Spiny lobster management

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SPINY LOBSTERS: FISHERIES AND CULTURE

SPINY LOBSTERS: FISHERIES AND CULTURE SECOND EDITION EDITED BY

B.F. PHILLIPS Curtin University of Technology, P . O . Box U1987, Perth, Western Australia 6845, Australia

J. KITTAKA Research Institute for Marine Biological Science, Research Institutes for Science and Technology, The Science University of Tokyo, Nemuro City, Fisheries Research Institute, Hokkaido 087-0166, Japan

Fishing News Books An imprint of Blackwell Science Blackwell Science

Copyright ( 2000 by Fishing News Books A division of Blackwell Science Ltd Editorial Offices: Osney Mead, Oxford OX2 OEL 25 John Street, London WClN 2BS 23 Ainslie Place, Edinburgh EH3 6AJ 350 Main Street, Malden, MA 02148 5018. USA 54 University Street. Carlton, Victoria 3053. Australia 10, rue Casimir Delavigne, 75006 Paris. France Other Editorial Offices: Blackwell Wissenschafts-Verlag GmbH Kunfurstendamm 57 10707 Berlin, Germany Blackwell Science KK M G Kodenmacho Building 7-10 Kodenmacho Nihombashi Chuo-ku, Tokyo 104, Japan The right of the Author to be identified as the Author of this Work has been asserted in accordance with the 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. First Edition published as Spiny Lobster Munugenwnt 1994 Second Edition 2000 Produced by and typeset in Times by Gray Publishing, Tunbridge Wells, Kent Printed and bound in Great Britain by MPG Books Ltd, Bodmin, Cornwall The Blackwell Science logo is a trade mark of Blackwell Science Ltd, registered at the United Kingdom Trade Marks Registry

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A catalogue record for this title is available from the British Library ISBN 0-85238-264-2 Library of Congress Cataloging-in-Publication Data Spiny lobsters : fisheries and culture / edited by B.F. Phillips and J. Kittaka. - 2nd ed. p. cm. Rev. ed. of: Spiny lobster management. 1994. Includes bibliographical references. ISBN 0-85238-264-2 (hardcover) 1. Lobster fisheries - Management. 2. Spiny lobsters. I. Phillips, Bruce F. 11. Kittaka, J. 111. Spiny lobster management. SH380 .S65 2000 639'.54dc2 1. 00-02937 1 For further information on Fishing News Books, visit our website: http://www.blacksci.co.uk/fnb/

Contents Preface

ix

B.F. PHILLIPS AND J. KITTAKA

List of Contributors Introduction: Ecology and Fishery Biology of Spiny Lobsters

xi 1

R.N. LIPCIUS and D.B. EGGLESTON

PART 1: FISHERIES: METHODS, MANAGEMENT A N D STATUS 1

The Status of Australia’s Rock Lobster Fisheries

43 45

B.F. PHILLIPS, C.F. CHUBB and R. MELVILLE-SMITH

2

New Zealand’s Rock Lobster Fisheries

78

J.D. BOOTH

3

Fisheries for Spiny Lobsters in the Tropical Indo-West Pacific

90

J.L. MUNRO

4

The Lobster Fishery in the North-western Hawaiian Islands

98

J.J. POLOVINA

5

The Commercial Fisheries for Jasus and Palinurus Species in the South-east Atlantic and South-west Indian Oceans

105

D.E. POLLOCK, A.C. COCKCROFT, J.C. GROENEVELD and D.S. SCHOEMAN

6

The State of the Lobster Fishery in North-east Brazil

121

A.A. FONTELES-FILIIO

7

The Cuban Spiny Lobster Fishery

135

J.A. BAISRE

8

The Atlantic Spiny Lobster Resources of Central America

153

N.M. EHRHARDT

9

The Spiny Lobster Fisheries in Mexico

169

P. BRIONES-FOURZAN and E. LOZANO-ALVAREZ

10

Status of the Fishery for Panulirus argus in Florida

189

J.H. HUNT

11

The French Fisheries for the European Spiny Lobster Palinurus elephas

200

H.J. CECCALDI and D. LATROUITE

V

vi

Contents

12

The Galapagos Spiny Lobster Fishery

210

R.H. BUSTAMANTE, G.K. RECK, B.I. RUTTENBERG and J . POLOVINA

13

The Spiny Lobster Fishery in Japan and Restocking

22 1

M. NONAKA, H. FUSHIMI and T. YAMAKAWA

PART 2: RESEARCH FOR MANAGEMENT: CASE STUDIES 14

Reproductive Biology: Issues for Management

243 245

C.F. CHUBB

15

Puerulus and Juvenile Ecology M.J. BUTLER IV and

16

276

W.F.HERRNKIND

Stock Identity of the Red (Jasus edwardsii) and Green (Jasus verreauxi) 302 Rock Lobsters Inferred from Mitochondria1 DNA Analysis J.R. OVENDEN and D.J. BRASHER

17

Spiny Lobster Catches and the Ocean Environment

32 1

B.F. PHILLIPS, A.F. PEARCE, R. LITCHFIELD and S. GUZMAN DEL PRO0

18

Measurement of Catch and Fishing Effort in the Western Rock Lobster Fishery

334

N. CAPUTI, C.F. CHUBB, N.G. HALL and R.S. BROWN

19

Predicting the Catch of Spiny Lobster Fisheries

357

B.F. PHILLIPS, R. CRUZ, N. CAPUTI and R.S. BROWN

20

Bioeconomic Modelling of the New Zealand Fishery for Red Rock Lobsters (Jams edwardsii)

376

P.A. BREEN, D.J. GILBERT and K. CHANT

21

Modelling for Management: The Western Rock Lobster Fishery

386

N.G. HALL and R.S. BROWN

22

The Artificial Shelters (Pesqueros) used for the Spiny Lobster (Panulirus argus) Fisheries in Cuba

400

R. CRUZ and B.F. PHILLIPS

23

The Use of Artificial Shelters (Casitas) in Research and Harvesting of Caribbean Spiny Lobsters in Mexico

420

P. BRIONES-FOURZAN, E. LOZANO-ALVAREZ and D.B. EGGLESTON

24

Recreational Spiny Lobster Fisheries - Research and Management

447

R. MELVILLE-SMITH, B.F. PHILLIPS and J. PENN

PART 3: AQUACULTURE AND MARKETING 25

Prospectus for Aquaculture J. KITTAKA and J.D. BOOTH

463 465

Contents 26

Maturation

vii 474

K. NAKAMURA

27

Breeding

48 5

A.B. MACDlARMlD and J. KITTAKA

28

Culture of Larval Spiny Lobsters

508

J. KITTAKA

29

Water Quality and Microflora in the Culture Water of Phyllosomas

533

M.A. IGARASHI and J . KITTAKA

30

Spiny Lobster Growout

556

J.D. BOOTH and J. KITTAKA

31

Diseases of Spiny Lobsters

586

L.H. EVANS, J.B. JONES and J.A. BROCK

32

Functional Morphology of the Digestive System

60 1

S. MIKAMI and F. TAKASHIMA

33

Nutrition and Food

61 1

A. KANAZAWA

34

Colour and Taste

625

S. KONOSU and K. YAMAGUCHI

35

Shipping

633

H. SUGITA and Y. DEGUCHI

36

Export Marketing of Australian and New Zealand Spiny Lobsters

64 1

R.N. STEVENS and D . SYKES

37

Marketing and Distribution in Japan

6 54

M. TSURUTA and J. KITTAKA

PART 4: PERSPECTIVES 38

Perspectives

665 667

B.F. PHILLIPS

Index

673

Preface Spiny lobsters, or rock lobsters as they are also known, are among the world’s most valuable and highly prized seafood. They are featured on menus from Japan to Europe. In addition, their size, abundance and position in the food web make them important ecologically. Fishing pressure on spiny lobster populations can be intense, leading to a need for wise management decisions. In this book, the management of spiny lobsters is addressed from biological and economic perspectives. Spiny lobsters are captured and marketed in over 90 countries. The world catch is currently in excess of 77 thousand metric tonnes (t) per year, with a landed value of approximately US $500 million. The principal producing countries are Australia, New Zealand, South Africa, Cuba, Brazil, Mexico and the USA, with over 70% of the catch coming from the Caribbean and south-east Atlantic region and the eastern Indian Ocean. The product is usually marketed frozen but the highest prices are obtained for live spiny lobsters, for which the Japanese are willing to pay more than US $100 per kilogram. Most spiny lobsters inhabit coral or rocky areas in shallow waters. This, combined with their large size and frequently dense populations, makes them important ecologically and relatively easy to study. Over the past 20 years we have learned a great deal about patterns of reproduction, migration, population growth and response to human exploitation in a number of species. In particular, an excellent picture of the ecology and fisheries biology of the Western Australian spiny lobster has been drawn, and research in Florida, Cuba, New Zealand and South Africa has begun to show similar patterns. Demand for spiny lobsters has escalated over the past two decades, spurring the need both for better management and for research on which to base that management. In Cuba, for instance, the catch rose from 500 t in 1965 to 13 500 t in 1985, but now oscillates between 8000 and 10 000 t. In South Africa, once the largest producer of spiny lobsters world-wide, the catch has dropped dramatically in recent years. In Western Australia, very strict management regulations and enforcement since virtually the beginning of the fishery has prevented overexploitation on the scale seen in other fisheries. A new and exciting development is now on the horizon and spiny lobster aquaculture appears to be a real possibility. Culture of the clawed North American lobster has been proposed and attempted for some 50 years, but is still not an economic reality. This, and the long larval life of spiny lobsters, has created an air of pessimism, resulting in few serious attempts to culture spiny lobster. However, recent developments in Japan and New Zealand have demonstrated the complete culture through the larval and puerulus stages. Australia is now investing in research into spiny lobser aquaculture and, although there continue to be problems to be ix

overcome, it may be only a few years before the major source of spiny lobsters is from aquaculture. We open the book with a brief review, by way of an introduction to spiny lobsters: general biology, types, distribution, fishing techniques, etc. Status reports of the major fisheries then follow, but the emphasis is on the latest management strategies, developments in aquaculture, marketing and economics. A special feature of the book is its reviews of the research activities and the marketing process in Japan. In the present book, we asked the authors to examine what was new and directly related to management, culture etc., and not to re-do the comprehensive Biology and Management of Lobsters (Cobb & Phillips, 1980, Academic Press). Thus, for instance, a chapter in this book on ‘Reproduction’ focuses only on those aspects of major importance to fisheries managers, and attempts to show how they are important. In the section on aquaculture, several authors examine maturation and breeding from that viewpoint. The authors of the book come from many parts of the world, and the varying approaches to science and writing styles are clear. The industry, the management and the science all have global connections. This is the first book that tries to bring the varied approaches together. We thank the authors for their efforts and hope that the communication fostered by their efforts will be rewarding. The book was first published in 1994 under the title of Spiny Lobster Management. It completely sold out, and there were demands for additional copies. Instead of simply reprinting the 1994 book, it was decided to revise all the chapters for a new edition. We have also taken the opportunity to include a chapter on recreational fishing and to add additional material on aquaculture, including the chapter on water quality and microflora (Chapter 29). The chapters on marketing have been replaced with new chapters which we feel are of greater relevance. In revising the chapters that we wrote as individuals, and in examining the chapters revised by the other authors, we were continually amazed at the changes that have occurred in the past 5 years. This completely vindicates the decision to revise all the chapters for the new edition and indicates the necessity for a ‘new’ book.

Acknowledgements Many people contributed to the development and production of this book. They are not acknowledged individually because of space availability, but all the authors wish to thank the many colleagues who assisted them with their contributions.

B.F. Phillips J. Kittaka

List of Contributors

J.A. Baisre Ministerio de la Industria Pesquera Barlovento Santa Fe 19 500 La Habana Cuba

J.D. Booth National Institute of Water and Atmospheric Research P.O. Box 14-901, Kilbirnie Wellington 6003 New Zealand D.J. Brasher 53c Devonshire Drive London SE 10 8 5 2 UK P.A. Breen National Institute of Water and Atmospheric Research P.O. Box 14-901, Kilbirnie Wellington 6003 New Zealand P. Briones-Fourzan Universidad Nacionai Autonoma de Mexico Instituto de Ciencias del Mar y Limnologia Unidad Academica Puerto Morelos Ap. Postal 1152 Cancun QR 77500 Mexico J. A. Brock Aquaculture Development Program Department of Agriculture Room 400 1177 Alakea Street Honolulu, Hawaii 968 13 USA

xi

xii

List of Coiztr.ihzrtor.v

R.S. Brown Bernard Bowen Fisheries Research Institute Western Australian Marine Research Laboratories P.O. Box 20 North Beach Western Australia 6020 Australia

R.H. Bustamante Charles Darwin Research Station Galipagos Islands Ecuador

M.J. Butler IV Department of Biological Sciences Old Dominion University Norfolk, VA 23529-0266 USA N. Caputi Bernard Bowen Fisheries Research Institute Western Australian Marine Research Laboratories P.O. Box 20 North Beach Western Australia 6020 Australia H.J. Ceccaldi Ecole Pratique des Hautes Etudes Centre d’Etudes des Ressources Animales Marines (CERAM) FacultC des Sciences et Technique de Saint-Jer6me Ave. Escadrille Normandie-Niemen F-13397 Marseille Cedex 20 France

K. Chant New Zealand Ministry of Economic Development P.O. Box 1473 Wellington 6001 New Zealand


C.F. Chubb Bernard Bowen Fisheries Research Institute Western Australian Marine Research Laboratories P.O. Box 20 North Beach Western Australia 6020 Australia A.C. Cockcroft Marine and Coastal Management Department of Environmental Affairs and Tourism P. Bag X2 Rogge Bay 8012 South Africa

R. Cruz Centro de Investigaciones Marinas Calle 16 144 entre Ave. lera y 3era Miramar Playa Ciudad de la Habana Cuba Y. Deguchi Department of Marine Science and Resources Nihon University Fujisawa Kanagawa 252-8510 Japan D. Eggleston Department of Marine, Earth and Atmospheric Sciences North Carolina State University Raleigh, NC 27695-8208 USA N.M. Ehrhardt Division of Marine Biology and Fisheries Rosenstiel School of Marine and Atmospheric Science University of Miami Florida, USA

...

xlll

xiv

List of' Contributors

L.H. Evans Aquatic Science Research Unit Muresk Institute of Agriculture Curtin University of Technology P.O. Box U1987 Perth Western Australia 6845 Australia A.A. Fonteles-Filho Instituto de Cigncias do Mar Universidade Federal do CearB Av. Da Aboliqdo 3207-Fortaleza CE 60165-081 Brazil

H. Fushimi Fukuyama University Sanzo Gakuen-cho Fukuyama Hiroshima 729-0292 Japan D.J. Gilbert National Institute of Water and Atmospheric Research P.O. Box 14-901, Kilbirnie Wellington 6003 New Zealand J.C. Groeneveld Marine and Coastal Management Department of Environmental Affairs and Tourism P. Bag X2 Rogge Bay 8012 South Africa

S. Guzman del Prbo Instituto Politecnico Nacional Escuela Nacional de Ciencias Biologicas Laboratorio de EcoligiB Marine Ap. Postal 26-375 02860 Mexico DF


N.G. Hall Bernard Bowen Fisheries Research Institute Western Australian Marine Research Laboratories P.O. Box 20 North Beach Western Australia 6020 Australia

W.F. Herrnkind Department of Biological Science Florida State University Tallahassee, FL 32306 USA J.H. Hunt Florida Department of Environmental Protection Marine Research Institute South Florida Research Laboratory 2796 Overseas Highway Suite 119 Marathon, FL 33050-2227 USA M.A. Igarashi Department of Fisheries Engineering University of Ceara Fortaleza-CE Brazil

J.B. Jones Fish Health Section Fisheries Western Australia P.O. Box 20 North Beach Western Australia 6020 Australia A. Kanazawa Faculty of Fisheries University of Kagoshima 4-50-20 Shimoarata Kagoshima 890-0088 Japan

xv

J. Kittaka Research Institute for Marine Biological Science Research Institutes for Science and Technology The Science University of Tokyo Nemuro City, Fisheries Research Institute Hokkaido 087-0166 Japan

S. Konosu 1-26-6 Kugahara Ota-ku Tokyo 146-0085 Japan D. Latrouite Institut Franqais de Recherches pour 1'Exploitation de la Mer Direction Ressources Vivantes, Ressources Halieutiques BP 70 F-29280 Plouzank France

R.N. Lipcius Virginia Institute of Marine Science The College of William and Mary Gloucester Point, VA 23062 USA R. Litchfield SIR Pty Ltd Sydney Australia

E. Lozano-Alvarez Universidad Nacional Autonoma de Mkxico Instituto de Ciencias del Mar y Limnologia Unidad Academica Puerto Morelos Ap. Postal 1152 Cancun QR 77500 Mt5xico


A.B. MacDiarmid National Institute of Water and Atmospheric Research P.O. Box 14-901, Kilbirnie Wellington 6003 New Zealand

R. Melville-Smith Bernard Bowen Fisheries Research Institute Western Australian Marine Research Laboratories P.O. Box 20 North Beach Western Australia 6020 Australia S. Mikami Australian Fresh Corporation c/o QDPI Bribie Island Aquaculture Research Centre P.O. Box 2066 Bribie Island Queensland 4507 Australia J.L. Munro International Centre for Living Aquatic Resources Management (ICLARM) Caribbean/Eastern Pacific Office Suite 158 Inland Messenger Service Road Town, Tortola British Virgin Islands K. Nakamura Faculty of Fisheries University of Kagoshima 4-50-20 Shimoarata Kagoshima-shi 890-0056 Japan

M. Nonaka Tokyo University of Fisheries 4-5-7 Kounan Minato-Ku Tokyo 108-8477 Japan

xvii

x vi ii List

of' Cont rihict ors

J.R. Ovenden Southern Fisheries Centre P.O. Box 76 Deception Bay Queensland 4508 Australia

A.F. Pearce CSIRO Division of Marine Research P.O. Box 20 North Beach Western Australia 6020 Australia J. Penn Bernard Bowen Fisheries Research Institute Western Australian Marine Research Laboratories P.O. Box 20 North Beach Western Australia 6020 Australia

B.F. Phillips Curtin University of Technology P.O. Box U1987 Perth Western Australia 6845 Australia

D.E. Pollock Marine and Coastal Management Department of Environmental Affairs and Tourism P. Bag X2 Rogge Bay 8012 South Africa J.J. Polovina Honolulu Laboratory Southwest Fisheries Science Center National Marine Fisheries Service, NOAA 2570 Dole Street Honolulu, Hawaii 96822-2396 USA


G.K. Reck Institute of Applied Ecology University of San Francisco of Quito Quito Ecuador

B.I. Ruttenberg School of Forestry and Environmental Studies Yale University New Haven, CT USA D.S. Schoeman Marine and Coastal Management Department of Environmental Affairs and Tourism P. Bag X2 Rogge Bay 8012 South Africa R.N. Stevens Western Australian Fishing Industry Council P.O. Box 55 Mount Hawthorn Western Australia 691 5 Australia

