cephalopods of the world vol 1 nautilus, sepioids

7/24/2019 Cephalopods of the World Vol 1 Nautilus, Sepioids

1/294

FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1

ISSN 1020-8682

CEPHALOPODS OF THE WORLD

AN ANNOTATED AND ILLUSTRATED CATALOGUE OF

SPECIES KNOWN TO DATE

Volume 1. Chambered Nautiluses and Sepioids(Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae)


2/294


3/294


4/294

The designations employed and the presentation of materialin this information product do not implythe expression of anyopinion whatsoever on the part of the Food and AgricultureOrganization of the United Nations concerning the legalstatus of any country, territory, city or area or of itsauthorities, or concerning the delimitation of its frontiers orboundaries.

All rights reserved. Reproduct ion and dissemination of material in this

information product for educational or other non-commercial purposes are

authorized without any prior written permission from the copyright holders

provided the source is fully acknowledged. Reproduction of material in this

information product for resale or other commercial purposes is prohibited

without written permission of the copyright holders. Applications for such

permission should be addressed to the Chief, Publishing Management

Service, Information Division, FAO, Viale delle Terme di Caracalla, 00100

Rome, Italy by e-mail to [email protected]

FAO 2005

ISBN 92-5-105383-9


5/294

The First Volume of this New Edition of the Cephalopods of the World Catalogue

is warmly dedicated to the memory of

Kir N. Nesis

an accomplished scientist and teuthologist,

a true gentleman

and a cherished friend and colleague.


6/294


7/294

PREPARATION OF THIS DOCUMENT

This document has been prepared by the Marine Resources Service, Fishery Resources Division, FAO FisheriesDepartment. It is part of the regular programme activities and a partial fulfilment of the Organization's role with regards

to the marine fisheries resources identification and biodata (FAO Programme Element 232A3). It received support throughcontributions from the Ministry of Agriculture and Forestry Policies of the Government of Italy and from the Ministry of

Foreign Affairs of the Kingdom of Norway to the FAO Global Partnerships for Responsible Fisheries (FISCODE).This publication is the first of three volumes of the second edition of the original FAO Catalogue of Cephalopods of theWorld (Roperet al., 1984), and it constitutes Volume I of Number 4 in the new series:FAO Species Catalogue for FisheriesPurposes, that evolved as an independent series in 2001 from the formerFAO Fisheries SynopsisNo. 125.

Because the new Catalogue has expanded apace with recent research and fisheries information and revisions, it now isnecessary to publish it as three free-standing volumes. Each volume has separate pagination, terminology/glossary,systematic sections, list of species and a volume-specific bibliography. This allows readers to use each volumeindependently without having to consult the other volumes for technical terms, measurements or bibliographic purposes.We hope that this added flexibility will provide convenience and utility for users of the Catalogue.

Programme managers: Jordi Lleonart and Michel Lamboeuf (FAO, Rome).

Scientific and technical editors: Patrizia Jereb (ICRAM, Rome) and Clyde F.E. Roper (Smithsonian Institution, NMNH,Washington DC, USA).

Scientific reviser:Nicoletta De Angelis (FAO, Rome).

Technical, editorial assistance: Ingrid Roper (Smithsonian Institution, NMNH, Volunteer, Washington DC, USA).

Scientific assistance:Michael J. Sweeney (New Mexico State University, Las Cruces, New Mexico, USA).

Scientific illustrator:Emanuela DAntoni (FAO, Rome).

Page composition and indexing:Michle S. Kautenberger-Longo (FAO, Rome).

Digitization of distribution maps:Fabio Carocci (FAO, Rome) and Elena V. Orlova (St Petersburg, Russia).

Cover illustration:Emanuela DAntoni (FAO, Rome).

Distribution

Authors

FAO Fisheries OfficersRegional Fisheries Councils and CommissionsSelector SC

Cephalopods of the World v

Jereb, P.; Roper, C.F.E.(eds)

Cephalopods of the world. An annotated and illustrated catalogue of cephalopod species known to date. Volume 1.Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae).

FAO Species Catalogue for Fishery Purposes. No. 4, Vol. 1. Rome, FAO. 2005. 262p. 9 colour plates.

ABSTRACT

This is the first volume of the entirely rewritten, revised and updated version of the original FAO Catalogue ofCephalopods of the World (1984). The present Volume is a multiauthored compilation that reviews six families:Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidaeand Spirulidae, with 23 genera and the 201 speciesknown to the date of the completion of the volume. It provides accounts for all families and genera, as well as

illustrated keys to all taxa. Information under each species account includes: valid modern systematic name andoriginal citation of the species (or subspecies); main synonyms; English, French and Spanish FAO names for thespecies; illustrations of dorsal and ventral aspect of the whole animal (as necessary) and other distinguishingillustrations; field characteristics; diagnostic features; geographic and vertical distribution, including GIS map; size;habitat; biology; interest to fishery; local names when available; a remarks section (as necessary) and literature. Thevolume is fully indexed and also includes sections on terminology and measurements, an extensive glossary, anintroduction with an updated review of the existing biological knowledge on cephalopods (including fisheriesinformation and catch data for recent years) and a dedicated bibliography.


8/294

vi FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1

For bibliographic reference the different sections should be quoted as follows:

Jereb, P.2005. Family Nautilidae. InP. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated and illustrated

catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae,Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes. No. 4,

Vol. 1. Rome, FAO. pp. 5155.

Jereb, P., Roper, C.F.E. & Vecchione, M.2005. Introduction.InP. Jereb & C.F.E. Roper, eds.Cephalopods of the world. Anannotated and illustrated catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids

(Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for FisheryPurposes. No. 4, Vol. 1. Rome, FAO. pp. 119.

Reid, A. 2005. Family Sepiadariidae. In P. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated and illustratedcatalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae,

Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes. No. 4,


Reid, A.2005. Family Idiosepiidae. InP. Jereb & C.F.E. Roper, eds.Cephalopods of the world. An annotated and illustrated

catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae,Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes. No. 4,


Reid, A.2005. Family Spirulidae.In P. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated and illustrated

catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae, Sepiidae,

Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes. No. 4,Vol. 1. Rome, FAO. pp. 211212.

Reid, A. & Jereb, P.2005. Family Sepiolidae.InP. Jereb & C.F.E. Roper, eds. Cephalopods of the world. An annotated andillustrated catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids (Nautilidae,

Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery Purposes.

No. 4, Vol. 1. Rome, FAO. pp. 153203.

Reid, A., Jereb, P. & Roper, C.F.E.2005. Family Sepiidae.In P. Jereb & C.F.E. Roper, eds.Cephalopods of the world. An

annotated and illustrated catalogue of species known to date. Volume 1. Chambered nautiluses and sepioids(Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae). FAO Species Catalogue for Fishery

Purposes. No. 4, Vol. 1. Rome, FAO. pp. 57152.


