal of Shellfish Research, 25(1), 187–194. Godoy C., Jerez, G. & Ponce, F. (1992) The introduction of abalone into Chile. In: Abalone of the World: Biology, Fisheries and Culture (eds S.M. Tegner & S. Guzmán del Próo) pp. 485–488. London, Blackwell Scientific Publishers. Hone, P.W., Madigan, S.M. & Fleming, A.E. (1999) Abalone hatchery manual for Australia. South Australian Research and Development Institute, Adelaide. Hoshikawa, H., Sakai, Y. & Kijima, A. (1998) Growth characteristics of the hybrid between pinto abalone, Haliotis kamtschatkana Jonas, and ezo abalone, H. discus hannai Ino, under high and low temperature. Journal of Shellfish Research, 17(3), 673–677. Hulata, G. (2001) Genetic manipulations in aquaculture: a review of stock improvement by classical and modern technologies. Genetica, 111, 155–173. Jia, Jiansan & Chen, Jiaxin (2001) Sea farming and sea ranching in China. FAO Fisheries Technical Paper T418. Jones, D.F. (1917) Dominance of linked factors as a means of accounting for heterosis. National Academy of Science (USA) 3, 310–312. Koike, Y., Sun, Z. & Takashima, F. (1988) On the feeding and growth of juvenile hybrid abalones. Suisanzoshoku, 36(3), 231–235. Lafarga-De la Cruz, F., Aguilera-Muñoz, F., Díaz Pérez, N. & Gallardo-Escaráte, C. (2010) Hatchery performance of interspecific hybrids between California red abalone (Haliotis rufescens) and Japanese abalone (H. discus hannai) to diversify Chilean abalone aquaculture. Aquaculture (submitted). Landau, M. (1991) Introduction to Aquaculture. Benjamin/Cummings Publishing Company, Inc. Lawrence, C.S. (2004) All-male hybrid (Cherax albidus × Cherax rotundus) yabbies grow faster than mixed-sex (C. albidus × C. albidus) yabbies. Aquaculture, 236(1–4), 211–220. Leighton, D.L. & Lewis, C.A. (1982) Experimental hybridisation in abalones. International Journal of Invertebrate Reproduction, 5, 273–282. Liu, X. & Zhang, G. (2007) Genetic analysis of segregation distorsion of AFLP markers in an F1 population of the Pacific abalone. Marine Sciences, 31(10), 70–76. Ludwig, G.M. (2004) Tank culture of larval sunshine bass, Morone chrysops (Rafinesque) × M. saxatilis (Walbaum), at three feeding levels, (2003), Aquaculture Research, 34(14), 1277–1285. Maguire, G.B. (2001) Farming Abalone. Fisheries Western Australia, 7, 1–8. Morse, D.E., Duncan, H., Hooker, N. & Morse, A. (1976) Hydrogen peroxide induces spawning in mollusks, with activation of prostaglandin endoperoxide synthetase. Science, 196, 298–300.

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Nell, J.A. (2002) Farming triploid oysters. Aquaculture, 210, 69–88. Nobre, A.M., Robertson-Anderson, D., Neori, A. & Sankar, K. (2010) Ecological-economic assessment of aquaculture options: comparison between abalone monoculture and integrated multi-trophic aquaculture of abalone and seaweeds. Aquaculture, 306, 116–126. O’Brien, D. (1996a) Preliminary analysis of the production costs and returns involved in abalone farming. In: Proceedings of the Abalone Aquaculture Workshop, December, 1995, Albany, Western Australia (ed. A. Forster), pp. 67–83. Aquaculture Development Council and Fisheries Department of Western Australia. O’Brien, D. (1996b) Growing abalone at sea – the latest ideas from Tasmania and further afield. In: Proceedings of the Abalone Aquaculture Workshop, December, 1995, Albany, Western Australia (ed. A. Forster), pp. 28–35. Aquaculture Development Council and Fisheries Department of Western Australia. Oakes, F.R. & Ponte, R.D. (1996) The abalone market: Opportunities for cultured abalone. Aquaculture, 140(1–2), 187–195. Park, J., Kim, H-B., Kim, P-K. & Jo, J-Y. (2008) The growth of disk abalone, Haliotis discus hannai at different culture densities in a pilot-scale recirculating aquaculture system with a baffled culture tank. Aquacultural Engineering, 38, 161–170. Pereira, L. & Rasse, S. (2007) Evaluation of growth and survival of juveniles of the Japanese abalone Haliotis discus hannai in two culture systems suspended in tanks. Journal of Shellfish Research, 26, 769–776. Plant, R., Mozquiera, A., Day, R. & Huchette, S. (2003) Conditioning and spawning blacklip abalone (Haliotis rubra). Proceedings from the 9th annual abalone aquaculture workshop, Queenscliff, Australia. Qi, Z., Liu, H., Li, B., et al. (2010) Suitability of two seaweeds, Gracilaria lemaneiformes and Sargassum pallidum, as feed for the abalone Haliotis discus hannai Ino. Aquaculture, 300, 189–193. Roberts, R.D. & Lapworth, C. (2001) Effect of delayed metamorphosis on larval competence, and postlarval survival and growth, in the abalone Haliotis iris (Gmelin). Journal of Experimental Marine Biology & Ecology, 258, 1–13. Seki, T. (1997) Biological studies on seed production of northern Japanese abalone. Bulletin of the Tohoku National Fisheries Research Institute, 59, 1–71. Shull, G.H. (1908) The composition of a field of maize. American Rare Breed Association, 4, 296–301. Subpesca (2006) Subsecretaría de Pesca. Resolución Final No. 4282. 14 de Diciembre del 2005, Valparaíso, Chile. Takahashi, K. & Koganezawa, A. (1988) Mass culture of Ulvella lens as a feed for abalone Halitosis discus hannai. NOAA Technical report NMFS, 70, 25–36. Wan, F., Bao, Z., Zhang, Q. & Wang, X. (2004) Comparative Studies on the Molecular Genetic Diversities among Haliotis discus hannai, H. discus discus and their hybrids. High Technology Letters, 10(3), 93–96. Wan, J.X., Wang, J., Pan, B., et al. (2001) RAPD analysis of the genetic change in parent abalone and their hybrids. Periodical of Ocean University of China, 31(4), 506–512. Wang, R. & Jiachun, F. (1999) Artificial breeding of red abalone, Haliotis rufescens, and cross breeding with Pacific abalone, H. discus hannai Ino. Journal of Dalian Fisheries College/Dalian Shuichan Xueyuan Xuebao, 14(3), 64–66.

9

Seaweed Culture with Special Reference to Latin America

Julieta Muñoz, Vivek Kumar and Ravi Fotedar

9.1 INTRODUCTION Seaweeds, also referred as macroalgae, are macroscopic multicellular marine photosynthetic organisms usually classified into three groups: Rhodophyta (red algae), Phaeophyta (brown algae) and Chlorophyta (green algae) (Lobban & Harrison 1994). Macroalgae are benthic organisms with a cosmopolitan distribution within the photic zone of the oceans (Lüning 1990) that play an important role in marine communities as primary producers and in the transference of energy to higher trophic levels of aquatic ecosystems (Dawes 1998). As components of aquatic ecosystems, macroalgae reflect the health of the environment through their population parameters such as density and abundance (Stevenson et al. 1996). In addition, macroalgae also provide shelter and substratum for other organisms, creating microhabitats and nursery areas (Chapman et al. 1987). For additional information the reader is referred to the excellent review by Chopin and Sawhney (2009), particularly on the significance of seaweeds in coastal ecosystems. This chapter also includes a review of current and potential seaweed cultivation in Latin America.

9.2

SEAWEED UTILISATION

From the commercial point of view, seaweeds represent an important natural resource as food, fertilisers, fodder, pigments, secondary metabolites and phycocolloids (i.e. agar, carrageenan and alginates) (Gellenbeck & Chapman 1983). These applications have created an industry which annually provides a wide variety of products with a total value of US$5.5–6 billion (McHugh 2001; Critchley et al. 2006). The major utilisation of these plants is as a source of food in Asia, particularly in countries like Japan, Korea and China, where seaweed cultivation has become a major industry (Critchley et al. 2006). In most Western countries, food and animal consumption is restricted due to various regulations (Renn 1997) and there has not been any major programme to promote and develop seaweed cultivation (Guiry 2008). China is the largest producer of

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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edible seaweeds, harvesting about 4.2 million wet tons of Laminaria japonica (‘kombu’) (Lüning & Pang 2003). The Republic of Korea grows about 800,000 wet tons of three different species, and about 50% of this is for Undaria (‘wakame’), grown in a similar fashion to Laminaria in China. Japanese production is around 600,000 wet tons and 75% of this is for Porphyra (‘nori’). Nori is the highest value product, at about US$16,000/ dry ton, compared to kombu at US$2,800/dry ton and wakame at US$6,900/dry ton (McHugh 2003). Industrial utilisation is largely confined to extraction for phycocolloids and to a lesser extent for active secondary metabolites extraction or as source of fertilisers. Seaweeds are a source of raw material for the extraction of phycocolloids, which are widely utilised in the food, pharmaceutical and cosmetic industry (Chapman & Chapman 1980). Gelidium and Gracilaria are the main sources of agar (Armisen & Galatas 2000), while Eucheuma spp. and Kappaphycus alvarezii are a source of carrageenan (Ask et al. 2003; Muñoz et al. 2004). Gelidium species are small, slow-growing plants, and while cultivation in ponds and tanks is possible, to date it has not been economically viable (McHugh 2003). Gracilaria chilensis is a major source for world agar production. Chile dominates the global production with an annual production of 70,000 wet tons with 90% being cultivated (Buschmann et al. 2006). The Philippines and China account for most of the production of Eucheuma denticulatum, E. cottonii and K. alvarezii, with 2,009,000 wet tons produced in 2008 through aquaculture (FAO 2010). Alginates are extracted from brown seaweeds of Ascophyllum, Durvillaea, Ecklonia, Laminaria, Lessonia, Macrocystis and Sargassum. However, most of the alginate extracted is obtained by harvesting these seaweeds from natural populations (Gacesa 1988). These species cannot be grown by vegetative means, but must go through a reproductive cycle involving an alternation of generations. For alginate production, this makes cultivated brown seaweeds too expensive when compared to the costs of harvesting and transporting wild seaweeds. The only exception is for Laminaria japonica, which is cultivated in China for food but sometimes surplus material is diverted to the alginate industry in China (refer to Tseng (1987) and FAO (1989) for further details on Laminaria cultivation). Active metabolites from several species of marine macro- and microalgae are reported to have antibacterial, antifungal, antialgal and/or antimacrofouling properties that are effective in the prevention of biofouling (Plouguerné et al. 2010). Different species have been identified as a potential source of marine metabolites, including Bifurcaria bifurcate (Maréchal et al. 2004), Dictyota pfaffii (Barbosa et al. 2007), Cystoseira baccata (Mokrini et al. 2008), Sargassum vulgare (Plouguerné et al. 2010), Fragilaria pinnata (Silkina et al. 2009), Ceramium rubrum, Mastocarpus stellatus and Laminaria digitata (Dubber & Harder 2008). Recently, extracts of Sargassum duplicatum have shown promising results as immunostimulants of the shrimp Litopenaneus vannamei (Yeh et al. 2010). However, the application of seaweeds as a source of biochemical products is still underdeveloped. Further investigations will include studies on metabolic pathways, physiology and large-scale production of targeted species as raw material for aquaculture. Species of Ascophyllum, Ecklonia and Fucus are commonly utilised as soil conditioners and fertiliser (Blunden 1991). They have a suitable content of nitrogen and potassium, but are much lower in phosphorus than traditional animal manures and the typical N:P:K ratios in chemical fertilisers (Metting et al. 1990). The large amounts of insoluble carbohydrates in brown seaweeds act as soil conditioners improving aeration and soil structure, especially in clay soils, and have good moisture retention properties. Furthermore, studies using extracts of the brown algae Ascophyllum nodosum as soil moisturiser have shown that they

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can improve growth rates and reduce pests, consequently increasing crop yields as well as improving the overall quality of the product (Blunden et al. 1996; Leach et al. 1999). In some terrestrial plants, the administration of A. nodosum extract has shown positive growth results and similar effects have been tested in seaweeds to improve the productivity of Kappaphycus alvarezii by controlling its epiphytes and diseases (Loureiro et al. 2010) and as a culture medium for its propagation (Hurtado et al. 2009). Seaweed extracts have given positive results in many applications, but further investigations are required to determine their full potential in this field, including studies on cost and return analysis. As a result of global warming research, attention has turned to seaweeds as source of fuel and as carbon sinks (Packer 2009). This field of algae utilisation is newer compared to other applications (i.e. food, phycocolloids) and the presence of large areas of cultivated seaweed has been seen as a tool to mitigate global warming (Muraoka 2004). Much of the existing research in energy and fuels from algae has been devoted to biodegradation, methane production and biodiesel production from microalgae (Scragg et al. 2002; Miao et al. 2004) and macroalgae (Ross et al. 2008) and showw contrasting results. However, according to Walker (2009) the evidence suggests that the net energy gain of liquid biofuels, derived either from algae or terrestrial crops, is either very modest or non-existent and will hardly be a solution to carbon dioxide emissions. There are two main areas where seaweeds have the potential for use in bioremediation. The first is the treatment of sewage and agricultural wastes to reduce the total nitrogen and phosphorus-containing compounds before releasing these waters into rivers or oceans. The second is for the removal of toxic metals from industrial wastewater. The use of seaweeds as biofilters has been seen as the most promising application of seaweeds. Recent investigations by various groups have shown that selected species of seaweeds possess good biosorption capacities for a range of heavy metal ions (Pan et al. 2000; Hashim & Chu 2004; Lodeiro et al. 2006a, 2006b; Rubín et al. 2006; Senthilkumar et al. 2006; Tsui et al. 2006). Species such as Enteromorpha prolifica (Michalak & Chojnacka 2010), Codium fragile (Kang et al. 2008), Ulva reticulata (Senthilkumar et al. 2006) and Sargassum sp. (PintoPadilha et al. 2005) have shown great potential in bioremediation but their application at large scale has not been tested. Similarly, Gracilaria cliftonii (Kumar et al. 2010) and G. chilensis (Hedt 2005; Berry 2007) have been identified as potential candidates for seaweed aquaculture as biofilters in an alternative to utilise inland saline water bodies in Australia. However, research on inland saline water aquaculture has focused mainly on other organisms (i.e. prawns, fish) (Prangnell & Fotedar 2005; Tantulo & Fotedar 2007; Partridge & Lymbery 2008). The use of seaweeds as biofilters in integrated multi-trophic aquaculture systems (IMTA) will be considered further in this chapter.

9.3

AQUACULTURE

The seaweed industry worldwide uses 7.5–8.0 million tons of seaweeds (fresh weight) (Critchley et al. 2006) annually, with a majority of it derived from aquaculture, as the demand for seaweed polysaccharides, like agar and carrageenan, and other seaweed-based products exceeds the supply of seaweed raw material from natural stocks. At present there are approximately 200 species of seaweeds used worldwide (Zemke-White & Ohno 1999), of which the following species or genera are intensively cultivated: Laminaria japonica, Undaria pinnatifida, Porphyra sp., Eucheuma sp., K. alvarezii, Gracilaria sp., Monostroma sp. and Enteromorpha sp. (Lüning & Pang 2003).

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Seaweed cultivation not only provides an alternative to seaweed resources which have been overexploited but also facilitates the selection of germplasm with desired traits (Reddy et al. 2008). Different alternatives, such as simple and cost-effective cultivation methods, use of selected material as seed stock, and good farm management, are some common practices to enhance the economic prospects of seaweed cultivation. Furthermore, in vitro cell culture techniques have also been employed as they facilitate development and propagation of genotypes of commercial importance. The selection of the methodology of cultivation is specific to targeted taxa. Although some seaweeds require one-step farming through vegetative propagation (such as Kappaphycus, Gracilaria), others need two-step or multi-step farming (such as Porphyra, Laminaria, Undaria) (Sahoo & Yarish 2005). Commercial seaweed cultivation has been carried out successfully for over 40 years in countries like Chile, the Philippines, Indonesia, Japan and China, and production has increased sharply (Doty & Alvarez 1975). However, production has been focused on species belonging to Gracilaria and Gelidium as source of agar and the carragenophytes Eucheuma, Kappaphycus and Chondrus for carrageenan extraction. Gracilaria species are the major agar source worldwide (McHugh 2003), and most of the research has focused on taxonomic studies, selection of species for agar yield and quality, cultural management, production systems, and plant nutrition (Santelices & Doty 1989; Friedlander et al. 1993; Dawes 1995; Glenn et al. 1998).

9.3.1

Traditional cultivation

Traditional cultivation methods and selection procedures for seedling material rely on vegetative propagation of the plant (Buschmann et al. 2001). Fast-growing or robust tissue sections are utilised as the ‘seed’ for the next growing season (Doty 1985). However, selection of vegetative fragments from the same farming population for years and the lack of natural populations as new source of seed stock, decreases the genetic diversity and superiority of germplasm, resulting in weak and excessive branching and low stress resistance, as well as reducing the production and quality of phycocolloid (Dawes et al. 1993). On the other hand, the increasing demand for phycocolloids worldwide requires the introduction of more high-quality and eurythermal cultivars. Genetic engineering techniques are therefore expected to improve seed stocks of seaweeds (Renn 1997). An alternative technique to vegetative propagation consists of propagation from released spores attached to ropes (Alveal et al. 1997; Mantri et al. 2009), which avoids the aging effect when replacing or replenishing old stands. An advantage of spore-seeding cultivation is that it requires only a small amount of reproductive material to produce a large quantity of seeded ropes. When the spore-seeding method was developed by Alveal et al. (1997) for Gracilaria chilensis, it showed a high production potential (6.5 kg m−1) after 15 months of cultivation. Nevertheless, the method was only tested under protected estuarine conditions. Recently, Halling et al. (2005) compared the biomass production of Gracilaria grown from spore-seeded ropes (spore propagation method) and found comparable results to that of suspended cultivation (vegetative propagation method). After one month of culture, the suspended ropes had a significantly higher productivity than the spore-seeded ropes. The biomass losses observed by Halling et al. (2005) were attributed to the culture method, which was not suitable for the exposed open coastline of Chile, where the study was undertaken.

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Under controlled conditions, the fertile life-stages of G. chilensis (cystocarpic and tetrasporic) of the spore-originated thalli have lower specific growth rates (SGR) compared to the sterile (vegetative) phase. This confirms reduction in thalli growth due to extra energy costs for producing gametes and spores (Kain & Destombe 1995). Furthermore, sporeoriginated thalli have a higher polymorphism compared to vegetative thalli, which could be the result of different genotypes. This could generate a wide variety of characteristics and as a result a higher capacity for adapting to changes in environmental conditions (Santelices et al. 1999). A combination of the two methods would be advantageous, as spore-seeded ropes could be used for the first outgrowth in protected bays (nursery areas), while net tubes could be used for continued production in open waters. Spore-inoculated ropes of G. chilensis may be useful for continuous supply of stocking material for such cultivation methods; however, its performance in open coastal water is limited due to biomass loss. The vegetative seeded ropes (Halling et al. 2005) had a significant advantage over algal bundles, used in previous studies. Similar observations were made by Mantri et al. (2009) with G. dura propagated through carpospores. Spore cultivation of Macrocystis pyrifera in Chile has been considered as an alternative to alleviate the actual harvesting pressure of the natural stocks due to the establishment of abalone farming in the area. Macchiavello et al. (2010) cultivated M. pyrifera for a year under laboratory and field conditions, with two different cultivation methodologies. They observed a maximum frond length of 175 cm and 22 kg m−1 of rope after 120 to 150 days of cultivation in the sea.

9.3.2

Commercial cultivation

9.3.2.1 Gracilaria sp. The methods employed at the commercial level for Gracilaria sp. cultivation are based on the vegetative propagation of the seaweed. Several methods have been developed, with two being the most important. The first one is the direct method, which consists of a direct burial of the thalli into the sandy bottom using different types of tools, and the second method (plastic tube method) consists of fastening bundles of thalli to plastic tubes, filled with sand, which anchor the algae to the sea bottom (Alveal 1986). Most of the studies of suspended Gracilaria cultivation consist of bundles of vegetative thalli tied to cultivation ropes (Pizarro & Barrales 1986; Westermeier et al. 1993). This practice is labour- and time-consuming, and inefficient, as large parts of the rope are left unproductive. Abreu et al. (2009) have recently compared the two traditional cultivation methods of G. chilensis. They demonstrated that Gracilaria growth performance was higher on the suspended cultures near fish nutrient effluents, with daily mean growth rates reaching 4% (± 0.29) and with a mean biomass production of over 1,600 g m2 month−1 (±290), which was double the growth rates observed for Gracilaria grown in area without fish effluents or on unimpacted sites. Similarly, the productivity of bottom-cultured Gracilaria was highly reduced by biomass losses. This reduction in productivity has been attributed to nutrient removal and a decrease in photosynthetic performance of cultivated Gracilaria. The longline cultivation unit proved to be the most efficient technology for nutrient removal with monthly removal of up to 9.3 g (± 1.6) nitrogen (N) per metre of longline. Although they provided a good comparison of the most important cultivation methods available, results may vary from one area to another and among species.

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9.3.2.2 Kappahycus and Eucheuma The methods utilised for the cultivation of Kappahycus/Eucheuma are similar to those used for Gracilaria cultivation. Although many systems have been developed from field and laboratory experiments, only one type, the tie–tie system (Doty & Alvarez 1975) manifested in three basic forms: fixed off-bottom, floating long line and rafts, has survived because it was simple and because farm materials are readily available and are inexpensive (Ask & Azanza, 2002). 9.3.2.3 Porphyra sp. Porphyra sp. is the most important seaweed produced for edible purposes, with nearly 1 million wet tons produced through aquaculture (McHugh 2003). Cultivation of Porphyra has evolved significantly since its introduction in Japan in the 1960s. The cultivation methods of Porphyra are similar in all countries, with minor modifications (Sahoo & Yarish 2005; Sahoo et al. 2007). Due to its complex life cycle, the farming system for Porphyra can be divided into 5 distinct phases: (1) culture of conchocelis, (2) collection of conchospores and seeding of culture nets, (3) nursery of sporelings, (4) harvesting and (5) processing (Chen & Xu 2010). 9.3.2.4

Laminaria

Laminaria is cultivated mainly in China, Korea and Japan. In 2008 China produced 4,765,076 wet tons through aquaculture (FAO 2010). In a similar way to the cultivation technology for Undaria, Laminaria is cultivated through a technique called ‘forced cultivation’ (Tseng 1987; Tseng & Borowitzka 2003). The cultivation of Laminaria consists of four phases: (1) collection and settlement of zoospores, (2) production of seedlings, (3) transplantation and outgrowing and (4) harvesting (Sahoo & Yarish 2005). During the past decade, research on Laminaria cultivation has focused on aspects related to its commercial cultivation such as sporogenesis (Jun Pang & Lüning 2004), effect of environmental factors (Mizuta et al. 2003; Shi et al. 2005; Roleda et al. 2006), hybridisation and genetic improvement (Wang et al. 2005; Li et al. 2008b).

9.4

INTEGRATED AQUACULTURE

Research on seaweed biofilters for treating effluents from mariculture practices started in the mid-1970s (Ryther et al. 1975; Langton et al. 1977). Since the start of the 21st century research in this field has attracted considerable interest and is now one of the most novel applications of seaweed (Chopin et al. 2001a; Troell et al. 2003; Neori et al. 2004). The integration of seaweeds with marine fish culture has been examined and studied in Canada, Japan, Chile, New Zealand, Scotland and the USA (Petrell et al. 1993; Petrell & Alie 1996; Troell et al. 1997; Chopin et al. 2001a; Chopin et al. 2001b; Chopin et al. 2003; Troell et al. 2003; Neori et al. 2003, 2004; Halling et al. 2005; Lu & Li 2006; Crab et al. 2007; Stenton-Dozey 2007; Sanderson et al. 2008; Abreu et al. 2009). Species like Ulva sp. and Enteromorpha sp. have been identified as ideal candidates for filtering fish effluents due to their capacity to rapidly absorb and metabolise nitrogen, removing up to 80% of dissolved ammonium in aquaculture wastewaters (Neori et al. 2000), their high growth

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rates, low epiphytism susceptibility, and their worldwide distribution (Jimenez del Rio et al. 1996; Neori et al. 2000; Msuya & Neori 2002). The technology for integrated multi-trophic aquaculture (IMTA) strategies exists, but has not been implemented at commercial scales (Buschmann et al. 2009). Furthermore, seaweed biofiltration of fish farm effluents has not been adopted by the aquaculture industry, and except for Asia, Ulva is not grown commercially because it has a low market value (Mata et al. 2010) and because integrated aquaculture systems require considerable capital, large-scale facilities, and an enormous input of energy (Tsutsui et al. 2009). The selection of seaweed species for their commercial use as biofilters depends principally on two aspects. The interest of investors for a new species to be incorporated depends firstly on the commercial value and, secondly, on the physiological capabilities for growth in culture conditions together with its capacity to remove dissolved nutrients (Buschmann 2001; Chopin et al. 2001b; Buschmann et al. 2008). Besides Ulva and Enteromorpha, a range of algal species has also been targeted, often based on their ability to value-add an additional product (Hernández et al. 2002; Schuenhoff et al. 2006; Yang et al. 2006; Zhou et al. 2006). Gracilaria species can be efficiently cultivated for the production of useful algal biomass and removal of nutrients from shrimp pond effluents (Marinho-Soriano et al. 2009a,b). According to Marinho-Soriano et al. (2009a) relative growth rates (RGR) for G. birdiae were between 3.6 and 1.6 d−1, with a mean of 2.6% d−1. The biofiltration capacity of G. birdiae was confirmed by the significantly reduced concentration of the three nutrients analysed (phosphates (PO−4), ammonium (NH+4) and nitrates (NO-3)) over the study period. The concentration of PO−4 decreased by 93.5%, NH+4 by 34% and NO−3 by 100% after the 4-week experimental period. Similarly, the potential of the red alga Kappaphycus alvarezii to remove nutrients was tested to treat effluents of the fish Trachinotus carolinus (Hayashi et al. 2008), showing positive results. In Australia, de Paula et al. (2008) investigated the biofiltering capacities of Cladophora coelothrix and Chaetomorpha indica, often referred as ‘green tide’ algae because of their excessive nuisance growth under eutrophic conditions (Taylor et al. 2001; Raven & Taylor 2003). They have fast growth and efficient nutrient uptake, as well as a tolerance of low dissolved oxygen, low pH and high levels of nutrients in the water (Morand & Merceron 2005), all of which are essential for pond-based integrated aquaculture. However, the implementation of such species into integrated aquaculture systems is dependent on their success (growth) under in situ environmental parameters (Troell et al. 2003). In Chile, the red seaweed Gracilaria chilensis (Gracilariales, Rhodophyta; Bird et al. 1986), has been shown to be suitable for integration with salmon in both land-based tanks and in suspended cages (Troell et al. 1997; Buschmann 2001). Gracilaria growth in such systems was shown to be more than doubled compared to monoculture, due to fertilisation effects (Buschmann et al. 2001). However, suspended culture methods that are effective and easy to manage need to be developed in order to make integrated aquaculture commercially viable. In Chile such methods are important as recirculation and reduction of wastes may become a prerequisite to the further expansion of the salmon industry (Chopin et al., 2001a). The diversity of species trialled in bioremediation systems to date is very limited and the majority of these studies have focused on temperate aquaculture systems (Troell et al. 2009). The inability of traditionally high value genera such as Gracilaria, Porphyra and more recently targeted high yield species such as Asparagopsis (Dawes et al. 1999; Matos et al. 2006) to survive the broad environmental fluctuations of tropical pond-based systems provides an impetus for the selection of alternative bioremediation species. Table 9.1

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259

Table 9.1 Seaweeds as biofilters in integrated aquaculture systems. Seaweed

Integrated species

Reference

Ulva rigida Ulva lactuca U. rotundata Enteromorpha intestinalis Asparagopsis armata Kappaphycus alvarezii

Fish Fishpond Dicentrarchus labrax Dicentrarchus labrax

Mata et al. 2010 Neori et al. 2003 Hernández et al. 2002 Hernández et al. 2002 Mata et al. 2010 Qian et al. 1996 Hayashi et al. 2008 Shpigel & Neori 1996

G. gracilis G. fisheri

Fish Pinctada martensii Trachinotus carolinus Sparus aurata Tapes philippinarum Haliotis tuberculata Litopenaeus vannamei L. vannamei Oncorhynchus mykiss O. kisutch Saccostrea commercialis Penaeus japonicus Dicentrarchus labrax Shrimp

G. lemaneiformis Gracilariopsis bailinae Sargassum sp.

Sebastodes fuscescens Chanos chanos Penaeus latisulcatus

Gracilaria sp. Gracilaria sp. G. birdiae G. caudata G. chilensis G. edulis

Neori et al. 1998 Marinho-Soriano et al. 2009a Marinho-Soriano et al. 2009b Troell et al. 1997 Jones et al. 2001 Jones et al. 2002 Hernández et al. 2002 Chirapart & Lewmanomont 2004 Zhou et al. 2006 Guanzon et al. 2004 Mai et al. 2008

provides examples of different seaweeds utilised as biofilters in Integrated Aquaculture Systems.

9.5

POST-HARVEST: AGAR EXTRACTION

The post-harvest methodology for Gracilaria sp. consist of different phases, including, cleaning, drying, packing, phycocolloid extraction and marketing (Sahoo & Yarish 2005). The general methodology established for agar extraction consists of leaching the dry Gracilaria in boiling water, filtering off the extract and separating the agar by freezing and thawing to eliminate the water (Armisen & Galatas 2000). Although the general steps in the agar extraction process from Gracilaria species are known (Marinho-Soriano 2001; Freile-Pelegrin & Murano 2005; Andriamanantoanina et al. 2007; Pereira-Pacheco et al. 2007; Arvizu-Higuera et al. 2008; Orduña-Rojas et al. 2008; Li et al. 2009; Villanueva et al. 2010), the extraction variables and methodologies differ. Therefore, it is necessary to standardise the extraction process to optimise the agar yield and quality. In general, agar extraction is done in a waterbath or autoclave and there is no account mentioned of any preference of one over another, except for Buriyo and Kivaisi (2003) who reported that agar yield is higher using an autoclave than a waterbath. Generally, Gracilaria species produce agars with low quality due to their high sulphate concentrations. However, the gel properties of many Gracilaria agars can be improved by alkali treatment, which converts L-galactose-6-sulphate to 3,6-anhydro-L-galactose (Duckworth et al. 1971; Freile-Pelegrin & Robledo 1997; Freile-Pelegrin & Murano 2005), which is responsible for the enhancement of the gel-forming ability.

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Recently, Kumar and Fotedar (2009) evaluated the agar extraction process of G. cliftonii to maximise the agar yield. The investigation determined the effects of different variables, i.e. soaking, time, soaking temperature, seaweed–water ratio, and time and temperature of extraction, on chemical and physical properties of agar from G. cliftonii (gel strength, gelling temperature, melting point and sulphate content). In addition, the effect of alkali treatments on the agar characteristics of G. cliftonii at different soaking temperatures was also investigated. Kumar and Fotedar (2009) observed that the agar yield from G. cliftonii was maximised when extraction process was carried out with 1 hour soaking time at 30°C with seaweed to water ratio of 1:150 and extracted for 3 hours at 100°C. The alkali– temperature combinations significantly influenced agar yield and properties, and irrespective of temperature, alkali treatments of G. cliftonii at 3% and 5% significantly increased the gel strength of its agar. New methodologies of extractions have been recently reported in the literature: agar extracted under cold conditions as demonstrated by Maciel et al. (2008) and the microwaveassisted extraction (MAE) method (Sousa et al. 2010). The first method involves storing the seaweed at -20°C, dissolving in distilled water and stirring for 15 hours at room temperature (25–28°C). The water-soluble fraction is separated from the insoluble fraction by filtration and centrifugation. The second method of agar extraction (MAE) was tested for Gracilaria vermiculophylla grown under integrated aquaculture conditions. Sousa et al. (2010) also evaluated the influence of the MAE operational parameters (extraction time, temperature, solvent volume and stirring speed) on the physical and chemical properties of agar (yield, gel strength, gelling and melting temperatures, as well as, sulphate and 3,6-anhydro-L-galactose contents). They observed that the quality of the extracted agar, from G. vermiculophylla, was similar to the traditional extraction method (2 hours at 85°C) and drastically reduced the extraction time, solvent consumption and waste disposal requirements. Agar yield of G. vermiculophylla was 14.4 ± 0.4%, with gel strength of 1331 ± 51 g/cm2, 40.7 ± 0.2°C gelling temperature, 93.1 ± 0.5°C melting temperature, 1.73 ± 0.13% sulfate content and 39.4 ± 0.3% 3,6-anhydro-L-galactose content. Furthermore, this study suggested the feasibility of cultivation of G. vermiculophylla in IMTA systems for agar production (Sousa et al. 2010). In current industrial practice of agar extraction, sodium hypochlorite and other chemicals are being used as bleaching agents during the extraction process (Li et al. 2009). Several drawbacks exist in the traditional chemical-bleaching agar extraction process. Harsh chemical reagents increase the extraction cost. The difficulty in maintaining a constant pH during the chemical bleaching procedure leads to unstable agar quality. The chloride gas produced during the process suffuses the manufacturing plant and surroundings, and in addition to the waste liquid produced after chemical-bleaching, threatens the workers’ health (Warburton 2005). Furthermore, the after-effect of chloride gas and its effluents pose a major threat to the environment. Li et al. (2008a) evaluated different extraction process (native, alkali-modified, chemicalbleached and photobleached) on the physical and chemical properties of agar products of G. asiatica and G. lemaneiformis. Among the different processed agars, gel strength of the agar extracted with the photobleached methodology, was 1913 g/cm2, the highest among the different processed agars. They suggested that the agar photobleaching extraction process is a feasible method for Gracilaria species and has a potential application in the industry. Furthermore, Li and Zhang (2008) reported the optimisation of key process parameters (i.e. alkali modification concentration, photobleaching duration, algal length and screen filter opening size) in order to scale up this new methodology of agar extraction.

Seaweed Culture with Special Reference to Latin America

9.6

261

CULTIVATION IN LATIN AMERICA

Latin America’s coastline is characterised by its extensiveness and diversity of marine resources. The ocean masses that surround Latin America and the Caribbean countries contribute significantly to the region’s diversity and productivity. Considering features such as currents, the extent of the shelf, dynamics of the shore, banks, reefs and estuaries, Lemay (1998) divided the region into four marine and coastal regions, each with its distinct features: South Western Atlantic: This region includes Argentina, Brazil and Uruguay with a transition zone extending along Suriname and Guyana into the Caribbean. In the south, the region is characterised by an extensive continental shelf off the coast of Argentina and Uruguay, a cold current flowing from the south and strong tides along the coast of Patagonia. Wider Caribbean: This region encompasses the areas of the Caribbean Sea, the Gulf of Mexico and Pacific coast of Central America. These are typical tropical waters with temperatures averaging 27°C and seasonal fluctuations not exceeding 3°C, ideal conditions for the formation of coral reefs, mangroves and seagrass beds where productivity is highest. South Eastern Pacific: This region stretches along the west coasts of Central and South America from the Mexico–Guatemala border to southern Chile, with the Galapagos and Cocos Islands and Juan Fernandez Islands extending well into the Pacific. An important characteristic of this region is the cold, nutrient-rich current, which support the highly productive fisheries of Chile and Peru. Central Pacific: This region encompasses the Pacific coast of Mexico from its border with Guatemala north to Baja California, Mexico. This is a warm temperate marine region fed by an extension of the Equatorial Counter Current, which flows northward along the coast of Central America. This is a region recognised for its high marine biodiversity associated with lagoons, coral reefs, mangroves and a network of estuaries, which concentrate productivity along the coastal zone.

9.6.1

Potential species for commercial exploitation

The coastal waters of Latin America have a diverse seaweed flora (Rhodophyceae, Phaeophyceae and Clorophyceae) that include many economically important genera and are utilised as food, fodder or as raw material for phycocolloid extraction. However, most of the efforts in commercial cultivation have focused on Gracilaria species for agar extraction and Eucheuma and the introduced Kappaphycus alvarezii for carrageenan extraction.

9.6.2

Production figures

Although there is no recent information available on current seaweed production by Latin American countries, McHugh (2001) and Zertuche-Gonzalez (1993b) provide valuable information on the economics of seaweeds in Latin America. The seaweed cultivation and processing industry in Latin America plays a major role worldwide, as 17% of the seaweeds processed by the industry come from this region. From

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the red seaweed production point of view Latin America contributes 37% of the biomass, Chile being the major contributor with approximately 13% of the total (McHugh 1991). Latin America contributes significantly to the agar industry. Four countries contribute 39% of the agarophytes utilised worldwide for the agar industry. Argentina, Brazil and Chile produce 10%, 4% and 50%, respectively, of Gracilaria production, while Chile and Mexico contribute 2% and 7%, respectively, of Gelidium production. Other countries, like Peru and Santa Lucia, contribute to a minor degree (Zertuche-Gonzalez 1993b). According to the FAO’s Fisheries and Aquaculture yearbook (2009), the production of aquatic plants by Chile was 23,298 tons with a value of US$37,278, accounting for most of the production from Latin America. Besides the phycocolloid industry, there are two major important markets emerging in the region. One is the utilisation of Gracilaria as food for abalone (Haliotis sp.) cultivation (Hahn 1989) and second for direct human consumption, which, except in Santa Lucia, is an application underdeveloped in the region (Smith et al. 1984). Although several attempts at carragenophyte cultivation have been made in the region, only Panama (Batista-Vega et al. 2003) and Brazil (Hayashi et al. 2007) have developed the commercial cultivation of Eucheuma/Kappaphycus at different scales.

9.6.3

Cultivation methods

The techniques utilised for seaweed cultivation in Latin America are based on the methods utilised for the cultivation of Gracilaria and Eucheuma/Kappaphycus species in Asian countries (Ask & Azanza 2002). The methods developed for Gracilaria cultivation include mariculture, pond culture and tanks (Santelices & Doty 1989), with mariculture and pond culture considered as the most profitable. The pond culture widely used in Asian countries such as Taiwan, China and Thailand has not been practised in Latin America, but it is considered to be the most viable method for Gracilaria cultivation for highly exposed areas (Zertuche-González et al. 1987a). The methods employed at the commercial level for Gracilaria cultivation are based on the vegetative propagation of the seaweed. The methods commonly used in pilot cultivation studies are based on those developed and established in Chile (Zertuche-González & Garcia-Esquivel 1989; Westermeier et al. 1991). In general they involve intertidal and subtidal vegetative cultivation of the species. The methods used for the cultivation of Kappahycus/Eucheuma are similar to those used in Asian countries, e.g. the Philippines and Indonesia, with some modifications. Although many systems were developed from field and laboratory experiments, only one type, the tie–tie system (Doty & Alvarez 1975) manifested in three basic forms: fixed off-bottom, floating longline and rafts, has survived. The system predominated primarily because it was simple and because farm materials are readily available and inexpensive (Ask & Azanza, 2002). At the experimental level Kappaphycus alvarezii has been cultivated in Brazil utilising rafts (Hayashi et al. 2007) and in Panama (Batista-Vega et al. 2004) and Mexico (Muñoz et al. 2004) with the longline method.