H. Sugita Department of Marine Science and Resources Nihon University Fujisawa Kanagawa 252-8510 Japan D. Sykes New Zealand Rock Lobster Industry Council P.O. Box 24901 Wellington New Zealand

xix

xx


F. Takashima Tokyo University of Fisheries 4-5-7 Konnan Minato-ku Tokyo 108-8477 Japan

M. Tsuruta Clean Bio Consulting Co. Ltd. Shirio-cho 438 Inba-gun Chiba-ken Japan K. Yamaguchi 1608, Hikawa Okutama Tokyo 198-0212 Japan T. Yamakawa Fisheries Research Institute of Mie Hamajima, Shima Mie 517-0404 Japan

SPINY L0BSTERS:FISHERIES AND CULTURE B.F. PHILLIPS&J. KITTAKA CoDvriaht 02000 bv Fishina News Books

Introduction

Ecology and Fishery Biology of Spiny Lobsters R.N. LIPCIUS Virginia Institute of Marine Science, The College of William and Mary, Gloucester Point, VA 23062, USA

D.B. EGGLESTON Department of Marine, Earth and Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695-8208, USA

Introduction Spiny (or rock) lobsters (Crustacea: Decapoda: Palinuridae) are ubiquitous in tropical and temperate seas (Fig. 1). Their value as a resource for food, for revenue, and for recreational and aesthetic value is undeniable. Spiny lobsters support some of the largest commercial fisheries in the world (Table 1), while also sustaining artisanal fisheries on remote islands or where they are in low abundance. Ecologically, palinurids are important links in marine food webs ranging from the deep ocean to shallow littoral habitats. In shallow coastal zones, palinurids are major predators of various benthic species (e.g. snails, clams and urchins) and important prey of larger predators (e.g. sharks and finfish). Their widespread occurrence and exploitation reflect the evolutionary and ecological success of the Palinuridae, and underscore the need for comprehensive ecological investigations and effective conservation strategies. In this chapter, an updated general overview is given of the biology, ecology and fisheries for palinurid lobsters, based on the earlier review (Lipcius & Cobb, 1994). Thus, the chapter serves as an introduction to the book, and as background information for subsequent chapters, which are up-to-date reviews of palinurid fisheries world-wide, and case studies dealing with key ecological and fisheries issues. This chapter is not exhaustive; rather, it gives the reader sufficient information to understand and appraise subsequent chapters. For additional information, the reader may consult the two-volume set on the biology and management of lobsters (Cobb & Phillips, 1980a, b), particularly those chapters dealing with palinurid fisheries and ecology (Bowen, 1980; Herrnkind, 1980; Kanciruk, 1980; Morgan, 1980; Phillips & Sastry, 1980; Phillips et al., 1980). Volume 43(11), 1986, of the Canadian Journal of Fisheries and Aquatic Sciences contains contributed papers from the 1985 International Workshop on Lobster Recruitment, and portrays advances in our understanding of lobster ecology and management, with emphasis on recruitment issues. Of the contributions in that volume, 12 are specifically devoted to palinurids. Distribution, identification and commercial trade of lobsters are detailed by Williams (1986, 1988) and Holthuis 1

2

Spiny Lobsters: Fisheries and Culture

Fig. 1 World-wide distribution of palinurid lobsters.

(1991). Cobb & Wang (1985) provide a comprehensive overview of the fisheries biology of clawed lobsters, spiny lobsters and freshwater crayfishes. More recent collections of spiny lobster biology and fisheries include the earlier edition of Spiny Lobster Management ( Ed. by B.F. Phillips, J.S. Cobb and J. Kittaka, 1994), as well as special issues of Crustaceana [Vol. 66(3), 1994] and Marine and Freshwater Research [Vol. 48(8), 1997].

Palinurid systematics, evolution and morphology Systematics and evolution The Palinuridae comprises over 47 species (Holthuis, 1991; George, 1997), of which about 33 species support commercial fisheries (Williams, 1988) (Tables 1 and 2). The family Palinuridae consists of decapod crustaceans in the superfamily Palinuroidea, which encompasses the Synaxidae (e.g. Palinurellus spp.) and the Scyllaridae (i.e. slipper lobsters), in addition to the palinurids (Williams, 1988; Holthuis, 1991; George, 1997). The Palinuroidea, along with the other lobster superfamilies (e.g. Nephropoidea, clawed lobsters), are contained within the suborder Macrura Reptantia, which consists of reptant (i.e. crawling) decapods, in contrast to the natant (i.e. swimming) decapods such as shrimp. Interestingly, Holthuis (1991) notes that the genera Panulirus and Linuparus are anagrams derived from Palinurus by White (1847) when he split the genus Palinurus into three genera. The palinurid

Hemisphere

N±S

Zone

Tropical

regius Panulirus spp.

East Africa, Indonesia New Guinea, East Africa Pacific Islands Thailand, India, Pakistan, SE Asia NW Africa Tropical

Geographical location

homarus ornatus penicillatus polyphagus

Jasus Caribbean, Florida, Brazil Ecuador, Panama Brazil

Palinurus

argus gracilis laevicauda

Panulirus

Table 1 Major fisheries for palinurid lobstersa

10 677

38 020

World catchb (mt)

14.3%

50.8%

Percentage of palinurid catch

Ecology and Fishery Biology of Spiny Lobsters 3

N

Subtropical

S

Hemisphere

continued.

Zone

Table 1

stimpsoni cygnus pascuensis

longipes marginatus

interruptus japonicus

inflatus

Panulirus

charlestoni delagoae

Palinurus

verreauxi

Jasus

East Australia, New Zealand

Hong Kong Western Australia Easter Islands Cape Verde Islands SE Africa, Mozambique

West Mexico, Guatemala, Honduras California Japan, South China Sea China, Japan Hawaii


3689

42 294c

11 450

1267

World catchb (mt)

4.9%

0.1% 0.4%

15.3%

1.7%


4 Spiny Lobsters: Fisheries and Culture

N±S

S

Temperate, subtropical

Temperate

Panulirus

b

Modified after Morgan (1980). Annual mean from 1991±1995 (FAO, 1997). c Mean from 1991±1993. c Mean from 1991±1993 and 1995.

a

Total palinurid catch

Hemisphere

continued.

Zone

Table 1

gilchristi

mauritanicus and elephas

Palinurus

St. Paul and New Amsterdam Islands Tristan de Cunha, St Helena

paulensis tristani

New Zealand Juan Fernandez, Chile SW Africa, Spain South Australia

West Africa, Mauritania, Mediterranean Basin, Western Europe South Africa


edwardsii frontalis lalandii edwardsii

Jasus

0.5% 100.1%

363 74 817

4.0% 0.1% 2.8%

1.3%

962d 3014 24 2129

3.9%


2886

World catchb (mt)

Ecology and Fishery Biology of Spiny Lobsters 5

6


Table 2 Palinurid species Genus Palinurus

Species

Other names

Location

Langouste

Cape Verde Island SW Indian Ocean NE Atlantic S South Africa

mauritanicus

Crayfish, spiny lobster Crawfish Gilchrist's crayfish, spiny lobster Langouste

argus cygnus

Florida spiny lobster, bug W Atlantic Western rock lobster W Australia

echinatus gracilis guttatus homarus homarus

Spiny lobster Blue lobster, langosta azul Spotted spiny lobster Green-scalloped rock lobster Deep-scalloped rock lobster Red-scalloped rock lobster

Central Atlantic Central E Pacific Caribbean Indian Ocean

inflatus interruptus japonicus laevicauda

Langosta Californian spiny lobster Ise-ebi Longosta

Mexico California Japan NE South America

longipes femoristriga

White-whiskered rock lobster Spotted-legged rock lobster Hawaiian lobster Ornate rock lobster

W Pacific

charlestoni delagoae elephas gilchristi

Panulirus

homarus megasculpta homarus rubellus

longipes longipes marginatus ornatus

W Arabian Sea SW Indian Ocean

Indian Ocean Hawaii Indo-West Pacific

pascuensis penicillatus

Longosta, crayfish Double-spined rock lobster

Easter Island Indo-West Pacific

polyphagus

Long-whiskered rock lobster Langouste royale

Indo-West Pacific

Hong Kong rock lobster Painted rock lobster Ryoma-ebi

South China Sea Indo-West Pacific Japan, Mauritius West Indies W Indian Ocean

regius

Justitia

E Atlantic

stimpsoni versicolor japonica longimanus mauritiana

E Atlantic

Ecology and Fishery Biology of Spiny Lobsters

7

Table 2 continued Genus

Species

Other names

Location

Jasus

caveorum edwardsii

Crayfish, rock lobster

Eastern South Pacific New Zealand

edwardsii frontalis lalandii paulensis tristani verreauxi

Linuparus

Palinustrus

Projasus

Cape crayfish, rock lobster Longouste Crayfish, rock lobster

S coast Australia Juan Fernandez W South Africa St Paul's Island Tristan da Cunha Tasman Sea, New Zealand

somniosus sordidus

E South Africa Australia, S China Sea

trigonus

Australia, Japan

mossambicus truncatus

E Africa Caribbean

waguensis Puerulus

Southern rock lobster

Wagu-ebi

Japan

angulatus carinatus

E Africa, New Guinea E Indian Ocean

sewelli velutinus

Arabian Sea Indonesia

bahamondei parkeri

SE Pacific Parker's crayfish

E South Africa

Adapted from Phillips et al. (1980).

lobsters are referred to by various common names. `Spiny' and `rock' lobsters belong to the same group of genera (i.e. the Palinuridae), but reflect different local traditions in naming. For consistency, we use `spiny lobster(s)' when referring to any of the palinurids. The classic studies by George & Main (1967) and George (1969) on evolutionary relationships in the Palinuridae were based on morphological characters; they divided palinurid lobsters into the Stridentes and Silentes. The four major morphological features included the relative size and disposition of the supraorbital processes, the elevation of the eyestalks, the structure of the pleopod on the second abdominal segment of the female and the general shape of the carapace. Coincident

8


with an evolutionary habitat trend from deeper to shallow waters, there was a morphological trend towards lateral separation and elevation of supraorbital processes, elongation and elevation of eyestalks, enlargement of the specified pleopod and rounding of the carapace, as exemplified in Panulirus. Presumably these features were adaptive in avoidance of predation in shallow, well-lit habitats, and to increase the effective area for aeration of the egg mass (George & Main, 1967). Within the palinurids, there appear to be two basic distribution patterns, one a circumpolar high-latitude pattern (e.g. Jasus), and the other a circumequatorial lowlatitude pattern (e.g. Panulirus). The highest diversity of palinurids occurs in the warm, shallow-water regions, probably as a result of the greater variety of habitats. Pollock (1990, 1992, 1993) examined the mechanisms producing speciation and a broad distribution in Jasus and Panulirus. The wide distribution of both genera was presumed to result from the long-distance dispersal capabilities of the teleplanic phyllosome larvae combined with circumoceanic gyral circulation routes. Speciation probably resulted from alterations in ocean current systems, particularly those associated with changes in sea level, and the emergence and subsidence of seabed ridges, rises and seamounts, such as those which occurred during the Pleistocene. In particular, the intensity and location of gyral flow, which tends to limit larval dispersal and thereby facilitates allopatric speciation, varied between glacial and interglacial periods and probably determined rates of speciation in palinurids (Pollock, 1993). Species integrity has probably been maintained by behavioural barriers to larval or post-larval recruitment, such that larvae and post-larvae are attracted to physical and chemical cues associated with the natal environment (Pollock, 1990, 1992). More recently, George (1997) has provided a thorough review of the evolution of Jasus and Panulirus (Fig. 2, Table 3). George postulates that the collective effects of tectonic plate movements, global changes in climate and oceanic currents, which influence larval transport, and alterations in habitat characteristics have promoted genetic differentiation and speciation in palinurids. `Some species responded to shifts in currents, some drifted apart with the continents, some adapted to remote islands and seamounts, some moved polewards as the climate cooled, some adapted to newly cretaed habitats and others became partially isolated by geographic barriers' (Fig. 2) (George, 1997). These conclusions regarding the evolution of palinurids are in keeping with our recent understanding of ecologically important forces, specifically the influential roles of recruitment processes (Phillips et al., 1994a, b; Lipcius et al., 1997; McWilliam & Phillips, 1997), habitat relationships (Herrnkind et al., 1994, 1997; Acosta & Butler, 1997) and spatial dynamics (Punt & Kennedy, 1997), as well as genetic relationships (Brasher et al., 1992; Ovenden et al., 1992, 1997; Ovenden & Brasher, 1994; Silberman et al., 1994).


9

Continental shelf fronting open ocean

Widespread equatorial

0

0

Isolated island and seamount

Restricted Sea

Riverinfluenced shelf

Fig. 2 Linkages between geographical location, shelf and oceanic circulation, and postulated larval transport in palinurids. Adapted from George (1997).

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Table 3 Ecological groupings for Jasus and Panulirus Group

Ecological type

Description

Species

A

Widespread equatorial

Panulirus: P. penicillatus, P. longipes subspecies, P. versicolor, P. homarus homarus, P. echinatus

B

Continental shelf fronting open ocean

Shallow, mostly clear water and low runoff on islands or continents. Large-scale oceanic dispersal of larvae. Deep (0±100 m), clear water and low runoff from land. Adjacent oceanic waters well defined by gyral, eddy or upwelling systems. Larvae retained within 1000±1500 km of coast.

C

Isolated island and seamount

D

Restricted sea

E

River influenced shelf

Deep (0±200 m) clear water and low runoff. Geographically positioned in a path of strong unidirectional currents. Larvae probably retained in back eddies and Taylor columns. Shallow, clear waters and low runoff. Enclosed by geographic boundaries. Larvae mostly retained within boundaries. Shallow, turbid water and heavy to moderate runoff. Larvae probably retained in local `estuarine' waters.

Panulirus: P. cygnus, P. japonicus, P. marginatus, P. interruptus, P. gracilis, P. h. rubellus, P. h. megasculpta, P. argus (Brazil) Jasus: J. verreauxi, J. verreauxi-NZ, J. edwardsii, J. e. novaehollandiae, J. lalandii Panulirus: P. pascuensis Jasus: J. tristani, J. paulensis, J. caveorum, J. frontalis

Panulirus: P. guttatus, P. argus (Caribbean), P. inflatus

Panulirus: P. polyphagus, P. laevicauda, P. regius, P. stimpsoni, P. ornatus

Adapted from George (1997).

Morphology Spiny lobsters are among the largest of crustaceans; the total body length sometimes attains 60 cm, as in the green rock lobster, Jasus verreauxi (Kensler, 1967; Holthuis, 1991). This body length is approximately equivalent to a carapace length of 24 cm; carapace length (i.e. the distance from the base of the supraorbital horns to the posterior edge of the carapace) is the usual quantitative measure of body length in spiny lobsters (Cobb & Wang, 1985). The body parts of a representative lobster are compartmentalized into a cephalothorax, which consists of the fused head and thorax, and an abdomen,


11

with their respective appendages (Fig. 3; Holthuis, 1991). The cephalothorax comprises 14 fused somites, each with a pair of appendages; the first six somites constitute the cephalon and the last eight the thorax (Fig. 3). Appendages on the cephalothorax include the eyes, which may be movable, reduced and immovable, or altogether absent (Holthuis, 1991); the antennae and antennules, which provide protection, mechanoreception and chemoreception (Ache & Macmillan, 1980; Zimmer-Faust & Case, 1982, 1983; Zimmer-Faust, 1991); the mouth parts, which include the mandibles, maxillae and maxillipeds; and five pairs of walking legs. The cephalothorax is encased dorsally by a carapace extending from the last thoracic somite to the eyes, sometimes forming a projecting rostrum beyond the eyes. Laterally, the carapace encloses the branchial chamber, which protects the gills. In some palinurids, an antennular plate carries spines and is formed into a soundemitting stridulatory apparatus in the Stridentes group of palinurids (e.g. Panulirus); both structures are useful taxonomically (George & Main, 1967; Holthuis, 1991).

Fig. 3

Morphology of a palinurid lobster. Adapted from Holthuis (1991).

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Those palinurids without a stridulatory apparatus are distinguished as the Silentes group. The ventral portion of the cephalothorax forms a sternum, which bears the gonopores at the bases of the third pair of pereiopods in females and the fifth pair in males (Fig. 3). Six separate somites comprise the abdomen, each protected by chitinous coverings on the dorsal, ventral and lateral (pleura) portions. The pleura are characteristically shaped or ornamented, and enclose the pleopods, which are used as swimmerets and form appendages on the first five abdominal somites (Fig. 3). The first two pairs of pleopods are often formed into copulatory organs in males, having a stiff style-like appearance, whereas in mature females the pleopods become setose to enclose the external egg mass (Lipcius & Herrnkind, 1987). Posteriorly, the sixth abdominal somite forms the tail fan, consisting of the heavily calcified uropods and the telson, which represents either a plate-like median appendage of the sixth somite or a seventh abdominal somite (Holthuis, 1991). The powerful abdominal musculature and blade-like structure of the tail fan effect the swift, backward escape response characteristic of many decapods (Cobb & Wang, 1985).

Life history and ecology General ecology Spiny lobsters inhabit temperate and tropical seas (Fig. 1), but most species and the highest abundances are found in the tropics (Holthuis, 1991). Habitats include the intertidal through the deep sea down to almost 3000 m depth (Fig. 4, Table 3), with shelter provided within substrates encompassing rocky crevices, mud and sand bottoms, and vegetated beds. Individuals usually seek shelter during daytime (Herrnkind, 1980; Cobb, 1981; Lipcius & Herrnkind, 1982; Cobb & Wang, 1985), although mating behaviour and mass migrations sometimes disrupt the normal diel rhythms in activity (Herrnkind, 1980; Lipcius et al., 1983; Lipcius & Herrnkind, 1985; MacDiarmid et al., 1991), which may be endogenously driven (Williams & Dean, 1989). Habitation of shelters is often communal, probably dictated by gregarious behaviour adapted for protection from diurnally active predators (Berrill, 1975; Cobb, 1981; Zimmer-Faust & Spanier, 1987; Eggleston & Lipcius, 1992). The aggregative behaviour may be driven by chemical (Zimmer-Faust et al., 1985) or visual stimuli, and may differ depending on the quality of available habitats (Trendall & Bell, 1989; Eggleston & Lipcius, 1992). Spiny lobsters are ecologically dominant because of the same characteristics which make them commercially valuable, i.e. their large size and abundance. Spiny lobsters prey nocturnally upon a diverse assemblage of benthic and infaunal species, including molluscs (e.g. snails and clams), smaller crustaceans, echinoderms, polychaetes (Fielder, 1965; Heydorn, 1969; Berry, 1971b; Herrnkind et al., 1975;


13

Fig. 4 Spatial distribution of palinurid lobsters by water depth, temperature and latitude. Adapted from George & Main (1967).