9/294

Acknowledgements

The authors are pleased to heartily acknowledge the contributions of colleagues who have supplied information or readdrafts of this Second Edition of the Cephalopod Catalogue. Without their good efforts this work would have been a lesscomprehensive, and consequently a less useful tool.

In particular, for this Volume, we want to thank: Eero Aro (Finnish Game and Fisheries Research Institute, Helsinki,Finland), Eduardo Balguerias (Centro Oceanografico de Canarias-IEO, Santa Cruz de Tenerife, Spain), GiambattistaBello(Acquario Provinciale, Bari, Italy),Alexei Birkun (Crimean State Medical University, Simferopol, Ukraine), TeresaBorges (Centre of Marine Sciences, University of Algarve, Lisbon, Portugal),Julia Filippova (Russian Federal Institute ofMarine Fisheries and Oceanography, Moscow, Russia),Ian Gleadall(Tohoku Bunka Gakuen University, Sendai, Japan),Angel Guerra (Instituto de Investigaciones Marinas, Vigo, Spain), Eugenia Lefkaditou (Institute of Marine BiologicalResources-NCMR, Athens, Greece),Marek Lipinski (Marine and Coastal Management-DEAT, Cape Town, South Africa),Piero Mannini(Marine Resources Services-FIR, FAO, Rome, Italy), Joanne Murphy(University of Aberdeen, Aberdeen,Scotland, UK), Chingiz Nigmatullin (Atlantic Research Institute of Fisheries and Oceanography, Kaliningrad, Russia),Joo Pereira(Instituto Nacional de Investigao Agrria e das Pescas, Lisbon, Portugal), Sergio Ragonese(Istituto perlAmbiente Marino CostieroCNR, Mazara del Vallo, Italy), Francisco Rocha (Instituto de Investigaciones Marinas, Vigo,Spain), Alp Salman (Ege University, Izmir, Turkey), Bruce Saunders (Bryn Mawr College, Bryn Mawr, Pennsylvania,USA), Ignacio Sobrino (Centro Oceanografico de Cdiz-IEO, Cadiz, Spain), Tooraj Valinassab (Iranian FisheriesResearch Organization, Teheran, Iran),Peter Ward (University of Washington, Seattle, USA).

Very special thanks are due to: Sigurd von Boletzky(Observatoire Ocanologique de Banyuls-CNRS, Banyuls-sur-Mer,France) for his knowledgeable help in solving some systematic problems; Peter Boyle, for his valuable editing of ourIntroductory Review;Renata Boucher(Musum National dHistoire Naturelle-CNRS, Paris, France) and Pilar Sanchez(Instituto de Ciencias del Mar, Barcelona, Spain) for their help with the French and Spanish vernacular names;NeethirajanNeethiselvan (Fisheries College and Research Institute, Tamil Nadu, India) who so kindly provided information andphotographic material on some Indian species;Mark Norman (Museum Victoria, Melbourne, Australia) for his invaluablehelp and support during many phases of this work; Takashi Okutani (Japan Marine Science and Technology, Yokouka City,Japan) for his knowledgeable support in reviewing and completing the sections on Japanese species; Martina Roeleveld(South African Museum, Cape Town, South Africa) who so kindly supplied information, data and photographic material onsome southern African species; Michael J. Sweeney (New Mexico State University, Las Cruces, New Mexico, USA) fortechnical support, nomenclatural listings, sources of information and literature searches.

Pere Oliver(Instituto Espaol de Oceanografa, Palma de Mallorca, formerly of FAO, Rome) initiated this revised edition,followed by Michel Lamboeuf (FAO, Rome) and Jordi Lleonart (FAO, Rome), who were our contacts at FAO. This

publication would not have been forthcoming without their very much appreciated support and encouragement. Weespecially thankMichel Lamboeuf for his help with the use of the FAO FISHSTAT database and for his revision of thegraphic sections thereafter.

Of course, a compilation of this nature must rely heavily on already-published works; we acknowledge these publicationswith gratitude. In particular, we acknowledge here that many illustrations from these works have been used for thepurposes of this Catalogue, for which we are most appreciative.

We also acknowledge with deep thanks the members of the FAO technical staff who so efficiently contributed to thepreparation of this Volume: Emanuela DAntoni for her excellent job in creating the many illustrations needed for theCatalogue and for greatly enhancing many illustrations from the literature; Nicoletta De Angelis and MichleKautenbergerfor their skilful collaboration in completing this highly technical and complex document andFabio Caroccifor his help in preparing the distribution maps.

Colour photographs are included in this new version of the Catalogue to enrich the quality and utility of the book. Thereforewe acknowledge with gratitude those who contributed with great generosity by offering photographic material:M. Demestre,J.W. Forsythe,M. Lamboeuf,J. Lleonart,M. Norman,W.F. Rathjen,P. Sanchez and W.B. Saunders.

Last, but not least, very special thanks are due to Ingrid H. Roper, for her technical and editorial assistance and invaluablesupport during many stages of the preparation of this Catalogue.

Cephalopods of the World vii


10/294

viii FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1

TABLE OF CONTENTS

PREPARATION OF T HIS DOCUMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v

1. INTRODUCTION. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

1.1 Plan of the Catalogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

1.2 General R emarks o n Cephalopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3

1.3 Interest to Fishery a nd Role in the Ecosystem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

1.4 IllustratedGlossary ofTechnicalTerms and Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

1.5 Key t o R ecent C ephalopod G roups and Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

2. CHAMBERED NAUTILUSES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50

Family N AUTILIDAE d e Blainville, 1 825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

Nautilus macromphalus Sowerby, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52

Nautilus pompilius Linnaeus, 1 758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53

SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLYFEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Nautilus belauensis Saunders, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54Nautilus repertus Iredale, 1 944 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Nautilus stenomphalus Sowerby, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54

Allonautilus scrobiculatus (Lightfoot, 1 786) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

Allonautilus perforatus (Conrad, 1 847) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55

3. CUTTLEFISHES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56

Family S EPIIDAE K eferstein, 1 866 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

Key t o genera i n the f amily S epiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

Metasepia pfefferi (Hoyle, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

Metasepia tullbergi (Appellf, 1 886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61

SepiaaculeataVan Hasselt, 1835 (in Frussac and dOrbigny, 18341848) . . . . . . . . . . . . . . . . . 63

SepiaandreanaSteenstrup, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65

SepiaapamaGray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67

Sepiaarabica Massy, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

SepiaaustralisQuoy and G aimard, 1 832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70

SepiabandensisAdam, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72

SepiaberthelotidOrbigny, 1835 (in Frussac and dOrbigny 18341848) . . . . . . . . . . . . . . . . . . 73

Sepiabraggi Verco, 1 907 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75

Sepia brevimanaSteenstrup, 1 875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77

SepiacultrataHoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78

Sepiaelegans Blainville, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

SepiaellipticaHoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81

SepiaelobyanaAdam, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82

SepiaesculentaHoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84

Sepia grahamiReid, 2001. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86

Sepiahedleyi Berry, 1 918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87

Sepia hierreddaRang, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88

SepiakobiensisHoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89

Sepia latimanusQuoy a nd G aimard, 1 832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91