9.6.4

Overview of countries

Although the harvesting of seaweeds has been very significant in Chile, exploitation of seaweeds in Latin America is still underdeveloped. Most of the information available is

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263

from eight Latin American countries: Argentina, Brazil, Colombia, Cuba, Chile, Ecuador, Mexico and Peru, and from Caribbean countries including St Lucia, Jamaica, Barbados, Trinidad and Tobago, San Vicente and Venezuela). In most of these countries the cultivation of seaweeds has remained merely at the level of experimental and pilot cultivation. Some details on seaweed production by some Latin American countries are presented below. 9.6.4.1 Argentina In Argentina the most important activity is the harvesting of seaweeds for the phycocolloid industry, and no major attempts at seaweed commercial cultivation have been carried out. The important species harvested are Gracilaria, Macrocystis, Gigartina, Lessonia, Porphyra, Ulva and Codium, with Gracilaria (3,060 tons) and Macrocystis (328 tons) being the most important in terms of production for 1994 and 1996 (Boraso-de-Zaixso et al. 2006). Gracilaria gracilis is used for the production of agar, seaweed flour and pellets. Gigartina skottsbergii is used for the production of carrageenan, seaweed flour and pellets. This seaweed is mixed with Eucheuma cottonii (Kappaphycus alvarezii) imported from the Philippines for the production of carrageenan for the internal market. Sarcothalia sp., mainly S. crispata, is sometimes collected and processed with Gigartina skottsbergii, which is a potential raw material for the production of carrageenans and is exported as flour and pellets. Macrocystis pyrifera, Porphyra sp., Ulva sp. and Codium sp. are harvested for the production of flour and pellets. Carrageenan and carrageenophyte seaweed flour are produced for the domestic market and are sometimes exported to South American countries such as Uruguay and Bolivia. Seaweed flour from Macrocystis pyrifera is now exported to the USA (Boraso-de-Zaixso et al. 2006). The experimental cultivation of Gracilaria verrucosa with the bottom-planting method has been carried out by Boraso-de-Zaixso (1987) in Golfo Nuevo using different methods. The best results were obtained with ropes, with five inocula of 200 g/m of rope. Harvests were carried out by hand and cutting with scissors. Accumulated yield of 120% above the inocula weight was obtained after 5 months. Similarly, Porphyra columbina harvested from natural populations for edible purposes has been cultivated under laboratory conditions (Piriz 1989). 9.6.4.2 Brazil In Brazil considerable research has been carried out on seaweed cultivation of local and introduced species, e.g. Kappaphycus alvarezii. Commercial cultivation of K. alvarezii has begun (Hayashi pers.comm), and there are attempts to cultivate this species in integrated systems with the fish Trachinotus carolinus (Hayashi et al. 2008). Regarding the commercial exploitation of seaweeds, only a mixture of Gracilaria cornea, G. caudata and Gracilariopsis tenuifrons, for agar, and Hypnea musciformis, for kappa-carrageenan, are being exploited in significant amounts in Brazil. However, there has been a trend to decrease this exploitation in recent years. One of the companies that produced agar recently closed, thereby reducing the total amount of algal polysaccharide production to about 80 tons per year. Additionally, there has been artisanal utilisation of Porphyra spiralis and P. acanthophora for the local production of nori sheets. Ulva sp. has been collected for use in the cosmetic industry and there has been sporadic harvesting of Sargassum sp. for medicinal

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uses. Gracilaria domingensis has been collected and exported to Japan for food (Oliveira 2006). All of these activities have been on a modest economic scale. Experiments have been made with different species of Gracilaria (Câmara Neto 1987), Hypnea musciformis (Berchez & Oliveira 1990) and Pterocladiella capillacea (Oliveira & Berchez 1993). In all these instances the basic technique utilised was the rope system. Experiments to grow several species in tanks were made by Oliveira (1990) but there is no economic perspective for this approach. Experimental cultivation on Monostroma sp. and Agardhiella subulata have also been performed (Oliveira 2006). 9.6.4.3 Caribbean The development of mariculture in the Caribbean has been slow in general. There have been several initiatives for the commercial production of seaweeds, both for export of raw material for industrial extraction and for the local market used for the production of traditional drinks and desserts (e.g. seamoss) (Smith & Rincones 2006). Experimental cultivation trials on Gracilaria sp., Eucheuma sp. and Gracilariopsis sp. have been performed with different outcomes. The most successful methods are based on vegetative propagation of Gracilaria sp. on ropes attached to bamboo rafts or tied between fixed stakes, following the methods used in the Philippines for Eucheuma and Kappaphycus (Smith & Renard 2002). The most successful cultivation in the Caribbean is the case of Kappaphycus alvarezii in Panama. 9.6.4.4 Panama Although the marine algae of Panama are not well studied, they are used by some ethnic groups as fertilisers, as food and for medical purposes. Phycocolloid extracts of algae such as agar, carrageenan and alginate are imported by some industries in Panama for the phycocolloid industry (Batista-Vega & Yee 2006). The company Gracilarias de Panama S.A. has been involved with the development of seaweed cultivation in Panama. Kappaphycus alvarezii has been introduced and is cultivated at a small scale. The production has been exported as raw material to Europe and processed locally (Batista Vega, personal communication). Also, Gracilaria dominguensis has been cultivated using two cultivation techniques: stake and rock planting (Batista-Vega 2004). 9.6.4.5 Colombia There have been a number of recent studies by university investigators related to the ecology and biology of potential commercial algae in Colombia. These studies have provided basic information of algal distribution, seaweed biomass, chemistry of phycocolloids and potential uses of species for algal derived products (McHugh 2001). Gracilaria cornea, Eucheuma isiforme and Kappaphycus alvarezii have been tested for experimental cultivation (Peña-Salamanca & Álvarez-León 2006). 9.6.4.6 Cuba Research on the identification of natural resources for commercial exploitation in Cuba has been carried out by different researchers. Briothamnion triquetrum, Digenea simplex,

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265

Gelidiella acerosa, Hypena acerosa and Gracilaria sp. are considered the most promising for commercial exploitation (Diaz-Piferrer 1961). Soloni (1952) investigated the chemical composition of B. triquetum, Gracilaria ferox, G. confervoides, G. lacinulata, Laurencia papillosa and Eucheuma isiforme. Cuba has successfully cultivated Eucheuma/Kappaphycus, but the country does not have the financial infrastructure to expand the cultivation to enable export, or to produce enough for its own carrageenan production. Therefore it has required the support of outside companies interested in joint ventures (McHugh 2001). 9.6.4.7 Mexico Mexico is rich in seaweed resources and many species have been identified as potential species for commercial exploitation (see Table 9.2 and Fig. 9.1). There is no commercial cultivation of seaweeds in Mexico and all seaweed industry has been confined to the Baja California Peninsula in the northern part of the country.

Table 9.2 Potential seaweeds for some Latin American countries. Country

Economic species

Reference

Argentina

Macrocystis pyrifera, Lessonia vadosa, Gracilaria gracilis, Gigartina skottsbergii, Sarcothalia spp., Porphyra columbina, Gymnogongrus sp., Ahnfeldtia plicata, Durvillaea antarctica Gracilaria, Laminaria, Caulerpa, Monostroma, Enteromorpha, Porphyra, Pterocladiella capillacea, Gelidium floridanum, Gracilaria cornea, G. caudate, Gracilariopsis tenuifrons, Hypnea musciformis, Agardhiella ramosissima, Meristiella gelidium, Chondrochantus teedii, C. acicularis, Laminaria abyssalis, L. brasiliensis, Sargassum spp. G. mammillaris, Gracilariopsis sp., Gelidiella acerosa, Bryothamnion triquetrum, Gelidium serrrulatum, Eucheuma isiforme, Hypnea musciformis, Hydropuntia cornea, H. crassissima, Gracilaria domingensis, Eucheuma isiforme Porphyra columbina, Gelidium chilense, Gelidium lingulatum, Gelidium rex, Callophyllis variegate, Ahnfeltia plicata, Mazzaella laminarioides, Sarcothalia crispate, Mastocarpus papillatus, Ahnfeltiopsis furcellata, Gigartina skottsbergii, Mazzaella membranacea, Chondracanthus chamissoi Gracilaria chilensis, Lessonia trabeculata, Lessonia nigrescens, Macrocystis pyrifera, M. integrifolia, Durvillaea Antarctica, Ulva spp. Macrocystis pyrifera, Gelidium robustum, Chondracanthus canaliculatas, Gracilariopsis lemaneiformis, Sargassum sp., Halymenia floresia, Echeuma isiforme, Gracilaria cornea Gracilaria crassissima, Gracilaria mamillaris Sargassum sp., S. natans, Eucheuma sp., turbinaria, Hypnea musciformis, Laurencia papillose, Acantophora specifera, Caulerpa sp. Porphyra columbina, Gigartina chamissoi, Macrocystis integrifolia, Lessonia nigrescens, Gelidium howei, Agardhiella tenera, Gymnogongrus furcellatus, Prionitis decipiens, Rhodoglossum denticulatum, Chondrus candiculatus, G. crinale var. luxurians, Pterocladia pyramidale, Gracilaria spp., Gracilariopsis spp., G. lemaneiformis, Hypnea valentiae

Boraso de Zaixso et al. 2006

Brazil

Caribbean

Chile

Mexico

Panama

Peru

Oliveira 2006

Smith & Renard 2002; Smith & Rincones 2006 Alveal 2006

Robledo 2006

Batista-Vega & Yee 2006 Acleto 2006

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Fig. 9.1

Harvesting of different coloured varieties of Kappaphycus alvarezii cultivated in Mexico.

Macrocystis pyrifera, Gelidium robustum, Gigartina canaliculata and Gracilariopsis sp. have been exploited since the 1950s and recently Gracilaria pacifica exploitation has started (Zertuche-Gonzalez 1993a; Robledo 2006). Gelidium robustum is harvested and utilised for the extraction of agar by a Mexican company, AGARMEX. Most of the production is exported to the United States, Japan and France (Zertuche-Gonzalez 1993a). The exploitation of Macrocytis has decreased with the closing of the main buyer KELCO®; however, there is a local market as feed for the abalone aquaculture and as soil stabilisers (Zertuche-Gonzalez pers. comm.). Also, there is small market for Eisenia arborea and Laminaria sp. for direct human consumption. Currently E. arborea is being cultivated for experimental purposes (Zertuche-Gonzalez pers. comm.). Laboratory and experimental cultivation trials of the native Eucheuma isiforme (FreilePelegrin & Robledo 2006) (Fig. 9.2) and Gracilaria cornea (Guzmán-Urióstegui & Robledo 1999) and the introduced Kappaphycus alvarezii (Muñoz et al. 2004) (Fig. 9.3 and Fig. 9.4) have been performed in the Yucatan Peninsula. In the California Peninsula, laboratory and field experiments have been undertaken on the native Eucheuma uncinatum (ZertucheGonzález et al. 1987b) and the introduced Chondrus crispus (Zertuche-González et al. 2001).

9.7

CONCLUSIONS

Although the harvesting of seaweeds has long been of considerable importance in Chile due to the existence of large stocks of native Gracilaria, the exploitation of seaweeds is still very recent in Latin America compared to Asian countries. Latin America is rich in

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Fig. 9.2 Eucheuma isiforme cultivated under experimental conditions at Dzilam de Bravo, Yucatan, Mexico. (Please see plate section for colour version of this figure.)

(a)

(b)

(c)

Fig. 9.3 Kappaphycus alvarezii color strains cultivated in Mexico. (a) green; (b) red; (c) brown. (Please see plate section for colour version of this figure.)

seaweed resources and the introduction of exotic species for seaweed cultivation is an important aspect that should be carefully analysed for each particular project. Seaweed exploitation and cultivation could be an important economic source for the region if market studies and assessment of commercial interests in investment are undertaken before funds are committed.

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Fig. 9.4

9.8

Cultivation of Kappaphycus under laboratory conditions.

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10

Marine Ornamental Fish Culture

Suresh Job

10.1 INTRODUCTION Ornamental aquarium species supply a multi-million dollar industry involving the farming or harvest, sale and use of live aquatic animals for display in aquaria. The aquarium hobby is one of the most popular hobbies globally, the third most popular in developed countries such as the United Kingdom (after keeping dogs and cats) according to the website www.ornamentalfish.org. The aquarium trade is a significant global industry. Global exports of ornamental fish (including both freshwater and marine species) in 2005 were valued at over US$237 million, and global imports were worth over US$282 million (UNEP-WCMC 2008). The majority (approximately 90%) of the trade in ornamental species comprises freshwater species, with marine ornamentals making up about 10% of the total (Sugiyama et al. 2004). Marine ornamental species involved in the aquarium trade are mostly associated with coral reefs from tropical coastal seas. An estimated 20–24 million marine ornamental fish and invertebrates are traded globally each year (Wabnitz et al. 2003). The global value of the marine aquarium industry is estimated to be worth between US$200 and US$330 million a year at the retail level (Wabnitz et al. 2003). Marine aquarium fish are a highvalue commodity, with an estimated average value of US$247 per kg (Job 2005). They are 100 to 200 times more valuable than average food fish (Olivier 2001; Job 2005). Aquarium animals are among the highest value-added products that can be harvested sustainably from coral reefs. Over 1,000 species of marine ornamentals are traded annually in order to supply this trans-global hobby (Wood 2001; Wabnitz et al. 2003). The tropical Indo-Pacific region, with its high marine biodiversity, is the major supply centre for marine ornamentals. Indonesia, the Philippines and Sri Lanka, for example, supply the vast majority of the world’s marine aquarium fishes (Wood 2001), accounting for about 75% of the total volume traded. Other important source areas include Australia, Brazil, Fiji, Hawaii, the Maldives, Puerto Rico, Sri Lanka, Thailand, Vietnam, the South Pacific island states, and the Red Sea and Persian Gulf states (Wood 2001). The US and European Union (EU) are the major markets for marine ornamentals. The US accounts for approximately 40–50% of the international market, with the EU accounting

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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for a slightly smaller percentage (Wood 2001; Wabnitz et al. 2003). Of the EU countries, Germany, the UK, France and the Netherlands account for over 65% of ornamental fish imports in terms of value. The other significant markets for marine ornamentals are those in Asia, particularly Japan and Hong Kong. Around half of the specimens traded are fish, with coral and non-coral invertebrates each representing around 25% of product traded (Wabnitz et al. 2003). Approximately 90–98% of marine ornamentals in the trade are wild-caught (Wabnitz et al. 2003; Sugiyama et al. 2004). This contrasts sharply with the freshwater aquarium fish trade where 2–10% of fishes traded are wild-caught (Sugiyama et al. 2004). The value of the ornamentals trade goes beyond its 0.5% share of the international fish trade. The sector plays a valuable role in providing employment opportunities and income to rural, coastal and insular communities in many places. The trade in ornamentals also supports the market for a multi-million dollar industry in aquarium tanks, filter systems and other accessories (O’Sullivan et al. 2008). Although at present the ornamental industry is dominated by freshwater species, the trade in marine aquarium species has been showing a growing trend since the end of the 20th century and the increasing popularity of marine aquaria may become the dominant trend during the 21st century (Olivier 2001).

10.1.1

Marine ornamental fish aquaculture

The marine ornamentals aquaculture industry has only truly begun to progress in a significant way over the past decade or so. The US leads the world in terms of the commercial aquaculture of marine ornamentals, but there is also significant commercial production in the EU, Asia and Australia. There are also countless small backyard operations that supply fish and invertebrates to local pet stores. Many of these smaller operators tend to be avid hobbyists, who have progressed to the point where they are keen to breed fishes and/or invertebrates. An increasing number of research scientists and fisheries/aquaculture managers are also starting to focus on marine ornamental species. The rapid rate at which coral reefs are declining globally has led to growing concern over the impact of the marine ornamentals trade on marine biodiversity (Wood 2001; Burke et al. 2002; Wabnitz et al. 2003). Commercial aquaculture production of marine ornamentals, particularly in tropical developing countries, could provide a complementary means of reducing overexploitation of ornamental species to industry regulation, while ensuring the sustainability of the trade and promoting socio-economic development (Tlusty 2002; Job 2005). Tank-bred marine aquarium fishes are already in high demand by hobbyists in the major markets. Tank-bred marine aquarium fish in the major markets, for example, generally command prices approximately 25% higher than equivalent wild-caught fish. Increased awareness of coral reef degradation issues amongst hobbyists will continue to ensure that tank-bred fish remain in high demand.

10.1.2

Key constraints

A small number of key families dominate the marine ornamentals trade in terms of volume. In general, about six or seven families of fish represent more than 60% of all fish commercially harvested. The main marine fish groups that dominate the aquarium market are: Pomacentridae (damselfishes and clownfishes), Acanthuridae (surgeonfishes), Balistidae

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(triggerfishes), Labridae (wrasses), Pomacanthidae (angelfishes), Chaetodontidae (butterflyfishes) and Syngnathidae (seahorses) (Olivier 2001). There are reports of around 100 species of marine ornamental fish having been bred in captivity, largely on a hobbyist or research scale (Table 10.1). Of these, approximately 30–35 species are currently in commercial production, albeit still on a relatively small scale. The most commonly aquacultured fish families are the anemonefishes and damselfishes (Pomacentridae), dottybacks (Pseudochromidae), cardinalfishes (Apogonidae), seahorses (Syngnathidae) and gobies (Gobiidae). The development of marine ornamentals aquaculture has been stymied to a large extent by the lack of published culturing protocols for most species in the trade. While a number of species are currently being produced by commercial ventures, most of the information

Table 10.1 Marine ornamentals species reported to have been bred in captivity Species

Common name

Clownfishes: Amphiprion akallopisos Amphiprion akindynos Amphiprion bicinctus Amphiprion clarkii Amphiprion ephippium Amphiprion frenatus Amphiprion melanopus Amphiprion nigripes Amphiprion ocellaris Amphiprion ocellaris Amphiprion percula Amphiprion percula Amphiprion perideraion Amphiprion polymnus Amphiprion rubrocinctus Amphiprion sandaracinos Amphiprion sebae Amphiprion tricinctus Premnas biaculeatus Premnas biaculeatus Hybrid: Amphiprion sebae x Amphiprion polymnus Hybrid: Premnas biaculeatus x Amphiprion ocellaris

Skunk clownfish Barrier Reef clownfish Two-band clownfish / Red Sea clownfish Clark ’s clownfish Flame clownfish Tomato clownfish Coral Sea clownfish / Red & Black clownfish Rose Skunk clownfish Black False Percula clownfish False Percula clownfish True Black Percula clownfish True Percula clownfish Pink Skunk clownfish Saddleback clownfish Australian tomato clownfish Orange Skunk clownfish Sebae clownfish Three-band clownfish Gold-Stripe maroon clownfish White-Stripe maroon clownfish White-tipped clownfish Cocoa clownfish

Dottybacks: Cypho purpurascens Ogibyina queenslandiae Pseudochromis aldabraensis Pseudochromis cyanotaenia Pseudochromis diadema Pseudochromis dilectus Pseudochromis flavivertex Pseudochromis fridmani Pseudochromis fuscus Pseudochromis olivaceus Pseudochromis polynemus

Flame dottyback Queensland dottyback Neon dottyback Bluelined dottyback Diadem dottyback Redhead dottyback/Sri Lankan dottyback Sunrise dottyback Orchid dottyback Golden dottyback Olive dottyback Long-finned dottyback (Continued)

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Table 10.1 (Continued ) Species

Common name

Clownfishes: Pseudochromis Pseudochromis Pseudochromis Pseudochromis Pseudochromis

porphyreus sankeyi splendens springeri steenei

Magenta dottyback Striped dottyback Splendid dottyback Springer ’s dottyback Flamehead dottyback

Grammas: Gramma loreto Cardinalfish: Apogon compressus Pterapogon kauderni Sphaeramia nematoptera

Blue-eye cardinalfish Banggai cardinalfish Pyjama cardinalfish

Longfins: Assessor flavissimus Assessor macneilli Calloplesiops altivelis

Yellow assessor Blue assessor Marine comet

Seahorses & pipefishes: Doryrhampus dactyliophorus Hippocampus barbouri Hippocampus erectus Hippocampus kuda Hippocampus procerus Hippocampus reidi Hippocampus zosterae

Banded pipefish Barbour ’s seahorse Lined seahorse Yellow kuda seahorse Emperor seahorse/ High-crown seahorse Brazilian seahorse Dwarf seahorse

Gobies: Amblygobius rainfordi Coryphopterus personatus Cryptocentrus cinctus Cryptocentrus leptocephalus Cryptocentrus lutheri Elacatinus evelynae Elacatinus figaro Elacatinus multifasciatus Elacatinus oceanops Elacatinus punticulatus Gobiodon citrinus Gobiodon okinawae Hybrid: Elacatinus horstii x Elacatinus lousiae Hybrid: Elacatinus oceanops x Elacatinus figaro

Rainfordi goby / Court Jester goby Masked goby Yellow/blue prawn goby Pink-speckled prawn goby Luther ’s prawn goby Sharknose goby Yellow-line goby Greenbanded goby Neon goby Red-head goby Citron goby Yellow clown goby Gold-stripe goby Hybrid cleaner goby

Blennies: Meiacanthus Meiacanthus Meiacanthus Meiacanthus Meiacanthus Meiacanthus Meiacanthus Meiacanthus

Forktail blenny Bundoon blenny Striped fangblenny Mozambique fangblenny Blackline fangblenny Canary blenny Disco blenny Green canary blenny

astrodorsalis bundoon grammistes mossambicus nigrolineatus oualanensis smithi tongaensis

Angelfishes: Pomacanthus maculosis Centropyge spp.

Royal gramma

Maculosis angelfish Centropyge angelfish

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generated is of a proprietary nature, and is not publicly available. One of the key issues in expanding marine ornamentals aquaculture is ensuring that culturing protocols are widely available and easily accessible. Much of the scientific research addressing the challenges of marine ornamentals aquaculture has only occurred over the past decade, with the bulk of it having been done since 2000. As a result, most aspects of marine ornamentals aquaculture are still poorly understood. In this chapter the developments in the culture of marine ornamental fish are reviewed.

10.2 10.2.1

BROODSTOCK AND EGGS Pomacentridae

Damselfishes and anemonefishes are amongst the best known of coral reef fishes. In the wild, the pomacentrids are diurnally active and shelter within the reef matrix at night. Damselfishes and anemonefishes are demersal spawners, i.e. their eggs are laid on a hard substrate (Thresher 1984). Pairing strategies vary in the Pomacentridae, ranging from stable pairs in the anemonefish to harem systems in many damselfish species where the males spawn with multiple females (Job et al. 1997). A number of pomacentrid species have been cultured in experimental and commercial systems (Fig. 10.1) for the ornamentals trade, with at least 12 species currently in commercial production. Current commercial production is primarily based on the anemonefish, with few damselfish species produced on a regular basis.

Fig. 10.1

Small-scale commercial marine ornamentals aquaculture system showing broodstock tanks.

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10.2.1.1 Spawning broodstock In general, broodstock of most cultured marine ornamentals species tend to spawn naturally if their environmental and nutritional requirements are met (e.g. Job et al. 1997; Avella et al. 2007; Olivotto et al. 2003, 2008a,b), and hormonal induction is unnecessary. Key factors to consider include water quality, temperature, photoperiod, tank size, tank dimensions, habitat, and suitable spawning substrates. Temperature, photoperiod and water quality are the most important environmental triggers (Holt & Riley 2001) to successful spawning of marine ornamentals in aquaculture systems, with the other factors being of greater or lesser importance depending on species. Table 10.2 lists the optimum ranges for some of the key water quality parameters for breeding tropical marine ornamental fishes. Some of the basic requirements for breeding anemonefish species have been described in Hoff (1996) and Wilkerson (1998). In contrast, published information on breeding damselfishes remains scarce (Danilowicz & Brown 1992; Job et al. 1997; Olivotto et al. 2003). Anemonefish are kept in pairs in broodstock tanks of approximately 70–200 L in volume (Job & Bellwood 1996, 2000; Job et al. 1997; Avella et al. 2007; Olivotto et al. 2008a,b). Depending on species, damselfish are either kept as pairs in broodstock tanks of 80–200 L (Olivotto et al. 2003) or as groups in broodstock tanks of over 200 L volume. The larger tanks generally tend to be used for damselfish species where the dominant males spawn with a number of females. These species tend to be maintained in groups, with the sex ratio at approximately 1 male: 4 females (Job et al. 1997). Clay flowerpots (Fig. 10.2), clay roofing tiles or rocks are placed in the broodstock tanks as spawning substrates. These artificial spawning substrates are designed to allow checking for eggs and easy removal for transport to the rearing tanks. Egg clutches are usually laid on this substrate, and guarded by the parent fish. Spawning tanks generally tend to be fairly bare aside from the spawning substrate in order to facilitate ease of tank cleaning. In anemonefish, both parents guard the eggs, although the male tends to do more of the egg cleaning and fanning. Anemonefish do not require anemones in order to spawn (Wilkerson 1998). In most damselfish species, the male guards and maintains the eggs, and keeps the female away from the spawning site. Broodstock are generally fed to satiation 2–3 times a day with a varied diet of chopped shrimp, fish, squid, flakes, mysids, krill etc. Under these conditions, new anemonefish

Table 10.2 Broodstock tank water quality parameters for tropical marine ornamentals species. Parameter

Broodstock

Temperature (°C) pH DO Salinity (ppt) Ammonia (mg/L) Nitrites (mg/L) Nitrates (mg/L) Alkalinity/Carbonate hardness (ppm) Redox potential (mv) Photoperiod

26–30 8.0–8.4 >85% saturation 30–36 <0.02 <0.02 <20 140–178 (8.0–10.0 dKH) 300–400 12L:12D to 16L:8D

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Fig. 10.2 Yellow-stripe Premnas biaculeatus broodstock with a terracotta flowerpot as a spawning substrate. (Please see plate section for colour version of this figure.)

broodstock begin spawning within 6–12 months of being placed in the broodstock tank. Damselfishes spawn readily in captivity, and can begin spawning within 3 months of being placed in the broodstock tank (Olivotto et al. 2003). On average, anemonefish spawn every 12–15 days (Job et al. 1997; Avella et al. 2007; Olivotto et al. 2008a, b). Egg duration at 28–30 °C varies from 6 days in Amphiprion clarkii to 7 days in A. ocellaris and Premnas biaculeatus and 9 days in A. melanopus (Job & Bellwood 1996, 2000; Avella et al. 2007; Olivotto et al. 2008a,b). Egg duration in most damselfish species averages approximately 3–4 days at 28–30 °C (Job & Bellwood, 2000). Egg clutches of most anemonefish species contain between 300 and 1,000 eggs per clutch, depending on species (Arvedlund et al. 2000; Callan 2007; Olivotto et al. 2008a). Premnas biaculeatus, for example, generally produce close to 1,000 eggs per clutch, while Amphiprion ocellaris generally produce less than 500 eggs per clutch depending on female parent size (S. Job, personal observation). The number of eggs in damselfish egg clutches range from approximately 300 eggs (Olivotto et al. 2003) to over 1,000 eggs (Danilowicz & Brown 1992), depending on species. 10.2.1.2 Egg hatching The spawning substrate with the attached eggs is removed from the broodstock tank on the day of hatching and relocated to the larval rearing tank, which has similar water quality parameters to the spawning tanks (Job et al. 1997; Olivotto et al. 2003, 2008a,b; Avella et al. 2007). Once in the rearing tank, the eggs need to be kept well oxygenated as low dissolved oxygen levels leads to high egg mortality. Eggs are kept adequately oxygenated

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by either using standard aquarium aeration (air bubbles driven through aquarium airstones) or by ensuring that water with high oxygen levels flows over the eggs on a constant basis (Danilowicz & Brown 1992; Job et al. 1997; Olivotto et al. 2003, 2008a,b; Avella et al. 2007). Eggs are able to withstand moderate aeration without being dislodged and the air stream is usually placed as close as possible to the eggs. Aeration is reduced just prior to hatching to minimise damage to hatching larvae. Hatching is triggered by decreasing light intensity (McAlary & McFarland 1993), and eggs hatch within 30–60 minutes of the lights being turned off (Danilowicz & Brown 1992; Hoff 1996; Job et al. 1997; Olivotto et al. 2003, 2008a,b; Avella et al. 2007). Partial hatches are occasionally observed, where about 10% of the eggs hatch a day earlier than the rest (S. Job, personal observation). Hatching rates are well over 90% for most anemonefish and damselfish egg clutches (Danilowicz & Brown 1992; Job et al. 1997; Olivotto et al. 2003, 2008a,b).

10.2.2

Apogonidae

Cardinalfishes (family Apogonidae) are ubiquitous on coral reefs (Randall et al. 1990). Most adult cardinalfish are planktivores, and readily adapt to aquarium life. In the wild, adult cardinalfish are nocturnal and seek shelter during the day (Helfman 1993). Cardinalfishes are mouth brooders, where the eggs are cared for until the larvae hatch. The larvae are pelagic in almost all species, and are released on hatching. The known exceptions are the two species from the genus Pterapogon (the Banggai cardinalfish, Pterapogon kauderni and the Sailfin cardinalfish, Pterapogon mirifica), where the eggs generally hatch into fully developed non-pelagic juveniles. A relatively small number of apogonid species have been cultured in experimental and commercial systems, and approximately three species are currently in commercial production for the ornamentals trade. 10.2.2.1 Spawning broodstock Most cardinalfish species are maintained in groups of up to 20 individuals in large tanks of over 200 L volume, due to the difficulty in correctly sexing them (Job et al, 1997). An adequate number of separate shelters need to be provided in the spawning tank to minimise any aggression between the fish. Shelters generally resemble caves or hollows that enable the fish to defend or hide in. Terracotta flowerpots tend to be commonly used. Pair fidelity and duration of pairing in apogonids varies with species (Kuwamura 1985; Okuda 1999). In the wild, pairing occurs during the spawning season in species that form social groups, but not in more solitary species (Kuwamura 1985). In aquaculture broodstock tanks, some species form pairs that last at least through the spawning season, while other cardinalfish species do not form distinct pairs in the broodstock tanks, and the males remain solitary while brooding (S. Job, personal observation). Spawning in this mainly nocturnal family apparently occurs primarily during daylight hours in many species (Kuwamura 1985; Helfman 1993). In the apogonids, it is the male that broods the eggs (Fig. 10.3) (Thresher 1984; Randall et al. 1990). Males with eggs in their mouths display slightly flared opercula when viewed from above and are easily distinguished. Confirmation is provided by a cessation of feeding. In species that form distinct pairs, the female fish stays close to the brooding male and defends him from other fish.

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Fig. 10.3 Male Banggai cardinalfish, Pterapogon kauderni, brooding eggs in its mouth. (Please see plate section for colour version of this figure.)

10.2.2.2 Egg hatching The best egg hatching results for cardinalfish species with pelagic larvae are obtained by removing the parent fish (the male) from the breeding tank with the eggs still in its mouth and placing it in a suspended net cage within the larval rearing tank. This generally ensures close to full hatching. The parent is then removed in the morning after the eggs have hatched. However, removing the parent from the breeding tank without causing it to spit out the eggs is difficult in some species (Job et al. 1997). An alternative method for hatching cardinalfish eggs involves using a modified funnel submerged within the larval rearing tank (Fig. 10.4) (Job et al. 1997). The egg mass is kept gently suspended in the funnel by introducing a gentle flow of water at the base of the funnel. The flow rate is set such that the egg mass is constantly being re-suspended within the funnel but not carried out of it. Due to the gap between the upper margin of the funnel and the water surface, the larvae, which swim to the surface immediately after hatching, are carried out of the funnel as they hatch. This method generally results in minimal loss of eggs or larvae. The roughly circular apogonid egg masses hatch anywhere between 0.5–3 hrs after the lights are turned off. In the case of the Banggai cardinalfish and Sailfin cardinalfish, which release juveniles, the male fish is transferred into a separate rearing tank prior to the juveniles being released. The male is then returned to the breeding tank after the juveniles are released. Alternatively, a screen is used to separate the broodstock tank into two sections. The juvenile fish are able to pass through the screen, which reduces the risk of predation by the adult fish. The juveniles can then be collected and transferred into the rearing tank.

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Valve

Water outflow Egg mass

Water inflow

Funnel Sieve

Hatching tank Fig. 10.4

Egg hatching system for cardinalfish species.

Egg duration in cardinalfish fish species with pelagic larvae ranges from approximately 4 days in species such as Apogon cyanosoma to 7 days in species such as A. compressus (Job et al. 1997; Job & Bellwood 2000). Egg duration in the Banggai cardinalfish is approximately 17–21 days. The size of the larvae at hatching also differs substantially between species. In general, the longer the egg duration, the larger the larvae (Job & Bellwood 2000).

10.2.3

Pseudochromidae

The bright colours and curious nature of the dottybacks (family Pseudochromidae) (Fig. 10.5) have made them popular with many tropical marine aquarium hobbyists despite their aggressive personalities. There are over 70 species of dottybacks and they are found throughout the Indo-Pacific region as a family. However, the distributions of individual species can be relatively restricted. Some of the Australian and Red Sea species, for example, are endemic to relatively small areas. Dottybacks are diurnal carnivores that spend much of their time sheltering in the reef matrix. In the wild, they feed on a range of small invertebrates and fish. Most species are relatively small in size, and reach an average of 5–10 cm in length. Most dottybacks are protogynous hermaphrodites, i.e. they are capable of changing sex from females into males.

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Fig. 10.5 The most commonly cultured dottyback species, the Orchid dottyback, Pseudochromis fridmani. (Please see plate section for colour version of this figure.)

There tends to be minimal sexual dichromatism in most dottyback species, and males and females generally differ primarily in size (sexual dimorphism). There are, however, some notable exceptions. Cypho purpurascens from the Great Barrier Reef, for example, shows distinct sexual dichromatism (Thresher 1984; Olivotto et al. 2006a). Approximately six species are currently in commercial production. The high level of aggression towards con-specifics in many species is one important factor that hampers commercial production. In general, Red Sea dottyback species appear to be less aggressive than Indo-Pacific species, and the vast majority of current commercial production is based on Red Sea species (Moe 1997; Olivotto et al. 2006a). 10.2.3.1 Spawning broodstock Dottybacks are generally maintained as pairs in tanks of approximately 100–200 L (Moe 1997; Olivotto et al. 2006a). Differentiating the sexes is difficult in most dottyback species, and pairing is done by placing two small or medium-sized fish that are either juveniles or females together in the broodstock tank. As they are protogynous hermaphrodites, the larger (dominant) fish generally becomes male while the other fish will become female. This method generally works fairly well if the size difference between the two fish is sufficient (Moe 1997; S. Job, personal observation). Obviously, this method is unlikely to work if two large individuals are placed together as they may both be males. Terracotta flowerpots, short lengths of PVC pipe (Fig. 10.6) and rock caves have all been used successfully in broodstock tanks as spawning substrates (Moe 1997; Olivotto et al. 2006a; S. Job, personal observation). These artificial spawning substrates are designed

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Fig. 10.6 Male Pseudochromis steenei using a PVC pipe as a spawning den. (Please see plate section for colour version of this figure.)

to provide a cave-like shelter for the fish to lay the eggs in and protect, while being easy to check for eggs. The water quality parameters required for the broodstock are similar to that required for other marine ornamentals species (Table 10.2). While spawning, the pair of dottybacks will lie side by side in the male’s den, and a spherical ball of eggs (approximately 2 cm in diameter) is laid and fertilised (Moe 1997; Olivotto et al. 2006a). Once spawning is complete, the male chases the female away and takes over complete responsibility for looking after the eggs. The male generally curls his body around the eggs, and seldom leaves them except for feeding. Even then, many males feed less frequently when guarding their eggs. The male also agitates the egg mass from time to time. This agitation is probably a method of oxygenating the egg mass, and of keeping detritus off them. Once the broodstock settle into a pattern, spawning generally occurs approximately every 2–3 weeks (S. Job, personal observation). 10.2.3.2 Egg hatching The eggs take approximately 4–6 days to hatch (at 27–29 °C) (Moe 1997; Olivotto et al. 2006a). The eggs are generally left with the parent fish until the day they are expected to hatch. The dottyback eggs are generally removed from the breeding tank on the day of hatching, and are incubated in the rearing tank until the eggs hatch. Eggs that are ready to hatch are readily recognisable by the highly reflective metallic-looking eyes. Alternatively, the eggs are allowed to hatch in the broodstock tank, and the phototactic larvae are collected in a beaker using a flashlight and transferred into the larval rearing tank (Olivotto et al. 2006a).

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Fig. 10.7

289

Pseudochromis fridmani eggs on day 4 after fertilisation. Eggs due to hatch that night.

A number of incubation methods have been used to successfully hatch the eggs in the larval rearing tanks on the day of hatch (Moe 1997). All effectively revolve around keeping the eggs suspended or moving and aerated. One simple way is to place the egg mass in a mesh cage in the larval rearing tank (Fig. 10.7). The eggs hatch within 45–60 minutes of the lights being turned off, and the newly hatched larvae swim out of the mesh cage. An airstone is placed just below the eggs to ensure that oxygen levels remain high within the egg mass. The aeration also helps to direct the larvae out of the mesh cage. Once the larvae have hatched, the mesh cage is removed. Approximately 500 eggs are laid in a clutch for Pseudochromis flavivertex, and hatching success can be as high as 99% (Olivotto et al. 2006a). The average egg diameter for dottybacks is approximately 1.2–1.5 mm (Thresher 1984; Olivotto et al. 2006a). The larvae measure approximately 3.5–3.8 mm in standard length at hatching (Olivotto et al. 2006a). Larvae hatch with limited remnant yolk that is completely exhausted within 24 hours, and need to be fed exogenously shortly after hatching (Olivotto et al. 2006a).

10.2.4

Syngnathidae

In the wild, seahorses (family Syngnathidae) are found worldwide in a diverse range of marine habitats, including seagrass beds, coral reefs, mangroves and estuaries (Lourie et al. 1999). Seahorses are planktivorous in the wild, and prefer live moving prey even when maintained in aquaria (Wilson & Vincent 1998). Seahorses have been the subject of extensive research over the past decade due to concerns over population declines in some parts of the world thought to be due partly to habitat degradation and overexploitation

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(Vincent 1996; Job et al. 2002). Seahorses are heavily exploited for the traditional medicines, curios and marine ornamentals trades. There has been strong interest in seahorse aquaculture over the past few years, and commercial production of seahorses occurs more or less regularly in a number of countries around the world, including the US, EU, China, Vietnam, Sri Lanka, Australia and New Zealand. At least six species are currently in production. However, despite this progress seahorses are still considered a difficult species to produce reliably on a commercial scale. 10.2.4.1 Spawning broodstock Seahorses are kept either as pairs or in groups in the spawning tanks (Lin et al. 2008; Olivotto et al. 2008c; Murugan et al. 2009). A male to female ratio of 1:1 generally provides the best results as competition for mates can occur where sex ratios are not balanced (Faleiro et al. 2008). Tank volumes used for spawning seahorse broodstock range from 150 L to 2000 L, depending on species (Lin et al. 2008; Olivotto et al. 2008c; Planas et al. 2008; Murugan et al. 2009). In general, larger seahorse species require relatively deep tanks of approximately 60 cm to 90 cm depth for successful egg transfer from the female to the male (Woods 2000a). Seahorse broodstock tanks are usually kept fairly bare with the provision of adequate holdfasts for the seahorses. Commonly used holdfasts include nylon rope, plastic plants, bleached branching coral pieces or other similar material. Seahorses for the marine ornamentals trade come from tropical, subtropical and temperate habitats. As such, broodstock tank temperatures vary depending on species. Tropical species are generally maintained at 27–30 °C (Olivotto et al. 2008c; Lin et al. 2008; Murugan et al. 2009), whereas temperate species are kept at approximately 17– 20 °C (e.g. Woods 2003a; Palma et al., 2008). The other water quality parameters are broadly similar between tropical and temperate species (Table 10.2). Light intensity for broodstock tanks varies from less than 1000 lux to 3000 lux, and natural photoperiods are often used. Seahorses have an extensive courtship ritual both in the wild (Lourie et al. 1999; Foster & Vincent 2004) and in spawning tanks (e.g. Woods 2000a; Faleiro et al. 2008; Planas et al. 2008). At the end of this ritual, the pair rises up in the water column, and the eggs are transferred from the female into the male’s pouch. The male then carries the eggs until the juveniles are released. The pregnant male is transferred into a separate holding / hatching tank either immediately after spawning or just before the young are due to hatch (Lin et al. 2008). Young seahorses are at a relatively advanced stage of development when they are released, and are generally considered to hatch as juveniles rather than larvae. The males are then transferred back into the broodstock tank after the young are released from the pouch. Under good conditions, seahorses spawn every 18–20 days, depending on species (Olivotto et al. 2008c). Gestation period varies between and within species, and is also influenced by temperature. In general, it ranges from 14–22 days (Job et al. 2002; Foster & Vincent 2004; Lin et al. 2008). The number of young per batch also varies substantially within and between species, and can range from less than 100 young per clutch in young broodstock to approximately 2,000 young per clutch in some species (Lin et al. 2008; Planas et al. 2008; Murugan et al. 2009). On average, between 300 and 500 young are released per clutch (Lin et al. 2008; Planas et al. 2008; Murugan et al. 2009).