Pollock, 1978; Engle, 1979; Berry & Smale, 1980; Andree, 1981; Joll & Phillips, 1984; Edgar, 1990) and algae (Joll & Crossland, 1983), while concurrently serving as prey for larger predators including various species of finfish, sharks and octopus (Cobb & Wang, 1985; Herrnkind & Butler, 1986; Howard, 1988; Eggleston et al., 1990, 1992, 1997; Smith & Herrnkind, 1992). Commercially important palinurids do not overlap significantly in spatial distribution when examined as a function of depth and latitude (Fig. 4) (George & Main, 1967; Cobb & Wang, 1985), although congeners can coexist in the same habitats (e.g. Panulirus argus and P. guttatus in Caribbean reefs; Herrnkind & Lipcius, 1989). Jasus and Panulirus, the two shallow-dwelling genera, typically inhabit temperate and tropical habitats, respectively (Fig. 4, Table 3). The remaining genera in the Palinuridae inhabit deep-water habitats (Cobb & Wang, 1985). Berry (1971a) examined habitat use in palinurid lobsters off the east coast of southern Africa, where seven of the eight extant genera have been recorded (i.e. Panulirus, Palinurus, Jasus, Projasus, Palinustus, Puerulus and Linuparus). The five

14


species of Panulirus, which inhabit shallow areas down to 18 m depths, appear to be separable ecologically on the basis of water turbidity, temperature, depth and tidal range. The remaining genera are deep-water inhabitants and evidently separable on the basis of substratum type. For instance, Palinurus gilchristi occupies rocky habitats, whereas Palinurus delagoae occurs outside rocky areas. Similarly, Puerulus is captured on muddy bottoms, whereas Linuparus inhabits rocky areas. George (1974) also reviewed habitat utilization of Panulirus in the Indo-West Pacific, showing the characteristic separation of species on the basis of habitat features (Fig. 5). One of the most impressive and characteristic features of the palinurid life history involves long-distance migrations, sometimes conducted in dramatic en masse fashion (Herrnkind, 1980). Migrations are often seasonal and can occur inshore± offshore or alongshore, as in the mass migrations. For instance, Panulirus argus migrates en masse in various locations throughout the Caribbean region after the first cold front in autumn (Herrnkind & Cummings, 1964; Herrnkind, 1969, 1980). During the movement, lobsters migrate in queues (i.e. single-file lines) both day and night for 2±3 days, with up to 64 individuals in a single queue. Other species performing long-distance migrations include J. edwardsii along the south-eastern New Zealand coast (Street, 1971), J. verreauxi in the North Cape region of New Zealand, Panulirus ornatus in the Gulf of Papua and Torres Strait (Moore & MacFarlane, 1977) and Panulirus cygnus during the exodus of `white' juveniles to deeper waters (George, 1958; Chittleborough, 1970; Phillips, 1983). These migrations

Fig. 5 Habitat specialization in palinurid lobsters. Adapted from George (1974).


15

are variously associated with reproduction (P. ornatus and Jasus), redistribution of juveniles to adult habitats (P. cygnus and, possibly, P. argus), or avoidance of physically stressful environmental conditions (P. argus). Spiny lobsters have been implicated as key predators in a variety of benthic habitats (Tegner & Levin 1983; Edgar, 1990), and their selective predation is apparently responsible for profound effects on species composition and sizefrequency distributions of invertebrates such as sea urchins, mussels and gastropods (Griffiths & Seiderer, 1980; Tegner & Levin, 1983; Joll & Phillips, 1984; Barkai & McQuaid, 1989; Edgar, 1990). Given the extended, nocturnal activity patterns of lobsters on the foraging grounds, it is likely that lobsters and other reef associates have a significant effect on benthic community structure and a negative effect on secondary production of certain benthic species. Predator impacts on benthic macrofauna may also vary seasonally with lobster abundance, and probably diminish with distance from shelter sites (Joll & Phillips, 1984; Jernakoff, 1987; Jernakoff et al., 1987). In addition, the effects of predator removal (i.e. spiny lobsters) through exploitation may impact the commmunity structure of ecosystems directly or in a cascading fashion (e.g. by removal of a dominant competitor), particularly if the spiny lobster is a keystone predator (sensu Paine, 1966). In contrast, role reversals can occur wherein prey of spiny lobsters inhibit the persistence or establishment of populations of spiny lobster. For instance, where rock lobsters (Jasus lalandii) are abundant in South African waters they control the abundance and species composition of mussels and whelks; in other habitats whelks are able to kill and consume immigrant rock lobsters, effectively preventing the establishment of a local population (Barkai & McQuaid, 1988). Large, transient environmental impacts to ecosystems that support spiny lobsters can also result in mass mortality. For example, periodic coastal upwelling and anoxia along the Namibian and South African coast result in mass mortality of Jasus lalandii (Grobler & Noli-Peard, 1997). Hypoxia in this same region appears to limit movement and growth rates of J. lalandii (Grobler & Noli-Peard, 1997).

Life history The palinurid life history reflects the predominant developmental pattern in marine crustaceans, i.e. release of a pelagic larva after embryonic development (Sastry, 1983a). Whereas most crustaceans with a pelagic larval stage release a nauplius larva (Sastry, 1983a), the palinurids pass the naupliar phase within the egg before hatching the distinctive phyllosome larvae. The pelagic larval phase is one component of a life history which, similarly to most crustaceans, does not easily conform to evolutionary theory incorporating r- and K-selection (Sastry, 1983b). Rather, the palinurid life history incorporates aspects of both r- and K-selection, including r-selected characters such as a large number of offspring and absence of parental care, and K-selected features such as delayed maturity and iteroparity (Sastry, 1983b). These

16


compromises are apparently in response to the differing pressures exerted on various phases of the life history, such as food or shelter limitation, predation pressure and physical transport processes. Spiny lobsters exhibit five major phases within the life cycle: adult, egg, phyllosoma (larval stages), puerulus (post-larval stage) and juvenile (Phillips et al., 1980), although the juvenile phase has recently been separated into an algal or early benthic phase (Marx & Herrnkind,1985a; Herrnkind & Butler, 1986) followed by the characteristic older juvenile phase, similar in habits to adults (Fig. 6). Adults frequently aggregate during the day in crevices of coral and rocky reefs (Berrill, 1975; Cobb, 1981; Zimmer-Faust & Spanier, 1987). At sunset spiny lobsters emerge from their dens to forage nocturnally in nearby habitats such as reef flats and seagrass beds (Herrnkind et al., 1975; MacDonald et al., 1984). Shortly before reaching adulthood many spiny lobsters undergo movements from the nursery habitat to the characteristic deeper reef habitats (Herrnkind, 1980), where reproduction occurs (Lipcius, 1985). Fertilization is external (Cobb & Wang, 1985), even in Jasus (MacDiarmid, 1988); the male deposits a spermatophoric mass on the female's sternum. The spermatophore is subsequently rasped several hours prior to spawning, thereby releasing sperm for fertilization of the eggs as they are extruded on to the abdomen and pleopods (Lipcius & Herrnkind, 1987). Egg masses are generally spawned and hatched in the spring and summer by females located in offshore reef areas. Subsequently, the early phyllosoma (larval) stages are transported offshore by wind-driven surface currents into oceanic habitats (Phillips & McWilliam, 1986).

Larval ecology Recent studies in marine systems have stressed the importance of recruitment processes in the population dynamics of marine species (Gaines & Roughgarden, 1987; Doherty & Fowler, 1994; Lipcius et al., 1997; Dixon et al., 1999). Interannual fluctuations in populations of benthic invertebrates and demersal fishes depend on recruitment and post-settlement dynamics, and are regulated by planktonic larval and post-larval availability, settlement rates in suitable habitats, and post-settlement movements and mortality rates (Connell, 1985). Settlement rates further vary as a function of hydrodynamics, larval density, delivery and mortality rates, likelihood of settlement due to habitat quality, and settlement behavior (Lipcius et al., 1990; Herrnkind et al., 1994). Because oceanographic processes influence the larval and post-larval phases in different ways, and because most commercially important species have open populations, such a synthesis for palinurids and other marine species requires an approach that integrates the fields of meteorology, oceanography (physical, chemical and biological), marine ecology and fisheries biology. Along with coral and slipper lobsters, palinurids are the only decapod crustaceans possessing phyllosoma larvae in the life history (Phillips & Sastry, 1980). Phyllosomes (derived from the Greek phyllos, a leaf, and soma, a body) are


Fig. 6

17

Schematized life cycle of palinurid lobsters.

dorsoventrally flattened, transparent and leaf-like larvae adapted for passive horizontal transport assisted by vertical migration. There are usually seven to 13 phyllosoma stages, with each representing one or more instars (Phillips & Sastry, 1980); sometimes a pre-phyllosome stage has been described, and is variously termed the `naupliosoma', `prenaupliosoma' and `prephyllosoma'. The pre-phyllosome stage lasts for no more than a few hours before ecdysis to the first phyllosoma stage; thereafter, larval durations differ greatly among palinurids, ranging from a few

18


months to almost 2 years before the last phyllosoma stage, approximately 35 mm in length, metamorphoses to the puerulus stage (Phillips & Sastry, 1980). Such variation is particularly evident in the cultured larvae of palinurids (Kittaka, 1994), such that the larval phase (i.e. the time from hatching until metamorphosis to the puerulus) encompassed 132 days in Palinurus elephas (Kittaka & Ikegami, 1988), 306 days in Jasus lalandii (Kittaka, 1988) and 340±91 days in Panulirus japonicus (Kittaka & Kimura, 1989). Studies of larval transport of palinurids have recently begun integrating oceanic or coastal circulation patterns with larval distribution (Fig. 2, Table 3) (George, 1997) and post-larval settlement patterns (Johnson, 1971; Phillips & McWilliam, 1986; Pearce & Phillips, 1988; Herrnkind et al., 1994; Lipcius et al., 1997; Eggleston et al., 1998). Quantitative information on the vertical migrations of larvae is essential to an understanding of transport mechanisms, larval sources, and recruitment processes in palinurids (Phillips & McWilliam, 1986; Yeung & McGowan, 1991). The importance of vertical migration in phyllosome transport is apparent in the larval behaviour of Panulirus cygnus (Rimmer & Phillips, 1979). The vertical migration behaviour of early phyllosomes (stages I±III) of P. cygnus places them near the surface. There they are transported offshore into the south-eastern Indian Ocean via surface wind drift at a speed of about 5.25 km/day (Phillips et al., 1979). This direction opposes the general circulation of the upper 300 m layer, which flows towards the coast of Western Australia at about 3.3 km/day. Mid- and late-stage P. cygnus phyllosomes (stages IV-IX) avoid the surface layer through an increased sensitivity to light (Rimmer & Phillips, 1979); thus, they become more subject to subsurface circulation, which returns them to the coast of Western Australia (Phillips, 1981). A different pattern exists for larvae of the Caribbean spiny lobster, P. argus, in south-eastern Florida, although the data are not extensive. In general, phyllosomes were distributed in the surface layer irrespective of larval developmental stage (Yeung & McGowan, 1991). However, there appear to be regional differences in ontogenetic vertical distribution patterns of phyllosomes, as well as the extent of local retention of spiny lobster larvae. Recently, P. argus phyllosomes, including late stages, were found in the coastal waters of south-west Cuba, where a gyre circulation persists (Yeung & McGowan, 1991). This gyre may retain phyllosome larvae locally, and thereby enhance the return of pueruli to south-western Cuba, probably resulting in the densest population of P. argus in the Caribbean. In contrast, on the northwestern coast of Cuba there is no gyre, but rather a rapid north-flowing current with low larval abundances. Off the Florida Keys, the Pourtales Gyre does not persist long enough (i.e. approximately 1±2 months) to retain phyllosomes during the entire larval development. In Exuma Sound, Bahamas, gyral flow apparently drives the spatial distribution of larvae and settlement patterns of post-larvae of the Caribbean spiny lobster (Lipcius et al., 1997). Thus, patterns of larval retention in palinurids apparently vary both within and between species, and require extensive ecological information (e.g. physical transport processes in oceanic habitats) to determine the key sources of variation in the larval phase.


19

Post-larval ecology After 6±24 months in the plankton, the last planktonic larval stage metamorphoses into the puerulus, a transparent, free-swimming, non-feeding post-larval stage that moves inshore where it settles to the benthos (Phillips & Olsen, 1975; Phillips, 1981; Calinski & Lyons, 1983; Nishida et al., 1990). Metamorphosis is probably dictated by the nutritional state of the phyllosoma, rather than by environmental cues (McWilliam & Phillips, 1997). Pueruli apparently navigate to the shallow-water juvenile nurseries by means of a complex receptor system formed by the antennae and a pinnate setal system, which enables orientation to cues associated with coastlines (Phillips & Macmillan, 1987). Once in the nursery grounds, pueruli prefer to settle in architecturally complex habitats, such as in the red alga Laurencia by post-larval P. argus (Marx & Herrnkind, 1985a; Herrnkind & Butler, 1986; Herrnkind et al., 1994), rocky crevices by J. edwardsii (Booth, 1979), and small holes near algae by P. japonicus (Yoshimura & Yamakawa, 1988) and P. interruptus (Serfling & Ford, 1975). The preference for these habitats is mediated by their structural complexity and not by food availability, which becomes influential for the post-puerulus juveniles (Herrnkind et al., 1988, 1994). Floating, artificial settlement substrates have proven useful in quantifying postlarval settlement patterns in several species, including Panulirus argus (Witham et al., 1968; Little, 1977; Little & Milano, 1980; Marx, 1986; Bannerot et al., 1991; Forcucci et al., 1994; Acosta et al., 1997; Eggleston et al., 1998), P. cygnus (Phillips, 1972, 1986), P. japonicus (Nonaka et al., 1980), P. marginatus (MacDonald, 1986) and J. edwardsii (Booth, 1979, 1989; Booth & Tarring, 1986; Booth & Bowring, 1988; Breen & Booth, 1989; Hayakawa et al., 1990; Booth et al., 1991). In the central Bahamas, post-larval settlement on `Witham-type' artificial settlement substrates was strongly correlated with the concentration of pueruli in the water column, and flux past a given point (Eggleston et al., 1998). In Florida Bay, post-larval settlement on modified `Witham-type' artificial settlement substrates correlated with planktonic abundance and settlement of postlarvae at regional scales of tens of kilometres, but not at local scales of tens of metres (Butler & Herrnkind, 1992). Abundance of P. argus juveniles in Florida Bay also correlated with post-larval supply, as measured with artificial settlement substrates, to the region 8 months earlier (Forcucci et al., 1994). Quantitative data from settlement on artificial settlement substrates have proven useful in successfully predicting future population size and fishery exploitation (Morgan et al., 1982; Phillips, 1986; Breen & Booth, 1989). Although shelter is generally thought to limit the abundance of early benthic phase lobsters in some areas (Ford et al., 1988; Butler et al., 1995), post-larval settlement may affect local abundance of juvenile lobsters. For example, a high post-larval supply of P. argus to the Middle Florida Keys, USA, fortuitously coincided with a massive dieoff of sponge refugia which, when combined with the availability of alternative, previously underused shelter (solution holes, coral heads, etc.), offset any major negative effects of sponge loss on juvenile population size (Butler et al., 1995;

20


Herrnkind et al., 1997). Further investigations are necessary to delineate the respective roles of pre- versus post-settlement processes on population abundance and structure. Studies that have used artificial settlement substrates to measure relative rates of settlement generally show that spatiotemporal variation in settlement patterns are often driven by distinct periodic (e.g. lunar phase, seasonality) and stochastic (wind speed and direction, coastal sea-level, etc.) factors. For example, post-larval influx of P. argus typically occurs during nocturnal flood tides during the first quarter of the lunar phase, with consistently highest settlement observed during the autumn (September±November) (Acosta et al., 1997; Eggleston et al., 1998). New-moon spring tides provide the potential for relatively strong tidal transport owing to increased tidal ranges and because recruitment during the darkest phase of the lunar cycle could reduce susceptibility to visual predators (Acosta, 1997). Seasonal differences in post-larval influx may reflect seasonal variation in current patterns that could influence larval and post-larval retention and advection, as well as seasonal spawning (Eggleston et al., 1998). In terms of stochastic factors influencing post-larval supply, wind-induced along-shore transport explained approximately 30% and 50% of the variation in settlement anomalies of P. argus in the Florida Keys and central Bahamas, respectively (Acosta et al., 1997; Eggleston et al., 1998). The unexplained variation (50±70%) in post-larval influx of P. argus to the Florida Keys and central Bahamas could be due to a variety of biological and physical factors, including local and regional patchiness of pueruli, variable larval developmental rates and vertical migration behaviour of larvae, regional scale meteorological influences on water levels, and the strength of coastal upwelling and down-welling. Post-larval influx in the western Australian rock lobster (P. cygnus) appears to be driven primarily by stochastic factors that may vary periodically over decadal or longer time scales. For example, post-larval influx of P. cygnus is correlated with the Southern Oscillation Index, coastal sea level, sea surface temperature, salinity and rainfall, as an index of storms (Phillips et al., 1991). In P. cygnus, post-larval settlement is high in years when El NinÄo Southern Oscillation (ENSO) events are minimal, resulting in a strong Leeuwin current which transports pueruli towards coastal nursery habitats (Pearce & Phillips, 1988). Depth of settlement is generally in the shallows, although pueruli can settle in deep habitats, potentially enhancing encounter rates with suitable habitats (Booth et al., 1991; Heatwole et al., 1991).

