ISSN 1020-8682

FAO Species Catalogue for Fishery Purposes No. 3

THREADFINS OF THE WORLD(Family Polynemidae)
AN ANNOTATED AND ILLUSTRATED CATALOGUE OFPOLYNEMID SPECIES KNOWN TO DATEFAO Species Catalogue for Fishery Purposes No. 3

FIR/Cat.3

THREADFINS OF THE WORLD(Family Polynemidae)
AN ANNOTATED AND ILLUSTRATED CATALOGUE OFPOLYNEMID SPECIES KNOWN TO DATE

byHiroyuki Motomura
Ichthyology, Australian Museum
SydneyAustralia

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS
Rome, 2004

The designations employed and the presentation of material inthis publication do not imply the expression of any opinion
whatsoever on the part of the Food and Agriculture Organizationof the United Nations concerning the legal status of any country,
territory, city or area or of its authorities, or concerning thedelimitation of its frontiers or boundaries.

ISBN xx-x-xxxxxx-x

All rights reserved. No part of this publication may be reproduced, stored in a
retrieval system, or transmitted in any form or by any means, electronic,
mechanical, photocopying or otherwise, without the prior permission of the
copyright owner. Applications for such permission, with a statement of the
purpose and extent of the reproduction, should be addressed to the Director,
Publications Division, Food and Agriculture Organization of the United
Nations, Via delle Terme di Caracalla, 00100 Rome, Italy.

© FAO Rome 2004

Threadfins of the World

iii

PREPARATION OF THIS DOCUMENTT

his document was prepared under the FAO Fisheries Department Regular Programme by the Species Identification
Programme in the Marine Resources Service of the Fishery Resources Division. It is the third number in the new series:
FAO Species Catalogue for Fishery Purposes.

The family Polynemidae (threadfins) is important in commercial and sport fisheries of tropical and subtropical seas,
especially in the Middle and Near East, South Asia, Southeast Asia, Oceania, and west Africa. However, detailed fisheryinformation is usually not available, largely because of difficulties with the correct identification of species. This difficulty is
because of past taxonomic confusion and the similarity of overall body appearance and colour pattern of some species.
Therefore, this catalogue is timely as an accurate and complete worldwide guide to the identification of polynemids. It is
intended to help fisheries workers gather statistics and examine aspects of their biology that will be useful for the
management of this fishery resource. The author has studied polynemids taxonomically on the basis of a large number of
specimens, including all of available known types, in museum collections and markets throughout the world. He has
published numerous scientific articles on polynemids, which were the source for most of the taxonomic and ecologicalinformation in this catalogue.
In view of the importance of overall body appearance for identification, photographs for all species have been included.
Project managers: Michel Lamboeuf and Jordi Lleonart (FAO, Rome).
Scientific editor: Kent Carpenter (Old Dominion University, Norfolk, VA, USA).
Scientific reviser: Nicoletta De Angelis (FAO, Rome).
Editorial assistance: Michèle Kautenberger-Longo (FAO, Rome).
Scientific illustrator: Emanuela D’Antoni (FAO, Rome).
Page composition and indexing: Michèle Kautenberger-Longo (FAO, Rome).
Digitization of distribution maps: Fabio Carocci (FAO, Rome) and Elena V. Orlova (St. Petersburg).
Cover illustration: Emanuela D’Antoni (FAO, Rome).

Motomura, H.
Threadfins of the world. An annotated and illustrated catalogue of polynemid species known to date. Family
Polynemidae.
FAO Species Catalogue for Fishery Purposes. No. 3. Rome, FAO. 2004. 117 p., 151 figs, 6 pls.

ABSTRACTThis is the third number in the new FAO series of worldwide annotated and illustrated catalogues of major groups oforganisms that enter marine fisheries. The present volume on the family Polynemidae includes 41 species belonging
to 8 genera. There is an introductory section with general remarks on habitat and fisheries of the family, a glossary oftechnical terms, an illustrated key to each genus and all species, and a detailed account for all species. Species
accounts include an illustration of each species, scientific and vernacular names, and information on habitat, biology,
fisheries, size, relevant literature, and distribution. Following the species accounts are a list of nominal species in thefamily, a table of species by major marine fishing areas, and colour plates. A list of all nominal species and theirpresent allocations is given. The work is fully indexed and there is a comprehensive list of references to pertinent
literature.

Distribution
AuthorFAO Fisheries Officers
Regional Fisheries Councils and Commissions
Selector SC

iv

FAO Species Catalogue for Fishery Purposes No. 3

Acknowledgements
Many persons have assisted with the preparation of this catalogue and my research on polynemid fishes. I am particularly
grateful to Kent E. Carpenter (Department of Biological Sciences, Old Dominion University, Virginia) who read the initialmanuscript and offered helpful comments, Pere Oliver (Spanish Institute of Oceanography, Madrid), Michel Lamboeuf(FIRM: Species Identification and Data Programme, Marine Resources Service, Fisheries Resources Division, Food andAgriculture Organization of United Nations, Rome), Luca Garibaldi (FIDI: Fishery Statistician, Fishery Information, Data andStatistics Unit, FAO), Emanuela D’Antoni, Nicoletta De Angelis, Michèle Kautenberger-Longo (FIRM: Species Identification
and Data Programme, Marine Resources Service, Fisheries Resources Division, FAO) and other FAO staff whose
endeavours have made possible this catalogue, and Graham S. Hardy (Ngunguru, New Zealand).
I am grateful to the curators and technical staff of the museums that have loaned or donated specimens and radiographs andprovided me with literature or information on specimens. In particular, I thank Barbara Brown and Xenia Freilich (AMNH:
Department of Ichthyology, Division of Vertebrate Zoology, American Museum of Natural History, New York); Jeffrey M. Leis,
Thomas Trnski, Mark A. McGrouther, Kerryn L. Parkinson, John R. Paxton, John Pogonoski, Sally E. Reader, Amanda Hay,
Domine Clark, Matthew Lockett, Brooke M. Carson-Ewart, and Douglass F. Hoese (AMS: Ichthyology, Australian Museum,
Sydney); Mark H. Sabaj, Mike Littmann, and Dominique D. Dagit (ANSP: Department of Ichthyology, Academy of Natural
Sciences, Philadelphia); Kwang-Tsao Shao (ASIZP: Laboratory of Fish Ecology and Evolution, Institute of Zoology,
Academia Sinica, Taipei); James Maclaine [BMNH: Fish Section, Department of Zoology, British Museum (Natural History),
London]; John E. Randall and Arnold Suzumoto (BPBM: Ichthyology, Natural Sciences Department, Bishop Museum,
Honolulu); Kunio Sasaki, Yoshihiko Machida and Hiromitsu Endo (BSKU: Department of Biology, Faculty of Science, Kochi
University, Kochi); William N. Eschmeyer, Tomio Iwamoto, David Catania, and Jon Fong (CAS: Department of Ichthyology,
California Academy of Sciences, San Francisco); Teguh Peristiwady (CRDOA: Center for Research and Development of
Oceanology, Indonesian Institute of Science, Ambon); Alastair Graham, Peter R. Last, Gordon Yearsley and Daniel Gledhill(CSIRO: Division of Marine Research, Commonwealth Science and Industrial Research Organization, Hobart); Mary Anne
Rogers and Kevin Swagel (FMNH: Fish Division, Field Museum of Natural History, Chicago); Seishi Kimura (FRLM:
Fisheries Research Laboratory, Mie University, Mie); Hitoshi Ida and Makoto Okamoto (FSKU: School of Fisheries
Sciences, Kitasato University, Iwate); Kazuhiro Nakaya, Mamoru Yabe, and Hisashi Imamura (HUMZ: Laboratory of Marine
Biodiversity, Graduate School of Fisheries Sciences, Hokkaido University, Hakodate); Hiroshi Senou (KPM-NI: Kanagawa
Prefectural Museum of Natural History, Odawara); Kris Rhode and Andy Bentley (KU: Division of Ichthyology, Natural History
Museum and Biodiversity Research Center, University of Kansas, Lawrence); Karsten E. Hartel (MCZ: Ichthyology
Department, Museum of Comparative Zoology, Harvard University, Cambridge); Claude Weber (MHNG: Department of
Herpetology and Ichthyology, Museum of Natural History, Geneva); Patrice Pruvost and Jean-Claude Hureau, Nicolas Bailly
and Philippe Béarez (MNHN: Laboratoire d’Ichthyologie Générale et Appliquée, Muséum National d’Histoire Naturelle,
Paris); Yukio Iwatsuki (MUFS: Division of Fisheries Sciences, Faculty of Agriculture, Miyazaki University, Miyazaki); Martin
F. Gomon and Dianne Bray (NMV: Ichthyology, Museum of Victoria, Melbourne); Ernst Mikschi and Helmut Wellendorf(NMW: Fischsammlung, Naturhistorisches Museum Wien, Wien); Sven O. Kullander (NRM: Department of Vertebrate
Zoology, Swedish Museum of Natural History Stockholm); Keiichi Matsuura and Gento Shinohara (NSMT: Fishes Division,
Department of Zoology, National Science Museum, Tokyo); Helen K. Larson, Barry C. Russell, Gavin Dally, and Steven
Gregg (NTM: Natural Sciences, Museum and Art Gallery of the Northern Territory, Darwin); Ukkrit Satapoomin (PMBC:
Phuket Marine Biological Center, Phuket); Pornsilp Pholpunthin and Vachira Lheknim (PSUZC: Department of Biology,
Faculty of Science, Prince of Songkla University, Songkla); Jeffrey W. Johnson (QM: Ichthyology, Queensland Museum,
Brisbane); Martien J. P. van Oijen (RMNH: Rijksmuseum van Natuurlijke Histoire, Leiden); Phillip C. Heemstra and M. Eric
Anderson (SAIAB: South African Institute for Aquatic Biodiversity, formerly RUSI, Grahamstown); Michelle G. van der Merwe
and Liz Hoenson (SAM: Marine Biology, Life Sciences, South African Museum, Cape Town); Hang-Ling Wu (SFU: Shanghai
Fisheries University, Shanghai); Michael Gudo (SMF: Sektion Ichthyologie, Forschungsinstitut Senckenberg, Frankfurt);
Douglas W. Nelson (UMMZ: Division of Fishes, Museum of Zoology, University of Michigan, Michigan); Tetsuo Yoshino(URM: Department of Marine Sciences, Faculty of Science, University of the Ryukyus, Okinawa); Susan L. Jewett, Lisa
Palmer, Sandra J. Raredon, Shirleen Smith, and Jeffrey T. Williams (USNM: Division of Fishes, Department of Vertebrate
Zoology, Smithsonian Institute, National Museum of Natural History, Washington DC); Sue Morrison (WAM: Fish
Department, Western Australian Museum of Natural Science, Perth); Masayoshi Hayashi (YCM: Yokosuka City Museum,
Yokosuka); Isaäc J.H. Isbrücker (ZMA: Sectie Ichthyologie, Zoölogisch Museum Amsterdam, Universiteit van Amsterdam,
Amsterdam); Hans-Joachim Paepke (ZMB: Ichthyologischen Abteilung, Museum für Naturkunde, Humboldt-Universität,
Berlin); Horst Wilkens (ZMH: Zoologisches Institut und Zoologisches Museum der Universität Hamburg, Hamburg); Peter
K.L. Ng and Kelvin K.P. Lim (ZRC: Raffles Museum of Biodiversity Research, Department of Biological Sciences, National
University of Singapore, Singapore); A.K. Karmakar (ZSI: Fish Section, Zoological Survey of India, Calcutta);
K. Venkataraman (ZSI/MBS: Marine Biological Station, Zoological Survey of India, Chennai); P.T. Cherian and K. Rema Devi
(ZSI/SRS: Fish Section, Southern Regional Station, Zoological Survey of India, Chennai); Kazuo Sakamoto (ZUMT:University Museum, Department of Zoology, University of Tokyo, Tokyo); Ross M. Feltes (State University of New Jersey,
New Brunswick); Matthew Pember (Centre for Fish and Fisheries Research, Murdoch University, Murdoch); Reiji Masuda(Fisheries Research Station, Kyoto University, Maizuru).

Threadfins of the Worldv

I extend my gratitude to Shinji Tsukawaki and Takahiro Kamiya (Kanazawa University, Kanazawa), M.N. Venugopal andAnirban Chakraborty (Department of Fishery Biology, College of Fisheries, Mangalore), Arnucit Choochern (Bangkok),
B.V. Seshagiri Rao and B. Ratnamala (Department of Zoology, D.N.R. College, Bhimavaram), M. Iqbal Djawad (Faculty ofMarine and Fishery Sciences, Hasanuddin University, Makassar), Yasuwo Fukuyo and Mitsunori Iwataki (Asian Natural
Environmental Science Center, University of Tokyo, Tokyo), Muna Faraj, Mohammad Al-Sarawi and Mahmoud Al-Khabbaz(Living Resources Department, Environment Public Authority, Kuwait) for their assistance in field work at various times.
I am grateful for the financial support of the Food and Agriculture Organization of United Nations, the Research Fellowshipsof the Japan Society for the Promotion of Science for Young Scientists, the Fujiwara Natural History Foundation, theSasakawa Scientific Research Grant from the Japan Science Society, and the Ito Foundation for the Advancement of
Ichthyology.
Special thanks are due to Gerald R. Allen and Mark Allen (WAM), Pedro Niny Duarte (Department of Oceanography and
Fisheries, University of Azores, Azores), Phillip C. Heemstra, M. Eric Anderson, and James Stapley (SAIAB), Hitoshi Ida and
Makoto Okamoto (FSKU), Johnny Jensen (Kastrup, Denmark), Jeffrey W. Johnson (QM), Seishi Kimura (FRLM), Hiroshi
Kobayashi (Tokyo, Japan), Maurice Kottelat (Cornol, Switzerland), Keiichi Matsuura (NSMT), National Research Institute of
Far Seas Fisheries (NRIFSF, Shimizu, Japan); John E. Randall (BPBM), D. Ross Robertson (STRI: Smithsonian TropicalResearch Institute, Ancon, Panama), Mark H. Sabaj (ANSP), Hiroshi Senou (KPM-NI), Kwang-Tsao Shao (ASIZP), Erwin
Schraml (Augsburg, Germany) and Eiji Yamamoto (Fukuoka, Japan), who made available their colour transparencies ofpolynemid fishes, and to the illustrator, Emanuela D’Antoni (FIRM) who kindly prepared the drawings for this catalogue.
Finally, I thank Yuko Motomura who assisted in innumerable ways.

vi

FAO Species Catalogue for Fishery Purposes No. 3

Table of Contents
PREPARATION OF THIS DOCUMENT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iii
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . iv
1. INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1 Habitat and Biology . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.2 Fisheries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.3 Classification of the Family Polynemidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
1.4 Illustrated Glossary of Technical Terms and Measurements. . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1.5 Plan of the Catalogue . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
2. SYSTEMATIC CATALOGUE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.1 Diagnostic Features of the Family Polynemidae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.2 Notes on the Identification of Polynemids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2.3

Illustrated Key to the Genera of Polynemids . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10

2.4 Information by Genus and Species. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

Eleutheronema Bleeker, 1862 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Key to the Species of Eleutheronema . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

Eleutheronema rhadinum (Jordan and Evermann, 1902) . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Eleutheronema tetradactylum (Shaw, 1804) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Eleutheronema tridactylum (Bleeker, 1849) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Filimanus Myers, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Key to the Species of Filimanus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

Filimanus heptadactyla (Cuvier, 1829) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
Filimanus hexanema (Cuvier, 1829) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Filimanus perplexa Feltes, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25
Filimanus sealei (Jordan and Richardson, 1910) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Filimanus similis Feltes, 1991 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Filimanus xanthonema (Valenciennes, 1831) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Galeoides Günther, 1860. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Galeoides decadactylus (Bloch, 1795). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
Leptomelanosoma Motomura and Iwatsuki, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Leptomelanosoma indicum (Shaw, 1804) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Parapolynemus Feltes, 1993 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Parapolynemus verekeri (Saville-Kent, 1889). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Pentanemus Günther, 1860 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Pentanemus quinquarius (Linnaeus, 1758). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Polydactylus Lacepède, 1803 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Key to the Indo-Pacific Species of Polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40

Threadfins of the Worldvii

Key to the Eastern Pacific Species of Polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Key to the Atlantic Species of Polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43

Polydactylus approximans (Lay and Bennett, 1839) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Polydactylus bifurcus Motomura, Kimura and Iwatsuki, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . 46
Polydactylus longipes Motomura, Okamoto and Iwatsuki, 2001. . . . . . . . . . . . . . . . . . . . . . . . 48
Polydactylus macrochir (Günther, 1867) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Polydactylus macrophthalmus (Bleeker, 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Polydactylus malagasyensis Motomura and Iwatsuki, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . 54
Polydactylus microstomus (Bleeker, 1851) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Polydactylus mullani (Hora, 1926) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Polydactylus multiradiatus (Günther, 1860) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Polydactylus nigripinnis Munro, 1964 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Polydactylus octonemus (Girard, 1858) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Polydactylus oligodon (Günther, 1860) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Polydactylus opercularis (Gill, 1863) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Polydactylus persicus Motomura and Iwatsuki, 2001 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 66
Polydactylus plebeius (Broussonet, 1782) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Polydactylus quadrifilis (Cuvier, 1829) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Polydactylus sexfilis (Valenciennes, 1831) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Polydactylus sextarius (Bloch and Schneider, 1801) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Polydactylus siamensis Motomura, Iwatsuki and Yoshino, 2001. . . . . . . . . . . . . . . . . . . . . . . . 75
Polydactylus virginicus (Linnaeus, 1758) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Polynemus Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Key to the Species of Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79

Polynemus aquilonaris Motomura, 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
Polynemus dubius Bleeker, 1854 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Polynemus hornadayi Myers, 1936 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 84
Polynemus kapuasensis Motomura and van Oijen, 2003 . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Polynemus melanochir dulcis Motomura and Sabaj, 2002 . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Polynemus melanochir melanochir Valenciennes, 1831 . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Polynemus multifilis Temminck and Schlegel, 1843 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Polynemus paradiseus Linnaeus, 1758 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
3. LIST OF NOMINAL SPECIES OF POLYNEMIDAE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
4. LIST OF SPECIES BY MAJOR FISHING AREAS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97
5. BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
6. INDEX OF SCIENTIFIC AND VERNACULAR NAMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 109
7. LIST OF COLOUR PLATES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 117

Threadfins of the World1

1. INTRODUCTIONT

he family Polynemidae comprises 41 species in 8
genera (Fig. 1). In many parts of the tropics, the marine
polynemids, including Eleutheronema , Galeoides ,Leptomelanosoma, Pentanemus and Polydactylus, areimportant food fishes (some species attaining 2 m in total
length), with the largest quantities taken commercially.
However, separate catch statistics are not reported for most
species and landings are usually summarized as avernacular name which often corresponds to two or more
species. This lack of species-specific catch data is due tothe difficulty in identifying many of the species. For theproper management of this resource correct specificidentifications are necessary.
Polynemids are easily identified as a family due to theirsimilarity in overall body appearance, and condition of theeye and pectoral-fin rays. They have a conical snout, inferior
mouth, adipose eyelid (firm transparent gelatinous tissue)
covering the eye, 2 well-separated dorsal fins, 3 to 16
thread-like (individually-separated) lower pectoral-fin rays
(pectoral filaments) and the caudal fin deeply forked to
lunate. However, these external morphological similarities
have led to much confusion in specific identifications. The
pectoral filaments of polynemids are the most distinctive
feature, specific identifications to date having been based
mainly on the number and length of the former. Taxonomic
confusion in polynemids may, however, result fromidentifications based on pectoral filament s only. For
example, Polydactylus species characterized by a large
black anterior lateral-line spot have until recently been
considered to include 3 species, P. microstomus (5 pectoral
filaments), P. mullani (7 filaments) and P. sextarius (6filaments). The latter, however, an Indo-West Pacificspecies originally believed to range from South Africa to
Papua New Guinea, is now known to comprise 3 species:
P. malagasyensis (distributed off the east coast of Africa andMadagascar), P. persicus (Persian Gulf), and P. sextarius(India to Papua New Guinea), all with 6 pectoral filaments.
The purposes of this polynemid catalogue are to provide ameans of identifying the various species and to present a
synopsis of the literature on the biology and fisheries ofthese species. The genera Eleutheronema (3 species),
Galeoides (1 species) and Leptomelanosoma (1 species),
plus the Indo-Pacific Polydactylus (14 species) and eastern
Pacific Polydactylus (2 species) have been recently revisedby the author of this catalogue and colleagues (Motomura
and Iwatsuki, 2001a; Motomura, Iwatsuki and Kimura,
2001b; Motomura et al., 2002a; Motomura, 2002;
Motomura, Kimura and Iwatsuki, 2002) the results of thosestudies being incorporated here. Treatments of the genera
Filimanus (6 species) and Parapolynemus (1 species)
were greatly facilitated by the revisions of Feltes (1991,
1993). Taxonomic information given here for Polynemus (8species) and western Atlantic Polydactylus (3 species) was
taken from on-going revisions of those genera by the author,
and publications on the living marine resources of thewestern central Pacific and western central Atlantic (FAO
Species Identification Guides for Fisheries Purposes). The
taxonomic information provided for all of the polynemid
species covered here is based on the examination by theauthor of all known available type specimens and a large

number of non-type specimens, representing broad
geographical ranges of each species.

1.1 Habitat and Biology
Polynemids are epibenthic fishes found in the tropical andsubtropical waters of all oceans. Most species occur incoastal waters and estuaries, although some (e.g.
Polynemus species and Polydactylus macrophthalmus)
live entirely in fresh-water rivers. Polynemids generally
occur on sandy and muddy bottoms in depths less than
150 m, although juveniles are found in seagrass beds andtidepools.
As major predators of coastal and estuarine ecosystems,
most polynemids generally feed on a variety of fishes andcrustaceans. A few small species (e.g. Filimanus species)
have numerous gillrakers adapted for plankton feeding.
Young stages of most polynemids also feed on plankton,
some species later becoming piscivorous with growth. The
number of gillrakers tends to decrease with fish size in all of
the Eleutheronema species, those on the anterior parts ofboth the upper and lower limbs being replaced during fish
growth by tooth plates with villiform teeth. Furthermore,each gillraker on both the upper and lower limbs becomesshorter with fish growth, indicative of a change from plankton
feeding to piscivorous habits.
The pectoral fins of polynemids are their most distinctive
feature, being divided into an upper part with the rays joinedby membranes and a lower part with 3 to 16 separate rays
(pectoral filaments). The pectoral filaments have been
considered to operate as a sense organ, enabling
polynemids to search for food in muddy water (Motomura,
Sado and Kimura, 2002). In particular, Parapolynemus andPolynemus species, generally occurring in fresh-water
rivers and estuaries, have extremely long pectoral filamentsthat exceed their total length. Pectoral filaments are usually
spread forward to enable searching for prey while swimmingand probably act as a substitute for eyesight. An exception,
Leptomelanosoma indicum, also generally occur in muddy
water in large river mouths (e.g., Ganges and Chao Phraya
rivers) but does not have long pectoral filaments. Instead it
has an unique swimbladder with numerous appendages in 2
rows along the entire, approximate midlateral surface.
Because these appendages are fully inserted into the lateralwalls of the abdominal cavity, the tips being located near theouter layer of epidermal tissue, the swimbladder isconsidered to have the capacity for sensing water vibrations
caused by movements of prey. Underwater observations of
L. indicum indicate that it usually spreads the pectoral
filament s, but remains motionless on the bottom, abehaviour pattern unknown in other members of the family.
In addition to the apparent ability of L. indicum to sense
subtle water vibrations while themselves remaining
motionless, there exist s also the possibility that theswimbladder appendages transmit sounds and may enablecommunication with conspecific individuals. Specializationsof both the pectoral filament s and swimbladder inpolynemids have probably arisen as an adaptation to theirmuddy habitats.

2

FAO Species Catalogue for Fishery Purposes No. 3

Fig. 1 Classification of the family Polynemidae

Threadfins of the World
Judging from the few species that have been studied, mostpolynemids exhibit protandry, their sex changing from maleto female with fish growth. However, spawning behaviour ofpolynemids has never been reported. Although thespawning grounds of most polynemids are also unknown,
they probably occur in coastal waters and estuaries. In thenorthern Bay of Bengal, Polynemus paradiseus may ascend
higher up rivers than usual for breeding purposes (David,
1954).
Information on the larval development of polynemids is
minimal. Eggs are spherical and pelagic (de Sylva in Moser
et al., 1984). Data for Eleutheronema tetradactylum,
Polydactylus sexfilis, and Polynemus paradiseus suggestthat polynemids are 0.8 to 2.0 mm long at hatching, have alarge yolk sac, unpigmented eyes, an unformed mouth andpigment that may intensify during yolk absorption (Jonesand Menon, 1953; Kowtal, 1972; May, Akiyama andSanterre, 1979). Leis and Trnski in Leis and Carson-Ewart(2000) described larval developmental stages of
Eleutheronema tetradactylum and Polydactylus
multiradiatus from Australian waters (Figs 2 and 3), andsummarized larval morphology of Indo-Pacific polynemids:a) 2.5 mmb) 4.1 mm

3moderate to deep bodied, compressed; gut coiled andtriangular, reaching between 44 and 62% of body length; gas
bladder conspicuous, located near apex of gut mass; head
initially round, slightly elongated by 7 mm; mouth initiallyoblique, reaching to level of midpupil, becoming horizontal
and inferior, and extending beyond level of posterior marginof eye by about 6 mm; many small villiform teeth on upperand lower jaws at 2.5 mm; adipose eyelid not present until atleast 15 mm, but present by settlement stage; a very small
spine appearing at anterior tip of maxilla, disappearing
shortly thereafter (see Figs 2c and 3b); dorsal- and anal-fin
anlagen appearing in preflexion larvae from 3.0 to 4.1 mm,
soft rays developing during flexion; feeble spines of firstdorsal and anal fins starting to form in early post-flexion
larvae; pectoral-fin elements beginning to differentiate inlate flexion to early post-flexion larvae; pectoral-fin rays fully
ossified by 6.5 mm, at which time ossification of pectoral
filaments initiated; all pectoral-fin elements ossified shortly
after 7 mm; pectoral-fin base initially level with top of gut, but
beginning to move ventrally shortly after flexion, eventually
located near ventral margin of body by about 12 mm, except
in Parapolynemus and Polynemus species; scales
beginning to form laterally on trunk and tail from aboutc) 5.9 mmd) 7.3 mm

Fig. 2 Larval development of Eleutheronema tetradactylum (from Leis and Trnski, 2000)a) 3.0 mmc) 6.9 mmb) 5.0 mmd) 11.6 mm

Fig. 3 Larval development of Polydactylus multiradiatus (from Leis and Trnski, 2000)

4

FAO Species Catalogue for Fishery Purposes No. 3

12 mm; all scales present by 15 mm. There is no distinct
transformation to the adult form, development is direct andgradual. A small spine on the anterior tip of the maxilla is the
only apparent specialization for larval life.

Table 1
FAO Reported world catch of polynemids inmetric tonnes by area in 2001
Main Areas

1.2 FisheriesSpecies

Polynemids are of considerable importance in commercial
fisheries in tropical and subtropical seas worldwide (Tables 1to 3), and in the sport fishery off Australia and the west coastof Africa. A few s pecies ( e.g. Eleutheronema
tetradactylum and Polydactylus sexfilis) are used inaquaculture (Table 4). Fresh-water polynemids (e.g.
Polynemus aquilonaris and P. multifilis) are sometimes
used as aquarium fishes.
Polynemids contribute to the global fish catch, with about
93 000 t reported to FAO for 2001 (Table 1). FAO compiled
statistics (Fishstat Plus 2000), may however greatly
underestimate the total catch, because most polynemids arecaught by local fisheries in tropical regions, where collection
and reporting of detailed statistics at the species level
encounter difficulties and several countries still report theircatches by large groups of species. In these circumstances,the catch data presented by individual species items arelikely to be underestimated and an unknown proportion of
the catches for a species or group of species might havebeen reported by the national office under miscellaneous
fishes. In particular, the catch statistics of Eleutheronema
tetradactylum have been reported only from Kuwait,
Pakistan and Taiwan Province of China (Table 1), although
the species is widely distributed from the Persian Gulf toAustralia and Papua New Guinea, and is one of the mostimportant fisheries species for countries throughout thoseregions. The largest commercial catch of polynemids has
been reported from the Western Central Pacific (Table 2,
FAO Fishing Area 71). Most catch statistics reported from
fresh-water rivers in India (Table 2, FAO Fishing Area 4)probably involved Polynemus paradiseus, although catches
were recorded as Polynemidae spp. From 1970 to 2001,
there has been a general increase in landings of polynemids(Table 3).

Inland Atlantic Indian Pacific Grandwaters Ocean Ocean Ocean Total

Eleutheronema
tetradactylum

55

1 715

1/

1 770

Galeoides
decadactylus

17 10117 101

Pentanemus
quinquarius

3 947

3 947

Polydactylus
quadrifilis

12 447

12 447Polynemidae
spp.

66

2 258

15 89439 43457 652Grand Total

66

35 75315 94941 14992 917

1/

Catches of a Polynemidae species reported by Taiwan Province ofChina probably belong to Eleutheronema rhadinum (Motomura etal., 2002a) instead of E. tetradactylum.