Sepia longipesSasaki, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93

Sepia lorigera Wlker, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94

Sepia lycidas Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95

SepiamadokaiAdam, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97

Sepia murrayiAdam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98

Sepiaofficinalis Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99

SepiaomaniAdam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101

Sepiaopipara (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102


11/294

SepiaorbignyanaFrussac in dOrbigny, 1 826 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103

SepiapapuensisHoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105

SepiapharaonisEhrenberg, 1 831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106

Sepiaplangon Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109

Sepiaprabahari Neethiselvan and Venkataramani, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . 110

SepiaprashadiWinckworth, 1 936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111Sepiaramani Neethiselvan, 2 001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113

Sepia recurvirostraSteenstrup, 1 875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114

Sepiarozella (Iredale, 1 926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116

SepiasavignyiBlainville, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117

Sepia smithi Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118

Sepia stelliferaHomenko and Khromov, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119

Sepiasulcata Hoyle, 1 885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120

Sepia trygonina (Rochebrune, 1 884) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122

Sepia vermiculata Quoy and G aimard, 1 832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124

SepiavietnamicaKhromov, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125

Sepia vossi Khromov, 1 996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127Sepiawhitleyana (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128

Sepiazanzibarica Pfeffer, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129

Sepiella inermis (Van Hasselt, 1835) (in Frussacand dOrbigny, 18341848) . . . . . . . . . . . . . . . 130

Sepiella japonicaSasaki, 1 929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132

Sepiella ornata (Rang, 1837) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134

SepiellaweberiAdam, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135

SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLYFEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

SepiaacuminataSmith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

SepiaadamiRoeleveld, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

SepiaangulataRoeleveld, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

SepiaappellofiWlker, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137

Sepia aureomaculataOkutani and H orikawa, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138

Sepiabartletti (Iredale, 1954) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138

SepiabathyalisKhromov, Nikitina and Nesis, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138

Sepia baxteri (Iredale, 1940) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138

Sepiabidhaia Reid, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

SepiaburnupiHoyle, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

SepiacarinataSasaki, 1 920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

Sepia chirotremaBerry, 1 918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139

Sepiaconfusa Smith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140

SepiacottoniAdam, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140

SepiadollfusiAdam, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140

SepiadubiaAdam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140

SepiaelongatadOrbigny, 18391842 (in Frussacand dOrbigny18341848). . . . . . . . . . . . . . . 141

Sepia erostrataSasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

Sepia faurei Roeleveld, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

Sepia foliopezaOkutani and Tagawa, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141

SepiagibbaEhrenberg, 1 831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

Sepiahieronis (Robson, 1924) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

Sepia incerta Smith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

Sepia insignisSmith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142

Sepia irvingiMeyer, 1 909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

Sepia ivanovi Khromov, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

Sepia joubini Massy, 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

Sepiakiensis Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143

Cephalopods of the World ix


12/294

x FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1

SepiakoiladosReid, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144

Sepia limata (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144

Sepia mascarensis Filippova a nd K hromov, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144

Sepiamestus Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144

Sepia mira (Cotton, 1932) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

SepiamirabilisKhromov, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145Sepia novaehollandiaeHoyle, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

SepiapapillataQuoy a nd G aimard, 1 832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145

SepiapardexSasaki, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

Sepia peterseniAppellf, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

SepiaplanaLu and Reid, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

Sepia plathyconchalisFilippova a nd Khromov, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146

Sepiapulchra Roeleveld and Liltved, 1985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

Sepia reesiAdam, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

Sepiarhoda (Iredale, 1 954) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

Sepiarobsoni (Massy, 1927) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147

SepiasayaKhromov, Nikitina and Nesis, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148SepiasentaLu and Reid, 1 997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

SepiasewelliAdam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

SepiasimonianaThiele, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148

Sepiasokotriensis Khromov, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

SepiasubplanaLu and Boucher-Rodoni, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Sepiasubtenuipes Okutani and H orikawa, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Sepia talaKhromov, Nikitina and N esis, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149

Sepia tanybracheia Reid, 2 000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

Sepia tenuipes Sasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

Sepia thurstoniAdam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

Sepia tokioensisOrtmann, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150

Sepia tuberculata Lamarck, 1 798 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

Sepia typica (Steenstrup, 1 875) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

SepiavercoiAdam, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

Sepiella cyanea Robson, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151

SepiellamangkangungaReid and Lu, 1998 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

Sepiella ocellataPfeffer, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

Sepia filibrachia Reid and Lu, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152

Family S EPIOLIDAE L each, 1 817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153

Keyto subfamiliesand genera in the familySepiolidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153

Subfamily S EPIOLINAE Appellf, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158

Sepiola affinis Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158Sepiola atlanticaOrbigny, 1 8391842 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159

Sepiola birostrataSasaki, 1 918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161

Sepiola intermediaNaef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162

Sepiola ligulata Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163

Sepiola parvaSasaki, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165

Sepiola robusta Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166

Sepiola rondeletiLeach, 1834 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167

Sepiola trirostrataVoss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169

Euprymnaberryi Sasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170

Euprymnamorsei (Verrill, 1 881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171

Euprymna tasmanica (Pfeffer, 1 884) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173Rondeletiola minor(Naef, 1912) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174

Sepietta neglectaNaef, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176

Sepietta obscuraNaef, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177

Sepietta oweniana (Orbigny, 18391841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178


13/294


Sepiola aurantiacaJatta, 1 896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

Sepiola knudseniAdam, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

Sepiola pfefferi Grimpe, 1921 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

Sepiola rossiaeformisPfeffer, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

Sepiola steenstrupianaLevy, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180

Euprymna albatrossaeVoss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

EuprymnahoyleiAdam, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Euprymna hyllebergiNateewathana, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Euprymnapenares (Gray, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

EuprymnaphenaxVoss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Euprymna scolopesBerry, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181

Euprymna stenodactyla (Grant, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182

Sepietta petersi (Steenstrup, 1 887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182

Inioteuthis capensis Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182

Inioteuthis japonica (Orbigny, 1 845) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182

Inioteuthis maculosa Goodrich, 1 896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182

Subfamily R OSSIINAE Appellf, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

Rossia macrosoma (Delle C hiaie, 1 830) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183

Rossia pacifica pacifica Berry, 1 911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185

Rossia tortugaensis Voss, 1956 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187

Semirossia equalis (Voss, 1950) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188

Semirossia tenera (Verrill, 1880) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189

Neorossia caroli (Joubin, 1 902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190

Austrorossia antillensis (Voss, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192

Austrorossia australis Berry, 1 918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193

Austrorossia bipapillata (Sasaki, 1920) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194


Rossia brachyura Verrill, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

Rossia bullisi Voss, 1956 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

Rossia glaucopis Loven, 1845 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

Rossia megaptera Verrill, 1 881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