Juvenile ecology Once settled, the puerulus metamorphoses into the first benthic instar approximately 6±7 mm in carapace length (CL). Juvenile spiny lobsters can exhibit up to three ecologically distinct phases following settlement: algal phase, post-algal phase, and subadult. For instance, algal phase (i.e. early benthic phase, sensu Wahle & Steneck,


21

1991; 5±15 mm CL) P. argus typically reside solitarily in or under large clumps of intricately branched red algae (Marx & Herrnkind, 1985a), which provide food and refuge (Marx & Herrnkind, 1985b; Herrnkind & Butler, 1986; Herrnkind et al., 1988). Similarly, early benthic phase P. cygnus occupy small holes on the face, in ledges and in caves on coastal limestone reefs, particularly holes with associated seagrass or algae (Jernakoff, 1990). Young juvenile P. interruptus shelter in Phyllospadix seagrass beds with dense cover over rocky bottom (Parker, 1972; Serfling, 1972; Engle, 1979) and early benthic phase P. japonicus reside in small crevices in rocks or algal clumps (Yoshimura & Yamakawa, 1988). As juvenile lobsters reach a size of 20±45 mm CL, they begin to move out of algal clumps to small crevices in algal-covered rock rubble (Andree, 1981), before becoming gregarious with larger juvenile lobsters during the day under crevices provided by rocks, sponges, octocoral or other structures (Herrnkind et al., 1975; Andree, 1981; Marx & Herrnkind, 1985a; Herrnkind & Lipcius, 1989; Forcucci et al., 1994; Butler & Herrnkind, 1997). For P. argus, this type of habitat may be rare in many areas of the Caribbean, where fringing mangroves (Acosta & Butler, 1997) with associated margins of seagrass, macroalgae and crevice shelters (Lipcius et al., 1998) function as nursery habitat for juvenile lobsters. At night, lobsters forage on small molluscs and crustaceans in the surrounding habitat (Andree, 1981; Herrnkind et al., 1994; Cox et al., 1997). Small, post-algal lobsters roam only metres from their daytime shelter (Herrnkind & Butler, 1986; Yoshimura & Yamakawa, 1988), but become nomadic and forage widely at about 45 mm CL (approximately 1 year post-settlement) (Herrnkind, 1980, 1983). As maturity approaches, approximately 2 years postsettlement (75 mm CL), spiny lobsters migrate to the reef tract. Size-dependent shifts in sociality have been identified for P. argus (Marx & Herrnkind, 1985), P. cygnus (Phillips et al., 1977), P. interruptus (Zimmer-Faust & Spanier, 1987), J. edwardsii (Macdiarmid, 1994) and P. ornatus (Dennis et al., 1997), but the mechanisms for such shifts are relatively poorly studied. Variation in the behaviour of post-algal lobsters may reflect the niche shift from full-time algal dwelling to diurnal crevice sheltering. For post-algal P. argus, however, there is no difference in predation risk between algal and crevice habitats, which may have explained the size at which this shift in habitat and sociality should occur (Childress & Herrnkind, 1994, 1997). The ontogenetic shift from solitary to gregarious shelter use by post-algal P. argus appears to be mediated by a size-dependent shift in receptivity to chemical attractants released by conspecifics, as well as by a massdependent release of the attractant leading to scale-dependent attraction (Ratchford & Eggleston, 1998). For example, algal-phase P. argus (100 km in Exuma Sound, Bahamas. Adult abundance was lowest at a site with the highest post-larval supply and little macroalgal settlement habitat; hence, it was tentatively classified as a sink. High post-larval supply was due to a large-scale gyre that appeared to concentrate and advect post-larvae towards the nominal sink. The remaining three sites, including one marine reserve, had higher adult abundances despite lower post-larval supply, and were therefore tentatively classified as sources. It appears that some sites with suitable settlement and nursery habitat are sources of spawning stock for P. argus in Exuma Sound, whereas others with poor habitat are sinks despite sufficient postlarval influx (Lipcius et al., 1997). In Australia, Ford et al. (1988) found that lobster survival was higher on reefs where lobster densities were reduced than on control reefs; however, the caveat remains that migration rates confounded interpretation of the results. Field experiments and examination of stage-based population abundance and habitat measurements across hydrodynamically realistic spatiotemporal scales, coupled with population modelling, appear requisite to the delineation of the ecological processes regulating population size of juvenile spiny lobsters. Moreover, similar to investigations with the post-larval phase, quantitative indices based on juvenile abundance may be useful in predicting population abundance and fishery catch (Chittleborough & Phillips, 1975; Caputi & Brown, 1986; Phillips, 1986; Breen & Booth, 1989; Cruz & Phillips, 1994).

24


Adult ecology There is considerable variation in the adult segment of the life cycle among species of palinurids (Berry & Heydorn, 1970; Aiken & Waddy, 1980; Lyons et al., 1981; Cobb & Wang, 1985), particularly in the size-specific relationships between moulting, mating and egg extrusion (Lipcius, 1985; MacDiarmid, 1989a). Courtship and mating appear to be controlled to a large degree by a receptive female's choice of a suitable male partner, particularly larger individuals (Lipcius et al., 1983; Lipcius & Herrnkind, 1985) and not so much by the male's ability to force a female to copulate (Berry, 1970; Silberbauer, 1971; McKoy, 1979; Lipcius & Herrnkind, 1985). Intermale aggression also influences mate choice, whereby larger males inhibit smaller male J. edwardsii from courting females (MacDiarmid, 1989b). Mature males characteristically moult well before the mating period to be in intermoult and fully hardened in preparation for mating (Lipcius, 1985). Fertilization is external (Cobb & Wang, 1985), even in Jasus (MacDiarmid, 1988), whereby the male deposits a spermatophoric mass on the female's sternum; the structure is subsequently rasped several hours prior to spawning to release the sperm for fertilization of the eggs as they are extruded onto the abdomen and pleopods (Lipcius & Herrnkind, 1987). Egg masses are generally spawned and hatched in the spring and summer by females located in offshore reef areas (Lyons et al., 1981; MacDiarmid, 1991), although autumnal reproduction may also occur after a midsummer lull (Kanciruk & Herrnkind, 1976; Herrnkind & Lipcius, 1989). Females of many species mate some time after their moult to maturity and are, therefore, not restricted in the time available for mating. For instance, P. argus, P. cygnus and P. homarus mate from a few days to several weeks or months after moulting (Berry, 1971a, b; Morgan, 1980; Phillips et al., 1980; Lyons et al., 1981; Lipcius, 1985). There are, however, limits in that mature females about to extrude eggs do not always resorb eggs in the absence of mating, or postpone egg extrusion indefinitely (Lipcius & Herrnkind, 1985; Kittaka & MacDiarmid, 1994). In other species, such as those in the genus Jasus, mating occurs shortly after the female moults (Heydorn, 1969; Berry, 1970; Cobb & Wang, 1985; Kittaka & MacDiarmid, 1994). Some of the most novel advances in palinurid reproduction deal with the potential for sperm limitation (MacDiarmid & Butler, 1999), which offers a contrast to the long-standing belief that most lobster and crab populations are limited primarily by female reproductive output (see Fogarty, 1995). If verified, the condition of sperm limitation requires managers to assess sex-specific mortality rates and impacts upon population dynamics and stock resilience. Moreover, there are size-specific patterns in the timing of moulting and reproduction (Lipcius, 1985; MacDiarmid, 1989a, 1991). Larger adult females generally spawn eggs and release larvae earlier in the reproductive period, and produce more annual broods than smaller, adult females, which moult early in the reproductive period. Similarly, smaller adult males of many species moult early in the mating season while larger males mate rather than moult. Larger females


25

produce two to four broods annually, depending on the species (e.g. Panulirus homarus produces up to four annually; Berry, 1973), whereas smaller females spawn at least once annually (Lyons et al., 1981; Lipcius, 1985). Incubation ranges from a few weeks to several months prior to release of the planktonic phyllosome larvae (Cobb & Wang, 1985). These size-specific patterns are apparently constrained by a combination of environmental and physiological factors, and potentially maximize the lifetime reproductive output of individuals (Lipcius, 1985). Environmental control of reproduction and moulting involves photoperiod and temperature, such that long daylengths and warmer temperatures enhance courtship, spawning frequencies and female gonadal development, but not aggression or male gonadal development (Lipcius & Herrnkind, 1987). Moulting rates are typically elevated by warm temperatures, but are apparently not affected significantly by photoperiod. Hatching appears to be rhythmic, with a hatching peak near sunrise for J. edwardsii (MacDiarmid, 1985). In some species, such as J. edwardsii, females form dense aggregations in areas of strong tidal water flow, probably to facilitate dispersal of newly hatched larvae (McKoy & Leachman, 1982). Shelter fidelity among spiny lobsters has been widely accepted, but little studied. Lobster movements monitored by ultrasonic telemetry or diver surveys demonstrated that lobsters have the ability to relocate a shelter or one nearby (Herrnkind et al., 1975; Cobb 1981). Only three studies have quantified the degree of shelter fidelity by tracking shelter use of several individuals over successive days (Herrnkind et al., 1975; MacDiarmid et al., 1991; Ratchford, 1999). Shelter fidelity was 42% and 38% for populations of P. argus in deep water (>10 m) in the US Virgin Islands and shallow water (1±2 m) in the central Bahamas, respectively (Herrnkind et al., 1975; Ratchford, 1999). Shelter fidelity was also 40% for J. edwardsii in New Zealand (MacDiarmid et al., 1991). Ratchford (1999) found that P. argus used three or four known shelters, moved up to 27 m among known shelters over a 4-week period, and moved 10±185 m overnight when shifting shelters. Similarly, Herrnkind et al. (1975) reported that lobsters used three or four shelters within 140 m of their study, and typically moved 30±90 m overnight. A major issue dealing predominantly with the adult phase concerns stock structure. In many species, identification of a fishery stock (i.e. a manageable segment of a population) is a relatively simple task, as in the western rock lobster, P. cygnus, which is limited by geographical boundaries and the relatively onshore± offshore orientation of coastal habitats. In contrast, other species are dispersed over diverse habitats linked to differing degrees by oceanic currents and geographic features. For instance, the Caribbean spiny lobster, P. argus, is widespread in coastal habitats from Bermuda to Brazil. Numerous currents traverse this region, as well as eddies and gyres, which may be long-lived or ephemeral (Yeung & McGowan, 1991). Previous attempts to identify stock structure in palinurids with electrophoretic techniques have met with limited success due to the relatively low levels of allozyme variation in J. edwardsii and J. novaehollandiae (Smith et al., 1980), P. marginatus (Shaklee & Samollow, 1984) and P. argus (Menzies & Kerrigan, 1979; Menzies,

26


1981). More recently, genetic variability of P. argus collected throughout the Caribbean and Florida and measured through an analysis of mitochondrial DNA was extremely low, suggesting substantial gene flow between hydrodynamically connected populations during their extended larval phase (Silberman et al., 1994). An analysis of mitochondrial DNA and the use of the polymerase chain reaction method for DNA amplification appears to be one of the most promising techniques in stock identification for palinurids, as evidenced in P. argus (Silberman et al., 1994), J. verreauxi (Brasher et al., 1992) and J. edwardsii (Ovenden et al., 1992).

Fishery ecology This chapter emphasizes an ecological approach towards attainment of an understanding of patterns and processes underlying population fluctuations in palinurids. Classical fisheries approaches (e.g. utilizing stock-recruit and yield models) have been described previously (Morgan, 1980; Cobb & Wang, 1985) and will not be emphasized here. Although emphasizing the American lobster, Homarus americanus, the comprehensive review by Fogarty (1995) serves as an excellent resource for recent and classical quantitative methods in fisheries management, including discussions of stage-based and spatially explicit models. The reader is urged to consult treatments of both approaches (i.e. ecological and stock assessment) and integrate these to achieve a thorough understanding of population dynamics in spiny lobsters. All lobster populations fluctuate substantially in abundance and at various scales in time and space (Cobb & Wang, 1985). These fluctuations result from diverse biological and physical forces acting on all life stages and in habitats ranging from shallow-water nurseries to the open ocean. The lack of understanding of the key controls of population fluctuations, both biotic and abiotic, has, in many instances, prevented us from attaining the primary goals of fisheries science, i.e. prediction of abundance, understanding sources of variation and development of effective fishery management strategies. The classical solution to these problems involves the utilization of stock-recruitment and other population dynamic models (Morgan, 1980; Cobb & Wang, 1985; Caputi, 1989; Chubb, 1994), with recent advances through the use of novel approaches such as risk analysis and spatially explicit models in a Bayesian framework (Punt & Kennedy, 1997). These analyses have yielded significant stock-recruitment models for some crustacean fisheries (Rothschild & Brunenmeister, 1984; Caputi, 1989; Lipcius & Van Engel, 1990; Chubb, 1994), although the variation about the relationships often limits their utility. However, recent advances in stock assessment methods (e.g. Punt & Kennedy, 1997) and biological techniques, such as age determination (Sheehy et al., 1998), have greatly enhanced the ability to manage palinurid fisheries effectively. The reader may refer to the references noted in the Introduction, as well as Fogarty (1995), as an entry into the world of stock assessment.


27

A complementary ecological approach is needed for the determination of population fluctuations in palinurids (Paulik, 1973; Fogarty & Idoine, 1986; Rothschild, 1986; Fogarty, 1995; Botsford et al., 1997). As exemplified in the Paulik diagram of stock-recruitment relationships (Fig. 7), the influences and controlling factors of populations can occur at various life-history stages. Stochastic forces, habitat quality and density-dependent regulation are postulated jointly to control the population dynamics of palinurids. Stochastic variation, such as that due to meteorological or oceanographic processes and their resultant density-independent survival and dispersal, strongly influence early life-history stages, causing substantial variation in the survival of larvae and post-larvae (Fig. 7, Quadrant 2). Thereafter, regulatory processes such as density-dependent mortality due to predation (Fig. 7, Quadrant 4) regulate survival in the juvenile phases, as exemplified by shelter limitation in P. argus and densitydependent mortality in P. cygnus. The critical or bottleneck stages (Caddy, 1986; Caddy & Stamatopoulos, 1990) for palinurids are likely to include the early benthic and later benthic phases of juveniles, which appear to be limited to some degree by settlement substrate and larger dens or crevices, respectively. Investigations of habitat relationships must account for the influence of the seascape (Dobson et al., 1997), which includes features such as habitat fragmentation, isolation and

Fig. 7 Paulik diagram representing the various life-history phases of a palinurid lobster and the likely regulatory mechanisms of population variation. Adapted from Rothschild (1986).

28


interconnectedness. The impact of these phases, and density-dependent regulation in general, is influenced greatly by the degree of stochastic variation imposed upon the larvae and post-larvae by physical processes, such as wind-driven oceanic transport. If variation in larval and post-larval abundance is high, then density-dependent regulation may only be important at extremely high or low levels of abundance in the juvenile phase (termed density-vague control); if variation is low, then densitydependent regulation may be the most critical control of population abundance through its action upon the juvenile phase. Future experimental investigations should emphasize identification of the relative influence of stochastic, habitat-related and density-dependent processes in the life history of palinurids.

Fisheries Palinurids sustain major commercial fisheries (Table 1) while simultaneously supporting local, small-scale fisheries in remote coastal locations and islands. Capture methods for palinurids are diverse, including the use of traps, pots, skin diving, spears, SCUBA (self-contained underwater breathing apparatus) and nets of various sorts (Bowen, 1980; Phillips et al., 1980). Many palinurids form the basis for specialized fisheries, such as P. argus in the Caribbean, P. cygnus in Western Australia and Jasus in New Zealand. Other species, such as Palinurus, are caught incidentally or as part of mixed-species fisheries. Although some species are not caught in large numbers, their high market value, which characterizes most palinurids, makes their capture and sale profitable, even if only to local hotels for the tourist market. Several deep-sea species require specially outfitted vessels for their capture, in many instances precluding large-scale commercial fisheries owing to the expenses involved in location and capture. Most palinurids are caught with lobster pots or traps in relatively shallow seas, although many are also trawled over muddy or sandy bottoms. Recreational fisheries also abound, with most lobsters caught by hand or spear while skin or SCUBA diving in shallow waters.

World catch of palinurid lobsters World-wide, the average yearly catch of marine crustaceans (5 210 920 mt) constitutes about 6.0% of the landings of marine species: 87 391 320 mt (annual average for 1991±1995, FAO, 1997). The annual world catch of 212 290 mt for palinurid, nephropid, homarid and scyllarid lobsters represents approximately 4% of the annual world catch for marine crustaceans, which comprises crabs, lobsters, galatheids, shrimps, prawns, krill and miscellaneous species. The annual palinurid catch averaged 74 817 mt annually from 1991 to 1995 (Table 4), which comprised 35.2% of the world catch for lobsters. Considering other lobsters, except for galatheids, homarids constituted 34.6% of the world lobster catch, nephropids


29

28.9% and scyllarids 1.3% (Table 4). Major species of exploited lobsters include the American lobster, H. americanus (70 555 mt), the Norway lobster, Nephrops norvegicus (60 013 mt), and the Caribbean spiny lobster, Panulirus argus (38 020 mt). As a contrast, the highest annual mean catches (1991±1995) for marine crabs are those of the gazami crab, Portunus trituberculatus, from the Yellow Sea, China Sea, Sea of Japan and Pacific Ocean equaling 215 149 mt; of the blue crab, Callinectes sapidus, from the Atlantic Ocean, Caribbean Sea and Gulf of Mexico, equalling 109 662 mt; and of the blue swimming crab, Portunus pelagicus, from the Pacific and Indian Oceans, equalling 57 722 mt (FAO, 1997). Most production of palinurid lobsters originates in South Africa, Australia, New Zealand, Cuba, Brazil, the USA and Mexico (Bowen, 1980). Of the commercially fished palinurid genera, Panulirus contributed 29.0% of lobster catches, Jasus 4.3% and Palinurus 3.1% (Table 4). Key species in the world palinurid catch from 1991 to 1995 included the Caribbean spiny lobster, P. argus, with 50.8%; the western rock lobster, P. cygnus, with 15.3%; the green rock lobster, J. verreauxi, with 4.9%; the red rock lobster, J. edwardsii, with 4.0%; and the Cape rock lobster, J. lalandii, with 2.8% (Table 1) (FAO, 1997).

Table 4 World catch for lobsters Mean catcha (mt)

%

Family

Genus

Palinuridae

Panulirus Jasus Palinurus

61 414 9219 4184

29.0 4.3 2.0

Subtotal:

74 817

35.3

Nephrops

60 013 1238

28.3 0.6

Subtotal

61 251

28.9

Homaridae

Homarus

73 452

34.6

Scyllaridae

Various genera

2770

1.3

Nephropidae

Total: Derived from FAO (1997). a Mean from 1991±1995.

212 290

30


Fisheries management and conservation Management of palinurid fisheries employs diverse approaches (Annala & Sullivan, 1997), ranging from input (e.g. effort) controls such as limited entry to output (e.g. catch quotas) controls such as individual transferable quotas (Table 5). The types of approach used and their effectiveness varies world-wide, as exemplified by the range of management strategies implemented in the Pacific and Indian Oceans (Table 5); similar variety in management strategies characterizes palinurid fisheries in other ocean basins. As an example of successful management, the J. edwardsii fishery off Gisborne, New Zealand, has witnessed substantial increases in catch per unit effort (CPUE) (Fig. 8a) after implementation of seasonal closures, reduction in minimum size, catch limits and temporal shifts in the timing of catches to wintertime (Breen & Kendrick, 1997). In contrast, the fishery for J. lalandii in Namibia suffered virtual collapse after both management-related and environmental perturbations (Fig. 8b). Prior to 1968, the fishery enjoyed relatively high catch rates, which may have lulled

Fig. 8 (a) CPUE of Jasus edwardsii in Gisborne, New Zealand, increased substantially after the implementation of a comprehensive management scheme in 1993. Adapted from Breen & Kendrick (1997). (b) CPUE of Jasus lalandii in Namibia decreased significantly after elimination of a minimum size limit in 1968, and further after episodic hypoxia caused mortality in 1988. Adapted from Grobler & Noli-Peard (1997).


31

management into a false sense of security, resulting in elimination of the minimum size limit in 1968 (Grobler & Noli-Peard, 1997). Catch rates (CPUE) dropped substantially within 2 years, and subsequently became negligible after episodic hypoxia further drove the stock to near collapse (Fig. 8b). These contrasting situations offer an excellent lesson on the need to adopt a risk-averse approach to fishery management, and hopefully avert fishery collapse, which sometimes results from the joint effects of overexploitation and poor environmental conditions. In an earlier treatment (Lipcius & Cobb, 1994), the following were deemed reasonable guidelines for the management of spiny lobster fisheries (Bowen, 1980; Brown & Caputi, 1986; Lyons, 1986): (1) accurate catch-effort and length-frequency data are required for sound regulation; (2) restrictions upon catch and effort through size limits, catch quotas or seasons, and limited or delayed entry should be mandated and enforceable to be effective; (3) development of accurate recruitment indices (e.g. Western rock lobster fishery) based on long-term post-larval or juvenile abundance data sets appears successful in projecting catch levels in subsequent years (Caputi & Brown, 1986; Phillips, 1986); and (4) there needs to be a communication network involving fishermen, industry, management and the research community. In addition to the measures described earlier (Table 5), a risk-averse approach is central to Table 5 Management strategies used with various spiny lobster fisheries Region Management strategy

India

South Africa

West Australia

New Zealand

Input controls Limited entry Vessel size Gear restrictions Pot limits Escapement gaps Closed areas Closed seasons Minimum legal size Condition Maximum legal size Output controls

* * *

* *

Total allowable catch Individual transferable quota Bag limits (recreational) Adapted from Annala & Sullivan (1997).