1.3 Classification of the Family Polynemidae
The family Polynemidae have long been considered to berelated to the Mugilidae and Sphyraenidae, but there is noconvincing evidence to support that relationship (Johnson,1993). In fact, Johnson (1993) stated that evidence at thattime supported a sister-group hypothesis for Polynemidae
and Sciaenidae, suggesting that they might be included in asuperfamily (Polynemoidea). However, otolith evidence
does not support such a placement (Grove and Lavenberg,1997) and the taxonomic status and relationships of the
Polynemidae remain poorly known.

Table 2
FAO Reported world catch of polynemids in metric tonnes by FAO Fishing Areas in 2001
FAO Fishing Areas
Species
4

34

47

57

55

Eleutheronema tetradactylum

1/51Galeoides decadactylus

17 101

Pentanemus quinquarius

3 947

Polydactylus quadrifilis

12 44761
1 715

Polynemidae spp.

66

424

1 834

2 827

13 067

Grand Total

66

33 919

1 834

2 882

13 06771

87

39 4241039 424

10

1/

1 715

Catches of a Polynemidae species reported by Taiwan Province of China probably belong to Eleutheronema rhadinum(Motomura et al., 2002a) instead of E. tetradactylum.

196

329

Pentanemus quinquarius

2 070

151

49 4642 063

118

42 76267 136

Pentanemus quinquarius

Polydactylus quadrifilis

Polynemidae spp.

Total

1/

13 83814 322

Galeoides decadactylus

92 75850 953

9 453

1 515

14 47516 3621988

44 08435 356

100

5 205

3 423

1972

91 53767 2783872

10 09313 2911989

58 47849 384

100

3 355

5 639

1973

66 52641 990147

870

11 39412 1251990

67 24658 613

100

3 974

4 559

1974

76 07047 995

4 905

2 620

12 542

8 008

1991

59 96352 017

106

3 556

4 284

1975

88 58654 636

8 120

4 882

9 893

11 0551992

55 17943 131

910

5 938

5 200

1976

72 35047 024

2 401

4 634

7 228

11 0631993

62 20547 003

1 545

8 304

5 353

1977

72 55150 907

3 670

4 218

8 500

5 256

1994

72 24753 960

2 214

8 515

7 558

1978

79 14054 624

3 598

3 734

12 903

4 281

1995

65 74083 64251 837

7 359

3 005

17 768

3 673

1996

54 32237 129450450
47 897342

10 551

5 850

1980

290

10 451

6 652

1979

87 82950 76910 7053 356

18 385

4 614

1997

56 29438 605

377

432

9 031

7 849

1981

85 40253 842

10 0234 430

12 890

4 217

1998

65 56144 145281

483

12 366

8 286

1982

81 20853 098

10 7704 162

10 764

2 414

1999

61 97143 111451

438

11 630

6 341

1983

Catches of a Polynemidae species reported by Taiwan Province of China probably belong to Eleutheronema rhadinum (Motomura et al., 2002a) instead of E. tetradactylum

77 995

12 4727 871

Eleutheronema tetradactylum1/

1987

49 2031986

39 548

Total

Species

34 155Polynemidae spp.

42 030

4 218

3 624

Galeoides decadactylus

Polydactylus quadrifilis

2 759

1 440

Eleutheronema tetradactylum1/

1971

1970

Species

Table 3
FAO Reported world catch of polynemids in metric tonnes for the year 1970 to 2001

90 48453 707

12 237

3 612

13 948

6 980

2000

62 86643 104421

994

11 933

6 414

1984

92 91757 65212 4473 947

17 101

1 770

2001

65 05842 440749

973

13 312

7 584

1985

Threadfins of the World
5

6

FAO Species Catalogue for Fishery Purposes No. 3
Table 4
FAO Reported aquacultural catch of polynemids in metric tonnes for the years 1972 to 2001
Species

Country

Area 1992

Eleutheronema tetradactylum

Thailand71

Eleutheronema tetradactylum

Singapore

71

Taiwan (Province ofChina)

61

Eleutheronema tetradactylum

1/

1/

15

1993

1994

16

16

1995

1996

1997

1998

409

155

4

1999

2000

2001

3

1

2

16

38

16

5

Aquaculture production of a Polynemidae species reported by Taiwan Province of China probably belongs to Eleutheronoma rhadinum
instead of E. tetradactylum.

1.4 Illustrated Glossary of Technical Terms andMeasurements
Anal-fin base length - The straight-line distance from theanal-fin origin to the posterior basal end of the anal-fin base(Fig. 4).
Body depth - The vertical distance from first dorsal-fin origin
to ventral surface (Fig. 4).
Body width - The width (thickness) between pectoral-finbases.

Caudal fin - The tail fin (Fig. 4).
Caudal peduncle - The part of the tail joins the caudal fin to
the body.
Caudal-peduncle depth - The least depth of the peduncle(Fig. 4).
Caudal-peduncle length - The straight-line distance fromthe rear end of the anal-fin base to a vertical at the caudal-finbase (Fig. 4).

total length
standard length
1st dorsal fin

d 2 ndba o rsal
se -filen ngth

2nd dorsal fin
body depth

d
hea

len

gth

caudalpeduncle
depth

scales above lateral line

pectoral-fin length

pored lateral line
scales below
lateral line

caudal-peduncle
lengthcaudal fin
anal fin
pe

ct

or

pectoral-fin
base
al

-fi

la

m

pectoral fin
pelvic fin

en

tl

en

gt

h

ana

l-fi

n

e
b as

len

gth

pectoral-fin filamentFig. 4 External morphology and measurements

Threadfins of the World

7

Cranium - Bony part of the head, enclosing the brain. It is
composed of many different bones (Fig. 5).
lateral frontal
ethmoidsphenotic

parietal pterotic

nasal
epiotic
exoccipitalGillrakers - The long bony protuberances along the anterior(inner) edge of the gill arches (Fig. 8). Gillraker counts areoften given as 2 numbers (X + Y), where X is the number ofrakers on the upper limb of the first arch, and Y is the numberof rakers on the lower limb, with the raker at the angle(junction of upper and lower limbs) included in the lower-limb
count.
rudiments
upper-limb rakers
lower-limb rakers

supraoccipital

rudiments

ethmoid

Fig. 5 Dorsal view of cranium of Leptomelanosoma
indicum (from Motomura and Iwatsuki, 2001a)
Dorsal fin - A median fin along the back. Polynemids have 2
dorsal fins; the first is supported by spinous rays while thesecond has 1 spinous ray anteriorly and soft rays posteriorly(Fig. 4).
Ectopterygoids - The paired bones on each side of the roofof the mouth immediately posterior to the palatines (Fig. 6).
vomerbony tooth plates

gill filaments

Fig. 8 Gillrakers(after Heemstra and Randall, 1993)
Head length - The distance from the most distant end of theopercle to the anterior tip of the snout (Fig. 4).
Interorbital length - The least distance between the orbits.
Maxilla - The posterior, toothless bone of the upper jaw;
partly covered by the upper lip when the mouth is closed(Fig. 7).
Orbit diameter - The horizontal distance between the bonymargins of the eye (Fig. 7).

premaxilla
palatineectopterygoid

Fig. 6 Ventral view of dentition of premaxilla
and roof of oral cavity

Eye diameter - The horizontal distance between the fleshy
margins of the eye (Fig. 7).

Palatines - The paired bones on each side of the roof of themouth immediately posterior to the vomer (Fig. 6).
Pectoral fin - The fin on each side of the body immediately
behind the gill opening (Fig. 4). Pectoral-fin ray counts only
include rays interconnected by a membrane; the lower freerays are given separately.
Pectoral-fin base - The distance from the uppermost pointof the pectoral-fin base to the lowermost point of thepectoral-filament base (Fig. 4).
Pectoral-fin length - The distance from the posterior tip of
the pectoral-fin ray to the uppermost point of the pectoral-finbase (Fig. 4).
Pectoral filaments - The free rays below the pectoral-fin
rays (Fig. 4). Pectoral-filament counts are made from theanteriormost (ventralmost) element.

orbit
diametereyediameter

Pectoral-filament length - The distance from the posterior
tip of the pectoral filament to the base of the filament (Fig. 4).

ut
sno gthnelPelvic fin - Paired fins ventrally on the body behind thepectoral fins (Fig. 4).

preopercle
maxillaupper-jaw length

Fig. 7 Parts of head

Pored lateral-line scales - The series of pored scales thatrun from the upper end of the gill cavity to the caudal fin(Fig. 4).
Premaxilla - The paired, tooth-bearing bones of the upperjaw (Fig. 6).
Preopercle - The rounded or angular bone on the front partof the opercle; the posterior edge is exposed (Fig. 7).

8

FAO Species Catalogue for Fishery Purposes No. 3

Ray - The rigid or flexible struts that support the fins. Softrays are segmented and flexible. Spinous rays are
unsegmented, simple (unbranched) and pungent (stiff andsharp pointed). The number of spines are designated byroman numerals (I, II, III, IV . . .) and the number of soft rays
are designated by Arabic numerals (1, 2, 3, 4 . . .). The last
ray of the second dorsal and anal fins of polynemids is split
to its base and looks like 2 rays, but it is counted as a single
ray because it is supported by a single pterygiophore.
Scales above and below lateral line - A transverse seriesof scale rows; scales below lateral line are counted from theorigin of the anal fin in an oblique row to the lateral-line scale,but not including the lateral-line row; scales above lateralline are counted from the origin of the first dorsal fin in an
oblique row to the lateral-line scale, but not including thelateral-line row (Fig. 4).
Second dorsal-fin base length - The straight-line distancefrom the second dorsal-fin origin to the posterior basal end of
the second dorsal-fin base (Fig. 4).
Snout length - The distance from the anterior margin oforbit to the anterior tip of the snout (Fig. 7).

Vertebrae - Axial skeleton bone; divided into 2 sections,precaudal (abdominal) and caudal vertebrae.
Vomer - A median bone which lies on the roof of the mouth(Fig. 6).

1.5 Plan of the Catalogue
A diagnosis of the family Polynemidae is given, followed byaids to identification of polynemids (keys to genera andspecies), and the information by genus and species(arranged alphabetically by genus and species). Since most
species occur only in a single major region, the key tospecies of the largest genus, Polydactylus, is divided into
regional keys to shorten them and make them easier to use.
Each genus account includes the type species and genericsynonyms. For genera with more than 1 species, thediagnosis and general comments on the habitat, biology,
distribution and interest to fisheries of the included species
are listed. The information pertaining to species is divided
into the following sections:(1)

Scientific Name: The reference to the originaldescription, type locality, and type information of thespecies is given. Type information includes themuseum catalogue number of the type specimen(abbreviations for m useums defined in‘Acknowledgements’).

(2)

Synonyms: Primary synonyms and alternate
combinations of generic and specific names are listed.

(3)

FAO Name: The FAO English name is considered thestandard to be used for fishery purposes. This should
avoid confusion caused by the existence of multiple
names for the same species or the same name forseveral species. The FAO name is not intended tosupplant the use of local names, but rather to serve asa worldwide reference. FAO French and Spanish
names are given for each species.

(4)

Diagnostic Features: Distinctive characters of thespecies are given as an aid for identification. These
diagnoses should be consulted to confirm species
identified using the illustrated key.

(5)

Geographical Distribution: The general geographic
range is given and illustrated on a map. The map shading
includes known areas of occurrence and intermediateareas where a species is expected to be found.

(6)

Habitat and Biology: Information on habitat, feeding
preferences, reproduction and behaviour is given.

(7)

Size: The maximum known size for each species.

(8)

Interest to Fisheries: General information on theextent, type of fisheries and utilization is given.
Detailed catch statistics (landings) are not available forall individual species.

(9)

Local Names: Published local names are given. A
local name is often applied to several species.

Standard length - The straight-line distance from the tip of
the snout to a vertical line passing through the base of thecaudal fin (taken to be the point of flexure of the caudal fin)
(Fig. 4).
Supraneural bones - Unpaired bones above the neuralspines of anterior vertebrae between the skull and the originof the dorsal fin (Fig. 9).
dorsal-fin spines

supraneural bones

neuralspines

1st proximal
pterygiophore

Fig. 9 Supraneural bones, anterior neural spines andfirst proximal pterygiophore
Swimbladder - A gas filled sac in the dorsal part of the body
cavity.
Synonym - An alternate name given to a particular species,
genus, family etc. The oldest (senior) synonym that is given
to a species is usually considered the valid (correct)
scientific name for that species.
Total length - The straight-line distance from the tip of thesnout to a vertical line passing through the posterior tip of thecaudal-fin ray (Fig. 4).

(10) Literature: Recent references that contain importanttaxonomic information are given.

Upper-jaw length - The distance from the anterior tip of thepremaxilla to the posterior margin of the maxilla (Fig. 7).

(11) Remarks: Taxonomic details and information that is
not appropriate in the previous paragraphs areincluded here.

Threadfins of the World

9

2. SYSTEMATIC CATALOGUE

2.1 Diagnostic Features of the Family Polynemidae
FAO Names: En - Threadfins; Fr - Barbures; Sp - Barbudos.
Diagnostic Features: Body elongate to moderately deep, compressed. Snout obtusely conical, overhanging. Adipose
eyelid (firm transparent gelatinous tissue) covering eye; 5 infraorbitals (3 in Parapolynemus). Mouth ventral, near-horizontal
and large; lip on upper jaw absent or poorly developed; maxilla extending beyond level of posterior margin of eye;supramaxilla absent; teeth villiform in broad bands on jaws, vomer, palatines and ectopterygoids (vomerine teeth absent insome species); canine, molariform or incisiform teeth absent. Posterior margin of preopercle serrated (without serrations in
Parapolynemus and Pentanemus); 7 branchiostegal rays, 1 ray present on epihyal. Two well-separated dorsal fins; firstdorsal fin with VIII spines (VII spines in 2 Polynemus species); second dorsal fin with I spine and 11 to 18 soft rays; anal fin
with III spines and 10 to 18 soft rays (II spines in 2 Polynemus species and 24 to 30 soft rays in Pentanemus; the last dorsaland anal-fin soft rays usually split to their base but counted as a single ray); pectoral fins divided into an upper part with 12 to
19 rays joined by membrane and a lower part with 3 to 11 separate rays (pectoral filaments) (14 to 16 separate rays in 1
Polynemus species); pelvic fin with I spine and 5 soft rays; scaly process (axillary scale) present at base of pelvic fin; caudal
fin deeply forked; small scales covering most of dorsal, pectoral, anal and caudal fins; trisegmental pterygiophores absent.
Scales weakly ctenoid, extending onto head; lateral line simple, extending from upper end of gill opening to posterior marginof caudal-fin membrane (lateral line bifurcating on caudal-fin base in 5 Polydactylus species and divided into 3 on caudal-finbase and membrane in 2 Eleutheronema species). Vertebrae 10 precaudal and 14 caudal (15 caudal in Eleutheronema
and Polynemus); supraneural bones 0 to 3.
The family Polynemidae, as here constituted, comprises 8 genera: Eleutheronema , Filimanus , Galeoides ,Leptomelanosoma, Parapolynemus, Pentanemus, Polydactylus and Polynemus.

2.2 Notes on the Identification of Polynemids
Because of the similarities in morphology and colour pattern of many polynemid species, correct identifications are often
difficult. Unlike typical reef fishes, most polynemid fishes are not easily identified on the basis of live or fresh colours,
although a few species have a distinct large black spot on the lateral body surface (e.g. Polydactylus malagasyensis,
P. mullani and P. sextarius) and several dark stripes along the longitudinal scale rows above and below the lateral line (e.g.
Polydactylus bifurcus and P. plebeius). Live coloration of most polynemids, except e.g. Polydactylus plebeius andP. sexfilis, has never been reported and the fresh colour of several species (e.g. Eleutheronema tridactylum andP. siamensis) is also still unknown. Although the number and length of pectoral filaments is distinctive enough for
identification of a few species, most polynemids can only be identified by a combination of morphological and meristic
characters, such as fin-ray scale and gill-raker counts, and proportional measurements. The keys provided in this catalogue
are based as much as possible on simple morphological and meristic characters, and proportional measurements.
Fin-ray counts are usually done with a bright light shining through the fin, which is fleshy and scaly in most polynemids.
Pectoral-fin counts are more easily made on the inner side of the fin. The last dorsal- and anal-fin rays are split to their basebut counted as a single ray. Pored lateral-line scale counts are facilitated by lying the fish on its right side (head pointing left)and directing a bright light at a low angle to the body surface. They may be assisted by using cyanin blue solution in 70%
ethyl alcohol.

10
2.3

FAO Species Catalogue for Fishery Purposes No. 3
Illustrated Key to the Genera of Polynemids

1a. Pectoral-fin insertion well below midline of body; eye diameter variable, 1.3 or less in snout length(Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Pectoral-fin insertion near midline of body; eye diameter 1.3 or more in snout length (Fig. 11) . . . . . . . . ® 7
eye diameter1.3 or less in
snout lengtheye diameter 1.3 ormore in snout length

pectoral-fin insertion well
below midline of body

pectoral-fin insertion near
midline of bodyFig. 10 Polydactylus bifurcus

Fig. 11 Polynemus paradiseus

2a. Anterior part of lower jaw with small teeth extending onto lateral surface, adjacent portion of lip absent(Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eleutheronema(Persian Gulf to Japan and Australia)

2b. Lip present (well or poorly developed) on anterior part of lower jaw (Fig. 13) . . . . . . . . . . . . . . . . . ® 3aadipose eyelid

maxillalipbmaxillalip

Fig. 13 Left lateral view of upper and lower
jaws of a) Leptomelanosoma indicum andb) Polydactylus virginicus(after Motomura and Iwatsuki, 2001a)Fig. 12 Left lateral view of upper and lower
jaws of Eleutheronema tetradactylum(after Motomura et al., 2002a)3a. Anal-fin base length greater than head length; anal-fin rays 24 to 30 (Fig. 14) . . . . . . . . . . . . Pentanemus(west coast of Africa)

3b. Anal-fin base length less than head length; anal-fin rays 10 to 18 (Fig. 15) . . . . . . . . . . . . . . . . . . ® 4anal-fin base length greater than head length

he

a

en
dl

anal-fin base length less than head length

gth
he

l
ad

en

gth

10-18 anal-fin
soft rays

anal-fin base length24-30 anal-fin
soft rays

Fig. 14 Pentanemus quinquarius

anal-fin base length

Fig. 15 Polydactylus bifurcus

Threadfins of the World

11

4a. Pectoral-fin base length (including pectoral-filament base) greater than or equal to upper-jaw length(Fig. 16); lateral line simple, extending to lower end of upper caudal-fin lobe (Fig. 17); swimbladderpresent, extending beyond anal-fin origin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Galeoides(west coast of Africa)
4b. Pectoral-fin base length (including pectoral-filament base) less than upper-jaw length (Fig. 18); lateralline simple, extending to upper end of lower caudal-fin lobe or middistal margin of caudal fin, or bifurcated on caudal-fin base, extending to posterior margins of upper and lower caudal-fin lobes (Fig. 19);
swimbladder absent, or if present, not extending beyond anal-fin origin . . . . . . . . . . . . . . . . . . . . ® 5pectoral-fin base greater than or equal to upper-jaw lengthcaudal fin

uppe
len r-jawgthlateral line

pectoral-fin base (including pectoral-filament base) length

Fig. 16 Galeoides decadactylus

pectoral-fin base less than upper-jaw length

uppe
len r-jaw
gthFig. 17 Lateral-line squamation on caudal-fin
membrane of Galeoides decadactylus

caudal fin

lateral line
pectoral-fin base (including pectoral-filament base) length

Fig. 18 Polydactylus bifurcus

a)

b)

c)

Fig. 19 Lateral-line squamation patterns on
caudal-fin membrane of a) Filimanus perplexa,b) Polydactylus bifurcus and c) Polydactylus plebeius

5a. Space separating premaxillary teeth bands 2 or more times width of each band (Fig. 20); basisphenoid
not in contact with prootic (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filimanus(Pakistan to Solomon Islands)
5b. Space separating premaxillary teeth bands less than width of each band (Fig. 22); basisphenoid in contact with prootic (Fig. 23) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6

premaxilla

premaxilla

basisphenoidbasisphenoid

prootic

prootic

Fig. 20 Ventral view ofdentition of premaxilla
and roof of oral cavityof Filimanus

Fig. 21 Lateral view of
cranium of Filimanus(after Feltes, 2001)

Fig. 22 Ventral view ofdentition of premaxilla
and roof of oral cavity of
Polydactylus virginicus

Fig. 23 Lateral view of
cranium of Polydactylus(after Feltes, 2001)

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FAO Species Catalogue for Fishery Purposes No. 3

6a. Tips of upper and lower caudal-fin lobes
filamentous (Fig. 24); sphenotics exposed
dorsally between anterior margins of parietal
and pterotic (Fig. 25); swimbladder withmany appendages inserted into lateral wallsof abdominal cavity (Fig. 26) . . . Leptomelanosoma(Pakistan to Indonesia)
6b. Tips of upper and lower caudal-fin lobes notfilamentous (Fig. 27); sphenotics not exposed dorsally (Fig. 28); swimbladder absent
or present (without appendages) . . . . Polydactylus

tips of upper and lower caudal-fin
lobes filamentous(worldwide)

sphenotic

parietalFig. 24 Leptomelanosoma indicum

pteroic

anterior

posteriorFig. 26 Dorsal view of swimbladder of
Leptomelanosoma indicum

Fig. 25 Dorsal view of cranium of
Leptomelanosoma indicum(from Motomura and Iwatsuki, 2001a)parietalpteroic

tips of upperand lower
caudal-fin lobes
not filamentous

Fig. 27 Polydactylus bifurcus

Fig. 28 Dorsal view of cranium of
Polydactylus virginicus(from Motomura and Iwatsuki, 2001a)

7a. Posterior margin of preopercle serrated(Fig. 29) . . . . . . . . . . . . . . . Polynemus

preopercle
serratedpreopercle
unserrated(India to Viet Nam and Indonesia)
7b. P o s t e r i o r m a r g i n o f p r e o p e r c l eunserrated (Fig. 30) . . . . . . Parapolynemus(southern Papua New Guinea and northern Australia)

Fig. 29 Head of
Polynemus paradiseus

Fig. 30 Head of
Parapolynemus verekeri

Threadfins of the World13

2.4 Information by Genus and Species

Eleutheronema Bleeker, 1862
Eleutheronema Bleeker, 1862a: 110 (type species: Polynemus tetradactylus Shaw, 1804).
Synonyms: None.
Diagnostic Features: Body and head elongate. Adipose eyelid well developed; eye diameter greater than snout length.
Anterior parts of lower jaw with small teeth extending onto lateral surface, adjacent portion of lip absent; width of tooth bandon upper and lower jaws greater than space (on symphysis) separating tooth bands on opposing premaxillae; teeth villiform
in broad bands on jaws, vomer, palatines and ectopterygoids, tooth plates on palatines shorter than those on ectopterygoids.
Posterior margin of maxilla extending well beyond level of posterior margin of adipose eyelid. Posterior margin of preopercle
serrated. Basisphenoid in contact with prootic; sphenotics not visible dorsally between anterior margins of parietal andpterotic. First dorsal fin with VIII spines; second dorsal fin with I spine and 13 to 15 soft rays; anal fin with III spines and 14 to
16 soft rays; anal-fin base less than head length; pectoral fin with 15 to 19 rays, all unbranched; pectoral-fin insertion well
below midline of body; pectoral-fin base (including base of pectoral filaments) less than upper-jaw length; pectoral filaments
3 or 4, not extending beyond level of posterior tip of pelvic fin; caudal fin deeply forked, but upper and lower caudal-fin lobes
not filamentous. Pored lateral-line scales 71 to 95; scale rows above lateral line 8 to 14, below 12 to 17. Gillrakers 2 to 8 on
upper limb, 2 to 10 on lower limb, 4 to 18 total; gillrakers on anterior parts of upper and lower limbs replaced during fish
growth by tooth plates with villiform teeth, each gillraker on both upper and lower limbs becoming shorter with fish growth.
Vertebrae 10 precaudal and 15 caudal; supraneural bones 0 to 2. Swimbladder absent.
Habitat and Biology: With the exception of E. tetradactylum, no information on habitat and biology for members of thegenus has been reported, although the genus comprises 3 species. Information on E. tetradactylum is given below under
the species’ account.
Geographic Distribution: The genus is distributed in the Indo-West Pacific, but has never been recorded in the Red Sea oroff the east coast of Africa (including Madagascar).
Interest to Fisheries: The species of Eleutheronema, except E. tridactylum, are among the most important commercial
fishes in the Indo-West Pacific, being the highest priced fishes in local fish markets. Eleutheronema tridactylum is of little
commercial importance because of its small size.
Species: The genus comprises 3 species:

Eleutheronema rhadinum: East Asia, Japan, China and Viet Nam.
Eleutheronema tetradactylum: Indo-West Pacific, Persian Gulf to Australia.
Eleutheronema tridactylum: Southeast Asia, Thailand to Indonesia.
Remarks: Bleeker (1862a) proposed Eleutheronema (name only) for Polynemus tetradactylus, but gave neither
diagnoses nor descriptions of the genus. Subsequently, Bleeker (1862b) described the genus in detail, including thefollowing characters: body oblong, compressed; scales very small (around 70 in lateral line); lip absent except in corner oflower jaw; vomer, palatines and ectopterygoids with teeth; posterior margin of preopercle serrated; pectoral filaments 3 or 4;anal-fin rays 15 to 17. Two species, Polynemus tetradactylus and Polynemus tridactylus, were included by Bleeker(1862b).
Feltes (1993) redefined the genera of the family Polynemidae, including a key, and suggested that the posterior extension of
tooth plates from the lateral margins of the primary vomerine tooth plate was diagnostic for Eleutheronema. However, thischaracter is not found in E. tridactylum at any life stage and in juveniles (less than about 70 mm standard length) of
E. tetradactylum. Therefore, the character should be not treated as diagnostic for the genus. The genus was recently
redefined by Motomura et al. (2002a).

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FAO Species Catalogue for Fishery Purposes No. 3

Key to the Species of Eleutheronema
1a. Pectoral filaments 3; vomer without tooth plates on either side throughout life (Fig. 31a); second
dorsal-fin soft rays 13 (rarely 14); gillrakers 4 to 10 (mode 8) (Fig. 32, Plate Ic) . . . Eleutheronema tridactylum(Thailand to Indonesia)
1b. Pectoral filaments 4; vomer with deciduous tooth plates on both sides (Fig. 31b), except in juveniles(less than about 70 mm standard length); second dorsal-fin soft rays 14 (rarely 13 or 15); gillrakers 6 to
18 (mode 12 or 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
vomer
13 (rarely 14) 2nd
dorsal-fin soft rays

deciduous
toothplatesa)b)

Fig. 31 Ventral view of dentition of premaxilla androof of oral cavity of a) E. tridactylum
and b) E. tetradactylum(from Motomura et al., 2002a)

4-10
gillrakers

3 pectoral filaments

Fig. 32 Eleutheronema tridactylum

2a. Pored lateral-line scales 82 to 95; 11 to 14 (mode 12) scale rows above lateral line, 15 to 17 (16) below;
pectoral-fin membranes black when fresh (Fig. 33, Plate Ia) . . . . . . . . . . . . . . Eleutheronema rhadinum(Japan, China and Viet Nam)
2b. Pored lateral-line scales 71 to 80; 9 to 12 (mode 10) scale rows above lateral line, 13 to 15 (14) below;
pectoral-fin membranes vivid yellow when fresh, except in large specimens (over about 350 mm standard length) (Fig. 34, Plate Ib) . . . . . . . . . . . . . . . . . . . . . . . . . . . Eleutheronema tetradactylum(Persian Gulf to Australia)
11-14 scale rows
above lateral line

pectoral fin black

82-95 pored
lateral-line scales

15-17 scale rows
below lateral line

Fig. 33 Eleutheronema rhadinum

9-12 scale rows
above lateral line

pectoral fin vivid yellow

71-80 pored
lateral-line scales

13-15 scale rows
below lateral line

Fig. 34 Eleutheronema tetradactylum

Threadfins of the World

15

Eleutheronema rhadinum (Jordan and Evermann, 1902)

Fig. 35; Plate Ia

Polydactylus rhadinus Jordan and Evermann, 1902: 351, fig. 20 [type locality: Taiwan Province, China; neotype (ASIZP
60745, 152 mm standard length) designated by Motomura et al., 2002a].
Synonyms: None.
FAO Names: En - East Asian fourfinger threadfin; Fr - Barbure à quatre doigts d'Asie; Sp - Barbudo cuatro dedos de Asia.