Rossia moelleri Steenstrup, 1 856 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

Rossia mollicella Sasaki, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196

Rossia pacifica diegensis Berry, 1 912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Rossia palpebrosa Owen, 1 834 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Austrorossia enigmatica (Robson, 1 924) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Austrorossia mastigophora (Chun, 1915) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Semirossia patagonica (Smith, 1881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

Neorossia leptodons Reid, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197

SubfamilyHETEROTEUTHINAE Appellf, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198

Heteroteuthis (Heteroteuthis) dispar(Rppell, 1 844) . . . . . . . . . . . . . . . . . . . . . . . . . . . 198

Stoloteuthis leucoptera (Verrill, 1878) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199

Sepiolina nipponensis (Berry, 1911) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201


Heteroteuthis (Heteroteuthis) weberi Joubin, 1 902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

Heteroteuthis (Stephanoteuthis) dagamensis Robson, 1 924 . . . . . . . . . . . . . . . . . . . . . . . 203

Heteroteuthis (Stephanoteuthis) hawaiiensis (Berry, 1 909) . . . . . . . . . . . . . . . . . . . . . . . . 203

Heteroteuthis (Stephanoteuthis) serventyiAllan, 1945 . . . . . . . . . . . . . . . . . . . . . . . . . . 203

Nectoteuthis pourtalesi Verrill, 1 883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

Cephalopods of the World xi


14/294

Iridoteuthis iris (Berry, 1909) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

Iridoteuthis maoria Dell, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203

FamilySEPIADARIIDAE Fischer, 1882 in 18801887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204

Key t o g enera i n t he family S epiadariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204

SepiadariumaustrinumBerry, 1921 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205

Sepiadarium kochii Steenstrup, 1 881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLYFEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

SepiadariumauritumRobson, 1 914 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

Sepiadariumgracilis Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

SepiadariumnipponianumBerry, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

Sepioloidea lineolata (Quoy a nd G aimard, 1832) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

Sepioloidea pacifica (Kirk, 1882) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207

Family I DIOSEPIIDAE A ppellf, 1 898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208


Idiosepius biserialis Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209Idiosepius macrocheirVoss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209

Idiosepius minimus (Orbigny, 1835) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209

Idiosepius notoides Berry, 1921 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209

Idiosepius paradoxus (Ortmann, 1 888) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209

Idiosepius picteti (Joubin, 1894) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210

Idiosepius pygmaeus Steenstrup, 1 881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210

Idiosepius thailandicus Chotiyaputta, Okutani and Chaitiamvong, 1991 . . . . . . . . . . . . . . . . . . 210

Family S PIRULIDAE O wen, 1 836 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211

Spirula spirula (Linnaeus, 1 758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211

4. LIST OF NOMINAL SPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213

5. LIST OF SPECIES BY MAJOR FISHING AREAS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218

6. REFERENCES. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225

7. INDEX OF SCIENTIFIC AND VERNACULAR NAMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249

8. LIST OF COLOUR PLATES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261

xii FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1


15/294

1. INTRODUCTION

by Patrizia Jereb, Clyde F.E. Roper and Michael Vecchione

W

ith the increasing exploitation of finfish resources,and the depletion of a number of major fish stocks

that formerly supported industr ial-scale f isheries,increasing attention continues to be paid to the so-calledunconventional marine resources, which includenumerous species of cephalopods. Cephalopod catcheshave increased steadily in the last 30 years, from about1 million metric tonnes in 1970 to more than 3 million tonnesin 2001. This increase confirms a potential development ofthe fishery predicted by G.L. Voss in 1973, in his firstgeneral review of the worlds cephalopod resourcesprepared for FAO. The rapid expansion of cephalopodfisheries in the decade or so following the publication ofVosss review, meant that a more comprehensive andupdated compilation was required, particularly for cephalopod fishery biologists, zoologists and students. TheFAO Species Catalogue, Cephalopods of the World byC.F.E. Roper, M.J. Sweeney and C.E. Nauen was publishedin 1984 to meet this need.

The number of cephalopod species that enter commercialfisheries has continued to grow significantly since 1984, asa result of a still-growing market demand and the expansionof fisheries operations to new fishing areas and to deeperwaters. It has been suggested that the cephalopod l i fe - s tr a te g y m a y g u ar a n te e s u r vi v a l a g ai n s tenvironmentally stressful conditions, including thosecaused by heavy fishing. However, as cephalopod fisheriesexperienced further extensive development, parallelconcern developed regarding potential overexploitation.

Thus, a broad consensus developed among fisherybiologists to apply the experience gained from errors madein finfish management to avoid possible failures incephalopod exploitation. To help prevent potential failures,refined species identification capabilities are required, aswell as a more detailed and accurate compilation ofinformation on cephalopod species, distribution, biology,f isheries and catch statist ics. Consequently, FAOrecognized that a new edition of the Cephalopods of theWorld catalogue was needed. To achieve this expandedgoal, several authors with particular areas of specializationwere assembled to enhance the accuracy, coverage andutility of this revised catalogue.

In our attempt to make this document as comprehensiveand as useful as possible, the taxonomic coverage of thisedition of the catalogue is organized into 3 levels of interest:

Level 1 : species of cephalopods currently exploitedcommercially and species utilized at the subsistence andartisanal levels;

Level 2 : species of occasional and fortuitous interest tofisheries; this includes species considered to have apotential value to fisheries, based on criteria such asedibility, suspected abundance, accessibility, marketability,etc.;

Level 3 :species with no current interest to fisheries, which

are listed only with the basic information available.

The inclusion of such a wide range of species is necessaryto provide the most comprehensive inventory of species

possibly useful to mankind, regardless of their currentcommercial status. For example, this work should be usefulfor the ever-expanding search for development andutilization of natural products, pharmaceuticals, etc.

The catalogue is based primarily on information available inpublished literature. However, yet-to-be-published reportsand working documents also have been used whenappropriate, especially from geographical areas where alarge body of published information and data are lacking,and we are particularly grateful to colleagues worldwidewho have supplied us with fisheries information, as well asbibliographies of local cephalopod literature.

The fishery statistics presented herein are taken from theFAO official database, FISHSTAT, now available on the

Worldwide web (FISHSTAT Plus 2000). This information issupplemented by field observations made by the authors inmany parts of the world, both in preparation of the 1984volume, as well as for the current edition. These field visitsprovided opportunities to examine fresh material at landingsites, markets and laboratories, as well as to obtainfirst-hand information about local cephalopod fisheries fromregional fisheries workers.

During the 20-plus years separating the two editions, therapid development of cephalopod fisheries worldwide andthe simultaneous increase in the population of fisheriesscientists, their research and publications, made availablean enormous amount of new data and research results.