* *

* * * *

* *

* *

* * * *

*

* *

*

* * *

32


successful fishery management, a combination of effective fishery management strategies and comprehensive research investigations is required (e.g. Breen & Kendrick, 1997; Pitcher et al., 1997), and the use of closed areas (e.g. marine reserves; Childress, 1997) should be considered as a valuable tool where other effort and catch controls have had limited success. Such measures are essential to the sustainable resource use of palinurids in the face of the growing threats due to anthropogenic global environmental change.

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Parker, K.P. (1972) Recruitment and behavior of puerulus larvae and juveniles of the California spiny lobster, Panulirus interruptus (Randall). M.S. thesis, San Diego State University, San Diego, CA, USA. Parrish, F.A. & Polovina, J.J. (1994) Habitat thresholds and bottlenecks in production of the spiny lobster (Panulirus marginatus). Bull. Mar. Sci., 54, 151±63. Paulik, G.J. (1973) Studies of the possible form of the stock-recruitment curve. Rapp. Proc.-Verb. Reun., 164, 302±15. Pearce, A.F. & Phillips, B.F. (1988) ENSO events, the Leeuwin Current, and larval recruitment of the western rock lobster. J. Cons. Int. Explor. Mer., 45, 13±21. Phillips, B.F. (1972) A semi-quantitative collector of the puerulus larvae of the western rock lobster Panulirus longipes cygnus George (Decapoda, Palinuridae). Crustaceana, 22, 147±54. Phillips, B.F. (1981) The circulation of the southeastern Indian Ocean and the planktonic life of the western rock lobster. Oceanogr. Mar. Biol. Ann. Rev., 19, 11±39. Phillips, B.F. (1983) Migrations of pre-adult western rock lobsters, Panulirus cygnus, in Western Australia. Mar. Biol., 76, 311±18. Phillips, B.F. (1986) Prediction of commercial catches of the western rock lobster Panulirus cygnus. Can. J. Fish. Aquat. Sci., 43, 2126±30. Phillips, B.F. (1990) Estimating the density and mortality of juvenile western rock lobsters (Panulirus cygnus) in nursery reefs. Can. J. Fish. Aquat. Sci., 47, 1330±8. Phillips, B.F., Brown, P.A., Rimmer, D.W. & Reid, D.D. (1979) Distribution and dispersal of the phyllosoma larvae of the western rock lobster Panulirus cygnus in the southeastern Indian ocean. Aust. J. Mar. Freshwat. Res., 20, 773±83. Phillips, B.F., Cobb, J.S. & George, R.W. (1980) General biology. In The Biology and Management of Lobsters, Vol. I (Ed. by J.S. Cobb & B.F. Phillips), pp. 1±82. Academic Press, New York, USA. Phillips, B.F., Cruz, R., Brown, R.S. & Caputi, N. (1994a) Predicting the catch of spiny lobster fisheries. In Spiny Lobster Management (Ed. by B.F. Phillips, J.S. Cobb & J. Kittaka), pp. 285± 301. Blackwell Scientific Publications, Cambridge, MA, USA. Phillips, B.F. & Macmillan, D.L. (1987) Antennal receptors in puerulus and postpuerulus stages of the rock lobster Panulirus cygnus (Decapoda: Palinuridae) and their potential role in puerulus navigation. J. Crust. Biol., 7, 122±35. Phillips, B.F. & McWilliam, P.S. (1986) The pelagic phase of spiny lobster development. Can. J. Fish. Aquat. Sci., 43, 2153±63. Phillips, B.F. & Olsen, L. (1975) Swimming behaviour of the puerulus larvae of the western rock lobster. Aust. J. Mar. Freshwat. Res., 26, 415±17. Phillips, B.F., Pearce, A.F., Litchfield, R. & Guzman del ProÂo, S.A. (1994b) Spiny lobster catches and the ocean environment. In Spiny Lobster Management (Ed. by B.F. Phillips, J.S. Cobb & J. Kittaka), pp. 250±61. Blackwell Scientific Publications, Cambridge, MA, USA. Phillips, B.F. & Sastry, A.N. (1980) Larval ecology. In The Biology and Management of Lobsters, Vol. II (Ed. by J.S. Cobb & B.F. Phillips), pp. 11±57. Academic Press, New York, USA. Pitcher, C.R., Dennis, D.M. & Skewes, T.D. (1997) Fishery-independent surveys and stock assessment of Panulirus ornatus in Torres Strait. Mar. Freshwat. Res., 48, 1059±68. Pollock, D.E. (1978) Growth and production rates of the rock lobster Jasus lalandii (H. MilneEdwards). Ph.D. thesis, University of Witwatersrand, South Africa. Pollock, D.E. (1990) Paleoceanography and speciation in the spiny lobster genus Jasus. Bull. Mar. Sci., 46, 387±405. Pollock, D.E. (1992) Paleoceanography and speciation in the spiny lobster genus Panulirus in the Indo-Pacific. Bull. Mar. Sci., 51, 135±46. Pollock, D.E. (1993) Speciation in spiny lobsters ± clues to climatically-induced changes in ocean circulation patterns. Bull. Mar. Sci., 53, 937±44.

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Punt, A.E. & Kennedy, R.B. (1997) Population modelling of Tasmanian rock lobster, Jasus edwardsii. Mar. Freshwat. Res., 48, 967±80. Ratchford, S.G. & Eggleston, D.B. (1998) Size- and scale-dependent chemical attraction contribute to an ontogenetic shift in sociality. Anim. Behav., 56, 1027±34. Rimmer, D.W. & Phillips, B.F. (1979) Diurnal migration and vertical distribution of phyllosoma larvae of the western rock lobster, Panulirus cygnus George. Mar. Biol., 54, 109±24. Rothschild, B.J. (1986) Dynamics of Marine Fish Populations, 277 pp. Harvard University Press, Cambridge, MA, USA. Rothschild, B.J. & Brunenmeister, S.L. (1984) The dynamics and management of shrimp on the northern Gulf of Mexico. In Penaeid Shrimps ± Their Biology and Management (Ed. by D.A. Gulland & B.J. Rothschild), pp. 145±72. Fishing News Books, Farnham, UK. Sastry, A.N. (1983a) Pelagic larval ecology and development. In The Biology of Crustacea, Vol. 7 (Ed. by F.J. Vernberg & W.B. Vernberg), pp. 213±82. Academic Press, New York, USA. Sastry, A.N. (1983b) Ecological aspects of reproduction. In The Biology of Crustacea, Vol. 7 (Ed. by F.J. Vernberg & W.B. Vernberg), pp. 179±270. Academic Press, New York, USA. Serfling, S.A. (1972) Recruitment, habitat preference, abundance and growth of the puerulus and early juvenile stages of the California spiny lobster, Panulirus interruptus (Randall). M.S. thesis, San Diego State University, San Diego, CA, USA. Serfling, S.A. & Ford, R.F. (1975) Ecological studies of the puerulus larval stage of the California spiny lobster Panulirus interruptus Randall. Fish. Bull., U.S., 73, 360±77. Shaklee, J.B. & Samollow, P.B. (1984) Genetic variation and population structure in a spiny lobster, Panulirus marginatus, in the Hawaiian archipelago. Fish. Bull., U.S., 82, 693±702. Sheehy, M., Caputi, N., Chubb, C. & Belchier, M. (1998) Use of lipofuscin for resolving cohorts of western rock lobster (Panulirus cygnus). Can. J. Fish. Aquat. Sci., 55, 925±36. Silberbauer, B.I. (1971) The biology of the South African rock lobster Jasus lalandii (H. MilneEdwards). 1. Development. S. Afr. Div. Sea Fish., Invest. Rep., 92, 1±70. Smith, K.N. & Herrnkind, W.F. (1992) Predation on early juvenile spiny lobsters Panulirus argus (Latreille): influence of size and shelter. J. Exp. Mar. Biol. Ecol., 157, 3±18. Smith, P.J., McKoy, J.L. & Machin, P.J. (1980) Genetic variation in the rock lobsters Jasus edwardsii and Jasus novaehollandiae. N.Z. J. Mar. Freshwat. Res., 14, 55±63. Spanier, E. & Zimmer-Faust, R.K. (1988) Some physical properties of shelter that influence den preference in spiny lobsters. J. Exp. Mar. Biol. Ecol., 121, 137±49. Street, R.J. (1971) Rock lobster migration off Otago. N.Z. Comm. Fish., June,16±17. Tegner, M.J. & Levin, L.A. (1983) Spiny lobsters and sea urchins: analysis of a predator-prey interaction. J. Exp. Mar. Biol. Ecol., 73, 125±50. Trendall, J. & Bell, S. (1989) Variable patterns of den habitation by the ornate rock lobster, Panulirus ornatus, in the Torres Strait. Bull. Mar. Sci., 45, 564±73. White, A. (1847) List of the specimens of Crustacea in the collection of the British Museum. i±viii, 1±143. Williams, A.B. (1986) Lobsters ± identification, world distribution, and U.S. trade. Mar. Fish. Rev., 48, 1±36. Williams, A.B. (1988) Lobsters of the World ± An Illustrated Guide, 186 pp. Osprey Books, Huntington, NY, USA. Williams, B.G. & Dean, I.C. (1989) Timing of locomotor activity in the New Zealand rock lobster, Jasus edwardsii. N.Z. J. Mar. Freshwat. Res., 23, 215±24. Witham, R.R., Ingle, R.M. & Joyce, E.A., Jr (1968) Physiological and ecological studies of Panulirus argus from the St. Lucie estuary. Fla. Bd. Cons. Tech. Ser., 53, 1±31. Yeung, C. & McGowan, M.F. (1991) Differences in inshore-offshore and vertical distribution of phyllosoma larvae of Panulirus, Scyllarus and Scyllarides in the Florida Keys in May±June, 1989. Bull. Mar. Sci., 49, 699±714.


41

Yoshimura, T. & Yamakawa, H. (1988) Microhabitat and behavior of settled pueruli and juveniles of the Japanese spiny lobster Panulirus japonicus at Kominato, Japan. J. Crust. Biol., 8, 524±31. Zimmer-Faust, R.K. (1991) Chemical signal-to-noise detection by spiny lobsters. Biol. Bull., 181, 419±26. Zimmer-Faust, R.K. & Case, J.F. (1982) Odors influencing foraging behavior of the California spiny lobster, Panulirus interruptus, and other decapod Crustacea. Mar. Behav. Physiol., 9, 35± 58. Zimmer-Faust, R.K. & Case, J.F. (1983) A proposed dual role of odor in foraging by the California spiny lobster, Panulirus interruptus (Randall). Biol. Bull., 164, 341±53. Zimmer-Faust, R.K. & Spanier, E. (1987) Gregariousness and sociality in spiny lobsters: implications for den habitation. J. Exp. Mar. Biol. Ecol., 105, 57±71. Zimmer-Faust, R.K., Tyre, J.E. & Case, J.F. (1985) Chemical attraction causing aggregation in the spiny lobster, Panulirus interruptus (Randall), and its probable ecological significance. Biol. Bull., 169, 106±18.

Part 1 Fisheries: Methods, Management and Status


Chapter 1

The Status of Australia's Rock Lobster Fisheries B.F. PHILLIPS Curtin University of Technology, P.O. Box 1987, Perth, Western Australia 6845, Australia

C.F. CHUBB and R. MELVILLE-SMITH Bernard Bowen Fisheries Research Institute, Western Australian Marine Research Laboratories, P.O. Box 20, North Beach 6020, Western Australia, Australia

1.1

Introduction

Seven Panulirus and Jasus rock lobster species are found in Australian waters, four of which support significant commercial and recreational fisheries. Panulirus cygnus is found on the lower west coast, Panulirus ornatus in northern Australia, particularly the Torres Strait and far north Queensland, Jasus edwardsii (formerly J. novaehollandiae) in southern Australia: South Australia, Victoria and Tasmania, and Jasus verreauxi on the central east coast, New South Wales (Fig. 1.1). The commercial whole weight catches of the four species were P. cygnus 13 600 t, P. ornatus 460 t (1990), J. edwardsii 4730 t (1997/98) and J. verreauxi 150 t (1997/ 98), giving a total of about 18 900 t, making Australia's rock lobster production the largest in the world. This chapter summarizes the level of knowledge on the four important commercial species and the status of each of the stocks. 1.2

The tropical rock lobster P. ornatus in the Torres Strait

The fishery for P. ornatus in the Torres Strait is reserved for exploitation by the traditional inhabitants of Torres Strait and is the basis of their most important commercial fishery. First assessments of the stocks showed that exploitation was low and it was recommended that increased involvement by Islander divers could be encouraged. However, lower recruitment and increased fishing mortality have now indicated that new management measures may, in future, need to be considered to address uncapped effort potential in the fishery (Pitcher et al., 1997). 1.2.1

Distribution and life history

Panulirus ornatus is widely distributed throughout the Indo-West Pacific and around the northern Australian coast, the Torres Strait and southern coast of Papua New Guinea (George, 1968, 1972), and prefers calm turbid waters (George, 1968). Most 45

46


Fig. 1.1

The main distribution of Australian rock lobster species.

research has been undertaken in the Torres Strait region, where a commercial fishery for this species operates (Fig. 1.2). As with all Panulirus species, P. ornatus has a relatively long larval phase (estimated to be 6 months) in the open ocean before the puerulus stage settles into small holes and crevices in the reefs and seabed. Growth is rapid with animals attaining the minimum 100 mm tail length (80 mm carapace) about 1 year after settling (1.5 years old). When P. ornatus is about 2.5 years old (2 years after settlement) there is a synchronized moult in August±October and virtually all the females and the majority of the males migrate north-east into the Gulf of Papua

The Status of Australia's Rock Lobster Fisheries

Fig. 1.2

47

Detail of the Torres Strait and the migration of Panulirus ornatus.

(Pitcher, 1991). Just before or during the migration the lobsters undergo reproductive development and large numbers finally arrive on the breeding grounds of the coastal reefs of the eastern Gulf of Papua (Fig. 1.2), while others probably migrate further east to the reefs at the northern end of the Great Barrier Reef (Pitcher, 1991). The females move offshore from the breeding grounds to release their eggs and then move back inshore again. The larvae are thought to become entrained in an ocean gyre in the north-west Coral Sea that could distribute the larvae back into the Torres Strait (Pitcher, 1991). It is not known how important the larval contribution of the breeding populations of the northern Great Barrier Reef and north-east Queensland coast is to overall recruitment. However, these populations are mainly in deep water (75 m), beyond the access of the commercial dive fishery (Pitcher, 1991). There is evidence that the post-migration breeding lobsters at Yule Island (Fig. 1.2) are in very poor physical condition and there is virtually total mortality of postspawning animals (Pitcher et al., 1991). This means that the level of puerulus settlement on the fishing grounds each year may depend on the success of the previous year's migration and breeding (Pitcher, 1991).

48


1.2.2

History of the fishery

There has been an artisanal fishery on P. ornatus in the Torres Strait and the east coast of Papua New Guinea for hundreds of years. Commercial fishing began in the area in the late 1960s and is restricted to the indigenous Torres Strait Islander people, who derive from it a significant portion of their income (Channells, 1986). Diving is the method of fishing, with fishers free diving (1±4 m depth) or using hookah (4±15 m depth) from small outboard powered dinghies and returning their catches to land-based processors or processing vessels. There are 277 licensed boats in the fishery. Twelve small mobile freezer vessels also operate in the fishery, acting as mother ships for teams of divers. The lobsters were originally sold as tails on the Australian and overseas (mainly USA) markets (Channells, 1986). However, a market has now developed for live lobsters in Asia. Island communities operate 300±400 dinghies throughout the fishery, with the principal fishing grounds being around Thursday Island, Orman Reef and Warrior Reefs (Fig. 1.2), which are the areas most accessible to the major population centres. Fishing occurs year round, with peak catches coming in March±August and little fishing activity during October±December (Pitcher, 1991). Catches (whole weight assuming a 40% tail weight recovery) from Torres Strait have varied from 170 to 310 t between 1970 and 1980. Catches peaked in 1986 at 875 t but were between 375 and 625 t from 1987 to 1991 (Fig. 1.3). Recruitment to the fishery varies considerably, and the estimated catch in 1999 is only 159 t (CSIRO, 1999).

1.2.3

Management controls

Management of the stock is affected by a minimum legal size of 100 mm tail length or an 80 mm carapace length measurement. There are no controls on fishing effort

Fig. 1.3

Torres Strait Panulirus ornatus catch (whole weight) from 1978 to 1998.


49

per se, but with access now limited to Torres Strait Islanders, it is unlikely that there would be a large expansion in the number of fishers. The only other source of fishing effort was trawlers that targeted the spawning migration in the Gulf of Papua. Trawling was banned in 1984. The fishery is managed under a treaty (Article 22) between Australia and Papua New Guinea as the Torres Strait Protection Zone, in which the traditional way of life and livelihood of the inhabitants is to be protected (Anon., 1992a). 1.2.4

Status of the stock

Since 1989 the abundance of P. ornatus over the approximate 25 000 square km of the Torres Strait fishery was estimated by using a dive/transect technique (Pitcher et al., 1992, 1997). Panurilus ornatus was found throughout the area, except in the centre of the fishery where the habitat is unsuitable silt and mud. Densities varied from two to over 100 animals per hectare. It was estimated that there are 11±17 million lobsters in the Torres Strait, with about 8 million being legal size (Pitcher, 1991; Pitcher et al., 1992). The annual fishery-independent surveys of the relative stock abundance, and catch sampling, have contributed to the development of a simple cohort dynamics model of the fishery for a range of fishing mortalities. It estimates the potential yield 1 year in advance ± information valuable for managers considering development options and negotiating catch-sharing agreements and access rights (Pitcher et al., 1997). It has been estimated that the long-term potential yield from the fishery could be 730 t (Anon., 1998a). However, the low catches in the last few years may require a review of this assessment. 1.2.5

Future research

Puerulus collectors (Phillips, 1972) are being tested to see whether they offer a suitable method for an even earlier assessment of recruitment to the fishery, as has been used for the western rock lobster (Caputi et al., 1995). Ecological studies on the early juveniles, growth and mortality will be undertaken to expand the general understanding of the species and the impact of fishing (Pitcher, 1991). Future research will develop the model by incorporating information from ongoing surveys, catch recording and logbook data from Australia and Papua New Guinea fisheries (Pitcher et al., 1997).