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10.2.5

291

Gobies

Gobies (Gobiidae) are amongst the most varied and widely distributed of marine fish. They are found in most marine habitats, as well as, in brackish water and freshwater environments. The two main groups of marine gobies (Gobiidae) that have been bred in captivity are the coral gobies (Gobiodon spp.) and the cleaner gobies (Elacatinus spp.). Three or four species of gobies are currently in commercial production. 10.2.5.1 Spawning Gobies are demersal spawners, and the parents guard the eggs until the pelagic larvae hatch. Gobies are generally kept as pairs in the broodstock tanks (Olivotto et al. 2005; Wittenrich et al. 2007). Gobies will spawn on a range of artificial and natural substrates including PVC pipe, ceramic tiles, live rock, etc. In most cases, the eggs are laid on the ceiling of the spawning shelter, and are attached via adhesive threads (Olivotto et al. 2005; Wittenrich et al. 2007). The male fish generally guards the eggs, even where the female fish sleeps in the same shelter at night. Most species that are spawned in captivity (primarily Gobiosoma spp. and Elacatinus spp.) lay between 200 and 350 eggs per clutch (Olivotto et al. 2005; Wittenrich et al. 2007). However, Priolepis nocturna may lay up to 3,000 eggs in a single clutch (Wittenrich et al. 2007). The eggs are generally removed from the broodstock tank and transferred into the larval rearing tank prior to hatching (Olivotto et al. 2005; Wittenrich et al. 2007). The eggs hatch approximately 5–7 days post-fertilisation, again depending on species (Olivotto et al. 2005; Wittenrich et al. 2007). Pairs spawn regularly and reliably in captivity, and new clutches are laid every few weeks. 10.2.5.2 Egg hatching Goby eggs are generally hatched out using similar methods to that described above for the other demersal spawning species groups (Olivotto et al. 2005; Wittenrich et al. 2007). In gobies, the eggs are transferred to the larval rearing tank on the day of hatching. Aeration at 100 ml/min is placed next to the eggs (Olivotto et al. 2005; Wittenrich et al. 2007). The eggs generally hatch within 1.5 hours of the lights being turned off, and the aeration is turned down to approximately 50 ml/min once the eggs have hatched. Eggs that are aerated in the absence of the male are able to survive and hatch, while eggs that are not aerated suffer from high mortality rates (Wittenrich et al. 2007). Egg survival is high in the gobies, and is generally over 90% (Olivotto et al. 2005; Wittenrich et al. 2007). The size of newly hatched larvae varies depending on species, and can range from just under 2 mm total length (TL) to over 3 mm TL (Olivotto et al. 2005; Wittenrich et al. 2007). Larvae hatch with limited remnant yolk, which is quickly depleted within 24 hours. In most cases, larvae begin feeding from day 1 post-hatch.

10.2.6

Marine angelfish

Breeding marine angelfish species has long been the ‘holy grail’ of marine ornamentals aquaculture. While there are some anecdotal reports of successful marine angelfish breeding in the hobbyist literature, commercial-scale production remains elusive. However,

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small-scale irregular production of some Centropyge species has begun recently (Baensch 2002, 2003a,b). Information in the scientific literature on the captive breeding of marine angelfish through to metamorphosis remains scarce (Olivotto et al. 2006b; Callan 2007). 10.2.6.1 Spawning The spawning requirements of some marine angelfish species have been described particularly for Centropyge sp. (Thresher 1984; Baensch 2003a; Olivotto et al. 2006b; Callan 2007). Marine angelfish are pelagic spawners and require sufficiently large and deep tanks of at least 50 cm depth for successful spawning (Baensch 2003a). The lemonpeel angelfish, Centropyge flavissimus, has been spawned successfully in 300 L tanks (Olivotto et al. 2006b). Live rock is usually added to the tank to simulate the reef habitat. The flame angelfish, C. loriculus, has been spawned in 200 L tanks that were kept bare except for PVC fittings that served as shelters (Callan 2007). In aquaculture environments, broodstock are kept in pairs rather than harems (e.g. 1 male with 2 females), as the second female may get overly harassed within the confines of the broodstock tanks (Olivotto et al. 2006b). Broodstock are fed with a varied diet that usually includes shrimp, fish and flakes (Callan 2007). Broodstock spawn when the water temperature and photoperiod are set to mimic summer conditions of 27–30 °C and 13L:11D or 14L:10D (Olivotto et al. 2006b; Callan 2007). Cycling of water temperature and photoperiod to simulate winter, spring, summer and autumn may facilitate the onset of spawning. The other water quality requirements are similar to those of other marine ornamentals species (Table 10.2). 10.2.6.2 Egg hatching Broodstock spawn on an almost daily cycle, and produce approximately 200–300 eggs per spawn (Olivotto et al. 2006b; Callan 2007). The floating eggs are collected in an external egg collecting basket after spawning, and transferred into the larval rearing tank (Callan 2007). Eggs of pelagic spawners are kept suspended in the larval rearing tank using airstones. Egg and larval characteristics are very similar among Centropyge species. Fertilised eggs are buoyant, transparent, colourless and spherical, and contain a single oil globule (Hioki et al. 1990). Centropyge eggs vary from 0.65 to 0.75 mm in diameter, and generally hatch 14–16 hours post-fertilisation at 27 °C (Baensch 2003b; Olivotto et al. 2006b; Callan 2007). Newly hatched larvae have been reported to range in size from 1.3 mm for C. ferrugatus (Hioki et al. 1990) to 2.3 mm for C. flavissimus (Olivotto et al. 2006b). Larvae grow rapidly during the first day. Flame angelfish larvae, for example, grew from 1.12 mm in length at hatch to 2.22 mm by 32 hours post-hatch (Rhody 2006 in Callan 2007). This rapid growth during the first day may explain some of the reported size differences at hatch of Centropyge species. Newly hatched larvae are poorly developed, lacking functional eyes, jaws or alimentary tract. However, by 72 hours post-hatch the larvae have fully developed eyes and digestive tract and are able to begin exogenous feeding (Rhody 2006 in Callan 2007).

10.2.7

Other species

Other species that have been cultured, albeit irregularly, include the Royal gramma (e.g. Gramma loreto and Gramma brasiliensis), Assessors (e.g. Assessor flavissimus), and

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Blennies (e.g. Meiacanthus spp.). Information on culturing these species is scarce in the scientific literature, although some anecdotal information is available in the hobbyists’ literature.

10.3

BROODSTOCK CONDITIONING

Aside from the provision of good water quality and adequate habitat, broodstock diet and nutrition is perhaps the most critical factor in the conditioning of broodstock in captivity (Cerda et al. 1994; Hoff 1996; Wilkerson 1998; Bruce et al. 1999; Emata et al. 2000; Wittenrich et al. 2007). In seahorses, for example, reproductive efficiency (number of juveniles released) of broodstock is strongly influenced by diet (Lin et al. 2007; Murugan et al. 2009). The number of juveniles released was higher in Hippocampus trimaculatus broodstock fed with live amphipods (Eriopisa spp.) (533 juveniles per clutch average) compared to animals fed with live sergestid shrimp (Acetes spp.) (457 juvenile per clutch average) (Murugan et al. 2009). The difference in that case was thought to be due to the higher protein content of amphipods compared to sergestid shrimp. Similarly, live Acetes shrimp produced better results in Hippocampus kuda broodstock in terms of fecundity, reproductive efficiency and gonadosomatic index (GSI) when compared to either live or frozen mysids or a mixture of live Acetes and frozen mysids (Lin et al. 2007). Seahorses fed with live Acetes shrimp displayed the shortest testes and ovary developmental durations, highest GSI and highest fecundity and reproductive efficiency, as well as highest juvenile survival rates for the young. Broodstock diet also has a high impact on egg quality, and thus on larval quality (Kerrigan 1997; Carnevali et al. 1998; Emata et al. 2003; McCormick 2003; Mazorra et al. 2003; Furuita et al. 2004; Papanikos et al. 2004; Salze et al. 2005). In seahorses, for example, egg quality (in terms of highly unsaturated fatty acid (HUFA) content and ratios) declines in captivity when seahorses are fed solely with enriched Artemia (Planas et al. 2008). Broodstock condition factor and wet weight also decline when seahorses are fed solely with Artemia, but improve when seahorses are fed with frozen mysids or frozen shrimp (Palma et al. 2008). As with other marine finfish species, marine ornamentals broodstock diets need to be relatively high in lipids, particularly in terms of highly unsaturated fatty acids (HUFAs) (Fernandez-Palacios et al.1995; Bell & Sargent 2003; Furuita et al. 2000, 2002, 2003; Mazorra et al. 2003; Papanikos et al. 2004; Li et al. 2005; Salze et al. 2005). There is also some evidence that high levels of proteins and/or amino acids may be useful in conditioning species such as seahorses (Murugan et al. 2009). High levels of anti-oxidants and vitamins may also contribute to improving egg and larval quality (Tucker 1998; Emata et al. 2000). As with most other finfish species, larger and older broodstock tend to produce greater numbers of eggs (Thresher 1984; Berkeley et al. 2004). Given the wide range of marine ornamentals species cultured and their varied natural diets, optimising broodstock nutrition for various species groups still remains a key challenge for the development of the industry. The general practice is to feed broodstock with as varied a diet as practicable (Job et al. 1997, 2002; Olivotto et al. 2005, 2006a,b, 2008a,b,c; Wittenrich et al. 2007; Lin et al. 2008; Murugan et al. 2009). In general, a varied mix of shrimp, fish, squid, mysid shrimp, Artemia sp. and fish flakes tends to be used.

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In addition, marine algae or even garden vegetables such as peas and lettuce may be included for omnivorous and herbivorous species, and live feeds may be included for species such as seahorses. There is a need, however, for more targeted research that identifies the dietary requirements of various marine ornamentals species groups in order to maximise egg quality and quantity. Broodstock are fed to satiation twice or three times a day (Olivotto et al. 2005, 2006a,b; Wittenrich et al. 2007). In addition to diet, temperature has also been shown to have a marked effect on gonad development, GSI, fecundity and reproductive efficiency of Hippocampus kuda (Lin et al. 2006). The shortest gonad development time and highest GSI was observed at 28 °C. The temperature range from 26–28 °C is the optimal temperature for fecundity and spawning of H. kuda (Lin et al. 2006). This temperature range is also likely to be optimal for most other tropical marine ornamentals species.

10.4

LARVAL CULTURE

Much of the scientific research on marine ornamentals has focused mainly on the larval stage as this is still considered the most difficult stage from an aquaculture perspective (Danilowicz & Brown 1992; Job & Bellwood 1996, 2000; Job & Shand 2001; Arvedlund et al 2000; Avella et al. 2007; Olivotto et al. 2003, 2006a,b, 2008a,b). Larval feeding and nutrition has been the primary focus of the work done to date, and most other aspects of larval culture of marine ornamentals remain poorly understood.

10.4.1

Physical environment

Larvae of marine ornamental fish have been reared in a range of tank types, sizes and shapes, ranging from small experimental tanks of a few litres in volume to tanks of over 200 L (Danilowicz & Brown 1992; Job et al. 1997; Olivotto et al. 2006; Wittenrich et al. 2007). Small tank sizes can potentially result in developmental artefacts in fish larvae (Blaxter 1988a; Schoedinger & Epifanio 1997), and also require greater water turnover rates due to the limited volume. On the other hand, larger tank sizes are inefficient due to the relatively small number of larvae per clutch for many species. They also require unnecessarily high inputs of live microalgae and zooplankton to achieve the required feed densities despite the relatively small numbers of larvae. Tanks of approximately 100–200 L in volume are an effective compromise, depending on species. Fibreglass, plastic and glass tanks have all been used for culture of marine ornamentals. In the case of glass tanks, the outsides and base of the rearing tanks are either painted black or completely covered with black panels (Job et al. 1997; Olivotto et al. 2003, 2005). In the case of fibreglass tanks, a blue or black background is normally used. A dark background is thought to improve the contrast of the prey items and increase larval feeding efficiency (Hinshaw 1985; Hoff 1996; Woods 2000b). Glass tanks tend to be rectangular or square in shape with flat bottoms, while fibreglass tanks tend to be conical-base circular tanks. High water quality in the larval rearing tanks is essential for obtaining good survival and growth rates. In broad terms, water quality parameters are maintained at very similar levels to those in the broodstock tanks (Olivotto et al. 2003, 2006a,b, 2008a,b). Temperature is generally maintained at 26–30 °C. Photoperiod used ranges from 14L:10D (Wittenrich et al. 2007) to 24L:0D (Olivotto et al. 2005). Light is usually provided by broad-spectrum

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fluorescent tubes or incandescent lights suspended above the tank, with a light intensity of approximately 15.4 μE.m-2.s-1 at the water surface (Job & Bellwood 2000). Gentle continuous aeration through standard aquarium airstones is used in the larval tanks. Fish larvae react more readily to prey that are moving noticeably, even if that movement is induced by turbulence from the aeration rather than intrinsic (S. Job, personal observation). Moderate levels of turbulence may improve the feeding efficiency of fish larvae by increasing prey encounter rates (MacKenzie & Miller 1994; MacKenzie & Kiorboe 1995). Some species such as the Centropyge angelfish, however, may require minimal turbulence during the first week after hatching (Olivotto et al. 2006b). Larvae of the yellow-tail damselfish, Chrysiptera parasema, also appear to be sensitive to turbulence, and no aeration is used in the larval rearing tank (Olivotto et al. 2003). Most marine ornamental fish larval culture is done using greenwater techniques during the early stages, with the exception of species without pelagic larvae such as seahorses and Banggai cardinalfish (Job et al. 1997, 2002, 2006; Olivotto et al. 2003, 2006a,b, 2008a,b). A wide range of microalgal species have been used in greenwater culture, including Nannochloropsis oculata, Chlorella spp., Isochrysis spp. and Tetraselmis sp. (Danilowicz & Brown 1992; Job et al. 1997; Avella et al. 2007; Olivotto et al. 2003, 2008a,b). Microalgal densities have ranged from 2,000 cells/mL to 50,000 cells/mL (Danilowicz & Brown 1992; Olivotto et al. 2008a). In general, N. oculata tends to be the dominant algal species used for greenwater culture. Greenwater culture is thought to help to maintain high water quality by consuming the nitrogenous waste products of fish larvae, and may improve prey contrast and visibility (Naas et al. 1992; Palmer et al. 2007). Greenwater techniques also reduce incidences of larvae pressing against the tank walls, which may reduce feeding efficiency and survival. Larval rearing tanks are usually set up either as static tanks in the early larval stages or on a slow drip continuous water exchange. Where static cultures are used, water quality is maintained through a daily manual water change of approximately 10–30%, or with an overnight gentle flushing (Job et al. 1997). New water is filtered before entering the rearing tank in order to ensure high water quality. The bottom of the tank is generally cleaned daily using a small siphon hose to prevent the accumulation of detritus. In most cases, tanks are switched to flow through or re-circulating systems once the larvae are past the rotifer feeding stage. A filter constructed from sections of PVC pipe with holes covered in nylon mesh (210–500 μm mesh) is connected to the inside of the rearing tank outlet to prevent larvae from being flushed from the tank. Flow rates for the water change range from 30 L.hr-1 in the early larval stages to 60 L.hr−1 in the oldest stages. The optimum photoperiod for rearing marine ornamental larvae is uncertain, and may vary depending on species and other factors. The tomato clownfish, Amphiprion melanopus, shows faster growth at 16L:8D compared to 24L:0D or 12L:12D (Arvedlund et al. 2000). This is similar to what has been observed with other marine fish species such as garramundi, Lates calcifer (Barlow et al. 1995). In contrast, larvae of the yellow-tail damselfish, C. parasema, only survive to settlement and beyond under 24L:0D (Olivotto et al. 2003). Larvae reared under 13L:11D did not survive past 3 days, and larvae reared under 16L:8D did not survive past 7 days (Olivotto et al. 2003). Similar results have been observed with first-feeding larvae of other species such as the sunrise dottyback, Pseudochromis flavivertex (Olivotto et al. 2006a), and the rabbitfish, Siganus guttatus (Duray & Kohno 1988). The specific reasons for these apparent species-specific differences are uncertain as 24L:0D cycles are unlikely to be experienced by tropical fish larvae in the wild.

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In experimental systems, larvae have been reared successfully in small tanks at a density of 5 larva/L (Olivotto et al. 2006a). Commercial operations tend to use larger tanks of 150–200 L, and stock larvae at approximately 3.0 larvae/L. Young damselfish larvae may be very sensitive to dissolved oxygen (DO) levels, and levels below 5 ppm may represent a critical point for some species (Olivotto et al. 2003). This has implications for the density of rotifers used as rotifer densities above 20 rotifers/mL could lead to oxygen depletion, and high larval mortality rates. Mortality rates in marine ornamentals larvae are highest in the first 24 hours after hatching, possibly due to physical damage to the larvae during hatching in captivity (Wilkerson 1998; Olivotto et al. 2006a). Survival rates tend to slowly taper off after the first 24 hrs (Olivotto et al. 2005).

10.4.2

Sensory capabilities

Larval fish are visual feeders with most feeding activity concentrated during the day (Blaxter 1986; Job & Bellwood 1996, 2000; Job & Shand 2001). The visual system of most larvae is functional just before or soon after yolk sac absorption (Blaxter 1988b). Work on the sensory capabilities of marine ornamentals larvae has demonstrated that the sensory capabilities of the larvae increase substantially with development and growth. Newly hatched larvae have relatively poor visual acuity (the ability to detect prey) (Job & Bellwood 1996), poor light sensitivity (Job & Bellwood 2000), and a narrower spectral range (Job & Shand 2001; Job & Bellwood 2007) compared to older larvae. In practice, this means that young larvae require relatively high light intensities and high prey densities in order to be able to detect and capture prey such as rotifers. In comparison, older larvae are able to detect prey at low light intensities and at lower prey densities. Older larvae also have greater reactive distances and a wider binocular field (Job & Bellwood 1996). Furthermore, older larvae are able to use a wider light spectrum, including UV light, to detect and capture prey. The result of these increased visual abilities is that older larvae are more efficient at detecting and tracking prey, and are able to feed at lower light levels than younger larvae. Thus, careful attention needs to be paid to factors such as light intensity and prey density with younger larvae. The ability of larvae to successfully capture prey also increases with larval development and growth. Capture success (i.e. the proportion of successful strikes) depends on the abilities of the larvae and on the escape abilities of the prey. In young larvae, prey capture rates are low and only a fraction of attacks are successful, although this varies strikingly between different taxa (Drost 1987; Coughlin 1994; Job & Bellwood 1996). The poor success may be caused by poor aiming accuracy or by active escape movements of the prey (Drost 1987). Capture success improves rapidly with larval age/size and is very high in later larval stages (Job & Bellwood 1996). The increase in capture success is due to an increase in sensory and locomotory abilities (Job & Bellwood 1996; Fisher et al. 2000). Thus, older larvae seldom miss prey that they strike at.

10.4.3

Larval feeding

Most marine ornamental fish larvae begin feeding on the first day after hatching. The tolerance of larvae to delayed feeding varies between species, depending on the size of the larvae and the amount of yolk reserves. In general, most marine ornamentals larvae need

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to be fed within 24 hours of hatching. The main exceptions are species that hatch out in an undeveloped form and are unable to feed until 24–48 hours after hatching (e.g. Centropyge angelfish (Olivotto et al. 2006b); some damselfish (Danilowicz & Brown 1992)) and species that hatch out as juveniles (e.g. seahorses). Newly released seahorse juveniles, for example, can tolerate 24–96 hours without food, but perform better if fed within 24–48 hours of being born (Sheng et al. 2007). Marine fish larvae require live zooplankton as the first prey items. The two main zooplankton used in marine finfish aquaculture are rotifers (Brachionus plicatilis and Brachionus rotundifornis) and brine shrimp (Artemia spp.). Many of the commonly cultured species such as the anemonefish and dottybacks survive adequately on these traditional aquaculture live feeds (Olivotto et al. 2006a, 2008a, 2008b). The main exceptions are species with very small larvae such as the Centropyge angelfish and some goby species (Holt 2003; Olivotto et al. 2006b; Wittenrich et al. 2007), and species such as seahorses (Job et al. 2002, 2006). For species that can be cultured using rotifers and Artemia spp., rotifers are used as the first food from hatching until the larvae are large enough to feed on Artemia spp. Artemia spp. are then used until the fish are developed enough to be weaned onto artificial or frozen feeds (usually at the juvenile stage). There is usually a 3–5 day overlap between the rotifers and the Artemia spp. to enable the larvae to slowly wean onto the new prey item. The specific age at which larvae are transitioned from rotifers to Artemia spp. varies depending on species, and is based primarily on larval size (Avella et al. 2007; Olivotto et al. 2006a, 2008a, 2008b). Juveniles are then slowly weaned onto pelleted feeds in commercial operations. Prey densities are usually maintained at between 10–15 prey/mL for rotifers, and 5 prey/ ml for Artemia spp. Live feeds are usually added incrementally to the tanks in the morning until the appropriate prey density is obtained, and topped up over the day where necessary. 10.4.3.1 Live feed enrichment Research on the nutritional requirements of anemonefish and damselfish larvae has demonstrated clearly the critical importance of HUFA enrichment of live feeds for larval survival and growth (Avella et al. 2007; Olivotto et al. 2003, 2008a,b). This is consistent with what has been observed to date with other marine fish species. Live feeds need to be separately enriched with specific enrichment products before use (Sargent et al., 1999; Sorgeloos et al. 2001). A range of enrichment products have been utilised, and appear to achieve broadly similar outcomes provided they have adequate levels of docosahexanoic acid (DHA), eicosapentanoic acid (EPA) and arachidonic acid (AA). While the importance of Artemia enrichment is well known, recent work has highlighted the importance of enriching the rotifers. In the yellow-tail damselfish, Chrysiptera parasema, for example, not enriching the rotifers with specific enrichment products resulted in total mortality within the first 48 hours, even where the rotifers had been fed with microalgae (Olivotto et al. 2003). This is consistent with what has been observed with the pseudochromid species, Pseudochromis flavivertex (Olivotto et al. 2006a). This is in contrast to current commercial practice with many marine foodfish species, where only the Artemia stage is enriched with specific enrichment products. In most cases, rotifers are only enriched with microalgae such as Nannochloropsis oculata, or a mix of microalgae (usually a mix of N. oculata and T-Isochrysis).

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Enrichment of Artemia spp. is well established in the aquaculture industry, and has been shown to increase growth and survival in numerous marine fish species (Sargent et al. 1999; Sorgeloos et al. 2001). Not enriching the Artemia resulted in reduced growth in Amphiprion ocellaris anemonefish, even where the rotifers had been enriched (Avella et al. 2007). Furthermore enriching both rotifers and Artemia resulted in reduced numbers of misbanded fish. Given that mis-banding is a significant issue for the industry, this suggests that enriching both live feeds may help to improve the quality of fish produced. However, there was no detectable effect of Artemia enrichment on survival (Avella et al. 2007). Recent work on Pseudochromis flavivertex supports the argument that both rotifers and Artemia need to be enriched for optimum results (Olivotto et al. 2006a). Larval survival and growth are impaired when rotifers are separately enriched with specific commercial products but the Artemia are not enriched as well (Olivotto et al. 2006a). Lack of enrichment for both rotifers and Artemia resulted in total mortality within the first week of the larval stage under experimental conditions (Olivotto et al. 2006a). Enrichment of live feeds also leads to more rapid settlement, with 85% of Pseudochromis flavivertex larvae completing metamorphosis and settlement by day 30 after hatch. In comparison, only 8% of larvae fed on enriched rotifers and unenriched Artemia completed metamorphosis and settlement by day 30 after hatch (Olivotto et al. 2006a). Inadequate live feed enrichment also leads to higher mortality rates in juveniles, particularly when exposed to stress. In the Banggai cardinalfish, for example, inadequate Artemia enrichment results in sudden fright syndrome, where juveniles faint and die on handling (Vagelli 2004). 10.4.3.2 Copepods and other non-traditional zooplankton One of the critical bottlenecks in the commercial production of a wider range of marine ornamental species is the challenge associated with getting larvae to survive on just traditional aquaculture live feeds such as rotifers and Artemia. A substantial amount of work has been done in recent years on producing calanoid or harpaticoid copepods, as well as other zooplankton species for use as first feed organisms (Stottrup 2000; McKinnon et al. 2003; Shields et al. 2005). Copepod stages are the primary prey of fish larvae in the wild, and there appears to be a preference for copepods in many larval fish species in captivity (Olivotto et al. 2006b). Recent work has evaluated the use of harpacticoid and calanoid copepods as live feeds for anemonefish larvae (Olivotto et al. 2008a,b). The results suggest that the harpacticoid copepod, Tisbe spp. is not adequate as a sole live feed organism for anemonefish larvae (Olivotto et al. 2008a). Larvae fed solely on Tisbe spp. nauplii followed by copepodites and adult copepods did not survive past day 6, but Tisbe spp. were beneficial as a supplement to rotifers and Artemia, and resulted in higher survival and growth (Olivotto et al. 2008a). Calanoid copepods, in contrast, are adequate as a sole live feed, and result in markedly higher survival and growth rates (Olivotto et al. 2008b). Survival in larvae fed on copepods was 90%, compared to 43% in larvae fed on rotifers and Artemia. The differences in the results between the harpacticoid and calanoid copepod species are thought to be due to differences in their behaviours rather than nutritional value (Olivotto et al. 2008a,b). Harpacticoid copepodite and adult stages are benthic, with only the nauplii stage being pelagic. In contrast all life history stages of the calanoid copepod are pelagic. This makes them more accessible as a live feed for fish larvae.

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Ciliates and wild-caught zooplankton have also been used as first foods, and have yielded promising results with some fish species (Job & Bellwood 2000; Job et al. 2002). In a study using wild-caught zooplankton, larvae of the goby, Priolepis nocturna were fed on the dinoflagellate Gymnodinium breve and tintinnids (Wittenrich et al. 2007). In Gobiosoma evelynae, survival was highest in larvae first fed enriched ciliates (Euplotes sp.) followed by enriched rotifers (Brachionus rotundiformis) as compared to larvae fed first on enriched B. rotundiformis followed by B. plicatilis (Olivotto et al. 2005). Goby larvae appear to prefer prey that are smaller in size than rotifers. Despite the progress in developing and utilising non-traditional live feeds such as copepods, the larvae of many pelagic spawners such as angelfish remain challenging to culture reliably (Olivotto et al. 2006b).

10.4.3.3 Prey selectivity Larvae are size selective when feeding, and select for increasingly larger prey organisms as they grow (Job & Bellwood 2000; Job et al. 2002). In work using wild-caught zooplankton (60–80% copepod nauplii and copepodites), mean prey size was maintained at approximately 4% of larval standard length (SL). For example, larvae below 3.5 mm SL were fed zooplankton between 53 and 125 μm (Job & Bellwood 2000). This was increased to 53– 210 μm for 3.5–5 mm larvae, 125–350 μm for 5–9 mm larvae, and 210–420 μm for larvae greater than 9 mm. In general, larvae prefer copepod nauplii or rotifers as a first food, followed by copepodites and then adult copepods or Artemia (Olivotto et al. 2008a,b). There are subtle variations between fish species, and some species display a marked preference for copepods compared to rotifers or Artemia, while other species prefer rotifers as a first food followed by copepod nauplii, then copepodites, then copepod adults and finally on-grown Artemia (Murugan et al. 2009). Many of the dietary shifts appear to be driven by prey size, but there may also be prey species selectivity in some fish. Different species of fish larvae follow different feeding strategies. Some species such as the damselfishes and anemonefishes are cruise predators (MacKenzie & Kiorboe 1995) and actively swim throughout the day, stopping only to feed. In contrast, cardinalfish larvae are pause-travel predators (MacKenzie & Kiorboe 1995) that spend considerable amounts of time stationary while searching for prey.

10.4.3.4 Artificial feeds Following the live feed stage, larvae are weaned onto an increasingly larger array of artificial feeds. This generally occurs from the juvenile stage onwards, i.e. after the fish have undergone metamorphosis and settlement. The weaning process is frequently a period of relatively high mortality as larvae that do not adapt readily to artificial feeds perish. The juveniles are first gradually weaned from Artemia nauplii onto pelleted feeds. Different species adjust to taking pelleted foods at different rates, and the mortality rate can be reduced by ensuring that the period of overlap between live feeds and artificial feeds is sufficiently long for most larvae to adapt. Some species such as seahorses are difficult to wean completely onto artificial pelleted feeds, and juvenile seahorses are usually weaned onto frozen diets such as frozen mysid shrimp instead (Woods 2003a; Lin et al. 2009a).

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10.5 10.5.1

JUVENILES Metamorphosis and settlement

Larval duration varies between species, and ranges from approximately 10 days in many anemonefish species to approximately 60 days in some angelfish species (Brothers et al. 1983; Wellington & Victor 1989). Larval durations are broadly similar in the laboratory compared to the field (Job & Bellwood 2000). Some species of marine ornamentals undergo a marked metamorphosis prior to settlement (e.g. anemonefishes and damselfishes (Job & Bellwood 1996, 2000); and dottybacks (Moe 1997), while other species undergo a gradual transition from the larval to the juvenile form (e.g. cardinalfishes (Job & Bellwood 2000)). Settlement in any given batch of cultured larvae occurs over a period of a few days, with some individuals settling faster than others (Job et al. 1997; Olivotto et al. 2006a). Metamorphosis and settlement in Pseudochromis flavivertex occurs between days 23 and 35 post-hatch at 27 °C with a photoperiod of 24L:0D (Olivotto et al. 2006a). Metamorphosis and settlement in P. fridmani (Fig. 10.8) occurs between days 18 and 28 post-hatch at 29 °C with a photoperiod of 14L:10D (S. Job, unpublished data). In both species, the majority of larvae settle within the first seven days. In Gobiosoma evelynae, larvae undergo metamorphosis and settlement between days 30 and 40 post-hatch (Olivotto et al. 2005). During this stage, the larvae transition from the pelagic to the demersal environment, and develop the juvenile coloration. The changes in coloration generally occur a couple of days after the fish settle on the bottom of the tanks (Fig. 10.9 & Fig. 10.10). The age at which metamorphosis and settlement occurs is relatively plastic in some marine ornamentals species such as the damselfish. Damselfish larvae will settle earlier if suitable settlement substrates such as live coral heads are provided (Danilowicz & Brown 1992).

Fig. 10.8

Captive-bred P. fridmani juveniles. (Please see plate section for colour version of this figure.)

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Fig. 10.9 Yellow-tail damselfish, Chrysiptera parasema, juveniles shortly after settlement, showing the adult colouration. (Please see plate section for colour version of this figure.)

Fig. 10.10 figure.)

Juvenile Amphiprion ocellaris clownfish. (Please see plate section for colour version of this

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Fig. 10.11 Newly released banggai cardinalfish, Pterapogon kauderni, juveniles. (Please see plate section for colour version of this figure.)

Seahorses and Banggai cardinalfish (Fig. 10.11) are an exception as juveniles are released from the parent fish (Job et al. 2002; Vagelli 2004). Seahorses and Banggai cardinalfish do not have a pelagic larval stage. Instead, the young are released at the juvenile stage. Newborn seahorses are still pelagic in their behaviour (Job et al. 2002), while Banggai cardinalfish are demersal from hatching (Vagelli 2004). In both cases, the young fish are relatively large at hatching. Once all the fish have undergone metamorphosis and settlement, they are generally transferred into larger growout tanks. Market size in most commonly cultured species is reached in 4–6 months. The size at which the market will accept aquacultured fish varies with species and between markets (countries and regions), and ranges from 3–5 cm. In general, market acceptance for small fish is increasing in many places. Closed cycle production occurs for many species that are cultured. In general, F1 fish begin to breed within 6–12 months of age. Egg production tends to be limited in small fish, and increases with fish size.

10.5.2

Seahorse juvenile rearing

10.5.2.1 Physical environment Much of the recent focus in seahorse aquaculture has been on the rearing of juveniles through to market size (Job et al. 2002; Lin et al. 2008, 2009a,b; Hora & Joyeux 2009; Murugan et al. 2009). Seahorse juveniles (Fig. 10.12) are pelagic for the first 14–15 days, and begin to attach to holdfasts after that time (Job et al. 2002; Hora & Joyeux 2009). Light

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Fig. 10.12 figure.)

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Juvenile seahorse at the pelagic stage. (Please see plate section for colour version of this

intensities used in rearing and experimental tanks range from 4.8 μEs-1 m-2 (MartinezCardenas & Purser, 2007) to 136 μEs-1 m-2 (Wong & Benzie 2003). Optimal light intensity for culturing may vary between seahorses of different ages. Young pelagic juveniles (0–6 days post-hatch) show greater survival at light intensities above 1,500 lux, with 2,000 lux (the highest light intensity tested) giving the best survival and growth rates in Hippocampus trimaculatus (Murugan et al. 2009). In contrast, sub-adult H. erectus displayed better growth rates in terms of wet weight gain at 1000 lux, and poorer growth at lower as well as higher light intensities (Lin et al. 2009b). However, growth in terms of SL was highest at a light intensity of 1500 lux (Lin et al. 2009b). Growth of older (sub-adult) H. whitei juveniles is not be affected by light intensities ranging from 24– 136 μEs-1 m-2 (Wong & Benzie 2003). The effect of tank background colour on feeding rate, growth and survival of juvenile seahorses is still uncertain. Martinez-Cardenas and Purser (2007) suggested that tank background colour has no significant effect on feeding rate, growth or survival in H. abdominalis from 3 days after hatch to 56 days after hatch. In contrast, Woods (2000a) suggests that 7-day-old H. abdominalis displayed a greater feeding rate in clear or white tanks rather than black tanks. The difference could in part be due to the type of enrichment used for the prey (Artemia). Woods (2000a) used a microalgae-based enrichment that would have resulted in the Artemia gut being darker coloured than in the study by Martinez-Cardenas & Purser (2007). Temperature has a marked effect on growth and survival (Lin et al. 2008). Highest level of growth and survival was obtained in H. erectus cultured at 28–29 °C. Lower temperatures down to 24 °C, as well as higher temperatures up to 33 °C resulted in lower growth and survival (Lin et al. 2008). At 28 °C, H. erectus reached 6.32 cm after 9 weeks, with a survival rate of 71% (Lin et al. 2008). In the temperate species, H. whitei, juveniles reared at

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26 °C were 48% heavier and 28% longer than seahorses reared at 17 °C (Wong & Benzie 2003). However, the condition of the seahorses declined with increasing temperature. The gonadosomatic index (GSI) was highest at 20 °C (Wong & Benzie 2003). Optimum salinity range varies between seahorse species. In H. erectus, growth rates were highest at a salinity of 31–33 ppt, and declined with higher or lower salinities (Lin et al. 2009b). H. kuda, which is able to tolerate a wide range of salinities in the wild, displays high growth and survival over a wide salinity range from 15–30 ppt, and can tolerate salinities of up to 50 ppt (Hilomen-Garcia et al. 2003). H. trimaculatus juveniles are able to tolerate salinities down to 26 ppt, while adults can tolerate salinities of as low as 17 ppt (Murugan et al. 2009). 10.5.2.2 Feeding Newly hatched seahorses are usually fed soon after hatching. Recent work suggests that the point of no return (point at which fish larvae will not feed even when offered food) is reached after 116.7 hours in H. trimaculatus and after 115.6 hours in H. kuda (Sheng et al. 2007). The first feeding rates of H. trimaculatus juveniles are highest (at 80%) when they are starved for 24 hours, intermediate if starved for 0 hours or 48 hours, and decline sharply after 48 hours (Sheng et al. 2007). The first feeding rates for H. kuda remain high (>= 80%) for the first 72 hours, and then decline markedly (Sheng et al. 2007). The feeding intensity in H. trimaculatus juveniles was highest after they had been starved for 24 hours. Feeding intensity was highest in H. kuda juveniles after they had been starved for 48 hours (Sheng et al. 2007). In general, young seahorses require various copepod stages in addition to rotifers and Artemia as a live feed in order to optimise grow and survival (Olivotto et al. 2008c; Hora & Joyeux 2009; Murugan et al. 2009). Young H. reidi seahorses fed with a combination of Tisbe spp. copepod nauplii and rotifers, followed by Artemia and Tisbe spp. copepodites, for example, showed substantially higher survival rates than seahorses fed solely on rotifers followed by Artemia (Olivotto et al. 2008c). However, juveniles fed solely on the harpacticoid copepod, Tisbe spp., showed a mean survival of only 4% at 7 days post-hatch. Thus, the harpacticoid copepod, Tisbe spp., is best used as a live feed supplement. In contrast, calanoid copepod stages are adequate as a sole prey item for syngnathid juveniles (Payne et al. 1998; Payne & Rippingale 2000). However, most calanoid copepod species are more difficult to produce reliably on a large scale than harpacticoid copepods (Olivotto et al. 2008b). Alternatively, seahorse juveniles can be reared successfully on size-sorted wild-caught zooplankton, followed by enriched Artemia (Job et al. 2002; Hora & Joyeux 2009). As with other marine fish species, Artemia used as prey for seahorses need to be enriched with high HUFA products (Wong & Benzie 2003). H. whitei fed on enriched Artemia were 28% heavier that those fed on unenriched Artemia when reared at 20 °C, and 19% heavier when reared at 26 °C (Wong & Benzie 2003). A range of commercial high-HUFA products have been tested with sub-adult H. abdominalis (Woods 2003b). The results suggested that the differences between them are minor, with all the enrichment products producing good growth and survival. Young seahorses show prey selectivity in terms of both prey species and prey size as they develop and grow (Job et al. 2002, 2006; Murugan et al. 2009). H. trimaculatus, for example, prefer rotifers immediately after hatching followed by copepod nauplii, then copepodites and finally adult copepods (Murugan et al. 2009). Artemia were not a preferred

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prey item. Prey densities are generally maintained at between 1–10 prey/mL, depending on the type of prey and the size and age of the juveniles (Job et al. 2002, 2006; Hora & Joyeux 2009). Higher prey densities tend to be used for younger juveniles. Digestive period was highest in young juveniles (3 hrs 20 min), and declined with development and growth (Murugan et al. 2009). Recent research has suggested that juvenile seahorses should not be weaned from live to frozen feeds too early as this results in reduced growth rates (Lin et al. 2009a). It has been suggested that H. erectus should be weaned onto frozen feeds at a weight of 0.54– 0.70 g wet weight or 6.59–7.46 cm SL (Lin et al. 2009a). The growth rate differences between juveniles that are weaned at different sizes are further magnified with continued growth, thus exacerbating the initial size differences. Similar results have been observed in H. abdominalis, where weaning 2-month old juveniles onto frozen copepods resulted in reduced growth and survival rates (Woods 2003a). Weaning seahorse juveniles onto artificial feeds has also proved challenging, with negative effects on growth (Woods 2003a). Feeding rates on frozen and artificial feeds are also lower than with live feeds (Woods 2003a). 10.5.2.3 Survival and growth Reported survival rates of cultured seahorses vary substantially across studies, and between and within species. Survival rates reported in H. abdominalis range from approximately 21% (Woods 2000a) to 80% over a 2-month period (Woods 2000b) and 67–87% over a 6-week period (Martinez-Cardenas & Purser 2007). Similarly, reported survival in H. reidi ranged from 10–35% at 21 days after hatching (Olivotto et al. 2008c) to 88.3% at 109 days after hatching (Hora & Joyeux 2009). Reported survival rates in other seahorse species include 65% after 26 weeks in H. trimaculatus (Murugan et al. 2009), 71% after 9 weeks in H. erectus (Lin et al. 2008), between 40% and 73% at 14 weeks in H. kuda (Job et al. 2002), and 90% after 9 weeks in H. comes (Job et al. 2006). As with other marine fish species, mortality in seahorse juveniles appears to be confined to the first few weeks after release from the male’s pouch, and may also be impacted by rapid dietary changes (Hora & Joyeux 2009). Growth rates of juvenile seahorses vary between species, with marked differences between temperate and tropical species (Job et al. 2006). In broad terms, tropical species such as H. erectus and H. comes reach a height of 33.29–39.40 mm at 35 days of age, depending on water temperature and interspecies differences (Job et al. 2006). In contrast, the temperate seahorse species, H. abdominalis, cultured at approximately 16–17.5 °C achieved a standard length of 36–43.04 mm at 56 days of age (Woods 2000a; MartinezCardenas & Purser 2007). Similarly the tropical species, H. trimaculatus, reached 119.9 mm in SL in 26 weeks compared to H. abdominalis, which reached 110.7 mm in 52 weeks (Woods 2000b; Murugan et al. 2009). The average growth rate varies even between tropical species, and ranges from 0.66 mm/ day in H. comes (Job et al. 2006) to 0.77 mm/day in H. reidi (Hora & Joyeux 2009) to 1.27 mm/day in H. trimaculatus (Murugan et al. 2009) and 0.90–1.53 mm/day in H. kuda (Job et al. 2002). The differences in growth rates were probably at least partially due to differences in the temperatures at which the various species were reared and differences in their final adult sizes (18.7 cm for H. comes, 19 cm for H. erectus, 17 cm for H. kuda and 35 cm for H. abdominalis) (Lourie et al. 2004). Furthermore, juveniles from different parents or broods within a species also show differences in growth rates (Lin et al. 2008).

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In general, however, tropical seahorse species appear to have substantially faster growth rates than temperate species. In sub-adult H. erectus, growth rates increased with decreased stocking density from 1.5 individuals/L to 0.25 individuals/L (Lin et al. 2009b). Pouches first became visible at approximately 4 months after hatching, and males tended to develop their pouches (a sign of reproductive maturity) earlier at higher stocking densities (Lin et al. 2009b). Furthermore, males tended to grow faster than females in the trial. Similar trends were observed in H. abdominalis juveniles tested at stocking densities ranging from 1–5 animals/L, where growth rates and condition factors increased as stocking density decreased (Woods 2003c). In contrast, there was no detectable effect of stocking density in 3-month-old H. whitei when tested over a narrow range of densities from 0.5–1 individuals/L (Wong & Benzie 2003). In H. trimaculatus, the first external signs of sexual maturity were observed at 95 days after hatching in males (pouch development), and at 115 days in females (dropping eggs) (Murugan et al. 2009). In H. reidi, the first visible signs of sexual maturity were apparent at approximately 60 days after hatch (Hora & Joyeux 2009). Similar results have been observed in other seahorse species as well. Gender segregation has no significant effect on the growth of H. abdominalis (Woods 2003c).