Sometimes it is difficult to evaluate the reliability ofpublished data, especially with regard to the identification ofspecies in areas where the cephalopod fauna has not beensufficiently studied. It is entirely understandable that fieldworkers isolated from good library and museum/collectionfacilities have difficulties in correctly identifying the speciesthey encounter in the field. Moreover, the discovery of newspecies, the more accurate delimitation of known species,or even the introduction of nomenclatural changes, maycause confusion and lead to the use of scientific names thatare incorrect by modern standards. Although great carewas exercised to evaluate and correct such publishedinformation used in the catalogue, some incorrectinterpretations may have occurred. Another difficulty, in the

taxonomic literature especially, is that information on theeconomic importance of species is rather scarce or of avery general nature. Also, we may have overlookedimportant information published only in relevant localfisheries literature that is unavailable on a broader scale. Allof these potential difficulties, however, have beensignificantly mitigated during the preparation of the newedition because of the availability on-line of fisheriesdatabases and bibliographic search capabilities.

With regard to the limitations mentioned above, we heartilyrequest that readers who detect any errors in theinformation presented, or who have any additionalinformation and data that will enhance the accuracy andutility of this book, please contact and inform one of theauthors or the Species Identification and Data Programme(SIDP) of the Marine Resources Service, FisheriesResources Division, Fisheries Department, FAO Rome.

Cephalopods of the World 1


16/294


17/294

1.2 General Remarks on Cephalopods

The group known as cephalopods (class Cephalopoda) isthe most complex in the phylum Mollusca, and indeed, in allof the invertebrate phyla. It includes exclusively marineanimals that live in all oceans of theworld with theexceptionof the Black Sea, from the Arctic Sea to the Antarctic Ocean

and from the surface waters down into the deep sea.

Cephalopods first appeared as a separate molluscantaxonomic entity, the nautiloids, in the Upper Cambrianperiod (over 500 million years ago), but more than half ofthese ancestors were already extinct by the end of theSilurian, 400 million years ago, when only the nautilusessurvived. Meanwhile, other forms arose in the latePalaeozoic (between 400 and 350 million years ago),including those of the Subclass Coleoidea, but most ofthem became extinct by the end of the Mesozoic, about150 million years ago. The only members of the subclassColeoidea that exist today are the forms that developed inthe Upper Triassic and Lower Jurassic (between 200 and

150 million years ago).Although there is a long fossil record of many differentgroups, all living cephalopods belong to two subclasses:the Coleoidea, which includes the major groups known assquids, cuttlefishes sensu lato, octopods and vampires,and the Nautiloidea, containing two genera, Nautilusand

Allona utilus, the only surviving cephalopods with anexternal shell.

At the present time the status and understanding of theS y s te m a t ic s a n d C l a s si f i c at i o n o f t h e R e c e n tCephalopoda is under considerable discussion. Thefamilies of living cephalopods are, for the most part, wellresolved and relatively well accepted. Species-level taxa

usually can be placed in well-defined families. The higherc l a s s i f i c a t i o n , h o we v e r , s t i l l i s n o t r e s o l v e d . T h eclassification above the family level is controversial and abroad consensus still needs to be achieved. This situationis not unexpected for a group of organisms that hasundergone explosive research attention in recent decades.

Consequently, rather than accept and promote anyparticular scheme of classification, before consensus andstabil ity are achieved, we will use an operationalbreakdown that is satisfactory for the objectives of thisCatalogue. For practical purposes we separate thecephalopods into several groups, without assigning orimplying taxonomic relationships. Figure 1 diagrams

several of the classification schemes currently underdiscussion.

In this work the following groups are used, as illustrated inFigure 2

1/:

Nautiluses

Cuttlefishes

Bobtail squids

Bottletail squids

Pygmy squids

Rams horn squid

Myopsid squids

Oegopsid squids

Vampires

Cirrate octopodsIncirrate octopods

Unresolved taxa:

Spirula

Idiosepius

Bathyteuthis

Chtenopteryx

Sepiadariidae

Plural versus singular usage of cephalopod common groupnames is standardized as follows:

squid, cuttlefish, octopod, octopus, vampire, nautilusrefer to one individual or one species;

squids, cuttlefishes, octopods, octopuses, vampires,nautilusesrefer to two or more individuals and/or species.These terms are also used to indicate the major groups.

The termcuttlefishesis also used sensu lato to indicatethe following groups: Cuttlefishes, Bobtail squids, Bottletailsquids, Pygmy squids and the Rams horn squid.

We differentiate between the members of the family

Octopodidae, which are calledoctopus/octopuses and themembers of the whole group (Incirrate and Cirrate or anycombination of non-Octopodidae taxa), which are calledoctopod/octopods.

Cephalopods occur in all marine habitats of the world:benthic forms are found on coral reefs, grass flats, sand,mud and rocks; epibenthic, pelagic and epipelagic speciesoccur in bays, seas; and epipelagic, mesopelagic,bathypelagic and benthopelagic species are all present inthe open ocean. Salinity is considered to be a limiting factorin cephalopod distribution; they are generally restricted tosalinity concentrations between 27 and 37. However,

Lolliguncula brevis, which lives and reproduces in watersof 17, demonstrates a capacity for a higher degree of

salinity tolerance (Hendrix et al ., 1981). Some speciesinhabit the Red Sea and the southern coasts of the IberianPeninsula (Guerra, 1992), where the salinity is higher than37 and other species have been found in waters wheresalinity ranges between 25 and 18 (Sea of Marmara;Unsal et al., 1999). The habitat depth range extends fromthe intertidal to over 5 000 m. Many species of oceaniccephalopods undergo diel vertical migrations: they occur atdepths of about 200 to 700 m during the day, then at theonset of twilight and increasing darkness, they ascend intothe uppermost 200 m for the night. A deeper-living layer ofdiel migrators occurs from about 1 000 m to 600 m duringthe daytime. The abundance of cephalopods varies,depending on group, habitat and season, from isolated,

territorial individuals (primarily benthic octopods andsepioids), through small schools of squids with a few dozenindividuals, to huge schools of neritic and oceanic specieswith millions of specimens.

The size of adultcephalopods ranges from less than 1 cm(Jackson, 1989) to the giant squid at approximately 20 m intotal length, including the tentacles. The largest specimensmay weigh well over 500 kg, but the average size ofcommercial species is 20 to 40 cm mantle length and about0.1 to 2.0 kg total weight.

Cephalopods are soft-bodied, bilaterally symmetricalanimals with a well-developed head and a body thatconsists of the muscular mantle, the mantle cavity that

houses the internal organs, and, when present, the externalfins. The head bears an anterior circumoral (surrounding


1/ The endings used in the group names do not imply any level ofclassification.