1.3

The eastern rock lobster Jasus verreauxi of New South Wales

Although the New South Wales rock lobster J. verreauxi catch is small (reported commercial catch approximately 100 t), it forms the basis of an important fishery.

50


The eastern rock lobster is considered the premier seafood of New South Wales, enjoying a high public profile, and fetching substantially higher prices than rock lobsters imported from other Australian states.

1.3.1


Jasus verreauxi is found along the entire New South Wales coast (Fig. 1.4), from inshore to the edge of the continental shelf (200 m depth). Mating takes place in spring and early summer and females carry eggs between September and January. The average size of mature females is approximately 167 mm carapace length (CL) and they have an estimated age of about 8 years (Montgomery, 1991; Anon., 1992b). The breeding stock is found predominantly in the northern sector of the fishery (i.e. north of Port Stephens, Fig. 1.4), in water deeper than 50 m (Montgomery, pers. comm.).

Fig. 1.4

Movement of Jasus verreauxi along the New South Wales coast.


51

The larval stage lasts for approximately 9 months before the puerulus stage settles on the inshore reefs and the benthic juvenile phase is commenced. J. verreauxi takes about 3±5 years from hatching to reach the legal minimum size of 104 mm CL. The animals are 6±7 years old before sexual maturity is attained. For animals around the legal size, 2±3 moults take place each year with carapace increments of about 6±7 mm each time (Anon., 1992b). Montgomery (pers. comm.) has hypothesized that there is a general movement of premature animals from the juvenile inshore areas to the deep-water breeding grounds and that lobsters from the southern section of the fishery undertake a prebreeding migration to the northern end of the species' distribution where the bulk of the breeding population is situated (Fig. 1.4). This migration takes place when the animals are about 4±5 years old. Once in the northern end of the distributional range, lobsters move seasonally to the edge of the Continental Shelf probably to mate, and then return to shallow inshore reefs to spawn. Further research needs to be undertaken to verify the extent of the movement and migration of J. verreauxi.

1.3.2


Jasus verreauxi has been fished in New South Wales since 1873. Reported catches peaked at nearly 400 t in the late 1920s, before dropping off in the 1930s and early 1940s (Fig. 1.5). A resurgence in landings took place in post-World War II years when many returned servicemen entered the general New South Wales fishing industry. Landings peaked once again at around 400 t in the late 1940s and early 1950s, but have since declined to an average of between 100 and 200 t, worth approximately $4.0 million per annum (Anon., 1998b). The fishery was initially focused inshore and in the northern sector from Crowdy Head to Coffs Harbour (Fig. 1.4). Fishermen used small boats to set traps around headlands and shallow reefs. In 1969, populations were discovered offshore from Sydney in deep water and the fishery expanded offshore (170±200 m depth) and southward from the ports of Ulladulla and Batemans Bay. The fishery has undergone significant changes in management since the early 1990s, with limited access introduced in 1992 and the introduction of a total allowable commercial catch (TACC) in 1994 (Montgomery et al., 1996).

1.3.3

Management controls

Controls in the fishery include a minimum legal size of 104 mm CL, a maximum legal size of 200 mm, a ban on the taking of egg-bearing females, restrictions on trap dimensions, restriction of the mesh size on traps to 50 mm and individual catch quotas (Montgomery et al., 1996). Recreational fishers are restricted to a bag limit of two per day and are allowed to use only one trap, or dive and capture with their

52


Fig. 1.5 New South Wales Jasus verreauxi catch in tonnes whole weight from 1884 to 1998 after Liggins et al. (1999). The landings include estimates of unreported commercial and recreational catches.

hands, without the use of compressed air. These regulations confine the recreational fishery to shallow inshore waters, although this was not the intention of their introduction (Montgomery, pers. comm.). There is no closed season (Anon., 1997a).

1.3.4

Status of the stock

Catch rates have been in decline over most of the life of the fishery, but stabilized in the early 1990s (Montgomery, 1995) and have increased since 1993/94 when a TACC was introduced (Anon., 1998a). It is considered that substantial quantities of the commercial catch went unreported in earlier years (Fig. 1.5), as many sales were made by commercial fishermen direct to restaurants and retail outlets and these went unrecorded. It is suggested (Montgomery et al., 1997), that catch and effort reporting by commercial fishermen has been more honest since the introduction of a TACC. A biomass dynamics model developed for the fishery (Montgomery et al., 1998) has suggested that there was a decline in biomass over most years between 1900 and 1993/94, but it has increased since then. The same assessment suggests that the unexploited biomass is likely to range around 36±41% of the biomass of the exploitable virgin stock. Most future scenarios for the fishery based on modelling


53

predictions suggest that TACCs of up to 150 t will result in an increase in stock biomass early into the twenty-first century (Montgomery et al., 1997). TACCs were set at 125 t in 1998/99 and were raised to 140 t in 1999/2000 (Montgomery pers. comm.). Of concern in this fishery is the depleted level of the spawning stock. The legal minimum size of 104 mm CL is well below 166 mm CL, the size at which the animals are first considered to breed (Montgomery, 1992). Length±frequency sampling (Montgomery, 1995) showed that only 10% of animals sampled in the commercial catch were larger than size at first breeding and that more than 64% of the annual total landings came from areas where only immature animals were sampled. It is suggested that the spawning stock may be as low as 3% of the exploitable unfished biomass (Montgomery et al., 1997). Puerulus collectors have been developed and deployed in this fishery since 1992 (Montgomery & Kittaka, 1994; Montgomery et al., 1996). The pattern of settlement has been shown, based on six sampling sites, to be highest in the central and southern part of the grounds between Sydney and Eden (Montgomery et al., 1997). The same study has shown that there are significant interannual differences between sampling sites. These data will form the baseline of a time series that may eventually provide an index of recruitment to the fishery. Another time series that may provide information in the future on the state of the brood stock as well as recruitment to the fishery, is a fishery-independent trapping survey that was initiated in 1995 (Montgomery et al., 1996, 1997). The number of recreational fishers has yet to be accurately established, but it has been suggested that possibly 60 000 people participated in this pastime in the 1996/ 97 season (Montgomery et al., 1997). A survey of recreational abalone divers, which picked up catches made by recreational lobster fishers as a by-product (Andrew et al., 1997), estimated that that group landed around 26 t of lobsters in the 1997 season.

1.3.5

Future research

An important future objective is to develop a length-based model for the fishery. Before this objective can be met, it will be necessary to collect additional information on the sizes of lobsters in the population, the size of the spawning biomass, growth and movement (Montgomery et al., 1997). Work on improving knowledge in these areas is underway.

1.3.6

Future management

The objective of management of the eastern rock lobster fishery as stated in the draft management plan is to increase the stock biomass (Montgomery et al., 1997). Based

54


on model outputs, Montgomery et al. (1997), all indications are that this objective will be achieved by the TACCs that have been set since the 1993/94 season.

1.3.7

Summary

Jasus verreauxi on the New South Wales coast is assessed to have been heavily exploited in the past. The current management strategy is aimed at rebuilding the depleted stocks and early results suggest that this initiative has achieved some success.

The southern rock lobster Jasus edwardsii (formerly known as Jasus novaehollandiae)

1.4

Jasus edwardsii has the widest distribution of any of the rock lobster species in Australia. The species supports significant commercial fisheries in South Australia, Tasmania, Victoria and southern Western Australia, with an insignificant amount taken in southern New South Wales (Fig. 1.1).

1.4.1


Jasus edwardsii is distributed around southern mainland Australia, Tasmania and New Zealand. In Australia, specimens have been found as far north as Geraldton in Western Australia (28 450 s, 114 070 E) and Coffs Harbour in northern New South Wales (30 180 S, 153 080 E), with the bulk of the population being on the coast of South Australia, Tasmania and southern Victoria from depths of 1±200 m (Figs. 1.1, 1.6). A small population is also found on the south coast of Western Australia east of 115 S. From morphological and genetic work (Booth et al., 1990, Ovenden et al., 1991), the Australian populations and those of New Zealand are the same stocks. Females are fertilized externally by the male depositing a spermatophore on their sternal plates (MacDiarmid, 1988) between April and July. The females carry eggs on the underside of their abdomen (tail) until hatching, which peaks in October. The larger the female the larger the number of eggs they carry (e.g. around 150 000 to over 640 000 eggs for females with a 97 mm and 150 mm carapace, respectively (Hobday & Ryan, 1997). The eggs hatch to a naupliosoma larvae that moults after about 12 h to the first phyllosoma stage. Puerulus settlement studies suggest that the phyllosoma may spend 8±22 months at sea prior to settlement (Kennedy, 1990; Booth, 1994), during which time they would become widely distributed in the southern ocean (Winstanley, 1977). The size at which 50% of the females breed varies considerably across the animal's range (Table 1.1). Females appear to reach sexual maturity (Table 1.1) at about 4

The Status of Australia's Rock Lobster Fisheries Table 1.1

55

Size of maturity of Jasus edwardsii in Australia

Location (Fig. 1.1)

Carapace length (mm) at which 50% of females mature

Western South Australiaa South-eastern South Australia Western Victoriab Eastern Victoriab King Island (Tasmania)c Southern Tasmaniac

100±114 a

88±96 90 112 115 100 mm decline by approximately 1 mm/year for every 20 m increase in depth of habitat, at depths of 20 m and greater. Males and females have similar growth rates until the females reach sexual maturity and their growth rate slows (McGarvey et al., 1999).

1.4.2


The catches of J. edwardsii reflect the distribution of the main part of the stock in South Australia, Tasmania and Victoria. Fishers use baited traps that they usually pull daily depending on weather and catch rates. In Tasmania, in waters deeper than 40 m, fishing is sometimes conducted during the day as well as at night during certain times of the season (i.e. traps are pulled more than once per day). In South Australia J. edwardsii was captured for domestic consumption as early as 1870. However, it was not until the introduction of freezer shipments to the USA in the late 1940s that commercial exploitation began in earnest. The growth of the fishery and changes in the distribution of fishing were most rapid between the late 1940s and 1966/67 (Copes, 1978). Fishing effort in 1949/50 was 109 000 pot lifts and by 1966/67 had escalated to 3 152 000 pot lifts; however, the catch had only increased from 1135 t to 2837 t over the same period (Copes, 1978; Staniford, 1986). This pattern of development is typical of that which occurred in the rock lobster fisheries in the other states (Victoria and Tasmania) and for P. cygnus in Western

56


Australia. Initially, there was strong growth in catch from the fishery and high catch rates; however, these declined sharply as the number of vessels and traps used in the fishery increased rapidly. The spectacular decline in catch rates in South Australia from 10.4 kg per trap lift in 1949/50 to 0.9 kg per trap lift in 1966/67 focused industry's and government's attention on the need to restrict further entry to the fishery. In 1967, the fishery became limited entry, no additional entrants were allowed and the number of traps used per vessel was restricted (Copes, 1978; Staniford, 1986). The fishery was divided into two management zones: a southern zone, from the Victorian border to the mouth of the Murray River, and a northern zone, from the mouth of the Murray River to the Western Australian border (Fig. 1.6). In the 1993/94 season the southern zone moved to TACC management system with individual transferable quotas. The northern zone has retained the effort-controlled management system, but has introduced a system of voluntary time closures to compensate for increases in effective fishing effort resulting from improved fishing efficiency. Both management systems appear to be working efficiently. Currently there are 183 boats and 11 923 traps in the southern zone and 71 boats and 3950 traps in the northern zone (Table 1.2). About 65% of the approximately 2500 t catch is taken in the southern zone. Because the grounds in the northern zone of the fishery are scattered over a wider area than in the southern zone, fishermen undertake fishing trips of several days to a week's duration, keeping their catch in live wells on their boats. In the southern zone virtually all fishing is day trips, although depending on catch rates and weather conditions, traps may not be pulled for 2±3 days. There are also about 6200 licensed recreational fishers in South Australia using up to two traps per person per day for a bag limit of four lobsters per person per day. Table 1.2 Number of boats and traps and maximum number of traps permitted to be fished per boat in the limited entry Jasus edwardsii fisheries in Western Australia (WA), South Australia (SA), Tasmania, Victoria and New South Wales (NSW) Location

Numbers of boats (entitlements)

WA (Esperance zone) WA (outer zones) SAa (S zone)

11 31 183

SAa (N zone) Tasmaniab Victoriac (E zone) (W zone) NSWd

71 314 69 90 177

Total number of traps

Maximum traps per boat

Minimum traps

568 3860 11 923

90 Unlimited 80

30 ± 40

3950 10 505 2615 5388 Unlimited

60 50 60 Unlimited Unlimited

25 15 15 10 ±

Source: aPrescott (pers. comm.), bKennedy & Gardner (pers. comm.), cMolloy, Hobday & Flint (pers. comm.), dMontgomery (pers. comm.).


57

Unlicensed recreational fishers are permitted to use up to three hoop nets or dive for up to four rock lobsters per day. Recreational trap fishermen were estimated to have taken 62 t of lobsters in 1991±92. (Prescott, pers. comm.) In Tasmania and Victoria the fishery for J. edwardsii developed along similar lines to that in South Australia, with a small fishery initially providing product for a limited local market. In 1882, Tasmania introduced its first management regulations for the fishery: season closures (Kennedy, 1989) and size limits (currently 110 mm and 105 mm CL for males and females, respectively). In 1926, limits were put on the number of traps that a boat could use. With the rapid expansion of the fishery after 1945, Tasmania introduced limited entry in 1967 to maintain the profitability of the industry and to limit the scope of future effort increases (Kennedy, pers. comm.). The fishery moved from effort to catch quota management for the 1998/99 fishing season, with the TACC being set at 1502.5 t for that season (Anon., 1997b). There are currently 314 boats in Tasmania fishing 10 505 traps (Table 1.2) for a catch of approximately 1500 t (Table 1.3). Many fishermen have access to other fisheries as well as rock lobsters (e.g. scallops, shark) but derive most of their income from rock lobster. Recreational fishing for rock lobster is a popular pastime for a small sector of the population. In June 1996 there were 6153 recreational licences issued to fishers allowing them to take rock lobsters with pots and 3465 by diving (Anon., 1997c). The recreational catch for the 1997/98 season was estimated to be 58 t (Lyle & Smith, 1998). In Victoria the fishery is divided into two zones at 143 400 E (Fig. 1.6), an eastern zone with 69 boats and 2615 traps and a western zone with 90 boats and 5388 traps (Table 1.2). About 80% of the total catch of 400±500 t (Table 1.3) comes from the western zone, where the densities of lobsters are higher. Western zone fishers have larger more powerful boats and an average trap licence of 60, whereas the eastern zone fishers have an average trap licence of 38 and are more heavily involved in other

Fig. 1.6

The fisheries for the southern rock lobster Jasus edwardsii in southern Australia.

58


Table 1.3 Catch (t whole weight) and fishing effort (thousands of trap lifts) for the Jasus edwardsii fishery in Western Australia, South Australia, Tasmania and Victoria Western Australia

South Australiaa

Tasmaniab

Victoriac

Year

Catch (t)

Catch (t)

Catch (t)

Thousand trap lifts

Catch (t)

Thousand trap lifts

1964/65 1965/66 1966/67 1967/68 1968/69

0.08 0.07 N/A N/A N/A

1513 1787 1946 1752 1700

1969/70 1970/71 1971/72 1972/73

1 3 5 10

1390 1607 1496 1583

1533 1609 1464

1148 1194 1081

1973/74 1974/75 1975/76 1976/77

17 21 20 11

18 23 14

1514 2031 2248 1891

2228 2364 2298

1195 1610 1394 1473

858 1053 1023 1119

N/A N/A N/A

N/A N/A N/A

1977/78 1978/79 1979/80 1980/81

18 17 23 12

24 22 22 11

1857 1938 1908 2810

2204 2337 1991 2523

1432 1569 1613 1854

1089 1067 1066 1136

N/A N/A 564 680

N/A N/A 738 853

1981/82 1982/83 1983/84 1984/85 1985/86

18 21 18 21 18

20 32 33 31 35

2720 2572 2412 2217 2204

2668 2766 2828 2605 2600

1747 2104 1982 2289 1970

1199 1427 1304 1585 1568

628 603 557 520 439

816 803 771 751 725

1986/87 1987/88 1988/89 1989/90

22 20 19 25

33 27 33 33

2207 2468 2275 2525

2519 2782 2554 2489

1862 1816 1896 1828

1617 1686 1693 1786

432 420 369 416

728 688 713 803

1990/91 1991/92 1992/93 1993/94

35 65 73 98

45 101 90 144

2666 3162 2818 2598

2640 2858 2507 2364

1738 1902 1797 1496

1848 2021 2055 1791

389 473 481 528

814 882 995 1003

1994/95 1995/96 1996/97 1997/98 1998/99d

96 100 81 82 76

198 201 159 151 148

2613 2587 2543 2623 2729

2217 2317 2475 2483 2258

1445 1841 1757 1588

1799 1908 1884 1798

508 482 464 501

1030 976 996 962

Thousand trap lifts

Thousand trap lifts

Source: aPrescott (pers. comm.), bKennedy & Gardner (pers. comm.), cMolloy, Hobday & Flint (pers. comm.), dProvisional estimates only.


59

fisheries besides rock lobster, e.g. shark (D. Molloy, Fisheries Department, Victoria, pers. comm.). Recreational lobster fishing is permitted in Victoria using hoop nets or by diving. Fishers are required to hold an amateur fishing licence, which enables them to catch a range of species, including up to four lobsters per person per day. The only survey of recreational fishers to date in Victoria (Hobday et al., 1998) suggested an annual catch of 18 t made by divers who completed a survey which targeted dive shops. This figure did not include catches made by hookah and snorkel divers, or by hoop net fishers, and would therefore have substantially underestimated the recreational lobster catch for the state (Hobday et al., 1998). The fishery for J. edwardsii in Western Australia is on the western extremity of the species distribution and has always been small (100 t or less; Table 1.3). The fishery is centred at Esperance (Fig. 1.6) and dates back to 1965 when fishermen supplied the local market and the larger towns of the inland goldfields. In the early years of the fishery (prior to 1986) most fishers treated the rock lobster fishery as a small adjunct to their main fishing operations (e.g. shark and tuna). In 1984 the tuna fishery, in which most fishers participated, came under quota management and nearly all the fishermen sold their allocations to the large operators in South Australia. Fishers then had more time and capacity to fish and generated additional fishing pressure on the J. edwardsii stocks on the traditional inshore grounds around Esperance, causing catch rates in this area to decline. Fishers began to move further afield, mainly eastwards and into deeper water. In 1987, the fishery became limited entry, with fishermen gaining access and a trap quota based on their catch and fishing effort in the preceding 3 years. Towards the end of the 1980s, deep-water grounds were found which improved the declining catch rates. In the early 1990s the Esperance zone of the fishery expanded westwards to Albany and Augusta and eastwards into the Great Australian Bight, resulting in a large increase in the total catch (Table 1.3). These areas produced good catch rates for a short period, but were rapidly depleted and although still being fished, are of diminishing importance. The Esperance zone is managed separately to the two outer zones and is restricted to 11 licensees with a combined pot holding of 568 pots. This zone of the fishery continues to be fished more intensively now than at any stage prior to the 1990s and appears to be withstanding the increased fishing pressure. The outer zones, by comparison, have 31 licence holders with a combined pot holding of 3860 pots. It is widely acknowledged that there is too much effort in these outer zones and that spiny lobster catch rates in these areas will remain depressed until effort levels are drastically reduced.

