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)

FAOSpeciesCatalogue for FisheryPurposesNo. 4, Vol. 1 FIR/Cat. 4/1
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)
edited by
P. Jereb
Istituto Centrale per la Ricerca Scientifica e Tecnologica Applicata al Mare
Rome, Italy
and
C.F.E. Roper
Smithsonian Institution, National Museum of Natural History
Washington, DC, United States of America
withthe support of the
Government of Italy
(Ministero per le Politiche Agricole e Forestali,
Direzione Generale per la Pesca e l?Acquacoltura)
FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
Rome, 2005
The designations employed and the presentation of material
in this information product do not imply the expression of any
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Organization of the United Nations concerning the legal
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authorities, or concerning the delimitation of its frontiers or
boundaries.
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? FAO 2005
ISBN 92-5-105383-9
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.

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 through
contributions 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 the
World (Roper et al., 1984), and it constitutes Volume I of Number 4 in the new series: FAO Species Catalogue for Fisheries
Purposes, that evolved as an independent series in 2001 from the former FAO Fisheries Synopsis No. 125.
Because the new Catalogue has expanded apace with recent research and fisheries information and revisions, it now is
necessary 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 volume
independently 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 D?Antoni (FAO, Rome).
Page composition and indexing: Mich?le S. Kautenberger-Longo (FAO, Rome).
Digitization of distribution maps: Fabio Carocci (FAO, Rome) and Elena V. Orlova (St Petersburg, Russia).
Cover illustration: Emanuela D?Antoni (FAO, Rome).
Distribution
Authors
FAO Fisheries Officers
Regional Fisheries Councils and Commissions
Selector 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 of
Cephalopods of the World (1984). The present Volume is a multiauthored compilation that reviews six families:
Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae and Spirulidae, with 23 genera and the 201 species
known 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 and
original citation of the species (or subspecies); main synonyms; English, French and Spanish FAO names for the
species; illustrations of dorsal and ventral aspect of the whole animal (as necessary) and other distinguishing
illustrations; 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. The
volume is fully indexed and also includes sections on terminology and measurements, an extensive glossary, an
introduction with an updated review of the existing biological knowledge on cephalopods (including fisheries
information and catch data for recent years) and a dedicated bibliography.
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. 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. 51?55.
Jereb, P., Roper, C.F.E. & Vecchione, M. 2005. Introduction. 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. 1?19.
Reid, A. 2005. Family Sepiadariidae. 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. 204?207.
Reid, A. 2005. Family Idiosepiidae. 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. 208?210.
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. 211?212.
Reid, A. & Jereb, P. 2005. Family Sepiolidae. 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. 153?203.
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. 57?152.
Acknowledgements
The authors are pleased to heartily acknowledge the contributions of colleagues who have supplied information or read
drafts of this Second Edition of the Cephalopod Catalogue. Without their good efforts this work would have been a less
comprehensive, 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), Giambattista
Bello (Acquario Provinciale, Bari, Italy), Alexei Birkun (Crimean State Medical University, Simferopol, Ukraine), Teresa
Borges (Centre of Marine Sciences, University of Algarve, Lisbon, Portugal), Julia Filippova (Russian Federal Institute of
Marine 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 Biological
Resources-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),
Jo?o Pereira (Instituto Nacional de Investiga??o Agr?ria e das Pescas, Lisbon, Portugal), Sergio Ragonese (Istituto per
l?Ambiente Marino Costiero?CNR, 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 C?diz-IEO, Cadiz, Spain), Tooraj Valinassab (Iranian Fisheries
Research Organization, Teheran, Iran), Peter Ward (University of Washington, Seattle, USA).
Very special thanks are due to: Sigurd von Boletzky (Observatoire Oc?anologique de Banyuls-CNRS, Banyuls-sur-Mer,
France) for his knowledgeable help in solving some systematic problems; Peter Boyle , for his valuable editing of our
Introductory Review; Renata Boucher (Mus?um National d?Histoire Naturelle-CNRS, Paris, France) and Pilar Sanchez
(Instituto de Ciencias del Mar, Barcelona, Spain) for their help with the French and Spanish vernacular names; Neethirajan
Neethiselvan (Fisheries College and Research Institute, Tamil Nadu, India) who so kindly provided information and
photographic material on some Indian species; Mark Norman (Museum Victoria, Melbourne, Australia) for his invaluable
help 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 on
some southern African species; Michael J. Sweeney (New Mexico State University, Las Cruces, New Mexico, USA) for
technical support, nomenclatural listings, sources of information and literature searches.
Pere Oliver (Instituto Espa?ol de Oceanograf?a, 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. We
especially thank Michel Lamboeuf for his help with the use of the FAO FISHSTAT database and for his revision of the
graphic sections thereafter.
Of course, a compilation of this nature must rely heavily on already-published works; we acknowledge these publications
with gratitude. In particular, we acknowledge here that many illustrations from these works have been used for the
purposes 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 the
preparation of this Volume: Emanuela D?Antoni for her excellent job in creating the many illustrations needed for the
Catalogue and for greatly enhancing many illustrations from the literature; Nicoletta De Angelis and Mich?le
Kautenberger for their skilful collaboration in completing this highly technical and complex document and Fabio Carocci
for 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. Therefore
we 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 invaluable
support during many stages of the preparation of this Catalogue.
Cephalopods of the World vii
viii FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
TABLE OF CONTENTS
PREPARATION OF THIS DOCUMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . v
1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1 Plan of the Catalogue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1.2 General Remarks on Cephalopods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
1.3 Interest to Fishery and Role in the Ecosystem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
1.4 Illustrated Glossary of Technical Terms and Measurements . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
1.5 Key to Recent Cephalopod Groups and Families . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
2. CHAMBEREDNAUTILUSES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Family NAUTILIDAE de Blainville, 1825 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Nautilus macromphalus Sowerby, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Nautilus pompilius Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Nautilus belauensis Saunders, 1981 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Nautilus repertus Iredale, 1944 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Nautilus stenomphalus Sowerby, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Allonautilus scrobiculatus (Lightfoot, 1786) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Allonautilus perforatus (Conrad, 1847) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
3. CUTTLEFISHES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Family SEPIIDAE Keferstein, 1866 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Key to genera in the family Sepiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
Metasepia pfefferi (Hoyle, 1885) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Metasepia tullbergi (Appell?f, 1886) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Sepia aculeata Van Hasselt, 1835 (in F?russac and d?Orbigny, 1834?1848) . . . . . . . . . . . . . . . . . 63
Sepia andreana Steenstrup, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Sepia apama Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Sepia arabica Massy, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Sepia australis Quoy and Gaimard, 1832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Sepia bandensis Adam, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Sepia bertheloti d?Orbigny, 1835 (in F?russac and d?Orbigny 1834?1848) . . . . . . . . . . . . . . . . . . 73
Sepia braggi Verco, 1907 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Sepia brevimana Steenstrup, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Sepia cultrata Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Sepia elegans Blainville, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Sepia elliptica Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Sepia elobyana Adam, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Sepia esculenta Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Sepia grahami Reid, 2001. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Sepia hedleyi Berry, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Sepia hierredda Rang, 1835 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Sepia kobiensis Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Sepia latimanus Quoy and Gaimard, 1832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Sepia longipes Sasaki, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Sepia lorigera W?lker, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Sepia lycidas Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Sepia madokai Adam, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Sepia murrayi Adam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Sepia officinalis Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Sepia omani Adam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sepia opipara (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Sepia orbignyana F?russac in d?Orbigny, 1826 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Sepia papuensis Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Sepia pharaonis Ehrenberg, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia plangon Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Sepia prabahari Neethiselvan and Venkataramani, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . 110
Sepia prashadi Winckworth, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Sepia ramani Neethiselvan, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Sepia recurvirostra Steenstrup, 1875 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Sepia rozella (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Sepia savignyi Blainville, 1827 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Sepia smithi Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Sepia stellifera Homenko and Khromov, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Sepia sulcata Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Sepia trygonina (Rochebrune, 1884) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
Sepia vermiculata Quoy and Gaimard, 1832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Sepia vietnamica Khromov, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Sepia vossi Khromov, 1996 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Sepia whitleyana (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
Sepia zanzibarica Pfeffer, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
Sepiella inermis (Van Hasselt, 1835) (in F?russac and d?Orbigny, 1834?1848) . . . . . . . . . . . . . . . 130
Sepiella japonica Sasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
Sepiella ornata (Rang, 1837) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Sepiella weberi Adam, 1939 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia acuminata Smith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia adami Roeleveld, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia angulata Roeleveld, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia appellofi W?lker, 1910 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia aureomaculata Okutani and Horikawa, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Sepia bartletti (Iredale, 1954) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Sepia bathyalis Khromov, Nikitina and Nesis, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Sepia baxteri (Iredale, 1940) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Sepia bidhaia Reid, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Sepia burnupi Hoyle, 1904 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Sepia carinata Sasaki, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Sepia chirotrema Berry, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Sepia confusa Smith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia cottoni Adam, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia dollfusi Adam, 1941 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia dubia Adam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia elongata d?Orbigny, 1839?1842 (in F?russac and d?Orbigny 1834?1848) . . . . . . . . . . . . . . . 141
Sepia erostrata Sasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Sepia faurei Roeleveld, 1972 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Sepia foliopeza Okutani and Tagawa, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Sepia gibba Ehrenberg, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia hieronis (Robson, 1924) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia incerta Smith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia insignis Smith, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia irvingi Meyer, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia ivanovi Khromov, 1982 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia joubini Massy, 1927 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia kiensis Hoyle, 1885 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Cephalopods of the World ix
x FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia koilados Reid, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Sepia limata (Iredale, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Sepia mascarensis Filippova and Khromov, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Sepia mestus Gray, 1849 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Sepia mira (Cotton, 1932) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia mirabilis Khromov, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia novaehollandiae Hoyle, 1909 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia papillata Quoy and Gaimard, 1832 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia pardex Sasaki, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia peterseni Appell?f, 1886 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia plana Lu and Reid, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia plathyconchalis Filippova and Khromov, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia pulchra Roeleveld and Liltved, 1985 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sepia reesi Adam, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sepia rhoda (Iredale, 1954) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sepia robsoni (Massy, 1927) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sepia saya Khromov, Nikitina and Nesis, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia senta Lu and Reid, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia sewelli Adam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia simoniana Thiele, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia sokotriensis Khromov, 1988 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Sepia subplana Lu and Boucher-Rodoni, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Sepia subtenuipes Okutani and Horikawa, 1987 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Sepia tala Khromov, Nikitina and Nesis, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Sepia tanybracheia Reid, 2000 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia tenuipes Sasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia thurstoni Adam and Rees, 1966 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia tokioensis Ortmann, 1888 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia tuberculata Lamarck, 1798 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Sepia typica (Steenstrup, 1875) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Sepia vercoi Adam, 1979 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Sepiella cyanea Robson, 1924 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Sepiella mangkangunga Reid and Lu, 1998 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Sepiella ocellata Pfeffer, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Sepia filibrachia Reid and Lu, 2005 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Family SEPIOLIDAE Leach, 1817 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Key to subfamilies and genera in the family Sepiolidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Subfamily SEPIOLINAE Appell?f, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Sepiola affinis Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Sepiola atlantica Orbigny, 1839?1842 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Sepiola birostrata Sasaki, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Sepiola intermedia Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 162
Sepiola ligulata Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Sepiola parva Sasaki, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
Sepiola robusta Naef, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
Sepiola rondeleti Leach, 1834 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
Sepiola trirostrata Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Euprymna berryi Sasaki, 1929 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
Euprymnamorsei (Verrill, 1881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Euprymna tasmanica (Pfeffer, 1884) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Rondeletiola minor (Naef, 1912) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
Sepietta neglecta Naef, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
Sepietta obscura Naef, 1916 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Sepietta oweniana (Orbigny, 1839?1841) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola aurantiaca Jatta, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola knudseni Adam, 1984 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola pfefferi Grimpe, 1921 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola rossiaeformis Pfeffer, 1884 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola steenstrupiana Levy, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Euprymna albatrossae Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna hoylei Adam, 1986 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna hyllebergi Nateewathana, 1997 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna penares (Gray, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna phenax Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna scolopes Berry, 1913 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna stenodactyla (Grant, 1833) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Sepietta petersi (Steenstrup, 1887) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Inioteuthis capensis Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Inioteuthis japonica (Orbigny, 1845) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Inioteuthis maculosa Goodrich, 1896 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Subfamily ROSSIINAE Appell?f, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Rossia macrosoma (Delle Chiaie, 1830) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Rossia pacifica pacifica Berry, 1911 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Rossia tortugaensis Voss, 1956 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Semirossia equalis (Voss, 1950) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Semirossia tenera (Verrill, 1880) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Neorossia caroli (Joubin, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Austrorossia antillensis (Voss, 1955) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Austrorossia australis Berry, 1918 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Austrorossia bipapillata (Sasaki, 1920) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia brachyura Verrill, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia bullisi Voss, 1956 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia glaucopis Loven, 1845 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia megaptera Verrill, 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia moelleri Steenstrup, 1856 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia mollicella Sasaki, 1920 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia pacifica diegensis Berry, 1912 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Rossia palpebrosa Owen, 1834 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Austrorossia enigmatica (Robson, 1924) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Austrorossia mastigophora (Chun, 1915) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Semirossia patagonica (Smith, 1881) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Neorossia leptodons Reid, 1992 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Subfamily HETEROTEUTHINAE Appell?f, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Heteroteuthis (Heteroteuthis) dispar (R?ppell, 1844) . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Stoloteuthis leucoptera (Verrill, 1878) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
Sepiolina nipponensis (Berry, 1911) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Heteroteuthis (Heteroteuthis) weberi Joubin, 1902 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Heteroteuthis (Stephanoteuthis) dagamensis Robson, 1924 . . . . . . . . . . . . . . . . . . . . . . . 203
Heteroteuthis (Stephanoteuthis) hawaiiensis (Berry, 1909) . . . . . . . . . . . . . . . . . . . . . . . . 203
Heteroteuthis (Stephanoteuthis) serventyi Allan, 1945 . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Nectoteuthis pourtalesi Verrill, 1883 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Cephalopods of the World xi
Iridoteuthis iris (Berry, 1909) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Iridoteuthis maoria Dell, 1959 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Family SEPIADARIIDAE Fischer, 1882 in 1880?1887 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Key to genera in the family Sepiadariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Sepiadarium austrinum Berry, 1921 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Sepiadarium kochii Steenstrup, 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiadarium auritum Robson, 1914 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiadarium gracilis Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiadarium nipponianum Berry, 1932 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepioloidea lineolata (Quoy and Gaimard, 1832) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepioloidea pacifica (Kirk, 1882) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Family IDIOSEPIIDAE Appell?f, 1898 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius biserialis Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius macrocheir Voss, 1962 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius minimus (Orbigny, 1835) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius notoides Berry, 1921 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius paradoxus (Ortmann, 1888) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius picteti (Joubin, 1894) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Idiosepius pygmaeus Steenstrup, 1881 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Idiosepius thailandicus Chotiyaputta, Okutani and Chaitiamvong, 1991 . . . . . . . . . . . . . . . . . . 210
Family SPIRULIDAE Owen, 1836 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Spirula spirula (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
4. LISTOFNOMINALSPECIES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
5. LISTOFSPECIES BY MAJORFISHINGAREAS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
6. REFERENCES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 225
7. INDEX OFSCIENTIFICANDVERNACULARNAMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 249
8. LISTOFCOLOURPLATES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
xii FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
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 industrial-scale fisheries,
increasing attention continues to be paid to the so-called
?unconventional marine resources?, which include
numerous species of cephalopods. Cephalopod catches
have increased steadily in the last 30 years, from about
1 million metric tonnes in 1970 to more than 3 million tonnes
in 2001. This increase confirms a potential development of
the fishery predicted by G.L. Voss in 1973, in his first
general review of the world?s cephalopod resources
prepared for FAO. The rapid expansion of cephalopod
fisheries in the decade or so following the publication of
Voss?s review, meant that a more comprehensive and
updated compilation was required, particularly for
cephalopod fishery biologists, zoologists and students. The
FAO Species Catalogue, ?Cephalopods of the World? by
C.F.E. Roper, M.J. Sweeney and C.E. Nauen was published
in 1984 to meet this need.
The number of cephalopod species that enter commercial
fisheries has continued to grow significantly since 1984, as
a result of a still-growing market demand and the expansion
of fisheries operations to new fishing areas and to deeper
waters. It has been suggested that the cephalopod
? l i fe-st rategy? may guar antee surv ival against
environmentally stressful conditions, including those
caused by heavy fishing. However, as cephalopod fisheries
experienced further extensive development, parallel
concern developed regarding potential overexploitation.
Thus, a broad consensus developed among fishery
biologists to apply the experience gained from errors made
in finfish management to avoid possible failures in
cephalopod exploitation. To help prevent potential failures,
refined species identification capabilities are required, as
well as a more detailed and accurate compilation of
information on cephalopod species, distribution, biology,
fisheries and catch statistics. Consequently, FAO
recognized that a new edition of the ?Cephalopods of the
World? catalogue was needed. To achieve this expanded
goal, several authors with particular areas of specialization
were assembled to enhance the accuracy, coverage and
utility of this revised catalogue.
In our attempt to make this document as comprehensive
and as useful as possible, the taxonomic coverage of this
edition of the catalogue is organized into 3 levels of interest:
Level 1 : species of cephalopods currently exploited
commercially and species utilized at the subsistence and
artisanal levels;
Level 2 : species of occasional and fortuitous interest to
fisheries; this includes species considered to have a
potential value to fisheries, based on criteria such as
edibility, 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 necessary
to provide the most comprehensive inventory of species
possibly useful to mankind, regardless of their current
commercial status. For example, this work should be useful
for the ever-expanding search for development and
utilization of ?natural produc ts?, pharmaceuticals, etc.
The catalogue is based primarily on information available in
published literature. However, yet-to-be-published reports
and working documents also have been used when
appropriate, especially from geographical areas where a
large body of published information and data are lacking,
and we are particularly grateful to colleagues worldwide
who have supplied us with fisheries information, as well as
bibliographies of local cephalopod literature.
The fishery statistics presented herein are taken from the
FAO official database, FISHSTAT, now available on the
Worldwide web (FISHSTAT Plus 2000). This information is
supplemented by field observations made by the authors in
many parts of the world, both in preparation of the 1984
volume, as well as for the current edition. These field visits
provided opportunities to examine fresh material at landing
sites, markets and laboratories, as well as to obtain
first-hand information about local cephalopod fisheries from
regional fisheries workers.
During the 20-plus years separating the two editions, the
rapid development of cephalopod fisheries worldwide and
the simultaneous increase in the population of fisheries
scientists, their research and publications, made available
an enormous amount of new data and research results.
Sometimes it is difficult to evaluate the reliability of
published data, especially with regard to the identification of
species in areas where the cephalopod fauna has not been
sufficiently studied. It is entirely understandable that field
workers isolated from good library and museum/collection
facilities have difficulties in correctly identifying the species
they encounter in the field. Moreover, the discovery of new
species, the more accurate delimitation of known species,
or even the introduction of nomenclatural changes, may
cause confusion and lead to the use of scientific names that
are incorrect by modern standards. Although great care
was exercised to evaluate and correct such published
information used in the catalogue, some incorrect
interpretations may have occurred. Another difficulty, in the
taxonomic literature especially, is that information on the
economic importance of species is rather scarce or of a
very general nature. Also, we may have overlooked
important information published only in relevant local
fisheries literature that is unavailable on a broader scale. All
of these potential difficulties, however, have been
significantly mitigated during the preparation of the new
edition because of the availability on-line of fisheries
databases and bibliographic search capabilities.
With regard to the limitations mentioned above, we heartily
request that readers who detect any errors in the
information presented, or who have any additional
information and data that will enhance the accuracy and
utility of this book, please contact and inform one of the
authors or the Species Identification and Data Programme
(SIDP) of the Marine Resources Service, Fisheries
Resources Division, Fisheries Department, FAO Rome.
Cephalopods of the World 1
For further reading and information on cephalopod biology,
fishery and resources, additional references and websites
are listed at the end of references.
1.1 Plan of the Catalogue
This catalogue is organized by families and their
appropriate genera within major cephalopod groups, then
alphabetically by species.
Level 1 , the most important species for fisheries, consists of
detailed information in all 12 categories listed below.
Level 2 , which comprises those species of occasional,
fortuitous or potential interest to fisheries, consists of
whatever information is available and appropriate in the 12
categories. Level 3 , those species for which there is no
current interest to fisheries, consists of basic information
(i.e. scientific name, size, geographical distribution,
literature). The format within the species sections includes
the first two levels of treatment (Level 1 and Level 2)
presented together. Species included in Level 3 are
presented at the end of each family.
Consequently, each major group and family is introduced
with general descriptive remarks, illustrations of diagnostic
features, highlights of the biology and relevance to
fisheries. The information that pertains to each species in
Levels 1 and 2 is arranged by categories as follows:
(1) scientific name; (2) synonymy; (3) misidentifications;
(4) FAO names; (5) diagnostic features with illustrations;
(6) maximum known size; (7) geographical distribution with
map; (8) habitat and biology; (9) interest to fisheries;
(10) local names; (11) remarks and (12) literature.
(1) Scientific Name: Reference to author, date and
publication is given for the original description of each
species.
(2) Frequent Synonyms: Principal synonyms and name
combinations are listed.
( 3) Misidentifications: Misidentifications as other species
are reported here and discussed in detail under section 11,
Remarks, along with other nomenclatural points.
(4) FAO Names: English, French and Spanish names for
each species, used primarily within FAO, are selected on
the basis of the following criteria: (i) each name must apply
to one species only, in a worldwide context; (ii) the name
must conform to FAO nomenclatural spelling; (iii) the name
should apply only to a cephalopod species, and should not
lead to confusion with species names in other major animal
groups. Wherever possible, these names are selected
based on vernacular names (or parts of names) already in
existence within the areas where the species is fished. FAO
species names, of course, are not intended to replace local
species names, but they are considered necessary to
overcome the considerable confusion caused by the use of
a single common name for many different species, or
several names for the same species.
(5) Diagnostic Features: Principal distinctive characters of
the species are given as an aid for identification,
accompanied by pert inent i l lustrat ions. Species
identifications should be attempted only after verification of
the family through use of the illustrated key to families.
Reference to FAO Species Identification Guides is given
wherever relevant.
(6) Size: The known mantle length (or total length in some
cases) of both males and females is provided where
possible. Sizes or measurements might not be completely
comparable because they were taken mostly from
preserved or fixed specimens, but measurements of
commercially important species often come from fresh
material. Because of the elasticity of tentacles and arms
total length is not a very accurate measurement. Where
both total length and mantle length are given, the respective
figures do not necessarily pertain to the same specimen but
may have been obtained from different sources. The
available information on the size attained by some species
often is very meagre, so the maximum reported size cited
here might be considerably smaller than the actual
maximum size. Maximum weight is given when available.
( 7) Geographical Distribution: The entire known
geographic range of the species, including areas of
seasonal occurrence, is given in the text and shown on a
map. In cases where only scattered records of occurrence
are available, question marks have been used to indicate
areas of suspected or unconfirmed distribution.
(8) Habitat and Biology: The known depth range of the
species and information on salinity and temperature of its
habitat are given where available. For the sake of
exactness actual depth data are reported, as given in the
literature referenced. Information on biological aspects,
such as migrations, spawning seasons and areas,
longevity, food, and predators, is also included.
(9) Interest to Fisheries: This paragraph gives an account
of the areas where the species is fished and of the nature of
the fishery; its importance is either qualitatively estimated
(minor, moderate, major or potential) or actual figures of
annual landings are provided. Data on utilization (fresh,
dried, cooked, frozen, canned, etc.) are also given where
available. Here, too, the quality and quantity of the available
information varies considerably among the species.
(10) Local Names: These are the names used locally for the
various species. The present compilation is necessarily
incomplete, since only a fraction of the local names applied
to specific entities is actually published. In many cases,
local names are available only for species that support
traditional fisheries. Apart from possible omissions due to
limitations of literature available, some of the names
included may be somewhat artificial, i.e. through
transliteration of indigenous words into English. The local
species name is preceded by the name of the country
concerned in capital letters and, where necessary, by
geographical specifications in lower case letters.
(11) Remarks: Important information concerning the
species, but not specifically linked to any of the previous
categories, is given here. For example, in some cases the
taxonomic status of certain scientific names requires further
discussion. Other nomenclatural problems are discussed in
this section, such as the use of subspecies names.
(12) Literature: This includes references to the most
important publications relevant to the species, the
emphasis being on biology and fisheries. Additional
references are included in the bibliography. In the case of a
few uncommon species, only systematic papers are
available.
2 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
1.2 General Remarks on Cephalopods
The group known as cephalopods (class Cephalopoda) is
the most complex in the phylum Mollusca, and indeed, in all
of the invertebrate phyla. It includes exclusively marine
animals that live in all oceans of the world with the exception
of 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 molluscan
taxonomic entity, the nautiloids, in the Upper Cambrian
period (over 500 million years ago), but more than half of
these ancestors were already extinct by the end of the
Silurian, 400 million years ago, when only the nautiluses
survived. Meanwhile, other forms arose in the late
Palaeozoic (between 400 and 350 million years ago),
including those of the Subclass Coleoidea, but most of
them became extinct by the end of the Mesozoic, about
150 million years ago. The only members of the subclass
Coleoidea that exist today are the forms that developed in
the Upper Triassic and Lower Jurassic (between 200 and
150 million years ago).
Although there is a long fossil record of many different
groups, all living cephalopods belong to two ?subclasses?:
the Coleoidea, which includes the major groups known as
squids, cuttlefishes sensu lato, octopods and vampires,
and the Nautiloidea, containing two genera, Nautilus and
Allonautilus, the only surviving cephalopods with an
external shell.
At the present time the status and understanding of the
Systematics and Classification of the Recent
Cephalopoda is under considerable discussion. The
families of living cephalopods are, for the most part, well
resolved and relatively well accepted. Species-level taxa
usually can be placed in well-defined families. The higher
classification, however, sti l l is not resolved. The
classification above the family level is controversial and a
broad consensus still needs to be achieved. This situation
is not unexpected for a group of organisms that has
undergone explosive research attention in recent decades.
Consequently, rather than accept and promote any
particular scheme of classification, before consensus and
stability are achieved, we will use an ?operational
breakdown? that is satisfactory for the objectives of this
Catalogue. For practical purposes we separate the
cephalopods into several groups, without assigning or
implying taxonomic relationships. Figure 1 diagrams
several of the classification schemes currently under
discussion.
In this work the following groups are used, as illustrated in
Figure 21/:
Nautiluses
Cuttlefishes
Bobtail squids
Bottletail squids
Pygmy squids
Ram?s horn squid
Myopsid squids
Oegopsid squids
Vampires
Cirrate octopods
Incirrate octopods
Unresolved taxa:
Spirula
Idiosepius
Bathyteuthis
Chtenopteryx
Sepiadariidae
Plural versus singular usage of cephalopod common group
names is standardized as follows:
squid, cuttlefish, octopod, oc topus, vampire, nautilus
refer to one individual or one species;
squids, cuttlefishes, octopods, octopuses, vampires,
nautiluses refer to two or more individuals and/or species.
These terms are also used to indicate the major groups.
The term ?cuttlefishes? is also used ? sensu lato? to indicate
the following groups: Cuttlefishes, Bobtail squids, Bottletail
squids, Pygmy squids and the Ram?s horn squid.
We differentiate between the members of the family
Octopodidae, which are called octopus/octopuses and the
members of the whole group (Incirrate and Cirrate or any
combination of non-Octopodidae taxa), which are called
octopod/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 species
occur in bays, seas; and epipelagic, mesopelagic,
bathypelagic and benthopelagic species are all present in
the open ocean. Salinity is considered to be a limiting factor
in cephalopod distribution; they are generally restricted to
salinity concentrations between 27 and 37?. However,
Lolliguncula brevis, which lives and reproduces in waters
of 17?, demonstrates a capacity for a higher degree of
salinity tolerance (Hendrix et al., 1981). Some species
inhabit the Red Sea and the southern coasts of the Iberian
Peninsula (Guerra, 1992), where the salinity is higher than
37? and other species have been found in waters where
salinity ranges between 25 and 18? (Sea of Marmara;
Unsal et al., 1999). The habitat depth range extends from
the intertidal to over 5 000 m. Many species of oceanic
cephalopods undergo diel vertical migrations: they occur at
depths of about 200 to 700 m during the day, then at the
onset of twilight and increasing darkness, they ascend into
the uppermost 200 m for the night. A deeper-living layer of
diel migrators occurs from about 1 000 m to 600 m during
the daytime. The abundance of cephalopods varies,
depending on group, habitat and season, from isolated,
territorial individuals (primarily benthic octopods and
sepioids), through small schools of squids with a few dozen
individuals, to huge schools of neritic and oceanic species
with millions of specimens.
The size of adult cephalopods ranges from less than 1 cm
(Jackson, 1989) to the giant squid at approximately 20 m in
total length, including the tentacles. The largest specimens
may weigh well over 500 kg, but the average size of
commercial species is 20 to 40 cm mantle length and about
0.1 to 2.0 kg total weight.
Cephalopods are soft-bodied, bilaterally symmetrical
animals with a well-developed head and a body that
consists of the muscular mantle, the mantle cavity that
houses the internal organs, and, when present, the external
fins. The head bears an anterior circumoral (surrounding
Cephalopods of the World 3
1/ The endings used in the group names do not imply any level of
classification.
4 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Roper et 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
Young et 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
Uniductia
Spirulida
Myopsida
Loliginida
Sepiida
Octobrachiomorpha
Vampyromorpha
Octopoda
Cirrata
Incirrata
Fig. 1 Some conflicting suprafamilial classifications of living coleoid cephalopods
Cephalopods of the World 5
Fig. 2 Living cephalopods
the mouth) crown of mobile appendages (arms, tentacles).
This characteristic feature reflects the origin of the name
Cephalopoda, which derives from the union of the two
Greek words: ?kefale?, head, and ?pous?, feet. The name
was erected by Schneider in 1784, and it became
permanently in use within the scientific context with the
publication of Cuvier?s work (1798). Arms and tentacles
bear suckers and/or hooks (except in Nautilus), which are
powerful tools to seize prey. The mouth has a pair of
chitinous jaws (the beaks) and, as in other molluscs, a
chitinous tongue-like radula (band of teeth) occurs in most
cephalopod species. The ancestral mollusc shell is
variously modified, reduced, or absent in living coleoids. It
is a calcium carbonate structure in cuttlefishes (the
cuttlebone of sepiids and the ram?s horn shell of Spirula),
reduced to a rigid structure composed of chitin in squids
(the gladius or pen, sometimes quite flexible) and to a
cartilaginous structure in finned octopods. In some
sepiolids no vestige of shell is found. A true external shell
occurs only in the primitive nautiluses (restricted to the
Indo-Pacific), although a shell-like egg case is produced
and carried by female argonauts (pelagic octopods often
misnamed ?paper nautilus?).
The loss of the external shell allowed the development of a
powerful muscular mantle that became the main
locomotory organ for fast swimming, via water jetting from
the funnel. The funnel (siphon) is a unique, multifunctional,
muscular structure that aids in respiration and expulsion of
materials in addition to locomotion. Oxygenated water is
drawn through the mantle opening around the head (neck)
into the mantle cavity, where it bathes the gills for
respiration. Mantle muscular contraction expels the
deoxygenated water from the mantle cavity through the
ventrally located funnel. The discharge jet serves to
eliminate nephridial and digestive wastes, as well as to
complete the respiratory cycle and for locomotion. Female
reproductive products (eggs, egg masses) also are
discharged through the funnel. Most coleoids produce ink,
a dark, viscous fluid also expelled through the funnel. The
ink may take the form of a mucoidal ?pseudomorph? (false
body) to decoy potential predators, or of a cloud to obscure
the escaping cephalopod.
One pair of gills (ctenidia) is present, except in Nautilus and
Allonautilus, which have two pairs for respiration, i.e. to
extract the oxygen from the water. However, in contrast
with coleoids, Nautilus makes use of anaerobic respiration
during periods of high activity and can survive in water with
very low oxygen content. Coleoids also use anerobic
muscle layers. Cutaneous respiration also occurs in some
cephalopods.
The cephalopod circulatory system is distinctive within the
Mollusca. It is a closed system (blood contained within
vessels), similar in many respects to that of vertebrates and
fulfilling the demand for the more efficient circulation
required by an active locomotory system. There is a
principal, or systemic, heart, two branchial hearts and
developed arterial, venous and capillary systems supplying
blood to the muscles and organs. From the gills, the
oxygenated blood goes through the efferent branchial
vessels to the systemic heart, where it is expelled from the
ventricle through three aortas: the cephalic or dorsal aorta,
which supplies the head and the anterior part of the gut; the
posterior, minor or abdominal aorta that supplies the mantle
and fins along with the posterior part of the gut and the
funnel; and the gonadal aorta that develops gradually with
sexual maturation of the animal. The blood is collected
through sinuses and capillaries into the veins, through
which it goes to the branchial hearts that pump it through
the filaments of the gills. The circulating respiratory pigment
used for oxygen transport is copper-contain ing
haemocyanin, a system of rather lower efficiency than the
iron-containing haemoglobin of vertebrates. In living
cephalopods blood sinuses are much reduced and
replaced functionally by muscles. The circulatory system
therefore has to work against the per ipheral
muscle-induced pressure, which increases with increasing
activity (maximum during jet-swimming). It also has to cope
with the resistance of the small diameter of the final
capillary blood vessels, and the low oxygen carrying
capacity of the blood (less than 4.5% by volume). In spite of
these limitations, the system has other functional
modifications (see for example Wells and Smith, 1987;
Martin and Voight, 1987) that achieve the capacity to deliver
oxygen at a rate comparable to that of active fishes,
enabling cephalopods to accomplish extraordinary
performances.
The excretory system also differs markedly from that of
other molluscs and, along with the closed circulatory
system and the branchial circulation, enables unique
relationships between blood and the final secretion, the
urine. The excretory system differs between living
nautiloids and coleoids and also among coleoids, but the
general organization is similar, consisting basically of the
renal sac with the renal appendages (organs comparable to
vertebrate kidneys), the pericardial glands, the branchial
hearts and the gills. Cephalopods are ammoniotelic, and
ammonium ions are continuously released by the gill
epithelium and by renal appendages into the surrounding
water. Ammonium ions are used by buoyant squids to
replace denser chloride ions in fluids in the coelom and in
the body tissues. Because this solution is less dense (and
hence more buoyant) than seawater, it provides lift.
The nervous system is highly developed in recent
cephalopods, with a large brain and peripheral connections,
contrasting with the original molluscan circumesophageal
nerve ring. Among its most remarkable features is the giant
fibre system of squids and cuttlefishes that connects the
central nervous system with the mantle muscles. This
system consists of three orders of cells and fibres and
ensures the immediate and simultaneous contraction of
mantle, fins and retractor muscles of both sides, rather than
an anterior to posterior sequential contraction that would be
counter-productive for water movement (expulsion). Also
remarkable is the eye development of most coleoids, for
which vision plays a major role in life. Their eyes are large,
have a design generally similar to that of fishes and other
vertebrates (e.g. a lens focuses images on the retina), and
all the available evidence suggests the ocular/visual
performance to be comparable to that of vertebrates. The
cephalopods also developed a system to keep the focused
image stationary on the retina while the animal turns, by
moving the eyes in coordination with the head/body
movement. This is extremely important for hunters that rely
on sight, and it is accomplished by connections of the eye
muscles with the statocysts, a mechanism similar to the
vestibulo-optic system of fishes. The statocyst system
provides cephalopods with information on their orientation,
as well as changes in position and direction of movement.
The statocyst system is highly developed in coleoids, where
it consists of separate cavities located in the cartilaginous
skull, posteroventral to the brain. The statocysts contain
6 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
nervous cells and receptors differentiated to detect both
linear acceleration, with the aid of calcareous stones called
statoliths, and angular acceleration. Many coleoids also
have extra-ocular photoreceptors (photosensitive vesicles)
about which little is known; in mesopelagic squids they
appear to monitor light intensity in order to enable the
animals to match the counter-illumination with the ambient
light by their own photophores (light-producing organs).
Cephalopods are provided with numerous mechano- and
chemoreceptors and recent evidence indicates that in
some species, like Loligo vulgaris, Sepia officinalis and
Octopus sp., ciliate cells form lines in several parts of the
body, a system analogous to the lateral-line system in
fishes.
Most Coleoidea are able to change colour by using a complex
system of chromatophores under nervous control. The
chromatophores are pigment-filled sacs present in the skin,
and capable of remarkable expansion and contraction. This
system 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 taxonomic
group and habitat. Most species also have iridocytes (shiny,
reflective platelets) in the skin. Most cephalopod behaviour
includes rapid changes in overall colour and colour patterns,
as well as changes in the texture of the skin, from smooth to
heavily papillate, tubercular, or with erected flaps. While
shallow-living cephalopods are able to conceal themselves by
chromatophore-produced colour patterns, chameleon-like
colour changes and textural presentations, many deep-sea
forms camouflage themselves by producing bioluminescent
light from photophores which eliminate their silhouettes
against the downwelling sunlight in the dimly-lit mid-depths.
Locomotion is achieved by any of, or a combination of, the
following methods, depending on the taxonomic group:
1) jet propulsion; 2) flapping or undulating the fins on the
mantle; 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 steering
during jet propulsion. Many families of midwater squids
have evolved to ?low energy life styles? and achieve neutral
buoyancy by producing and storing in tissues or in different
organs substances/elements with specific properties, such
as oils or solutions of ammonium ions. This capability
enabled coleoid cephalopods to inhabit open water, even in
the great depths in the ocean, the greatest volume of living
space on earth.
Cephalopods are voracious, active predators that feed
upon crustaceans, fishes, other cephalopods and, in the
case of some benthic octopods, on bivalved molluscs. The
speed of cephalopods, their high mobility and powerful
visual systems, along with strongly-muscled arms and
tentacles, both equipped with suckers and/or hooks, make
them extremely efficient hunters. A common hunting
technique in sepioids and loliginids involves extremely
rapid shooting forward of the tentacles to capture the prey,
while in some oegopsid squids the tentacles may be used
like long, sucker-covered fishing lures. Some octopods use
their web to envelop crabs and occasionally may wait until
prey touches them before attacking. The captured prey of
cephalopods is brought to the mouth and killed by bites of
the strong, chitinous beaks, equipped with powerful
muscles. Sometimes the prey are first paralysed with a
fast-acting toxin, immobilization being a strong advantage
in case of large and/or very active prey such as large crabs.
The Incirrate octopods and Sepia species usually poison
their prey before eating them, while squids do not seem
able to produce such strongly toxic secretions. Digestion in
cephalopods is rapid and efficient and cephalopod
metabolism is essentially proteinic: there is little or no
digestion/assimilation of carbohydrates and lipids. Food
conversion is highly efficient, especially in octopuses,
where up to 50% of the food eaten can be converted into
body mass. More active cephalopods like squids, however,
need several times the amount of food required by
octopuses and can eat from 3 to 15% of their body weight
each day.
All cephalopods are dioecious (separate sexes) and many,
though not all, exhibit external sexual dimorphism, either in
morphological or in size differences. Females frequently
are larger than males, a phenomenon which reaches its
extreme in some pelagic octopods (such as Argonauta)
where males are truly dwarf forms. Males of many species
possess one, occasionally two, modified arm(s) (the
hectocotylus) for transferring spermatophores to females
during mating. The males of some species also exhibit
modifications to other arms, in addition to the hectocotylus.
The hectocotylus may be simple or complex and can
consist of modified suckers, papillae, membranes, ridges
and grooves, flaps. The one or two limbs function to transfer
the spermatophores (tubular sperm packets) from the
male?s reproductive tract to an implantation site on the
female. The spermatophores may be implanted inside the
mantle cavity (where they may penetrate the ovary), into the
oviducts themselves, around the mantle opening on the
neck, on the head, in a pocket under the eye, around the
mouth or in other locations. Females of a few species also
develop gender-specif ic structures (e.g. arm-t ip
photophores) when mature. Mating is often preceded or
accompanied by courtship behaviour that involves striking
chromatophore patterns and display. Copulatory behaviour
varies significantly among species, in colour and textural
display, proximity of male and female, duration of display
and spermatophore transfer, and the location of
implantation of the spermatophores on the female.
The gonads form a single mass at the posterior end of the
mantle cavity, and female gonoducts may be paired (in
oegopsids and incirrate octopods) or single, as in other
coleoids and in the nautiluses. Cephalopod reproductive
systems are highly complex structures with ducts, glands
and storage organs. Female octopods have oviducal
glands, while decapods, in addition, have nidamental
glands and, in some families, accessory nidamental glands.
Spermatophores are produced in the mul t i -uni t
spermatophoric gland and stored in the Needham?s sac,
from which they are released through the terminal part of
the duct, the penis. This term is not strictly accurate in many
groups, because the spermatophores are passed to, or
taken by, the hectocotylized arm(s), which in turn transfer(s)
the spermatophore(s) to the female. Some families do not
possess a hectocotylus. Instead, the terminal portion of the
male reproductive tract forms a functional penis that can be
greatly enlarged and elongated, often extending out of the
mantle cavity and past the head. It is likely that these
structures directly transfer the spermatophores to the
female. The number and size of spermatophores vary
greatly, 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. The
spermatophores evert, with the resultant extrusion of the
Cephalopods of the World 7
sperm packet caused by the penetration of water inside the
spermatophoric cavity, where the osmotic pressure is
higher. The resulting extruded sperm packet is named
spermatangium (or sperm bulb or body). Sperm are able to
survive several months once stored in the female, at least in
some species, and fertilization of mature ova may take
place either in the ovary, the oviducal glands, the mantle
cavity or the cone formed by the outstretched arms while
the eggs are laid. Males of the pelagic octopod Argonauta
(and its relatives) insert sperm into a detachable
hectocotylized arm. Mating occurs by the arm severing and
crawling into the female?s mantle cavity.
Fertilized eggs are embedded in one or more layers of
protective coatings produced by the oviducal and
nidamental glands and generally are laid as egg masses.
Egg masses may be benthic or pelagic, varying among the
major taxonomic groups. Some oceanic mesopelagic and
deep-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 containing
from a few to several hundred eggs. Deposited in
multifinger masses (sometimes called ?sea mops?), these
eggs are attached to rocks, shells or other hard substrates
on the bottom in shallow waters. Many oceanic squids lay
their eggs into large sausage-shaped or spherical
gelatinous masses containing tens or even hundreds of
thousands of eggs that drift submerged in the open sea.
Other pelagic species lay individual eggs, not enmeshed in
gelatinous masses.
Cuttlefishes sensu lato lay few, relatively large (around 10
to 40 mm in diameter), grape-like eggs that are attached to
hard substrates. Some species camouflage the eggs from
predators by making them dark using a coating of ink
deposited 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 (see
Boletzky, 1982, 1986), and they are laid individually or in
small clusters of a very few eggs.
Benthic incirrate octopods lay their eggs singly or in
grape-like clusters and strands. Most species attach the
eggs to hard surfaces such as rock or shells while some
carry the eggs within their arms and webs. The eggs vary in
size from a few millimetres to 40 mm long. They are
attached to each other and/or to the substratum by cement
produced by the oviducal gland. They lack the gelatinous
outer matrix found in squid and cuttlefish eggs and the outer
shell found in the Cirrate octopod eggs: the protective
function has been ?replaced? by the brooding behaviour of
the female parent. The female of the pelagic octopod
Argonauta constructs a thin, decorative shell-like egg
case, which encases her mantle and into which she
periodically lays and attaches festoons of minute eggs.
The mode of reproduction and egg-laying still is unknown
for many species, especially those of oceanic and deep-sea
environments.
Development of cephalopod embryos is direct, without true
metamorphic stages. Hatchlings of species with large eggs
look like miniatures of the adult, while hatchlings of species
with small eggs undergo gradual changes in proportions
during development. The young of some species, however,
differ conspicuously from the adults. Thus, the term
?paralarva? has been introduced for these early stages of
cephalopods that differ morphologically and ecologically
from older stages. Hatchlings from large-egged benthic
octopods are either benthic crawl-away young or
temporarily planktonic, quickly settling back to the adult
benthic habitat. Small-egged benthic octopuses produce
planktonic hatchlings with very simple skin patterns of large
chromatophores. The paralarvae of many deep-sea
species of squids and octopods occur in the upper 100 m of
the open ocean; then they exhibit an ontogenetic descent,
gradually descending to deeper depths with increasing size
until the adult depth is attained. Time of embryonic
development varies widely, from a few days to many
months, depending on the species and the temperature
conditions. Hatching may occur synchronously from a
single clutch or be extended over a period of 2 or 3 weeks.
In spite of the large number of studies and research carried
out on cephalopods, especially in recent decades, the life
history of the majority of species is still unknown, and our
knowledge of the life cycles of the members of this
interesting class remains fragmentary. Information comes
from studies in the field as well as from observations in the
laboratory. However, little is known of life history for species
that are not targets of regular fisheries, and only a handful
of cephalopod species have been reared successfully in the
laboratory.
Studies and monitoring of growth in cephalopods are
complicated by the high variability in individual growth
rates. This makes it difficult to apply conventional methods,
e.g. length frequency analysis, used for more traditional
resources such as fishes and crustaceans. Determination
of age in Recent coleoids is also difficult, because they
have 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 of
statoliths that has resulted in an increased knowledge of
age, in squids at least. This has led to changes in our
perspectives about the physiology and ecology of many
teuthoids, but more research is required before a full
understanding is achieved (see Jereb et al., 1991; Okutani
et al., 1993; Jackson, 1994 and Lipinski and Durholtz, 1994
for reviews and discussions). Principle results obtained
from the research generally confirm a very high growth rate
in cephalopods, comparable to that of the fastest-growing
fishes.
Nautilus species have a life span of about 20 years, during
which they spawn intermittently when first mature, then
annually when fully mature, laying a few large eggs at a
time. The life expectancy of most coleoids appears to range
from a few months to one or two years. Many small oceanic
squids, such as pyroteuthids may complete their life cycles
in less than six months, while some minute forms of
sepioids have a life span of only 2 to 3 months. Recent
evidence, however, suggests that larger species of squids
and octopods, for example the giant squid (Architeuthis
spp.) and the giant octopus (Enteroctopus dofleini), as well
as those that live in coldest habitats, may live for several
years.
A general consensus exists that spawning is a terminal
event, 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
8 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
strategies (Rocha et 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 the
typical example of this life strategy.
B) Multiple event spawning , formerly termed iteroparity,
with the following possibilities:
1 - polycyclic spawning: egg-laying occurs in separate
batches during the spawning season and growth
occurs between production of egg batches and
subsequent spawning events (e.g. Nautilus species);
2 - multiple spawning, with group-synchronous
ovulation, monocyclic spawning and growth between
egg batches (e.g. Sthenoteuthis oualaniensis);
3 - intermittent terminal spawning with group-
synchronous ovulation, monocyclic spawning and no
growth between egg batches (e.g. Sepia officinalis,
Loligo vulgaris, Illex coindetii);
4 - continuous spawning, with asynchronous ovulation,
monocyclic spawning and growth between egg
batches (e.g. Idiosepius pygmaeus, Opisthoteuthis
spp., Argonauta spp.).
All coleoids die after their spawning period.
The total number of living species of cephalopods that
have been described is fewer than 1 000; over 720 are
listed in the present catalogues. However, many species
still wait to be described and this is particularly true for the
octopuses, where at present at least another 100
undescribed species are estimated to exist.
The status of the systematics of cephalopods has changed
in the last 30 years, as research and associated scientific
discussions have increased substantially. However,
phylogenetic relationships among most families within the
major groups remain uncertain, and new species are
described fairly frequently as new habitats are explored and
as families are gradually better-understood.
The major groups of living cephalopods, i.e. squids,
cuttlefishes, bobtail and bottletail squids, octopods,
vampires and chambered naut i luses, are easi ly
distinguished by external characteristics.
The squids have an elongate, cylindrical body with
posterolateral fins on the mantle (rarely, the fins extend for
the length of the mantle); 10 circumoral appendages
anteriorly on the head, not connected at bases with a web
(except Histioteuthidae); 8 arms with 2 (occasionally 4 or
more) series of stalked suckers with chitinous rings (and/or
chitinous hooks in some groups) that extend along the
entire arm length; 2 longer tentacles with an organized
cluster of 2 or more series of stalked suckers (and/or hooks)
at the distal section (tentacular club); the proximal
tentacular stalks usually are devoid of suckers or hooks.
The cuttlefishes have broad sac-like bodies with lateral fins
that are narrow and extend along the length of the mantle;
posterior lobes of the fins free (subterminal) and separated
by the posterior end of the mantle; 10 circumoral
appendages, the longest 2 (tentacles) are retractile into
pockets on the ventrolateral sides of the head; the 8
remaining arms frequently with 4 series of stalked suckers
with chitinous rings, never hooks (otherwise 2 series); eyes
covered with a transparent membrane and eyelids present;
dorsal, internal, finely chambered shell, thick, chalky,
calcareous (cuttlebone).
The bobtail and bottletail squids have sac-like bodies with
short, round and flap-like fins, attached laterally about
midway along the mantle, with pronounced anterior lobes
and free posterior lobes; 10 circumoral appendages as
above, 8 short arms, 2 retractable tentacles, each with a
well defined club; large eyes covered with a transparent
membrane; shell thin, chitinous or lost.
The octopods have a short, sac-like body, either with no
lateral fins or with separate, paddle-like fins in the deep-sea
cirrate octopods; 8 circumoral arms only (no tentacles) with
bases connected by a membranous web, and unstalked
suckers in one or two series, without chitinous rings or
hooks, along the length of the arms.
The vampires have 8 circumoral arms, plus an additional
pair of long, thin, suckerless filaments, retractile into
pockets, located between arms I and II; one pair of
posterior, elongate-oval fins (2 pairs in juveniles);
numerous small photophores over body, posteriorly a pair
of large, external photophores; single series of suckers
bordered with cirri on each arm.
The chambered nautiluses are characterized by an
external, smooth, coiled, chambered shell, numerous
circumoral appendages without suckers (up to 100
appendages), a ?funnel? of rolled muscular flaps or lobes
not fused in the ventral midline and simple eyes without
lenses.
More detailed descriptions of these groups are presented in
the appropriate systematic section.
Cephalopods are important experimental animals in
biomedical research with direct applications to humans.
Because of their highly developed brains and sensory
organs, they are valuable in behavioural and comparative
neuroanatomical studies. In addition, the extremely large
single nerve axons of some cephalopods, the largest in the
animal kingdom, are used extensively in neurophysiological
research.
The bite of cephalopods, especially octopuses, can be
painful at the least to humans, poisonous or secondarily
infected, or, rarely, lethal. Several human deaths have
been recorded in the western Pacific Ocean and Australia
as a result of poisonous bites from the small blue-ringed
octopus, Hapalochlaena spp.; the lethal toxin is a neural
blocker considered a form of tetrodotoxin. Another
documented threat by squids to humans is from the large
ommastrephid squid, Dosidicus gigas, which forms large
aggressive schools that are known to have attacked
fishermen that have fallen in the water, causing several
confirmed deaths. Scuba divers also have been attacked.
Therefore, cephalopods must be handled carefully.
Cephalopods of the World 9
1.3 Interest to FisheryandRole in theEcosystem
Cephalopods are very important for human consumption.
They have been fished on an artisanal basis for several
thousand years and were already well known and highly
valued as food by ancient Greeks, Romans and Chinese. In
more recent times, increasingly heavy demand for
cephalopods by the Japanese led to the development of a
commercial fishery that started in the 1960s and rapidly
expanded to become global in scope. Fishing pressure on
cephalopods has increased as stocks of finfishes have
been depleted worldwide, and cephalopod resources now
are exploited throughout the world oceans.
A catch of around 3.3 million tonnes was reported for 2001
by FAO statistics (FAO, 2000); this figure represents about
3.6% of the world total marine catch for the same year.
Estimated value for this catch was around US$5 billion, a
value that positions cephalopods in fifth place in the
monetary value scale, following miscellaneous coastal
fishes, shrimps, tunas, cods and hakes.
Table 1 reports the world catch of cephalopods since 1950,
by major fisheries ?categories?. The discrepancies in the
last digit of the totals in the tables are caused by rounding
off the individual figures. Squids, both inshore and oceanic,
are by far the main component in the world cephalopod
fishery production, accounting for about 70% of the total
catch, followed by cuttlefishes (species of Sepia, Sepiella
and allied genera), then by octopuses (mainly Octopus spp.
and Eledone spp.). The impressive increase in production
during the last 20 years is due mainly to the increase of
squid catches, from a little more than 1 million tonnes in
1980 to over 2.2 million tonnes for 2001; this value
represents an increase of 250%. Such a significant
increase in production was mostly related to the ?discovery?
and increasing exploitation of squid resources in the
southwest Atlantic (Fig. 3b), principally for Illex argentinus,
(Fig. 4b), as well as an increase in the production of other
major squid target species, mainly Todarodes pacificus in
the northwest Pacific and Dosidicus gigas in the eastern
Pacific (Figs 3d?e and 4b). Illex argentinus catches
exceeded 1 million tonnes in 1999 (Table 2), a record peak
which placed this species at the eleventh position in value
of the total world marine-species production of that year.
Fluctuations in squid catches are responsible for the major
fluctuations in total cephalopod landings, changes usually
related to a combination of environmental, marketing
and/or political reasons. For example, the low cephalopod
catch in 1998 in the eastern Pacific, reflected the poor catch
of Dosidicus gigas as a result of the El Ni?o weather system
in the southern and central areas that year. By contrast, the
low catch in the southwest Atlantic and in the northwest
Pacific in the same year was mainly a result of low fishing
activities by the main squid-producing countries (i.e. Korea,
Japan and Taiwan Province of China). This was caused by
a lower market demand due to very high catches of the
previous year, together with higher fishing fees set by
Argentina on foreign trawlers fishing in its waters that year.
Cuttlefishes sensu lato production also increased strikingly
over the last two decades, doubling between 1980 and
2001 (Table 1; Fig. 4d). The main producer is China (over
50% of the total), followed by Thailand (western Pacific and
central Indo-Pacific areas) and by Morocco (eastern central
Atlantic), where production increased steadily in recent
years.
Octopus catches have increased as well, but not as
conspicuously (Table 1; Fig. 4e). The main producer is
Morocco, which accounted for slightly more than 35% of the
total reported octopus production in 2001, followed by
Japan (about 14% of total world production). Mexico ranks
first in the ? Octopus vulgaris? landings, with over 20 000
tonnes reported for 2001, followed by Spain (nearly 15 000
tonnes); Mexican production accounts for more than
one-th i rd (38.7 % ) of tota l ? Octopus vulgaris ?
species-complex reported production.
Cephalopod fisheries and landings are unevenly distributed
in the world?s oceans, as shown in Table 3 (Antarctic zone
captures are not reported in the figure because they are
minimal; these catches are discussed in the individual
species sections). More than half of the total cephalopod
catch (about 58% in 2001) is taken in the northwest Pacific
and the southwest Atlantic combined. Other important
areas are the western central Pacific, eastern Atlantic and
eastern Pacific, but smaller-scale fishing activities in other
areas also developed consistently in the last decades (see,
for example, the Indian Ocean fisheries). Cephalopod
fisheries are especially intense in Japan, southeastern Asia
and China and in the Mediterranean Sea, where, however,
cephalopod catches show a decreasing trend for all three
main groups in the last decade (Fig. 3g).
Japan remains the leading cephalopod-producing country;
over 560 000 tonnes were caught by Japanese fleets in
2001, which represents about 17% of the total world
cephalopod catch (Table 4). However, China, Korea and
Argentina follow closely. In fact, Korea was the main
producer in 1999, principally as a consequence of the peak
in Illex argentinus production.
International commerce of cephalopod products also
increased strikingly during the last 20 years. Imported and
exported cephalopod products were estimated at around
325 000 and 240 000 tonnes, respectively, in 1980,
compared with more than 1.3 million tonnes estimated for
2001 (Table 5).
Estimated cephalopod import value reached US$ 2 700
billion in 2001. This represents about 4.6% of the value of
the total world fisheries commodities imports for the same
year, while the value of cephalopod exports was a little
higher than US$ 2 500 billion, i.e. about 4.5% of the total
world fisheries commodities export value.
Argentina was the lead exporter in the 2001 statistics in
terms of quantity, followed by Spain, then by Morocco and
the United States (Table 5). However, the highest monetary
values of exports are those attained by Morocco (frozen
Octopus spp. and squids, both Loligo spp. and Todarodes
sagittatus), followed by Thailand (frozen cuttlefishes and
squids, as well as cephalopod ?preparations?) and Viet Nam
(dried squids, followed by various frozen cephalopods,
including Octopus spp.).
European countries are the main market at present for
imported cephalopod products, with Spain, Italy, Greece,
Portugal and France accounting for 49.6% of the world total
(Table 5). Spain has been the leading European importer
(by tonnage) in the world since 1997. The highest import
values, however, remain those spent by Japan, mainly for
frozen Octopus spp., followed by various other cephalopod
commodities.
10 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
(Source: FAO, 2000)
Cephalopods of the World 11
Squids Total 1 / Loligo
spp.
Ommastrephidae Non-identified Squids Octopus es Cuttlefish es
Non-identified
Cephalopods
World Cephalopod
Total
mt % mt mt mt mt % mt % mt % mt
1950 494.5 85 12.4 431.4 50.7 35.8 6.20 47.2 8.10 3.0 0.50 580.5
1951 548.5 85 9.9 483.1 55.5 39.9 6.20 55.6 8.60 3.0 0.50 647.0
1952 686.4 85 15.7 605.3 65.4 49.7 6.20 67.9 8.40 3.0 0.40 807.0
1953 505.9 81 14.8 429.4 61.7 53.6 8.60 62.6 10.00 3.0 0.50 625.1
1954 470.4 78 13.4 400.2 56.8 56.6 9.40 69.6 11.60 3.0 0.50 599.7
1955 477.2 78 16.9 393.8 66.5 60.0 9.80 72.7 11.80 4.0 0.70 613.9
1956 402.4 75 16.7 313.7 72.0 60.9 11.30 70.4 13.10 4.0 0.70 537.7
1957 479.6 76 17.8 393.5 68.3 67.4 10.70 77.3 12.30 4.6 0.70 628.9
1958 468.2 75 18.4 391.4 58.4 68.4 10.90 86.7 13.80 4.8 0.80 628.0
1959 611.9 78 17.8 512.1 82.0 70.7 9.00 95.0 12.10 4.3 0.50 781.9
1960 612.2 78 23.1 510.7 78.4 76.0 9.70 94.6 12.00 4.5 0.60 787.3
1961 575.1 76 24.1 481.2 69.8 79.9 10.60 94.3 12.50 4.1 0.50 753.4
1962 708.1 78 24.6 592.4 91.1 93.0 10.30 100.6 11.10 4.1 0.50 905.8
1963 830.2 81 24.8 705.4 100.0 95.1 9.30 96.2 9.40 4.8 0.50 1 026.3
1964 448.8 66 27.2 348.4 73.2 97.9 14.50 125.5 18.50 5.0 0.70 677.2
1965 616.3 70 33.9 498.9 83.5 116.4 13.20 144.4 16.40 4.8 0.50 881.9
1966 602.8 71 33.5 481.5 87.7 111.1 13.00 133.6 15.70 5.7 0.70 853.1
1967 684.6 70 31.8 539.6 113.2 146.1 14.90 146.8 14.90 5.0 0.50 982.5
1968 902.1 72 37.1 764.2 100.7 188.9 15.10 154.4 12.40 4.8 0.40 1 250.2
1969 697.5 69 34.9 548.6 114.1 163.8 16.10 151.2 14.90 4.6 0.50 1 017.1
1970 680.3 69 76.4 501.7 102.1 153.1 15.50 151.4 15.30 6.3 0.60 991.1
1971 640.1 66 79.3 427.0 133.8 150.6 15.40 178.8 18.30 7.6 0.80 977.1
1972 805.4 66 112.3 550.9 142.2 212.1 17.50 181.3 15.00 13.5 1.10 1 212.4
1973 717.5 68 111.3 434.4 171.9 178.9 16.80 150.0 14.10 15.6 1.50 1 062.0
1974 718.0 66 117.9 388.1 211.9 228.0 20.80 139.5 12.70 10.3 0.90 1 095.8
1975 797.8 67 120.6 458.5 218.7 249.1 20.80 137.3 11.50 13.7 1.10 1 197.9
1976 782.5 64 104.6 429.9 248.0 232.0 19.10 186.5 15.40 12.9 1.10 1 213.9
1977 800.6 64 117.6 357.5 325.5 206.2 16.50 223.0 17.90 17.5 1.40 1 247.3
1978 873.4 64 114.5 418.3 340.6 203.7 15.00 263.3 19.30 21.4 1.60 1 361.7
1979 1 059.8 70 114.5 571.6 373.7 157.9 10.40 285.8 18.80 19.6 1.30 1 523.1
1980 1 091.0 71 118.5 536.3 436.1 181.7 11.70 252.9 16.40 20.9 1.40 1 546.5
1981 944.7 68 124.2 448.0 372.5 222.2 16.10 198.6 14.40 17.5 1.30 1 383.1
1982 1 114.2 69 136.5 457.1 520.6 202.3 12.50 281.0 17.40 18.6 1.10 1 616.1
1983 1 085.5 65 175.0 392.8 517.6 250.3 14.90 316.5 18.90 22.4 1.30 1 674.6
1984 1 125.8 68 150.5 511.9 463.3 213.3 12.80 294.8 17.70 32.3 1.90 1 666.2
1985 1 260.2 70 178.9 530.9 550.4 211.1 11.80 280.8 15.70 36.2 2.00 1 788.3
1986 1 205.7 69 250.5 467.2 488.0 250.5 14.20 275.3 15.70 26.5 1.50 1 758.1
1987 1 756.5 76 328.4 871.1 557.0 245.5 10.70 247.5 10.80 47.6 2.10 2 297.1
1988 1 644.0 74 293.3 846.9 503.8 253.7 11.30 268.6 12.00 69.1 3.10 2 235.4
1989 1 973.8 75 357.7 968.9 647.2 275.1 10.40 279.3 10.60 115.9 4.40 2 644.0
1990 1 720.0 73 296.6 822.2 601.2 294.4 12.40 299.1 12.60 57.5 2.40 2 371.1
1991 1 856.1 72 293.7 1 072.4 489.9 319.3 12.50 301.0 11.80 84.7 3.30 2 561.1
1992 2 095.3 76 310.2 1 362.4 422.7 297.4 10.70 267.4 9.70 106.9 3.90 2 767.0
1993 1 973.6 73 317.1 1 390.4 266.1 296.6 11.00 310.5 11.60 107.1 4.00 2 687.8
1994 1 959.5 70 318.4 1 322.3 318.8 275.4 9.80 383.6 13.70 184.9 6.60 2 803.4
1995 2 001.8 68 328.0 1 313.0 360.8 304.3 10.40 421.7 14.40 210.6 7.20 2 938.4
1996 2 276.2 72 305.2 1 607.4 363.7 305.6 9.70 381.4 12.10 186.5 5.90 3 149.7
1997 2 487.9 72 269.2 1 909.8 308.9 273.4 7.90 469.4 13.60 225.9 6.50 3 456.6
1998 1 883.5 66 295.4 1 243.6 344.5 319.6 11.20 441.5 15.50 213.0 7.50 2 857.6
1999 2 582.1 72 268.3 1 860.2 453.6 352.5 9.80 453.1 12.60 210.0 5.80 3 597.7
2000 2 560.7 70 304.9 1 773.8 482.0 312.2 8.50 497.4 13.60 284.8 7.80 3 655.2
2001 2 242.6 67 288.9 1 574.5 379.2 317.2 9.50 533.3 15.90 253.7 7.60 3 346.8
Table 1
Total world cephalopod catch in thousand metric tonnes since 1950, by major fisheries categories
1/ Squids are broken down into Loliginids, Ommastrephids and unidentified squids to show each group?s relative importance.
12 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Cephalopodspecies
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
Il
le
x
a
rg
e
n
ti
n
u
s
16
.0
33
.3
79
.7
56
.2
10
6.
1
20
4.
9
27
1.
3
54
6.
3
56
4.
3
55
8.
5
41
0.
1
55
9.
5
60
9.
8
63
8.
5
50
5.
7
52
0.
9
65
6.
5
98
0.
3
69
3.
5
11
45
.0
93
0.
8
74
3.
0
To
d
a
ro
d
e
s
p
a
c
if
ic
u
s
40
5.
4
29
0.
4
27
4.
2
24
5.
9
28
6.
5
21
4.
3
14
1.
5
26
2.
0
22
7.
8
31
9.
8
32
1.
5
40
3.
0
54
5.
2
54
8.
4
50
4.
4
51
3.
4
71
5.
9
60
3.
4
37
8.
6
49
7.
9
57
0.
4
52
8.
5
Do
si
d
ic
u
s
g
ig
a
s
19
.1
9.
8
1.
2
0.
1
0.
4
15
.9
1.
2
0.
3
1.
7
10
.4
14
.9
45
.8
10
9.
4
12
4.
4
19
5.
2
13
6.
3
14
2.
2
16
2.
5
27
.5
13
4.
8
18
2.
4
22
3.
8
N
o
to
to
d
a
ru
s
sl
o
a
n
i
0.
8
48
.8
48
.9
55
.7
90
.1
67
.1
28
.1
32
.7
38
.5
53
.0
29
.8
35
.1
64
.4
45
.1
79
.4
94
.1
53
.7
64
.6
55
.6
31
.4
25
.6
45
.1
Om
m
a
st
re
p
h
e
s
b
a
rt
ra
m
i
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
49
.9
55
.0
36
.1
47
.4
23
.9
Il
le
x
il
le
c
e
b
ro
su
s
88
.7
50
.2
31
.3
12
.0
10
.6
6.
9
6.
7
18
.1
5.
9
17
.7
25
.6
19
.7
23
.9
25
.6
30
.7
18
.6
29
.0
34
.8
26
.6
9.
9
11
.2
5.
7
To
d
a
ro
d
e
s
sa
g
it
ta
tu
s
6.
4
15
.5
21
.8
22
.5
18
.1
21
.8
9.
8
11
.4
8.
4
8.
3
8.
3
7.
5
7.
8
6.
5
5.
8
5.
2
5.
9
5.
6
6.
6
4.
9
4.
9
4.
3
Il
le
x
c
o
in
d
e
ti
i
0.
0
0.
0
0.
1
0.
4
0.
1
0.
1
0.
2
0.
3
0.
2
1.
2
0.
5
0.
6
0.
8
0.
8
0.
7
0.
6
0.
4
0.
4
0.
2
0.
3
0.
4
0.
3
M
a
rt
ia
li
a
h
y
a
d
e
si
0.
0
0.
0
0.
0
0.
0
0.
0
0.
0
8.
4
0.
0
0.
0
0.
2
11
.6
1.
4
1.
0
1.
3
0.
4
24
.0
3.
8
8.
4
0.
1
0.
0
0.
7
0.
0
Ommastrephids
536.3
44
8.0
457.
1
392.8
511
.9
530.9
467.
2
871.
1
846.
9
968.
9
822.2
1
072.4
1
362.4
1
390.
4
1
322.3
1
313.
01
607.4
1
909.8
1
243.
61
860.2
1
773.8
1
574.
5
Lo
li
g
o
sp
p
.
93
.1
99
.6
11
4.
0
13
1.
6
11
3.
9
14
8.
4
18
9.
5
24
5.
4
22
4.
8
23
4.
7
21
8.
3
22
1.
4
21
5.
9
24
4.
3
23
4.
6
21
6.
8
21
6.
7
22
7.
6
21
8.
1
19
9.
8
21
5.
0
21
3.
6
Lo
li
g
o
g
a
h
i
0.
0
0.
0
0.
0
16
.0
13
.0
9.
2
39
.7
68
.6
44
.0
89
.2
57
.1
45
.8
71
.8
44
.3
55
.5
85
.2
68
.5
21
.7
51
.7
42
.5
67
.0
57
.7
Lo
li
g
o
p
e
a
le
i
23
.6
22
.5
20
.8
25
.7
22
.4
17
.0
17
.1
11
.7
19
.1
23
.0
16
.3
19
.6
19
.7
22
.2
22
.5
18
.9
12
.5
16
.2
18
.9
18
.7
16
.9
14
.2
Lo
li
g
o
re
y
n
a
u
d
i
1.
9
2.
1
1.
7
1.
7
1.
2
4.
3
4.
2
2.
7
5.
4
10
.7
5.
0
7.
0
2.
8
6.
3
5.
8
7.
0
7.
5
3.
7
6.
7
7.
2
6.
0
3.
4
Lo
lig
o
species
11
8.5
124
.2
136.
5
175.
0
150.
5
178.
9
250.5
328.4
293.3
357.7
296.6
293.7
310.
2
317.
1
318.
4
328.0
305.
2
269.2
295.4
268.3
304.
9
288.9
N
o
n-id
ent
if
ie
d
squ
id
s
436.
1
372.5
520.6
517.
6
463.
3
550.
4
488.
0
557.0
503.8
647.
2
601.
2
489.
9
422.
7
266.1
318.
8
360.8
363.
7
308.9
344
.5
453.
6
482.
0
379.2
SQUIDS
1
091.
0
944
.7
1
11
4.
21
085.5
1
125.
81
260.
21
205.7
1
756.5
1
644
.0
1
973.8
1
720.0
1
856.
1
2095.3
1
973.6
1
959.5
2001
.8
2276.
22
487.
91
883.5
2582.1
2560.7
22
42.
6
O
ct
o
p
o
d
id
ae
11
3.
2
13
1.
5
13
7.
8
14
6.
1
14
5.
4
14
4.
5
18
3.
3
18
0.
1
19
2.
4
21
0.
8
22
9.
2
23
5.
7
21
8.
0
22
3.
6
20
5.
2
23
5.
0
23
0.
8
21
0.
8
24
5.
4
29
5.
0
25
9.
7
26
1.
8
Oc
to
p
u
s
v
u
lg
a
ri
s
66
.4
88
.7
61
.9
99
.8
65
.1
62
.9
64
.4
62
.5
57
.8
61
.8
62
.2
80
.2
75
.8
69
.4
66
.3
66
.8
72
.9
60
.3
72
.5
56
.1
50
.5
53
.1
E
le
d
o
n
e
sp
p
.
2.
1
2.
0
2.
6
4.
3
2.
7
3.
7
2.
7
2.
9
3.
6
2.
5
3.
0
3.
4
3.
6
3.
5
3.
9
2.
5
1.
9
2.
3
1.
8
1.
4
2.
0
2.
4
OCTOPUSES
181
.7
222.2
202.3
250.3
213.
3
211
.
1
250.5
245.
5
253.7
275.1
294.
4
319.
3
297.4
296.6
275.4
304.
3
305.
6
273.4
319.
6
352.5
312.
2
317.
2
S
ep
ii
d
ae
,
S
ep
io
li
d
ae
24
8.
1
19
4.
7
275.8
31
1.
3
28
8.
2
27
3.
2
26
6.
7
23
6.
4
25
3.
3
26
2.
1
28
3.
1
28
9.
2
25
4.
1
29
7.
9
37
1.
0
41
1.
4
37
0.
0
45
5.
7
42
9.
1
43
8.
3
48
4.
7
51
9.
3
S
e
p
ia
o
ff
ic
in
a
li
s
4.
8
4.
0
5.
2
5.
2
6.
6
7.
6
8.
6
11
.1
15
.3
17
.2
16
.0
11
.8
13
.2
12
.6
12
.5
10
.4
11
.4
13
.8
12
.4
14
.9
12
.7
14
.0
CUTT
LEFISHES
252.9
198.
6
281.
0
316.
5
294.
8
280.
8
275.3
247.
5
268.6
279.3
299.1
301.
0
267.4
310.
5
383.6
421
.7
381.
4
469.
4
44
1.
5
453.
1
497.
4
533.3
N
o
n-id
ent
if
ie
d
ce
p
h
al
o
p
o
d
s
20.9
17.
5
18.
6
22.4
32.3
36.
2
26.5
47.
6
69.1
11
5.9
57.5
84.
7
106.
9
107.
1
184
.9
210.
6
186.
5
225.9
213.
0
210.
0
284.
8
253.7
WORLDT
OTAL
1
546.
51
383.1
1
616.
1
1
674.
61
666.2
1
788.
31
758.1
2297.1
2235.4
264
4.
02
371.
1
2
561.
1
2767.0
2687.8
2803.4
2938.4
3
14
9.8
34
56.7
2857.6
3597.7
3655.2
3
346.
8
Table
2
To
ta
l
wo
rl
d
ce
p
h
al
o
p
o
d
ca
tc
h
in
th
o
u
sa
nd
met
ri
c
to
nnes
si
nce
1
980,
by
majo
r
sp
ec
ie
s
(S
ou
rc
e:
FA
O
,2
00
0)
Cephalopods of the World 13
Area
1980
1981
1982
1983
1984
1985
1986
1987
1988
1989
1990
1991
1992
1993
1994
1995
1996
1997
1998
1999
2000
2001
%
NWPacific
88
2.
3
69
1.
9
76
1.
6
73
8.
8
71
7.
3
70
9.
3
59
6.
0
70
6.
6
69
1.
4
87
7.
9
86
1.
7
81
5.
6
91
3.
8
87
3.
7
93
7.
9
1
00
6.
5
1
16
5.
5
1
15
4.
1
1
03
3.
6
1
11
7.
8
1
22
2.
0
1
13
7.
5
34
.0
SWAtlantic
31
.0
54
.1
20
8.
4
20
5.
2
25
2.
1
35
2.
9
41
0.
5
76
3.
9
68
1.
4
76
4.
3
55
0.
9
69
6.
9
73
4.
3
71
9.
5
58
6.
2
64
5.
2
74
7.
9
10
16
.6
75
0.
0
1
19
2.
7
1
00
1.
0
80
2.
9
24
.0
WCPacific
11
9.
6
13
0.
0
15
3.
6
18
3.
1
17
0.
2
18
2.
6
19
5.
5
20
4.
9
21
5.
7
23
9.
1
25
9.
8
27
0.
0
27
9.
8
31
4.
2
35
9.
3
35
5.
8
34
3.
9
36
0.
0
36
6.
1
36
4.
6
43
5.
3
39
0.
5
11
.7
ECAtlantic
13
8.
0
19
0.
5
17
5.
9
21
1.
6
15
6.
6
15
4.
0
19
4.
3
16
7.
7
16
5.
0
18
4.
5
20
2.
4
23
8.
8
18
8.
4
19
5.
0
17
9.
1
19
1.
2
19
0.
6
15
0.
4
18
3.
7
25
0.
3
21
6.
4
21
2.
9
6.
4
SEPacific
0.
3
1.
2
3.
0
2.
8
3.
4
19
.5
5.
9
4.
8
7.
2
9.
8
20
.6
72
.5
11
7.
5
13
5.
5
19
9.
6
10
9.
2
36
.0
32
.1
11
.4
82
.8
12
7.
6
17
5.
0
5.
2
ECPacific
30
.7
32
.3
19
.8
16
.0
17
.7
24
.7
35
.5
67
.9
76
.6
80
.8
80
.2
64
.6
40
.8
38
.2
58
.8
11
1.
3
20
1.
5
21
3.
1
30
.4
15
0.
6
20
2.
5
16
7.
1
5.
0
WIndianOcean
23
.7
11
.9
11
.8
10
.1
20
.0
27
.3
24
.0
32
.0
42
.0
45
.1
37
.5
70
.5
77
.2
81
.7
10
2.
0
11
0.
6
94
.9
14
8.
6
11
2.
4
12
1.
3
11
9.
0
14
0.
1
4.
2
EIndianOcean
27
.4
22
.6
23
.4
25
.4
34
.3
32
.9
39
.1
53
.6
60
.5
92
.5
51
.8
62
.3
84
.1
73
.3
71
.3
88
.4
10
4.
6
10
9.
4
11
5.
7
11
1.
8
12
4.
1
11
6.
0
3.
5
SWPacific
80
.2
63
.3
71
.1
81
.0
11
6.
8
88
.8
72
.1
73
.6
75
.0
10
7.
1
67
.9
63
.7
11
3.
0
69
.7
11
6.
7
13
9.
6
73
.3
84
.8
71
.0
43
.6
35
.5
57
.3
1.
7
Mediterranean &
BlackSea
51
.7
47
.1
49
.4
55
.3
56
.7
65
.7
68
.4
67
.7
83
.3
76
.9
72
.5
67
.8
71
.7
65
.4
68
.7
62
.5
59
.3
60
.1
54
.3
50
.7
53
.8
53
.8
1.
6
NEAtlantic
29
.2
37
.4
41
.9
48
.5
37
.2
38
.2
25
.4
43
.4
42
.4
49
.7
55
.5
41
.2
49
.9
49
.0
44
.0
54
.0
53
.4
51
.0
49
.9
53
.1
58
.8
44
.5
1.
3
WCAtlantic
8.
4
8.
9
9.
3
12
.3
10
.2
10
.4
13
.3
11
.5
12
.0
18
.2
19
.3
20
.3
21
.5
18
.8
20
.0
20
.7
31
.1
20
.8
24
.1
20
.1
24
.4
23
.2
0.
7
NWAtlantic
11
1.
3
72
.0
52
.1
39
.9
33
.1
23
.9
23
.7
23
.5
22
.0
37
.6
39
.0
36
.3
40
.7
46
.4
52
.0
35
.6
38
.9
48
.7
44
.7
26
.8
26
.4
18
.4
0.
6
SEAtlantic
8.
5
11
.1
10
.1
10
.0
8.
3
12
.1
12
.6
20
.2
11
.4
14
.7
7.
8
9.
9
8.
2
7.
1
6.
5
7.
4
8.
2
4.
1
7.
8
8.
3
7.
5
4.
7
0.
1
NEPacific
4.
3
8.
9
24
.7
34
.7
32
.4
46
.0
41
.7
55
.8
49
.7
45
.7
43
.9
30
.9
26
.0
0.
3
1.
4
0.
3
0.
7
2.
8
2.
3
3.
1
0.
7
3.
0
0.
1
TO
TA
L
1
546.
51
383.1
1
616.
1
1
674.
61
666.
21
788.3
1
758.1
2297.1
2235.4
264
4.
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1
2561
.1
2767.0
2687.8
2803.4
2938.
4
31
49.
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456.
62
857.6
3597.7
36
55.2
334
6.8
(S
ou
rc
e:
FA
O
,2
00
0)
Table
3
W
o
rl
d
ce
p
h
al
o
p
o
d
ca
p
tu
re
p
ro
d
u
ct
io
ni
nt
h
o
u
sa
nd
met
ri
c
to
nnes
,
by
F
A
O
ar
ea
s
d
u
ri
ng
th
e
y
ea
rs
1
980?2001
14 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Country 1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001
Japan 622 235 595 516 772 777 639 683 647 090 599 685 728 759 700 036 446 927 558 527 668 147 566 371
China 69 918 70 167 70 836 122 159 193 561 225 421 175 835 246 690 379 056 417 031 482 802 504 287
Korea,
Republic
of
333 709 411 673 475 625 432 160 381 040 421 492 449 352 481 960 310 631 591 087 426 679 412 513
Argentina 27 778 46 375 77 746 194 653 197 971 200 400 292 851 414 041 291 993 342 934 279 914 229 874
Thailand 135 072 154 402 150 315 153 237 144 436 156 397 173 183 173 648 188 156 174 382 171 990 170 945
Taiwan
Province
of China
223 868 280 878 207 948 219 926 191 257 187 820 171 847 250 811 237 060 297 528 258 907 165 521
Morocco 74 143 92 162 82 368 91 685 83 047 90 134 91 324 63 963 68 553 112 829 147 727 140 829
Viet Nam 23 000 28 000 32 000 33 000 87 000 103 000 92 000 92 500 103 000 110 000 180 000 130 000
India 30 907 55 273 72 682 70 423 95 109 103 739 85 120 117 624 95 834 93 709 96 408 114 681
USA 43 497 63 307 50 975 73 646 97 879 104 119 108 878 101 506 45 240 117 124 143 808 105 125
World
Total 2 371 064 2 561 100 2 766 970 2 687 779 2 803 421 2 938 392 3 149 764 3 456 670 2 857 620 3 597 670 3 655 150 3 346 828
(Source: FAO, 2000)
Table 4
World cephalopod capture (in metric tonnes) ranked by the 10 main producing countries from 1990 to 2001
2001 Exports 2001 Imports
Quantity (Thousand tonnes) Value (Millions US$) Quantity (Thousand tonnes) Value (Millions US$)
Argentina 158.5 Morocco 399.7 Spain 286.2 Japan 812.8
Spain 126.1 Thailand 319.0 Japan 198.7 Spain 567.3
Morocco 111.9 Viet Nam 269.7 China 195.9 Italy 421.9
USA 108.1 Spain 254.7 Italy 186.2 USA 143.6
China 101.4 China 213.1 Korea, Republic of 66.2 China 131.0
Thailand 92.4 Argentina 149.4 USA 62.1 Korea, Republic of 124.2
Korea, Republic of 76.6 India 106.1 Thailand 49.8 Thailand 65.9
Taiwan Province of China 62.5 USA 85.9 Greece 33.5 Portugal 61.0
India 60.1 Korea, Republic of 71.7 Portugal 31.2 Greece 56.2
Viet Nam 59.8 Mauritania 65.5 France 25.8 France 55.4
World Total 1 310.8 World Total 2 506.0 World Total 1 329.4 World Total 2 750.9
(Source: FAO, 2000)
Table 5
World cephalopod export and import quantities and values in 2001 for the 10 leading countries
Cephalopods of the World 15
Fig. 3 Cephalopod capture (tonnes) by main FAO fisheries areas
(Source: FAO, 2000)
Western Atlantic
0
100
200
300
400
500
600
700
800
900
1000
1100
1200
1300
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
South
North
Central
B Eastern Atlantic
0
100
200
300
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Central
North
South
C
Western Pacific
0
100
200
300
400
500
600
700
800
900
1000
1100
1200
1300
1400
1500
1600
1700
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
North
Central
South
D Eastern Pacific
0
100
200
300
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
North
South
Central
E
Indian Ocean
0
100
200
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Eastern
Western
F Mediterranean and Black Sea
0
25
50
75
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
G
0
500 000
1000 000
1500 000
2 000 000
2 500 000
3 000 000
3 500 000
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
A
World Total
16 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 4 Total world cephalopod capture (tonnes) by main species groups or species
(Source: FAO, 2000)
0
500 000
1000 000
1500 000
2 000 000
2 500 000
3 000 000
3 500 000
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Squids (Ommastrephidae)
Squids (Loliginidae)
Squids (unspecif ied)
Octopuses
Cuttlef ishes
Cephalopods unspec.
World cephalopod catchesA
0
500 000
1000 000
1500 000
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Illex argentinus
Todarodes pacificus
Dosidicus gigas
Illex illecebrosus
Other Ommastrephidae
Ommastrephidae
B
0
100 000
200 000
300 000
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Loligo spp
Loligo gahi
Loligo reynaudi
Loligo pealei
Loliginidae
C
0
100 000
200 000
300 000
400 000
500 000
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Sepiidae, Sepiolidae
Sepia officinalis
Cuttlefishes
D
0
100 000
200 000
300 000
1950 1955 1960 1965 1970 1975 1980 1985 1990 1995 2000
Octopodidae
Octopus vulgaris
Eledone spp
Octopuses
E
Numerous fishing techniques and methods to capture
cephalopods have been developed over time. These were
extensively reviewed, for example, by Rathjen (1984, 1991)
and Roper and Rathjen (1991). They include hand
collection, small traps, spears, lures, jigs, lampara nets,
chemical flushing, midwater trawls and otter trawls.
Jigging is the most widely used method, which accounts for
almost half of the world cephalopod catch: all squids, primarily
ommastrephids, but also a few loliginids. This technique is
employed primarily at night, when many species of squids are
attracted to the fishing vessel by lights. Figure 5 shows the
distribution of the world?s light fishery for some of the most
important squid species. Jigs, which feature numerous,
variously-arranged, barbless hooks (Plate IX, 60), are lowered
and retrieved by jigging machines that simulate the constant
swimming behaviour of natural prey, inducing the squids to
attack them. While simple hand-jigging machines are still
used in small-scale, artisanal fisheries, large modern vessels
for industrial fishing activities are equipped with scores of
automated, computer-controlled jigging machines, each
capable of catching several tonnes per night (Plate VIII, 50;
Plate IX, 56). Occasionally, cuttlefishes also are captured by
hand jig.