18/294

4 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1

Roperet al.(1984) Order Suborder

Teuthoidea Myopsida

Oegopsida

Sepioidea

Vampyromorpha

Octopoda

Cirrata

Incirrata

Engeser and Bandel (1988) Superorder Order Suborder

Decapoda

Spirulida

higher decapods(name not given)

Teuthina

Sepiina

Vampyromorphoidea

Vampyromorpha

Octopoda Cirrata

Incirrata

Clarke (1988) Order Suborder

Sepioidea

Sepiolioidea

Teuthoidea Myopsida

Oegopsida

Vampyromorpha

Octopoda

Sweeney and Roper (1998) Superorder Order Suborder

Decabrachia

Spirulida

Sepiida

Sepiolida

Teuthida Myopsina

Oegopsina

Octobrachia

Vampyromorphida

Octopodida Cirrina

Incirrina

Younget al.(1998) Division Superorder Order Suborder

Neocoleoidea

Decapodiformes

Oegopsida

Myopsida

Sepioidea

Sepiida

Sepiolida

Spirulida

Incertae sedis

Octopodiformes

Vampyromorpha

Octopoda Cirrata

Incirrata

Boletzky (1999) Grade Superorder Order

Decabrachia

Spirulida

Sepiida

Sepiolida

Idiosepiida

Teuthida

Vampyropoda

Pseudooctobrachia Vampyromorpha

Octobrachia Cirroctopoda

Octopoda

Haas (2002)ranks not given

1 2 3 4 5

Neocoleoidea

Decabrachiomorpha

Oegopsida

UniductiaSpirulida

Myopsida Loliginida

Sepiida

Octobrachiomorpha

Vampyromorpha

Octopoda Cirrata

Incirrata

Fig. 1 Some conflicting suprafamilial classifications of living coleoid cephalopods


19/294


Fig. 2 Living cephalopods


20/294

the mouth) crown of mobile appendages (arms, tentacles).This characteristic feature reflects the origin of the nameCephalopoda, which derives from the union of the twoGreek words: kefale, head, and pous, feet. The namewas erected by Schneider in 1784, and i t becamepermanently in use within the scientific context with thepublication of Cuviers work (1798). Arms and tentacles

bear suckers and/or hooks (except inNautilus), which arepowerful tools to seize prey. The mouth has a pair ofchitinous jaws (the beaks) and, as in other molluscs, achitinous tongue-like radula (band of teeth) occurs in mostcephalopod species. The ancestral mollusc shell isvariously modified, reduced, or absent in living coleoids. Itis a calcium carbonate structure in cuttlefishes (thecuttlebone of sepiids and the rams horn shell ofSpirula),reduced to a rigid structure composed of chitin in squids(the gladius or pen, sometimes quite flexible) and to acartilaginous structure in finned octopods. In somesepiolids no vestige of shell is found. A true external shelloccurs only in the primitive nautiluses (restricted to theIndo-Pacific), although a shell-like egg case is produced

and carried by female argonauts (pelagic octopods oftenmisnamed paper nautilus).

The loss of the external shell allowed the development of apowerful muscular mantle that became the mainlocomotory organ for fast swimming, via water jetting fromthe funnel. The funnel (siphon) is a unique, multifunctional,muscular structure that aids in respiration and expulsion ofmaterials in addition to locomotion. Oxygenated water isdrawn through the mantle opening around the head (neck)into the mantle cavity, where it bathes the gills forrespiration. Mantle muscular contraction expels thedeoxygenated water from the mantle cavity through theventrally located funnel. The discharge jet serves to

eliminate nephridial and digestive wastes, as well as tocomplete the respiratory cycle and for locomotion. Femalereproductive products (eggs, egg masses) also aredischarged through the funnel. Most coleoids produce ink,a dark, viscous fluid also expelled through the funnel. Theink may take the form of a mucoidal pseudomorph (falsebody) to decoy potential predators, or of a cloud to obscurethe escaping cephalopod.

One pair of gills (ctenidia) is present, except inNautilus andAllonautilus, which have two pairs for respiration, i.e. toextract the oxygen from the water. However, in contrastwith coleoids,Nautilusmakes use of anaerobic respirationduring periods of high activity and can survive in water withvery low oxygen content. Coleoids also use anerobicmuscle layers. Cutaneous respiration also occurs in somecephalopods.

The cephalopodcirculatory system is distinctive within theMollusca. It is a closed system (blood contained withinvessels), similar in many respects to that of vertebrates andfulfilling the demand for the more efficient circulationrequired by an active locomotory system. There is aprincipal, or systemic, heart, two branchial hearts anddeveloped arterial, venous and capillary systems supplyingblood to the muscles and organs. From the gills, theoxygenated blood goes through the efferent branchialvessels to the systemic heart, where it is expelled from theventricle through three aortas: the cephalic or dorsal aorta,

which supplies the head and the anterior part of the gut; theposterior, minor or abdominal aorta that supplies the mantleand fins along with the posterior part of the gut and thefunnel; and the gonadal aorta that develops gradually with

sexual maturation of the animal. The blood is collectedthrough sinuses and capillaries into the veins, throughwhich it goes to the branchial hearts that pump it throughthe filaments of the gills. The circulating respiratory pigmentu s e d f o r o x y g e n t r a n s p o r t i s c o p p e r - c o n t a i n i n ghaemocyanin, a system of rather lower efficiency than theiron-containing haemoglobin of vertebrates. In living

cephalopods blood sinuses are much reduced andreplaced functionally by muscles. The circulatory systemt he re f or e h as t o w or k a ga in s t t h e p er i ph er almuscle-induced pressure, which increases with increasingactivity (maximum during jet-swimming). It also has to copewith the resistance of the small diameter of the finalcapillary blood vessels, and the low oxygen carryingcapacity of the blood (less than 4.5% by volume). In spite ofthese l imitat ions, the system has other functionalmodifications (see for example Wells and Smith, 1987;Martin and Voight, 1987) that achieve the capacity to deliveroxygen at a rate comparable to that of active fishes,enabling cephalopods to accomplish extraordinaryperformances.

The excretory system also differs markedly from that ofother molluscs and, along with the closed circulatorysystem and the branchial circulation, enables uniquerelationships between blood and the final secretion, theurine. The excretory system differs between livingnautiloids and coleoids and also among coleoids, but thegeneral organization is similar, consisting basically of therenal sac with the renal appendages (organs comparable tovertebrate kidneys), the pericardial glands, the branchialhearts and the gills. Cephalopods are ammoniotelic, andammonium ions are continuously released by the gillepithelium and by renal appendages into the surroundingwater. Ammonium ions are used by buoyant squids to

replace denser chloride ions in fluids in the coelom and inthe body tissues. Because this solution is less dense (andhence more buoyant) than seawater, it provides lift.