Fig. 35 Eleutheronema rhadinum

Diagnostic Features: A large species. Body elongate, body depth at first dorsal-fin origin 23 to 27% (mean 24%) of
standard length; head length 28 to 30% (mean 29%) of standard length. Posterior margin of maxilla extending well beyond
level of posterior margin of adipose eyelid; upper-jaw length 15 to 16% (mean 16%) of standard length; depth of posterior
margin of maxilla 3% of standard length; anterior parts of lower jaw with small teeth extending onto lateral surface, adjacentportion of lip absent; tooth plate extension onto lateral surface of lower jaw 8 to 9% (mean 8%) of standard length; teeth
villiform in broad bands on vomer, palatines and ectopterygoids; vomer with deciduous tooth plates on both sides;
ectopterygoids and 2 deciduous tooth plates becoming wider and larger, respectively, with continued fish growth. Posterior
margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness (base of second spine
slightly more robust than others in large specimens); second dorsal fin with I spine and 14 (rarely 13 or 15) soft rays; anal fin
with III spines and 14 to 16 (mode 15) soft rays, anal-fin base longer than second dorsal-fin base; pectoral fin with 17 or 18(mode 18, rarely 19) unbranched rays, its length 20 to 22% (mean 21%) of standard length, posterior tip just short of level ofposterior tip of pelvic fin; pectoral filaments 4, first filament shortest, not reaching to level of pelvic-fin origin; second filament
extending slightly (or well) beyond level of pelvic-fin origin; third filament extending beyond (or just short of) level of pelvic-fin
origin; fourth filament longest, its length 15 to 27% (mean 21%) of standard length, not reaching to level of posterior tip of
pectoral fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 32 to 38%(mean 35%) and lower lobe 29 to 36% (mean 33%) of standard length. Pored lateral-line scales 82 to 95 (mode 95); lateralline simple, extending from upper end of gill opening to upper end of lower caudal-fin lobe, or bifurcating on caudal-fin base,upper branch extending to lower end of upper caudal-fin lobe and lower branch secondarily bifurcating on middle of lower
caudal-fin lobe; scale rows above lateral line 11 to 14 (mode 12), below 15 to 17 (mode 16). Gillrakers 5 to 8 (mode 5) on
upper limb, 5 to 9 (mode 7) on lower limb, 10 to 17 (mode 12) total; gillrakers on anterior parts of upper and lower limbs
replaced during fish growth by tooth plates with villiform teeth, each gillraker on both upper and lower limbs becoming shorter
with fish growth. Vertebrae 10 precaudal and 15 caudal; supraneural bones 1 to 2. Swimbladder absent. Colour: Uppersides of head and trunk with slight darkish silver tinge, becoming lighter on lower sides; anterior margins of first and second
dorsal fins blackish, remaining parts translucent and slightly blackish, respectively (both fins uniform dense black in largefish); pectoral fin dense black; pectoral filaments white; pelvic fin white; base of anal fin slightly yellowish, other parts white(pelvic fin uniformly black in large fish); base and posterior margin of caudal fin yellowish and dense black, respectively, otherparts blackish.

16

FAO Species Catalogue for Fishery Purposes No. 3

Geographical Distribution: Distributed in East Asia
(Japan, China and Viet Nam), being endemic to thatarea (Fig. 36). The species generally occurs on
continental shelves (Chinese mainland, Taiwan
Province of China, Japanese mainland), having at notime been reported from the vicinity of oceanic islands(e.g. Ryukyu Islands and Ogasawara Islands). The
record from Japan is based only on a single large
specimen (MUFS 18880, 923 mm total length) from
Aomori, northernmost Honshu Island.
Habitat and Biology: No data are available.
Size: Maximum standard length at least 739 cm(Motomura, Senou and Iwatsuki, 2001; Motomura
et al., 2002a).
200 m

Interest to Fisheries: Reliable fishery information forthe species is not available.
Local Names: CHINA: Shi-jih-ma-fa-yu; JAPAN:
Minami-konoshiro; TAIWAN PROVINCE OF CHINA:
Shi-jih-ma-fa-yu; Gor-ah.

Fig. 36 Eleutheronema rhadinum
Known distribution

Literature: Motomura, Senou and Iwatsuki (2001, as Eleutheronema tetradactylum); Motomura et al., 2002a; Motomura(2003b).
Remarks: Polydactylus rhadinus was described by Jordan and Evermann (1902) from Taiwan Province of China, China, on
the basis of a single specimen (273 mm total length) which has apparently been lost [a neotype was designated by Motomura
et al. (2002a)]. Since the original description, the species has been treated as a junior synonym of Eleutheronema
tetradactylum (e.g. Weber and de Beaufort, 1922; Herre, 1953; Kagwade, 1970). However, E. rhadinum is clearly
distinguished from the latter by the number of pored lateral-line scales (82 to 95 versus 71 to 80 in the latter) and coloration of
pectoral-fin membranes (black versus vivid yellow when fresh), although proportional measurements of the 2 species are
similar. Accordingly, E. rhadinum was recently regarded as a valid species by Motomura et al. (2002a).

Eleutheronema rhadinum differs from Eleutheronema tridactylum by having vomer with 2 deciduous tooth plates in
specimens at least over ca. 70 mm standard length (vs. vomer without tooth plates in the latter) and higher counts of second
dorsal-fin soft rays [14 (rarely 13 or 15, 3 of 19 specimens) versus 13 (rarely 14, 1 of 34 specimens)], pectoral filaments (4versus 3) and gillrakers [10 to 17 (mode 12) versus 4 to 10 (mode 8)]. Furthermore, E. rhadinum tends to have a slightly
deeper posterior margin of the maxilla (3% of standard length) and a slightly shorter tooth plate on the lateral surface of thelower jaw [8 to 9% (mean 8%) of standard length] than E. tridactylum [2 to 3% (mean 2%) and 9 to 10% (mean 9%) of
standard length, respectively], although the proportional length measurements overlapped among the 2 species.

Eleutheronema tetradactylum (Shaw, 1804)

Fig. 37; Plate Ib

Polynemus tetradactylus Shaw, 1804: 155 [original locality: Vizagapatnam, India, based on “Maga Jellee” of Russell, 1803:
67, fig. 183; type locality: Gariahat, Calcutta, India, based on a neotype (NSMT-P 60912, 189 mm standard length)
designated by Motomura et al., 2002a].
Synonyms: Polynemus teria Hamilton, 1822: 224 (type locality: estuaries of Ganges River, India; no types known, seeMotomura et al., 2002a). Polynemus coecus Macleay, 1878: 354, pl. 9, fig. 1 [type locality: Darwin, Northern Territory,
Australia; 2 syntypes (AMS I. 9791, 369 mm standard length; AMS I. 16295-001, 360 mm standard length)].

Threadfins of the World

17

FAO Names: En - Fourfinger threadfin; Fr - Barbure à quatre doigts; Sp - Barbudo de cuatro dedos.

Fig. 37 Eleutheronema tetradactylum
Diagnostic Features: A large species. Body elongate, body depth at first dorsal-fin origin 23 to 27% (mean 25%) of
standard length; head length 28 to 31% (mean 29%) of standard length. Posterior margin of maxilla extending well beyond
level of posterior margin of adipose eyelid; upper-jaw length 14 to 17% (mean 15%) of standard length; depth of posterior
margin of maxilla 3 to 4% (mean 3%) of standard length; anterior parts of lower jaw with small teeth extending onto lateralsurface, adjacent portion of lip absent; tooth plate extension onto lateral surface of lower jaw 7 to 9% (mean 8%) of standard
length; teeth villiform in broad bands on vomer, palatines, and ectopterygoids; vomer with deciduous tooth plates on bothsides, except in juveniles (less than about 70 mm standard length); ectopterygoids and 2 deciduous tooth plates becoming
wider and larger, respectively, with continued fish growth. Posterior margin of preopercle serrated. First dorsal fin with VIII
spines, all spine bases of similar thickness (second spine slightly more robust than others in large specimens, over about
300 mm standard length); second dorsal fin with I spine and 14 (rarely 13 or 15) soft rays; anal fin with III spines and 14 to 16(mode 15) soft rays, anal-fin base longer than second dorsal-fin base; pectoral fin with 16 or 18 (mode 17, rarely 15 or 19)
unbranched rays, its length 20 to 23% (mean 21%) of standard length, posterior tip just short of level of posterior tip of pelvic
fin; pectoral filaments 4, first filament shortest, not reaching to level of pelvic-fin origin; second filament extending slightly (orwell) beyond level of pelvic-fin origin; third filament (rarely longest) extending beyond level of pelvic-fin origin; fourth filament
longest, its length 15 to 28% (mean 24%) of standard length, reaching to (not reaching or extending slightly beyond) level ofposterior tip of pectoral fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe
30 to 38% (mean 34%) and lower lobe 26 to 36% (mean 32%) of standard length. Pored lateral-line scales 71 to 80 (mode73); lateral line simple, extending from upper end of gill opening to upper end of lower caudal-fin lobe, or bifurcating on
caudal-fin base, upper branch extending to lower end of upper caudal-fin lobe and lower branch secondarily bifurcating on
middle of lower caudal-fin lobe; scale rows above lateral line 9 to 12 (mode 10), below 13 to 15 (mode 14). Gillrakers 3 to 8
(mode 6) on upper limb, 3 to 10 (mode 7) on lower limb, 6 to 18 (mode 13) total; gillrakers on anterior parts of upper and lower
limbs replaced during fish growth by tooth plates withvilliform teeth, each gillraker on both upper and lower
limbs becoming shorter with fish growth. Vertebrae 10
precaudal and 15 caudal; supraneural bones 1 to 2.
Swimbladder absent. Colour: Upper sides of head
and trunk with slight darkish silver tinge, becoming
lighter on lower sides; anterior margins of first andsecond dorsal fins blackish, remaining part stranslucent and slightly blackish, respectively; pectoralfin vivid yellow (dusky yellow in large specimens overabout 350 mm standard length); pectoral filaments
white; anterior margin of pelvic fin yellow, other parts
white; base of anal fin yellow, other parts yellowish
white; base of caudal fin yellowish, other part s
blackish.
Geographical Distribution: Distributed in the
Indo-West Pacific where it ranges from the Persian
Gulf to Papua New Guinea and northern Australia(from mouth of Ashburton River, Western Australia tomouth of Mary River, Queensland) (Fig. 38). The
species is not known from East Asia (where it isreplaced by the endemic E. rhadinum).

200 m

Fig. 38 Eleutheronema tetradactylum
Known distribution

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FAO Species Catalogue for Fishery Purposes No. 3

Habitat and Biology: Generally occurs on continental shelves on muddy and sandy substrata, and frequently enters
brackish waters, especially as juveniles. According to Malhotra (1953), larvae from 7 to 30 mm total length feed mainly on
copepods and mysids, shrimps and prawn larvae having secondary importance as prey. Juveniles (31 to 60 mm total length)
feeds mainly on prawns, shrimps and mysids, and adult fish prey on other fishes, including mugilids, sillaginids andsciaenids, and sometimes smaller conspecifics.
Patnaik (1967) referred to the occurrence of hermaphroditism in 2 examples (404 and 405 mm total length) of the species
collected from Chilka Lake, India, and Stanger (1974) studied sexuality and growth of the species in Australian waters.
Eleutheronema tetradactylum grows very rapidly during the first 6 months. At 1 year old, the species averages 300 mm fork
length, and at 3 years old, 450 mm fork length. Males range from 240 to 470 mm fork length, hermaphrodites from 250 to
460 mm fork length, and females from 280 to 720 mm fork length. On the northeast coast of Queensland, Australia, most
specimens reported (from 450 to 500 mm fork length) were females. Hermaphroditism develops in 1- to 2-year-old fishes,
females first appearing as 2- to 3-year-olds. According to Stanger (1974), males probably commence sex reversal
immediately after spawning, the condition possibly persisting until after the next spawning period. The progression fromhermaphrodite to female is completed by the following season.
In waters off Townsville, Queensland, Australia, spawning begins in late October (Stanger, 1974), specimens collected fromthe northeast coast of Queensland in November and December having either recently spawned or being ready to spawn(Kailola and Stewart in Kailola et al., 1993). Although the species spawns only once each year in Queensland waters(Stanger, 1974), in the northern Arabian Sea, 2 peak spawning periods occur, during January to April and July to September(Karandikar and Palekar, 1950).
Kagwade (1970) believed that the species first matures at a total length of between 360 and 390 mm in Indian waters. Earlier,
Karandikar and Palekar (1950) had described ova sizes in the species from India: immature ovaries contained ova
measuring up to 0.4 mm diameter, maturing ovaries with ova between 0.45 and 0.68 mm diameter, and mature ovaries with
ova between 0.7 and 1.0 mm diameter. Oil globules in the mature ova measured 0.25 to 0.30 mm diameter. Sarojini and
Malhotra (1952) traced the development of larvae of the species from 6.0 mm until the juvenile phase. The swimbladder, welldeveloped in larvae, had become vestigial in juveniles and adults. Body scales were apparent on larvae of about 15 mm.
Size: Maximum total length about 2 m; weight 145 kg (Kailola and Stewart in Kailola et al., 1993).
Interest to Fisheries: One of the most important fisheries species for Kuwait, India, Thailand, Viet Nam, Malaysia,
Singapore and Indonesia, and is also a significant component of fisheries off Bangladesh, Myanmar, Cambodia and northern
Australia. The species is mainly caught by gill net, sometimes by trawl, averaging about 3% of the total landings in the gill net
fishery in the Gulf of Carpentaria, Australia, between 1980 and 1987 (Kailola and Stewart in Kailola et al., 1993). The species
is also an important sport fish in Australian waters, being caught by rod and reel or handlines. In Queensland, Australia, thefishery for the species is significant from Port Douglas to Gladstone on the east coast, as well as in the Gulf of Carpentaria
around the ports of Weipa and Karumba (Kailola and Stewart in Kailola et al., 1993).
Local Names: AUSTRALIA: Blue threadfin, Blue salmon, Cooktown salmon, Rockhampton salmon, Giant threadfin,
Colonial salmon, Blunt-nosed salmon, Blind tassel-fish, Bluenose salmon; BANGLADESH: Tailla; CAMBODIA: Trey pream;
INDIA: Salliah, Saccolih; INDONESIA: Baling, Kesumbang, Kurau, Lausan, Sumbal; ISLAMIC REPUBLIC OF IRAN:
Rashgoo; KUWAIT: Sheem; MALAYSIA: Genohong, Ikan salangan, Jenohong, Jubal senohong, Kubal, Kurau, Kurau
janggut, Kuru janggut, Mancong, Senangin, Senohong, Sinanghi, Selanghi; MYANMAR: Kakuyan, Nga-let-kwa, Za yaw gyi;
PHILIPPINES: Bikau, Hugao, Hugau, Kagau, Kugao, Mamali, Pugao; SRI LANKA: Kalawa, Kalemeen, Pozhakkala,Yevakala; VIET NAM: Cá chét.
Literature: Motomura et al. (2002a).
Remarks: Eleutheronema tetradactylum was originally proposed by Shaw (1804) as Polynemus tetradactylus for the“Maga Jellee” of Russell (1803), whose description of the species included a figure (fig. 183), but lacked a formal scientificname and gave no indication of any type specimens. Subsequently, Hamilton (1822) overlooked Shaw’s (1804) description
and described Polynemus teria from estuaries of the Ganges River, India, stating it to be the same species as Russell’s(1803) “Maga Jellee.” The description of P. teria also failed to identify any type specimens. In fact, the present whereaboutsof all of Hamilton’s (1822) types, including the type of P. teria, are unknown; they are not held in BMNH or other British or
Indian museums (Hora, 1929). Polynemus teria is clearly justified as a junior synonym of E. tetradactylum because “4pectoral filaments” were included in the original description of P. teria which is consistent with the pectoral filament condition
in E. tetradactylum. Also, Hamilton (1822) considered P. teria to be the same as Russell’s (1803) “Maga Jellee”(= E. tetradactylum).

Polynemus coecus was described by Macleay (1878) from Darwin, Northern Territory, Australia, on the basis of 2
specimens. Examination of these syntypes (AMS I. 9791 and I. 16295-001) showed them both to be conspecific with
E. tetradactylum. Their meristic and morphological characters are included in Motomura et al. (2002a: table 2).
Günther (1860), Day (1876) and Weber and de Beaufort (1922) reported Polynemus salliah Cantor, 1838 as a juniorsynonym of Polynemus tetradactylus or E. tetradactylum. However, Cantor (1838) wrote only “To the genus Polynemus, I
shall add a species, called by the natives Salliah, or Saccolih,” not following the Principles of Binominal Nomenclature(Article 5.1, ICZN-1999). Furthermore, when Cantor’s (1838) “Salliah or Saccolih” was first reported as a scientific name(Polynemus salliah Cantor, 1838), it was treated as a junior synonym of Polynemus tetradactylus, then considered to beThreadfins of the World19

valid. Accordingly, Polynemus salliah is not available under Articles 11.4 and 11.6 (ICZN-1999) (see Motomura, Senou andIwatsuki, 2001; Motomura et al., 2002a).

Eleutheronema tetradactylum previously reported from East Asia (e.g. Shen, 1984; Motomura, Senou and Iwatsuki, 2001)
is now identified as E. rhadinum.
Two patterns of lateral-line squamation on the caudal-fin membrane exist in E. tetradactylum. The lateral line ofE. tridactylum is unbranched, extending from the upper end of the gill opening to the upper end of the lower caudal-fin lobe,
whereas it is either branched or unbranched in E. tetradactylum. All specimens (18 specimens examined by the author) of
E. tetradactylum collected from northern and eastern Australia, and southern Papua New Guinea had unbranched laterallines, whereas all specimens (93 specimens examined by the author), except 2 specimens, of that from other localities had
the lateral line divided into 3 lines on the caudal-fin membrane (see Motomura et al., 2002a: fig. 7). Although in
E. tetradactylum, the relative frequency of specimens with an unbranched lateral line ranged from 100% (from Australia andPapua New Guinea) to 2% (from other localities), there were no other differences apparent. Therefore, the differences inlateral-line squamation are believed to represent intraspecific (geographic) variation. A single example with the lateral line
divided into 4 lines on the caudal-fin membrane was observed by the author at Mangalore central fish market in Mangalore,
Karnataka, India, but the specimen was not retained because of its large size (about 1.5 m standard length). The existenceof 3 or 4 (the latter a malformation?) branches of the lateral line on the caudal-fin membrane of some E. tetradactylum is
unique among polynemids, although bifurcation of the lateral line on the caudal-fin base is known in some Polydactylus
species, e.g., P. approximans, P. bifurcus and P. virginicus. Other members of polynemids have an unbranched lateralline, extending to the upper end of the lower caudal-fin lobe, the lower end of the upper caudal-fin lobe or the middistal
margin of the caudal-fin membrane.

Eleutheronema tridactylum (Bleeker, 1849)

Fig. 39; Plate Ic

Polynemus tridactylus Bleeker, 1849: 57 [type locality: Jakarta, Java, Indonesia; holotype (RMNH 6012, 255 mm standard
length) determined from 11 Bleeker specimens by Motomura et al., 2002a].
Synonyms: None.
FAO Names: En - Threefinger threadfin; Fr - Barbure à trois doigts; Sp - Barbudo de tres dedos.

Fig. 39 Eleutheronema tridactylum
Diagnostic Features: A medium-sized species. Body elongate, body depth at first dorsal-fin origin 22 to 27% (mean 24%)of standard length; head length 26 to 30% (mean 28%) of standard length. Posterior margin of maxilla extending wellbeyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 16% (mean 15%) of standard length; depth ofposterior margin of maxilla 2 to 3% (mean 2%) of standard length; anterior parts of lower jaw with small teeth extending onto
lateral surface, adjacent portion of lip absent; tooth plate extension onto lateral surface of lower jaw 9 to 10% (mean 9%) of
standard length; teeth villiform in broad bands on vomer, palatines and ectopterygoids; vomer lacking deciduous tooth plates
on both sides throughout life; ectopterygoids becoming wider with continued fish growth. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 13 (rarely 14)
soft rays; anal fin with III spines and 14 or 15 (mode 15) soft rays, anal-fin base longer than second dorsal-fin base; pectoral
fin with 16 to 18 (mode 17) unbranched rays, its length 21 to 23% (mean 22%) of standard length, posterior tip just short of

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FAO Species Catalogue for Fishery Purposes No. 3

level of posterior tip of pelvic fin; pectoral filaments 3, first filament shortest, reaching to (or extending slightly beyond) level ofpelvic-fin origin; second filament extending well beyond level of pelvic-fin origin; third filament longest, its length 21 to 29%(mean 25%) of standard length, not reaching to level of posterior tip of pectoral fin; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 29 to 37% (mean 34%) and lower lobe 25 to 33% (mean 30%) of
standard length. Pored lateral-line scales 72 to 79 (mode 75); lateral line simple, extending from upper end of gill opening toupper end of lower caudal-fin lobe; scale rows above lateral line 8 to 10 (mode 9), below 12 to 16 (mode 14). Gillrakers 2 to 4
(mode 3) on upper limb, 2 to 6 (mode 5) on lower limb, 4 to 10 (mode 8) total; gillrakers on anterior parts of upper and lower
limbs replaced during fish growth by tooth plates with villiform teeth, each gillraker on both upper and lower limbs becoming
shorter with fish growth. Vertebrae 10 precaudal and 15 caudal; supraneural bones 0 to 2. Swimbladder absent. Colour:
Upper sides of head and trunk green, becoming silvery white or yellowish white on lower sides; dorsal, pectoral, and caudal
fins dark with black borders.
Geographical Distribution: Currently known fromThailand (Gulf of Thailand and Phuket Island,Andaman Sea), Malaysia (Malay Peninsula andSarawak, Kalimantan) and Indonesia (Sumatra, Java,
Kalimantan and Moluccas) (Fig. 40).
Habitat and Biology: No data are available.
Size: Maximum standard length at least 25 cm(Motomura et al., 2002a).
Interest to Fisheries: Of little commercial importancebecause the population of the species appears to be
too small. The species has been taken with
E. tetradactylum.
Local Names: INDONESIA: Suro.
Literature: Feltes in Carpenter and Niem (2001);
Motomura et al. (2002a).
Remarks: Initially, Bleeker (1845) gave only the name“Polynemus tridactijlus,” there being no distinguishingfeatures included. However, the species was later
described in detail by Bleeker (1849) on the basis of asingle specimen, about 350 mm total length which
represents the largest confirmed record of the species.

200 m

Fig. 40 Eleutheronema tridactylum
Known distribution

Eleutheronema tridactylum is easily distinguished from both E. rhadinum and E. tetradactylum by having vomer without
tooth plates (versus vomer with 2 deciduous tooth plates in E. rhadinum and E. tetradactylum) and lower counts of second
dorsal-fin soft rays [13 (rarely 14) versus 14 (rarely 13 or 15 in E. rhadinum and E. tetradactylum)], pectoral filaments (3versus 4 in E. rhadinum and E. tetradactylum) and gillrakers [4 to 10 (mode 8) versus 10 to 17 (mode 12) and 6 to 18 (mode13) in E. rhadinum and E. tetradactylum, respectively].

Filimanus Myers, 1936
Filimanus Myers, 1936: 379 [type species: Polynemus melanochir (not of Valenciennes): Myers, 1936 (later described as
Filimanus perplexa Feltes, 1991)].
Synonyms: None.
Diagnostic Features: Body oblong to moderately deep, compressed. Adipose eyelid well developed; eye diameter greaterthan snout length. Lip on lower jaw well developed and dentary teeth restricted to dorsal surface; width of tooth band on
upper and lower jaws less than space (on symphysis) separating tooth bands on opposing premaxillae; teeth villiform innarrow bands on jaws, palatines and ectopterygoids, tooth plates on palatines shorter than those on ectopterygoids; vomer
with inconspicuous tooth plate. Posterior margin of maxilla extending beyond level of posterior margin of adipose eyelid.
Posterior margin of preopercle serrated. Basisphenoid not in contact with prootic; sphenotics not visible dorsally between
anterior margins of parietal and pterotic. First dorsal fin with VIII spines; second dorsal fin with I spine and 11 to 13 soft rays;
anal fin with III spines and 10 to 15 soft rays; anal-fin base less than head length; pectoral fin with 13 to 16 rays, allunbranched; pectoral-fin insertion well below midline of body; pectoral-fin base (including base of pectoral filaments) lessthan upper-jaw length; pectoral filaments 5 to 8, not extending beyond level of posterior tip of caudal fin; caudal fin deeply
forked, but upper and lower caudal-fin lobes not filamentous. Pored lateral-line scales 43 to 52; scale rows above lateral line
5 to 8, below 9 to 12. Gillrakers 35 to 49. Vertebrae 10 precaudal and 14 caudal; supraneural bones 2 or 3.

Threadfins of the World

21

Habitat and Biology: Filimanus species are usually found in 1 to 80 m on open sandflats and muddy substrates. Theyfrequently enter estuaries and are mostly taken in trawls.
Geographic Distribution: The genus is distributed in the Indo-West Pacific where it ranges from Pakistan to thePhilippines, Papua New Guinea and the Solomon Islands.
Interest to Fisheries: Because of their small size, Filimanus species are generally of little commercial importance.
Species: The genus comprises 6 species:

Filimanus heptadactyla: Western Pacific, Thailand to Papua New Guinea.
Filimanus hexanema: Java, Indonesia.
Filimanus perplexa: Andaman Sea to western Indonesia.
Filimanus sealei: Western Pacific, Philippines to Solomon Islands.
Filimanus similis: Indian Ocean, Pakistan to Andaman Sea.
Filimanus xanthonema: East coast of India to western Indonesia.
Remarks: Myers (1936) described Filimanus as a new genus on the basis of a single specimen (USNM 72742, 114 mmstandard length), but had misidentified the latter as Polynemus melanochir Valenciennes in Cuvier and Valenciennes,
1831. Polynemus melanochir is, in fact, a valid species belonging to Polynemus, the specimen used by Myers (1936) in his
description of Filimanus representing an undescribed species, subsequently described by Feltes (1991) as F. perplexa[based on 26 specimens (68 to 157 mm standard length), including the specimen used in the original description ofFilimanus]. In Opinion 1761 (ICZN) established F. perplexa as the type species of Filimanus. Feltes (1991) included 5
species in Filimanus in addition to the type species.

Key to the Species of Filimanus
1a. Pectoral filaments extending well beyond level of midpoint of anal-fin base; anal-fin soft rays 13 to 15(mode 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Pectoral filaments not extending beyond level of midpoint of anal-fin base; anal-fin soft rays 10 to 12(mode 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Pectoral filaments 6; body depth at first dorsal-fin origin 29 to 32% (mean 30%) of standard length(Fig. 41, Plate Ie) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filimanus hexanema(Java, Indonesia)
2b. Pectoral filaments 7; body depth at first dorsal-fin origin 32 to 37% (mean 36%) of standard length(Fig. 42, Plate If) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filimanus perplexa(Andaman Sea to western Indonesia)
body depth 29-32% of standard length

body depth 32-37% of standard length

6 pectoral filaments
7 pectoral filaments

Fig. 41 Filimanus hexanema

Fig. 42 Filimanus perplexa

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FAO Species Catalogue for Fishery Purposes No. 3

3a. Pectoral filaments 8 (rarely 7); gillrakers
40 to 48 (mode 46) (Fig. 43, Plate Ig). . . . . . . . . . . . . . . . . Filimanus sealei(Philippines to Solomon Islands)
3b. Pectoral filaments 6 or 7 (rarely 5);
gillrakers 35 to 49 . . . . . . . . . . . . . . . ® 4

40-48
gillrakers

8 (rarely 7) pectoral filaments

Fig. 43 Filimanus sealei

4a. Pectoral filaments 6 (occasionally 5), or
asymmetrically 5 and 6, or 6 and 7;
gillrakers 36 to 46 (mode 41) (Fig. 44,
Plate IIa) . . . . . . . . Filimanus xanthonema(east coast of India to western Indonesia)

4b. Pectoral filaments 7 . . . . . . . . . . . . . ® 5
36-46gillrakers

6 (occasionally 5) pectoral
filaments or asymmetrically
5 and 6, or 6 and 7

Fig. 44 Filimanus xanthonema

5a. Gillrakers 35 to 41 (mode 39) (Fig. 45, Plate Id)
5b. Gillrakers 40 to 49 (mode 43) (Fig. 46, Plate Ih)

35-41
gillrakers. . . . . . . . . . . . . . . . . . . . . Filimanus heptadactyla(Thailand to Papua New Guinea). . . . . . . . . . . . . . . . . . . . . . . . Filimanus similis(Pakistan to Andaman Sea)

40-49
gillrakers

Fig. 45 Filimanus heptadactyla

Fig. 46 Filimanus similisThreadfins of the World

23

Filimanus heptadactyla (Cuvier, 1829)

Fig. 47; Plate Id

Polynemus heptadactylus Cuvier in Cuvier and Valenciennes, 1829: 390 (type locality: Jakarta, Java, Indonesia, based on a
drawing; no types known, see Feltes, 1991).
Synonyms: None.
FAO Names: En - Sevenfinger threadfin; Fr - Barbure à sept doigts; Sp - Barbudo de siete dedos.