10.6

COMMERCIAL PRODUCTION

Despite significant interest in the development of commercial marine ornamental aquaculture globally, there are only a small number of marine ornamental enterprises that have remained in operation for 10 years or more. While most of the focus in the hobby and research literature appears to be on the technical challenges associated with producing marine ornamentals, there are a number of other fundamental issues that need to be addressed in order to make more widespread commercial marine ornamentals aquaculture a reality.

10.6.1

Market knowledge

It is essential to develop a thorough understanding of the marine ornamentals trade, particularly with regard to realistic market demand for individual species. One of the key challenges facing new marine ornamentals aquaculturists is gathering accurate information on the volumes traded for the target species. It is essential to gather information from as many sources as possible, and over a period of time. It frequently helps to also gather information from different levels in the trade (exporters, importers, wholesalers and retailers), where possible. This allows cross-validation of the market data. Another aspect of knowing the market is to understand the relationship between supply and pricing for individual species. The high prices commanded by many of the less common species, for example, are often a reflection of their rarity and the small numbers available. Increases in the supply of such species through aquaculture tend to result in price declines, and thus declining profit margins. Some of the less common true black percula anemonefish variants, for example, are substantially more valuable than the common percula anemonefish due to their relative rarity. When supplies are increased through aquaculture, however, either prices drop or demand plateaus. It is therefore critical to have a good understanding of the true market demand for less common species, and to regulate production accordingly.

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Production of controlled numbers of rare species can be lucrative, as long as production volumes are managed appropriately. A more problematic situation arises where the demand for some species may in fact be relatively inelastic, i.e., lower prices do not result in a significant increase in sales. In such cases, there are likely to be non-price related factors that constrain market size. This is frequently the case for aggressive species such as some of the larger dottybacks (e.g. Pseudochromis steenei) where fish compatibility is more likely to limit market size. Only a limited number of hobbyists keep predator tanks that are suitable for such species.Another aspect of knowing the market is being aware of major international developments that affect market demand. Tropical seahorses, for example, were reasonably easily available until their listing on Appendix 2 of CITES (Convention on the International Trade in Endangered Species) in 2004. The CITES listing temporarily resulted in supply from the wild declining while source countries developed species management plans. As a result, the prices for tank-bred seahorses increased sharply. This situation is changing as exports from the wild resume, and will undoubtedly continue to change in the future. Another example is provided by the aftermath of the release of the Pixar/Disney film Finding Nemo. This popular film resulted in a dramatic increase in demand for clownfishes (Amphiprion ocellaris and A. percula anemonefish in particular). Being aware of such changes while they are just looming on the horizon allows enterprises to prepare for these events, and thus take advantage of them, keeping in mind that such changes in demand may be relatively transient.

10.6.2

Product knowledge

Another important issue is to be familiar with the biological characteristics of the target species. Each species has its own little foibles and quirks, even within families. In some cases, these quirks can be quite critical. In species such as the dottybacks, their aggressiveness even after being paired can result in the loss of fish through injury. This is a serious issue as good broodstock are invaluable, and can be hard to replace. Knowing the fish well enough to design the broodstock tanks such that aggressiveness is minimised is therefore important as it helps to minimise such losses. In other cases, the characteristics of the juveniles can affect their marketability. In the case of the Solomon Islands variant of the true black percula anemonefish (Amphiprion percula) (Fig. 10.13), for example, the juveniles tend to develop their distinctive black coats several months after hatching in most cases. Thus fishes at market size (approximately 3–5 cm) usually only have relatively small areas that are already black. As a result, many new hobbyists are reluctant to pay the higher prices that this variant usually commands (as compared to standard percula anemonefish). This characteristic tends to reduce the demand for juvenile fishes of these species. This issue can be managed by working in collaboration with retail outlets. Retail outlets can help improve sales by having full-coloured adults in their display tanks or by having photographs of the adults on the fronts of the tanks that house the fish. This allows new hobbyists to know what the fish will eventually look like. Another, less cost-effective, option is to simply grow the fishes until they are large enough for the black coats to have developed.

10.6.3

Species diversity

The marine ornamentals trade is characterised by its high species diversity. Over 1,000 species are found in the trade. One of the challenges with marine ornamentals aquaculture

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Fig. 10.13 Solomon Islands variant of the true black percula anemonefish, Amphiprion percula. (Please see plate section for colour version of this figure.)

is that the actual numbers of fish traded for most species tend to be relatively low, compared to many other aquaculture industry segments. One of the problems facing new marine ornamentals enterprises is that the range of species produced in the early stages tends to be relatively low. In addition, many new enterprises focus on particular species groups. Anemonefish, for example, are probably the most commonly cultured species, and form the ‘bread and butter ’ species group of the trade. However, there tend to be numerous anemonefish producers, and competition can be high. Moreover, while the market for clownfishes as a family is large, this tends to be dominated by the ocellaris/percula anemonefish. So, with many clownfish species, new entrepreneurs can end up with products that are not particularly profitable. On the other hand, many medium-value species such as the dottybacks, while substantially more valuable than clownfishes, are traded in considerably smaller volumes. These fishes can be quite aggressive towards other fish, and may not be suitable for community tanks with more timid species. This tends to limit their popularity with many hobbyists. With these species, new entrepreneurs end up with products that have better profit margins, but are not generally sold in large numbers. One practical solution is to produce multiple species, and from multiple families. In many cases, this has the additional advantage of allowing aquaculturists to achieve economies of scale. Most importantly perhaps, it makes it easier for them to sell their products to retail outlets that are usually reluctant to stock large numbers of any one species.

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Another advantage of having greater species/family diversity is that such diversity ensures that revenue will remain steady even if individual species sales volumes vary. The demand for many individual species can fluctuate over the year and/or between years. Demand tends to be relatively stable for key species such as ocellaris/percula anemonefish and some seahorse and dottyback species, but may vary cyclically for others. These fluctuations in demand are a normal part of the marine aquarium trade, and indeed of the broader pet industry. Enterprises with only a few species may thus face serious difficulties during periods when their particular product mix is not in demand. Having a wider range of species for sale, however, would help circumvent this issue. In the long run, therefore, species/ family diversity is critical to the success of a marine ornamentals aquaculture enterprise.

10.6.4

Quality control

In the early days of marine ornamentals aquaculture, the quality of the fishes produced tended to be variable at best. Tank-bred fish soon developed a reputation for having poor coloration compared to wild-caught fishes, and being prone to diseases. With the development of better culturing protocols, tank-bred fish have lost their reputation for being sickly, and indeed are now considered hardier that wild-caught fishes in most cases. However, the reputation for being less vividly coloured than wild-caught fishes has been harder to lose. While most established commercial ventures have demonstrated that tank-bred fishes can in fact display similar coloration to wild-caught fishes, the fishes produced by many backyard operators and hobbyists can be quite drab even today. Such problems can be prevented through good nutrition and rigorous broodstock selection. A second issue is the occurrence of abnormal coloration patterns in some cultured fish. The presence of a small percentage of fishes in a batch with broken banding patterns is relatively common in some anemonefish species, for example. Having more than 5–10% of fish produced with broken bands can be detrimental to sales, as many hobbyists prefer fishes with full bands. Having said that, however, some hobbyists do find broken-banded fishes attractive (Fig. 10.14), and there is a small market for them. As a result, some producers have given their broken-banded fishes more marketable names (e.g. ‘designer banding’). Completely eradicating broken banding patterns has proven difficult with some species (Olivotto et al. 2008a,b). However, minimising the occurrence of such banding problems is far more achievable. Broken bands appear to be caused by a complex array of genetic, environmental and nutritional factors. In general, having good quality broodstock with full bands, and ensuring that the larvae and juveniles are reared under optimal conditions (particularly with regard to water quality and nutrition), will go a long way in minimising broken banding patterns. It should be noted that broken bands are generally only an issue in a few anemonefish species. Indeed, incompletely developed bands may be a natural characteristic of the juveniles of some species. True percula anemonefish juveniles, for example, start off with only the head-band, with the other two bands developing more slowly with growth. As a result, market-size juveniles may not possess fully developed tail bands in many cases. The most serious quality control issue is perhaps the presence of deformed fish due to poor nutrition or water quality management. Such fishes can suffer from a range of deformities, from incompletely developed gill covers to scoliosis or lordosis (spinal curvature). While severely deformed fishes are seldom sold, less seriously deformed fishes occasionally

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Fig. 10.14 Ocellaris clownfish displaying partial broken banding on one side. (Please see plate section for colour version of this figure.)

appear on the market, usually tucked in with normal fish. Supplying the market with deformed fishes will rapidly destroy a producer ’s reputation and credibility. Worse, it tends to give the industry as whole a bad reputation as well. Deformed fish should be culled. Most deformities can be prevented through good nutrition.

10.6.5

Broodstock selection

Good broodstock are the foundation of a successful marine ornamentals aquaculture enterprise. Ideally, broodstock should lay large egg batches, produce large larvae, display bright coloration, be resistant to disease, be non-aggressive and spawn readily. In the first instance, most enterprises are likely to start with wild-caught broodstock. Given that the quality of wild-caught broodstock is likely to be quite variable, particularly in terms of spawning characteristics, it is usually advisable to obtain a larger than necessary number of broodstock. This provides an enterprise with a range of potential broodstock from which they can select the best. In the long run, the best option is to use the best tank-bred fishes as future broodstock (FI broodstock). The potential F1 broodstock should be carefully selected for key characteristics such as fast growth rates, resistance to disease, bright vibrant coloration and mild temperaments (unaggressive). In general, poor quality broodstock are likely to produce poor quality offspring. The young from the F1 broodstock (F2s), thus, are more likely on average to display the characteristics for which their parents were selected. One of the challenges in developing F1 broodstock is that it is a long-term process. F1 broodstock take time to mature and become functional broodstock. In some species such

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as the orchid dottyback (and many other dottyback species), the fishes can take as little as 6 months to start spawning, although egg numbers tend to be relatively low for the first few months until the fish become larger. These fishes may become suitable as broodstock in as little as 12 months. In other species, however, F1 fishes may take two years or longer before they become suitable. The benefits of these F1 broodstock, however, far outweigh the costs over the longer term.

10.6.6

Competition

The main competition is usually against imported wild-caught fish. The main attractions of fish imported from overseas are the relatively low price and the high species diversity. The main disadvantages with imported fish are the poor and unpredictable quality of the fish, seasonal restrictions in supply and the reluctance of experienced hobbyists to purchase imported fish. As such, the combination of low price and good supply, but higher mortality rates for imported fish has attracted retail outlets that favour a rapid turnover model, where fish are sold as quickly as possible and at relatively low prices. The key advantage that aquacultured fish have over imported fish is their high quality. Many experienced hobbyists and established retail outlets prefer to purchase aquacultured fish due to the unpredictable quality of many imported fish. The price premium that customers are actually willing to pay, however, is generally less than what is reflected in surveys (e.g. 30% premium). In many cases, customers will only select for aquacultured fish if they are of very high quality and there is only a marginal cost difference. As such, marine ornamentals producers need to ensure that their fish are of excellent quality and reasonably priced in order to maintain a competitive advantage.

10.6.7

Issues and trends

Most of the early issues with tank-bred fish having pale colours have been resolved to some extent in established commercial enterprises through improved nutrition and water quality management practices, and the colours of aquacultured fish are now broadly similar to those of wild-caught fish in most cases. The beta-carotene derivative, astaxanthin, has been shown to be a critical factor in ensuring that the colours of aquacultured anemonefish are as bright and vivid as those of wild-caught fish (Hoff 1996). There is still a need to improve colours aside from the orange and red complex that is driven by beta-carotene derivatives. The biochemical drivers for some of these colours are known, but practical cost-effective implementation remains problematic. Another issue is the occurrence of broken banding patterns in species such as the anemonefish. While improved nutrition and water quality management has helped alleviate this to a large extent, some issues still remain and the primary drivers for broken banding patterns remain uncertain. One of the key bottlenecks is the lack of accessible information, as much of the existing work is being done by commercial enterprises and the available information is proprietary in nature. There is a substantial difference, for example, in the information that is publicly available for foodfish species as opposed to marine ornamental species. Furthermore, there are substantial gaps in the available information for marine ornamentals aquaculture. Information on the use of immunostimulants or probiotics to improve survival rates, for example, is still lacking.

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Other recent trends include the breeding of hybrid species such as the ‘cocoa anemonefish’ (which are a cross between A. ocellaris and Premnas biaculeatus) and the production of species variants (e.g. an almost completely white coloured A. ocellaris).

10.7

CONCLUSIONS

The overall popularity of marine aquaria has increased steadily over the past few decades. Where marine aquaria were once considered difficult and best left to experts, advancements in aquarium technology have now brought them within reach of even new hobbyists. Knowledge of filtration systems, light requirements and food requirements has led to an increase in demand for a high diversity of organisms for marine aquaria. Further, the evolution and popularity of ‘reef ’ displays has required collectors to provide an increasingly diverse range of live animals (O’Sullivan et al. 2008). On the supply side, growing concerns over the health of coral reefs has led to increasing levels of regulation of the collection and export of marine ornamentals from source countries. In addition, the deterioration of coral reefs worldwide from factors such as habitat degradation, coral bleaching, crown-of-thorns starfish infestations and coral diseases has reduced the effective habitat for a number of coral reef species. While aquaculture has helped fill the growing demand for marine ornamentals for a limited range of species, there is still a growing gap between supply and demand for most marine ornamental species in the trade. In addition, more and more customers are becoming concerned about the environmental sustainability of the trade, and are actively choosing marine ornamentals that are perceived as being from environmentally sustainable sources (especially aquacultured animals), provided that prices are broadly comparable. Marine ornamentals are attractive as an aquaculture product due to the high value of the fish, their short time to market, and the comparatively low infrastructure costs (Job 2005). Such high value at small size is largely unmatched in marine finfish aquaculture. The small size of most aquarium fish species also means that marine aquarium fish aquaculture facilities can be relatively small in terms of their physical footprint and infrastructure requirements, thus resulting in lower capital and operating costs. Many hobbyists also prefer tank-bred fish due to their superior quality. Having been ‘born and raised’ in an aquarium environment, tank-bred fish are fully adapted to life in aquaria. They readily take to standard aquarium foods such as flakes, are disease and stress resistant and thrive under aquarium conditions. This is in sharp contrast to many wildcaught fish that tend to be fussy about foods, highly stressed in aquaria and easily succumb to disease. Moreover, wild fish that are caught with cyanide generally suffer from damage to vital organs, and many die within a few months in aquaria. The high quality of tank-bred fish has made them firm favourites with experienced hobbyists. Tank-bred fish are gradually becoming the only reliable source of key species for the global marine aquarium fish trade in the face of increasingly constrained supplies from the wild. The trend towards greater levels of regulation of the collection of fishes from coral reefs is likely to increase with time. These restrictions on collection have already increased the demand for tank-bred fishes of species such as seahorses. Despite the business potential and popularity of tank-bred fish globally, the development of the marine ornamentals aquaculture industry has been relatively slow. One of the reasons for this is that marine ornamentals aquaculture is a multi-species industry, with all the

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concomitant challenges that this brings. As such, it is far different from standard foodfish aquaculture, and is more similar in many ways to the freshwater ornamentals aquaculture industry. Establishing a successful marine ornamentals aquaculture enterprise requires being able to produce a wide range of key species, having very high quality products, and having excellent marketing and market research skills.

10.8

REFERENCES

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Drost, M.R. (1987) Relationship between aiming and catch success in larval fishes. Canadian Journal of Fisheries & Aquatic Science, 44, 304–315. Duray, M. & Kohno, H. (1988) Effects of continuous lighting on growth and survival of first-feeding larval rabbitfish, Siganus guttatus. Aquaculture, 109, 311– 321. Emata, A., Borlongan, I. & Damaso, J. (2000) Dietary vitamin C and E supplementation and reproduction of milkfish Chanos chanos Forsskal. Aquaculture Research, 31, 557–564. Emata, A., Ogata, H., Garibay. E. & Furuita, H. (2003) Advanced broodstock diets for the mangrove red snapper and potential importance of arachidonic acid in eggs and fry. Fish Physiology & Biochemistry, 28, 489–491. Faleiro, F., Narciso, L. & Vicente, L. (2008) Seahorse behavior and aquaculture: how to improve Hippocampus guttulatus husbandry and reproduction? Aquaculture, 282, 33–40. Fernandez-Palacios, H., Izquierdo, M.S., Robiana, L., Valencia, A., Salhi, M. & Vergara, J., M. (1995) Effect of n-3 HUFA level in broodstock diets on egg quality of gilthead sea bream (Sparus aurata L.). Aquaculture, 132, 325–337. Fisher, R., Bellwood, D.R. & Job, S.D. (2000) Development of swimming abilities in reef fish larvae. Marine Ecology Progress Series, 2002, 163–175. Foster, S.J. & Vincent, A.C.J. (2004) Life history and ecology of seahorses: implications for conservation and management. Journal of Fish Biology, 65, 1–61. Furuita, H., Tanaka, H., Yamamoto, T., Shiraishi, M. & Takeuchi, T. (2000) Effects of n-3 HUFA levels in broodstock diet on the reproductive performance and egg and larval quality of the Japanese flounder, Paralicthys olivaceus. Aquaculture, 187, 387–398. Furuita, H. Tanaka, H., Yamamoto, T., Suzuki, N. & Takeuchi, T. (2002) Effects of high levels of n-3 HUFA in broodstock diet on egg quality and egg fatty acid composition of Japanese flounder, Paralicthys olivaceus. Aquaculture, 210, 323–333. Furuita, H., Yamamoto, T., Shima, T., Suzuki, N. & Takeuchi, T. (2003) Effect of arachidonic acid levels in broodstock diet on larval and egg quality of Japanese flounder Paralicthys olivaceus. Aquaculture, 220, 725–735. Furuita, H., Tanaka, H., Yamamoto, T., Shiraishi, M. & Takeuchi, T. (2004) Effects of high dose of vitamin A on reproduction and egg quality of Japanese flounder Paralichthys olivaceus. Fisheries Science, 67, 606–613. Helfman, G.S. (1993) Fish behaviour by day, night and twilight. In: Behaviour of Teleost Fishes (ed. T.J. Pitcher), 2nd edn. Chapman & Hall, London. Hilomen-Garcia, G.V., Reyes, R.D. & Garcia, C.M.H. (2003) Tolerance of seahorse Hippocampus kuda (Bleeker) juveniles to various salinities. Journal of Applied Ichthyology, 19, 94–98. Hinshaw, J.M. (1985) Effects of illumination and prey contrast on survival and growth of larval yellow perch Perca flavescens. Transactions of the American Fisheries Society, 114, 540–545. Hoff, F.H. (1996) Conditioning, spawning and rearing of fish with emphasis on Marine Clownfish. Aquaculture Consultants Inc., Dade City, FL. Hioki, S., Katsumi, S. & Tanaka, Y. (1990) Development of eggs and larva in the angelfish, Centropyge ferrugatus. Japan Journal of Ichthyology, 37(1), 34–38. Holt, G.J. (2003) Research on culturing the early life history stages of marine ornamental species. In: Marine Ornamental Species: Collection, Culture and Conservation (eds J.C. Cato & C.L. Brown), pp. 251–254. Iowa State Press, Ames, IA. Holt, G.J. & Riley, C.M. (2001) Laboratory spawning of coral reef fishes: Effects of temperature and photoperiod. Proceedings of the 28th US–Japan Natural Resources Aquaculture Panel: Spawning and Maturation of Aquaculture Species. UJNR Technical Report No. 28, 33–38. Hora, M.S.C. & Joyeux, J-C. (2009) Closing the reproductive cycle: Growth of the seahorse Hippocampus reidi (Teleostei, Syngnathidae) from birth to adulthood under experimental conditions. Aquaculture, 292, 37–41. Job, S.D. (2005) Integrating marine conservation and sustainable development: Community-based aquaculture of marine aquarium fish. SPC Live Reef Fish Information Bulletin 13, 24–29. http://www.spc.int/ coastfish/news/LRF/13/index.htm Job, S.D. & Bellwood, D.R. (1996) Visual acuity and feeding in larval Premnas biaculeatus. Journal of Fish Biology, 48, 952–963. Job, S.D. & Bellwood, D.R. (2000) Light sensitivity in larval fishes: implications for vertical zonation in the pelagic zone. Limnology & Oceanography, 45, 362–371. Job, S.D. & Bellwood, D.R. (2007) Ultraviolet photosensitivity and feeding in larval and juvenile coral reef fishes. Marine Biology, 151, 495–503.

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Job, S.D. & Shand, J. (2001) Spectral sensitivity of larval and juvenile coral reef fishes: implications for feeding in a variable light environment. Marine Ecology Progress Series, 214, 257–277. Job, S.D., Arvedlund, M. & Marnane, M. (1997) Culture of coral reef fishes. Austasia Aquaculture, 11, 56–59. Job, S.D, Do, H.H., Meeuwig, J.J. & Hall, H.J. (2002) Culturing the oceanic seahorse, Hippocampus kuda. Aquaculture, 214, 333–341. Job, S.D., Buu, D. & Vincent, A.C.J. (2006). Growth and survival of the tiger tail seahorse Hippocampus comes. Journal of the World Aquaculture Society, 37, 322–327. Kerrigan, B.A. (1997) Variability in larval development of the tropical reef fish Pomacentrus amboinensis (Pomacentridae): the parental legacy. Marine Biology, 127, 395–402. Kuwamura, T. (1985) Social and reproductive behaviour of three mouthbrooding cardinalfishes, Apogon doederleini, A. niger and A. notatus. Environmental Biology of Fishes, 13, 17–24. Lee, K. & Dabrowski, K. (2004) Long-term effects and interactions of dietary vitamins C and E on growth and reproduction of yellow perch, Perca flavescens. Aquaculture, 230, 377–389. Li, Y., Chen, W., Sun, Z., Chen, J. & Wu, K. (2005) Effects of n-3 HUFA content in broodstock diet on spawning performance and fatty acid composition of eggs and larvae in Plectorhynchus cinctus. Aquaculture, 245, 263–272. Lin, Q., Lu, J.Y. & Gao, Y.L. (2006) The effect of temperature on gonad, embryonic development and survival rate of juvenile seahorses, Hippocampus kuda Bleeker. Aquaculture, 254, 701–713. Lin, Q., Gao, Y.L., Sheng, J.Q., Chen, Q.X., Zhang, B. & Lu, J.Y. (2007) The effect of food and the sum of effective temperature on the embryonic development of the seahorse, Hippocampus kuda Bleeker. Aquaculture, 262, 481–492. Lin, Q., Lin, J. & Zhang, D. (2008) Breeding and juvenile culture of the lined seahorse, Hippocampus erectus Perry, 1810. Aquaculture, 277, 287–292. Lin, Q., Lin, J., Zhang, D. & Wang, Y. (2009a) Weaning of juvenile seahorses Hippocampus erectus Perry, 1810 from live to frozen food. Aquaculture, 291, 224–229. Lin, Q., Lin, J., Zhang, D. & Wang, Y. (2009b) Effects of light intensity, stocking density, feeding frequency and salinity on the growth of sub-adult Hippocampus erectus Perry 1810. Aquaculture, 292, 111– 116. Lourie, S.A., Vincent, A.C. & Hall, H.J. (1999). Seahorse: An Identification Guide to the World’s Species and their Conservation, p. 214. Project Seahorse, London. Lourie, S.A., Foster, S.J., Cooper, E.W.T. & Vincent, A.C.J. (2004) A Guide to the identification of seahorses. Project Seahorse and TRAFFIC North America. University of British Columbia and World Wildlife Fund, Washington DC, USA. MacKenzie, B.R. & Kiorboe, T. (1995) Encounter rates and swimming behaviour of pause-travel and cruise larval fish predators in calm and turbulent laboratory environments. Limnology & Oceanography, 40, 1278–1289. MacKenzie, B.R. & Miller, T.J. (1994) Evidence for a dome-shaped relationship between turbulence and larval fish ingestion rates. Limnology & Oceanography, 39, 1790–1799. Martinez-Cardenas, L. & Purser, G.J. (2007) Effect of tank colour on Artemia ingestion, growth and survival in cultured early juvenile pot-bellied seahorses (Hippocampus abdominalis). Aquaculture, 264, 92–100. Mazorra, C., Bruce, M., Bell, J.G., et al. (2003) Dietary lipid enhancement of broodstock reproductive performance and egg and larval quality in Atlantic halibut (Hipppoglassus hippoglassus). Aquaculture, 227, 21–33. McAlary, F.A. & McFarland, W.N. (1993) The effect of light and darkness on hatching in the pomacentrid Abudefduf saxatilis. Environmental Biology of Fishes, 37, 237–244. McCormick, M.I. (2003) Consumption of coral propagules after mass spawning enhances larval quality of damselfish through maternal effects. Oecologia, 136, 37–45. McKinnon, A.D., Duggan, S., Nichols, P.D., Rimmer, M.A., Semmems, G. & Robino, B. (2003) The potential of tropical paracalanid copepods as live feeds in aquaculture. Aquaculture, 223, 89–106. Moe, M.A. (1997) Breeding the Orchid dottyback, Pseudochromis fridmani: an Aquarist’s Journal. Green Turtle Publications, Islamorada, FL.. Murugan, A., Dhanya, S., Sreepada, R.A., Rajagopal, S. & Baalasubramanian, T. (2009) Breeding and mass-scale rearing of three spotted seahorse, Hippocampus trimaculatus Leach under captive conditions. Aquaculture, 290, 87–96. Naas, K.E., Naess, T. & Harboe, T. (1992) Enhanced first feeding of halibut larvae (Hippoglossus hippoglosus L.) in green water. Aquaculture, 105, 143–156.

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O’Sullivan, D., Clark, E. & Morison, J. (2008) The Australian ornamental fish industry in 2006/07. FRDC project no. 2007/238. Fisheries Research and Development Corporation, Department of Agriculture, Fisheries and Forestry, Canberra. Okuda, N. (1999) Sex roles are not always reversed when the potential reproductive rate is higher in females. American Naturalist, 153, 540–548. Olivier, K. (2001) FAO/Globefish Research Programme, Vol. 67. United Nations Food and Agriculture Organisation, Rome. Olivotto, I., Cardinali, M., Barbaresi, L., Maradonna, F. & Carnevali, O. (2003) Coral reef fish breeding: the secrets of each species. Aquaculture, 224, 69–78. Olivotto, I., Zenobi, A., Rollo, A., Migliarini, B., Avella, M. & Carnevali, O. (2005) Breeding, rearing and feeding studies in the cleaner goby Gobiosoma evelynae. Aquaculture, 250, 175–182. Olivotto, I., Rollo, A., Sulpizio, R., Avella, M., Tosti, L. & Carnevali, O. (2006a) Breeding and rearing the sunrise dottyback Pseudochromis flavivertex: the importance of live prey enrichment during larval development. Aquaculture, 255, 480–487. Olivotto, I., Holt, S.A., Carnevali, O. & Holt, J.G. (2006b) Spawning, early development and first feeding in the lemonpeel angelfish Centropyge flavissimus. Aquaculture, 253, 270–278. Olivotto, I., Capriotti, F., Buttino, I., et al. (2008a) The use of harpacticoid copepods as live prey for Amphiprion clarkia larviculture: effects on larval survival and growth. Aquaculture, 274, 347–352. Olivotto, I., Buttino, I., Borroni, M., Piccinetti, C.C., Malzone, M.G. & Carnevali, O. (2008b) The use of the Mediterranean calanoid copepod Centropages typicus in yellowtail clownfish (Amphiprion clarkii) larviculture. Aquaculture, 284, 211–216. Olivotto, I., Avella, M.A., Sampaolesi, G., Piccinetti, C.C., Navarro Ruiz, P. & Carnevali, O. (2008c). Breeding and rearing the longsnout seahorse Hippocampus reidi: rearing and feeding studies. Aquaculture, 283, 92–96. Palma, J., Stockdale, J., Correia, M. & Andrade, J.P. (2008) Growth and survival of adult long snout seahorse (Hippocampus guttulatus) using frozen diets. Aquaculture, 278, 55–59. Palmer, P.J., Burke, M.J., Palmer, C.J. & Burke, J.B. (2007) Developments in controlled green-water larval culture technologies for estuarine fishes in Queensland, Australia and elsewhere. Aquaculture, 272, 1–21. Papanikos, N., Phelps, R.P, Williams, K., Ferry, A. & Maus, D. (2004) Egg and larval quality of natural and induced spawns of red snapper Lutjanus campechnus. Fish Physiology & Biochemistry, 28, 487–488. Payne, M.F. & Rippingale, R.J. (2000) Rearing West Australian seahorse, Hippocampus subelongatus, juveniles on copepod nauplii and enriched Artemia. Aquaculture, 188, 353–361. Payne, M.F., Rippingale, R.J. & Longmore, R.B. (1998) Growth and survival of juvenile pipefish (Stigmatopora argus) fed live copepods with high and low HUFA content. Aquaculture, 167, 237–245. Planas, M., Chamorro, A., Quintas, P. & Vilar, A. (2008) Establishment and maintenance of threatened long-snouted seahorse, Hippocampus guttulatus, broodstock in captivity. Aquaculture, 284, 19–28. Randall, J.E., Allen, G.R. & Steene, R.C. (1990) Fishes of the Great Barrier Reef and Coral Sea. Crawford House Press, Bathurst. Salze G., Tocher, D., Roy, W. & Robertson, D. (2005) Egg quality determinants in cod (Gadus morhua L.): egg performance and lipids from farmed and wild broodstock. Aquaculture Research, 36, 1488–1499. Sargent, J., McEvoy, L., Estevez, A., et al. (1999) Lipid nutrition of marine fish during early development: current status and further directions. Aquaculture, 170, 217–229. Schoedinger, S.E. & Epifanio, C.E. (1997) Growth, development and survival of larval Tautoga onitis (Linnaeus) in large laboratory containers. Journal of Experimental Marine Biology & Ecology, 210, 143–155. Sheng, J.Q., Lin, Q., Chen, Q.X., Gao, Y.L., Shen, L. & Lu, J.Y. (2006) Effects of food, temperature and light intensity on the feeding behavior of three-spot juveniles, Hippocampus trimaculatus Leach. Aquaculture, 256, 596–607. Sheng, J., Lin, Q., Chen, Q., Shen, L. & Lu, J. (2007) Effect of starvation on the initiation of feeding, growth and survival rate of juvenile seahorses, Hippocampus trimaculatus Leach and Hippocampus kuda Bleeker. Aquaculture, 271, 469–478 Shields, R.J., Kotani, T., Molnar, A., Marion, K., Kobashigawa, J. & Tang, L. (2005) Intensive cultivation of a subtropical paracalanid copepod, Parvocalanus sp., as prey for small marine fish larvae. In:

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11 Tilapia Luan Dinh Tran, Trung Van Dinh, Thoa Phu Ngo and Ravi Fotedar

11.1 INTRODUCTION Tilapia is the common name applied to three genera and more than 70 species of fish in the family Cichlidae: Oreochromis, Sarotherodon and Tilapia (Macintosh & Little 1995). Several characteristics distinguish these three genera, but possibly the most critical relates to reproductive behaviour. All Tilapia species are nest builders; fertilised eggs are guarded in the nest by a brood parent. Species of both Sarotherodon and Oreochromis are mouth brooders; eggs are fertilised in the nest but parents immediately pick up the eggs in their mouths and hold them through incubation and for several days after hatching. In Oreochromis species only females practise mouth brooding, while in Sarotherodon spp., the male or both male and female are mouth brooders. Tilapias are among the most important warm-water fish species used for aquaculture production and originated from Africa and the Middle East (Feryer & Iles 1972). Since then tilapia farming has grown considerably in terms of area and production (Fitzsimmons 2000). One advantage of tilapia is that they can feed on a wide range of food from natural organisms to artificial pellets (Bowen 1982; Jauncey & Ross 1982). To date, several tilapia species and hybrids have been widely distributed throughout the tropics, subtropics and temperate continents for culture purposes (Eknath 1995). The species that are most important for aquaculture are in the genus Oreochromis, including the Nile tilapia, O. niloticus, the Mozambique tilapia, O. mossambicus, the blue tilapia, O. aureus, and O. urolepis hornorum (Watanabe et al. 2002). Nile tilapia (Oreochromis niloticus) is the most important, constituting 90% of all tilapia cultured outside Africa (FAO 2004a). Most of the culturing of Nile tilapia in developing countries is carried out in polyculture systems with carps or shrimp, and production is on semi-intensive or small-scale levels. In recent times, the production and areas of tilapia culture have been increasing due to development of quality seed and relevant production techniques (Little 2004). Large gaps in tilapia productivity between small-scale and commercial levels, as well as variation in production conditions, will require different research approaches to meet the future demands. Evaluation of the Nile tilapia strain was conducted at several locations after the introduction of tilapia. Macaranas et al. (1997) reported that the Chitralada strain of O. niloticus

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

Tilapia 319 Table 11.1 Global aquaculture production of Oreochromis niloticus (adapted from FAO Fishery Statistics). Year 1950 1960 1970 1980 1990 2000 2009

Production in tons (1000) 5 10 20 80 220 820 2,340

showed the best reproduction, growth and survival between four strains evaluated. The Egypt and Ivory Coast strains were more reproductive than the Victoria and Segana strains, and they showed similar growth performance (Osure & Phelps 2006). The increasing interest in tilapia culture and the almost unanimous acceptance that cultured tilapia stocks needed genetic assessment and improvement led to the birth of the regional research and development programme ‘Genetic Improvement of Farmed Tilapia’ (GIFT), under the leadership of the World Fish Centre (WFC), based in Penang, Malaysia (then referred to as the International Centre for Living Aquatic Resources Management or ICLARM, based in Manila, Philippines) (de Silva et al. 2004). After Genetically Improved Farmed Tilapia (GIFT) dissemination, this strain indicated a superior growth rate and it was recommended for aquaculture in Asian countries (Sifa et al. 1999; Dey et al. 2000; Dan & Little 2000). Most results of on-farm trials with GIFT showed higher growth rates than other strains (Dey et al. 2000). Moreover, new O. niloticus strains such as GET EXCEL, GSTs and GMT have served well in tilapia aquaculture (Tayamen 2004; Zimmermann & Natividad 2004). In general, Nile tilapia plays an important role in aquaculture systems. In the twenty-first century, tilapias are likely to become the most important of all cultured fishes. Production of tilapia is widely distributed in 85 countries around the world. Their yield has increased rapidly (Table 11.1) and the worldwide production of Tilapia exceeded 2.5 million tons in 2007 (FAO 2010). Total tilapia production is mainly Nile tilapia (Oreochromis niloticus). All new countries entering tilapia production concentrate on this species. Total tilapia production in 2010 can be forecast to be 3.5 million tons with the bulk coming from aquaculture, the majority of the increase likely to come from Nile tilapia and be produced in China with almost 1 million tons production in 2005 (FAO 2010). The dramatic development of tilapia aquaculture production has been due to the application of advanced technology on seed production; culture technology; marketing, nutrition and genetic improvement aspects as well as better environment and disease management. This chapter reviews the contribution of these factors to the fast development of Tilapia aquaculture. Several other publications provide excellent reviews on some aspects of tilapia culture including those by de Silva et al. (2004), El-Sayed (2006), and Lim and Webster (2006).

11.2

SEED PRODUCTION

At present, monosex (male) tilapia fingerlings can be produced in pond, tank and hapa systems (Green 2006). Incubation facilities (Fig. 11.1) are required where fertilised eggs

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Fig. 11.1 Tilapia eggs incubator system at the National Broodstock Centre – Vietnam.

and sac fry are collected. Production of fertilised eggs and sac fry requires more intensive management than harvest of swim-up fry from earthen ponds, but daily seed production is higher and total output can be greater. The reliable and successful technologies for tilapia fingerling production focus on broodstock management, genetic selection, monosex culture, spawning and initial larval rearing, larval feed production and disease management in hatchery (Green et al. 1997; Green 2006).

11.2.1

Broodstock selection and management

The quantity and quality of the offspring depends on the quality of broodstock. In order to ensure good quality and quantity of offspring, the broodstock selection process should consider the following characteristics (Gunasekera et al. 1996):

• •

Brood fish should be of genetically pure origin. The important thing is that fish of unknown or questionable origins must be destroyed. Furthermore, brood fish can be selected from populations under a selection and improvement programme. Brood fish should not be too small, too young or too old because small fish are less fecund and less efficient at egg incubation and fry protection. The perfect weight for broodstock ranges from 0.3 to 0.5 kg. The fecundity decreases with maternal age and

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• •

successive spawning. Nile tilapia can mature sexually at 6 months of age at a weight of approximately 40 g (Macintosh & Little 1995). Brood fish should be in good shape, without deformities and injuries. Inferior and unwanted tilapia must be prevented from entering broodstock ponds; therefore, incoming water should be filtered continuously.

In addition, quality of the seed depends not only on brood fish selection but also broodstock management. Obtaining quality broodstock requires the knowledge of managing and selecting breeders. It is imperative to keep records of the breeders to manage the stocks. These records need to be collected for each generation, and include numbers, sex, holding facility, growth rate, survival, fecundity and deformities. The key to good management is obtaining and maintaining good quality broodstock. The hatchery should obtain parent stocks of known origin, usually from the authorised research stations. The broodstock should have good performance in term of good growth rate, lowest feed conversion ratio, disease resistance, etc. Broodstock are protected through management practices and this prevents the loss of quality. The following is a list of the requirements for successful management of broodstock. If a broodstock provider lacks any of these requirements, it will be difficult to manage stocks properly.

• • • • •

The hatchery operator and the assistant staff need to be well trained in overall hatchery operation, including genetic aspects of broodstock management. Enough ponds, tanks, and hapas are needed to hold broodstock fish of different varieties, ages and sexes and keep them separate so there is no mixing and uncontrolled breeding between them. The hatchery facilities need to be safe from flooding, theft or any other disturbances that can result in broodstock fish getting lost or mixed up together or mixed with wild stocks. Unwanted fish in the pond must be destroyed before introducing new stock, to prevent mixing of stocks. The breeding pond should be away from main roads, free from any noise stress because the noise affects the brooding female and eggs may be expelled from her mouth.

11.2.2

Nursing and harvesting of fingerlings

Tilapia fry feeds mainly on detritus and neuston, and juveniles feed on detritus and periphyton. In ponds, the natural foods available to individual fish depend on a number of factors such as soil fertility, type and amount of fertilisers added, and the number and weight of fish stocked. The use of live food for initial larval stage of tilapia could enhance the growth performance and survival rate. In intensive culture, the feeding practice must consider the mouth size and feed size especially for the initial larval stages. During larval stages, the fish require a daily ration of about 20–30% of their body weight, divided into six to eight feedings. The fry, which have left their mother ’s mouth a few days after hatching and begun to eat food, are reared for 15–25 days until they become fingerlings (about 1–3 g). Fry are very delicate and small. Their movement and ability to feed are weak. Their diet is restricted and they have a high metabolic rate. Rearing should be carefully managed to maximise

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survival rate and produce healthy fingerlings. Growth rate is very rapid during fry and fingerling stages. During these stages they have different biological characteristics from adults, especially in terms of feeding habits, growth and habitat preferences. The growth rate declines as the fish get older. At the fingerling stage, the growth rate is 5 to 6 times less than at the fry stage.

11.3

CULTURE PRACTICES

It is well accepted that tilapia males grow much faster than females, and therefore farmers prefer all male tilapia to female fish for culture (Little et al. 2003). Another major problem of pond-cultured tilapia is excessive reproduction, and subsequent stunting of fish due to overcrowding. The culture of tilapia males also overcomes the problem of massive recruitment resulting from excessive reproduction. Nowadays, all-male tilapia have been used for all tilapia culture models including pond culture, cage culture and indoor culture.

11.3.1

Pond culture

Pond culture (Fig. 11.2) is the most commonly used method of growing tilapia. One of the big advantages with a pond culture is that it closely resembles the life of wild tilapia and makes it possible for the fish to feed on naturally occurring food. Unfortunately, tilapia loves to spawn in ponds and the number of fry and fingerlings can rapidly reach large quantities if male and female fish are kept together. This will result in a situation where fry

Fig. 11.2

Pond culture of tilapia at the National Broodstock Centre – Vietnam.

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Fig. 11.3 Cage or happa culture of tilapia in Vietnam.

and fingerlings compete for food with the adults, resulting in a lower growth speed for the fish in the pond. One way of solving the problem is to cultivate only male fish in the pond. Ponds also tend to become a part of the natural landscape, for good and for bad. When animals such as birds and snails are attracted to the pond, they can bring parasites that cause trouble for tilapia.