1.4.3

Major management controls

Boat and trap restrictions All states in Australia control their J. edwardsii fisheries through limited entry. This sets strict limits on the number of boats and the traps that can be used. Table 1.2

60


shows the number of boats and traps licensed to fish J. edwardsii in each state. In all states, the licence (endorsement to fish J. edwardsii) can be transferred from one fisher to another. No additional boats or traps are allowed. Maximum, and in some cases minimum, trap entitlements have also been set (Table 1.2) and most states (Western Australia, South Australia and Tasmania) have defined the size and design of traps, including the incorporation of escape gaps or large mesh to reduce the handling and hence the mortality of undersized animals. Seasonal closures Table 1.4 lists the various seasonal closures for male and female J. edwardsii. Tasmania (Kennedy, 1989) and Victoria (Hobday & Ryan, 1997) have longer closed seasons for females to give them added protection. Minimum size Various minimum sizes (carapace length) have been set for males and females in the five states, owing to differences in the size at first maturity and non-biological historic decisions, which in some cases were based on a size that the market would accept. No states have maximum sizes. Table 1.5 shows the minimum legal size for J. edwardsii in each state. In Tasmania the minimum size for females of 105 mm carapace protects virtually all breeding females on the south coast but is well under the size at 50% maturity for females in the north of the fishery at King and Flinders Islands. Recreational fishing Recreational fishing for J. edwardsii occurs in all states. Table 1.6 lists the main regulations governing the recreational fisheries. Recreational fishing is usually confined to shallow, sheltered waters. Table 1.4 Seasonal closure for the Jasus edwardsii fishery (males and females) in Western Australia (WA), South Australia (SA), Tasmania, Victoria and New South Wales (NSW) Locationa

Males

Females

WA

1 July to 14 November

1 July to 14 November

SA (S zone) SA (N zone) Tasmania

1 May to 30 September 1 June to 31 October 15 September to second Saturday in November, plus the last 5 days in February (commercial only) 1 September to 15 November None

1 May to 30 September 1 June to 31 October 1 May to second Saturday in November, plus the last 5 days in February (commercial only) 1 June to 15 November None

Victoria NSW a

Sources as for Table 1.2.


61

Table 1.5 Legal minimum carapace length (mm) for male and female Jasus edwardsii in Western Australia (WA), South Australia (SA), Tasmania, Victoria and New South Wales (NSW) Locationa

Males

Females

WA SA (S zone) SA (N zone) Tasmania

98.5 98.5 102 110

98.5 98.5 102 105

Victoria NSW

110 110

105 105

a

Sources as for Table 1.2.

1.4.4

Current status of the J. edwarsdii stocks

Throughout its range, J. edwardsii is considered to be fully exploited. There is currently no evidence that the breeding stock has been reduced to a level that is affecting recruitment to the fishery; however, levels of egg production have declined to low levels in most areas of the fishery. Egg production in Tasmania is considered to be below 10% of pristine in the northern areas, but above 80% in some parts of the southern ground (Punt & Kennedy, 1997). In Victoria egg production is considered to lie between 6 and 19% and in South Australia between 10 and 20% of Table 1.6 Rules governing the recreational fishery for Jasus edwardsii in Western Australia (WA), South Australia (SA), Tasmania, Victoria and New South Wales (NSW) Regulation

WA

SAa

Tasmaniab

Victoriac

NSWd

Possession limit Fishing gear

16 per boat

8 per boat

10

No limit

2

2 traps

1 trap, 4 ring nets

No traps, 2 ring nets

8

2 traps or 3 ring/hoop/drop nets 4

5

4

2

Snorkel

Snorkel

Snorkel

Snorkel

Snorkel

Equipment

SCUBA Hookah Hand only



Hand only

Bag limit

8

SCUBA Hookah No pointed objects 4

5

4

2

Types and no. per person Bag limit Diving methods Air supply

Source: aPrescott (pers. comm.), bKennedy, Gardner & Ford (pers. comm.), cMolloy, Hobday & Flint (pers. comm.), dMontgomery (pers. comm.).

62


the historic unfished situations (Hobday et al., 1998; Prescott, pers comm.). Longterm trends in egg production are being monitored by researchers in South Australia, Tasmania and Victoria to ensure that the already depleted state of the brood stock is not further eroded. 1.4.5

Current research

Routine commercial catch and length±frequency monitoring data, together with large-scale southern lobster tag release and recapture projects, have provided the basis for considerable stock-assessment modelling work undertaken in South Australia (McGarvey et al., 1997), Victoria (Treble, 1996; Hobday et al., 1998) and Tasmania (Frusher et al., 1997; Punt & Kennedy, 1997; Punt et al., 1997). Much of this work has focused on the calculation of exploitation rates in the lobster fisheries of the southern states, as this is crucial to future management strategies. In addition, work has been undertaken in all southern states to establish the state of egg production and in two of the states (South Australia and Tasmania) on the estimation of size transition matrices for the lobster populations. A large-scale programme to monitor the level of puerulus settlement across the southern states of South Australia, Victoria and Tasmania is being undertaken to obtain an understanding of the spatial and temporal distribution of pueruli. Later, this may provide a mechanism for predicting the commercial catch in advance of the event and an indication of factors contributing to annual variation. The longer-term trends in the index of puerulus abundance may also provide the first indication of any stock recruitment problems, i.e. a reduction in the size of the breeding stock to the point where it results in a reduction in the level of puerulus settlement. Data to date show marked seasonal and interannual patterns in settlement for different regions (Frusher et al., 1997). 1.4.6

Conclusions

Jasus edwardsii is fully exploited throughout its range and in most areas the breeding stock is considered to be well below 20% of the levels in the fishery prior to exploitation. There are no indications to date that recruitment is being affected by the size of the brood stock. Most of the fishery (southern South Australia, Tasmania) is now under a TACC form of management and consideration is being given to implementing this form of management in Victoria (Hobday, pers. comm.).

1.5

The western rock lobster (Panulirus cygnus) fishery

The western rock lobster (P. cygnus) now supports the largest rock lobster fishery in the world, with seasonal catches averaging 10 800 t over the past 19 years (1980/81


63

to 1998/99) and the 1998/99 season producing a record 13 t. The 596 vessels licensed to participate in the fishery share between $200 and $300 million gross per season, making it Australia's most valuable single species fishery. Because of the high prices paid for P. cygnus and the good returns that a fisher can expect, the stock has experienced very high and increasing exploitation during the 1980s and 1990s. The stock now is fully exploited.

1.5.1


Panulirus cygnus is found in commercial quantities from just east of Cape Leeuwin (34 240 S) in the south to Shark Bay (24 450 S) in the north (Fig. 1.7). The juveniles populate the shallow (40 m), directly west and north-west of their juvenile reefs (Fig. 1.7), to where the bulk of the breeding stock is situated (Morgan, 1977; Phillips, 1983; Chubb et al., unpubl.). Some of the migrating lobsters continue past the deep-water breeding habitat to the edge of the continental shelf, combine with lobsters moving from deeper water areas where they probably settled as pueruli and then move northwards, broadly following the 140±180 m depth contour (Fig. 1.7). As the migrating lobsters migrate north, some move back inshore to suitable habitat in depths of 40±100 m. Abrolhos Islands `white' lobsters are joined by some coastal `whites' on the `Big Bank run', where lobsters migrate northwards out of the Abrolhos zone (Fig. 1.7) (Anon., 1989). It is not known how far north the animals migrate, but large quantities of legal-size lobsters are taken during the migration and the mainly sublegal-size remnants may be in poor physical condition, in some seasons, by the time they are west of Shark Bay (Fig. 1.7) (Chubb et al., 1994; Chubb et al., unpubl.). One to 2 years after reaching legal size (i.e. about 6±7 years after hatching), the female lobsters of the coastal populations commence breeding at a CL of 90±100 mm (50% mature) (Chittleborough, 1976; Chubb, 1991). They are estimated to produce about 50% of total egg production (Chubb, 1991). At the Abrolhos Islands, females commence breeding well below the legal minimum size, with 50% being mature at 65 mm CL (Chubb, 1991). The reason for the smaller size at sexual maturity at the Abrolhos Islands is unknown, but it may be influenced by the very high densities of lobsters in this region and the fact that the Abrolhos is a very different habitat area from the coast, comprising coral atolls and reefs. The islands also are directly in the path of the warm tropical southward-flowing Leeuwin Current, which maintains significantly higher seawater temperatures than those experienced on the coast.

1.5.2


Aboriginal people were the first to use P. cygnus as a food source, to support their hunter±gatherer existence. A small commercial fishery developed soon after European settlement and in 1897 the first regulation for the fishery was proclaimed, namely a minimum legal size (weight), which has remained virtually unchanged to the present day. By the 1930s, a small fleet of boats was operating out of Fremantle and Geraldton (Fig. 1.7) supplying some 250 t annually to the local market. It was not until the end of World War II (1945), when the export of frozen lobster tails to the USA commenced, that exploitation began in earnest. The export of 1000 t in 1948 grew to 8000 t by 1958. Since then, the 10 seasons 1970/71 to 1979/80 produced average catches of 9250 t with an average of 10 800 t over the next 19 seasons (1980/ 81 to 1998/99). A record catch of 13 000 t was taken in 1998/99 with catches in

66


excess of 12 000 t on four occasions between 1987/88 and 1992/93 (see Caputi et al., chapter 18, for a table of catch and fishing effort). During the developmental years of the fishery (1945±1962) there were no restrictions on licences and gear and the number of new entrants, boats and pots (traps) grew rapidly. By the late 1950s and early 1960s, the fishing effort was still rising rapidly but the catch had stabilized at 7±8 000 t. Fishers saw their catch rates and individual catches declining and feared for their economic viability. In 1963, after representation from the fishing industry, the government introduced a policy of `limited entry' and closed the fishery to new entrants. In 1965 the number of pots that fishers could use also was strictly controlled, limiting the total number of pots to 76 623 and boats to 845. Since the introduction of limited entry, both nominal fishing effort (pot lifts) and the effectiveness of that effort continued to increase as fishers built larger vessels and worked more days per month, gaining experience and improving their gear and fish-finding technology. In an attempt to offset these increases in fishing effort and efficiency, management, in consultation with industry, introduced measures to shorten the fishing season, strictly define pot design and dimensions and reduce the number of pots and boats in the fishery (Brown, 1991; see Caputi et al., Chapter 18, for further details). These measures have been successful in slowing the rate of increase in effective fishing effort, but not stopping it. An active recreational fishery, operating in the inshore waters (5811

++ + 2 >7334

++ + 4 >8952

++ + 3 >8790

++ + 20 >10 115

Source: FAO (1998) plus other sources.a±d +, Modest catches; ++, substantial catches of unknown magnitude. a About 4 t/year around Zanzibar in 1973 (Mutagyera, 1975). b Pitcher (1993). c Australian Fisheries Service. d Zann (1984).

to the Indo-West Pacific region. Some significant producers have no recorded catches in the FAO statistics (FAO, 1998). Prescott (1988) suggested that catch rates in terms of catch per person-hour, either by swimming or wading along the reef edge at appropriate tides, should give reasonable estimates of relative abundance of stocks (other than P. ornatus). Depletion experiments conducted on relatively limited sectors of reefs could give some estimates of populations and biomass per kilometre of reef front but clearly would be inapplicable over wider areas of shelf with generalized cover of corals. None of the available parameter estimates is very securely based and, for example, is sufficiently divergent to have generated much uncertainty about the best management measures to adopt for the Tongan fishery (Prescott, 1990).

3.2

Conclusions

Spiny lobster catches in the tropical Indo-West Pacific are relatively small, but are a commercially important component of the catches of the small-scale fishers in the tropical Indo-West Pacific. The principal means of capture is by spearing but there is sufficient evidence of the vulnerability of P. penicillatus and P. longipes to traps to warrant further investigation.

96


Because of the relatively low volume and dispersed nature of landings, estimates of production in most countries are incomplete or non-existent and are probably gross underestimates in most cases. Few sustained attempts have been made to estimate the biological parameters regulating production. Consequently, apart from P. ornatus in the Torres Strait and Gulf of Papua (see references in Phillips et al., Chapter 1), no well-founded management plans are in place for any stock in the entire tropical Indo-West Pacific region.

References Berry, P.F. (1970) Mating behaviour, oviposition and fertilization in the spiny lobster, Panulirus homarus (Linnaeus). Invest. Rep. Oceanogr. Res. Inst., Durban, South Africa, 24, 16 pp. Berry, P.F. (1971) The spiny lobsters (Palinuridae) of the east coast of southern Africa: distribution and ecological notes. Invest. Rep. Oceanogr. Res. Inst., Durban, South Africa, 27, 23 pp. Bhatia, U. (1974) Distribution of spiny lobsters along the west coast of Thailand with observations on their fishing grounds. Phuket Mar. Biol. Ctr. Res. Bull., No. 5, 20 pp. Charbonnier, D. & Crosnier, A. (1961) Quelques donnees sur la peche des langoustes a Madagascar. Peche Marit., No. 994, 16±18. De Bruin, G.H.P. (1969) The ecology of spiny lobsters, Panulirus spp. of Ceylon waters. Bull. Fish Res. Stn. Ceylon, No. 20, 171±89. Ebert, .A. & Ford, R.F. (1986) Population ecology and fishery potential of the spiny lobster, Panulirus penicillatus at Enewetak Atoll, Marshall Islands. Bull. Mar. Sci., 38, 56±67. FAO (1998) Yearbook of fishery statistics. 1996 catches and landings. Food and Agriculture Organization of the United Nations, Rome (online database). Felfoldy-Ferguson, K. (1988) The collection and uses of inshore reef fisheries information to assess and monitor the shelf fisheries of the Kingdom of Tonga, using the ICLARM approach: summary of the first year's activities and results. South Pacific Commission Workshop on Pacific Inshore Fishery Resources. Noumea New Caledonia: March 1988. Background Paper No. 41, 13 pp. George, R.W. (1972) South Pacific islands rock lobster resources. FAO Rep. FI:RAS/69/102/9, 41 pp. Juinio, A.R. & Gomez, E.D. (1986) Spiny lobster fishery in eastern Samar, Philippines. In The First Asian Fisheries Forum. (Ed. by J.L. Maclean, L.B. Dizon & H.V. Hosillos), pp. 381±4. Asian Fisheries Society, Manila, Philippines. MacDonald, C. (1979) Management aspects of the biology of the spiny lobsters, Panulirus marginatus, P. penicillatus, P. versicolor and P. longipes femoristriga in Hawaii and the Western Pacific. Report to Western Pacific Regional Fishery Management Council, Univ. Hawaii. Acct. No. F-78±237-F-051-B-123, 126 pp. MacDonald, C.D. (1982) Catch composition and reproduction of the spiny lobster Panulirus versicolor at Palau. Trans. Am. Fish. Soc., 111, 694±9. Mohamed, K.H. & George, M.J. (1968) Results of tagging experiments on the Indian spiny lobster, Panulirus homarus ± movement and growth. Indian J. Fish., 15, 15±26. Munro, J.L. (1988) Growth and mortality rates and state of exploitation of spiny lobsters in Tonga. South Pacific Commission, Workshop on Pacific Inshore Fisheries Resources. Noumea New Caledonia: March 1988. Working paper No. 51, 34 pp. Mutagyera, W.B. (1975) A preliminary report on the spiny lobster fishery in Zanzibar. Afr. J. Trop. Hydrobiol. Fish., 4, 51±9

Fisheries for Spiny Lobsters in the Tropical Indo-West Pacific

97

Pitcher, R.C. (1993) Spiny lobster. In Nearshore Marine Resources of the South Pacific (Ed. by A. Wright & L. Hill), pp. 539±607. Institute of Pacific Studies, Suva, Fiji; Forum Fisheries Agency, Honiara, Solomon Islands & International Centre for Ocean Development, Canada. Prescott, J. (1980) Report on the South Pacific Commission Lobster Project in Solomon Islands. South Pacific Commission, Noumea, New Caledonia, 24 pp. Prescott, J. (1988) Tropical spiny lobster: an overview of their biology, the fisheries and the economics with particular reference to the double-spined rock lobster, P. penicillatus. South Pacific Commission, Workshop on Pacific Inshore Fishery Resources, Noumea, New Caledonia: March 1988. Working paper No. 18, 35 pp. Prescott, J. (1990) A survey of the lobster resources in the Ha'apai group, Kingdom of Tonga. South Pacific Forum Fisheries Agency, Rep. No. 90/93, 31 pp. Pyne, R.R. (1970) Tropical spiny lobsters Panulirus spp. of Papua and New Guinea. Search, 1, 248± 53. Rongmuangsart, S. & Luvira, O. (1973) Studies on the biology and population dynamics of the spiny lobster, Panulirus polyphagus (Herbst), of the West Coast of Thailand, with notes on experimental rearing of P. versicolor (Latrielle) in the laboratory. Phuket Marine Biological Center, Res. Bull. No. 2, 22 pp. Williams, A.B. (1988) Lobsters of the World. Osprey Books, New York, USA, 186 pp. Zann, L.P. (1984) A preliminary investigation of the biology and fisheries of the spiny lobsters (Palinuridae) in the Kingdom of Tonga. Institute of Marine Resources, University of the South Pacific Rep., 55 pp.


Chapter 4

The Lobster Fishery in the North-western Hawaiian Islands J.J. POLOVINA Honolulu Laboratory, Southwest Fisheries Science Center, National Marine Fisheries Service, NOAA, 2570 Dole Street, Honolulu, Hawaii 96822-2396, USA

4.1

Introduction

The North-western Hawaiian Islands (NWHI) are an isolated range of islands, islets, banks and reefs extending 1500 nautical miles north-west, from Nihoa Island to Kure Atoll (Fig. 4.1). Lobster concentrations in the NWHI were documented by research cruises in 1976 and commercial trapping had begun by 1977 (Uchida & Tagami, 1984). The fishery targets two species: the endemic spiny lobster Panulirus marginatus Quoy and Gaimard 1825 and the common slipper lobster Scyllarides squammosus Milne-Edwards 1837. Two other species ± the ridgeback slipper lobster S. haanii de Haan 1841 and the Chinese slipper lobster Parribacus antarcticus Lund 1793 ± are caught incidentally in low abundance. Since 1983, the lobster fleet has ranged from nine to 16 vessels (15±35 m long), each averaging three trips per year (Fig. 4.2). The vessels set about 800 traps per day and remain at sea for almost 2 months per trip. Landings and effort increased rapidly in the early 1980s to a maximum landing for spiny and slipper lobsters combined of about 2 million in 1985 with an effort of about 1 million trap-hauls (Fig. 4.3). However, by the late 1980s, as a result of the process of fishing down a previously unexploited population, combined spiny and slipper landings had dropped to about 1 million lobsters. In the early 1990s a change from open access to limited entry and harvest quota management, together with an environmental regime change which adversely impacted spiny lobster recruitment, has resulted in recent landings in the range of 200 000±300 000 lobsters (Fig. 4.3). The ex-vessel revenue of the fishery has ranged from about US $5±6 million in the late 1980s to US $1±2 million in the mid-1990s. For most of the 1980s and early 1990s the lobsters were landed and marketed as frozen tails, but beginning in 1997 a few boats explored landing and marketing live lobsters. Since 1988, 60±80% of the landings have been spiny lobster (Table 4.1). Fathoms Plus shellfish traps are used by all fishermen (Fig. 4.4). This trap is dome-shaped, single-chambered and made of moulded black polyethylene which measures 98 770 295 mm, with a mesh size of 45 45 mm (inside dimensions). Each trap has two entrance cones located on opposite sides. Each trap also has two escape vent panels each consisting of four circular vents, 67 mm in diameter, located on opposite sides to facilitate the escape of immature lobsters. The traps are typically baited with chopped mackerel (Scomber sp.), fished 98


Figure 4.1

99

The Hawaiian Archipelago, including the North-western Hawaiian Islands.

in strings of several hundred traps per string, and most frequently set in depths of 20±50 m.