Fig. 47 Filimanus heptadactyla
Diagnostic Features: A small species. Body depth at first dorsal-fin origin 28 to 33% (mean 31%) of standard length; headlength 28 to 33% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of maxilla
extending beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 16% (mean 15%) of standard length; depthof posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted to dorsal surface;teeth villiform in narrow bands on palatines and ectopterygoids; vomer with an inconspicuous tooth plate. Posterior margin of
preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 12 or
13 (mode 12) soft rays; anal fin with III spines and 11 or 12 (mode 11) soft rays, anal-fin base approximately equal to or greaterthan second dorsal-fin base; pectoral fin with 14 to 16 (mode 15) unbranched rays, its length 23 to 29% (mean 26%) of standard
length, posterior tip just short of level of posterior tip of pelvic fin; pectoral filaments 7, first filament shortest, not reaching to levelof posterior margin of pelvic fin; third and fourth filaments usually longest, its length 28 to 40% (mean 34%) of standard length,
reaching to level of pelvic-fin origin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin
lobe 28 to 40% (mean 35%) of standard length. Poredlateral-line scales 46 to 49 (mode 48); lateral line simple,
extending from upper end of gill opening to mid-distal
margin of caudal-fin membrane; scale rows above lateralline 6 or 7 (mode 6), below 9 or 10 (mode 10). Gillrakers(total) 35 to 41 (mode 39). Vertebrae 10 precaudal and14 caudal; supraneural bones 3. Swimbladder simple.
Colour: Upper sides of head and trunk with brown tinge,
becoming golden on lower sides; posterior margins offirst, second dorsal, anal and caudal fins blackish,remaining areas translucent or yellowish; pectoral fin
mostly black; pectoral filaments white or yellowish white;
pelvic-fin anterior margin yellow, elsewhere white;anal-fin base yellow, elsewhere yellowish white.
Geographical Distribution: Distributed in the western
Pacific where it ranges from Thailand, Malaysia (eastcoast of Malay Peninsula) and Indonesia (Java,
Kalimantan, and Moluccas) to Papua New Guinea (Gulfof Papua) (Fig. 48).

200 m

Fig. 48 Filimanus heptadactyla
Known distribution

24

FAO Species Catalogue for Fishery Purposes No. 3

Habitat and Biology: See above under genus account. No other data are available.
Size: Maximum standard length at least 13 cm (Feltes, 1991).
Interest to Fisheries: None.
Local Names: INDONESIA: Laas, Laos; PAPUA NEW GUINEA: Seven-fingered threadfin.
Literature: Feltes (1991); Feltes in Carpenter and Niem (2001).
Remarks: Polydactylus heptadactylus was described as a new species on the basis of a drawing (reproduced in Feltes,
1991: 313, fig. 9) from Jakarta, Java, Indonesia. Subsequently, many researchers (e.g. Marathe and Bal, 1958; Nayak,
1959b; Kagwade, 1970) have misidentified F. similis, Polydactylus mullani and P. sextarius as F. heptadactyla.
Polydactylus mullani and P. sextarius can be easily distinguished from F. heptadactyla by the presence of a large blackanterior lateral-line spot (absent in the latter). Filimanus heptadactyla, distributed in the western Pacific, is similar toF. similis, distributed in the eastern Indian Ocean, in having 7 pectoral filaments not reaching to level with the midpoint of theanal-fin base. However, the former differs from F. similis in having lower gill-raker counts [35 to 41 (mode 39) versus 40 to
49 (mode 43) in the latter].

Filimanus hexanema (Cuvier, 1829)

Fig. 49; Plate Ie

Polynemus hexanema Cuvier in Cuvier and Valenciennes, 1829: 389 (type locality: Jakarta, Java, Indonesia, based on a
drawing; no types known, see Feltes, 1991).
Synonyms: Polynemus senarius Gronow in Gray, 1854: 176 [type locality: Indian Ocean, but probably erroneous, seeFeltes, 1991; holotype (BMNH 1853.11.12.59, 99 mm standard length)].
FAO Names: En - Javanese threadfin; Fr - Barbure de Java; Sp - Barbudo javanés.

Fig. 49 Filimanus hexanema

Threadfins of the World

25

Diagnostic Features: A small species. Body depth at first dorsal-fin origin 29 to 32% (mean 30%) of standard length; headlength 28 to 31% (mean 29%) of standard length. Snout blunt, interorbital region of frontals not markedly concave. Posterior
margin of maxilla extending beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 15% (mean 15%) of
standard length; lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in narrow bands on
palatines and ectopterygoids; vomer with an inconspicuous tooth plate lacking teeth. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 11 or 12(mode 12) soft rays; anal fin with III spines and 14 soft rays, anal-fin base longer than second dorsal-fin base; pectoral fin
with 14 or 15 (mode 14) unbranched rays, its length 31 to 42% (mean 36%) of standard length, posterior tip extending
beyond level of posterior tip of pelvic fin (sometimes reaching to level of anal-fin origin); pectoral filaments 6, first filament
shortest, just short of level of anal-fin origin; second filament extending beyond level of posterior part of anal-fin base; thirdand fourth filaments usually longest, its length 66 to 81% (mean 74%) of standard length, extending beyond level of midpointof anal-fin base (sometimes extending beyond level of midpoint of caudal peduncle); caudal fin deeply forked, upper andlower caudal-fin lobes not filamentous, upper caudal-fin lobe 30 to 42% (mean 34%) of standard length. Pored lateral-line
scales 49 or 50 (mode 50); lateral line simple, extending from upper end of gill opening to mid-distal margin of caudal-fin
membrane; scale rows above lateral line 6 or 7 (mode 6), below 9 or 10 (mode 10). Gillrakers (total) 43 to 48 (mode 45).
Vertebrae 10 precaudal and 14 caudal; supraneural bones 2. Colour: Body uniformly silver; posterior margins of first,
second dorsal, anal, and caudal fins blackish, remaining parts translucent or yellowish; pectoral fin with scatteredmelanophores; pectoral filaments white; anterior margin of pelvic fin white, other parts translucent.
Geographical Distribution: Known only from western
Java, Indonesia (off Jakarta, Serang and Labuhan)(Fig. 50).
Habitat and Biology: Nothing has been published on
the biology of this rare species.
Size: Maximum standard length at least 12 cm (Feltes,
1991).
Interest to Fisheries: None.
Local Names: None known.
Literature: Feltes (1991); Motomura et al. (2001a);
Feltes in Carpenter and Niem (2001).
Remarks: F. hexanema appears to be one of the rarest
polynemids. It is similar to F. perplexa in having longpectoral filaments, extending beyond level with the
midpoint of the anal-fin base, but differs from the latterin having lower counts of pectoral filaments (6 versus 7
in the latter) and gillrakers [44 to 48 (mode 45) versus
47 to 55 (mode 50)], and a shallower body depth [29 to
32% (mean 30%) of standard length versus 32 to 37%(mean 36%) of standard length].

Filimanus perplexa Feltes, 1991

200 m

Fig. 50 Filimanus hexanema
Known distribution

Fig. 51; Plate If

Filimanus perplexa Feltes, 1991: 307, fig. 5 [type locality: northwest of Denpasar, Bali, Indonesia; holotype (BMNH
1988.4.6.1, 148 mm standard length); 23 paratypes (BMNH 1988.4.6.2, 155 mm standard length; BMNH 1988.4.6.3,
146 mm standard length; BMNH 1988.4.6.4, 121 mm standard length; BMNH 1988.4.6.5, 156 mm standard length; BMNH
1988.4.6.6, 104 mm standard length; BMNH 1988.4.6.7, 105 mm standard length; BMNH 1988.4.6.8, 148 mm standard
length; BMNH 1988.4.6.9, 157 mm standard length; BMNH 1988.4.6.10, 104 mm standard length; CAS 59177, 134 mm
standard length; FMNH 97100, 2 specimens, 109 to 118 mm standard length; MCZ 30869, 104 mm standard length; MNHN
A. 3054, 68 mm standard length; NMW 12479, 79 mm standard length; NMW 12548, 3 specimens, 84 to 107 mm standard
length; NMW 12549, 2 specimens, 71 to 86 mm standard length; NMW 22598, 107 mm standard length; USNM 72742,
114 mm standard length; USNM 280288, 115 mm standard length)].
Synonyms: None.

26

FAO Species Catalogue for Fishery Purposes No. 3

FAO Names: En - Splendid threadfin; Fr - Barbure magnifique; Sp - Barbudo espléndido.

Fig. 51 Filimanus perplexa
Diagnostic Features: A small species. Body moderately deep, body depth at first dorsal-fin origin 32 to 37% (mean 34%)of standard length; head length 30 to 34% (mean 32%) of standard length. Snout blunt, interorbital region of frontals
slightly concave. Posterior margin of maxilla extending beyond level of posterior margin of adipose eyelid; upper-jaw
length 15 to 18% (mean 17%) of standard length; lip on lower jaw well developed, dentary teeth restricted to dorsal surface;teeth villiform in narrow bands on palatines and ectopterygoids; vomer with an inconspicuous tooth plate. Posterior marginof preopercle serrated. First dorsal fin with VIII spines, second spine slightly more robust than others; second dorsal fin
with I spine and 11 soft rays; anal fin with III spines and 13 to 15 (mode 14) soft rays, anal-fin base longer than second
dorsal-fin base; pectoral fin with 13 or 14 (mode 14) unbranched rays, its length 31 to 49% (mean 35%) of standard length,
posterior tip extending beyond level of posterior tip of pelvic fin (sometimes reaching to level of anal-fin origin); pectoral
filaments 7, first filament shortest, just short of (or reaching to) level of anal-fin origin; third and fourth filaments usually
longest, its length 56 to 118% (mean 92%) of standard length, extending well beyond level of midpoint of anal-fin base(sometimes extending beyond level of fork of caudal fin); caudal fin deeply forked, upper and lower caudal-fin lobes notfilamentous, upper caudal-fin lobe 31 to 56% (mean 44%) of standard length. Pored lateral-line scales 49 to 51 (mode 50);
lateral line simple, extending from upper end of gill opening to mid-distal margin of caudal-fin membrane; scale rows above
lateral line 5 to 8 (mode 6), below 9 or 10 (mode 10). Gillrakers (total) 47 to 55 (mode 50). Vertebrae 10 precaudal and 14
caudal; supraneural bones 2. Swimbladder simple. Colour: Body and fins fawn to yellow; fins yellowish white; base ofpectoral filaments white, becoming light brown on posterior tips; pectoral fin with scattered melanophores (intensity ofpectoral-fin pigmentation variable).
Geographical Distribution: Known only from Padang(east side of Sumatra), Nias (west side of Sumatra), Javaand Bali, Indonesia, and Phuket, Thailand (Fig. 52). The
record from Andaman Sea is based only on a single
specimen (PMBC 5913, 111 mm standard length).
Habitat and Biology: See above under genus account.
No other data are available.
Size: Maximum standard length at least 16 cm (Feltes,
1991).
Interest to Fisheries: None.
Local Names: INDONESIA: Char-wen-e-kerning.
Literature: Feltes (1991); Motomura, Satapoomin and
Iwatsuki. (2000); Feltes in Carpenter and Niem (2001).
200 m

Remarks: F. perplexa was described as a new species
by Feltes (1991). Bleeker (1849, 1855, 1856, 1857a,
1859, 1860) erroneously referred to this species as
Polynemus melanochir, that name being used forFig. 52 Filimanus perplexa
F. perplexa by many subsequent authors (e.g. Volz,
Known distribution
1903; Bean and Weed, 1912; Myers, 1936). Although
Feltes (1991) described F. perplexa on the basis of 26 specimens, he listed only 24 type specimens in his paper. The present
whereabouts of remaining 2 type specimens are unknown. Comparisons of F. perplexa with a related species, F. hexanema,
are given in the account of the latter.

Threadfins of the World

27

Filimanus sealei (Jordan and Richardson, 1910)

Fig. 53; Plate Ig

Polydactylus opercularis Seale and Bean, 1907: 234 [type locality: Zamboanga, Mindanao, Philippines; holotype (USNM
57844, 124 mm standard length); junior secondary homonym of Trichidion opercularis Gill, 1863]. Polydactylus sealei
Jordan and Richardson, 1910: 16 (replacement name for P. opercularis Seale and Bean).
Synonyms: None.
FAO Names: En - Eightfinger threadfin; Fr - Barbure à huit doigts; Sp - Barbudo de ocho dedos.

Fig. 53 Filimanus sealei
Diagnostic Features: A small species. Body depth at first dorsal-fin origin 29 to 35% (mean 32%) of standard length; headlength 29 to 34% (mean 32%) of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of maxilla
extending beyond level of posterior margin of adipose eyelid; depth of posterior margin of maxilla less than eye diameter;
upper-jaw length 14 to 16% (mean 16%) of standard length; lip on lower jaw well developed, dentary teeth restricted to dorsal
surface; teeth villiform in narrow bands on palatines and ectopterygoids; vomer with an inconspicuous tooth plate. Posterior
margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine
and 11 or 12 (mode 12) soft rays; anal fin with III spines and 11 soft rays, anal-fin base approximately equal to or greater thansecond dorsal-fin base; pectoral fin with 14 or 15 (mode 14) unbranched rays, its length 24 to 28% (mean 25%) of standard
length, posterior tip just short of level of posterior tip of pelvic fin; pectoral filaments 8 (rarely 7), first filament shortest; third andfourth filaments usually longest, its length 37 to 55% (mean 42%) of standard length, reaching to (or just short of) level ofanal-fin origin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 33 to 42%(mean 38%) of standard length. Pored lateral-line scales 46 to 50 (mode 49); lateral line simple, extending from upper end of gill
opening to mid-distal margin of caudal-fin membrane; scale rows above lateral line 6 or 7 (mode 7), below 9 or 10 (mode 10).
Gillrakers (total) 40 to 48 (mode 46). Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Colour: Upper sides of head
and trunk with blue tinge, becoming silvery white on lower
sides; fins yellow with black at margins.
Geographical Distribution: In the western Pacific,
where it ranges from the Philippines to the Solomon
Islands, including Papua New Guinea and the Bismarck
Archipelago (Fig. 54). A single specimen (ANSP
123498, 108 mm standard length) supposedly collectedfrom Western Australia in 1950, lacked detailed locality
and other collection data. Accordingly, Australian
distribution of the species still requires confirmation.
Habitat and Biology: See above under genus account.
No other data are available.
Size: Maximum standard length at least 15 cm (Feltes,
1991).
Interest to Fisheries: None.

200 m

Local Names: PAPUA NEW GUINEA: Eight-fingered
threadfin.

Fig. 54 Filimanus sealei
Known distribution

28

FAO Species Catalogue for Fishery Purposes No. 3

Literature: Feltes (1991); Feltes in Carpenter and Niem (2001).
Remarks: Polydactylus opercularis (Gill, 1863) was originally described as Trichidion opercularis on the basis of a single
specimen (apparently lost) from the west coast of central America. Although the former was regarded as a valid species,
Seale and Bean (1907) overlooked Gill’s (1863) description and described Polydactylus opercularis on the basis of a single
specimen (USNM 57844, 124 mm standard length) from Zamboanga, Mindanao, Philippines. Jordan and Richardson(1910) recognized this designation as a homonym and provided the replacement name of P. sealei.

Filimanus sealei can be easily distinguished from other congeners by the number of pectoral filaments (usually 8 versus
usually 6 in F. hexanema and F. xanthonema, and 7 in F. heptadactyla, F. perplexa and F. similis).

Filimanus similis Feltes, 1991

Fig. 55; Plate Ih

Filimanus similis Feltes, 1991: 318, fig. 12 [type locality: Beruwala, Sri Lanka; holotype (USNM 304495, 99 mm standard
length); 43 paratypes (AMS I. 211033-004, 3 specimens, 124 to 127 mm standard length; ANSP 74846, 2 specimens, 69 to
73 mm standard length; BMNH 1860.3.19.56, 105 mm standard length; BPBM 19048, 2 specimens, 101 mm standard
length; CAS SU 14591, 116 mm standard length; CAS SU 22901, 99 mm standard length; FMNH 58982, 113 mm standard
length; LACM 38134-17, 19 specimens, 82 to 104 mm standard length; USNM 149704, 3 specimens, 70 to 98 mm standard
length; USNM 278215, 82 mm standard length; USNM 278202, 2 specimens, 61 to 69 mm standard length; USNM 278242,
7 specimens, 87 to 103 mm standard length)].
Synonyms: None.
FAO Names: En - Indian sevenfinger threadfin; Fr - Barbure sept doigts des Indes; Sp - Barbudo de siete dedos índico.

Fig. 55 Filimanus similis
Diagnostic Features: A small species. Body depth at first dorsal-fin origin 30 to 34% (mean 32%) of standard length; headlength 29 to 34% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of maxilla
extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw length 13 to 17% (mean 15%) of standard
length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted todorsal surface; teeth villiform in narrow bands on palatines and ectopterygoids; vomer with an inconspicuous tooth plate.
Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal
fin with I spine and 11 to 13 (mode 12) soft rays; anal fin with III spines and 10 to 12 (mode 11) soft rays, anal-fin baseapproximately equal to or greater than second dorsal-fin base; pectoral fin with 14 to 16 (mode 15) unbranched rays, its
length 24 to 31% (mean 27%) of standard length, posterior tip just short of level of posterior tip of pelvic fin; pectoral filaments
7, first filament shortest, not reaching to level of posterior tip of pelvic fin; third and fourth filaments usually longest, its length
25 to 48% (mean 39%) of standard length, reaching to (sometimes extending slightly beyond) level of anal-fin origin; caudal
fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 31 to 44% (mean 38%) of standard

Threadfins of the World29

length. Pored lateral-line scales 45 to 49 (mode 48); lateral line simple, extending from upper end of gill opening to mid-distal
margin of caudal-fin membrane; scale rows above lateral line 5 to 7 (mode 6), below 9 to 11 (mode 10). Gillrakers (total) 40 to
49 (mode 43). Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Colour: Upper sides of head and trunk withbrown tinge, becoming golden on lower sides; posterior margins of first, second dorsal, anal, and caudal fins blackish,remaining parts yellowish; pectoral fin mostly black; base of pectoral filaments white, becoming yellowish white on posterior
tips; base of pelvic fin white, other parts yellowish.
Geographical Distribution: In the eastern Indian
Ocean where it ranges from Pakistan to the Andaman
Sea (Fig. 56). The species has currently beenrecorded from Pakistan, India, Sri Lanka and Thailand(Phuket, west coast of Malay Peninsula).
Habitat and Biology: See above under genus
account. No other data are available.
Size: Maximum standard length at least 13 cm.
Interest to Fisheries: A local bottom trawl fisheryexists in southern parts of India.
200 m

Local Names: MYANMAR: Za yaw.
Literature: Feltes (1991).

Fig. 56 Filimanus similis

Remarks: F. heptadactyla had been considered to be
Known distribution
widely distributed in the Indo-Pacific. However, Feltes(1991) recognized that the species was restricted to
the Pacific Ocean, including Indonesia, and described F. similis as a new species for Indian Ocean examples previously
identified as F. heptadactyla. Comparisons of F. similis with F. heptadactyla are given in the account of the latter.

Filimanus xanthonema (Valenciennes, 1831)

Fig. 57; Plate IIa

Polynemus xanthonemus Valenciennes in Cuvier and Valenciennes, 1831: 517 [type locality: Pondicherry, India; lectotype(MNHN A. 3033, 1 of 2 specimens, 109 mm standard length) designated by Feltes, 1991; paralectotype (MNHN A. 3033, 1 of
2 specimens, 107 mm standard length)].
Synonyms: Polynemus diagrammicus Bleeker, 1849: 60 (type locality: Jakarta, Java, Indonesia; type material probably
lost, see Feltes, 1991). Polynemus pfeifferi Bleeker, 1853a: 249 [type locality: Priaman, Sumatra, Indonesia; holotype
probably included in RMNH 6008 (5 specimens, 39 to 105 mm standard length), see Feltes, 1991]. Polydactylus
konadaensis Mishra and Krishnan, 1993: 285 [type locality: Konada, Andhra Pradesh, India; holotype (ZSI/MBS-F-631,
105 mm standard length); paratype (ZSI/MBS-F-632, 97 mm standard length)].
FAO Names: En - Yellowthread threadfin; Fr - Barbure à doigts oranges; Sp - Barbudo de dedos amarillos.

Fig. 57 Filimanus xanthonema

30

FAO Species Catalogue for Fishery Purposes No. 3

Diagnostic Features: A small species. Body depth at first dorsal-fin origin 27 to 35% (mean 31%) of standard length; headlength 28 to 35% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of maxilla
extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 17% (mean 15%) of standard
length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted
to dorsal surface; teeth villiform in narrow bands on palatines and ectopterygoids; vomer with an inconspicuous tooth plate.
Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal
fin with I spine and 11 to 13 (mode 12) soft rays; anal fin with III spines and 10 to 12 (mode 11) soft rays, anal-fin baseapproximately equal to or greater than second dorsal-fin base; pectoral fin with 13 to 15 (mode 15) unbranched rays, its
length 23 to 28% (mean 26%) of standard length, posterior tip just short of level of posterior tip of pelvic fin; pectoral filaments
usually 6 on each side but occasionally 5 on each side, or asymmetrically 5 and 6 or 6 and 7, first filament shortest, notreaching to level of posterior tip of pelvic fin; third and fourth filaments usually longest, its length 27 to 47% (mean 38%) of
standard length, reaching to (sometimes extending slightly beyond) level of anal-fin origin; caudal fin deeply forked, upperand lower caudal-fin lobes not filamentous, upper caudal-fin lobe 26 to 44% (mean 36%) of standard length. Poredlateral-line scales 43 to 52 (mode 47); lateral line simple, extending from upper end of gill opening to mid-distal margin of
caudal-fin membrane; scale rows above lateral line 5 to 8 (mode 6), below 9 to 12 (mode 10). Gillrakers (total) 36 to 46(mode 43). Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Colour: Upper sides of head and trunk with browntinge, becoming golden on lower sides; posterior margins of first, second dorsal, anal, and caudal fins blackish, remaining
parts yellowish; pectoral fin mostly black; base of pectoral filaments white, becoming yellowish white on posterior tips; baseof pelvic fin white, other parts yellowish.
Geographical Distribution: F. xanthonema is
distributed in the eastern Indian and western Pacific
Oceans where it ranges from east coast of India toLombok Island, Indonesia (Fig. 58).
Habitat and Biology: Available collection data forF. xanthonema indicate that the species occurs on
muddy bottoms from depths of 1 to 30 m. According to
Hida (1967), F. xanthonema exhibits protandry, sexchanging from male to female within a narrow size
range. Males range from 9.0 to 11.0 cm standard
length with a peak at 10.0 cm, females ranging from10.0 to 11.5 cm with a peak at 10.5 to 11.0 cm.
Hermaphrodites range from 9.0 to 11.5 cm with a peakat 10.0 cm. Filimanus xanthonema feeds on small
crustaceans, especially Mysidacea and shrimps in the
Bay of Bengal (Hida, 1967).
Size: Maximum standard length at least 14 cm (Feltes,
1991).

200 m

Interest to Fisheries: None.
Local Names: MYANMAR: Za yaw.
Literature: Feltes (1991); Motomura et al. (2001b);
Feltes in Carpenter and Niem (2001).

Fig. 58 Filimanus xanthonema
Known distribution

Remarks: F. xanthonema was originally described as Polynemus xanthonemus on the basis of 2 specimens registered as
MNHN A. 3033. The larger (109 mm standard length, with 6 pectoral filaments on each side) of the 2 specimens was
designated as lectotype by Feltes (1991), the paralectotype (107 mm standard length) having asymmetrically 6 and 7
pectoral filaments. Polynemus diagrammicus, from Jakarta, Java, Indonesia, and P. pfeifferi, from Priaman, Sumatra,Indonesia, were regarded as junior synonyms of F. xanthonema by Feltes (1991). Recently, Polydactylus konadaensis,
from Konada, Andhra Pradesh, India, was also reduced to the synonymy of F. xanthonema (see Motomura et al., 2001b).

Filimanus xanthonema can be easily distinguished from other congeners by the number and length of pectoral filaments.
The species usually has 6 pectoral filaments on each side (occasionally 5 on each side, or asymmetrically 5 and 6 or 6 and
7), not reaching to level with the midpoint of the anal-fin base, whereas F. hexanema and F. perplexa have 6 and 7 pectoral
filaments, respectively, extending beyond the level of the midpoint of the anal-fin base in both species. Filimanus
heptadactyla, F. sealei and F. similis have 7, 8 (occasionally 7) and 7 pectoral filaments, respectively, not reaching to levelwith the midpoint of the anal-fin base.

Threadfins of the World

31

Galeoides Günther, 1860
Galeoides Günther, 1860: 319, 332 [type species: Polynemus polydactylus Vahl, 1798, presently regarded as a juniorsynonym of Galeoides decadactylus (Bloch, 1795)].
Synonyms: None.
Species: The genus comprises a single species:

Galeoides decadactylus: West coast of Africa, Algeria to Namibia.
Remarks: To date, Galeoides has been considered to comprise 2 species, G. decadactylus (type species of the genus) andG. microps (e.g. Fowler, 1935; Myers, 1936; Springer, 1982), the latter being a poorly known threadfin originally described by
Steindachner (1869a) on the basis of a single specimen (NMW 77568, 171 mm standard length) from China (probably
erroneous). However, examination of the holotype of G. microps showed it to belong to the genus Polynemus, being a juniorsynonym of P. melanochir melanochir (see account of P. m. melanochir). Accordingly, Galeoides represents a monotypic
genus (Motomura, Mikschi and Iwatsuki, 2001).

Galeoides has the following unique characters among the family: lateral line simple, extending from upper end of gill opening
to lower end of upper caudal-fin lobe; a black spot present below anterior part of lateral line; and swimbladder extending
beyond anal-fin origin.

Galeoides decadactylus (Bloch, 1795)

Fig. 59; Plate IIb

Polynemus decadactylus Bloch, 1795: 26, pl. 401 [type locality: Guinea; holotype (ZMB 569, 175 mm standard length)].
Synonyms: Polynemus polydactylus Vahl, 1798: 164 (type locality: near Tangier, Morocco; no types known). Polynemus
enneadactylus Cuvier in Cuvier and Valenciennes, 1829: 392 (type locality: Tangier, Morocco; no types known). Polynemus
astrolabi Sauvage, 1881: 102 [type locality: Mauritius (clearly erroneous, see Motomura, Iwatsuki and Kimura, 2001b);
holotype (MNHN 9727, 190 mm standard length)].
FAO Names: En - Lesser African threadfin; Fr - Petit capitaine; Sp - Barbudo enano africano.

Fig. 59 Galeoides decadactylus
Diagnostic Features: A medium-sized species. Body moderately deep, body depth at first dorsal-fin origin 29 to 35%(mean 32%) of standard length; head length 31 to 36% (mean 33%) of standard length. Adipose eyelid well developed; eyediameter greater than snout length. Lip on lower jaw well developed, dentary teeth restricted to dorsal surface; width of toothband on upper and lower jaws greater than space (on symphysis) separating tooth bands on opposing premaxillae; teeth
villiform in broad bands on jaws, palatine and ectopterygoids, tooth plates on palatines longer than those on ectopterygoids;
tooth plates on ectopterygoids conspicuously small; vomerine tooth plate covered with skin and teeth absent. Posterior
margin of maxilla reaching to (or just short of) level of posterior margin of adipose eyelid; upper-jaw length 12 to 14% (mean32

FAO Species Catalogue for Fishery Purposes No. 3

13%) of standard length; maxillary scales absent. Posterior margin of preopercle serrated. Basisphenoid in contact withprootic; sphenotics not visible dorsally between anterior margins of parietal and pterotic. First dorsal fin with VIII spines;
second dorsal fin with I spine and 13 or 14 (mode 13) soft rays; anal fin with III spines and 10 or 11 (mode 11) soft rays,
anal-fin base less than second dorsal-fin base; pectoral fin with 12 to 15 (mode 15) unbranched rays, its length 21 to 25%(mean 23%) of standard length, posterior tip just short of level of posterior tip of pelvic fin; pectoral-fin insertion well below
midline of body; pectoral-fin base (including base of pectoral filaments) greater than or equal to upper-jaw length; pectoral
filaments 9 to 11 (mode 9); first filament shortest, not reaching to level of pelvic-fin origin; uppermost filament longest, its
length 21 to 33% (mean 27%) of standard length, reaching to or extending beyond level of pelvic-fin origin; caudal fin deeply
forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 31 to 43% (mean 38%) and lower caudal-fin
lobe 33 to 36% (mean 34%) of standard length. Pored lateral-line scales 45 to 50 (mode 46); lateral line simple, extending
from upper end of gill opening to lower end of upper caudal-fin lobe; scale rows above lateral line 5 or 6 (mode 5), below 7 to 9(mode 8). Gillrakers 9 to 14 on upper limb, 15 to 23 on lower limb, 24 to 36 total; gillrakers decreasing with fish growth.
Vertebrae 10 precaudal and 14 caudal; supraneural bone 1. Swimbladder simple, extending beyond anal-fin origin. Colour:
Upper sides of head and trunk with brown tinge, becoming silver on lower sides; posterior margins of first and second dorsal
fins, and caudal fins dense black, remaining parts blackish; pelvic and anal fin white; pectoral fin mostly black; base ofpectoral filaments white, becoming blackish on posterior tips; a black spot, its diameter approximately equal to eye diameter,
present below anterior part of lateral line; several brown stripes along longitudinal scale rows above and below lateral line
(disappeared in preserved specimens).
Geographical Distribution: Generally distributed from Morocco toAngola, west coast of Africa. In addition, the species has been reported
from Algeria, northern Africa and Namibia, southern Africa (Fig. 60).
Habitat and Biology: Usually taken on muddy bottoms in shallow
coastal waters from depths of 10 to 70 m, and frequently found inestuaries and lagoons. Generally feeds on crustaceans and small fishes(Longhurst, 1957: tables 1, 9). Longhurst (1960) examined the stomach
contents of 630 specimens collected from the open sea and 1 437
specimens from estuaries, and found that detritus amounted to 31% offood contained in the stomachs of the estuarine specimens (versus 0%
in open sea specimens).
Longhurst (1965) studied the biology, including sexuality and growth, ofG. decadactylus in Lagos, Nigeria, finding that about 25% of females
developed directly from the juvenile stages, the others developing
female gonads after passing through a non-functional hermaphroditic
stage arising in apparently normal males. He noted the sex compositionof 25 357 adult specimens as: males 63.73%, hermaphrodites 22.22%
and females 14.03%. Spawning of the species occurs in all months,
peaking in the dry season, but almost ceasing during the rainy season in
Nigerian waters. Derivation of lengths at first maturity for the species iscomplicated by the nature of the reproductive cycle. Lengths ranged
from 150 mm total length for males to 255 mm for secondary females,
the species growing rapidly during the first year. At 1 year old, thespecies averages 207 mm total length, 317 mm at 2 years old, and
390 mm at 3 years old. According to Longhurst (1965), it appears
improbable that many individuals survive a fourth year of life.
Aboussouan (1966) described and illustrated larval developmental
stages of G. decadactylus (as G. polydactylus) from Senegal.
Size: Maximum total length 45 cm, common to 30 cm (Allen in Fischer
et al., 1981).