Trawling is the secondmost productive fishery method to
catch cephalopods (Plate VIII, 51). Formerly, many species
of squids, octopods and cuttlefishes sensu lato were caught
as bycatch in trawl fisheries for finfishes and shrimps, but
some specialized small-scale fisheries trawling for
cephalopods also existed (e.g. for cuttlefishes in the
Mediterranean Sea). The amount of cephalopods taken as
bycatch in bottom trawls for finfish fisheries drove
increasing attention to the resource by the 1980s; this led to
the development of the (principally) midwater trawl fisheries
specifically targeting squids, particularly the South Atlantic/
Subantarctic fishery for Illex argentinus. Trawling is a very
efficient technique to catch benthic and epibenthic species,
but soft-bodied animals like cephalopods are often
damaged by the other species in the catch, particularly in
benthic and epibenthic otter trawls. Even in fisheries in
which squid-specific trawling occurs, the huge catches of
squids per tow often result in crushed and damaged
product. Consequently, trawled squid product generally is
less valuable than jig-caught squids. However, modern
oceanic trawlers can process on board many metric tonnes
of cephalopods per day, which helps insure a high-quality
product. Bottom trawling can be very dangerous for benthic
habitats because of the physical damage it causes to the
seabed and associated fauna and because of its lack of
selectivity. Consequently, less intense exploitation by this
traditional fishing technique and an approach toward
diversification of methods and redistribution of the fisheries
through different areas were encouraged and still are highly
recommended, especially in situations where small-scale
fisheries still exist and new, more efficient methods can be
implemented.
The remaining cephalopod catches come from several
differentmethods (Plate VIII, 47, 49, 52-54; Plate IX, 55, 59).
Cuttlefishes are captured by dredges, seines, jigs and pots
specifically developed for these species and very popular in
some regions, e.g. the Mediterranean Sea. Nearshore, neritic
squids are caught by purse seines, octopuses are taken by
pots, dragnets, hooks and spears, while Nautilus spp. are
caught in deep-set wicker or wire traps.
The utilization of cephalopods for human consumption is
extensive and diverse. Products range from fresh food,
eaten raw as ?sashimi? in Japan and, in recent years,
worldwide, and fresh-cooked, as well as various types of
processed product (dried, canned, frozen, reduced to meal,
etc.). The high protein and low fat content of cephalopods
make them an important and healthy element in the human
Cephalopods of the World 17
Fig. 5 Distribution of the world? s light fisheries for ommastrephids (illustrations based on night-time satellite imagery)
(from Rodhouse et al., 2001)
a) Kuroshio Current Province
( Todarodes pacificus and Ommastrephes bartrami )
b) Southwest Atlantic Province
( Illex argentinus )
18 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
diet. With the increasing demand for food for human
consumption, cephalopod resources probably will receive
even more attention in the future.
Considering the present level of exploitation of the
commercially-fished cephalopod populations, a further
increase in such fishery production is likely to occur first by
expansion of the fisheries into the less-fished regions of the
oceans, e.g. the Southern Ocean, probably the ?last frontier?
in the field of marine fisheries. There, a standing stock of
squid biomass as high as 100 million tonnes was estimated
by scientists, based on an estimate of 30 million tonnes
consumed by vertebrate predators (see Rodhouse et al.,
1994 for details), even though squid captures are rarely
successful. Therefore, a priority for the future research in
the field of Antarctic cephalopod biology will be to assess
the squid biomass there, quantitatively and qualitatively,
with the objective of determining a sustainable fishery
production. However, polar cephalopods probably are
longer living and slower growing than species currently
harvested. Therefore, caution must be exercised in
assumptions and decisions for management of polar
cephalopod fisheries.
In the future, it is likely that attention will be focused on
finding other species and families to replace fish stocks that
become severely reduced by overfishing. Even though
clear evidence reveals the existence of large cephalopod
resources available for exploitation in the open oceans,
based on the estimated consumption by predators (see
Clarke, 1996; Piatkowski et al., 2001 for reviews), many
oceanic squids are distasteful for human consumption as
their tissues have a high ammonium content. Research is
being carried out on how to remove this factor on a
commercial scale, but results will take time and catches will
need to be processed before utilization. A number of
ommastrephid squids that lack ammonium are considered
to be underexploited. These include: Sthenoteuthis
pteropus, Ommastrephes bartrami, Martialia hyadesi,
Todarodes sagittatus, Sthenoteuthis oualaniensis,
Nototodarus philippinensis and Dosidicus gigas, and the
circumpolar, subantarct ic Todarodes filippovae .
Exploitation of these species would provide large tonnages
of high quality cephalopods and would require only minor
development in catching techniques. However it will be
necessary to determine where these species congregate
for feeding and spawning activities. An analysis of biomass,
production and potential catch for the 21 species of
Ommastrephidae is presented in Nigmatullin (2004).
Although a number of other oceanic squid families have
large populations and high quality flesh, they are not
currently exploited on a commercial scale except for a few
seasonal fisheries. These include members of the families
Thysanoteuthidae, Gonatidae and Pholidoteuthidae, for
example. Increased exploitation of these groups, however,
would also require some research and development of
catching techniques. Commercial exploitation of the
cosmopolitan family Histioteuthidae also could be
considered, since at least one large commercial-level catch
has been made in the North Atlantic (see Okutani, personal
communication, in Clarke, 1996). However, the increased
exploitation of these oceanic squid species might have
unpredictable, far-reaching negative effects on the
mesopelagic ecosystem. Therefore, great caution must be
exercised in developing this kind of fishery.
Almost all of our knowledge of the general biology of
cephalopods, in fact, is limited to the shelf-living species as
well as to those ommastrephids that move onto the shelf at
certain seasons. These represent only about 15% of all
cephalopod species (Clarke, 1996). Even so, many gaps
still exist in our knowledge about their life cycles, especially
as far as the relationships among species are concerned
(e.g. prey-predator balances). Some populations of
harvested species have shown sudden, occasionally
catastrophic, declines before adequate biological data
could be gathered and analysed. Squid stocks, for
example, experienced true collapses at least in two
well-known and documented cases. These were the
northwest Pacific Todarodes pacificus fishery failure in the
1970s and the northwest Atlantic Illex illecebrosus fishery
collapse in the 1980s. While the T. pacificus fishery has
recovered, the I. illecebrosus fishery has remained
insignificant. These collapses are thought to have occurred
mainly as a consequence of temporarily unfavourable
envi ronmental condi t ions or actual long-term
environmental changes, probably aggravated by heavy
fishing pressure (Dawe and Warren, 1993).
A significant challenge thus exists to deepen our knowledge
and learn the details of distribution, life history and biology
of exploited species in order to allow rational utilization of
the stocks. The necessity for research as a key factor
towards attaining this goal has been stressed by many
authors (e.g. Lipinski et al., 1998) and it is especially
important in the fields of life-cycle clarification, stock
structure and genetics, role in the food web and interactions
with the environment. The last topic seems of particular
interest wi th in the more general context of
climate/environmental global changes, since the unusual
biological characteristics and short l ife cycles of
cephalopods are strongly linked to immediate, temporal
environmental circumstances. Therefore, cephalopods are
potentially very good ?indicator species? to predict or reflect
changes in environmental conditions, both locally and on a
broader scale (O?Dor and Dawe, 1998; Arkhipkin et al.,
2001; Bendik, 2001; Dawe et al., 2000, 2001; Jereb et al.,
2001; Laptikhowsky and Remelso, 2001; Roberts, 2001).
Perhaps even more significant is the challenge that exists
for future exploitation of new species or populations. The
role of cephalopods in the ecosystem, in fact, is more
complex than it was thought to be only a few decades ago.
Cephalopods can be considered subdominant predators
that tend to increase in biomass when other species,
particularly their predators and competitors for food,
become depleted, as a result of a combination of heavy or
excessive fishing, other human impacts, oceanographic
fluctuations and competition for food (see Caddy, 1983, and
Caddy and Rodhouse, 1998 for a detailed analysis of the
transi t ion f rom f inf ish- targeted f isher ies to
cephalopod-targeted fisheries). A thoroughly studied case
is that of the snapper fishery on the Endeavour Bank
(Balguerias et al., 2000), which was replaced by a
cephalopod fishery between 1960 and 1970. In turn,
cephalopods are major food items in the diets of
innumerable species of fishes, toothed whales (e.g. sperm
whales, beaked whales, dolphins, porpoises), pinnipeds
(seals, sea lions) and seabirds (penguins, petrels,
albatrosses).
Muscular cephalopods derive their energy from
crustaceans, fishes and other cephalopods. At the same
time, they are a very efficient food store for the large,
oceanic predators, by rapidly converting oceanic resources
into high energy food. On the other hand, neutrally buoyant
ammoniacal squids, which probably greatly outnumber the
muscular squids in biomass, also provide food to many of
the same predators, but not over the continental shelf and
with consistently lower energy per unit body mass. We
know virtually nothing about the details of feeding, growth,
life cycles, periodicities, distribution and spawning in
ammoniacal species.
In spite of our relatively poor knowledge, it is now clear that
cephalopods are a dominant component within marine
ecosystems and that their abundance ultimately may
influence the abundance of their predators and prey
populations. Recent studies of the effects of consumption of
important pelagic squids and fishes by predatory fishes on
the northeastern shelf of the United States (Overholtz et al.,
2000), concluded that changes in predator abundance may
have important implications for the long-term fishery yields
of pelagic species.
What seems consistent with our present knowledge is that
removal of cephalopods through fisheries would have a
consistent impact on the environment: populations of small
midwater fishes would increase, while top predators like
cetaceans, seabirds, seals and even some fish populations
would decrease.
Taking into consideration these factors, increasing effort
should be focused on improving our scientific knowledge of
this group. Cephalopod catches need increased
moni tor ing, especia l ly in those areas of major
envi ronmental f luctuat ions and where f isher ies
management is complicated by multiple countries fishing
the same resource. Cooperation, collaboration and
commitment are required to better understand these
important and fascinating animals.
Cephalopods of the World 19
20 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
1. 4 Illustrated Glossary of Technical Terms and Measurements
a) Schematic illustration of a squid
a r m l e n g t h
h e a d
f i n l e n g t h
t a i l
m a n t l e l e n g t h
b) Schematic illustration of a cuttlefish
f i n l e n g t h
m a n t l e l e n g t h
t o t a l l e n g t h
Fig. 6
a r m l e n g t h
h e c t o c o t y l u s l e n g t h i n m a l e s
h e a d
m a n t l e l e n g t h
t o t a l l e n g t h
c) Schematic illustration of an octopus
Abdominal septum ? Median septum that traverses the
mantle cavity parallel to the body axis. It extends from the
visceral mass to the ventral mantle wall. The ventral mantle
artery runs along the dorsal edge of this septum.
Aboral ? Away from or opposite to the mouth.
Abyssal ? The great depths of the ocean: from 2 000 to
6 000 m.
Accessory nidamental glands ? Glands of unknown
function; consist of tubules containing symbiotic bacteria.
Found in all decapodiformes except oegopsid squids.
Adult ? A female that has mature eggs (these frequently are
stored in the oviducts), or a male that has produced
spermatophores (these are stored in Needham?s sac).
Afferent blood vessel ? Artery vessel carrying blood toward
an organ.
Afferent nerve ? Nerve carrying impulses toward the brain
or specific ganglia.
Anal flaps ? A pair of fleshy papillae involved in directing
releases of ink, one flap situated at each side of the anus
(Fig. 7).
Anal pads ? Ovoid pads of unknown function, apparently
glandular, one located on each side of the anus in some
squids (e.g. bobtail squids).
Anterior ? Toward the head-end or toward the arm-tips of
cephalopods.
Anterior salivary glands ? Glands on or in the buccal mass
that aid in preliminary digestion.
Antitragus ? Knob that projects inward from the posterior
surface of the central depression in the funnel-locking
cartilage of some squids (Fig. 8).
Anus ? Terminal opening of the digestive tract in the
anterior mantle cavity, sometimes extending to inside the
funnel, through which digestive waste products, as well as
ink, are expelled.
Apomorphic ? Derived from a more ancestral condition.
Loosely considered the ?advanced? condition.
Arm ? One of the circumoral appendages of cephalopods.
Arms are designated by the numbers I to IV, starting with I
as the dorsal (or upper) pair. In decapodiformes each
appendage of the fourth original pair is modified to form a
tentacle. The second pair of arms is modified in vampires
into a long filamentous structure. It has been lost in cirrate
and incirrate octopods.
Arm formula ? Comparative length of the 4 pairs of arms
expressed numerically in decreasing order: the largest arm
is indicated first and the shortest last, e.g. IV>III>II>I. If
IV>III=II>I, then arm IV is the longest, followed by arm III
which is the same size as arm II and both are longer than
arm I. In octopods, the non-hectocotylized arm III is used in
this formula.
Armature ? The grappling structures of the arms and
tentacular clubs, including suckers and/or hooks.
Bactritida ? A fossil taxon that existed about 428 to 216
million years ago and is thought to be the ancestral stock of
the Ammonoidea and Coleoidea.
Bathypelagic ? The deep midwater region of the ocean.
Beak ? One of the 2 chitinous jaws of cephalopods, bound
in powerful muscles. The dorsal beak is referred to as the
?upper? beak and it inserts within the ?lower? (ventral) beak to
tear tissue with a scissors-like cutting action.
Belemnoidea ? A fossil group of cephalopods that is
thought to be the sister group of the Coleoidea. Belemnoids
are distinguished by the presence of hook-like structures on
the arms rather than suckers.
Benthopelagic ? A free-swimming animal that lives just
above the ocean floor but rarely rests on the ocean floor.
Bilateral symmetry ? The symmetry exhibited by an
organism or an organ if only one plane can divide the animal
structure into 2 halves that are mirror images of each other.
Bioluminescence ? The production of light by living
organisms, sometimes called ?living light?. The light is
produced through a chemical reaction that generally takes
place in complex organs called photophores or light organs.
Brachial ? Pertaining to the arms.
Brachial crown ? The combination of arms and tentacles
that surround the mouth.
Cephalopods of the World 21
Fig. 7 Terminal portion of the digestive tract
ink sac
anal flaps
Fig. 8 Funnel-locking cartilage
antitragus
tragus
Brachial lobe (of the brain) ? The anteriormost part of the
brain located ventral to the oesophagus. The large axial
nerve cords that run down the centres of the arms connect
to this lobe. The proper name is ?anterior suboesophageal
mass?.
Brachial photophore ? Photophore located on the arms.
Brachial pillar ? A narrow, elongate anterior region on the
paralarval or juvenile head of some families, between the
eyes and the base of the brachial crown; especially well
developed in young cranchiid squids.
Brain ? Medial portion of the central nervous system that
includes the suboesophageal and supraoesophageal
masses but generally does not include the large optic lobes.
Branchial ? Pertaining to the gills.
Branchial canal ? A large opening at the base of each gill
lamella and between the primary afferent and efferent blood
vessels of the gill. A branchial canal is absent in nautiluses,
cuttlefishes, bobtail squids, bottletail squids, pygmy squids,
ram?s horn squids, and cirrate octopods.
Branchial gland ? Elongate or spheroidal gland adjacent
and parallel to the gill attachment to the mantle wall.
B rooding ? Incubation of eggs by the female. A
characteristic feature of incirrate octopods, but also found
in some squids (e.g. Gonatidae).
Buccal ? Pertaining to the mouth.
Buccal connective ? Thin muscular band that attaches the
buccal support of the buccal membrane to the base of the
adjacent arm. The position of attachment of the connective
on the fourth arms was recognized in the early twentieth
century as an important character for phylogenetic
relationships among decapodiformes (Fig. 9).
Buccal crown ? Umbrella-like structure that surrounds the
mouth and in turn is enveloped by the brachial crown. It
consists of buccal supports and the buccal membrane. The
buccal crown is present in most decapodiformes but
absent from all octopodiformes.
Buccal lappet ? A small, subtriangular flap at the tip of each
buccal support of the buccal membrane; thought to be
homologous with the inner ring of tentacles that surrounds
the mouth of nautiluses. May bear suckers (Fig. 9).
Buccal mass ? Muscular bulb at the anteriormost part of
the digestive system that consists of the mouth, beaks,
radula, muscles and pairs of salivary glands.
Buccal membrane ? The muscular membrane that
encircles the mouth like an umbrella (Fig. 9). It connects to
the buccal supports to form the buccal crown. The
pigmentation of the buccal membrane often differs from that
of the adjacent oral surfaces of the arms.
Buccal membrane connectives ? See buccal connective
(Fig. 9).
Buccal suckers ? Small suckers on the buccal
lappets/membrane of some species (Fig. 9).
Buccal support ? Muscular rod fused to buccal membrane
as supporting rib (Fig. 9); 6 to 8 in number.
Buoyancy (neutral, positive, negative) ? The tendency to
float in seawater. A neutrally buoyant object does not rise or
sink but maintains its position in the water; a positively
buoyant object will rise and a negatively buoyant object will
sink.
Bursa copulatrix ? The wrinkled area present in most
bobtail squids near the female genital opening for the
attachment of the spermatophores; it is more differentiated
in members of the subfamily Sepiolinae and forms a
characteristic structure called ?bursa?, which is used during
copulation (Fig. 10). It differs conspicuously among species
and can be used as a key character to identify Sepiola and
Sepietta species females.
Caecal sac ? The sac-like, thin-walled posterior portion of
the caecum in the digestive tract that lacks the internal,
ciliated leaflets characteristic of the anterior portion of the
caecum.
Caecum ? Region of the digestive tract of all cephalopods
between the stomach and intestine. It is the primary site of
food absorption.
22 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 9 Buccal anatomy of decapods
tentacle
buccal
suckers
buccal
lappet
buccal
membrane
buccal connective
(dorsally attached)
a r m I I
a r m I ( d o r s a l )
a r m I V ( v e n t r a l )
a r m I I I
buccal
connective
(ventrally
attached)
buccal support
(beneath membrane)
Fig. 10 Mantle cavity of female Sepiola rondeletii
showing the bursa copulatrix
bursa copulatrix
Calcified ? Chalky, calcareous material of calcium salts
(calcium carbonate), formed by deposition.
Calamus ? The conical papilla or projection at the base of
the ligula on the hectocotylus of octopods, at the distal
terminus of the sperm groove, distal to the last sucker
(Fig. 11) (see Ligula).
Cambrian period ? Oldest period of the modern geological
timescale.
Carpal cluster (= Carpal pad) ? An usually distinct group of
suckers and knobs on the carpus of the tentacular club
(Fig. 12).
Carpal knobs ? Small, rounded, hemispherical, muscular
protuberances on the carpus to which carpal suckers from
the opposite club adhere during the locking of the clubs
(Fig. 12).
Carpal suckers ? Small suckers on the carpus of the club
that adhere to the carpal knobs on the opposite carpus
during the locking of the clubs (Fig. 12).
Carpus ? The proximal zone of small suckers and knobs on
the base of the tentacular club in some families (Fig. 12).
Cartilaginous structures or ?scales? ? Cartilage-like
structures in the skin of certain squids; may be overlapping
and scale-like, or multifaceted platelets, knobs or papillae
(Fig. 13).
Cement body ? Structure in the spermatophore that allows
adhesion of the discharged spermatophore to a female
(Fig. 38).
Cephalic cartilage ? Cartilage-like tissue that envelops the
poster ior part of the brain of cephalopods and
encompasses the statocysts. Anteriorly the cartilage thins
and entwines with muscular tissue, which makes a
well-defined limit difficult to distinguish. The cartilage has a
large central foramen through which the oesophagus
passes and minor foramina for nerves and blood vessels.
Cephalic vein ? Large vein that drains blood from the head
region; it lies along the ventral surface of the visceral sac,
beside or dorsal to the intestine. The cephalic vein
terminates by dividing into the two venae cavae, each of
which passes through the ?kidney? (nephridium), the
branchial heart and into the gill.
Cephalopoda ? The class wi th in the Mol lusca
characterized by bilateral symmetry, internal ?shell? or
absence of shell (except nautiluses), anterior head
appendages and funnel, posterior mantle, mantle cavity
with organs, and shell and fins when present.
Character state ? A particular condition of a taxonomic
character. For example, the character ?sucker? may include
the two states: sucker with a horny ring or sucker without a
horny ring.
Chemotactile ? Refers to chemical and touch sensitivity.
Chitin(ous) ? A horny polysaccharide substance
(fingernail-like) that forms the sucker rings, hooks and
beaks.
Chorion ? A tough secreted membrane that encapsules the
egg.
Chromatophores ? Pigment-filled muscular sacs in the
skin under individual nervous control that collectively
provide the background colour, colour patterns and colour
dynamics (play) of cephalopods.
Circumoral appendages ? The eight arms of
decapodiformes and octopodiformes, plus the two
tentacles of decapodiformes and the very numerous arms
of nautiluses. All arise from the head and encircle the mouth
(Fig. 9).
Cephalopods of the World 23
Fig. 11 Distal tip of hectocotylus of cirrate and
incirrate octopods
ligula calamus sucker
sperm grooveligula length
Fig. 12 Tentacular club of squid
keel
hooks
keel
carpal knob
carpal sucker
carpus
Fig. 13 Two types of cartilagineous structures or
? s c a l e s ?
Cirri (singular cirrus): 1) Arm: elongate, fleshy, finger-like
papillae along the lateral edges of the oral surface of the
arms, especially in cirrate octopods (Fig. 14).
2) Body : fleshy protuberances of the skin that can be
erected as papillae, usually over the eyes (Fig. 15).
Club-fixing apparatus ? Arrangement of suckers and
matching knobs on the carpal region of the tentacular club
that permits the two clubs to be locked together (Fig. 12).
Coelom ? An internal body cavity of mesodermal origin that
is lined by an epithelium. Cephalopods have two coeloms,
the viscero-pericardial coelom and the nephridial coelom.
Collar ? Muscular, flange-like structure that extends from
the nuchal cartilage to the funnel; it forms a one-way valve
that allows water to enter the mantle cavity but closes as the
mantle contracts, thereby forcing exhalant water out
through the funnel.
Cone, conus ? The spoon-like, cup-like, spiked or simple
conical posterior terminus of the gladius or cuttlebone;
homologous to the phragmacone of fossil squids (Fig. 16).
Conus fields ? The sides of the conus that continue
anteriorly along the vanes of the gladius.
Cornea ? Smooth, thin, turgid, transparent skin without
muscles that covers the eyes to protect the eye lenses of
incirrate octopods and some decapods (myopsids and
sepioids) (Fig. 17).
24 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 14 Cirri on arms of cirrate octopods
Fig. 15 Cirri dorsal to eyes of incirrate octopods
cirri
Fig. 16 Gladii of some squids
rachis
vane
conus, rostrum
or spine
Fig. 17 A composite diagram illustrating
basic squid features
I I I I I I
I V
sheathed hooks
carpus
(fixing
apparatus)
tentacle
manus
dactylus
myopsid
eye
mantle
mantle-locking
cartilage
fin
(posteriorly
concave)
photophores
funnel-locking
cartilage
funnel-mantle fusion
funnel
oegopsid eye
(orbital pore)
funnel groove
buccal membrane
keel
suckers
fin lobe
ventral view
tail
Cephalopods of the World 25
Counter illumination ? The production of bioluminescent
light by an animal to conceal its silhouette against a lighted
background. The process can allow an animal to become
virtually invisible under dim directional light.
Cretaceous ? The last period of the Mesozoic Era.
Crop ? Expansion (i.e. a broadening or a side pocket) of
the oesophagus for storing ingested food, prior to entering
stomach. Present in nautiluses and most octopods.
Cusp ? A point or projection on a tooth of the radula or on a
cartilagenous tubercule in the skin.
Cuttlebone (= sepion) ? The calcareous (chalky) oblong,
supporting protective and buoyancy shield in the dorsal
part of the mantle of cuttlefishes (Fig. 18).
Dactylus ? The distal, terminal section of the tentacular
club, often characterized by suckers of reduced size
(Fig. 17).
Decapodiformes ? Higher-level taxon that includes all
10-limbed cephalopods: cuttlefishes, bobtail squids,
bottletail squids, pygmy squids, ram?s horn squids,
myopsid squids and oegopsid squids (Fig. 2 Living
Cephalopods). Because of the long history of referring to
these cephalopods by the common name ? decapods?, the
latter is maintained as the common name for the
Decapodiformes.
Decapods ? Common name for the Decapodiformes.
Demersal ? Organisms that live close to the ocean floor.
Diel vertical migration ? Vertical animal migration during
twilight periods. Many mesopelagic animals migrate to
shallow depths at sunset, where they spend the night
feeding. Then they descend at sunrise from near-surface
waters to spend the day hiding at greater, darker depths.
Some animals migrate vertically over 1 000 m, others
migrate less than 100 m.
Digestive gland ? Primary organ in cephalopods that
secretes digestive enzymes. It is also important in
absorption and excretion (Fig. 19).
Digestive gland duct appendages ? Outpockets of the
ducts leading from the digestive gland that are covered with
glandular epithelium (Fig. 19).
Distal ? Away from the central region of the body or point of
origin; toward the peripheral parts (opposite of proximal).
Doratopsis ? The peculiar paralarval stage that is
characteristic of all members of the oegopsid squid family
Chiroteuthidae.
Fig. 18 Cuttlebone
a) ventral view b) lateral view
inner cone
striae
outer
cone
rostrum
or spine
Fig. 19 Digestive system
(after Bidder, 1966)
a) squid b) cuttlefish
c) octopus
caecum
digestive
gland
stomach
intestine
digestive
gland duct
appendages
caecum
digestive
gland
intestine
stomach
digestive gland
digestive gland
duct appendages
intestinecaecum
digestive
gland duct
stomach
crop
esophagus
Dorsal ? The uppermost or back surface of a cephalopod,
opposite the ventral surface where the funnel is located
(Fig. 20).
Efferent vein ? Vein that carries blood away from the heart
or an organ.
Efferent nerve ? Nerve carrying impulses away from the
brain or specific ganglia.
Egg mass ? A large number of eggs encapsulated in a
gelatinous matrix or a large number of such structures that
are attached together. The pelagic egg mass of an oceanic
squid can be a large, fragile, gelatinous ball that carries
many thousands of eggs. In contrast, the egg mass of a
neritic squid (loliginid) can be composed of hundreds of
very tough, encapsulated eggs in strings, attached together
at their bases and to the substrate.
Ejaculatory apparatus ? Portion of the spermatophore
involved in the vigorous extrusion of the sperm mass
(Fig. 38).
Epipelagic zone ? The uppermost pelagic zone of the
ocean.
Epithelial pigmentation ? The pigmentation contained in
epithelial cells that are unable to change their shape in the
absence of muscles and nerves. Colour in most
cephalopods, however, is created by pigment granules that
are contained in specialized organs, the chromatophores,
that can change shape rapidly by muscular action under
nervous control (see Chromatophores).
Exploitation rate (E) ? When fishing mortality (F) and
natural mortality (M) operate concurrently, the exploitation
rate represents the fraction of dead animals due to the
fishery (i.e. caught by the fishery), which is, F/Z where Z
denotes the total (i.e. M+F) mortality rate.
Eye (position and size ) ? Eyes are the primary sensory
organs of cephalopods; they usually are large and located
one on each side of the head. However, some species have
small eyes, eyes on stalks or telescopic eyes.
Eyelid sinus (= optic sinus = orbital sinus) ? Indentation,
often complex, of the anterior margin of the eyelid (Fig. 28).
Family ? The taxon above the genus level, comprised of the
most closely related genera.
Fin(s) ? The pair of muscular flaps that arise along the
dorsolateral surface of the mantle of decapodiformes,
vampires and cirrate octopods; used for locomotion,
steering and stabilization (Fig. 17).
Fin angle ? The angle between the longitudinal axis of the
mantle and the posterior border of one fin (Fig. 21).
Fin attachment ? A fin attaches to the shell, the mantle, the
opposite fin or some combination of these.
Fin cartilage ? Cartilage associated with the fins of all
fin-bearing cephalopods.
Fin length ? Length from anterior lobe, or anteriormost
attachment of lobe, to posteriormost attachment of fin to
mantle or tail. Extremely long, spike-like tails usually do not
include fin tissue.
Fin lobe ? The portion of a fin that extends anteriorly from
the fin?s anterior point of attachment, or posteriorly from the
fin?s posterior point of attachment of the fin, to the mantle
(Fig. 17). This often is called the ?free? lobe.
Fin position ? Fins are located anterior to the termination of
the muscular mantle (subterminal position) or mostly
posterior to it (terminal position) or in an area of overlap
between the two.
Fin shape ? Fins are classified, somewhat arbitrarily, by
their shape as sagittate, rhomboid, circular/elliptical,
lanceolate, ear-shaped, ribbed, lobate or skirt-like.
Fixing apparatus ? The mechanism of suckers and knobs
on the carpal region of the tentacular club that permits the
two clubs to be locked together during capture of prey
(Figs 12 and 17) (see Carpus).
Foot ? See Molluscan foot .
26 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 20 Schematic lateral view of octopus features
eye ocellus
funnel
web
suckers
hectocotylus
ligula
apertureouter gill
lamellae
(internal)
dorsal
ventral
I V
I I I
I I
I
armsmantle
Fig. 21 Fin angle on squid
fin angle
Foveola ? Transverse, membranous fold of skin that forms
a pocket in the anterior end of the funnel groove of some
oegopsid squids (Fig. 22) (see Side pockets ).
Funnel ? The ventral, subconical tube through which water
is expelled from the mantle cavity during locomotion and
respiration (reproductive and waste products and the ink
also pass through the funnel) (Figs 17 and 20). Archaic
term: siphon.
Funnel-adductor muscles ? Muscles that support the
lateral attachment of the funnel to the head.
Funnel groove ? The depression in the posteroventral
surface of the head in which lies the anterior portion of the
funnel (Fig. 17).
Funnel-locking cartilage ? The cartilaginous groove, pit,
pocket or depression on each ventrolateral side of the
posterior part of the funnel that joins with the mantle
component to lock the funnel and mantle together during
locomotion and respiration, so that water is expelled only
through the funnel and not around the mantle opening
(Figs 17 and 23) (see Mantle-locking cartilage ).
Funnel organ ? The glandular structure fused to the
internal surface of the funnel, generally a single W-shaped
form in octopodiformes and a dorsal inverted V-shaped
component with opposed ventral oblong components in
decapodiformes (Fig. 24).
Funnel-retractor muscles ? Large muscles that attach to
the corners of the funnel and run posteriorly to attach to the
sides of the shell sac (generally near the base of the gills)
or, in some species, insert on the interior mantle wall.
Funnel valve ? The semi lunar muscular flap in the dorsal
inner surface near the distal opening of the funnel in some
species (Fig. 24).
Genus ? The taxon below the family level and above the
species level.
Gill ? Primary organ for the exchange of respiratory gases
with seawater (Fig. 30).
Gill lamellae ? The leaf-like convoluted individual
components of the gill through which gas exchange occurs
(Figs 25 and 30).
Gladius (= pen) ? The feather or rod-shaped chitinous
supporting structure in the dorsal midline of squids ; the
homologue of the shell of ancestral forms (Fig. 16).
Gladius length (GL) ? Sometimes used as a measurement
of the body (= mantle) length when direct measurement of
the mantle is unreliable (usually due to deformation).
Gonoduct(s) ? Tubular structure(s) of the reproductive
system which serves to transport reroductive products from
the gonad(s) to the exterior (see Oviducts ).
Hatchling ? Young cephalopod newly hatched from the egg.
Head length (HL) ? A standard measurement within
species growth stages and for species comparisons;
measured from posterior limit to V-notch at base of Arms I
(Fig. 6).
Head-mantle fusion ? Zone of fusion of head and mantle; it
varies among groups/families; of systematic and biological
significance.
Cephalopods of the World 27
Fig. 22 Funnel groove
a ) Toda r o d i n a e b ) O m m a s t r e p h i n a e
foveola
side pockets
Fig. 23 Funnel-locking cartilage; examples of shapes
and structures
Fig. 24 Funnel organ and funnel valve components on
inner surface of funnel of squids
funnel valve
Fig. 25 Gill lamella, a single element from
a complex structure
Hectocotylus ? One (or more) modified arm in male
cephalopods used to transfer spermatophores to the
female; modifications may involve suckers, sucker stalks,
protective membranes, trabeculae (Figs 11 and 26) (see
Calamus, Ligula).
Hood ? Leathery cap or trapdoor of nautiluses that can be
retracted to protect the animal within the shell.
Holotype ? The single specimen designated by the original
author to represent the new species name. It is an
international standard of reference that provides objectivity
and stability for the species name.
Hooks ? Chitinous, claw-like structures ontogenetically
derived from the suckers on the arms and/or clubs of some
oegopsid squids (Fig. 12).
Horizontal arm septa ? Septa extending the length of the
arms (i.e. parallel to the arm axis), that roughly divide the
arms into oral and aboral regions. This feature is a
characteristic of the arms of cirrate octopods and the
incirrate octopods of the family Bolitaenidae . The
functional significance is unknown.
Horny rings of suckers ? Suckers of decapods have two
types of hard, horny rings. One, the inner ring, lies around
the inner walls of the acetabulum (cup) and often bears
teeth. The other, the outer ring, is composed of numerous
minute platelets and lies on the surface of the infundibulum
(outer rim) (Fig. 40).
Inferior frontal lobe system ? A system of lobes in octopod
brains: the paired posterior buccal, lateral inferior frontal
and subfrontal lobes, and the single median inferior frontal
lobe. They form a functional unit concerned with the
chemotactile information from the arms and its use.
Ink sac ? The structure that manufactures and stores the
ink of cephalopods; it lies parallel with the intestine and
empties via a duct into the rectum (Figs 7 and 30).
Intestine ? Distal region of the alimentary canal between
the stomach/caecum complex and the anus (Fig. 19).
Juvenile ? Life history stage between the hatchling and the
nearly-mature subadult stages.
Keel ? (1) A flattened, muscular extension along the aboral
surface of some arms to render them more hydrodynamic
(Fig. 17); (2) 1 or 2 expanded muscular membranes along
the tentacular club of some groups (Fig. 12); (3) the lateral
ridge of skin around the lateral margin in incirrate octopods
(Fig. 27).
Lateral ? Pertaining to the side(s) of an organism or
structure, away from the centre or midline.
Lateral funnel-adductor muscles ? See Funnel-adductor
muscles.
Lateral membranes of arms IV ? See Tentacular sheath.
Lateral-line analogue ? Sensory structure analogous to the
lateral line of fishes. The lateral-line analogue, which
senses vibrations transmitted by seawater, is located along
a series of lines on the dorsal surface of the head, with
some sensory cells extending onto the bases of the arms.
Lateral ridge ? A narrow, horizontal, muscular structure
along the lateral side of the mantle of incirrate octopods.
Also referred to as a keel (Fig. 27).
Length at 50% maturity ? Mantle length at which 50% of
specimens examined in a representative sample is sexually
mature, according to the maturity scale and the statistical
model used.
Lens (in photophores) ? Structure in a photophore that can
focus or disperse bioluminescent light.
Light guides ? Structures in photophores that specifically
direct light via internal reflection.
Light organ (= photophore) ? A simple or complex
structure that produces bioluminescence (cool light) by
intrinsic (self generated) or extrinsic (bacterial) means
(Figs 17 and 28).
Ligula ? The spatulate to spoon-shaped, terminal structure
of the hectocotylus of many incirrate octopods, that
contains the calamus basally (proximally) and usually a
series of transverse ridges and grooves on the oral surface
(Fig. 11) (see Calamus, Hectocotylus). Spermatophores
28 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 26 Hectocotylized arm ( Illex oxygonius )
only enlarged
trabeculae, no
suckers
suckers
trabeculae
(transverse fleshy supports)
modified portion
starts here
Fig. 27 Incirrate octopod
keel
Fig. 28 Light organs (photophores) on ventral
surface of squid eyeball; eyelid or orbital sinus
eye
photophores
eyelid
sinus
Cephalopods of the World 29
transferred along the sperm groove of the hectocotylized
arm are presumably gripped by the ligula. Details of the
function of the ligula are unknown.
Lips ? Two concentric, muscular, glandular rings of skin that
surround the mouth and beaks.
Living chamber ? Largest and most recently formed
chamber of nautilus shell in which the animal resides (also
refers to numerous shells of fossil cephalopods).
Macrotritopus ? A complex of paralarval forms in the
incirrate octopod family Octopodidae, characterized by the
third pair of arms being extremely elongate.
Mantle ? The fleshy (muscular) tubular or sac-like body of
cephalopods; provides propulsion through jet-like
expulsion of water; contains the viscera (Figs 17 and 20).
Mantle cavity ? Space enclosed by the mantle. In
cephalopods the mantle cavity contains the visceral sac,
gills, anus, openings of the gonoducts, nephridial pores and
various muscles and septa (Fig. 30).
Mantle length (ML) ? The standard measure of length in
coleoid cephalopods. In decapods ML is measured along
the dorsal midline from the mantle margin to the posterior
tip of the body (Fig. 6). In octopods ML is measured from a
line joining the mid-point of the eyes (rather than the
anterior mantle margin, since the latter is obscured by the
head/mantle fusion) to the posteriormost area of the mantle
(Fig. 6).
Mantle-locking cartilage ? The cartilaginous
ridge, knob or swelling on each side of the
ventrolateral, internal surface of mantle that locks
into the funnel component of the locking apparatus
during locomotion (Figs 17 and 23) (see
Funnel-locking cartilage ).
Manus ? Central or ?hand? portion of club between
the dactylus distally and the carpus proximally
(Fig. 17).
Mature ? In cephalopods this term refers to sexual
maturity which is determined for females by the
presence of ova (mature eggs) free in the coelom
or oviducts (Fig. 30), and for males by the
presence of spermatophores in Needham?s sac
(Fig. 29) (see Adult).
Medial(n) ? Pertaining to a structure located
toward, on, or along the dorsal or ventral midline.
Mesopelagic zone ? The middle-depth zone of the
pelagic realm of the ocean.
Mollusca ? One of the major invertebrate phyla.
Some of the common molluscs are snails and clams. The
Cephalopoda is a class within the Mollusca.
Molluscan foot ? A major structure in molluscan
morphology. In gastropods the foot is the muscular sole that
the animal crawls with. In cephalopods the funnel, and
possibly the arms and tentacles are derived from the
molluscan foot. The evolutionary origin of the latter is still
uncertain. They may represent outgrowths of the head
(favoured by anatomical evidence of the nerve
connections) or modifications of the molluscan foot that
have migrated around the mouth ( favoured by
embryological evidence, the migration of arm primordia).
Monophyletic group ? A natural group (taxon) that shares
a common ancestor.
Myopsida ? A high-level taxon (order) within the
Decapodiformes. In recent classification, the Myopsida
(including the family Loliginidae) have been considered the
sister group of the Oegopsida and the two groups together
compose the Teuthoidea (squids). In contrast, at the
beginning of the twentieth century, the Myopsida included
the Loliginidae and the Sepioidea (cuttlefish and relatives)
and the presence of a cornea covering the eye lens defined
the group. At present the phylogenetic relationships within
the Decapods are unresolved.
Neck ? The region that separates the posterior end of the
cephalic cartilage and head musculature. Only those
cephalopods with elongate heads (e.g. the oegopsid squid
family Chiroteuthidae) have distinct necks.
Needham?s sac (= spermatophore/spermatophoric sac) ?
The elongate, membranous organ of males where
completed, functional spermatophores are stored. It opens
into the mantle cavity or directly into the water through the
penis (Fig. 29).
Nephridial coelom ? The cavity of the renal (kidney) sac. It
connects with the exterior via the renal pore and with the
viscero-pericardial coelom via a pair of slender ducts from
the latter.
Nephridial papillae ? Small raised openings to the renal
cavities.
Neritic ? The region of the ocean that overlies the
continental shelf.
Fig. 29 Male squid reproductive apparatus
Needham?s or
spermatophoric sac
spermatophoric
organ
testis
sperm
duct
spermato-
phoric
duct
penis
ampulla
penis
spermatophoric
complex
(after Grieb, 1976 )
Nidamental glands ? Large glandular structures in females
of most decapods and nautiluses that lie in and open
directly into the mantle cavity. The glands are composed of
numerous lamellae that are involved in secretion of egg
cases or the jelly of egg masses (Fig. 30).
Nominal species ? A species that has been formally
described and is based on a morphological type. It is an
available name but not necessarily a valid species.
Nuchal cartilage ? See Nuchal-locking apparatus .
Nuchal crest ? Prominent transverse ridge in most
decapods that extends across the dorsal head and down
the lateral head surfaces at its posterior end (Fig. 31).
Nuchal folds ? Fixed folds or pleats of the head integument
that adjoin the nuchal crest posteriorly and are
perpendicular to it. The function of the folds is uncertain
(Fig. 31).
30 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
nidamental
glands
ovary
gill
ink sac
intestine
oviducts,
paired
no accessory
nidamental
glands
accessory
nidamental
glands
present
oviduct,
single
c) Ommastrephidaeb) Loliginidae
accessory
nidamental
gland
nidamental gland
a) Sepiidae
intestine, anus
oviduct,
single
gill
nephridial pores
ovary
Fig. 30 Internal organs
[a, b and c - Decapodiformes; d, e and f - Octopodiformes]
spermatheca oviduct
central
cavity
spermato-
phoric
gland
ampulla
vas
deferens
seminal
vesicle
prostate
diverticulum penis
penisbag-like
portion
glandular
portion
spermato-
phoric
sac
testis
gill
ovary
branchial
heart
anus oviduct and
oviducal
gland
d) female Octopus e) oviducal gland
(after Froesch and Marthy, 1975)
f) male reproductive tract
(after Mann, 1984)
Nuchal-locking apparatus ? An oblong, cartilaginous-
locking structure in decapods located mid-dorsally just
posterior to the head. It is composed of the nuchal cartilage,
which also forms an attachment site for collar and head
retractor muscles, and an interlocking, complementary
cartilage on the mantle that underlies the gladius. The
apparatus keeps the head and mantle aligned dorsally
during mantle contractions (Fig. 31).
Nuchal membrane (= occipital membrane) ? A thin
membrane that connects the main nuchal folds at their
posterior ends (Fig. 31).
Nuchal organ ? Smal l sensory organ with
photoreceptor-like sensory cells that is located in the
nuchal region of apparently all coleoid cephalopods.
Nuchal region ? The dorsolateral area around posterior
part of the head and the area immediately posterior to it,
normally covered by the anterior mantle wall.
Occipital crest ? See Nuchal crest.
Occipital folds ? See Nuchal folds.
Occipital membrane ? See Nuchal membrane.
Ocellus ? A pigmented spot or patch that usually consists of
a central locus of concentrated chromatophores with one or
more outer concentric rings of chromatophores. Ocelli
occur on some octopuses, and their normally vivid
iridescence and pigmentation cause them to stand out
against the background coloration of the skin. Also called
? false eyespot? (Fig. 20).
Octopodiformes ? Higher-level taxon that includes all
8-limbed cephalopods: vampires, cirrate octopods, incirrate
octopods. (Fig. 2 Living Cephalopods). Because of the long
history of referring to these cephalopods by the common
name ? octopods?, the latter is used as the common name
for the Octopodiformes.
Octopods ? Common name for Octopodiformes.
Ocular photophore ? Photophore(s) that lie(s) on the
eyeball (Fig. 28).
Oegopsida ? A high- level taxon with in the
Decapodiformes. In recent classification, the Oegopsida
(oceanic squids) has been considered the sister group of
the Myopsida and the two groups together compose the
Teuthoidea (squids). At present the composition and
affinities of the Oegopsida are unresolved.
Oesophagus ? The portion of the digestive tract between
the buccal mass and the stomach (Fig. 19). Often a portion
of the oesophagus is expanded to form a crop for food
storage (see Crop).
Olfactory papilla ? A pit, or bump-like to finger-like
protuberance on the posterolateral surface of each side of
the head; of olfactory function.
Olfactory organ ? A chemosensory organ present in all
coleoid cephalopods thought to be the homologue of the
rhinophore of Nautilus.
Ontogenetic descent ? The progressive descent into a
deeper-water habitat as a mesopelagic cephalopod grows
older and larger. This distribution pattern is particularly
common in many pelagic chiroteuthid and cranchiid squids.
Opening/closing trawl ? A trawl whose mouth is open
during fishing at a selected depth but is closed during
descent and retrieval.
Optic lobes of brain ? Large lobes of the brain associated
with the eyes. In octopods and some squids the optic lobes
may be separated from the rest of the brain by an optic stalk
of varying length.
Optic sinus ? See Eyelid sinus.
Oral ? Toward or pertaining to the mouth.
Orbital pore ? Minute pore in the anterior part of the
transparent tissue (cornea) that covers the eyes of
decapods except for the oegopsid squids; remnant of the
primary eyelids (Fig. 17).
Orbital sinus ? See Eyelid sinus.
Order ? The taxonomic category above the family level.
Outer cone ? Rim that surrounds the phragmocone in
cuttlebones.
Oviduct(s) ? Female gonoduct(s). The oviduct conducts
eggs from the visceroper icard ia l coelom, that
encompasses the ovary, to the mantle cavity and often is
used to store eggs. In some argonautid octopods eggs are
fertilized and undergo either partial (Argonauta) or
complete (Ocythoe) embryonic development within the
oviduct (Fig. 30).
Oviducal gland ? Glandular structure that surrounds the
anterior end of the primary oviduct and secrets some of the
external coatings around spawned eggs.
Paralarva ? The term that indicates the first free-living life
history stage (typically planktonic) for those cephalopods
that differ in morphology and ecology from older juveniles.
Pedicel (= sucker stalk ) ? A short, muscular stalk that
supports a sucker in sepioids and teuthoids (Fig. 32).
Cephalopods of the World 31
Fig. 31 Nuchal folds and nuchal crest
nuchal
folds
nuchal
membrane
a) Onychoteuthis b) Onykia
nuchal-locking
apparatus
nuchal
crest
32 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Pelagic ? (1) Free swimming in open ocean; (2) the region
of the ocean away from the ocean floor.
Pen ? See Gladius.
Penis ? The long, muscular terminal section of the male
gonoduct that serves to transfer spermatophores to the female
(Fig. 29). Apparently, in species with a hectocotylus, the penis
transfers spermatophores to the hectocotylus which in turn
transfers them to the female. In species without a hectocotylus,
the penis often is greatly elongate, capable of extending
beyond the mantle opening and apparently can transfer
spermatophores directly to the female.
Photocytes ? Cells that produce bioluminescence in
photophores.
Photophore ? An organ that produces and distributes
bioluminescence or ?living light?, either intrinsically through
biochemical reaction or extrinsically through luminescent
bacteria (Figs 17 and 28) (see Light organ).
Phylum ? The major, formative, principal taxonomic level;
above ?Class?.
Phragmocone ? System of plates that comprise the
cuttlebone.
Pocket, tentacular ? An open depression in the
anteroventral surface of the head between the bases of
arms III and IV of decapods, except myopsid and oegopsid
squids, into which the ejectable feeding tentacles are
retracted when not in use (Fig. 33).
Polarity (Evolutionary) ? The direction of evolution. That is,
one state is ?primitive? (plesiomorphic) and another is
?derived? (apomorphic).
Polarize (Evolutionary) ? To determine the direction of
evolution. That is, to determine which state is ?primitive?
(plesiomorphic) and which is ?derived? (apomorphic).
Posterior ? Toward the closed, tail-end of the mantle, away
from the head and arms.
Primary conus ? A solid conus on the gladius that is not
formed by the in-folding of the lateral vanes.
Protective membrane ? Thin web-like integument along
the lateral angles of the oral surface of the arms and clubs
lateral to the suckers, supported by muscular rods called
trabeculae (Fig. 34) (see Trabeculae).
Proximal ? Situated nearest or next to the centre of the
body or nearest the point of origin or attachment of a
muscle, appendage, etc. (opposite of distal).
Pseudomorph ? An ejected mass of ink and mucous that
approximates the size and shape of the cephalopod that
released it; i.e. a false body that fixes the attention of a
predator while the cephalopod escapes.
Rachis ? The thickened central axis that usually extends
the entire length of the gladius. Free rachis is the portion
that does not support vanes (Fig. 16) (see Gladius, Vane).
Radula ? The chitinous, ribbon-like band in the mouth of
cephalopods that contains up to seven transverse rows of
teeth that aid in transport of food into the oesophagus
(Fig. 35). Has a significant higher taxonomic value.
Fig. 32 Hectocotylized arm of males
pedicels
fringed
flaps
lamellae
(modified
trabeculae)
Fig. 33 Diagramatic sketch of arms and tentacles of
decapods other than myopsid and oegopsid squids
IV
III II I
tentacular
club
stalk
pocket
Fig. 34 Trabeculae, protective membranes and
suckers on arm of squid
sucker
trabeculae
protective
membrane
Fig. 35 Radula
Cephalopods of the World 33
Recent ? Geological term referring to an organism or
species that is living or has lived within the past 10 000
years, or to an object formed or events that have occurred
within the past 10 000 years.
Renal appendages ? Structures that form the nephridium
(= kidney). The renal appendages are out-pockets of the
veins within the renal sac (primarily the venae cavae) that
are covered with renal epithelium. The renal sac empties
into the mantle cavity via the nephridial (or renal) pore.
Renal pore ? The opening(s) of the renal cavities into the
mantle cavity, through which urine is discharged.
Rhinophore ? A short sensory tentacle in nautiluses. Four
in total, one on both sides of each eye.
Rhynchoteuthion ? Paralarval s tage of the
Ommastrephidae characterized by the fusion of the
tentacles into a trunk-like proboscis.
Rostrum (= spine) ? A spike-like posterior projection of the
gladius or cuttlebone, exterior to the conus (Figs 16 and 18).
Secondary conus ? A conical region at the posterior end of
the gladius that is formed by an in-rolling and fusion of the
vanes. The ventral line of fusion usually is apparent. The
secondary conus may be rather short or exceed half the
gladius length (Fig. 36).
Secondary eyelid ? An eyelid fold that covers the ventral
part of the cornea in cuttlefishes.
Secondary fin ? A non-muscular fin-shaped structure found
in some oegopsid squids, located posterior to the true or
primary fin (Fig. 37). The secondary fin may act as a
buoyancy organ. Vampire juveniles also have secondary
fins, which, however, are lost with growth.
Secondary web ? The narrow membrane that connects the
primary web to the arms in some cirrate octopods; e.g.
Cirroteuthidae.
Sepion (= cuttlebone) ? The calcareous, laminate, dorsal
supportive buoyancy structure in the mantle of cuttlefishes
(Fig. 18).
Semelparous ? A reproductive strategy in which females
spawn once then die. Sometimes called terminal or
?big-bang? spawners. Many cephalopods are semelparous
but in some species reproduction is prolonged (up to 50% of
the ontogenesis). Nautiluses are iteroparous and spawn
repeatedly over a period of years.
Sepioid gills ? Gills of some cirrate octopods that look
superficially like gills of sepioids.
Sepioidea ? A high-level taxon within the Decapodiformes.
Typically, this taxon includes the Sepiidae, Idiosepiidae,
Sepiolidae, Sepiadariidae and Spirulidae, but the
monophyletic nature of the group has been questioned. At
present the phylogenetic relationships and classification
within the Decapodiformes are unresolved.
Shell sac ? The sac that secretes the shell in the
Coleoidea, composed of ectodermal epithelium that
invaginates during embryonic development to form an
internal sac.
Side pockets ? Small membranous folds of the integument
that form small, shallow pockets lateral to the foveola in the
funnel groove (Fig. 22) (see Foveola ).
Siphuncle ? The tube-like posterior extension of the body
wall, coelom and covering sheath that penetrates into the
phragmocone chambers in cephalopods with a chambered
shell (Spirula, nautiluses and cuttlefishes). It regulates
gas exchange into the phragmocone chambers.
Spadex ? Male reproductive structure used to transfer
spermatophores to the female in nautiluses. It is formed by
the fusion of 4 circumoral tentacles.
Species ? Populations of animals that interbreed or are
potentially capable of interbreeding in nature. Considerable
debate exists over the general definition of a species and
how the theoretical definition should be applied in practice.
With regard to the latter problem, cephalopod species
generally are defined by distinct morphological traits not
exhibited by any other species. This practice is valid if
interbreeding does not occur. However, the amount of
interbreeding (i.e. hybridization) that actually occurs in
nature and contributes to or diminishes speciation is
virtually unknown in cephalopods.
Sperm cord ? The coiled rope of sperm that lies within the
spermatophore (Fig. 38).
Sperm duct (= seminal duct) ? The duct of males which
joins the testis with the spermatophoric organ (Fig. 29).
Sperm groove ? Sulcus along the ventral side of the
hectocotylus used to transfer the spermatophores (Fig. 11).
Sperm mass ? The mass of sperm held within the
spermatangia of everted spermatophores.
Fig. 36 Secondary conus Fig. 37 Secondary fin
Grimalditeuthis bomplandii
Sperm receptacle ? A bulbous structure in the buccal
region or at the openings of the oviducts in females of
certain cephalopods for deposition of spermatangia.
Spermatangium (pl. spermatangia) ? Extruded, exploded,
evaginated spermatophore/s, often in the form of a round
bulb.
Spermathecae ? Specialized sperm-storage structures
found in the skin of some female decapodiformes or as
pockets of the oviducal gland in octopods.
Spermatophore ? A tubular structure manufactured by
male cephalopods for packaging sperm; capable of holding
millions of sperm, it is transferred and attached to the
female until fertilization begins (Fig. 38). It forms a
spermatangium after the spermatophoric reaction occurs
and the spermatophore has everted.
Spermatophore pad ? A fleshy patch of tissue, usually in
the mantle cavity of some female cephalopods (e.g.
loliginids), to which spermatangia adhere after mating and
remain until fertilization occurs.
Spermatophoric complex ? The unit formed by the sperm
duct, the spermatophoric organ, the spermatophoric sac,
the spermatophoric duct and the penis (Fig. 29).
Spermatophoric duct ? The duct of males through which
the spermatophores, once formed, pass from the
spermatophoric organ to the spermatophoric sac (Fig. 29).
Spermatophoric organ ? Male organ where the
spermatophores are formed (Fig. 29).
Spermatophoric reaction ? The evagination of a
spermatophore with the extrusion of the sperm mass, caused
by the penetration of water inside the spermatophoric cavity,
where the osmotic pressure is higher.
Spermatophoric sac ? See Needham?s sac (Fig. 29).
Spine ? See Rostrum (Fig. 18).
Squid ? Common name given to members of the
Teuthoidea and some members of the Sepiolidae.
Squid, general terminology ? Diagrammatic drawing with
external features labelled, ventral view (Fig. 17).
Stalk of tentacle ? See Tentacle stalk .
Stalked eyes ? See Eye (position and size).
Statocyst ? A sense-organ that detects gravity, angular
accelerations and low-frequency sound. The statocyst is
embedded within the cephalic cartilage and contains the
statoliths.
Statolith ? A calcareous stone in the statocyst that detects
linear acceleration, angular acceleration and orientation
(Fig. 39). Statoliths of many species can be used to
estimate age.
Stellate ganglion ? Major ganglion of the peripheral
nervous system of neocoleoid cephalopods that controls
nerves to the mantle muscles.
Stomach ? The muscular organ of the digestive system
where primary digestion occurs (Fig. 19). The stomach
generally is lined with cuticular ridges to aid in grinding food,
and is supplied with digestive enzymes from the digestive
gland. The stomach may be greatly expandable in size and
serve as a storage area until food can be fully processed.
Striae ? Numerous thin septa visible in the ventroposterior
end of cuttlebone, making the cuttlebone appear as if
transversely striated (Fig. 18).
34 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 38 Spermatophore
ejaculatory apparatus
cement body
sperm cord
Fig. 39 Diagrams of a generalized teuthoid statolith (anterior view) with a) basic terms and
b) basic dimensions labelled
(after Clarke, 1978)
b)
do
rs
al
do
me
le
ngt
h
total length
ventrolateral
length
dorsolateral
length
a)
dorsal dome
dorsal spur
medial fissure
spur
superior lobe
wing
anterior rostral lobe
rostral angle
inferior lobe of
lateral dome
attachment area
lateral
dome
lateral rostral lobe
rostrum
ventral indentation
dorsal indentation
anterior
dorsal ridge
Stylets ? A pair of rod-like structures considered remnants
of the molluscan shell in incirrate octopods. Generally in the
form of a slender, cartilage-like, pointed rod tightly
surrounded by the shell sac and buried in the mantle
muscle at a dorsolateral position.
Subadult ? Stage at which all of the characters that typically
define the species are present, but the reproductive system
is not mature and functional. It follows the juvenile stage
and precedes the adult stage. A subadult stage is defined in
cephalopods since the adult phase frequently is
abbreviated.
Subequal ? Nearly equal. Generally refers to the length of
the arms when these appear to be approximately the same
length. Arm lengths cannot be measured very accurately
due to variation in their states of contraction.
Sucker/s ? Muscular, suction-cup structure/s on the arms and
tentacles (occasionally on the buccal membrane) of
cephalopods. It consists of a cup-shaped portion, the
acetabulum, and a flat, distal ring, the infundibulum, that
contacts the substrate. Some are stalked, placed on muscular
rods that contract (squids and cuttlefishes); some are sessile,
embedded without stalks on the oral surface of the arms
(octopuses) (Fig. 40a). They usually are counted either in
longitudinal rows or in transverse (oblique) series (Fig. 40b).
Sucker ring ? Chitinous, often serrated or toothed, ring that
encircles the opening of suckers of squids and cuttlefishes
(Fig. 41).
Sucker series ? The longitudinal rows of suckers on the
arms or tentacles of decapods. Series (= transverse rows)
contrasts with rows (= longitudinal rows) in describing
sucker arrangement. In octopods, sucker ?series? generally
is synonymous with ?rows?, the ?longitudinal? component of
the term is implied.
Sucker stalk ? The muscular support and connective
structure between the sucker and the arm. In octopods it is
a cylindrical structure about the same diameter as the
sucker and in decapods it is constricted into a conical pillar.
Sucker teeth ? Sharp, blunt, or rounded teeth on the inner
horny sucker rings of some decapods.
Sulcus ? A median longitudinal groove, sometimes flanked
by two low ridges on the ventral side of the cuttlebone.
Superior buccal lobes ? Lobes of the central nervous
system that sit dorsal to the oesophagus where the latter
enters the buccal mass.
Swimbladder ? Gas-filled structure found in the dorsal
region of the pelagic octopod, Ocythoe.
Swimming membrane (= keel ) ? An elongate, flat,
muscular vane along the aboral surface of arms and clubs
of decapods that functions to streamline and supports the
appendages during swimming (Figs 12 and 17).
Synonym ? One of two or more names applied to the same
taxon/species.
Systematics ? The classification of organisms into
hierarchal groups based on phylogenetic relationships.
Tail ? Posterior narrow extension of the body posterior to
the fins. The end of the fins and the beginning of the tail
often overlap. An operational definition for point of
demarcation for the purposes of measurement is: the point
where a hypothetical line, continuous with the broad
posterior edge of the fin, crosses the midline of the body
(Fig. 17).
Taxa, taxon ? A taxonomic group of any rank. A taxonomic
unit.
Tentacles ? Modified fourth pair of appendages in decapods,
used for prey capture and capable of considerable extension
and contraction. Distal ends contain clubs with suckers and/or
hooks; stalks are frequently devoid of suckers. Tentacles can
retract into pockets, or merely contract (Figs 17 and 33).
Although the tentacles are derived evolutionarily from the
fourth pair of arms, the term ?arms IV? is reserved for the
subsequent arm pair, the ventral arms, which are
evolutionarily the fifth pair of arms.
Tentacle absence ? Tentacles can be absent because the
species lacks tentacles, they are accidentally lost during
capture, or they are naturally lost at a particular stage of
development.
Tentacle pads ? Poorly understood and complex pad-like
photophores that are found on the tentacular stalks of some
squids of the family Chiroteuthidae.
Tentacle stalk ? Region of the tentacle proximal to the club.
Tentacle terminology ? See Figs 17 and 33.
Tentacular club ? The distal, terminal, usually expanded,
part of the tentacle that bears suckers and/or hooks. Used
for capturing prey (Figs 12, 17 and 33).
Cephalopods of the World 35
Fig. 40 Squid and octopus sucker and squid sucker
orientation
squid
a) sucker
octopus
b) club
suckers in
longitutinal
rows
8 suckers
in (oblique)
transverse
series
acetabulum
infundibulum
Fig. 41 Sucker ring
teeth
Tentacular retractor muscles ? Muscles that serve to coil
the tentacle when retracted, in contrast with those muscles
that serve to shorten (contract) the tentacle.
Tentacular sheath ? (1) The keels of arms IV are off-set
laterally and often enlarged to fully or partially conceal,
protect or encase the adjacent tentacles. The latter function
is most ful ly developed in the chiroteuthids and
mastigoteuthids; ( 2) tubular muscular sheaths of
nautiluses into which the tentacles can be retracted.
Terminal fins ? Fins with more than 50% of their length
posterior to the muscular mantle. These fins, therefore, are
at the ?terminal? or posterior end of the body and generally
are supported by an elongate secondary conus of the
gladius.
Terminal organ ? Alternative name for penis, as true
definition of a penis is ?organ of insertion?. In most
cephalopods, the hectocotylized arm is used for
spermatophore insertion, or placement, in the females.
Terminal pad (of tentacular club) ? A small, distinct pad or
circlet of small terminate suckers at the distal tip of the club.
Teuthoidea ? The higher taxon that includes all squid-like
decapods; now archaic. The monophyly of this taxon is
questionable.
Total length (TL) ? Length measured from the posterior tip
of the mantle to the anterior tip of the outstretched tentacles
(squids and cuttlefishes) or arms (octopuses) (Fig. 6).
Trabeculae ? Muscular rods that support the protective
membranes on the arms and clubs of cephalopods
(Fig. 34). Occasionally membranes are reduced and/or
trabeculae are elongated, so they extend beyond the edge
of the membrane, papilla-like.
Tragus ? Particular inward projecting knob in the funnel-
locking apparatus of some squids. It is the knob found on
the medial surface of the central depression (Fig. 8).
Truncate teeth ? Teeth on the inner chitinous rings of
decapod suckers that do not terminate in a point but rather
in a broad, flat tip.
Umbilicus ? The central core of the chambered nautilus
shell, representing the juvenile shell with its initial coils;
often a depression (Fig. 42). It can be open, so as to show
the inner most coils, or closed, often covered by a thickened
layer, the callus.
Vane ? Thin, lateral expansion of the gladius that arises
from the rachis (Fig. 16) (see Rachis).
Ventral ? The lowermost or belly surface of a cephalopod,
the surface on which the funnel is located. Opposite the
dorsal surface (Figs 17 and 20).
Visceral sac ? The body region posterior to the head
surrounded by the mantle. The body wall in this region that
encases the viscera usually is rather thin-walled, hence the
name ?visceral sac?. The visceral sac is also called the
?visceral dome?.
Visceropericardial coelom ? The largest coelom in all
cephalopods, except in the incirrate and cirrate octopods,
where it is greatly reduced. Generally it encloses the gonad,
and partially encapsulates the stomach, caecum and
ventricle, among other structures. The visceropericardial
coelom also communicates with the other cephalopodan
coelom, the nephridial coelom, and the mantle cavity via a
pair of ducts that open at the bases of the nephridial
papillae.
Water pores ? (1) Large cephalic orifices at base arms of
some pelagic octopods, e.g. Tremoctopus (Fig. 43);
(2) 8 small openings to the water pouches located at the
base of the arms on the ventral web of the benthic octopod
Cistopus.
Water pouches ? Glandular pouches with muscular pore
openings situated in the oral webs between the base of
each arm in the incirrate octopus genus Cistopus.
Web ? A membranous sheet of greater or lesser extent that
extends between the arms of many octopods, giving an
umbrella-like appearance when the arms are spread out,
e.g. on cirroteuthids (Figs 20 and 43). It is reduced or
absent in most decapods.
36 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 42 Nautilus (lateral view)
umbilicus
Fig. 43 Tremoctopus (dorsal view)
web
water pores
1.5 Key to Recent Cephalopod Groups and Families
1a. More than ten (63 to 94) circumoral appendages; suckers absent; chambered, coiled external shell . . . N a u t i l o i d e a
F a m i l y N a u t i l i d a e (Fig. 44)
1b. Eight or ten circumoral appendages; suckers (and/or hooks) present; no external shell . . . . . . C o l e o i d e a 
 2
2a. Suckers stalked with chitinous rings; 10 circumoral appendages, 8 arms and 2 ventrolateral
tentacles (tentacles may be lost in some species). Mantle cavity communicates with the exterior via
3 openings . . . . . . . . . C u t t l e f i s h e s , B o b t a i l s q u i d s , B o t t l e t a i l s q u i d s , P i g m y s q u i d s , R a m ? s H o r n s q u i d ,
M y o p s i d s q u i d s , O e g o p s i d s q u i d s 
 3
2b. Suckers without stalks, bases sometimes constricted in finned (cirrate) octopods and vampires,
without chitinous rings; 8 arms, no ventrolateral tentacles. Mantle cavity communicates with the
exterior via one opening, rarely 2 . . . . . . . . . . . . . . . . . . . . . . . . . . O c t o p o d s a n d V a m p i r e s 
 4
3a. Internal shell straight, laminate, calcified (Sepiidae), coiled, chambered (Spirulidae), rudimentary,
straight and chitinous (Sepiolidae) or absent (Sepiadariidae, Idiosepiidae); tentacles contractile
and retractile into pockets between arms III and IV; fins not joined posteriorly; mantle edge near
mantle cartilages straight . . . . . . . . . . . . . C u t t l e f i s h e s , B o b t a i l s q u i d s , B o t t l e t a i l s q u i d s , P i g m y s q u i d s ,
R a m ? s H o r n s q u i d 
 5
3b. Internal shell straight, feather- or rod-shaped, chitinous; tentacles contractile, not retractile, no
pockets; fins usually joined posteriorly; mantle edge near mantle cartilages with small projections or
?angles? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . M y o p s i d s q u i d s , O e g o p s i d s q u i d s 
 7
4a. Internal shell a chitinous, thin, broad plate; a pair of small filamentous appendages in pouches
between bases of arms I and II; light organ (photophore) present at base of each fin and medial to
each eye dorsally; colour black to dark red or brown . . . . . . . . . . . . . . . . . . . . V a m p i r e s ( monotypic)
F a m i l y V a m p y r o t e u t h i d a e (Fig. 45 )
4b. Internal shell vestige either small cartilaginous rods or a U-shaped support or absent; secondary
filamentous appendages absent; suckers sessile, without stalks and without chitinous rings; colour
extremely variable but never black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . O c t o p o d s 
 3 6
Cephalopods of the World 37
Fig. 44 Nautilidae ( Nautilus)
lateral view
Fig. 45 Vampyroteuthidae ( Vampyroteuthis)
dorsal view
5a. Internal shell calcified, coiled, chambered . . . . . . . . . . . . . . . . . . . . . . F a m i l y S p i r u l i d a e (Fig. 46 )
5b. Internal shell calcified, straight, laminate, chalky . . . . . . . . . . . . . . . . . . . . F a m i l y S e p i i d a e (Fig. 47 )
5c. Internal shell (gladius) chitinous (except Euprymna); one only or both dorsal arms or one
dorsolateral arm hectocotylized . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y S e p i o l i d a e (Fig. 48 )
5d. Internal shell (gladius) absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 6
6a. Left ventral arm hectocotylized; dorsal border of mantle fused with head . . . . . F a m i l y S e p i a d a r i i d a e (Fig. 49 )
6b. Both ventral arms hectocotylized; dorsal border of mantle free (unfused) from head
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y I d i o s e p i i d a e (Fig. 50 )
38 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 46 Spirulidae ( Spirula)
dorsal view
Fig. 47 Sepiidae ( Sepia)
dorsal view
Fig. 48 Sepiolidae ( Rossia)
dorsal view
F i g . 4 9 S e p i a d a r i i d a e ( S e p i o l o i d e a )
dorsal view
Fig. 50 Idiosepiidae ( Idiosepius)
dorsal view
7a. Eye covered by transparent
membrane (cornea) (Fig. 51a)
. . . . . . . . . . . . . . . . M y o p s i d s q u i d s 
 8
7b. Eye without cornea; lens in open
contact with seawater (Fig. 51b)
. . . . . . . . . . . . . . O e g o p s i d s s q u i d s 1 / 
 9
8a. Four longitudinal rows (series) of
suckers on manus of tentacular
clubs; fins united at posterior
end of mantle; medial posterior
border of fins concave (Fig. 52)
. . . . . . . . . . . . . . . . F a m i l y L o l i g i n i d a e
8b. Two longitudinal rows (series) off
suckers on manus of tentacular
clubs; fins not united at posterior
end of mantle; medial posterior
borders of fins convex (Fig. 53)
. . . . . . . . . . . . . . . . . Pickfordiateuthis2 /
9a. Funnel free from mantle; funnel-mantle locking apparatus present . . . . . . . . . . . . . . . . . . . . . . 
 1 0
9b. Funnel fused to mantle on each side3/; no funnel-mantle locking apparatus present . . . . . . . . . . . . . 
 3 5
Cephalopods of the World 39
Fig. 51
myopsid eye
(cornea)
oegopsid eye
(open)
a ) b )
Fig. 52 Loliginidae ( Loligo)
ventral view tentacular club
Fig. 53 Pickfordiateuthidae ( Pickfordiateuthis)
dorsal view tentacular club
2 rows of
suckers
1/ A new family of oegopsid squid has been recognized from Australian waters by C.C. Lu (personal communication). The description for
publication is currently in press.
2/
Pickfordiateuthis , the sole genus in a formally recognized family, recently has been placed in the family Loliginidae; it is included in the
Key to indicate its unique characters within the Loliginids.
3/ Fusion of the mantle to the funnel also occurs in adults of two Ommastrephid species: Sthenoteuthis oualaniensis and Eucleoteuthis
luminosa.
10a. Funnel-locking apparatus a simple, straight groove and ridge1/ (Fig. 54) . . . . . . . . . . . . . . . . . . . 
 1 1
10b. Funnel-locking apparatus not a simple, straight groove and ridge . . . . . . . . . . . . . . . . . . . . . . . 
 2 8
11a. Arms with hooks or with suckers in 4 longitudinal rows on proximal half of ventral arms . . . . . . . . . . . 
 1 2
11b. Arms without hooks and with suckers in 2 longitudinal rows on proximal half of ventral arms . . . . . . . . . 
 1 6
12a. Armature (suckers, hooks) of arms in 2 rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 1 3
12b. Armature (suckers, hooks) of arms in 4 rows (Fig. 55) . . . . . . . . . . . . . . . . . . . . . . F a m i l y G o n a t i d a e
13a. Tentacles and clubs absent in adults although present in larvae or occasionally in juveniles
(Taningia) but, when present, always with rudimentary clubs armed with few suckers (Fig. 56)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y O c t o p o t e u t h i d a e
13b. Tentacles present; fully developed clubs present (Fig. 57) . . . . . . . . . . . . . . . . . . . . . . . . . . 
 1 4
14a. Photophores on viscera and on mantle or surface of head or arms . . . . . . . . F a m i l y P y r o t e u t h i d a e (Fig. 57 )
14b.Photophores on mantle and surface of head and arms but not on viscera . . . . . . . . . . . . . . . . . . 
 1 5
40 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 54 Funnel-locking apparatus
e ) f )a ) d )c )b )
tragus
antitragus
Fig. 55 Gonatidae ( Gonatus )
armature elements (suckers and
hooks) in 4 rows
ventral view
Fig. 56 Octopoteuthidae ( Taningia )
dorsal view
rudimentary
or absent
Fig. 57 Pyroteuthidae ( Pterygioteuthis)
ventral view
1/ The classification ?simple and straight? includes some locking apparatuses that show considerable variation. For example, in the
Octopoteuthidae and the Histioteuthidae the central groove is fairly broad and may curve slightly. The homogeneity of this classification
becomes apparent when this type of locking cartilage is compared to the more highly specialized types (Fig. 54).
15a. Photophores on tentacles but not on eyeballs . . . . . . . . . . . . . . . . F a m i l y A n c i s t r o c h e i r i d a e (Fig. 58 )
15b. Photophores on ventral eyeballs but not on tentacles . . . . . . . . . . . . . . F a m i l y E n o p l o t e u t h i d a e (Fig. 59 )
16a. Buccal membrane connectives attach to ventral sides of arms IV (Fig. 60b) . . . . . . . . . . . . . . . . . 
 1 7
16b. Buccal membrane connectives attach to dorsal sides of arms IV1/ (Fig. 60a) . . . . . . . . . . . . . . . . . 
 2 3
17a. Hooks present on tentacular clubs (Fig. 61a and b); tentacles and clubs are lost in mature females
(Fig. 61c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y O n y c h o t e u t h i d a e
17b. Hooks lacking on tentacular clubs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 1 8
18a. Cartilaginous scales present on mantle (may be minute); tentacular clubs with 4 longitudinal rows of
suckers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 1 9
18b. Cartilaginous scales lacking; tentacular clubs with more than 4 longitudinal rows of suckers on some
areas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 2 0
Cephalopods of the World 41
Fig. 58 Ancistrocheiridae
( Ancistrocheirus)
dorsal view
Fig. 59 Enoploteuthidae
( Abralia )
ventral view
Fig. 60 Oral view
arm II I
arm IV
(ventral)
arms I (dorsal)
buccal connective
(ventrally attached)
arm II
buccal connective
(dorsally attached)
ba
stalk of
tentacle
hooks
b) tentacular cluba) dorsal view c) ventral view
(mature female)
Fig. 61 Onychoteuthidae ( Onychoteuthis)
no tentacles
1/ This character is difficult to detect in some histioteuthids that have secondary modifications to the buccal membrane connectives.
19a. Tentacles present with numerous laterally compressed club suckers . . . . . F a m i l y P h o l i d o t e u t h i d a e (Fig. 62 )
19b. Tentacles lost in adults; tentacles in juveniles small and weak, with a few (about 6) poorly
differentiated suckers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y L e p i d o t e u t h i d a e (Fig. 63 )
2 0a. Fin nearly as long as mantle, supported by strong, transverse, muscular ribs; minute suckers
present on oral surface of buccal lappets (Fig. 64) . . . . . . . . . . . . . . . . . . . F a m i l y C h t e n o p t e r y g i d a e
2 0b. Fins less than half the body length and without supporting ribs; no suckers on buccal lappets . . . . . . . . 
 2 1
21a. Tentacular clubs with 6 uniform longitudinal rows of suckers; a long, spike-like tail present greater
than fin length (Fig. 65) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y B a t o t e u t h i d a e
21b. Tentacular clubs with 2 or 4 longitudinal rows of suckers . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 2 2
42 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 62 Pholidoteuthidae ( Pholidoteuthis)
dorsal view
cartilaginous scales
on mantle
Fig. 63 Lepidoteuthidae ( Lepidoteuthis)
dorsal view
cartilaginous scales
on mantle
Fig. 64 Chtenopterygidae ( Chtenopteryx )
ventral view
Fig. 65 Batoteuthidae ( Batoteuthis)
ventral view tentacular club
2 2a. Tentacular clubs with 4 longitudinal rows of suckers on distal portion, numerous rows on proximal
portion; no long, spike-like tail (Fig. 66) . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y B r a c h i o t e u t h i d a e
2 2b. Tentacular clubs with 2 longitudinal rows of very widely spaced, tiny suckers; mantle broad, bluntly
rounded posteriorly; fin short, wide, transversely oval (Fig. 67) . . . . . . . . . . . . . F a m i l y W a l v i s t e u t h i d a e
2 3a. Ventral surface of eye with a row of photophores; buccal membrane with 8 separate lappets (Fig. 68)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y L y c o t e u t h i d a e
2 3b. No photophores on eyes; buccal membrane with 7 lappets or less . . . . . . . . . . . . . . . . . . . . . . 
 2 4
24a. Surface of mantle, head and arms covered with numerous photophores (usually large and distinct)
(Fig. 69) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y H i s t i o t e u t h i d a e
24b. Surface of mantle and head without photophores (arms may have a few photophores) . . . . . . . . . . . . 
 2 5
Cephalopods of the World 43
Fig. 66 Brachioteuthidae ( Brachioteuthis)
dorsal view tentacular club
Fig. 67 Walvisteuthidae ( Walvisteuthis)
dorsal view tentacular club
Fig. 68 Lycoteuthidae ( Lycoteuthis)
ventral view
photophores
Fig. 69 Histioteuthidae ( Histioteuthis)
ventral view
photophores
44 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
2 5a. Minute suckers present on oral surface of buccal membrane (Fig. 70) . . . . . . . . . . F a m i l y B a t h y t e u t h i d a e
2 5b. No suckers on oral surface of buccal lappets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 2 6
2 6a. Many small to minute suckers (or suckers and knobs) at proximal end of manus (Figs 71 and 72) . . . . . . . . 
 2 7
2 6b. No cluster of small suckers at proximal end of manus (Fig. 73) . . . . . . . . . . . . F a m i l y P s y c h r o t e u t h i d a e
2 7a. Posterior borders of fins slightly convex; carpal knobs in a single dorsal row or absent; adults attain
small size (Fig. 71) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y N e o t e u t h i d a e
2 7b. Posterior borders of fins concave; carpal knobs in a cluster alternating with carpal suckers; adults
attain gigantic size (Fig. 72) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y A r c h i t e u t h i d a e
Fig. 70 Bathyteuthidae ( Bathyteuthis)
dorsal view
tentacular
club
oral view of arms and
buccal membrane
suckers on
buccal lappets
photophores
Fig. 71 Neoteuthidae ( Alluroteuthis)
ventral view tentacular club
cluster of
suckers
only 1 row
of knobs or
none
Fig. 72 Architeuthidae ( Architeuthis)
dorsal view tentacular club
k nobs
alternating
with suckers
cluster of
suckers
Fig. 73 Psychroteuthidae ( Psychroteuthis)
dorsal view tentacular club
no cluster
of suckers
Cephalopods of the World 45
2 8a. Funnel-locking cartilage with a longitudinal and a transverse groove 
-shaped or


-shaped
(Fig. 54b and c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 2 9
2 8b. Funnel-locking cartilage oval, triangular, or oval with inward projecting knobs (Fig. 54d, e and f) . . . . . . . 
 3 0
2 9a. Funnel-locking cartilage with a longitudinal groove crossed by a transverse groove at its posterior
end, 
-shaped; fins less than 60% of mantle length (Fig. 74) . . . . . . . . . . . . . . F a m i l y O m m a s t r e p h i d a e
2 9b. Funnel-locking cartilage with a longitudinal groove from which a shorter groove branches medially,


-shaped; fins more than 80% of mantle length (Fig. 75) . . . . . . . . . . . . . . . F a m i l y Thysa n o t e u t h i d a e
3 0a. Funnel-locking cartilage oval with 1 or 2 knobs directed toward the centre of the concavity (Fig. 54d) . . . . . . 
 3 1
3 0b. Funnel-locking cartilage oval or subtriangular, without knobs (Fig. 54e and f) . . . . . . . . . . . . . . . . 
 3 2
31a. Club with only 4 longitudinal rows (series) of suckers (Fig. 76) . . . . . . . . . . . . . . F a m i l y C h i r o t e u t h i d a e
31b. Club with many (more than 15) longitudinal rows (series) of minute suckers (Fig. 77) . F a m i l y M a s t i g o t e u t h i d a e
Fig. 74 Ommastrephidae ( Ommastrephes )
ventral view
funnel-locking
cartilage
Fig. 75 Thysanoteuthidae ( Thysanoteuthis )
dorsal view
funnel-locking
cartilage
Fig. 76 Chiroteuthidae ( Chiroteuthis)
ventral view tentacular club
4 rows of
suckers
Fig. 77 Mastigoteuthidae ( Mastigoteuthis)
ventral view
suckers minute,
more than 15
rows
3 2a. Suckers on arms in 4 to 6
longitudinal rows (series) . . . . . . . . . 
 3 3
3 2b. Suckers on arms in 2 rows; tail
short (less than half of mantle
length) or absent . . . . . . . . . . . . . 
 3 4
3 3a. Six longitudinal rows of suckers on
arms I to III, 4 longitudinal rows of
suckers on arms IV; tail extremely
long (greater than mantle length),
as a spike-like extension of the
gladius; no fins on tail (Fig. 78)
. . . . . . . . . . . . F a m i l y J o u b i n i t e u t h i d a e
3 3b. Three or 4 longitudinal rows of
suckers proximally on all arms; fins
terminal, extremely long and
broad, extend far posterior to
mantle; gladius incorporated in
fins to posterior tip (Fig. 79)
. . . . . . . . . . . . . F a m i l y M a g n a p i n n i d a e
34a. Suckers on tentacular club in 4
longitudinal rows (series); mantle
free dorsally (Fig. 80) . . F a m i l y C y c l o t e u t h i d a e
34b. Suckers on tentacular club in 8 or
more longitudinal rows (series);
mantle fused dorsally to head
(Fig. 81) . . . . . . F a m i l y P r o m a c h o t e u t h i d a e
46 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 78
Joubiniteuthidae
( Joubiniteuthis)
dorsal view
tail
Fig. 79 Magnapinnidae
( Magnapinna )
gladius
ventral view
Fig. 80 Cycloteuthidae ( Discoteuthis)
ventral view tentacular club
4 rows of
suckers
Fig. 81 Promachoteuthidae ( Promachoteuthis )
ventral view tentacular club
3 5a. Mantle free dorsally, articulates with head by ridge and groove (Fig. 82) . . . . . . . F a m i l y G r i m a l d i t e u t h i d a e
3 5b. Mantle fused dorsally with head (Fig. 83) . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y C r a n c h i i d a e
3 6a. Fins present on mantle; cirri on arms . . . . . . . . . . . . . . . . . . . . . . . . . . . C i r r a t e O c t o p o d s 
 3 7
3 6b. Fins absent and cirri absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I n c i r r a t e O c t o p o d s 
 3 9
3 7a. Web attaches directly to arms . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y O p i s t h o t e u t h i d a e (Fig. 84 )
3 7b. Contractile intermediate membrane (?secondary web?) present between each arm and the primary
web . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 3 8
3 8a. Shell (cartilaginous fin support) simple, U-shaped . . . . . . . . . . . . . . . F a m i l y S t a u r o t e u t h i d a e (Fig. 85 )
3 8b. Shell more complex, with expanded fin-attachment areas . . . . . . . . . . . . F a m i l y C i r r o t e u t h i d a e (Fig. 86 )
Cephalopods of the World 47
Fig. 82 Grimalditeuthidae
( Grimalditeuthis)
ventral view
Fig. 83 Cranchiidae
( Helicocranchia )
dorsal view
Fig. 84 Opisthoteuthidae
( Opisthoteuthis)
top view
Fig. 85 Stauroteuthidae ( Stauroteuthis)
dorsal view
Fig. 86 Cirroteuthidae ( Cirrothauma )
ventral view
cirri and suckers
seen through
transparent web
3 9a. Muscle tissue of body gelatinous, often transparent in life . . . . . . . . . . . . . . . . . . . . . . . . . . 
 4 0
3 9b. Muscle tissue of body firm (may be covered by gelatinous subdermal layer) . . . . . . . . . . . . . . . . . 
 4 3
40a. Suckers biserial distal to edge of web (Fig. 87) . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y A l l o p o s i d a e
40b. Suckers uniserial along entire length of arms (Fig. 88) . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 4 1
41a. Eyes tubular, mantle fused to funnel base forming 2 openings to mantle cavity (Fig. 89) . F a m i l y A m p h i t r e t i d a e
41b.Eyes not tubular, though not necessarily round, single opening to mantle cavity . . . . . . . . . . . . . . . 
 4 2
42a. Digestive gland very elongate, spindle-shaped, pointed at one end . . . . . . F a m i l y V i t r e l e d o n e l l i d a e. . . . . . F a m i l y V i t r e l e d o n e l l i d a e (Fig. 90 )
42b. Digestive gland oblong, not pointed at end . . . . . . . . . . . . . . . . . . . . . . F a m i l y B o l i t e n i d a e (Fig. 88 )
43a. Funnel-locking apparatus absent; water pores on head absent; males not very much smaller than
females, with left or right ventrolateral arm hectocotylized (never in pocket), with spoon-shaped,
non-filamentous tip; females without dorsal arm flaps or permanent reticulate sculpturing of ventral
mantle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . F a m i l y O c t o p o d i d a e (Fig. 91 )
43b. Funnel-locking apparatus present; dwarf males very much smaller than females; hectocotylus (left
or right ventrolateral arm) temporarily coiled in sac below eye, with extremely long filamentous tip . . . . . . . . 
 4 4
48 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 87 Alloposidae ( Alloposus )
ventral view
2 rows of
suckers
Fig. 88 Bolitaenidae ( Japetella )
lateral view
1 row of
suckers
Fig. 89 Amphitretidae ( Amphitretus)
dorsolateral view
Fig. 90 Vitreledonellidae ( Vitreledonella)
lateral view
single opening
Fig. 91 Octopodidae ( Octopus )
dorsal view
web
funnel
eye
mantle
44a. Water pores present on head at bases
of both dorsal and ventral arms; dorsal
and dorsolateral arms of females
jo ined by very deep, th in web
. . . . . . . . . . . . . F a m i l y Tremo c t o p o d i d a e (Fig. 92 )
44b.Dorsal water pores absent; web, when
present, not as above . . . . . . . . . . . . . . . . . 