11.3.2

Cage culture

Tilapia cage culture (Fig. 11.3) involves raising fish in cages of nylon nettings and bamboo or wood frames that are floated, submerged or fixed at the bottom. It utilises bodies of water such as dams, rivers, lakes, bays, reservoirs and coves. This is one of the effective technologies used in growing tilapia. Tilapia cage culture model has many advantages such as: easier handling, inventory and harvesting of fish, better control of fish population, efficient control of fish competitors and predators, effective use of fish feeds, reduced mortality, high stocking rate, total harvesting and swift or immediate return of investment, less manpower requirement, and minimum supervision. Depending on investment level and the topography where the fish cage is placed, two types of cage culture can be applied: fixed or floating. The success of tilapia cage culture depends on several factors including water quality, water level, the species used, stocking size and density, cage size and shape, feed quality and feeding frequency.

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11.3.3

Indoor tilapia culture

The indoor tilapia culture model involves technology to raise tilapia in composite (Fig. 11.4) or cement (Fig. 11.5) tanks with a water circulating system. Tilapia is cultured in tanks and raceways of varying sizes and shapes (circular, rectangular, square and oval). An important characteristic of tank design is the effective removal of solid waste; a circular tank with a central drain is the most efficient design. This type of culture has many advantages when compared with the pond culture model. It is easy to control the environmental factors in a tank (e.g. when it comes to water temperature, pH level and oxygen content). When tilapia are stocked at high density and optimal water conditions are present, even small parcels of land can yield large harvests. In addition, it is also easier to avoid negative effects from birds, snake and snails compared to a pond. Birds and snails can harbour parasites capable of infesting fish. Feeding and harvesting is typically less labour-intensive in tanks than in ponds. In ponds where both male and female tilapia are kept, uncontrolled breeding can easily turn into a problem. When tilapias are stocked in high density in tanks, the natural breeding behaviour is disrupted. The adult fish can put more energy into growing when they do not have to compete for food with their offspring. However, there are some disadvantages of indoor tilapia culture when compared with pond culture. Pond-grown tilapia can make use of naturally occurring food, while tankgrown tilapia have no access to a natural food source and have to be fed to a much higher

Fig. 11.4 Culture of tilapia using composite tanks at the National Broodstock Centre – Vietnam.

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Fig. 11.5 Raising Tilapia in cement tanks in Vietnam.

extent. The food for tank-grown tilapia has to be complete, i.e. it must contain the proper amounts of all necessary vitamins and minerals. When tilapia are grown in tanks, most farmers select a high stocking density. A high stocking density creates a stressful environment for the fish and therefore the risk of ill health is high. The risks will be increased further if the farmer fails to provide the fish with optimal water conditions and a satisfactory diet. Recirculation systems also require a substantial investment and must be properly managed.

11.4

HARVESTING AND VALUE ADDED PRODUCTS

Harvesting methods depend on pond size, culture systems and levels of technology applied. Complete harvests are necessary in ponds and are accomplished by seining in combination with draining. A complete harvest is not possible by seining alone as tilapias are adept at escaping seine nets. The pond should be dried between production cycles or treated with pesticides to kill tilapia fry to avoid carryover to the next production cycle (FAO 2010). Small-scale tilapia farmers in many parts of Africa and Asia cannot easily get harvesting nets and other equipment required for complete harvesting of their ponds. To overcome this problem, many farmers adopt partial harvesting techniques, using locally available gear. Partial harvesting is usually designed to remove large fish and provide smaller fish with more space for growth (Hepher & Pruginin 1982). Partial harvests of tanks, raceways and recirculation systems, which maximise production, are accomplished with grader bars to remove the largest fish (FAO 2010). Large-scale tilapia producers adopt more advanced harvesting tools, such as winches, because they generally prefer batch harvesting.

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Tilapia is generally marketed in rural areas and local markets in developing countries, either fresh or iced, with little handling and processing. However, the global tilapia market is growing sharply, not only in producing regions (mainly Southeast Asia, the Far East and Africa), but also in many non-producing regions, such as the USA, Europe and Australia. More attention should, therefore, be paid to tilapia handling and processing in order to maintain quality and meet the quality standards required by external markets. Poor handling and holding conditions, inadequate processing and the use of inappropriate processing methods can seriously affect the quality of fish and increase post-harvest losses. Live tilapia has become an important product in the market in many parts of the world where consumers prefer to buy live rather than iced or frozen fish (Singh & Daud 2001). It is becoming common to find live tilapia in display tanks and aquaria in seafood restaurants and supermarkets in countries like Singapore, Thailand and Malaysia (Singh & Daud 2001). However, live food systems require effective support systems, including live holding containers, specially equipped trucks, live holding centres and other infrastructure components. Consumers prefer to pay higher prices in order to get live fish. In Malaysia, for example, the price of live red tilapia is 37–40% higher than that of chilled fish (Singh & Daud 2001). According to El-Sayed (2006), the system involves transferring the fish into tanks at a temperature of 21 °C for about 10 hours to reduce stress. The water in the tanks is gradually cooled to 11 °C, with the addition of ice, to reduce fish activity. The fish are then packed in plastic bags with water and oxygen at a fish : water ratio of 1:1. The bags are tied and placed in polystyrene boxes, which are sealed and transferred to refrigerated trucks maintained at 15 °C and transported to their final destination. The need for proper handling and processing of tilapia is important both for the fishing industry and for the consumers. Improvement of the processing and handling of tilapia in terms of quality, product range and volume results in increased economic activity and employment. It is also one way of stabilising fish marketing by providing an outlet for surplus and peak catch even during emergency harvest, thereby ensuring high fishing activities and stable prices. It can also contribute to the efforts related to nutritional goals (El-Sayed 2006).

11.5

GENETIC IMPROVEMENT OF TILAPIA

Genetic improvement programmes have contributed significantly to increasing the productivity of cultured aquatic species (Dey & Gupta 2000; Gjedrem 2000; Lymbery et al. 2000). Breeding programmes for a number of species have been carried out, such as for carps, shrimps, oysters and other marine species. However, Nile tilapia has recently been investigated as a high-potential species in aquaculture in both tropical and subtropical regions. A number of selection experiments and testing programmes that aimed to increase the growth rate of tilapia culture in ponds have been conducted for O. niloticus (Hulata et al. 1986; Eknath et al. 1993; Brzeski & Doyle 1995; Bentsen et al. 1998; Bolivar &Newkirk 2002; Ponzoni et al. 2005; Rutten et al. 2005; Charo-Karisa et al. 2006; Luan et al. 2008; Rezk et al. 2009). These selective breeding programmes have typically been done in favourable environments where growth is expected to be high, and the results indicate an additive genetic variance that can be exploited through such programmes. Moreover, no evidence of genotype environment interaction (G x E) has been reported for harvest body weight traits in different freshwater environments, except that a minor interaction of little practical importance was found for harvest weight by Bentsen et al. (1998). However, recently a

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G x E was clearly found for harvest body weight and survival of O. niloticus in fresh and brackish water environments (Luan et al. 2008). Genetic parameters of salinity tolerance or selective breeding in brackish and marine environments have not been documented for O. niloticus, except those reported by Luan et al. (2008). Most research has been carried out to evaluate growth rates and to define suitable culture species in different salinity levels (Kamal & Mair 2005), or to develop a strain suitable for saline water (Rosario et al. 2004; Tayamen et al. 2004). Furthermore, most studies consider hybrid red tilapia and its potential for marine farming. In fact, the less saline tolerant O. niloticus appear to grow as well as hybrid red tilapia in fresh water at moderate salinities (Suresh & Lin 1992). Hence, testing and genetic improvement of growth performance and survival of O. niloticus in brackish water environment may be possible. The genetic mechanism and control of cold tolerance is still poorly understood and little is known about the differences in cold tolerance within and between tilapia species and strains. Testing of cold tolerance for O. niloticus strains and hybrids have been conducted, but these have been mainly limited to evaluating the magnitude of genetic parameters and potential for selection programme (Behrends et al. 1996; Charo-Karisa et al. 2005). Some studies however, have reported low heritabilities for this trait (<0.10) (Behrends et al. 1996; Cnaani et al. 2000; Charo-Karisa et al. 2005). Hence, both selective breeding and improved husbandry practices maybe used to improve cold tolerance of O. niloticus.

11.6

ENVIRONMENT AND DISEASE MANAGEMENT

Environment and disease are the most important aspects affecting yield of aquaculture systems, not only with tilapia species but also with other aquatic species. For instance, poor water quality parameters such as high organic matter, high ammonia, low dissolved oxygen and high bacterial load can create a sub-optimal environment that can be stressful for the fish and lead to a higher incidence of parasitic outbreaks. Each culture system has its own characteristics. For instance, tanks or cages, which hold high densities of fish, are good environments for the transmission of ectoparasites with a direct life cycle such as monogenean trematodes. Earthen ponds provide a more complex environment with vegetation where parasites such as crustacean copepods or leeches can lay eggs. The mud can be a reservoir for cysts of dinoflagellates such as Amyloodinium or invertebrates acting as intermediate hosts such as snails for digenean trematodes. The bigger the pond, the more difficult it is to control the parasite population as it is more open to fish predators which can seed eggs and other parasite stages. Recirculation systems have their own set of problems relative to parasitism: due to the build-up of sediment and slow turnover of water, recirculation systems also favour the growth and concentration of parasites. Therefore, particular vigilance is necessary when introducing fish or fish eggs in these systems. Komar and Wendover (2007) have comprehensively listed some of the common pathogens, disease symptoms, epidemiology and the possible treatments encountered in tilapia culture systems (Table 11.2).

11.7

MARKETING OF TILAPIA

Tilapia markets have shown two patterns of development. In the developing, mostly tropical countries, tilapia was first grown as a low cost, alternative protein source by and for

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Table 11.2 Common diseases of Tilapia (Komar & Wendover 2007, with permission from Aquaculture Health).

Trichodina spp.

Disease signs: Erratic swimming; Opened operculum; Scraping against walls; Jumping out of water; Erosion of fins, skin ulcers; Gill hyperplasia Epidemiology /risk factors: Massive mortality in hatchery and nursery phases Possible treatments: Salt bath; formalin bath; H2O2 bath and KMnO4 bath Disease signs: Appearance of white spots on skin; Thick mucus on skin; Stunted growth and mortality Epidemiology /risk factors: Most severe in larval stages. Problem in recirculation system Possible treatments: Repeated formalin bath and increased salinity

Ichthyophthirius multifilis Disease signs: Decreased appetite; Flashing; Accumulation of mucus Epidemiology /risk factors: Brackish water 10–15 ppt. Does not occur in fresh water Possible treatments: H2O2 bath

Amyloodinium spp. Disease signs: Grubs (yellow or white) on the skin. Skin haemorrhage and death if mass penetration of the parasite Epidemiology /risk factors: Occur in pond farming when snails and birds are present Possible treatments: Remove or eradicate snails from the pond Prevent bird access to the farm

Clinostomum spp. Disease signs: Skin darkening; Fin erosion; Excessive mucus; Rapid movement of operculum; Emaciation in young fish Epidemiology /risk factors: Juvenile and fingerling stages Possible treatments: Formalin bath; H2O2 bath

Dactyolgyrus spp. Disease signs: Skin irritation; Loss of condition; Associated secondary skin bacterial infection Epidemiology /risk factors: Severe in larval stages and fingerlings Possible treatments: Organophosphates

Argulus sp.

(Continued)

Tilapia 329 Table 11.2 (Continued ) Disease signs: Rub against sides of container Whitish spots of curled up worms embedded in the skin Epidemiology /risk factors: Can affect mouth breeding Possible treatments: Organophosphates

Lernea spp. Disease signs: High number of leeches on an adult fish induce anemia Epidemiology /risk factors: Severe in early stages Affect fish already weakened by another disease Possible treatments: Organophosphates

Leeches

rural farmers. Over time, urban markets have developed, but still as a relatively low priced fish product (Engle 1997; Funez et al. 2001). As production procedures have improved, restaurants and middle class consumers have started to purchase tilapia and it has moved into the mainstream of seafood products. In the highly industrialised countries, small markets developed in the 1970s and 1980s amongst restaurants and grocery shops of immigrant communities (Fitzsimmons 2001). These markets have been supplied with live fish by local farms and with low cost frozen fish from their home countries. Fresh fillets from tropical producers began to work their way into these ethnic restaurants and then migrated into upper echelon restaurants that were looking for new products and appreciated the qualities of fresh tilapia. From there, tilapia simultaneously moved into the casual dining restaurant chains and the hypermarkets and club stores. These are the major markets that are feeding the explosive growth in consumption. Virtually all the casual dining chains in the United States now feature one or more tilapia dishes on their menus. The reasonable cost, year-round supply and flexibility in preparation are ideal for their needs. Europe is the next ‘hot spot’ for tilapia markets. Although Europe’s immigrant communities have been fuelling growing imports for several years (FAO 2004b), most Europeans are just discovering tilapia. The major supermarket chains are testmarketing the fish. Tilapia production surged around the world during the 1990s and early 2000s. Tilapias are now the second most popular farmed fishes after the carps, with global production well over 1,526,000 tons in 2003. This has been accomplished by developing large consumer markets in North America and Europe, which are purchasing fish grown in tropical countries with year-round production and relatively low costs. These markets have evolved in a similar manner, starting with immigrant communities and moving into high cost, white tablecloth restaurants. With this rapid development in markets have come a variety of new product forms and commercial by-products. This has been a critical factor in the continued rise of tilapia in the ranks of popular seafoods and is a result of thoughtful marketing and product development, as well as increasing demands from retailers and consumers.

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11.8

CONCLUSION

Tilapia has become one of the most important farm-raised fish and is playing an increasing role in the international food trade. Due to the large number of rearing systems and microeconomic variables, it is difficult to generalise which types of systems will be profitable for specific countries. Based on current production trends, a variety of systems will likely be effective, from extensive in the tropics to highly intensive in temperate zones, where proximity to markets and maximal growth will offset higher production costs. However, in general, access to freshwater supplies will become one of the most pressing environmental and economic issues in both the developed and industrialised world (Hankins 2000). Hence, in both tropical and temperate zones future development of tilapia aquaculture will depend on the ability of production systems to produce more fish with less water, less food, and less time to lower production costs and reduce pollutants to the environment: a process known as intensification. Complete diets, aeration, water reuse and disease control will be important factors in this regard. To improve production efficiency, a number of challenges and research issues are of greatest concern to tilapia culturists in the world. These include a loss of species/strains, improved growth, fillet yield, environmental tolerance, disease resistance, off-flavours, waste management, and marketing. The most significant methods for improving production efficiency are through faster growth and improved feed utilisation, because these reduce time to market, increase system throughput, and reduce waste, all of which will lower the cost of production. The development of genetically improved production stocks can be achieved through selective breeding approaches, such as individual or family selection (Gjedrem 1999). The GIFT (Genetic Improvement of Farmed Tilapia) strain in Southeast Asia has demonstrated that growth rates of Tilapia can be significantly improved through a breeding program involving selection of diverse genetic groups of O. niloticus (Eknath & Acosta 1999). Chromosome manipulation techniques in tilapia have shown promise for improving growth rates and profitability. Genetically male tilapia produced from novel YY males (a technology that involves hormonally induced sex reversal), has shown potential for improved growth and profitability. The development of highly inbred lines and hybrids with superior growth can also be accelerated through gynogenesis. Finally, biotechnological approaches such as insertion of new genes into the Tilapia genome have high potential (Hallerman 2000). Transgenic Tilapia containing an exogenous growth hormone (GH) gene construct have been shown to exhibit dramatic growth improvement (Cabezas et al. 1997). At present, it is unclear if such transgenes can be transmitted over many generations and what additional effects they may have on the physiology of the animal. The genetic improvement programs will need to develop production stocks that are suited for each environment. Farms undertaking selective breeding programs must make significant investments for facilities to maintain broodstock, labour, feed, data handling, estimation of breeding values, and selection of broodstock. Therefore, success in genetic improvement programs will require longterm support and collaboration between partners from industry, university, and government (Hallerman 2000). As aquaculture production continues to intensify in the future, the use of intensive recirculating systems will expand. Meanwhile, the intensive indoor tilapia culture is constrained by high capital and operating costs, moderate growth rates, poorly utilised production capacity and consequent low system throughput. Therefore, advances in filter design and construction hold promise for reduced capital and operating costs for tilapia industry.

Tilapia 331

Tilapia products will continue to increase in availability and variety. Demand is certain to increase as recognition of its attributes spreads and as supply is provided to more markets. Production costs will continue to drop as farmers switch to faster-growing, more efficient stocks and as nutritional knowledge improves and producers learn to reduce feed costs. Huge markets are still available for expansion and the sophistication of the marketing firms has improved markedly. Tilapia has been referred to as the ‘aquatic chicken’. Presently, the status of tilapia is still in the earliest stages of the industry when compared to what has been accomplished by the poultry producers. It is a great time to be in the tilapia world.

11.9

REFERENCES

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Engle, C. (1997) Marketing tilapias. In: Tilapia Aquaculture in the Americas (eds B.A. Costa-Pierce & J.E. Rakocy). Baton Rouge, Louisiana: World Aquaculture Society. FAO (2004a) State of the world fisheries and aquaculture. Food and Agricultural Organisation, Rome. FAO (2004b) Tilapias as alien aquatics in Asia and the Pacific: A review. FAO Fisheries Technical Papers – T453. ISBN: 9251052271. FAO (2010) Cultured aquaculture species – Nile Tilapia. Accessed 2010. Cited from: http:// www.thefishsite.com/articles/911/cultured-aquaculture-species-nile-tilapia Feryer, G. & Iles, T.D. (1972) The Cichlid Fishes of the Great Lakes of Africa, their Biology and Distribution. Oliver & Boyd, Edinburgh. Fitzsimmons, K. (2000) Tilapia: The most important aquaculture species of the 21st century In: Tilapia Aquaculture in the 21st Century (eds K. Fitzsimmons & F.J. Carvalho), Proceedings of the Fifth International Symposium on Tilapia Aquaculture, Vol. 1. Ministry of Agriculture, Rio de Janeiro, Brazil. Fitzsimmons, K. (2001) China and international tilapia markets. In: Aquatic Products Processing and Trading Symposium: China, Japan, Korea (ed. D. Gang). Qingdao, China, China Aquatic Products Processing and Marketing Association. Funez, N.O., Neira, I. & Engle, C. (2001) Supermarket outlets for tilapia in Honduras: an overview of survey results. In: 6to. Simposio Centroamericano de Acuacultura Proceedings: Tilapia sessions, 22–24 August 2001 (ed. D. Meyer), pp. 82–86. Tegucigalpa, Honduras. Gjedrem, T. (1999) Aquaculture needs genetically improved animals. Global Aquaculture Advocate, 2, 69–70. Gjedrem, T. (2000) Genetic improvement of cold–water fish species. Aquaculture Research, 31, 25–33. Green, B.W. (2006) Fingerling production systems. In: Tilapia: Biology, Culture, and Nutrition (eds C.E. Lim & C.D. Webster). London: The Haworth Press. Green, B.W., Verrica, K.L. & Fitzpatrick, M.S. (1997) Fry and fingerling production. In: Dynamics of Pond Aquaculture (eds H.S. Egna & C.E. Boyd). CRC Press, Boca Raton, Florida. Gunasekera, R.M., Shim, K.F. & Lam, T.J. (1996). Influence of protein content of broodstock diets on larval quality and performance in Nile tilapia, Oreochromis niloticus (L.). Aquaculture, 146, 245–259. Hallerman, E.M. (2000) Genetic improvement of fishes for commercial recirculating aquaculture system: a case study involving tilapia. In: Proceedings of the International Conference on Recirculating Aquaculture Systems, 2000 Blacksburg, Virginia. Virginia Polytechnic and State University. Hankins, J.A. (2000) Perspective on the role of government, industry, and research in advancing the environmental compatibility and sustainability of aquaculture. In: Proceedings of the International Conference on Recirculating Aquaculture, 2000 Blacksburg, Virginia Virginia Polytechnic and State University. Hepher, B. & Pruginin, Y. (1982) Tilapia culture in ponds under controlled conditions. In: Pullin, R.S.V & Lowe-McConnell, R.H. (eds.), The Biology and Culture of Tilapias: proceedings of the International Conference on the Biology and Culture of Tilapias, 2–5 September 1980, Bellagio, Italy. pp. 185–203. Hulata, G., Wohlfarth, G.W. & Halevy, A. (1986) Mass selection for growth rate in the Nile tilapia (Oreochromis niloticus). Aquaculture, 57, 177–184. Jauncey, K. & Ross, B. (1982) A guide to Tilapia feeds and feeding. University of Stirling, Scotland. Kamal, A.H. M.M. & Mair, G.C. (2005) Salinity tolerance in superior genotypes of tilapia, Oreochromis niloticus, Oreochromis mossambicus and their hybrids. Aquaculture, 247, 189–201. Komar, C. & Wendover, N. 2007. Illustration of major parasites of Tilapia, associated clinical signs, epidemiology and possible treatments. http://www.thefishsite.com/articles/294/ parasitic-diseases-of-tilapia Lim, C. & Webster, C.D. (eds) (2006). Tilapia: Biology, Culture and Nutrition. The Haworth Press, New York. Little, D.C. (2004) Delivering better quality tilapia seed to farmers. In: New Dimensions in Farmed Tilapia (eds R. Bolivar, G. Mair & K. Fitzsimmons). Proceedings of the 6th International Symposium on Tilapia in Aquaculture, Manila, Philippines. Little, D.C., Bhujel, R.C. & Pham, T.A. (2003) Advanced nursing of mixed-sex and mono-sex tilapia (Oreochromis niloticus) fry, and its impact on subsequent growth in fertilized ponds. Aquaculture, 221, 265–276. Luan, T.D., Olesen, I., Ødegård, J., Kolstad, K. & Dan, N.C. (2008) Genotype by environment interaction for harvest body weight and survival of Nile tilapia (Oreochromis niloticus) in brackish and fresh water ponds. In: Eighth International Symposium on Tilapia in Aquaculture (eds H. Elghobashy, K. Fitzsimmons & A.S. Diab). Vol. 1, pp. 231–240. Cairo, 12–14 August.

Tilapia 333 Lymbery, A.J., Doupe, R. G., Jenkins, G. & Thorne, T. (2000) Genetic improvement in the aquaculture industry. Aquaculture Research, 31, 145–149. Macaranas, J.M., Mather, P.B., Lal, S.N., Vereivalu, T., Lagibalavu, M. & Capra, M.F. (1997) Genotype and environment: A comparative evaluation of four tilapia stocks in Fiji. Aquaculture, 150, 11–24. Macintosh, D.J. & Little, D.C. (1995) Nile tilapia (Oreochromis niloticus). In: Brood Stock Management and Egg and Larval Quality (eds N.R. Bromage & R.J. Roberts). Oxford, Blackwell Science. Osure, G.O. & Phelps, R.P. (2006). Evaluation of reproductive performance and early growth of four strains of Nile tilapia (Oreochromis niloticus, L.) with different histories of domestication. Aquaculture, 253, 485–494. Ponzoni, R.W., Hamzah, A., Tan, S. & Kamaruzzaman, N. (2005) Genetic parameters and response to selection for live weight in the GIFT strain of Nile Tilapia (Oreochromis niloticus). Aquaculture, 247, 203–210. Rezk, M.A., Ponzoni, R.W., Khaw, H.L., Kamel, E.A., Dawood, T. & John, G. (2009) Selective breeding for increased body weight in a synthetic breed of Egyptian Nile tilapia, Oreochromis niloticus: Response to selection and genetic parameters. Aquaculture, 293, 187–194. Rosario, W.R., Georget, C., Chevassus-Au-Louis, B., et al. (2004) Selection from an interspecific hybrid population of two strains of fast growing and salinity tolerant tilapia In: New Dimensions in Farmed Tilapia (eds R. Bolivar, G. Mair & K. Fitzsimmons). Proceedings of the 6th International Symposium on Tilapia in Aquaculture, Manila, Philippines. Rutten, M.J.M., Bovenhuis, H. & Komen, H. (2005) Genetic parameters for fillet traits and body measurements in Nile tilapia (Oreochromis niloticus L.). Aquaculture, 246, 125–132. Sifa, L., Chenhong, L., Dey, M. & Dunham, R. (1999) Seinability of four strains of Nile tilapia, Oreochromis niloticus, in Chinese ponds. Aquaculture, 174, 223–227. Singh, T. & Daud, W.J.W. (2001) Live handling and marketing of tilapia. In: Tilapia: production, marketing and technological developments (eds S. Subasinghe & T. Singh), pp. 88–93. Proceedings of the Tilapia 2001 International Technical and Trade Conference on Tilapia, 28–30 May 2001, Kuala Lumpur, Malaysia. Suresh, A.V. & Lin, C.K. (1992) Tilapia culture in saline waters: a review. Aquaculture, 106, 201–226. Tayamen, M.M. (2004) Nationwide dissemination of GET EXCEL tilapia in the Philippines. In: New Dimensions in Farmed Tilapia (eds R. Bolivar, G. Mair & K. Fitzsimmons). Proceedings of the 6th International Symposium on Tilapia in Aquaculture, Manila, Philippines. Tayamen, M.M., Abella, T.A., Reyes, R.A., et al. (2004) Development of Tilapia for saline water in the Philippines. In: New Dimensions in Farmed Tilapia (eds R. Bolivar, G. Mair & K. Fitzsimmons), pp. 463–478. Proceedings of the 6th International Symposium on Tilapia in Aquaculture, Manila, Philippines. Watanabe, W.O., Losordo, T.M., Fitzsimmons, K. & Hanley, F. (2002) Tilapia production systems in Americas: Technological advances, trends, and challenges. Reviews in Fisheries Science, 10, 465–498. Zimmermann, S. & Natividad, J. (2004) Comparative pond performance evaluation of GenoMar ASA and GenoMar Supreme tilapiaTM GST1 and GST3 groups. In: New Dimensions in Farmed Tilapia (eds R. Bolivar, G. Mair & K. Fitzsimmons). Proceedings of the 6th International Symposium on Tilapia in Aquaculture, Manila, Philippines.

12

Carp Polyculture in India

Dilip Kumar

12.1 INTRODUCTION India is the second largest contributor to global aquaculture and in 2007 it produced about 3.355 million tons of fish and shellfish (FAO 2008). Recent data from India indicate that aquaculture production rose to 3.59 million tons in 2008–2009. This included production from mariculture, coastal aquaculture and freshwater aquaculture and contributed about 47% to the country’s total fish production of 7.61 million tons (Fig. 12.1). Aquaculture in India is highly dominated by freshwater aquaculture (3.43 million tons) followed by coastal aquaculture (0.15 million tons) and mariculture (0.01 million tons), which is just emerging. Coastal aquaculture mainly concerns shrimp and contributes just over 4.4% to the total aquaculture production, compared with the 95% contributed by freshwater aquaculture. Freshwater aquaculture is almost synonymous with carp culture in India as well as its neighbouring countries. Carp culture mainly involves members of the Order Cypriniformes and Family Cyprinidae (Khanna, 1988, Talwar & Jhingran 1991) belonging to two main groups, i.e. the three Indian major carps such as catla (Catla catla), rohu (Labeo rohita) and mrigal (Cirrhinus mrigala), and three domesticated exotic carps such as silver carp (Hypophthalmichthys molitrix), grass carp (Ctenopharyngodon idella) and common carp (Cyprinus carpio). Other species such as giant freshwater prawn (Macrobrachium rosenbergii) and native catfish (Clarias batrachus) are also becoming popular. Clandestine introduction of certain exotic fish species such as African catfish (Clarias gariepinus) and the Thai catfish (Pangasius sutchi) has led to these being cultured in certain pockets. Carp culture in India was restricted to a homestead backyard pond culture activity in the eastern Indian states of West Bengal, Orissa and Assam until the late 1950s. Seed of Indian major carps collected from riverine sources were used as the only input for pond fish culture. After stocking they were allowed to grow on natural fish food organisms in rain-fed, non-drainable ponds, resulting in very low levels of yield. Though the importance of fish culture as an economically promising enterprise was subsequently realised, the nonavailability of quality fish seed and lack of knowledge about scientific fish culture constrained the development of carp farming until the 1960s. This chapter reviews the development of composite fish culture in India.

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

Carp Polyculture in India 8 7 6 5 4 3 2 1 0

Marine

96-97

97-98

Freshwater

335

Total (million tons)

99-00 2000-01 2001-02 2002-03 2003-04 2004-05 2005-06 2006-07 2007-08 2008-09*

Fig. 12.1 Trends in fish production of India (adapted from the Department of Animal Husbandry, Dairying & Fisheries Annual Report, 2008–09, Ministry of Agriculture, Govt. of India, New Delhi). * signifies 2009 data is not for the full year.

12.2

FRESHWATER AQUACULTURE RESOURCES IN INDIA

India has vast and varied freshwater resources for aquaculture in the form of ponds, tanks, lakes, reservoirs, rivers, irrigation canals, floodplains, derelict water bodies, etc. Ponds and tanks alone account for about 2.41 million hectares; floodplain, lakes and derelict water bodies comprise about 0.8 million ha; reservoirs spread over an area of 2.907 million ha; and rivers and canals have a total length of just under 0.2 million km. There is also about 1.24 million hectares suitable for brackish water aquaculture. Owing to these resources India is considered to be one of the richest countries in terms of aquaculture resources. Pond and tanks are the most common freshwater resources used for aquaculture. Ponds are relatively smaller while tanks are bigger in size and excavated for harvesting/holding rainwater with the objective of providing water for irrigation. In general these ponds are simple excavated earthen structures, which are non-drainable and mostly rain-fed. In some areas, being closer to irrigation / drainage canals, these ponds and tanks also receive canal water. As a result there is wide fluctuation of water levels throughout the year, with the maximum during the rainy season (July–August) and the minimum during summer (May–June). Ox-bow lakes and floodplain areas are also potential resources for freshwater aquaculture. These water bodies are rain-fed, flood-fed or both. In general ox-bow lakes are perennial and used extensively for culture-based fisheries. These are usually common property resources and in most cases used for fish culture by groups of local fishing communities. Floodplains in the forms of chaurs, mauns and haors (vast expanses of water) are usually created during the rainy season on individually owned agricultural lands. Chaurs are common in many parts of India while haurs are common in Assam and Bangladesh. During the rainy season vast areas of low-lying agricultural land are flooded and the water remains for several months depending upon the contour, landscape and rainfall. While these areas dry up completely during summer, the deeper areas of such floodplains retain water throughout the year. Such residual water bodies are very important from an aquatic biodiversity point of view, as these are refuge for the broodstock of many indigenous smaller fish species. Floodplains are used for short-term culture of carp. However, so far the potential of floodplains for aquaculture has been tapped only partially. It requires extensive social mobilisation initiatives to develop a workable model where the resource-owning communities work collectively to get an additional crop of fish from submerged areas in addition to their usual individually cultivated crop. Reservoirs also offer vast potential, which is yet to be harnessed.

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12.3

DEVELOPMENT OF AQUACULTURE

Freshwater aquaculture in India started with the use of indigenous traditional knowledge. Because floods are a regular phenomenon, the local communities used to construct houses on raised platforms for which earth was collected by digging adjacent lands. As a result pits of varying depths were created close to each dwelling. During the rainy / flood season these pits were filled with water that lasted for several months. Initially these pits were also used for trapping fish fingerlings during floods. After the floodwaters receded, the communities would catch fish for their own consumption and sometimes also leave them to grow and catch for important occasions such as festivals and during visits by friends and relatives. Subsequently such pits or ponds were also stocked with seeds caught from natural sources. The farmers knew about the spawning behaviour of indigenous carp, which helped in catching seeds from nearby rivers. This practice led to the development of indigenous rearing of Indian major carps and other native species. The technological breakthrough of induced breeding of carps through hypophysation in 1957 revolutionised freshwater aquaculture in the country (Chaudhuri & Alikunhi 1957). With an assured supply of quality seed (spawn) through induced breeding technology and the development of nursery and rearing practices, appropriate stocking material (fingerlings) were made available for aquaculture. Subsequently, a package of practices for the culture of compatible species of carps, three Indian major carps (C. catla, L. rohita and C. mrigala) and three Chinese carps (H. molitrix, C. idella and C. carpio), popularly known as composite fish culture, was developed by the Pond Culture Division of the Central Inland Fisheries Research Institute (CIFRI) (Chaudhuri et al. 1976). The technology of composite fish culture defined various packages of practices to be followed for culture of major carps in earthen non-drainable ponds. This included pre-stocking pond preparation including liming and manuring, stocking and post-stocking practices. Its further refinement through multi-location trials under the All India Co-ordinated Research Project (AICR) on Composite Fish Culture Projects (Sinha 1971) revolutionised pond aquaculture in India. The All India Co-ordinated Research Projects on Composite Fish Culture and Fish Seed Production launched by the Indian Council of Agricultural Research (ICAR), through the CIFRI in Barrackpore, operating at 12 centres all over the country till 1984, virtually laid the foundation of scientific carp farming. Carp culture since then has expanded in terms of area coverage and intensity of operation in states like Andhra Pradesh, Assam, Tripura, Punjab, Haryana, Uttarakhand, Uttar Pradesh, Bihar, Jharkhand, Madhya Pradesh, Chhattisgarh, Manipur, Maharashtra, etc. In many states this has become a commercial farming enterprise. It is also interesting to note that carp culture in India and neighbouring countries is practised exclusively in un-drainable earthen ponds which are largely rain-fed. A detailed account of fish culture practice in non-drainable ponds is given in Kumar 1992.

12.4

COMMONLY CULTURED SPECIES

Carp is still the dominant fish species in aquaculture in India, contributing about 95.62% of the total aquaculture production followed by brackish-water shrimp (4.36%) and coldwater fishes (0.01%). Among carp, Indian major carps contribute more than 85%, exotic carps 13% and the remainder is accounted for by other minor species. Species-wise, fish production in different states of India is presented in Table 12.1.

24,100 0 580 35,000 1,587 70 10 0 22,335 956 0 11,600 6,869 0 0

76,670 578,670

1,300

64,393 149,177 450 121 18,273 53,650 82

75,800 997

124,195 63,570 68,466

Assam Andhra Pradesh Arunachal Pradesh Bihar Chhatisgarh Delhi Goa Gujarat Haryana Himachal Pradesh Jharkhand Jammu & Kashmir Karnataka Kerala Madhya Pradesh

Exotic carps

Indian major carps

131,064 63570 68,466

75,800 12,597

99,393 150,764 520 131 18,273 75,985 1,038

1,880

100,770 578,670

Total carp

6,869.133 0 0

0 6,504.85

9,000 7,934.5 188 0 9,570 300 276.7

580

103,743 287,243

Catfish and other minor fishes

Inland fish production (tons) in different states of India, by species.

State

Table 12.1

5,784 16,000 0

0 0

0 0 0 289 0 0 0

0

0 26,341

Brackish water fishes

0 0 0

0 168

0 0 0 0 0 0 64

11

0 0

Coldwater fishes

143,717.1 79,570 68,466

75,800 19,269.85

108,393 158,698.5 708 420 2,7843 7,6285 1,378.7

2,471

204,513 892,254

Total in tons

(Continued)

3.48 1.92 1.66

1.83 0.47

2.62 3.84 0.02 0.01 0.67 1.84 0.03

0.06

4.95 21.58

% age to total

Carp Polyculture in India 337

NA NA 1,076 2,996 18,886 37,687 175 340 66 211 3,699.06 172,541 52,000 NA 405,694.1 12.8

NA 14,000 1,521 2,645 139,180 30,543 1,000 20,000 0 80,829 27,743 832,756 244,000

NA 2,764,741 87.2

Mizoram Manipur Meghalaya Nagaland Orissa Punjab Puducherry Rajasthan Sikkim Tamil Nadu Tripura West Bengal Uttar Pradesh Uttarakhand India % of total NA 3,170,435 76.68

NA 14,000 2,597 5,641 158,066 68,230 1,175 20,340 66 81,040 31,442.06 1,005,297 296,000

107,619.5

Total carp

NA 788,371 19.07

NA 4,800 1,385 184 0 0 230 3,760 0 67,618.71 5,548.62 201,680 49,400

21,555.5

Catfish and other minor fishes

NA 175,430 4.24

NA 0 0 0 22,969 0 800 0 0 15,846 0 87,401 0

0

Brackish water fishes

NA = no data available Source: Based on information on hatcheries supplied to CIFE by the Directorate of Fisheries of the various states

12,910

94,709.5

Maharashtra

Exotic carps

Indian major carps

State

Table 12.1 (Continued )

NA 580 0.01

NA 0 0 0 0 0 0 0 5 0 0 332 0

0

Coldwater fishes

NA 4,134,816 100

129,175 NA 18,800 3,982 5,825 181,035 68,230 2,205 24,100 71 164,504.7 36,990.68 1,294,710 345,400

Total in tons

NA 100.00

3.12 NA 0.45 0.10 0.14 4.38 1.65 0.05 0.58 0.00 3.98 0.89 31.31 8.35

% age to total

338 Recent Advances and New Species in Aquaculture

Carp Polyculture in India

339

The state of West Bengal is the largest producer as well as the biggest consumer of carp. The state contributes about 32% of total production, followed by Andhra Pradesh (15.6%), Uttar Pradesh (8.6%), Maharashtra (more than 8%) and others. Three species of Indian major carps (IMC), i.e. catla (Catla catla), rohu (Labeo rohita) and mrigal (Cirrhinus mrigala), followed by three exotic Chinese carps: silver carp (Hypophthalmichthys molitrix), grass carp (Ctenopharyngodon idella) and common carp (Cyprinus carpio), are the commonly cultured carps in India. In some eastern states Labeo calbasu is also produced along with other major carps, due to high demand for it in the local market. In recent years there has been growing interest in the diversification of freshwater aquaculture. Accordingly, a number of research programmes are being undertaken by research institutes, universities and even some of the state fisheries departments to diversify freshwater aquaculture as well as brackish water aquaculture, which depends mainly on one species of shrimp, Penaus monodon. Due to increasing demand and the high price of some of the indigenous fish species, efforts are being made to develop seed production and culture technologies of some species such as Anabas testudeneus, Labeo bata, Amblypharyngodon mola, Ompok pabda, etc. Culture of native catfish such as Clarias batrachus and Heteropneutes fossilis is also becoming popular. Efforts are being made to develop commercial seed production technology for these two species. Due to growing demands by farmers the government of India has recently allowed the culture of Thai catfish, Pangasius sutchi. Culture of certain clandestinely introduced species such as African catfish Clarias gariepinus is also being practised extensively in several states of India.

12.5

AQUACULTURE PRACTICES/SYSTEMS

Ongoing aquaculture practices in India include mainly extensive culture of major carps, polyculture of carps and freshwater prawn, culture of native catfish Clarias batrachus and exotic African catfish (Clarias gariepinus) and Thai catfish (Pangasius sutchi). However, carp polyculture is still the mainstay of freshwater aquaculture. Other practices, though considered to be emerging, are already reaching significant levels. Brackish-water aquaculture is still confined to the culture of tiger shrimp (Penaeus monodon). Recently Penaeus vannamei has also been introduced but its practice is still limited to carefully selected farms. Culture of giant sea bass (Lates calcarifer) is also emerging due to the development of induced breeding and seed rearing technology by CIBA (Central Institute of Brackishwater Aquaculture) and the Rajiv Gandhi Aquaculture Centre of the Marine Products Export Development Authority (MPEDA). Aquaculture activities are divided into two distinct segments: seed production and table-size production. Accordingly, Indian aquaculture will be described under these two headings.