200 m

Fig. 60 Galeoides decadactylus
Known distribution

Interest to Fisheries: In Nigeria, G. decadactylus is an importantcomponent in the commercial trawl fishery, constituting between 10 and 20% of the total landings by weight, the percentage
differing from port to port (Longhurst, 1963).
Local Names: GUINEA: Barbinho, Sanis; MAURITANIA: Plexiglass, Sibet mba, Tiekem; SÃO TOMÉ AND PRINCIPE:
Barbudo; SENEGAL: Barbo, Barudo de dez barbas, Cekéém, Lesser African threadfin, Sikket mbàw; SIERRA LEONE:
Shineose.
Literature: Allen in Fischer et al. (1981); Daget and Njock in Daget et al. (1986); Njock in Quéro et al. (1990); Motomura,
Iwatsuki and Kimura (2001b).
Remarks: Polynemus enneadactylus and P. polydactylus have been treated as junior synonyms of G. decadactylus (e.g.
Daget and Njock in Daget et al., 1986; Njock in Quéro et al., 1990). In the faunal work, Fricke (1999) synonymized
Polynemus astrolabi with Polydactylus sextarius. However, the holotype (MNHN 9727, 190 mm standard length) of
Polynemus astrolabi differs from the specimens including the holotype (ZMB 565, 125 mm standard length) of Polydactylus

Threadfins of the World33

sextarius in having 9 pectoral filaments (6 in the latter), unbranched pectoral-fin rays (almost branched in the latter) and 1
supraneural bone (3 in latter), and in lateral-line squamation on the caudal-fin membrane. Recently, Polynemus astrolabi
was regarded as a junior synonym of G. decadactylus by Motomura, Iwatsuki and Kimura (2001b).
Although Sauvage (1881, 1891) reported Polynemus astrolabi (= G. decadactylus) from Mauritius, Indian Ocean, thespecies is distributed along the west coast of Africa. Because the Equatorial current, which is the primary current influencing
waters off Mauritius, flows from the equator to the Southwest Indian Ridge, it appears to be highly unlikely that the species
can be transported from the west coast of Africa to Mauritius. Therefore, the type locality (Mauritius) of Polynemus astrolabi
is considered to be erroneous.

Leptomelanosoma Motomura and Iwatsuki, 2001
Leptomelanosoma Motomura and Iwatsuki, 2001a: 13 (type species: Polynemus indicus Shaw, 1804).
Synonyms: None.
Species: The genus comprises a single species:

Leptomelanosoma indicum: Indo-West Pacific, Pakistan to Indonesia.
Remarks: Leptomelanosoma has been recently proposed for the polynemid fish, Polynemus indicus, previously identified
as a member of Polydactylus. Leptomelanosoma differs from other genera in having the following characters: swimbladderwith many appendages inserted into lateral walls of abdominal cavity (unique character among the family); anterior one-thirdof lower jaw with small teeth extending onto lateral surface, adjacent portion of lip poorly developed throughout life [thischaracter also present in large specimens of Parapolynemus and Polydactylus opercularis]; ethmoid not covered dorsallyby frontals; sphenotics visible dorsally between anterior margins of parietal and pterotic (unique character among the family);upper and lower caudal-fin lobes very long, filamentous (this character also present in Parapolynemus); greyish black body(this character also appears in Polydactylus longipes).

Leptomelanosoma indicum (Shaw, 1804)

Fig. 61; Plate IIc

Polynemus indicus Shaw, 1804: 155 [type locality: Vizagapatnam, India, based on “Maga Booshee”of Russell (1803: 68, fig.
184); no types known, see Motomura and Iwatsuki, 2001a].
Synonyms: Polynemus sele Hamilton, 1822: 226 (type locality: estuaries of Ganges, India; no types known, see Motomura
and Iwatsuki, 2001a). Polynemus uronemus Cuvier 1829a: 155 (type locality: unknown; new name for P. indicus Shaw,
1804; no types known, see Motomura and Iwatsuki, 2001a). Polynemus gelatinosus McClelland, 1843: 181, pl. 6 (typelocality: Bengal, India; new name for P. sele Hamilton, 1822; no types known, see Motomura and Iwatsuki, 2001a).
FAO Names: En - Indian threadfin; Fr - Barbure indien; Sp - Barbudo indio.

Fig. 61 Leptomelanosoma indicum34

FAO Species Catalogue for Fishery Purposes No. 3

Diagnostic Features: A large species. Body and head elongate, body depth at first dorsal-fin origin 22 to 25% (mean 23%)of standard length; head length 30 to 34% (mean 32%) of standard length. Adipose eyelid moderately developed; eyediameter approximately equal to or greater than snout length. Anterior one-third of lower jaw with small teeth extending onto
lateral surface, adjacent portion of lip poorly developed; width of tooth band on upper and lower jaws greater than space (onsymphysis) separating tooth bands on opposing premaxillae; teeth villiform in broad bands on jaws, vomer, palatines andectopterygoids, tooth plates on palatines longer than those on ectopterygoids; tooth plates on palatines well developed.
Posterior margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 15%(mean 15%) of standard length; maxilla covered with deciduous scales. Posterior margin of preopercle serrated.
Basisphenoid in contact with prootic; sphenotics visible dorsally between anterior margins of parietal and pterotic. First
dorsal fin with VIII spines; second dorsal fin with I spine and 12 or 13 (mode 13) soft rays; anal fin with III spines and 10 or 11(mode 11) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 12 to 14 (mode 14) rays, its length 19 to
22% (mean 21%) of standard length, posterior tip not reaching to level of posterior tip of pelvic fin; pectoral-fin insertion well
below midline of body; pectoral-fin base (including base of pectoral filaments) less than upper-jaw length; pectoral filaments
5; first filament shortest, not reaching to level of pelvic-fin origin; fifth filament longest, its length 28 to 45% (mean 37%) of
standard length, extending well beyond level of posterior tip of pelvic fin; caudal fin deeply forked, upper and lower caudal-fin
lobes very long, filamentous, upper caudal-fin lobe 39 to 71% (mean 58%) and lower caudal-fin lobe 36 to 78% (mean 60%)of standard length. Pored lateral-line scales 69 to 72 (mode 70); lateral line simple, extending from upper end of gill opening
to upper end of lower caudal-fin lobe; scale rows above lateral line 7 or 8 (mode 8), below 9 to 12 (mode 10). Gillrakers 8 or 9
(mode 8) on upper limb, 10 to 12 (mode 11) on lower limb, 18 to 21 (mode 19) total. Vertebrae 10 precaudal and 14 caudal;
supraneural bones 2. Swimbladder with many appendages inserted into lateral walls of abdominal cavity. Colour: Headand upper sides of trunk tinged slightly blackish brown, becoming deep black on lower sides; snout and abdominal regions
blackish; membranes of first and second dorsal fins and caudal fin blackish, distal part of these fins black; pectoral-fin
membrane deep black; origin of pectoral filaments dusky yellowish, grading to blackish posteriorly; pelvic-fin origin dusky
yellowish, other parts dusky white.
Geographical Distribution: Although
Leptomelanosoma indicum has been
considered to be widely distributed in
the Indo-West Pacific, ranging fromSouth Africa to Indonesia (e.g. Smith,
1949; Smith in Smith and Heemstra,
1986; Menon and Babu Rao in Fischer
and Banchi, 1984; Feltes in Carpenterand Niem, 2001), report s of thespecies from the east coast of Africa
must presently remain unsubstantiated, owing to the examination ofa specimen (SAIAB 19493, 325 mmstandard length) from South Africa,
previously identified and reported as
Polydactylus indicus (= L. indicum),showing it to represent Polydactylus
plebeius. The species is currently
confirmed from Pakistan to Papua
New Guinea (Fig. 62). The record from
Papua New Guinea (Kerema Bay, Gulf
of Papua) is based only on a single
specimen (CSIRO A. 3028, 120 mmstandard length).

200 m

Fig. 62 Leptomelanosoma indicum
Known distribution

Habitat and Biology: Mainly occurs near estuaries, sometimes entering rivers, and is occasionally taken from depths to
100 m. Young of the species feed on small crustaceans, including shrimps and prawn larvae, but become piscivorous withincreasing growth.
Kagwade (1969) noted the occurrence of hermaphroditism in L. indicum from India, reporting the sex composition of 83
specimens as: hermaphrodites 39.76%, males 19.28% and females 40.96%. According to Nayak (1959a), 1-year-olds
attain 345 mm total length, 2-year-olds 545 mm total length and 3-year-olds 745 mm total length (based on trawl samples
from Dwarka, India). Kagwade (1970) considered that specimens from 910 to 1000 mm total length (the dominant sizegroup in the fishery) were 5-year-olds.
Nayak (1959a) found L. indicum to have 2 peak spawning periods, during April to June and October to December, aroundMumbai, northwestern India, the first period being the major one. On the other hand, Kuthalingam (1960) recorded intensive
spawning of the species around Chennai, southeastern India, during October to November. Karekar and Bal (1960)
reported the size of L. indicum at first maturity to be about 800 mm standard length.
Nayak (1959a) and Karekar and Bal (1960) described the following ova sizes of L. indicum from India: immature ovaries
with ova measuring up to 0.3 mm diameter, maturing ovaries with ova between 0.31 and 0.62 mm diameter, and mature
ovaries with ova between 0.63 and 1.10 mm diameter. Oil globules in mature ova were 0.26 to 0.40 mm diameter. According
to Kuthalingam (1960) and Kagwade (1970), planktonic eggs in this species averaged 1.3 mm diameter, with a large oilThreadfins of the World

35

globule of 0.5 mm diameter. Scale rudiments were apparent on larvae measuring 24.8 mm, post-larvae reaching thejuvenile stage at 31.8 mm.
Size: Maximum total length at least 1.4 m (Kagwade, 1970).
Interest to Fisheries: One of the most important species in fisheries of India, Sri Lanka, Bangladesh, Myanmar, Thailand,
Malaysia and Singapore, being caught by gill nets, trawls, handlines and beach seines. According to Nayak (1959a) and
Kagwade (1970), during 1950 and 1951 and 1956 and 1957, catches of L. indicum in India accounted for between 0.89 and26.0% of the total landings.
Local Names: BANGLADESH: Lakhua; INDIA: Indian threadfin; INDONESIA: Kurau hitam; MALAYSIA: Kurau, Kurow,
Mancong, Senangin; MYANMAR: Ka ku yan; SRI LANKA: Tahlunkala.
Literature: Motomura and Iwatsuki (2001a).
Remarks: Polynemus indicus (= Leptomelanosoma indicum) was proposed by Shaw (1804), for the “Maga Booshee” ofRussell (1803), whose description of the species included a correct figure (fig. 184), but lacked a formal scientific name andgave no indication of any type specimens. Although the description of P. indicus by Shaw (1804) and that earlier by Russell(1803) were very poor, Russell’s drawing clearly indicates 5 pectoral filaments, long caudal-fin lobes and a greyish black
body. Leptomelanosoma indicum can be easily distinguished from other polynemid species by the combination of thecharacters. Subsequently, Hamilton (1822) described Polynemus sele as a new species from estuaries of the Ganges River,India. This description also gave no indication of any type specimens. However, the diagnostic characters of 5 pectoral
filaments and long caudal-fin lobes given in the original description of P. sele are consistent with those of L. indicum. An
initial unpublished plate of P. sele, subsequently published by McClelland (1843, pl. 6), also shows 5 pectoral filaments, longcaudal-fin lobes and a greyish black body. Furthermore, Hamilton (1822) wrote; “The Maga booshee [= P. indicus] of
Russell has certainly a strong resemblance to the Sele [= P. sele] of the Ganges.” Although P. sele was said to differ from the“Maga Booshee” of Russell (1803) in having 7 first dorsal-fin spines (versus 8 spines in the latter) and 14 second dorsal-fin
rays (15 rays), the first spine of the first dorsal fin was apparently overlooked by Hamilton (1822) due to its very small size.
Cuvier in Cuvier and Valenciennes (1829) described Polynemus uronemus, as a new name for P. indicus. Furthermore,
McClelland (1843) described P. gelatinosus, from Bengal, India, as a new name for P. sele, and included Hamilton’s (1822)
unpublished plate. Both new names, P. uronemus and P. gelatinosus, are junior synonyms.

Parapolynemus Feltes, 1993
Parapolynemus Feltes, 1993: 207 (type species: Polynemus verekeri Saville-Kent, 1889).
Synonyms: None.
Species: The genus comprises a single species:

Parapolynemus verekeri: Northern Australia and southern Papua New Guinea.
Remarks: Parapolynemus has been recently proposed for the polynemid fish, Polynemus verekeri. According to Feltes(1993), Parapolynemus differs from other genera in having the following unique characters: highly porous skeleton; urohyal
with several longitudinal ridges on lateral surface of lamella; 3 infraorbitals; haemal spine of eleventh vertebra with prominent
medial anterior-posterior expansion.

Parapolynemus verekeri (Saville-Kent, 1889)

Fig. 63; Plate IId

Polynemus verekeri Saville-Kent, 1889: 234 [type locality: Ord River, Cambridge Gulf, northern Australia; lectotype (AMS I.
2770, 72 mm standard length) designated by Feltes, 1993; paralectotype (AMS I. 2771, 54 mm standard length)].
Synonyms: Polynemus intermedius Nichols, 1954: 3 [type locality: Merauke River, Papua New Guinea; holotype (AMNH
20114, 79 mm standard length); 16 paratypes (AMNH 20115, 74 mm standard length; AMNH 20116, 15 specimens, 37 to 79
mm standard length)].

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FAO Species Catalogue for Fishery Purposes No. 3

FAO Names: En - Dwarf paradise fish; Fr - Barbure paradis nain; Sp - Barbudo paraíso enano.

Fig. 63 Parapolynemus verekeri
Diagnostic Features: A small species. Body and head elongate, body depth at first dorsal-fin origin 20 to 32% (mean 25%) of
standard length; head length 26 to 32% (mean 29%) of standard length. Interorbital concave in larger specimens. Adipose
eyelid poorly developed; eye diameter less than snout length. Lip on lower jaw well developed, dentary teeth restricted to dorsal
surface in smaller specimens, but in larger specimens (over about 70 mm standard length) anterior one-third of lower jaw withsmall teeth extending onto lateral surface, adjacent portion of lip poorly developed; width of tooth band on upper and lower jaws
less than space (on symphysis) separating tooth bands on opposing premaxillae; teeth villiform in broad bands on jaws,
palatines and ectopterygoids, tooth plates on palatines longer than those on ectopterygoids; vomerine teeth absent. Posterior
margin of maxilla extending well beyond level of posterior margin of adipose eyelid; maxilla covered with deciduous scales.
Posterior margin of preopercle not serrated, but with a single spine posteroventrally; upper-jaw length 16 to 19% (mean 18%) of
standard length. Basisphenoid absent; sphenotics not visible dorsally between anterior margins of parietal and pterotic. First
dorsal fin with VIII spines; second dorsal fin with I spine and 11 to 14 (mode 13) soft rays; anal fin with III spines and 10 to 12(mode 11) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 12 to 14 (mode 13) rays, its length 37 to
51% (mean 44%) of standard length, posterior tip extending beyond level of posterior part of anal-fin base; pectoral-fin insertion
near midline of body; pectoral-fin base (including base of pectoral filaments) less than upper-jaw length; pectoral filaments 6 or
7 (mode 7); first filament shortest, extending beyond level of anal-fin origin; fifth filament longest, its length 47 to 104% (mean79%) of standard length, extending well beyond level of posterior tip of caudal fin; caudal fin deeply forked, upper and lower
caudal-fin lobes very long, filamentous, upper caudal-fin lobe 37 to 86% (mean 60%) of standard length. Pored lateral-line
scales 50 to 60 (mode 56); lateral line simple, extending from upper end of gill opening to upper end of lower caudal-fin lobe;
scale rows above lateral line 4 to 6 (mode 5), below 8 to 11 (mode 10). Gillrakers 30 to 43 (mode 40). Vertebrae 10 precaudal
and 14 caudal; supraneural bones 2. Swimbladder not apparent. Colour: Head and body yellow; upper sides of trunk tinged
slightly blackish brown; fins bright orange; pectoral filaments intense vermilion.
Geographical Distribution: Currently known from southern
Papua New Guinea (Merauke River and Gulf of Papua) tonorthern Australia (Cambridge Gulf of Western Australia to
Port Stuart of Northern Territory) (Fig. 64).
Habitat and Biology: Occurs in muddy estuaries and lower
portions of rivers (Saville-Kent, 1889; Munro, 1964, 1967).
According to Feltes in Carpenter and Niem (2001), hundredsof adults approaching breeding condition were taken in trawl
on Medusa Banks at a depth of 9 m in January or February
1968.
Size: Maximum standard length at least 11 cm (Feltes, 1993).
Interest to Fisheries: None.
Local Names: PAPUA NEW GUINEA: Streamered tasselfish.
Literature: Feltes (1993); Feltes in Carpenter and Niem (2001).
200 m

Remarks: P. verekeri is one of the smallest polynemids.
Munro (1964, 1967) suggested the possibility of Polynemus
intermedius being a junior synonym of Parapolynemus
verekeri. Feltes (1993) later confirmed that synonymy on thebasis of the types.

Fig. 64 Parapolynemus verekeri
Known distribution

Threadfins of the World

37

Pentanemus Günther, 1860
Pentanemus Günther, 1860: 330 (type species: Polynemus quinquarius Linnaeus, 1758).
Synonyms: None.
Species: The genus comprises a single species:

Pentanemus quinquarius: West coast of Africa, Senegal to Angola.
Remarks: Günther (1860) proposed Pentanemus for Polynemus quinquarius on the basis of internal and external
characters. Pentanemus differs from other genera in having the following characters: anal-fin rays 24 to 30 (less than 18 in
all other genera); anal-fin base greater than head length (unique character among the family); width of tooth band on upperand lower jaws less than space (on symphysis) separating tooth bands on opposing premaxillae (this character also present
in Filimanus); posterior margin of preopercle not serrated (this character also present in Parapolynemus).

Pentanemus quinquarius (Linnaeus, 1758)

Fig. 65; Plate IIe

Polynemus quinquarius Linnaeus, 1758: 317 [type locality: America (but clearly erroneous); no types known, see Dagetand Njock in Daget et al., 1986].
Synonyms: Polynemus artedii Bennett, 1831: 146 (type locality: Atlantic coast of north Africa; no types known). Polynemus
macronemus Pel, 1851: 9 [type locality: Gold Coast, Gulf of Guinea; 4 syntypes (RMNH 6016, 116 to 169 mm standard
length)].
FAO Names: En - Royal threadfin; Fr - Capitaine royal; Sp - Barbudo real.

Fig. 65 Pentanemus quinquarius

Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 26 to 33% (mean 30%) of standard
length; head length 27 to 31% (mean 30%) of standard length. Adipose eyelid poorly developed; eye diameter greater thansnout length. Lip on lower jaw well developed, dentary teeth restricted to dorsal surface; width of tooth band on upper andlower jaws narrower than space (on symphysis) separating tooth bands on opposing premaxillae; teeth villiform in broad
bands on jaws, palatines and ectopterygoids, tooth plates on palatines shorter than those on ectopterygoids; tooth plates onpalatines conspicuously small; vomerine tooth plate covered with skin, teeth absent. Posterior margin of maxilla extending
beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 15% (mean 14%) of standard length; maxillary

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FAO Species Catalogue for Fishery Purposes No. 3

scales absent. Posterior margin of preopercle not serrated. Basisphenoid in contact with prootic; sphenotics not visible
dorsally between anterior margins of parietal and pterotic. First dorsal fin with VIII spines; second dorsal fin with I spine and
14 or 15 (mode 15) soft rays; anal fin with III spines and 24 to 30 (mode 28) soft rays, anal-fin base longer than second
dorsal-fin base; pectoral fin with 14 to 16 (mode 15) unbranched rays, its length 30 to 42% (mean 36%) of standard length,
posterior tip reaching to or just short of level of midpoint of anal-fin base; pectoral-fin insertion well below midline of body;
pectoral-fin base (including base of pectoral filaments) less than upper-jaw length; pectoral filaments 5; first filament
shortest, just reaching to or extending beyond level of anal-fin origin; second to fifth pectoral filaments extending well beyond
level of posterior tips of caudal-fin lobes; third pectoral filament longest, its length 242 to 296% (mean 266%) of standard
length; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 36 to 46% (mean42%) and lower caudal-fin lobe 38 to 47% (mean 43%) of standard length. Pored lateral-line scales 68 to 76 (mode 72);
lateral line simple, extending from upper end of gill opening to mid-distal margin of caudal-fin membrane; scale rows above
lateral line 8 or 9 (mode 9), below 15 or 16 (mode 16). Gillrakers 18 to 23 on upper limb, 28 to 32 on lower limb, 47 to 53 total.
Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present, well developed. Colour: Upper sides ofhead and trunk with golden tinge, becoming silver on lower sides; margins of first and second dorsal fins, and caudal fins
black, remaining parts blackish; pelvic and anal fin white; pectoral fin yellow with melanophores; base of pectoral filaments
white, becoming blackish on posterior tips.
Geographical Distribution: Generally distributed
from Senegal to Angola, west coast of Africa (Fig. 66).
Günther (1860) reported the species from Cuba, West
Indies, on the basis of a single “half grown” specimen,
but apparently no other western Atlantic specimens
have been taken.
Habitat and Biology: Generally taken on muddy
bottoms in shallow coastal waters from depths of 10 to
70 m, and frequently found in estuaries and lagoons.
Generally feeds on crustaceans and small fishes(Longhurst, 1960: table 1).
Longhurst (1965) studied the biology, including
sexuality and growth, of P. quinquarius in Lagos,
Nigeria. The species has a normal bisexual
reproductive cycle, the sex composition of 25 292 adultspecimens: males 44.96%, hermaphrodites less than
0.01%, and females 55.04%. Spawning occurs in all
months, peaking in the dry season and almost ceasing
during the rainy season in Nigerian waters. Matures atless than 6 months of age (about 150 mm total length).
At 3 months, the species averages 100 mm total
length, 175 mm at 6 months and about 250 mm at 1
year. According to Longhurst (1965), it is unlikely thatmany individuals survive a second year of life.
Size: Maximum total length 35 cm, common to 25 cm(Allen in Fischer et al., 1981).

200 m

Fig. 66 Pentanemus quinquarius
Known distribution

Interest to Fisheries: One of the most important fisheries’ species off the west coast of Africa. Caught mainly by trawl, butsometimes by gill net and beach seine.
Local Names: ANGOLA: Barbudo; CAPE VERDE: Barbudo-real; GHANA: Sikor, Threadfin; GUINEA: Gbalakassa;
SENEGAL: Capitaine, Njaane, Njaane jaara; SIERRA LEONE: Bearbear, Beardfish.
Literature: Allen in Fischer et al. (1981); Daget and Njock in Daget et al. (1986); Njock in Quéro et al. (1990).
Remarks: Polynemus artedii and P. macronemus have been treated as junior synonyms of Pentanemus quinquarius (e.g.
Daget and Njock in Daget et al., 1986; Njock in Quéro et al., 1990). Although Bennett (1831) gave no indication of any typespecimens, he stated that Polynemus artedii has 25 anal-fin soft rays, which is diagnostic of Pentanemus quinquarius (thiscount not found in all other members of the family). Examination of types of Polynemus macronemus also showed them tobe conspecific with Pentanemus quinquarius.

Threadfins of the World

39

Polydactylus Lacepède, 1803
Polydactylus Lacepède, 1803: 419 [type species: Polydactylus plumierii Lacepède, 1803, presently regarded as a juniorsynonym of Polydactylus virginicus (Linnaeus, 1758)].
Synonyms: None.
Diagnostic Features: Body oblong to moderately deep. Adipose eyelid well developed; eye diameter greater than snout
length. Lip on lower jaw well developed, dentary teeth restricted to dorsal surface except in adults of Polydactylus
opercularis (in adults of P. opercularis, anterior one-third of lower jaw with small teeth extending onto lateral surface,
adjacent portion of lip poorly developed); width of tooth band on upper and lower jaws greater than space (on symphysis)
separating tooth bands on opposing premaxillae; teeth villiform in broad bands on jaws, palatines and ectopterygoids;
vomerine tooth plate without teeth in some species. Posterior margin of maxilla just short of, reaching to, or extending
beyond level of posterior margin of adipose eyelid. Posterior margin of preopercle serrated. Basisphenoid in contact withprootic; sphenotics not visible dorsally between anterior margins of parietal and pterotic. First dorsal fin with VIII spines;
second dorsal fin with I spine and 11 to 15 soft rays; anal fin with III spines and 10 to 18 soft rays; anal-fin base less than headlength; pectoral fin with 12 to 18 rays; pectoral-fin insertion well below midline of body; pectoral-fin base (including base ofpectoral filaments) less than upper-jaw length; pectoral filaments 4 to 9, not extending beyond level of posterior tip of caudal
fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous. Pored lateral-line scales 45 to 94; scale rows
above lateral line 5 to 11, below 8 to 16. Gillrakers 21 to 38. Vertebrae 10 precaudal and 14 caudal; supraneural bones 2
or 3. Swimbladder present or absent.
Habitat and Biology: Polydactylus species are generally found on sandy or muddy bottoms in coastal waters andestuaries, and in depths of 1 to 150 m. Only a single species, Polydactylus macrophthalmus, occurs in complete
fresh-water rivers on the Indonesian islands. Most species of Polydactylus are epibenthic predators feeding on crustaceans
and fishes on or near the bottom. The reproduction of a few species has been studied, and they appear to be protandrous,
sex changing from male to female with growth.
Geographic Distribution: The genus is widely distributed in tropical and subtropical latitudes of all oceans, but has neverbeen recorded in the Red Sea or Mediterranean. Fourteen species occur in the Indo-Pacific, 2 species are known from theeastern Pacific, 3 species are found in the western Atlantic, and 1 species occurs in the eastern Atlantic.
Interest to Fisheries: The species of Polydactylus are among the most important commercial fishes in tropical fisheries of
the world. Especially, in the middle East, South Asia and Southeast Asia, they are among the highest priced species in fish
markets. A few species (e.g. Polydactylus quadrifilis and P. macrochir) are known as important sport fishes. Polydactylus
sexfilis is cultivated in Hawaii.
Species: The genus comprises 20 species:
Polydactylus approximans: California to Peru (eastern Pacific)

Polydactylus bifurcus: Indonesia (Indo-Pacific)
Polydactylus longipes: Philippines (western Pacific)
Polydactylus macrochir: Northern Australia and southern Papua New Guinea (Indo-Pacific)
Polydactylus macrophthalmus: Indonesia (western Pacific)
Polydactylus malagasyensis: East coast of Africa (Indian Ocean)
Polydactylus microstomus: Sri Lanka to New Caledonia (Indo-Pacific)
Polydactylus mullani: Arabian Sea (Indian Ocean)
Polydactylus multiradiatus: Northern Australia and southern Papua New Guinea (Indo-Pacific)
Polydactylus nigripinnis: Northern Australia and southern Papua New Guinea (Indo-Pacific)
Polydactylus octonemus: New York to Yucatán (western Atlantic)
Polydactylus oligodon: Florida to Brazil (western Atlantic)
Polydactylus opercularis: California to Peru (eastern Pacific)
Polydactylus persicus: Persian Gulf (Indian Ocean)
Polydactylus plebeius: South Africa to Polynesia (Indo-Pacific)
Polydactylus quadrifilis: Eastern Atlantic
Polydactylus sexfilis: Mauritius to Pitcairn Island (Indo-Pacific)
Polydactylus sextarius: India to Papua New Guinea (Indo-Pacific)
Polydactylus siamensis: Thailand (Indo-Pacific)
Polydactylus virginicus: New Jersey to Brazil (western Atlantic)

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FAO Species Catalogue for Fishery Purposes No. 3

Remarks: Polydactylus plumierii, described as a new genus and species by Lacepède (1803), is presently regarded as a
junior synonym of Polydactylus virginicus. Polydactylus differs from other genera in having the following combination ofcharacters: eye diameter 1.3 or less in snout length (1.3 or more in Parapolynemus and Polynemus); width of tooth band on
upper and lower jaws greater than space separating tooth bands on opposing premaxilla (narrower in Filimanus andPentanemus); lip of anterior parts of lower jaw well or poorly developed (absent in Eleutheronema); basisphenoid in contact
with prootic (not in contact in Filimanus); pectoral-fin base, including base of pectoral filaments, less than upper-jaw length(greater in Galeoides); anal-fin soft rays less than 18 (more than 24 in Pentanemus); swimbladder absent or simple, notextending beyond anal-fin origin (swimbladder with many appendages in Leptomelanosoma and extending beyond anal-fin
origin in Galeoides).