 4 5
45a. Females with broad, membranous flap
that secretes and holds a thin,
shel l- l ike egg case; males with
hectocotylus in non-stalked sac
beneath eye . . . . . . . . F a m i l y A r g o n a u t i d a e (Fig. 93 )
45b. Females with permanent reticulate
sculpturing on ventral mantle; dorsal
(first) arms of females lack broad,
membranous flap; no shell-like egg
case; males with hectocotylus in
stalked sac beneath eye (Fig. 94) . . . F a m i l y O c y t h o i d a e
Cephalopods of the World 49
Fig. 92 Tremoctopodidae ( Tremoctopus )
dorsal view
(female)
dorsal water
pores
web
Fig. 93 Argonautidae ( Argonauta )
lateral view of female
membrane
I
III
II
Fig. 94 Ocythoidae ( Ocythoe )
lateral view
permanent
sculpturing
50 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
2. CHAMBERED NAUTILUSES
by Patrizia Jereb
L iving nautiluses are limited to a few species belonging to 1 family and 2 genera: the sole survivors of a once extremelyspecious subclass. They are unique among living cephalopods in possessing a coiled, pearly, external shell (Fig. 95).
This is punctuated with chambers and the animal lives in the outermost chamber with its body attached to the sides of the
chamber by the adductor muscles. This outermost chamber is connected to the innermost (apical) chamber by a tube
(ectosiphuncle or siphuncular tube) which contains a core of tissue (endosiphuncle). Together, the tube and the internal
tissue form the siphuncle, which serves as a wick to remove the fluid from the chambers. This enables the animal to
regulate its buoyancy through control of fluid and gas in the chambers. Nautiluses have up to 47 pairs of circumoral
arm-like appendages, also called ?tentacles?, arranged in 2 ri ngs around the mouth and 2 pairs lateral to the eyes. These
appendages lack suckers and have several different sensory functions. Above the tentacles is a large fleshy wedge, called
the ?hood?. This is used as a trapdoor to seal the shell closed if the animal is attacked. On each side, between the hood and
the tentacles, are the eyes, which are simple and lack lenses, so that seawater flows in and out. Below the tentacles is a
rolled flap of skin which acts as a funnel, enabling jet propulsion powered by the large adductor muscles that contract,
pulling the body into the shell like a piston.
Until recently, our knowledge of these ?living fossils? was limited. They were thought to be rare animals occurring at only a
few remote Pacific island sites. Now they are known to occur through much of the tropical Indo-Pacific region, where they
live close to the bottom, primarily over reef slopes, from near the surface to about 500 m depth. Scavengers and
opportunistic predators, they seem to compete successfully with deep-water crustaceans and fishes.
Fig. 95 Schematic cross-section of a Nautilus
arm-like
appendages
hood
dorsal region
of mantle
obturating muscle
of the funnel
funnel
ventral region
of mantle
anus
digestive gland
septa separating chambers
chambers
umbilicus
ectosiphuncle
stomach
ovary
shell
gills
radula
(tooth-plate)
mouth
2.1 F a m i l y N A UTILI D A E de Blainville, 1825
Nautilidae de Blainville, 1825, Manuel de Malacologie et de Conchologie , 647 pages, Paris, 386.
FAO Names: En ? Chambered nautiluses; Fr ? Nautiles; Sp ? Nautilos.
Diagnostic Features: Coiled, pearly, ext ernal shell punctuated with chambers ; 2 pairs of gills; up to 47 pairs of arm-like
appendages around mouth; suckers and hooks lacking ; eyes simple, without lenses; funnel (or infundibulum) consisting
of 2 lobes which fold together to form a tube-like structure. Chromatophores and ink sac absent. Shell compressed and
involute to umbilicate and even perforate. An umbilical callus may be present. Shell sculpture of sinuous growth lines and
possibly fine spiral ridges in some species. Shell coloration variable by species. Irregular red to yellow-brown stripes
typically radiate from the umbilicus; they cover the entire shell in juveniles, but are restricted to the dorsal, chambered,
portion of shell in adults. Shell broader and larger in mature males than in mature females. Radula with 13 elements,
including 2 lateral teeth and 2 inner marginal support plates not seen in coleoids; jaws with calcite denticulation. Four
tentacles in males fused to form the spadix, used to transfer spermatophores to females during copulation. Males are
larger than females.
Size: Shell up to 229 mm in diameter, average animal weight up to 1 675 g. Shell size varies depending on species and
geography.
Habitat and Biology: Nautiluses are primarily mobile benthic bottom dwellers, associated with coral reefs, ranging from near
the surface to about 500 m depth, but their optimal range seems to be from 150 to 300 m. Factors controlling the upper limits
include predation by fishes and temperature: temperatures exceeding 25?C may be let hal for these animals. This explains why
most shallow water sightings of nautiluses occur at night and during cold seasons. Maximum depth limits are determined by
shell implosion (about 800 m) and chamber flooding (about 300 to 400 m). Males outnumber females and both live for many
years after maturity is reached at an estimated age of 5 to 15 years. Recent mark-recapture studies indicate that nautiluses may
live for more than 20 years. Growth is slow and no more somatic growth occurs after the animals reach sexual maturity. Egg
capsules have not been seen in natural habitats, but in captivity eggs are laid singly, attached to hard substrates and take up to
14 months to hatch in warm water (21? to 24?C). It is hypothesized that in nature they are laid in relatively shallow water (80 to
100 m) and that after hatching the young move to deeper, colder waters. The egg, similar in shape and size to a whole garlic, is
equipped with 1 or 2 capsules, separated by a narrow space full of seawater. The outer shell is white, tough but flexible, and
provided with a series of small holes that allow seawater to circulate between the 2 capsules. In aquaria, young hatchlings feed
almost immediately on shrimp or other food items. Decapod crustaceans seem to be a standard food for adults, but echinoid
fragments, fish bones, coleoid beaks and nautilus tentacles have also been observed in the crops of examined animals.
Nautiluses are highly mobile; tagged animals have been seen to move over 150 km in one year. They also move to shallow
waters at night and return to deeper waters during the day, where, however, they remain active.
Interest to Fisheries: At least 2 of these species are of commercial value as food, largely at the artisanal and subsistence
levels, and nautilus shells are sold commercially in the shell trade (e.g. in Indonesia, Fiji, New Caledonia, and the
Philippines). Nautiluses are also collected alive for public display and home aquaria and for research. They are caught
using baited fish traps.
Remarks: Although 11 species have been named within the genus Nautilus, only 4 are well established and can be
distinguished by differences in shell and (in some instances) soft part morphology: Nautilus belauensis, N. macromphalus,
N. pompilius and N. stenomphalus. One species, N. repertus, has questionable status. Recently, Ward and Saunders (1997)
erected a new genus, Allonautilus, for the most unusual member of the group, A. scrobiculatus, and one morphologically
similar, but extremely rare, and not well known form, A. perforatus.
Literature: Flower (1964), Solem and Roper (1975), Saunders (1981a), Roper et al. (1984), Saunders (1987), Saunders and
Landman (1987), Saunders (1998), Ward and Saunders (1997), Norman (2000).
Key to genera in the family Nautilidae
1a. Umbilicus small, or moderate, i.e. from 5 to about 16% of shell diameter; whorl cross-section oval (Fig. 96) . . . Nautilus
1b. Umbilicus larger, approximately 20% of shell d iameter; whorl cross-section quadrate (Fig. 97) . . . . . . Allonautilus
Cephalopods of the World 51
Fig. 96 Nautilus
cross-section
Fig. 97 Allonautilus
cross-section
52 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Nautilus macromphalus Sowerby, 1849 F i g . 9 8
Nautilus macromphalus Sowerby, 1849. Monograph of the genus Nautilus. In: Thesaurus Concyliorum, 2: 463?465
[type locality: unknown].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Bellybutton nautilus; Fr ? Nautile
bouton; Sp ? Nautilo ombligo.
Diagnostic Features: Umbilicus open, a deep, round
shouldered concavity, approximately 15 to 16% of shell
diameter at its widest point; inner coils of the shell
visible. No thickened layer (callus) present.
Size: Maximum shell diameter around 160 mm.
Geographic Distribution: Southwestern Pacific Ocean;
live animals known only from off northeastern Australia,
New Caledonia and Loyalty Islands, while drift shells
have been recorded as far south as Lizard Island,
Queensland (eastern Australia) (Fig. 99).
Habitat and Biology: Nautilus macromphalus inhabits
continental shelf and slope waters associated with coral
reefs, from the surface to a depth of about 500 m. In
some areas of southern New Caledonia, this species
rises to very shallow waters at night (i.e. less than 20 m
depth), so divers can observe animals foraging.
Interest to Fisheries: Consumed locally, it is
object of artisanal fishery. It also supports a
small fishery for public and private aquaria and
research trade. Collected alive at a depth of
about 65 m on the outer slope of the barrier reef
in New Caledonia; in the Coral Sea it is caught
by traps, at depths between 300 and 400 m.
Literature: Dunning (1998), Saunders (1987),
Norman (2000).
Fig. 98 Nautilus macromphalus
Fig. 99 Nautilus macromphalus
Known distribution
?? 200 m
Nautilus pompilius Linnaeus, 1758 F i g . 1 0 0 ; P l a t e I , 1 ? 5
Nautilus pompilius Linnaeus, (1758). ?Systema Naturae.? 10 (1): 709. (H olmiae.) [=Stockholm] [Reprint Leipzig, 1894]
[type locality: Ambon, Indonesia].
Frequent Synonyms: None.
Misidentifications: None.
FAONames: En ? Emperor nautilus; Fr ? Nautile flamm?;
Sp ? Nautilo com?n.
Diagnostic Features: Umbilicus small, visible as shiny
silver and black patch, closed; callus usually present (with
rare exceptions). No inner coils visible. Shell colour
patterns variable: irregular brown to reddish brown stripes
radiate from the umbilicus to venter in the usual
coloration, but this striping can be reduced to various
degrees, leaving the umbilicus and even much of the
flanks white.
Size: Shell diameters typically between 170 and 180 mm
around Fiji and the Philippines, larger in the Western
Australian population (i.e. mean diameter up to 222 mm).
Geographical Distribution: Indo-West Pacific;
Andaman Islands, Ambon, the Philippines, New
Guinea to Fiji; northeastern and northwestern
Australia. Absent from around New Caledonia,
where i t is replaced by Nautilus
macromphalus. Sympatric with Allonautilus
scrobiculatus off New Guinea and Nautilus
stenomphalus off northeastern Australia.
Replaced by Nautilus belauensis around Palau
(Fig. 101).
Habitat and Biology: Nautilus pompilius is the
most widely distributed and best known
Nautilus species. It inhabits deeper continental
shelf and slope waters around coral reefs, from
near the surface to a depth of about 750 m. The
species has been bred in captivity and animals
are frequently kept in public aquaria.
Interest to Fisheries: This species supports a
substantial shell trade, mostly from beach-drift
specimens, and subsistence and artisanal
fisheries. Captured in bamboo fish traps at
depths from 60 to 240 m, the meat is sold in local
markets and the shells go to the shell trade. The
outer layers of the shells are sometimes
removed, leaving the outer surface a silvery
mother-of-pearl layer.
Remarks: The Western Australian population includes the largest specimens known, with a mean shell diameter of
222 mm and mean weight of 1 675 g. Here, shell striping varies from the typical form to reduced yellowish stripes that leave
the umbilicus and flanks white. The questionable species N. repertus was described by Iredale (1944) on the basis of its
large size (about 228 mm shell diameter) and the reduced orange-brown coloration observed on 2 drifted specimens from
Western Australia. Historically, this has not generally been considered adequate to distinguish the species, given the
considerable range in variation in both shell size and colour pattern in N. pompilius. However, increasing documented
evidence of variation in shell colour pattern and dimensions between different regions, suggests that Nautilus pompilius
populations are isolated from each other, and may, at least, represent distinct subspecies. The application of recently
developed biochemical and genetic analysis may shed light on the validity of N. repertus and on the possible existence of
N. pompilius subspecies. Northern Australian records of animals with shells up to 240 mm in diameter may represent an
additional related species.
Literature: Saunders and Davis (1985), Saunders (1987), Saunders and Ward (1987a), Swan and Saunders (1987),
Dunning (1998), Saunders (1998).
Cephalopods of the World 53
Fig. 100 Nautilus pompilius
Fig. 101 Nautilus pompilius
Known distribution
?? 200 m
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE
Nautilus belauensis Saunders, 1981 P l a t e s I , 6 a n d I I , 7 ? 9
Nautilus belauensis Saunders, 1981, A new species of Nautilus from Palau, The Veliger, 24(1): 1?7, 3 plts, 2 text figs
[type locality: Palau, Western Caroline Islands].
Size: Shell diameter to 226 mm, animal weight to around 1308 g. The second largest species of Nautilus.
Geographical Distribution: Palau, Western Caroline Islands.
Remarks: The first species bred in aquaria.
Literature: Saunders,1981b, Saunders and Spinosa (1978, 1979), Norman (2000).
Nautilus repertus Iredale, 1944
Nautilus repertus Iredale, 1944, Australian pearly Nautilus. Australian Zoologist , 10(3): 294?298 [type locality: Rottnest
Island and Pelsart Island, Western Australia].
Size: Shell diameter up to 228 mm.
Geographical Distribution: Type locality and General distribution: Rottnest Island and Pelsart Island, Western Australia.
Remarks: Doubtful species. See Remarks in Nautilus pompilius.
Literature: Dunning (1998), Saunders (1998), Norman (2000).
Nautilus stenomphalus Sowerby, 1849 P l a t e I I , 1 0
Nautilus stenomphalus Sowerby, 1849. Monograph of the genus Nautilus. In: Thesaurus Concyliorum, 2: 463?465 [type
locality: Great Barrier Reef, eastern Australia].
Size: Shell diameter to around 170 mm.
Geographical Distribution: Great Barrier Reef, eastern Australia.
Remarks: Very similar in size and weight to Nautilus pompilius, from which it can be distinguished by the absence of the
callus and the reduced coloration, lacking in the umbilicus region. The hood is covered by elevated papillae, in a very
characteristic pattern. The first live captures were reported by Saunders and Ward (1987b).
Literature: Saunders and Ward (1987b), Saunders (1998), Norman, 2000.
54 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Allonautilus scrobiculatus ( Lightfoot, 1786 ) P l a t e I , 4 ? 5
Nautilus scrobiculatus Lightfoot, 1786, A catalogue of the Portland Museum, lately the property of the Duchess Dowager
of Portland, Deceased. London, 194 p. [Type localit y: Ndrova Island, Manus, Papua New Guinea].
Size: Shell diameter to around 180 mm.
Geographical Distribution: Tropical western Pacific: Papua New Guinea, Manus Province, Bismark Archipelago and
Milne Bay. Drift shells are known as far south as the Solomon Islands.
Remarks: The rarest and most distinctive species among nautiluses. Other than for the larger umbilicus, Allonautilus
scrobiculatus differs from the other nautiluses in the very peculiar features of the shell, including creases and encrusting
layer on the periostracum (see Fig. 97). Gills and male reproductive system also differ from those of Nautilus species.
Literature: Ward and Saunders (1997), Norman (2000).
Allonautilus perforatus (Conrad, 1847)
Nautilus perforatus Conrad, 1847, Notes on shells, with description of new genera and species. Journal of the Academy
of Natural Sciences, Philadelphia, Second Series, 1: 210?214 [type locality: unknown].
Size: Shell diameter to around 180 mm.
Geographical Distribution: Bali, Indonesia.
Remarks: Very rare form of questionable validity, only known from shells. Shell very similar to that of Allonautilus
scrobiculatus.
Literature: Ward and Saunders (1997).
Cephalopods of the World 55
3. CUTTLEFISHES
by Amanda Reid, Patrizia Jereb and Clyde F.E. Roper
The families formerly referred to as belonging to theOrder Sepioidea (Roper et al., 1984), include the
Sepiidae, which are of significant commercial value to
artisanal and industrial fisheries; the Sepiolidae, which
are exploited by many artisanal and subsistence
fisheries; the Sepiadariidae, which are not fished at
present (one species might be of potential interest); the
Idiosepiidae and the Spirulidae, which are of no
commercial interest. These families are treated herein
as separate entities.1/
Cuttlefishes sensu lato are characterized by the
following features: shell calcareous (Sepia, Spirula) or
chitinous (sepiolids); 10 circumoral appendages; 2
tentacles retractile into pockets; suckers with chitinous
rings; posterior fin lobes free, not connected at midline;
eye covered with a transparent membrane, false eyelids
present (except Spirula); 1 pair of gills, without
branchial canal between afferent and efferent branchial
blood vessels; digestive gland (liver) divided or bilobed;
each tooth of radula with a single projection; buccal
membrane present; olfactory organ a ciliated pit.
The combined catch of the exploited cuttlefishes made
up about 12 to 16% of total cephalopod catches in the
last 10 years, roughly fluctuating between 300 000 and
over 500 000 metric tonnes (FAO, 2000). The most
important genus exploited is Sepia.
Table 6 summarizes the catch data available for the
most important world fishing areas and identifies the
major cuttlefish-harvesting countries, which also are the
primary consumers and traders. These are mainly the
southeast Asian countries, the northwest Indian Ocean
regions and the nations that border the Mediterranean
Sea. In spite of the progress made in the last decade to
ensure the quality of data and information collected,
many of the figures above should still be considered as
very conservative, because the general tendency is to
underestimate the artisanal component of landings,
which in fact supports a conspicuous fraction of the
fishery but usually is not properly recorded in national
statistics.
In the industrial fisheries, cuttlefishes usually are taken
only as bycatch to other target species in bottom trawls,
even in cases where they make up a sizeable portion of
the catch. In the artisanal fisheries, on the other hand,
they are actively sought with highly selective gear and
fishing techniques based on knowledge of the biology
and behaviour of the species. Such techniques include
the use of aggregation devices (e.g. light, substrates for
egg deposition), live or artificial lures (for example, live
sexually mature females are used to attract males), and
a variety of fishing gear, such as harpoons, spears,
trammel nets, pound nets, hoop nets, lines, jigs, baited
pots, etc.
The flesh of most cuttlefishes is tender and excellent for
human consumption, as well as in terms of nutrients and
protein quality. It is marketed fresh, frozen, canned and dried
(?surume?).
56 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Area Country 2001 %
Atlantic, Eastern
Central
Morocco 17 544 39
China 8 238 18
Mauritania 4 744 10
Area 51 total 45 553 ?
Atlantic,
Northeast
France 13 814 72
United Kingdom 2 705 14
Portugal 1 348 7
Area 27 total 19 310 ?
Atlantic,
Southeast
Spain 26 84
Russian Federation 5 16
Angola 0
Area 47 total 31 ?
Indian Ocean,
Eastern
Thailand 20 819 57
Malaysia 9 810 27
Indonesia 5 360 15
Area 57 total 36 516 ?
Indian Ocean,
Western
Yemen 9 330 42
Pakistan 5 256 24
Oman 3 854 17
Area 51 total 22 086 ?
Mediterranean
and Black Sea
Tunisia 7 148 38
Italy 6 131 32
Greece 1 623 9
Area 37 total 18 904 ?
Pacific,
Northwest
China 310 129 95
Japan 8 297 3
Taiwan Province of China 4 546 1
Area 61 total 324 915 ?
Pacific,
Southwest
Australia 219 100
Area 81 total 219 ?
Pacific, Western
Central
Thailand 44 122 67
Malaysia 11 657 18
Indonesia 7 850 12
Area 71 total 65 766 ?
Cuttlefishes and
bobtail squids
World Total 533 300
(Source: FAO, 2000)
Table 6
Cuttlefishes and bobtail squids captured in 2001 by FAO
areas, showing the three leading countries in that year and
their share of total area capture
1/ The order of presentation of the sepioid families, sensu lato, in the text does not necessarily reflect phylogenetic or other systemic
sequence.
3.1 F a m i l y S E P I I D A E Keferstein, 1866 by Amanda Reid, Patrizia Jereb and Clyde F.E. Roper
Sepiidae Keferstein, 1866, Bronn?s Klass. Ordn., Thierreichs , 1862?66: 1441.
FAO Names : En ? Cuttlefishes; Fr ? Seiches; Sp ? Sepias, J?bias, Chocos, Choquitos.
Diagnostic Features: Small to medium-sized cephalopods. Mantle robust, slightly flattened dorsoventrally, may be broad
or slender; oval, oblong or nearly circular in outline; anterior dorsal mantle margin projected forward, not fused with head.
Fins narrow, located dorsolaterally on mantle, approximately equal to mantle length; posterior fin lobes free, not
connected to each other. Head robust, slightly narrower than mantle; eyes prominent, covered by a transparent membrane
and a conspicuous secondary fold on the eyelid. Mouth surrounded by 10 appendages (8 arms and 2 tentacles). Arms with
2 to 4 suckers in transverse rows. Males of some species with hectocotylized ventral arm(s) IV for holding and transferring
spermatophores; when present, the hectocotylus usually consists of a region of reduced suckers; hectocotylized region
also may be swollen and crenulated by transverse folds. Tentacular clubs with 4 or more suckers in transverse rows;
tentacles retractile into pockets on the ventrolateral sides of the head between arms III and IV . Arm and club suckers
with chitinous rings. Mantle-locking apparatus angular or curved in shape. Internal calcareous cuttlebone located dorsally
in the mantle underneath the skin; cuttlebone length usually equal to mantle length (Metasepia is an exception); cuttlebone
shape ranges from lanceolate to oval or diamond-shaped; dorsal side a calcareous plate (dorsal shield); ventral side finely
laminate, porous, and comprised of thin, transverse septa supported by transverse calcareous rods. One pair of gills; no
branchial canal between afferent and efferent branchial blood vessels. ?Liver? (digestive gland) divided or bilobed. Buccal
membrane present, with or without suckers; each radula tooth unicuspid (with a single projection). Olfactory organ a
ciliated pit.
Cephalopods of the World 57
Fig. 102 Diagram of basic cuttlefish features
left arm IV in male
hectocotylized
modified
portion
tentacular club
medial suckers
(moderately
enlarged)
cuttlebone
ventral view lateral view
inner cone
striae
rostrum
or spine
IV III II I
tentacular club
stalk
fin
spine
buccal membrane
mantle
pocket
funnel
2 rows
4 rows
ventral view
outer
cone
Size: Up to 500 mm mantle length, and 12 kg in weight.
Habitat and Biology: Cuttlefishes inhabit the continental shelf and upper slope to a maximum depth of approximately
1000 m. They are primarily bottom-dwellers over a range of habitats, including rocky, sandy, and muddy substrates,
seagrass, seaweed and coral reefs. They are slower swimmers than the more streamlined squids. Cuttlefishes are able to
attain neutral buoyancy by regulating the relative amounts of gas and fluid in the chambers of the cuttlebone, and they are
able to hover in midwater, with fins acting as stabilizers. The cuttlebone length, width, septal spacing and structural
morphology are correlated to maximum habitat depth: the deepest living species known, including Sepia australis,
S. elegans, S. orbignyana and S. hieronis, all are capable of descending to depths in excess of 400 m; all share the
characteristics of cuttlebones with closely packed septa and modified sutures. Large species such as Sepia latimanus,
S. officinalis and S. pharaonis are restricted to much shallower depths, usually less than 200 m, and show very different
septal spacing and sutures than the deeper water species. Some species migrate seasonally in response to temperature
changes and aggregate, usually in shallow water, at spawning time. Within a species, individuals may attain sexual
maturity at very different sizes, depending upon the combined effects of a number of factors including temperature, light
and diet. Eggs, relatively few in number, are individually attached to various substrates in clusters; the time required for
development depends upon temperature. The lifespan is between 18 and 24 months, although males of some species may
live longer. Post-spawning mortality is high in females. Cuttlefishes feed on a wide range of invertebrates and bony fishes.
Literature: Hoyle (1886), Adam (1939a), Adam and Rees (1966), Roeleveld (1972), Adam (1979), Nesis (1987), Ward
(1991), Oka et al. (1989), Okutani (1995), Khromov et al. (1998) and Lu (1998a).
Key to genera in the family Sepiidae
1a. Cuttlebone diamond-shaped in outline (Fig. 103); cuttlebone much shorter than mantle, located in
the anterior 1/2 to 2/3 of mantle; dorsal anterior edge of mantle without tongue-like projection
(Fig. 104a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Metasepia
1b. Cuttlebone outline elliptical to lanceolate; cuttlebone length approximately equal to mantle length;
dorsal anterior edge of mantle usually with tongue-like projection (Fig. 104b) . . . . . . . . . . . . . . . . . 
 2
58 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 103 Metasepia pfefferi
(illustration: K. Hollis/ABRS)
cuttlebone (ventral view)
Fig. 104 Head and anterior mantle (dorsal view)
(illustrations: K. Hollis/ABRS)
a) Metasepia pfefferi b) Sepia, Sepiella
tongue-like
projection
2a. A gland and gland pore located on the ventral side of the posterior end of the mantle (Fig. 105);
mantle-locking apparatus with triangular projection (Fig. 106a); cuttlebone inner cone with very
short limbs; outer cone a wide, spatulate, chitinized border around posterior end of cuttlebone
(Fig. 107a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepiella
2b. Gland and gland pore absent; mantle-locking apparatus semicircular, without triangular projection
(Fig. 106b); cuttlebone inner cone with relatively long limbs; outer cone usually calcareous, not
obviously spatulate posteriorly (Fig. 107b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepia
Cephalopods of the World 59
Fig. 105 Sepiella
(illustration: K. Hollis/ABRS)
posterior end of mantle
(ventral view)
gland
pore
Fig. 106 Mantle-locking apparatus
a) Sepiella b) Sepia
triangular
projection
Fig. 107 Cuttlebones
(illustrations: K. Hollis/ABRS)
a) Sepiella
outer cone
chitinous,
flared
posteriorly
outer cone
chitinous,
not flared
posteriorly
b) Sepia
ventral view
60 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Metasepia pfefferi ( Hoyle, 1885 ) F i g . 1 0 8 ; P l a t e I I , 1 1
Sepia (Metasepia) pfefferi Hoyle, 1885, Annals and Magazine of Natural History , Series 5, 16: 199 [type locality:
Challenger Stn. 188, 09?59'S 139?42' E, South of Papua, Arafura Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Flamboyant
cuttlefish; Fr ? Seiche flamboyante;
Sp ? Sepia llamativa.
Diagnostic Features: Mantle oval,
very broad. Arms broad, blade-like;
arms I shorter than rest. Protective
membranes in both sexes narrow.
Arm suckers tet raser ia l .
Hectocotylus present on left ventral
arm: oral surface of modified region
wide, swol len, f leshy, wi th
transversely grooved ridges and
deep median furrow. Club
sucker-bearing surface flattened,
with 5 or 6 suckers in transverse
rows ; suckers differ markedly in
size: 3 or 4 median suckers greatly
enlarged, occupying most of
median portion of club. Swimming
keel of club extends well proximal to
carpus; dorsal and ventral protective
membranes not joined at base of
club but fused to tentacular stalk.
Dorsal and ventral membranes
differ in length; extend proximal to
carpus along sta lk . Dorsal
membrane forms shallow cleft at
junction with stalk. Cuttlebone much
shorter than mantle, located in
anterior 2/3 to 3/4 of mantle ; outline
rhomboidal; acuminate, acute,
anteriorly and posteriorly; dorsal
surface yellowish, evenly convex;
texture smooth, not pustulose;
dorsal median rib absent. Chitin
present as a thin film over entire
dorsal surface of cuttlebone. Spine
absent, or small, chitinous. Striated
zone concave; last loculus strongly
convex, thick in anterior third; sulcus
deep, wide, extends along striated
zone only. Anterior str iae are
inverted V-shape; limbs of inner
cone are very short, uniform width,
narrow, U-shape thickened slightly
posteriorly; outer cone absent.
Dorsal mantle has 3 pairs of large,
flat, flap-like papillae . Head with
papillae over eyes . Colour: Base
colour of dark brown with mobile
patterns of white and yel low;
purple-pink along arms.
Size: Up to 60 mm mantle length.
lateral view dorsal view
Fig. 108 Metasepia pfefferi
lateral view ventral viewdorsal view
c u t t l e b o n e
(illustrations: K. Hollis/ABRS)
Geographical Distribution: Tropical Indo-Pacific:
northern Australia from Mandurah, Western Australia,
32?33'S 115?04'E, northeas tward to Moreton Bay,
southern Queensland, 27?25' S 151?43'E. New Guinea
(Fig. 109).
Habitat and Biology: Depth range from 3 to 86 m. Sand
and muddy substrate. Active during the day, hunting
fishes and crustaceans. Capable of excellent
camouflage while stalking prey. If disturbed or attacked,
it quickly changes to a dramatic colour pattern of dark
brown, black, white and yellow patches and may show
bright red on the arm tips. It has been observed to
?amble? along the sea floor in this colour pattern,
rhythmically waving the wide protective membranes on
the arms. Eggs are laid singly in crevices or ledges in
coral, rock or wood. One female has been observed
squeezing her eggs through the hole of a coconut husk.
The round white eggs turn clear as they develop.
Juveniles are capable of the same bright colour pattern
as adults from birth.
Interest to Fisheries: None as food. But this species is
spectacular in its colour and textural displays and would
make an excellent aquarium species.
Literature: Adam and Rees (1966), Roper and
Hochberg (1987), Roper and Hochberg (1988), Lu
(1998a).
Metasepia tullbergi (Appell?f, 1886 ) F i g . 1 1 0
Sepia tullbergi Appell?f, 1886, Svenska Vetenskaps-Akademiens Handlingar ,
21(13): 26 [type locality: Japan, Nagasaki].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Paintpot cuttlefish; Fr ? Seiche encrier; Sp ? Sepia tintero.
Cephalopods of the World 61
Fig. 109 Metasepia pfefferi
Known distribution
?? 200 m
dorsal view
dorsal view lateral view ventral view
c u t t l e b o n e
Fig. 110 Metasepia tullbergi
(after Sasaki, 1929 )
(after Sasaki, 1929 )
t e n t a c u l a r
c l u b
62 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Diagnostic Features: Mantle broad, oval. Ventral surface of mantle with 10 to 13 pores on each ventrolateral surface. Fins
joined posteriorly. Arm suckers tetraserial. Hectocotylus present on left ventral arm: suckers normal proximally, 10 to 12
rows reduced suckers medially, followed by some enlarged suckers, then reduced suckers to arm tip; reduced suckers
much smaller than normal arm suckers. Oral surface of modified region is wide, swollen, fleshy, and has
transversely-grooved ridges. Suckers of hectocotylus in 2 dorsal and 2 ventral series displaced laterally, with gap between
them. Club crescent-shaped, sucker-bearing surface flattened, with 4 to 6 suckers in transverse rows; suckers differ
markedly in size: 4 or 5 suckers in second dorsal series enlarged and 3 or 4 suckers in third series from dorsal side
slightly enlarged. Swimming keel of club extends well proximal to carpus; dorsal and ventral protective membranes not
joined at base of club; dorsal membrane broad, much wider than ventral membrane. Cuttlebone much shorter than mantle,
located in anterior 2/3 to 3/4 of mantle; outline rhomboidal, broad; bone strongly convex in lateral view; acuminate, acute,
anteriorly and posteriorly; dorsal surface evenly convex. Chitin present as a thin film over entire dorsal surface of
cuttlebone; bone extends posteriorly into sharp spine with chitinous plate on dorsal side . Cuttlebone nearly completely
chitinized anteriorly, calcareous posteriorly; sulcus shallow, narrow, extends along striated zone only; limbs of inner cone
are very short; outer cone absent. Head and dorsal and lateral mantle rugose with numerous papillae and tubercles; pair of
broken longitudinal ridges on dorsum. Dorsum dark in colour lateral to fleshy ridges, with bright yellow patches on head ,
dorsal mantle and arms. Protective membranes on arms red.
Size: Up to 70 mm mantle length, from 30 to 40 g total
weight.
Geographical Distribution: Indo-Pacific: Japan from
southern Honshu, Sea of Japan, Yellow Sea, East
China Sea to Taiwan Province of China and Hong
Kong, South China Sea, Philippines, Gulf of Thailand
(Fig. 111).
Habitat and Biology: Depth range from 20 to 100 m.
This nerit ic demersal species occurs on the
continental shelf on sandy to muddy substrate; it also
has been collected from among sea pens on rocky
substrate. Eggs hatch in summer in rocky areas at
about 20 m depth. Hatchlings migrate to deeper,
sandy-mud areas (down to 80 m) in August to
September; mature individuals migrate to shallower
rocky areas from March to May to spawn.
Interest to Fisheries: Currently there is no known
fishery for this small, colourful species, but it does
occur occasionally as bycatch. Its relevance in
artisanal fisheries is undetermined. If its supply were
steady, it undoubtedly would be a popular, colourful
aquarium species.
Local Names: CHINA: Mak dau; JAPAN: Hana-ika.
Literature: Okutani et al. (1987), Nomura et al. (1997). Fig. 111 Metasepia tullbergi
Known distribution
?? 200 m
Sepia aculeata Van Hasselt, 1835 ( in F?russac and d?Orbigny, 1834?1848) F i g . 1 1 2 ; P l a t e I I , 1 2 ? 1 4
Sepia aculeata Van Hasselt, 1835, pl. 5 bis (in F?russac and d?Orbigny 1834?1848), Histoire naturelle g?n?rale et
particuli?re des C?phalopodes Ac?tabuliferes vivants et fossiles [type locality: Java].
Frequent Synonyms: Sepia indica d?Orbigny, 1848.
Misidentifications: None.
FAO Names: En ? Needle cuttlefish; Fr ? Seiche aiguille;
Sp ? Sepia de aguja.
Diagnostic Features: Mantle oblong. Male and female arm lengths subequal. Hectocotylus present on left ventral arm: 3
rows of normal size suckers proximally, 5 or 6 rows of reduced suckers medially, then normal size suckers to arm tip.
Suckers of hectocotylus in the 2 dorsal series aremuch smaller than those in the 2 ventral series ; suckers in both dorsal
and ventral series displaced laterally, with gap between; arm with deep median furrow in adults. Club sucker-bearing
surface convex; males with 10 to 12, females with 13 or 14 club suckers in transverse rows; all club suckers of similar,
minute, size . Dorsal and ventral protective membranes not joined at base of club, extend proximal to carpus along stalk.
Buccal membrane with a few minute suckers. Cuttlebone with distinct dorsal median and lateral ribs. Chitin present as wide
glaze-like patch posteriorly; a narrow chitinous rim borders lateral margins of cuttlebone. Spine long, pointed, without keel.
Anterior striae are inverted U-shape . Inner cone limbs are narrow anteriorly, broaden poste riorly, then are raised into a
thick, round ledge ; outer cone narrow anteriorly, broadens posteriorly. Dorsal mantle with longitudinal row of up to 8
ridge-like papillae and associated, numerous, large papillae along each side, adjacent to base of each fin. Head and arms
with scattered, small tubercles. Colour of preserved specimens pale brownish. Dorsal mantle has white blotches, spots or
patches, or it has a transverse saddle mark, or a fine, darkly pigmented, transverse reticulate pattern. Fins with pale
reflective line along base . Live animals are light brown. Arms I to III have a longitudinal orange-red pigmented stripe along
their aboral surfaces. Dorsal mantle with pale transverse saddle mark and pattern of bold, broad, zebra stripes during the
spawning season; 3 circular, white patches occur mid-dorsally within transverse saddle mark; posteriormost patch
composed of a ring of white spots.
Size: Maximum mantle length 230 mm; weight 1.3 kg.
Cephalopods of the World 63
Fig. 112 Sepia aculeata
dorsal view
t e n t a c u l a r c l u b
c u t t l e b o n e (illustrations: K. Hollis)
ventral viewdorsal view
64 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Geographical Distribution: Indo-Pacific: India
(Arabian Sea north to off the Gulf of Kutch) to the
Andaman Sea and Gulf of Thailand, South China Sea,
Taiwan Province of China, East China Sea, Indonesia
and possibly the Timor Sea (Fig. 113).
Habitat and Biology: Sepia aculeata is a demersal,
neritic species that occurs to 60 m depth. Along the
Indian coasts maturing and mature animals are
present throughout the year, which indicates a
prolonged spawning and breeding season; activity
peaks occur from January to April and in the second
half of the year in the eastern waters, and in April, July
and December in the southwestern waters, off Cochin.
In the Hong Kong area, a prespawning concentration
occurs on the continental shelf, followed by movement
inshore to spawn at depths of 5 to 20 m from March to
May at water temperatures of 18? to 24?C. In the Gulf
of Thailand, spawning occurs all year at depths of 10 to
50 m, with March to April and July to September the
peak months. Smallest mature individuals in this
region and in the Bay of Bengal (eastern Indian
Ocean) were found at 70 mm mantle length, while size
at 50% maturity varies between 75 and 100 mm for
males and 100 and 120 mm for females, respectively.
In the Arabian Sea (western Indian Ocean) the size at 50% maturity is a little higher, i.e. over 120 and 130 mm for males
and females, respectively. Growth rates are similar in females and males and the sex ratio is 1:1. Commercially caught
animals from the Gulf of Thailand and the Andaman Sea range between 60 and 130 mm mantle length, those caught from
the east coast of India range between 50 and 190 mm and on the west coast females of 200 mm mantle length also are
captured.
Interest to Fisheries: This species is the thirdmost important commercial cuttlefish around Hong Kong where it is caught
with setnets and seines during the spawning season. It is also one of the most important commercial cuttlefish in southwest
India, where is mainly caught by trawl, with peak landings in October and November. The species is fished commercially in
southern China, Taiwan Province of China, Sri Lanka and Thailand, where it is caught by otter trawl, pair trawl, squid
light-lures, traps and push nets. Captures by traps in Thai waters are most abundant in January and February, when most
animals are fully mature and females are predominantly caught.
Local Names: CHINA: Jam Mak Yue; JAPAN: Ami-mom-nkouika.
Literature: Adam and Rees (1966), Silas et al. (1986), Siraimeetan (1990), Chotiyaputta (1993), Rao et al. (1993),
Chantawong and Suksawat (1997), Chotiyaputta and Yamrungreung (1998), Nateewathana (1999).
Fig. 113 Sepia aculeata
Known distribution
?? 200 m ?
Sepia andreana Steenstrup, 1875 F i g . 1 1 4
Sepia andreana Steenstrup, 1875, Danske Videnskabernes Selskabs Skrifter, 5 Raekke, Naturvidenskabelig og
Mathematisk , 10(7): 473 [type locality: Japan, Hakodate].
Frequent Synonyms: None.
Misidentifications: Sepia kobiensis Hoyle, 1885.
FAO Names: En ? Andrea cuttlefish; Fr ? Seiche
andreana; Sp ? Sepia andreana.
Diagnostic Features: Mantle oblong; dorsal anterior margin triangular, acute; ventral mantle margin emarginate. Fins
narrow, end in small, auriculate lobes posteriorly, with a small V-shape interstice where posterior spine is located. Male
arms II greatly elongate, 3 times longer than other arms (except in young specimens), and bluntly rounded distally, not
tapered. Female arm lengths subequal. Arm sucker arrangement differs between sexes. In males, sucker rows on modified
arms II tetraserial on proximal third, biserial distally, becoming rudimentary and sparse at tips; arms I and III tetraserial
proximally, biserial only on extreme distal arm tips; arms IV suckers tetraserial. In females, suckers on arms I tetraserial
proximally, biserial on slightly expanded distal tips; arms II and III tetraserial proximally, distal third with biserial suckers,
arms IV suckers tetraserial; female distal biserial suckers minute, displaced laterally, with gap between, and with distinct
medial gap between rows. Hectocotylus present on left ventral arm: 10 rows of normal size suckers proximally, distal half
of arm with rudimentary suckers on swollen peduncular bases. Club crescent-shaped, sucker-bearing surface convex
with 5 or 6 suckers in transverse rows ; suckers differ in size: 4 or 5 slightly enlarged suckers in longitudinal series
towards posterior end of club. Swimming keel of club extends proximally slightly beyond carpus; dorsal and ventral
protective membranes not joined at base of club but fused to tentacular stalk, terminate at posterior end of carpus. Dorsal
membrane forms deep cleft at junction with stalk. Buccal membrane without suckers; in females extends ventrally with 2
spermathecae. Cuttlebone outline lanceolate; dorsal median rib indistinct; spine straight; striated zone and last loculus
convex; sulcus shallow, narrow, extends entire length of cuttlebone. Anterior striae shallow m-shape ; inner cone narrow,
U-shape; outer cone limbs are expanded posteriorly into 2 short ?wings?, directed v entrally, to form a recurved cup-like
structure. Dorsal mantle with a longitudinal row of approximately 6 ridge-like papillae along each side, adjacent to base of
each fin. Colour: Pale brownish. Arms II have oblique bands of chromatophores, arms I to III have a longitudinal
orange-red pigmented stripe along their aboral surfaces . Dorsal mantle has yellow spots and chromatophores
concentrated medially above cuttlebone. Ventral mantle pale with a narrow, longitudinal, iridescent band on each side
adjacent to fins.
Cephalopods of the World 65
Fig. 114 Sepia andreana
dorsal view
dorsal view ventral view
c u t t l e b o n et e n t a c u l a r c l u b h e a d o f m a l e
(after Voss, 1963 )
(illustration: A. Reid) (illustration: K. Hollis)
(after Okutani, 1995 )
(illustration: A. Reid)
66 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Size: Up to 120 mm mantle length, 200 g total weight.
Geographical Distribution: Western Pacific: Japan
(Pacific coasts of southern Hokkaido, Tsugaru Strait,
northern and central Honshu), Japan Sea, Yellow Sea,
along south China coast, South China and East China
Seas to northern Philippines. (Fig. 115).
Habitat and Biology: Depth to 50 m. This demersal
species occurs in coastal waters. In the Yellow Sea the
spawning season is very long, with two spawning
peaks, one in spring and one in autumn, the main
spawning occurring in March and April. It feeds on
fishes and small crustaceans.
Interest to Fisheries: Taken as bycatch in trawl and
setnet fisheries. It consistently supports a fishery in the
Yellow Sea.
Remarks: Sepia andreana may be confused with
S. kobiensis Hoyle, 1885. Mature male S. andreana
can be distinguished by the greatly elongate second
arm pair, with biserial, rather than tetraserial, suckers
on the distal half. All arms are of similar length in
S. kobiensis and suckers on all arms are tetraserial.
The hectocotylus sucker arrangement differs between
the two species: Sepia andreana has 10 proximal rows
of normal size suckers, and the remainder of the arm
has rudimentary suckers on swollen peduncular
bases; Sepia kobiensis has 6 to 12 series of normal suckers proximally, 7 to 10 series medially are reduced, with those on
the distal remainder of the arm normal in size. The club suckers differ only slightly in size in S. kobiensis, while in
S. andreana they differ markedly in size. Adult Sepia kobiensis are also slightly smaller. Sepia andreana was misidentified
in Voss (1963) and Roper et al. (1984). The species referred to in those works could be S. kobiensis. Sepia andreana lives
in cooler water than the distributions in these publications would suggest.
Literature: Okutani et al. (1987), Cheng (1997), Kubodera and Yamada (1998).
Fig. 115 Sepia andreana
Known distribution
?? 200 m
Sepia apama Gray, 1849 F i g . 1 1 6 ; P l a t e I I I , 1 5 ? 2 0
Sepia apama Gray, 1849, Catalogue of the Mollusca in the British Museum. Part I. Cephalopoda Antepedia , 103 [type
locality: South Australia, Port Adelaide, 34?50'S 138?30'E].
Frequent Synonyms: Sepia palmata Owen, 1881; Amplisepia verreaux (non Rochebrune, 1884) Iredale, 1926;
Amplisepia parysatis Iredale, 1954.
Misidentifications: None.
FAO Names: En ? Giant Australian cuttlefish; Fr ? Seiche g?ante australienne; Sp ? Sepia gigante australiana.
Diagnostic Features: Very large, robust species. Mantle broadly oval. Fins wide, extend anteriorly slightly beyond mantle
margin, rounded posteriorly, with wide gap between them. Funnel long, broad-based, extends to level of anterior rim of eye.
Funnel organ dorsal elements inverted V-shape, with a small anterior papilla; ventral elements oval with acute anterior tips.
Head short, broad, narrower than mantle. Male and female arms subequal in length; protective membranes wide, well
developed. Arm suckers tetraserial. Hectocotylus present on left ventral arm: proximal end of arm with 6 to 10 rows of
slightly reduced, equal sized suckers ; oral surface of modified region not wide, fleshy, but normal, as on opposite arm.
Club crescent-shaped, moderate length; sucker-bearing surface flattened, with 4 or 5 suckers in transverse rows ;
suckers differ markedly in size: some median suckers greatly enlarged ; dorsal marginal longitudinal series of suckers
slightly larger than those in ventral marginal series. Swimming keel of club very broad, extends well proximal to carpus;
dorsal and ventral protective membranes joined at base of club; separated from stalk by membrane; dorsal membrane
Cephalopods of the World 67
Fig. 116 Sepia apama
dorsal view
(illustrations: K. Hollis/ABRS)c u t t l e b o n e
lateral view ventral viewdorsal view
adult subadult
t e n t a c u l a r c l u b
(illustration: A. Reid)
(after Lu, 1998 )
forms shallow cleft at junction with stalk. Buccal membrane without suckers; in females with single median spermathecae
in ventral part. Eggs spherical. Cuttlebone outline oval, or oblong (juveniles broadly oval, wider in anterior half, becoming
elongate in adults); bone bluntly rounded anteriorly; acuminate, rounded posteriorly; dorsal surface creamy white, evenly
convex posteriorly, flat anteriorly; dorsal surface slightly granulose with irregular longitudinal ridges; indistinct dorsal
median rib present in large specimens, broadens anteriorly, bordered laterally by indistinct grooves; lateral ribs indistinct.
Chitin borders lateral and anterior margins of cuttlebone. Spine present in juveniles, lost in adults. Striated zone flat, or
slightly concave; last loculus flat in large animals, or convex in smaller ones; shallow, wide, sulcus extends along striated
zone only, or sulcus absent. Anterior striae are inverted U-shape in smaller speci mens, becoming straight in large
specimens. Inner cone limbs are narrow anteriorly, broaden slightly posteriorly; in large specimens thickened into rough
callus on inner margin of outer cone, which narrows anteriorly, broadens posteriorly. Three flat, semicircular, flap-like
papillae posterior to eyes. Colour: Reddish brown. All arms have whitish transverse bars and spots bordered by darker
pigment. Dorsal mantle with fine, irregular, reticulate, white bands and spots, some fused to form irregular transverse
bands; bands joined laterally into longitudinal broken white band adjacent to fins. Fins pale, with narrow white band along
outer margin.
Size: Maximum size up to 500 mm mantle length and weight in excess of 10.5 kg. This is the worlds? largest cuttlefish.
Geographical Distribution: Southern Indo-Pacific:
southern Australia from Moreton Bay 27?25'S
153?20'E, possibly as far north as Shoalwater Bay,
Queensland (22?30'S) to Point Cloates, Western
Australia (22?43'S 113?40'E) . Lord Howe and Norfolk
Islands. Australian endemic (Fig. 117).
Habitat and Biology: Depth range from 1 to 100 m.
Sepia apama is a neritic demersal species with cryptic
habits; it occurs in coral areas, seagrass beds and on
open trawl grounds. It feeds on fishes, crabs and other
crustaceans. Dur ing a highly r i tual ized and
stereotyped courtship, males display broad bands of
dark chromatophores that pulsate across the dorsal
mantle as part of their visual display. Males often guard
females to ward off other males. Small males have
been observed to mimic the coloration and behaviour
of females, using this diversion to sneak close to
females to mate with them without the awareness of
the larger aggressive males. Mating takes place
head-to-head and spermatophores are placed in the
female?s spermathecae on the buccal membrane
ventral to the mouth. Southern Australian animals
spawn from April to September. Mass spawning occurs in the Spencer Gulf from April to June, with peak spawning during
May to June. The lemon-shaped eggs are laid in subtidal crevices and hatch after 3 to 5 months. Low incubation
temperatures around 12?C (which mitigat e potential problems of diffusive gas exchanges) may be a factor that limits the
distribution of the species to cool southern waters.
Interest to Fisheries: The species is taken as bycatch in prawn and mixed species trawl fisheries, by hook-and-lines or
speared by divers. It is commonly available in fish markets along the southern coast of Australia, sold for human
consumption and as bait. Few details on quantities or origin of catches are available.
Local Names: AUSTRALIA: Giant Australian cuttlefish.
Remarks: Recent morphological and molecular analyses have shown that specimens of Sepia apama from the east coast
and southern Australia comprise two very divergent populations. However, these populations are not geographically
isolated. The data suggests that populations of S. apama may have been geographically isolated in the past and have
come into secondary contact.
Literature: Adam and Rees (1966), Lu (1998a), Cronin and Seymour (2000), Norman (2000), Kassahn et al. (2003).
68 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 117 Sepia apama
Known distribution
?? 200 m
?
Sepia arabica Massy, 1916 F i g . 1 1 8
Sepia arabica Massy, 1916, Records of the Indian Museum , 12(part 5)(16): 228 [type locality: Laccadive Sea, 11?14'30"N
74?57'15"E and Persian G ulf, 26?20'N 53?54'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Arabian
cuttlefish; Fr ? Seiche d?Arabie;
Sp ? Sepia ar?biga.
Diagnostic Features: Mantle oval. Fins widest in posterior third; anterior origin posterior to mantle margin, posteriorly with
wide gap between them. Head slender, narrower than mantle. Male and female arms subequal in length. Arm suckers
tetraserial in both sexes, suckers small and widely spaced. Arm sucker rims smooth. Hectocotylus present on left ventral
arm: with greatly reduced suckers , much smaller than normal arm suckers; oral surface of modified region wide, swollen,
fleshy, with transversely grooved ridges; suckers in 2 dorsal and 2 ventral series displaced laterally, with gap between
them. Both ventral arms of males with sucker-bearing surfaces folded together and nearly completely covered by the
protective membranes. Club crescent-shaped, small; with 5 or 6 small, similar sized suckers in transverse rows.
Swimming keel of club extends proximally slightly beyond carpus; dorsal and ventral protective membranes not joined at
base of club; dorsal membrane much wider than ventral membrane. Cuttlebone outline lanceolate; bone bluntly rounded
anteriorly, very narrow, strongly tapered posteriorly ; recurved ventrally ; granulose dorsally; dorsal median rib indistinct,
broadens anteriorly. Chitin present as wide bands bordering lateral sides of cuttlebone, extending to median rib. Spine
absent; dorsoposterior end of cuttlebone with short median longitudinal ridge. Striated zone and last loculus convex;
sulcus shallow, narrow, extends entire length of cuttlebone. Anterior striae V-shape ; inner cone lateral limbs are separated
from outer cone by smooth zones; inner cone limbs are uniform width, narrow V-shape posteriorly; slightly raised into
rounded posterior ridge; inner cone posteriorly with irregular ribs radiating into outer cone (ribs calcareous medially and
chitinous toward margin of outer cone); outer cone narrow anteriorly, broadens posteriorly; outer cone limbs are expanded
into 2 rounded ?wings? , directed ventrally, to form a recurved cup-like structure. Dorsal mantle has pale circular tubercles
between median row of dark patches dorsal to base of fins and has a series of elongate papillae along each side, adjacent
Cephalopods of the World 69
Fig. 118 Sepia arabica
dorsal view
t e n t a c u l a r c l u b
c u t t l e b o n e
ventral view enlargement of posteriorend of cuttlebone,
ventral view
lateral viewdorsal view
70 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
to base of each fin. Head with large, fleshy ear-shaped projections posterior to eyes . Colour: Reddish purple. Head with
chromatophores concentrated over eye orbits. Dorsal mantle chromatophores are irregularly distributed in patches; base
of fins on posterior half with 10 to 12 patches of concentrated reddish purple chromatophores.
Size: Up to 88 mm mantle length.
Geographical Distribution: Indian Ocean: Red Sea,
Gulf of Aden, Persian Gulf, western and southern
India, Laccadive Islands (Fig. 119).
Habitat and Biology: Depth range from 80 to 272 m.
Interest to Fisheries: Presently undetermined. The
species has been reported from bottom trawl resource
surveys.
Literature: Adam and Rees (1966), Filippova et al.
(1995), Nateewathana (1996).
Sepia australis Quoy and Gaimard, 1832 F i g . 1 2 0
Sepia australis Quoy and Gaimard, 1832. Voyage de d?couvertes de l ?Astrolabe pendant les annees
1826?1827?1828?1829, Zoologie , 2(1): 70 [Type locality: Agulhas Bank, South Africa].
Frequent Synonyms: Sepia capensis d?Orbigny, 1845; Sepia sinope Gray, 1849.
Misidentifications: None.
FAO Names: En ? Southern cuttlefish;
Fr ? Seiche australe; Sp ? Sepia austral.
Fig. 119 Sepia arabica
Known distribution
?? 200 m
t e n t a c u l a r c l u b
ventral view
Fig. 120 Sepia australis
dorsal view
c u t t l e b o n e
Diagnostic Features: Fins rounded posteriorly, with wide gap between them. Arm suckers tetraserial; median suckers
larger than marginal suckers (more so in males). Hectocotylus present on left ventral arm: 2 rows of normal size suckers
proximally, 6 or 7 rows of greatly reduced suckers medially , then normal size suckers to arm tip; suckers in 2 dorsal and
2 ventral series displaced laterally, with gap between them. Club crescent-shaped, short, with 5 suckers in transverse
rows ; suckers differ markedly in size with 3 greatly enlarged suckers towards proximal end of club. Swimming keel of club
extends proximally slightly beyond carpus; dorsal and ventral protective membranes not joined at base of club .
Cuttlebone outline oval; bone bluntly rounded anteriorly; tapers abruptly posteriorly, acuminate, acute, sharply pointed;
strongly recurved ventrally; dorsal surface creamy white; evenly convex; texture smooth. Spine and posterior end
(approximately one quarter) of bone covered with ochre-coloured, smooth, glaze-like substance. Dorsal median rib
indistinct; broadens slightly anteriorly; ribs bordered laterally by distinct grooves; lateral ribs distinct. Chitin surrounds
entire margin of cuttlebone. Spine long, straight, directed dorsally; dorsoposterior end of cuttlebone with short, median
longitudinal ridge anterior to, and separate from spine. Striated zone and last loculus convex; sulcus deep, narrow,
flanked by rounded ribs , extends entire length of cuttlebone. Anterior striae are inverted U-shape . Inner cone limbs are
uniform width, narrow, U-shape posteriorly; slightly raised into rounded posterior ledge; thickened slightly; dull, not shiny;
outer cone chitinous, not calcified, narrow. Colour: Purplish brown. Fins pale with broad orange-pink band along base
dorsally; ventral pigment present; chromatophores concentrated near fins as for dorsum.
Size: Up to 85 mm mantle length, 50 g total weight.
Geographical Distribution: Southeastern Atlantic and
western Indian Ocean: from Namibia (27?S) to Port
Elizabeth, South Africa (few records east of Port
Elizabeth). Possibly Red Sea, Gulf of Aden and
Somalia. Occurrence in China (as S. sinope Gray
1849), is extremely doubtful (Fig. 121).
Habitat and Biology: Depth range from 45 to 345 m
(most abundant from 60 to 190 m). Sepia australis is
able to thr ive in areas of depleted oxygen
concentrations (to as low as 1.5 ml l-1) as is found in
shallow northern waters of the west coast of southern
Africa; here the best catches are associated with
oxygen concentrations between 1.5 ml l-1 to 3.5 ml l-1
and temperatures around 9?C. Off the south coast of
South Africa, mature animals occur in early winter and
the main spawning grounds are likely to be located in
deeper water on the western side of the Agulhas Bank.
Sepia australis is an important prey item of fur seals,
hake, skates and other groundfishes. It preys mostly
upon stomatopod crustaceans. On the west coast of
South Africa, the largest recorded female measured
85 mm (weight 50 g) and the largest recorded male
was 62 mm (weight 23 g), while on the south coast, the
largest recorded female measured 67 mm (weight 15
g) and the largest recorded male 57 mm (weight 11 g).
Interest to Fisheries: This is one of the most common
sepiid species found off the South African coast. The
importance of the species, defined on the basis of
?survey abundance? (including commercial fisheries)
and its trophic links with other organisms, indicate that
Sepia australis is sufficiently abundant to be exploited
by fisheries. This, along with its tasty flesh, gives it
excellent commercial potential. Unknown factors are
dispersal, availability and market considerations.
Literature: Adam and Rees (1966), Roeleveld
(1972), Lipinski et al. (1991), S?nchez and
Villanueva (1991), Lipinski (1992), Lipinski et al.
(1992a), Lipinski et al. (1992b), Roeleveld et al.
(1993), Augustyn et al. (1995).
Cephalopods of the World 71
Fig. 121 Sepia australis
Known distribution
?? 200 m
?
?
?
?
72 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia bandensis Adam, 1939 F i g . 1 2 2 ; P l a t e I V , 2 1 ? 2 2
Sepia bandensis Adam, 1939b. Bulletin du Mus?e royal d?Histoire naturelle de Belgique , 15(18): 1 [type locality:
Indonesia: Banda Sea, Banda Neira].
Frequent Synonyms: Sepia baxteri (Iredale, 1940) and Sepia bartletti (Iredale, 1954) are possible synonyms.
Misidentifications: None.
FAO Names: En ? Stumpy cuttlefish; Fr ? Seiche trapue; Sp ? Sepia achaparrada.
Diagnostic Features: Club with 5
suckers in transverse rows , central 3
suckers enlarged. Swimming keel
extends beyond base of club. Dorsal
and ventral protective membranes
joined at base of club , separated from
stalk by a membrane. Cuttlebone
outline broad, oval; bone bluntly
rounded anteriorly and posteriorly;
dorsal surface evenly convex; calcified
with reticulate sculpture; dorsal median
and lateral ribs absent. Spine reduced
to tiny, blunt tubercle. Sulcus shallow,
narrow, extends along striated zone
only. Anterior striae are inverted
U-shape . Inner cone limbs are narrow
anteriorly, broaden posteriorly, slightly
convex; outer cone narrow anteriorly,
broadens posteriorly. Dorsal mantle
has longitudinal row of ridge-like
papillae along each side, adjacent to
base of each fin and scattered short
r idges dorsal to cut t lebone
(corresponding to whitish bars). Head
and arms with short, scattered, bar-like
papi l lae posit ioned dorsal ly and
laterally. Colour: Light brown, or
greenish yellow-brown when fresh, and
with whitish mottle. Head with scattered white spots. Dorsal mantle has white blotches concentrated into short, longitudinal
bars. It often shows a pair of brown patches on the posterior end of the mantle, often accompanied by white patches over
the eye orbits. Fins pale with row of small luminescent blue spots at base.
Size: Up to 70 mm mantle length. Animals from the Alas Strait, Indonesia reported at about 40 g (males) and 45 g
(females).
Geographical Distribution:
Tropical Indo-Pacific: Philippines,
Malaysian Borneo, Java,
Sulawesi, Alas Strait, eastern
Indonesia, New Guinea. Possibly
northern Australia and Marshall
Islands (Fig. 123).
Habitat and Biology: Coastal
shallow waters. Found over coral
reefs and sand. Night active. This
species has been observed
?walking?, rather than swimming,
using arms III [or IV] and a pair of
raised flaps on the ventral mantle.
It is often found in association
with sea cucumbers and sea
stars.
Literature: Adam and Rees
(1966), Gofhar (1989).
t e n t a c u l a r c l u b
Fig. 122 Sepia bandensis
ventral viewdorsal view
c u t t l e b o n e
(illustrations: K. Hollis)
Fig. 123 Sepia bandensis
Known distribution
?? 200 m
?
?
Sepia bertheloti d?Orbigny, 1835 ( in F?russac and d?Orbigny 1834?1848) F i g . 1 2 4
Sepia bertheloti d?Orbigny, 1835. Histoire naturelle g?n?rale et particuli? re des C?phalopodes Ac?tabuliferes vivants et
fossiles, pl. 11 [type locality: Canary Islands: Tenerife].
Frequent Synonyms: Sepia verrucosa L?nnberg, 1896; Sepia mercatoris Adam, 1937.
Misidentifications: None.
FAO Names: En ? African cuttlefish; Fr ? Seiche africaine; Sp ? Sepia africana.
Diagnostic Features: Dorsal anterior mantle margin triangular, acute; ventral mantle margin emarginate. Fins wide, with
wide gap between them posteriorly. Arms IV elongate, particularly in males. Arm suckers tetraserial. Male median arm
suckers with greater diameter than marginal ones. Hectocotylus present on left ventral arm: 1 or 2 rows of normal size
suckers proximally, 9 to 13 rows of greatly reduced suckers medially ; suckers in 2 dorsal and 2 ventral series displaced
laterally, with gap between them. Dorsal protective membranes well developed on hectocotylized arm, covering
suckers . Club straight, slender, with 5 or 6 suckers in transverse rows ; suckers differ in size: several suckers of inner 2
or 3 rows larger than rest . Swimming keel of club terminates at posterior end of carpus. Dorsal and ventral protective
membranes not joined at base of club but fused to tentacular stalk. Dorsal and ventral membranes same length,
terminate at posterior end of carpus. Buccal membrane in females extends ventrally with 2 spermathecae. Cuttlebone
outline oblong; acuminate, acute, anteriorly; bluntly rounded posteriorly; dorsal surface evenly convex; entire surface
calcified with reticulate granulose sculpture concentrated medially and on lateral margins in irregular longitudinal ridges;
calcification thickest posteriorly; dorsal median rib indistinct; broadens slightly anteriorly; lateral ribs absent. Chitin borders
lateral and anterior margins of cuttlebone. Spine moderate length, keel(s) absent. Striated zone and last loculus convex;
sulcus shallow, narrow, extends entire length of cuttlebone. Anterior striae shallow m-shape, or inverted U-shape. Inner
cone limbs are narrow, strap-like anteriorly, broaden slightly posteriorly; outer cone chitinous, spatulate, expanded .
Dorsal mantle with series of elongate papillae along each side, medial to base of each fin. Colour: Purplish brown. Arms
have a transverse zebra-stripe pattern and a longitudinal orange-red pigmente d stripe along their aboral surfaces.
Dorsal mantle has small spots (predominantly posteriorly) and narrow, cream-coloured, broken transverse lines. Fins pale
with narrow lustrous orange-pink band al ong base inmales bordered by purplish band and 1 or 2 longitudinal rows of
numerous short bars.
Cephalopods of the World 73
h e c t o c o t y l u s c u t t l e b o n et e n t a c u l a r c l u b
(after Adam, 1952 )
ventral view
Fig. 124 Sepia bertheloti
dorsal view
(after Adam, 1952 )
74 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Size: Males up to 175 mm mantle length; females to
134 mm mantle length.
Geographical Distribution: Eastern Atlantic: off
northwestern Africa, from Canary Islands and off
Western Sahara, to Angola (southwestern Africa),
14?S. Endeavour Bank (Fig. 125).
Habitat and Biology: Depth range from 20 to 156 m.
On the Endeavour Bank, Sepia bertheloti migrates to
shallow waters to spawn during summer and autumn.
The size distribution observed in the captures shows a
larger range for this species than for the co-occurring
S. officinalis Linnaeus, 1758 in summer.
Interest to Fisheries: Sepia bertheloti is trawled off
the Canary Islands, where greatest concentrations are
encountered between 70 and 140 m depth. It is
commonly captured off the western Endeavour Bank,
where it shares the same depth distribution with the
co-occurring species S. hierredda Rang, 1835.
Separate statistics are difficult to obtain because the
two species are often marketed together, but
information from the Spanish fishery in Saharan
waters indicate that it represented 11% of cuttlefishes
captured over the last decade, with S. hierredda
comprising the remaining 89%. It is also a sizeable
component of cephalopod catches in Mauritania
waters, where it represented about 35% of the
cuttlefishes captured in recent years (the remaining
65% again made up by S. hierredda). Off Senegal, it is
reported to represent a minor portion of the total
cuttlefish catch taken by trawls. It is marketed fresh or
deep-frozen for export.
Remarks: Sexual dimorphism is pronounced: the mantle is relatively wider in females and the arms are much longer in
males, especially the ventral pair. The soft parts are very similar to those of S. officinalis, although the ventral arms of the
males are much longer and the largest tentacular suckers relatively smaller. However, the cuttlebone differs markedly
between the two species. Recent genetic studies show the distinctiveness of this species in relation to S. officinalis and
S. hierredda.
Literature: Adam and Rees (1966), Delgado de Molina et al. (1993), Hernandez-Garcia and Castro (1994).
Fig. 125 Sepia bertheloti
Known distribution
?? 200 m
Cephalopods of the World 75
Sepia braggi Verco, 1907 F i g . 1 2 6
Sepia braggi Verco, 1907, Transactions of the Royal Society of South Australia , 31: 213 [type locality: South Australia,
Glenelg, 34?58'S 138?32'E].
Frequent Synonyms: None.
Misidentifications: Sepia bidhaia Reid, 2000; Sepia cottoni Adam, 1979; Sepia limata (Iredale, 1926); Sepia rhoda
(Iredale, 1954); Sepia vercoi Adam, 1979.
FAO Names: En ? Slender cuttlefish; Fr ? Seiche gracile; Sp ? Sepia gr?cil.
Diagnostic Features: Dorsal anterior mantle margin triangular, acute. Fins widest in posterior third, rounded posteriorly,
with wide gap between them. Head short, slender, narrower than mantle. Male and female arms differ in relative lengths;
male (mature) arms III elongate; female arms II and III longer than arms I and IV (arms II usually longer than arms III).
Distal arm tips strongly attenuate in both sexes, suckers enclosed by protective membranes. Arm suckers tetraserial
proximally, biserial at distal arm tips (biserial region over greater proportion of arms II and III than of arms I and IV);
biserial suckers reduced and displaced laterally, with gap between them on arms II of females and arms III of males.
Hectocotylus absent. Club slightly recurved, short, sucker-bearing surface flattened, with 4 to 6 suckers in transverse
rows ; suckers differ in size: 5 or 6 median suckers twice diameter of rest. Swimming keel of club extends proximally slightly
beyond carpus; dorsal and ventral protective membranes not joined at base of club but fused to tentacular stalk. Dorsal
and ventral membranes same length, extend proximal to carpus along stalk. Dorsal membrane broad, much wider than
ventral membrane and forms shallow cleft at junction with stalk. Gills with 26 to 30 lamellae per demibranch. Buccal
membrane without suckers. Buccal membrane in females without spermathecae. Spermatophores 2.4 to 4.4 mm long;
smallest male recorded with well-developed spermatophores in spermatophoric sac was 24.1 mm mantle length. Eggs
oval, up to 9.4 mm long, 5.5 mm wide. Cuttlebone outline lanceolate, broadest in anterior third at anterior end of striated
zone; acuminate, acute, anteriorly and posteriorly; strongly recurved ventrally; dorsal surface pinkish; evenly convex;
calcified medially, slightly granulose with irregular longitudinal ridges (particularly posteriorly). Spine and posterior end
(approximately one quarter) of bone covered with ochre-coloured, smooth, glaze-like substance. Dorsal median rib distinct
anteriorly, indistinct posteriorly; rib broadens anteriorly; lateral ribs absent. Chitin present as wide bands bordering lateral
margins of cuttlebone (covers about half of dorsal surface of cuttlebone). Spine short, pointed, curves dorsally; keel(s)
absent; fine, radiating ribs between outer cone and spine. Striated zone and last loculus convex; sulcus deep, narrow,
dorsal view lateral view ventral view
c u t t l e b o n e
(illustrations: K. Hollis)
Fig. 126 Sepia braggi
dorsal view
t e n t a c u l a r c l u b
(illustration: A. Reid)
76 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
flanked by rounded ribs, extends entire length of cuttlebone. Anterior striae are inverted U-shape, incurved medially,
slightly V-shape following sulcus . Inner cone limbs are strap-like anteriorly, narrower posteriorly, U-shape; slightly raised
into rounded posterior ridge; thickened, shiny (yellowish), inner cone posteriorly with irregular calcareous ribs radiating into
outer cone; outer cone calcified, narrow, limbs are expanded posteriorly into 2 short ?wings? , directed ventrally, to form a
recurved cup-like structure. Dorsal mantle has scattered short ridges dorsal to cuttlebone and up to five orange-pink ridges
at base of each fin. Head with short, scattered papillose bars posterior to eyes; arm papillae present; same as those on
head. Colour: In preserved specimens pale buff to pinkish brown. Arms I have a darkly pigmented purplish band on each
side of dorsal midline. Arms II to IV have a median longitudinal purplish band along their aboral surfaces. All arms
have short transverse bars and blotches lateral to these longitudinal bands. Arm and club sucker rims midbrown. Fins
without markings along base.
Size: Males up to 49 mm mantle length; females up to
80 mm mantle length.
Geographical distribution: Southern Indo-Pacific:
southern Australia from southern New South Wales,
36?23'S 150?07'E, to Wester n Australia, 31?51'S 115?
35'E. Australian endemic (Fig. 127).
Habitat and Biology: Depth range from 30 to 86 m.
Sepia braggi is a demersal species. A report of many
fresh bones washed up on Gunnamatta Beach
(Victoria) 26?27 May 2001, is perhaps is indicative of a
spawning event.
Interest to Fisheries: Taken as bycatch with other
cuttlefishes along southern Australia, then used locally
as bait and food.
Remarks: The pigmentation on the arms is very
distinctive and clearly visible even in small specimens.
While arms III are obviously elongate in adult males,
they are similar in length to the remaining arms in
immature males (less than approximately 30 mm
mantle length). Sepia braggi males differ from S.
limata Iredale, 1926 and S. rhoda (Iredale, 1954)
males in having elongate third arms. Sepia braggi does not have a hectocotylus; the left ventral arms are hectocotylized in
S. limata and S. rhoda. The arrangement of the arm suckers differs among these three species. Female S. rhoda have
elongate second arms. While the second and third arms of S. braggi females are slightly longer than the other arms, they
do not differ to the same extent as those of mature S. rhoda females. The anterior striae on cuttlebones are M-shape in S.
rhoda and the posterior inner cone is very broad, while in S. braggi the anterior striae are an inverted U-shape and the
inner cone narrow. The cuttlebone is broader in S. limata than the S. braggi cuttlebone. Sepia braggi of both sexes differ
from all other Australian narrow-boned species in its distinctive arm pigmentation.
Literature: Adam and Rees (1966), Lu (1998a), Reid (2000).
Fig. 127 Sepia braggi
Known distribution
?? 200 m
Sepia brevimana Steenstrup, 1875 F i g . 1 2 8
Sepia brevimana Steenstrup, 1875, Danske Videnskabernes Selskabs Skrifter, 5 Raekke, Naturvidenskabelig og
Mathematisk , 10(7): 475 [type locality: Indian Ocean].
Frequent Synonyms: Sepia rostrata (in part) F?russac and d?Orbigny, 1848.
Misidentifications: Sepia esculenta Hoyle, 1885; Sepia
stellifera Homenko and Khromov, 1984.
FAO Names: En ? Shortclub cuttlefish; Fr ? Seiche petites
mains; Sp ? Sepia mazicorta.
Diagnostic Features: Mantle broad, oval; dorsal anterior
margin triangular, acute. Arm suckers tetraserial. Club
small, with 6 to 8 suckers in transverse rows ; all club
suckers of similar small size. Swimming keel of club extends
well proximal to carpus; dorsal and ventral protective
membranes not joined at base of club; dorsal membrane
much wider than ventral membrane . Cuttlebone outline
oval; bone very angular, V-shape anteriorly ; dorsal surface
flat, texture rough, with irregular calcified projections. Spine
long, pointed, with dorsal and ventral keels. Sulcus shallow,
narrow, extends along striated zone only, flanked by rounded
ribs. Anterior striae are inverted blunt V-shape. Inner cone
limbs broaden posteriorly, thickened, rose-coloured or
yellowish orange ; outer margin of inner cone raised into a
flattened ledge posteriorly; outer cone calcified, narrow
anteriorly, broadens posteriorly.
Size: Up to 110 mm mantle length.
Geographical Distribution: Indo-Pacific:
northern Indian Ocean, Andaman Sea,
Andaman and Maldive Islands, Singapore to the
Gulf of Tonkin, South China Sea, Hong Kong,
Java, Sulu and Celebes Seas. Possibly
Saya-de-Malha Bank (Fig. 129).
Habitat and Biology: Depth range from 10 to
100 m. Sepia brevimana is a shelf species,
restricted to coastal waters. Around India,
spawning occurs throughout the year. Several
spawning peaks have been observed between
July and February in eastern Indian waters.
Hatchlings grow to adult size in 11 to 13 months,
depending on temperature conditions. Off
Chennai (Madras), animals attain 29 to 34 mm
within 6 months, 56 to 58 mm at the end of 12
months and about 75 mm after 18 months.
Length at 50% maturity for males is 56 mm off
Madras and 62 mm off Waltair, and for females,
59 and 63 mm, respectively. Maximum length in
the Indian trawl fishery is 85 mm off Madras, and
95 mm off Waltair (northeast India), with common size ranges from about 40 to 70 mm. In the Gulf of Thailand, 40 to 60 mm
mantle length animals are caught commonly, with the maximum size attained 90 mm mantle length. The sex ratio of males
to females caught is 1:2.
Interest to Fisheries: This species appears as a bycatch in the eastern Indian trawl fishery off Chennai and Waltair, but
separate catch data are not reported. Sepia brevimana is important to the commercial fishery of Thailand. Most
cuttlefishes in this area are caught using otter trawl, some using pair trawl, and small catches are made using squid
light-lures, traps and push nets; bottom otter and pair trawls are used offshore, while push nets and lift nets are used in
inshore and coastal waters. It is also fished in the South China Sea.
Remarks: This species may be confused with S. stellifera Homenko and Khromov, 1984, but differs in having 6 to 8, rather
than 10, oblique rows of suckers on the club. The sulcus of the cuttlebone in S. brevimana is shallow and narrow. In
S. stellifera, the sulcus is deep and wide. Sepia brevimana is similar to S. esculenta Hoyle, 1885, but differs in having fewer
Cephalopods of the World 77
Fig. 128 Sepia brevimana
ventral view
t e n t a c u l a r c l u b c u t t l e b o n e
60?
60?
70?
70?
80?
80?
90?
90?
100?
100?
110?
110?
120?
120?
 10?  10?
0? 0?
10? 10?
20? 20?
Fig. 129 Sepia brevimana
Known distribution
?? 200 m
?
?
78 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
club suckers (there are 10 to 16 suckers in transverse rows in S. esculenta), no fleshy papillae along the base of the fins,
and by the coloured inner cone.
Literature: Adam and Rees (1966), Voss and Williamson (1971), Silas et al. (1982), Silas et al. (1986), Chantawong and
Suksawat (1997).
Sepia cultrata Hoyle, 1885 F i g . 1 3 0
Sepia cultrata Hoyle, 1885, Annals and Magazine of Natural History , (series 5)16: 198 [type locality: Australia, Twofold
Bay, 36?59'S 150?20'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Knifebone cuttlefish;
Fr ? Seiche ? os en couteau; Sp ? Sepia de
sepi?n de cuchillo.
Diagnostic Features: Male and female
arm lengths subequal. Arm suckers
tetraserial. Hectocotylus present on left
ventral arm: approximately 7 rows of
normal size suckers proximally ; 5 or 6
ro w s of greatly reduced suc kers
medially. Suckers of hectocotylus in 2
dorsal series are smaller than those in 2
ventral series (in some specimens, 2
dorsal rows appear merged as if in a single
row); oral surface of modified region
swollen; suckers in 2 dorsal and 2 ventral
series displaced laterally, with gap between
them. Club crescent-shaped, sucker-
bearing surface flattened, with 5 or 6 small
suckers in transverse rows. Swimming
keel of club extends well proximal to
carpus; dorsal and ventral protective
membranes not joined at base of club ;
dorsal and ventral membranes differ in length ; dorsal membrane extends proximal to carpus along stalk , ventral
membrane terminates at proximal end of carpus . Buccal membrane without suckers. Cuttlebone outline oval; bone
triangular, obtuse anteriorly; acuminate, acute, posteriorly; dorsal surface pinkish ; dorsal surface convex posteriorly, flat
anteriorly; some granulation on lateroposterior margins bordered by a longitudinal ridge; dorsal median rib present; rib
broadens anteriorly; lateral ribs indistinct. Spine straight. Striated zone concave in extreme posterior part, slightly
convex in anterior part ; last loculus slightly concave; sulcus absent. Anterior striae slightly convex to straight . Inner
cone limbs are uniform width, very narrow, U-shape posteriorly ; thickened. Dorsal mantle with scattered short papillae
along each side close to fins and with longitudinal row of ridge-like papillae adjacent to base of each fin; ventral mantle with
longitudinal row of 4 or 5 narrow ridges along each side close to fins. Colour: Pale buff pinkish brown.
Size: Up to 120 mm mantle length.
Geographic Distribution: Southern Indo-Pacific:
southern Australia from southern Queensland,
26?35'S 153?45'E, southwestward to Western
Australia, Houtman Abrolhos, 28?49'S 114?04'E,
including Tasmania, 42?43'S 148?22'E (Fig. 131).
Habitat and Biology: Depth range from 132 to 803 m
with the majority of catches from 300 to 500 m; outer
shelf and upper bathyl.
Interest to Fisheries: Species taken as bycatch of
prawn and mixed species trawl fisheries. Possibly
more than one species is identified under this name.
This requires further investigation.
Literature: Adam and Rees (1966), Lu (1998a).
Fig. 130 Sepia cultrata
dorsal view
t e n t a c u l a r c l u b
ventral view
c u t t l e b o n e
Fig. 131 Sepia cultrata
Known distribution
?? 200 m
Sepia elegans Blainville, 1827 F i g . 1 3 2
Sepia elegans Blainville, 1827, Dictionnaire des Sciences Naturelles , 48: 284 [type locality: Sicily ( S. elegans );
Mediterranean Sea (as S. biserialis and S. italica ); Island of Noirmoutiers and La Rochelle (as S. rupellaria ).
Frequent Synonyms: Sepia rupellaria F?russac and d?Orbigny, 1835?1848; Sepia biserialis Blainville, 1827; Sepia
italica Risso, 1854.
Misidentifications: Sepia bertheloti d?Orbigny, 1835.
FAO Names: En ? Elegant cuttlefish; Fr ? Seiche ?l?gante; Sp ? Sepia elegante.
Diagnostic Features: Small species. Mantle oblong; dorsal anterior margin triangular, acute; ventral mantle margin
emarginate. Male and female arms subequal in length. Arm sucker arrangement differs between sexes: in males, arm I
suckers biserial for a few rows, rest tetraserial ; arms II and III suckers tetraserial proximally , biserial at extreme distal
tips; arm IV suckers variously arranged (approximately 10 pairs biserial suckers on arms I to III, arms IV with 2 to 4 rows
biserial suckers); in females, suckers biserial proximally, tetraserial distally (5 rows biserial suckers arms I to III, 2 to 4 rows
biserial suckers arms IV). Male non-hectocotylized median arm suckers with greater diameter than marginal ones.
Hectocotylus present on left ventral arm: 1 or 2 rows of normal size suckers proximally, 9 to 11 rows of reduced minute
suckers medially , then normal size suckers to arm tip; suckers in 2 dorsal and 2 ventral series displaced laterally. Club
short, oval; sucker-bearing surface flattened, with 6 to 8 suckers in transverse rows ; suckers differ markedly in size: 3 or 4
greatly enlarged suckers toward posterior end of club and several dorsal suckers enlarged, but not as large as medial
suckers. Buccal membrane in females with single median spermathecae in ventral part. Cuttlebone outline oblong; convex
in lateral view; acuminate, acute, anteriorly and posteriorly; recurved ventrally; dorsal surface evenly convex; last loculus
convex. Anterior striae are inverted U-shape. Inner cone limbs are uniform width, narrow V-shape posteriorly ; outer cone
narrow throughout; outer cone limbs are expanded posteriorly into 2 curved ?wings? , directed ventrally, to form a recurved
cup-like structure. Spine very short . Dorsoposterior end of cuttlebone with short, rugose, calcareous keel. Ventral mantle
with longitudinal row of 6 narrow ridges along each side close to fins; anteriormost pair and posterior 2 pairs shorter than
rest. Colour: Reddish brown . Head with a few scattered chromatophores. Arms without markings. Dorsal mantle pale,
peppered with scattered purple-black chromatophores. Fins and ventral mantle pale. Ridges whitish.
Cephalopods of the World 79
dorsal viewFig. 132 Sepia elegans
t e n t a c u l a r c l u b c u t t l e b o n e
ventral view
(after Adam, 1952 )
Size: Males up to 72 mm mantle length; females up to
89 mm mantle length. Total weight between 50 and 60 g.
Geographical Distribution: Eastern Atlantic and
Mediterranean Sea: around British Isles, western
Scotland, Ireland, Dingle Bay, Co. Kerry and the English
Channel, from 50?N through the Mediterranean
(including Ligurian Sea, Tyrrhenian Sea, Aegean Sea,
Sea of Marmara and Levantine Sea), west Africa:
Endeavour Bank to 15?S, and possibly Agulhas Bank
(Fig. 133).
Habitat and Biology: Depth to 500 m, although records
below 450 m are sporadic. Sepia elegans is a sublittoral
species. It spends the winter in deep water (200?400 m),
then migrates into the shallows in spring and summer to
spawn. It feeds mainly on molluscs, small crustaceans,
fishes and polychaetes. In the Sea of Marmara
(Mediterranean Sea), this species has been found in
brackish waters (salinity between 18 and 25?), which
indicates a high degree of to lerance. In the
Mediterranean Sea, mature males and females are
present throughout the year, which suggests a
continuous spawning period. As a consequence of this,
a continuous recruitment also occurs, with observed
peaks in several Mediterranean areas. Smallest mature
males have been observed at 20 mm, and females at
30 mm mantle length. The eggs are laid in clusters of 12
to 25 (diameter 5 mm) attached to alcyonarians (sea
fans), shells, etc., on muddy bottoms. After hatching,
juveniles immediately adopt a benthic lifestyle. A rough
estimate of growth in mantle length of 2.8 mm per month
for males and 3.0 mm for females is reported for animals
in the Sici l ian Channel. A negative allometric
length?weight relationship has been found for both
sexes in all seasons and different locations, with females
heavier than males at any given mantle length. Females
also have longer tentacular clubs and heavier stomach
contents than males. Off west Africa, spawning extends
almost throughout the year in shallow inshore waters,
with peaks in summer and autumn. Maturity is attained
at about 1 year of age. Males may carry about 95
spermatophores and females about 250 eggs. Spawning
occurs at temperatures of 13? to 18?C. In Portuguese
waters mature animals are also present throughout the
year, but with minimum numbers in summer. The
lifespan of this species is about 12 to 18 months.
Interest to Fisheries: Sepia elegans is taken mainly as a bycatch in Mediterranean and west African trawl fisheries. It is
most abundant at about 150 m depth, hence a little deeper than S. officinalis Linnaeus, 1758 and S. bertheloti d?Orbigny,
1835. Separate statistics are not reported for this species, which, however, represent a very significant percentage of the
catches in some areas of its distributional range. In the Mediterranean Sea it is marketed along with S. orbignyana
F?russac in d?Orbigny, 1826 and small S. officinalis and constitutes a valuable resource locally. In the Sicilian Channel,
research studies showed an exploitation rate of 0.73 for the species, which suggests a very intense fishing pressure on this
resource. It is marketed fresh and frozen.
Local Names: ITALY: Seppia elegante, Castagnola; SPAIN: Casta?o.
Remarks: This species resembles S. australis Quoy and Gaimard, 1832, but differs in the arrangement of arm suckers,
club and hectocotylus. Sepia elegans differs from S. officinalis in having a reddish, rather than brown, coloration and
because it generally inhabits deeper water and does not bury in the sand during the day.
Literature: Mangold-Wirz (1963), Adam and Rees (1966), Guerra and Castro (1989), Bello (1990a), Jereb and Ragonese
(1991), Ragonese and Jereb (1991), Wurtz et al. (1991), Guerra (1992), Filippova et al. (1995), Sanjuan et al. (1996),
Neige and Boletzky (1997), Sanchez et al. (1998), Belcari (1999a), Unsal et al. (1999), Salman et al. (2002), Sobral (2002).
80 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 133 Sepia elegans
Known distribution
?? 200 m
?
Sepia elliptica Hoyle, 1885 F i g . 1 3 4
Sepia elliptica Hoyle, 1885, Annals and Magazine of Natural History , (series 5)16: 189 [type locality: Arafura Sea, south of
Papua, 09?59'S 139?42'E, and 08?56'S 136?05'E].
Frequent Synonyms: None.
Misidentifications: Sepia esculenta Hoyle, 1885; Sepia madokai Adam, 1939a; Sepia
stellifera Homenko and Khromov, 1984.
FAO Names: En ? Ovalbone cuttlefish;
Fr ? Seiche ? sepion ovale; Sp ? Sepia de
sepi?n oval.
Diagnostic Features: Mantle oval;
dorsal anterior margin triangular, acute.
Male and female arm lengths subequal;
protective membranes narrow. Arm
suckers tetraserial. Hectocotylus present
on left ventral arm: 7 or 8 rows of normal
size suckers proximally ; 7 rows of
reduced suckers medially , then normal
size suckers distally to arm tip. Suckers of
hectocotylus in 2 dorsal series are
smaller than ventral reduced suckers ;
reduced suckers only slightly smaller than
normal arm suckers; oral surface of
modified region not wide, fleshy, but
normal, as on opposite arm. Club
sucker-bearing surface flattened, with 10
to 12 minute suckers in transverse rows ;
suckers all similar size. Swimming keel of
club extends well proximal to carpus;
dorsal and ventral protective membranes
joined at base of club; dorsal membrane
forms shallow cleft at junction with stalk. Cuttlebone outline oval ; bone very angular, V-shape anteriorly ; bluntly rounded
posteriorly; dorsal surface creamy white; dorsal surface evenly convex; texture smooth, not pustulose; dorsal median rib
indistinct, broadens anteriorly; lateral ribs indistinct. Chitin surrounds entire margin of cuttlebone. Spine short, pointed,
curves dorsally, keel(s) absent. Striated zone concave; last loculus convex; sulcus deep, wide. Anterior striae are inverted
U-shape . Inner cone limbs are narrow anteriorly, broaden posteriorly; outer margin of inner cone raised into flat posterior
ledge; ledge whitish (sometimes with a thin rim of chitin
on outer margin); ledge not thickened ; outer cone
calcified.