Th e nervous system is highly developed in recentcephalopods, with a large brain and peripheral connections,contrasting with the original molluscan circumesophagealnerve ring. Among its most remarkable features is the giantfibre system of squids and cuttlefishes that connects thecentral nervous system with the mantle muscles. Thissystem consists of three orders of cells and fibres andensures the immediate and simultaneous contraction ofmantle, fins and retractor muscles of both sides, rather thanan anterior to posterior sequential contraction that would becounter-productive for water movement (expulsion). Alsoremarkable is the eye development of most coleoids, forwhich vision plays a major role in life. Their eyes are large,have a design generally similar to that of fishes and othervertebrates (e.g. a lens focuses images on the retina), andall the available evidence suggests the ocular/visualperformance to be comparable to that of vertebrates. Thecephalopods also developed a system to keep the focusedimage stationary on the retina while the animal turns, bymoving the eyes in coordination with the head/bodymovement. This is extremely important for hunters that relyon sight, and it is accomplished by connections of the eyemuscles with the statocysts, a mechanism similar to thevestibulo-optic system of fishes. The statocyst systemprovides cephalopods with information on their orientation,

as well as changes in position and direction of movement.The statocyst system is highly developed in coleoids, whereit consists of separate cavities located in the cartilaginousskull, posteroventral to the brain. The statocysts contain



21/294

nervous cells and receptors differentiated to detect bothlinear acceleration, with the aid of calcareous stones calledstatoliths, and angular acceleration. Many coleoids alsohave extra-ocular photoreceptors (photosensitive vesicles)about which little is known; in mesopelagic squids theyappear to monitor light intensity in order to enable theanimals to match the counter-illumination with the ambient

light by their own photophores (light-producing organs).Cephalopods are provided with numerous mechano- andchemoreceptors and recent evidence indicates that insome species, like Loligo vulgaris, Sepia officinalis andOctopussp., ciliate cells form lines in several parts of thebody, a system analogous to the lateral-line system infishes.

Most Coleoidea are able to change colour by using a complexsystem of chromatophores under nervous control. Thechromatophores are pigment-filled sacs present in the skin,and capable of remarkable expansion and contraction. Thissystem responds to current situations in the environment, e.g.background coloration and threatening predators, and it is

critical for survival, especially for shallow-water benthic forms.Coloration capabilities are variable depending on taxonomicgroup and habitat. Most species also have iridocytes(shiny,reflective platelets) in the skin. Most cephalopod behaviourincludes rapid changes in overall colour and colour patterns,as well as changes in the texture of the skin, from smooth toheavily papillate, tubercular, or with erected flaps. Whileshallow-living cephalopods are able to conceal themselves bychromatophore-produced colour patterns, chameleon-likecolour changes and textural presentations, many deep-seaforms camouflage themselves by producing bioluminescentlight from photophores which eliminate their silhouettesagainst the downwelling sunlight in the dimly-lit mid-depths.

Locomotionis achieved by any of, or a combination of, thefollowing methods, depending on the taxonomic group:1) jet propulsion; 2) flapping or undulating the fins on themantle; 3) crawling along the bottom on the arms;4) medusoid swimming with arms and interbrachial webs.The fins on the mantle also provide balance and steeringduring jet propulsion. Many families of midwater squidshave evolved to low energy life styles and achieve neutralbuoyancy by producing and storing in tissues or in differentorgans substances/elements with specific properties, suchas oils or solutions of ammonium ions. This capabilityenabled coleoid cephalopods to inhabit open water, even inthe great depths in the ocean, the greatest volume of livingspace on earth.

Cephalopods are voracious, active predators that feedupon crustaceans, fishes, other cephalopods and, in thecase of some benthic octopods, on bivalved molluscs. Thespeed of cephalopods, their high mobility and powerfulvisual systems, along with strongly-muscled arms andtentacles, both equipped with suckers and/or hooks, makethem extremely efficient hunters. A common huntingtechnique in sepioids and loliginids involves extremelyrapid shooting forward of the tentacles to capture the prey,while in some oegopsid squids the tentacles may be usedlike long, sucker-covered fishing lures. Some octopods usetheir web to envelop crabs and occasionally may wait untilprey touches them before attacking. The captured prey ofcephalopods is brought to the mouth and killed by bites of

the strong, chitinous beaks, equipped with powerfulmuscles. Sometimes the prey are first paralysed with afast-acting toxin, immobilization being a strong advantagein case of large and/or very active prey such as large crabs.

The Incirrate octopods and Sepia species usually poisontheir prey before eating them, while squids do not seemable to produce such strongly toxic secretions. Digestion incephalopods is rapid and efficient and cephalopodmetabolism is essentially proteinic: there is little or nodigestion/assimilation of carbohydrates and lipids. Foodconversion is highly efficient, especially in octopuses,

where up to 50% of the food eaten can be converted intobody mass. More active cephalopods like squids, however,need several times the amount of food required byoctopuses and can eat from 3 to 15% of their body weighteach day.

All cephalopods are dioecious (separate sexes) and many,though not all, exhibit external sexual dimorphism, either inmorphological or in size differences. Females frequentlyare larger than males, a phenomenon which reaches itsextreme in some pelagic octopods (such as Argonauta)where males are truly dwarf forms. Males of many speciespossess one, occasionally two, modified arm(s) (thehectocotylus) for transferring spermatophores to females

during mating. The males of some species also exhibitmodifications to other arms, in addition to the hectocotylus.The hectocotylus may be simple or complex and canconsist of modified suckers, papillae, membranes, ridgesand grooves, flaps. The one or two limbs function to transferthe spermatophores (tubular sperm packets) from themales reproductive tract to an implantation site on thefemale. The spermatophores may be implanted inside themantle cavity(where they maypenetrate theovary), into theoviducts themselves, around the mantle opening on theneck, on the head, in a pocket under the eye, around themouth or in other locations. Females of a few species alsod e v e l o p g e n d e r - s p e c i f i c s t r u c t u r e s ( e . g . a r m - t i pphotophores) when mature. Mating is often preceded or

accompanied by courtship behaviour that involves strikingchromatophore patterns and display. Copulatory behaviourvaries significantly among species, in colour and texturaldisplay, proximity of male and female, duration of displaya n d s p e r m a t o p h o r e t r a n s f e r , a n d t h e l o c a t i o n o f implantation of the spermatophores on the female.

The gonadsform a single mass at the posterior end of themantle cavity, and female gonoducts may be paired (inoegopsids and incirrate octopods) or single, as in othercoleoids and in the nautiluses. Cephalopod reproductivesystems are highly complex structures with ducts, glandsand storage organs. Female octopods have oviducalglands, while decapods, in addition, have nidamentalglands and, in some families, accessory nidamental glands.S p e r m a t o p h o r e s a r e p r o d u c e d i n t h e m u l t i - u n i tspermatophoric gland and stored in the Needhams sac,from which they are released through the terminal part ofthe duct, the penis. This term is not strictlyaccurate in manygroups, because the spermatophores are passed to, ortaken by, the hectocotylized arm(s), which in turn transfer(s)the spermatophore(s) to the female. Some families do notpossess a hectocotylus. Instead, the terminal portion of themale reproductive tract forms a functional penis that can begreatly enlarged and elongated, often extending out of themantle cavity and past the head. It is likely that thesestructures directly transfer the spermatophores to thefemale. The number and size of spermatophores varygreatly, depending on the species and group (for reviews on

spermatophore structures and function see Mann et al.,1966, 1970; Mann, 1984). Once in contact with seawater,the so called spermatophoric reaction begins. Thespermatophores evert, with the resultant extrusion of the



22/294

sperm packet caused by the penetration of water inside thespermatophoric cavity, where the osmotic pressure ishigher. The resulting extruded sperm packet is namedspermatangium (or sperm bulb or body). Sperm are able tosurvive several months once stored in the female, at least insome species, and fertilization of mature ova may takeplace either in the ovary, the oviducal glands, the mantle

cavity or the cone formed by the outstretched arms whilethe eggs are laid. Males of the pelagic octopodArgonauta(and i ts relat ives) insert sperm into a detachablehectocotylized arm. Mating occurs by the arm severing andcrawling into the females mantle cavity.