Key to the Indo-Pacific Species of Polydactylus
1a. A large black spot present anteriorly on lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .® 2
1b. No large black spot anteriorly on lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
2a. Pectoral filaments 5; body and fins tinged with yellowish silver (Fig. 67, Plate IIId) . . . Polydactylus microstomus(India to New Caledonia)

2b. Pectoral filaments 6 or 7; body and fins tinged with silver . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Pectoral filaments 7; second spine of first dorsal fin more robust than other dorsal-fin spines (Fig. 68,
Plate IIIe) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus mullani(northern Arabian Sea)

3b. Pectoral filaments 6; all first dorsal-fin spines of similar thickness . . . . . . . . . . . . . . . . . . . . . . . ® 4
2nd dorsal-fin spine
more robust thanother spines

5 pectoral filaments

body and fins tinged
with yellowish silver
7 pectoral filaments

Fig. 68 Polydactylus mullani

Fig. 67 Polydactylus microstomus

4a. Gillrakers 25 to 30 (mode 28); second
dorsal-fin spine short (mean 6% of
standard length); swimbladder atrophied(Fig. 69, Plates Ve-f) . . . . . . Polydactylus sextarius

2nd dorsal-fin spine length 6%(mean) of standard length(India to Papua New Guinea)
4b. Gillrakers 29 to 35 (mode 31); second
dorsal-fin spine long (mean 7% of
standard length); swimbladder welldeveloped . . . . . . . . . . . . . . . . . . . . . ® 5
swimbladder atrophied
25-30 gillrakers

Fig. 69 Polydactylus sextarius

Threadfins of the World

41

5a. Pectoral-fin rays 14; scale rows below lateral line 10 (rarely 9); palatines inwardly turned anteriorly; pectoral fin long (mean 24% of standard length); pectoral filaments short (mean 29% of standard length)(Fig. 70, Plate IIIc) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus malagasyensis(east coast of Africa and Madagascar)
5b. Pectoral-fin rays 12 to 14 (mode 12); scale rows below lateral line 9 (rarely 8); palatines straight anteriorly; pectoral fin short (mean 19% of standard length); pectoral filaments long (mean 32% of standard
length) (Fig. 71, Plate IVc) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus persicus(Persian Gulf)
pectoral-fin
length 24%(mean) ofstandard length

palatines
inwardly
turnedanteriorly

10 (rarely 9) scale rows
below lateral line

14 pectoral-fin rays
pectoral-filament length 29%(mean) of standard length

pectoral-fin
length 19%(mean) ofstandard length

9 (rarely 8) scale rows
below lateral line

palatines
straight
anteriorly

12-14 (mode 12)
pectoral-fin rays

pectoral-filament length 32%(mean) of standard length

Fig. 70 Polydactylus malagasyensis

Fig. 71 Polydactylus persicus

6a. Pectoral filaments 5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6b. Pectoral filaments 6 or 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
7a. Posterior margin of maxilla extending considerably beyond level of posterior margin of adipose eyelid;
occipital profile concave in adults (Fig. 72, Plate IIIa) . . . . . . . . . . . . . . . . . . Polydactylus macrochir(northern Australia and southern Papua New Guinea)
7b. Posterior margin of maxilla reaching to or extending only slightly beyond level of posterior margin of adipose eyelid; occipital profile nearly straight throughout life . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Lateral line bifurcated on caudal-fin base, extending to posterior margins of upper and lower caudal-fin
lobes; second spine of first dorsal fin more robust than other spines (Fig. 73, Plate IIg) . . . Polydactylus bifurcus(Indonesia)
8b. Lateral line unbranched on caudal-fin base, extending to posterior margins of lower caudal-fin lobe;
thickness of all spines of first dorsal fin similar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
2nd spine more robust
than other spines
occipital profile
concave in adults

occipital profile
nearly straightlateral line bifurcatedon caudal-fin baseposterior margin of maxilla extending
considerably beyond level of posterior
margin of adipose eyelid

Fig. 72 Polydactylus macrochirFig. 73 Polydactylus bifurcus

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FAO Species Catalogue for Fishery Purposes No. 3

9a. Pored lateral-line scales 60 to 68 (mode 63); gillrakers 24 to 32 (mode 26); scale rows above lateral line
8 or 9 (mode 8), below 12 or 13 (mode 12); upper jaw short (mean 15% of standard length) (Fig. 74,
Plates IVd-h) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus plebeius(South Africa to French Polynesia)
9b. Pored lateral-line scales 54 to 58 (mode 54); gillrakers 22 to 24 (mode 23); scale rows above lateral line
7, below 10 or 11 (mode 11); upper jaw long (mean 17% of standard length) (Fig. 75, Plate Vg). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus siamensis(Thailand)
8-9 scale rows
above lateral line

upper-jaw
length 15%(mean) ofstandard
length24-32
gillrakers

54-58 pored lateral-line scales

7 scale rows
above lateral line

60-68 pored lateral-line scales

upper-jaw
length 17%(mean) ofstandard
length12-13 scale rows below lateral line

Fig. 74 Polydactylus plebeius

10-11 scale rows below lateral line
22-24
gillrakers

Fig. 75 Polydactylus siamensis

10a. Longest pectoral filament extending beyond level of midpoint of caudal peduncle . . . . . . . . . . . . . . ® 11
10b. Longest pectoral filament not reaching to level of midpoint of caudal peduncle . . . . . . . . . . . . . . . ® 12
11a. Pectoral filaments 6; pored lateral-line scales 54 or 55 (mode 55); scale rows above lateral line 6, below
11; gillrakers 31 or 32 (mode 31) (Fig. 76, Plate IIh) . . . . . . . . . . . . . . . . . . . . Polydactylus longipes(Philippines)
11b. Pectoral filaments 7; pored lateral-line scales 87 to 94 (mode 88); scale rows above lateral line 10 to 12(mode 11), below 15 or 16 (mode 16); gillrakers 25 to 29 (mode 27) (Fig. 77, Plate IIIb). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus macrophthalmus(Indonesia)
6 scale rows
above lateral line

54-55 pored lateral-line scales

10-12 scale rows
above lateral line

87-94 pored lateral-line scales

25-29
gillrakers

31-32
gillrakers
6 pectoral
filaments

7 pectoral
filaments

11 scale rows below lateral line

15-16 scale rows below lateral line

Fig. 77 Polydactylus macrophthalmus

Fig. 76 Polydactylus longipes

12a. Anal-fin soft rays 16 to 18 (mode 16);
pectoral filaments 7 (rarely 6) (Fig. 78,
Plate IIIf) . . . . . . . . . Polydactylus multiradiatus(northern Australia and southern Papua New Guinea)
12b. Anal-fin soft rays 11 or 12; pectoral
filaments 6 . . . . . . . . . . . . . . . . . . . . ® 13

7 (rarely 6) pectoral
filaments

16-18 anal-fin soft rays

Fig. 78 Polydactylus multiradiatus

Threadfins of the World43

13a. Vomer without teeth; pored lateral-line scales 46 to 50 (mode 47); swimbladder absent; orbit diametersmall (mean 6% of standard length) (Fig. 79, Plate IIIg) . . . . . . . . . . . . . . . . . Polydactylus nigripinnis(northern Australia, and southern Indonesia and Papua New Guinea)
13b. Vomer with teeth; pored lateral-line scales 61 to 67 (mode 64); swimbladder present; orbit diameter
large (mean 9% of standard length) (Fig. 80, Plates Vb-d) . . . . . . . . . . . . . . . . . . Polydactylus sexfilis(Mauritius to Pitcairn Island)
orbit diameter 6% (mean)of standard length

vomer
without teeth

46-50 pored
lateral-line scales

orbit diameter 9%(mean) ofstandard length61-67 pored
lateral-line scales

vomer withteeth
swimbladder absent

swimbladder present

Fig. 80 Polydactylus sexfilisFig. 79 Polydactylus nigripinnis

Key to the Eastern Pacific Species of Polydactylus
1a. Pectoral filaments 6 (rarely 5 or 7); pored lateral-line scales 56 to 63 (mode 60); gillrakers 26 to 31(mode 30); lateral line bifurcated on caudal-fin base, extending to posterior margins of upper and lower
caudal-fin lobes (Fig. 81, Plate IIf) . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus approximans(California to Peru)
1b. Pectoral filaments 9 (rarely 8); pored lateral-line scales 66 to 74 (mode 70); gillrakers 31 to 37 (mode34); lateral line unbranched on caudal-fin base, extending to posterior margins of lower caudal-fin lobe
(Fig. 82, Plate IVb) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus opercularis(California to Peru)
66-74 pored
lateral-line scales

56-63 pored
lateral-line scales

26-31
gillrakers

lateral line unbranchedon caudal-fin baselateral line bifurcatedon caudal-fin base
6 (rarely 5 or 7) pectoral filaments

Fig. 81 Polydactylus approximans

31-37
gillrakers

9 (rarely 8) pectoral filaments

Fig. 82 Polydactylus opercularis

Key to the Atlantic Species of Polydactylus
1a. Pectoral filaments 4 (Fig. 83, Plate Va). . . . . . . . . . . . . . . Polydactylus quadrifilis(Senegal to the Congo, east coast of Africa)
1b. Pectoral filaments 7 or 8 (rarely 9) . . . . . . . ® 2
4 pectoral filaments

Fig. 83 Polydactylus quadrifilis

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FAO Species Catalogue for Fishery Purposes No. 3

2a. Pectoral filaments 8 (rarely 9);
gillrakers 34 to 38 (mode 36) (Fig. 84,
Plate IIIh) . . . . . . . . . . Polydactylus octonemus(New York to Yucatán, western Atlantic)
2b. Pectoral filaments 7 (rarely 8);
gillrakers 22 to 30 (mode 27 or 28) . . . . . . . . ® 3
3a. Pored lateral-line scales 67 to 73(mode 70); anal-fin soft rays 13 to 15(mode 14) (Fig. 85, Plate IVa) . Polydactylus oligodon(Florida to Brazil, western Atlantic)

34-38
gillrakers

3b. Pored lateral-line scales 54 to 63(mode 58); anal-fin soft rays 11 to 14(mode 13) (Fig. 86, Plate Vh) . Polydactylus virginicus

8 (rarely 9) pectoral filaments

Fig. 84 Polydactylus octonemus(New Jersey to Brazil, western Atlantic)
54-63 pored
lateral-line scales

67-73 pored
lateral-line scales

13-15 anal-fin soft rays

Fig. 85 Polydactylus oligodon

Polydactylus approximans (Lay and Bennett, 1839)

11-14 anal-fin soft rays

Fig. 86 Polydactylus virginicus

Fig. 87; Plate IIf

Polynemus approximans Lay and Bennett, 1839: 57 [type locality: San Blas and Mazatlán, Mexico, based on a brief sketch
and description by Mr Collie; type material apparently lost, see Motomura, Kimura and Iwatsuki, 2002].
Synonyms: Polynemus californiensis Thominot, 1886: 161 [type locality: California, USA; holotype (MNHN 1884-487,
179 mm standard length)].
FAO Names: En - Blue bobo; Fr - Barbure bleu; Sp - Barbudo azul.

Fig. 87 Polydactylus approximans

Threadfins of the World

45

Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 25 to 32% (mean 30%) of standard
length; head length 28 to 35% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight in young,becoming somewhat concave with growth. Posterior margin of maxilla not reaching to or extending slightly beyond level ofposterior margin of adipose eyelid; maxilla not covered with scales; upper-jaw length 13 to 15% (mean 14%) of standard
length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted todorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids; shape of vomerine tooth plate
crescentic or elliptic. Posterior margin of preopercle serrated. First dorsal fin with VIII spines, thickness of all first dorsal-fin
spine bases similar; second dorsal fin with I spine and 11 to 13 (mode 12) soft rays; anal fin with III spines and 13 to 15 (mode14) soft rays, anal-fin base greater than second dorsal-fin base; pectoral fin with 14 to 16 (mode 15) rays (all rays
unbranched), its length 24 to 32% (mean 28%) of standard length, posterior tip not reaching to or extending beyond level ofposterior tip of pelvic fin; pectoral filaments 6 (rarely 5 or asymmetrically 6 and 7), first filament shortest, just reaching to or
extending slightly beyond level of pelvic-fin origin; second to fourth pectoral filaments extending beyond level of pelvic-fin
origin; length of fifth pectoral filament variable, from not reaching level with posterior margin of pelvic fin to just reaching levelwith anal-fin origin; sixth pectoral filament longest, its length 35 to 48% (mean 41%) of standard length, just short of level ofanal-fin origin or reaching to near midpoint of anal-fin base; caudal fin deeply forked, upper and lower caudal-fin lobes notfilamentous, upper caudal-fin lobe 37 to 43% (mean 40%) and lower lobe 35 to 40% (mean 37%) of standard length. Poredlateral-line scales 56 to 63 (mode 60); lateral line bifurcated on caudal-fin membranes, extending to posterior margins ofupper and lower caudal-fin lobes; scale rows above lateral line 7 or 8 (mode 7), below 11 to 13 (mode 12). Gillrakers 11 to 14(mode 13) on upper limb, 15 to 18 (mode 17) on lower limb, 26 to 31 (mode 30) total. Vertebrae 10 precaudal and 14 caudal;
supraneural bones 3. Swimbladder present, small. Colour: Head and body grey dorsally, silver ventrally; anterior margins
of first and second dorsal fins black, remaining parts dusky; pectoral fin dusky; base of pectoral filaments white, becoming
blackish on posterior tips; base and posterior margin of pelvic fin white, remaining parts dusky; posterior margin of anal fin
white, remaining parts dusky; margins of caudal fin black, remaining parts dusky.
Geographical Distribution: Currently known from theeastern Pacific Ocean, where it ranges from Monterey
Bay, California, USA to Callao, Peru (Fig. 88), being
relatively common in the Central American region inlatitudes between 5° and 20°N. Grove and Lavenberg(1997) reported a single small example (LACM
W53-178) collected from the Galápagos Islands, and isconsidered to occur rarely around the islands. Although
Pequeño (1989) listed P. approximans andP. opercularis from Chile, such records remain
unsubstantiated.
Habitat and Biology: Although P. approximans
generally inhabits muddy or sandy bottoms in coastal
waters and estuaries, larvae and juveniles (less thanabout 40 mm standard length) sometimes occur at thewater surface several hundred kilometres offshore(Grove and Lavenberg, 1997). Most of the collection
data available indicated the specimens as having beentaken from depths less than 30 m. According to Groveand Lavenberg (1997), the species is omnivorous,
feeding on worms, sand crabs, shrimp, clams, andsometimes fish like small anchovy.

200 mFig. 88 Polydactylus approximans
Known distribution

Size: Maximum total length at least 35 cm (Allen and Robertson, 1994).
Interest to Fisheries: Esteemed as a food fish throughout the tropical eastern Pacific.
Local Names: COLOMBIA: Barbeta; EQUADOR: Aguapuro; MEXICO: Barbudo azul; NICARGUA: Barbudo seis barbas.
Literature: Motomura, Kimura and Iwatsuki (2002).
Remarks: P. californiensis, originally described by Thominot (1886) from California, USA on the basis of a single specimen(MNHN 1884-487, 179 mm standard length), was recently synonymized under P. approximans, originally described by Lay
and Bennett (1839) on the basis of a brief sketch and description by Mr Collie (Motomura, Kimura and Iwatsuki 2002).
Although Lay and Bennett (1839) quoted the absence of a swimbladder in the species from the brief description by Mr Collie,
the presence of a swimbladder was confirmed by Motomura, Kimura and Iwatsuki (2002). The swimbladder was apparently
overlooked by Mr Collie due to its very small size (about 13% of standard length).

Polydactylus approximans and 6 Indo-Pacific Polydactylus species, P. longipes, P. malagasyensis, P. nigripinnis,
P. persicus, P. sexfilis and P. sextarius, are characterized by having 6 pectoral filaments. However, P. approximans isdistinguished from the other 6 species by having a bifurcated lateral line (unbranched in the latter). The lateral-line conditionof P. approximans is also found in an Indian Ocean species, P. bifurcus, and all 3 currently-recognized Atlantic west coast
species, P. octonemus, P. oligodon and P. virginicus. Polydactylus approximans differs from the above 4 species in

46

FAO Species Catalogue for Fishery Purposes No. 3

having usually 6 pectoral filaments (5 in P. bifurcus, usually 7 in P. oligodon and P. virginicus, and usually 8 in
P. octonemus.

Polydactylus approximans has the maxilla without scales, a character state also found in the 3 Atlantic west coast species
and an African west coast species, P. quadrifilis. Other congeners (distributed in the Indo-Pacific and eastern Pacific) havethe maxilla with deciduous scales, the size and number of scales being both intra- and interspecifically variable.
Polydactylus approximans and P. opercularis occur in the eastern Pacific Ocean. The former can be easily distinguishedfrom the latter by having lower counts of pectoral filaments [6 on each side (rarely 5 on each side or asymmetrically 6 and 7)
versus 9 on each side (rarely 8 on each side or asymmetrically 8 and 9) in the latter], pored lateral-line scales [56 to 63 (mode60) versus 66 to 74 (mode 70)], scale rows above and below the lateral line [7 or 8 (mode 7) and 11 to 13 (mode 12) versus 7
to 10 (mode 8) and 12 to 15 (mode 13), respectively] and gillrakers [upper series 11 to 14 (mode 13), lower 15 to 18 (mode17) and total 26 to 31 (mode 30) versus 13 to 18 (mode 16), 17 to 20 (mode 19) and 31 to 37 (mode 34), respectively], andhigher counts of anal-fin soft rays [13 to 15 (mode 14) versus 12 to 14 (mode 13)].
In addition to the above meristic characters, P. approximans differs from P. opercularis in having the following distinct
morphological characters: lip on lower jaw well developed, dentary teeth restricted to dorsal surface (anterior parts of lower
jaw with villiform teeth extending onto lateral surface, adjacent portion of lip poorly developed in larger P. opercularis);
maxilla not covered with scales (covered with small deciduous scales in P. opercularis); lateral line bifurcated on caudal-fin
membranes, extending to posterior margins of upper and lower caudal-fin lobes (lateral line unbranched, extending to upper
end of lower caudal-fin lobe in P. opercularis); swimbladder present (absent in P. opercularis).

Polydactylus bifurcus Motomura, Kimura and Iwatsuki, 2001

Fig. 89; Plate IIg

Polydactylus bifurcus Motomura, Kimura and Iwatsuki, 2001: 299, figs. 1-3 [type locality: Kuta beach, Lombok Island,
Indonesia; holotype (NSMT-P 60494, 144 mm standard length)].
Synonyms: None.
FAO Names: En - Slender fivefinger threadfin; Fr - Barbure svelte; Sp - Barbudo esbelto.

Fig. 89 Polydactylus bifurcus
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 26 to 28% (mean 27%) of standard
length; head length 27 to 28% (mean 28%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla extending slightly beyond level of posterior margin of adipose eyelid; depth of posterior margin of maxilla
less than eye diameter; upper-jaw length 14% of standard length; lip on lower jaw well developed, dentary teeth restricted todorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, second spine more robust than others; second dorsal fin with I spine and 13 softrays; anal fin with III spines and 11 or 12 (mode 11) soft rays, anal-fin base approximately equal to second dorsal-fin base;pectoral fin with 15 rays (several rays branched), its length 19 to 21% (mean 20%) of standard length, posterior tip notreaching to level of posterior tip of pelvic fin; pectoral filaments 5, first filament shortest, not extending beyond level ofThreadfins of the World47

posterior tip of pelvic fin; second, third, and fifth filaments not extending beyond level of posterior tip of pelvic fin; fourthfilament longest, its length 35 to 38% (mean 36%) of standard length, extending well beyond level of posterior tip of pelvic fin,but not reaching to level of anal-fin origin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper
caudal-fin lobe 37 to 39% (mean 38%) and lower lobe 37% of standard length. Pored lateral-line scales 69 to 72; lateral line
bifurcated on caudal-fin base, extending to posterior margins of upper and lower caudal-fin lobes; scale rows above lateralline 8 or 9 (mode 8), below 10 to 12 (mode 12). Gillrakers 12 or 13 (mode 13) on upper limb, 17 or 18 (mode 17) on lower
limb, 30 total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present, well developed. Colour:
Upper sides of head and trunk tinged silvery green, becoming more silver on lower sides; first and second dorsal fins darkgreen; pectoral fin white, becoming dark green on posterior tip; pectoral filaments and pelvic fin white; anterior part of anal fin
dark green, remainder of fin white; upper, lower and posterior margins of caudal fin black, remainder of fin dark green (lower
lobe darkest); 8 or 9 dark stripes along longitudinal scale rows above lateral line, 8 or 9 faint stripes below.
Geographical Distribution: Currently known only
from the south coast of the Greater Sunda Islands:Nias, Java and Lombok Island (Fig. 90).
Habitat and Biology: Collection data for the holotype
indicated that it was taken in shallow water (1.5 m) overa muddy bottom (Motomura, Kimura and Iwatsuki,2001). No other data are available.
Size: Maximum standard length at least 27 cm(Motomura, 2002).
Interest to Fisheries: None.
Local Names: None known.
Literature: Motomura, Kimura and Iwatsuki (2001);
Motomura (2002).
200 m

Remarks: P. bifurcus can be distinguished from allother Indo-Pacific Polydactylus species by having thelateral line bifurcated on the caudal-fin base, extending
Fig. 90 Polydactylus bifurcus
to the posterior margins of the upper and lower
Known distribution
caudal-fin lobes (an unbranched lateral line in all other
Polydactylus). However, 1 of 2 eastern Pacific species, P. approximans, and all of 3 Atlantic west coast species,
P. octonemus, P. oligodon and P. virginicus, have a similar lateral line condition to P. bifurcus. Polydactylus bifurcus
differs from the above 4 species in having 5 pectoral filaments (usually 6 in P. approximans, 8 in P. octonemus, and 7 inP. oligodon and P. virginicus). Polydactylus approximans rarely has 5 pectoral filaments, but the species is clearly
distinguished from P. bifurcus by the numbers of the pored lateral-line scales and anal-fin soft rays (55 to 60 and 13 or 14
versus 69 to 72 and 11 or 12, respectively, in the latter). The lateral lines of the other eastern Pacific species, P. opercularis,and 1 West African species, P. quadrifilis, are unbranched and extend to the upper end of the lower caudal-fin lobe.
Five species have been recognized as valid Indo-Pacific Polydactylus species with 5 pectoral filaments: P. bifurcus,
P. macrochir, P. microstomus, P. plebeius and P. siamensis. Polydactylus bifurcus further differs from P. microstomus inhaving higher counts of pored lateral-line scales and scales above and below the lateral line (69 to 72 and 8 or 9 / 10 to 12,
respectively versus 46 to 49 and 6 or 7 / 8 to 10, in the latter), fourth pectoral filaments longest (fifth longest in the latter),
teeth present on the vomer (absent in the latter) and a large black spot absent anteriorly on the lateral line (present, in thelatter).
Like P. bifurcus, P. plebeius and P. siamensis are characterized by several dark stripes along the longitudinal scale rows
above and below the lateral line and 5 pectoral filaments. However, P. bifurcus is distinguished from the others by having
higher counts of the pored lateral-line scales (69 to 72 versus 60 to 66, rarely 68 and 54 to 58 in P. plebeius and P. siamensis,
respectively), fourth pectoral filaments longest (fifth longest in P. plebeius and P. siamensis) and a very strong second spineof the first dorsal fin (more robust than other spines of first dorsal fin versus similar to other spines).

Polydactylus bifurcus is similar to P. macrochir in having higher counts of pored lateral-line scales and scales above andbelow the lateral line, fourth pectoral filaments longest, teeth present on the vomer, and a very strong second spine of thefirst dorsal fin. However, the former differs from the latter in having the posterior margin of the maxilla extending slightly
beyond the level of the posterior margin of the adipose eyelid (extending considerably beyond in the latter) and lower countsof gillrakers (30 versus 32 to 35 in P. macrochir).
Polydactylus bifurcus appears to be one of the rarest polynemids and only 3 specimens are present in museum collections(NSMT-P 60494, holotype, 144 mm standard length; ZMA 114-500, 272 mm standard length; ZMA 116-717, 199 mmstandard length).

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FAO Species Catalogue for Fishery Purposes No. 3

Polydactylus longipes Motomura, Okamoto and Iwatsuki, 2001

Fig. 91; Plate IIh

Polydactylus longipes Motomura, Okamoto and Iwatsuki, 2001: 1087, fig. 1 [type locality: Davao market, Mindanao Island,
Philippines; holotype (USNM 363173, 159 mm standard length); 2 paratypes (FSKU-P 19840, 134 mm standard length;
MUFS 20290, 152 mm standard length)].
Synonyms: None.
FAO Names: En - Long-limb threadfin; Fr - Barbure longs-doigts; Sp - Barbudo de dedos largos.

Fig. 91 Polydactylus longipes

Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 29 to 32% (mean 30%) of
standard length; head length 35% of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of
maxilla reaching to level of posterior margin of adipose eyelid; upper-jaw length 17 to 18% (mean 17%) of standard length;
depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted to dorsal
surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle serrated.
First dorsal fin with VIII spines, thickness of all first dorsal-fin spine bases similar; second dorsal fin with I spine and 12 softrays; anal fin with III spines and 11 soft rays, anal-fin base slightly less than second dorsal-fin base; pectoral fin with 13 rays
(all rays unbranched), its length 28 to 29% (mean 28%) of standard length, posterior tip reaching to or extending slightly
beyond level of posterior tip of pelvic fin; pectoral filaments 6, first filament shortest, extending well beyond level of pelvic-fin
origin; second pectoral filament reaching to or just short of level of posterior tip of pectoral fin; third pectoral filament reaching
to or just short of level of anal-fin origin; fourth pectoral filament extending beyond midpoint of anal-fin base; fifth pectoral
filament extending slightly beyond midpoint of caudal peduncle; sixth pectoral filament longest, its length 71 or 72% (mean71%) of standard length, extending well beyond midpoint of caudal peduncle; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 40 to 44% (mean 42%) and lower lobe 38 to 40% (mean 38%) of
standard length. Pored lateral-line scales 54 or 55 (mode 55); lateral line simple, extending from upper end of gill opening toupper end of lower caudal-fin lobe; scale rows above lateral line 6, below 11. Gillrakers 13 or 14 (mode 13) on upper limb, 18
on lower limb, 31 or 32 (mode 31) total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present,
well developed. Colour: Upper sides of head and trunk tinged greyish black, becoming lighter grey on lower sides; opercularregion black; all spines of first dorsal fin black, tip of membrane between third and fifth spines black, interspinous membraneotherwise transparent; pectoral-fin membranes greyish black; posterior-most portions of second dorsal and anal fins white,
all spines and first and second soft rays dusky, remainder greyish black; tip of anterior margin of pelvic fin white, remainder offin dusky; pectoral filaments and caudal fin greyish black.

Threadfins of the World

49

Geographical Distribution: Currently known only
from Mindanao Island, Philippines (Fig. 92).
Habitat and Biology: No data are available.
Size: Maximum standard length at least 16 cm(Motomura, Okamoto and Iwatsuki, 2001).
Interest to Fisheries: None.
Local Names: None known.
Literature: Motomura, Okamoto and Iwatsuki (2001);
Motomura (2002).
Remarks: Although P. longipes is similar to
P. macrophthalmus in having long pectoral filaments
which extend well beyond the midpoint of the anal-finbase, P. longipes differs from the latter in having 6
200 m
pectoral filaments (7 in the latter). In P. longipes theupper 2 pectoral filaments extend beyond the midpointof the caudal peduncle, not reaching the caudal-fin
Fig. 92 Polydactylus longipesbase. In P. macrophthalmus the upper 3 filaments
Known distribution
extend beyond the caudal-fin base, the fifth extending
beyond the caudal-fin lobe tip. Furthermore,
P. longipes is clearly distinguished from P. macrophthalmus by lower dorsal-fin soft ray counts (12 versus 13 or 14 in thelatter), fewer pored lateral-line scales (54 or 55 versus 87 to 94 in P. macrophthalmus) and scales above and below thelateral line (6 and 11 versus 10 to 12 and 15 or 16, respectively), and higher gill-raker counts (31 or 32 versus 25 to 29 in
P. macrophthalmus).

Polydactylus species with 6 pectoral filaments now include 6 valid species from the Indo-Pacific: P. longipes,
P. malagasyensis, P. nigripinnis, P. persicus, P. sexfilis and P. sextarius. Polydactylus nigripinnis further differs fromP. longipes in having higher pectoral-fin ray counts [16 to 18 (mode 17) versus 13 in the latter], lower counts of pored
lateral-line scales and gillrakers [46 to 50 (mode 47) and 24 to 29 (mode 27) versus 54 or 55 (mode 55) and 31 or 32 (mode31), respectively], and vomerine teeth absent (present in P. longipes).