Size: Up to 175 mm mantle length.
Geographical Distribution: Tropical Indo-Pacific:
northern Australia from Western Australia, Exmouth
Gulf, 22?23'S 114?06'E, to Queensland, Capricorn
Island Group, 23?30'S 152?00'E, including Gulf of
Carpentaria. Viet Nam and India (doubtful records)
(Fig. 135)
Habitat and Biology: Depth range from 16 to 142 m.
Sepia elliptica occurs mainly in coastal waters. This
species was collected between 10 and 62 m in Gulf of
Carpentaria surveys. A broad size range of both sexes
was found to occur in the Gulf over most of the year,
indicative of an extended spawning season. Several
experimental studies were carried out on growth of this
species in captivity; results showed that both water
temperature and feeding levels affect somatic growth
but with different effects on the muscular tissue
development and protein composition. No direct
relationship was found between individual growth rates
and food availability.
Cephalopods of the World 81
c u t t l e b o n e
Fig. 134 Sepia elliptica
t e n t a c u l a r c l u b h e c t o c o t y l u s
(after Hoyle, 1886 )
dorsal ventral
subadult
ventral view
Fig. 135 Sepia elliptica
Known distribution
?? 200 m
? ?
82 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Interest to Fisheries: Sepia elliptica is taken as bycatch of prawn and mixed species trawl fisheries in most of its
distributional area. The observed population structure in the Gulf of Carpentaria (Australia) by experimental trawl surveys
and the relative high catch rates of trawlers in some areas suggest that S. elliptica is a promising resource for fishery there.
Remarks: This species may be confused with S. esculenta Hoyle, 1885 but it can be distinguished by the following
characters: the hectocotylus of S. esculenta has 5 or 6 series of normal size-suckers, followed by 6 series of reduced
suckers; in S. esculenta the dorsal and ventral protective membranes are not joined at the base of the club; in S. esculenta,
the cuttlebone is bluntly rounded anteriorly, the striae are inverted V-shape and the inner cone limbs are thickened
posteriorly. Duc (1978, 1993) refers to S. elliptica in Vietnamese waters; however, this probably is a misidentification of
S. esculenta. The occurrence of S. elliptica in Indian waters is yet to be confirmed.
Literature: Adam and Rees (1966), Dunning et al. (1994), Lu (1998a), Moltschaniwskyj and Martinez (1998), Martinez
et al. (2000), Moltschaniwskyj and Jackson (2000).
Sepia elobyana Adam, 1941 F i g . 1 3 6
Sepia elobyana Adam, 1941a, Memoires du Mus?e royal d?Histoire naturelle de Belgique , (2)21: 121 [type locality: Africa:
west Africa, Gulf of Guinea, Eloby Island, 01?01'N 09?29'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Guinean cuttlefish; Fr ? Seiche de Guin?e; Sp ? Sepia guineana.
Fig. 136 Sepia elobyana
dorsal view
(after Adam, 1952 )
h e c t o c o t y l u s tentacular club
c u t t l e b o n e
dorsal view ventral view
Diagnostic Features: Small species. Mantle broad, oval; ventral mantle margin emarginate, with distinct lateral angles.
Fins wide. Male and female arms subequal in length; protective membranes wide, well developed, particularly on modified
portions of arms I and II in males. Distal tips of arms I and II attenuate in both sexes , with about 20 rows of globular suckers
in males. Arm sucker arrangement differs between sexes: in males, arms I to III suckers tetraserial proximally, biserial
distally; in females, arms I and II suckers tetraserial proximally, biserial distally . Male median arm suckers larger than
dorsal and ventral marginal series on proximal end of arms. Hectocotylus present; both ventral arms modified . Left
ventral arm with 7 or 8 rows of greatly reduced suckers proximally, suckers normal size distally with suckers in 2 dorsal
series smaller than 2 ventral rows on modified portion of arm. Suckers in 2 dorsal and 2 ventral series displaced laterally,
with gap between, 2 ventral series close together, rows alternate, dorsal and ventral marginal suckers at base of protective
membranes. Male right ventral arm suckers normal proximally for 2 transverse rows, followed distally by 4 or 5 rows of
transformed suckers: median and somet imes lateral sucker rings wide (coverin g whole diameter of sucker) and thick ;
teeth and infundibulum absent from these suckers. Club with 8 suckers in transverse rows ; suckers all of similar, small,
size. Cuttlebone outline oval; acuminate, acute, anteriorly; bluntly rounded posteriorly; dorsal surface evenly convex;
calcified with reticulate sculpture; dorsal median rib and lateral ribs absent. Spine a blunt knob . Striated zone separated
from outer cone by broad, smooth, marginal zones; sulcus shallow, wide, bordered by some irregular grooves, extends
along striated zone only. Anterior striae are inverted U-shape (slightly irregular due to grooves); limbs of inner cone extend
anteriorly to end of striated zone. Inner cone limbs are narrow anteriorly, broaden posteriorly. Dorsal mantle has scattered
tubercles and reticulate pattern of ridges.
Size: Up to 53 mm mantle length.
Geographical Distribution: Eastern Atlantic: along
west African coast from Senegal to Gulf of Guinea and
Gabon (southern limit undetermined) (Fig. 137).
Habitat and Biology: Unknown.
Interest to Fisheries: Possibly taken with other
cuttlefishes off west Africa, but not distinguished from
similar species.
Remarks: Very rarely reported species.
Literature: Adam and Rees (1966).
Cephalopods of the World 83
Fig. 137 Sepia elobyana
Known distribution
?? 200 m
84 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia esculenta Hoyle, 1885 F i g . 1 3 8
Sepia esculenta Hoyle, 1885, Annals and Magazine of Natural History , (series 5)16: 188 [type locality: Japan].
Frequent Synonyms: None.
Misidentifications: Sepia elliptica Hoyle, 1885; S. hoylei Ortmann, 1888.
FAO Names: En ? Golden cuttlefish; Fr ? Seiche dor?e; Sp ? Sepia dorada.
Diagnostic Features: Mantle broadly oval; dorsal anterior margin triangular, obtuse; ventral mantle margin emarginate.
Arm suckers tetraserial. Hectocotylus present on left ventral arm: 5 or 6 rows of normal size suckers proximally, 6 rows
of reduced suckers medially , then normal size suckers to arm tip. Club sucker-bearing surface flattened, with 10 to 16
suckers in transverse rows ; all club suckers of similar, minute, size (except for 2 enlarged suckers in anteriodorsal
corner). Dorsal and ventral protective membranes not joined at base of club, extend proximal to carpus along stalk. Buccal
membrane without suckers. Cuttlebone outline elliptical; bone bluntly rounded anteriorly and posteriorly; dorsal surface
convex posteriorly, flat anteriorly; dorsal median and lateral ribs indistinct. Chitin present as wide patch posteriorly and a
narrow chitinous rim borders lateral margins of cuttlebone. Spine present. Striated zone concave; last loculus flat; sulcus
deep, wide . Anterior striae are inverted V-shape . Inner cone limbs are narrow anteriorly, br oaden posteriorly; raised into
flat, thickened posterior ledge ; outer cone calcified, narrow anteriorly, broadens posteriorly. Dorsal mantle with
longitudinal row of 6 or 7 yellowish, fleshy tubercles at base of each fin. Colour: Light brown with whitish mottle. Dorsal
mantle has faint light-coloured wavy transverse stripes and dark spots or blotches, sometimes studded with yellowish
tubercles. Arms I to III have spots and a longitudinal orange-red pigmented stripe along their aboral surfaces. Fins with
pale golden iridescent line along base, both dorsally and ventrally.
Size: Up to 180 mm mantle length, 600 g total weight.
Fig. 138 Sepia esculenta
c u t t l e b o n et e n t a c u l a r c l u b
(after Okutani et al., 1987 )
(after Okutani et al., 1987 )
ventral view
dorsal view
Geographical Distribution: Indo-Pacific: East China
Sea, Japan from central Honshu (Sea of Japan and
Pacific sides) south to China and Hong Kong, Taiwan
Province of China, South China Sea (north of central
Philippines), Viet Nam and possibly Singapore and
western Indonesia. The southernmost extent of its
range is yet to be determined (Fig. 139).
Habitat and Biology: Depth range from 10 to 100 m,
mainly inner shelf. This is a demersal neritic species
found on sand, sometimes burrowing into the
substrate. After overwintering in deeper water, animals
migrate into shallower coastal waters where they
spawn when the water temperature increases in
spring. Males often guard females to ward off
competing males. Before mating begins, the brilliance
of the body colours of males increases dramatically,
with coloration changing rhythmically from green to red
and gold and the iridescent lines along the base of the
fins appear to be fluorescent. Mating takes place
head-to-head and spermatophores are placed in the
female?s buccal membrane ventral to the mouth. The
eggs are of an amber colour immediately after laying,
covered by a pear-shaped egg case, milky white and
translucent. They are deposited on macrophytic algae
and other substrates, such as sunken branches. The
sticky, clear, outermost layer of the egg cases
accumulates sand and other debris for camouflage as
the embryonic development proceeds. Studies in
captivity indicate that females prefer long, fine, and
immobile materials as spawning substrates. The
spawning season extends from spring to early
summer. Hatching occurs after about 30 to 80 days, depending on the water temperature. Newly hatched animals already
have schooling and benthic tendencies and bury themselves in the sand. Sepia esculenta feeds mainly on crustaceans and
small fishes.
Interest to Fisheries: Sepia esculenta supports localized and subsistence fisheries in the Philippines, with animals
collected in waters including the Visayan and Samar Seas, the Lingayen Gulf and Carigara Bay. It is the dominant Sepia
species landed around the Shantung and Kiangsu provinces of China and off western Japan (East China Sea). Studies off
Korea determined that maximum fishing conditions were characterized by a temperature range from 10? to 15?C and
bottom salinity between 33.2 and 34.45?. The principa l capture techniques are otter trawls, pound nets, hoop nets and
hook-and-line. The flesh is highly prized as food, especially in Japan, South Korea and China. In Japan, large-sized
animals are consumed as sashimi, while smaller animals are marketed packed and frozen ready for cooking. This species
has been reared to market size in experiments under laboratory conditions, at growth rates well above those in natural
populations.
Local Names: CHINA: Gam woo chak, Jam mak yue; JAPAN: Hariika, Kouika, Maika, Sumiika.
Remarks: Sepia esculenta can be confused with S. elliptica Hoyle, 1885. In S. esculenta, the inner cone ledge is thick and
directed anterioventrally. In S. elliptica, the ledge is much thinner, flatter and it points anteriorly, covering the posterior end
of the phragmocone. The cuttlebone striae of S. elliptica are U-shaped and the bone is very angular, V-shaped anteriorly.
The hectocotylus of S. elliptica differs in having 7 or 8 series of normal sized suckers, followed by 7 series of reduced
suckers.
Literature: Adam and Rees (1966), Choe (1966), Baik and Park (1985), Fujita et al. (1997), Okutani et al. (1987), Natsukari
and Tashiro (1991), Watanuki et al. (1993).
Cephalopods of the World 85
Fig. 139 Sepia esculenta
Known distribution
?? 200 m?
?
Sepia grahami Reid, 2001 F i g . 1 4 0
Sepia grahami Reid, 2001b, Proceedings of the
Linnean Society of New South Wales , 123: 160 [type
locality: Australia: New South Wales, 29?33'S
153?25'E to 29?32' S 153?25'E].
Frequent Synonyms: None.
Misidentifications: Sepia mestus Gray, 1849; Sepia
rozella (Iredale, 1926).
FAO Names: En ? Ken?s cuttlefish; Fr ? Seiche de
Ken; Sp ? Sepia de Ken.
Diagnostic Features: Mantle oval. Fins wide,
rounded posteriorly. Male and female arms subequal
in length; protective membranes narrow; distal arm
tips not attenuate. Arm suckers tetraserial.
Hectocotylus absent. Club short, slightly recurved,
sucker-bearing surface flattened, with 4 or 5 suckers
in transverse rows (usually 4, rarely 5) ; suckers
differ only slightly in size: 3 or 4 suckers slightly
enlarged. Swimming keel of club extends well
proximal to carpus; dorsal and ventral protective
membranes not joined at base of club; dorsal
membrane extends proximal to carpus along stalk,
ventral membrane terminates at proximal end of
carpus. Gills with 23 to 26 lamellae per demibranch.
Cuttlebone oval; dorsal surface creamy white, with
slight pinkish tinge; dorsal median rib absent, lateral
ribs indistinct. Spine with ventral keel . Striated zone
concave, sulcus shallow, narrow, indistinct .
Anterior striae are inverted U-shape ; last loculus flat.
Inner cone limbs are narrow strap-like anteriorly,
broaden posteriorly. Inner cone limbs are raised into
rounded ledge posteriorly, ledge thickened,
yellowish or ochre coloured . Dorsal mantle with
longitudinal rows of up to 6 ridges along each side,
close to fins; prominent ear-shape lobe dorsal to eye
and 2 lobes ventral to eye, anteriormost lobe largest.
Colour : Pinkish brown. Mantle with dorsal
?eyespots? .
Size: Males up to 66 mm mantle length;
females up to 82 mm mantle length.
Geographical Distribution: Southwestern
Pacific: Australia, New South Wales, from
southeast of Yamba area (29?32' 153?25'E) to
off Tathra (36?44'S 150?05'E) (Fig. 141).
Habitat and Biology: Depth range from 2 to
84 m.
Interest to Fisheries: There is no official
information on this species but it is commonly
confused with S. rozella (Iredale, 1926) and
S. mestus Gray, 1849. Sepia rozella commonly
appears in fish markets in Sydney.
Literature: Reid (2001b).
86 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 140 Sepia grahami
t e n t a c u l a r c l u b c u t t l e b o n e
ventral view
Fig. 141 Sepia grahami
Known distribution
?? 200 m
Cephalopods of the World 87
Sepia hedleyi Berry, 1918 F i g . 1 4 2
Sepia hedleyi Berry, 1918, Biological Results of the Fishing Experiments carried on by the F.I.S. ?Endeavour? 1909?14,
4(5): 258 [type locality: South Australia, Investigator St rait area (35?25'S 137?22'E), s outh of Kangaroo Island (35?50'S
137?15'E)].
Frequent Synonyms: Sepia dannevigi Berry, 1918; Decorisepia
rex Iredale, 1926.
Misidentifications: Sepia vossi Khromov, 1996 (as S. rex ).
FAO Names: En ? Hedley?s
cuttlefish; Fr ? Seiche d?Hedley;
Sp ? Sepia de Hedley.
Diagnostic Features: Mantle
broad, oval . Fins widest in
poster ior th i rd, rounded
posteriorly, with narrow gap
between them. Head short ,
broad, narrower than mantle.
Male and female arms subequal
in length; protective membranes
narrow. Arm suckers tetraserial;
male non-hectocotylized arm
suckers smaller than female arm
suckers in diameter. Hectocotylus
present on left ventral arm: 6 to 8
rows of normal size suckers
proximally, 9 or 10 rows of
reduced suckers medially .
Suckers of hectocotylus in 2
dorsal series are much smaller
than those in 2 ventral series ;
oral surface of modified region
wide, swol len, f leshy, wi th
transversely grooved ridges; 2 dorsal and 2 ventral series of suckers displaced laterally, with gap between; suckers in 2
ventral series aligned in a single row . Club crescent-shaped, moderate length, sucker-bearing surface flattened, with 9 to
12 suckers in transverse rows; all club suckers of similar, small, size. Swimming keel of club extends well proximal to
carpus; dorsal and ventral protective membranes not joined at base of club but are fused to tentacular stalk. Dorsal and
ventral membranes same length; extend proximal to carpus along stalk. Gills with 29 or 30 lamellae per demibranch.
Buccal membrane without suckers; in females extends ventrally with 2 spermathecae. Spermatophores 5.8 to 7.9 mm
long. Eggs spherical, 2.5 to 3.1 mm diameter. Cuttlebone outline oblong; acuminate, acute, anteriorly and posteriorly;
dorsal surface creamy white; dorsal surface evenly convex; granulose; spine and posterior tip of bone covered with smooth
glaze-like substance. Dorsal median rib distinct; sides approximately parallel; ribs bordered laterally by distinct grooves;
lateral ribs indistinct. Chitin surrounds entire margin of cuttlebone. Spine short, pointed, straight, parallel to bone, keel(s)
absent. Striated zone flat; last loculus convex; sulcus shallow, narrow, extends entire length of cuttlebone. Anterior striae
are inverted U-shape . Inner cone limbs are uniform width, narrow V-shape posteriorly, thickened, shiny; outer cone
calcified, narrow anteriorly, broadens posteriorly.
Dorsal mantle has longitudinal row of up to 6 ridge-like
papillae along each side, adjacent to base of each fin.
Colour: Buff pinkish brown. Arms without markings.
Dorsal ridges orange-pink in colour.
Size: Males up to 83 mm mantle length; females up to
108 mm mantle length.
Geographical Distribution: Southern Indo-Pacific:
Australia, Queensland, from off the Great Barrier Reef,
22?35.3'S 153?46.7'E, sout hwards around southern
Australia to Western Australia, southwest of Shark Bay,
27?07'S 112?49'E. Austra lian endemic (Fig. 143).
Habitat and Biology: Depth range from 47 to 1 092 m.
Interest to Fisheries: Sepia hedleyi is taken as bycatch
of prawn and mixed species trawl fisheries.
Literature: Lu (1998a), Reid (2001a).
Fig. 142 Sepia hedleyi
ventral view
c u t t l e b o n e
dorsal view
t e n t a c u l a r c l u b
Fig. 143 Sepia hedleyi
Known distribution
?? 200 m
88 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia hierredda Rang, 1835 F i g . 1 4 4
Sepia hierredda Rang, 1835, (in F?russac and d?Orbigny 1834?1848), Histoire naturelle g?n?rale et particuli?re des
C?phalopodes Ac?tabuliferes vivants et fossiles , pl. 13 [type locality: western Africa].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Giant African
cuttlefish; Fr ? Seiche g?ante
africaine; Sp ? Sepia gigante
africana.
Diagnostic Features: Hectocotylus
present on left ventral arm: 6 rows
of normal size suckers proximally,
8 to 14 rows of reduced suckers
distally. Club long, slightly recurved,
with 5 or 6 suckers in transverse
rows ; suckers differ in size: 5 or 6
median suckers twice diameter of
rest; club swimming keel extends
slightly proximal to sucker-bearing
surface. Cuttlebone outline oblong,
lateral margins concave in anterior
third; acuminate, acute, anteriorly;
bluntly rounded posteriorly; dorsal
median rib present, broadens
anteriorly; median rib bordered
laterally by distinct grooves; lateral
ribs present. Spine present. Anterior
striae shallow m-shape . Inner cone
limbs are narrow anteriorly, broaden
posteriorly; outer cone broadens
posteriorly. Colour: Pattern very
similar to that of Sepia officinalis.
Size: Up to nearly 500 mm mantle length; total weight over
7 500 g.
Geographical Distribution: Southeastern Atlantic: Africa,
Cape Blanc (21?N), Mauritania (19?N) to Tigres Bay, Angola
(16?30'S). Endeavour Bank (Fig. 145).
Habitat and Biology: Spawning occurs from February to
September and the lifespan is 24 months. Migrations of
S. hierredda occur off the west African coast. Females
mature at 130 mm mantle length. In those areas where the
distribution of S. hierredda overlaps that of S. officinalis, the
former species tends to be restricted to waters shallower
than 50 m, while the distribution of S. officinalis extends to
depths in excess of 100 m.
Interest to Fisheries: Sepia hierredda is the most
commercially important cuttlefish in the east central Atlantic
waters (from Cape Blanc, 21?N, to Cape Bojador, 26?N). It
represents the dominant cuttlefish caught off Western
Sahara and in Mauritania waters. Specific statistics are not
available, but S. hierredda made up 90 and 65%,
respectively, of the cuttlefishes caught in these areas by the
Spanish fishery in recent years.
Fig. 145 Sepia hierredda
Known distribution
?? 200 m
Fig. 144 Sepia hierredda
ventral view
c u t t l e b o n e
dorsal view
t e n t a c u l a r c l u b
(after Adam, 1952 )
Remarks: Sepia hierredda often was considered to be a subspecies of S. officinalis Linnaeus, 1758. However, the results
of recent morphological and allozyme analyses strongly support the fact that S. officinalis and S. hierredda are different
species. The 2 species can be distinguished as follows: the number of transverse rows of suckers is higher in S. hierredda
than in S. officinalis; for animals with the same mantle length, the length of the striated zone on the cuttlebone is shorter in
S. officinalis from the northeastern Atlantic than in S. hierredda (for animals from the Canary Islands (central eastern
Atlantic), however, this character is not useful since the striated zone is of the same length in both species); the mantle is
narrower and the arms are shorter in S. hierredda than in S. officinalis. The 2 species also differ at 13 allozyme loci. Sepia
hierredda and S. officinalis are sympatric off the northwestern coast of Africa.
Literature: Hatanaka (1979), Bakhayokho (1983), Khromov et al. (1998), Guerra et al. (2001).
Sepia kobiensis Hoyle, 1885 F i g . 1 4 6
Sepia kobiensis Hoyle, 1885, Annals and Magazine of Natural History , (series 5)16: 195 [type locality: Japan: Yokohama
market (as var. S. andreanoides ) and Bay of Kobe for S. kobiensis ; Toyama Bay (as var. toyamensis); Beppu, Oita
Prefecture (as var. beppuana); Enoura, Shizuoka Prefecture (as var. crassa); Korea Strait (as var. albatrossi)].
Frequent Synonyms: Sepia andreanoides Hoyle, 1885.
Misidentifications: Sepia andreana Steenstrup, 1875.
FAO Names: En ? Kobi cuttlefish; Fr ? Seiche kobi; Sp ? Sepia kobi.
Cephalopods of the World 89
Fig. 146 Sepia kobiensis
ventral view
c u t t l e b o n e
dorsal view
t e n t a c u l a r c l u b
90 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Diagnostic Features: Small species. Mantle oblong; dorsal anterior margin triangular, acute. Male and female arm lengths
subequal, short. Arm suckers tetraserial, median rows larger than marginal rows; suckers displaced laterally, with gap
between on arms I to III . Hectocotylus present on left ventral arm: 6 to 12 rows of normal size suckers proximally, 7 to 10
rows of greatly reduced suckers medially , then normal size suckers to arm tip; oral surface of modified region wide,
swollen, fleshy, with transversely g rooved ridges and deep median furrow ; suckers in 2 dorsal and 2 ventral series
displaced laterally, with gap between them. Club crescent-shaped, narrow; with 4 or 5 suckers in transverse rows ;
suckers differ slightly in size: 4 or 5 suckers toward proximal end of club slightly larger than others. Swimming keel of club
extends proximally slightly beyond carpus; dorsal and ventral protective membranes not joined at base of club. Cuttlebone
outline lanceolate; acuminate anteriorly and posteriorly; dorsal surface pinkish, or yellowish; entire surface calcified with
very fine granulose sculpture partly arranged in irregular longitudinal ridges and bordered laterally by 2 narrow calcareous
ridges; dorsal median rib indistinct, broadens anteriorly. Chitin (trace only) borders lateral margins of cuttlebone. Spine
long, pointed, directed dorsally. Striated zone and last loculus convex; sulcus shallow, narrow, extends entire length of
cuttlebone. Anterior striae are inverted m-shape . Inner cone limbs are uniform width, narrow, U-shape posteriorly,
thickened; outer cone calcified; limbs are expanded posteriorly into 2 short ?wings? , directed ventrally, to form a recurved
cup-like structure. Colour: Reddish brown. Head with V-shape reddish stripe on dorsal margins of eye orbits and with
orange stripes extending fromposterior end of head to basal portions of arms I to III. Dorsal mantle has reddish spots.
Size: Up to 90 mm mantle length (usually 70 mm mantle length) and 80 g total weight.
Geographical Distribution:
Indo-Pacific: off Japan,
f rom southern part of
Hokkaido, south to Kyushu
(both coasts), Yellow Sea,
East China Sea, Taiwan
Province of China, Gulf of
Tonkin, South China Sea,
Phi l ippines, Gul f of
Thailand and the northern
part of the Indian Ocean
from the Persian Gulf,
Arabian Sea and Bay of
Bengal, Myanmar. Possibly
the Banda Sea (Fig. 147).
Habitat and Biology: Depth
range subtidal to 200 m.
Interest to Fisheries: Sepia
kobiensis is taken in small
quantities between 80 and
160 m in the Hong Kong
area, and it is also taken as
bycatch in small- scale
fisheries off southern Japan
and in the Inland Sea, mainly with fixed nets, trawls and beach seines. It is an object of fisheries in China where it is
abundant in shallow waters and locally utilized.
Local Names: JAPAN: Hime-kouika.
Remarks: Sepia kobiensis is probably a species complex. This requires investigation, particularly considering its
significance in fisheries. This species has been confused with S. andreana Steenstrup, 1875, from which it is distinguished
by several characters. All arms in adult S. kobiensis are similar in length, while in adult male S. andreana, the second pair
of arms is greatly elongate and bears biserial, rather than tetraserial suckers. The hectocotylus sucker arrangement in
Sepia kobiensis follows a pattern of normal, reduced, then normal-sized suckers from the proximal to distal end of the arm,
while in S. andreana, some normal-sized suckers occur at the base of the hectocotylus, and the rest are reduced and very
rudimentary. The club suckers differ markedly in size in S. andreana, while those of S. kobiensis differ only slightly in size.
Literature: Adam and Rees (1966), Okutani et al. (1987).
Fig. 147 Sepia kobiensis
Known distribution
?? 200 m
?
Sepia latimanus Quoy and Gaimard, 1832 F i g . 1 4 8 ; P l a t e I V , 2 3 ? 2 7
Sepia latimanus Quoy and Gaimard, 1832, Voyage de d?couvertes de l?Astrolabe pendant les annees
1826?1827?1828?1829, Zoologie , 2(1): 68 [type locality: New Guinea, Port Dorey, 00?51'S 134?01'E].
Frequent Synonyms: Ponderisepia eclogaria Iredale, 1926; Sepia harmeri Robson, 1928; Sepia rappiana F?russac,
1834 in F?russac and d?Orbigny (1834?1848); Sepia mozambica Rochebrune, 1884; Sepia hercules Pilsbry, 1894.
Misidentifications: None.
FAO Names: En ? Broadclub cuttlefish; Fr ? Seiche grandes mains; Sp ? Sepia mazuda.
Diagnostic Features: Mantle oval. Arm suckers tetraserial. Hectocotylus absent. Club crescent-shaped, sucker-bearing
surface flattened, with 5 or 6 suckers in transverse rows ; suckers differ markedly in size: some greatly enlarged.
Swimming keel of club extends well proximal to carpus; dorsal and ventral protective membranes joined at base of club ;
separated from stalk bymembrane ; dorsal membrane forms shallow cleft at junction with stalk. Buccal membrane without
suckers. Cuttlebone bluntly rounded anteriorly and posteriorly; dorsal surface convex posteriorly, flat anteriorly; granulose
(texture loosely arranged along growth lines); dorsal median rib and lateral ribs indistinct, or absent. Spine short, pointed,
stout, keels absent. Striated zone concave; last loculus convex; sulcus shallow, narrow , extends entire length of
cuttlebone. Anterior striae are inverted V-shape . Inner cone limbs are uniform width, narrow, U-shape posteriorly;
thickened into a rounded, shiny ridge; outer cone calcified, narrow anteriorly, broadens posteriorly, deep cup-like. Dorsal
mantle covered with numerous large papillae and with series of elongate papillae along each side, adjacent to base of each
fin. Head papillose. Colour: Light brown, yellowish or dark brown (dark in males during courtship and breeding), with
whitish mottle, blotches or spots. Arms have longitudinal white bands on their margins that appear as broad white
blotches when the arms are extended, and arms I to III have broad, longitudinal brownish bands medially, extending onto
head. Dorsal mantle has transverse saddle mark, small white and brown spots, narrow brown transverse bands, and bold,
white, transverse stripes and spots dorsally (spots and blotches are present in most colour patterns; saddle-mark shown
only occasionally; transverse stripes shown by males during the breeding season). In live animals, the eyes are typically
yellow around the ventral margins. Fins pale with white, transverse stripes extending onto mantle and narrow, white, band
along outer margin.
Cephalopods of the World 91
dorsal view
Fig. 148 Sepia latimanus
c u t t l e b o n e
(illustration: K. Hollis)
t e n t a c u l a r c l u b
(after Okutani et al., 1987 )
ventral view
Size: Up to 500 mm mantle length and
weight to 10 kg.
Geographical Distribution: Indo-Pacific:
from southern Mozambique, throughout
the periphery of the Indian Ocean,
Malacca Strait, Melanesian Islands,
South China Sea, Philippine Sea and
East China Sea, Taiwan Province of
China and Japan to southern Kyushu.
Indonesia to northwestern and
northeastern Australia (from Western
Australia, Shark Bay, 25?25'S 113?35'E,
across northern Australian waters to
southern Great Barrier Reef, about 23?S,
Queensland), the Coral Sea, Palau,
Guam, New Caledonia, Fiji and Cocos
(Keel ing) Is lands. Records from
Madagascar and southeastern Australia
doubtful (Fig. 149).
Habitat and Biology: Depth to 30 m,
weight to 10 kg. This shallow water
species inhabits tropical coral reefs.
Mating occurs on the west coast of Guam and off Okinawa in shallow water (30 m) from January to May, and the eggs hatch
in 38 to 40 days. In the Alas Strait, Indonesia, males grow up to 170 mm (500 g), females 240 mm (1.3 kg) mantle length.
Sepia latimanus is day active. During hunting it appears to mesmerize prey by displaying rhythmic colour bands along the
body. It feeds on fishes and crustaceans. During the breeding season, males establish a territory, defending a coral head
(typically Porites ) among which females lay eggs following mating. Courtship is highly ritualized and stereotyped, and it
incorporates striking visual displays. Males often guard females to ward off other males. Mating takes place head-to-head
and spermatophores are placed in the females? buccal membrane ventral to the mouth. The eggs harden after laying and
are very difficult to extract from the coral. They hatch after 4 to 6 weeks, and the young immediately hide among the coral
and coral rubble. Juveniles often mimic mangrove leaves, their coloration and posture complete with the stem, ribs and
scattered black spots.
Interest to Fisheries: Sepia latimanus is an important fisheries species throughout its range, taken by trawl, setnet, jig,
handline and spear. It supports local fisheries in western Japan and the Philippines where it is caught with jigs, handlines,
setnets and spears and it is common as bycatch in southeast Asian trawl fisheries. Sepia latimanus is fished in small
quantities in the Ryukyus, China, near Taiwan Province of China and in the waters of Indo-China. In the Philippines, the
large cuttlefishes, S. pharaonis Ehrenberg, 1831 and S. latimanus, are split open dorsally, the cuttlebone and viscera
removed, and the animals are dried in the sun without salt.
Local Names: INDONESIA: Sotong-besar; JAPAN: Kobushime, Kubushime.
Remarks: Sepia latimanus probably comprises a species complex. This requires investigation, particularly considering its
fisheries significance. Khromov (1996) suggests that Duc?s (1978) record of S. hercules Pilsbry, 1894 (a junior synonym of
S. latimanus) from Viet Nam actually is S. pharaonis.
Literature: Adam and Rees (1966), Okutani et al. (1987), Hanlon and Messenger (1996).
92 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 149 Sepia latimanus
Known distribution
?? 200 m
Sepia longipes Sasaki, 1913 F i g . 1 5 0
Sepia longipes Sasaki, 1913, Doubutsugaku Zasshi [Zoological Magazine, Tokyo] 25(292): 78 [type locality: Japan:
Choshi, Ibaraki Prefecture (Chiba Peninsula)].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Longarm cuttlefish; Fr ? Seiche pieuvre; Sp ? Sepia brazolargo.
Diagnostic Features: Dorsal anterior mantle margin triangular, acute; ventral mantle margin weakly convex. Fins wide,
with very narrow gap between them. Male arms I greatly elongate, attenuate, whip-like, about twice as long as other
arms, and arms II slightly elongate (relative lengths greater in large specimens); arms I with biserial suckers on
attenuated section. Distal part of arms I thickened, bordered with long, broad trabeculae; protective membranes
demarcated from proximal part by strong constriction; pedicels enlarged on distal portion of arm. Female arm lengths
subequal (mostly shorter than mantle length). In males, suckers on arms I tetraserial proximally, biserial distally, arms II
to IV tetraserial. Female arm suckers tetraserial. Sucker rings mainly smooth. Hectocotylus present; both ventral arms
modified. Left ventral arm: sucker size normal proximally, distal 40% of arm length with reduced suckers ; distal 40%
furrowed longitudinally. Right ventral arm: suckers reduced distally after fourteenth row (not as much as on the left); oral
surface of modified region (both ventral arms) wide, swollen, fleshy, with transversely grooved ridges. Club with 3 or 4
suckers in transverse rows ; suckers differ markedly in size: 4 or 5 greatly enlarged suckers in longitudinal series towards
posterior end of club. Cuttlebone outline lanceolate; bone bluntly rounded anteriorly; acuminate posteriorly; dorsal surface
pinkish, granulose; dorsal median rib distinct, continuing to longitudinal groove on last loculus, rib broadens anteriorly.
Spine moderate length, pointed, spindle-shape, pale pinkish. Striated zone convex, with a high median ridge; sulcus
shallow, narrow, extends entire length of cuttlebone; sulcus flanked by rounded ribs bor dered laterally by shallow
grooves . Anterior striae shallow m-shape . Inner cone limbs are uniform width, narrow V-shape posteriorly; slightly raised
into rounded, shiny posterior ridge; outer cone chitinous laterally, calcareous in expanded posterior part; outer cone limbs
are expanded posteriorly into 2 short ?wings? , directed ventrally, to form a recurved cup-like structure. Colour: Pale
brownish or purplish brown. Arms I to III have a longitudinal orange-red pigmente d stripe along their aboral surfaces.
Dorsal mantle has chromatophores concentrated medially above cuttlebone and scattered reddish spots.
Cephalopods of the World 93
Fig. 150 Sepia longipes
ventral viewdorsal view
c u t t l e b o n et e n t a c u l a r c l u b dorsal view, male
94 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Size: Up to 250 mm mantle length; 1 kg total weight.
Geographical Distribution: Northwestern Pacific:
Japan, Chiba Peninsula to southern Kyushu and East
China Sea (Fig. 151).
Habitat and Biology: Depth range from 100 to 300 m.
Interest to Fisheries: Occasionally trawled in
southwestern Japan.
Local Names: JAPAN: Tenaga-kouika.
Remarks: This species is similar to S. lorigera W?lker,
1910, but differs in having a broader body, the first arm
pair attenuate, whip-like and the cuttlebone with a high
median ridge and a moderate spine.
Literature: Okutani et al. (1987).
Sepia lorigera W?lker, 1910 F i g . 1 5 2
Sepia lorigera W?lker, 1910, Abhandlungen der mathematische-physikalische Klasse der
K?eniglich Bayerischen Akademie der Wissenschaften , 3(Suppl. 1): 12 [type locality: Japan:
Misaki].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Spider cuttlefish; Fr ? Seiche araign?e; Sp ? Sepia loriga.
Diagnostic Features: Mantle oval (width about 40% of mantle length); dorsal anterior margin
triangular, acute. Fins narrow, equal in width throughout. Male arms I greatly elongate (more
than 1 1/2 times longer than mantle length, 3 times as long as other arms); arms I narrow
proximally, flared distally, p rotective membranes broad ; middle portion of arm I slender
with enlarged protective membrane supported by trabeculae. Female arm lengths subequal.
Non-hectocotylized arm sucker arrangement same in both sexes: suckers on arms I to III
tetraserial proximally, biserial at distal tips ; arms IV suckers tetraserial (in males, distal
suckers arms I and II rudimentary, minute, on swollen pedicels). Hectocotylus present on left
ventral arm: sucker size normal proximally, suckers reduced in size on distal 1/3 and sucker
pedicels swollen . Club with 4 suckers in transverse rows; suckers differ markedly in size : 3
or 4 extremely large suckers surrounded by several moderately large suckers. Swimming
keel of club extends proximally slightly beyond carpus. Cuttlebone outline lanceolate;
acuminate, acute, anteriorly and posteriorly; dorsal surface pinkish; dorsal median rib
indistinct, sides approximately parallel, bordered by grooves. Spine long, pointed. Anterior
striae shallow m-shape . Inner cone limbs are uniform width, narrow, U-shape posteriorly;
slightly raised into wide, V-shape, slightly thickened ridge; outer cone limbs are expanded
posteriorly into 2 short ?wings?, directed ventra lly, to form small recurved cup-like structure.
Colour: Reddish brown. Head with scattered red spots and chromatophores concentrated
medially and over eye orbits. Arms have reddish spots. Ventral mantle pale, with narrow
longitudinal bands of brownish pigment w ith curved teardrop-shape ends on each side,
adjacent to fins .
Size: Up to 250 mm mantle length; 1 kg total weight.
Fig. 151 Sepia longipes
Known distribution
?? 200 m
Fig. 152 Sepia lorigera
ventral view
c u t t l e b o n e
Geographical Distribution: Northwestern Pacific:
southern and southwestern Japan from Sagami Bay to
the East China Sea via the Pacific coast of Shikoku.
South China Sea, Viet Nam (Fig. 153).
Habitat and Biology: Depth range from 100 to 300 m.
Interest to Fisheries: Object of minor fishery,
occasionally caught by trawlers off southwestern
Japan.
Local Names: JAPAN: Usubeni-kouika.
Remarks: The relative lengths of the attenuated arms
increase with the size of the animal. This species is
similar to the Australian species S. tanybracheia Reid,
2000, but in S. tanybracheia the dorsal arms are not as
proportionally elongate and the enlarged club suckers
are not as large as those of S. lorigera. The distinctive
ventral pigment bands are not present in the Australian
species.
Literature: Okutani et al. (1987).
Sepia lycidas Gray, 1849 F i g . 1 5 4
Sepia lycidas Gray, 1849. Catalogue of the Mollusca in the British Museum. Part I. Cephalopoda Antepedia , 103 [type
locality: Guangzhou (China)].
Frequent Synonyms: Sepia subaculeata Sasaki, 1914.
Misidentifications: None.
FAO Names: En ? Kisslip cuttlefish; Fr ? Seiche baisers;
Sp ? Sepia labiada.
Cephalopods of the World 95
Fig. 153 Sepia lorigera
Known distribution
?? 200 m
Fig. 154 Sepia lycidas
c u t t l e b o n et e n t a c u l a r c l u b dorsal view
(after Okutani et al., 1987 )
ventral view
Diagnostic Features: Mantle elliptical; dorsal anterior margin triangular, acute; ventral mantle margin emarginate. Fins
end posteriorly in lobes, with only a small slit between them. Arm suckers tetraserial. Hectocotylus present on left ventral
arm: 6 rows of normal size suckers proximally, 4 rows of reduced suckers distally. Club with 8 suckers in transverse rows
(more than 200 in number), all club suckers of similar, small size. Swimming keel of club shorter than carpus, extends from
proximal 1/3 to distal tip of club. Dorsal and ventral protective membranes not joined at base of club , extend as
membranous ridges along entire stalk. Buccal membrane with a few, minute suckers (single sucker on most lobes).
Cuttlebone outline oblong; bone bluntly rounded anteriorly and posteriorly; dorsal surface evenly convex; dorsal median rib
absent. Chitin present as wide patch posteriorly and a narrow chitinous rim borders lateral margins of cuttlebone. Spine
short, pointed. Sulcus deep, wide . Anterior striae are inverted V-shape . Inner cone limbs are thickened posteriorly,
broaden into a rounded ridge; outer cone calcified; narrow anteriorly, broadens posteriorly. Colour: Reddish brown or
purple. Dorsal mantle has scattered ocellate patches and narrow, irregular, light-coloured, transverse stripes (pattern
more prominent in males than females). Wide stripe adjacent to fins.
Size: Up to 380 mm mantle length. Maximum weight 5 kg.
Geographical Distribution : Indo-Paci f ic :
southwestern Japan, south of Boso Peninsula from
southern Honshu and Chingtao, Korea, to East China
Sea, Taiwan Province of China and South China Sea,
Philippine Sea, Viet Nam and Borneo. In the Gulf of
Thailand it occurs south of 10?N, but never appears in
the inner and eastern coasts. Commonly distributed in
the Andaman Sea. Western limit of range unknown
(Fig. 155).
Habitat and Biology: Sepia lycidas is a neritic
demersal species with a depth range of 15 to 100 m. In
the Gulf of Thailand and the Andaman Sea, most
animals are caught between 20 and 40 m. The sex
ratio of male to female animals caught in the Gulf of
Thailand is 1:2. Sepia lycidas spends the winter at
around 60 to 100 m, then moves into shallow waters to
spawn in spring and early summer. In the South China
Sea, it is abundant between 60 and 100 m depth in the
prespawning period (November?February), then it
migrates inshore to spawn in depths of 15 to 30 m from
March to May. Immediately after laying, the eggs are
shaped like hen eggs, and are amber coloured, like
those of S. esculenta Hoyle, 1885. The egg case is
pear-shaped and one of its pointed ends is bifurcated,
to attach the egg to small twigs and other solid
substrates. The newly hatched young individuals
already show the adult benthic tendencies and bury
themselves into the sand, but no schooling behaviour
has been observed.
Interest to Fisheries: Sepia lycidas is an important commercial species in Japan, China, South Korea, Viet Nam and
Thailand. Most cuttlefishes are caught off Thailand using otter trawl, with smaller catches made using pair trawl and to a
lesser extent, squid light-lures, traps and push nets; bottom otter and pair trawls are used offshore, while push nets and lift
nets are used in inshore and coastal waters. This species is the secondmost important commercial cuttlefish in Hong Kong
and Japan, caught as bycatch in trawls and with setnets and jigs, or by using live cuttlefishes as lures during the spawning
season, and by hook, baited with live prawns or crabs, in other seasons. The mantle flesh is thick and tasty and is,
therefore, highly esteemed. The species has been reared successfully in aquaculture experiments under the name Sepia
subaculeata. Research indicates that waste materials obtained during the processing of this species have potential in
supplementing the skin of land vertebrates as a source of collagen.
Local Names: CHINA: Fa gai na, Mak gung, Yi muk woo chak; JAPAN: Kaminari-ika, Mongouika.
Literature: Adam and Rees (1966), Choe (1966), Voss and Williamson (1971), Okutani et al. (1987), Natsukari and Tashiro
(1991), Nagai et al. (2001).
96 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 155 Sepia lycidas
Known distribution
?? 200 m
?
?
Sepia madokai Adam, 1939 F i g . 1 5 6
Sepia madokai Adam, 1939a, Siboga - Expeditie. Resultats des expeditiones zoologi ques, botaniques,
oceanographiques et entreprises aux Indes Neerlandaises Orientales en 1899?1900 , 55b: 77 [type locality: Japan, Tokyo
Bay and Kagoshima].
Frequent Synonyms: Sepia robsoni Sasaki, 1929 (non S. robsoni
Massy, 1927).
Misidentifications: Sepia acuminata Smith, 1916; Sepia hedleyi
Berry, 1918.
FAO Names: En ? Madokai?s cuttlefish; Fr ? Seiche madokai;
Sp ? Sepia madokai.
Diagnostic Features: Mantle elliptical; dorsal anterior margin
triangular, acute; ventral mantle margin emarginate, without distinct
lateral angles. Hectocotylus present on left ventral arm: sucker size
normal proximally, 10 rows reduced suckers medially , then normal
size suckers to arm tip. Suckers of hectocotylus in 2 dorsal series are
smaller than those in 2 ventral series . Club crescent-shaped,
sucker-bearing surface flattened, with 8 to 10 equal-sized, small
suckers in transverse rows . Cuttlebone length approximately equal
to mantle length; outline oblong, nearly rhomboidal; acuminate,
acute, anteriorly; bluntly rounded posteriorly; dorsal surface pinkish;
dorsal median and lateral ribs indistinct. Spine present, keel(s)
absent. Anterior striae are inverted U-shape . Inner cone limbs are of
uniform width, narrow, U-shape posteriorly , thickened, very
slender; not raised into ledge posteriorly; outer cone calcified, narrow
anteriorly, broadens posteriorly. Colour: Pale brownish. Dorsal
mantle has white blotches or spots.
Size: Up to 100 mm mantle length, 100 g total weight.
Geographical Distribution: Northwestern Pacific:
southwestern Japan, south of Tokyo Bay on the Pacific
Coast and south of Noto Peninsula on the Japan Sea
side to Kyushu. Tsushima Strait, East and South China
Seas, Taiwan Province of China and Viet Nam
(Fig. 157).
Habitat and Biology: Depth range from 20 to 200 m.
Sepia madokai is a demersal species, most common in
bays. It is believed that in years characterized by
intensification of warm currents and high temperatures
in coastal waters this thermophilic cuttlefish may
migrate to areas in the northern Sea of Japan.
Interest to Fisheries: This species is common in the
inland sea area of Japan, where it is fished with bottom
drift nets and trawls. Due to its small size, it has limited
commercial value. It often is caught with Sepia
esculenta, particularly in inland seas and bays off
Japan. It is a minor object of fisheries in Japan and
China (e.g. Peter the Great Bay).
Local Names: JAPAN: Kouika-modoki, Hari-ika.
Remarks: Sepia madokai has been confused with
S. acuminata Smith, 1916 and as S. rex (a synonym of
S. hedleyi Berry, 1918). In S. madokai, the reduced suckers of the hectocotylus differ in size, with those of the 2 dorsal rows
much smaller than those of the 2 ventral rows. In S. hedleyi all reduced hectocotylus suckers are similar in size. The club
has 8 oblique rows of suckers in S. madokai, rather than 9 to 12 as seen in S. hedleyi. The cuttlebone is bluntly rounded
posteriorly, the dorsal median ribs faint and the inner cone limbs broaden posteriorly in S. madokai; in S. hedleyi, the
cuttlebone is acuminate posteriorly, the median ribs are distinct, and the inner cone limbs are uniform width along the
length of the cuttlebone.
Literature: Adam and Rees (1966), Okutani et al. (1987), Shevtsov (1996), Kubodera and Yamada (1998), Lu (1998b).
Cephalopods of the World 97
Fig. 156 Sepia madokai
ventral view
(after Okutani et al., 1987 )
t e n t a c u l a r c l u b
110?
110?
120?
120?
130?
130?
140?
140?
10? 10?
20? 20?
30? 30?
40? 40?
Fig. 157 Sepia madokai
Known distribution
?? 200 m
98 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia murrayi Adam and Rees, 1966 F i g . 1 5 8
Sepia murrayi Adam and Rees, 1966, John Murray Expedition 1933? 3 4, Scientific Reports , 11(1): 63 [type locality: Gulf of
Oman, 25?35'00"N 56?42'18"E t o 25?43'00"N 56?39'18"E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Frog cuttlefish; Fr ? Seiche
grenouille; Sp ? Sepia rana.
Diagnostic Features: Mantle oblong; dorsal anterior
margin triangular, acute. Fins wide, ending in lobes
posteriorly, with a narrow gap between them. Protective
membranes in both sexes wide, well developed. Distal
arm tips bluntly pointed in females, suckers enclosed by
protective membranes. In females, suckers on arms I and
II biserial, suckers arms III and IV tetraserial proximally,
biserial distally (arms III may be biserial throughout);
female distal biserial suckers minute and displaced
laterally, w ith distinct gap between . Club
crescent-shaped, short, curved posteriorly; with 5 suckers
in transverse rows ; all club suckers of similar, small,
size . Swimming keel of club extends proximally slightly
beyond carpus; dorsal and ventral protective membranes
not joined at base of club but fused to tentacular stalk,
extend proximal to carpus along stalk as narrow ridges;
dorsal membrane much wider than ventral membrane
(nearly as wide as sucker-bearing surface). Cuttlebone
outline lanceolate; strongly recurved ventrally; entire
surface calcified with reticulate granulose sculpture
concentrated posteriolaterally and posteriorly in irregular longitudinal ridges; dorsal median rib indistinct, flanked by shallow
grooves; lateral ribs distinct. Chitin borders lateral margins of cuttlebone. Spine straight, directed dorsally, keels absent. Sulcus
shallow, narrow, flanked by rounded ribs, extends entire length of cuttlebone. Anterior striae shallow m-shape ; inner cone
lateral limbs are separated from outer cone by smooth zones. Inner cone limbs are narrow anteriorly, broaden posteriorly,
U-shape, raised into flat posterior V-shape ledge ; inner cone posteriorly with irregular calcareous ribs radiating into outer cone;
outer cone calcified, narrow, limbs are expanded posteriorly into 2 very short ?wings?, directed ventrally, to form a recurved
cup-like structure.
Size: Up to 41 mm mantle length.
Geographical Distribution: Northwestern Indian Ocean:
Gulf of Oman, Gulf of Aden, Somalia (Fig. 159).
Habitat and Biology: A neritic demersal species; depth
range still undetermined (the only record is 106 m).
Interest to Fisheries: Reported from a bottom trawl
resource survey in the Gulf of Aden; its relevance to
artisanal or industrial fisheries is undetermined.
Remarks: The males of this species are unknown. Sepia
murrayi differs from S. kobiensis Hoyle, 1885 in having
subequal rather than variously-sized suckers. The dorsal
calcareous granules are also more pronounced in
S. kobiensis. The inner cone does not have a ventral
ledge in S. kobiensis and the bone lacks the radiating
calcareous ribs on the outer cone. The cuttlebone is
similar to S. burnupi Hoyle, 1904 and S. trygonina
(Rochebrune, 1884), but in S. murrayi the striae differ
and the limbs of the inner cone are more divergent and
with much sharper edges. The inner cone of S. murrayi
is similar to that of S. omani Adam and Rees, 1966, but in
S. omani the spine is keeled and the soft parts differ.
Literature: Adam and Rees, 1966.
Fig. 158 Sepia murrayi
ventral viewlateral view
t e n t a c u l a r c l u b c u t t l e b o n e
Fig. 159 Sepia murrayi
Known distribution
?? 200 m
Sepia officinalis Linnaeus, 1758 F i g . 1 6 0 ; P l a t e s I V , 2 8 a n d V , 2 9 ? 3 0
Sepia officinalis Linn?, 1758. Systema Naturae per Regna Tria Naturae, Secundum Classes, Ordines, Genera, Species
cum Characteribus, Differentiis, Synonymis, Locis , 658 [type locality: eastern North Atlantic].
Frequent Synonyms: Sepia filliouxi Lafont, 1869; Sepia mediterranea Ninni, 1884.
Misidentifications: None.
FAO Names: En ? Common cuttlefish; Fr ? Seiche commune; Sp ? Sepia com?n.
Diagnostic Features: Fins wide, extend anteriorly slightly beyond mantle margin. Arm suckers tetraserial. Hectocotylus
present on left ventral arm: 6 rows of normal size suckers proximally ; 4 to 8 rows of reduced suckersmedially . Club with
5 or 6 suckers in transverse rows ; suckers differ in size: 5 or 6 median suckers twice diameter of rest . Cuttlebone outline
oblong; acuminate, acute, anteriorly; bluntly rounded posteriorly. Spine short, pointed, surrounded by chitinous shield.
Sulcus present on last loculus only, absent from striated zone; sulcus shallow, narrow. Anterior striae are inverted
U-shape, or shallow m-shape . Inner cone limbs are narrow anteriorly, broaden posteriorly; outer cone chitinous; outer
cone narrow anteriorly, broadens posteriorly, spatulate, lateral limbs are flared ventrolaterally. Colour: Light brown .
Head with scattered white spots and with dark pigment around eye orbits. Arms I to III have a broad, longitudinal brownish
band medially, extending onto head. Dorsal mantle has bold transverse zebra stripe pattern during the breeding season ;
paired dorsal eye spots absent. Fins with narrow white band along outer margin and with small white spots, becoming
larger toward junction of mantle and fins. Mature males with arms IV emboldened by white and black zebra bands and
white arm spots .
Size: Up 490 mm mantle length, weight up to 4 kg in temperate waters and to 300 mm mantle length and 2 kg in the
subtropics.
Cephalopods of the World 99
t e n t a c u l a r c l u b
dorsal view
h e c t o c o t y l u s
(after Jatta, 1896 )
Fig. 160 Sepia officinalis
c u t t l e b o n e
ventral view
100 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Geographical Distribution: Eastern Atlantic and
Mediterranean Sea: eastern North Atlantic, from the
Shetland Islands and southern Norway (not present in
the Baltic Sea, except for occasional incursions with
the northeasternmost Atlantic waters) south through
the Mediterranean Sea (including Aegean Sea, Sea of
Marmara and Levantine Sea) to northwestern Africa,
with the southern boundary coinciding approximately
with the border between Mauritania and Senegal
(16?N). Endeavour Bank (Fig. 161).
Habitat and Biology: Depth range from subtidal
waters to 200 m, most abundant in upper 100 m, with
larger animals found at greater depths. Sepia
officinalis is a neritic, demersal species, found on the
continental shelf, predominantly on sandy or muddy
substrates. The species undergoes seasonal
migrations between inshore waters during spring and
summer and medium shelf grounds (about 100 m
depth) during autumn and winter. For example, in the
Mediterranean, large individuals leave deeper water
early in spring to migrate to shallower water, with
males preceding females. This group is followed by a
succession of smaller animals throughout the summer.
In autumn, gradual decent to deeper water begins.
Following an elaborate and ritualized courtship, which
incorporates stereotyped visual displays and ?mate
guarding?, spawning occurs in shallow water, with
peaks at water temperatures from 13? to 15?C. In the
western Mediterranean, this occurs between April and
July; off Senegal and on the Endeavour Bank, spawning occurs between January and April. Males carry up to 1 400
spermatophores and females (depending on their size) between 150 and 4 000 eggs. Mating takes place head-to-head
and spermatophores are placed in the females? buccal membrane ventral to the mouth. Eggs, 8 to 10 mm in diameter and
blackened with ink, are attached in grape-like clusters to seaweed, shells, debris and other substrates. They hatch after 30
to 90 days depending upon water temperature. The total length of hatchlings is between 7 and 8 mm. The growth rate
varies directly with temperature and inversely with size. Very young animals cannot live at depths much greater than about
50 to 80 m, because their cuttlebones cannot withstand water pressures higher than about 6 to 9 atmospheres. From
hatchlings to adults, S. officinalis exhibits a light-induced burying behaviour; most individuals spend the daytime hidden in
sand. The life cycle under natural conditions covers 12 to 24 months, varying with environmental conditions. Young
hatched in early summer from the spring brood usually spawn in the autumn of the following year; those from the autumn
brood spawn in the spring of their second year of life. Adult males may predominate because of massive postspawning
mortality among large females. Food consists of small molluscs, crabs, shrimps, polychaetes, other cuttlefishes and
juvenile demersal fishes. Prey is often ambushed when these cuttlefishes are partially buried and hidden in sand. The first
and second arm pairs commonly protrude from the substrate and are darkened, perhaps acting as lures. Cannibalism is
common and may be a strategy to overcome temporary prey shortages. The growth rate is rapid, given their short lifespan.
Predators include sharks, sparids and other demersal fishes and cephalopods. Sepia officinalis has been reared
successfully in aquaria and has aquaculture potential if live prey can be substituted by cheaper food items. Recent
information on the distribution of this species in Portuguese waters (i.e. Ria de Aveiro and adjacent coasts) indicates that
Sepia officinalis can tolerate brackish waters. The age and mantle length of specimens distributed in brackish areas
decrease with decreasing salinity. Younger specimens have greater ecophysiological plasticity and tolerate greater
environmental instability: this allows them to colonize the upper zones of the lagunar system, thus avoiding intraspecific
competition.
Interest to Fisheries: Sepia officinalis is one of the most important species for cephalopod fisheries in many countries. It is
probably close to its maximum sustainable production in several areas of its distribution since negative trends in captures
have been observed in recent years in some heavily fished areas (e.g. in the Mediterranean). Separate official statistics for
this species indicate that the main producer country in the year 2001 was Tunisia in the Mediterranean Sea, followed by
Greece; Spain and Portugal were also important producers, fishing S. officinalis in the northeastern Atlantic. However, the
species is also known to contribute extensively to the ?cuttlefishes? sensu lato official figures both in the Mediterranean
(where Italy is the main producer for this category) and in Atlantic waters. In the English Channel, the value of cuttlefish
landings increased greatly during recent decades, such that cuttlefishes have become a consistent component of earnings
both for French and United Kingdom fishermen. This has focused attention toward a managed exploitation of the resource.
Highest catches are recorded for Tunisia in the Mediterranean and off west Africa by Spanish and Moroccan fleets. In the
industrial fisheries, S. officinalis is primarily trawled, either as a target species, or as bycatch to demersal finfishes. The
artisanal fisheries, however, utilize a variety of selective gear, such as spears, pots and traps, often combined with the use
of light. Sometimes mature females are used as lures to attract males. Trammel nets are also efficient gear to catch
cuttlefishes and they are frequently used inshore in many Mediterranean fishing areas. Common cuttlefish usually is
marketed fresh or frozen and it is a highly valued food item, especially in Japan, Korea, Italy and Spain.
Fig. 161 Sepia officinalis
Known distribution
?? 200 m
Remarks: Five subspecies along the European?African coast were designated by Adam (1940). Khromov et al. (1998)
recognized S. o. hierredda Rang, 1837 and S. o. vermiculata Quoy and Gaimard, 1832 as valid, but consider S. o. filliouxi
Lafont, 1868 and S. o. mediterranea Ninni, 1884 as indistinct from S. officinalis (this designation restricts the distribution of
the species to the subtropical and temperate waters of the eastern Atlantic in the northern hemisphere). Sepia officinalis
and S. hierredda now are known to be distinct species (see Remarks S. hierredda). In the Mediterranean, young
S. officinalis can be distinguished from S. orbignyana F?russac, 1826 and S. elegans Blainville, 1827 by their brown,
rather than red, skin colour, the shape of the bone and the club sucker arrangement. The biology of S. officinalis is well
known. It has been the most extensively studied of all cuttlefish species.
Local Names: ALGERIA: Choubai, Choueb?, Seiba, Seich, Sepia, Seppio; Soupia; EGYPT: Sobbeit; FINLAND:
Mustekala, Sepia; BULGARIA: Sepija; CYPRUS: Corsica: Seppia; FRANCE: Casseron, Chakod, Chibia, Margade,
Seiche; GERMANY: Gemeiner Tintenfisch, Sepie; GREECE: Soupia; ISRAEL: Dyonon refui; ITALY: Seppia comune;
Scarpetta, Scarpitta; Scarpitelle, Seccetella, (juveniles), Secce, Seppa, Seppia, Siccia, Sepa, Sepia imperiale; JAPAN:
Mongo-ika, Yoroppa kouika; LEBANON: Sabbidije; LIBYA: Shoubia; MADEIRA: Choco; MALTA: Sicca; MONACO: Supia;
MOROCCO: Chubei, Seiche; NETHERLANDS: Gewone Inktvis, Zeekat; PORTUGAL: Checo, Choco; ROMANIA: Sepia;
RUSSIAN FEDERATION: Kora katitza; SENEGAL: Seiche; SPAIN: Aluda, Choco, Coca, Jibia, Jibi?n, Luda, Rellena,
Relleno, Sipia, Sipionet; TUNISIA: Choubei, Chouebi, Seche, Sibia, Sipia, Soubia; TURKEY: S?bye; UK: Cuttlefish;
YUGOSLAVIA: Sipa.
Literature: Tompsett (1939), Mangold-Wirz (1963), Boletzky (1983a), Guerra and Castro (1988), Hanlon and Messenger
(1988), Guerra (1992), Augustyn et al. (1995), Hanlon and Messenger (1996), Sanjuan et al. (1996), Neige and Boletzky
(1997), Khromov et al. (1998), Belcari (1999b), P?rez-Losada et al. (1999), Denis and Robin (2001), Guerra et al. (2001),
Belcari et al. (2002), Jereb (2002), Salman et al. (2002), Sobral (2002), Vrgoc
 et al. (2004). Numerous other references
pertaining to this species are available.
Sepia omani Adam and Rees, 1966 F i g . 1 6 2
Sepia omani Adam and Rees, 1966. The John
Murray Expedition , 11(1): 92 [type locality: Gulf of
Oman].
Frequent Synonyms: None.
Misidentifications: Sepia vossi Khromov, 1996.
FAO Names: En ? Oman cuttlefish; Fr ? Seiche
d?Oman; Sp ? Sepia de Oman.
Diagnostic Features: Mantle oval. Arm suckers
tetraserial ; webs wide between dorsal and
dorsolateral arms. Hectocotylus present on left
ventral arm: 2 or 3 rows of normal size suckers
proximally, suckers reduced medially for 40% of arm
length, then normal size suckers to arm tip; reduced
suckers much smaller than normal arm suckers; oral
surface of modified region wide, swollen, fleshy, with
transversely grooved ridges; suckers in 2 dorsal and
2 ventral series displaced laterally, with gap between;
2 ventra l ser ies crowded together. Club
sucker-bearing surface flattened, with 3 or 4 suckers
in transverse rows ; suckers differ markedly in size:
3 to 5 suckers in middle of third longitudinal row
extremely enlarged . Swimming keel of club extends
proximally slightly beyond carpus; dorsal and ventral
protective membranes not joined at base of club , do
not continue along stalk. Dorsal membrane forms
deep cleft at junction with stalk. Cuttlebone
acuminate, acute, anteriorly and posteriorly; dorsal
median rib present, sides almost parallel, flanked by
shallow grooves; lateral ribs distinct. Spine long, pointed. Anterior striae shallow m-shape ; sulcus shallow, narrow,
extends entire length of cuttlebone. Inner cone limbs are uniform width, narrow, U-shape posteriorly, thickened; inner cone
posteriorly with irregular calcareous ribs radiating into outer cone; outer cone calcified, narrow anteriorly, broadens
posteriorly. Colour: Light brown. Dorsal mantle has dark brown transverse stripes.
Cephalopods of the World 101
Fig. 162 Sepia omani
dorsal viewventral view
t e n t a c u l a r c l u b c u t t l e b o n e
102 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Size: Up to 100 mm mantle length.
Geographical Distribution: Northern Indian Ocean:
Gulf of Oman, off Pakistan and western India. Limits of
range unknown (Fig.163).
Habitat and Biology: This is a neritic demersal
species. Depth range from 50 to 210 m.
Interest to Fisheries: At present still undetermined.
Probably the species occurs in mixed species trawls,
where it is not separated from other sepiids.
Remarks: Records from the Indo-Pacific refer to
S. vossi Khromov (1996).
Literature: Filippova et al. (1995).
Sepia opipara ( Iredale, 1926 ) F i g . 1 6 4
Glyptosepia opipara Iredale, 1926, The Australian Zoologist , 4(3): 191 [type locality: Australia: Queensland, Masthead
Island, Capricorn Group, 23?32'S 151?44'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Magnificent
cuttlefish; Fr ? Seiche magnifique;
Sp ? Sepia magn?fica.
Diagnostic Features: Mantle broad,
oval. Male and female arm lengths
subequal; protective membranes
narrow. Arm suckers tetraserial.
Hectocotylus present on left ventral
arm: 5 or 6 rows of normal size
suckers proximally, 6 or 7 rows of
reduced suckers medially , then
normal size suckers distally to arm
tip; oral surface of modified region
not wide, fleshy, but normal, as on
opposite arm. Club sucker-bearing
surface flattened, with 3 or 4
suckers in trans verse ro ws ;
suckers differ markedly in size:
middle of club with 4 or 5 greatly
enlarged suckers , second proximal
one the largest, obliquely 1 small
lateral sucker and 1 minute marginal
sucker dorsal to the big sucker, 1
small lateral and 4 minute marginal
suckers ventral to it ; suckers
proximal and distal to the big suckers are small. Swimming keel of club extends well proximal to carpus; dorsal and ventral
protective membranes not joined at base of club but fused to tentacular stalk . Dorsal and ventral membranes of equal
length; extend proximal to carpus along stalk. Dorsal membrane forms deep cleft at junction with stalk. Cuttlebone outline
oblong; bone bluntly rounded anteriorly; acuminate, acute, posteriorly; dorsal surface pinkish ; dorsal surface flat
medially and laterally; texture rough, with irregular calcified projections; dorsal median rib distinct; sides approximately
parallel, flanked by broad grooves; lateral ribs distinct. Chitin borders lateral and anterior margins of cuttlebone. Spine
Fig. 163 Sepia omani
Known distribution
?? 200 m
?
?
Fig. 164 Sepia opipara
ventral viewdorsal view
t e n t a c u l a r c l u b
c u t t l e b o n e
(illustration: K. Hollis)
(after Iredale, 1926 )
short, pointed, curves dorsally, with ventral keel .
Striated zone flat; last loculus convex; sulcus shallow,
narrow. Anterior striae are inverted U-shape ; limbs of
inner cone extend anteriorly to end of striated zone.
Inner cone limbs are uniform width, narrow, U-shape
posteriorly, thickened; outer cone calcified, narrow
anteriorly, broadens posteriorly.
Size: Up to approximately 150 mm mantle length.
Geographical Distribution: Southern Indo-Pacific:
northern Australia from Dirk Hartog Island, 25?45'S
113 ?03 'E (Western Austra l ia) , to southern
Queensland, 36?57'S 151?45'E (Fig. 165).
Habitat and Biology: Depth range from 83 to 184 m.
Interest to Fisheries: Species taken as bycatch of
prawn and mixed species trawl fisheries. The size of
this species and its broad distribution indicate that it
has potential for exploitation.
Literature: Lu (1998a).
Sepia orbignyana F?russac in d?Orbigny, 1826 F i g . 1 6 6
Sepia orbignyana F?russac in d?Orbigny, 1826, Annales des Sciences Naturelles, Paris, series 1, 7: 156 [type locality:
France: La Rochelle].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Pink cuttlefish;
Fr ? Seiche ros?e; Sp ? Sepia con
punta.
Cephalopods of the World 103
120?
120?
130?
130?
140?
140?
150?
150?
 40?  40?
 30?  30?
 20?  20?
 10?  10?
Fig. 165 Sepia opipara
Known distribution
?? 200 m
t e n t a c u l a r c l u b
dorsal view
(after Adam, 1952 )
Fig. 166 Sepia orbignyana
c u t t l e b o n e
dorsal view ventral view
Diagnostic Features: Mantle oval. Male and female arms subequal in length. Arm suckers tetraserial. Male medial
non-hectocotylized arm suckers with greater diameter than marginal ones. Hectocotylus present on left ventral arm: 1 or 2
rows of normal size suckers proximally, greatly reduced suckers medially, then normal size suckers distally to arm tip.
Suckers of hectocotylus in 2 dorsal and 2 ventral series displaced laterally, with gap between them. Club short, oval , with 5
or 6 suckers in transverse rows ; suckers differ markedly in size: 3 large suckers medially with one slightly smaller
sucker on each side of these . Swimming keel of club extends proximal to carpus; dorsal and ventral protective
membranes joined at base of club . Buccal membrane in females with single median spermathecae in ventral part.
Cuttlebone outline oblong; acuminate, acute, anteriorly; bluntly rounded posteriorly; strongly recurved ventrally ; dorsal
surface pinkish; dorsal surface flat medially, curved, convex laterally; granulose; dorsal median rib absent; bone with
dorsal median groove present , shallow, narrow, extending full length of cuttlebone. Spine long, pointed (prominent),
straight, directed dorsally, with ventral keel . Striated zone and last loculus convex; sulcus shallow, narrow, extends entire
length of cuttlebone; sulcus flanked by rounded ribs borde red laterally by shallow grooves . Anterior striae shallow
m-shape, or wavy. Inner cone limbs are uniform width, narrow V-shape posteriorly, thickened slightly; outer cone limbs are
slightly expanded posteriorly, directed ventrally to form a recurved cup-like structure. Colour: Reddish brown.
Size: Males up to 96 mm mantle length; females up to 120 mm mantle length.
Geographical Distribution: General distribution: eastern
Atlantic and Mediterranean Sea: from the Irish Sea, English
Channel and Bay of Biscay to southern Angola (17?S), and
throughout the Mediterranean Sea (including the Adriatic
and Aegean Seas, Sea of Marmara and Levantine Sea).
Saharan Bank (Fig. 167).
Habitat and Biology: The depth range for this species is
from 15 to 570 m, and it is most abundant between 50 and
250 m. Sepia orbignyana is a demersal species that lives
mainly on sandy and sandy-muddy bottoms. It is frequently
sympatric (and confused) with S. elegans Blainville, 1827. In
the Sea of Marmara the species can occur in brackish
waters. In Mediterranean waters, males and females are
usually found together throughout the year and the
spawning period is probably continuous, with peaks of
activity from spring to autumn. A predominance of mature
individuals in spring is also reported for the species in
Portuguese waters. No onshore spawning migrations have
been reported. Due to the extended reproductive period,
recruitment is also continuous but variable, with seasonal
density peaks. Growth rates of females are higher than
those of males, and females also attain larger sizes. In the
Mediterranean, the smallest mature males are 35 mm, and
the smallest females 65 mm mantle length. Mature males,
aged 6 or 7 months, carry about 100 spermatophores;
females of 9 or 10 months of age, carry around 400 eggs.
Egg diameter increases with the size of the females. The
eggs (maximum size 7 to 8.5 mm diameter) are laid in
clusters of 30 to 40 and are attached to sponges on muddy
bottoms. Diet consists mainly of crustaceans, but fish and
cephalopods make up a minor component.
Interest to Fisheries: Sepia orbignyana is one of the most
abundant cephalopod species in some areas of its distributional range (e.g. within the Mediterranean: Aegean, southern
Adriatic and Tyrrhenian Seas and in the Sicilian Channel). It is taken mainly as a bycatch throughout the Mediterranean
and in the west African trawl fisheries. Separate statistics are not reported, but S. orbignyana represents a very significant
percentage of the catches in some areas. In the Mediterranean Sea it is marketed along with S. elegans and small
S. officinalis and constitutes a valuable resource locally. In the Sicilian Channel, research studies showed an exploitation
rate of 0.60 for this species, which suggests an intense fishing pressure on this resource. It is marketed fresh and frozen.
Local Names: ITALY: Seppia pizzuta; SPAIN: Sepia puntiaguda, Chopito.
Remarks: Sepia orbignyana differs from S. elegans in having more than 100 suckers on the club. The 2 species also differ
in the cuttlebone form and structure. Sepia elegans lacks a cuttlebone spine, while a spine is present in S. orbignyana. In
addition, Sepia orbignyana differs from S. officinalis Linnaeus, 1758 in having a reddish, rather than brown coloration, it
generally inhabits deeper water and does not bury in the sand during the day.
Literature: Mangold-Wirz (1963), Adam and Rees (1966), Bello (1990a), Jereb and Ragonese (1991), Ragonese and
Jereb (1991), Wurtz et al. (1991), Guerra (1992), D?Onghia et al. (1996), Sanjuan et al. (1996), Neige and Boletzky (1997),
Belcari (1999c), Salman et al. (2002).
104 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
 20?
 20?
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
 10?  10?
0? 0?
10? 10?
20? 20?
30? 30?
40? 40?
50? 50?
60? 60?
Fig. 167 Sepia orbignyana
Known distribution
?? 200 m
Sepia papuensis Hoyle, 1885 F i g . 1 6 8 ; P l a t e V , 3 1
Sepia papuensis Hoyle, 1885, Annals and Magazine of Natural History , series 5, 16: 197 [type locality: Challenger Station
188, 09?59'S 139?42'E, Araf ura Sea, South of Papua].
Frequent Synonyms: ? Sepia galei Meyer, 1909; Solitosepia submestus Iredale, 1926; Solitosepia occidua Cotton,
1929; Solitosepia genista Iredale, 1954; Solitosepia lana Iredale, 1954; S. prionota Voss, 1962a.
Misidentifications: None.
FAO Names: En ? Papuan cuttlefish; Fr ? Seiche de
Papouasie; Sp ? Sepia de Papua.
Diagnostic Features: Mantle oval. Male and female
arm lengths subequal; protective membranes wide,
well developed as a series of semicircular lappets. Arm
sucker arrangement differs between sexes: in males,
sucker rows on arms I to III tetraserial proximally,
biserial at distal tips, arms IV suckers tetraserial ; in
females, arm suckers tetraserial. Hectocotylus
absent. Club sucker-bearing surface flattened, with 5
or 6 suckers in transverse rows ; suckers differ
markedly in size . Swimming keel of club extends well
proximal to carpus; dorsal and ventral protective
membranes not joined in small specimens, joined at
base of club in large specimens; dorsal protective
membrane much longer than ventral protective
membrane, extending proximal to carpus along stalk.
Dorsal membrane forms deep cleft at junction with
stalk. Cuttlebone length approximately equal to mantle
length; outline oblong; bone bluntly rounded anteriorly
and posteriorly; dorsal surface creamy white; dorsal
surface convex medially, flat laterally; texture smooth,
not pustulose; dorsal median and lateral ribs distinct;
median rib broadens anteriorly. Chitin borders lateral
and anterior margins of cuttlebone. Spine long,
pointed, curves dorsally, with ventral keel . Striated
zone and last loculus concave; sulcus shallow, wide,
extends entire length of cuttlebone. Anterior striae are
inverted U-shape . Inner cone limbs are narrow
anteriorly, broaden posteriorly, thin; outer cone
calcified; narrow anteriorly, broadens posteriorly.
Dorsal mantle with series of elongate papillae along
each side, adjacent to base of each fin, rest of mantle
covered with numerous small papillae. Papillae in 2
series: i) a conspicuous series that consists of a pair of
large, triangular, flattened primary papillae that
emerge from mantle white spots; ii) just posterior and
lateral to mantle white spots lies a pair of papillate
ridges located along the boundary between a white
midline stripe and darker lateral fields. Head with 2
pairs large papillae located over anterior and posterior
ends of each eye, numerous scattered, small papillae
and one on each eyelid. Arms I to III with papillae.
Colour: Light brown. Dorsal mantle has white blotches
or spots. Paired dorsal eye spots present.
Size: Up to 110 mm mantle length.
Geographical Distribution: Indo-West Pacific:
Philippine Islands, Indonesia, Bali, Ternate, Arafura
and Coral Seas, northern Australia from southern
Western Australia (Freemantle, 32?03'S 115?44'E) to
southern NSW (36?35'S 150? 16'E), including Gulf of
Carpentaria (Fig. 169).
Cephalopods of the World 105
Fig. 168 Sepia papuensis
c u t t l e b o n e
(after Lu, 1998 )
t e n t a c u l a r c l u b
(illustration: K. Hollis)
subadult ventral view
100?
100?
110?
110?
120?
120?
130?
130?
140?
140?
150?
150?
 30?  30?
 20?  20?
 10?  10?
0? 0?
10? 10?
20? 20?
Fig. 169 Sepia papuensis
Known distribution
?? 200 m
?
?
?
?
?
106 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Habitat and Biology: Depth range from 10 to 155 m. Sepia papuensis is a shelf species found on silt, sand or muddy
substrates. This species appears to be night-active and has been observed foraging in beds of seaweed and seagrass.
Small individuals have been observed to extend flattened arms to mimic the shape of the seaweed Halimeda as
camouflage. Behavioural studies have shown that the species exhibits a very wide range of colour patterns associated with
substrate colour and texture, and in response to potential attack by predators.
Interest to Fisheries: Species taken as bycatch of prawn and mixed species trawl fisheries. This species was collected in
small numbers in demersal trawl and dredge surveys in the Gulf of Carpentaria, Australia, in 1990 and 1991 and probably
forms part of the bycatch of Taiwanese trawl fisheries in the region.
Literature: Adam and Rees (1966), Roper and Hochberg (1987), Lu (1998a).
Sepia pharaonis Ehrenberg, 1831 F i g . 1 7 0 ; P l a t e V , 3 2 ? 3 4
Sepia pharaonis Ehrenberg, 1831, In: Ehrenberg, C.G. (1831). Cephalopoda in Mare Rubro viventia. In ?Symbolae
Physicae, seu Icones et descriptiones Corporum Nauralium?quae ex itineribus per Libyan, Aegyptum?
Habessiniam?Pars Zoologica?. ( P.C. Hemprich and C.G. Eh renberg, eds) [4]: 4 [type locality: Massaouah, Gulf of Suez].
Frequent Synonyms: Sepia torosa Ortmann, 1888;
Sepia rouxii d?Orbigny, 1839?1842; Sepia formosana
Berry, 1912a; Crumenasepia hulliana Iredale, 1926;
Crumenasepia ursulae Cotton, 1929; Sepia rouxi
d?Orbigny, 1841; Sepia formosana Berry, 1912; Sepia
formosana Sasaki, 1929; Sepia tigris Sasaki, 1929.
Misidentification: None.
FAO Names: En ? Pharaoh cutt lef ish;
Fr ? Seiche pharaon; Sp ? Sepia fara?n.
Fig. 170 Sepia pharaonis
dorsal view
c u t t l e b o n e
ventral view
h e c t o c o t y l u s t e n t a c u l a r c l u b
Diagnostic Features: Mantle oval. Male and female arm lengths subequal. Arm suckers tetraserial. Hectocotylus present
on left ventral arm: 10 to 12 rows of normal size suckers proxi mally, 6 rows of reduced suckers medially , then normal
size suckers distally to arm tip. Suckers of hectocotylus in 2 dorsal series aremuch smaller than those in 2 ventral series ;
oral surface of modified region wide, swollen, fleshy, with transversely grooved ridges; with shallow median furrow; suckers
in 2 dorsal and 2 ventral series displaced laterally, with gap between them. Club sucker-bearing surface flattened, with 8
suckers in transverse rows ; suckers differ markedly in size: 5 or 6 median suckers enlarged (3 or 4 of these are greatly
enlarged). Swimming keel of club terminates at proximal end of carpus. Dorsal and ventral protective membranes not
joined at base of club ; dorsal and ventral membranes same length; extend proximal to carpus along stalk. Dorsal
membrane forms shallow cleft at junction with stalk. Buccal membrane with a few, minute suckers (each lappet bearing 1
or 2 small suckers). Cuttlebone outline oblong; bone bluntly rounded anteriorly; acuminate, acute, posteriorly; dorsal
surface creamy white; dorsal surface evenly convex; texture smooth; dorsal median rib distinct, rib broadens anteriorly;
lateral ribs indistinct. Chitin borders lateral and anterior margins of cuttlebone. Spine short, pointed, curves dorsally, keel(s)
absent. Striated zone concave; last loculus flat; sulcus deep, wide, extends entire length of cuttlebone; sulcus flanked by
rounded ribs. Anterior striae are inverted U-shape ; limbs of inner cone extend anteriorly to end of striated zone. Inner
cone limbs are narrow anteriorly, broaden posteriorly with distinctive thick bulbous swelling ; outer cone calcified; narrow
anteriorly, broadens posteriorly. Dorsal mantle with series of elongate papillae along each side, adjacent to base of each
fin, or covered with numerous small papillae. Colour: Pale brownish or reddish purple. Head and arms with transverse
zebra-stripe pattern. Dorsal mantle has white blotches or spots, transverse saddle mark, and has a transverse zebra-stripe
pattern (saddle mark in females; stripes especially in males; small specimens may show stripe markings or few markings).
Fins with longitudinal white band at base, bordered by narrow band of ground-coloured pigment along each side; white
stripe solid on anterior 3/4 of body, interrupted by blocks of ground-coloured pigment on posterior 1/4 of mantle.
Size: Up to 420 mm mantle length; weight to 5 kg.
Geographical Distribution: Indian
Ocean and western Pacific: including
the Red Sea and Arabian Sea south to
Zanzibar and Madagascar, Andaman
Sea to South China Sea, East China
Sea, Taiwan Province of China, Japan
(Kyushu and possibly southern Honshu),
eastern Indonesia and northern Australia
(from Monte Bello Island, Western
Australia, 20?26'S 115?37'E, to at least
Townsvi l le , Queensland, 19 ?16 'S
146?41'E, including Gulf of Carpentaria).
Wadge Bank (Fig. 171).
Habitat and Biology: Sepia pharaonis is
a neritic demersal species which occurs
down to 130 m. In the Gulf of Thailand
and the Andaman Seas, animals are
found from the coastal shallows to 100 m
depth, with most caught between 10 and
40 m. Around Hong Kong, animals
migrate to shallower waters during the
mating season, where large numbers of
adults congregate in 40 to 80 m on the
continental shelf from November to
February. During February and March,
they move to the coast where spawning
takes place from April to May in water
temperatures between 18 and 24?C.
Eggs are laid in clusters and attached to
plants, shells and other hard substrates
in approximately 5 to 20 m depth.
In the Gulf of Thailand, spawning occurs all year round, with peak months during January and February and from July to
September. The ratio of males to females is 1:3. During the spawning season, 4 or 5 adults congregate, sometimes near a
boulder. During mating, the males adopt a very bold striped pattern. Both males and females raise their dorsal arms
vertically and the female elevates lappets on her mantle: 6 on the base of each fin and 2 over each eye. The males
approach either head on, or with ventral arms spread, they swim from behind and over her back. Mating takes place head
on, each holding the other with the arms. Mating may occur more than once with a single pair. Females lay white eggs that
are attached singly to hard surfaces.
Cephalopods of the World 107
Fig. 171 Sepia pharaonis
Known distribution
?? 200 m
In Iranian waters, S. pharaonis is the dominant cephalopod species. The main spawning season occurs between the
southwest and northeast monsoon periods in November and December. In Yemen, S. pharaonis is one of the species with
greatest commercial value; experts have estimated the standing stocks and an annual exploitable yield has been
recommended.
Sepia pharaonis rises to the lower part of the water column to feed at night, mainly on crustaceans and a variety of small
demersal fishes. During maturation, there is a shift of emphasis from somatic growth to gonadal development and
vitellogenesis. Research observations suggest that energy and nutrients for maturation are supplied mainly by diet rather
than stored resources: the species does not use protein from muscle tissue for developing and growing its reproductive
tissues. Sepia pharaonis appears to be an intermittent multiple spawner. In captive animals, the life cycle is less than 10
months. Estimated growth rates for wild animals show higher values for females than for males.
Interest to Fisheries: This species supports industrial or artisanal fisheries throughout its range. With S. esculenta Hoyle,
1885, it is the most abundant cuttlefish species caught in the Philippines and the Samar and Visayan Seas, with the highest
catches reported in the Lingayen Gulf and Carigara Bay. In Iran, the fishing activity occurs during the spawning season,
when adults migrate from deeper waters to shallower waters in the littoral zone. Sepia pharaonis is caught by bottom
trawlers in the Oman Sea, and by traps in the Persian Gulf and is one of the most important cuttlefish species fished in both
areas. The species is important to the commercial cephalopod fishery of Thailand, being highly abundant in the Gulf and
the Andaman Sea, where it is the most common species of cuttlefish caught. The species contributes about 90% of the
cuttlefishes caught off Australia by Chinese pair trawlers. Off the North West Shelf and Timor Seas, sepiids (mainly
S. pharaonis) tend to replace squids as the dominant cephalopods caught. Domestic fisheries in these waters take this
species as bycatch of prawn and mixed species trawl fisheries. In the Hong Kong area, it is the most abundant cuttlefish
species and it is of greatest commercial importance in this area and along the whole coast of Kwangtung and Fukien, with
about 400 tonnes landed annually in Hong Kong. Animals in this region are caught by spearing, lure-hooking and trawling.
In southern Thailand, in addition to otter and pair trawls, the trammel net and hook-and-line are commonly used for
catching S. pharaonis, with bottom otter and pair trawls used offshore, and push nets and lift nets used in inshore and
coastal waters. Squid traps, in which egg clusters are placed to entice squids to enter, are also widely used and
cuttlefishes, all mature animals ready to spawn, are a major bycatch. These traps accounted for 5% of total Thai
cephalopod catch in 1994 (i.e. over 7 000 tonnes). Off the southwest coast of India, a modified type of hook, a baited
hand-jig, is used to catch this species. Sepia pharaonis has been grown successfully in culture, and techniques are
currently being improved in Thailand to culture these animals commercially. The flesh is thick, tender and excellent for
human consumption.
Local Names: CHINA: Mak mo, Foo baan woo chack; JAPAN: Torafu-kouika, Mongouika.
Remarks: The diagnostic features given above were obtained from a range of sources and examination of Australian
specimens. They should be treated with caution because animals currently referred to as S. pharaonis are probably
representatives of a species complex. One species from southeast India, S. ramani Neethiselvan, 2001, has been
described recently. Sepia ramani differs from S. pharaonis in having a longer club with 15 to 24 subequal enlarged
suckers. Sepia pharaonis has 6 enlarged medial club suckers, 3 or 4 of which are much larger than the rest. Sepia ramani
has 14 to 16 transverse rows of normal size suckers on the proximal end of the hectocotylized arm, instead of 10 to 12
rows, as in S. pharaonis. In other traits, these 2 species are very similar. The hectocotylus differs between S. pharaonis
from Japan and Australia. Given its fisheries importance, the status of this putative species needs to be examined and
specimens from throughout its range compared with those from the type locality to accurately determine its population
structure and/or species boundaries.