12.5.1 Seed production Lack of local availability of quality fish seed in adequate quantity and with a year-round reliable supply is considered to be one of the major constraints limiting the speedy development of aquaculture in India as well as neighbouring countries. Seed for aquaculture in India comes from three sources – natural resources like rivers, bundhs and hatcheries. In the mid-1960s riverine seed collection accounted for over 90% of the total seed produced. However, during the 1980s the contribution of bundh breeding in West Bengal

340

Recent Advances and New Species in Aquaculture

Fig.12.2

Chinese circular carp hatchery.

and Madhya Pradesh rose to over 60% of the total seed produced in the country. Bundhs are special type of shallow earthen ponds having one side modified for receiving rainwater from the catchment area. Inflow of water in such ponds simulates riverine condition during themonsoon, resulting in sympathetic spawning of carps. Initially rivers were the mainstay for seed but were gradually replaced by bundhs and subsequently by hatcheries. With the development and growth of carp hatcheries bundh breeding has lost its relevance. Currently there are over 1,600 Chinese-type circular carp hatcheries (Fig. 12.2) in the country, which produced about 85,000 million seed (95% of the total carp spawn produced in the country) in 2008–09. 12.5.1.1 Carp hatcheries In 2010, hatcheries accounted for almost all of India’s spawn production. Fry production of about 211 million in 1964–65 rose to 31,688 million in 2007. This quantum jump in fish seed production is attributed mainly to the development of induced breeding technology and introduction of hapa-based hatching. Subsequently hapa were replaced by jar hatcheries with better-controlled facilities and more reliable results. By the early 1990s jar hatcheries were replaced by Chinese-style circular carp hatcheries. In the meantime there was further improvement in induced spawning technology, which replaced use of crude pituitary extract with combination of LHRH analogue and domperidone, resulting in more reliable results. Despite the upsurge in the number of carp hatcheries there is still a shortage of seed. However, the use of hatching hapa is still prevalent in some areas. Currently there are

Carp Polyculture in India

341

Table 12.2 Carp hatcheries in India. States

Assam Andhra Pradesh Arunachal Pradesh Bihar Chhatisgarh Delhi Goa Gujarat Haryana Himachal Pradesh Jharkhand Jammu & Kashmir Karnataka Kerala Madhya Pradesh Maharashtra Mizoram Manipur Meghalaya Nagaland Orissa Punjab Puducherry Rajasthan Sikkim Tamil Nadu Tripura West Bengal Uttar Pradesh Uttarakhand India (total)

No. of circular hatcheries 227 86 1

Spawn (million)

9,226 6,000 36

Hapabased hatcheries 400 0 0

4,415 0 0

22 3

400 40

0 0

0 0

540 18.2

50 20 53

400 20 1,895.7

21 0 0

60 0 0

206.8 0 523.52

28 0 3 2 2 107 81 1 7 1 28 7 573 198

1,897.57 0 597 0

53 0 19 0 10 0 92 0 0 7 0 6 0 0

356 0 0 0 0

183.17 0 179 0 48.5 782.27 52.8 0 43.8 1

NA 1,615

NA 1,174

9 0 0 0.5 0 0 0 NA 4,866

Fingerling production (million)

400 1,500 18

259 24,151 0 1.5 58.57 4,982 0

90 1 900 1,400 28,130 2,190.1 NA 85,758

3.8 0 0 0 0 NA 21.61

Fry (million)

37 50 0 3 10 15 0

2,992.5 90

553 0 0 0 0 10 3

Spawn (million)

0 5,137 0.4 1 47.02 332.2 3.61

0 14,000 1297.6 NA 24,965

60 0 0.54 0 809.49 0.1 0.5 21.21 NA 0 34 4.5 4.65 2.5 NA 14.76 0 125 0 0 142.45 21.1 0 0 0.2 800 286.96 8,400 6.54 NA 10,730

NA = no data available Source: Based on information on hatcheries supplied to CIFE by the Directorate of Fisheries of the various states

about 1,700 Chinese-type circular hatcheries, of which 573 are in the state of West Bengal is the largest. Details on the number of hatcheries and spawn production are presented in Table 12.2. 12.5.1.2 Seed production by state India produced only 409 million fry during the year 1973–74, which increased to 31,688 million fry during 2006–07. This production figure appears impressive but the distribution of fish seed production is largely confined to states like West Bengal, Andhra Pradesh and

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Table 12.3 Fish seed production in India by state (in millions). States

2004–05

2005–06

2006–07

% total

Andhra Pradesh Arunachal Pradesh Assam Bihar Chhatisgarh Goa Gujarat Haryana Himachal Pradesh Jammu & Kashmir Jharkhand Karnataka Kerala Madhya Pradesh Maharashtra Manipur Meghalaya Mizoram Nagaland Orissa Punjab Rajasthan Sikkim Tamil Nadu Tripura Uttar Pradesh Uttarakhand West Bengal Others India

421 26.1 2,741.47 318.31 457.78 0.89 508.93 249.57 19.17 16 80 175.22 12.17 334.6 47.45 118 0.9 16 44 604.79 133 255 2.5 529.28 401.86 1,037.32 35.82 12,200 3.81 20,790.64

851.9 27 3,207.99 344.94 505.54 0.69 611.59 282.08 21.27 17 17.27 215.72 12.48 438.62 182.14 123 0.74 18 45 554.14 139.03 299 2.5 529.28 546.53 1,085.75 31.69 12,500 3.76 22,614.72

9,912.7 30 2,062 330.46 591.68 0.67 621.89 331.14 16.98 17.3 16.21 195.43 12.7 485.87 549.73 129 0.77 18.2 48 546.72 153.38 342 2.5 529.28 420 1,091.56 28.21 13,200 3.63 31,688.01

31.28 0.09 6.51 1.04 1.87 0 1.96 1.05 0.05 0.05 0.05 0.62 0.04 1.53 1.73 0.41 0 0.06 0.15 1.73 0.48 1.08 0.01 1.67 1.33 3.44 0.09 41.66 0.01 100

Source: Based on information from the Central Institute of Fisheries Education (CIFE), Indian Council of Agricultural Research

Assam. They jointly contribute about 80% of total fish seed production in the country (Table 12.3). Among the different states, West Bengal is ranked first in fish seed production contributing about 42% of total fish seed production in the country. The state of Andhra Pradesh, which witnessed a revolution in carp culture and seed production during the 1980s, is placed second in fish seed production and contributes about 31% of national fish seed production (Govt. of India 2007–08). State-by-state data for seed production during the year 2008–09 and estimates reported by Directorate of Fisheries of different states are presented in Tables 12.4 and 12.5. Data thus collected indicates that except for West Bengal, Andhra Pradesh, Assam and Tripura all other states are deficit in fish seed production of carps and hence they resort to the import of fish seed from other states. In case of brackish water fish seed production, the states of Tamil Nadu and Andhra Pradesh have a surplus. All other states meet their requirements by importing post-larvae (PL) from these states. In the case of coldwater fisheries, none of the states has a surplus in fish seed production.

Assam Andhra Pradesh Arunachal Pradesh Bihar Chhatishgarh Delhi Goa Gujarat Haryana Himachal Pradesh Jharkhand Jammu & Kashmir Karnataka Kerala MP Maharashtra

States

Table 12.4

0.9 200 80.6 0.1 0.5 21.2 0 0 34 9.8 0 2.4 0 103.19

250 513.7 0.2 1 47 332.2 10

540 26

206.8 12.17 523.5 569.69

515.3 0

Fingerling

2.2

2,914.6 1,530

Fry

Production (million)

206.8 14.57 523.5 750.55

574 35.8

450 594.3 0.3 1.5 68.2 332.2 10

3.1

3,429.9 1530

Total

637.7 2,002.4

465

800 50

1,600 542.5 1.7 1 47 346 50

13

2,631.3 1,255

Fry

0 40 0 1,820

80 19

800 85.4 0.3 0.9 79.8 NA NA

0.4

600 0

Fingerling

465 40 637.7 3,822.5

880 69

2,400 627.9 2 1.9 126.8 346 50

13.4

3231.3 1,255

Total

Requirement (million)

Production and requirement of carp seed in India during 2008–09, by state.

−258.2 12.17 −1,14.2 −1,432.71

−260 −24

−1,350 −28.8 −1.5 0 0 −13.8 −40

−10.8

283.3 275

Fry

0 −37.6 0 −1,716.81

−46 −9.2

−600 −4.8 −0.2 −0.4 −58.6 NA NA

0.5

−84.7 0

Fingerling

Surplus/deficit

(Continued)

−258.2 −25.43 −114.2 −3,071.95

−306 −33.2

−1,950 −33.6 −1.7 −0.4 −58.6 −13.8 −40

−10.3

198.6 275

Total

Carp Polyculture in India 343

18.2 178.9 0 48.6 782.3 52.8 0 340.1 3.3 529 420 13,370 1,297.6 35.82 24,555.68

Fry

(Continued )

0 125 0.011 0 142.5 21.1 0 0 0.7 160 287 8,010 6.5 0 9,720.801

Fingerling

Production (million)

18.2 303.9 0.011 48.6 924.8 73.9 0 340.1 4 689 707 21,380 1,304.1 35.82 34,354.15

Total 0 196 NA 68.5 880 102.8 22 576.6 4 529 420 8,050 1470 NA 22,761.5

Fry 0 137 NA 0 440 21.1 8 0 1 252 143.3 4,775 30 0 9,333.2

Fingerling 0 333 0 68.5 1,320 123.9 30 576.6 5 781 563.3 1,2825 1,500 NA 32,094.8

Total

Requirement (million)

18.2 −17.1 NA −19.9 −97.7 −50 −22 −236.5 −0.7 0 0 5,320 −172.4 NA 1,758.36

Fry

0 −12 NA 0 −297.5 0 −8 0 −0.3 −92 143.7 3,235 −23.5 0 387.59

Fingerling

Surplus/deficit

18.2 −29.1 0.011 −19.9 −395.2 −50 −30 −236.5 −1 −92 143.7 8,555 −195.9 NA 2,223.531

Total

NA = no data available Note: Production figure for the states of Tamil Nadu, Tripura, Mizoram and Kerala are taken from the figure for the year 2006–07 published by the Department of Animal Husbandry, Dairying and Fisheries, Govt. of India Source: Adapted from information supplied by the Directorates of Fisheries of the various states to CIFE, Mumbai

Mizoram Manipur Meghalaya Nagaland Orissa Punjab Puducherry Rajasthan Sikkim Tamil Nadu Tripura West Bengal UP Uttarakhand India

States

Table 12.4

344 Recent Advances and New Species in Aquaculture

Carp Polyculture in India

345

Table 12.5 Production and requirement of coldwater fish seed in India 2008–09, by state. States

Arunachal Pradesh HP Jammu & Kashmir Sikkim West Bengal India

Production (million)

Requirement (million)

Surplus/ deficit (millions)

Fry

Fry

0 0.4 21.2

Fingerling 0.9 0 10

Total

0.9

2

0.4 31.2

5 35

Fingerling

Total

Fry

Fingerling

Total

1.2

3.2

-2

-0.3

-2.3

-4.6 -13.8

0 -10

-4.6 -23.8

0 20

5 55

0.5 0.05

0.1 0.025

0.6 0.075

0.5 0.5

0.1 0.25

0.6 0.75

0 -0.45

0 -0.225

0 -0.68

22.15

11.025

33.175

43

21.55

64.55

-20.85

-10.53

-31.38

Source: Based on information supplied by the Directorates of Fisheries of the different states to CIFE

12.5.1.3 Movement of fish seed across Indian states Due to the concentration of fish seed production centres in the country, fish seed are marketed across states and transported long distances from the state of West Bengal up to Tamil Nadu in the south, Jammu and Kashmir in the north and Manipur in the northeast. Information on the export and import of fish seed by different states during 2008–09, collected from the Directorate of Fisheries of the different states of India, are presented in Tables 12.6 and 12.7. These data indicate that fish seed are exported to neighbouring countries like Bhutan, Nepal and Bangladesh. It is interesting to note that fish seed is being exported not only by states that have a surplus of fish seed, but also by states that have a deficit, taking advantage of seasonal availability and the proximity of seed production centres to seed deficit areas in other states. However, all stages of fish seed: spawn, fry and fingerlings, are traded across states. West Bengal is the largest exporter of freshwater fish seed but it does import shrimp seed from the states of Tamil Nadu, Andhra Pradesh, Orissa and Kerala. 12.5.1.4 Seed quality considerations The rapid growth of aquaculture in Asia in general and India in particular has been mainly due to the availability of fish seed to farmers. Even though the seed of the major cultivated species is now produced in large quantities in hatcheries, poor quality is still perceived as a major constraint for the expansion of aquaculture in India. The availability of quality fish seed at the right time and at the right location is seen as a prerequisite for sustainable aquaculture development. Although carp, both indigenous and exotic, catfish and murrels (Ophicephalus spp.) make up the bulk of fish raised in the country, several other species like mahseer (Tor spp.), trout, several minor carps (Cirrhinus reba H., Labeo bata H., L. calbasu H., L. kontius Fedon, and C. cirohosa Bloch), etc., are also important. A number of agencies (both government and private) are involved in fish seed production and distribution. In addition, seed production by farmers themselves is now widely practised in India. Quality deterioration due to genetic inbreeding depression has recently been seen as a growing problem. In genetic terms, quality seed may be defined as those having better food conversion efficiency, high growth rate potential, better adaptability to changing

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Table 12.6 Export of fish seed from the various Indian states. States

Species

Assam

IMC & EC

283.3

Andhra Pradesh Bihar

IMC & EC

300

IMC Exotic carp Chhatishgarh IMC

190 10

Goa Gujarat Jammu & Kashmir Jharkhand

IMC & EC IMC

0 100 90

0 0

50 0

10 0

10000 15000

0 0

Manipur Punjab

IMC Exotic carp IMC IMC & EC

0

5000

0

Rajasthan Tripura

IMC & EC IMC

0 0

175 0

0 143.62

Tamil Nadu

P. monodon 700 million PL stage IMC 5,190

West Bengal

Uttar Pradesh IMC

Spawn (million)

0

Fry (1,000)

Fingerling (1,000)

11,400 7,600 600 400 174,100

55,000

0

Destination

Arunachal Pradesh, Meghalaya, Nagaland, Mizoram, Manipur, Tripura Maharashtra, Tamil Nadu, Karnataka, Orissa Uttar Pradesh, Nepal, Jharkhand Uttar Pradesh, Nepal, Jharkhand Himachal Pradesh, Orissa, Maharashtra Maharashtra Maharashtra, Rajasthan Sikkim, Bhutan, Uttarakhand Chhatishgarh Orissa Nagaland Haryana, Rajasthan, Himachal Pradesh and Jammu & Kashmir Haryana, Punjab North Eastern States (NEH), Bangladesh Andhra Pradesh, Kerala, Gujarat, West Bengal, Maharashtra Andhra Pradesh, Assam, Bihar, Chhatishgarh, Gujarat, Jharkhand, Maharashtra, Manipur, Orissa, Rajasthan, Tripura, Uttarakhand, Bangladesh, Nepal, Bhutan, Karnataka, Tamil Nadu Bihar, Haryana, Himachal Pradesh

Source: Based on information supplied to CIFE by the Directorate of Fisheries of the governments of the various states

environmental conditions and resistance to diseases. Appropriate breeding programmes for maintaining and improving the genetic quality have yet to be introduced in most of the seed farms and hatcheries. Seed collected from natural resources are now used mainly for raising broodstocks with the objective of improving / maintaining the genetic quality of native major carps. Significant development in this regard is the development of an improved rohu popularly known as Jayanti rohu by the Central Institute of Freshwater Aquaculture (CIFA) through a long-term programme of genetic selection. India expects to introduce seed certification soon. Guidelines for seed certification include certain prerequisites for certification of hatcheries and other regulations. These guidelines have been prepared by the Indian Council of Agricultural Research (ICAR), which awaits formal introduction by the Government. The National Fisheries Development Board (NFDB) is working on strategy for achieving fish seed self-sufficiency through decentralised fish seed production and ensuring year-round availability of adequate seed of the desired quality and size.

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Table 12.7 Import of fish seed by the different states of India. State

Species

Andhra Pradesh Arunachal Pradesh Bihar

Pangasius

Spawn (million)

Fry (1,000)

Fingerling (1,000)

State

25,000

0

West Bengal

IMC

0

13.4

0

Assam

IMC EC IMC IMC IMC P. monodon IMC IMC & EC

20 2 0 1.3 0 PL stage 0 0

5,000 5,000 212,260

0 0 0

0 20,000 10,000 13,800

40

IMC &EC

0

3,002.89

0

West Bengal West Bengal West Bengal West Bengal Karnataka Tamil Nadu, Karnataka West Bengal Uttar Pradesh & West Bengal Punjab

IMC EC

1500

30,000 500

0

West Bengal West Bengal

IMC

0

127,534.8

0

IMC

0

249,600

8,000

0 PL stage

13,600 14,500

2,500

0 0 25 0 0 0

Rajasthan

EC M. rosenbergi IMC IMC IMC IMC &EC IMC EC Others IMC & EC

100,000

2,072.34 15,000 45,000 50,000 15,000 5,000 2,000 237,012

0 0 5,000 2,000 1,000 800

Sikkim Tamil Nadu

EC IMC

0

2,000 0

72,000

Uttar Pradesh West Bengal

IMC P. monodon

0 PL stage

Maharashtra, Andrha Pradesh, West Bengal, Chhatishgarh Andrha Pradesh, West Bengal, Gujarat Chhatishgarh Andrha Pradesh, West Bengal Andrha Pradesh, West Bengal, Gujarat Assam Assam, Manipur West Bengal West Bengal Tamil Nadu Tamil Nadu Tamil Nadu West Bengal, Gujarat, Uttar Pradesh West Bengal Andrha Pradesh and West Bengal West Bengal Andrha Pradesh, Tamil Nadu, Kerala, Orissa

Chhatishgarh Delhi Goa Gujarat Haryana Himachal Pradesh Jharkhand Jammu & Kashmir Madhya Pradesh Maharashtra

Meghalaya Nagaland Orissa Punjab Puducherry

30,000

0

0 510

Source: Based of information supplied by the Directorate of Fisheries Govt. of different states to CIFE

In neighbouring Bangladesh, which is an equally important country for carp production, efforts are being made to development and implement a fish seed certification system. In the year 2007–8 the country produced about 423,986 kg of spawn from its 950 hatcheries. Details are given in Table 12.8. A detailed account of freshwater fish seed resources and supply is detailed in Bondad-Reantaso (2007).

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Recent Advances and New Species in Aquaculture

Table 12.8 Production of fish and shrimp seeds in Bangladesh in 2007–8. Type of hatcheries

Carp & other fish (Pangasius spp., Anabas, Clarias) Shrimp (monodon) Prawn (M. rosenbergii)

Number of hatcheries

Annual production

Wild-caught

950

423,986 kg*

1,872 kg*

59

5,115 million PL 1,100 million PL

1,000 million PL

70

* Usually 3.5 to 4 million carp spawn weigh 1 kg Source: personal communication with Ministry of Agriculture, Bangladesh

12.5.1.5 Carp seed rearing facilities and practices Non-drainable dug-out earthen ponds are extensively used for the rearing of fish seed throughout India and even in Bangladesh. Usually two stages of seed rearing are in progress, from spawn to fry and fry to fingerlings. Spawn are stocked in well prepared nursery ponds and reared generally for two to four weeks. Such ponds, known as nursery ponds, are relatively small and shallow. Nursery pond preparation involves steps such as dewatering and drying of ponds, liming, manuring, application of oil emulsion / soft organophosphorus insecticides to control predatory insects, etc. Manuring is done by application of organic and/or inorganic fertilisers. Commonly applied organic manures are raw cow dung or poultry manure. These manures are applied by broadcasting or by mixing and sprinkling throughout the pond. If it is not possible to dewater the pond, pesticides like mahua oil cake or bleaching powder are applied to get rid of unwanted fish and predators. Spawn is stocked after a week of manuring or about a fortnight after the application of mahua oil cake. Some farmers also apply soaked and partially fermented oil cake to manure the pond. They believe that application of oil cake as manure prevents the growth of predatory larger forms of crustaceans – copepods and cladocerans. Spawn is stocked at the rate of 5 to 10 million per hectare depending upon availability of spawn and the farmer ’s market strategy. Usually spawn are reared from two to four weeks to reach fry stage. Fry are then transferred to bigger ponds (usually 0.05 to 0.1 ha) for rearing of fry up to fingerling stage. Rearing ponds are prepared in the same way as nursery ponds except for eradication of aquatic insects or larger-sized copepods and cladocerans. Stocking of fry is usually at the rate of about 0.1 to 0.5 million depending upon the farmer ’s choice and market demand related to size of fingerlings (Fig. 12.3). In recent years many farmers have been carrying out one-step rearing of seed. Even bigger ponds are stocked heavily and thinned periodically by selling fry and subsequently fingerlings and even yearlings. Such practices are becoming increasingly popular due to increasing demand for yearlings for stocking in table fish production. There are specialised farms that produce fry, fingerlings and yearlings. Spawn are reared up to fry, fingerling and yearling sizes in ponds usually ranging between 0.2 and 0.8 ha in size. Most of the seed farms sell fry and fingerlings to table-size production farms but some seed farms maintain fingerlings for a period of one year to sell yearlings of 50–150 g. In such cases, the pond area ranges between 1 and 2 ha. The yearlings thus reared constitute major stocking material for culture ponds in Andhra Pradesh and are now becoming popular

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Fig. 12.3 Preferred size of fingerling for stocking.

in other areas as well. Some table-size fish growers/farms also keep separate ponds of 1–2 ha to meet their own year-round requirement for advanced fingerlings/yearlings. Nylon twine/strings are hung across the entire pond at short distances, about 1 m above the water level, to avoid predation by birds. 12.5.1.6 Seed transportation and marketing Spawn are transported in oxygen-packed polythene bags which are kept in tins or cardboard boxes, while fry, fingerlings and yearlings are transported in 0.5–1.0 ton LDPE tanks and loaded in trucks/tractors. Commercial oxygen cylinders are also carried in sufficient numbers to provide continuous aeration of water in carrier tanks. This method is also followed for long-distance transportation of fry, fingerlings and yearlings (Fig. 12.4). The transportation time may vary from 6 to 24 hours. The survival rate is reported to be very high (80–90%). This method is also followed for long-distance transport by road up to 800–1,200 km. During the course of over two decades of commercial carp operations in the Krishna and W. Godavari districts of Andhra Pradesh, an efficient and organised marketing chain has also been established. Such a marketing chain is unique to this area. Fish traders arrive at the pond site with trucks loaded with ice, and fish packing trays. After harvesting fish are weighed at the site, chilled and packed. At the packing unit, fish is repacked in trays of 40 kg each with crushed ice in 1:1 ratio. Before loading, a thick layer of rice husk is spread on the floor of the truck to provide insulation. Trays packed with fish ice are stacked in the truck and husk is placed in between the trays. The entire load is then covered with tarpaulin and fastened. This method ensures safe preservation for up to one week.

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Recent Advances and New Species in Aquaculture

Fig. 12.4 Transport of advanced fingerlings/yearlings.

12.5.2 Table-size production – systems and practices The fisheries sector contributed Rs.25.4 billion (about US$0.6 billion at then current exchange rates) to the national income, which was 0.8% of the total gross domestic product (GDP) and 4.97% of the agricultural GDP of India during 2007–08 (Govt. of India 2010). In the present context when the country is focusing on inclusive growth that can close the widening gap between India’s urban and rural sectors, much is expected from the fisheries sector. About two-thirds of India’s people live in villages and depend largely on agriculture and allied sub-sectors – animal husbandry, dairy and fisheries – for their livelihood. It is increasingly realised that there is no other alternative than to boost agriculture and the allied food production sector for poverty reduction through development of rural livelihoods, improving the quality of life of rural population and achieving food and nutritional security. In this context, compared to other food production sectors, aquaculture and enhanced fisheries have still considerable untapped potential for both horizontal and vertical expansion. The government introduced the Mahatma Gandhi National Rural Employment Guarantee Act (MGNREGA) in 2005. This is a ‘mega programme’ with the objective of providing a guaranteed 100 days of employment to rural workers. It gives priority to the creation of productive permanent assets in the form of ponds, irrigation tanks, rural roads, etc. This mega programme is expected to create a huge number of water harvesting and holding structures, in addition to the existing aquaculture resources.

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Currently about 95% of India’s aquaculture production is through inland aquaculture. A viable technology for freshwater fish culture that evolved over many years has contributed significantly to the growth of the country’s inland fish production.

12.5.2.1 Systems of culture 12.5.2.1.1 Pond culture Fish culture in non-drainable rain-fed ponds is the most common and extensively practised system in India. Depending upon the availability of water, and the depth and quality of the soil, these ponds are seasonal to perennial in nature. Perennial ponds are used for table-size fish production while seasonal ponds are used for seed production and short-duration culture of food fish. In eastern and some northeastern states of India like Orissa, West Bengal, Tripura, Assam, etc., the majority of households have small multipurpose homestead ponds. Water is used for bathing, washing clothes and utensils as well as for animals, besides the culture of fish. A similar practice is also prevalent in neighbouring Bangladesh. Such homestead ponds are relatively small – ranging from few hundred square metres to about 0.1 to 0.2 ha in area. In addition to homestead ponds there are community village ponds in most parts of India. These ponds are larger in size, spreading out to one to several hectares and excavated with multipurpose objectives. Such community ponds hold water for irrigation, recharging the groundwater table, for bathing, washing, drinking water for animals and also water to be used during emergencies such as outbreak of fire, etc. In some parts of India, especially a few districts of Andhra Pradesh, Punjab and Haryana ponds are also fed with water from irrigation or drainage canals. In most cases these ponds are constructed exclusively for commercial aquaculture. Such ponds are usually larger, spreading to several hectares and in some cases with drainage facilities and inlet structures. 12.5.2.1.2 Pen culture Culture of fish in pens is developing rapidly. Pens are enclosures created in larger sheets of shallow water such as floodplains, ox-bow lakes, etc. Locally available fencing materials such as bamboo are used for erecting fences. In some areas net is also used in addition to bamboo fencing to prevent the escape of smaller fingerlings. Pen culture is used for raising both food fish and fingerlings. 12.5.2.1.3 Cage culture Though it has been practised to some extent for a long time, cage culture is yet to become popular in India. Economics factor are the main constraint to the popularisation of cage culture of carp. Since carp are a low-value fish species and culture of fish in cages is based mainly on feed, the resulting margin of profit is usually not encouraging. In recent years cage culture has been used to rear fingerlings for reservoir stocking. In such cases fingerling rearing in cages is not economically viable, but it becomes economically viable under overall economics of reservoir fisheries. Usually such cages are constructed using locally available framing materials such as wood, bamboo, etc.; plastic drums and synthetic netting materials 6 × 3 × 1.5 m in area with 1 mm mesh size are also commonly used.

12.6

DEVELOPMENTS IN CULTURE PRACTICES

Carp culture is synonymous with freshwater aquaculture in India. Though the current practices of carp culture follow the same basic principles of composite fish culture there

352

Recent Advances and New Species in Aquaculture

are variations in scale of operations and extent of input use. Based on the scale of operations the current practices may be classified into two major types: household-based smallscale culture and large-scale commercial aquaculture. The former is widely practised in states where carp are the favoured food of the locals, while commercial-scale operations are most prevalent in states such as Andrha Pradesh, Punjab and Haryana, where the produce are marketed to other states and even exported to a certain extent. Though freshwater fish culture in general is considered as an important and powerful tool for promoting rural development, small-scale culture is also widely accepted as significant contributor to food and nutritional security and poverty reduction.

12.6.1 Small-scale aquaculture Small-scale aquaculture accounts for the bulk of freshwater fish production. It includes low cost manure-based systems, attaining production of 1 to 3 tonnes/ha, and manureand feed-based systems with production over 2 tonnes/ha. The practice includes steps like drying, desilting of ponds, application of lime and initial manuring with locally available organic manure such as raw cow dung, compost, poultry droppings, etc. In cases where farmers encounter difficulties in drying the pond completely they apply de-oiled cake of mahua (Madhuca longifolia) at the rate of 250 ppm to remove predatory and unwanted fish species. In such case the initial manuring is ignored as the mahua cake itself acts as organic manure. 12.6.1.1 Low- cost manure -based aquaculture The majority of subsistence farmers cannot afford to apply supplementary feed as the feed alone costs anything between 60 and 70% of the total cultural cost. To fit into the budget of subsistence-level farmers, manure-based low-cost aquaculture has been introduced in Bangladesh and also parts of India. Usual pond preparation practice including liming and initial or base manuring is followed. However, to increase the efficiency of manuring the total initial dose of organic and inorganic manures are mixed with an adequate amount of water and spread uniformly over the entire pond. Alternatively, green manuring is also followed by sowing seeds of dhaincha (Madhuca longifolia) at the rate of 50–60 kg/ha prior to the rains. Once rain comes, the crop of well grown dhaincha plants are uprooted and buried in pond water. Within a fortnight they are completely decomposed and serve as good organic manure. In case of mahua oil cake application and in cases where water turns deep green within a week of liming, initial manuring is avoided. Such conditions prevail in older ponds where abundant nutrients are available in thick bottom sediments. In such ponds raking of bottom sediment at frequent intervals in the afternoon is recommended. Ponds are stocked with yearlings to get better growth. Stocking density is followed at the rate of 3,000–5,000 per ha and with a combination of three Indian major carps. In many cases multi-stocking is also done depending upon the availability of desired quality and quantity of seed. Partial harvesting also goes on simultaneously. Depending upon the demand and price and availability of aquatic weeds, Chinese carps such as silver carp, common carps and grass carp are also included. Since common carp cause erosion of pond dykes due to their burrowing habit, many farmers avoid common carp. Common carp is also avoided because of the need for the fish to attain maturity and reproduce at an even size range of 200–300 g. This is difficult in the highly inbred

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353

common carp stock. Care is taken to make the best use of summer months. Pond preparation is done during the winter months when the growth of fish is minimal. Care is also taken that the pond is not kept empty during the summer months when the growth of fish is at a maximum. Except for manure, no other inputs are applied in the pond. To enhance the efficiency manure is applied in liquid form and on a daily basis. A mixture of raw cow dung (5–6 kg), urea (100 g), triple super phosphate (50 g) and muriate of potash (25 g) is mixed with an adequate amount of water and spread over the entire pond daily. Manure is applied during the daytime when dissolved oxygen level is at a high level. On overcast days and when the water turns deep green, manuring is suspended. Harvesting (Fig. 12.5) is conducted depending the growth attained and also market demand. Usually larger fish are removed and sold (Fig. 12.6) or consumed and the pond is re-stocked with a similar number of fingerlings. Farmers following the package of practices seriously usually attain a production of about 2.5 to 3.5 tons/ha/year. Due to the low costs of production the margin of profit is much higher and the level of risk is minimal. 12.6.1.2 Manure - and feed-based aquaculture More aware and relatively well off and advanced farmers follow semi-intensive aquaculture practice where in addition to manure, feed is also applied. They follow the pre-stocking pond preparation more strictly. Predators and weed fishes are eradicated by dewatering and drying of the pond. Alternatively, pesticide of plant origin such as mahua oil cake is applied at 250 ppm. Depending on availability, bleaching powder is also used in some areas.

Fig. 12.5 Harvesting in carp culture pond.

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Recent Advances and New Species in Aquaculture

Fig. 12.6 Catla (Catla catla) being packed in ice prior to sale.

Bleaching powder is applied at the rate of 50 ppm. This is followed by application of lime at the rate of about 250 kg/ha. A relatively higher stocking is followed, usually ranging from 6,000–10,000 fingerlings per hectare. A combination of the three Indian major carps is most common although major Chinese carp species – silver carp, grass carp and common carp – are added in the combination. Local availability of seed and market demand and cost influence the species combination. After manuring, feed is applied at the rate of about 2–3% per day in the form of a mixture of mustard or groundnut oil cake and rice or wheat bran. These are mixed usually at a 1:1 ratio. Recently some commercially available feeds have also come into use. In addition to feed, manuring is conducted on a monthly or fortnightly basis. A combination of both organic and inorganic manure is applied in the pond. A total of 10–25 tons of organic manure, usually in the form of raw cow dung, is applied per hectare per year, depending upon the texture and quality of pond bottom. Usually higher doses are applied in newly dug out ponds and ponds with a sandy soil bottom. In older ponds the dose is reduced and even in ponds where the water turns deep green after liming, manuring is avoided. The organic manure is mixed with nitrogenous fertiliser (urea, ammonium phosphate or calcium nitrate). Usually a total of about 120 kg of nitrogen is applied per hectare per year in split doses (monthly or fortnightly) along with the organic manure. Similarly a total of 60 kg of phosphorus is also applied in split doses. Phosphorus is applied in the form of commonly available phosphoric fertilisers – single super phosphate or triple super phosphate. Potash is applied in the form of muriate of potash at the rate of 20–30 kg per hectare per year whenever needed. The total quantity of organic and inorganic manures is divided into monthly instalments and the first instalment is applied as an initial dose

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during the pond preparation stage. Manuring is suspended on rainy or overcast days and also when the pond water turns deep green.

12.6.2 Commercial carp aquaculture Commercial carp culture is becoming increasingly popular. It is practised by entrepreneurs as well as relatively better-off and progressive farmers who either own or have access to bigger water bodies or a number of ponds. However, the majority of commercial farms are located in Andhra Pradesh near Kolleru Lake, one of the largest natural freshwater lakes in India. It covers an area of 308.55 km2, extending into the districts of West Godavari and Krishna in Andhra Pradesh. The paddy fields surrounding this lake used to flood every year and the rice yield was very poor. During the 1970s a few innovative farmers of Pothumarru village started fish culture in community ponds and obtained substantial profits as compared to the income obtained from rice farming. Since then aquaculture has gained prominence in the area. Although profits were initially low due to the lack of local demand for carp, the potential of the industry was realised in view of the demand and high price for fish in Kolkata market. During the 1980s large-scale conversion of paddy fields into fish ponds was witnessed especially in the peripheral areas of Kolleru Lake spread over West Godavari and Krishna. With the indirect concurrence of the government the entry of the dynamic and enterprising farmers to fish culture led to proliferation of fish ponds to an extent of about 60,000 ha with each growout pond ranging between 4 and 60 ha. The farmers followed the technology of composite fish culture and could achieve sustained annual fish production at the rate of about 5–10 tons/ha. Some of the more innovative farmers were even able to attain a yield of 10–15 tons/ha. However, they were aware of the high risk associated with cases of fish mortality due to outbreak of diseases and environmental problems. With increasing experience, farmers kept on modifying their practices to ensure a higher income. Currently they follow a ‘greenwater ’ system by using mainly locally available inputs such as chicken manure, inorganic fertilisers, mixed mash feed of de-oiled rice bran and oil cake. It is estimated that during the year 2008 carp production reached 450,000 tons exclusively from this region (Edwards 2008). Ponds used for commercial carp culture are usually bigger in size ranging between 4 and 60 ha, the average size being 6 to 8 ha. Larger ponds of over 8 ha are of the trench type, with a shallow central portion retaining a water level of about 1.2–1.6 m and 2–2.5 m in the trenches. Though there are no fixed norms for the shape most of these ponds are rectangular or square in shape. Prevalent pond soils are black cotton with more clay content. Irrigation canals originating from Godavari and Krishna rivers serve as the primary water source for fish ponds in this area. However, there has been large-scale and indiscriminate expansion of ponds by the conversion of paddy fields into fish ponds, and some farms also depend on drainage canals using runoff water from paddy fields as the major source of water. The network of drainage canals serve as feeder as well as drainage channels to a large number of commercial fish ponds.

12.6.2.1 Pond preparation Pond preparation starts with draining and drying of ponds. This is followed by liming and manuring. In the case of organic manures, poultry manure is applied at a rate of 7–10 tons/

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ha as initial manuring. Inorganic fertilisers such as super phosphate and DAP/urea are also applied at a rate of 200 kg and 50–75 kg/ha, respectively. Post-stocking manuring is done in phases using poultry droppings at a rate of 2.5–4.0 tons/ha depending on water colour (availability of plankton). No definite periodicity is maintained. Usually subsequent manuring is done as needed. It should be noted that farmers do attempt to maximise fish production and profitability by enhancing the efficiency of the technology and inputs used, reducing costs of production and minimising the role and share of middlemen. This practice of carp culture largely depends upon the natural fish food organisms and feed is applied to supplement growth. 12.6.2.2 Species composition Initially the practice of commercial aquaculture in this region started with the classical practice of culturing three Indian major carp and three Chinese carp. Subsequently the farmers discarded the Chinese carp, except for grass carp, which they included at a lower stocking ratio. This modification was introduced for financial resons. During the past decade farmers resorted mainly to the culture of three Indian major carp were catla (30%), rohu (60%) and mrigala (10%). Subsequently, due to problems encountered in the harvesting of bottom-dweller mrigal the species was gradually discarded. Usually farmers had to resort to complete dewatering of the ponds to ensure complete harvesting. Farmers feel that dewatering ponds ranging between 20 and 100 acres in size is a colossal waste of water, as well as energy, money and time. In addition, mrigal does not have a market price comparable to catla or rohu. Of late, the stocking ratio of catla has also been reduced to around 10%, as fish over 2 kg commands high value in the local markets. Reducing stocking density invariably results in larger sized catla. The commonly practised stocking composition is catla 10% and rohu 90%. 12.6.2.3 Stocking density Usually no fixed stocking density is followed. It depends on the ponds, their situation, prevailing water depth, availability of stocking materials, etc. However, the density usually varies between 6,000 and 10,000/ha. The practice of stocking of ponds by fingerlings has been replaced by stocking with stunted yearlings with a size range of 50–150 g. The rate of stocking also depends upon the season of stocking, harvesting time and market trends prevailing in the Howrah, Eastern and Northeast markets. Other factors taken into account include desired target production, input supply, capacity of the farmer, etc. 12.6.2.4 Feeds and feeding practices Supplementary feeds commonly applied are largely locally compounded using ingredients like de-oiled rice bran and groundnut oil cake. Alternatively, broken rice, cotton-seed cake, broken maize, jowar (Sorghum bicolor), etc., are also used. A composition of 85% de-oiled rice bran (DOB) and 15% groundnut oil cake (GOC) is commonly used for supplementary feed for carp culture. However, composition is modified a month before harvesting by increasing GOC to 20%. Cotton-seed cake is given to offset the escalating cost of GOC. Crushed maize and jowar are added as a cheap source of carbohydrates. In recent years floating as well as sinking pellet feeds have also been introduced. A few farmers tried a combination of traditional mash feed comprising DOB and GOC along with pellet feeds. However, in this method pellet feed constitutes about 30% of the total feed

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Fig. 12.7 Bag feeding.

applied. Generally feed is applied at the rate of 4–5% of the fish biomass for the first few months and then reduced to 3% and finally 1.5–2% during the last few months of culture period. The most common method of application of feed is bag feeding (Fig. 12.7). Mash (powdered) feed is put in used plastic bags and tied to bamboo poles fixed in the pond. About 10–12 holes of 1 inch diameter are made on the bags to facilitate the slow release of feed in the pond. About 10–15 feed bags are spread out evenly over a pond of one hectare to ensure proper distribution of feed. These bags are hung on poles and kept immersed in water. In larger ponds of 20 ha and above a new method is being followed. A nylon rope is tied from one bank to the other across the centre of the pond, and all feed bags are suspended from this rope at equal distances apart. This method has several advantages – it saves on the cost of bamboo poles, distance is cut short and a person can fix all the bags to the rope quickly by moving across the pond. Farmers generally use small locally made boats to apply feed, since the water level is kept close to 2 m. 12.6.2.5 Pond environment management Irrigation /drainage canal water is the main source of water for aquaculture in this area. This source of water is available for about 10 months of the year. While water is pumped into the culture ponds from the adjoining canal, sluices are used for draining/dewatering. Water exchange practice is not adopted unless there is a severe disease problem. Water is maintained at a depth of around 2 m. However, in trenches it may be deeper. During harvesting the water level is reduced to about 1 m for ease and efficiency of drag netting.

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12.6.2.6 Culture period Usually the culture period lasts for 8–10 months. Some farmers carry out partial harvesting (Fig. 12.8) after 6 months or when the fish exceed 1.2 kg and can already fetch a good price. In such cases restocking with yearlings follows harvesting and the culture is continued. Commonly the harvest size of rohu (Fig. 12.9) is 1.2–1.5 kg and of catla 2.0–2.5 kg (Fig. 12.10). 12.6.2.7 Fish yield and cost of production Fish yield of about 7.5–12 tons per hectare per year is common in the area. Farm gate prices were fairly constant for several years. However, a marked increase has been noticed recently varying from Rs 45–52 per kg (approximately US$1.00–1.10 per kg). It should be noted that the farm gate price is very dependant on markets in the Eastern and Northeastern states which are about 1,000–2,000 km away and where the prices range from Rs100–150 per kg depending upon the size of the fish. Cost of production ranges from Rs35–40 per kg.

12.6.2.8 Fish health management Commonly encountered diseases in commercial carp culture operations are caused by parasites and microbes. Parasitic infections caused by Argulus, Lernaea, Gyrodactylus and Dactylogyurus, Myxobolus, etc., are frequently encountered. Among the microbial disease columnaris, hemorrhagic septicaemia and dropsy are common. Outbreak of viral disease has not been reported so far. However, none of these diseases causes mass-scale mortality.

Fig. 12.8 Partial harvesting by netting to assess the stock.

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Fig. 12.9 Freshly harvested rohu (Labeo rohita).

Prophylactic treatment in the form of pond treatment and medicated feed is common. Among the parasitic disease argulosis is most frequently encountered in brood fish and table-sized rohu. Organophosphorus pesticides such as Malathion and Sumithion were used in past years to prevent, control and eradicate this disease by three applications at weekly intervals. However, it has been reported that Argulus has developed resistance to such pesticides. Pyrethroids such as Deltamethrin, Cypermethrin, Decis, etc., are used extensively to control Argulus. Lernaea causes problems in fingerling rearing. Larnaea infection is more frequently encountered in catla fingerlings. A commonly used treatment is an application of formalin as well as pesticides. Bacterial diseases are prevented and controlled using antibiotics such as Doxycycline, Endofloxacin, etc. Farmers have found that most of the formulated/packed antibiotics are ineffective. As a result farmers are now resorting to using the active ingredients mixed with the feed. Some of the farmers also apply probiotics and sanitisers for water and health management. On an average about Rs7,000–10,000 per ha is spent on fish health management.