Polydactylus sexfilis differs from P. longipes in having higher counts of pectoral-fin rays [15 or 16 (mode 16) versus 13 in thelatter], pored lateral-line scales [60 to 67 (mode 64) versus 54 or 55 (mode 55 in P. longipes)], and scales above and below
the lateral line [8 or 9 (mode 9) and 12 to 14 (mode 13) versus 6 and 11, respectively].
Polydactylus malagasyensis, P. persicus and P. sextarius are also clearly distinguished from P. longipes as follows: allpectoral-fin rays (except upper 1 or 2) branched (all unbranched in the latter), vomerine teeth absent (present), and a large
black spot present anteriorly on the lateral line (absent).
Polydactylus longipes appears to be one of the rarest polynemids, being known only from the 3 type specimens.

Polydactylus macrochir (Günther, 1867)

Fig. 93; Plate IIIa

Polynemus macrochir Günther, 1867: 60 [type locality: New South Wales, Australia (but probably Queensland, seeMotomura et al., 2000); holotype (BMNH 1866.2.13.17, 171 mm standard length)].
Synonyms: Polynemus sheridani Macleay, 1884: 21 (type locality: Mary River, Queensland, Australia; type material
apparently lost, see Motomura et al., 2000).

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FAO Species Catalogue for Fishery Purposes No. 3

FAO Names: En - King threadfin; Fr - Barbure gros-doigts; Sp - Barbudo de manos grandes.

Fig. 93 Polydactylus macrochir
Diagnostic Features: A large species. Body depth at first dorsal-fin origin 23 to 28% (mean 25%) of standard length; headlength 28 to 34% (mean 30) of standard length. Snout pointed; occipital profile nearly straight in young but concave in adults.
Posterior margin of maxilla extending considerably beyond level of posterior margin of adipose eyelid; upper-jaw length 13 to
17% (mean 15%) of standard length; depth of posterior margin of maxilla greater than eye diameter in adults (over about
200 mm standard length); lip on lower jaw well developed and dentary teeth restricted to dorsal surface; teeth villiform inbroad bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VIII
spines, second spine more robust than others; second dorsal fin with I spine and 11 to 13 (mode 12) soft rays; anal fin with III
spines and 10 to 12 (mode 11) soft rays, anal-fin base approximately equal to second dorsal-fin base; pectoral fin with 14 or
15 (mode 14) rays (all rays unbranched), its length 22 to 27% (mean 24%) of standard length, posterior tip just reaching to or
not reaching to level of posterior tip of pelvic fin; pectoral filaments 5, first filament shortest, extending well beyond level ofpelvic-fin origin; second pectoral filament extending well beyond level of posterior tip of pelvic fin; third and fifth pectoral
filaments extending beyond level of anal-fin origin; fourth pectoral filament longest, its length 40 to 53% (mean 46%) of
standard length, extending well beyond level of anal-fin origin; caudal fin deeply forked, upper and lower caudal-fin lobes notfilamentous, upper caudal-fin lobe 33 to 41% (mean 35%) and lower lobe 31 to 46% (mean 33%) of standard length. Poredlateral-line scales 70 to 76 (mode 72); lateral line simple, extending from upper end of gill opening to upper end of lower
caudal-fin lobe; scale rows above lateral line 8 to 11 (mode 9), below 12 to 15 (mode 12). Gillrakers 13 to 15 (mode 14) on
upper limb, 18 to 20 (mode 20) on lower limb, 32 to 35(mode 34) total. Vertebrae 10 precaudal and 14 caudal;
supraneural bones 3. Swimbladder present, welldeveloped. Colour: Upper sides of head and trunk
tinged golden silver, becoming more silver on lower
sides; first and second dorsal fins pale brown; pectoraland pelvic fins vivid yellow; anterior part of anal fin
yellow, other parts yellowish white; pectoral filaments
white or yellowish white; posterior margin of caudal fin
greyish black, other parts greyish.
Geographical Distribution: Endemic to northern
Australia and southern Papua New Guinea (Fig. 94).
The species is currently known from Broome, Western
Australia to Brisbane River, Queensland, northern
Australia, and from south coast of Irian Jaya, Indonesia
to south coast of Papua New Guinea. Girija Kumari,Ratnamala and Seshagiri-Rao (1985) reported that 9
specimens of Polydactylus macrochir (as Polynemus
sheridani) were collected from Bay of Bengal, India.
However, the description of those specimens given by
Girija Kumari, Ratnamala and Seshagiri-Rao (1985),
which includes 19 gillrakers, caudal-fin lobes
filamentous and greyish body colour, are closer to thediagnosis of Leptomelanosoma indicum.

200 m

Fig. 94 Polydactylus macrochir
Known distribution

Threadfins of the World

51

Habitat and Biology: Inhabits turbid coastal waters, estuaries and mangrove creeks as well as mangrove-lined rivers and istaken from depths of 0.2-6 m. Feeds mainly on crustaceans and fishes.
The species exhibits protandry. According to Kailola and Stewart in Kailola et al. (1993), individuals between 700 and
1 000 mm fork length may be transitional hermaphrodites. However, most less than 800 mm fork length are males and most
more than 950 mm fork length are females. In the Gulf of Carpentaria and along the northeastern Queensland coastline,
Australia, transitional P. macrochir are found in the months from June to September. In north Australian waters, it is known
to spawn over the summer months from October to February or March, with a peak during December.
Size: Although P. macrochir was stated to reach a weight of about 45 kg (Macleay, 1884), such examples are not seen today(Marshall, 1964). Kailola and Stewart in Kailola et al. (1993), recorded the species at 140 cm fork length and 32 kg in the Gulf
of Carpentaria, and more than 150 cm fork length in Princess Charlotte Bay. Validated ageing studies indicated that the
species can live for more than 20 years, with an estimated maximum attainable size of 170 cm fork length and more than
40 kg (Kailola and Stewart in Kailola et al., 1993).
Interest to Fisheries: P. macrochir is one of the most important species in the fisheries at Northern Territory, Queensland
and Western Australia. The species is caught mainly by coastal set gillnets, but also by fixed tidal traps, beach seines, ringnets and handlines. In the southern Gulf of Carpentaria, Australia, the fishery for the species is largely based on male 3- to
6-year-old fish (Kailola and Stewart in Kailola et al., 1993). From the Gulf of Carpentaria gillnet fishery between 1980 and
1987, the species averaged 30% by weight of total landings (Kailola and Stewart in Kailola et al., 1993). It is also an
important sport fish in northern Australian waters. They are caught with rod and reel or handlines, from shore or boat.
Local Names: AUSTRALIA: Burnett salmon, Gold threadfin, King salmon, King threadfin, Sheridan’s threadfin; PAPUANEW GUINEA: Longfinned threadfin.
Literature: Motomura et al. (2000); Motomura (2002).
Remarks: Polydactylus sheridani has been regarded as a valid species (e.g. Marshall, 1964; Munro, 1967; Menon, 1974;
Grant, 1982). However, the characters of P. sheridani, which includes maxilla large, posterior margin of maxilla extending
beyond posterior margin of eye and second spine of first dorsal fin very strong, as described by Macleay (1884), were foundto be consistent with those of the holotype of P. macrochir. Comparison of a range of material with the latter and the originaldescription of P. sheridani, led to the conclusion that P. macrochir was a senior synonym of P. sheridani (see Motomura
et al., 2000).

Polydactylus macrochir and 4 other Indo-West Pacific species included in this genus: P. bifurcus, P. microstomus,
P. plebeius and P. siamensis, are characterized by 5 pectoral filaments. Polydactylus macrochir can be easily
distinguished from the 4 species by the posterior margin of the maxilla extending considerably beyond the level of theposterior margin of the adipose eyelid (slightly short of, reaching to, or extending slightly beyond in the other 4 species).

Polydactylus macrochir further differs from P. microstomus in lacking a large black spot anteriorly on the lateral line(present in the latter). In overall body appearance, P. macrochir is similar to P. bifurcus, P. plebeius and P. siamensis.
However, P. macrochir differs from P. bifurcus in having higher counts of gillrakers [32 to 35 (mode 34) versus 30 in thelatter] and a simple lateral line (bifurcated in P. bifurcus). Polydactylus macrochir is distinguished from P. plebeius andP. siamensis by having higher counts of pored lateral-line scales [70 to 76 (mode 72) versus 60 to 68 (mode 63) in P. plebeius
and 54 to 58 (mode 54) in P. siamensis], higher counts of gillrakers [32 to 35 (mode 34) versus 24 to 32 (mode 26) in
P. plebeius and 22 to 24 (mode 23) in P. siamensis], occipital profile concave in adults (nearly straight throughout life inP. plebeius and P. siamensis), and the second spine of the first dorsal fin more robust than the other spines of the first dorsal
fin (all spines similar in P. plebeius and P. siamensis).

Polydactylus macrophthalmus (Bleeker, 1858)

Figs 95-96; Plate IIIb

Polynemus macrophthalmus Bleeker, 1858b: 10 [type locality: Palembang, Musi River, Sumatra, Indonesia; lectotype(RMNH 6015, 129 mm standard length) designated by Motomura et al., 2001c; paralectotype (RMNH 33967, 117 mmstandard length); date of publication of original description determined by Motomura et al., 2001c].
Synonyms: None.

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FAO Species Catalogue for Fishery Purposes No. 3

FAO Names: En - River threadfin; Fr - Barbure de rivière; Sp - Barbudo de rio.

Fig. 95 Polydactylus macrophthalmus (adult)

Fig. 96 Polydactylus macrophthalmus (juvenile)

Diagnostic Features: A medium- to large-sized species. Body depth at first dorsal-fin origin 22 to 28% (mean 25%) of
standard length; head length 29 to 33% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight inyoung but concave in adults. Posterior margin of maxilla extending well beyond level of posterior margin of adipose eyelid;
upper-jaw length 13 to 15% (mean 14%) of standard length; depth of posterior margin of maxilla approximately equal to orless than eye diameter; lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad
bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VIII spines,
second spine more robust than others; second dorsal fin with I spine and 13 or 14 (mode 14) soft rays; anal fin with III spinesand 10 or 11 (mode 11) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 13 or 14 (mode 14) rays (allrays unbranched), its length 24 to 26% (mean 24%) of standard length, posterior tip not reaching to level of posterior tip ofpelvic fin; pectoral filaments 7, upper 3 filaments extending beyond caudal-fin base; first to third pectoral filaments notreaching to level of posterior tip of pelvic fin; fourth pectoral filament extending slightly beyond anal-fin origin; fifth pectoral
filament longest, its length 161 to 192% (mean 176%) of standard length, extending beyond posterior tips of caudal-fin lobes;
caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 38 to 46% (mean 42%) andlower lobe 36 to 40% (mean 38%) of standard length. Pored lateral-line scales 87 to 94 (mode 88); lateral line simple,
extending from upper end of gill opening to upper end of lower caudal-fin lobe; scale rows above lateral line 10 to 12 (mode11), below 15 or 16 (mode 16). Gillrakers 10 to 12 (mode 11) on upper limb, 15 to 17 (mode 16) on lower limb, 25 to 29 (mode27) total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present, well developed. Colour:
Upper sides of head and trunk tinged silvery blue, becoming more silver on lower sides; posterior margins of first dorsal,
second dorsal and caudal fins greyish black, other parts hyaline; tips of pelvic and anal fins white, other parts transparent;
pectoral fin hyaline; pectoral filaments white.

Threadfins of the World

53

Geographical Distribution: Currently known only
from 3 rivers on 2 Indonesian islands: Kapuas River,
Kalimantan, and Musi and Batanghari Rivers, Sumatra(Fig. 97). These rivers were part of a single large river,the ancient Central or North Sunda River, on the
Sundaland during the Pleistocene, the last cold period
being about 12 000 years ago (see Morley and Flenleyin Whitmore, 1987: fig. 5.5; Motomura et al., 2001c:
fig. 3). Geographical evidence indicates that the
species was well adapted to the fresh-water basins inthis extensive Pleistocene River, whereas other
Polydactylus species are generally found in salt water
habitats. The Sundaland presently being submerged
to a depth of about 100 m, P. macrophthalmus is now
restricted to the 3 presently-known localities in theSouth China Sea.
Habitat and Biology: P. macrophthalmus has much
200 mlonger pectoral filaments and a more slender body thanother Polydactylus species, which generally inhabit
marine waters. Such morphological characters are
considered as an adaptation to a fresh-water
Fig. 97 Polydactylus macrophthalmus
existence, owing to their occurrence in Polynemus
Known distribution
species restricted to fresh-water and/or estuarine
habitats. The character of the long pectoral filaments is probably useful as a sense organ to search for food in muddy
waters. Furthermore, they had been collected from Jambi and Palembang, located about 75 km up the Batanghari and Musirivers, respectively (i.e. completely fresh-water basins). Accordingly, P. macrophthalmus is considered to be more heavily
dependent upon a fresh-water habitat than other congeners.
Size: Maximum standard length at least 52 cm (Feltes in Carpenter and Niem, 2001).
Interest to Fisheries: Esteemed as a food fish at least along the Kapuas River, Kalimantan, Indonesia. The species has
been often exported to Japan as an aquarium fish.
Local Names: None known.
Literature: Motomura et al. (2001c); Motomura (2002).
Remarks: P. macrophthalmus has long been treated as a member of Polynemus since its original description (e.g. Weberand de Beaufort, 1922; Myers, 1936; Roberts, 1989; Kottelat et al., 1993). However, the species differs from Polynemus
species in having the pectoral-fin insertion well below the midline on the lateral body (versus near midline), a basisphenoid
present (versus absent) and 24 vertebrae (versus 25), and clearly conform to the following diagnostic characters of
Polydactylus: tooth plate on vomer simple; width of tooth band on upper and lower jaws wider than space separating toothbands on opposing premaxilla; basisphenoid in contact with prootic; pectoral-fin base including base of pectoral filaments
less than upper-jaw length; swimbladder simple, not extending beyond anal-fin origin. Accordingly, the species was includedin the genus Polydactylus by Motomura et al. (2001c).

Polydactylus macrophthalmus can be easily distinguished from other congeners (except P. longipes) by the very longupper 3 pectoral filaments, extending beyond the caudal-fin base (not reaching caudal-fin base in the latter). Polydactylus
macrophthalmus is similar to P. longipes in having long pectoral filaments, but the former differs from the latter in having 7
pectoral filaments (6 in the latter). Further comparisons of P. macrophthalmus with P. longipes are given in the account of
the latter.
The Indo-Pacific Polydactylus with 7 pectoral filaments includes only 3 species: P. macrophthalmus, P. multiradiatus and
P. mullani. In addition to the difference in pectoral-filament length, P. macrophthalmus differs from P. multiradiatus inhaving lower anal-fin ray counts [10 or 11 (mode 11) versus 16 to 18 (mode 16) in the latter], higher pored lateral-line scalecounts [87 to 94 (mode 88) versus 49 to 56 (mode 52) in P. multiradiatus] and a concave occipital profile in adults (nearlystraight in P. multiradiatus). Furthermore, P. macrophthalmus is clearly distinguished from P. mullani by the higher countsof pored lateral-line scales, and scales above and below the lateral line [87 to 94 (mode 88) and 10 to 12 (mode 11) / 15 or 16(mode 16), respectively versus 46 to 50 (mode 48) and 5 to 7 (mode 6) / 9 or 10 (mode 10), respectively, in the latter], lower
gill-raker counts [25 or 29 (mode 27) versus 31 to 35 (mode 32) in P. mullani], a concave occipital profile in adults (nearlystraight in P. mullani), vomerine tooth present (absent), unbranched pectoral-fin rays (branched except uppermost 1 or 2
rays in P. mullani), and a large black spot absent anteriorly on the lateral line (present in P. mullani).

Polydactylus macrophthalmus is similar to P. macrochir in having a concave occipital profile in adults and the second
spine of the first dorsal fin more robust than other spines of the first dorsal fin. However, P. macrophthalmus differs from thelatter in having higher counts of pectoral filaments (7 versus 5 in the latter) and pored lateral-line scales [87 to 94 (mode 88)
versus 70 to 76 (mode 72) in P. macrochir], and lower gill-raker counts [25 to 29 (mode 27) versus 32 to 35 (mode 34) in
P. macrochir].

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FAO Species Catalogue for Fishery Purposes No. 3

Polydactylus malagasyensis Motomura and Iwatsuki, 2001

Fig. 98; Plate IIIc

Polydactylus malagasyensis Motomura and Iwatsuki, 2001b: 338, figs. 1, 7a, 8a [type locality: estuary of Mananjary River(about 100 m from sea), Mananjary, eastern Madagascar; holotype (AMNH 88029, 125 mm standard length); 35 paratypes(AMNH 231222, 3 specimens, 95 to 141 mm standard length; AMS I. 28114009, 2 specimens, 102 to 104 mm standard
length; ANSP 54807, 59 mm standard length; ANSP 77390, 127 mm standard length; ANSP 86372, 2 specimens, 94 to
136 mm standard length; CAS 66577, 3 specimens, 99 to 117 mm standard length; CAS 131390, 2 specimens, 144 to
148 mm standard length; MUFS 20381, 104 mm standard length; NRM 10479, 3 specimens, 135 to 149 mm standard
length; NRM 10480, 4 specimens, 74 to 87 mm standard length; USNM 171045, 126 mm standard length; USNM 278209,
113 mm standard length; USNM 301505, 125 mm standard length; USNM 307631, 129 mm standard length; USNM 358684,
3 specimens, 124 to 139 mm standard length; USNM 363484, 3 specimens, 63 to 126 mm standard length; SAM 34057, 3
specimens, 121 to 134 mm standard length)].
Synonyms: None.
FAO Names: En - African blackspot threadfin; Fr - Barbure tâche noire d'Afrique; Sp - Barbudo mancha negra africano.

Fig. 98 Polydactylus malagasyensis

Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 31 to 35% (mean 34%) of
standard length; head length 31 to 35% (mean 33%) of standard length. Snout pointed; occipital profile nearly straight.
Posterior margin of maxilla extending slightly beyond or reaching to level of posterior margin of adipose eyelid; upper-jaw
length 13 to 14% (mean 13%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw
well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on palatines and ectopterygoids;
vomerine tooth plate covered with skin and teeth absent; palatines inwardly turned anteriorly. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, second spine slightly more robust than or similar to others; second dorsal fin with I
spine and 12 or 13 (mode 13) soft rays; anal fin with III spines and 12 soft rays, anal-fin base approximately equal to or
greater than second dorsal-fin base; pectoral fin with 14 rays (all rays unbranched, except uppermost 1 or 2), its length 21 to
26% (mean 24%) of standard length, posterior tip not reaching to level of posterior tip of pelvic fin; pectoral filaments 6, first
filament shortest, reaching to level of pelvic-fin origin; second to fifth pectoral filaments extending slightly beyond level ofpelvic-fin origin; sixth pectoral filament longest, its length 27 to 31% (mean 29%) of standard length, just short of or extending
slightly beyond level of posterior tip of pectoral fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous,
upper caudal-fin lobe 35 to 40% (mean 39%) and lower lobe 34 to 38% (mean 36%) of standard length. Pored lateral-line
scales 46 to 51 (mode 47); lateral line simple, extending from upper end of gill opening to upper end of lower caudal-fin lobe;
scale rows above lateral line 5 or 6 (mode 6), below 9 or 10 (mode 10). Gillrakers 12 to 16 (mode 13) on upper limb, 17 to 19(mode 18) on lower limb, 29 to 34 (mode 31) total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3.
Swimbladder present, well developed. Colour: Golden olive above, but mainly silvery; a large black spot present anteriorly
on lateral line.

Threadfins of the World

55

Geographical Distribution: Currently known only
from Kenya, Mozambique, South Africa andMadagascar (Fig. 99). It is likely to be distributed over a
wider area, including United Republic of Tanzania andSomalia, east coast of Africa.
Habitat and Biology: Taken both in estuaries (less5.5 m) and offshore (16 to 62 m). No other data
available.
Size: Maximum standard length at least 15 cm(Motomura and Iwatsuki, 2001b).
Interest to Fisheries: None.
Local Names: MOZAMBIQUE: Barudo de mancha;
UNITED REPUBLIC OF TANZANIA: Kupe, Mkizi komomaji; SOUTH AFRICA: Sesvinger-draadvin, Sixfinger
threadfin.
Literature: Motomura and Iwatsuki (2001b); Motomura(2002).
Remarks: P. malagasyensis, previously identified as
P. sextarius (e.g. Smith in Smith and Heemstra, 1986;
Menon and Babu Rao in Fischer and Bianchi, 1984),
was recently described as a new species on the basis of
36 specimens (Motomura and Iwatsuki, 2001b).

200 m

Five Polydactylus species: P. malagasyensis ,Fig. 99 Polydactylus malagasyensis
P. microstomus , P. mullani , P. persicus andKnown distribution
P. sextarius, are characterized by having all pectoral-fin
rays branched, except the uppermost 1 or 2, the vomer
without teeth and a large black spot anteriorly on the lateral line. Characters of the pectoral fin and coloration of this group are
unique among the family Polynemidae. In addition to the group, only a single species, P. nigripinnis, has the vomer without
teeth in the genus.

Polydactylus malagasyensis is similar to other 4 species in overall body appearance. However, the species can be easily
distinguished from P. microstomus and P. mullani by the number of pectoral filaments [6 versus 5 (rarely asymmetrically 5
and 6) and 7 (rarely asymmetrically 6 and 7), respectively, in the latter].
The 3 species, with 6 pectoral filaments: P. malagasyensis, P. persicus and P. sextarius, have been considered asconspecific (e.g. Menon in Fischer and Whitehead, 1974; Menon and Babu Rao in Fischer and Bianchi, 1984). However,P. malagasyensis differs from P. sextarius in having a well-developed swimbladder (length about 40 to 45% of standard
length versus an atrophied, string-like swimbladder, about 20% of standard length in the latter; see Motomura and Iwatsuki,
2001b: fig. 7), higher counts of gillrakers [29 to 34 (mode 31) versus 25 to 30 (mode 28) in P. sextarius]. Furthermore, thesecond dorsal-fin spine length in P. malagasyensis [7 to 9% (mean 7%) of standard length] is significantly longer than that of
P. sextarius [5 to 8% (mean 6%) of standard length].

Polydactylus malagasyensis differs from P. persicus in having higher counts of pectoral-fin rays [14 versus 12 to 14 (mode12) in the latter] and scales below the lateral line [9 or 10 (mode 10) versus 8 or 9 (mode 9) in P. persicus], in addition to thepalatine being inwardly turned anteriorly (straight in P. persicus; see Motomura and Iwatsuki, 2001b: fig. 8). Whereas theposterior tip of the uppermost pectoral filament of P. malagasyensis extends slightly beyond or does not reach to theposterior tip of the pectoral fin, that of P. persicus extends well beyond the latter, owing to P. malagasyensis having both alonger pectoral fin [21 to 26% (mean 24%) of standard length versus 18 to 20% (mean 19%) of standard length in P. persicus]
and shorter pectoral filaments [27 to 31% (mean 29%) of standard length versus 26 to 36% (mean 32%) of standard length in
P. persicus]. Furthermore, the pectoral-fin base length, including the pectoral-filament base, of P. malagasyensis [10 to 12%(mean 12%) of standard length] is significantly greater than that of P. persicus [10 to 11% (mean 10%) of standard length].

Polydactylus microstomus (Bleeker, 1851)

Fig. 100; Plate IIId

Polynemus microstoma Bleeker, 1851b: 217 [type locality: Bulukumba, Sulawesi, Indonesia; holotype (RMNH 6044, 53 mm
standard length) determined from 9 Bleeker specimens by Motomura and Iwatsuki, 2001b].
Synonyms: Polydactylus zophomus Jordan and McGregor in Jordan and Seale, 1907: 11, fig. 4 [type locality: Cavite,
Luzon Island, Philippines; holotype (CAS 120113, 138 mm standard length) determined by Motomura and Iwatsuki, 2001b;paratype (USNM 55598, 2 specimens including a paratype and a non-type, 137 to 151 mm standard length)].

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FAO Species Catalogue for Fishery Purposes No. 3

FAO Names: En - Smallmouth threadfin; Fr - Barbure à petite bouche; Sp - Barbudo de boca pequeña.

Fig. 100 Polydactylus microstomus
Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 26 to 36% (mean 32%) of
standard length; head length 29 to 36% (mean 33%) of standard length. Snout pointed; occipital profile nearly straight.
Posterior margin of maxilla reaching to or slightly short of level of posterior margin of adipose eyelid; upper-jaw length 12 to
14% (mean 13%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw welldeveloped, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on palatines and ectopterygoids;
vomerine tooth plate covered with skin and teeth absent; palatines inwardly turned anteriorly. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, second spine slightly more robust than or similar to others; second dorsal fin with I
spine and 12 to 14 (mode 13) soft rays; anal fin with III spines and 11 or 12 (mode 12) soft rays, anal-fin base approximately
equal to or less than second dorsal-fin base; pectoral fin with 13 or 14 (mode 13, rarely 12 or 15) rays (all rays unbranched,
except uppermost 1 or 2), its length 17 to 20% (mean 18%) of standard length, posterior tip not reaching to level of posterior
tip of pelvic fin; pectoral filaments 5, first filament shortest, not reaching to or just reaching to level of pelvic-fin origin; second
to fourth pectoral filaments just short of or extending slightly beyond level of pelvic-fin origin; fifth pectoral filament longest,
its length 21 to 30% (mean 26%) of standard length, just short of or extending slightly beyond level of posterior tip of pectoral
fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 31 to 41% (mean 36%)
and lower lobe 31 to 39% (mean 35%) of standard length. Pored lateral-line scales 46 to 49 (mode 47); lateral line simple,
extending from upper end of gill opening to upper end of lower caudal-fin lobe; scale rows above lateral line 6 or 7 (mode 6),
below 9 or 10 (mode 10, rarely 8). Gillrakers 10 to 14 (mode 13) on upper limb, 13 to 18 (mode 16) on lower limb, 24 to 33(mode 29) total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present, well developed.
Colour: Head and upper sides of trunk tinged yellowish silver, becoming lighter silver on lower sides; snout
semi-translucent; posterior margins of first and second dorsal fins and caudal fin slightly blackish, remaining partstranslucent yellowish white; pectoral finmembrane yellowish; pectoral filaments
faintly white; anterior margins and origins
of pelvic and anal fins faintly white,remaining parts yellow; a large black spot
anteriorly on lateral line.
Geographical Distribution: Currently
known from the Indian Ocean, where it
ranges from Tamil Nadu, east of thesouthernmost tip of India, Sri Lanka,
Myanmar and Phuket Island, Thailand, tothe West Pacific where it ranges from
Taiwan Province of China to New
Caledonia, being relatively common in theeastern part of Indonesia and Philippines(Fig. 101). However, examples of
P. microstomus hav e at no t im e(apparently) been collected from
Australian waters.

200 m

Fig. 101 Polydactylus microstomus
Known distribution

Threadfins of the World

57

Habitat and Biology: Inhabits turbid coastal waters, estuaries and mangrove creeks, as well as mangrove-lined rivers.
Most of the species are taken from depths of less than 2 m. However, Schroeder (1980) reported the species in depths ofless than 20 m in the Philippines. Furthermore, a specimen (USNM 300898, 132 mm standard length) from Myanmar was
purportedly collected from a depth of 55 m.
Size: Maximum standard length at least 16 cm (Motomura and Iwatsuki, 2001b).
Interest to Fisheries: None.
Local Names: PHILIPPINES: Akin-akin, Kuwa-kuwa, Mamaling babai, Mamaling bato; PAPUA NEW GUINEA:
Small-mouthed threadfin.
Literature: Motomura and Iwatsuki (2001b); Motomura (2002).
Remarks: P. zophomus, originally described from Cavite, Luzon Island, Philippines, has been frequently regarded as a valid
species (Jordan and Richardson, 1908; Seale, 1910), although type specimens of the species have apparently never been
compared directly with those of P. microstomus. Examination of the types of both species showed that they represented asingle species (Motomura and Iwatsuki, 2001b). Therefore, P. zophomus is a junior synonym of P. microstomus.
Günther (1860) described 2 specimens with 5 pectoral filaments and a black blotch on the lateral line near its origin as
Polynemus plebejus, but the specimens are clearly Polydactylus microstomus. The species name, microstoma, given by
Bleeker (1851b), is changed to microstomus in accordance with the masculine gender of the genus name.

Polydactylus microstomus and 4 Polydactylus species: P. malagasyensis, P. mullani, P. persicus and P. sextarius, arecharacterized by having all pectoral-fin rays branched, except the uppermost 1 or 2, the vomer without teeth and a large
black spot anteriorly on the lateral line. The above 5 species are similar to each other in overall body appearance. However,P. microstomus can be easily distinguished from the other 4 species by the number of pectoral filaments [5 (rarely
asymmetrically 5 and 6) versus 7 (rarely asymmetrically 6 and 7) in P. mullani and 6 in the remaining 3 species].

Polydactylus mullani (Hora, 1926)

Fig. 102; Plate IIIe

Polynemus sextarius mullani Hora, 1926: 453 [type locality: Mumbai, India; holotype (ZSI-F 10747, 157 mm standard
length); 3 paratypes (ZSI-F 10748, 106 mm standard length; ZSI-F 10749, 97 mm standard length; ZSI-F 10750, 92 mm
standard length) reregistered by Motomura and Iwatsuki, 2001b].
Synonyms: None.
FAO Names: En - Arabian blackspot threadfin; Fr - Barbure à tâche noire d'Arabie; Sp - Barbudo de mancha negra árabe.