Literature: Adam and Rees (1966), Silas et al. (1982), Valinassab (1983), Nair (1986), Okutani et al. (1987), Gutsal (1989)
Chu et al. (1992), Kukharev et al. (1993), Khaliluddin (1995), Chantawong and Suksawat (1997), Chotiyaputta and
Yamrungreung (1998), Lu (1998a), Rocha et al. (1998), Gabr et al. (1999).
108 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia plangon Gray, 1849 F i g . 1 7 2 ; P l a t e V , 3 5
Sepia plangon Gray, 1849, Catalogue of the Mollusca in the British Museum. Part I. Cephalopoda Antepedia : 104 [type
locality: Australia, New South Wales, Port Jackson, 33?51'S 151?16'E].
Frequent Synonyms: Solitosepia plangon adhaesa
Iredale, 1926.
Misidentifications: None.
FAO Names: En ? Striking cuttlefish; Fr ? Seiche
impressionnante; Sp ? Sepia impresionante.
Diagnostic Features: Mantle
broad, oval. Male and female
Arm lengths subequal;
protective membranes narrow.
Arm suckers tetraserial.
Hectocotylus present on left
ventral arm: 5 rows of normal
size suckers proximally, 5 rows
of reduced suckers medially ,
then normal size suckers
distally to arm tip. Suckers
equal in size , reduced suckers
only slightly smaller than
normal arm suckers; oral
surface of modified region not
wide, fleshy, but normal, as on
opposite arm. Club sucker-
bearing surface flattened, with
5 suckers in transverse rows ;
suckers differ markedly in size:
some greatly enlarged (median
series largest followed by those
immediately bordering ventral
series). Swimming keel of club
extends well proximal to
carpus; dorsal and ventral protective membranes joined at base of club but fused to tentacular stalk. Dorsal and ventral
membranes same length; extend proximal to carpus along stalk. Dorsal membrane forms deep cleft at junction with stalk .
Cuttlebone outline oblong; acuminate, acute, anteriorly; bluntly rounded posteriorly; dorsal surface pinkish; dorsal surface
convex medially, flat laterally; granulose; dorsal median rib distinct; sides approximately parallel; lateral ribs indistinct. Chitin
borders lateral and anterior margins of cuttlebone. Spine long, pointed, straight, parallel to bone, with ventral keel . Striated zone
concave; last loculus flat; sulcus deep, narrow , extends entire length of cuttlebone; sulcus flanked by rounded ribs (giving
striae a wavy appearance). Anterior striae are inverted V-shape . Inner cone limbs are uniform width , narrow, U-shape
posteriorly; outer cone calcified; narrow anteriorly, broadens posteriorly. Colour: Reddish purple, light brown, or dark brown, or
sometimes with whitish mottle. During the breeding season, the head and dorsal mantle of males have narrow, irregular
light-coloured transverse stripes. Fins pale with broad light greenish band along base.
Size: Up to 135 mm mantle length.
Geographical Distribution: Southwestern Pacific:
eastern Australia from Gulf of Carpentaria to Sydney,
NSW 33?53'S 151?13'E (Fig. 173).
Habitat and Biology: Depth range intertidal to 83 m.
Sepia plangon frequently is found among seaweed
and seagrass. It is active during the day, foraging for
crustaceans and fishes. It often can be seen resting on
the sea floor, raising the anterior end of the body by
using its arms as stilts. Females are plain coloured,
without pattern, during courtship.
Interest to Fisheries: Species taken as bycatch of
prawn and mixed species trawl fisheries.
Literature: Adam and Rees (1966), Lu (1998a).
Cephalopods of the World 109
Fig. 172 Sepia plangon
c u t t l e b o n e
ventral view
t e n t a c u l a r c l u b
dorsal view
Fig. 173 Sepia plangon
Known distribution
?? 200 m
110 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia prabahari Neethiselvan and Venkataramani, 200 2 F i g . 1 7 4
Sepia prabahari Neethiselvan and Venkataramani, 2002, Indian Journal of Marine Sciences , 31(1): 45 [type locality:
southeast coast of India, Tuticorin, 08?47'N 78?9'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Small striped cuttlefish; Fr ? Petite seiche ray?e; Sp ? Sepia listada peque?a.
Diagnostic Features: Male arms elongate, robust; arms I
and IV elongate, whip-like (more pronounced in mature
males); in females arms approximately subequal in length;
arm formula in both sexes IV, I, III, II; arm suckers
tetraserial. Hectocotylus present on left arm IV: 8 transverse
rows of normal size suckers proximally, followed by 7 rows
of modified suckers, then rest normal to arm tip. Suckers in
2 dorsal series greatly reduced, suckers in 2 ventral
series normal in size on modified portion of arm; 2 dorsal
and 2 ventral series displaced laterally, with fleshy ridge
between. Club short, with 6 suckers in transverse rows ;
suckers all of similar minute size . Swimming keel
terminates at posterior end of carpus. Dorsal protective
membrane broader than ventral protective membrane.
Dorsal and ventral protective membranes extend slightly
beyond carpus, not joined at base of club . Cuttlebone
elliptical; broader in females than males; rugose dorsally,
with indistinct median and lateral ribs. Spine curved dorsally,
without keels. Sulcus deep, broad, extends length of
striated zone. Anterior striae are inverted V-shape. Inner
cone limbs are narrow anteriorly, br oaden posteriorly,
then are raised into a thick, round ledge ; outer cone
male female
Fig. 174 Sepia prabahari
male female
h e c t o c o t y l u s
c u t t l e b o n e
ventral view
narrow anteriorly, broadens posteriorly, fused to inner cone. Colour: Dorsal mantle, head and arms dark brown with transverse
zebra-stripe pattern (more prominent in males than females, and more obvious in fresh animals). Fins with pale, narrow,
longitudinal line along base. Golden yellow nidamental glands visible in mantle cavity of female over 60 mm mantle length.
Cuttlebone inner cone white.
Size: Up to 130 mm mantle length.
Geographical Distribution: Indian Ocean: Indian
Coast, Gulf of Mannar. Depth ranges up to 100 m
(Fig. 175).
Habitat and Biology: No detailed information is
available yet for this newly described species.
Interest to Fisheries: Sepia prabahari is fished year
round along the Thoothukkudi southeast coast (India).
Remarks: Sepia prabahari appears to breed
continuously throughout the year.
Local Names: INDIA: Chinna vari kanavai.
Literature: Neethiselvan and Venkataramani, 2002.
Sepia prashadi Winckworth, 1936 F i g . 1 7 6
Sepia prashadi Winckworth, 1936, Proceedings of the Malacological Society of London , 22(1): 16 [type locality: India,
Chennai (Madras), Bay of Bengal].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Hooded cuttlefish; Fr ? Seiche capuchon;
Sp ? Sepia caperuza.
Cephalopods of the World 111
Fig. 175 Sepia prabahari
Known distribution
?? 200 m
Fig. 176 Sepia prashadi
dorsal view
dorsal viewventral view
c u t t l e b o n et e n t a c u l a r c l u b
Diagnostic Features: Male and female arms subequal in length; protective membranes narrow. Arm suckers tetraserial.
Male median arm suckers with greater diameter thanmarginal ones over most of arm length. Hectocotylus present on left
ventral arm: 4 rows of normal size suckers proximally, 12 to 14 rows of reduced suckers medially , then normal size
suckers to arm tip; oral surface of modified region wide, swollen, fleshy, with transversely grooved ridges. Suckers of
hectocotylus in 2 ventral series are displaced laterally, with gap between on proximal part of modified region, becoming
closer together distally. Club short, oval; sucker-bearing surface flattened, with 3 to 5 suckers in transverse rows ; suckers
differ markedly in size: median 4 suckers extremely large , surrounded by moderately large suckers. Swimming keel of
club extends proximally slightly beyond carpus; dorsal and ventral protective membranes not joined at base of club .
Cuttlebone outline oblong; bone triangular, obtuse anteriorly; bluntly rounded posteriorly; granulose; dorsal median rib
distinct, broadens slightly anteriorly; ribs bordered laterally by distinct grooves ; lateral ribs distinct. Chitin borders lateral
and anterior margins of cuttlebone. Spine long, pointed, straight, directed dorsally, with dorsal and ventral keel. Striated
zone convex; striated zone separated from outer cone by narrow, smooth marginal zones; sulcus shallow, narrow, flanked
by rounded ribs, extends entire length of cuttlebone. Anterior striae shallow m-shape . Inner cone limbs are narrow
anteriorly, broaden posteriorly; raised into rounded, thickened ledge ; outer cone chitinous laterally, calcareous in
expanded posterior part. Dorsal mantle has transverse zebra stripe pattern in breeding males.
Size: Up to 140 mm mantle length.
Geographical Distribution: Indian Ocean: from off
northeastern India (Calcutta) to the Gulf of Oman, Red
Sea, Gul f of Suez, southern Mozambique,
Madagascar, Mauritius, Andaman?Nicobar Seas
(Fig. 177).
Habitat and Biology: A demersal, shallow water,
small-sized species ranging in depth from the coastline
to 200 m. Off Waltair (northeastern coast of India), the
species occurs throughout the year, even though
sporadically. For this area, preliminary studies reported
mature individuals at 72 and 67 mm mantle length for
females and males, respectively. In the waters off
Madras (southeast India), S. prashadi is found from
January to April, along with upwelled deeper water
fishes. In the Gulf of Suez a life span of 18 months was
estimated.
Interest to Fisheries: Taken by trawls in the Red Sea
and along the Indian east coast off Waltair and Madras
(northeast India), S. prashadi is most abundant from
the beginning of the year to June and in some years,
from October to December. Off Madras, it is taken in
small quantities in local upwelling areas as bycatch to
finfishes normally occurring in deeper waters.
Separate catch statistics are not reported for this
species. Exploratory surveys in the Andaman?Nicobar
Seas indicate a wide distribution of cephalopod
resources both in the coastal and oceanic waters. It
was suggested that suitable techniques, such as light
fishing with lift nets, would result in better exploitation
of demersal species, among which S. prashadi is well
represented. Based on stock assessment studies in
the Gulf of Suez, S. prashadi was underexploited in the
mid-1990s.
Literature: Adam and Rees (1966), Silas et al. (1982), Silas et al. (1986), Okutani et al. (1987), Sreenivasan and Sarvesan
(1990), Emam (1994), Filippova et al. (1995), Khaliluddin (1995), Nateewathana (1999).
112 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 177 Sepia prashadi
Known distribution
?? 200 m
Sepia ramani Neethiselvan, 2001 F i g . 1 7 8
Sepia ramani Neethiselvan, 2001, Indian Journal of Marine Sciences , 30: 82 [type locality: southeast coast of India,
Tuticorin, 08?47'N 78?09'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Large striped cuttlefish; Fr ? Grande seiche ray?e; Sp ? Sepia listada grande.
Diagnostic Features: Male and female arms
approximately subequal in length; arm formula in both
sexes IV, III, II, I; arm suckers tetraserial. Hectocotylus
present on left arm IV: 14 to 16 transverse rows of
normal size suckers proximally, followed by 7 to 10
rows of modified suckers, then rest normal to arm tip.
Suckers in 2 dorsal seriesminute, suckers in 2 ventral
series normal in size on modified portion of arm; 2
dorsal and 2 ventral series displaced laterally, with
fleshy ridge between. Club long; 15 to 24 suckers
enlarged, all enlarged suckers similar in size .
Swimming keel narrow, terminates at posterior end of
carpus. Dorsal and ventral protective membranes
extend slightly beyond carpus, not joined at base of
club. Buccal membrane with minute suckers.
Cuttlebone slender, elliptical; rugose dorsally, with
indistinct median and lateral ribs. Spine short, stout,
without keels. Sulcus deep, broad, extends length of
striated zone. Anterior striae are inverted V-shape.
Inner cone relatively short, limbs are narrow anteriorly,
broaden posteriorly, wit h distinctive, thick, bulbous
swelling ; outer cone narrow anteriorly, broadens
posteriorly. Colour: Dorsal mantle, head and arms dark
brown with transverse zebra stripe pattern (more
prominent in males than females, and more obvious in
fresh animals). Fins with pale, narrow, longitudinal line
along base.
Cephalopods of the World 113
male female
Fig. 178 Sepia ramani
male female
h e c t o c o t y l u s c u t t l e b o n e
t e n t a c u l a r
c l u b
ventral view
114 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Size: Up to 375 mm mantle length.
Geographical Distribution: Indian Ocean:
southern Indian Coast, Gulf of Mannar. Depth
ranges up to 100 m (Fig. 179).
Habitat and Biology: No detailed information is
available yet for this recently described species.
Interest to Fisheries: Sepia ramani is a
component of the commercia l f ishery
throughout the year along the Thoothukkudi
coast (southeast India).
Remarks: Sepia ramani is morphologically very
similar to Sepia pharaonis Ehrenberg, 1831.
Sepia pharaonis has 5 or 6 enlarged club
suckers, of which 3 or 4 suckers are greatly
enlarged, in contrast to S. ramani with 15 to 24
enlarged suckers that are all of similar size. In
addition, there are 10 to 12 transverse rows of
normal suckers on the base of the hectocotylus
in S. pharaonis, and 14 to 16 transverse rows of
normal suckers in S. ramani. The spawning
season of S. ramani seems to extend from
September to December.
Local Names: INDIA: Periya vari kanavai.
Literature: Neethiselvan, 2001.
Sepia recurvirostra Steenstrup, 1875 F i g . 1 8 0
Sepia recurvirostra Steenstrup, 1875, Danske Videnskabernes Selskabs Skrifter, 5 Raekke, Naturvidenskabelig og
Mathematisk , 10(7): 479 [type locality: South China Sea].
Frequent Synonyms: Sepia singaporensis Pfeffer, 1884.
Misidentifications: None.
FAONames: En ? Curvespine cuttlefish; Fr ? Seiche hame?on; Sp ? Sepia
ganchuda.
Fig. 179 Sepia ramani
Known distribution
?? 200 m
dorsal viewt e n t a c u l a r c l u b
Fig. 180 Sepia recurvirostra
c u t t l e b o n e
ventral view
Diagnostic Features: Mantle broad, oval. Non-hectocotylized arm sucker arrangement same in both sexes: suckers arms
I to III tetraserial proximally, biserial at distal tips ; arms IV suckers tetraserial . Hectocotylus present on left ventral arm:
sucker size normal proximally, reduced medially, then normal size suckers to arm tip. Suckers of hectocotylus in 2 dorsal
series smaller than those in 2 ventral series ; reduced suckers much smaller than normal arm suckers; suckers evenly
spaced on modified portion of arm. Club sucker-bearing surface flattened, with 5 or 6 suckers in transverse rows ; suckers
differ markedly in size: 5 or 6 median suckers enlarged . Swimming keel of club extends proximally slightly beyond carpus;
dorsal and ventral protective membranes not joined at base of club; dorsal membrane forms deep cleft at junction with
stalk. Buccal membrane without suckers. Cuttlebone outline oblong; bone very angular, V-shape anteriorly ; bluntly
rounded posteriorly; dorsal median and lateral ribs present. Chitin present as wide patch posteriorly and a narrow chitinous
rim borders lateral margins of cuttlebone. Spine curves ventrally. Last loculus concave; sulcus shallow, narrow, extends
along striated zone only. Anterior striae are inverted U-shape . Inner cone limbs are uniform width, narrow, U-shape
posteriorly, thickened posteriorly into a rough chitinous callus ; outer cone calcified, narrow anteriorly, broadens
posteriorly. Dorsal mantle pale with opalescent blue transverse stripes. Fins with pale reflective opalescent blue line
along base.
Size: Up to 170 mm mantle length and 0.4 kg weight.
Geographical Distribution: Western Pacific: Yellow
Sea, China, Hong Kong, East China Sea, Taiwan
Province of China, South China Sea, Philippines,
Celebes Sea, Java Sea, Gulf of Thailand, Singapore.
Indian Ocean: Andaman Sea and Myanmar, Bay of
Bengal (Fig. 181).
Habitat and Biology: Depth range from 10 to 140 m.
Sepia recurvirostra is a demersal species that inhabits
the continental shelf. In the Gulf of Thailand, spawning
occurs all year, with peak times from November to
February and July to September. The ratio of males to
females caught in the Gulf is 1:3.
Interest to Fisheries: The species has some
commercial importance in Hong Kong, where it is
caught in multispecies trawls. It is a commercial
species in the Gulf of Thailand, South and East China
Seas, and Japan. In Thailand, most cuttlefishes are
caught using otter trawl, with smaller catches reported
from pair trawl, squid light-lures, traps and push nets,
with bottom otter and pair trawls used offshore, and
push nets and lift nets used in inshore and coastal
waters. A few cuttlefishes are caught using purse seine
and hook and line. In the Gulf of Thailand, most
captured animals are between 40 and 130 mm mantle
length.
Local Names: CHINA: Jam mak yue; JAPAN:
Ajia-kouika.
Literature: Adam and Rees (1966), Chantawong and
Suksawat (1997).
Cephalopods of the World 115
90?
90?
100?
100?
110?
110?
120?
120?
 10?  10?
0? 0?
10? 10?
20? 20?
30? 30?
Fig. 181 Sepia recurvirostra
Known distribution
?? 200 m
116 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia rozella ( Iredale, 1926 ) F i g . 1 8 2
Solitosepia rozella Iredale, 1926, The Australian Zoologist , 4(3): 190 [type locality: New South Wales, Manly Beach,
33?48'S 151?17'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Rosecone cuttlefish; Fr ? Seiche au
c?ne ros?; Sp ? Sepia de cono rosado.
Diagnostic Features: Mantle oval; dorsal anterior
margin triangular, acute. Fins wide, rounded posteriorly,
with narrow gap between them. Head elongate; slender,
narrower than mantle. Male and female arm lengths
subequal. Arm suckers tetraserial; female arms I
protective membranes well developed, enveloping
suckers; arm tips attenuate. Male non-hectocotylized
arm suckers normal in size (not greatly enlarged).
Hectocotylus absent; the only modification being the
basal 40% of the left ventral arm is slightly thicker and
the oral surface of arm is wider than the corresponding
portion of the right ventral arm, with a shallow median
furrow. Club short, oval; sucker-bearing surface
flattened, with 4 or 5 suckers in transverse rows ;
suckers differ slightly in size: 4 or 5 enlarged suckers in
longitudinal series towards posterior end of club.
Swimming keel of club extends well proximal to carpus;
dorsal and ventral protective membranes joined at base
of club (V-shape); separated from tentacular stalk by
membrane; dorsal and ventral membranes same length;
dorsal membrane much wider than ventral membrane;
dorsal membrane forms deep cleft at junction with stalk.
Buccal membrane without suckers. Eggs spherical.
Cuttlebone oblong in outline; bone bluntly rounded anteriorly and posteriorly; recurved ventrally; dorsal surface creamy
white; dorsal surface evenly convex; entire surface calcified, granulose, sculpture reticulate on posterolateral half of bone.
Spine and posterior tip of bone covered with smooth glaze-like substance. Dorsal median rib and lateral ribs indistinct.
Chitin borders lateral margins of cuttlebone. Spine long, pointed, straight, directed dorsally, with ventral keel and ventral
notch at base. Striated zone concave; last loculus flat; sulcus deep, wide, distinct V-shape medially, flanked by rounded
ribs. Anterior striae are inverted V-shape . Inner cone limbs are narrow anteriorly, broaden posteriorly, rose coloured;
outer margin of inner cone raised into rounded, thickened posterior ledge ; outer cone calcified; narrow anteriorly,
broadens posteriorly; outer cone lateral limbs are flared ventrolaterally. Dorsal mantle covered with numerous small
papillae; ventral mantle with longitudinal row of 6 narrow ridges along each side close to fins; anteriormost pair and
posterior 2 pairs shorter than rest. Head papillose dorsally and laterally. Colour: Purplish brown. Head with
chromatophores concentrated over eye orbits. Arms without markings. Paired dorsal eye spots absent. Fins pale, without
markings at base. Dorsal ridges reddish purple.
Size: Up to 140 mm mantle length.
Geographical Distribution: Southwestern Pacific:
northeastern Australia, from southern Queensland,
27?42'S 153?35'E, to New South Wales, northeast of
Tathra, 36?41'S 150?02'E (Fig. 183).
Habitat and Biology: Depth range from 5 to 183 m.
Interest to Fisheries: Species taken as bycatch of
prawn and mixed species trawl fisheries.
Literature: Lu (1998a).
Fig. 182 Sepia rozella
c u t t l e b o n et e n t a c u l a r c l u b
ventral view
Fig. 183 Sepia rozella
Known distribution
?? 200 m
Sepia savignyi Blainville, 1827 F i g . 1 8 4
Sepia savignyi Blainville, 1827, Dictionnaire des Sciences Naturelles , 48: 285 [type locality: Red Sea].
Frequent Synonyms: None.
Misidentifications: Sepia plathyconchalis Filippova and
Khromov, 1991.
FAO Names: En ? Broadback cuttlefish; Fr ? Seiche gros
dos; Sp ? Sepia robusta.
Diagnostic Features: Mantle broad, oval; ventral mantle
margin emarginate. Fins wide. Male and female arms
subequal in length. Arm suckers tetraserial. Club straight,
slender; sucker-bearing surface convex, with 8 suckers in
transverse rows ; suckers differ slightly in size; several
suckers of inner 2 or 3 rows very slightly larger than
rest. Swimming keel of club terminates at proximal end of
carpus (approximately). Dorsal and ventral protective
membranes not joined at base of club , extend proximal to
carpus along stalk. Buccal membrane with a few minute
suckers. Cuttlebone outline oval; bone slightly acuminate,
acute, anteriorly; bluntly rounded posteriorly; calcified with
reticulate sculpture; dorsal median rib and lateral ribs
indistinct; median rib broadens anteriorly. Chitin borders
lateral and anterior margins of cuttlebone. Spine short,
pointed. Striated zone flat, or slightly concave posteriorly;
slightly convex anteriorly on each side of sulcus; striated
zone separated from outer cone by broad, smooth
marginal zones; last loculus flat; sulcus shallow, wide,
extends along striated zone only. Anterior striae are
inverted U-shape ; limbs of inner cone extend anteriorly to
approximately two-thirds length of striated zone. Inner
cone limbs are narrow anteriorly, broaden posteriorly;
outer margin of inner cone slightly raised as a slightly
rounded ridge; not thickened; dull, not shiny; outer cone
calcified, narrow anteriorly, broadens posteriorly. Colour:
Light brown.
Size: Up to 190 mm mantle length.
Geographical Distribution: Western Indian Ocean: Red
Sea, Gulf of Aden, Arabian Sea, Persian Gulf and
Saya-de-Malha Bank. Not recorded south of Socotra
Island (Fig. 185).
Habitat and Biology: Depth range from 20 to 50 m.
Interest to Fisheries: Not determined at present, but the
size and distribution of the species indicate that S. savignyi
is probably harvested in mixed species fisheries even
though not specifically targeted.
Remar ks : This species can be confused with
S. plathyconchalis Filippova and Khromov, 1991, from
which it differs by having a ventral median sulcus and a
normal striated zone in the cuttlebone. The striated zone is
very narrow in S. plathyconchalis.
Literature: Adam and Rees (1966), Filippova et al. (1995).
Cephalopods of the World 117
Fig. 184 Sepia savignyi
c u t t l e b o n et e n t a c u l a r c l u b
ventral view
Fig. 185 Sepia savignyi
Known distribution
?? 200 m
Sepia smithi Hoyle, 1885 F i g . 1 8 6 ; P l a t e V , 3 6
Sepia smithi Hoyle, 1885, Annals and Magazine of Natural History , series 5, 16: 190 [type locality: Arafura Sea, South of
Papua, 09?59'S 139?42'E].
Frequent Synonyms: Acanthosepion pageorum Iredale, 1954.
Misidentifications: Sepia whitleyana (Iredale, 1926).
FAO Names: En ? Smith?s cuttlefish; Fr ? Seiche de Smith; Sp ? Sepia de Smith.
Diagnostic Features: Mantle broad, oval. Fins
wide, anterior origin almost at mantle margin,
rounded posteriorly, with narrow gap between
them. Head elongate; slender, narrower than
mantle. Male and female arms subequal in length.
Arm suckers tetraserial. Hectocotylus present;
both ventral arms modified . Left ventral arm: 8
rows of normal size suckers (approximately)
proximally, 5 to 8 rows of greatly reduced
suckersmedially , then normal size suckers distally
to arm tip. Suckers of hectocotylus in 2 ventral
series are smaller than those in 2 dorsal series ;
oral surface of modified region not wide, fleshy, but
normal; without distinct median furrow. Male right
ventral arm: approximately 6 to 8 rows normal size
suckers proximally, 5 rows reduced suckers
medially, then normal size suckers to arm tip.
Suckers of hectocotylus in dorsal row are smaller
than those in 2 ventral series in modified region;
oral surface of modified region very wide, fleshy,
sponge-like in texture with pronounced median
depression; suckers in 2 dorsal series aligned in a
single row. Both ventral arms of males: suckers in 2
ventral series are displaced laterally, with gap
between, widely spaced (more obviously so on
right ventral arm). Club straight, slender, or slightly
recurved, long, sucker-bearing surface convex,
with 13 to 22 suckers in transverse rows ; all club
suckers of similar, minute, size . Swimming keel of
club terminates at proximal end of carpus. Dorsal
and ventral protective membranes not joined at
base of club but fused to tentacular stalk. Dorsal
and ventral membranes same length, terminate at
posterior end of carpus. Dorsal membrane forms
shallow cleft at junction with stalk. Buccal membrane without suckers; in females with single median spermathecae in
ventral part. Eggs spherical. Cuttlebone outline oblong; bone triangular, obtuse anteriorly; bluntly rounded posteriorly;
dorsal surface creamy white, or yellowish; dorsal surface evenly convex; entire surface calcified, granulose with sculpture
arranged in series of blunt, transverse, inverted V-shape ridges. Spine and posterior end (approximately one quarter) of
bone covered with ochre-coloured smooth g laze-like substance . Dorsal median rib and lateral ribs indistinct; median rib
broadens slightly anteriorly, bordered laterally by indistinct grooves. Chitin surrounds entire margin of cuttlebone. Spine
short, pointed, straight, directed dorsally, keel(s) absent . Striated zone concave; last loculus flat; sulcus shallow, wide,
extends along striated zone only. Anterior striae are inverted U-shape. Inner cone limbs are broad, strap-like anteriorly,
U-shape posteriorly; outer margin of inner cone raised into rounded, thickened, shiny, posterior ledge . Dorsal mantle
covered with numerous small papillae and with series of 7 or 8 elongate orange-pink papillae along each side, adjacent to
base of each fin. Head papillose dorsally and laterally. Colour: Pale buff pinkish brown, light brown, or greyish brown with
white blotches and spots, sometimes joined to form irregular transverse bands. Arms I to III have a longitudinal band of
orange-red pigment along their aboral surfaces . Arm and club sucker rims pale brown, or yellow brown. Dorsal mantle
has pinkish spots, or blotches and scattered purplish spots and blotches. Paired dorsal eye spots are present (both
sexes) . Fins in both sexes with longitudinal whitish band lateral to series o f orange-pink ridges overlain with regularly
spaced transverse purplish blotches .
Size: Males up to 140 mm mantle length; females up to 150 mm mantle length.
118 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 186 Sepia smithi
c u t t l e b o n et e n t a c u l a r c l u b
(illustrations: K. Hollis)
ventral view
Geographical Distribution: Indo-Pacific: northern
Australia from Shark Bay, Western Australia, 25?25'S
113?35'E, along the norther n coast, then southward to
Moreton Bay, Queensland, 27?25'S 153?20'E. Timor,
Arafura and Coral Seas (Fig. 187).
Habitat and Biology: Depth range from 33 to 138 m. In
the Gulf of Carpentaria, S. smithi has been caught
where bottom depths varied between 7 and 55 m,
typically on sand and mud. Sepia smithi has been
observed at night lying on the sea floor or burying into
the sand, its pale colours and skin textures matching
the sand surface. This lack of nocturnal activity
suggests the species may be day-active.
Interest to Fisheries: Species taken as bycatch of
prawn and mixed species trawl fisheries.
Remarks: Sepia smithi often is confused with
S. whitleyana (Iredale, 1926). The 2 species differ in
the nature of the cuttlebone inner cone, which is round,
shiny and thickened in S. smithi and thin, narrow and not shiny in S. whitleyana . The bases of the fins of S. whitleyana lack
the longitudinal whitish bands that are present in S. smithi. Male S. smithi lack the distinctive longitudinal white bars that
are present in male S. whitleyana . The cuttlebone is wider in S. whitleyana than in S. smithi, particularly in the anterior third
of the bone
Literature: Lu (1998a).
Sepia stellifera Homenko and Khromov, 1984 F i g . 1 8 8
Sepia stellifera Homenko and Khromov, 1984, Zoologicheskii Zhurnal , 63: 1150 [type locality: east Arabian Sea, 13?59'N
73?58'E].
Frequent Synonyms: None.
Misidentifications: Sepia brevimana Steenstrup, 1875.
FAO Names: En ? Starry cuttlefish; Fr ? Seiche ?toil?e;
Sp ? Sepia estrellada.
Cephalopods of the World 119
Fig. 187 Sepia smithi
Known distribution
?? 200 m
dorsal view
Fig. 188 Sepia stellifera (after Homenko and Khromov, 1984)
h e c t o c o t y l u s c u t t l e b o n et e n t a c u l a r c l u b
(after Homenko and Khromov, 1984)
ventral view
Diagnostic Features: Club with 10 suckers in transverse rows ; all club suckers of similar, small size. Dorsal and ventral
protective membranes not joined at base of club. Hectocotylus present; left ventral arm modified : suckers normal
proximally, reduced medially, then normal size suckers proximally to arm tip. Suckers of hectocotylus in 2 ventral series are
displaced laterally, with gap between; oral surface of hectocotylus with transversely grooved ridges. Cuttlebone outline
oval; bone very angular, V-shape anteriorly (subrhomboidal in anterior fourth); dorsalmedian rib very pronounced ; lateral
ribs present. Spine long, pointed, with dorsal and ventral keel . Sulcus deep, wide, extends along striated zone only;
sulcus with 3 deep, broad grooves and 2 ribs between them ; limbs of inner cone extend anteriorly to end of striated zone;
outer cone present. Colour: Dorsal mantle with many small, bright, brown-pink spots, encircled with green-blue rings, the
whole pattern resembling a starry sky.
Size: Up to 120 mm mantle length.
Geographical Distribution: Indian
Ocean: Arabian Sea and west coast of
India to Cape Comorin, Bay of Bengal,
Andaman Sea, Gulf of Thailand. Possibly
Saya-de-Malha Bank. Eastern extent of
range unknown (Fig. 189).
Habitat and Biology: Depth to 200 m.
Interest to Fisheries: This species is
fished commercially in India and may
figure in statistical data with Sepia
brevimana Steenstrup, 1875.
Remarks: Sepia stellifera is similar to
S. brevimana Steenstrup, 1875. It differs
from S. brevimana in having 10 oblique
club sucker rows, rather than 6 to 8, and
the sulcus is deep and wide, rather than
narrow and shal low as seen in
S. brevimana.
Literature: Homenko and Khromov, 1984.
Sepia sulcata Hoyle, 1885 F i g . 1 9 0
Sepia sulcata Hoyle, 1885, Annales and
Magazine of Natural History , (5)16: 192 [type
locality: Indonesia: Arafura Sea, off Kai Island,
05?49'15"S 132?14'15"E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Grooved cuttlefish; Fr ? Seiche
stri?e; Sp ? Sepia estriada.
Diagnostic Features: Sucker rows on arms I and
III in males biserial proximally, tetraserial distally
(approximately 8 to 11 rows biserial suckers
arms I, 6 to 8 rows biserial suckers arms II and III);
suckers on arms IV biser ia l at base
(approximately 2 rows), tetraserial distally. Sucker
rows on arms I to III in females biserial proximally
for 6 or 7 rows, tetraserial distally , suckers on
arms IV biserial proximally for 2 or 3 rows, rest
tetraserial. Distal arm tips strongly attenuate. Left
ventral arm hectocotylized: proximally 14 rows
(approximately) of greatly reduced suckers; 2
dorsal series smaller than 2 ventral series ; 2
dorsal and 2 ventral series widely spaced in
mature males. Oral surface wide, fleshy with deep
longitudinal furrow between swollen protective
membranes. Club with 5 to 7 small, subequal
120 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 189 Sepia stellifera
Known distribution
?? 200 m
?
?
Fig. 190 Sepia sulcata
ventral viewdorsal view
c u t t l e b o n et e n t a c u l a r c l u b
suckers in transverse rows. Dorsal and ventral protective membranes not joined at base of club. Cuttlebone oblong,
strongly acuminate, acute, anteriorly and posteriorly. A thick, cartilaginous subdermal layer adheres closely to dorsal
side of cuttlebone. Dorsal median and lateral ribs distinct; sulcus shallow, narrow, flanked by prominent rounded ribs
bordered laterally by shallow grooves. Inner cone limbs form narrow, raised ledge posteriorly . Dorsal mantle with up to
10 short, longitudinal, orange-pink ridges along each side close to fins; ventral mantle with 6 longitudinal ridges along
each side close to fins.
Size: Males up to 68 mm mantle length; females up to 97 mm mantle length.
Geographical Distribution: Indo-Pacific: Indonesia,
Arafura Sea, off Kai Islands, 05?49'15"S 132?14'15"E,
to Australia: Western Australia, North West Shelf,
19?58'S 115?13'E (Fig. 191).
Habitat and Biology: Depth range from 150 to 404 m.
Habitat mud, silt, rock.
Interest to Fisheries: Due to its size and distribution,
this species may occur in multispecies trawl catches.
Remarks: The depth range occupied by this species is
broad. There is some evidence to suggest that there
might be a migration into relatively shallower waters
for spawning. The cuttlebone of this species shows
some similarities with that of S. australis Quoy and
Gaimard, 1832 from South Africa. In both species, the
cuttlebone is narrow and oblong, distinctly pointed
anteriorly, with pronounced dorsal median and lateral
ribs and, on the ventral side, the lateral ribs in both
species are bordered by grooves. The ventral
grooves, and sulcus are much deeper and more
pronounced in S. australis than in S. sulcata, with the
sulcus extending along the last loculus in S. australis,
unlike that of S. sulcata that is present only in the
striated zone. Both species have a short, median ridge
on the dorsal side of the cuttlebone, anterior to the
spine. The posterior end of the cuttlebone is broader
in S. sulcata; in S. australis the outer cone is poorly
developed and the inner cone is not raised into a
ledge. Sepia australis is darkly pigmented, unlike
S. sulcata, and it has a distinctive, unbroken
longitudinal ridge at the base of the fins. The arm
suckers in S. australis are arranged in four series,
unlike those seen in S. sulcata.
Literature: Adam and Rees (1966), Reid (2000).
Cephalopods of the World 121
Fig. 191 Sepia sulcata
Known distribution
?? 200 m
Sepia trygonina ( Rochebrune, 1884) F i g . 1 9 2
Doratosepion trygoninum Rochebrune, 1884, Bulletin des Sciences par la Societe Philomatique de Paris , 7(8): 97 [type
locality: Red Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Trident cuttlefish;
Fr ? Seiche trident; Sp ? Sepia tridente.
Diagnostic Features: Mantle oblong, acuminate posteriorly. Male and female arms differ in relative lengths: male arms I
shorter than rest, female arm lengths subequal . Arm sucker arrangement differs between sexes: in males, arm suckers
tetraserial; in females, suckers on arms I and IV tetraserial, suckers arms II and III tetraserial proximally, biserial on
distal third of arms; suckers displaced laterally, with gap between , on distal ends of female arms II and III. Hectocotylus
present on left ventral arm: 6 rows normal size suckers proximally, suckers absent medially , then normal size suckers to
arm tip. Oral surface of modified region of hectocotylus concave and covered by large transversely grooved protective
membranes; ventral membrane wider than dorsal membrane , thick and curved inwards (similar to S. elongata). Club
short, oval; with 5 suckers in transverse rows; suckers differ markedly in size: 4 or 5 greatly enlarged suckers in
longitudinal series towards posterior end of club. Dorsal and ventral protective membranes not joined at base of club.
Swimming keel of club extends proximally slightly beyond carpus. Buccal membrane in females without spermathecae.
Cuttlebone outline lanceolate; strongly recurved ventrally; dorsal surface pinkish ; calcified medially, thickest posteriorly,
slightly granulose with irregular longitudinal ridges; dorsal median rib indistinct. Chitin present as wide bands bordering
lateral margins of cuttlebone. Spine curves dorsally, keel(s) absent . Sulcus shallow, wide , extends entire length of
cuttlebone; sulcus flanked by rounded ribs . Anterior striae are inverted U-shape; inner cone lateral limbs overlie
calcareous striated zone, anteriorly bordered and separated from outer cone by striated zone. Inner cone limbs are uniform
width, narrow, U-shape posteriorly ; slightly raised into rounded posterior ridge; outer cone limbs are expanded into 2 short
?wings?, directed ventrally, to form a recurved cup-like structure. Colour: Purplish brown. Base of fins in males with dark
purple band; adjacent to band (or partially on it) is a series of small oval or circular cream-coloured patches, often raised as
low tubercles.
Size: Up to 140 mm mantle length.
122 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
dorsal view, male
t e n t a c u l a r c l u b
Fig. 192 Sepia trygonina
c u t t l e b o n e
ventral view
Geographical Distribution: Indian Ocean: from
Saya-de-Malha Bank, 15?35'N?11?30'S, region of
Mascarene Ridge and Zanzibar to Red Sea, Persian
Gulf and southern India (Fig. 193).
Habitat and Biology: Depth from 20 to 410 m.
Interest to Fisheries: The fishery potential of the
species is presently undetermined. However, the size
and distribution of S. trygonina suggest that it probably
occurs in the local fisheries catches.
Remarks: The female of this species is very similar to
Sepia sokotriensis Khromov, 1988. The cuttlebone is
very similar to S. burnupi Hoyle 1904, but differs in
having a less well-defined dorsomedial rib. There is
sexual dimorphism in the relative lengths and
armament of the arms. In S. trygonina, the arms of
males are not as modified as in S. burnupi, and
females differ in having widely spaced suckers on the
distal ends of the lateral arms. The club and
hectocotylus resemble that of S. elongata d?Orbigny,
1839-1842, but the cuttlebone of S. elongata is thicker
and the inner cone and the striae differ.
Literature: Adam and Rees (1966), Filippova and
Khromov (1991), Filippova et al. (1995).
Cephalopods of the World 123
Fig. 193 Sepia trygonina
Known distribution
?? 200 m
Sepia vermiculata Quoy and Gaimard, 1832 F i g . 1 9 4
Sepia vermiculata Quoy and Gaimard, 1832, Voyage de decouvertes de l?Astrolabe pendant les annees
1826?1827?1828?1829, Zoologie , 2(1): 64 [type locality: Africa: South Africa, Cape of Good Hope].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Patchwork cuttlefish; Fr ? Seiche r?ticul?e;
Sp ? Sepia reticulada.
Diagnostic Features: Mantle broadly oval, anterior margin somewhat produced dorsally. Fins broad and rounded, well
separated posteriorly. Head short and broad. Arms joined by a shallow web, ventral arms longer than dorsal arms. Arms III
and IV keeled, arms II sometimes keeled, arms I usually not keeled; arm tips somewhat attenuated. Arm suckers
tetraserial; all suckers with finely toothed rings, distal teeth longer than proximal ones. Protective membranes well
developed. Hectocotylus present on left ventral arm: approximately 6 normal suckers basally, followed by 8 to 13 rows of
modified suckers , modified suckers much smaller and separated by transverse ridges on arm; distal portion of arm
normal. Tentacular club length approximately 1/3 of mantle length, with 3 or 4 suckers in transverse rows, small distally;
median suckers of proximal part variously enlarged , 1 1/2 to 3 times as large as marginal suckers. Rings of large club
suckers smooth. Swimming keel slightly shorter than club. Cuttlebone broadly oval, tapering slightly anteriorly and
posteriorly; spine present, surrounded by chitinous covering; dorsal surface tuberculate, with fairly broad chitinous
margins; no marked dorsal ridge. Striated zone about half of total cuttlebone length, with shallow sulcus; anterior border of
striated zone convex on either side of median ridge .
Size: Up to 287 mm mantle length.
124 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
dorsal view
Fig. 194 Sepia vermiculata
t e n t a c u l a r c l u b
dorsal view ventral view
c u t t l e b o n e
Geographical Distribution: Southwestern
Indian Ocean: South Africa, 30?42'S 15?59'E, to
central Mozambique, 19?S; Saya-de-Malha
Bank and region of Mascarene Ridge (Fig. 195).
Habitat and Biology: Little information is
available at present, but the species deserves
investigation, due to its potential to fisheries and
aquaculture. This is the only Sepia in southern
Africa that enters lagoons and river mouths (e.g.
Langebaan and Kysna lagoons, Kowie River),
and it is found from very shallow waters (few
metres) to 290 m.
Interest to Fisheries: No separate information
is available at present, but the species is
believed to have potential to fisheries, due to its
size and distribution.
Literature: Adam and Rees (1966), Roeleveld
(1972), Sanchez (1998), Fi l ippova and
Khromov (1991), Filippova et al. (1995).
Sepia vietnamica Khromov, 1987 F i g . 1 9 6
Sepia vietnamica Khromov, 1987, Asian Marine Biology , 4: 35 [type locality: South China Sea, Gulf of Tonkin, 18?00'N
107?08'E].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Viet Nam cuttlefish; Fr ? Seiche du Viet Nam; Sp ? Sepia de Viet Nam.
Diagnostic Features : Mant le broad, oval ,
dorsoventrally compressed. Fins wide, anterior origin
almost at mantle margin, rounded posteriorly. Male
and female arms subequal in length; ventral arms
slightly longer than rest. Arm suckers tetraserial
proximally, biserial at extreme distal tips , medial
suckers larger than lateral suckers on ventral arms.
Hectocotylus present on left ventral arm: sucker size
normal proximally for 8 to 11 rows , followed by 2 or
3 rows reduced suckers , suckers of normal size
medially, then distal third with reduced suckers to tip.
Suckers of hectocotylus in 2 dorsal series are
smaller than those in 2 ventral series ; dorsal
suckers partially covered by wide protective
membrane; reduced suckers much smaller than
normal arm suckers; arm with deep median furrow;
suckers in 2 dorsal and 2 ventral series displaced
latera l ly, wi th gap between them. Club
crescent-shaped, small; sucker-bearing surface
flattened, with 5 suckers in transverse rows ;
suckers differ slightly in size: 4 or 5 suckers enlarged
in second ventral row, with suckers decreasing in
size from this row to margins. Swimming keel of club
extends proximally slightly beyond carpus; dorsal
and ventral protective membranes not joined at
base of club. Buccal membrane without suckers.
Cuttlebone outline lanceolate, widest in anterior 1/3;
bone bluntly rounded anteriorly; acuminate, acute,
posteriorly; dorsal surface creamy white; dorsal
surface evenly convex; dorsal median rib indistinct;
Cephalopods of the World 125
Fig. 195 Sepia vermiculata
Known distribution
?? 200 m
Fig. 196 Sepia vietnamica
ventral view
c u t t l e b o n e
(after Khromov, 1987 )
t e n t a c u l a r c l u b
126 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
lateral ribs absent. Chitin (trace only) borders lateral margins of cuttlebone. Spine long, pointed, straight, keel(s) absent.
Striated zone and last loculus convex; sulcus deep, narrow, extends entire length of cuttlebone. Anterior striae M-, or
m-shape ; limbs of inner cone are very short. Inner cone limbs are uniform width, narrow V-shape posteriorly, thickened;
slightly raised into rounded posterior ridge; outer cone narrow throughout; outer cone limbs are expa nded posteriorly
into 2 short ?wings?, directed ventrally , to form a recurved cup-like structure . Dorsal mantle with longitudinal row of
ridge-like papillae along each side, adjacent to base of each fin (males only?); up to 6 fin papillae (approximately) in each
row; papillae present on aboral surfaces of arms I to III. Colour: Dark brown. Head with 2 small, crescent-shaped orange
spots near dorsal projection of mantle margin and above eyes . Arms I to III have spots on aboral surface as for mantle.
Fins pale with rows of large wine-coloured spots (in males and only at certain times).
Size: Up to 70 mm mantle length.
Geographical Distribution : Indo-Paci f ic :
northwestern South China Sea, off Viet Nam and
Taiwan Province of China. From original description,
type specimens were collected off Viet Nam between
11?34'N and 20?00'N. The northern and southern
extents of the range of this species have yet to be
determined (Fig. 197).
Habitat and Biology: Depth range from 23 to 104 m.
Type specimens were collected between 33 and 70 m.
Interest to Fisheries: No separate information is
available at present, but the species is likely to occur in
the catches within its distributional range.
Remarks: This species differs from Sepia lorigera,
W?lker 1910 in having subequal arms, while the 2
dorsal arms of S. lorigera are long in males. The
posterior region of the inner cone of the cuttlebone is
expanded in S. lorigera, and is an unexpanded furrow
in this species. There is no hectocotylus in S. lorigera.
Literature: Lu (1998b).
Fig. 197 Sepia vietnamica
Known distribution
?? 200 m
?
?
Sepia vossi Khromov, 1996 F i g . 1 9 8
Sepia vossi Khromov, 1996, Ruthenica , 5(2): 143 [type locality: east coast of Viet Nam].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Voss? cuttlefish; Fr ? Seiche de Voss; Sp ? Sepia de Voss.
Diagnostic Features: Arms short, subequal
in length; arm suckers tet raser ia l .
Hectocotylus present on left arm IV: 3 rows
of normal size suckers proximally, followed
by greatly reduced suckers, then rest
normal to arm tip. Suckers in 2 dorsal and 2
ventral series displaced laterally, with gap
bet ween, w idely spaced , crowded
together; oral surface wide, fleshy with
transversely grooved ridges. Club short,
broad with 3 to 5 suckers in transverse
rows ; suckers differ markedly in size: 3 to 5
suckers in middle of third longitudinal
row extremely large . Swimming keel
extends sl ight ly beyond club
sucker-bearing surface. Dorsal and ventral
protective membranes not joined at base
of club. Cuttlebone pinkish, blunt-pointed
anteriorly and posteriorly; median rib
distinct, lateral ribs indistinct. Spine with
dorsal and ventral keels. Sulcus shallow,
narrow, extends entire length of cuttlebone;
sulcus flanked by shallow furrows. Anterior
striae are inverted U-shape. Inner cone
limbs are uniform width, narrow, rounded,
into V-shape posterior ledge; outer cone
recurved, cup-like. Colour: Dorsal mantle
light brown with dark brown transverse
stripes.
Size: Up to 100 mm mantle length.
Geographical Distribution: Indo-Pacific: Hong Kong to southern
Viet Nam. Depth range from 2 to 140 m. Neritic demersal species
(Fig. 199).
Interest to Fisheries: Separate statistics are not available, but the
species is probably captured along with other demersal species.
Remarks: Treated under Sepia omani Adam and Rees, 1966, in
Voss and Williamson (1971) and S. rex (a synonym of S. hedleyi
Berry, 1918) in Khromov (1988).
Literature: Khromov, 1996.
Cephalopods of the World 127
Fig. 198 Sepia vossi
c u t t l e b o n e
tentacular club hectocotylus
(after Voss and Williamson, 1971)
ventral view
Fig. 199 Sepia vossi
Known distribution
?? 200 m
Sepia whitleyana ( Iredale, 1926 ) F i g . 2 0 0
Acanthosepion whitleyana Iredale, 1926, The Australian Zoologist , 4(3): 195 [type locality: New South Wales, Port
Macquarie, 31?27'S 152?55'E].
Frequent Synonyms: None.
Misidentifications: Sepia smithi Hoyle, 1885.
FAONames: En ? Whitley?s cuttlefish; Fr ? Seiche de Whitley; Sp ? Sepia de Whitley.
Diagnostic Features: Mantle broad, oval. Fins rounded posteriorly, with narrow gap
between them. Male and female arms subequal in length. Arm suckers tetraserial.
Hectocotylus present; both ventral armsmodified . Male left ventral arm: 6 to 8 rows
of normal size suckers proximally, 5 or 6 rows of reduced suckers medially , then
normal size suckers to arm tip. Suckers of hectocotylus in 2 dorsal series are much
smaller than those in 2 ventral series ; oral surface of modified region wide, slightly
fleshy (but not to same extent as right ventral arm). Male right ventral arm: 5 to 7 rows
normal size suckers proximally, 5 or 6 rows reduced suckersmedially , then normal
size suckers to arm tip; oral surface of modified region wide, fleshy, sponge-like in
texture with distinct median furrow ; suckers in 2 dorsal and 2 ventral series
displaced laterally, with wide gap between them ; reduced suckers all approximately
same size or dorsal and ventralmost rows may be smaller than median 2 rows on
proximal end of modified region. Club long; sucker-bearing surface flattened, with 12
to 21 small suckers in transverse rows . Swimming keel of club terminates at
proximal end of carpus. Dorsal and ventral protective membranes not joined at base
of club but fused to tentacular stalk. Dorsal and ventral membranes same length,
terminate at proximal end of carpus; approximately equal width; dorsal membrane
forms shallow cleft at junction with stalk. Buccal membrane without suckers; females
have single, median spermathecae in ventral part. Cuttlebone outline oblong;
acuminate, acute, anteriorly; bluntly rounded posteriorly; dorsal surface creamy
white; dorsal surface convex medially, flat laterally; granulose, or calcified medially.
Spine and posterior tip of bone covered with smooth glaze-like substance . Dorsal
median rib and lateral ribs absent or indistinct; sides of median rib approximately
parallel. Chitin surrounds entire margin of cuttlebone. Spine short, pointed, straight,
directed dorsally. Spine keel(s) absent . Striated zone deeply concave; last loculus
convex; sulcus deep, wide, extends along striated zone only; sulcus flanked by rounded ribs. Anterior striae are inverted
U-shape; limbs of inner cone short, extend anteriorly to approximately one-third the length of the striated zone. Inner cone
limbs are narrow anteriorly, broad U-shape posteriorly; slightly raised into flat, very n arrow, posterior ledge; ledge not
thickened, dull, not shiny ; outer cone calcified, narrow anteriorly, broadens posteriorly. Dorsal mantle with longitudinal
row of up to 9 ridge-like papillae along each side, adjacent to base of each fin. Colour: Bluish grey. Head with a few
scattered chromatophores. Arms I to III have a
longitudinal band of pale pigment along their aboral
surfaces. Arm and club sucker rims pale brown, or
yellow brown. Dorsal mantle has pinkish spots, or
blotches and narrow longitudinal whitish bands in males
only. Paired dorsal eye spots present. Fins without
markings at base , or with approximately six short, dark
transverse bars along each side.
Size: Up to 174 mm mantle length (both sexes).
Geographical Distribution: Southwestern Pacific:
eastern Australia from the Gulf of Carpentaria, 16?51'S
139?51'E, southward to o ff Port Stephens, 32?37'S
152?25'E. Western extent of range unknown (Fig. 201).
Habitat and Biology: Depth range from 0 to 128 m.
Interest to Fisheries: Species taken as bycatch of
prawn and mixed species trawl fisheries.
Remarks: Sepia whitleyana is often confused with
S. smithi Hoyle, 1885. See S. smithi Remarks for
differences.
Literature: Lu (1998a).
128 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 200 Sepia whitleyana
c u t t l e b o n e
ventral view
Fig. 201 Sepia whitleyana
Known distribution
?? 200 m
?
Cephalopods of the World 129
Sepia zanzibarica Pfeffer, 1884 F i g . 2 0 2
Sepia zanzibarica Pfeffer, 1884, Abhandlungen aus dem Gebiete der Naturwissenschaften, Hamburg , 8(1): 9 [type
locality: east Africa, Zanzibar].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Zanzibar cuttlefish; Fr ? Seiche de
Zanzibar; Sp ? Sepia de Zanzibar.
Diagnostic Features : Arm suckers tet raser ia l .
Hectocotylus present on left ventral arm: 6 rows of
reduced suckers medially , then normal size suckers
distally to arm tip; reduced suckers much smaller than
normal arm suckers; oral surface of modified region wide,
swollen, fleshy, with transversely grooved ridges; suckers
in 2 dorsal and 2 ventral series displaced laterally, with gap
between; 2 ventral series close together, rows alternate .
Club short, oval; with 6 suckers in transverse rows ;
suckers differ slightly in size; several suckers of inner 2 or 3
rows slightly larger than rest. Swimming keel of club
extends proximally slightly beyond carpus; dorsal and
ventral protective membranes not joined at base of club .
Buccal membrane with a few, minute suckers. Cuttlebone
outline oblong; bone bluntly rounded anteriorly and
posteriorly; strongly recurved ventrally; dorsal surface
evenly convex; entire surface calcified, granulose,
sculpture reticulate on posterolateral half of bone; dorsal
median rib indistinct; rib broadens anteriorly; lateral ribs
indistinct. Chitin borders lateral and anterior margins of
cuttlebone. Spine short, pointed, curves dorsally, keel(s)
absent. Striated zone concave; last loculus convex; sulcus
deep, wide , extends along striated zone only. Anterior
striae are inverted V-shape ; limbs of inner cone extend
anteriorly to approximately halfway along striated zone;
inner cone lateral limbs are separated from outer cone by
smooth zones. Inner cone limbs are narrow anteriorly,
broaden slightly posteriorly; raised into rounded
posteriorly thickened ledge ; outer cone calcified; narrow
anteriorly, broadens posteriorly.
Size: Up to 250 mm mantle length.
Geographical Distribution: Western Indian Ocean: from
the Gulf of Aden to South Africa, Port Elizabeth. Common
in open regions of the Indian Ocean, near islands including
Madagascar, Saya-de-Malha Bank, region of Mascarene
Ridge and Socotra (Fig. 203).
Habitat and Biology: Depth range from 20 to 125 m.
Interest to Fisheries: This species is commercially
important in the Gulf of Aden and is likely to constitute a
sizeable component of mixed species fisheries along the
eastern African coastal and open oceanic waters.
Literature: Adam and Rees (1966), Roeleveld (1972),
Filippova and Khromov (1991), Filippova et al. (1995).
Fig. 202 Sepia zanzibarica
c u t t l e b o n e
ventral view
t e n t a c u l a r c l u b
Fig. 203 Sepia zanzibarica
Known distribution
?? 200 m
Sepiella inermis (Van Hasselt, 1835 ) ( in F?russac and d?Orbigny, 1834?1848) Fig. 204; Plate VI, 3 7
Sepia inermis Van Hasselt, 1835 (in F?russac and d?Orbigny 1834?1848). Histoire naturelle g?n?rale et particuli?re
C?phalopodes Ac?tabuliferes vivants et fossiles , pl. 6bis [type locality: Indian Ocean].
Frequent Synonyms: Sepia (Sepiella) microcheirus Gray, 1849; Sepia affinis Eydoux and Souleyet, 1852; Sepiella
maindroni de Rochebrune, 1884.
Misidentifications: None.
FAO Names: En ? Spineless cuttlefish; Fr ? S?pia inerme; Sp ? Sepia inerme.
Diagnostic Features: Mantle oblong. Posterior gland and gland pore pigmented reddish. Protective membranes on arms
in both sexes wide, well-developed. Arm suckers tetraserial. Hectocotylus present on left ventral arm: 10 rows of reduced
suckers proximally. Club with 12 to 24 suckers in transverse rows (very variable depending upon geographic area).
Swimming keel of club slightly shorter than carpus; dorsal and ventral protective membranes not joined at base of club;
extend proximal to carpus along stalk as low ridges. Cuttlebone outline oval, broad; cuttlebone width 33 to 43% cuttlebone
length; strongly convex in lateral view ; granulose dorsally; dorsal median rib distinct. Spine absent. Striated zone and last
loculus convex; sulcus extends entire length of cuttlebone. Inner cone limbs are uniform width, narrow, inner cone U-shape
posteriorly, thickened, raised in the centre as a rounded knob; outer cone chitinous, spatulate, expanded. Colour: Greyish
brown. Dorsal mantle has more than 7 reddish patches adjacent to base of fins.
Size: Maximum mantle length about 125 mm.
130 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 204 Sepiella inermis
h e c t o c o t y l u s tentacular club
c u t t l e b o n e
lateral view
dorsal view
dorsal view ventral view
(after Adam, 1939 )
(after Adam, 1939 )
Geographical Distribution: Indian
Ocean: mouth of Zambezi River,
southern Red Sea, Gulf of Aden, Persian
Gulf, to Andaman Sea and southern
South China Sea, Gulf of Tonkin and
eastern Indonesia. The northern and
eastern extent of the distribution of this
species is not known (Fig. 205).
Habitat and Biology: Sepiella inermis is
a demersal, shallow water species found
to 40 m depth. Length at maturity varies
with stocks: on the east coast, off Waltair,
it is about 53 mm for males and 52 mm
for females and off Madras, about 56 mm
and 60 mm; off Cochin, however, on the
west coast, animals mature at larger
sizes: 81 mm and 83 mm, for males and
females, respectively. Females grow
larger than males and growth is
allometric in both sexes. Spawning
extends almost throughout the year, with
seasonal peaks corre lat ing wi th
environmental conditions. Off Kakinada and Waltair (northeast India) spawning occurs from June to September and from
November to December; off Madras, in September, December and March; off Portonovo from March to October; off Cochin
(southwest coast) in April and from September through October. The majority of spawning cuttlefishes are aged 9 to 12
months on the east coast and about 18 months on the west coast. Capsules containing several eggs are attached to
various substrates in very shallow waters. Growth varies according to environmental conditions. Hatchlings may attain 29
to 35 mm mantle length after 6 months. At the end of one year, they can grow to sizes ranging from 53 mm (Kakinada) to
61 mm (Cochin), while after 18 months they reach 74 mm (Waltair and Kakinada) to 82 mm (Madras). Longevity on the east
coast is estimated at 24 months, while on the west coast it is believed to be over 24 months. The diet of spineless cuttlefish
consists primarily of small demersal fishes and crustaceans; cephalopods are a minor food component. This species has
been reared successfully in aquaculture experiments conducted in Thailand since 1978 with the objective to develop
commercial mass culture techniques. Three generations have been reared in culture, fed on live mysids and postlarval
penaeid shrimp. The eggs are laid singly, are black and round, and have an incubation period that averages 12.6 days at
28?C. In culture, hatchlings were planktonic for 5 days bef ore changing to a benthic stage. Females mature at 60 days,
spawn at 87 days and produce about 500 eggs. Both sexes die after spawning, at an average age of 116 days. Maximum
observed life span was 149 days, with size of around 80 mm and weight around 65 grams. Size at first maturity was 50 mm
for males and 31 mm for females in experimental cultures in India.
Interest to Fisheries: The species is an object of several small fisheries and is caught in quantities in the Arabian Sea near
southern India and in the waters of Indo-China. Maximum lengths in the Indian trawl catches are around 110 mm on the
east coast and over 120 mm on the west coast off Cochin. Sepiella inermis is important to the commercial cuttlefish fishery
in Thailand, where it is highly abundant in the Gulf and in the Andaman Sea. In the Gulf of Thailand, most animals caught
commercially range between 20 and 80 mm mantle length, with the maximum size recorded at 105 mm mantle length.
Cuttlefishes in this region usually are caught using otter trawl or pair trawl, but squid light lures, traps, and to a lesser
extent, push nets, purse seines and hook-and-line also are used, with bottom otter and pair trawls used offshore, and push
nets and lift nets used in inshore and coastal waters. Sepiella inermis is one of the main commercial species in India and
Sri Lanka. In the Andaman Sea it is collected using push nets. Off East Java, in Probolinggo waters, this species has been
collected using bagan (lift net) in inshore areas. In shallow Saurashtra waters off Veraval (Gujarat, India), a continuous
decline in the annual landings and average annual stock observed from trawl fishery operations (1979?1983) was
attributed to high fishing pressure. This led to proposals to develop suitable regulatory measures for this fishery.
Remarks: Sepiella inermis is possibly a species complex; this should be examined given its fisheries significance. It is very
similar to S. cyanea Robson, 1924b.
Literature: Adam and Rees (1966), Silas et al. (1982), Unnithan (1982), Silas et al. (1986), Sudjoko (1987), Kasim (1988),
Appannasastry (1989), Filippova et al. (1995), Khaliluddin (1995), Chantawong and Suksawat (1997), Nabhitabhata
(1997).
Cephalopods of the World 131
Fig. 205 Sepiella inermis
Known distribution
?? 200 m
Sepiella japonica Sasaki, 1929 F i g . 2 0 6
Sepiella japonica Sasaki, 1929, Journal of the College of Agriculture, Hokkaido Imperial University , 20 (suppl.): 219 [type
locality: Japan Sea, Toyama Bay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Japanese spineless cuttlefish; Fr ? S?pia inerme japonaise; Sp ? Sepia inerme japonesa.
Diagnostic Features: Mantle oval; dorsal anterior margin triangular, obtuse. Posterior gland and gland pore present
between, and just below, the posterior fin lobes. Posterior half of ventral mantle sometimes stained brown with secretion
from pore. Fins widen gradually posteriorly. Arms relatively short. Arm suckers tetraserial. Hectocotylus present on left
ventral arm: 12 rows of reduced suckers proximally ; reduced suckers much smaller than normal arm suckers, ventral 2
rows of suckers close together, dorsal 2 rows separated. Suckers of hectocotylus in 2 dorsal and 2 ventral series are
displaced laterally, with gap between them. Club with 16 to 32 suckers in transverse rows (usually >20); all club suckers of
similar, minute, size. Cuttlebone outline broad, oval; cuttlebone width 30 to 35% cuttlebone length ; bone strongly convex
in lateral view; dorsal surface evenly convex; granulose; dorsal median rib and lateral ribs indistinct. Spine absent. Striated
zone and last loculus convex; sulcus shallow, narrow, extends entire length of cuttlebone. Anterior striae are inverted
U-shape striated zone with faint, radiating grooves, the median groove is more pronounced than the others. Inner cone
limbs are uniform width, narrow, U-shape posteriorly (rudimentary); inner cone with rounded posterior knob; outer cone
chitinous, spatulate, expanded. Colour: Greyish brown. Dorsal mantle has white blotches or spots. Fins with pale
reflective line along base . No spots or wine-coloured patches at base of fins. Ventral mantle pale but usually stained pale
red.
Size: Up to 180 mm mantle length, 600 g total weight.
132 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 206 Sepiella japonica
c u t t l e b o n e
ventral viewdorsal view
h e c t o c o t y l u s dorsal view
(after Sasaki,1929 )
(after Sasaki,1929 )
Geographical Distribution: Northwestern Pacific:
Russian Federation, Possiet Bay in southern
Primorye; Japan, from the Kanto region of Honshu, to
South Korea, East China Sea, South China Sea,
Taiwan Province of China, Kwang-chow (Canton),
Hong Kong (Fig. 207).
Habitat and Biology: Depth to 50 m. In the Hong Kong
area, the species is present in low numbers to the 50 m
contour, but peak occurrences, when huge numbers of
this species occur, have been reported. It spawns in
May off Japan, and February to March around Hong
Kong, varying with temperature. It is believed that in
years characterized by intensification of warm currents
this species migrates to areas in the northern Sea of
Japan (up to Vostok Bay and Peter the Great Bay,
Russian Federation).
Interest to Fisheries: Sepiella japonica is important in
the fisheries of Japan, South Korea, Taiwan Province
of China and China, caught in large numbers in bottom
trawls. In Japan, most of the catch is dried and
marketed as ?surume?. It is the dominant species of
cuttlefishes caught around the Chekiang and Kiangsu
provinces of China, with over 10 000 tonnes caught
annually, and is a component of the fishery in the
waters around Jiaozhou Bay. Around Hong Kong it is
normally not important for the fishery, except when
infrequent population peaks occur.
Local Names: CHINA: Ngor Huet Mak, Mo Jam Woo Chak; JAPAN: Harinashi-kouika, Shirikusari, Shiriyakeika.
Remarks: According to Khromov (1996), Duc?s (1978, 1993, 1997) records from Viet Nam possibly refer to S. inermis Van
Hasselt, 1835. The species has been reared successfully in aquaculture experiments under the name Sepiella maindroni
(Choe, 1966).
Literature: Adam and Rees (1966), Choe (1966), Ueda (1985), Okutani et al. (1987), Natsukari and Tashiro (1991),
Morozov (1997), Zhang et al. (1997).
Cephalopods of the World 133
Fig. 207 Sepiella japonica
Known distribution
?? 200 m
Sepiella ornata ( Rang, 1837 ) F i g . 2 0 8
Sepia ornata Rang, 1837, Magasin de Zoologie , 7(Cl.V.): 76 [type locality: Gulf of Guinea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Ornate cuttlefish; Fr ? S?pia orn?e; Sp ? Sepia ornada.
Diagnostic Features: Mantle oblong. Arm suckers tetraserial. Hectocotylus present on left ventral arm: 10 rows of greatly
reduced suckers proximally; suckers equal in size across rows; oral surface of modified region wide, swollen, fleshy, with
transversely grooved ridges. Suckers of hectocotylus in 2 dorsal and 2 ventral series are displaced laterally, with gap
between them. Club with 10 to 14 suckers in transverse rows . Cuttlebone much shorter than mantle, located in anterior
2/3 to 3/4 of mantle; outline oblong; cuttlebone width 24 to 27% cuttlebone length in males and 27 to 30% cuttlebone
length in females; bone strongly convex in lateral view ; acuminate, acute, anteriorly; bluntly rounded posteriorly. Spine
absent. Sulcus shallow, narrow, extends along striated zone only, flanked by rounded ribs; limbs of inner cone are short,
extend anteriorly to junction of striated zone and posterior termination of last loculus. Inner cone limbs are uniform width,
narrow V-shape posteriorly; outer cone chitinous, spatulate, expanded, recurved ventrally. Colour: Fin bases with row of
large wine coloured spots .
Size: Up to 100 mm mantle length.
134 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 208 Sepiella ornata
c u t t l e b o n e
ventral viewdorsal view
h e c t o c o t y l u s
dorsal view
lateral view
(after Adam,1939 )
Cephalopods of the World 135
Geographical Distribution: Eastern Atlantic: off west Africa
from Cape Blanco to Cape Frio (Fig. 209).
Habitat and Biology: Depth range from 20 to 150 m, but
most abundant in waters deeper than 50 m.
Interest to Fisheries: It is taken mostly as bycatch in bottom
trawls mixed with Sepia species, the highest yields coming
from waters deeper than 50 m.
Local Names: JAPAN: Hana-ika.
Literature: Okutani (1980), Roper and Sweeney (1981).
Sepiella weberi Adam, 1939 F i g . 2 1 0
Sepiella weberi Adam, 1939a, Part III. Revision du Genre Sepiella (Gray) Steenstrup, 1880. Siboga-Expeditie , 556: 98
[type locality: Timor, 8?35'S 126?00'E, and Soemba, 10?S 119?56'E, eastern Indonesia].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Web?s cuttlefish; Fr ? S?pia de Web; Sp ? Sepia de Web.
Fig. 209 Sepiella ornata
Known distribution
?? 200 m
Fig. 210 Sepiella weberi
dorsal view
c u t t l e b o n e
ventral viewdorsal view lateral view
(illustrations: K. Hollis/ABRS)
Diagnostic Features: Male and female arms subequal in length. Arm suckers tetraserial; 2 median series of suckers in
males are larger than dorsal and ventral marginal series, and larger than female arm suckers. Hectocotylus present on left
ventral arm: 11 or 12 rows of reduced suckers proximally ; suckers in 2 dorsal series smaller than those in 2 ventral series;
reduced suckers much smaller than normal arm suckers; oral surface of modified region wide, swollen, fleshy, with
transversely grooved ridges. Suckers of hectocotylus in 2 dorsal and 2 ventral series are displaced laterally, with gap
between, 2 ventral series close together, rows alternate. Club crescent-shaped, sucker-bearing surface convex, with 7 to
10 suckers in transverse rows . Dorsal and ventral protective membranes not joined at base of club. Swimming keel and
club protective membranes are equal in length to sucker-bearing surface of club . Gills with 27 or 28 lamellae per
demibranch. Buccal membrane without suckers. Cuttlebone outline oblong; cuttlebone width 21 to 31% cuttlebone length
in males, 30 to 33% in females; bone strongly recurved ventra lly; dorsal median rib indistinct; rib broadens anteriorly;
lateral ribs absent. Striated zone and last loculus convex; sulcus shallow, narrow, flanked by rounded ribs, extends entire
length of cuttlebone. Anterior striae are inverted U-shape, or wavy; limbs of the inner cone are short, thickened slightly;
shiny; outer cone chitinous, spatulate, expanded; without indentation on posterolateral margin. Colour: Purplish brown.
Dorsal mantle pale, peppered with scattered purple-black chromatophores. Fins with 5 or 6 oval orange-pink spots at
base in both sexes, spots slightly larger in males.
Size: Males up to 60 mm mantle length; females up to 70 mm mantle length.
Geographical Distribution: Indo-Pacific: Indonesia
(Timor?Soemba), 08?35'S 126?00'E to 10?S 119?56'E,
to northwestern Australia, 12?39'S 127?03'E to 19?35'S
117?14'E. South China Sea: Viet Nam (unconfirmed
records) (Fig. 211).
Habitat and Biology: Depth range from shallow waters
to 88 m (Australia). In contrast all other Sepiella
species, the ventral side of the cuttlebone is not
strongly convex medially in S. weberi .
Interest to Fisheries: Specific reports are not available,
but the species may occur in multispecies trawl catches
due to its size and distribution.
Literature: Lu (1998a), Reid and Lu (1998).
136 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 211 Sepiella weberi
Known distribution
?? 200 m
?
?
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES FOR WHICH ONLY
FEW RECORDS EXIST TO DATE
Sepia acuminata Smith, 1916
Sepia acuminata Smith, 1916, Proceedings of the Malacological Society of London , 12(1): 21 [type locality: South Africa:
Port Elizabeth and Natal, Tongaat Beach].
Size: Males up to 100 mm mantle length; females up to 120 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: southeast Africa from South Africa, Port Elizabeth?Somalia,
01?30'N?30?S, and Madagascar. Depth range from 44 to 369 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova et al. (1995).
Sepia adami Roeleveld, 1972
Sepia adami Roeleveld, 1972, Annals of the South African Museum , 59(10): 224 [type locality: South Africa: S 79?E off
Cape Natal].
Size: Females up to 59 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: known only from the type locality. Depth to 99 m.
Literature: Roeleveld (1972).
Sepia angulata Roeleveld, 1972
Sepia angulata Roeleveld, 1972, Annals of the South African Museum , 59(10): 242 [type locality: Africa: South Africa,
Bloubergstrand, 33?48'S 18?27'E].
Size: Up to 75 mm cuttlebone length.
Geographical Distribution: Southeastern Atlantic: Bloubergstrand, 33?48'S 18?27'E, to Still Bay, 34?23'S 21?25'E. Known
only from cuttlebones.
Literature: Roeleveld (1972).
Sepia appellofi W?lker, 1910
Sepia appellofi W?lker, 1910, Abhandlungen der mathematische-physikali sche Klasse der K?eniglich Bayerischen
Akademie der Wissenschaften, 3(Suppl. 1): 14 [type locality: Japan: Misaki].
Size: Up to 90 mm mantle length.
Geographical Distribution: Northwestern Pacific: Japan, Kyushu to south Honshu, Tsushima Strait. Depth to 350 m.
Literature: Adam and Rees (1966).
Cephalopods of the World 137
Sepia aureomaculata Okutani and Horikawa, 1987
Sepia aureomaculata Okutani and Horikawa, 1987. In: T. Okutani, M. Tagawa and H. Horikawa, Cephalopods from
continental shelf and slope around Japan : 62 [type locality: Japan: Tosa Bay].
Size: Up to 160 mm mantle length.
Geographical Distribution: Northwestern Pacific: Japan, Kii Channel, Tosa Bay, Hyuga-Nada and East China Sea. Depth
range from 190 to 350 m.
Literature: Okutani et al. (1987).
Sepia bartletti ( Iredale, 1954)
Blandosepia bartletti Iredale, 1954, Australian Zoologist , 12(1): 67 [type locality: Misima and the Conflict Group,
Louisiade Archipelago (southeast of Papua New Guinea)].
Size: Up to 74 mm mantle length.
Geographical Distribution: Western Pacific: known only from the type locality.
Remarks: Depth range unknown. Status uncertain, known only from type and six other specimens (all cuttlebones)
reported by Iredale, 1954. Possible synonym of Sepia bandensis Adam, 1939b.
Literature: Iredale (1954).
Sepia bathyalis Khromov, Nikitina and Nesis, 1991
Sepia bathyalis Khromov, Nikitina and Nesis, 1991, Zoologicheskii Zhurnal , 70(6): 13 [type locality: Madagascar, 22?19' to
22?23'S 43?06'E].
Size: Up to 80 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: northwestern and southwestern Madagascar. Depth range from
300 to 500 m.
Literature: Khromov et al. (1991).
Sepia baxteri ( Iredale, 1940)
Blandosepia baxteri Iredale, 1940, Australian Zoologist , 9(4): 442 [type locality: Lord Howe Island, 31?33'S 159?05'E].
Size: Up to 74 mm mantle length.
Geographical Distribution: Southwestern Pacific: Lord Howe Island, 31?33'S 159?05'E. Known only from type
cuttlebones. Depth range unknown. Possible synonym of Sepia bandensis Adam, 1939b.
Literature: Adam and Rees (1966).
138 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia bidhaia Reid, 200 0
Sepia bidhaia Reid, 2000, Invertebrate Taxonomy , 14: 4 [type locality: Australia: Queensland, 17?55.8'S 146?58.2'E].
Size: Males up to 37 mm mantle length; females up to 57 mm mantle length.
Geographical Distribution: Southwestern Pacific: Australia, Queensland, off the Great Barrier Reef, 17?20'S 146?41.5'E
to 22?07'S 153?19'E. Depth range from 200 to 304 m.
Literature: Reid (2000).
Sepia burnupi Hoyle, 1904
Sepia burnupi Hoyle, 1904, Journal of Conchology , 11(1): 27 [type locality: South Africa: Natal, Umkomaas].
Size: Up to 90 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: southeast Africa, from Port Elizabeth to southern Mozambique.
Saya-de-Malha Bank. Depth range from 40 to 240 m. Rare species.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova et al. (1995).
Sepia carinata Sasaki, 1920
Sepia carinata Sasaki, 1920, Proceedings of the United States National Museum , 57(2310): 192 [type locality: Japan:
Sagami Sea, 35?04'50''N 139?38'18''E].
Size: Up to 60 mm mantle length.
Geographical Distribution: Western Pacific: southern Japan, Sagami Bay, South China Sea, Viet Nam. Depth range lower
sublittoral, to 128 m.
Remarks: This is likely to be a very rare species as it has not been recorded from Japanese waters since its original
description.
Literature: Sasaki (1920).
Sepia chirotrema Berry, 1918
Sepia chirotrema Berry, 1918. Biological Results Fishing Experiments ?Endeavour? 1909?1914 , 4(5): 268 [type locality:
South Australia, Investigator Strait area, 35?25' S 137?22'E, south of Kangaroo Island, 35?50'S 137?15'E].
Size: Up to approximately 200 mm mantle length.
Geographical Distribution: Southern Indo-Pacific: Southern Australia from Investigator Strait, 35?25'S 137?22'E, to
Western Australia, Dirk Hartog Island, 25?45'S 113?03'E. Depth range from 120 to 210 m.
Literature: Lu (1998a).
Cephalopods of the World 139
Sepia confusa Smith, 1916
Sepia confusa Smith, 1916, Proceedings of the Malacological Society of London , 12(1): 24 [type locality: South Africa:
Natal, Port Elizabeth, Tongaat Beach].
Size: Up to 150 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: southeastern Africa from Port Elizabeth to south Mozambique,
Zanzibar and Pemba, 5? to 30?S. Saya-de-Malha Bank. M adagascar. Record from the Arabian Sea (Zuev, 1971)
erroneous, while record from Saya-de-Malha Bank (Nesis, 1993) has not been confirmed by more recent expeditions.
Depth range from 53 to 352 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova et al. (1995).
Sepia cottoni Adam, 1979
Sepia cottoni Adam, 1979, Records of the Western Australian Museum , 7(2): 193 [type locality: Australia: Western
Australia, west of Lancelin, 31?54'S 114?55'E].
Size: Up to 55 mm mantle length.
Geographical Distribution: Southeastern Indian Ocean: Australia, Western Australia, northwest of Broome, 17?31.5'S
121?27'E, to Rottnest Island, Armstrong Poin t, 32?S 115?30'E. Depth range from 83 to 183 m.
Literature: Reid (2000).
Sepia dollfusi Adam, 1941
Sepia dollfusi Adam, 1941b, Bulletin du Mus?e royal d?Histoire naturelle de Belgique , 17(62): 12 [type locality: southern
Red Sea: P?rim Island].
Size: Up to 110 mm mantle length.
Geographical Distribution: Red Sea and southern part of Suez Canal. Depth range unknown.
Literature: Adam and Rees (1966), Gabr (1999).
Sepia dubia Adam and Rees, 1966
Sepia dubia Adam and Rees, 1966, John Murray Expedition 1933? 3 4, Scientific Reports , 11(1): 119 [type locality: Africa:
South Africa, False Bay, 34?11'S 18?27'E].
Size: Up to 17 mm mantle length.
Geographical Distribution: Southeastern Atlantic: known only from the type locality. Depth 25 m.
Literature: Roeleveld (1972).
140 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia elongata d?Orbigny, 1839 ?1842 ( in F?russac and d?Orbigny 1834?1848)
Sepia elongata d?Orbigny, 1839?1842 (in F?russac and d?Orbigny), Histoire naturelle g?n?rale et particuli?re
C?phalopodes Ac?tabuliferes vivants et fossiles , pl. 24 [type locality: Red Sea, near Cossier].
Size: Up to 97 mm mantle length.
Geographical Distribution: Northwestern Indian Ocean: Red Sea to Somalia. Depth range unknown.
Literature: Adam and Rees (1966).
Sepia erostrata Sasaki, 1929
Sepia erostrata Sasaki, 1929, Journal of the College of Agriculture, Hokkaido Imperial University , 20(supplement): 183
[type locality: Japan: Manazuru, Kanagawa Prefecture].
Size: Up to 90 mm mantle length.
Geographical Distribution: Northwestern Pacific: off western mainland Japan, Sagami Bay to Kii Peninsula. Subtidal,
inshore.
Literature: Okutani et al. (1987).
Sepia faurei Roeleveld, 1972
Sepia faurei Roeleveld, 1972, Annals of the South African Museum , 59(10): 251 [type locality: South Africa, 88 km
southeast of Cape Seal, 35?00'S 23?07'E].
Size: Up to 21 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: South Africa, S 14?E(?) of Cape Seal 88 km, 35?00'S 23?07'E
(to the east of the Cape of Good Hope). Depth to 168 m.
Literature: Roeleveld (1972).
Sepia foliopeza Okutani and Tagawa, 1987
Sepia foliopeza Okutani and Tagawa, 1987, In: T. Okutani, M. Tagawa and H. Horikawa, Cephalopods from continental
shelf and slope around Japan: 66 [type locality: East China Sea].
Size: Up to 110 mm mantle length.
Geographical Distribution: Western Pacific: East China Sea and Taiwan Province of China. Depth range unknown.
Literature: Okutani et al. (1987), Lu (1998b).
Cephalopods of the World 141
Sepia gibba Ehrenberg, 1831
Sepia gibba Ehrenberg, 1831, In: C.G. Ehrenberg, 1828?1845. Symbolae Physicae, seu Icones et descriptiones
Corporum Nauralium novorum aut minus cognitorum, P.C. Hemprich et C.G. Ehrenberg, Pars Zoologica : 5 [type locality:
Red Sea].
Size: Up to 100 mm mantle length.
Geographical Distribution: Red Sea. Depth range unknown, at least as shallow as 1 m.
Literature: Adam and Rees (1966).
Sepia hieronis ( Robson, 1924)
Rhombosepion hieronis Robson, 1924a, Proceedings of the Zoological Society of London , 1924(2): 645 [type locality:
South Africa, Cape Town, 32? 32' to 33?03'S 17?29' to 17?42'E].
Size: Up to 70 mm mantle length.
Geographical Distribution: Southeastern Atlantic and southwestern Indian Ocean: west Africa, southern Namibia, from
approximately 27?S to Port Alfred, Sout h Africa, and east Africa from 17?S to Kenya and Mozambique. Saya-de-Malha
Bank. Depth range from 43 to 500 m (most abundant at 110 to 250 m).
Literature: Adam and Rees (1966), Augustyn et al. (1995), Filippova et al. (1995).
Sepia incerta Smith, 1916
Sepia incerta Smith, 1916, Proceedings of the Malacological Society, London , 12: 23 [type locality: South Africa: Tongaat
Beach, Natal; Port Elizabeth, Cape Province].
Size: Up to 150 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: south and east Africa, from Port Elizabeth to Mozambique (north
to 18?S). Saya-de-Malha Bank. Depth range from 90 to 345 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova et al. (1995).
Sepia insignis Smith, 1916
Sepia insignis Smith, 1916, Proceedings of the Malacological Society, London , 12: 25 [type locality: Africa: South Africa,
Natal, Tongaat, on beach, 29?35'S 31?07'E (cuttlebones only)].
Size: Up to 60 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: Africa, south Africa, from Cape of Good Hope to Natal. Depth to
42 m.
Literature: Adam and Rees (1966), Roeleveld (1972).
142 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia irvingi Meyer, 1909
Sepia irvingi Meyer, 1909, Die Fauna S?dwest-Australiens , 2(19): 333 [type locality: Western Australia, Cockburn Sound,
Garden Island, 32?11'S 115?43'E, Port Royal (King G eorge Sound), 35?03'S 117?58'E, and Warnbro Sound, 32?20'S
115?43'E].
Size: Up to 100 mm mantle length.
Geographical Distribution: Southeastern Indian Ocean: Western Australia, from Cockburn Sound to North West Shelf,
18?49'S 118?29'E. Depth range from 130 to 170 m.
Literature: Lu (1998a).
Sepia ivanovi Khromov, 1982
Sepia ivanovi Khromov, 1982, Zoologichesky Zhurnal , 61(1): 137 [type locality: southeast Africa: Kenya, Mombasa,
04?03'S 40?00'E].
Size: Up to 70 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: probably throughout southeast Africa, Kenya, Mozambique to
mouth of Zambezi River. Depth to 50 m.
Literature: Khromov (1982).
Sepia joubini Massy, 1927
Sepia joubini Massy, 1927, Annals of the South African Museum , 25: 161 [type locality: Cape Natal, Tugela River Mouth,
Cape Natal, The Bluff, Durban].
Size: Females up to 64 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: South Africa, off Tugela River Mouth, 29?11'S 31?25'E, to Cape
Natal. Off southern Mozambique, 24?S to 25?S. Sa ya-de-Malha Bank. Depth range from 66 to 170 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova et al. (1995).
Sepia kiensis Hoyle, 1885
Sepia kiensis Hoyle, 1885, Annals and Magazine of Natural History , (series 5)16: 194 [type locality: Arafura Sea, off Kai
Island].
Size: Up to 37 mm mantle length
Geographical Distribution: Indo-Pacific: Kai Islands, possibly to Timor and northern Australia. Depth up to 256 m.
Literature: Adam and Rees (1966).
Cephalopods of the World 143
Sepia koilados Reid, 200 0
Sepia koilados Reid, 2000, Invertebrate Taxonomy , 14: 24.
Size: Males up to 68 mm mantle length; females up to 58 mm mantle length.
Geographical Distribution: Type locality: Australia: Western Australia, North West Shelf, 18?36'S 118?2'E to 18?39'S
118?4'E. General distributi on: southeastern Indian Ocean: Western Australia, North West Shelf: 18?31'S 118?09'E to
19?12.6'S 116?25.8'E. D epth range from 182 to 203 m.
Literature: Reid (2000).
Sepia limata ( Iredale, 1926 )
Arctosepia limata Iredale, 1926, The Australian Zoologist , 4: 193 [type locality: Australia: New South Wales, Manly Beach,
33?48'S 151?17'E].
Size: Males up to 35 mm mantle length; females up to 42 mm mantle length.
Geographical Distribution: Southwestern Pacific: Australia, southern Queensland to New South Wales, 26?30'S
153?44'E to 34?40'S 150?51'E. Depth range from 17 to 183 m.
Literature: Reid (2000).
Sepia mascarensis Filippova and Khromov, 1991
Sepia mascarensis Filippova and Khromov, 1991, Zoologichesky Zhurnal , 70(8): 63 [type locality: Indian Ocean,
Saya-de-Malha Bank].
Size: From 67 to 124 mm mantle length.