Figure 4.2 A typical lobster vessel fishing in the North-western Hawaiian Islands.


Figure 4.3 Total lobster landings (millions of lobsters) and trapping effort (millions of traphauls) for the combined slipper and spiny lobster fishery in the North-western Hawaiian Islands, 1983±1997.

Figure 4.4 A lobster trap with an escape panel commonly used by the lobster fishery in the North-western Hawaiian Islands.


101

Table 4.1 Annual landings of spiny and slipper lobsters (thousands), trapping effort (thousand trap-hauls), CPUE (lobsters per trap-haul) and the percentage of spiny lobster in the landings, 1983±1990a Year

Spiny lobster

Slipper lobsterb

Total lobsters

Trapping effort

CPUE

Spiny lobster (%)

1983c 1984 1985 1986

158 677 1022 843

18 207 900 851

176 884 1902 1694

64 371 1041 1293

2.75 2.38 1.83 1.31

90 78 53 50

1987 1988 1989 1990

393 888 944 591

352 174 222 187

745 1062 1166 777

806 840 1069 1182

0.92 1.26 1.09 0.66

53 84 81 76

1991 1992 1993d 1994 1995

131 248 ± 85 34

35 163 ± 46 3

166 411 ± 131 37

295 685 ± 168 64

0.56 0.60 ± 0.79 0.58

79 60 ± 65 92

1996 1997

165 176

22 134

187 310

115 178

0.94 1.74

88 57

a Data were provided to the National Marine Fisheries Service as required by the Crustacean Fishery Management Plan of the Western Pacific Regional Fishery Management Council. b Slipper lobster landings for 1984±1987 are 72% of those reported, so they are comparable to landings after 1987 when a minimum size allowed the retention of about 72% of the catch. c April±December 1983. d Fishery closed.

4.2

Management

The NWHI lobster fishery has been managed under federal jurisdiction with a fishery management plan (FMP) administered by the Western Pacific Regional Fishery Management Council (WPRFMC) since 1983. The NWHI fishery is managed with a limited-entry system for a maximum of 15 vessels, an annual fleet harvest quota and a closed season from January to June to protect the spawning biomass before the summer spawning. An annual harvest quota is set for each region as 13% of the July estimated exploitable population (DiNardo & Wetherall, 1999). The constant harvest rate of 13% was selected based on a decision by managers that the corresponding annual fishing mortality have a probability of less than 0.10 of exceeding the fishing mortality, which results in a spawning potential ration of 20% (DiNardo & Wetherall, 1999). The region-wide quota is allocated geographically to prevent overfishing at the closest banks. There is no minimum size or prohibition on the retention of egg-bearing females. In fact, while all traps are required to have

102 Spiny Lobsters: Fisheries and Culture escape vents to reduce handling and release-induced mortality on immature lobsters, all lobsters which are caught are counted against the quota and must be landed. Finally, all vessels must submit logbooks recording daily catch and trapping effort. Currently, fishermen and managers are examining whether an individual quota would be an improvement over the current fleet quota.

4.3

Stock assessment

Stock assessment has used the annual catch of spiny and slipper lobsters and trapping effort data from the commercial logbooks since 1983 (Table 4.1). During 1983±1997, the proportion of spiny and slipper lobsters in the catches varied because of targeting by fishermen and variations in abundance. Trapping is a multispecies effort and logbooks do not specify when effort targets spiny or slipper lobster. Stock assessment of the lobster resource is hindered by the relatively short catch and effort time series, the inability to age lobsters and changes in management regulations. For example, the increase in catch per unit effort (CPUE) in 1996 and 1997 is primarily due to a management change eliminating a minimum size in favor of requiring the retention of all catch. However, the following population dynamics model has been used successfully since the early 1990s. The monthly total number of exploitable lobsters (Nt) is expressed as a function of the number of exploitable lobster in the previous month (Nt 1) adjusted for annual instantaneous natural mortality (m), monthly catch (Ct 1), and constant annual recruitment (R) as: Nt Nt 1 e

m=12

Ct

1

R=12:

4:1a

The model-based estimate of Nt was then converted to a CPUE value by multiplying by the catchability (q): CPUEt qNt :

4:1b

Model-based parameters (m, q, R) were estimated using an iterative non-linear least squares method that minimizes the residual sums of squares between observed and estimated monthly CPUE. For several years the season harvest quota was set for the entire fishing ground, computed as 13% of the total number of exploitable lobsters (Nt) in the fishing ground, estimated from this model for the beginning of the fishing season. However, beginning in 1998 the harvest quota was determined separately for each of four regions in the fishing ground by application of the assessment model on a regional basis. Estimated regional exploitable population sizes were obtained and regional harvest quotas assigned as 13% of those numbers. The population dynamics model finds evidence of a 50% drop in mean annual recruitment after 1990, consistent with atmospheric and oceanographic data and NWHI ecosystem data indicating a regime shift in the late 1980s (Polovina et al., 1994, 1995; Polovina & Haight, in press).

The Lobster Fishery in the North-western Hawaiian Islands 4.4

103

Research

After the initial research cruises documenting lobster concentrations in the NWHI in 1976, research focused on the biology of the spiny lobster P. marginatus. Tagging studies at Kure Atoll and French Frigate Shoals estimated a von Bertalanffy growth curve for growth (in carapace length) to have a parameter k of 0.31/year with an asymptotic carapace length of 13.2 cm (MacDonald, 1984). Also obtained was a mean natural mortality estimate of 0.37/year, along with estimates for the ages at the onset of sexual maturity (2.7 years for males and 1.7 years for females; MacDonald, 1984). In 1998 over 3000 spiny lobsters were tagged at Necker Island as part of a study to update estimates of population parameters. Trapping surveys mapped the spatial distribution of P. marginatus in the NWHI and indicated that the highest catch rates ranged from depths of 55±73 m in the south-eastern portion of the NWHI to 19±54 m in the north-western portion of the Hawaiian Archipelago (Uchida & Tagami, 1984). Research conducted during 1984±1987 developed escape vents to reduce the catch and hence mortality of sublegal spiny lobster ( 75 mm CL) at about 5% of its preexploitation level, and spawning biomass (females > 65 mm CL) at some 20% of its believed pristine level. Of major concern is that the decrease in spawning biomass might be associated with a decline in recruitment, and hence future sustainable yields. Model results tend to confirm this fear, suggesting that recruitment during recent decades has been substantially depressed in comparison with the average for the pristine stock, presumably as a result of the smaller parent population. This implies that the drop in biomass since the mid-1980s may have resulted in a further decline in recruitment. To compensate for this, scientific consensus stipulated that any medium-term strategy for the utilization of J. lalandii should incorporate a stock rebuilding strategy. The culmination of the extensive modelling conducted during the mid-1990s was the development of an operational management procedure (OMP). The OMP is a standardized procedure for incorporating and analysing data to be used when setting annual TACs. Revolving around a relatively simple mathematical formula, the OMP requires as input the previous year's TAC and indices of commercial CPUE, FIMS CPUE and somatic growth rate. Following comprehensive robustness tests and consultation with both the industry and other role players, this procedure was accepted and implemented for the 1997/98 assessment. The OMP fulfils many of the precautionary principle guidelines (Cockcroft & Payne, 1999) and in its aims includes a 20% increase in resource biomass by the year 2006, thereby satisfying the need for stock rebuilding. The management problems caused by low growth were compounded during the review period by a series of rock lobster mass mortalities resulting from localized strandings along the South African west coast. These events were all associated with the decay of intense phytoplankton blooms and ranged in magnitude from a 60-t stranding in 1994 (Matthews & Pitcher, 1996) to a 2000-t stranding over a protracted period in April and May 1997. The latter was the worst rock lobster mortality ever recorded in South Africa (Cockcroft et al., 2000) and was followed by a further 200-t stranding at the same location in April, 1999 (Marine and Coastal Management, unpubl. data). Notwithstanding the previous discussion, the recent event that has had the most profound ramifications both for the rock lobster industry and for management fora, has been the implementation of a new fisheries policy for South Africa, the Marine Living Resources Act of 1998. This Act aims to maintain sustainability and industrial stability, while introducing a far greater level of equity than was previously apparent (Cockcroft & Payne, 1999). Although it is too early to evaluate the full impact of these changes, ongoing monitoring programmes have been instituted for this purpose.

114 Spiny Lobsters: Fisheries and Culture 5.5

The South Atlantic Fishery for Jasus tristani

Jasus tristani is endemic to the Vema Seamount and to the isolated mid-ocean islands of Gough, Tristan da Cunha, Nightingale and Inaccessible in the South-east Atlantic (Fig. 5.1). While the small resource at the Vema Seamount was rapidly depleted after the inception of the fishery there in the early 1960s (Pollock, 1994), catches from the other populations remain substantial. Although British authorities provide management advice, the Islands are largely self-sufficient and depend heavily on revenue generated by the rock lobster fishery. In his most recent review of the fishery for J. tristani, Pollock (1994) suggested that this resource could sustain annual catches of approximately 450 t, given the TAC regulations introduced during 1991, the minimum legal size of 70 mm CL and restrictions on gear. However, since that time, the TAC has infrequently been landed in full and catches have declined somewhat, reaching 311 t in the 1997/98 season (Table 5.1). Nevertheless, this does not necessarily reflect a decline in population strength; the TACs have stabilized at about 323 t and the size-composition data have not recently displayed the marked diminution evident during the early phases of the fishery (Pollock, 1994). Instead, the influence of weather patterns on fishing behaviour and the change to a new concession holder in 1997, with a concomitant modification of the fishing season, have combined to confound the conventional indicators of fishing performance. The perception of scientists involved with research on this resource is that stocks are gradually recovering (James Glass, Head of Natural Resources, Tristan da Cunha, pers. comm.).

5.6

The South African trap-fishery for Palinurus gilchristi

Palinurus gilchristi is endemic to the southern coast of South Africa (Fig. 5.1), where it occurs on rocky substrata inshore of the shelf break between Cape Point (18 E) and East London (28 E). Locally known as the south coast rock lobster, it is found in commercial densities along the coast and up to 250 km offshore on the Agulhas Bank. It is a deep-water species and is fished between depths of 50 and 200 m. Females generally carry eggs during the austral winter (June±October), when the bottom water temperature is somewhat warmer than it is in summer (Groeneveld & Rossouw, 1995). After the eggs hatch in spring (October±November), the majority of both male and female rock lobster undergo their annual moult (November± February). However, the frequency of moulting is size dependent, and small specimens (20 years) and relatively slow-growing species (Groeneveld, 1997). Growth rates are faster on the western fishing grounds than on fishing grounds east of 27 E, a pattern mirrored by both the overall average size and the average size of female rock lobster at sexual maturity (Table 5.3). These trends are thought to be caused by a gradient in competition for limited resources, which results from the higher rock lobster abundance in the east, coupled with the restricted space available on the narrower continental shelf in that area. Commercial exploitation of P. gilchristi commenced in 1974 and expanded rapidly during the following 2 years. With numerous local and foreign fishing vessels converging on the newly discovered fishing grounds, catches had escalated to 2000 t by 1975 (Pollock & Augustyn, 1982). The subsequent recognition of P. gilchristi as a sedentary species of the continental shelf, belonging to South Africa alone, forced the withdrawal of foreign fishing vessels from the South African EEZ in 1976. Many of the remaining local fishing vessels were later forced to withdraw from the fishery when catches declined suddenly to 260 t in 1979/80. The resulting period of reduced effort allowed the resource (and catches) to recover somewhat (Fig. 5.4). Although the initial phases of the fishery were regulated only by limiting the number of traps permitted per vessel, an annual TAC was eventually introduced as a management tool in 1984 and trap limitations were abolished in 1988. Being based primarily on the recent performance of the fishery, the TAC remained stable at approximately 1025 t per year until the 1993/94 fishing season. During 1994, a more rigorous procedure was developed for the assessment of the resource. This used a Bayesian approach to fit a surplus production model to Table 5.3 Size at sexual maturity (mm CL), average size (mm CL) and growth parameters for Palinurus gilchristi on the western (Agulhas Bank, St Francis Bay and Port Elizabeth) and eastern (Port Alfred) fishing grounds

Size at 50% maturity: Presence of ovigerous setae

Western grounds

Eastern grounds

Male

Male

Female

Source

Female

±

65

±

59

±

71

±

62

Groeneveld & Melville-Smith (1995)

Presence of eggs Average CL (mm): 1978±1980 1988±1992

84.3 ±

82.9 75.3

69.5 ±

67.5 69.4

Pollock & Augustyn (1982) Groeneveld & Roussouw (1995)

Growth parameters: L1 (mm)

111.2

96.1

95.8

78.5

Groeneveld (1997)

K/year

0.092

0.129

0.050

0.065

Groeneveld (1997)


Fig. 5.4 Trends in catch and relative indices of effort and CPUE from the Palinurus gilchristi fishery (effort and CPUE were normalized from set hours and kg/trap, respectively).

commercial catch and effort data, growth parameters from a long-term tagging programme and catch-at-size information. Both CPUE trends and the surplus production model indicated a continuous decline in resource biomass since the 1989/ 90 fishing season (Fig. 5.4). In response, a programme of phased TAC reductions was initiated in 1994/95, resulting in a TAC for the 1997/98 season of 865 t. Unfortunately, these reductions failed to impact on the trend of declining CPUE, which now stands at 50% of its value in 1989/90 (Fig. 5.4). Although CPUE is often used directly as an index of relative abundance, it is possible that the relationship between these variables may be obscured by changes in fishing techniques. In 1974, fishing gear comprised individually buoyed traps deployed by fairly small vessels, which could remain at sea for limited periods. The subsequent evolution of a long-line system (which employs up to 200 plastic traps attached to each of several bottom lines), the introduction of larger vessels (up to 60 m in length), the deployment of increasing numbers of traps for longer periods (Fig. 5.4), and the improvement in navigational equipment have substantially increased the efficiency of fishing operations. As a result, although the mass of rock lobster landed per trap (CPUE) may have decreased, the total catch is made more cheaply. This may, at least in part, explain the continual decline in CPUE. To investigate this possibility, an experimental trap-reduction programme was designed and implemented for the 1998/99 commercial season. The programme is pivotal to the fishery; should results be negative, further reductions in the TAC will be necessary.

Fisheries in the South-east Atlantic and South-west Indian Oceans

117

The trap and trawl fisheries for Palinurus delagoae off South Africa and Mozambique

5.7

The distribution of P. delagoae (Fig. 5.1) stretches southwards along the eastern African coast from northern Mozambique (17 S) to southern KwaZulu-Natal (32 S), South Africa, including the south-eastern coast of Madagascar between these latitudes (Holthuis, 1991; Cockcroft et al., 1995). Like P. gilchristi, P. delagoae occurs in a temperate (12 C), deep-water (100±600 m) environment, and is slow growing (K = 0.059 0.076/year; L1 = 155 ± 165 mm CL) and long lived (M = 0.09 ± 0.15/year), where K and L1 are parameters of the Von Botalouffy equation, and M is the instantaneous rate of natural mortality (Groeneveld, in press). The population is spatially segregated according to size, with small specimens ( 3 t of rock lobster in any year. Of these, 15 LFRs process > 150 t annually. Rock lobster for export must processed and packed in premises holding Export Fish Packhouse certificates issued by the Ministry of Agriculture (MAFQual). LFRs are subject to self-audit of product flow reported to Ministry of Fisheries Compliance division. MFish Compliance routinely conducts intensive inspection and audit to ensure adherence to the reporting requirements of the QMS. LFRs have strict liability to ensure that lobsters are purchased only from legitimate commercial fishermen and that all lobsters meet regulatory requirements in terms of MLS, condition, hygiene and health and safety issues. The export of rock lobster is deregulated and in addition to lobster processors there is a small number of export brokers and marketers offering their services to the New Zealand industry. The New Zealand Rock Lobster Exporters Association (NZRLEA) membership comprises the majority of long established lobster processing and exporting companies. The New Zealand Seafood Industry Council (SeaFIC) issues fish Export Licences (FELs).

36.5

Pack styles

Lobsters are processed into a number of different forms and pack styles. Live Lobster: lobsters are held alive, typically for 3 days, in flow-through or recirculating seawater holding tanks. This allows the lobster to purge their gut contents, and processors to cull those that will not survive packing and transport. The lobsters are immobilized prior to packing by lowering their body temperature. They are graded and packed in 8-kg, 9-kg or 10-kg net weight cartons, usually with a 1-2% margin to allow for weight loss in transit. Larger lobsters are packed in random-weight cartons. Whole raw lobster (WRL), to be packed as WRL lobster must be alive when entering the process. Lobsters are anaesthetized and suffocated in iced fresh water, graded, wrapped and packed in 10-kg cartons. The cartons are blast frozen to -4O"C, and stored below -25°C. They are shipped in sea containers, or occasionally by air. Whole cooked lobster (WCL): unique to Western Australia, the process is similar to that for WRL, other than the lobster are boiled in heavily salted water for between 8 and 20 min, depending on size, then rapidly chilled and thoroughly washed and drained before wrapping, packing and freezing. WCL are also consigned in chilled form, by air, particularly to France. Lobster tails: the first export product for rock lobster, lobster tails are processed from lobster that are anaesthetized and suffocated in iced fresh water. The head is removed, and the hindgut vacuum extracted. The tails are then thoroughly washed, drained, graded and packed and frozen as above. As tails are consigned

648 Spiny Lobsters: Fisheries and Culture almost exclusively for the US market they are packed in cartons in Imperial weights of 10 lb (4.54 kg) and 25 lb (11.34 kg). Lobster heads: a byproduct from tail production; the heads are not washed or wrapped, but graded and placed in a 10-kg lined carton and frozen. Paste: this is another byproduct from tail production. Lobster heads are finely macerated and the resultant paste is plate-frozen or air-blast-frozen into blocks. Lobsters are delivered by either air or sea freight. Live lobsters are routinely airfreighted, and will survive comfortably for up to 30 h with minimal (

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