Fertilized eggs are embedded in one or more layers ofprotective coatings produced by the oviducal andnidamental glands and generally are laid as egg masses.Egg masses may be benthic or pelagic, varying among themajor taxonomic groups. Some oceanic mesopelagic anddeep-sea species, however, lay individual eggs.

Eggs of neritic, inshore squids, except in Sepioteuthis,

generally are very small (only a few millimetres in diameter)and frequently are laid in finger-like pods each containingfrom a few to several hundred eggs. Deposited inmultifinger masses (sometimes called sea mops), theseeggs are attached to rocks, shells or other hard substrateson the bottom in shallow waters. Many oceanic squids laytheir eggs into large sausage-shaped or sphericalgelatinous masses containing tens or even hundreds ofthousands of eggs that drift submerged in the open sea.Other pelagic species lay individual eggs, not enmeshed ingelatinous masses.

Cuttlefishes sensu lato lay few, relatively large (around 10to 40 mm in diameter), grape-like eggs that are attached tohard substrates. Some species camouflage the eggs frompredators by making them dark using a coating of inkdeposited by the female at egg laying.

Cirrate octopods lay rather large (from 10 to 25 mm length)single eggs, enclosed in a tough protective coating (seeBoletzky, 1982, 1986), and they are laid individually or insmall clusters of a very few eggs.

Benthic incirrate octopods lay their eggs singly or ingrape-like clusters and strands. Most species attach theeggs to hard surfaces such as rock or shells while somecarry the eggs within their arms and webs. The eggs vary insize from a few millimetres to 40 mm long. They areattached to each other and/or to the substratum by cement

produced by the oviducal gland. They lack the gelatinousouter matrix found in squid and cuttlefish eggs and the outershell found in the Cirrate octopod eggs: the protectivefunction has been replaced by the brooding behaviour ofthe female parent. The female of the pelagic octopod

Argonauta constructs a thin, decorative shell-like eggcase, which encases her mantle and into which sheperiodically lays and attaches festoons of minute eggs.

The mode of reproduction and egg-laying still is unknownfor many species, especially those of oceanic and deep-seaenvironments.

Development of cephalopod embryos is direct, without true

metamorphic stages. Hatchlings of species with large eggslook like miniatures of the adult, while hatchlings of specieswith small eggs undergo gradual changes in proportionsduring development. The young of some species, however,differ conspicuously from the adults. Thus, the term

paralarva has been introduced for these early stages ofcephalopods that differ morphologically and ecologicallyfrom older stages. Hatchlings from large-egged benthicoctopods are either benthic crawl-away young or temporarily planktonic, quickly settling back to the adultbenthic habitat. Small-egged benthic octopuses produceplanktonic hatchlings with very simple skin patterns of large

chromatophores. The paralarvae of many deep-seaspecies of squids and octopods occur in the upper 100 m ofthe open ocean; then they exhibit an ontogenetic descent,gradually descending to deeper depths with increasing sizeuntil the adult depth is attained. Time of embryonicdevelopment varies widely, from a few days to manymonths, depending on the species and the temperatureconditions. Hatching may occur synchronously from asingle clutch orbe extended over a period of 2 or 3 weeks.

In spite of the large number of studies and research carriedout on cephalopods, especially in recent decades, the lifehistoryof the majority of species is still unknown, and ourknowledge of the life cycles of the members of this

interesting class remains fragmentary. Information comesfrom studies in the field as well as from observations in thelaboratory. However, little is known of life history for speciesthat are not targets of regular fisheries, and only a handfulof cephalopod species have been reared successfullyin thelaboratory.

Studies and monitoring of growth in cephalopods arecomplicated by the high variability in individual growthrates. This makes it difficult to apply conventional methods,e.g. length frequency analysis, used for more traditionalresources such as fishes and crustaceans. Determinationof age in Recent coleoids is also difficult, because theyhave few hard structures that show daily marks (rings) that

would enable direct estimates of age. In the last 15 years,progress has been made on the study and analysis ofstatoliths that has resulted in an increased knowledge ofage, in squids at least. This has led to changes in ourperspectives about the physiology and ecology of manyteuthoids, but more research is required before a fullunderstanding is achieved (see Jereb et al., 1991; Okutaniet al., 1993; Jackson, 1994 and Lipinski and Durholtz, 1994for reviews and discussions). Principle results obtainedfrom the research generally confirm a very high growth ratein cephalopods, comparable to that of the fastest-growingfishes.

Nautilusspecies have alife span of about 20 years, duringwhich they spawn intermittently when first mature, then

annually when fully mature, laying a few large eggs at atime. The life expectancy of most coleoids appears to rangefrom a few months to one or two years. Many small oceanicsquids, such as pyroteuthids may complete their life cyclesin less than six months, while some minute forms ofsepioids have a life span of only 2 to 3 months. Recentevidence, however, suggests that larger species of squidsand octopods, for example the giant squid (Architeuthisspp.) and the giant octopus (Enteroctopus dofleini), as wellas those that live in coldest habitats, may live for severalyears.

A general consensus exists that spawning is a terminalevent, in spite of the high variability in the duration of

individual spawning periods (5 to 50% of ontogenesis;Nigmatullin, 2002) as well as the type of spawning, e.g.from one-time, total spawning, to prolonged, intermittent,multiple batches. A recent review of coleoid reproductive



23/294

strategies (Rochaet al., 2001) defined five comprehensive,flexible strategies:

A) Single event spawning, formerly termed semelparity,consists of synchronous ovulation, and monocyclic,simultaneous terminal spawning; Octopus vulgaris is thetypical example of this life strategy.

B) Multiple event spawning, formerly termed iteroparity,with the following possibilities:

1 - polycyclic spawning: egg-laying occurs in separatebatches during the spawning season and growthoccurs between production of egg batches andsubsequent spawning events (e.g.Nautilus species);

2 - mult iple spawning, with group-synchronousovulation, monocyclic spawning and growth betweenegg batches (e.g.Sthenoteuthis oualaniensis);

3 - i n t e r mi t t e n t t e r m i na l s p a wn i n g wi t h g r o u p-synchronous ovulation, monocyclic spawning and nogrowth between egg batches (e.g. Sepia officinalis,

Loligo vulgaris,Illex coindetii);

cephalopods of the world vol 1 nautilus, sepioids

Documents