Geographical Distribution: Western Indian Ocean: Saya-de-Malha Bank, 11?31'S 61?00'E, Mascarene Ridge,
Cargados-Carajos Shoals. Depth range from 87 to 325 m.
Literature: Khromov et al. (1991), Filippova et al. (1995).
Sepia mestus Gray, 1849
Sepia mestus Gray, 1849. Catalogue of the Mollusca in the British Museum. Part I. Cephalopoda Antepedia , 108 [type
locality: Australia].
Size: Males up to 77 mm mantle length; females up to 124 mm mantle length.
Geographical Distribution: Southwestern Pacific: Queensland: Escape Reef, 15?47'S 145?47'E t o New South Wales,
Jervis Bay, Murrays Beach, 35?08'S 150?46'E. Records from C hina and Viet Nam are now known to be misidentifications.
Sepia mestus is an Australian endemic. Depth range from 0 to 22 m.
Literature: Lu (1998a), Reid and Lu (2005).
144 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia mira (Cotton, 1932 )
Tenuisepia mira Cotton, 1932, Records of the South Australian Museum , 4(4): 546 [type locality: Queensland,
North-West Islet, Capricorn Group, 23?18'S 151?42'E].
Size: Up to 55 mm mantle length.
Geographical Distribution: Southwestern Pacific: Australia, New South Wales, Clarence R., 29?19'S 153?29'E, to off
Wooli, 29?49'S 153?27'E. Depth range from 20 to 72 m.
Literature: Lu (1998a), Reid (1998).
Sepia mirabilis Khromov, 1988
Sepia mirabilis Khromov, 1988, Zoologicheskii Zhurnal , 67: 785 [type locality: Arabi an Sea, 12?22'N 54?28'E].
Size: Up to 70 mm mantle length.
Geographical Distribution: Western Indian Ocean: near Sokotra Island, probably to east Africa. Depth to 50 m.
Literature: Khromov (1988).
Sepia novaehollandiae Hoyle, 1909
Sepia novaehollandiae Hoyle, 1909, Proceedings of the Royal Physical Society of Edinburgh , 17(6): 266 [type locality:
South Australia, Kangaroo Island, 35?50'S 138?03'E].
Size: Up to 77 mm mantle length.
Geographical Distribution: Southern Indo-Pacific: Australia from Shellharbour, NSW, 34?35'S 150?52'E to Western
Australia, North West Shelf, 18?57'S 118?45'E. Depth range from 15 to 348 m.
Literature: Adam and Rees (1966), Lu (1998a).
Sepia papillata Quoy and Gaimard, 1832
Sepia papillata Quoy and Gaimard, 1832, Voyage de decouvertes de l 'Astrolabe pendant les annees
1826?1827?1828?1829, Zoologie , 2(1): 61 [type locality: Africa: South Africa, Cape of Good Hope].
Size: Up to 140 mm mantle length.
Geographical Distribution: Southeastern Atlantic and southwestern Indian Ocean: South Africa, from L?deritz Bay,
26?11'S 15?10'E, to Natal coast off Tugela and Umvoti River s, 29?11'S 31?25'E. Mascarene Ridge. Depth range from 26 to
210 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova and Khromov (1991), Augustyn et al. (1995), Filippova
et al. (1995).
Cephalopods of the World 145
Sepia pardex Sasaki, 1913
Sepia pardex Sasaki, 1913, Doubutsugaku Zasshi [Zoological Magazine, Tokyo] , 25(292): 74 [type locality: Japan: Chiba
Prefecture, Kajiyama].
Size: Up to 230 mm mantle length.
Geographical Distribution: Western Pacific: Japan, along the Pacific coast from the Chiba Peninsula and along the Japan
Sea coast from Toyama Bay to South Korea, Taiwan Province of China and the East China Sea. Depth range unknown.
Literature: Okutani et al. (1987), Lu (1998b). Also cited as Sepia pardalis Sasaki, 1914.
Sepia peterseni Appell?f, 1886
Sepia peterseni Appell?f, 1886, Svenska Vetenskaps-Akademiens Handlingar , 21(13): 23 [type locality: Japan].
Size: Up to 120 mm mantle length.
Geographical Distribution: Western Pacific: Japan, south of central Honshu to southern Kyushu, South Korea. Depth
range from 20 to 100 m; inner shelf.
Literature: Okutani et al. (1987).
Sepia plana Lu and Reid, 1997
Sepia plana Lu and Reid, 1997, Records of the Western Australian Museum , 18: 279 [type locality: southeastern Indian
Ocean: Australia: Western Australia, North West Shelf, 14?15'S 121?59'E].
Size: Males up to 99 mm mantle length; females up to 151 mm mantle length.
Geographical Distribution: Australia: Western Australia, North West Shelf, 12?04'S 122?59'E to 17?55.5'S 118?16.0'E.
Depth range from 396 to 505 m. Cuttlebones found off eastern Australia.
Literature: Lu and Reid (1997).
Sepia plathyconchalis Filippova and Khromov, 1991
Sepia plathyconchalis Filippova and Khromov, 1991, Zoologichesky Zhurnal , 70(8): 66 [type locality: Indian Ocean,
Saya-de-Malha Bank, 09?42'S 61?08'E].
Size: From 8 to 63 mm mantle length.
Geographical Distribution: Western Indian Ocean: Saya-de-Malha Bank; Cargados-Carajos Shoals; St Brandon Shoals.
Shallow water species, depth range from 25 to 63 m.
Literature: Filippova and Khromov (1991).
146 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia pulchra Roeleveld and Liltved, 1985
Sepia pulchra Roeleveld and Liltved, 1985, Annals of the South African Museum , 96(1): 2 [type locality: Africa: South
Africa, Cape Peninsula, Llandudno].
Size: Up to 22 mm mantle length.
Geographical Distribution: Southeastern Atlantic: South Africa, Cape Peninsula. Depth range from 15 to 50 m.
Literature: Boletzky and Roeleveld (2000).
Sepia reesi Adam, 1979
Sepia reesi Adam, 1979, Records of the Western Australian Museum , 7(2): 200 [type locality: Western Australia, Rottnest
Island, Salmon Bay].
Size: Up to 45 mm mantle length.
Geographical Distribution: Southeastern Indian Ocean: cuttlebone known only from the type locality.
Literature: Adam (1979).
Sepia rhoda ( Iredale, 1954)
Arctosepia rhoda Iredale, 1954, Australian Zoologist , 12(1): 75 [type locality: Australia: Western Australia, Point Cloates,
22?43'S 113?40'E].
Size: Males up to 61 mm mantle length; females up to 58 mm mantle length.
Geographical Distribution: Indo-Pacific: Australia, Northern Territory, Arafura Sea, 10?24'S 130?23'E to Western
Australia, NW Shelf, 20?47'S 114?48'E. Depth range from 64 to 184 m.
Literature: Reid (2000).
Sepia robsoni ( Massy, 1927 )
Rhombosepion robsoni Massy, 1927, Annals of the South African Museum , 25(4): 159 [type locality: Africa: South Africa,
Hout Bay].
Size: Up to 20 mm mantle length.
Geographical Distribution: Southeastern Atlantic: South Africa, Hout Bay. Known only from the type locality. Depth range
from 17 to 37 m.
Literature: Adam and Rees (1966), Roeleveld (1972).
Cephalopods of the World 147
Sepia saya Khromov, Nikitina and Nesis, 1991
Sepia saya Khomov, Nikitina and Nesis, 1991, Zoologicheskii Zhurnal , 70(6): 13 [type locality: western Indian Ocean,
Saya-de-Malha Bank, 10?30'S 61?10'E to 10?32'S 61?11'E].
Size: Up to 90 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: known only from the type locality. Depth range from 87 to 117 m.
Literature: Khromov et al. (1991).
Sepia senta Lu and Reid, 1997
Sepia senta Lu and Reid, 1997, Records of the Western Australian Museum , 18: 286 [type locality: Australia: Western
Australia, 14?07'S 122?06'E].
Size: Males up to 62 mm mantle length; females up to 83 mm mantle length.
Geographical Distribution: Southeastern Indian Ocean: Western Australia, North West Shelf, 12?04'S 122?51'E to
18?44'S 116?59'E. Possibly Indonesia. A very similar, probably c losely related animal has been found in the Philippines.
Depth range from 256 to 426 m.
Literature: Lu and Reid (1997).
Sepia sewelli Adam and Rees, 1966
Sepia sewelli Adam and Rees, 1966. Scientific Reports. The John Murray Expedition , 11(1): 61 [type locality: Africa: near
Cape Guardafui, Somalia, 11?57'12''N 50?35'00''E to 11?56'42''N 50?39'12''E].
Size: Up to 30 mm mantle length.
Geographical Distribution: Western Indian Ocean: from Cape Guardafui, Somalia, 11?57'N 50?35' E to 05?40'S 39?17'E,
to Zanzibar and probably Madagascar. Depth range from 37 to 238 m.
Literature: Adam and Rees (1966).
Sepia simoniana Thiele, 1920
Sepia simoniana Thiele, 1920, Deutsche Sudpolar-Expedition 1901?1903 , 16(Zoology 8): 436 [type locality: Africa: South
Africa Simon?s Bay].
Size: Up to 185 mm mantle length.
Geographical Distribution: Western Indian Ocean: from Cape Town to Agulhas Bank, north to northern Kenya and
southern Mozambique. Saya-de-Malha Bank. Depth to 190 m (usually less than 100 m).
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova and Khromov (1991), Augustyn et al. (1995), Filippova
et al. (1995).
148 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia sokotriensis Khromov, 1988
Sepia sokotriensis Khromov, 1988, Zoologichesky Zhurnal , 67(5): 788 [type locality: Peoples? Democratic Republic of
Yemen, Arabian Sea, Sokotra Island, 12?10'N 54?25'E].
Size: Up to 80 mm mantle length.
Geographical Distribution: Western Indian Ocean: near Sokotra Island, probably east Africa. Depth to 100 m.
Literature: Khromov (1988).
Sepia subplana Lu and Boucher-Rodoni, 2001
Sepia subplana Lu and Boucher-Rodoni, 2001, M ?moires du Mus?um National d?Histoire Naturelle , 185: 370 [type
locality: southwest Pacific, Bayonnaise Bank, 11?54'S 179?32'W].
Size: Male 60 mm mantle length; female 55 mm mantle length (known only from types).
Geographical Distribution: Southwestern Pacific: known only from the type locality. Depth range from 400 to 600 m.
Literature: Lu and Boucher-Rodoni (2001).
Sepia subtenuipes Okutani and Horikawa, 1987
Sepia subtenuipes Okutani and Horikawa, 1987. In: T. Okutani, M. Tagawa and H. Horikawa, 1987, Cephalopods from
continental shelf and slope around Japan : 57 [type locality: Japan].
Size: Up to 94 mm mantle length.
Geographical Distribution: Western Pacific: southwestern Japan: Tosa Bay, Shikoku, Kii Channel. East China Sea. Depth
range from 90 to 300 m.
Literature: Okutani et al. (1987).
Sepia tala Khromov, Nikitina and Nesis, 1991
Sepia tala Khromov, Nikitina and Nesis, 1991, Zoologischesky Zhurnal , 70(6): 17 [type locality: southwest Madagascar:
off Cape Tala, 22?19'S to 22?23'S, 43?06'E].
Size: Up to 80 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: southwestern Madagascar off Cape Tala. Known only from the
type locality. Depth range from 325 to 332 m.
Literature: Khromov et al. (1991).
Cephalopods of the World 149
Sepia tanybracheia Reid, 200 0
Sepia tanybracheia Reid, 2000, Invertebrate Taxonomy , 14: 47 [type locality: Australia: Western Australia, 12?36'S
123?45'E].
Size: Males up to 51 mm mantle length.
Geographical Distribution: Southeastern Indian Ocean: known only from type locality. Depth range from 200 to 205 m.
Literature: Reid (2000).
Sepia tenuipes Sasaki, 1929
Sepia tenuipes Sasaki, 1929, Journal of the College of Agriculture, Hokkaido Imperial University , 20(supplement): 193
[type locality: Japan: Ibaraki Prefecture, Isohama].
Size: Up to 105 mm mantle length.
Geographical Distribution: Western Pacific: southwestern Japan, eastern Honshu and western Japan Sea to south of
Kyushu, Korea and East China Sea. Depth range from 100 to 250 m.
Literature: Okutani et al. (1987), Kubodera and Yamada (1998).
Sepia thurstoni Adam and Rees, 1966
Sepia thurstoni Adam and Rees, 1966, John Murray Expedition 1933? 3 4, Scientific Reports , 11(1): 2 [type locality: India,
Madras, Ramesvaram Island, 09?18'N 79?19'E].
Size: Up to 110 mm mantle length.
Geographical Distribution: Indian Ocean: India, off Madras. Sri Lanka: off Negombo and Hambantota. Depth range from
20 to 40 m.
Literature: Adam and Rees (1966).
Sepia tokioensis Ortmann, 1888
Sepia tokioensis Ortmann, 1888, Zoologische Jahrbucher, (Systematik) , 3: 653 [type locality: Japan: Tokyo Bay].
Size: Up to 90 mm mantle length.
Geographical Distribution: Western Pacific: Japan, from Tsugaru Strait to Kyushu and the Ohsumi Islands including
Sagami Bay, Suruga Bay, western Japan Sea and Shimane Prefecture. Shelf.
Literature: Okutani et al. (1987) (as Sepia (Doratosepion) misakiensis ).
150 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia tuberculata Lamarck, 1798
Sepia tuberculata Lamarck, 1798, Bulletin des Sciences, par la Societe Philomatique , 2(5): 130 [type locality: unknown].
Size: Up to 82 mm mantle length.
Geographical Distribution: Africa: South Africa, Melbosstrand, 33?43' S 18?26'E to Knysna, 34? 03'S 23?03'E. Depth
to 3 m.
Literature: Adam and Rees (1966), Roeleveld (1972).
Sepia typica (Steenstrup, 1875 )
Hemisepius typicus Steenstrup, 1875, Danske Videnskabernes Selskabs Skrifter, 5 Raekke, Naturvidenskabelig og
Mathematisk , 10(7): 468 [type locality: Africa: Sout h Africa, Table Bay, 33?50'S 18?27'E].
Size: Up to 26 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean and southeastern Atlantic: South Africa, from Saldanha Bay,
33?00'S 17?56'E, to southern Mozam bique. Depth range from 2 to 290 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Augustyn et al. (1995), Filippova et al. (1995).
Sepia vercoi Adam, 1979
Sepia vercoi Adam, 1979, Records of the Western Australian Museum , 7(3): 190 [type locality: Western Australia, west of
Shark Bay, 25?31'S 112?29'E].
Size: Up to 46 mm mantle length (both sexes).
Geographical Distribution: Southeastern Indian Ocean: Western Australia, 18?31'S 118?09'E to 25? 31'S 112?29'E. Depth
range from 76 to 201 m.
Literature: Reid (2000).
Sepiella cyanea Robson, 1924
Sepiella cyanea Robson, 1924b, Report of the Fisheries and Marine Biological Survey of the Union of South Africa , 3: 13
[type locality: South Africa, Natal].
Size: Up to 80 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: Port Elizabeth and Durban, north to central Mozambique, 26?N
and Madagascar. Depth range from 13 to 73 m.
Literature: Adam and Rees (1966), Roeleveld (1972), Filippova et al. (1995).
Cephalopods of the World 151
Sepiella mangkangunga Reid and Lu, 1998
Sepiella mangkangunga Reid and Lu, 1998, The Beagle, Records of the Museums and Art Galleries of the Northern
Territory, 14: 74 [type locality: Australia: Northern T erritory, Stingray Head, 12?48'S 130?21'E].
Size: Males up to 58 mm mantle length; females up to 59 mm mantle length.
Geographical Distribution: Indo-Pacific: Australia, Northern Territory, 12?48'S 130?21'E to 13?14' S 130?57'E. Depth from
1.1 to 3.3 m.
Literature: Reid and Lu (1998).
Sepiella ocellata Pfeffer, 1884
Sepiella ocellata Pfeffer, 1884, Abhandlungen aus dem Gebiete der Naturwissenschaften, Hamburg , 8(1): 13 [type
locality: Java].
Size: Up to 50 mm mantle length.
Geographical Distribution: Indo-Pacific: Known only from the type locality. Depth range unknown. Known from a single
male specimen only. The status of this species is questionable.
Literature: Reid and Lu (1998).
Note:
As this catalogue was being printed, the description of a new species of Sepia became available (Reid and Lu, 2005). It is
hereby reported (in synthesis) for sake of completeness.
Sepia filibrachia Reid and Lu, 200 5
Sepia filibrachia Reid and Lu, 2005, Zootaxa , 911: 13 [type locality: Taiwan: Kaohsiung, Ling Yuan, 22?28?N 120?24?E].
Size: Males up to 62 mm, females up to 70 mm mantle length.
Geographical Distribution: Taiwan Province of China, Mu-Do-Yu, 23?44?N 119?35?E, Peng-Hu and Wu-chi, Taichung
24?17?N 120?30?E. South China Sea, Hainan Island, Haikou. 20?40?N 107?56?E; 18?40?N 106?47?E; 11?25?N 109?15?E. Viet
Nam, Gulf of Tonkin, Guryanova. Depth range 34?95 m.
Interest to Fisheries: The species is marketed in Taiwan Province of China.
Literature: Reid and Lu (2005).
152 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
3.2 F a m i l y S E P I O L I D A E Leach, 1817 by Amanda Reid and Patrizia Jereb
Sepiolidae Leach, 1817, Synopsis of the Orders, Families and Genera of the Class Cephalopoda , In: Leach and Nodder,
The Zoological Miscellany, 3(30):137?141.
FAO Names: En ? Bobtail squids; Fr ? S?pioles; Sp ? Globitos, Sepietas, Sepiolas, Sepiolinas.
Diagnostic Features: Mantle broad, oval; posterior mantle margin rounded. Dorsal mantle fused to head, or free from
head. Mantle cavity divided by thin septum. Fins wide; rounded, semicircular, or kidney-shaped, with pronounced anterior
lobes, or ?earlets?; attached about midway along mantle; fin at tachment short, fin length exceeds attachment length. Head
slightly broader than, or as wide as, mantle. Large eyes covered by corneal membranes. Mantle-locking cartilage simple,
linear. Arms short; protective membranes absent in both sexes. Webs between arms III and IV envelop tentacle bases on
outside only, without forming sac. Hectocotylus present: left dorsal arm, both dorsal arms, or dorsolateral arms modified.
Tentacles retractable, each bears a well defined club. Only the left oviduct is developed. Eggs large, development direct.
Internal gladius present, rudimentary, chitinous, or absent.
Size: Small, up to 80 mm, rarely 100 mm, mantle length.
Habitat and Biology: Benthic or neritic. Represented in tropical, temperate and subpolar waters of all oceans.
Interest to Fisheries: Separate statistics are not reported for this group, but many species are utilized locally as food even
though not commercially exploited. The flesh is generally very tasty though difficult to preserve. Currently of low
commercial value; in some areas of their distribution catches are very abundant and the resource is valued as a delicacy
(i.e. some areas of the Mediterranean Sea).
Literature: Fioroni (1981), Boletzky (1995).
Key to subfamilies and genera in the family Sepiolidae
1a. All arms except the fourth pair united by a broad web (Fig. 212); anterior edge of ventral mantle
extended into extensive ventral shield anteriorly, covering funnel and sometimes reaching eye level
(Fig. 213) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ( s u b f a m i l y H e t e r o t e u t h i n a e ) 
 3
1b. Arms not united by a broad web, or only third and fourth arms united by a broad web; anterior edge of
ventral mantle not forming a ventral shield . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 2
Cephalopods of the World 153
Fig. 212 Iridoteuthis iris
ventral view
Fig. 213 Heteroteuthis serventyi
lateral view of male
ventral
shield
154 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
2a. Dorsal mantle fused to head by cutaneous occipital band (Fig. 214); nuchal cartilage absent; left
dorsal arm hectocotylized; internal gladius rudimentary, or absent . . . . . . . . . . ( s u b f a m i l y S e p i o l i n a e ) 
 7
2b. Dorsal mantle not fused to head (Fig. 215); nuchal cartilage present; left or both dorsal arms
hectocotylized; gladius present . . . . . . . . . . . . . . . . . . . . . . . . . . . . ( s u b f a m i l y R o s s i i n a e ) 
 1 1
3a. Dorsal mantle fused to head (Fig. 216) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 4
3b. Dorsal mantle not fused to head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 
 6
4a. Body slightly flattened dorsoventrally; fins from 60 to 80% mantle length (Fig. 216) . . . . . . . . . . . Sepiolina
4b. Body broad, not flattened dorsoventrally; fin length usually >80% mantle length (Fig. 217) . . . . . . . . . . 
 5
Fig. 214 Euprymna tasmanica
dorsal view
hectocotylus
occipital
band
Fig. 215 Austrorossia australis
dorsal view of female
mantle not
fused to head
(illustration: K. Hollis/ABRS)
Fig. 216 Sepiolina nipponensis
dorsal view
Fig. 217 Stolotheutis leucoptera
dorsal view
5a. Occipital band (joining mantle to head) from 40 to 50% head width (Fig. 217); ventral shield
extending only to level of eyes (Fig. 218); fins attach to anterior part of mantle; posterior-fin margins
do not reach end of body; eyes small (Fig. 218) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stoloteuthis
5b. Occipital band almost equal to head width (Fig. 219); ventral shield exceeds level of anterior margin
of eye, reaching proximal end of arms (Fig. 220); fins attached to mantle at level of midline; posterior
fin margins reach, or extend posteriorly beyond mantle margin; eyes very large (Fig. 220) . . . . . . Iridoteuthis
6a. Fins attached close to mantle midline; anterior fin margins extend beyond anterior edge of mantle
(Fig. 221a); distal arm suckers with elongate, thick pedicels (Fig. 221b) . . . . . . . . . . . . . . . Nectoteuthis
6b. Fins attached to posterior half of mantle, not reaching anterior edge (Fig. 222); sucker pedicels
normal, not thick, elongate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heteroteuthis
Cephalopods of the World 155
Fig. 218 Stolotheutis leucoptera
ventral view
Fig. 219 Iridoteuthis sp.
dorsal view
Fig. 220 Iridoteuthis sp .
lateral view
Fig. 221 Nectoteuthis pourtalesi
a ) dorsal view b ) d i s t a l t i p
o f a r m
Fig. 222 Heteroteuthis serventyi
dorsal view
156 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
7a. Arm suckers usually tetraserial (biserial only in E. phenax ); distal suckers on male hectocotylized
arm greatly modified, with closely packed fleshy papillae formed from enlarged and elongate
swollen sucker pedicels (Fig. 223a); male third arms not bent inward (Fig. 223b) . . . . . . . . . . . Euprymna
7b. Proximal arm suckers biserial; hectocotylus modified as copulatory apparatus with recesses and
projections, not as above; male third arms bent toward mouth (Fig. 224) . . . . . . . . . . . . . . . . . . . 
 8
8a. Occiptal band wide, from 33 to 50% head width; light organ on ink sac . . . . . . . . . . . . . . . . . . . . 
 9
8b. Occipital band narrow, usually <33% head width; inc sac without light organs . . . . . . . . . . . . . . . . . 
 1 0
9a. Paired, kidney-shaped light organs on anterior surface of ink sac (Fig. 225a); tentacular club
suckers usually in 4 to 8 transverse rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepiola
9b. Unpaired, round light organ (Fig. 225b) deeply embedded in tissue on ventral side of ink sac;
tentacular club suckers in 16 transverse rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rondeletiola
Fig. 223 Euprymna tasmanica
b ) dorsal view
a ) h e c t o c o t y l u s
Fig. 224 Sepiola birostrata
dorsal view
(after Okutani, et al., 1987 )
Fig. 225 Mantle cavity
a ) Sepiola
paired
kidney-shaped
light organs
b ) Rondeletiola
unpaired round
light organ
(after Norman and Lu, 1997 )
10a. Both dorsal arms of males fused proximally; hectocotylized arm broad, spoon-like distal to
copulatory apparatus (Fig. 226); tentacular-club suckers in >10 transverse rows . . . . . . . . . . . . . Sepietta
10b. Hectocotylized arm not broad distal to copulatory apparatus (may be slightly broad in region of
copulatory apparatus); tentacular club suckers in 8 to 10 rows . . . . . . . . . . . . . . . . . . . . . Inioteuthis
11a. Light organ present on ink sac; left dorsal arm of male hectocotylized (Fig. 227) . . . . . . . . . . . . Semirossia
11b. No light organ on ink sac; both dorsal arms of males hectocotylized . . . . . . . . . . . . . . . . . . . . . 
 1 2
12a. Anal flaps present, well developed (Fig. 228a); ink sac well developed . . . . . . . . . . . . . . . . . . . . 
 1 3
12b. Anal flaps reduced, or absent (Fig. 228b); ink sac greatly reduced . . . . . . . . . . . . . . . . . . . Neorossia
13a. Male with both dorsal arms hectocotylized by an expanded ventral membrane or thickened fold
(R. moelleri ); arm suckers of males not greatly enlarged; tentacular clubs expanded with 5 to 16
transverse rows of medium-sized suckers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rossia
13b. Males with both dorsal arms hectocotylized by an expanded swollen membrane or fold on the
proximal ventral margin; some suckers of all male arms enlarged; tentacular clubs not expanded,
with 24 to 40 transverse rows of minute suckers . . . . . . . . . . . . . . . . . . . . . . . . . . . Austrorossia
Cephalopods of the World 157
Fig. 226 Sepietta oweniana
h e c t o c o t y l u s
Fig. 227 Semirossia tenera
o r a l v i e w o f m a l e
Fig. 228 anus (inside mantle cavity)
a ) Rossia
anal flaps
well
developed
b ) Neorossia
anal flaps
greatly
reduced
3.2.1 S u b f a m i l y S E P I O L I N A E Appell?f, 1898
Sepiola affinis Naef, 1912 F i g . 2 2 9
Sepiola affinis Naef, 1912b, Zoologischer Anzeiger , 40(2): 81 [type locality: Tyrrhenian Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Analogous bobtail squid; Fr ? S?piole analogue; Sp ? Sepiola an?loga.
Diagnostic Features: Mantle slightly conical, head robust with prominent eyes; fins wide; rounded, semicircular; do not
exceed length of mantle anteriorly or posteriorly; arm suckers biserial; tentacles delicate, club with 6 transverse rows of
very small suckers; a few of the dorsal sucker rows close to the stalk slightly enlarged. Hectocotylus present, left dorsal arm
modified: 3 small suckers at the base, followed distally by the swollen lobe of the copulatory apparatus consisting of a
primary lobe followed by a smaller lobe and a wrinkled tubercle; suckers biserial distal to copulatory apparatus, the dorsal
ones differing in size: first 4 to 6 small suckers
are followed by 3 or 4 very large ones ,
remaining suckers small, decreasing regularly
in size to distal tip of arm; ventral rows of
suckers uniformly small. Paired, kidney-shaped
light organs present inside mantle cavity on
each side of ink sac. Bursa copulatrix small .
Colour: Dark brown, chromatophores large,
violet brownish, dense.
Size: Up to 25 mm mantle length (both sexes).
Geographical Distribution: Mediterranean
Sea, including Strait of Sicily, Adriatic Sea and
north Aegean Sea. Depth range from 20 to
178 m (Fig. 230).
158 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 229 Sepiola affinis
m a n t l e c a v i t y o f f e m a l e
h e c t o c o t y l u s
dorsal view
bursa copulatrix
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
40?
40?
30? 30?
40? 40?
Fig. 230 Sepiola affinis
Known distribution
?? 200 m
Habitat and Biology: A Mediterranean endemic, Sepiola affinis has been found at depths slightly over 150 m, but it is
typically abundant in shallow waters (15?30 m), mostly on sandy or sandy-muddy substrates. It is often associated with
S. intermedia Naef, 1912, from which young females are very difficult to distinguish. The smallest mature males are
12 mm, and the smallest mature females are 18 mm mantle length. Eggs are relatively large (2.2 ? 1.8 mm) and
experimental observations indicate that spawning may occur in one single event as well as being continuous or
intermittent. The maximum observed spawning duration is 2 months and all animals die after spawning. During prey
capture the species exhibits a great variety of colour patterns, with rapid colour changes.
Interest to Fisheries: It is commonly caught by small-scale and artisanal fisheries and consumed locally together with
other species of the group. Separate statistics are not available.
Local Names: ITALY: Seppiola affine, Cappuccetto.
Literature: Naef (1923), Mauris (1989), Bello (1990b), Guerra (1992), Bello (1995), Gabel-Deickert (1995), D?Onghia et al.
(1996), Jereb et al. (1997), Salman et al. (2002).
Sepiola atlantica Orbigny, 1839 ?1842 F i g . 2 3 1
Sepiola atlantica Orbigny, 1839?1842 (in F?russac and d?Orbigny, 1834?1848), Histoire Naturelle G?n?rale et
Particuli?re C?phalopodes Ac?tabuliferes Vivants et Fossiles , pl. 4 [type locality: France: Bay of Biscay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Atlantic bobtail squid; Fr ? S?piole grandes
oreilles; Sp ? Sepiola atl?ntica.
Diagnostic Features: Fins short, do not exceed length of mantle anteriorly or posteriorly. Arms IV with biserial suckers
proximally, 4 to 8 rows of minute suckers at distal tips ; remaining arms with biserial suckers. Hectocotylus present; left
dorsal arm modified: proximal end with fleshy pad formed from enlarged and fused sucker pedicels; copulatory apparatus a
large swollen horn, with secondary lobes basally; dorsal row of suckers distal to copulatory apparatus with 3 or 4 slightly
enlarged suckers with swollen pedicels, 3 or 4 greatly reduced suckers, then 3 to 5 greatly enlarged suckers
approximately halfway along arm. Hectocotylized arm strongly bent in distal half. Club with 8 suckers in transverse rows.
Paired, kidney-shaped light organs (photophores) present inside mantle cavity on each side of ink sac.
Cephalopods of the World 159
dorsal view
Fig. 231 Sepiola atlantica
h e c t o c o t y l u s m a n t l e c a v i t y o f f e m a l e
bursa copulatrix
Size: Up to 21 mm mantle length (mature males and
females similar in size).
Geographical Distribution: Northeastern Atlantic:
65?N to 35?N, from Iceland, Faeroe Islands and
western Norway to Morocco. Mediterranean Sea
(single record only) (Fig. 232).
Habitat and Biology: Continental shelf to edge of
slope. Epibenthic, but has been collected in midwater
during both night and day. The ecology of the species
in Firemore Bay, Loch Ewe, Scotland has been well
studied. The resident population has a variable age
composi t ion, which suggests an extended
reproductive season. Juveniles occur throughout the
year, with peak recruitment in April and July to August.
Interest to Fisheries: Specific records are not
available, but the species is likely to occur in local
fisheries, due to its wide distribution and fairly common
occurrence on the north European continental shelf.
Remarks: According to Naef (1916), Grimpe (1925),
Bello (1986), Mangold and Boletzky (1988) and Bello
(1992) Sepiola atlantica is absent from the
Mediterranean Sea. However, Wurtz et al. (1995)
referred to one mature male captured in the
Tyrrhenian Sea at 90 m depth. Sepiola atlantica is
distinguished from other sepiolid species in the
northeastern Atlantic by the 4 to 8 rows of minute
suckers at the distal tips of arms IV and the presence
of the internal, ventrally positioned, bilobed light
organs.
Literature: Joubin (1902a), Naef (1912b), Herring
et al. (1981), Roper et al. (1984), Guerra (1986), Nesis
(1987), Guerra (1992), Yau and Boyle (1996), Collins
et al. (2001).
160 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 232 Sepiola atlantica
Known distribution
?? 200 m
?
Sepiola birostrata Sasaki, 1918 F i g . 2 3 3
Sepiola birostrata Sasaki, 1918, Doubutsugaku Zasshi [Zoological Magazine Tokyo], 30: 235. [In Japanese] [type
locality: Japan Sea, Toyama Bay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Butterfly bobtail squid; Fr ? S?piole papillon; Sp ? Sepiola mariposa.
Diagnostic Features: Mantle dome-shaped, half as
wide as long. Dorsal mantle fused to head. Fins wide,
ovate, short, do not exceed length of mantle anteriorly
or posteriorly; attached slightly anterior to mantle
midline. Arms III in both sexes stout and strongly curved
inward (more obviously so in males) .
Non-hectocotylized arm sucker arrangement same in
both sexes: arm suckers biserial. Dorsal and ventral
rows of suckers on arms I and II of males equal in size,
larger than suckers on arms III and IV. Hectocotylus
present, left dorsal arm modified: proximal end with 2
slender, fleshy, papillae formed from enlarged and
elongate sucker pedicels ; anteriormost papilla longest
and thickest, posterior papilla fleshy; both papillae blunt
distally. Club slightly recurved; short; with 4 suckers in
proximal transverse rows, up to 16 rows distally;
suckers small, dorsal marginal longitudinal series of
suckers slightly larger than those in ventral marginal
series. Paired, kidney-shaped light organs present
inside mantle cavity on each side of ink sac. Colour:
Mantle and head with many minute brown or black
chromatophores; arms III deep pink, arms I to III each
with single longitudinal row of large chromatophores,
arms IV with double row of small chromatophores.
Size: Up to 22 mm mantle length.
Geographical Distribution: Northwestern Pacific: Sea
of Okhotsk, southern Sakhalin, south Kurile Islands,
Primorye (Russian Federation), North and South
Korea, Japan, East China Sea (Fig. 234).
Cephalopods of the World 161
Fig. 233 Sepiola birostrata
(after Okutani et al., 1995 )
dorsal viewa r m s o f m a l e
tentacle
I V
I I I
I I
I (hc.)
Fig. 234 Sepiola birostrata
Known distribution
?? 200 m
Habitat and Biology: Neritic. Depth range to approximately 100 m.
Interest to Fisheries: No specific information is available, but the species is likely to occur, occasionally, in multispecies
fisheries.
Remarks: Sepiola birostrata usually differs from S. trirostrata Voss, 1962, by the absence of a third blunt, fleshy lobe at the
base of the pair of elongate papillae on hectocotylized arm. However, there appears to be some intraspecific variability in
S. birostrata: 3 lobes have been reported to occur in some specimens. All papillae on the hectocotylized arm of
S. birostrata arise from a common mound-like base. In contrast, the third lobe-like papilla of S. trirostrata does not arise
from the same base as the other two papillae. See Remarks under Sepiola parva Sasaki, 1913 for differences between
S. birostrata and S. parva.
Literature: Roper et al. (1984), Nesis (1987), Takayama and Okutani (1992), Okutani (1995), Kubodera and Yamada
(1998).
Sepiola intermedia Naef, 1912 F i g . 2 3 5
Sepiola intermedia Naef, 1912a, Zoologischer Anzeiger , 39(7): 270 [type locality: Tyrrhenian Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Intermediate bobtail squid; Fr ? S?piole interm?diaire; Sp ? Sepiola intermedia.
Diagnostic Features: Fins short, rounded, do not exceed length of mantle anteriorly or posteriorly. Arms II and IV inmales
with a few enlarged suckers . Tentacles very delicate, tentacle clubs with 6 transverse rows of small suckers. Hectocotylus
present; left dorsal arm modified: 3 normal suckers at the base (only slightly enlarged); copulatory apparatus consisting of
a swollen and wrinkled tubercle, curved inward, with the markedly enlarged lobe projecting free; dorsal row of suckers
distal to copulatory apparatus very variable in size: 2 markedly enlarged (of which the proximal is slightly larger); or 3
enlarged, of which the median is the largest; or 1 very small followed by 1 very large then 1moderately large sucker distal
to the basal apparatus; left dorsal arm much longer than the right one. Colour: Dark coffee brown, with a strong shade of
reddish brown; large chromatophores loosely scattered all over the mantle surface.
Size: Males up to 26 mm mantle length; females up to 28 mm mantle length.
162 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 235 Sepiola intermedia
m a n t l e c a v i t y o f f e m a l eh e c t o c o t y l u s
(after Naef, 1923 )
bursa copulatrix
Geographical Distribution: Northeastern
Atlantic and Mediterranean Sea: including
Ligurian Sea, Strait of Sicily, Adriatic Sea and
northern Aegean Sea. Possibly Gulf of C?diz.
Continental shelf, depth range from 8 to
110 m (Fig. 236).
Habitat and Biology: Sepiola intermedia
lives mainly on muddy bottoms between 60
and 200 m depth throughout its distributional
range. In Italian waters, a preference for
shallower, coastal and sandy bottoms has
been observed both in the Adriatic and the
Tyrrhenian Seas, where the species can be
very abundant and is often found associated
with Turritella communis , Astropecten
bispinosus and Ophiura texturata . The
biology of the species is poorly known, but the
presence of mature individuals throughout the year suggests there may be a prolonged spawning period. Adult Sepiola
intermedia have been collected in midwater, at night, by purse seine for blue fishes, confirming the strong swimming ability
of this small bobtail squid (as observed in other species of the group).
Interest to Fisheries: No separate statistics are available, but the species is collected by local small-scale fisheries
throughout its distributional range and represents a component of the bobtail squids captured in many Mediterranean
areas.
Local Names: ITALY: Seppiola intermedia, Cappuccetto.
Literature: Naef (1912b), Naef (1923), Belcari et al. (1989), Orsi Relini and Bertuletti (1989), Guerra (1992), Bello (1995),
Volpi et al. (1995), D?Onghia et al. (1996), Jereb et al. (1997), Salman et al. (1997).
Sepiola ligulata Naef, 1912 F i g . 2 3 7
Sepiola ligulata Naef, 1912a, Zoologischer Anzeiger , 39(7): 271 [type locality: Tyrrhenian Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Tongue bobtail squid; Fr ? S?piole languette; Sp ? Sepiola leng?ita.
Cephalopods of the World 163
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
40?
40?
30? 30?
40? 40?
Fig. 236 Sepiola intermedia
Known distribution
?? 200 m
?
Fig. 237 Sepiola ligulata
m a n t l e c a v i t y o f f e m a l eh e c t o c o t y l u s
(after Naef, 1923 )
bursa copulatrix
Diagnostic Features: Mantle ventral margin may be markedly produced, with a deep median indentation. Fins relatively
small, with a round lateral corner and relatively straight posterior margin. Tentacles delicate; club with 8 dense transverse
rows of minute suckers. Hectocotylus present, left dorsal arm modified; 3 small normal suckers at the base followed
distally by the copulatory apparatus: spatula-like thick and large lobe extending towards the distal part of the arm and an
adjacent appendage forming a stalked plate or shovel- like structure that extends toward the right arm ; distal to
copulatory apparatus, first suckers of the ventral row enlarged, with long, robust stalks. Bursa copulatrix large, extending
anteriorly to cover renal papillae and genital opening and extends through the mantle septum over a large part of the right
side of the mantle cavity. Colour: Bright, almost golden yellow, with yellowish orange, brownish red tones.
Size: Up to 25 mm mantle length.
Geographical Distribution: Mediterranean
Sea: including Ligurian Sea, Tyrrhenian Sea,
Strait of Sicily, Adriatic Sea and northern
Aegean Sea. Possibly eastern Atlantic around
Portugal and Spain. Depth range from 44 to
380 m (Fig. 238).
Habitat and Biology: The male copulatory
apparatus and the female bursa copulatrix are
very distinctive in S. ligulata, enabling it to be
readily distinguished from other species in the
genus. It is one of the smallest species of the
genus and is reported to be common throughout
its range, though never abundant. Sepiola
ligulata is an epibenthic species, living on
muddy bottoms and mainly on the continental
shelf, although it has also been found on the
slope. It is often captured with Sepietta oweniana (Orbigny, 1839?1841) and Rondeletiola minor Naef, 1912. Information
on the biology of S. ligulata is scarce, and is mostly derived from aquarium observations. Eggs are red and relatively large
(3.5 mm diameter) and growth is rapid with water temperatures between 12? and 20?C, allowing newly hatched individuals
(1.8 to 2.5 mm mantle length) to reach 9 mm mantle length in about six months. The smallest mature males are 11 mm, and
the smallest females 14 mm mantle length.
Interest to Fisheries: Never abundant, but often present in captures with the other species of the group, it is used as food
locally.
Local Names: ITALY: Seppiola linguetta.
Literature: Naef (1912b), Naef (1923), Boletzky et al. (1971), Orsi Relini and Bertuletti (1989), Bello (1990b), Guerra
(1992), Bello (1995), Sartor and Belcari (1995), Volpi et al. (1995), Wurtz et al. (1995), D?Onghia et al. (1996), Jereb et al.
(1997), Salman et al. (2002).
164 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
40?
40?
30? 30?
40? 40?
Fig. 238 Sepiola ligulata
Known distribution
?? 200 m
?
?
Sepiola parva Sasaki, 1913 F i g . 2 3 9
Sepiola parva Sasaki, 1913, Dobutsugaku Zasshi [Zoological Magazine Tokyo], 25: 252. [In Japanese] [type locality:
Japan, Tokyo Bay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Spotty bobtail squid; Fr ? S?piole mouchet?e; Sp ? Sepiola manchada.
Diagnostic Features: Mantle dome-shaped. Dorsal
mantle fused to head. Fins narrow, ear-shaped, short; do
not exceed length of mantle anteriorly or posteriorly. Head
broad, narrower than mantle. Non-hectocotylized arm
sucker arrangement same in both sexes: arm suckers
biserial. Hectocotylus present, left dorsal arm modified:
proximal end with fleshy hook-like papilla formed from
enlarged sucker pedicel; 2 rows of suckers on arm
proximal to hook-like papilla; distal end of hectocotylized
arm with sucker pedicels enlarged and tightly packed
forming two double rows of columnar structures; suckers
reduced with tiny, fleshy, slit-like openings. Club with 8
suckers in transverse rows; dorsal and ventral marginal
longitudinal series of suckers smaller than medial suckers
(medial suckers 1.6 times larger than marginal ones).
Internal gladius absent. Paired, kidney-shaped light
organs present inside mantle cavity on each side of ink
sac.
Size: Up to 10 mm mantle length.
Geographical Distribution: Northwestern Pacific:
southern Japan to northern Philippines (Fig. 240).
Habitat and Biology: Upper sublittoral. This species
seems to prefer the subtidal zone with hard substrates,
unlike S. birostrata Sasaki, 1918, which is abundant in
shallow water with sandy or muddy substrates.
Cephalopods of the World 165
Fig. 239 Sepiola parva
h e c t o c o t y l u s ventral view of male dorsal view of male
(after Okutani, 1995 )
Fig. 240 Sepiola parva
Known distribution
?? 200 m
Interest to Fisheries: The species is occasionally caught and consumed locally.
Remarks: This species closely resembles S. birostrata. The 2 species, however, can be distinguished by differences in the
dentition of the chitinous rings on the tentacle-club suckers, regardless of maturity and sex. In Sepiola parva, the
club-sucker rings have three rows of polygonal processes bearing bush-like papillae. Sepiola birostrata Sasaki, 1913 has
4 rows of polygonal processes; the inner 2 rows bear blunt, flat-topped pegs. The 2 or 3 hook-like processes on the
hectocotylized arm arise from a common elevated base in S. birostrata, unlike that of S. parva, in which a single process
arises directly from the arm flesh.
Literature: Takayama and Okutani (1992), Okutani (1995), Reid and Norman (1998).
Sepiola robusta Naef, 1912 F i g . 2 4 1
Sepiola robusta Naef, 1912a, Zoologischer Anzeiger , 39(7): 271 [type locality: Tyrrhenian Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Robust bobtail squid; Fr ? S?piole
robuste; Sp ? Sepiola robusta.
Diagnostic Features: Ventral mantle normal, not strongly produced anteriorly. Fins short, do not exceed length of mantle
anteriorly or posteriorly. Hectocotylus present, left dorsal arm modified: proximal end with fleshy pad formed from enlarged
and fused sucker pedicels; copulatory apparatus a complexly twisted lobe (very var iable) consisting of 3 hook-like
structures; horn of copulatory apparatus slightly recurved, but does not form a small hole; base of hectocotylus with one
sucker of ventral row markedly enlarged; distal end of hectocotylized arm with normal suckers (not enlarged or modified);
arm slightly widened distal to copulatory apparatus and sharply bent distally; arm without groove on inner side. Arm often
narrower at level of fifth and sixth sucker rows distal to copulatory apparatus. Club large with 8 suckers in transverse
rows; suckers in dorsal rows in widest part of club marked ly enlarged. Female bursa copulatrix moderately large
(exceeds length of gill insertion posteriorly) ; caecum absent; posterior left side of mantle cavity in females without
mantle constrictor muscle (cutaneous muscle extending from mantle septum to base of left gill). Paired, kidney-shaped
light organs (photophores) present inside mantle cavity on each side of ink sac.
Size: Males up to 25 mm mantle length; females up to 28 mm mantle length.
166 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 241 Sepiola robusta
ventral view of male
m a n t l e c a v i t y o f f e m a l e
d o r s a l a r m s o f m a l e s
( h e c t o c o t y l u s )
bursa copulatrix
Geographical Distribution: Mediterranean Sea: absent
in Marmara and Black Sea (Fig. 242).
Habitat and Biology: Outer shelf. Depth range from 26
to 498 m. Mating takes place when the male grasps the
female?s ?neck? region, and spermatophores are placed
in the females? bursa copulatrix. Males have been
observed to guard females during courtship.
Interest to Fisheries: Sepiola robusta is one of the most
frequently captured Mediterranean Sepiola, where it
contributes (although in low numbers) to the bobtail
squid catches by local trawlers.
Local Names: ITALY: Seppiola robusta, Cappuccetto.
Remarks: The doubtful record from Galizia (Spain,
northeastern Atlantic) noted by Guerra (1984) has not
subsequently been confirmed; the species is
considered to be a Mediterranean endemic (Guerra,
1992).
Literature: Naef (1923), Boletzky et al. (1971), Boletzky
(1983b), Nesis (1987), Guerra (1984), Belcari et al.
(1989), Orsi Relini and Bertuletti (1989), Guerra (1992),
Bello (1995), Jereb and Di Stefano (1995), Volpi et al.
(1995), Jereb et al. (1997).
Sepiola rondeleti Leach, 1834 F i g . 2 4 3
Sepiola rondeleti Leach, 1834 (in F?russac and d?Orbigny,
1834?1848), H istoire Naturelle G ?n ?rale et Particuli ?re
C?phalopodes Ac?tabuliferes Vivants et Fossiles , 230 [type
locality: Mediterranean Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Dwarf bobtail squid; Fr ? S?piole naine;
Sp ? Sepiola enana.
Cephalopods of the World 167
Fig. 242 Sepiola robusta
Known distribution
?? 200 m
Fig. 243 Sepiola rondeleti
m a n t l e c a v i t y o f f e m a l e
(after Naef, 1912)
d o r s a l a r m s o f m a l e
( h e c t o c o t y l u s ) dorsal view of female
bursa copulatrix
Diagnostic Features: Ventral mantle margin projects markedly anteriorly . Fins short, do not exceed length of mantle
anteriorly or posteriorly. Hectocotylus present, left dorsal arm modified: proximal end with fleshy pad formed from enlarged
and fused sucker pedicels; horn of copulatory apparatus recurved to form a small hole ; base of hectocotylus with equal
sized suckers; distal end of hectocotylized arm with some enlarged suckers; ventral row of suckers distal to copulatory
apparatus much smaller than those in dorsal row; arm without groove on inner side. Club with 8 suckers in transverse
rows; dorsal marginal series of sucker s enlarged on proximal end of club . Female bursa copulatrix large; extends
posteriorly beyond gill insertion; caecum present, small; barely protrudes into right half of mantle cavity. Posterior left side
of mantle cavity in females without mantle constrictor muscle (cutaneous muscle extends from mantle septum to base of
left gill). Internal gladius present, rudimentary, chitinous. Paired, kidney-shaped light organs present inside mantle cavity,
on each side of ink sac. Anterior ventral mantle margin with dense border of chromatophores. Outer surface of tentacular
clubs heavily pigmented with chromatophores.
Size: Males up to 25 mm mantle length; females up to 60 mm mantle length (but most commonly from 40 to 50 mm mantle
length).
Geographical Distribution: Northeastern Atlantic and
Mediterranean Sea: including Strait of Sicily, Aegean
Sea, Adriatic Sea, Sea of Marmara and Levantine
Sea. In the northeastern Atlantic it extends from the
North Sea to Senegal (Fig. 244).
Habitat and Biology: Sandy and muddy substrates,
common in Posidonia seagrass beds down to 35 m.
Epibenthic, or mesobenthic, Sepiola rondeleti has
been recorded from very shallow waters down to
around 450 m. Mating takes place when males grasp
the female?s ?neck? region, and spermatophores are
placed in her bursa copulatrix. In the western
Mediterranean, the spawning season extends from
March to November, with longevity estimated at 18
months. The species feeds on crustacea and small
fishes. All females over 30 mm mantle length are
mature.
Interest to Fisheries: Captured during bottom trawls
and occasionally with purse seines and drift nets. It is
consumed locally; the flesh is very tasty, though
difficult to preserve.
Local Names: ITALY: Seppiola di Rondelet,
Cappuccetto.
Remarks: Sepiola rondeleti is known to preferentially
inhabit waters shallower than 50 m throughout the
year. Deeper records from the Mediterranean were
long considered doubtful (Mangold-Wirtz 1963, Bello
1983?84). However, more recent evidence has
confirmed a greater depth distribution for this species
than previously thought, verifying the maximum
recorded depth of 450 m (Dieuzeide, 1955).
Literature: Joubin (1895, 1902a), Naef (1912b), Naef (1923), Boletzky et al. (1971), Bello (1983?84), Roper et al. (1984),
Guerra (1992), Bello (1995), Orsi Relini and Bertuletti (1989), D?Onghia et al. (1996), Jereb et al. (1997).
168 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 244 Sepiola rondeleti
Known distribution
?? 200 m
Sepiola trirostrata Voss, 1962 F i g . 2 4 5
Sepiola trirostrata Voss, 1962a, Proceedings of the Biological Society of Washington , 75: 172 [type locality: Philippines].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Knobby bobtail squid; Fr ? S?piole
bossel?e; Sp ? Sepiola nudosa.
Diagnostic Features: Fins short, do not exceed length
of mantle anteriorly or posteriorly. Arms III in both
sexes stout and strongly curved inward, more
obviously so in males. Suckers in ventral series of right
arm I and arms II of males larger than dorsal suckers.
Hectocotylus present, left dorsal arm modified:
proximal end with 2 slender fleshy papillae
(anteriormost papilla longest) and dorsolateral to
these a blunt tongue-like lobe , all formed from
enlarged and elongate sucker pedicels; 2 rows of
suckers on arm proximal to fleshy pad; distal end of
hectocotylized arm with sucker pedicels enlarged and
tightly packed to form 2 double rows of columnar
structures; suckers reduced with tiny, fleshy, slit-like
openings. Club with 4 large suckers in transverse
rows; suckers differ in size; dorsal marginal
longitudinal series of suckers larger than those in
ventral marginal series. Paired kidney-shaped light
organs present inside mantle cavity on each side of ink
sac. Colour: Mantle and head with many minute
brown or black chromatophores; arms III deep pink,
arms I to III each with single longitudinal row of large
chromatophores, arms IV with double row of small
chromatophores.
Size: Up to 12.5 mm mantle length.
Geographical Distribution: Indo-Pacific: Philippines,
Singapore (Fig. 246).
Cephalopods of the World 169
Fig. 245 Sepiola trirostrata
(after Voss, 1963 )
dorsal view of male
m a l e a r m a r r a n g e m e n t
tentacle
I I I left
I left
I I left
I right
I V left
hectocotylus
Fig. 246 Sepiola trirostrata
Known distribution
?? 200 m
170 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Interest to Fisheries: Presently undetermined.
Remarks: Differs from S. birostrata Sasaki, 1918 in the presence of the blunt, fleshy papilla, in addition to the 2 slender
papillae on the hectocotylus. The slender papillae arise from a common mound-like base, while the third lobe arises
separately. In this trait, S. trirostrata differs from those S. birostrata specimens that bear a third lobe on the hectocotylus.
Literature: Voss (1963).
Euprymna berryi Sasaki, 1929 F i g . 2 4 7
Euprymna berryi Sasaki, 1929, Journal of the College of Agriculture, Hokkaido Imperial University , 20(supplement): 143
[type locality: Japan, Honshu].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Humming-bird bobtail squid;
Fr ? S?piole colibri; Sp ? Globito colibri.
Diagnostic Features: Fins short, do not exceed length of mantle anteriorly or posteriorly. Non-hectocotylized arm sucker
arrangement same in both sexes: arm suckers tetraserial. Male non-hectocotylized arm suckers enlarged on some arms:
arms II and IV with approximately 10 moderately enlarged suckers in dorsal and ventral rows from proximal second to
fourth suckers; approximately 8 slightly enlarged suckers i n ventral row of arm III from proximal fifth to eighth sucker .
Hectocotylus present; left dorsal arm modified: arm thick, short, and blunt; proximal end with 2 fleshy papillae formed from
enlarged, or elongate sucker pedicels; distal end of hectocotylized arm with sucker pedicels enlarged and tightly packed
forming 2 double rows of 70 to 80 columnar structures; suckers reduced with tiny, fleshy, slit-like openings. Marginal
suckers on dorsal arms I are slightly larger than medial ones. Tentacular club suckers goblet-shaped. Swimming keel of
club extends well beyond carpus.
Size: Males up to 30 mm mantle length; females up to 50 mm mantle length.
(after Okutani et al. , 1995 )
dorsal view of male
Fig. 247 Euprymna berryi
a r m s o f m a l e
(oral view)
I (hc.)
I I I
I I
I V I V
I I I I
I I
Geographical Distribution: Indo-Pacific: Along coasts
of China, south to Hong Kong and Japan (but not in
Hokkaido), Taiwan Province of China, possibly
Andaman Islands and Sri Lanka (Fig. 248).
Habitat and Biology: Upper sublittoral and pelagic in
warm temperate waters; depth range to 107 m. This
species has been reared successfully in aquaculture
experiments. Females lay clumps of round, pale
orange eggs.
Interest to Fisheries: Consumed locally in China and
Taiwan Province of China, where this species is
abundant and sustains local small fisheries; currently
not commercially exploited in Hong Kong.
Local Names: CHINA: Leung yee jai; JAPAN:
Niyori-mimi-ika.
Remarks: The taxonomic status of members of this
genus is largely unresolved (Norman and Lu 1997).
Euprymna berryi is sympatric with, and very similar to
E. morsei Verrill (1881). Non-quantitative differences
between the 2 species include: among 4 rows of arm
suckers, only the ventral rows are enlarged in
E. morsei (particularly from third to fourth suckers on
arms II to IV), but both dorsal and ventral arms have enlarged suckers in E. berryi; the club suckers are spherical, or
cup-shaped in E. morsei, but goblet-shaped in E. berryi. The club suckers have an indistinct proximal border in E. berryi,
while those in E. morsei have distinct borders.
Literature: Roper et al. (1984), Okutani and Horita (1987), Okutani (1995), Norman and Lu (1997), Kubodera and Yamada
(1998), Lu (1998b).
Euprymna morsei (Verrill, 1881) F i g . 2 4 9
Inioteuthis morsei Verrill, 1881, Transactions of the Connecticut Academy of Sciences, 5(6): 417 [type locality: Japan,
Tokyo Bay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Mimika bobtail squid; Fr ? S?piole
mimika; Sp ? Globito mimika.
Cephalopods of the World 171
Fig. 248 Euprymna berryi
Known distribution
?? 200 m
? ?
dorsal view of female
Fig. 249 Euprymna morsei
a r m s o f m a l e
(oral view)
I
I I
I V
I
I I I
hectocotylus
Diagnostic Features: Mantle dome-shaped, plump. Dorsal mantle fused to head. Fins wide, rounded, semicircular; short,
do not exceed length of mantle anteriorly or posteriorly; posteriorly with wide gap between fins; anterior origin posterior to
mantle margin. Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers tetraserial. Ventral rows of
arms II to IV in males with enlarged s uckers, approximately 10 on each arm f rom proximal third or fourth suckers .
Hectocotylus present; distal half of left dorsal arm modified: sucker pedicels enlarged and tightly packed to form 2 double
rows of columnar structures; suckers reduced with tiny, fleshy, slit-like openings; proximal end of arm with single nipple-like
papilla. Marginal suckers arms I slightly larger than medial suckers. Tentacular-club suckers cup-shaped or spherical.
Internal gladius absent. Paired saddle-shaped light organs present inside mantle cavity on ink sac. Colour: Iridescent gold
to purple with large black chromatophores.
Size: Up to 40 mm mantle length.
Geographical Distribution: Indo-Pacific: southern
Japan (sympatric with E. berryi Sasaki, 1929), East
China Sea and the Philippines, south at least to
Indonesia. Possibly Bay of Bengal, India (record based
on females only) and Maldive Islands (Fig. 250).
Habitat and Biology: This neritic species has been
reported from the stomach contents of lancet fishes,
Alepisaurus ferox, captured in the Solomon Sea.
Interest to Fisheries: Harvested on a minor scale,
primarily as trawl bycatch. Low economic value, but
utilized locally.
Remarks: The taxonomic status of members of this
genus is largely unresolved (Norman and Lu 1997).
Literature: Joubin (1902a), Raj and Kalyani (1971),
Okutani et al. (1987), Okutani and Tsukada (1988),
Okutani (1995), Norman and Lu (1997), Kubodera and
Yamada (1998), Reid and Norman (1998).
172 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 250 Euprymna morsei
Known distribution
?? 200 m
?
?
?
Euprymna tasmanica (Pfeffer, 1884) F i g . 2 5 1 ; P l a t e V I , 3 9 ? 4 1
Sepiola tasmanica Pfeffer, 1884, Abhandlungen aus dem Gebiete der Naturwissenschaften, Hamburg , 8(1): 6 [type
locality: Australia: Bass Strait].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Southern bobtail squid; Fr ? S?piole du
Tasmanie; Sp ? Globito de Tasmania.
Diagnostic Features: Dorsal mantle fused to head. Fins wide, rounded, semicircular; short, do not exceed length of mantle
anteriorly or posteriorly. Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers tetraserial. Dorsal
and ventral rows of suckers on arms II to IV in males enlar ged; ventral marginal rows of arms II and III with 1 to 3
greatly enlarged suckers basally (8 to 11% mantle length); dorsal and ventral marginal rows of arms II to IV with more
than 10 enlarged suckers (diameter 4 to 7% mantle length). Hectocotylus present, distal half of left dorsal arm modified:
third and/or fourth proximal suckers in ventral row elongated into long papilla(e), each bearing a tiny sucker; base of
hectocotylus with 29 to 38 normal suckers; distal end of hectocotylized arm with sucker pedicels enlarged and tightly
packed to form 2 double rows of columnar structures; suckers reduced with tiny, fleshy, slit-like openings (30 to 38 in dorsal
rows, 29 to 38 in ventral rows). Tentacular club suckers (many hundreds) all of similar minute size. Saddle-shaped bacterial
light organ inside mantle cavity.
Size: Up to 40 mm mantle length.
Geographical Distribution: Southern Indo-Pacific:
eastern and southeastern Australia, from Brisbane to
Shark Bay, Western Australia (Fig. 252).
Habitat and Biology: This species lives in sandy and
muddy areas, often in association with seagrass beds.
It remains buried in sand during the day, and emerges
at night to forage for small crustaceans and fishes. It
can ?glue? sand grains to its dorsal body surface to aid
in camouflage. Spawning occurs in spring and
summer. Females lay pale orange eggs in loose
clumps, usually at the base of seaweed or seagrass.
Interest to Fisheries: Undetermined, but the species
is likely to be utilized on a local basis.
Remarks: The taxonomic status of members of this
genus is largely unresolved.
Literature: Norman and Lu (1997), Reid and Norman (1998).
Cephalopods of the World 173
Fig. 251 Euprymna tasmanica
t e n t a c u l a r c l u bh e c t o c o t y l u s
dorsal view of male
H
110?
110?
120?
120?
130?
130?
140?
140?
150?
150?
160?
160?
 40?  40?
 30?  30?
 20?  20?
 10?  10?
Fig. 252 Euprymna tasmanica
Known distribution
?? 200 m
174 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Rondeletiola minor (Naef, 1912) F i g . 2 5 3
Sepietta minor Naef, 1912a, Zoologischer Anzeiger , 39(7): 267 [type locality: Tyrrhenian Sea].
Frequent Synonyms: Sepietta minor Naef, 1912.
Misidentifications: None.
FAO Names: En ? Lentil bobtail squid; Fr ? S?piole bobie; Sp ? Globito peque?o.
Diagnostic Features: Mantle bullet-shaped, delicate consistency. Fins small, bluntly pointed laterally rather than curved;
short, do not exceed length of mantle anteriorly or posteriorly. Non-hectocotylized arm sucker arrangement same in both
sexes: arm suckers biserial. Hectocotylus present; left dorsal arm modified: proximal end with fleshy pad formed from
enlarged and fused sucker pedicels; copulatory apparatus a large, swollen, transverse, hood-shaped horn with small
accessory papilla; 3 small suckers proximal to fleshy pad; proximal half of ventral row of suckers distal to copulatory
apparatus with enlarged suckers; oral surface of modified region as for remaining arms, not wide, fleshy; suckers evenly
spaced on modified portion of arm, rows not widely separated. Club with 16 suckers in transverse rows. Paired, roundish
light organs present inside mantle cavity on ventral side of ink sac, deeply embedded in its tissue; closely connected to
nidamental gland in female. Base of right gill of female separated fromme dian and posterior part of mantle cavity by a
septum.
Size: Up to 23 mm mantle length.
Fig. 253 Rondeletiola minor
dorsal view of male
d o r s a l a r m s o f m a l e
( h e c t o c o t y l u s )
Geographical Distribution: Eastern Altantic and
Mediterranean Sea: northwest of Spain, Portugal and
the eastern, central and western Mediterranean Sea
(including Ligurian Sea, northern and southern
Tyrrhenian Sea, Strait of Sicily, Gulf of Taranto,
Adriatic Sea, north Aegean Sea, Sea of Marmara and
Levantine Sea) to the southeastern Atlantic Beguela
Current off Namibia (Fig. 254).
Habitat and Biology: Rondeletiola minor lives on
muddy bottoms. It is a sublittoral, demersal, or upper
bathyal species, with depth a range between 76 and
496 m. In the Sea of Marmara (Mediterranean Sea) the
species was found in brackish waters (salinity between
18 and 25?), which indicates a high degree of
tolerance to fresh water. Recent studies confirm a
common reproductive pattern for the species in the
western and eastern Mediterranean (Aegean Sea): an
extended spawning period is likely to occur, since
mature animals are found throughout the year.
Rondeletiola minor may ascend to the surface during
the reproductive period and has been collected at the
surface at night using artificial light.
Interest to Fisheries: Caught throughout its range and
consumed locally.
Local Names: ITALY: Seppiola minore, Cappuccetto.
Remarks: This species, often reported as rare in some
areas of its distributional range, has recently proved to
be fairly common in several areas of the western
Mediterranean and in the Aegean Sea.
Literature: Naef (1923), Guerra (1982), Roper et al.
(1984), Bello (1990b), Guerra (1992), Villanueva and
S?nchez (1993), Bello (1995), Jereb and DiStefano
(1995), Sartor and Belcari (1995), Villanueva (1995),
Volpi et al. (1995), Wurtz et al. (1995), D?Onghia et al.
(1996), Salman and Katagan (1996), Jereb et al.
(1997), Unsal et al. (1999).
Cephalopods of the World 175
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
 30?  30?
 20?  20?
 10?  10?
0? 0?
10? 10?
20? 20?
30? 30?
40? 40?
Fig. 254 Rondeletiola minor
Known distribution
?? 200 m
Sepiola trirostrata Voss, 1962 F i g . 2 4 5
Sepiola trirostrata Voss, 1962a, Proceedings of the Biological Society of Washington , 75: 172 [type locality: Philippines].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Knobby bobtail squid; Fr ? S?piole
bossel?e; Sp ? Sepiola nudosa.
Diagnostic Features: Fins short, do not exceed length
of mantle anteriorly or posteriorly. Arms III in both
sexes stout and strongly curved inward, more
obviously so in males. Suckers in ventral series of right
arm I and arms II of males larger than dorsal suckers.
Hectocotylus present, left dorsal arm modified:
proximal end with 2 slender fleshy papillae
(anteriormost papilla longest) and dorsolateral to
these a blunt tongue-like lobe , all formed from
enlarged and elongate sucker pedicels; 2 rows of
suckers on arm proximal to fleshy pad; distal end of
hectocotylized arm with sucker pedicels enlarged and
tightly packed to form 2 double rows of columnar
structures; suckers reduced with tiny, fleshy, slit-like
openings. Club with 4 large suckers in transverse
rows; suckers differ in size; dorsal marginal
longitudinal series of suckers larger than those in
ventral marginal series. Paired kidney-shaped light
organs present inside mantle cavity on each side of ink
sac. Colour: Mantle and head with many minute
brown or black chromatophores; arms III deep pink,
arms I to III each with single longitudinal row of large
chromatophores, arms IV with double row of small
chromatophores.
Size: Up to 12.5 mm mantle length.
Geographical Distribution: Indo-Pacific: Philippines,
Singapore (Fig. 246).
Cephalopods of the World 169
Fig. 245 Sepiola trirostrata
(after Voss, 1963 )
dorsal view of male
m a l e a r m a r r a n g e m e n t
tentacle
I I I left
I left
I I left
I right
I V left
hectocotylus
Fig. 246 Sepiola trirostrata
Known distribution
?? 200 m
170 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Interest to Fisheries: Presently undetermined.
Remarks: Differs from S. birostrata Sasaki, 1918 in the presence of the blunt, fleshy papilla, in addition to the 2 slender
papillae on the hectocotylus. The slender papillae arise from a common mound-like base, while the third lobe arises
separately. In this trait, S. trirostrata differs from those S. birostrata specimens that bear a third lobe on the hectocotylus.
Literature: Voss (1963).
Euprymna berryi Sasaki, 1929 F i g . 2 4 7
Euprymna berryi Sasaki, 1929, Journal of the College of Agriculture, Hokkaido Imperial University , 20(supplement): 143
[type locality: Japan, Honshu].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Humming-bird bobtail squid;
Fr ? S?piole colibri; Sp ? Globito colibri.
Diagnostic Features: Fins short, do not exceed length of mantle anteriorly or posteriorly. Non-hectocotylized arm sucker
arrangement same in both sexes: arm suckers tetraserial. Male non-hectocotylized arm suckers enlarged on some arms:
arms II and IV with approximately 10 moderately enlarged suckers in dorsal and ventral rows from proximal second to
fourth suckers; approximately 8 slightly enlarged suckers i n ventral row of arm III from proximal fifth to eighth sucker .
Hectocotylus present; left dorsal arm modified: arm thick, short, and blunt; proximal end with 2 fleshy papillae formed from
enlarged, or elongate sucker pedicels; distal end of hectocotylized arm with sucker pedicels enlarged and tightly packed
forming 2 double rows of 70 to 80 columnar structures; suckers reduced with tiny, fleshy, slit-like openings. Marginal
suckers on dorsal arms I are slightly larger than medial ones. Tentacular club suckers goblet-shaped. Swimming keel of
club extends well beyond carpus.
Size: Males up to 30 mm mantle length; females up to 50 mm mantle length.
(after Okutani et al. , 1995 )
dorsal view of male
Fig. 247 Euprymna berryi
a r m s o f m a l e
(oral view)
I (hc.)
I I I
I I
I V I V
I I I I
I I
Geographical Distribution: Indo-Pacific: Along coasts
of China, south to Hong Kong and Japan (but not in
Hokkaido), Taiwan Province of China, possibly
Andaman Islands and Sri Lanka (Fig. 248).
Habitat and Biology: Upper sublittoral and pelagic in
warm temperate waters; depth range to 107 m. This
species has been reared successfully in aquaculture
experiments. Females lay clumps of round, pale
orange eggs.
Interest to Fisheries: Consumed locally in China and
Taiwan Province of China, where this species is
abundant and sustains local small fisheries; currently
not commercially exploited in Hong Kong.
Local Names: CHINA: Leung yee jai; JAPAN:
Niyori-mimi-ika.
Remarks: The taxonomic status of members of this
genus is largely unresolved (Norman and Lu 1997).
Euprymna berryi is sympatric with, and very similar to
E. morsei Verrill (1881). Non-quantitative differences
between the 2 species include: among 4 rows of arm
suckers, only the ventral rows are enlarged in
E. morsei (particularly from third to fourth suckers on
arms II to IV), but both dorsal and ventral arms have enlarged suckers in E. berryi; the club suckers are spherical, or
cup-shaped in E. morsei, but goblet-shaped in E. berryi. The club suckers have an indistinct proximal border in E. berryi,
while those in E. morsei have distinct borders.
Literature: Roper et al. (1984), Okutani and Horita (1987), Okutani (1995), Norman and Lu (1997), Kubodera and Yamada
(1998), Lu (1998b).
Euprymna morsei (Verrill, 1881) F i g . 2 4 9
Inioteuthis morsei Verrill, 1881, Transactions of the Connecticut Academy of Sciences, 5(6): 417 [type locality: Japan,
Tokyo Bay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Mimika bobtail squid; Fr ? S?piole
mimika; Sp ? Globito mimika.
Cephalopods of the World 171
Fig. 248 Euprymna berryi
Known distribution
?? 200 m
? ?
dorsal view of female
Fig. 249 Euprymna morsei
a r m s o f m a l e
(oral view)
I
I I
I V
I
I I I
hectocotylus
Diagnostic Features: Mantle dome-shaped, plump. Dorsal mantle fused to head. Fins wide, rounded, semicircular; short,
do not exceed length of mantle anteriorly or posteriorly; posteriorly with wide gap between fins; anterior origin posterior to
mantle margin. Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers tetraserial. Ventral rows of
arms II to IV in males with enlarged s uckers, approximately 10 on each arm f rom proximal third or fourth suckers .
Hectocotylus present; distal half of left dorsal arm modified: sucker pedicels enlarged and tightly packed to form 2 double
rows of columnar structures; suckers reduced with tiny, fleshy, slit-like openings; proximal end of arm with single nipple-like
papilla. Marginal suckers arms I slightly larger than medial suckers. Tentacular-club suckers cup-shaped or spherical.
Internal gladius absent. Paired saddle-shaped light organs present inside mantle cavity on ink sac. Colour: Iridescent gold
to purple with large black chromatophores.
Size: Up to 40 mm mantle length.
Geographical Distribution: Indo-Pacific: southern
Japan (sympatric with E. berryi Sasaki, 1929), East
China Sea and the Philippines, south at least to
Indonesia. Possibly Bay of Bengal, India (record based
on females only) and Maldive Islands (Fig. 250).
Habitat and Biology: This neritic species has been
reported from the stomach contents of lancet fishes,
Alepisaurus ferox, captured in the Solomon Sea.
Interest to Fisheries: Harvested on a minor scale,
primarily as trawl bycatch. Low economic value, but
utilized locally.
Remarks: The taxonomic status of members of this
genus is largely unresolved (Norman and Lu 1997).
Literature: Joubin (1902a), Raj and Kalyani (1971),
Okutani et al. (1987), Okutani and Tsukada (1988),
Okutani (1995), Norman and Lu (1997), Kubodera and
Yamada (1998), Reid and Norman (1998).
172 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 250 Euprymna morsei
Known distribution
?? 200 m
?
?
?
Euprymna tasmanica (Pfeffer, 1884) F i g . 2 5 1 ; P l a t e V I , 3 9 ? 4 1
Sepiola tasmanica Pfeffer, 1884, Abhandlungen aus dem Gebiete der Naturwissenschaften, Hamburg , 8(1): 6 [type
locality: Australia: Bass Strait].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Southern bobtail squid; Fr ? S?piole du
Tasmanie; Sp ? Globito de Tasmania.
Diagnostic Features: Dorsal mantle fused to head. Fins wide, rounded, semicircular; short, do not exceed length of mantle
anteriorly or posteriorly. Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers tetraserial. Dorsal
and ventral rows of suckers on arms II to IV in males enlar ged; ventral marginal rows of arms II and III with 1 to 3
greatly enlarged suckers basally (8 to 11% mantle length); dorsal and ventral marginal rows of arms II to IV with more
than 10 enlarged suckers (diameter 4 to 7% mantle length). Hectocotylus present, distal half of left dorsal arm modified:
third and/or fourth proximal suckers in ventral row elongated into long papilla(e), each bearing a tiny sucker; base of
hectocotylus with 29 to 38 normal suckers; distal end of hectocotylized arm with sucker pedicels enlarged and tightly
packed to form 2 double rows of columnar structures; suckers reduced with tiny, fleshy, slit-like openings (30 to 38 in dorsal
rows, 29 to 38 in ventral rows). Tentacular club suckers (many hundreds) all of similar minute size. Saddle-shaped bacterial
light organ inside mantle cavity.
Size: Up to 40 mm mantle length.
Geographical Distribution: Southern Indo-Pacific:
eastern and southeastern Australia, from Brisbane to
Shark Bay, Western Australia (Fig. 252).
Habitat and Biology: This species lives in sandy and
muddy areas, often in association with seagrass beds.
It remains buried in sand during the day, and emerges
at night to forage for small crustaceans and fishes. It
can ?glue? sand grains to its dorsal body surface to aid
in camouflage. Spawning occurs in spring and
summer. Females lay pale orange eggs in loose
clumps, usually at the base of seaweed or seagrass.
Interest to Fisheries: Undetermined, but the species
is likely to be utilized on a local basis.
Remarks: The taxonomic status of members of this
genus is largely unresolved.
Literature: Norman and Lu (1997), Reid and Norman (1998).
Cephalopods of the World 173
Fig. 251 Euprymna tasmanica
t e n t a c u l a r c l u bh e c t o c o t y l u s
dorsal view of male
H
110?
110?
120?
120?
130?
130?
140?
140?
150?
150?
160?
160?
 40?  40?
 30?  30?
 20?  20?
 10?  10?
Fig. 252 Euprymna tasmanica
Known distribution
?? 200 m
174 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Rondeletiola minor (Naef, 1912) F i g . 2 5 3
Sepietta minor Naef, 1912a, Zoologischer Anzeiger , 39(7): 267 [type locality: Tyrrhenian Sea].
Frequent Synonyms: Sepietta minor Naef, 1912.
Misidentifications: None.
FAO Names: En ? Lentil bobtail squid; Fr ? S?piole bobie; Sp ? Globito peque?o.
Diagnostic Features: Mantle bullet-shaped, delicate consistency. Fins small, bluntly pointed laterally rather than curved;
short, do not exceed length of mantle anteriorly or posteriorly. Non-hectocotylized arm sucker arrangement same in both
sexes: arm suckers biserial. Hectocotylus present; left dorsal arm modified: proximal end with fleshy pad formed from
enlarged and fused sucker pedicels; copulatory apparatus a large, swollen, transverse, hood-shaped horn with small
accessory papilla; 3 small suckers proximal to fleshy pad; proximal half of ventral row of suckers distal to copulatory
apparatus with enlarged suckers; oral surface of modified region as for remaining arms, not wide, fleshy; suckers evenly
spaced on modified portion of arm, rows not widely separated. Club with 16 suckers in transverse rows. Paired, roundish
light organs present inside mantle cavity on ventral side of ink sac, deeply embedded in its tissue; closely connected to
nidamental gland in female. Base of right gill of female separated fromme dian and posterior part of mantle cavity by a
septum.
Size: Up to 23 mm mantle length.
Fig. 253 Rondeletiola minor
dorsal view of male
d o r s a l a r m s o f m a l e
( h e c t o c o t y l u s )
Geographical Distribution: Eastern Altantic and
Mediterranean Sea: northwest of Spain, Portugal and
the eastern, central and western Mediterranean Sea
(including Ligurian Sea, northern and southern
Tyrrhenian Sea, Strait of Sicily, Gulf of Taranto,
Adriatic Sea, north Aegean Sea, Sea of Marmara and
Levantine Sea) to the southeastern Atlantic Beguela
Current off Namibia (Fig. 254).
Habitat and Biology: Rondeletiola minor lives on
muddy bottoms. It is a sublittoral, demersal, or upper
bathyal species, with depth a range between 76 and
496 m. In the Sea of Marmara (Mediterranean Sea) the
species was found in brackish waters (salinity between
18 and 25?), which indicates a high degree of
tolerance to fresh water. Recent studies confirm a
common reproductive pattern for the species in the
western and eastern Mediterranean (Aegean Sea): an
extended spawning period is likely to occur, since
mature animals are found throughout the year.
Rondeletiola minor may ascend to the surface during
the reproductive period and has been collected at the
surface at night using artificial light.
Interest to Fisheries: Caught throughout its range and
consumed locally.
Local Names: ITALY: Seppiola minore, Cappuccetto.
Remarks: This species, often reported as rare in some
areas of its distributional range, has recently proved to
be fairly common in several areas of the western
Mediterranean and in the Aegean Sea.
Literature: Naef (1923), Guerra (1982), Roper et al.
(1984), Bello (1990b), Guerra (1992), Villanueva and
S?nchez (1993), Bello (1995), Jereb and DiStefano
(1995), Sartor and Belcari (1995), Villanueva (1995),
Volpi et al. (1995), Wurtz et al. (1995), D?Onghia et al.
(1996), Salman and Katagan (1996), Jereb et al.
(1997), Unsal et al. (1999).
Cephalopods of the World 175
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
 30?  30?
 20?  20?
 10?  10?
0? 0?
10? 10?
20? 20?
30? 30?
40? 40?
Fig. 254 Rondeletiola minor
Known distribution
?? 200 m
Sepietta neglecta Naef, 1916 F i g . 2 5 5
Sepietta neglecta Naef, 1916, Pubblicazioni della Stazione Zoologica di Napoli , 1: 9 [type locality: Tyrrhenian Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Elegant bobtail squid; Fr ? S?piole ?l?gante; Sp ? Sepieta elegante.
Diagnostic Features: Fins rounded, bluntly pointed laterally rather than curved; short, do not exceed length of mantle
anteriorly or posteriorly. Hectocotylus present, left dorsal arm modified: proximal end with fleshy pad formed from enlarged
and/or fused sucker pedicels; copulatory apparatus a dome-shaped lobe medially and short, pointed, horn with smaller papilla
between these; horn of copulatory apparatus slightly recurved, but does not form a small hole; base of hectocotylus proximal to
fleshy pad with 4 normal suckers (not modified); dorsal row of suckers distal to copulatory apparatus with first 3 or 4 suckers
markedly enlarged ; arm broad, spoon-like. Tentacles very thin, delicate. Club with 16 uniform-sized suckers in transverse
rows . Female bursa copulatrix large (extends posteriorly beyond gill insertion). Light organs absent.
Size: Up to 33 mm mantle length.
Geographical Distribution: Northeastern Atlantic and
Mediterranean Sea: southern coast of Norway and
Orkney Islands to Morocco; eastern and western
Mediterranean Sea (Ligurian Sea, Strait of Sicily,
Adriatic Sea, north Aegan Sea, Sea of Marmara and
Levantine Sea) (Fig. 256).
Habitat and Biology: Sepietta neglecta l ives
preferentially on muddy substrates at depths ranging
between 25 and 475 m. It is often associated with
Rossia macrosoma and Sepietta oweniana . It spawns
continuously throughout the year.
Interest to Fisheries: Even though less common,
S. neglecta is caught along with other species of the
genus and is sold and consumed locally.
Remar ks : This species c losely resembles
S. oweniana (Orbigny, 1839?1841) and it can be
difficult to distinguish females of the 2 species under a
certain size (i.e. S. oweniana is larger). This is
particularly true when the tentacle clubs are damaged
or lost during fisheries operations. The clubs differ
between the 2 species: the club is shorter, more
delicate and bears smaller suckers in S. neglecta, than
in S. oweniana . Males are easily identified by the
structure of the hectocotylus.
Literature: Naef (1923), Boletzky et al. (1971), Guescini
and Manfrin (1986), Orsi Relini and Bertuletti (1989),
Bello (1990b), Guerra (1992), Bello (1995), Jereb and
Di Stefano (1995), Volpi et al. (1995), Jereb et al. (1997),
Lefkaditou and Kaspiris (1998).
176 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 255 Sepietta neglecta
m a n t l e c a v i t y o f f e m a l ed o r s a l a r m s o f m a l e ( h e c t o c o t y l u s )
(after Naef, 1923 )
bursa copulatrix
Fig. 256 Sepietta neglecta
Known distribution
?? 200 m
Sepietta obscura Naef, 1916 F i g . 2 5 7
Sepietta obscura Naef, 1916, Pubblicazioni della Stazione Zoologica di Napoli , 1: 4 [type locality: Gulf of Naples].
Frequent Synonyms: None.
Misidentifications: Sepietta oweniana (Orbigny, 1839?1841); Sepiola rondeleti Leach, 1834.
FAO Names: En ? Mysterious bobtail squid; Fr ? S?piole myst?rieuse; Sp ? Sepieta misteriosa.
Diagnostic Features: Fins small, short, almost circular, margin broadly rounded. Tentacles relatively robust; club with 12
suckers in transverse rows, dorsalmost suckers markedly enlarged. Arms II and III with 3 to 5 normal suckers proximally,
followed by 2 larger suckers distally, then smaller ones continuously decreasing in size towards distal tip of the arm.
Hectocotylus present, left dorsal arm modified: 3 normal suckers at base of the arm, followed distally by the copulatory
apparatus: apparatus consists of a transverse ?crest? with four elevations, differing in development and outline, the outer
one usually more prominent, forming a separate, inward curved lobe , the inner one with a simple edge. Distal to the
copulatory apparatus, the first 2 suckers of the dorsal row are markedly enlarged, followed distally by smaller suckers
decreasing in size; stalks of first suckers of ventral rows are elongate and often connected with each other. Distal part of the
arm widened and spoon-shaped. Bursa copulatrix small, not extending beyond gill insertion. Colour: Reddish to dark
brown.
Size: Males up to 19 mm mantle length; females up to 30 mm mantle length.
Geographical Distribution: Mediterranean
Sea, including Ligurian Sea, Tyrrhenian Sea,
Strait of Sicily, Adriatic Sea, northern Aegean
Sea and Levantine Sea. Northeastern Atlantic
(off Portugal) (Fig. 258).
Habitat and Biology: Sepietta obscura lives
preferentially on sandy and muddy bottoms,
often colonized by Posidonia oceanica in
coastal areas. Its depth range is 27 to 376 m. It
is often associated with Sepiola affinis Naef,
1912, and both species seem to have
gregarious habits. A benthic species, Sepietta
obscura has, however, been captured by
pelagic nets, confirming its capacity to leave the
bottom and undergo signif icant vertical
migrations. The smallest mature individuals
measure 12 mm mantle length. The spawning season extends at least from spring to autumn in the Mediterranean.
Females in aquaria were observed to spawn intermittently over a period of 2 weeks, and the animals died afterwards. The
eggs are relatively large (3.7?4.5 mm diameter) but young hatchlings measure about 2 mm mantle length.
Interest to Fisheries: No statistics are available, but the species is frequently captured as a bycatch of trawl and purse
seine fisheries, and can be a relatively abundant component of the bobtails marketed locally.
Local Names: ITALY: Seppiola misteriosa.
Cephalopods of the World 177
Fig. 257 Sepietta obscura
m a n t l e c a v i t y o f f e m a l e(after Naef, 1923 )
h e c t o c o t y l u s
bursa copulatrix
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
40?
40?
30? 30?
40? 40?
Fig. 258 Sepietta obscura
Known distribution
?? 200 m
Remarks: Nesis (1987), in his world review, considered Sepietta obscura a junior name for S. petersi (Steenstrup, 1887).
However, Naef did not mention Steenstrup?s work in his original description of S. obscura (Naef, 1916). This is very unusual
considering Naef?s accuracy and his otherwise detailed and very numerous references to previous workers. It is possible
that he may have overlooked the description of S. petersi when determining the status of his supposed new Sepietta
species. However, considering the current need for a revision of this group, including a proper (re)description of Sepietta
petersi, S. obscura is here retained as the valid name for the species. Long considered a Mediterranean endemic, recent
records of 2 specimens of Sepietta obscura from the waters off Portugal extend the distributional range of this small bobtail
squid.
Literature: Naef (1923), Boletzky et al. (1971), Orsi Relini and Bertuletti (1989), Guerra (1992), Bello and Biagi (1995),
Gabel Deickert (1995), Wurtz et al. (1995), Pereira (1996), Jereb et al. (1997), Salman et al. (2002).
Sepietta oweniana (Orbigny, 1839 ?1841) F i g . 2 5 9
Sepiola oweniana Orbigny, 1839?1841 (in F?russac and d?Orbigny, 1834?1848), Histoire Naturelle G?n?rale et
Particuli?re C?phalopodes Ac?tabuliferes Vivants et Fossiles , pl. 3 [type locality: Tyrrhenian Sea (uncertain)].
Frequent Synonyms: Sepiola oweniana d?Orbigny, 1840.
Misidentifications: None.
FAO Names: En ? Common bobtail squid; Fr ? S?piole commune; Sp ? Sepieta com?n.