12.7

CULTURE OF PANGASIUS (PANGASIANODON HYPOPHTHALMUS)

Many farmers have adopted mono- or polyculture of Pangasius sutchi and P. hypophthalmus (Lakra & Singh 2010), which has recently been legally accepted for culture by the government of India.

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Fig. 12.10 Freshly harvested size of a catla (Catla catla).

Commercial-scale culture of this introduced exotic catfish is still limited mainly to Andhra Pradesh. However, the seed is still supplied from West Bengal with the main source probably being Bangladesh. Culture of this species has spread to over 24,000 ha (2009) in Andhra Pradesh alone with production of about 2,500,000 tons a year. Like carps it is also reared in seed tank up to advanced fingerlings of about 4–6 months old before stocking. The most common practice is monoculture of this catfish. However, some farmers also culture this species in combination with carp. In monoculture operations stocking of advanced fingerlings is done at the rate of about 15,000–20,000 per ha. Some farmers combine with carp at a rate of 200–500 per ha. Fish are fed with feed compounded using ingredients like broken rice (boiled), cotton-seed cake and maize, in addition to de-oiled rice bran. Recently commercially available soya-based floating pelleted feeds (Fig. 12.11) have also been introduced. Four commercial feed manufacturers have also put feeds on the market. The cost of these commercial feed ranges from Rs20–21/kg, which gives a feed conversion ratio (FCR) of about 1.1 to 1.2:1. The major advantage of using pelleted feed is the low incidence of disease occurrence, good pond water quality and no risk factors, unlike carp farming. This has lured many shrimp and carp farmers to switch over to catfish culture in a big way but the dearth of seed is slowing down the expansion of catfish aquaculture. Pangasius production (Fig. 12.12) is going through the phase of intensification with yield ranging from 20–50 tons/ha. Larger-size Pangasius are sold at the farm gate at Rs45 and 50/kg (Fig. 12.13), while the cost of production ranges from Rs28–35/kg (with mash feed) and Rs33–38/kg with pellet feed.

Fig. 12.11

Commercially available floating pelleted feed for Pangasius.

Fig. 12.12 Harvest of catfish (Pangasius).

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12.8

FRESHWATER PRAWN FARMING

Culture of the giant freshwater prawn Macrobrachium rosenbergii (Fig. 12.14) in freshwater aquaculture is also gaining momentum. Carp culture along with prawns offers a vast potential for generating a large quantity of good quality animal protein at low cost (Mohapatra et al. 2007). According to an estimate by MPEDA, in 2005–06 scampi (as giant freshwater prawn is popularly known) was cultivated on nearly 43,000 ha. During this period production rose to about 42,780 tons in India (Fig. 12.15), making it the second largest global producer. The state of Andhra Pradesh alone contributes about 87% of the total production.

12.8.1 Status in landlocked states Landlocked states such as Punjab, Haryana, Rajasthan, Uttar Pradesh, Bihar, Tripura, Assam, Chattisgarh and Madhya Pradesh have also started scampi culture. Initially this depended on seed collected from the wild or from hatcheries located in coastal states. However, many of the inland states have also established hatcheries. The Central Institute of Fisheries Education (CIFE) has also assisted states like Tripura, Assam, Nagaland and Manipur to establish scampi hatchery. CIFE has also established synthetic seawater-based and underground saline water fed hatcheries in Haryana and Madhya Pradesh. CIFE is also

Fig. 12.13 Market size freshly harvested catfish (Pangasianodon hypophthalmus).

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Fig. 12.14 Freshwater prawn (Macrobrachium rosenbergii).

40000

33,556

35000 Area (ha.) 30000

Production (tonnes)

25000 20000 15000 10000 5000 752

144

692.57

1934.76

3680 310

59.25

0 Kerala

Karnataka Maharashtra

Gujarat West Bengal

Orissa

Andhra Pradesh

Tamil Nadu

Fig. 12.15 Giant freshwater prawn (Scampi) production in the coastal states of India. Adapted from: Marine Product Export Development Authority (MPEDA) report.

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providing technical assistance to the Department of Fisheries of the government of Bihar in designing, construction and operation of scampi hatchery.

12.8.2 Nursery pond management Nursery rearing is essential to produce juveniles for stocking. Stocking of juveniles ensures better growth and high survival. Usually nursery ponds range between 0.2 and 0.4 ha in size. Ponds are prepared after draining and drying. Water is introduced after proper sieving. This is followed by application of organic fertilisers – cattle dung at a rate of 1.5–2 ton/ha or poultry droppings at 0.5–0.75 ton/ha along with inorganic fertilisers such as Di-ammonia phosphate (DAP) or urea at 50–75 kg/ha and super phosphate at 25–40 kg/ha for promoting luxuriant development of plankton. Hatchery produced seed (PL 5 to 7) are stocked after proper acclimatisation. At this stage PL usually cost about Rs150–250 per thousand. PL are stocked in prepared nursery at the stocking density of 150,000 and 200,000/ha. Survival of seed is estimated by stocking samples of 200 PLs in a hapa for 24 to 48 hours. Commercially available starter feed is broadcast 2 to 3 times a day. Hideouts made of coconut leaves are placed in the pond. Nursery rearing is generally carried out for 45 days or till the seed attain juvenile stage and weigh 1.5 g and above. Cast nets are used for harvesting.

12.8.3 Grading of juveniles Harvested juveniles are graded by size and seed of less than 1.5 g are released back to the nursery pond for further rearing. Selected juveniles are hand sexed to segregate the males, which are stocked in growout ponds. Female juveniles are normally discarded or sold at the throw way price of Rs20–35 per kg.

12.8.4 Growout pond management (allmale culture) Ponds are prepared as for nurseries. Manuring is conducted but unlike with carp culture, manures are applied in low doses. Usually growout scampi ponds are smaller than carp culture ponds and range between 0.4–1.0 ha and water level is maintained at 1–1.3 m. In monosex culture male juveniles are stocked in ponds at the rate of 1.5 and 2 per m2. Commercially produced pelleted feeds are available in the market from several manufactures such as CP, Avanthi, Higashi, Water Base, Godrej, Kargil, etc. In addition, several locally manufactured pelleted feeds are also available and extensively used in scampi culture. Grower feeds are applied 3 to 4 four times a day as prescribed doses printed on feed bags. No aerators are used in scampi culture.

12.8.5 Claw ablation After about two to three months of rearing, the water level is reduced by 0.3–0.5 m and blue-clawed (BC) males are picked up by cast netting. The second leg is chelated, which normally turns orange in colour at the anterior two segments with some blue colour on the spines. The blueclawed animals are usually larger, aggressive, dominant and territorial and sexually very active. Besides competing for feed they inhibit the growth of other, smaller

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prawns. The growth of BC is reported to cease but farmers ablate the claw and continue to culture. The blue claw is broken, leaving the last segment attached to the carapace and released back to the ponds. The orange-clawed (OC) prawns are released back into ponds. These prawns are known to possess high somatic growth potential and are sub-dominant as well as sexually inactive.

12.8.6 Culling and harvest After 4 months of culture first culling is undertaken by using a cast net. At this stage the BC grown above 60 g are harvested. A second culling is carried out after the fifth month of rearing and final or total harvesting is done normally at the end of the sixth month. Harvested prawns usually weigh between 60 and 90 g. A small stock of prawns (small males: SM), 15–25 g in weight, are left behind and shifted to other ponds for further rearing. On average farmers achieve a yield of 800–1,000 kg/ha. Culture of prawns with carp is not common, though some of the shrimp farmers who have suffered continuous losses have switched over to carp farming. Such farmers have started polyculture of carps with scampi. Polyculture is not adopted in traditional carp culture ponds as they are deep and the survival of prawns in such ponds is very low. It is mainly for this reason that polyculture of scampi with carps is usually done either in shrimp culture ponds or shallow ponds constructed especially for scampi culture. Culture ponds usually cover an area of about 0.5–1.0 ha. Ponds are drained, dried, limed and then filled with sieved water. Following this ponds are manured by application of cow dung or poultry litter at a rate of 5 tons/ha or 1–2 tons/ha respectively along with inorganic fertilisers such as DAP or urea at 75 kg/ha and super phosphate at 40 kg/ha. Water level in the pond is maintained between 1.3 and 1.5 m throughout the culture period. Phased manuring is done using inorganic fertilisers since the use of organic manures is believed to increase the chance of bottom pollution. However, as and when required poultry litter at 500 kg/ha in liquid form is broadcast in order to maintain sustained growth of plankton.

12.8.7 Stocking Yearlings of carp of 50–100 g size comprising 20% catla and 80% rohu are stocked at a density of about 2,500–3,000 per ha. Prawn juveniles of not less than 1.5 g are stocked at 1 and 2 per m2. Traditional mash feed comprising a mixture of de-oiled rice bran (80%) and groundnut oil cake (20%) is applied in the morning hours by the bag-feeding method. Grower pelleted feed is applied in the evening hours for the prawn. In this system the culling of prawns is avoided. The culture period lasts for 7–8 months. Under this system a yield of about 400–500 kg/ha of prawn with average weight of 50 g and fish about 2,200–3,000 kg/ha with average weight of about 800 g (rohu) and 1,200 g (catla) is obtained.

12.8.8 Prospects and problems in Macrobrachium farming Based on the results obtained in pockets of Andhra Pradesh it is concluded that scampi production has scope for further development and expansion in low-lying waterlogged areas and even alkaline and low saline and alkaline waters. There is also the possibility of

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polyculture of scampi with carp in seasonal tanks and reservoirs. Females discarded after males are stocked in growout ponds can also be used for stocking natural water bodies like seasonal tanks to supplement income and produce large-size females for raising broodstock. However, improvements in Macrobrachium farming will depend on overcoming the problems listed below:

• • • • • •

Poor survival of post-larvae to juvenile Stunted growth and deformities in sub-adults Growing incidence of swollen gills and white muscle disease (WMD) Inbreeding effects Environmental problems Differential growth at harvest

12.9

RECENT DEVELOPMENTS

The compatibility of olive barb, Puntius sarana, a medium-sized carp species, in polyculture with major carps has been studied in an effort to diversify carp polyculture (Jena et al. 2008). Utilisation of non-traditional resources, such as greenhouse pond environment to improve productivity during low temperature periods, has been considered (Mohapatra et al. 2007). Progress has also been made in induced breeding of carps. Pituitary extracts were replaced by human chorionic gonadotropin (HCG), as the quality and potency of pituitaries declined (Chonder 1985). Recent studies have identified Ovaprim as a substitute for pituitary extracts (Nandeesha et al. 1990; Rokade et al. 2006). Ovaprim contains analogue of salmon GnRH and dopamine inhibitor and is believed to utilise the hormonal mechanism of the fish to induce maturation and spawning (Rokade et al. 2006).

12.10

REFERENCES

Bondad-Reantaso M.G. (ed.) (2007). Assessment of freshwater fish seed resources for sustainable aquaculture. FAO Technical Paper No. 501. FAO, Rome. Chaudhuri, H. & Alikunhi, K.H. (1957) Observations on the spawning in Indian major carps by hormone injection. Current Science, 26(12), 381–382. Chaudhuri, H., Chakrabarty, R.D., Sen, P.R., Rao, N.G.S. & Jena, S. (1976) A new high in fish production in India with record yields by composite fish culture in freshwater ponds. Aquaculture, 6, 343–355. Chonder, S.L. (1985) HCG: a better substitute for pituitary gland for induced breeding of silver carp on commercial scale. In: Proceedings of the Second International Conference on Warm water Aquaculture Finfish, pp. 521–534. GSA, Hawaii. Department of Animal Husbandry, Dairying and Fisheries (2009) Annual Report (2008–09), Ministry of Agriculture, Government of India, New Delhi. Edwards, P. (2008) Comments on possible improvements to carp culture in Andhra Pradesh. Aquaculture Asia, XIII (3), 3–7. FAO (2008) The State of World Fisheries and Aquaculture. FAO, Rome. Jena, J., Das, P.C., Kar, S., Singh, T.K. (2008) Olive barb, Puntius sarana (Hamilton) is a potential candidate species for introduction into the grow-out carp polyculture system. Aquaculture, 280, 154–157. Khanna, S.S. (1988) An Introduction to Fishes, 3rd edn. Indian Universities Press, Allahabad, India. Kumar, D. (1992) Fish culture in undrainable ponds. A manual for extension. FAO Fisheries Technical Paper No. 325. FAO, Rome.

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Lakra, W.S. & Singh, A.K. (2010) Risk analysis and sustainability of Pangasianodon hypophthalmus culture in India. Aquaculture Asia, XV (1), 34–37. Accessed from: http://library.enaca.org/Aquaculture Asia/Articles/jan-march-2010/8-striped-catfish-risk-assessment.pdf Mohapatra, B.C., Singh, S.K., Sarkar, B., Majhi, D., Sarangi, N. (2007) Observation of carp Polyculture with giant freshwater prawn in solar heated fish pond. Journal of Fisheries & Aquatic Science, 2, 149–155. Nandeesha, M.C., Rao, K.G., Jayanna, R., et al. (1990) Induced spawning of Indian major carps through singl application of Ovaprim. In: The Second Asian Fisheries Forum (eds R. Hirano & M. Hanyu), pp. 581–585. Asian Fisheries Society, Manila, Phillipines. Rokade, P., Ganeshwade, R.M. & Somwane, S.R. (2006) A comparative account on the induced breeding of major carp Cirrhina mrigala by pituitary extract and ovaprim. Journal of Environmental Biology, 27, 309–310. Sinha, V.R.P. (1971) Review of composite fish culture techniques. In: Proceedings of All-India Coordinated Research Project on Composite Fish Culture. ICAR, Cuttuck. Talwar, P.K. & Jhingran, A.G. (1991) Inland Fishes of India and Adjacent Countries, Vol. 1., p. 541. Oxford and IBH Publishing Co, New Delhi.

13

Future Directions

Bruce Phillips, Ravi Fotedar, Jane Fewtrell and Simon Longbottom

13.1 INTRODUCTION The over-exploitation and decline of natural fisheries, because of an increasing human population, has led inevitably to a need for alternative sources of seafood. In many parts of the world, aquaculture has already established itself as an important source of food and other aquatic products. As a result of the decline in wild fisheries catch, the contribution from aquaculture to world seafood production has overtaken wild fisheries for the first time. Many scientists question whether this growth rate in aquaculture is sustainable. In order to address sustainability issues, it is imperative that the focus of aquaculture shifts from being only production-oriented to address the triple bottom line, which not only includes higher economic returns but also addresses environmental and social concerns. This chapter discusses certain issues that should be incorporated while strategically planning the future of this industry.

13.2

DEVELOPMENTS IN MANAGING THE ENVIRONMENTAL IMPACTS OF AQUACULTURE

The rapid increase in aquaculture production and the use of more intensive practices has led to concern over the environmental impacts of aquaculture (Pillay 2004). The level and type of impact will depend on the species involved, the system used and the nature of the surrounding environment. Generally, common impacts include: water use issues, pollution (particularly suspended solids, nutrients and chemicals), physical loss of habitat, hybridisation and loss of genetic diversity to wild populations through escapees and an increase in pathogens to the surrounding environment. These and other direct impacts will result in a multitude of flow-on effects. Some examples of recent developments in knowledge and technology that already, or could potentially, assist in the assessment, monitoring, prediction and mitigation of these impacts are discussed in the following sections. However before continuing, it would be remiss not to mention the potential environmental benefits that

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

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aquaculture possesses for relieving the pressure on wild fisheries, wild stock enhancement and supporting the reduction in trade of wild-caught species used for medicinal and ornamental purposes (Koldewey & Martin-Smith 2010; Crowe et al. 2002; Southward et al. 2005; Zmora et al. 2005; Pomeroy et al. 2006; Naish et al. 2007; Bell et al. 2008). Research into these areas should be encouraged to increase understanding and overcome issues such as interactions with and impacts on genetic diversity of wild stocks and improving methods for accurately measuring the success of fisheries enhancement programmes (Windsor & Hutchinson 1990; Bell et al. 2006; Støttrup & Sparrevohn 2007; Kitada et al. 2009).

13.2.1 Geographical information systems – site selection and monitoring Site selection is an essential factor to consider when establishing any aquaculture farm, both in terms of productivity and ecological sustainability. In the past, the major factor influencing site selection for inland aquaculture was proximity to a water source, which led to the exploitation of sites with a high water table. This resulted in the destruction or contamination of many onshore aquatic ecosystems, particularly in Southeast Asia. The environmental impacts of offshore aquaculture tend to be more inclined to manifest from cumulative processes over time and therefore, site selection to reduce environmental impacts, and predicting and monitoring the impacts can be difficult. This will become more complex as marine-based aquaculture moves further offshore to accommodate other uses of the coastal zone, as little information exists on the linkages between near and offshore systems (Skladany et al. 2007). The increasing emphasis on sustainability has shifted the criteria for the selection of sites for aquaculture, both onshore and offshore, and has highlighted the necessity for comprehensive environmental monitoring. Avoiding proximity to sensitive ecosystems and therefore immediate, localised damage, is a relatively simple process. However, predicting the long-term environmental impacts of aquaculture practices to the area outside the limits of the farm can prove complex. Recent technological advances in tools that develop our understanding of environmental processes, for example Geographical Information Systems (GIS), remote sensing and modelling software, are important in determining the farreaching ecological implications of aquaculture practices. GIS and remote sensing have been used in various aspects of aquaculture development for approximately 20 years and possess enormous potential to benefit the environmental management of aquaculture, particularly offshore cage culture. Comprehensive mapping and surveys of inland, coastal and offshore areas using remote sensing techniques prior to farm development provides information on the suitability of an area for various types of aquaculture (Ross et al. 1993; Nath et al. 2000; Salam et al. 2003; Perez et al. 2005). Spatial and temporal data has also been used in predictive models for proposed and established farms to manage the environmental impacts and to give valuable insight into the assimilative capacity of an area (Pérez et al. 2002; Corner et al. 2006; Ferreira et al. 2007; Hossain & Das 2010). Models have also been, and will continue to be, useful in predicting the effects that climate change will have on the environment. Should the predicted changes in sea level, rainfall, water temperatures and significant weather events occur, selection of sites for aquaculture will be affected (Handisyde et al. 2006). Factors such as limited understanding of methodologies and lack of support and communication between database operators have been identified as major issues that have

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hampered GIS development and its use in aquaculture (Nath et al. 2000; Perez et al. 2003). The progression of software to more user friendly and universally compatible versions has facilitated some of these issues but further advancement is required if GIS is to become a routine analytical tool for resource managers and other stakeholders.

13.2.2

Bio-engineering

The major impact that established aquaculture ventures have on the surrounding environment is from effluent, particularly from the nutrients and suspended material that it usually contains. Therefore, research focuses, and should continue to focus, on culture techniques and effluent treatment methods that reduce the concentration of contaminants in, and the volume of, effluent from aquaculture systems. Improving the quality of effluent in closed aquaculture systems will also result in a reduction in the need for water exchange, which would be of particular benefit in areas with limited access to water or in situations where water conservation is important. The importance and technological aspects of recirculating aquaculture systems (RAS) are discussed later in this chapter. Culture techniques such as integrated aquaculture have been used successfully to ameliorate aquaculture effluent in recirculating systems or prior to release into the environment. In particular, many Asian countries have been practising integrated aquaculture in simple forms for centuries. However, in most cases, the species chosen tend to be haphazard and dependant on availability with little variation of culture conditions to accommodate and, therefore, optimise conditions for all species included in the culture (Troell et al. 2009). In addition to improvement of effluent quality, integrated aquaculture has the potential to add value through a secondary crop, a benefit not usually present in other methods of bioremediation. To date, research into integrated aquaculture has included species combinations of finfish, invertebrates and macroalgae with varying levels of success. Macroalgae species of the genera Gracilaria (Troell et al. 1997; Matos et al. 2006; Zhou et al. 2006), Porphyra (Carmona et al. 2006) and Ulva (Neori et al. 2003; Schneider et al. 2005) have shown potential to reduce dissolved nutrient levels in aquaculture effluent. Filter feeders such as bivalve molluscs have also been shown to significantly reduce the levels of suspended particulate matter resulting from finfish culture (Reid et al. 2010). There is also evidence that sea cucumbers will utilise, and therefore reduce, the benthic organic deposition from oysters when grown in co-culture and have the potential for providing a valuable secondary crop (Paltzat et al. 2008). Continued research into species combinations and culture parameters (e.g. stocking densities), on a commercial scale, is important to maximise benefits gained by integrated aquaculture, particularly in open water and offshore systems where research has been hindered by challenging conditions and high costs. 13.2.2.1 Advances in production system designs through recirculating aquaculture systems Advances in methods of water treatment will be of benefit to the environmental management of aquaculture. The increasing intensification of aquaculture has resulted in the need for recirculating aquaculture systems (RAS) with efficient water treatment (Fig. 13.1). Recirculating aquaculture systems are generally regarded as production systems that continually re-use culture tank water after various forms of filtration. In comparison to

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Fig. 13.1 Recirculating Aquaculture System at Curtin Aquatic Research Laboratory (CARL), Curtin University, Perth, Australia. (Please see plate section for colour version of this figure.)

traditional methods of growing fish outdoors in open ponds or raceways these systems operate at higher densities in a ‘controlled’ environment using less water and occupying less area. The environmental benefits of recirculating aquaculture systems over pond, flow-through or cage systems include: a reduction in water use, an effluent lower in contaminants (e.g. a 500–1,000 fold reduction in nutrients), a decrease in the likelihood of escapees or pathogens being released into the surrounding environment and generally, a higher feed conversion efficiency (FCE). However, the associated costs with filtration and recirculating systems have been a major obstacle in their application, particularly in developing countries where aquaculture is most prevalent. It must be noted that technology-heavy RAS are expensive to construct, and in some cases run, in comparison to traditional methods. It has been estimated that the cost of producing freshwater food fish in a pond or flow-through system is currently two-thirds of that from a recirculating system. Likewise, the cost of raising marine fish in recirculating systems has been shown to increase the cost of production by 19% when compared to offshore cage culture (Gutierrez-Wing et al. 2006). However, this equation is constantly changing with restricted access to water, mass production of equipment in many countries and changing policy on emission taxes. It is beyond the scope of this chapter to argue the validity of RAS in the Australian aquaculture industry; rather the chapter seeks to highlight changes in system components and materials

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over the last 10 years. In theory, RAS could involve zero replacement of water (aside from evaporation) but in reality most Australian systems replace 2–20% per day (Losordo et al. 2009). This replacement water is to compensate for either sludge removed by filters (often 80% water) or direct water exchange (largely linked to the amount of investment in filtration technology). 13.2.2.2 Components of recirculating aquaculture systems Fish grown in RAS must be supplied with all the conditions necessary to grow and remain healthy. All recirculating production systems remove waste solids, oxidise ammonia and nitrite nitrogen, remove carbon dioxide, and aerate or oxygenate the water before returning it to the fish tank. More intensive systems or systems culturing sensitive species may require additional treatment processes such as fine solids removal, dissolved organics removal, or some form of disinfection. In terms of basic system components RAS have changed little in the last 10 years. What has become evident is the changes in material used to build these components and the availability (both in price and size) to the ‘common producer ’ of many high technology pieces such as ORP controlled ozone contactors (Fig. 13.2) and wireless tank monitoring systems to home computers. Culture tanks: RAS tanks can be broadly classified into raceway (flow from one end to the other) and circular (although this often refers to the flow rather than strictly tanks shape). Choice of tank style is generally governed by the primary use. For example, circular flow

Fig. 13.2 Ozone generation with ORP control.

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tanks are generally easier to setup as self-cleaning where as raceways are easier to grade fish in situ. Traditionally tanks have been concrete, steel-lined or fibreglass but in recent years plastic (HDPE) tanks have made a significant impact in the industry. HDPE can either be roto-molded or sheets welded together onsite to form whatever shape the owner desires. In this way the inefficient space utilisation of traditional round fibreglass tanks can be overcome. The recent widespread use of single-operator, plastic welders within the industry has also been a factor in the rise of HDPE, as the material can be much more quickly repaired or altered compared to other tank materials. In the Australian industry the trend in term of tanks is the direct importation of overseas designs from large, established firms such as Hesy Aquaculture BV. With improvements in global communication it is perfectly viable for even small businesses to source their entire systems from overseas yet still enjoy full service and backup from the supplier. Mechanical filters: The bacterial breakdown of fish waste and uneaten feed consumes dissolved oxygen and generates ammonia-nitrogen within a RAS. Therefore it is in the aquaculturist’s best interest to remove waste solids as quickly as possible. Waste solids can be classified into three categories (from an equipment point of view): settling, suspended, and fine/dissolved solids. The first two are of primary concern from a pollutant point of view, but dissolved organics and fine (sub-40 micron) solids become a problem as water exchange decreases. ‘Settleable’ solids are those that will generally settle out of the water within 1 hour under still conditions and are easily removed from the system. Settleable solids can be removed as they accumulate on the tank bottom through proper placement of drains. Generally ‘double drains’ and swirl separators are used to remove these solids quickly before they break down. Many pre-fabricated versions of double drains and swirl separators are available within the Australian market. Suspended solids are those that will not settle to the bottom of the culture tank under normal system flow conditions. They are therefore removed from the culture tank via normal water exchange and removed via drums filters (Fig. 13.3), belt filters, conventional sand filter housings (with varying media inside) and fluidised beds. Fine suspended solids smaller than 30 microns can contribute more than 50% of the total suspended solids load in a RAS. Dissolved organic solids (proteins) can also contribute significantly to the total oxygen demand of RAS if left untreated. Dissolved solids and fine suspended solids are generally removed using a process called foam fractionation or protein skimming. Foam fractionation introduces fine air bubbles (generally via venturi action) at the bottom of a closed column. As the bubbles rise through the water column, fine suspended solid particles attach to the surface of the bubbles, creating protein-rich foam at the top of the column. The foam buildup then flows to a waste collection tank. Biofilters: A quality mechanical filter can remove much of the uneaten food and faecal contribution to total ammonia-nitrogen (TAN). However, TAN excreted from fish gills and bacterial processes still remains within RAS. While there are technically several ways to remove TAN, the only real practical technique is with biological filtration; that is, using Nitrosomas/Nitrobacter spp. to convert ammonia to nitrate. The simplest form of biofilters are large submerged sumps containing any material for bacteria to adhere to. However, this method is inefficient as there is generally insufficient oxygen for the nitrification process to occur. The result is a much larger filter (and associated costs) when compared to modern compact floating bead filters and moving bed filters. Many companies in the Australian industry recognised this area as one that needed improving and now make or import proven designs. It must be noted that no one filter can be suitable for all types of RAS. For example, trickle down biofilters, while not as advanced or

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Fig. 13.3 Drum filters used for mechanical filtration.

efficient, do provide excellent degassing, therefore potentially eliminating some equipment (and cost). Aeration and degassing: As indicated above, filtration up to this point in an RAS consumes a lot of oxygen and increases carbon dioxide levels. This must obviously be remedied before the water returns to the culture tanks. Additionally, systems that use subsurface water as a supply often need to remove nitrogen, hydrogen sulphide, carbon dioxide or other gases from water before use. Aeration is generally supplied to culture tanks from proven equipment such as side channel blowers. While this is not the most efficient place to aerate it does provide a ‘backup’ of sorts should incoming water fail for some reason. More efficient aeration is achieved just before tank re-entry. Simple ‘packed column aerators’ are often used for this process and are effective at removing undesirable gases. Therefore, investigation into increasing the cost efficiency of recirculating systems, and in particular biofiltration, is becoming increasingly important. Adoption of methods commonly used in the wastewater treatment industry, for example moving bed reactors (Hem et al. 1994; Rusten et al. 2006; Yang et al. 2010) and airlift reactors may prove to be beneficial in this area (Losordo et al. 2009; Tal et al. 2009; Wik et al. 2009). Traditionally, biofiltration in aquaculture has relied on fixed film bacterial cultures, mainly due to their reliability and low maintenance requirements. However, recently the focus has shifted to activated suspension techniques (biofloc technology) which generally has less infrastructure

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Fig. 13.4 Oxygen diffuser used in intensive RAS.

requirements and may also provide a food source for the cultured animals. It therefore has potential as a cost efficient method for treatment of aquaculture effluent (Crab et al. 2007; De Schryver et al. 2008). Pure oxygen injectors: In intensive production systems oxygen consumption may exceed the capabilities of typical aeration equipment to diffuse atmospheric oxygen into the water. In these cases, pure gaseous oxygen diffusion (Fig. 13.4) is used. When pure oxygen is used with gas diffusion systems, the saturation concentration of oxygen in water is increased nearly fivefold to 43 mg/L at standard atmospheric pressure. This condition allows for more rapid transfer of oxygen into water even when the ambient tank dissolved oxygen concentration is maintained close to atmospheric saturation (>7 mg/L). There are three sources of oxygen used: compressed oxygen cylinders, liquid oxygen or onsite oxygen generators. Cylinders are only practical as a backup in most situations so the choice is between bulk liquid oxygen and an oxygen generator. This choice comes down to reliability and cost in relation to the specific RAS operation. Regardless of oxygen source, diffusing this relatively expensive commodity into the water is critical. As noted, aeration directly into culture tanks is not efficient therefore most Australian RAS operators using pure oxygen employ down-flow bubble contactors for efficient diffusing (up to 90%). Other equipment: Monitoring cameras and software, costing less than AUS$1,000, can be set to alarm the computer when there is no movement on screen; that is, no water flow.

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Wireless probes are available now for water quality relay back to computer. Most internet packages now come with web space sufficient to log, upload and check what is happening remotely.

13.2.3

Biotechnological

The major environmental contaminants of concern resulting from aquaculture systems are dissolved nutrients from feed and chemicals, mainly from the treatment or for the prevention of diseases. It is also accepted that the practice of using fishmeal and fish oil as a protein source in feed is environmentally unsound. Therefore, research and advances in feed formulations and alternative disease treatments continue to be of the utmost importance if the aquaculture industry is to be environmentally sustainable. Nutrients from uneaten feed and in faeces are the major contributors to the pollution of ecosystems from aquaculture and therefore methods that improve the FCE will facilitate better waste management. Simple modifications such as altering feeding regime for optimum performance and some new developments in improving FCE through behavioural, physiological and genetic manipulations are proving beneficial (Islam 2005). Breeding aquatic organisms for a particular trait that makes them more commercially viable is common in aquaculture. Worldwide at present there are at least 35 aquatic species that have undergone genetic modification resulting in improved commercial viability of the species. From an environmental perspective, the major issue that genetic modification (GM) introduces is the question of escape of GM individuals and the subsequent reproduction and competition with natural populations. Though sterilisation of GM individuals is an effective method to negate this impact, as yet there is no infallible, commercially viable sterilisation technique. Traceability of modified organisms is also important. However, for this to be commercially viable, cheaper methods than those that currently exist need to be investigated. The use of chemicals in aquaculture is widespread and they are commonly used as pesticides to control disease and reduce cage/tank fouling. However, little data exists on the effect that chemical residues in effluent, have on non-target species (Cole et al. 2009). Alternative methods of treating diseases and pests are being successfully developed, for example in vaccines, probiotics and prebiotics and other immunostimulants (Gillund et al. 2008; Kesarcodi-Watson et al. 2008; Wang et al. 2008; Kunttu et al. 2009; Zhou et al. 2009). Stress caused by high stocking densities and poor water quality induces many disease outbreaks leading to the necessity for chemotherapy. Therefore, as mentioned above, the developments in bioengineering that lead to higher water quality may reduce the need for chemicals in aquaculture. Recent figures show that over 60% of the global fishmeal and 80% of global fish oil production is used in feeds for aquaculture (Tacon & Metian 2008). To achieve environmental (and economic!) sustainability of the industry, a cost-effective alternative source of protein for use in aquafeeds is required that will not compromise other factors such as growth, survival, FCR, immunity to disease and end product quality. Research into the use of a range of plant-based protein sources is ongoing and has shown potential, particularly soybean meal (Quartararo et al. 1998; Smith et al. 2007; Fountoulaki et al. 2009; Salze et al. 2009). However, due to the relatively low crude protein level and amino acid profile in plant material, recent research has focused on the technology required to concentrate the level of protein gained from these sources and suitable dietary supple-

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ments (Lunger et al. 2007; Aksnes et al. 2008; Barrows et al. 2008; Gaylord & Barrows 2009). Recently, Kuhn et al. (2009) found that replacing fishmeal in the diet of the prawn Litopenaeus vannamei with microbial floc meal had no detrimental effect on the growth and survival. While finding a suitable alternative to animal products in aquafeeds should remain a priority, there has also been a call for a shift in focus from the culture of high-level carnivores in favour of the culture of omnivores and herbivores (New & Wijkström 2002).

13.2.4

Biomonitoring

For established aquaculture farms, monitoring programmes are an important tool for managing environmental impacts. In addition to traditional water and sediment quality monitoring, recent developments and innovative techniques include the use of bioindicators, such as invertebrate biodiversity and seagrass health, for monitoring and assessing impacts (Burford et al. 2003; Lin & Fong 2008; Pérez et al. 2008). To date research has concentrated on developing bioindicators for monitoring specific environmental impacts in localised areas. The next step is to investigate the cost effectiveness and benefits of incorporating these methods into standard monitoring programmes on a broad scale.

13.2.5

Government, policy, guidelines and standards

As relative newcomers to aquaculture, countries such as Australia can benefit from the knowledge gained from other countries’ experiences in the environmental management of aquaculture, particularly from Southeast Asia where the environmental consequences of the non-regulation of aquaculture developments and practices is apparent. With a focus on economic gain and with a lack of environmental regulations, aquaculture in this region expanded rapidly in the 1970s and 1980s (Hishamunda et al. 2009). This resulted in massive environmental degradation and the destruction of important aquatic ecosystems, especially mangroves in Thailand and the Philippines (Huitric et al. 2002). Since then, legislation and regulations have been introduced that have assisted in more suitable site selection for aquaculture and have proven successful in mitigating some of the environmental impacts. It is also becoming more common for companies to voluntarily adopt and, in many cases create their own Environmental Management Systems (EMS), Best Management Practices (BMP), Code of Practices (COP), standards and guidelines as awareness of the impact of aquaculture on the environment increases. Likewise, ecolabelling is becoming more popular as aquaculturists appreciate the benefits of being seen to provide an ‘environmentally friendly’ product. When introducing new policy to manage the impacts of aquaculture consideration must be given to, amongst other factors, the different scales of facilities, varying environmental conditions, over-restrictive policy, chemicals used, cost of compliance and social impacts (O’Bryen & Lee 2003). The logistics and practicalities of enforcing policy must also be considered. It is unusual for legislation be introduced that is specific to protecting the environment from aquaculture or to protecting aquaculture alone. For example, legislation such as the Environmental Protection and Biodiversity Conservation Act 1999 introduced in Australia is generally applied to any action that could have a potentially ‘significant effect on an

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aspect of the environment that has international significance’. Applying and enforcing legislation that governs site selection for aquaculture is relatively straightforward, and in most countries that practice aquaculture there is now at least some form of legislation governing this aspect; for example, legislation protecting the green belt in Indonesia. In Thailand, government policy also ensures that farms established prior to the introduction of legislation preventing pond development in mangroves can continue to operate under the stipulation of participation in mangrove rehabilitation programmes. Most developed countries have legislation that regulates the standards for aquaculture effluent that is to be released into the environment. However, most aquaculture occurs in developing countries where legislation regulating aquaculture effluent management is virtually nonexistent. Even if legislation was introduced, the vast area, number and variety of farms in these countries would make effective enforcement impossible. Also, in many forms of aquaculture (e.g. land-based ponds and marine cage culture) the application of standards that specify levels of constituents in effluent is difficult as a result of characteristic non point sources of pollution. In these cases BMPs have been found to be more successful than universally applicable legislation (O’Bryen & Lee 2003). The major issue facing the adoption of BMP for aquaculture, particularly in developing countries, is finding applicable incentives for farmers (Stanley 2000). The focus must be on providing them with evidence of the resultant increase in production and hence profitability, by following these practices. Although specific BMPs are dependent on the particular situation, some are broadly applicable to most types of aquaculture and, as a requirement of BMPs is continual reassessment, will improve as technology advances (Donovan 1997; Fisheries 2009). Environmental Management Systems remain one of the most powerful existing tools for management of the environmental impact of aquaculture, particularly with the incorporation of Environmental Impacts Assessment (EIA), Ecological Risk Assessments (ERA) and Environmental Management Plans (EMP). Whether it is applied as mandatory legislation, as in Australia, or is part of a self-regulated, voluntary process it enables the decision makers to assess the subsequent impacts of a proposed action on the environment. It is an efficient method for collecting, collating and presenting available information on the most efficient and recent developments and how they can be applied to mitigate the environmental impacts of aquaculture.

13.3

ECOLABELLING

Up until now the certification of fisheries has been confined to wild-capture fisheries. However, there are now a number of initiatives to develop certification and ecolabelling for aquaculture (Ward & Phillips 2008). Ecolabelling to indicate ‘environmentally friendly’ products began in 1977 with the establishment of the Blue Angel programme by the government of Germany (Müller 2002). Since then worldwide concern about sustainability issues has led to the emergence of ecolabelling schemes for many products, including those from forests (Forest Stewardship Council) and the oceans (Marine Stewardship Council (MSC)) (MSC 2002). The emergence of ecolabelling for natural resources, and particularly marine products, has been driven strongly by the involvement of non-governmental organisations (NGOs) (Sutton & Wimpee 2008). The ecolabelling of seafood has arisen in the past decade to become an

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important marketing tool in countries where consumers are sensitised to issues of environmental sustainability in food products. However, despite at least three decades of ecolabelling experience, the more recent ecolabelling of seafood presents a number of important issues, including the technical quality of criteria used to award ecolabels, the response of consumers to the rapid proliferation of different ecolabels that apply to similar products, and the potential for a distortion of the international trade in seafood to the detriment of developing countries (Deere 1999; Rotherham 2005). An ecolabel is a mark, a logo, a label or a product endorsement affixed to seafood product at the point of sale that implies to a purchaser that the product has been produced through ecologically sustainable procedures, and is from a source that is wellmanaged. Ecolabels are usually applied to each individual seafood product to provide a product endorsement that is visible at the point of retail sale. The product endorsement from an ecolabel is designed to convey to the consumer the simple message that they can confidently purchase the labelled product in preference to an unlabelled product if they wish to give their support to seafood produced in ways that have less ecological impact on fish stocks and the environment. The direct inference is that such products are more ecologically sustainable. An ecolabel may be applied to a product after it has been certified as being in compliance with the rules and criteria of an ecolabelling programme. An ecolabelling programme is a system used to create a market-based incentive to encourage products that can demonstrate they are produced in an ecologically sustainable manner. The incentive is created in the marketplace through the selective purchasing power of consumers, who preferentially purchase products marked with the ecolabel, and possibly pay a higher price for the ecolabelled product. This provides the seller and the ecolabelled product with a market advantage over non-ecolabelled products. The price increment, or possibly the increased volume of sales of the ecolabelled product, preferentially rewards the producers of the more ecologically sustainable products over those producing products without ecolabels. This possibly reduces the sales and returns to producers of less sustainable products, reduces the value or marketability of non-ecolabelled products and creates an incentive for producers to change harvesting or farming practices to be more ecologically sustainable and improve their environmental practices. Certification is the outcome of an assessment process that confirms (verifies) that a product complies with the sustainability standard and a set of criteria established by the incentive programme. A certification of compliance may be used as part of an ecolabel programme, and indeed a certificate may be issued, but not all certification systems lead to the award of an ecolabel. Some certification systems may not have any direct relationship to retail marketing issues, and may be used purely for industry or regulatory purposes (such as to demonstrate compliance with government requirements for safe food-processing procedures). So, while ecolabelling is normally based on some form of assessment process and a consequent certification, the process of assessment leading to a certification of compliance does not necessarily always lead to an ecolabel. The essential difference between certification and ecolabelling is the form of market-based incentive – influencing consumer purchases through the influence of the ecolabel at the point of retail sale and influencing purchasing patterns through the provision of other forms of product endorsement or buying recommendation. Buying guides and ratings are closely related to ecolabels, and while they constitute a different form of product endorsement, like ecolabels they also act to influence consumerpurchasing patterns and are designed to create market-based incentives. Buying guides and

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ratings provide information about a product to consumers or to resellers through advertising and seafood awareness programmes, using such tools as wallet/purse cards, brochures and websites, where ‘buy’ or ‘don’t buy’ advice is usually provided. In some systems the buying recommendation may be based on a formal and extensive underpinning decision process. Seafood recommendations for consumers based on environmental and ecological considerations were first developed in the mid-1990s, and the first seafood recommendation card for consumer use was issued in 1999 in the US Audubon Society’s magazine. Ecolabelling programmes are voluntary instruments – fisheries or aquaculture ventures can choose to submit their products for compliance assessment to determine if their products can carry the ecolabel. The voluntary nature of ecolabelling is an important aspect of consumer appeal, because this infers that products that do not meet the sustainability standard would not be submitted by the producers for assessment, and hence do not carry the ecolabel. Consumers may interpret this as meaning that only the ecolabelled seafood products available in their marketplace are indeed produced in a sustainable manner. Voluntary submission of products for ecolabelling carries with it the responsibility for meeting the various costs of conducting the compliance assessment and meeting any conditions or corrective actions that may be required to keep the product certified. These costs include the cost of the certification companies in conducting the assessment, the cost of preparing and presenting data and information about the products that match the requirements of the ecolabel programme and the ongoing costs of dealing with conditions and the costs of verification of continuing compliance. This matter of the cost of the assessment and verification system has often been raised (Deere 1999; Wessells et al. 2001) as a discriminatory factor that can be used to lock out both fisheries and aquaculture products from developing countries and small-scale ventures from the developed world. Indeed, some commentators consider that the expensive thirdparty assessment systems are verging on placement of national regulatory systems by imposing a more powerful (market-based) private sector set of management measures that are beyond the control of national governments, and transcend the more usual participatory and locally relevant management systems (Steinberg 1999; O’Rourke 2006). However, others consider that such private sector (so-called ‘non-state’) ecolabel systems can only succeed by working in partnership with government or community-led management measures, and so there is little risk of the ‘non-state’ systems replacing the ‘state’ systems (Janen 2007). Ecolabels are designed and propagated to reduce ecological impacts and improve the ecological friendliness of practices used in production, harvesting or growing of products, with a view to ultimately increasing the sustainability of all products across all the market. To achieve this, they must create ‘market-pull’ through differential appeal to consumers who are sensitive to the impact on sustainability inferred by the product endorsement of the ecolabel. The market-pull is created through establishing the credibility of the product and its ecolabel endorsement with potentially sensitive consumers, and a credible differential from competing products in the same marketplace. Nonetheless, for the increasing number of aquaculture ventures to be sustainable and to effectively manage the resulting environmental impacts, both direct and indirect impacts must be identified and, more importantly, understood. Often the observed effects of aquaculture on the environment are a result of cumulative processes, which creates difficulty in identifying the original cause. To date, research into the impact of aquaculture on the environment has been limited and has focused on the direct effects. Further research, with a

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more holistic approach, is necessary to provide information that will facilitate understanding and advances in the monitoring and mitigation of the impacts and assist in progress towards the sustainability of the industry.