Diagnostic Features: Mantle dome-shaped; posteriorly more rounded in females. Fins wide; rounded, semicircular; with
pronounced anterior lobes, or ?earlets? ; short, do not exceed length of mantle anteriorly or posteriorly; attached slightly
obliquely at mantle midline. Arm suckers biserial. Proximal end of arms I fused in males. Hectocotylus present; left dorsal
arm modified: proximal end with fleshy pad formed from enlarged and fused sucker pedicels; copulatory apparatus a fleshy
transverse swelling with long, hook-like inwardly curved horn, deep cleft medially, flask-like rugose bulb, and swelling at
dorsal edge; 2 rows of normal suckers on arm proximal to fleshy pad (usually 2 rows, sometimes 3, 5 or 6 suckers); first 2
or 3 suckers in dorsal row distal to copulatory apparatus gr eatly enlarged, followed by 2 to 4 smaller suckers, then 2
enlarged, then remaining suckers decrease in size to distal tip of arm; ventral rows with moderately enlarged suckers; oral
surface of modified region broad; concave. Club long, well developed, with 32 suckers in transverse rows; all suckers of
similar minute size, giving velvety appearance. Female bursa copulatrix large (exte nds posteriorly beyond gill
insertion). Light organs absent.
178 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 259 Sepietta oweniana dorsal view
t e n t a c u l a r c l u b h e c t o c o t y l u s
Size: Up to 50 mm mantle length in the North Sea;
males up to 35 mm, females up to 40 mm mantle
length in the Mediterranean Sea.
Geographical Distribution: Eastern Atlantic and
Mediterranean Sea: from Norway to the Faeroe
Islands; Morocco and Madeira Islands south to
Mauritania; Mediterranean Sea, including Ligurian
Sea, Strait of Sicily, Aegean Sea, Adriatic Sea, Sea of
Marmara and Levantine Sea. Indian Ocean:
Visakhapatam (single record only) (Fig. 260).
Habitat and Biology: Sepietta oweniana is an
epipelagic?mesopelagic species, occurring within a
wide depth range, i.e. from close to the surface (8 m)
down to over 1 000 m. In the North Atlantic it is most
common between 50 and 300 m and in the
Mediterranean between 100 to 200 and 400 m. It
prefers soft, muddy substrates throughout its
distributional range, and it is often found on shrimp
fishing grounds. Tolerance to salinity variations seems
lower than that observed in other bobtail squids and
S. oweniana has never been found in brackish waters.
Seasonal movements, related to reproduction, and
vertical movements, mostly l inked to trophic
relationships, have been reported for this species, both in the North Atlantic and in the Mediterranean. In the
Mediterranean Sea, mature animals are found throughout the year, suggesting an almost continuous spawning season.
Peaks in reproductive activity, however, are reported for several different areas of the Mediterranean. In the western
Mediterranean, for example, onshore migrations of large individuals occur in late winter to early spring, followed by
spawning in spring and early summer. In the Tyrrhenian Sea, summer spawning peaks have been observed, while in the
Aegean Sea, peaks of spawning occur in April to May and October to November. Mating takes place head-to-head and
spermatophores are placed in the female?s bursa copulatrix. Spawning usually occurs in relatively shallow coastal waters,
though records of egg masses in deeper waters (i.e. down to 200 m) do exist. Several subsequent laying events have been
observed in aquaria. The eggs, spherical to lemon-shaped and greyish white in colour, are deposited on various solid
substrates, both living and dead, with an interesting preference for ascidians (Microcosmus). The duration of egg
development depends upon the water temperature: at 20?C they take about 30 days to hatch, while at 10?C this time range
extends to about 2 months. Growth after hatching, however, seems relatively independent of water temperature, and quite
rapid. Studies in the laboratory indicate that animals grow and mature within 6 months following hatching. The whole life
cycle is, therefore, rather short and may range between 6 and 9 months, depending mainly on the time of embryonic
development. Field data indicates that the mantle length at 50% maturity varies between the Atlantic and the
Mediterranean female populations, and is greater in the Atlantic (i.e. 33 mm versus less than, or equal to, 30 mm). Within
the Mediterranean populations, females from the eastern and central basin are mature at a slightly smaller size compared
with those from the western basin (i.e. 28 and 26 mm respectively, versus 30 mm). Similar differences were not apparent in
males, where the size at 50% maturity varies between 20 and 24 mm both in the Atlantic and in the Mediterranean. Sepietta
oweniana has been cultured successfully in aquaria. Juveniles were fed on mysids ( Praunus flexuosus and P . inermis ),
amphipods (Ericthonius) and large copepods. Adults fed on Pranus flexuosus and the shrimps Palaemon elegans ,
Thoralus cranchii and Crangon crangon . Animals in wild populations feed mainly upon crustaceans; a specific
preference for the euphasiid Maganyctiphanes norvegica in north Atlantic waters and the decapod Pasiphaea sivado in
the northern Tyrrhenian Sea has been observed, supporting hypothesized trophic migrations of S. oweniana in response
to prey abundance and distribution. Feeding occurs primarily from dusk to dawn, with adult animals spending the day
buried in the bottom substrate. Sepietta oweniana is preyed upon by several demersal fish and occasionally by cetaceans.
Interest to Fisheries: This is one of the most abundant bobtail squids throughout its distributional range and one of the
most abundant cephalopods caught in some Mediterranean areas. It represents an important bycatch of many trawl
fisheries, both multispecific and shrimp targeting. Specific statistics are not available, but the species is commonly sold on
Mediterranean markets and it is valued as a delicacy in some areas (e.g. in southern Sicily). In the Mediterranean, catches
are generally most abundant in summer and a marked seasonality has been observed in some areas (e.g. the northern
Tyrrhenian Sea).
Local names: ITALY: Seppiola comune, Cappuccetto.
Remarks: Six specimens were recorded from Visakhapatam in the Indian Ocean (off the northeastern Indian coasts, in the
Bay of Bengal) by Mohan and Rao in 1978. However, no subsequent records exist for the area, or in other parts of the
Indian Ocean.
Literature: Joubin (1902a), Naef (1912b), Naef (1923), Mangold and Froesh (1977), Mohan and Rao (1978), Bergstrom
and Summers (1983), Bergstrom (1985), Roper et al. (1984), Orsi Relini and Massi (1988), Orsi Relini and Bertuletti
(1989), Guerra (1992), Bello (1995), Blanco et al. (1995) Jereb and Di Stefano (1995), Santos et al. (1995), Volpi et al.
(1995), D?Onghia et al. (1996), Bello (1997), Jereb et al. (1997), Salman (1998), Sartor et al. (1998), Belcari and Sartor
(1999a).
Cephalopods of the World 179
 20?
 20?
 10?
 10?
0?
0?
10?
10?
20?
20?
30?
30?
40?
40?
50?
50?
60?
60?
70?
70?
80?
80?
10? 10?
20? 20?
30? 30?
40? 40?
50? 50?
60? 60?
70? 70?
Fig. 260 Sepietta oweniana
Known distribution
?? 200 m ?
180 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES
FOR WHICH ONLY FEW RECORDS EXIST TO DATE
Sepiola aurantiaca Jatta, 1896
Sepiola aurantiaca Jatta, 1896, Fauna und Flora des Golfes von Neapel , 23: 130 [type locality: Tyrrhenian Sea.]
Size: Up to 20 mm mantle length (both sexes).
Geographical Distribution: Northeastern Atlantic, from southern Norway, and western Mediterranean Sea. Outer shelf
and upper bathyal. Depth range possibly from 200 to 400 m. Rare.
Literature: Naef (1912b), Naef (1923), Guerra (1992), Bello (1995).
Sepiola knudseni Adam, 1984
Sepiola knudseni Adam, 1984, Atlantide Report , 13: 157 [type locality: Atl antic Ocean: 06?17'N 03?27'E].
Size: Males up to 8.5 mm mantle length; females up to 18 mm mantle length.
Geographical Distribution: Eastern Atlantic: northwest and west Africa, from the Canary Islands to the Gulf of Guinea.
Inner continental shelf. Depth range from 32 to 90 m.
Literature: Adam (1984).
Sepiola pfefferi Grimpe, 1921
Sepiola pfefferi Grimpe, 1921, Zoologischer Anzeiger , 53(1?2): 4 [type locality: northeaste rn Atlantic: 53?53'N 00?32'E].
Size: Males up to 12 mm mantle length; females up to 13 mm mantle length.
Geographical Distribution: Northeastern Atlantic: Faeroe Islands and southern Norway to Brittany, France. Continental
shelf. Depth range unknown.
Literature: Grimpe (1921).
Sepiola rossiaeformis Pfeffer, 1884
Sepiola rossiaeformis Pfeffer, 1884, Abhandlungen aus dem Gebiete der Naturwissenschaften, Hamburg , 8(1): 8 [type
locality: Java].
Size: Type 6.0 mm mantle length.
Geographical Distribution: Indo-Pacific: unknown.
Literature: Pfeffer (1884).
Sepiola steenstrupiana Levy, 1912
Sepiola steenstrupiana Levy, 1912, Archives de Zoologie Experimentale et G ?n?rale, (series 5)9, Notes et Revue , 3: LVI
[type locality : ?Villafranca?.].
Size: Up to 30 mm mantle length.
Geographical Distribution: Mediterranean Sea: including central Tyrrhenian Sea, Adriatic Sea, Aegean Sea and
Levantine Sea. Red Sea, Gulf of Aden. Indian Ocean: Somalia.
Literature: Naef (1923), Guerra (1992), Bello (1995), Rocha et al. (1998).
Euprymna albatrossae Voss, 1962
Euprymna albatrossae Voss, 1962a, Proceedings of the Biological Society of Washington , 75: 171 [type locality:
Philippines].
Size: Up to 24 mm mantle length.
Geographical Distribution: Western Pacific: Philippines. Depth range unknown. Types collected using nightlight.
Literature: Voss (1963).
Euprymna hoylei Adam, 1986
Euprymna hoylei Adam, 1986, Bulletin de l?Institut royal des Sciences naturelles de Belgique , 56: 133 [type locality: Sulu
Archipelago].
Geographical Distribution: Tropical Indo-Pacific: western tropical Pacific and northwestern Australia.
Literature: Norman and Lu (1997).
Euprymna hyllebergi Nateewathana, 1997
Euprymna hyllebergi Nateewathana, 1997, Phuket Marine Biological Center Special Publication , 17(2): 466 [type
locality: Thailand: Andaman Sea, Trang Province, Kantang Fish Landing].
Size: Up to 35 mm mantle length.
Geographical Distribution: Eastern Indian Ocean: Thailand, Andaman Sea. Depth to 74 m.
Literature: Nateewathana (1997).
Euprymna penares ( Gray, 1849)
Fidenas penares Gray, 1849, Catalogue of the Mollusca in the British Museum. Part I. Cephalopoda Antepedia , 95 [type
locality: Singapore].
Geographical Distribution: Indo-Pacific: unknown.
Literature: Gray (1849).
Euprymna phenax Voss, 1962
Euprymna phenax Voss, 1962a, Proceedings of the Biological Society of Washington , 75: 171 [type locality: Philippines].
Size: Type 11 mm mantle length.
GeographicalDistribution: Indo-Pacific: Philippines, possibly East China Sea. Depth range unknown (collected at nightlight).
Literature: Kubodera and Yamada (1998).
Euprymna scolopes Berry, 1913
Euprymna scolopes Berry, 1913, Proceedings of the United States National Museum , 45(1996): 564 [type locality:
Hawaiian Islands].
Size: Up to 30 mm mantle length.
Geographical Distribution: Central Pacific: Hawaiian Islands. Shallow coastal waters.
Literature: Arnold (1972), Moynihan (1983a), Shears (1988), McFall-Ngai and Montgomery (1990), McFall-Ngai (1994),
Fleisher and Case (1995), Montgomery and McFall-Ngai (1995), Anderson and Mather (1996), Hanlon et al. (1997),
Norman and Lu (1997), Claes and Dunlap (1999).
Cephalopods of the World 181
182 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Euprymna stenodactyla ( Grant, 1833 )
Euprymna stenodactyla Grant, 1833, Transactions of the Zoological Society of London , 1: 85 [type locality: Mauritius].
Geographical Distribution: Indian Ocean: known with certainty only from Mauritius. Records from the Indo-west Pacific
(Mascarene Islands to Queensland, Australia and Polynesia) are doubtful.
Literature: Joubin (1902a), Okutani (1995), Norman and Lu (2000).
Sepietta petersi (Steenstrup, 1887 )
Sepiola petersi Steenstrup, 1887, Oversigt over det Kongelige Danske Videnskabernes Selskabs Forhandlinger ,
1887: 58 [type locality: Adriatic Sea].
Geographical Distribution: Mediterranean Sea and Atlantic coast of Morocco (doubtful).
Remarks: Doubtful species, that has been considered the senior name for S. obscura Naef, 1916 (see remarks in
S. obscura). A detailed redescription of all the anatomical features of this purported species is needed to solve the problem.
Literature: Steenstrup (1887).
Inioteuthis capensis Voss, 1962
Inioteuthis capensis Voss, 1962b, Transactions of the Royal Society of South Africa , 36(4): 255 [type locality: South
Africa].
Geographical Distribution: Southeastern Atlantic: South Africa, from L?deritz Bay to Mossel Bay.
Literature: Voss (1962b).
Inioteuthis japonica (Orbigny, 1845)
Sepiola japonica Orbigny, 1845 (in F?russac and d?Orbigny, 1834?1848), Histoire Naturelle G?n?rale et Particuli?re
C?phalopodes Ac?tabuliferes Vivants et Fossiles , 234 [type locality: Japan].
Size: Up to 20 mm mantle length.
Geographical Distribution: Western Pacific: Southern Japan, China and Taiwan Province of China.
Literature: Joubin (1902a).
Inioteuthis maculosa Goodrich, 1896
Inioteuthis maculosa Goodrich, 1896, Transactions of the Linnaean Society of London , (series 2, Zoology), 7(1): 2 [type
locality: Andaman Islands].
Size: Males up to 13 mm mantle length; females up to 14 mm mantle length.
Geographical Distribution: Indo-Pacific: northern Indian Ocean, Persian Gulf, India, Arabian Sea, Bay of Bengal,
Andaman Sea, Taiwan Province of China, Philippines, Indonesia.
Literature: Nateewathana (1997).
3.2.2 S u b f a m i l y R O S S I I N A E Appell?f, 1898
Rossia macrosoma ( Delle Chiaie, 1830 ) F i g . 2 6 1
Sepiola macrosoma Delle Chiaie, 1830, Memoire sulla storia e notomia degli Ani mali senza vertebre del Regno di Napoli .
4 volumes, atlas. Napoli, pl. 17 [type locality: Tyrrhenian Sea].
Frequent Synonyms: Sepiola macrosoma Delle Chiaie, 1829.
Misidentifications: None.
FAO Names: En ? Stout bobtail squid; Fr ? S?piole melon; Sp ? Globito robusto.
Diagnostic Features: Body smooth, soft. Males mature at smaller sizes and do not grow as large as females. Mantle
dome-shaped. Dorsal mantle free from head (not fused to head). Nuchal cartilage oval, broad . Fins short, do not exceed
length of mantle anteriorly or posteriorly. Arm webs broad between arms III and IV. Non-hectocotylized arm sucker
arrangement same in both sexes: arm suckers biserial basally, tetraserial medially and distally . Dorsal and ventral
sucker rows of arms II to IV of males enlarged; ventral marginal rows of arms II and III with 1 to 3 greatly enlarged suckers
basally (diameter 8 to 11% mantle length); dorsal and ventral marginal sucker rows of arms II to IV with more than 10
enlarged suckers (diameter 4 to 7% mantle length); suckers on median rows in males smaller than female arm suckers in
size. Hectocotylus present; both dorsal arms modified: ventrolateral edge of proximal oral surface of hectocotylized arms
bordered by swollen glandular crest, inner edge of which forms a deep furrow; glandular crest extends over entire arm
length; suckers decrease in size from proximal to distal end of arms; biserial proximally, tetraserial distally (marginal and
medial suckers similar in size, smaller than on rest of arm); arms with deep median furrow and with transversely grooved
ridges. Tentacular club expanded, broader than stalk, with 8 to 12 suckers in transverse rows ; suckers small, very similar
in size ; protective membrane borders entire club; swimming keel equal to club length . Anal flaps well developed. Ink sac
well developed. Epirenal bodies absent; anal pads absent . Colour: Light yellowish brown with greenish tinge to dark
reddish brown.
Cephalopods of the World 183
Fig. 261 Rossia macrosoma
dorsal view
a r mt e n t a c u l a r c l u b
Size: Up to 85 mm mantle length, usually between 20
to 60 mm mantle length.
Geographical Distribution: Eastern Atlantic and
Mediterranean Sea: Greenland Sea, off Greenland
and Iceland, Norway, Faeroe Islands, North Sea,
Britain to the Azores, Morocco and Senegal;
Mediterranean Sea except northern Adriatic Sea and
southeastern Levantine Sea (Fig. 262).
Habitat and Biology: Sandy and muddy substrate.
Demersal, depth range 32 to 899 m; animals have
been collected between the surface and 500 m at
night. This species is common at depths between 200
and 400 m in the Mediterranean Sea, where it is
associated wi th the wide transi t ional zone
(100?600 m) representing a region of overlapping
shelf and slope faunas. It usually prefers deeper
waters, especially in winter, ascending to shallower
waters to spawn. It is fairly frequent in catches but
generally never abundant, thus information on its
biology and ecology is still poor. Most of the present
knowledge is based on the observations made in the
western Mediterranean and studies in the laboratory.
In the western Mediterranean Rossia macrosoma
carries out seasonal migrations between deeper
offshore waters in winter and shallower coastal zones for the rest of the year. This migration is partitioned by size such that
the largest individuals arrive first in spring, followed by smaller animals in summer. Mature males, aged 7 to 8 months carry
85 to 100 spermatophores; females, 8 to 11 months, have about 120 to 150 eggs in their ovaries. The spawning season
probably extends throughout the year throughout the whole Mediterranean, with peaks in spring and autumn. Mating takes
place when the male grasps the female?s ?neck? region and then inserts the hectocotylus into her mantle cavity.
Spermatophores have been found attached to the oviducal openings in this species. Rossia macrosoma is polytelic, each
individual spawning several times. Eggs, measuring 7 to 8 mm in diameter, are laid in small clusters of 30 to 40, and
covered by a violet-red hard coat. They are deposited on bivalve shells e.g. of Pinna sp., or other solid substrates.
Hatching occurs after 45 days at temperatures of about 16? C. Females grow larger than males, and longevity is about 12
months. Smallest mature males are 30 mm mantle length; smallest females 35 mm mantle length.
Interest to Fisheries: Rossia macrosoma is a species of relatively minor commercial importance, taken as bycatch in
bottom trawls, mostly between 200 and 400 m. Its commercial value varies among Mediterranean countries. The flesh is
tasty but difficult to preserve. Separate statistics are not reported for this species, which, however, is rather common in fish
markets and sold fresh and frozen.
Local Names: FRANCE: S?piole; ITALY: Seppiola grossa, Babbunedda, Cape e chiuove, Capo di chiodo, Purpo seccia,
Vurpascele; MOROCCO: Sepiole; SPAIN: Choco, Chopito, Globito.
Literature: Joubin (1895, 1902b), Naef (1923), Mangold-Wirtz (1963), Boletzky and Boletzky (1973), Roper et al. (1984),
Bello (1990b), Guerra (1992), Bello (1995), Jereb and Di Stefano (1995), Okutani (1995), Sartor and Belcari (1995), Volpi
et al. (1995), Wurtz et al. (1995), D?Onghia et al. (1996), Jereb et al. (1998), Belcari and Sartor (1999b), Quetglas et al.
(2000).
184 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 262 Rossia macrosoma
Known distribution
?? 200 m
?
?
Rossia pacifica pacifica Berry, 1911 F i g . 2 6 3
Rossia pacifica pacifica Berry, 1911a, Proceedings of the United States National Museum , 40(1838): 591 [type locality:
Alaska].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? North Pacific bobtail squid; Fr ? S?piole du Pacifique bor?al; Sp ? Globito del Pac?fico boreal.
Diagnostic Features: Body smooth, soft. Males mature at smaller sizes and do not grow as large as females. Mantle
dome-shaped. Dorsal mantle free from head (not fused to head). Nuchal cartilage oval, narrow . Fins ear-shaped. Male
and female arms short, stout; not united by deep webs. Non-hectocotylized arm sucker arrangement same in both sexes:
suckers biserial basally, tetraserial medially and distally; male arm suckers enlarged on arms II to IV, larger than female
arm suckers. Hectocotylus present; both dorsal arms modified: ventrolateral edge of proximal oral surface of
hectocotylized arms bordered by swollen glandular crest, inner edge of which forms a deep furrow; glandular crest
extends over 2/3 arm length ; suckers smaller than on sessile arms (about 1/3 the size of those on lateral arms). Tentacular
club expanded, broader than stalk; sucker-bearing face flattened, with 6 to 8 suckers in transverse rows proximally ,
4 rows distally; suckers differ slightly in size, dorsal suckers largest; protective membrane borders entire club. Lower beak
with tooth on shoulder. Anal flaps well developed. Ink sac well developed. Epirenal bodies and anal pads absent. Internal
gladius present, chitinous; gladius spoon-shaped, short.
Size: Males up to 45 mm mantle length; females up to 90 mm mantle length.
Cephalopods of the World 185
Fig. 263 Rossia pacifica pacifica
t e n t a c u l a r c l u b
dorsal view
Geographical Distribution: Northern Pacific: Bering Sea, Sea of Okhotsk, Kamchatka, Kuril Islands, Japan to Korea,
Aleutian Islands and Gulf of Alaska south to British Columbia, San Diego and California (to about 28?N off Baja California)
(Fig. 264).
Habitat and Biology: Neritic; depth range 30 to 310 m (possibly to 550 m) in the western Pacific; 10 m and deeper in the
eastern Pacific (maximum depth undetermined). This species is night active; it remains buried in sand and mud during the
day, then emerges at night to forage. Shrimps, crabs, small fishes and cephalopods comprise over 80% of the diet of this
species in its natural environment. Females are polytelic: ova ripen in small clusters, each individual spawning several
times, probably throughout the year. In the northeastern Pacific, eggs are attached singly, or in small groups, to seaweed
and other objects on the sea floor. Egg masses are found in the Greater Puget Sound Region in calm water, with little tidal
flow, attached to smooth, hard, surfaces protected from siltation by overhanging rocks. The eggs are about 1 cm in
diameter and ovoid, with a flattened area of attachment; they are usually found at depths from 15 to 30 m, but are reported
from 250 m depth (1.6?C) in the Bering Sea. This species has been reared successfully in captivity; egg development took
nearly 5 months both in an open seawater system, with temperatures ranging between 6? and 15?C reflecting the ambient
temperature, and in a temperature controlled environment at 10?C (? 1?C). The time of embryonic development, however,
seems linked to the lunar cycle. Because this species is known to occur over a wide depth range and lacks a planktonic
juvenile stage, hatching of eggs laid far below the photic zone would be out of synchrony with those laid at shallow depths
based on light effects alone. Growth in captivity is strongly dimorphic, with females growing much faster than males after a
first phase of rather slow growth common to both sexes. The lifespan from hatching to death is 18 to 19 months.
Accordingly, the time taken for embryonic development was recently estimated to be about 5 months for animals of the
Bering, Okhotsk (Russia) and northern seas of Russia and the Polar Basin. However, results obtained from statolith
microstructure in a few specimens of Rossia pacifica from the northwest Bering Sea, would suggest a much shorter
lifespan (i.e. 4?5 months).
Interest to Fisheries: Rossia pacifica often is trawled in large quantities off the Sanriku-Hokkaido coasts of Japan and
other subarctic Pacific regions. It is believed to have inferior meat quality and therefore has low economic value, even
though the resource is large. Separate statistics are not reported for this species.
Local Names: JAPAN: Bouzuika.
Remarks: This species seems most closely related to the Arctic R. moelleri Steenstrup, 1856, suggesting its possible
origin by isolation during the Tertiary elevations of the Bering Strait. Rossia pacifica diegensis Berry, 1912 is found at
greater depths than R. pacifica pacifica; it is smaller and more delicate in structure, and it has relatively larger fins and arm
suckers, predominantly in 2 rows. Some large Rossia eggs (14?18 mm) have been found in 1 204 to 1 222 m (4.1?C) depth
in Santa Catalina Basin. These may belong to R. pacifica diegensis, or it may be that R. pacifica pacifica produces larger
eggs at lower temperature in this location. Alternatively, another Rossia taxon may exist in these waters, and R. pacifica
pacifica may actually be a cold water, large-egged form, with the second taxon an unnamed smaller-egged species found
in shallower water.
Literature: Sasaki (1920), Sasaki (1929), Mercer (1968), Boletzky (1970), Brocco (1971), Hochberg and Fields (1980),
Roper et al. (1984), Okutani et al. (1987), Summers and Colvin (1989), Summers (1992), Anderson and Shimek (1994),
Arkhipkin (1995), Okutani (1995), Mangold et al. (1998), Nesis (1999).
186 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 264 Rossia pacifica pacifica
Known distribution
?? 200 m
Rossia tortugaensis Voss, 1956 F i g . 2 6 5
Rossia tortugaensis Voss, 1956, Bulletin of Marine Science of the Gulf and Caribbean , 6(2): 103 [type locality: Caribbean
Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Tortuga bobtail squid; Fr ? S?piole de la Tortue; Sp ? Globito de Tortugas.
Diagnostic Features: Body soft, fleshy. Males mature at
smaller sizes and do not grow as large as females. Dorsal
mantle free from head (not fused to head). Fins ovate,
short. Male and female arms long. Non-hectocotylized
arm sucker arrangement same in both sexes: arm
suckers biserial. Suckers elongated, barrel-shaped,
without dentition. Hectocotylus present, both dorsal arms
modified: glandular crest extends over entire arm
length. Club large; sucker-bearing face flattened;
tentacular club expanded, broader than stalk, with 10
suckers in transverse rows ; suckers finely toothed;
swimming keel of club extends slightly proximal to
carpus. Anal flaps reduced. Ink sac well developed.
Epirenal bodies and anal pads absent.
Size: Up to 50 mm mantle length.
Geographical Distribution: Tropical western Atlantic:
Gulf of Mexico, Dry Tortugas, Florida and off Suriname
(Fig. 266).
Habitat and Biology: Depth range 520 m (Dry Tortugas)
to 760 m (Suriname). These are the only known records.
This species is polytelic: ova ripen in small clusters, each
individual spawning several times, probably throughout
the year.
Interest to Fisheries: Undetermined.
Literature: Roper et al. (1984), Okutani (1995).
Cephalopods of the World 187
Fig. 265 Rossia tortugaensis
dorsal view
Fig. 266 Rossia tortugaensis
Known distribution
?? 200 m
Semirossia equalis (Voss, 1950 ) F i g . 2 6 7
Rossia (Semirossia) equalis Voss, 1950, Revista de la Sociedad Malacol?gica ?Carlos de la Tarre? , 7(2): 73 [type locality:
Caribbean Sea].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Greater shining bobtail squid;
Fr ? S?piole cracheuse; Sp ? Globito reluciente.
Diagnostic Features: Fins wide, ovate, short, do not
exceed length of mantle anteriorly or posteriorly;
anterior fin lobes prominent. Male and female arms
long, slender. Non-hectocotylized arm sucker
arrangement same in both sexes: arm suckers
biserial, widely spaced; median arm suckers enlarged.
Suckers globular, barrel-shaped with untoothed rings.
Hectocotylus present, left dorsal arm modified:
suckers normal proximally, reduced distally; broad
lateral membrane present, extending from third
sucker pair for about 3/4 of arm length ; sucker
arrangement from proximal to distal end of arm:
10 series of normal suckers, 4 series of reduced
suckers ; oral surface of modified region swollen,
fleshy, sucker pedicels form palisade effect, with
transversely grooved ridges. Club moderate length,
expanded, broader than stalk, with 7 or 8 suckers in
transverse rows ; suckers differ in size, dorsal suckers
largest, those in two dorsalmost rows larger than rest;
length of swimming keel of club equal to length of
carpus. Light organs present inside mantle cavity on
ink sac. Colour: Deep purple; fins pigmented over
basal half.
Size: Up to 50 mm mantle length.
Geographical Distribution: Tropical western Atlantic:
eastern Gulf of Mexico to Florida, Cuba, the
Caribbean Sea and Suriname (Fig. 268).
Habitat and Biology: Muddy substrates. Benthic;
depth range from 130 to 260 m, but specimens have
been collected in the water column between the
surface and 115 m at night.
Interest to Fisheries: No specific information
available, but it is likely to be used as food locally.
Remarks: Differs from S. tenera (Verrill 1880) in its
larger size, smaller arm suckers and slightly enlarged
suckers on the dorsal side of the club.
Literature: Roper et al. (1984), Okutani (1995).
188 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 267 Semirossia equalis
dorsal view
Fig. 268 Semirossia equalis
Known distribution
?? 200 m
Semirossia tenera (Verrill, 1880 ) F i g . 2 6 9
Heteroteuthis tenera Verrill, 1880, American Journal of Science , (series 3) 20(41): 392 [type locality: northwestern
Atlantic Ocean].
Frequent Synonyms: Rossia tenera (Verrill, 1880); Heteroteuthis tenera Verrill, 1880.
Misidentifications: None.
FAO Names: En ? Lesser shining bobtail squid;
Fr ? S?piole calamarette; Sp ? Globito tierno.
Diagnostic Features: Small; body soft, fleshy. Mantle
dome-shaped, broad. Dorsal mantle free from head (not fused
to head). Fins ovate. Male and female arms moderate length.
Arms III and IV united by a web . Arm suckers greatly enlarged
in middle section of arms in males, abruptly decrease in size
distally near arm tip. Hectocotylus present; left dorsal arm
modified: suckers normal proximally, reduced distally; broad
lateral membrane present ventrally, and extends distally for
about 3/4 of arm length from third basal pair of suckers ;
sucker arrangement from proximal to distal end of arm: 7
series of normal suckers, 4 series of reduced suckers ; oral
surface of modified region swollen, fleshy, sucker pedicels
form a pallisade effect, with transversely grooved ridges.
Tentacular club moderately expanded, broader than stalk; club
with 6 or 7 suckers in transverse rows ; suckers differ in size,
dorsal suckers twice as large as rest ; suckers toothed around
entire margin; length of club swimming keel equal to length of
carpus. Light organs present inside mantle cavity, on ink sac.
Size: Up to 50 mm mantle length.
Geographical Distribution: Western North Atlantic:
widespread around the eastern coast of North America, from
Nova Scotia, the northern Gulf of Maine to the Gulf of Mexico
and the Caribbean Sea; possibly also present in the
southwestern Atlantic, along the coasts of Suriname, French
Guiana, Brazil and Uruguay (latter records questionable).
(Fig. 270).
Habitat and Biology: Sandy and muddy substrates. Demersal;
depth range from 85 to 135 m in New England area (USA).
Cephalopods of the World 189
Fig. 269 Semirossia tenera dorsal view, male
o r a l v i e w o f m a l e
Fig. 270 Semirossia tenera
Known distribution
?? 200 m
?
?
?
?
190 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Interest to Fisheries: Reported to be fished in the Gulf of San Matias and some localities on the south coast of Argentina
(though species identification needs to be verified).
Local Names: ARGENTINA: Calamarcito.
Literature: Joubin (1902b), Voss (1956), Roper et al. (1984), Okutani (1995).
Neorossia caroli ( Joubin, 1902 ) F i g . 2 7 1
Rossia caroli Joubin, 1902b, Bulletin de la Soci?t? Zoologique de France , 27: 138 [type locality: Eastern Atlantic: Azores
Islands].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Carol bobtail squid; Fr ? S?piole de Carol; Sp ? Globito de Carol.
Diagnostic Features: (Based on type and specimens from the eastern Atlantic.) Body soft, fleshy. Mantle broad, oval;
dorsal anterior margin slightly convex; posterior mantle margin rounded. Dorsal mantle free from head (not fused to head).
Nuchal cartilage oval . Fins wide, rounded, semicircular; anterior origin almost at mantle margin. Funnel organ dorsal
elements inverted V-shaped with small anterior papilla, median limbs with broad, blunt lobes; ventral elements oval with
acute anterior tips. Mantle-locking cartilage a simple straight ridge; funnel-locking cartilage a simple, straight depression.
Head slightly broader than mantle; eyes large. Male and female arms similar in relative lengths; arm formula 3,4,2,1, or
3,2,4,1. Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers biserial; largest suckers on arms II
and III larger than those on arms I and IV in both sexes . Hectocotylus present, both dorsal arms modified: ventrolateral
edge of proximal oral surface of hectocotylized arms bordered by swollen glandular crest, inner edge of which forms a deep
furrow; inside crest fleshy, with transverse grooves; oral surface broad, fleshy; suckers smaller than on remaining arms;
glandular crest extends over sucker rows 3 to 18 . Club slightly recurved, sucker-bearing face convex with 8 to 11 suckers
in transverse rows; all suckers of similar size; swimming keel of club extends well proximal to carpus. Anal flaps present,
reduced. Ink sac greatly reduced, non functional. Epirenal bodies present in males only. Internal gladius present,
chitinous; gladius elongate, slender, diamond-shaped anteriorly, tapers posteriorly; length approximately equal to mantle
length; rachis extends posteriorly for 2/3 length of vane; vane present only on posterior half of gladius .
Size: Males up to 51 mm mantle length; females up to 83 mm mantle length.
Fig. 271 Neorossia caroli
dorsal viewh e c t o c o t y l u s
Geographical Distribution: Eastern Atlantic and
Mediterranean Sea: from southwestern Iceland
and Ireland southward to the Gulf of Guinea and
Namibian coast of southern Africa; in the
Mediterranean Sea, from the northwestern
Mediterranean eastward to the Ligurian Sea,
northern and southern Tyrrhenian Sea, Strait of
Sicily, Adriatic Sea, northern Aegean Sea,
Levantine Sea and Algerian Sea. Doubtful
records from the southern slope of the Great
Newfoundland Bank, the slope off Nova Scotia,
and the Gulf of Mexico. Southwestern Atlantic:
(subspecies N. c. jeannae Nesis et al. (2001))
Patagonian slope north of Falkland Islands
(Fig. 272).
Habitat and Biology: Depth range from 40 to
1 744 m. Neorossia caroli is the most bathyal
among the species belonging to the family
Sepiolidae, collected down to the greatest depths
in the western Mediterranean basin (1 744 m)
and in the eastern Atlantic (1 535 m). It is a
demersal species living preferentially on deep
muddy bottoms characterized by Isidella
elongata populations, often overlapping with
Rossia macrosoma in the upper level of its
distributional range and frequently associated
with Sepietta oweniana and Rondeletiola minor.
In the western Mediterranean, it is the most
common cephalopod captured between 1 000
and 2 000 m, along with Bathypolypus sponsalis.
The occurrence of small individuals of both
N. caroli and B. sponsalis at greater depths than
larger individuals, suggests there may be an
upslope ontogenetic migration. Neorossia caroli
is most abundant between 400 and 600 to 700 m
both in the western and eastern Mediterranean, although it is also present on the upper slope (200?400 m), and there are
sporadic records of its occurrence in shallower waters (less than 100 m). Neorossia c. jeannae has been collected between
474 and 670 m. Mature individuals are found throughout the year, suggesting an extended spawning season, probably with
peaks in summer and/or autumn. The smallest mature males are 35 mm, and the smallest mature females 50 mm mantle
length. Lower sizes at first maturity have, however, been reported for specimens from the central Mediterranean (i.e. Strait
of Sicily: 30.5 mm for males and 35 mm for females). Eggs are large (8?10 mm diameter), covered by a hard
violet-coloured coating and are attached to hard substrates at various depths. The lifespan is probably between 12 and 24
months.
Interest to Fisheries: Of minor commercial importance, it is taken usually as trawl fishery bycatch. Separate statistics are
not reported for this species. It is sold fresh and frozen in fish markets with Rossia macrosoma and other bobtail squids.
Local names: ITALY: Seppiola grossa di fondo.
Remarks: Differs from its congener, N. leptodons Reid, 1992, in the shape of the radula teeth. The rhachidian teeth and
first lateral teeth have broad, rather than narrow, bases and the base usually is strongly indented. Neorossia c. jeannae
differs from N. c. caroli only in the shape of the size of the fins and nuchal cartilage (smaller in N. c. jeannae ) and shape of
the nuchal cartilage. Future study and comparison of specimens from the entire range of N. caroli will determine the status
of the two subspecies, and (possibly) help in revaluating the genus.
Literature: Joubin (1902b), Chun (1913), Joubin (1924), Mercer (1968), Boletzky (1971), Bello (1990b), Salcedo-Vargas
(1991), Guerra (1992), Reid (1992), Villanueva (1992), Bello (1995), Jereb and Di Stefano (1995), Sartor and Belcari
(1995), Volpi et al. (1995), Wurtz et al. (1995), D?Onghia et al. (1996), Jereb et al. (1998), Collins et al. (2001), Nesis et al.
(2001).
Cephalopods of the World 191
 80?
 80?
 60?
 60?
 40?
 40?
 20?
 20?
0?
0?
20?
20?
40?
40?
 50?  50?
 30?  30?
 10?  10?
10? 10?
30? 30?
50? 50?
70? 70?
Fig. 272 Neorossia caroli
Known distribution
?? 200 m
?
??
Austrorossia antillensis (Voss, 1955 ) F i g . 2 7 3
Rossia antillensis Voss, 1955, Bulletin of Marine Science of the Gulf and Caribbean , 5(2): 86 [type locality: Cuba].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Antilles bobtail squid; Fr ? S?piole mignonne; Sp ? Globito antillano.
Diagnostic Features: Body soft, fleshy. Males mature at smaller sizes and do not grow as large as females. Mantle oval,
short, saccular. Dorsal mantle free from head (not fused to head). Fins wide; ovate; short, do not exceed length of mantle
anteriorly or posteriorly. Head slightly broader than mantle; eyes large. Male and female arms moderate length.
Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers biserial, suckers sparsely arranged;
median arm suckers enlarged in males; larger than female arm suckers. Hectocotylus present, both dorsal arms modified:
ventrolateral edge of proximal oral surface of hectocotylized arms bordered by swollen glandular crest, inner edge of which
forms a deep furrow; glandular crest extends over
sucker rows 3 to 8 . Club slightly recurved, short;
sucker-bearing face convex; tentacular club not
expanded, same width as stalk; with 30 to 40 suckers
in transverse rows ; all suckers of similar minute size;
swimming keel of club extends slightly proximal to
carpus. Anal flaps well developed. Epirenal bodies
and anal pads absent. Ink sac well developed.
Colour: Pinkish brown, with scattered dark purple
chromatophores; fins pigmented the same as the
mantle; dorsal pigmentation extends to ventral surface
of fins.
Size: Up to 90 mm mantle length.
Geographical Distribution: Tropical western Atlantic:
Caribbean Sea, Cuba, Dry Tortugas, and the Gulf of
Mexico northward to the latitude of Tampa, Florida.
Suriname (Fig. 274).
192 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 273 Austrorossia antillensis
dorsal viewt e n t a c u l a r c l u b
Fig. 274 Austrorossia antillensis
Known distribution
?? 200 m
Habitat and Biology: Demersal; depth range from 540 to 700 m. This species is polytelic: ova ripen in small clusters, each
individual spawning several times, probably throughout the year.
Interest to Fisheries: Undetermined, but it is likely to be consumed locally when captured as a bycatch of demersal trawl
fisheries.
Remarks: Austrorossia antillensis is a large species in the genus. It is replaced by Rossia bullisi Voss, 1956 in more
northerly waters.
Literature: Boletzky (1970), Roper et al. (1984), Okutani (1995).
Austrorossia australis Berry, 1918 F i g . 2 7 5
Rossia (Austrorossia) australis Berry, 1918, Biological Results of the Fishing Experiments carried on by the F.I.S.
?Endeavour,? 1909?14 , 4(5): 253 [type locality: southwestern Australia : Great Australian Bight southwest of Eucla,
Western Australia].
Frequent Synonyms: None.
Misidentification: None.
FAO Names: En ? Big bottom bobtail squid; Fr ? S?piole australe; Sp ? Globito austral.
Diagnostic Features: Body smooth, soft. Males mature at smaller sizes and do not grow as large as females. Dorsal
mantle free from head (not fused to head). Nuchal cartilage oblong, rounded anteriorly, tapers to slightly narrower
posteriorly. Fins wide, ovate, short, do not exceed length of mantle anteriorly or posteriorly; attached within anterior 2/3 of
mantle. Non-hectocotylized arm sucker arrangement same in both sexes: arm suckers biserial; largest suckers on arms II
and III larger than those on arms I and IV in both sexes. Hectocotylus present, both dorsal arms modified: ventrolateral
edge of proximal oral surface of hectocotylized arms bordered by swollen glandular crest, inner edge of which forms a deep
furrow; glandular crest extends from sucker rows 4?6 to 8?11 . Club straight, slender, long; sucker-bearing face convex;
males 18 to 26 suckers in transverse rows; females 25 to 33 suckers in transverse rows ; all suckers of similar minute
size. Anal flaps well developed. Ink sac well developed. Epirenal bodies present inmales only; anal pads present in both
sexes . Internal gladius present, chitinous; length approximately equal to mantle length; rachis extends almost to posterior
tip of vane; vane extends entire length of gladius ; shape lanceolate. Colour: Uniform pinkish to purplish brown.
Cephalopods of the World 193
Fig. 275 Austrorossia australis dorsal view of female
n u c h a l - l o c k i n g c a r t i l a g e t e n t a c u l a r c l u b
(illustrations: K. Hollis/ABRS)
Size: Males up to 34 mm mantle length; females up to
63 mm mantle length.
Geographical Distribution: Indo-Pacific: Australia,
Queensland, Raine Island, 11?35'S 144?04'E, to
Western Australia, Great Australian Bight, 34?S
130?50'E (Fig. 276).
Habitat and Biology: Sandy and muddy substrates.
Benthic; depth range from 131 to 665 m. This species
is polytelic: ova ripen in small clusters, each individual
female spawns several times, probably throughout the
year.
Interest to Fisheries: Possibly minor fishery potential.
Remarks: A second Rossia species is found on the
northwest shelf of Western Australia. While it has been
shown to differ from R. australis, primarily in
quantitative characters, it cannot be distinguished
from the African Austrorossia mastigophora (Chun,
1915). Further work is needed to clarify the status of
this species.
Literature: Reid (1992), Okutani (1995), Reid and Norman (1998).
Austrorossia bipapillata (Sasaki, 1920 ) F i g . 2 7 7
Rossia bipapillata Sasaki, 1920, Proceedings of the United States National Museum , 57(2310): 190 [type locality: Japan,
Suruga Bay, Shizuoka Prefecture].
Frequent synonyms: None.
Misidentifications: None.
FAONames: En ? Big-eyed bobtail squid; Fr ? S?piole
? gros yeux; Sp ? Globito ojos grandes.
Diagnostic Features: Body smooth, soft. Males
mature at smaller sizes and do not grow as large as
females. Dorsal anterior margin of mantle triangular,
acute, blunt. Dorsal mantle free from head (not fused
to head). Nuchal cartilage oval . Fins rounded,
semicircular, short, do not exceed length of mantle
anteriorly or posteriorly. Funnel organ strongly
shouldered, with long extensions; ventral elements
oval with acute anterior tips. Male and female arms
subequal in length and long; female arm formula
3:2:1=4. Dorsal and lateroventral arms with low, but
distinct keels. Non-hectocotylized arm sucker
arrangement same in both sexes: arm suckers
biserial, globular. Club straight, slender, or coiled; with
25 to 30 suckers in transverse rows ; all suckers of
similar minute size. Anal flaps well developed. Ink sac
well developed. Epirenal bodies and anal pads
present in both sexes .
Size: Up to 57 mm mantle length.
194 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 276 Austrorossia australis
Known distribution
?? 200 m
Fig. 277 Austrorossia bipapillata
dorsal view
(after Okutani, 1995 )
Geographical Distribution: Western Pacific: East
China Sea, Japan (common on the lower shelf in
Suruga Bay to Tosa Bay), Taiwan Province of China,
Philippines (Fig. 278).
Habitat and Biology: Depth range 240 m in Suruga
Bay, 432 m in the East China Sea. A polytelic species:
ova ripen in small clusters and each individual female
spawns several times, probably throughout the year.
Interest to Fisheries: Of minor interest to fisheries, it is
usually caught as bycatch and is marketed fresh and
frozen.
Remarks: This species differs from Austrorossia
mastigophora (Chun, 1915) in the shape of the funnel
organ, which is not so sharply angled in
A. mastigophora , and in the arm formula
(Rossia mastigophora: 3:4:2:1). Austrorossia
bipapillata differs from R. mollicella Sasaki, 1920 in
possessing anal pads and in the number of
tentacle-club suckers (8 in each transverse row in
R. mollicella). It differs from R. pacifica Berry, 1912 in
having an oval, rather than elongate, parrallel-sided,
nuchal-locking cartilage, and in the number of club
suckers; the club suckers are arranged in 8 to 10 rows,
and anal pads are lacking in R. pacifica.
Literature: Sasaki (1929), Voss (1963), Okutani et al.
(1987), Okutani (1995), Kubodera and Yamada
(1998), Lu (1998b), Reid and Norman (1998).
Cephalopods of the World 195
Fig. 278 Austrorossia bipapillata
Known distribution
?? 200 m
196 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES
FOR WHICH ONLY FEW RECORDS EXIST TO DATE
Rossia brachyura Verrill, 1883
Rossia brachyura Verrill, 1883, Bulletin of the Museum of Comparative Zoology , 11(5): 110 [type locality: Caribbean Sea].
Geographical Distribution: Tropical western Atlantic: Greater and Lesser Antilles.
Literature: Joubin (1902b).
Rossia bullisi Voss, 1956
Rossia bullisi Voss, 1956, Bulletin of Marine Science of the Gulf and Caribbean , 6(2): 101 [type locality: Gulf of Mexico].
Size: Up to 45 mm mantle length.
Geographical Distribution: Tropical western Atlantic: northern Gulf of Mexico and Straits of Florida.
Literature: Roper et al. (1984).
Rossia glaucopis Loven, 1845
Rossia glaucopis Loven, 1845, Ofversigt af Kongl. Vetenskaps-Akademiens Forhandlingar , 2(5): 121 [type locality: Chile].
Geographical Distribution: Southeastern Pacific, Chile.
Literature: Joubin (1902b), Rocha (1997).
Rossia megaptera Verrill, 1881
Rossia megaptera Verrill, 1881, Transactions of the Connecticut Academy of Sciences, 5(6): 349 [type locality: northwest
Atlantic Ocean].
Size: Up to 40 mm mantle length.
Geographical Distribution: Northwestern Atlantic: Davis Strait and western Greenland, Hudson Canyon, off New York.
Depth range from 179 to 1 536 m.
Literature: Mercer (1968), Joubin (1902b), Okutani (1995).
Rossia moelleri Steenstrup, 1856
Rossia moelleri Steenstrup, 1856, Kongelige Danske Videnskabernes Selskabs Skrifter, 5 Raekke, Naturvidenskabelig
og Mathematisk , 4: 198 [type locality: Greenland].
Geographical Distribution: North Atlantic and Arctic Ocean: eastward to the Laptev Sea, westward to Amundsen Bay,
western and northeastern Greenland, northeastern Canada, Labrador, Spitzbergen, Jan Mayen and Kara Seas. Depth
range from 17 to 250 m.
Literature: Mercer (1968), Joubin (1902b), Okutani (1995), Nesis (1999).
Rossia mollicella Sasaki, 1920
Rossia mollicella Sasaki, 1920, Proceedings of the United States National Museum , 57(2310): 189 [type locality: Japan,
Wakayama Prefecture].
Size: Up to approximately 36 mm mantle length.
Geographical Distribution: Western Pacific: Japan, Pacific coast, south from Sendai Bay. Outer shelf and upper bathyal.
Depth range from 729 to 805 m.
Literature: Sasaki (1929), Okutani (1995).
Rossia pacifica diegensis Berry, 1912
Rossia pacifica diegensis Berry, 1912b, Bulletin of the Bureau of Fisheries , 30(1910): 292 [type locality: California].
Geographical Distribution: Eastern Pacific: USA, California, Santa Catalina Basin(?) (see Remarks in R. pacifica
pacifica).
Literature: Mangold et al. (1998).
Rossia palpebrosa Owen, 1834
Rossia palpebrosa Owen, 1834, In J. Ross, Narrative of a second voyage in search of a North West Passage, 1829?1833.
(Volume II, Appendix, Natural History) , xcii [type locality: ?Arctic Regions?].
Size: Up to approximately 45 mm mantle length.
Geographical Distribution: Amphi-North Atlantic: from the Canadian Arctic, Baffin Bay, Greenland south to South
Carolina (32?N ) in the western Atlantic; from Iceland, Spit zbergen, Scotland, Berents Sea and Kara Sea to the North Sea
and off Ireland (51?N) in the eastern Atlantic. Depth range from 75 to 549 m.
Literature: Joubin (1902b), Akimushkin (1963), Aldrich and Lu (1968), Mercer (1968), Boletzky (1970).
Austrorossia enigmatica ( Robson, 1924)
Rossia enigmatica Robson, 1924a, Proceedings of the Zoological Society of London , 1924(2): 635 [type locality: South
Africa].
Size: Types up to 27 mm mantle length.
Geographical Distribution: Southeastern Atlantic: South Africa, Namibia to Cape Province. Depth range from 276 to
400 m.
Literature: Voss (1962b), Roeleveld et al. (1992), Augustyn et al. (1995).
Austrorossia mastigophora (Chun, 1915)
Rossia mastigophora Chun, 1915. Wissebschaftliche Ergebnisse der Deutschen Tiefsee Expedition auf dem Dampfer
?Valdivia? 1898?1899 , 18(2): 405 [type locality: northwestern Indi an Ocean, 0?27'S 42?47'E, near East African coast].
Size: Males up to approximately 31 mm mantle length; females up to 46 mm mantle length.
Geographical Distribution: Western, southern and eastern Africa: from Guinea and Somalia to the Cape of Good Hope.
Chile (doubtful). Depth up to approximately 640 m.
Literature: Voss (1962b), Rocha (1997).
Semirossia patagonica (Smith, 1881)
Rossia patagonica Smith, 1881, Proceedings of the Zoological Society of London , 1881(1): 22 [type locality: Argentina:
Patagonia].
Geographical Distribution: Southwestern Atlantic and southeastern Pacific: southern part of South America from Chile,
Anegada Bay, Tierra del Fuego, Argentina, Falkland Islands.
Literature: Joubin (1902b), Rodhouse et al. (1992), Rocha (1997).
Neorossia leptodons Reid, 1992
Neorossia leptodons Reid, 1992, Bulletin of Marine Science , 49(3): 797 [type locality: South Australia: Great Australian
Bight, 37?18.81'S 138?36.3'E to 37?17. 76'S 138?35.01'E].
Size: Males up to 42 mm mantle length; females up to 77.5 mm mantle length.
Geographical Distribution: Southwestern Pacific: Australia, New South Wales, 32?08'S 153?07'E to South Australia,
33?58'S 131?22'E. Depth r ange from 130 to 1 110 m.
Literature: Reid (1992).
Cephalopods of the World 197
198 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
3.2.3 S u b f a m i l y H ETEROTEUTHIN A E Appell?f, 1898
Heteroteuthis (Heteroteuthis) dispar ( R ?ppell, 1844) F i g . 2 7 9
Sepiola dispar R?ppell, 1844, Giornale del Gabinetto Letterario di Messina , 5(27?28): 133 [type locality: Italy: Sicily].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Odd bobtail squid;
Fr ? S?piole diff?rente; Sp ? Globito
aberrante.
Diagnostic Features: Dorsal mantle free from head (not fused to head). Ventral mantle strongly produced anteriorly, nearly
covers funnel. Fins long; positioned posterior to the dorsal mid-point; anterior edge approximately at level of middle of
ventral side of mantle. Male and female arms differ in relative lengths : arms I and II equal in length and shorter than
arms III and IV. Right arms I and II connected from inner side by muscular band for half their length, depth of web between
first arms in mature males 33 to 50% of arm length. Distal tips of arms I and II in mature females without suckers, tip of
arm II slightly thickened with keel on oral side; maturemale arms III with enlarged suckers: 2 very large suckers (several
times larger than rest) attached at angle
of 90? to each other, followed distally by
three smaller suckers. Tentacles very long
with more than 8 transverse rows of club
suckers. Rounded light organ present
inside mantle cavity on ink sac.
Size: Up to 25 mm mantle length.
Geographical Distribution: Amphi-
Atlantic and Mediterranean Sea: from the
Bermuda Islands, Caribbean Sea, to La
Plata in the western Atlantic; from
southwestern Ireland southward to the
Azores Islands, Madeira Islands, Canary
Islands and Guinea, in the eastern
Atlantic; throughout the Mediterranean
Sea, including Ligurian Sea, northern and
southern Tyrrhenian Sea, Adriatic Sea,
Aegean Sea and Levantine Sea. Also
reported f rom Walters Shoals
(southwestern Indian Ocean) and the
Nazca and Sala y Gomez submarine
ridges (eastern Pacific) (Fig. 280).
Fig. 279 Heteroteuthis (Heteroteuthis) dispar
dorsal view of maleventral view of male
I I I
a r m s o f m a t u r e m a l e
I II
I V
(after Naef, 1923 [197 2 ] )
Fig. 280 Heteroteuthis (Heteroteuthis) dispar
Known distribution
?? 200 m
Fig. 281 Stoloteuthis leucoptera
dorsal viewventral view
lateral view
o r a l v i e w o f a r m s
male female
Habitat and Biology: Mesopelagic, or benthopelagic, but has also been collected on the bottom; depth range to 1 588 m.
Spawning occurs on the bottom, on the slope. The paralarvae live in the mesopelagic and bathypelagic zones, often far
from the coasts, with bottom depths ranging between 1 500 and 3 000 m. Adults live frequently in groups in the lower
epipelagic and mesopelagic zones, most commonly in depths between 200 and 300 m. This is one of the most common
pelagic species in the Mediterranean Sea, where it is often found in areas inhabited by populations of red shrimps.
Heteroteuthis dispar represent a sizeable component of the diet of several predators at the top of the food web, including
dolphins (e.g. Grampus griseus ) sharks (Etmopterux spinax, Galeus melastomus , Scyliorhinus canicula), swordfish
( Xiphias gladius ) and tunas ( Thunnus alalunga ). Populations from the eastern and western Atlantic are probably
isolated from each other.
Interest to Fisheries: Of no commercial interest because of the low quantities available to fisheries, Heteroteuthis dispar
is caught by pelagic nets as well as by trawlers, mainly targeting shrimp, and it is marketed along with other bobtail squids.
Remarks: Another species, Heteroteuthis atlantis, was described by Voss (1955). Nesis (1987) records H. atlantis as a
synonym of H. dispar, but other authors disagree (e.g. Guerra, 1992) and the name H. atlantis is still used in some
publications. Until the taxonomy of the genus is studied in detail, the validity of H. atlantis remains questionable.
Literature: Joubin (1902a), Naef (1923), Nesis (1987), Bello (1990), Guerra (1992), Nesis (1994), Bello (1995, 1996, 1997,
1999), Orsi Relini (1995), Sartor and Belcari (1995), Volpi et al. (1995), Wurtz et al. (1995), Parin et al. (1997), Lefkaditou
et al. (1999).
Stoloteuthis leucoptera (Verrill, 1878 ) F i g . 2 8 1
Sepiola leucoptera Verrill, 1878, American Journal of Science and Arts , (series 3) 16(46): 378 [type locality: USA: Gulf of
Maine].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Leucoptera
bobtail squid; Fr ? S?piole
leucopt?re; Sp ? Globi to
leuc?ptero.
Cephalopods of the World 199
200 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Diagnostic Features: Mantle with marked median bulge dorsally . Dorsal mantle fused to head, junction of mantle and
head broad, 40 to 50% head width ; site of fusion lies considerably more dorsally than middle of eye level. Ventral mantle
projects anteriorly only slightly, to about level of eyes. Ventral mantle broadly flattened into dark shield-like structure , with
median anterior indentation. Fins long; positioned about midway along mantle; fin attachment short, fin length exceeds
attachment length. Mantle-locking cartilage a simple straight ridge; funnel-locking cartilage a simple, straight narrow
depression. Eyes small. Male and female arms short, muscular. All arms except ventral pair united by broad web. First arm
pair in males with thickened, cushion-like lateral membra nes for 2/3 arm length and transverse bundles of finger-like
structures adjacent to these (function unknown). Arm sucker arrangement differs between sexes: in males, suckers
biserial proximally, tetraserial at distal tips; in females, arm suckers biserial. Male arms II with pair (3 rarely) of enlarged
suckers at level of fifth and sixth sucker rows . Club with 12 to 14 suckers in transverse rows. Club with lateral crest at
base. Anal flaps well developed. Rounded light organ on ink sac, with 2 pores medially. Colour: In life brown on dorsal side
of head, arms IV, central mantle and ventral shield, silvery laterally and posteriorly. Blue alongmargin of shield, head and
dorsum. Fins large, almost transparent.
Size: Males up to 17 mm mantle length; females up to 18 mm mantle length.
Geographical Distribution: Amphi-Atlantic and the Mediterranean Sea: from the Gulf of St Lawrence to the Straits of
Florida in the western Atlantic and in the Bay of Biscay in the Eastern Atlantic; Mediterranean Sea, including the northern
and southern Tyrrhenian Sea, Ligurian Sea and Gorgona Island. Also recorded from the Benguela Current off Namibia.
Possibly off eastern Tasmania (Fig. 282).
Habitat and Biology: Lower sublittoral and upper bathyal; depth range from 160 to 700 m. Possible diurnal activity in the
upper mesopelagic.
Interest to Fisheries: Undetermined.
Remarks: Whether all recorded specimens of this widely distributed taxon are conspecific is unknown and needs to be
investigated. A related, possibly identical, form has been found in the upper bathyal near the Kerguelens, Prince Edward
Islands and on the Discovery Bank (southern Indian Ocean). There are only a few records of this species from the
Mediterranean; it may be a recent introduction.
Literature: Joubin (1902a), Nesis (1987), Orsi Relini and Massi (1991), Guerra (1992), Villanueva and S?nchez (1993),
Bello (1995), Volpi et al. (1995), Wurtz et al. (1995).
Fig. 282 Stoloteuthis leucoptera
Known distribution
?? 200 m
?
Sepiolina nipponensis ( Berry, 1911) F i g . 2 8 3
Stoloteuthis nipponensis Berry, 1911b, Zoologischer Anzeiger , 37(2): 39 [type locality: Japan].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Japanese bobtail squid; Fr ? S?piole japonaise; Sp ? Sepiolina japonesa.
Diagnostic Features: Mantle oval, or dome-shaped. Dorsal mantle fused to head; junction of dorsal mantle and head
narrow, approximately 3 mm. Fins wide, ovate, short, lobe approximately 60% mantle lengt h. Non-hectocotylized arm
sucker arrangement same in both sexes: arm suckers biserial; female arm suckers numerous, small throughout; male
suckers enlarged on arms II and III and to a lesser extent on arms IV. Females with higher average sucker counts than
males. Hectocotylus present, both dorsal arms modified: oral surface of modified region swollen, fleshy; with transversely
grooved ridges; suckers in dorsal and ventral series widely spaced, small. Club straight, slender; sucker-bearing face
convex; with 13 to 16 suckers in transverse rows ; all suckers of similar minute size; swimming keel of club extends slightly
proximal to carpus. Rounded light organ inside mantle cavity, on ink sac, visible through mantle in fresh specimens; light
organ secretes a luminous cloud instead of ink. Colour: Body with numerous small chromatophores; ventral pigment
present, dark; ventral mantle margin encircled by silver y iridescent band, approximately 5 mm wide , chromatophores
peppered over band.
Cephalopods of the World 201
Fig. 283 Sepiolina nipponensis
(illustration: K. Hollis/ABRS)
dorsal view
o r a l v i e w o f a r m s o f m a l e
a r m s o f f e m a l e (oral view)
I
I V
I I I
I I
a r m s o f m a l e (oral view)
202 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Size: Up to 25 mm mantle length.
Geographical Distribution: Western Pacific: southern
Japan, Taiwan Province of China, the Philippines and
the Great Australian Bight (southern Australia)
(Fig. 284).
Habitat and Biology: Neritic.
Interest to Fisheries: Undetermined.
Remarks: Based on its distribution, it seems likely that
more than a single species is present. This requires
investigation.
Literature: Roper et al. (1984), Okutani et al. (1987),
Okutani (1995), Lu (1998b), Reid and Norman (1998).
Fig. 284 Sepiolina nipponensis
Known distribution
?? 200 m
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES
FOR WHICH ONLY FEW RECORDS EXIST TO DATE
Heteroteuthis (Heteroteuthis) weberi Joubin, 1902
Heteroteuthis weberi Joubin, 1902a, Bulletin de la Societe scientifique et medicale de l?Ouest , 11: 401 [type locality:
Indonesia].
Geographical Distribution: Indo-Pacific: central Indonesia.
Literature: Joubin (1902c), Nesis (1987), Reid and Norman (1998).
Heteroteuthis (Stephanoteuthis) dagamensis Robson, 1924
Heteroteuthis hawaiiensis var. dagamensis Robson, 1924b, Report of the Fisheries and Marine Biological Survey of the
Union of South Africa , 3: 11 [type locality: South Africa].
Geographical Distribution: Southeastern Atlantic and southwestern Indian Ocean: western, southern and southeastern
Africa.
Literature: Robson (1924b).
Heteroteuthis (Stephanoteuthis) hawaiiensis ( Berry, 1909 )
Stephanoteuthis hawaiiensis Berry, 1909, Proceedings of the United States National Museum , 37(1713): 409 [type
locality: Hawaiian Islands: near Kauai Island].
Size: Up to approximately 30 mm mantle length.
Geographical Distribution: Central and western Pacific: Hawaii, Bonin, Ryukyu Islands, Indonesia, Great Australian
Bight. Possibly southwest Pacific, Banc Combe, 12?14'S 177?28'W, 795 to 820 m.
Literature: Young (1995), Lu and Boucher-Rodoni (2001).
Heteroteuthis (Stephanoteuthis) serventyi Allan, 1945
Heteroteuthis serventyi Allan, 1945, Records of the Australian Museum , 21(6): 340 [type locality: Australia: New South
Wales, Jervis Bay].
Geographical Distribution: Southwestern Pacific: southeastern Australia.
Literature: Allan (1945).
Nectoteuthis pourtalesi Verrill, 1883
Nectoteuthis pourtalesi Verrill, 1883, Bulletin of the Museum of Comparative Zoology , 11(5): 108 [type locality: Barbados
Island].
Geographical Distribution: Tropical western Atlantic: Florida and the Antilles. Bathybenthic.
Literature: Joubin (1902a).
Iridoteuthis iris ( Berry, 1909 )
Stoloteuthis iris Berry, 1909, Proceedings of the United States National Museum , 37(1713): 410 [type locality: Hawaiian
Islands: off south coast of Molokai Island].
Size: Males up to 24 mm mantle length; females up to 28 mm mantle length.
Geographical Distribution: Northern central Pacific: Hawaiian Islands; southeast and northwest Hancock, Colahan and
Kammu seamounts, Ceram Sea (doubtful). Pelagic, found in the open ocean.
Literature: Harman and Seki (1990), Young (1995).
Iridoteuthis maoria Dell, 1959
Iridoteuthis maoria Dell, 1959, Zoology Publications from Vict oria University of Wellington , 25: 410 [type locality: New
Zealand].
Geographical Distribution: Southwestern Pacific: New Zealand, North Island, Cook Strait, Chatham Rise. Nazca and
Sala y Gomez submarine ridges (eastern Pacific).
Literature: Parin et al. (1997).
Cephalopods of the World 203
204 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
3.3 F a m i l y S E P I A D A R I I D A E Fischer, 1882 in 1880 ?188 7 by Amanda Reid
Sepiadariidae Fischer, 1882, Manuel de Conchyliologie et de Pal?ontologi e Conchyliologique ou histoire naturelle des
Mollusques vivants et fossiles , 1369 pages, 23 plates, London.
FAO names: En ? Bottletail squids, bottle squids; Fr ? S?piolettes; Sp ? Sepiolillas.
Diagnostic Features: Small. Dorsal mantle fused to head. Nuchal cartilage absent. Fins narrow, or short; ear-shaped, with
pronounced anterior lobes, or ?earlets?; fin attachment long. Mantle and funnel-locking cartilages absent in Sepiadarium
(mantle and funnel permanently fused) or with 2 components in Sepioloidea. Non-hectocotylized arm sucker arrangement
same in both sexes: biserial, sometimes tetraserial at middle and distal tips of arms. Hectocotylus present, left ventral arm
modified: distal end of hectocotylus without suckers, pedicels modified forming series of transverse lamellae. Tentacle
stalk held in deep sheath or web between bases of arms III and IV, web encircles base of tentacles on outside and inside
forming cutaneous sac. Internal gladius absent. Light organs absent. Dorsal mantle surface covered with large, white
leucophores surrounded by smaller red-brown chromatophores.
Size: Up to 40 mm mantle length.
General Distribution: Found only in the western central Pacific, including Australia and New Zealand.
Habitat and Biology: Benthic. All species bury in the sand during the day and emerge at night to feed. All produce mucous
from the underside of the body when disturbed.
Key to genera in the family Sepiadariidae
1a. Ventral mantle permanently connected to funnel by wide muscular band, funnel-locking apparatus
absent; fins positioned in posterior half of mantle (Fig. 285) . . . . . . . . . . . . . . . . . . . . . Sepiadarium
1b. Ventral mantle not permanently connected to funnel, funnel-locking apparatus present; fins extend
along most of mantle length (Fig. 286) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sepioloidea
Fig. 285 Sepiadarium
dorsal view
Fig. 286 Sepioloidea
dorsal view
Sepiadarium austrinum Berry, 1921 F i g . 2 8 7 ; P l a t e V I I , 4 2 ? 4 4
Sepiadarium austrinum Berry, 1921, Records of the South Australian Museum , 1(4): 354 [type locality: South Australia:
St Vincent Gulf].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Southern bottletail squid; Fr ? S?piolette
du sud; Sp ? Sepiolilla sure?a.
Diagnostic Features: Dorsal mantle fused to head. Mantle
cavity divided by thin septum. Ventral mantle fused to each
side of proximal end of funnel. Fins short, do not exceed
length of mantle anteriorly or posteriorly; attached within
posterior half of mantle. Head broad, as wide as mantle.
Male and female arms differ in relative lengths: arms I and II
longer than III and IV. Non-hectocotylized arm sucker
arrangement same in both sexes: biserial, sometimes
tetraserial at middle and distal tips of arms. Hectocotylus
present, left ventral arm modified: base of hectocotylus with
9 or 10 pairs of normal suckers; suckers replaced by
single series of conical lamellae on distal end of
hectocotylized arm ; arm grooved at distal tip, recurved
dorsally, without protective membrane. Club with 6 suckers
in transverse rows ; club with wide keel, not bound by
distinct folds or continuous membranes. Colour: Body
transparent, yellow or orange, w ith large ovoid
leucophores.
Size: Up to 30 mm mantle length.
Geographical Distribution: Southern Indo-Pacific: southern
Australia (Fig. 288).
Habitat and Biology: This species is found in sandy habitats
in sheltered waters. They bury in the sand during the day,
with only the eyes showing; they emerge at night to feed over
the sand and near seagrass beds. They can be maintained
easily in aquaria, fed on amphipods, isopods and other small
crustaceans. They are able to produce slime from glands on
the ventral surface of the body. Both sexes mate from an
early age, with immature females able to store sperm in a
buccal pouch. Sand-coated eggs are attached to the base of
seaweed or seagrass. The young settle soon after hatching.
Interest to Fisheries: Undetermined. This species may
have a certain economic interest due to its easy
survival in aquaria.
Literature: Lu and Dunning (1998), Norman (2000).
Cephalopods of the World 205
Fig. 287 Sepiadarium austrinum
dorsal view
(illustration: K. Hollis)
Fig. 288 Sepiadarium austrinum
Known distribution
?? 200 m
206 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepiadarium kochii Steenstrup, 1881 F i g . 2 8 9
Sepiadarium kochii Steenstrup, 1881, Danske Videnskabernes Selskabs Skrifter, 6 Raekke, Naturvidenskabelig og
Mathematisk , 1(3): 218 [type locality: Hong Kong, probably Deepwater Bay].
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Koch?s bottletail squid; Fr ? S?piolette de Koch;
Sp ? Sepiolilla de Koch.
Diagnostic Features: Small, mantle dome-shaped. Dorsal mantle fused
to head; length of fused portion approximately 1/4 mantle length. Mantle
cavity divided by thin septum. Ventral mantle fused to each side of
proximal end of funnel. Fins narrow; oblong; short, do not exceed length
of mantle anteriorly or posteriorly; posteriorly with wide gap between
them; attached along posterior half of mantle. Male and female arms
subequal in length; arms I and II longer than II I and IV.
Non-hectocotylized arm sucker arrangement same in both sexes:
biserial basally, tetraserial over distal 2 0 transverse rows .
Hectocotylus present; left
ventral arm modified: base of
hectocotylus suckers normal,
not modified; 18 to 20 fleshy
lamellae over distal 2/3 of
arm, bordered by fold-like
membrane; lamellae thick,
longi tudinal ly grooved
transverse pads. Club with
8 suckers in transverse
rows ; all suckers of similar
minute size; very densely
set. Colour: Dorsal mantle
surface covered with large,
white leucophores sur -
rounded by smaller red-
brown chromatophores .
Size: Up to 30 mm mantle length.
Geographical Distribution: Indo-Pacific: throughout
Indo-Malayan waters from Japan (Tokyo Bay, Pacific
side and Toyama Bay (Japan Sea side), Taiwan
Province of China and the South China Sea to India;
southern Indonesia to New Guinea and northern
Australia. The species has been reported from South
Australia, though this record is unconfirmed (Fig. 290).
Habitat and Biology: Upper sublittoral; depth range to
60 m. Inhabits soft sediments. Remains buried during
the day, then emerges at night to feed, primarily on
small crustacea.
Interest to Fisheries: Undetermined.
Literature: Roper et al. (1984), Okutani et al. (1987),
Okutani (1995), Lu (1998), Lu and Dunning (1998),
Norman and Reid (1998).
Fig. 289 Sepiadarium kochii
dorsal viewo r a l v i e w , m a l e
Fig. 290 Sepiadarium kochii
Known distribution
?? 200 m
?
?
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES
FOR WHICH ONLY FEW RECORDS EXIST TO DATE
Sepiadarium auritum Robson, 1914
Sepiadarium auritum Robson, 1914, Proceedings of the Zoological Society of London , 1814: 678 [type locality: Western
Australia: Monte Bello Islands, Hermite Island].
Size: Type specimen, 11 mm mantle length.
Geographical Distribution: Eastern Indian Ocean: northwestern Australia.
Literature: Robson (1914).
Sepiadarium gracilis Voss, 1962
Sepiadarium gracilis Voss, 1962a, Proceedings of the Biological Society of Washington , 75: 170 [type locality:
Philippines].
Geographical Distribution: Indo-Pacific: South China Sea, Western Philippines.
Literature: Voss (1962a).
Sepiadarium nipponianum Berry, 1932
Sepiadarium nipponianum Berry, 1932, The Philippine Journal of Science , 47(1): 42 [type locality: Japan].
Geographical Distribution: Western Pacific: Japan, southern Honshu, Shikoku, Kyushu.
Literature: Berry (1932).
Sepioloidea lineolata ( Quoy and Gaimard, 1832 ) P l a t e V I I , 4 5 ? 4 6
Sepiola lineolata Quoy and Gaimard 1832, Mollusques. Voyage de decouvertes de l?Astrolabe pendant les annees
1826?1827?1828?1829, Zoologie , 2(1): 82 [type locality: southeastern Australia: Jervis Bay].
Size: Up to 50 mm mantle length.
Geographical Distribution: . Southern Indo-Pacific: eastern, southern and western Australia.
Literature: Norman and Reid (1998), Okutani (1995) Norman (2000).
Sepioloidea pacifica ( Kirk, 1882 )
Sepiola pacifica Kirk, 1882, Transactions and Proceedings of the New Zealand Institute , 14(42): 283 [type locality: New
Zealand].
Geographical Distribution: Southern Pacific: New Zealand (western Pacific) and the Nazca and Sala y Gomez submarine
ridges (eastern Pacific).
Literature: Parin et al. (1997), Lu and Dunning (1998).
Cephalopods of the World 207
208 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
3.4 F a m i l y I D I O S E P I I D A E Appell?f, 1898 F i g . 2 9 1 by Amanda Reid
Idiosepiidae Appell?f, 1898, Cephalopoden von Ternate, 2: Untersuchungen ?ber Idiosepius, Sepiadarium und
verwandte Formen. Ein Beitrag zur Beleuchtung der Hektokotylisation und ihrer systematischen Bedeutung.
Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft , 24(4): 570?637.
FAO names: En ? Pygmy cuttlefishes; Fr ? Seiches pygm?es; Sp ? Sepias pigmeas.
Diagnostic Features: Small animals (less than 25 mm mantle length). Mantle elongate, obovate; posterior mantle margin
bluntly pointed at distal tip. Fins small, kidney-shaped, attached laterally to posterior end of mantle. Dorsal mantle not
fused to head. Nuchal cartilage absent. Glandular oval attachment organ on dorsal posterior end of mantle. Arms short;
arm suckers biserial. Arm suckers enlarged in males. Hectocotylus present, both ventral arms modified: right arm
flattened, wide, with protective membranes, sometimes with transverse ridges and grooves, distal tip of left ventral arm
bilobed; both arms mainly without suckers. Club with 2 to 4 suckers in transverse rows. Gladius absent. In females, right
oviduct non-functional; eggs small, benthic. Development includes pelagic stage, but no metamorphosis occurs. All live
near the coasts in shallow beds of seagrass or algae.
Interest to Fisheries: None.
Literature: Nesis (1987).
Fig. 291 Idiosepiidae
dorsal view
(after Voss, 1963 )
v e n t r a l h e c t o c o t y l i z e d a r m s
( o r a l v i e w )t e n t a c u l a r c l u b
left arm right arm
attachment
organ
SPECIES OF NO CURRENT INTEREST TO FISHERIES, OR RARE SPECIES
FOR WHICH ONLY FEW RECORDS EXIST TO DATE
Idiosepius biserialis Voss, 1962
Idiosepius biserialis Voss, 1962b, Transactions of the Royal Society of South Africa , 36(4): 258 [type locality: South Africa].
Size: Females up to 10.5 mm mantle length.
Geographical Distribution: Southwestern Indian Ocean: southern Africa. Records from Andaman Sea, Thailand probably
refer to I. thailandicus Chotiyaputta et al. (1991). Shallow waters inshore.
Literature: Hylleberg and Nateewathana (1991a), Okutani (1995), Norman and Lu (2000).
Idiosepius macrocheir Voss, 1962
Idiosepius macrocheir Voss, 1962b, Transactions of the Royal Society of South Africa , 36(4): 259 [type locality: South
Africa].
Geographical Distribution: Southwestern Indian Ocean: Southern Africa. Shallow waters inshore.
Literature: Voss (1962b).
Idiosepius minimus (Orbigny, 1835 )
Cranchia minimus Orbigny, 1835 (in F?russac and d?Orbigny, 1834?1848), Histoire Naturelle G?n?rale et Particuli?re
C?phalopodes Ac?tabuliferes Vivants et Fossiles, pl. 1, figs 4?5 [type locality: unreported].
Size: Up to 15 mm mantle length.
Geographical Distribution: ?Coast of Africa? fide (Berry 1932). Shallow, inshore waters.
Literature: Berry (1932).
Idiosepius notoides Berry, 1921
Idiosepius notoides Berry, 1921, Records of the South Australian Museum , 1(4): 361 [type locality: South Australia: Goolwa].
Size: Males up to 15.8 mm mantle length; females up to 25 mm mantle length.
Geographical Distribution: Southwestern Pacific: southern and eastern Australia. Shallow, inshore waters.
Literature: Burn (1959), Norman (2000).
Idiosepius paradoxus (Ortmann, 1888 )
Microteuthis paradoxus Ortmann, 1888, Japanische Cephalopoden. Zoologische Jahrbucher, (Systematik) , 3: 649 [type
locality: Japan: Bay of Tokyo, Kadsiyama].
Size: Up to 16 mm mantle length.
Geographical Distribution: Western Pacific: Japan, southern Hokkaido, Honshu, Kyushu. South Korea, northern
Australia. Shallow, inshore waters.
Literature: Joubin (1902a), Natsukari (1970), Dong (1992), Okutani (1995), Norman and Lu (2000).
Cephalopods of the World 209
210 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Idiosepius picteti ( Joubin, 1894)
Loligo picteti Joubin, 1894, Revue Suisse de Zoologie et Annales du Museum d?Historie Naturelle de Gen?ve , 2: 60 [type
locality: Indonesia: Amboina].
Size: Up to 17 mm mantle length.
Geographical Distribution: Indo-Pacific: eastern Indonesia.
Literature: Joubin (1894).
Idiosepius pygmaeus Steenstrup, 1881
Idiosepius pygmaeus Steenstrup, 1881, Danske Videnskabernes Selskabs Skrifter, 6 Raekke, Naturvidenskabelig og
Mathematisk , 1(3): 219 [type locality: South C hina Sea and 04?20'N 107?20'E].
Size: Up to 20 mm mantle length.
Geographical Distribution: Indo-Pacific: Japan, South China Sea, Philippines, Palau Islands, Indonesia, northern and
northeastern Australia. Shallow, inshore waters.
Literature: Allan (1945), Moynihan (1983b), Jackson (1986, 1989), Yamamoto (1988), Hylleberg and Nateewathana
(1991b), Lewis and Choat (1993), Okutani (1995), Semmens et al. (1995), Pecl and Moltschaniwshyj (1997), Reid and
Norman (1998), Norman and Lu (2000).
Idiosepius thailandicus Chotiyaputta, Okutani and Chaitiamvong, 1991
Idiosepius thailandicus Chotiyaputta et al., 1991, Venus, 50(3): 167 [type locality: Gulf of Thailand].
Geographical Distribution: Indo-Pacific: Thailand.
Literature: Okutani (1995), Nabhitabhata (1998).
3.5 F a m i l y S P I R U L I D A E Owen, 1836 by Amanda Reid
Spirulidae, Owen 1836, Descriptions of some new and rare Cephalopoda, collected by Mr. George Bennet, Corr. Memb.
Z. S. Proceedings of the Zoological Society of London , 37: 19?24
Spirula spirula ( Linnaeus, 1758 ) F i g . 2 9 2
Nautilus spirula Linnaeus, 1758, Systema Naturae per Regna tria Natu rae, Secundum Classes, Ordines, Genera,
Species cum Characteribus, Differentiis, Synonymis, Locis , 710.
Frequent Synonyms: None.
Misidentifications: None.
FAO Names: En ? Ram?s horn squid; Fr ? Spirule; Sp ? Esp?rula.
Diagnostic Features: Mantle cylindrical, thin, muscular; dorsal anterior margin triangular, acute, free from head (not fused
to head). Ventral mantle with 2 tongue-like projections from between which funnel protrudes. Fins narrow, ovate, with
fringed anterior margins; short, do not exceed length of mantle anteriorly or posteriorly; attached dorsolaterally on posterior
end of mantle (almost perpendicular to longitudinal axis of body). Mantle-locking cartilage a simple straight ridge;
funnel-locking cartilage a simple, straight depression. Eyes large; ventral eyelids present. Arms increase in length dorsally
to ventrally, with arms I short, arms IV longest. All arms except fourth pair united by broad webs. Non-hectocotylized arm
sucker arrangement same in both sexes: arm suckers tetraserial, or in 6 rows. Hectocotylus present, both ventral arms
modified: right hectocotylized arm grooved, concave, with spoon-like expansion, pointed tip and 2 finger-like
outgrowths; left hectocotylized arm round in cross-sectio n with 2 spoon-like and one finge r-like outgrowth with soft
papillae at distal tip. Tentacular club straight, slender; not expanded, same width as stalk; with 12 to 16 suckers in
transverse rows; all suckers of similar small size. Radula absent. Buccal membrane in females with spermathecae.
Spirally coiled internal shell present, located in posterior end of animal; shell comprised of over 30 chambers in adults.
Large photophore present between fins, surrounded by annular fold of skin. Colour: Dark reddish brown. Luminescent.
Size: Rarely exceeds 45 mm mantle length.
Cephalopods of the World 211
Fig. 292 Spirula spirula
dorsal view of male
(illustration: K. Hollis/ABRS)
i n t e r n a l s h e l l
Geographical Distribution: Tropical and subtropical oceanic waters worldwide, where the water temperature at 400 m is
10?C or warmer (Fig. 293).
Habitat and Biology: This is a mesopelagic species, inhabiting from 600 to 700 m during the day and found in depths less
than 300 m at night. Its common name is derived from the coiled shell, numbers of which are frequently washed ashore.
The capture of young at depths between 1 000 and 1 750 m suggests that females possibly lay eggs on the bottom of
continental slopes. Eggs are small and the smallest known paralarva has a 1.5 mm mantle length with two shell chambers.
The species attains sexual maturity at about 30 mm mantle length (after 12 to 15 months of life), and the life span is
estimated to be about 18 to 20 months. Spirula spirula is normally covered in a red-brown to silvery skin, which is often lost
in trawled animals. Live animals have been seen to retract their head and arms into their mantle and close the opening with
the pointed ?flaps? formed by the mantle. They take up a vertical position, head downwards, when at rest.
Interest to Fisheries: No direct interest to fishery exists, but the shells collected on the beach are sold in the shell trade.
Literature: Bruun (1943, 1955), Clarke (1970), Nesis (1987), Lu et al. (1992), Reid and Norman (1998).
212 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Fig. 293 Spirula spirula
Known distribution
?? 200 m
4. LIST OF NOMINAL SPECIES
The following list gives information (horizontally) in the order (i) the scientific name as it originally appeared, in alphabetical
order according to the specific name; (ii) the author(s); (iii) date of publication; and (iv) present allocation.