13.4

THE FUTURE

Many of the chapters in this volume have suggested areas for future research or development in different areas of aquaculture, but of the species discussed cobia is perhaps the one with the greatest potential. The expansion of cobia culture is still hindered by the lack of regular supply of seed as larval nutrition and impact of formulated diets on broodstock condition and subsequent larval health is still not well understood. Further, the spawning behaviour of cobia and environmental clues which trigger the spawning of cobia is very much site-specific and needs further research in order to ensure a constant supply of viable cobia fingerlings from the hatchery. The survival of 20-day old larvae is below commercially acceptable levels due to the lack of understanding of broodstock management and the nutritional requirements of larvae. Stress during transport from nursery tanks/inshore cages to growout cages and diseases during the nursery phase are other areas that need further research. As pressure on the intensification of cobia culture increases there will be a need for further research on the use of commercially viable dietary immunostimulants to improve the immunocompetence under stressful intense farming environments. As the demand for aquacultured seafood increases the challenges relating to product quality, the environment and sustainability become more pressing. The focus of research therefore needs to change from ‘production only’ to sustain ‘triple bottom line’. This change of research focus calls for greater resources to be spent on value-addition across the entire supply chain of seafood, quality assurance programmes and the setting up of ecolabelling protocols for the aquaculture industry.

13.5

REFERENCES

Aksnes, A., Mundheim, H., Toppe, J. & Albrektsen, S. (2008) The effect of dietary hydroxyproline supplementation on salmon (Salmo salar L.) fed high plant protein diets. Aquaculture, 275, 242–249. Barrows, F.T., Gaylord, T.G., Sealey, W.M., Porter, L. & Smith, C.E. (2008) The effect of vitamin premix in extruded plant-based and fish meal based diets on growth efficiency and health of rainbow trout, Oncorhynchus mykiss. Aquaculture, 283, 148–155. Beardmore, J.A. & Porter, J.S. (2003). Genetically modified organisms and aquaculture. FAO Fisheries Circular number 989, FAO Rome. Bell, J.D., Bartley, D.M., Lorenzen, K. & Loneragan, N.R. (2006) Restocking and stock enhancement of coastal fisheries: Potential, problems and progress. Fisheries Research, 80, 1–8. Bell, J.D., Purcell, S.W. & Nash, W.J. (2008) Restoring small-scale fisheries for tropical sea cucumbers. Ocean & Coastal Management, 51, 589–593. Burford, M.A., Costanzo, S.D., Dennison, W.C., et al. (2003) A synthesis of dominant ecological processes in intensive shrimp ponds and adjacent coastal environments in NE Australia. Marine Pollution Bulletin, 46, 1456–1469. Carmona, R., Kraemer, G.P. & Yarish, C. (2006) Exploring Northeast American and Asian species of Porphyra for use in an integrated finfish-algal aquaculture system. Aquaculture, 252, 54–65.

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Index

Note: numbers in bold indicate photographs, numbers in italic indicate tables or diagrams. abalone 231–249 in Australia 231, 232, 240–243, 248, 248 barrel culture 237, 238, 241, 242 in Chile 233–239, 236, 237, 238, 239 in China 231, 232, 232 conditioning 234–235, 240 culture of 231, 233–245 feeding 237, 238, 239 global production 231, 232 growout 236–239, 237, 237, 238, 239, 242 hatching 240–241 hybridisation 246–248 in Japan 231, 232, 232 market for 231–233 in New Zealand 243–245, 249, 243, 244 nursery 235, 236, 241 plate culture 235, 236, 239, 241 production technology 233–245 in Southeast Asia 231–232, 232 spawning 235, 240 ‘surfboard’ tank system 242 ‘Taiwanese’ culture method 242 in USA 231, 232, 233 Acanthopagrus burcheri (black bream) 14 Acanthuridae (surgeonfishes) 278 Acantophora specifera 265 African catfish see Clarias gariepinus agar 252, 253, 259–260 Agardhiella ramosissima 265 Agardhiella subulata 264 Agardhiella tenera 265 Ahnfeltia plicata 265 Ahnfeltiopsis furcellata 265 All India Co-ordinated Research Project (AICR) on Composite Fish Culture Projects 336

alternative sites for aquaculture 11–16 see also site selection Amblypharyngodon mola 339 ammonia (in water) 2, 41–42, 139, 327, 373 Amphiprion akallopisos (Skunk clownfish) 279 Amphiprion akindynos (Barrier Reef clownfish) 279 Amphiprion bicinctus (Two-band clownfish/ Red Sea clownfish) 279 Amphiprion clarkii (Clark’s clownfish) 279, 283 Amphiprion ephippium (Flame clownfish) 279 Amphiprion frenatus (Tomato clownfish) 279 Amphiprion melanopus (Coral Sea clownfish/ Red & Black clownfish) 279, 283, 295 Amphiprion nigripes (Rose Skunk clownfish) 279 Amphiprion ocellaris ([Black] False Percula clownfish) 279, 283, 301, 307, 310 Amphiprion percula (True Percula clownfish) 279, 307, 308 Amphiprion perideraion (Pink Skunk clownfish) 279 Amphiprion polymnus (Saddleback clownfish) 279 Amphiprion rubrocinctus (Australian tomato clownfish) 279 Amphiprion sandaracinos (Orange Skunk clownfish) 279 Amphiprion sebae (Sebae clownfish) 279 Amphiprion tricinctus (Three-band clownfish) 279 Anabas testudeneus 339 Andhra Pradesh, aquaculture in 336, 337, 339, 341, 342, 342, 343, 348, 355, 360, 363 angelfishes 280–281, 291–292 anemonefishes (clownfishes) see Pomacentridae

Recent Advances and New Species in Aquaculture, First Edition. Edited by Ravi K. Fotedar, Bruce F. Phillips. © 2011 Blackwell Publishing Ltd. Published 2011 by Blackwell Publishing Ltd.

388

Index

antibiotics 10, 42, 124, 130, 146, 148, 156, 160, 193, 220, 359 Apogonidae (cardinalfishes) 279, 284–289, 285, 286, 287 Apogon compressus 286 Apogon cyanosoma 286 AquaMat® 141 aquarium trade 277–281 Argentina, aquaculture in 261, 262, 263, 265 argulosis 359 Argyrosomus japonicus (mulloway) 13, 14 Arripis georgiana (Australian herring) 15 Artemia salina (brine shrimp) 14, 15, 41 Arunachal Pradesh, aquaculture in 337, 341, 342, 343, 345 Ascophyllum nodosum 253–254 Asian sea bass see Lates calcarifer (barramundi) Assam, aquaculture in 334, 337, 341, 342, 342, 343, 346 Assessor flavissimus 280, 292 Atlantic salmon see Salmo salar Australia abalone culture in 231, 232, 240–243 freshwater ornamental aquaculture 5–10 ISW aquaculture in 11–15, 11, 14–15 slipper lobster culture 85, 86, 90–91 spiny lobster culture 26–27, 28–29, 30–33, 49–52, 61–62, 63, 65–67 Australian bass see Macquaria novemaculeata Australian Fresh Corporation Pty 90, 104 Australian herring see Arripis georgiana Australian southern rock lobster see Jasus edwardsii bag feeding (of carp) 357, 357 Bahamas, spiny lobster production 57 Balmian bugs 86 Balistidae (triggerfishes) 278–279 Banan prawn see Penaeus merguiensis Banggai cardinalfish see Pterapogon kauderni barramundi see Lates calcarifer Bangladesh, carp production 347, 348, 348 bass, hybridisation of 247 Belize, spiny lobster production 46, 57, 64 Best Management Practices (BMP) 377, 378 Bidyanus bidyanus (silver perch) 15 Bifurcaria bifurcate 253 Bihar, aquaculture production in 336, 337, 341, 342, 343 bio-engineering 370–376 biomonitoring 377

Bio–Mos® 150 black bream see Acanthopagrus burcheri black foot paua see Haliotis iris; paua black tiger prawn see Penaeus monodon blennies 280, 293 blue tilapia see Oreochromis aureus Brazil seaweed production 262, 263–264, 265 spiny lobster 27–28 brine shrimp see Artemia salina Briothamnion triquetrum 264 British Virgin Islands 59 brown algae (Phaeophyta) 252, 253; see also seaweed Bryothamnion triquetrum 265 bundhs 339, 340 butterfly koi 6, 7, 7, 8, 9 Callophyllis variegate 265 cardinalfishes see Apogonidae Caribbean, aquaculture 195, 261, 263, 264, 265 carrageenan 252, 253, 255, 263 carp, common see Cyprinis carpio carp, European see Cyprinis carpio carp, Japanese coloured see koi carp, ornamental see koi carp (India) cage culture 351 commercial production 355–359 culture systems 350–351 feeding 351, 352, 354, 355, 356–357, 357 fingerlings 349, 350, 354 harvesting 353, 353, 358 hatchery 340–341, 340 health management 358–359 large-scale production 353–355 manuring 352, 353–354 output by state 337, 338 pen culture 351 polyculture of (India) 324–366 pond culture 351, 352–353, 353, 355–357 seed production 339–350 semi-intensive production 353–354 small-scale production 352–353 table-sized production 350–251 transport of 350, 354 yields 358 see also individual states catfish, native see Clarias batrachus Catla catla (catla carp) 334, 336, 354, 360 Central Inland Fisheries Research Institute (CIFRI) (India) 336

Index

Central Institute of Fisheries Education (CIFE) (India) 362, 364 Central Institute of Freshwater Aquaculture (CIFA) (India) 346 Centropyge spp. 292, 295 Centropyge flavissimus (lemonpeel angelfish) 292 Centropyge loriculus (flame angelfish) 292 Ceramium rubrum 253 Chaetodontidae (butterflyfishes) 279 Chaetomorpha indica (‘green tide algae’) 258 Chanos chanos (milkfish) 16, 259 chaurs 335 Cherax albidus 247 Cherax rotundus 247 Chhattisgarh, aquaculture in 336, 337, 341, 342, 343 Chelactus cultrifer 93 Chile abalone production in 233–239, 236, 237, 238, 239 seaweed production in 253, 258, 265 China see People’s Republic of China ‘Chinese’ carps 336, 339 see also Ctenopharyngodon idella; Cyprinus carpio; Hypophthalmichthys molitrix Chinese mitten crab see Eriocheir japonica sinensis Chondracanthus canaliculatas 265 Chondracanthus chamissoi 265 Chondracanthus teedii 265 Chondrus candiculatus 265 Chondrus crispus 266 Chromosome set manipulation (molluscs) 245–246 Chrysiptera parasema (yellow-tail damselfish) 295, 297, 301 Cichlidae 318 see also Tilapia CIFA see Central Institute of Freshwater Aquaculture CIFE see Central Institute of Fisheries Education CIFRI see Central Inland Fisheries Research Institute circular tanks 33, 34, 39, 372–373 Cirrhinus mrigala (mrigal carp) 334, 336; see also carp Cladophora coelothrix ‘green tide algae’ 258 Clarias batrachus (native catfish) 334, 339 Clarias gariepinus (African catfish) 334, 339 clawed lobster (Homarus americanus) 89 cobia see Rachycentron canadaum 179–196

389

Codium fragile 254 Colombia, seaweed production 264 common carp see Cyprinis carpio composite fish culture 336 containerised production (crustaceans) 125, 128–130, 129 crabs see mud crabs; portunid crabs Crassostrea gigas 14 Crenarctus bicuspidatus 93 cryopreservation (of spermatozoa) 153, 203 Ctenopharyngodon idella (grass carp) 334, 336, 339 Cuba, aquaculture in 46, 56, 256–265 Cypho purpurascens 287 Cyprinis carpio (common or European carp) 5, 14, 334, 336, 339 Cystoseira baccata 253 damselfishes see Pomacentridae Delhi, aquaculture production in 337, 341, 343 Dicentrarchus labrax 259 Dictyota pfaffii 253 Digenea simplex 264 disease avoidance see disease control disease control (in aquaculture) 1–5 disease exclusion 2–4 disease resistance (in aquaculture) 1–5, 10 dottybacks see Pseudochromidae doughnut-shaped rearing tank 96, 97 drug resistance 148 see also antibiotics Durvillaea antarctica 265 ear-shell see abalone ecolabelling 378–381 Eisenia arborea 266 Elacatinus spp. see Gobiidae Enteromorpha intestinalis 259 Enteromorpha prolifica 254 Enteromorpha spp. 257, 258, 259, 265 environmental impact of aquaculture 368–378 Environmental Management Systems (EMS) 377, 378 Eriocheir japonica sinensis (Chinese mitten crab) 116 Eucheuma cottonii 253, 263 Eucheuma denticulatum 253 Eucheuma isiforme 264, 265, 266, 267 Eucheuma uncinatum 266 Eucheuma spp. 253, 264, 265 European carp see Cyprinis carpio ezo abalone see Haliotis discus hannai

390

Index

Finding Nemo 307 fishmeal, as aquaculture feed 58, 141, 185–186, 376, 377 floating cages 47, 52, 55 floating raft cultivation (seaweed) 268 flocs 139, 377 Fucus spp. 253 Galapagos slipper lobster 86 Gelidiella acerosa 265, 265 Gelidium 253, 265, 266 Gelidium chilense 265 Gelidium floridanum 265 Gelidium howei 265 Gelidium lingulatum 265 Gelidium rex 265 Gelidium robustum 265, 266 Gelidium serrulatum 265 ‘Genetic Improvement of Farmed Tilapia’ (GIFT) 319 genetic improvement 5, 152, 376 genetic markers 152–153 geographical information systems (GIS) 369– 370, 397 giant clam see Hippopus hippopus giant freshwater prawn see Macrobrachium rosenbergii giant sea bass see Lates calcarifer (barramundi) GIFT see ‘Genetic Improvement of Farmed Tilapia’ Gigartina canaliculata 266 Gigartina chamissoi 265 Gigartina skottsbergii 263, 265 GLOBALGAP 156 Goa, aquaculture in 337, 341, 342, 343 Gobiidae (gobies) 279, 280, 291–292, 299 gobies see Gobiidae Gobiodon spp. 291 Gobiosoma evelynae 300 GOC see groundnut oil cake goldfish 5 government policy on aquaculture 377–378 Gracilaria spp. 253, 254, 256, 258, 259, 263, 264–265, 265, 370 Gracilaria asiatica 260 Gracilaria birdiae 258, 259 Gracilaria caudata 259, 263 Gracilaria chilensis (red algae) 239, 253, 256, 259, 265 Gracilaria cliftonii 254, 260 Gracilaria cornea 264, 265 Gracilaria crassisima 265 Gracilaria domingensis 264, 265

Gracilaria fisheri 259 Gracilaria gracilis 259, 265 Gracilaria lemaneiformis 259, 260, 265 Gracilaria mamillaris 265 Gracilaria vermiculophylla 260 Gracilaria verrucosa 263 Gracilariopsis bailinae 259 Gracilariopsis tenuifrons 263, 265 grammas 280, 292 Gramma brasiliensis 292 Gramma loreto 292 grass carp see Ctenopharyngodon idella green algae (Chlorophyta) 252; see also seaweed greenback flounder see Rhombosolea tapirina greenlip abalone see Haliotis laevigata greenwater culture system in barramundi aquaculture 208 in cobia aquaculture 188 in marine ornamental culture 295 in prawn aquaculture 141, 155 grey mullet see Mugil cephalus 365 Gujarat, aquaculture in 337, 341, 342, 343 Gymnogongrus furcellatus 265 Gymnogongrus spp. 265 Haliotidae see abalone Halliotis see abalone Haliotis asinina 232 Haliotis australis 232 Haliotis discus 232 Haliotis discus hannai (Japanese or ezo abalone) 232, 234, 241, 247 Haliotis diversicolor 232 Haliotis fulgens 232 Haliotis iris 232 Haliotis kamtschatkana (pinto abalone) 232, 247 Haliotis laevigata 14, 232, 240, 241, 242, 248 Haliotis midae 232 Haliotis rubra 232, 241 Haliotis rufescens (red abalone) 232, 233, 234, 248 Haliotis scalaris 248, 248 Haliotis tuberculata 232, 233, 259 Haliotis virginea 243 Halymenia floresia 265 haors 335 hariwake (koi) 9 Haryana, aquaculture in 16, 336, 337, 341, 342, 343, 362,

Index

Heteropneutes fossilis 339 Himachal Pradesh, aquaculture in 337, 341, 342, 343, 345 Hippocampus abdominalis 303, 304, 305, 306 Hippocampus erectus 280, 303, 304, 305, 306 Hippocampus kuda 280, 293, 294, 304 Hippocampus reidi 280, 304, 306 Hippocampus trimaculatus 293, 303, 304, 305, 306 Hippocampus whitei 303–304 Hippopus hippopus 14 ‘hi utsuri’ (koi) 8 Homarus americanus see clawed lobster horizontal resistance 4 host resistance, breeding for 1, 2, 3–4 host resistance, development of see host resistance, breeding for hybridisation 246–248, 368 Hydropuntia cornea 265 Hydropuntia crassissima 265 Hypnea acerosa 265 Hypnea musciformis 263, 264, 265 Hypnea valentiae 265 Hypophthalmichthys molitrix (silver carp) 334, 336, 339 Ibacus ciliatus 93 Ibacus novemdentatus 93 Ibacus peronii 93, 97 Ibacus spp. 86, 93 ICAR see Indian Council of Agricultural Research immunity see disease resistance IMC see Indian major carps immunostimulants as feed additives in aquaculture 10–11 in prawn culture 146, 148–151, 156, 158 see also prebiotics; probiotics IMTA see integrated multi-trophic aquaculture; integrated aquaculture systems India aquaculture output 334, 335, 337–338, 350 cage culture 351 carp polyculture in 334–366 commonly cultured species 336–339 freshwater resources 335 giant freshwater prawn production 362–366, 363, 363 inland fish production 337–338, 351 ISW aquaculture in 16, 334 pen culture 351 pond culture in 335, 351

391

seed production 340–350, 341, 342, 343–345, 346, 347 slipper lobster in 85–86, 89, 91 spiny lobster in 28, 46, 53–54, 62, 64 traditional aquaculture 336 Indian Council of Agricultural Research (ICAR) 336 Indian major carps (IMC) 339; see also Catla catla; Cirrhinus mrigala; Labeo rohita Indonesia, spiny lobster production 46, 59 infectious hypodermal hematopoietic necrosis virus (IHHNV) 146 inland saline ground water 15–16 inland saline water (ISW), aquaculture in 11–16, 11, 14 integrated aquaculture systems 258, 259 integrated multi-trophic aquaculture (IMTA) 258, 259 ISGW see inland saline ground water ISW see inland saline water Jammu & Kashmir, aquaculture in 337, 341, 342, 343, 345 Japan, carp cultivation in 5–10 spiny lobster cultivation in 25–26, 33–39, 34, 36, 46, 65 Japanese abalone see Haliotis discus hannai Japanese coloured carp see koi Jasus edwardsii (Australian southern rock lobster) 23, 23, 26, 27, 29–30, 32–33, 41, 51–52, 60–61 Jasus lalandii (South African rock (spiny) lobster) 22, 23, 61 Jayanti rohu 346 Jharkhand, aquaculture in 336, 337, 341, 342, 343 Kailis, M.G., Pty Ltd 30 Kappaphycus alvarezii 253, 256, 258, 259, 262, 264, 266, 267 Karnataka, aquaculture in 337, 341, 342, 343 Kerala, aquaculture in 337, 341, 342, 343 ‘kin gin rin’scale trait 7, 7 King George whiting see Sillaginodes punctatus kohaku 8 koi 5–10 Kolleru Lake 355 ‘kombu’ see Laminaria japonica Kuruma prawn see Penaeus japonicus

392

Index

Labeo bata 339 Labeo calbasu 339 Labeo rohita (rohu carp) 334, 336, 339, 359 see also carp Labridae (wrasses) 279 Laminaria abyssalis 265 Laminaria brasiliensis 265 Laminaria digitata 253 Laminaria japonica (‘kombu’) 253, 254 Laminaria spp. 257, 265 Larnaea 359 Lates calcarifer (Asian sea bass, barramundi, giant sea bass) aquaculture 199–224 in Australia 199, 222 biology 200–202 breeding 202–207, 203 broodstock 202 cage culture 212 culture systems 207– 209, 208 diseases 216–221, 218 distribution 199–200 extensive culture 207 feeding see nutrition and growth global production 200 growout 211–212 habitat 200 hatchery 202–205, 203, 205, 206 health management 216–221, 218 in India 339 inland saline water (ISW) cultivation 12, 13, 14, 15 intensive culture 207, 208–209 juveniles 209–211, 210, 211 larvae 205, 206, 209 life history strategy 200–201 nursery phase 210–211 nutrition and growth 213–216 physiology 202 pond culture 212–213 post-harvest 221 quality 221 recirculating systems 212 sales and marketing 221–222 selective breeding 222–223 spawning 201, 202–205, 203, 205 Latin America, seaweed cultivation in 261–268 Laurencia papillose 265 LD cycles see photoperiod Lessonia nigrescens 265 Lessonia trabeculata 265 Lessonia vadosa 265

light–dark cycles see photoperiod light, effect on larval rearing slipper lobster 98, 99, 105 spiny lobster 38–39 lipopolysaccharides 10, 148 Litopenaeus stylirostris 143 Litopenaeus vannamei (white prawn) 13, 136, 139, 155, 157, 259 lobster see clawed lobster; slipper lobster; spiny lobster Lobster Harvest Pty Ltd 30, 68 long-fin koi see butterfly koi longfins 280 lysozyme 145 Macquaria novemaculeata 14 Macrobrachium rosenbergii (giant freshwater prawn; scampi) Bangladesh 348 claw ablation 264–265 culling and harvest 365 growout nursery pond management 364 in polyculture 334, 362–336, 363, 363 problems in culture 366 saline production 16, 365 stocking 365 Macrocystis integrifolia 265 Macrocystis pyrifera 239, 249, 256, 263, 265, 266 Madhuca longifolia (mahua) 352, 353 Madhya Pradesh, aquaculture in 336, 337, 339–340, 341, 342, 343, 362 Maharashtra, aquaculture in 336, 338, 339, 341, 342, 343 Mahatma Gandhi National Rural Employment Guarantee Act (MGNREGA) 350 mahua see Madhuca longifolia Malaysia, spiny lobster cultivation in 57 mangroves, aquaculture in 11, 119, 377, 388 mangrove crabs 119 Manipur, aquaculture in 336, 338, 341, 342, 344 marine diatom see Phaeodacrylum tricomutum marine ornamental fish see ornamental fish, marine Mastocarpus papillatus 265 Mastocarpus stellatus 253 mauns 335 Mazzaella laminarioides 265 Mazzaella membranacea 265 MBV see Monodon bacillus virus

Index

Meghalaya, aquaculture in 338, 341, 342, 344 Meristiella gelidium 265 metallic koi 6, 6, 9 see also koi methnolics 10–11 Mexico, spiny lobster cultivation in 60 seaweed production 262, 265–266, 265, 266, 267, 268 MGNREGA see Mahatma Gandhi National Rural Employment Guarantee Act microalgae see seaweed micronutrients (in aquaculture) 156, 184 microsatellites 152–153 milkfish see Chanos chanos Mizoram, aquaculture in 338, 341, 342, 344 Monodon bacilovirus (MBV) 3, 146 Monostroma spp. 154, 264, 265 Moreton Bay bugs (Thenus sp.) 30 see also Thenus orientalis Morone chrysops (white bass) 247 Morone saxatilis (striped bass) 247 MOS (mannan oligosaccharide) 61–62 Mourilyan virus (MoV) 146, 155 Mozambique tilapia see Oreochromis mossambicus mrigal carp see Cirrhinus mrigala mud crab (Scylla spp.) aquaculture 115–131 cannabalism in 124, 125, 127–128 containerised production 128–130, 129, 131 diet 117, 118–119, 122–123, 123, 125, 126–127 farming, suitability for 121–122 fattening 118–119 feeding see diet; fattening growout 117, 125, 126 growth 119–120, 120, 125–126 habits 119 hard-shell production 116 hatchery 122–124, 130–131 juvenile phase 124–125, 127–128, 128 larval rearing 122–124, 123 life history 121 moult 120, 126 nursery 124–125 polyculture with 117 pond rearing 117, 118, 127–128, 130 reproduction 120–121, 121 seed crabs 115, 116 soft-shell production 116, 117–118 in Southeast Asia 130 spawning 122 technological development in aquaculture 122–125

393

Mugil cephalus (grey mullet) 16 mulloway see Argyrosomus japonicus Nagaland, aquaculture in 338, 341, 342, 344 Namibia, aquaculture 48, 59, 232 Nannochloropsis oculata 295 National Fisheries Development Board (NFDB) (India) 346 natural fisheries, over-exploitation of 368 New Zealand abalone culture 243–245, 249, 243, 244 spiny lobster rearing in 27, 32, 52–53, 60 NFDB see National Fisheries Development Board Nile tilapia see Oreochromis niloticus Nishikigoi see koi nitrogenous waste see ammonia nodavirus see viral nervous necrosis nori (Porphyra) 253 offshore aquaculture, environmental impacts of 369 ogon see metallic koi olive barb see Puntius sarana Ompok pabda 339 Onchorynchus mykiss (rainbow trout) 14, 259 Onchorynchus kisutch 259 Ophicephalus spp. 345 Orchid dottyback see Pseudochromis fridmani Oreochromis 318 see also Tilapia Oreochromis aureus (blue tilapia) 318 Oreochromis mossambicus (Mozambique tilapia) 318 Oreochromis niloticus (Nile tilapia) 318–319, 319, 326–327 see also tilapia Orissa, aquaculture in 334, 338, 341, 342, 344 ornamental aquaculture, freshwater 5–10 see also entries for individual species ornamental fish, marine aquaculture 277–313 broodstock and eggs 281–293, 281, 285, 287, 288, 282, 310–211 broodstock conditioning 293–294 captive-bred 279–280 commercial production 277–278, 306–312 constraints in breeding, 278–279, juveniles 300–306, 301, 302, 303 larval culture 294–299 quality control 309–310 spawning 283, 287–288, 288 tank water quality 382

394

Index

oxytetracycline (OTC) 124 see also antibiotics ozonation 33, 43, 44, 66, 130 Pacific oyster see Crassostrea gigas paddy (rice) fields, aquaculture in 11 Pagrus auratus (snapper) 15, 15 palinurids see spiny lobster Panama, seaweed production 262, 263, 265 Pangasianodon hypophthalmus 359, 362 Pangasius sutchi (Thai catfish) 334, 339, 359 Pangasius culture 359–361, 361 Panulirus argus 23, 28, 34, 58–59, 62 Panulirus cygnus (western rock lobster) 14, 23, 23, 24, 26, 28, 49–50, 63 Panulirus echinatus 27, 28 Panulirus elephas 23, 24, 45 Panulirus homarus 23, 28 Panulirus japonicus (Japanese spiny lobster) 23, 25–26, 29, 33–34, 34, 36, 37–38 Panulirus laevicauda 27, 28 Panulirus longipes bispinosus 23 Panulirus ornatus 23, 26, 28, 30–32, 48–49, 52, 55, 61–62, 106 Panulirus regius 28 Panulirus stimpsoni 28 Paua 243–245, 243, 244 see also abalone; Haliotis australis; H. iris; H. virginea Penaeid prawns 136–161 captive breeding see domesticated programmes culture systems 140–143, 147 diet 139, 141–142, 145, 158 diseases 144, 151, 152, 157–158, 159 domesticated programmes 154–161 genetics 151–154, 158–161 habitat 137–138 immunological aspects 148–151, 158, 160 inbreeding 140 intensive culture 140–141 physiology 143 probiotics 148–150 reproduction 142–143 salinity 138–139 temperature 139, 140 water quality 139, 140, 141 WSSV in 139, 145–146 Penaeus esculentus 141 Penaeus japonicus (Kuruma prawn) 14, 138, 155, 259 Penaeus latisulcatus (western king prawn) 14, 140, 259

Penaeus merguiensis (banan prawn) 14 Penaeus monodon (tiger prawn, black tiger prawn, giant tiger shrimp) 2, 12, 13, 14, 106, 117, 136, 137, 155–156, 157, 339 Penaeus semisulcatus 138 Penaeus vannamei (white shrimp) 2, 15 People’s Republic of China, aquaculture in 5, 11, 28, 60, 116, 199, 231, 319 peptidoglycan (PG) (immunostimulant) 10, 148 Peru 265 Petractus demani 93 Petractus rugosis 91, 93 PG see peptidoglycan Phaeodacrylum tricomutum (marine diatom) 15 Philippines, spiny lobster production 46, 58 photoperiod, effects on rearing abalone 235 marine ornamentals 292, 294, 295, 300 seahorses 290 spiny lobster 28, 38–39, 44 Pinctada martensii 259 planktonkreisel 34, 36, 43, 45 plate culture (abalone) 235, 236, 239, 241 Polyploid induction (molluscs) 245–246 polysaccharides 10, 148 Pomacanthidae (angelfishes) 279 Pomacentridae (damselfishes and clownfishes) 278, 279, 279–280, 281–284, 300, 307 Porphyra spp. 254, 257, 263, 370 Porphyra acanthophora 263 Porphyra columbina 263, 265 Porphyra spiralis 263 Portugal, spiny lobster rearing in 28 Portunidae (swimming crabs) see mud crabs Portunus pelagicus 117, 126 Portunus spp. see portunid crabs prawn giant freshwater see Macrobrachium rosenbergii tiger see Penaeus monodon see also penaeid prawns prebiotics 10, 148, 150, 158 Premnas biaculeatus 283, 283 Prionitis decipiens 265 probiotics 10, 140, 148–150, 158, 160 Pseudochromidae (dottybacks) 279, 279, 286– 289, 287, 308 Pseudochromis fridmani (orchid dottyback) 287, 289, 300, 300 Pseudochromis flavivertex (sunrise dottyback) 289, 295, 297, 298 Pseudochromis steenei 288, 307

Index

Pterocladia pyramidale 265 Pterocladiella capillacea 264 Pterapogon kauderni (Banggai cardinalfish) 284, 285, 285, 286, 302 Pterapogon mirifica (Sailfin cardinalfish) 284, 285 Puducherry, aquaculture in 338, 341, 344 Punjab, aquaculture in 336, 338, 341, 342, 344 Puntius sarana (olive barb) 366 QX disease 4 raceway tanks 96, 97, 102, 104–106, 105, 237, 242, 372 Rachycentron canadaum (cobia) barriers to culture 195–196 cage culture 190, 195 commercial potential 179, 381 culture of 179, 186–191, 190, 194–195 diseases 191–194 distribution 181 egg production 186–187 feeding 182–183, 185–186, 187, 189, 195 growout 189–191, 190 growth 181, 185 hatchery 186–188 induced spawning 186–188 juvenile culture 189, 195–196 larval rearing 188–189, 188 marketing 194 morphology 179–181, 180 nutritional profile 194 nutritional requirements 183–186, 189 see also feeding parasites 191–193 post-harvest 194 reproduction 182 Taiwan, culture in 187, 194, 195–196 temperature, effect on 185 rainbow trout see Onchorynchus mykiss Rajasthan, aquaculture in 338, 341, 342, 344 Random Amplified Polymorphic DNA (RAPD) 246–247 rearing tank see tank recirculating aquaculture systems (RAS) 129–130, 129, 239370–376, 371, 372, 374, 375 red abalone see Haliotis rufescens red algae (Rhodophyta) 252; see also seaweed red drum see Sciaenops ocellatus Rhodoglossum denticulatum 265 Rhombosolea tapirina (greenback flounder) 14

395

Saccostrea commercialis 259 Saccostrea glomerata 4, 14 Sagmariasus verreauxi 23, 26, 27, 32–33 Salmo salar (Atlantic salmon) 14 Sarcothalia spp. 263, 265 Sarcothalia crispata 263, 265 Sargassum spp. 259, 263, 265 Sargassum vulgare 253 Sarotherodon 318 see also Tilapia scampi see Macrobrachium rosenbergii Sciaenops ocellatus (red drum) 15 Scylla paramamosain 117, 120, 121, 127 Scylla serrata 117, 119, 120, 121, 126–127 Scylla spp. 119, 120 see also mud crabs Scyllarides astori 101 Scyllarides latus 101 Scyllarides nodifer 101 scyllarids (Scyllarinae) see slipper lobster Scyllarus aequinoctialis 97 Scyllarus americanus 97 Scyllarus arctus 93, 97 Scytlarides 101 sea-cages 51, 107, 108 see also floating cages sea-ear see abalone seahorses see Syngnathidae sea snail see abalone seaweed agar extraction 259–260 aquaculture 254–268 biofilters, use as 257, 258, 259 bioremediation, use in 254, 257 in Chile 253, 258, 261, 262, in China 253 commercial uses 252–254 cultivation see aquaculture fertiliser, use as 253–254 fuel, use as 254 integrated aquaculture 257–259, 259 in Latin America 261–268 post-harvest 259–260 propagation methods 255–256 use as seed collector 48 in Philippines 253 Sebastodes fuscescens 259 ‘seed’ crabs 115, 116 selective breeding (for disease control) 1–2, 4 Siganus guttatus (rabbitfish) 295 Sikkim, aquaculture in 338, 341, 342, 344, 345 Sillaginodes punctatus (King George whiting) 14 Sillago ciliate (sand whiting) 14 Sillago schomburgkii (yellow-fin whiting) 15

396

Index

silver carp see Hypophthalmichthys molitrix silver perch see Bidyanus bidyanus Singapore, spiny lobster production 46, 56 site selection 369–370 slipper lobster aquaculture potential 88–89, 90 in Australia 85, 90–91, 104–106 biology of, 87–88, 88 earthen ponds culture 106, 107 environmental factors in rearing 97–98, 97, 100 growout 102–109, 105 hatching 99–100 in India 89, 91, 103, 106, 107, 108 indoor culture 103–104 juveniles, growth of 101–102 larval development in 92–95, 93, 94–100, 100 life history of 87–88, 88, 91–94 marketing 89–90 nutrition in 94–95 recirculatory cultivation system 103, 104 seed, rearing from 100 soft-shell production 105–106 tank design for 96–97, 96, 100 tank culture 103, 104, 104 in Vietnam 107, 108 snapper see Pagrus auratus South African rock (or spiny) lobster see Jasus lalandii South Africa, spiny lobster cultivation in 56–57, 61 soybean (as aquaculture feed) 141, 185, 186, 215, 376 Sparus aurata 259 specific pathogen free (SPF) stock 3, 154 SPF see specific pathogen free spiny lobster aquaculture 22–68, 23, 24, 88 in Australia 26–27, 29, 30–33, 61–62, 63, 65–67 in Belize & Bahamas 46, 57 in Brazil 27–28 broodstock management 28–30 coloration 50 culture systems for 34–35, 34, 35 diet of 29–30, 39–41, 49, 51–52, 58, 61 ecosystem considerations 63–64 environmental parameters 35–41, 49–51 feeding behaviour 40–41 fisheries, conflict with 63–64 growout of 46–68, 47

in India 28, 46, 46, 53–54, 62, 64 in Indonesia 46, 59 in Japan 25–26, 33–34, 34, 39 juveniles, rearing of 46–47 larval stage 29–43 life history in wild 22–24, 40 in Malaysia 57 in Mexico 60 metamorphosis 44–45, 45 in Namibia 59 in New Zealand 27, 32, 52–53, 60, 63 in People’s Republic of China 28 in Philippines 46, 58 in Portugal 28 pueruli, rearing of 46–47, 47 see also growout seed, collection of 48–49 in South Africa 56–57, 61 in Southeast Asia 48–49, 67 tank design for 33–35, 34, 35, 39, 43–44 in Taiwan 46, 57 transport, problems with 50 in USA 58–59 in Vietnam 46, 46, 48, 54–56, 55 wild stocks, reseeding 64–65 world production of 46, 46 Streptococcus iniae 219–221 Sulfitobacter 32 Sydney rock oyster see Saccostrea glomerata Syngnathidae (seahorses) 279, 280, 289–291, 302–306, 303 Taiwan cobia culture 187, 194, 195–196 spiny lobster production 46, 57 Tamil Nadu, aquaculture in 338, 341, 342, 344 tanks see circular tanks; raceway tanks; slipper lobster, tank design; spiny lobster Tapes philippinarum 259 Taura syndrome virus (TSV) 2, 4, 157 Thai catfish see Pangasius sutchi Thenus orientalis 85–86, 86, 90, 93, 94, 97, 98, 99–100, 99, 100 Thenus spp. 99–101, 99 see also slipper lobster tiger prawn see Penaeus monodon tiger shrimp, giant see Penaeus monodon Tilapia in Africa 318, 325, 326 aquaculture 318–326, 322 broodstock selection and management 320–321 cage culture 323, 323

Index

disease management 327, 328–329 environment 327 genetic improvement 326–327, 330 global production 329 harvesting 325–326 indoor culture 324–325, 324, 325, 330 marketing 327, 329, 331 nursery and harvesting 321–322 pond culture 322–323, 322 seed production 319–320, 320 in Vietnam 318–331, 320, 322, 323, 324, 325 Trachinotus carolinus 259 triploidy (in molluscs) 245–246 Tripura, aquaculture in 336, 338, 341, 342, 344 Trochus niloticus 14 Trochus see Trochus niloticus Turks & Caicos, spiny lobster 59 Ulva lactuca 259 Ulva reticulata 254 Ulva rigida 259 Ulva rotundata 259 Ulva spp. 257, 258, 259, 263, 265 Undaria pinnatifida 254 United States inland saline water prawn culture 15–16 marine ornamental aquaculture 278 spiny lobster rearing in 58–59 Uttarakhand, aquaculture in 336, 338, 341, 342, 344 Uttar Pradesh, aquaculture in 336, 338, 339, 341, 342, 344, 362

397

vaccination 2, 3, 144 vertically revolving tank 35, 37 Vibrios 4, 31, 32, 42, 95, 109, 138, 144, 146, 148, 157–158, 193 Vibriosis see vibrios Vietnam spiny lobster production in 46, 46, 48, 54–56, 55 tilapia production in 318–331, 320, 322, 323, 324, 325 viral encephalopathy and retinopathy (VER) see viral nervous necrosis viral nervous necrosis (VNN) 216–219 wakame (Undaria) 253 West Bengal carp culture in 334, 339, 341, 342, 344, 345 fish output 338 giant freshwater prawn production 363 western king prawn see Penaeus latisulcatus western rock lobster see Panulirus cygnus West Texas, prawn culture in 15 white food paua see paua; Haliotis virginea white prawn see Litopenaeus vannamei white spot syndrome virus (WSSV) 2, 3, 4, 10–11, 106, 139, 145–146, 151 wild stock enhancement 269 WSSV see white spot syndrome virus yabby 247 yellow-fin whiting see Sillago schomburgkii yellow head virus (YHV) 2, 3, 146 YHV see yellow head virus