N O M I N A L S P E C I E S P R E S E NT ALL O C ATION
NAUTILIDAE
Nautilus belauensis Saunders, 1981 Nautilus belauensis
Nautilus macromphalus Sowerby, 1849 Nautilus macromphalus
Nautilus perforatus Conrad, 1847 Allonautilus perforatus
Nautilus pompilius Linnaeus (1758) Nautilus pompilius
Nautilus repertus Iredale, 1944 Nautilus repertus
Nautilus scrobiculatus Lightfoot, 1786 Allonautilus scrobiculatus
Nautilus stenomphalus Sowerby, 1849 Nautilus stenomphalus
SEPIIDAE
Acanthosepion whitleyana Iredale, 1926 Sepia whitleyana
Arctosepia limata Iredale, 1926 Sepia limata
Arctosepia rhoda Iredale, 1954 Sepia rhoda
Blandosepia bartletti Iredale, 1954 Sepia bartletti
Blandosepia baxteri Iredale, 1940 Sepia baxteri
Doratosepion trygoninum Rochebrune, 1884 Sepia trygonina
Glyptosepia opipara Iredale, 1926 Sepia opipara
Hemisepius typicus Steenstrup, 1875 Sepia typica
Rhombosepion hieronis Robson, 1924a Sepia hieronis
Rhombosepion robsoni Massy, 1927 Sepia robsoni
Sepia aculeata Van Hasselt, 1835 Sepia aculeata
Sepia acuminata Smith, 1916 Sepia acuminata
Sepia adami Roeleveld, 1972 Sepia adami
Sepia andreana Steenstrup, 1875 Sepia andreana
Sepia angulata Roeleveld, 1972 Sepia angulata
Sepia apama Gray, 1849 Sepia apama
Sepia appellofi W?lker, 1910 Sepia appellofi
Sepia arabica Massy, 1916 Sepia arabica
Sepia aureomaculata Okutani and Horikawa, 1987 Sepia aureomaculata
Sepia australis Quoy and Gaimard, 1832 Sepia australis
Sepia bandensis Adam, 1939b Sepia bandensis
Sepia bathyalis Khromov, Nikitina and Nesis, 1991 Sepia bathyalis
Sepia bertheloti d?Orbigny, 1835 Sepia bertheloti
Sepia bidhaia Reid, 2000 Sepia bidhaia
Sepia braggi Verco, 1907 Sepia braggi
Sepia brevimana Steenstrup, 1875 Sepia brevimana
Sepia burnupi Hoyle, 1904 Sepia burnupi
Sepia carinata Sasaki, 1920 Sepia carinata
Sepia chirotrema Berry, 1918 Sepia chirotrema
Sepia confusa Smith, 1916 Sepia confusa
Sepia cottoni Adam, 1979 Sepia cottoni
Sepia cultrata Hoyle, 1885 Sepia cultrata
Sepia dollfusi Adam, 1941b Sepia dollfusi
Sepia dubia Adam and Rees, 1966 Sepia dubia
Cephalopods of the World 213
Sepia elegans Blainville, 1827 Sepia elegans
Sepia elliptica Hoyle, 1885 Sepia elliptica
Sepia elobyana Adam, 1941a Sepia elobyana
Sepia elongata d?Orbigny, 1839?1842 Sepia elongata
Sepia erostrata Sasaki, 1929 Sepia erostrata
Sepia esculenta Hoyle, 1885 Sepia esculenta
Sepia faurei Roeleveld, 1972 Sepia faurei
Sepia filibrachia Reid and Lu, 2005 Sepia filibrachia
Sepia foliopeza Okutani and Tagawa, 1987 Sepia foliopeza
Sepia gibba Ehrenberg, 1831 Sepia gibba
Sepia grahami Reid, 2001b Sepia grahami
Sepia hedleyi Berry, 1918 Sepia hedleyi
Sepia hierredda Rang, 1835 Sepia hierredda
Sepia incerta Smith, 1916 Sepia incerta
Sepia inermis Van Hasselt, 1835 Sepiella inermis
Sepia insignis Smith, 1916 Sepia insignis
Sepia irvingi Meyer, 1909 Sepia irvingi
Sepia ivanovi Khromov, 1982 Sepia ivanovi
Sepia joubini Massy, 1927 Sepia joubini
Sepia kiensis Hoyle, 1885 Sepia kiensis
Sepia kobiensis Hoyle, 1885 Sepia kobiensis
Sepia koilados Reid, 2000 Sepia koilados
Sepia latimanus Quoy and Gaimard, 1832 Sepia latimanus
Sepia longipes Sasaki, 1913 Sepia longipes
Sepia lorigera W?lker, 1910 Sepia lorigera
Sepia lycidas Gray, 1849 Sepia lycidas
Sepia madokai Adam, 1939a Sepia madokai
Sepia mascarensis Filippova and Khromov, 1991 Sepia mascarensis
Sepia mestus Gray, 1849 Sepia mestus
Sepia mirabilis Khromov, 1988 Sepia mirabilis
Sepia murrayi Adam and Rees, 1966 Sepia murrayi
Sepia novaehollandiae Hoyle, 1909 Sepia novaehollandiae
Sepia officinalis Linn?, 1758 Sepia officinalis
Sepia omani Adam and Rees, 1966 Sepia omani
Sepia orbignyana F?russac in d?Orbigny, 1826 Sepia orbignyana
Sepia ornata Rang, 1837 Sepiella ornata
Sepia papillata Quoy and Gaimard, 1832 Sepia papillata
Sepia papuensis Hoyle, 1885 Sepia papuensis
Sepia pardex Sasaki, 1913 Sepia pardex
Sepia peterseni Appell?f, 1886 Sepia peterseni
Sepia (Metasepia) pfefferi Hoyle, 1885 Metasepia pfefferi
Sepia pharaonis Ehrenberg, 1831 Sepia pharaonis
Sepia plana Lu and Reid, 1997 Sepia plana
Sepia plangon Gray, 1849 Sepia plangon
Sepia plathyconchalis Filippova and Khromov, 1991 Sepia plathyconchalis
Sepia prabahari Neethiselvan and Venkataramani, 2002 Sepia prabahari
Sepia prashadi Winckworth, 1936 Sepia prashadi
Sepia pulchra Roeleveld and Liltved, 1985 Sepia pulchra
214 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia ramani Neethiselvan, 2001 Sepia ramani
Sepia recurvirostra Steenstrup, 1875 Sepia recurvirostra
Sepia reesi Adam, 1979 Sepia reesi
Sepia savignyi Blainville, 1827 Sepia savignyi
Sepia saya Khomov, Nikitina and Nesis 1991 Sepia saya
Sepia senta Lu and Reid, 1997 Sepia senta
Sepia sewelli Adam and Rees, 1966 Sepia sewelli
Sepia simoniana Thiele, 1920 Sepia simoniana
Sepia smithi Hoyle, 1885 Sepia smithi
Sepia sokotriensis Khromov, 1988 Sepia sokotriensis
Sepia stellifera Homenko and Khromov, 1984 Sepia stellifera
Sepia subplana Lu and Boucher-Rodoni, 2001 Sepia subplana
Sepia subtenuipes Okutani and Horikawa, 1987 Sepia subtenuipes
Sepia sulcata Hoyle, 1885 Sepia sulcata
Sepia tala Khromov, Nikitina and Nesis, 1991 Sepia tala
Sepia tanybracheia Reid, 2000 Sepia tanybracheia
Sepia tenuipes Sasaki, 1929 Sepia tenuipes
Sepia thurstoni Adam and Rees, 1966 Sepia thurstoni
Sepia tokioensis Ortmann, 1888 Sepia tokioensis
Sepia tuberculata Lamarck, 1798 Sepia tuberculata
Sepia tullbergi Appell?f, 1886 Metasepia tullbergi
Sepia vercoi Adam, 1979 Sepia vercoi
Sepia vermiculata Quoy and Gaimard, 1832 Sepia vermiculata
Sepia vietnamica Khromov, 1987 Sepia vietnamica
Sepia vossi Khromov, 1987 Sepia vossi
Sepia zanzibarica Pfeffer, 1884 Sepia zanzibarica
Sepiella cyanea Robson, 1924b Sepiella cyanea
Sepiella japonica Sasaki, 1929 Sepiella japonica
Sepiella mangkangunga Reid and Lu, 1998 Sepiella mangkangunga
Sepiella ocellata Pfeffer, 1884 Sepiella ocellata
Sepiella weberi Adam, 1939a Sepiella weberi
Solitosepia plagon adhaesa Iredale, 1926 Sepia plagon
Solitosepia rozella Iredale, 1926 Sepia rozella
Tenuisepia mira Cotton, 1932 Sepia mira
SEPIOLIDAE
Euprymna albatrossae Voss, 1962a Euprymna albatrossae
Euprymna berryi Sasaki, 1929 Euprymna berryi
Euprymna hoylei Adam, 1986 Euprymna hoylei
Euprymna hyllebergi Nateewathana, 1997 Euprymna hyllebergi
Euprymna phenax Voss, 1962a Euprymna phenax
Euprymna scolopes Berry, 1913 Euprymna scolopes
Euprymna stenodactyla Grant, 1833 Euprymna stenodactyla
Fidenas penares Gray, 1849 Euprymna penares
Inioteuthis capensis Voss, 1962 Inioteuthis capensis
Inioteuthis maculosa Goodrich, 1896 Inioteuthis maculosa
Inioteuthis morsei Verrill, 1881 Euprymna morsei
Sepietta minor Naef, 1912a Rondeletiola minor
Sepietta neglecta Naef, 1916 Sepietta neglecta
Cephalopods of the World 215
Sepietta obscura Naef, 1916 Sepietta obscura
Sepiola affinis Naef, 1912b Sepiola affinis
Sepiola atlantica Orbigny, 1839?1842 Sepiola atlantica
Sepiola aurantiaca Jatta, 1896 Sepiola aurantiaca
Sepiola birostrata Sasaki, 1918 Sepiola birostrata
Sepiola intermedia Naef, 1912a Sepiola intermedia
Sepiola japonica Orbigny, 1845 Inioteuthis japonica
Sepiola knudseni Adam, 1984 Sepiola knudseni
Sepiola ligulata Naef, 1912a Sepiola ligulata
Sepiola oweniana Orbigny, 1839?1841 Sepietta oweniana
Sepiola parva Sasaki, 1913 Sepiola parva
Sepiola petersi Steenstrup, 1887 Sepietta petersi
Sepiola pfefferi Grimpe, 1921 Sepiola pfefferi
Sepiola robusta Naef, 1912a Sepiola robusta
Sepiola rondeleti Leach, 1834 Sepiola rondeleti
Sepiola rossiaeformis Pfeffer, 1884 Sepiola rossiaeformis
Sepiola steenstrupiana Levy, 1912 Sepiola steenstrupiana
Sepiola tasmanica Pfeffer, 1884 Euprymna tasmanica
Sepiola trirostrata Voss, 1962a Sepiola trirostrata
Subfamily ROSSIINAE
Heteroteuthis tenera Verrill, 1880 Semirossia tenera
Neorossia leptodons Reid, 1992 Neorossia leptodons
Sepiola macrosoma Delle Chiaie, 1830 Rossia macrosoma
Rossia antillensis Voss, 1955 Austrorossia antillensis
Rossia (Austrorossia) australis Berry, 1918 Austrorossia australis
Rossia bipapillata Sasaki, 1920 Austrorossia bipapillata
Rossia brachyura Verrill, 1883 Rossia brachyura
Rossia bullisi Voss, 1956 Rossia bullisi
Rossia caroli Joubin, 1902b Neorossia caroli
Rossia enigmatica Robson, 1924a Austrorossia enigmatica
Rossia (Semirossia) equalis Voss, 1950 Semirossia equalis
Rossia glaucopis Loven, 1845 Rossia glaucopis
Rossia mastigophora Chun, 1915 Austrorossia mastigophora
Rossia megaptera Verrill, 1881 Rossia megaptera
Rossia moelleri Steenstrup, 1856 Rossia moelleri
Rossia mollicella Sasaki, 1920 Rossia mollicella
Rossia pacifica diegensis Berry, 1912b Rossia pacifica diegensis
Rossia pacifica pacifica Berry, 1911a Rossia pacifica pacifica
Rossia palpebrosa Owen, 1834 Rossia palpebrosa
Rossia patagonica Smith, 1881 Semirossia patagonica
Rossia tortugaensis Voss, 1956 Rossia tortugaensis
Subfamily HETEROTEUTHINAE
Heteroteuthis hawaiiensis var. dagamensis Robson, 1924b Heteroteuthis (Stephanoteuthis) dagamensis
Heteroteuthis serventyi Allan, 1945 Heteroteuthis (Stephanoteuthis) serventyi
Heteroteuthis weberi Joubin, 1902a Heteroteuthis (Heteroteuthis) weberi
Iridoteuthis maoria Dell, 1959 Iridoteuthis maoria
Nectoteuthis pourtalesi Verrill, 1883 Nectoteuthis pourtalesi
Sepiola leucoptera Verrill, 1878 Stoloteuthis leucoptera
216 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepiola dispar R?ppell, 1844 Heteroteuthis (Heteroteuthis) dispar
Stephanoteuthis hawaiiensis Berry, 1909 Heteroteuthis (Stephanoteuthis) hawaiiensis
Stoloteuthis nipponensis Berry, 1911b Sepiolina nipponensis
Stoloteuthis iris Berry, 1909 Iridoteuthis iris
SEPIADARIIDAE
Sepiadarium auritum Robson, 1914 Sepiadarium auritum
Sepiadarium austrinum Berry, 1921 Sepiadarium austrinum
Sepiadarium gracilis Voss, 1962a Sepiadarium gracilis
Sepiadarium kochii Steenstrup, 1881 Sepiadarium kochii
Sepiadarium nipponianum Berry, 1932 Sepiadarium nipponianum
Sepiola lineolata Quoy and Gaimard, 1832 Sepioloidea lineolata
Sepiola pacifica Kirk, 1882 Sepioloidea pacifica
IDIOSEPIIDAE
Cranchia minimus Orbigny, 1835 Idiosepius minimus
Idiosepius biserialis Voss, 1962b Idiosepius biserialis
Idiosepius macrocheir Voss, 1962b Idiosepius macrocheir
Idiosepius notoides Berry, 1921 Idiosepius notoides
Idiosepius pygmaeus Steenstrup, 1881 Idiosepius pygmaeus
Idiosepius thailandicus Chotiyaputta et al., 1991 Idiosepius thailandicus
Loligo picteti Joubin, 1894 Idiosepius picteti
Microteuthis paradoxus Ortmann, 1888 Idiosepius paradoxus
SPIRULIDAE
Nautilus spirula Linnaeus, 1758 Spirula spirula
Cephalopods of the World 217
218 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
5. LIST OF SPECIES BY MAJOR FISHING AREAS
SPECIES
p
a
g
e
GEOGRAPHICAL DISTRIBUTION
MAJOR FISHING AREAS FOR STATISTICAL PURPOSES
18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88
ARC WNA ENA WCA ECA MED WSA EDA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANW
Nautilus macromphalus 52 ?
Nautilus pompilius 53  
Nautilus belauensis 54 
Nautilus repertus 54 
Nautilus stenomphalus 54 
Allonautilus scrobiculatus 55 
Allonautilus perforatus 55 (?) 
Metasepia pfefferi 60  
Metasepia tullbergi 61  
Sepia aculeata 63    
Sepia andreana 65  
Sepia apama 67   
Sepia arabica 69 
Sepia australis 70  
Sepia bandensis 72  
Sepia bertheloti 73  
Sepia braggi 75  
Sepia brevimana 77   
Sepia cultrata 78   
Sepia elegans 79    
Sepia elliptica 81  
Sepia elobyana 82 
Sepia esculenta 84  
Sepia grahami 86  
Sepia hedleyi 87   
Sepia hierredda 88  
Sepia kobiensis 89    
Sepia latimanus 91    
Sepia longipes 93 
Sepia lorigera 94  
Sepia lycidas 95   
Sepia madokai 97  
Sepia murrayi 98 
Sepia officinalis 99   
Sepia omani 101 
Sepia opipara 102   
SPECIES
p
a
g
e
GEOGRAPHICAL DISTRIBUTION
MAJOR FISHING AREAS FOR STATISTICAL PURPOSES
18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88
ARC WNA ENA WCA ECA MED WSA EDA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANW
Sepia orbignyana 103    
Sepia papuensis 105   
Sepia pharaonis 106    
Sepia plangon 109  
Sepia prabahari 110  
Sepia prashadi 111  
Sepia ramani 113  
Sepia recurvirostra 114   
Sepia rozella 116  
Sepia savignyi 117 
Sepia smithi 118  
Sepia stellifera 119   
Sepia sulcata 120  
Sepia trygonina 122 
Sepia vermiculata 124  
Sepia vietnamica 125  
Sepia vossi 127  
Sepia whitleyana 128  
Sepia zanzibarica 129  
Sepiella inermis 130    
Sepiella japonica 132 
Sepiella ornata 134  
Sepiella weberi 135 
Sepia acuminata 137  
Sepia adami 137 
Sepia angulata 137 
Sepia appellofi 137 
Sepia aureomaculata 138 
Sepia bartletti 138 
Sepia bathyalis 138 
Sepia baxteri 138 
Sepia bidhaia 139 
Sepia burnupi 139  
Sepia carinata 139  
Sepia chirotrema 139 
Sepia confusa 140  
Sepia cottoni 140 
Sepia dollfusi 140 
Cephalopods of the World 219
220 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
SPECIES
p
a
g
e
GEOGRAPHICAL DISTRIBUTION
MAJOR FISHING AREAS FOR STATISTICAL PURPOSES
18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88
ARC WNA ENA WCA ECA MED WSA EDA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANW
Sepia dubia 140 
Sepia elongata 141 
Sepia erostrata 141 
Sepia faurei 141 
Sepia foliopeza 141  
Sepia gibba 142 
Sepia hieronis 142  
Sepia incerta 142  
Sepia insignis 142  
Sepia irvingi 143 
Sepia ivanovi 143 
Sepia joubini 143  
Sepia kiensis 143  
Sepia koilados 144 
Sepia limata 144  
Sepia mascarensis 144 
Sepia mestus 144  
Sepia mira 145 
Sepia mirabilis 145 
Sepia novaehollandiae 145  
Sepia papillata 145  
Sepia pardex 146 
Sepia peterseni 146 
Sepia plana 146 
Sepia plathyconchalis 146 
Sepia pulchra 147 
Sepia reesi 147 
Sepia rhoda 147  
Sepia robsoni 147 
Sepia saya 148 
Sepia senta 148 
Sepia sewelli 148 
Sepia simoniana 148  
Sepia sokotriensis 149 
Sepia subplana 149 
Sepia subtenuipes 149 
Sepia tala 149 
Sepia tanybracheia 150 
SPECIES
p
a
g
e
GEOGRAPHICAL DISTRIBUTION
MAJOR FISHING AREAS FOR STATISTICAL PURPOSES
18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88
ARC WNA ENA WCA ECA MED WSA EDA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANW
Sepia tenuipes 150 
Sepia thurstoni 150 
Sepia tokioensis 150 
Sepia tuberculata 151 
Sepia typica 151  
Sepia vercoi 151 
Sepiella cyanea 151  
Sepiella mangkangunga 152  
Sepiella ocellata 152  
Sepia filibrachia 152  
Sepiola affinis 158 
Sepiola atlantica 159   
Sepiola birostrata 161 
Sepiola intermedia 162 
Sepiola ligulata 163 
Sepiola parva 165  
Sepiola robusta 166 
Sepiola rondeleti 167   
Sepiola trirostrata 169 
Euprymna berryi 170 
Euprymna morsei 171   
Euprymna tasmanica 173   
Rondeletiola minor 174    
Sepietta neglecta 176   
Sepietta obscura 177  
Sepietta oweniana 178   
Sepiola aurantiaca 180  
Sepiola knudseni 180  
Sepiola pfefferi 180 
Sepiola rossiaeformis 180 (?) (?)
Sepiola steenstrupiana 180  
Euprymna albatrossae 181 
Euprymna hoylei 181  
Euprymna hyllebergi 181 
Euprymna penares 181 (?) (?)
Euprymna phenax 181 (?) 
Euprymna scolopes 181 
Euprymna stenodactyla 182 
Cephalopods of the World 221
222 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
SPECIES
p
a
g
e
GEOGRAPHICAL DISTRIBUTION
MAJOR FISHING AREAS FOR STATISTICAL PURPOSES
18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88
ARC WNA ENA WCA ECA MED WSA EDA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANW
Sepietta petersi 182  
Inioteuthis capensis 182 
Inioteuthis japonica 182 
Inioteuthis maculosa 182    
Rossia macrosoma 183   
Rossia pacifica pacifica 185   
Rossia tortugaensis 187 
Semirossia equalis 188 
Semirossia tenera 189  
Neorossia caroli 190   
Austrorossia antillensis 192 
Austrorossia australis 193   
Austrorossia bipapillata 194  
Rossia brachyura 196 
Rossia bullisi 196 
Rossia glaucopis 196 
Rossia megaptera 196 
Rossia moelleri 196   
Rossia mollicella 196 
Rossia pacifica diegensis 197 
Rossia palpebrosa 197  
Austrorossia enigmatica 197 
Austrorossia
mastigophora 197   
Semirossia patagonica 197  
Neorossia leptodons 197  
Heteroteuthis
(Heteroteuthis) dispar 198     
Stoloteuthis leucoptera 199        (?)
Sepiolina nipponensis 201   
Heteroteuthis
(Heteroteuthis) weberi 203  
Heteroteuthis
(Stephanoteuthis)
dagamensis
203  
Heteroteuthis
(Stephanoteuthis)
hawaiiensis
203   
Heteroteuthis
(Stephanoteuthis)
serventyi
203 
Nectoteuthis pourtalesi 203 
Iridoteuthis iris 203  
Iridoteuthis maoria 203  
SPECIES
p
a
g
e
GEOGRAPHICAL DISTRIBUTION
MAJOR FISHING AREAS FOR STATISTICAL PURPOSES
18 21 27 31 34 37 41 47 48 51 57 58 61 67 71 77 81 87 88
ARC WNA ENA WCA ECA MED WSA EDA ANC WIO EIO ANE WNP ENP WCP ECP WSP ESP ANW
Sepiadarium austrinum 205 
Sepiadarium kochii 206   
Sepiadarium auritum 207 
Sepiadarium gracilis 207 
Sepiadarium nipponianum 207 
Sepioloidea lineolata 207  
Sepioloidea pacifica 207  
Idiosepius biserialis 209 
Idiosepius macrocheir 209 
Idiosepius minimus 209 (?) (?) (?) (?)
Idiosepius notoides 209 
Idiosepius paradoxus 209  
Idiosepius picteti 210 
Idiosepius pygmaeus 210  
Idiosepius thailandicus 210  
Spirula spirula 211          
Cephalopods of the World 223
224 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
MAJOR FISHING AREAS FO R STATISTICAL PURPOSES
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LINKS AT PUBLICATION DATE:
Cephalopod International Advisory Council (CIAC) : http://www.nbs.ac.uk/public/mlsd/ciac/index.html
Coleoid Cephalopods through Time, International Symposium:
http://www.userpage.fu-berlin.de/%7epalaeont/palbio3/palbio3.htm
FAO catalogue Cephalopods of the World: ftp://ftp.fao.org/fi/document/sidp/ac479e_03cephalopods/ac479e00.pdf
FAO Statistical Database: http://faostat.fao.org/faostat/default.jsp?version=int&hasbulk=1
GlobeFish (Cephalopods): http://www.globefish.org/dynamisk.php4?id=2394
Integrated Taxonomic Information System : http://www.itis.usda.gov/index.html
National Museum of Natural History (NMFS) : http://www.mnh.si.edu/cephs/
National Resource Center for Cephalopods: http://www.nrcc.utmb.edu/
Review of the state of world marine fishery resources, World squid resources :
ftp://ftp.fao.org/docrep/fao/007/y5852e/Y5852E19.pdf
Sea Around Us Project : http://www.seaaroundus.org/project.htm
SeaMounts Online : http://seamounts.sdsc.edu/
Smithsonian Institution research Information System (SIRIS): http://siris-bibliographies.si.edu/#focus
Species 2000 : http://www.sp2000.org/
Squid Atlas, Geographical Information System (GIS) : http://www.nerc-bas.ac.uk/public/mlsd/squid-atlas/
SquidFish : http://www.squidfish.net/
The Cephalopod Page: http://is.dal.ca/~ceph/TCP/index.html
The Octopus News Magazine Online (TONMO) : http://www.tonmo.com/
Tree of Life (Cephalopods): http://tolweb.org/tree?group=Cephalopoda&contgroup=Mollusca
Cephalopods of the World 247
7. INDEX OF SCIENTIFIC AND VERNACULAR NAMES
Explanation of the System
Italics : Valid scientific names (double entry by genera and species)
Italics : Synonyms, misidentifications and subspecies (double entry by genera and species)
ROMAN : Family names
ROMAN : Scientific names of divisions, classes, subclasses, orders, suborders and subfamilies
Roman : FAO names
Roman : Local names
Cephalopods of the World 249
A
Acanthosepion pageorum . . . . . . . . . . . . . . . . . . . . . 118
Acanthosepion whitleyana . . . . . . . . . . . . . . . . . . . . 128
aculeata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 ?64
acuminata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 97, 137
adami, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
adhaesa, Solitosepia plangon . . . . . . . . . . . . . . . . . . 109
affinis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
affinis, Sepiola . . . . . . . . . . . . . . . . . . . . . . . 158 ?159, 177
African cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Ajia-kouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 115
albatrossae, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . 181
albatrossi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Allonautilus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 6, 51
Allonautilus perforatus . . . . . . . . . . . . . . . . . . . . . . 51, 55
Allonautilus scrobiculatus . . . . . . . . . . . . . . . . 51, 53, 55
Aluda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Ami-mom-nkouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
AMMONOIDEA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Amplisepia parysatis . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Amplisepia verreaux . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Analogous bobtail squid . . . . . . . . . . . . . . . . . . . . . . . 158
Andrea cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
andreana, Sepia . . . . . . . . . . . . . . . . . . . . . . 65 ?66, 89?90
andreanoides, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 89
angulata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
antillensis, Austrorossia . . . . . . . . . . . . . . . . . . . 192 ?193
antillensis, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Antilles bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . 192
apama, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 ?68
appellofi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Arabian cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
arabica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Architeuthis spp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Arctosepia limata. . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Arctosepia rhoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
argentinus, Illex . . . . . . . . . . . . . . . . . . . . . . . . . . . 10, 17
Argonauta spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . 7?9 , 31
Argonautid octopods . . . . . . . . . . . . . . . . . . . . . . . . 31, 36
Argonauts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Atlantic bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . 159
atlantica, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . 159 ?160
atlantis, Heteroteuthis . . . . . . . . . . . . . . . . . . . . . . . . 199
aurantiaca, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
aureomaculata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 138
auritum, Sepiadarium . . . . . . . . . . . . . . . . . . . . . . . . . 207
australis, Austrorossia . . . . . . . . . . . . . . . . . . . . . . . . . 193
australis, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
australis, Rossia (Austrorossia) . . . . . . . . . . . . . . . . 193
australis, Sepia . . . . . . . . . . . . . . . . . . 58, 70?71 , 80, 121
austrinum, Sepiadarium . . . . . . . . . . . . . . . . . . . . . . . 205
Austrorossia antillensis . . . . . . . . . . . . . . . . . . . . 192 ?193
Austrorossia australis . . . . . . . . . . . . . . . . . . . . . . . . . 193
Austrorossia bipapillata . . . . . . . . . . . . . . . . . . . 194 ?195
Austrorossia enigmatica . . . . . . . . . . . . . . . . . . . . . . . 197
Austrorossia mastigophora . . . . . . . . . . . . . 194?195, 197
B
Babbunedda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
bandensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 72 , 138
bartletti, Blandosepia . . . . . . . . . . . . . . . . . . . . . . . . 138
bartletti, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72, 138
bartramii, Ommastrephes . . . . . . . . . . . . . . . . . . . . . . . 18
bathyalis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Bathypolypus sponsalis . . . . . . . . . . . . . . . . . . . . . . . . 191
Bathyteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
baxteri, Blandosepia . . . . . . . . . . . . . . . . . . . . . . . . . 138
baxteri, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72, 138
belauensis, Nautilus . . . . . . . . . . . . . . . . . . . . . 51, 53?54
BELEMNOIDEA. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Belemnoids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Bellybutton nautilus . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
beppuana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
berryi, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . 170 ?172
bertheloti, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 73 , 79?80
bidhaia, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75, 139
Big bottom bobtail squid . . . . . . . . . . . . . . . . . . . . . . 193
Big-eyed bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . 194
bipapillata, Austrorossia . . . . . . . . . . . . . . . . . . . 194 ?195
bipapillata, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
birostrata, Sepiola . . . . . . . . . . . . 161 ?162, 165?166, 170
biserialis, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . . . 209
biserialis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Blandosepia bartletti . . . . . . . . . . . . . . . . . . . . . . . . . 138
Blandosepia baxteri . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Blue-ringed octopus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Bobtail squids . . . . . . . . . . . . . . . . . . 3, 9, 21?22, 25, 153
BOLITAENIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Bottle squids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Bottletail squids . . . . . . . . . . . . . . . . . . . 3, 9, 22, 25, 204
Bouzuika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 186
brachyura, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
braggi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 ?76
brevimana, Sepia . . . . . . . . . . . . . . . . . . . . . . 77 , 119?120
brevis, Lolliguncula . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Broadback cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Broadclub cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
bullisi, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . 193, 196
burnupi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . 98, 123, 139
Butterfly bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . 161
C
Calamarcito . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Cape e chiuove . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
capensis, Inioteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . 182
capensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Capo di chiodo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Cappuccetto. . . . . . . . . . . . . 159, 163, 167?168, 175, 179
carinata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Carol bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
caroli caroli, Neorossia . . . . . . . . . . . . . . . . . . . . . . 191
caroli jeannae, Neorossia . . . . . . . . . . . . . . . . . . . . . 191
caroli, Neorossia . . . . . . . . . . . . . . . . . . . . . . . . . 190 ?191
250 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
caroli, Neorossia caroli . . . . . . . . . . . . . . . . . . . . . . 191
caroli, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Casseron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Castagnola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Casta?o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
CEPHALOPODA . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 6, 29
Chakod . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Chambered nautiluses . . . . . . . . . . . . . . . . . . . . . . . 9, 51
Checo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Chibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Chinna vari kanavai . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Chiroteuthid squids . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
CHIROTEUTHIDAE. . . . . . . . . . . . . . . . . . . . . . 25, 29, 35
Chiroteuthids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
chirotrema, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Choco . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101, 184
Chocos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Chopito . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104, 184
Choquitos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Choubai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Choubei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Chouebi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Choueb?. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Chtenopteryx. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Chubei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Cirrate octopods. . . . . . . . . 3, 8, 21?22, 26, 28, 31, 33, 36
CIRROTEUTHIDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Cirroteuthids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Cistopus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Coca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
coindetii, Illex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
COLEOIDEA . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 7, 21, 33
Coleoids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6?7
Common bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . 178
Common cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
confusa, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Corsica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
cottoni, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75, 140
Cranchia minimus . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Cranchiid squids . . . . . . . . . . . . . . . . . . . . . . . . . . . 22, 31
crassa, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Crumenasepia hulliana . . . . . . . . . . . . . . . . . . . . . . . 106
Crumenasepia ursulae . . . . . . . . . . . . . . . . . . . . . . . 106
cultrata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Curvespine cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . 114
Cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29, 101
Cuttlefishes . 3, 6, 8?10, 17, 22, 25, 33, 35?36, 56?57 ?58
cyanea, Sepiella . . . . . . . . . . . . . . . . . . . . . . . . . . 131, 151
D
dagamensis, Heteroteuthis (Stephanoteuthis) . . . . . . . 203
dagamensis, Heteroteuthis hawaiiensis var. . . . . . . 203
dannevigi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
DECAPODIFORMES. . . . . . . . . 21?23, 25, 27, 29, 31, 33
Decapods. . . . . . . . . . . . . . . . . . . . . . 7, 24, 29?30, 35?36
Decorisepia rex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
diegensis, Rossia pacifica . . . . . . . . . . . . . . . . . . 186, 197
dispar, Heteroteuthis . . . . . . . . . . . . . . . . . . . . . . . . . 199
dispar, Heteroteuthis (Heteroteuthis) . . . . . . . . . . . . . 198
dispar, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
dofleini, Enteroctopus . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
dollfusi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Doratosepion trygoninum . . . . . . . . . . . . . . . . . . . . . 122
Dosidicus gigas . . . . . . . . . . . . . . . . . . . . . . . . . . 9?10, 18
dubia, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Dwarf bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
Dyonon refui . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
E
eclogaria, Ponderisepia . . . . . . . . . . . . . . . . . . . . . . . 91
Eledone spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
elegans, Sepia . . . . . . . . . . . . . . . . . . . . . 58, 79 , 101, 104
Elegant bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . 176
Elegant cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
elliptica, Sepia . . . . . . . . . . . . . . . . . . . . . . . 81 ?82, 84?85
elobyana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
elongata, Sepia . . . . . . . . . . . . . . . . . . . . . . . 122-123, 141
Emperor nautilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
enigmatica, Austrorossia . . . . . . . . . . . . . . . . . . . . . . . 197
enigmatica, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Enteroctopus dofleini . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
equalis, Rossia (Semirossia) . . . . . . . . . . . . . . . . . . . 188
equalis, Semirossia . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
erostrata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Esp?rula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
esculenta, Sepia . . . . . . . . . 77?78, 81?82, 84 , 96?97, 108
Euprymna albatrossae . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna berryi . . . . . . . . . . . . . . . . . . . . . . . . . 170 ?172
Euprymna hoylei. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna hyllebergi. . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna morsei . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Euprymna penares . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna phenax. . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna scolopes . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
Euprymna stenodactyla. . . . . . . . . . . . . . . . . . . . . . . . 182
Euprymna tasmanica . . . . . . . . . . . . . . . . . . . . . . . . . 173
F
Fa gai na . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
faurei, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Fidenas penares . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
filippovae, Todarodes . . . . . . . . . . . . . . . . . . . . . . . . . . 18
filibrachia, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
filliouxi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
filliouxi, Sepia officinalis . . . . . . . . . . . . . . . . . . . . . 101
Flamboyant cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . 60
foliopeza, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Foo baan woo chack . . . . . . . . . . . . . . . . . . . . . . . . . . 108
formosana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Frog cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
G
galei, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Gam woo chak. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Cephalopods of the World 251
Gemeiner Tintenfisch. . . . . . . . . . . . . . . . . . . . . . . . . . 101
genista, Solitosepia . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Gewone Inktvis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Giant African cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . 88
Giant Australian cuttlefish . . . . . . . . . . . . . . . . . . . 67 ?68
gibba, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
gigas, Dosidicus . . . . . . . . . . . . . . . . . . . . . . . . . 9?10, 18
glaucopis, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Globito . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Globito aberrante . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Globito antillano . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Globito austral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Globito colibri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
Globito de Carol . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Globito de Tasmania . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Globito de Tortugas . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Globito del Pac?fico boreal . . . . . . . . . . . . . . . . . . . . . 185
Globito leuc?ptero . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
Globito mimika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
Globito ojos grandes . . . . . . . . . . . . . . . . . . . . . . . . . 194
Globito peque?o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
Globito reluciente . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Globito robusto. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Globito tierno . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Globitos. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Glyptosepia opipara . . . . . . . . . . . . . . . . . . . . . . . . . 102
Golden cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
GONATIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 22
gracilis, Sepiadarium . . . . . . . . . . . . . . . . . . . . . . . . . 207
grahami, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Grande seiche ray?e . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Greater shining bobtail squid . . . . . . . . . . . . . . . . . . . 188
Grooved cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Guinean cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
H
Hana-ika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Hapalochlaena spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Hariika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Hari-ika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Harinashi-kouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
harmeri, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
hawaiiensis var. dagamensis, Heteroteuthis . . . . . . 203
hawaiiensis, Heteroteuthis (Stephanoteuthis) . . . . . . . 203
hawaiiensis, Stephanoteuthis . . . . . . . . . . . . . . . . . . 203
Hedley?s cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
hedleyi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 87 , 97, 127
Hemisepius typicus . . . . . . . . . . . . . . . . . . . . . . . . . . 151
hercules, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91?92
HETEROTEUTHINAE . . . . . . . . . . . . . . . . . . . . . . . . . 198
Heteroteuthis (Heteroteuthis) dispar . . . . . . . . . . . . . . 198
Heteroteuthis (Heteroteuthis) weberi . . . . . . . . . . . . . 203
Heteroteuthis (Stephanoteuthis) dagamensis . . . . . . . 203
Heteroteuthis (Stepha noteuthis) hawaiiensis . . . . . . . 203
Heteroteuthis (Stephanoteuthis) serventyi . . . . . . . . . 203
Heteroteuthis atlantis . . . . . . . . . . . . . . . . . . . . . . . . 199
Heteroteuthis dispar . . . . . . . . . . . . . . . . . . . . . . . . . 199
Heteroteuthis hawaiiensis var. dagamensis . . . . . . . 203
Heteroteuthis serventyi . . . . . . . . . . . . . . . . . . . . . . . 203
Heteroteuthis tenera . . . . . . . . . . . . . . . . . . . . . . . . . 189
Heteroteuthis weberi . . . . . . . . . . . . . . . . . . . . . . . . . 203
hieronis, Rhombosepion . . . . . . . . . . . . . . . . . . . . . . 142
hieronis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58, 142
hierredda, Sepia . . . . . . . . . . . . . . . . . . . . 74, 88 ?89, 101
hierredda, Sepia officinalis . . . . . . . . . . . . . . . . . . . . 101
Hime-kouika. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
HISTIOTEUTHIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Hooded cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
hoylei, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
hoylei, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
hulliana, Crumenasepia . . . . . . . . . . . . . . . . . . . . . . 106
Humming-bird bobtail squid . . . . . . . . . . . . . . . . . . . . 170
hyadesi, Martialia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
hyllebergi, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . 181
I
IDIOSEPIIDAE. . . . . . . . . . . . . . . . . . . . . . . . . 33, 56, 208
Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Idiosepius biserialis . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius macrocheir . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius minimus . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius notoides . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius paradoxus . . . . . . . . . . . . . . . . . . . . . . . . . 209
Idiosepius picteti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Idiosepius pygmaeus . . . . . . . . . . . . . . . . . . . . . . . . 9, 210
Idiosepius thailandicus . . . . . . . . . . . . . . . . . . . . 209?210
illecebrosus, Illex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Illex argentinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10, 17
Illex coindetii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Illex illecebrosus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
incerta, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Incirrate octopods. . . 3, 7?8, 21?22, 24, 28?29, 31, 35?36
indica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
inermis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
inermis, Sepiella . . . . . . . . . . . . . . . . . . . . . 130 ?131, 133
Inioteuthis capensis. . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Inioteuthis japonica . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Inioteuthis maculosa . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Inioteuthis morsei . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
insignis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
intermedia, Sepiola . . . . . . . . . . . . . . . . . . . 159,162 ?163
Intermediate bobtail squid . . . . . . . . . . . . . . . . . . . . . 162
Iridoteuthis iris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Iridoteuthis maoria . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
iris, Iridoteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
iris, Stoloteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
irvingi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
italica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
ivanovi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
J
Jam mak yue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85, 115
Jam Mak Yue. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Japanese bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . 201
252 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Cephalopods of the World 253
Japanese spineless cuttlefish . . . . . . . . . . . . . . . . . . 132
japonica, Inioteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . 182
japonica, Sepiella . . . . . . . . . . . . . . . . . . . . . . . . 132 ?133
japonica, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
jeannae, Neorossia caroli . . . . . . . . . . . . . . . . . . . . . 191
Jibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Jibia africana. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
J?bias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Jibi?n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
joubini, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
K
Kaminari-ika. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Ken?s cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
kiensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Kisslip cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Knifebone cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Knobby bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . 169
knudseni, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Kobi cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
kobiensis, Sepia . . . . . . . . . . . . . . . . . . . . . . 65?66, 89 , 98
Kobushime. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
Koch?s bottletail squid . . . . . . . . . . . . . . . . . . . . . . . . 206
kochii, Sepiadarium . . . . . . . . . . . . . . . . . . . . . . . . . . 206
koilados, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Kora katitza . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Kouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Kouika-modoki. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Kubushime. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
L
lana, Solitosepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Large striped cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . 113
latimanus, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . 58, 91
Lentil bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
leptodons, Neorossia . . . . . . . . . . . . . . . . . . . . . . 191, 197
Lesser shining bobtail squid . . . . . . . . . . . . . . . . . . . 189
Leucoptera bobtail squid . . . . . . . . . . . . . . . . . . . . . . 199
leucoptera, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . 199
leucoptera, Stoloteuthis . . . . . . . . . . . . . . . . . . . . . . . . 199
Leung yee jai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
ligulata, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . 163 ?164
limata, Arctosepia . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
limata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 75?76, 144
lineolata, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
lineolata, Sepioloidea . . . . . . . . . . . . . . . . . . . . . . . . . 207
Loliginid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
LOLIGINIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Loliginids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17, 34
Loligo picteti . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Loligo spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Loligo vulgaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7, 9
Lolliguncula brevis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Longarm cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
longipes, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
lorigera, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 , 126
Luda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
lycidas, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
M
macrocheir, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . 209
macromphalus, Nautilus . . . . . . . . . . . . . . . . . . 51?52 ?53
macrosoma, Rossia . . . . . . . . . . . . . . . 176, 183 ?184, 191
macrosoma, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . 183
maculosa, Inioteuthis . . . . . . . . . . . . . . . . . . . . . . . . . 182
madokai, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81, 97
Madokai?s cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Magnificent cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . 102
Maika. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
maindroni, Sepiella . . . . . . . . . . . . . . . . . . . . . . 130, 133
Mak dau. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Mak gung. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Mak mo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
mangkangunga, Sepiella . . . . . . . . . . . . . . . . . . . . . . . 152
maoria, Iridoteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Margade . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Martialia hyadesi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
mascarensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
mastigophora, Austrorossia . . . . . . . . . . . . 194?195, 197
mastigophora, Rossia . . . . . . . . . . . . . . . . . . . . 195, 197
Mastigoteuthids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
mediterranea, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 99
mediterranea, Sepia officinalis . . . . . . . . . . . . . . . . . 101
megaptera, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
mercatoris, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
mestus, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 144
Metasepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Metasepia pfefferi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Metasepia tullbergi . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
microcheirus, Sepia (Sepiella) . . . . . . . . . . . . . . . . . 130
Microteuthis paradoxus . . . . . . . . . . . . . . . . . . . . . . 209
Mimika bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . 171
minimus, Cranchia . . . . . . . . . . . . . . . . . . . . . . . . . . 209
minimus, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . . . 209
minor, Rondeletiola . . . . . . . . . . . . . . . 164, 174 ?175, 191
minor, Sepietta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
mira, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
mira, Tenuisepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
mirabilis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
misakiensis, Sepia (Doratosepion) . . . . . . . . . . . . . . 150
Mo Jam Woo Chak . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
moelleri, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . 186, 196
mollicella, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . 195?196
MOLLUSCA. . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 6, 23, 29
Mongo-ika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Mongouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96, 108
morsei, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
morsei, Inioteuthis . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
mozambica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
murrayi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Mustekala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Myopsid squids . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 25, 32
MYOPSIDA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29, 31
Myopsids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Mysterious bobtail squid . . . . . . . . . . . . . . . . . . . . . . 177
N
Nautile bouton . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Nautile flamm? . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Nautiles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
NAUTILIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Nautilo com?n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 53
Nautilo ombligo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
NAUTILOIDEA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
Nautiloids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Nautilos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Nautilus. . . . . . . . . . . . . . . . . . . . 3, 6, 8?9, 31, 51, 53?55
Nautilus belauensis . . . . . . . . . . . . . . . . . . . . . . 51, 53?54
Nautilus macromphalus . . . . . . . . . . . . . . . . . . 51?52 ?53
Nautilus perforatus . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Nautilus pompilius . . . . . . . . . . . . . . . . . . . . . . 51, 53 ?54
Nautilus repertus. . . . . . . . . . . . . . . . . . . . . . . . 51, 53?54
Nautilus scrobiculatus. . . . . . . . . . . . . . . . . . . . . . . . . 55
Nautilus spirula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Nautilus spp.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Nautilus stenomphalus . . . . . . . . . . . . . . . . . . . 51, 53?54
Nautiluses . . . . . . . . . . . . . 3, 7, 22?23, 25, 28, 30, 33, 50
Nectoteuthis pourtalesi . . . . . . . . . . . . . . . . . . . . . . . . 203
Needle cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
neglecta, Sepietta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
Neorossia caroli . . . . . . . . . . . . . . . . . . . . . . . . . 190 ?191
Neorossia caroli caroli . . . . . . . . . . . . . . . . . . . . . . . 191
Neorossia caroli jeannae . . . . . . . . . . . . . . . . . . . . . 191
Neorossia leptodons . . . . . . . . . . . . . . . . . . . . . . . . 1, 197
Neritic squid. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Ngor Huet Mak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
nipponensis, Sepiolina . . . . . . . . . . . . . . . . . . . . . . . . 201
nipponensis, Stoloteuthis . . . . . . . . . . . . . . . . . . . . . 201
nipponianum, Sepiadarium . . . . . . . . . . . . . . . . . . . . 207
Niyori-mimi-ika. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
North Pacific bobtail squid . . . . . . . . . . . . . . . . . . . . . 185
notoides, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Nototodarus philippinensis . . . . . . . . . . . . . . . . . . . . . . 18
novaehollandiae, Sepia . . . . . . . . . . . . . . . . . . . . . . . . 145
O
obscura, Sepietta . . . . . . . . . . . . . . . . . . . . . . . . . 177 ?178
occidua, Solitosepia . . . . . . . . . . . . . . . . . . . . . . . . . 105
Oceanic squids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
ocellata, Sepiella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
OCTOPODIDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 29
OCTOPODIFORMES . . . . . . . . . . . . . . . . . 22?23, 27, 31
Octopods . . . . . . . . . . . . . . . . . 3, 9, 23, 25, 29, 31, 34?36
Octopus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Octopus sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Octopus spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Octopus vulgaris . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9?10
Octopuses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7, 36
Ocythoe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31, 35
Odd bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Oegopsid squids . . . . . . . . . . . . . . . . 3, 21, 25, 27?29, 32
OEGOPSIDA. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29, 31
Oegopsids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
officinalis filliouxi, Sepia . . . . . . . . . . . . . . . . . . . . . 101
officinalis hierredda, Sepia . . . . . . . . . . . . . . . . . . . . 101
officinalis mediterranea, Sepia . . . . . . . . . . . . . . . . . 101
officinalis vermiculata, Sepia . . . . . . . . . . . . . . . . . . 101
officinalis, Sepia . . . 7, 9, 58, 74, 80, 88?89, 99 ?100, 104
Oman cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
omani, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 98, 101 , 127
Ommastrephes bartramii . . . . . . . . . . . . . . . . . . . . . . . 18
Ommastrephid squid . . . . . . . . . . . . . . . . . . . . . . . . . 9, 18
OMMASTREPHIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Ommastrephids . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17?18
opipara, Glyptosepia . . . . . . . . . . . . . . . . . . . . . . . . . 102
opipara, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Opisthoteuthis spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
orbignyana, Sepia . . . . . . . . . . . . . . . . . . 58, 80, 101, 103
ornata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
ornata, Sepiella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Ornate cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
oualaniensis, Sthenoteuthis . . . . . . . . . . . . . . . . . . . 9, 18
Ovalbone cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
oweniana, Sepietta . . . . . . . . . . . . 164, 176?178 ?179, 191
oweniana, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
P
pacifica diegensis, Rossia . . . . . . . . . . . . . . . . . . 186, 197
pacifica pacifica, Rossia . . . . . . . . . . . . . . . 185 ?186, 197
pacifica, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . 186, 195
pacifica, Rossia pacifica . . . . . . . . . . . . . . . 185 ?186, 197
pacifica, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
pacifica, Sepioloidea . . . . . . . . . . . . . . . . . . . . . . . . . . 207
pacificus, Todarodes . . . . . . . . . . . . . . . . . . . . . . . . 10, 18
pageorum, Acanthosepion . . . . . . . . . . . . . . . . . . . . . 118
Paintpot cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
palmata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
palpebrosa, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Paper nautilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
papillata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Papuan cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
papuensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 105 ?106
paradoxus, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . . 209
paradoxus, Microteuthis . . . . . . . . . . . . . . . . . . . . . . 209
pardalis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
pardex, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
parva, Sepiola . . . . . . . . . . . . . . . . . . . . . . . 162, 165 ?166
parysatis, Amplisepia . . . . . . . . . . . . . . . . . . . . . . . . . 67
patagonica, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . 197
patagonica, Semirossia . . . . . . . . . . . . . . . . . . . . . . . . 197
Patchwork cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . 124
penares, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
penares, Fidenas . . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
perforatus, Allonautilus . . . . . . . . . . . . . . . . . . . . . 51, 55
perforatus, Nautilus . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Periya vari kanavai. . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
peterseni, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
petersi, Sepietta . . . . . . . . . . . . . . . . . . . . . . . . . . 178, 182
petersi, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
254 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Petite seiche ray?e . . . . . . . . . . . . . . . . . . . . . . . . . . . 110
pfefferi, Metasepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
pfefferi, Sepia (Metasepia) . . . . . . . . . . . . . . . . . . . . . 60
pfefferi, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Pharaoh cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
pharaonis, Sepia . . . . . . . . . . . 58, 92, 106 ?108, 114, 147
phenax, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . . . 181
philippinensis, Nototodarus . . . . . . . . . . . . . . . . . . . . . 18
PHOLIDOTEUTHIDAE. . . . . . . . . . . . . . . . . . . . . . . . . . 18
picteti, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
picteti, Loligo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 210
Pink cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
plana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
plangon adhaesa, Solitosepia . . . . . . . . . . . . . . . . . . 109
plangon, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
plathyconchalis, Sepia . . . . . . . . . . . . . . . . . . . . . 117, 146
pompilius, Nautilus . . . . . . . . . . . . . . . . . . . . . . 51, 53 ?54
Ponderisepia eclogaria . . . . . . . . . . . . . . . . . . . . . . . . 91
pourtalesi, Nectoteuthis . . . . . . . . . . . . . . . . . . . . . . . . 203
prabahari, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 110 ?111
prashadi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
prionota, Solitosepia . . . . . . . . . . . . . . . . . . . . . . . . . 105
pteropus, Sthenoteuthis . . . . . . . . . . . . . . . . . . . . . . . . . 18
pulchra, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Purpo seccia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
pygmaeus, Idiosepius . . . . . . . . . . . . . . . . . . . . . . . 9, 210
Pygmy cuttlefishes . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
Pygmy squids . . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 22, 25
R
Ram?s horn . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Ram?s horn squid . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 211
Ram?s horn squids. . . . . . . . . . . . . . . . . . . . . . . . . . 22, 25
ramani, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113 ?114
rappiana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
recurvirostra, Sepia . . . . . . . . . . . . . . . . . . . . . . . 114?115
reesi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Rellena . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Relleno . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
repertus, Nautilus . . . . . . . . . . . . . . . . . . . . . . . 51, 53?54
rex, Deciruseoua . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
rex, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87, 97, 127
rhoda, Arctosepia . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
rhoda, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . 75?76, 147
Rhombosepion hieronis . . . . . . . . . . . . . . . . . . . . . . . 142
Rhombosepion robsoni . . . . . . . . . . . . . . . . . . . . . . . 147
robsoni, Rhombosepion . . . . . . . . . . . . . . . . . . . . . . . 147
robsoni, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97, 147
Robust bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . 166
robusta, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . 166 ?167
rondeleti, Sepiola . . . . . . . . . . . . . . . . . . . . . 167 ?168, 177
Rondeletiola minor . . . . . . . . . . . . . . . 164, 174 ?175, 191
Rosecone cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Rossia (Austrorossia) australis . . . . . . . . . . . . . . . . 193
Rossia (Semirossia) equalis . . . . . . . . . . . . . . . . . . . 188
Rossia antillensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
Rossia australis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
Rossia bipapillata . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
Rossia brachyura . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia bullisi . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193, 196
Rossia caroli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Rossia enigmatica . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Rossia glaucopis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia macrosoma. . . . . . . . . . . . . . . . 176, 183 ?184, 191
Rossia mastigophora . . . . . . . . . . . . . . . . . . . . . 195, 197
Rossia megaptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . 196
Rossia moelleri . . . . . . . . . . . . . . . . . . . . . . . . . . 186, 196
Rossia mollicella . . . . . . . . . . . . . . . . . . . . . . . . . 195?196
Rossia pacifica . . . . . . . . . . . . . . . . . . . . . . . . . . 186, 195
Rossia pacifica diegensis. . . . . . . . . . . . . . . . . . . 186, 197
Rossia pacifica pacifica . . . . . . . . . . . . . . . . 185 ?186, 197
Rossia palpebrosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Rossia patagonica . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Rossia spp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 194
Rossia tenera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
Rossia tortugaensis . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
rossiaeformis, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . 180
ROSSIINAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
rostrata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
rouxi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
rouxii, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
rozella, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 116
rozella, Solitosepia . . . . . . . . . . . . . . . . . . . . . . . . . . 116
rupellaria, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
S
Sabbidije . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
sagittatus, Todarodes . . . . . . . . . . . . . . . . . . . . . . . 10, 18
savignyi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
saya, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Scarpetta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Scarpitelle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Scarpitta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
scolopes, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . . 181
scrobiculatus, Allonautilus . . . . . . . . . . . . . . . . 51, 53, 55
scrobiculatus, Nautilus . . . . . . . . . . . . . . . . . . . . . . . . 55
Sea mops . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Secce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seccetella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seche . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seiba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seich . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seiche . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seiche ? os en couteau . . . . . . . . . . . . . . . . . . . . . . . . . 78
Seiche ? sepion ovale . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Seiche africaine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Seiche aiguille. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Seiche andreana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Seiche araign?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Seiche au c?ne ros? . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Seiche australe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Seiche baisers. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Seiche capuchon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Seiche commune . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Cephalopods of the World 255
Seiche d?Arabie . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Seiche d?Hedley . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Seiche d?Oman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seiche de Guin?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Seiche de Ken . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Seiche de Papouasie . . . . . . . . . . . . . . . . . . . . . . . . . 105
Seiche de Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Seiche de Voss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Seiche de Whitley . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
Seiche de Zanzibar . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
Seiche dor?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Seiche du Viet Nam. . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Seiche ?l?gante . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Seiche encrier . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Seiche ?toil?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Seiche flamboyante . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Seiche g?ante africaine . . . . . . . . . . . . . . . . . . . . . . . . 88
Seiche g?ante australienne . . . . . . . . . . . . . . . . . . . . . 67
Seiche gracile . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Seiche grandes mains . . . . . . . . . . . . . . . . . . . . . . . . . 91
Seiche grenouille . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Seiche gros dos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Seiche hame?on . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Seiche impressionnante . . . . . . . . . . . . . . . . . . . . . . . 109
Seiche kobi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Seiche madokai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
Seiche manifique . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Seiche petites mains. . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Seiche pharaon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Seiche pieuvre . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Seiche r?ticul?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Seiche ros?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Seiche stri?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Seiche trapue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Seiche trident . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
Seiches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Seiches pygm?es . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
Semirossia equalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
Semirossia patagonica . . . . . . . . . . . . . . . . . . . . . . . . 197
Semirossia tenera . . . . . . . . . . . . . . . . . . . . . . . . 188?189
senta, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sepia . . . . . . . . . . . . . . . . . . . . . . . 7, 10, 56, 85, 101, 125
Sepia (Doratosepion) misakiensis . . . . . . . . . . . . . . 150
Sepia (Metasepia) pfefferi . . . . . . . . . . . . . . . . . . . . . . 60
Sepia (Sepiella) microcheirus . . . . . . . . . . . . . . . . . . 130
Sepia achaparrada . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Sepia aculeata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63 ?64
Sepia acuminata. . . . . . . . . . . . . . . . . . . . . . . . . . . 97,137
Sepia adami. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia affinis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Sepia africana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Sepia albatrossi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Sepia andreana . . . . . . . . . . . . . . . . . . . . . . 65 ?66, 89?90
Sepia andreanoides . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Sepia angulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia apama . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67 ?68
Sepia appellofi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 137
Sepia arabica. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Sepia ar?biga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Sepia aureomaculata. . . . . . . . . . . . . . . . . . . . . . . . . . 138
Sepia austral . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Sepia australis . . . . . . . . . . . . . . . . . . . 58, 70 ?71, 80, 121
Sepia bandensis. . . . . . . . . . . . . . . . . . . . . . . . . . . 72 , 138
Sepia bartletti . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72, 138
Sepia bathyalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 138
Sepia baxteri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72, 138
Sepia bertheloti . . . . . . . . . . . . . . . . . . . . . . . . . 73 , 79?80
Sepia bidhaia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75, 139
Sepia biserialis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Sepia braggi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75 ?76
Sepia brazolargo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Sepia brevimana . . . . . . . . . . . . . . . . . . . . . . 77 , 119?120
Sepia burnupi . . . . . . . . . . . . . . . . . . . . . . . . 98, 123, 139
Sepia capensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Sepia caperuza . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
Sepia carinata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Sepia chirotrema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Sepia com?n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Sepia con punta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Sepia confusa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia cottoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75, 140
Sepia cultrata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Sepia dannevig i. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Sepia de aguja . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Sepia de cono rosado . . . . . . . . . . . . . . . . . . . . . . . . . 116
Sepia de Hedley . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Sepia de Ken . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Sepia de Oman . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sepia de Papua . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Sepia de sepi?n de cuchillo . . . . . . . . . . . . . . . . . . . . . 78
Sepia de sepi?n oval . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Sepia de Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
Sepia de Viet Nam . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Sepia de Voss . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
Sepia de Web . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
S?pia de Web . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Sepia de Whitley . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
Sepia de Zanzibar . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
Sepia dollfusi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia dorada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Sepia dubia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 140
Sepia elegans. . . . . . . . . . . . . . . . . . . . . . 58, 79 , 101, 104
Sepia elegante . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Sepia elliptica . . . . . . . . . . . . . . . . . . . . . . . 81 ?82, 84?85
Sepia elobyana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Sepia elongata . . . . . . . . . . . . . . . . . . . . . . . 122-123, 141
Sepia erostrata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Sepia esculenta . . . . . . . . . 77?78, 81?82, 84 , 96?97, 108
Sepia estrellada. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
Sepia estriada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Sepia fara?n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia faurei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Sepia filibrachia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
256 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Sepia filliouxi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Sepia foliopeza . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Sepia formosana . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia galei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Sepia ganchuda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Sepia gibba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia gigante africana . . . . . . . . . . . . . . . . . . . . . . . . . 88
Sepia gigante australiana . . . . . . . . . . . . . . . . . . . . . . . 67
Sepia gr?cil . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Sepia grahami . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Sepia guineana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Sepia harmeri. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Sepia hedleyi . . . . . . . . . . . . . . . . . . . . . . . . . . 87 , 97, 127
Sepia hercules . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91?92
Sepia hieronis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58, 142
Sepia hierredda . . . . . . . . . . . . . . . . . . . . . 74, 88 ?89, 101
Sepia hoylei . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Sepia imperiale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sepia impresionante . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Sepia incerta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia indica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Sepia inerme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
S?pia inerme japonaise . . . . . . . . . . . . . . . . . . . . . . . 132
Sepia inerme japonesa . . . . . . . . . . . . . . . . . . . . . . . . 132
Sepia inermis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
S?pia inerme . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Sepia insignis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 142
Sepia irvingi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia italica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Sepia ivanovi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia joubini . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia kiensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
Sepia kobi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Sepia kobiensis . . . . . . . . . . . . . . . . . . . . . . 65?66, 89 , 98
Sepia koilados . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Sepia labiada. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Sepia latimanus. . . . . . . . . . . . . . . . . . . . . . . . . . . . 58, 91
Sepia limata. . . . . . . . . . . . . . . . . . . . . . . . . . . 75?76, 144
Sepia listada grande . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Sepia listada peque?a . . . . . . . . . . . . . . . . . . . . . . . . . 110
Sepia llamativa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60
Sepia longipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Sepia loriga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Sepia lorigera . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94 , 126
Sepia lycidas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
Sepia madokai. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81, 97
Sepia magn?fica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Sepia mascarensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . 144
Sepia mazicorta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Sepia mazuda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Sepia mediterranea . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Sepia mercatoris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Sepia mestus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 144
Sepia mira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia mirabilis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia mozambica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Sepia murrayi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Sepia novaehollandiae . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia officinalis . . . . 7, 9, 58, 74, 80, 88?89, 99 ?100, 104
Sepia officinalis filliouxi . . . . . . . . . . . . . . . . . . . . . . 101
Sepia officinalis hierredda . . . . . . . . . . . . . . . . . . . . 101
Sepia officinalis mediterranea . . . . . . . . . . . . . . . . . 101
Sepia officinalis vermiculata . . . . . . . . . . . . . . . . . . 101
Sepia omani. . . . . . . . . . . . . . . . . . . . . . . . . . 98, 101 , 127
Sepia opipara . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
Sepia orbignyana. . . . . . . . . . . . . . . . . . . 58, 80, 101, 103
Sepia ornada . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Sepia ornata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
S?pia orn?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Sepia palmata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Sepia papillata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
Sepia papuensis . . . . . . . . . . . . . . . . . . . . . . . . . . 105 ?106
Sepia pardalis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia pardex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia peterseni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia pharaonis . . . . . . . . . . . . 58, 92, 106 ?108, 114, 147
Sepia plana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 146
Sepia plangon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Sepia plathyconchalis . . . . . . . . . . . . . . . . . . . . . 117, 146
Sepia prabahari . . . . . . . . . . . . . . . . . . . . . . . . . . 110 ?111
Sepia prashadi . . . . . . . . . . . . . . . . . . . . . . . . . . . 111?112
Sepia pulchra . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sepia puntiaguda. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Sepia ramani . . . . . . . . . . . . . . . . . . . . . . . . 108, 113 ?114
Sepia rana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 98
Sepia rappiana. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Sepia recurvirostra . . . . . . . . . . . . . . . . . . . . . . . 114?115
Sepia reesi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sepia rex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87, 97, 127
Sepia reticulata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Sepia rhoda . . . . . . . . . . . . . . . . . . . . . . . . . . . 75?76, 147
Sepia robsoni. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97, 147
Sepia robusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Sepia rostrata. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Sepia rouxi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia rouxii . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia rozella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 116
Sepia rupellaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Sepia savignyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117
Sepia saya . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia senta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia sewelli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia simoniana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Sepia singaporensis . . . . . . . . . . . . . . . . . . . . . . . . . . 114
Sepia sinope . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70?71
Sepia smithi . . . . . . . . . . . . . . . . . . . . . . . . . 118 ?119, 128
Sepia sokotriensis . . . . . . . . . . . . . . . . . . . . . . . . 123, 149
Sepia sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Sepia stellifera . . . . . . . . . . . . . . . . . . . . . 77, 81, 119 ?120
Sepia subaculeata . . . . . . . . . . . . . . . . . . . . . . . . . 95?96
Sepia subplana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Sepia subtenuipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Sepia sulcata . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 ?121
Sepia tala. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
Cephalopods of the World 257
Sepia tanybracheia . . . . . . . . . . . . . . . . . . . . . . . . 95, 150
Sepia tenuipes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia thurstoni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia tigris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia tintero . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Sepia tokioensis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Sepia torosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Sepia toyamensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Sepia tridente . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
Sepia trygonina . . . . . . . . . . . . . . . . . . . . . . . 98, 122 ?123
Sepia tuberculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Sepia tullbergi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Sepia typica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
Sepia vercoi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75, 151
Sepia vermiculata . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
Sepia verrucosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Sepia vossi . . . . . . . . . . . . . . . . . . . . . . . 87, 101?102, 127
Sepia whitleyana . . . . . . . . . . . . . . . . . . . . . 118?119, 128
Sepia zanzibarica . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
SEPIADARIIDAE . . . . . . . . . . . . . . . . . . . . . 3, 33, 56, 204
Sepiadarium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Sepiadarium auritum . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiadarium austrinum . . . . . . . . . . . . . . . . . . . . . . . 205
Sepiadarium gracilis . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiadarium kochii . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
Sepiadarium nipponianum . . . . . . . . . . . . . . . . . . . . . 207
Sepias . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Sepias pigmeas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 208
Sepie . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sepiella . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Sepiella cyanea . . . . . . . . . . . . . . . . . . . . . . . . . . 131, 151
Sepiella inermis . . . . . . . . . . . . . . . . . . . . . . 130 ?131, 133
Sepiella japonica . . . . . . . . . . . . . . . . . . . . . . . . . 132 ?133
Sepiella maindroni . . . . . . . . . . . . . . . . . . . . . . 130, 133
Sepiella mangkangunga . . . . . . . . . . . . . . . . . . . . . . . 152
Sepiella ocellata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 152
Sepiella ornata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 134
Sepiella sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 136
Sepiella weberi . . . . . . . . . . . . . . . . . . . . . . . . . . 135 ?136
Sepieta com?n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Sepieta elegante . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
Sepieta misteriosa . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Sepietas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
Sepietta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Sepietta minor . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
Sepietta neglecta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
Sepietta obscura . . . . . . . . . . . . . . . . . . . . . . . . . 177 ?178
Sepietta oweniana . . . . . . . . . . . . 164, 176?178 ?179, 191
Sepietta petersi . . . . . . . . . . . . . . . . . . . . . . . . . . 178, 182
Sepietta sp. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
SEPIIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33, 56?57
Sepiids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Sepija . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
SEPIOIDEA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29, 56
Sepioids. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24, 33
Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22, 167
Sepiola affinis . . . . . . . . . . . . . . . . . . . . . . . 158 ?159, 177
Sepiola an?loga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
Sepiola atlantica . . . . . . . . . . . . . . . . . . . . . . . . . 159 ?160
Sepiola atl?ntica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Sepiola aurantiaca . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola birostrata. . . . . . . . . . . . . 161 ?162, 165?166, 170
Sepiola dispar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
Sepiola enana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
Sepiola intermedia. . . . . . . . . . . . . . . . . . . . 159, 162 ?163
Sepiola japonica . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Sepiola knudseni . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola leng?ita . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Sepiola leucoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
Sepiola ligulata . . . . . . . . . . . . . . . . . . . . . . . . . . 163 ?164
Sepiola lineolata . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiola macrosoma . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Sepiola manchada . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
Sepiola mariposa . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
Sepiola nudosa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
Sepiola oweniana . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
Sepiola pacifica . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepiola parva. . . . . . . . . . . . . . . . . . . . . . . . 162, 165 ?166
Sepiola petersi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 182
Sepiola pfefferi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola robusta . . . . . . . . . . . . . . . . . . . . . . . . . . 166 ?167
Sepiola rondeleti . . . . . . . . . . . . . . . . . . . . . 167 ?168, 177
Sepiola rossiaeformis . . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola steenstrupiana . . . . . . . . . . . . . . . . . . . . . . . . 180
Sepiola tasmanica . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Sepiola trirostrata . . . . . . . . . . . . . . . . . . . . . . . . 162, 169
Sepiolas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
S?piole. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
S?piole ? gros yeux . . . . . . . . . . . . . . . . . . . . . . . . . . 194
S?piole analogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . 158
S?piole australe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
S?piole bobie . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 174
S?piole bossel?e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 169
S?piole calamarette . . . . . . . . . . . . . . . . . . . . . . . . . . 189
S?piole colibri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 170
S?piole commune . . . . . . . . . . . . . . . . . . . . . . . . . . . . 178
S?piole cracheuse . . . . . . . . . . . . . . . . . . . . . . . . . . . 188
S?piole de Carol . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
S?piole de la Tortue . . . . . . . . . . . . . . . . . . . . . . . . . . 187
S?piole diff?rente . . . . . . . . . . . . . . . . . . . . . . . . . . . . 198
S?piole du Pacifique bor?al . . . . . . . . . . . . . . . . . . . . 185
S?piole du Tasmania . . . . . . . . . . . . . . . . . . . . . . . . . . 173
S?piole ?l?gante . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 176
S?piole grandes oreilles . . . . . . . . . . . . . . . . . . . . . . . 159
S?piole interm?diaire . . . . . . . . . . . . . . . . . . . . . . . . . 162
S?piole japonaise . . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
S?piole languette . . . . . . . . . . . . . . . . . . . . . . . . . . . . 163
S?piole leucopt?re . . . . . . . . . . . . . . . . . . . . . . . 161 , 199
S?piole melon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
S?piole mignonne . . . . . . . . . . . . . . . . . . . . . . . . . . . . 192
S?piole mimika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 171
S?piole mouchet?e . . . . . . . . . . . . . . . . . . . . . . . . . . . 165
S?piole myst?rieuse . . . . . . . . . . . . . . . . . . . . . . . . . . 177
S?piole naine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
258 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
Cephalopods of the World 259
S?piole papillon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 161
S?piole robuste . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 166
S?pioles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
S?piolette de Koch . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
S?piolette du sud . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
S?piolettes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
SEPIOLIDAE . . . . . . . . . . . . . . . . . . . . . . . 33?34, 56, 153
Sepiolids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Sepiolilla de Koch . . . . . . . . . . . . . . . . . . . . . . . . . . . . 206
Sepiolilla sure?a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Sepiolillas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Sepiolina japonesa . . . . . . . . . . . . . . . . . . . . . . . . . . . 201
Sepiolina nipponensis . . . . . . . . . . . . . . . . . . . . . . . . . 201
Sepiolinas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 153
SEPIOLINAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22, 158
Sepioloidea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 204
Sepioloidea lineolata . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepioloidea pacifica . . . . . . . . . . . . . . . . . . . . . . . . . . 207
Sepioteuthis. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
Seppa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seppia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seppia comune . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seppia elegante. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
Seppia pizzuta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
Seppio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Seppiola affine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 159
Seppiola comune. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 179
Seppiola di Rondelet . . . . . . . . . . . . . . . . . . . . . . . . . . 168
Seppiola grossa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Seppiola grossa di fondo . . . . . . . . . . . . . . . . . . . . . . . 191
Seppiola intermedia. . . . . . . . . . . . . . . . . . . . . . . . . . . 163
Seppiola linguetta . . . . . . . . . . . . . . . . . . . . . . . . . . . . 164
Seppiola minore. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 175
Seppiola misteriosa . . . . . . . . . . . . . . . . . . . . . . . . . . . 177
Seppiola robusta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
serventyi, Heteroteuthis . . . . . . . . . . . . . . . . . . . . . . 203
serventyi, Heteroteuthis (Stephanoteuthis) . . . . . . . . . 203
sewelli, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
Shirikusari . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Shiriyakeika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 133
Shortclub cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Shoubia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sibia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sicca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Siccia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
simoniana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 148
singaporensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 114
sinope, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70?71
Sipa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sipia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Sipionet . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Slender cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Small striped cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . 110
Smith?s cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 118
smithi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 118 ?119, 128
Sobbeit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
sokotriensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . 123, 149
Solitosepia genista. . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Solitosepia lana . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Solitosepia occidua . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Solitosepia plangon adhaesa . . . . . . . . . . . . . . . . . . 109
Solitosepia prionota . . . . . . . . . . . . . . . . . . . . . . . . . 105
Solitosepia rozella . . . . . . . . . . . . . . . . . . . . . . . . . . . 116
Solitosepia submestus . . . . . . . . . . . . . . . . . . . . . . . . 105
Sotong-besar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
Soubia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Soupia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Southern bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . 173
Southern bottletail squid . . . . . . . . . . . . . . . . . . . . . . 205
Southern cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Spider cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Spineless cuttlefish. . . . . . . . . . . . . . . . . . . . . . . . . . . 130
Spirula. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 6, 33, 56
Spirula spirula . . . . . . . . . . . . . . . . . . . . . . . . . . . 211 ?212
spirula, Nautilus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
spirula, Spirula . . . . . . . . . . . . . . . . . . . . . . . . . . 211 ?212
Spirule . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
SPIRULIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . 33, 56, 211
sponsalis, Bathypolypus . . . . . . . . . . . . . . . . . . . . . . . 191
Spotty bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . 165
Squids . . . . . . . . . . . . . . 3, 6, 9?10, 17, 27, 29, 31, 35?36
Starry cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 119
steenstrupiana, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . 180
stellifera, Sepia . . . . . . . . . . . . . . . . . . . . 77, 81, 119 ?120
stenodactyla, Euprymna . . . . . . . . . . . . . . . . . . . . . . . 182
stenomphalus, Nautilus . . . . . . . . . . . . . . . . . . . 51, 53?54
Stephanoteuthis hawaiiensis . . . . . . . . . . . . . . . . . . . 203
Sthenoteuthis oualaniensis . . . . . . . . . . . . . . . . . . . . 9, 18
Sthenoteuthis pteropus . . . . . . . . . . . . . . . . . . . . . . . . . 18
Stoloteuthis iris . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 203
Stoloteuthis leucoptera . . . . . . . . . . . . . . . . . . . . . . . . 199
Stoloteuthis nipponensis . . . . . . . . . . . . . . . . . . . . . . 201
Stout bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Striking cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
Stumpy cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
subaculeata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . 95?96
submestus, Solitosepia . . . . . . . . . . . . . . . . . . . . . . . 105
subplana, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
subtenuipes, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
S?bye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
sulcata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . 120 ?121
Sumiika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 85
Supia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
T
tala, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 149
tanybracheia, Sepia . . . . . . . . . . . . . . . . . . . . . . . . 95, 150
tasmanica, Euprymna . . . . . . . . . . . . . . . . . . . . . . . . . 173
tasmanica, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Tenaga-kouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
tenera, Heteroteuthis . . . . . . . . . . . . . . . . . . . . . . . . . 189
tenera, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
260 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
tenera, Semirossia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 189
tenuipes, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Tenuisepia mira . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 145
TEUTHOIDEA . . . . . . . . . . . . . . . . . . . . . . . . . . 29, 31, 34
thailandicus, Idiosepius . . . . . . . . . . . . . . . . . . . . 209?210
thurstoni, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
THYSANOTEUTHIDAE . . . . . . . . . . . . . . . . . . . . . . . . . 18
tigris, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Todarodes filippovae. . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Todarodes pacificus . . . . . . . . . . . . . . . . . . . . . . . . 10, 18
Todarodes sagittatus . . . . . . . . . . . . . . . . . . . . . . . . 10, 18
tokioensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150
Tongue bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . 163
Torafu-kouika. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 108
torosa, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 106
Tortuga bobtail squid . . . . . . . . . . . . . . . . . . . . . . . . . 187
tortugaensis, Rossia . . . . . . . . . . . . . . . . . . . . . . . . . . 187
toyamensis, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Tremoctopus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Trident cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
trirostrata, Sepiola . . . . . . . . . . . . . . . . . . . . . . . . 162, 169
trygonina, Sepia . . . . . . . . . . . . . . . . . . . . . . 98, 122 ?123
trygoninum, Doratosepion . . . . . . . . . . . . . . . . . . . . 122
tuberculata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
tullbergi, Metasepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
tullbergi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
typica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 151
typicus, Hemisepius . . . . . . . . . . . . . . . . . . . . . . . . . . 151
U
ursulae, Crumenasepia . . . . . . . . . . . . . . . . . . . . . . . 106
Usubeni-kouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
V
Vampire squids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Vampires . . . . . . . . . . . . . . . . . . . . . . . . . . 3, 9, 21, 26, 31
vercoi, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75, 151
vermiculata, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 124
vermiculata, Sepia officinalis . . . . . . . . . . . . . . . . . . 101
verreaux, Amplisepia . . . . . . . . . . . . . . . . . . . . . . . . . . 67
verrucosa, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Viet Nam cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
vietnamica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Voss? cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 127
vossi, Sepia . . . . . . . . . . . . . . . . . . . . . . 87, 101?102, 127
vulgaris, Loligo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7, 9
vulgaris, Octopus . . . . . . . . . . . . . . . . . . . . . . . . . . . 9?10
Vurpascele. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
W
Web?s cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
weberi, Heteroteuthis . . . . . . . . . . . . . . . . . . . . . . . . 203
weberi, Heteroteuthis (Heteroteuthis) . . . . . . . . . . . . . 203
weberi, Sepiella . . . . . . . . . . . . . . . . . . . . . . . . . . 135 ?136
Whitley?s cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 128
whitleyana, Acanthosepion . . . . . . . . . . . . . . . . . . . . 128
whitleyana, Sepia . . . . . . . . . . . . . . . . . . . . . 118?119, 128
Y
Yi muk woo chak . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 96
Yoroppa kouika . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
Z
Zanzibar cuttlefish . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
zanzibarica, Sepia . . . . . . . . . . . . . . . . . . . . . . . . . . . . 129
Zeekat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 101
8. LIST OF COLOUR PLATES
P L ATE I
1. Nautilus pompilius
2. Nautilus pompilius male
3. Nautilus pompilius adult
4. Allonautilus scrobiculatus and Nautilus pompilius
5. Allonautilus scrobiculatus and Nautilus pompilius
6. Nautilus belauensis mature animal
P L ATE II
7. Nautilus belauensis juvenile
8. Nautilus belauensis
9. Nautilus belauensis
10. Nautilus stenomphalus
11. Metasepia pfefferi
12. Sepia aculeata juvenile
13. Sepia aculeata
14. Sepia aculeata
P L ATE II I
15. Sepia apama
16. Sepia apama
17. Sepia apama
18. Sepia apama
19. Sepia apama
20. Sepia apama
P L ATE IV
21. Sepia bandensis
22. Sepia bandensis
23. Sepia latimanus
24. Sepia latimanus large adult
25. Sepia latimanus
26. Sepia latimanus frontal view
27. Sepia latimanus juvenile
28. Sepia officinalis mature male
P L ATE V
29. Sepia officinalis mature male
30. Sepia officinalis juvenile
31. Sepia papuensis
32. Sepia pharaonis
33. Sepia pharaonis female
34. Sepia pharaonis male
35. Sepia plangon
36. Sepia smithi
P L ATE VI
37. Sepiella inermis
38. Sepiola
39. Euprymna tasmanica
40. Euprymna tasmanica adult
41. Euprymna tasmanica
P L ATE VI I
42. Sepiadarium austrinum
43. Sepiadarium austrinum
44. Sepiadarium austrinum
45. Sepioloidea lineolata
46. Sepioloidea lineolata
P L ATE VI I I
47. Beach seine and seine boat - India
48. Lift nets and fish market - India
48. Octopus maya catch from pots - Venezuela
50. Squid jigging machines at night - Japan
51. Trawl catch of squid - New England
52. Traditional waterwheel clay scoops recycled for
Octopus vulgaris fishery - Sant Carles de la R?pita,
Catalonia
53. Clay octopus pots linked by a line - Farwah Lagoon,
Libya
54. PVC pipe pots linked by a line for Octopus vulgaris
fishery - Deltebre, Catalonia
P L ATE IX
55. Clay pots for Octopus vulgaris fishery - L?Ampolla,
Catalonia
56. Squid catch on deck - Western North Atlantic
57. Squid drying in sun - Japan
58. Squid drying in sun - Japan
59. Closing up purse seine with squid - Japan
60. Soft body (green) and hard body (black) squid jigging
hooks on reel - Japan
Cephalopods of the World 261
262 FAO Species Catalogue for Fishery Purposes No. 4, Vol. 1
ADDITIONAL INFORMATION
Plate I
Fig. 3: This animal was caught in a baited trap set at 330 m which was then brought slowly to the surface. Milne Bay,
Papua New Guinea.
Fig 4: Photographed in shallow water off Ndrova Island, Manus, Admiralty Islands, Papua New Guinea, after being
trapped at about 270 m depth.
Fig. 5: Photographed at about 270 m off Ndrova Island, Manus, Admiralty Islands, Papua New Guinea. Large red
snapper Etelis carbunculus was also attracted to bait and entered trap.
Fig. 6: Mature animal trapped at about 30 m depth off Mutremdiu Point, Palau Islands, South Pacific.
Plate II
Fig. 7: Juvenile trapped at about 300 m depth off Mutremdiu Point, Palau Islands, South Pacific.
Fig. 8: Two specimens attracted to tuna bait at 217 m depth off Mutremdiu Point, Palau Islands, South Pacific.
Fig. 9: Two adults approaching exterior foreground with tentacles extended in typical search posture. Six other nautilus
are within trap; one pair is mating (lower left). Photo taken at 274 m depth off Mutremdiu Point, Palau Islands,
South Pacific.
Fig. 10: Specimen captured at about 250 m off Lizard Island, Great Barrier Reef, Australia.
Fig. 11: Specimen ?walking? on substrate with ventral arms and abdominal flaps at 20 m. Lizard Island, Great Barrier Reef,
Australia.
Fig. 12: Juvenile hovering above the substrate. Total length 15 cm. Papua New Guinea.
Fig. 13: Specimen sitting on bottom in highly cryptic body pattern. Milne Bay, Papua New Guinea.
Plate III
Fig. 15: Photographed in south Australia.
Fig. 16: Sneaker male (above right) in mottled colour pattern with egg-laying female (below) and mate-guarding male.
Fig. 17: Two males in combat display over an egg-laying female (visible bottom left).
Fig. 18: Three males compete to mate with a small female (far left).
Fig. 19: Frontal ?spade? display of male, often used to flip an oppenent over.
Fig. 20: Photographed in south Australia.
Plate IV
Fig. 21: North Sulawesi, Indonesia.
Fig. 23: Great Barrier Reef, Australia
Fig. 24: Large adult hanging beneath a tabletop coral head. Total length about 50 cm. Palau Islands, South Pacific.
Fig. 25: A somewhat cryptic body pattern. Palau Islands, South Pacific.
Fig. 26: Frontal view illustrates how a cuttlefish can literally hide its body behind its own head and arms. Photo taken in the
laboratory (represents a wild population from Palau Islands, South Pacific).
Fig. 27: Photo of juvenile taken in the laboratory (represents a wild population from Palau Islands, South Pacific).
Fig. 28: Photo of mature male taken in the laboratory (represents a wild population from eastern Atlantic, UK or France).
Plate V
Fig. 29: Mature male showing intense zebra display. Photo taken in the laboratory (represents a wild population from the
eastern Atlantic).
Fig. 30: Juvenile showing the green iridescence on the ventral mantle in contrast to the pattern on the dorsal mantle.
Photo was taken in the laboratory (represents a wild population from the eastern Atlantic).
Fig. 31: Western Australia.
Fig. 32: Male on top of female being guarded. Photo taken in the laboratory (represents a wild population of Gulf of
Thailand).
Fig. 33: Female swimming in large culture tank in general mottled body pattern with an iridescent blue stripe along fin
base. Photo was taken in the laboratory (represents a wild population from the Gulf of Thailand, western Pacific).
Fig. 34: Male in dramatic zebra stripe body pattern with arms spread as he postures toward other males nearby. Photo
taken in the laboratory.
Fig. 35: New South Wales, Australia.
Plate VI
Fig. 37: Singapore.
Fig. 40: Adult with fully developed light organ.
Fig. 41: Victoria, Australia.
Plate VII
Fig. 43: Victoria, Australia.
Fig. 44: Pair mating face-to-face (female stores sperm in a pouch around her mouth).
Fig. 46: Specimen using both eyes to take aim at a passing shrimp. Western Australia.
COLOUR PLATES
PLATE I
3. Nautilus pompilius adult
(J.W. Forsythe)
5. 2 Allonautilus scrobiculatus and 3 Nautilus pompilius
(W.B. Saunders)
2. Nautilus pompilius male
(M. Norman)
4. Allonautilus scrobiculatus (left) and Nautilus pompilius (right)
(W.B. Saunders)
6. Nautilus belauensis mature animal
(W.B. Saunders)
1. Nautilus pompiliu s
(M. Norman)
PLATE II
8. Nautilus belauensis
(W.B. Saunders)
7. Nautilus belauensis juvenile
(W.B. Saunders)
9. Nautilus belauensis
(W.B. Saunders)
10. Nautilus stenomphalus
(W.B. Saunders)
11. Metasepia pfefferi
(C.F.E. Roper)
12. Sepia aculeata juvenile
(J.W. Forsythe)
13. Sepia aculeata
(J.W. Forsythe)
14. Sepia aculeata
(J.W. Forsythe)
PLATE III
15. Sepia apama
(M. Norman)
16. Sepia apama
(M. Norman)
17. Sepia apama
(M. Norman)
18. Sepia apama
(M. Norman)
20. Sepia apama
(M. Norman)
19. Sepia apama
(M. Norman)
PLATE IV
21. Sepia bandensis
(M. Norman)
22. Sepia bandensis
(M. Norman)
23. Sepia latimanus
(M. Norman)
24. Sepia latimanus large adult
(J.W. Forsythe)
27. Sepia latimanus juvenile
(J.W. Forsythe)
28. Sepia officinalis mature male
(J.W. Forsythe)
25. Sepia latimanus
(J.W. Forsythe)
26. Sepia latimanus frontal view
(J.W. Forsythe)
PLATE V
29. Sepia officinalis mature male
(J.W. Forsythe)
30. Sepia officinalis juvenile
(J.W. Forsythe)
31. Sepia papuensis
(M. Norman)
32. Sepia pharaonis
(J.W. Forsythe)
35. Sepia plangon
(M. Norman)
36. Sepia smithi
(M. Norman)
33. Sepia pharaonis female
(J.W. Forsythe)
34. Sepia pharaonis male
(J.W. Forsythe)
PLATE VI
37. Sepiella inermis
(M. Norman)
38. Sepiola
(J.W. Forsythe)
39. Euprymna tasmanica
(M. Norman)
40. Euprymna tasmanica adult
(M. Norman)
41. Euprymna tasmanica
(M. Norman)
PLATE VII
42. Sepiadarium austrinum
(M. Norman)
43. Sepiadarium austrinum
(M. Norman)
44. Sepiadarium austrinum
(M. Norman)
45. Sepioloidea lineolata
(M. Norman)
46. Sepioloidea lineolata
(M. Norman)
PLATE VIII
47) Beach seine and seine boat - India
(C.F.E. Roper)
48) Lift nets and fish market - India
(C.F.E. Roper)
49) Octopus maya catch from pots - Venezuela
(W.F. Rathjen)
50) Squid jigging machines at night - Japan
(W.F. Rathjen)
53) Clay octopus pots linked by a line - Farwah Lagoon, Libya
(M. Lamboeuf)
54) PVC pipe pots linked by a line for Octopus vulgaris fishery -
Deltebre, Catalonia
(M. Demestre)
51) Trawl catch of squid - New England
(W.F. Rathjen)
52) Traditional waterwheel clay scoops recycled for Octopus vulgaris
fishery - Sant Carles de la R?pita, Catalonia
(P. S?nchez)
PLATE IX
55) Clay pots for Octopus vulgaris fishery - L?Ampolla, Catalonia
(J. Lleonart)
56) Squid catch on deck - Western North Atlantic
(W.F. Rathjen)
57) Squid drying in sun - Japan
(C.F.E. Roper)
58) Squid drying in sun - Japan
(C.F.E. Roper)
59) Closing up purse seine with squid -
Japan
(C.F.E. Roper)
60) Soft body (green) and hard body (black)
squid jigging hooks on reel - Japan
(C.F.E. Roper)

This is the first volume of the entirely rewritte n, revised and updated version of the original
FAO Catalogue of Cephalopods of the World (1984). The present volume is a multiauthored
compilation that reviews six families: Nautilidae, Sepiidae, Sepiolidae, Sepiadariidae, Idiosepiidae
and Spirulidae, with 23 genera and the 201 speci es known 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: va lid modern systematic name and original
citation of the species (or subspecies); mai n synonyms; English, French and Spanish FAO
names for the species; illustrations of dorsal and ventral aspect of the whole animal (as
necessary) and other distinguishing illustrations; field characteristics; diagnostic features;
geographic and vertical distribution, including GIS map; size; habitat; biology;
interest to fishery; local names when availabl e; a remarks section (as necessary) and literature.
The volume is fully indexed and also include s sections on terminology and measurements,
an extensive glossary, an introduction with an updated review of the existing biological
knowledge on cephalopods (includi ng fisheries information and catch data for recent years) and
a dedicated bibliography.
9 7 8 9 2 5 1 0 5 3 8 3 6
TC/M/A0150E/1/10.05/2000
ISBN 92-5-105383-9  ISSN 1020-8682