Fig. 102 Polydactylus mullani

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FAO Species Catalogue for Fishery Purposes No. 3

Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 28 to 35% (mean 33%) of
standard length; head length 33 to 37% (mean 35%) of standard length. Snout pointed; occipital profile nearly straight.
Posterior margin of maxilla just reaching to or not reaching to (in adults over about 60 mm standard length), or extending wellbeyond (in young) level of posterior margin of adipose eyelid; upper-jaw length 14 to 17% (mean 15%) of standard length;
depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted to dorsal
surface; teeth villiform in broad bands on palatines and ectopterygoids; vomerine tooth plate covered with skin and teeth
absent; palatines inwardly turned anteriorly. Posterior margin of preopercle serrated. First dorsal fin with VIII spines,
second spine more robust than others; second dorsal fin with I spine and 12 or 13 (mode 13) soft rays; anal fin with III spinesand 11 or 12 (mode 12) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 13 or 14 (mode 13) rays (allrays unbranched, except uppermost 1 or 2), its length 19 to 22% (mean 20%) of standard length, posterior tip not reaching tolevel of posterior tip of pelvic fin; pectoral filaments 7 (rarely asymmetrically 6 and 7); first (shortest) and second filaments,
not reaching to level of pelvic-fin origin; third to sixth pectoral filaments extending beyond level of pelvic-fin origin; seventh
pectoral filament longest, its length 28 to 45% (mean 33%) of standard length, extending well beyond level of posterior tip of
pectoral fin and not reaching to (in adults over about 60 mm standard length) or reaching to (in young) level of anal-fin origin;caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 32 to 52% (mean 39%) andlower lobe 32 to 50% (mean 38%) of standard length. Pored lateral-line scales 46 to 50 (mode 48); lateral line simple,
extending from upper end of gill opening to upper end of lower caudal-fin lobe; scale rows above lateral line 5 to 7 (mode 6),
below 9 or 10 (mode 10). Gillrakers 13 to 16 (mode 14) on upper limb, 18 to 21 (mode 18) on lower limb, 31 to 35 (mode 32)
total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present, well developed. Colour: Headand upper sides of trunk tinged darkly silver, becoming lighter silver on lower sides; snout semi-translucent; anterior andposterior margins of first and second dorsal, anal and caudal fins slightly blackish, other parts grey; pectoral-fin membranegrey with scattered melanophores; base of pectoral filaments white, becoming dark posteriorly; anterior margin and lower tipof pelvic fin grey, other parts white; a large black spot anteriorly on lateral line.
Geographical Distribution: Endemic and common in
the northern Arabian Sea (Fig. 103).
Habitat and Biology: Taken from depths of 14 to
115 m. Hida (1967) examined 401 specimens of thespecies and determined their sexes: thehermaphrodites (231 specimens, 57.6% of total) range
from 6.0 to 16.4 cm with a peak at 11.0 cm; the mature
females with large granulated ova (144 specimens,
35.9% of total) range from 10.0 to 18.4 cm with a peakat 12.5 cm; and immature females (25 specimens,
6.2% of total) and presumed male (1 specimen, 0.3% of
total) range from 9.0 to 15.9 cm without conspicuous
peaks. Polydactylus mullani feeds mainly on shrimps
and small fishes.
Size: Maximum standard length at least 19 cm(Motomura and Iwatsuki, 2001b).
Interest to Fisheries: One of the most importantcommercial species in the northern Arabian Sea.
Caught by bottom trawls. Hida (1967) reported that 3
700 examples were caught in 1 haul off southern
Kathiawar, India on 15 November 1963.
Local Names: None known.

200 m

Fig. 103 Polydactylus mullani
Known distribution

Literature: Motomura and Iwatsuki (2001b); Motomura (2002).
Remarks: Hora (1926) described Polynemus sextarius mullani as a new subspecies on the basis of 4 specimens (holotype
and 3 paratypes). The subspecies was raised to specific status (as Polydactylus mullani) by Motomura and Iwatsuki(2001b), because the species can be easily distinguished from other congeners, including P. sextarius.

Polydactylus mullani has been frequently misidentified as Filimanus heptadactyla (e.g. Kagwade, 1970, as Polynemus
heptadactylus; Talwar and Kacker, 1984, as Polydactylus heptadactylus). However, P. mullani differs from the latter inhaving wider teeth bands on the upper and lower jaws (compared with the space separating the teeth bands on opposing
premaxilla versus narrower teeth bands on the upper and lower jaws in F. heptadactyla), the basisphenoid in contact with theprootic (not in contact in F. heptadactyla), lower gill-raker counts [31 to 35 (mode 32) versus 35 to 41 (mode 39 in
F. heptadactyla)] and a large black spot present anteriorly on the lateral line (absent in F. heptadactyla). Furthermore,
whereas P. mullani is currently known only from the northern Arabian Sea in the Indian Ocean, F. heptadactyla is distributedin the western Pacific Ocean.

Polydactylus mullani and 4 Polydactylus species: P. malagasyensis, P. microstomus, P. persicus and P. sextarius, arecharacterized by having all pectoral-fin rays branched, except the uppermost 1 or 2, the vomer without teeth and a large
black spot anteriorly on the lateral line. Polydactylus mullani can be easily distinguished from the other 4 species by the
number of pectoral filaments [7 (rarely asymmetrically 6 and 7) versus 5 (rarely asymmetrically 5 and 6) in P. microstomus
and 6 in the remaining 3 species].

Threadfins of the World59

The body appearance of P. mullani varies remarkably with overall fish growth, compared with that of the other 4 species witha large black anterior lateral-line spot. The uppermost pectoral filament and posterior margin of the maxilla in young
specimens (less than about 60 mm standard length) of P. mullani reach (versus not reaching in adults) the anal-fin origin
and extend well beyond (versus just reaching or not reaching) posterior margin of the adipose eyelid, respectively.
Furthermore, the upper and lower caudal-fin lobes of young specimens (less than about 60 mm standard length) of
P. mullani are extremely long [50 to 52% (mean 51%) of standard length and 43 to 50% (mean 47%) of standard length,
respectively versus 32 to 42% (mean 38%) of standard length and 32 to 39% (mean 36%) of standard length, respectively, in
adults]. These growth-related features are not found in the other 4 species.

Polydactylus multiradiatus (Günther, 1860)

Fig. 104; Plate IIIf

Polynemus multiradiatus Günther, 1860: 324 [type locality: China (but probably erroneous, see Motomura, Johnson and
Iwatsuki, 2002); holotype (BMNH 1852.5.4.5, stuffed specimen, 147 mm standard length)].
Synonyms: Polynemus specularis De Vis, 1883: 285 (type locality: Brisbane River, Queensland, Australia; 2 or more
syntypes apparently lost, see Motomura, Johnson and Iwatsuki, 2002). Polydactylus auratus McKay, 1970: 8 [type locality:
Napier Broome Bay, Western Australia, Australia; holotype (WAM-P 16792-001, 115 mm standard length); 3 paratypes(WAM-P 16793-001, 106 mm standard length; WAM-P 16794-001, 129 mm standard length; WAM-P 16795-001, 99 mmstandard length)].
FAO Names: En - Australian threadfin; Fr - Barbure à dos sombre; Sp - Barbudo de lomo oscuro.

Fig. 104 Polydactylus multiradiatus
Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 31 to 38% (mean 35%) of
standard length; head length 28 to 33% (mean 30%) of standard length. Snout pointed; occipital profile nearly straight.
Posterior margin of maxilla not reaching to, just reaching to or extending slightly beyond level of posterior margin of adipose
eyelid; upper-jaw length 12 to 14% (mean 13%) of standard length; depth of posterior margin of maxilla less than eye
diameter; lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on vomer,
palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases ofsimilar thickness; second dorsal fin with I spine and 13 to 15 (mode 14) soft rays; anal fin with III spines and 16 to 18 (mode16) soft rays, anal-fin base greater than second dorsal-fin base; pectoral fin with 14 to 17 (mode 15) rays (all rays
unbranched), its length 25 to 30% (mean 28%) of standard length, posterior tip not reaching to level of posterior tip of pelvic
fin; pectoral filaments 7 (rarely 6 on each side, asymmetrically 6 and 7, or 7 and 8); first (shortest) and second pectoral
filaments not reaching to level of pelvic-fin origin; third pectoral filament reaching to or just short of level of pelvic-fin origin;fourth pectoral filament extending slightly beyond or reaching level of pelvic-fin origin; fifth and sixth (rarely longest) pectoral
filaments extending well beyond level of pelvic-fin origin; seventh pectoral filament longest, its length 26 to 32% (mean 30%)of standard length, not extending to posterior tip of pectoral fin; caudal fin deeply forked, upper and lower caudal-fin lobes notfilamentous, upper caudal-fin lobe 36 to 42% (mean 39%) and lower lobe 34 to 41% (mean 37%) of standard length. Poredlateral-line scales 49 to 56 (mode 52); lateral line simple, extending from upper end of gill opening to upper end of lower
caudal-fin lobe; scale rows above lateral line 7 or 8 (mode 8), below 14 or 15 (mode 14). Gillrakers 11 to 14 (mode 12) on
upper limb, 16 to 20 (mode 17) on lower limb, 27 to 33 (mode 30) total. Vertebrae 10 precaudal and 14 caudal; supraneuralbones 3. Swimbladder absent. Colour: Upper sides of head and trunk with slightly darkish silver to bronze tinge, becoming

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FAO Species Catalogue for Fishery Purposes No. 3

lighter silvery white on lower sides; snout semi-translucent with dusky tip; intensity of pigmentation of pectoral and upper partof first dorsal fin variable, but usually at least partially covered with dusky melanophores; posterior tip of second dorsal fin
blackish, other parts pale fleshy yellow; pectoral filaments white; anterior margin of anal fin white, other parts dusky whitish
to yellow; caudal fin semi-translucent greyish.
Geographical Distribution: Currently known only fromsouthern Indonesia (Timor and Arafura Seas) andnorthern Australia (Northern Territory, Queensland, andnorthern Western Australia and New South Wales)(Fig. 105). Judging from presently known locality data
for the species, the type locality (China), given by
Günther (1860) is most likely erroneous.
Habitat and Biology: P. multiradiatus has a similar
distribution to P. macrochir, occurring in southern
Papua New Guinea and northern Australia. However,collection data for P. macrochir indicated thatspecimens had been taken from depths of 0.2 to 6 m on
muddy or sandy bottoms, whereas most specimens of
P. multiradiatus (except a single young specimen,
AMS IB. 3089, 103 mm standard length, collected from
depth of 4 m), were taken from depths of 10 to 56 m,
also on muddy or sandy bottoms. This suggests that
P. macrochir and P. multiradiatus are segregatedvertically, although fully overlapping geographically.
On the basis of gonad examinations, it is clear that
P. multiradiatus is a protandrous hermaphrodite,changing sex from male to female between 120 and
140 mm standard length (Motomura, Johnson and
Iwatsuki, 2002). The body depth of individuals also
increased significantly during that period (Motomura,Johnson and Iwatsuki, 2002: fig. 2).

200 m

Fig. 105 Polydactylus multiradiatus
Known distribution

Sumpton and Greenwood (1990: table 1) listed the food items in the stomachs of juveniles (20 to 59 mm fork length) of
P. multiradiatus from the Logan-Albert estuarine system, Moreton Bay, Queensland, Australia. Smaller individuals of thespecies (20 to 29 mm fork length) fed predominantly on calanoid copepods, which formed 83% by volume of their diet. With
progressive increases in the size of the species, copepods became a less important dietary item, comprising only 28% of theaverage stomach volume at a size of 50 to 59 mm fork length.
Size: Maximum standard length at least 15 cm (Motomura, Johnson and Iwatsuki, 2002).
Interest to Fisheries: None.
Local Names: AUSTRALIA: Günther’s threadfin; PAPUA NEW GUINEA: Many-rayed threadfin.
Literature: Motomura, Johnson and Iwatsuki (2002b); Motomura (2002).
Remarks: Polynemus specularis was briefly described by De Vis (1883) on the basis of 2 or more specimens from theBrisbane River, Queensland, Australia. De Vis (1883) stated that the anal-fin soft rays of the species numbered 17 or 18,
which is consistent with Polydactylus multiradiatus (16 to 18), and different from the usual condition (10 to 13) among otherIndo-Pacific Polydactylus species. Polynemus specularis is therfore regarded as a junior synonym of Polydactylus
multiradiatus (Motomura, Johnson and Iwatsuki, 2002). Polydactylus auratus, described by McKay (1970) on the basis of
4 specimens from Napier Broome Bay, Western Australia, Australia, are also found to be conspecific with P. multiradiatus.
Their meristic and morphological characters are included in Motomura, Johnson and Iwatsuki (2002: table 1).
Grant (1982, 1995) provided figures (1985, pl. 273 and 1995, fig. 631a) of a threadfin identified as Polydactylus
heptadactylus. This species, now recognized as Filimanus heptadactyla, is characterized by 11 or 12 anal-fin soft rays,
whereas the threadfin figured by Grant (1982, 1995) clearly had 16 or more anal-fin soft rays. Therefore, Grant’s
P. heptadactylus is clearly P. multiradiatus.

Polydactylus multiradiatus can be easily distinguished from other Indo-Pacific congeners by the higher anal-fin soft ray
counts (16 to 18 versus 10 to 13 in all others). Three Indo-Pacific Polydactylus species, P. macrophthalmus,
P. multiradiatus and P. mullani, have 7 pectoral filaments. In addition to the difference in numbers of anal-fin soft rays,
P. multiradiatus differs from P. macrophthalmus in having lower pored lateral-line scale counts [49 to 56 (mode 52) versus
87 to 94 (mode 88) in the latter], shorter pectoral filaments (no filaments extending beyond posterior tip of pectoral fin versus
upper 3 filaments extending beyond caudal-fin base in P. macrophthalmus) and occipital profile nearly straight throughout
life (occipital profile concave in adults in P. macrophthalmus). Polydactylus multiradiatus is clearly distinguished fromP. mullani by the presence of vomerine teeth (absent in the latter) and absence of a large black spot anteriorly on the lateralline (present in P. mullani).

Threadfins of the World

61

Polydactylus nigripinnis Munro, 1964

Fig. 106; Plate IIIg

Polydactylus nigripinnis Munro, 1964: 156, fig. 4 [type locality: 4 km off east head of Purari River, Gulf of Papua, Papua
New Guinea; holotype (CSIRO A. 1828, 103 mm standard length); 3 paratypes (CSIRO A. 1827, 96 mm standard length;
CSIRO A. 1829, 108 mm standard length; CSIRO A. 1830, 100 mm standard length)].
Synonyms: None.
FAO Names: En - Blackfin threadfin; Fr - Barbure à nageoires noires; Sp - Barbudo de aletas negras.

Fig. 106 Polydactylus nigripinnis
Diagnostic Features: A small-sized species. Body depth at first dorsal-fin origin 30 to 36% (mean 33%) of standard length;
head length 32 to 36% (mean 33%) of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of
maxilla extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 16% (mean 15%) of
standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth
restricted to dorsal surface; teeth villiform in broad bands on palatines and ectopterygoids; vomerine tooth plate covered withskin and teeth absent. Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar
thickness; second dorsal fin with I spine and 12 or 13 (mode 13) soft rays; anal fin with III spines and 11 to 13 (mode 12) softrays, anal-fin base slightly greater than second dorsal-fin base; pectoral fin with 16 to 18 (mode 17) rays (all rays
unbranched), its length 29 to 34% (mean 31%) of standard length, posterior tip extending well beyond level of posterior tip ofpelvic fin; pectoral filaments 6; first (shortest) and second pectoral filaments not reaching to level of pelvic-fin origin; third to
fifth pectoral filaments extending beyond level ofpelvic-fin origin; sixth pectoral filament longest, its
length 26 to 35% (mean 32%) of standard length, justshort of level of posterior tip of pectoral fin; caudal fin
deeply forked, upper and lower caudal-fin lobes notfilamentous, upper caudal-fin lobe 36 to 45% (mean42%) and lower lobe 36 to 40% (mean 39%) of
standard length. Pored lateral-line scales 46 to 50(mode 47); lateral line simple, extending from upper
end of gill opening to upper end of lower caudal-fin
lobe; scale rows above lateral line 7, below 11 to 13(mode 11). Gillrakers 10 to 13 (mode 11) on upper
limb, 14 to 16 (mode 15) on lower limb, 24 to 29 (mode27) total. Vertebrae 10 precaudal and 14 caudal;
supraneural bones 3. Swimbladder absent. Colour:
Upper sides of head and trunk with slightly darkish
silver tinge, becoming lighter on lower sides; snout
semi-translucent; first and second dorsal fins yellowish
white; anterior margin of pelvic fin yellowish, otherparts white; anal fin white; pectoral fin dense black;pectoral filaments white; caudal fin darkish silver.
200 m
Geographical Distribution: Currently known only from
Papua New Guinea (Gulf of Papua), Australia(Cambridge Gulf, Western Australia to Van Diemen Gulf,
Northern Territory) and the Arafura Sea (Fig. 107).

Fig. 107 Polydactylus nigripinnis
Known distribution

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FAO Species Catalogue for Fishery Purposes No. 3

Habitat and Biology: Taken off river mouths in Australia and Papua New Guinea, and is also caught offshore in the Arafura
Sea. No other data are available.
Size: Maximum standard length at least 15 cm (Feltes in Carpenter and Niem, 2001).
Interest to Fisheries: None.
Local Names: AUSTRALIA: Black-finned threadfin; PAPUA NEW GUINEA: Black-finned threadfin.
Literature: Feltes in Carpenter and Niem (2001); Motomura (2002).
Remarks: P. nigripinnis and 5 other species: P. malagasyensis, P. microstomus, P. mullani, P. persicus and P. sextarius,
are all characterized by having the vomer covered with skin and without teeth. Polydactylus nigripinnis is easily
distinguished from the others by lacking a large black spot anteriorly on the lateral line (present in the other 5 species) andhaving unbranched pectoral-fin rays (branched, except uppermost 1 or 2 in the other 5 species).
In the Indo-Pacific, only 3 species, P. longipes, P. nigripinnis and P. sexfilis, have 6 pectoral filaments and no black spot
anteriorly on the lateral line. Polydactylus nigripinnis differs from P. sexfilis in having a vomer without teeth (present in thelatter), lower counts of pored lateral-line scales [46 to 50 (mode 47) versus 61 to 67 (mode 64) in P. sexfilis], swimbladderabsent (present in P. sexfilis) and smaller orbit diameter [5 to 7% (mean 6%) of standard length versus 8 to 11% (mean 9%)of standard length in P. sexfilis]. Comparisons of P. nigripinnis with P. longipes are given in the account of the latter.

Polydactylus octonemus (Girard, 1858)

Fig. 108; Plate IIIh

Polynemus octonemus Girard, 1858: 167 [type locality: Brazos Santiago and Galveston, Texas, USA; 3 syntypes presently
known (ANSP 22900, 62 mm standard length; USNM 739, 2 specimens, 62 to 79 mm standard length)].
Synonym: Trichidion octofilis Gill, 1861: 280 [type locality: New York, USA; holotype (USNM 15074, 166 mm standard
length)].
FAO Names: En - Atlantic threadfin; Fr - Barbure huit doigts atlantique; Sp - Barbudo ocho dedos atlántico.

Fig. 108 Polydactylus octonemus
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 28 to 32% (mean 29%) of standard
length; head length 27 to 33% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla just reaching to or extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw length
12 to 15% (mean 14%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw welldeveloped, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids.
Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal
fin with I spine and 11 to 13 (mode 12) soft rays; anal fin with III spines and 12 to 14 (mode 13) soft rays, anal-fin base greaterthan second dorsal-fin base; pectoral fin with 14 to 16 (mode 15) rays (all rays unbranched), its length 24 to 29% (mean 27%)of standard length, posterior tip not reaching to level of posterior tip of pelvic fin (or just reaching in young specimens lessthan about 90 mm standard length); pectoral filaments 8 (rarely 9); first pectoral filament shortest, extending beyond level ofThreadfins of the World63

pelvic-fin origin; second to fifth pectoral filaments extending beyond level of pelvic-fin origin in adults (in young specimens
less than about 90 mm standard length, third pectoral filament extending beyond level of posterior tip of pelvic fin, fourth
pectoral filament reaching to level of anal-fin origin; fifth pectoral filament extending beyond level of anal-fin origin); sixth
pectoral filament reaching to or extending slightly beyond level of posterior tip of pelvic fin in adults (extending beyond levelof anal-fin origin in young); seventh pectoral filament not reaching to or extending beyond level of anal-fin origin; eighth
pectoral filament longest, its length 39 to 51% (mean 43%) of standard length, not reaching to or extending beyond level ofanal-fin origin in adults (reaching to level of midpoint of anal-fin base in young); caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 39 to 42% (mean 40%) and lower lobe 33 to 40% (mean 37%) of
standard length. Pored lateral-line scales 56 to 64 (mode 59); lateral line bifurcated on caudal fin, extending to posterior
margins of upper and lower caudal-fin lobes; scale rows above lateral line 6 or 7 (mode 6), below 10 to 12 (mode 11).
Gillrakers 34 to 38 (mode 36). Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder present, welldeveloped. Colour: Upper sides of head and trunk with slightly darkish silver tinge, becoming lighter on lower sides; snout
semi-translucent; first and second dorsal fins dusky yellow; pectoral fin black; pectoral filaments translucent; posterior
margin of anal fin white, other parts dusky yellow; posterior margin of caudal fin blackish, other part dusky yellow.
Geographical Distribution: In the western Atlantic
Ocean ranging from New York, USA to the west coast of
Yucatán (Fig. 109).
Habitat and Biology: Generally occurs on sandy or
muddy bottoms and on beaches, frequently in the surf.
The species is taken at depths from 5 to 66 m in the
Gulf of Mexico (Springer and Bullis, 1956; Dentzau andChittenden, 1990), but is most abundant at depths of 5
to 16 m (Dentzau and Chittenden, 1990). In the Gulf of
Mexico, young specimens (less than about 66 mm)
have been taken at the surface over water depths from
1 035 to 2 736 m.
Dentzau and Chittenden (1990) studied the biology of
P. octonemus in detail. It generally matures between
165 and 210 mm total length (approaching 7 to 9months of age) and probably spawns in the Gulf of
Mexico, off Louisiana and Texas. Spawning primarily
occurs from mid-December to midMarch, spanning 45
to 120 days overall. Larvae (2- to 4-month-olds) arerecruited in waters less than 16 m, and then begin todisperse to deeper waters in early summer. The
species ranges from 135 to 165 mm total length at 6
months, from 165 to 215 mm at 9 months and from 180
to 205 mm at 1 year. The typical maximum life span isabout 1 year but may exceed that if individuals survivea pelagic stage after spawning.

200 m

Fig. 109 Polydactylus octonemus
Known distribution

Size: Maximum total length 23 cm, commonly to 20 cm (Dentzau and Chittenden, 1990).
Interest to Fisheries: Caught incidentally by seines and trawls and is of little commercial importance.
Local Names: None known.
Literature: Feltes in Carpenter (2003).
Remarks: Examination by the author of the holotype of Trichidion octofilis showed it to be conspecific with Polydactylus
octonemus, the former thereby being a junior synonym of the latter.

Polydactylus octonemus, P. oligodon and P. virginicus, all occurring in the western Atlantic, are very similar to each other inoverall body appearance. Polydactylus octonemus differs from the other 2 species in having higher counts of pectoral
filaments [8 (rarely 9) versus 7 (rarely 8) in the other 2 species] and gillrakers [34 to 38 versus 22 to 30 in the other 2 species].

Polydactylus oligodon (Günther, 1860)

Fig. 110; Plate IVa

Polynemus oligodon Günther, 1860: 322 [original locality: Rio de Janeiro, Brazil and Jamaica, based on 2 syntypes; type
locality: Rio de Janeiro, Brazil, based on a lectotype (BMNH 1853.4.18.23, 147 mm standard length) designated by Randall,
1966; paralectotype (BMNH 1848.1.12.890, stuffed specimen, 146 mm standard length)].
Synonyms: None.

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FAO Names: En - Littlescale threadfin; Fr - Barbure à petites écailles; Sp - Barbudo de escamas pequeñas.

Fig. 110 Polydactylus oligodon
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 27 to 30% (mean 28%) of standard
length; head length 30 to 33% (mean 32%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla just short of or reaching to level of posterior margin of adipose eyelid; upper-jaw length 13 to 15% (mean14%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentaryteeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of
preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and
11 or 12 (mode 12) soft rays; anal fin with III spines and 13 to 15 (mode 14) soft rays, anal-fin base greater than second
dorsal-fin base; pectoral fin with 15 or 16 (mode 16) rays (all rays unbranched), its length 22 to 24% (mean 23%) of standard
length, posterior tip not reaching to level of posterior tip of pelvic fin; pectoral filaments 7 (rarely 8); first pectoral filament
shortest, extending slightly beyond level of pelvic-fin origin; second and third pectoral filaments extending beyond level ofpelvic-fin origin; fourth pectoral filament reaching to level of posterior tip of pelvic fin; fifth pectoral filament reaching to or
extending beyond level of posterior tip of pelvic fin; sixth pectoral filament reaching to or extending slightly beyond level ofanal-fin origin; seventh pectoral filament longest, its length 31 to 41% (mean 38%) of standard length, extending beyond
level of anal-fin origin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 40 to
43% (mean 42%) and lower lobe 37 to 41% (mean 39%) of standard length. Pored lateral-line scales 67 to 73 (mode 70);
lateral line bifurcated on caudal fin, extending to posterior margins of upper and lower caudal-fin lobes; scale rows above
lateral line 7 to 9 (mode 9), below 11 to 14 (mode 13).
Gillrakers 22 to 30 (mode 27). Vertebrae 10 precaudal and14 caudal; supraneural bones 3. Swimbladder present.
Colour: Upper sides of head and trunk with slightly darkish
silver tinge, becoming lighter on lower sides; snout
semi-translucent; first and second dorsal fins translucent;pectoral fin white with a small number of melanophores;
pectoral filaments and pelvic fin white; base and posterior
margin of anal fin white, other parts translucent; posterior
margin of caudal fin blackish, other part whitish.
Geographical Distribution: In the western Atlantic Ocean
ranging from the east coast of Florida (at least to Fort
Lauderdale), USA to Santos, Brazil, including the Antilles,
Venezuela, Guyana, Suriname and French Guiana(Fig. 111). However, the species does not occur in the Gulf of
Mexico and western Caribbean Sea.
Habitat and Biology: Occurs close to the shore in surfalong exposed sandy beaches. No other data are available.
Size: Maximum total length 46 cm (Feltes in Carpenter,2003).
Interest to Fisheries: Caught incidentally by seines andtrawls and is of little commercial importance.
Local Names: BAHAMAS: Smallscale threadfin;
DOMINICA: Barbú.

200 m

Fig. 111 Polydactylus oligodon
Known distribution

Threadfins of the World

65

Literature: Feltes in Carpenter (2003).
Remarks: Although Polydactylus oligodon, originally described from Rio de Janeiro, Brazil and Jamaica on the basis of 2
specimens, had long been regarded as a junior synonym of P. virginicus by many authors, Randall (1966) recognized theformer as valid and designated a lectotype for the species. Randall (1966) noted differences between P. oligodon and
P. virginicus in the shape of the posterior margin of the maxilla, and certain meristic characters (including numbers oflateral-line scales, pectoral-fin rays and anal-fin rays) and proportional measurements (including length of anal-fin base). In
particular, the rounded shape of the posterior margin of the maxilla has been subsequently treated as a diagnostic character
for P. oligodon (versus truncate to concave in P. virginicus) in many publications (e.g. Randall in Fischer, 1978; Cervigón in
Cervigón et al., 1993; Randall, 1996). According to Feltes in Carpenter (2003) and confirmed by examination by the author,
however, that character shows considerable individual variation and it is difficult to clearly distinguish between P. oligodon
and P. virginicus on that basis.
Although P. oligodon and P. virginicus are very similar to each other and distinction between the 2 species in recent
literature is somewhat confused, P. oligodon can be distinguished from the latter by the higher counts of pored lateral-line
scales [67 to 73 (mode 70) versus 54 to 63 (mode 58) in the latter].

Polydactylus opercularis (Gill, 1863)

Fig. 112; Plate IVb

Trichidion opercularis Gill, 1863: 168 (type locality: west coast of Central America, probably Cape San Lucas, Baja
California, Mexico; holotype apparently lost, see Motomura, Kimura and Iwatsuki, 2002).
Synonyms: Polynemus melanopoma Günther, 1864: 148 [type locality: San José, Guatemala; holotype (BMNH
1864.1.26.321, 263 mm standard length)].
FAO Names: En - Yellow bobo; Fr - Barbure jaune; Sp - Barbudo amarillo.

Fig. 112 Polydactylus opercularis
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 25 to 29% (mean 27%) of standard
length; head length 29 to 33% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight throughout life.
Posterior margin of maxilla extending well beyond level of posterior margin of adipose eyelid; maxilla covered with small
deciduous scales; upper-jaw length 16 to 18% (mean 17%) of standard length; depth of posterior margin of maxilla less thaneye diameter; anterior parts of lower jaw with villiform teeth extending onto lateral surface, adjacent portion of lip poorly
developed in larger specimens; teeth villiform in broad bands on vomer, palatines and ectopterygoids; tooth plate on
palatines wider than those on ectopterygoids; shape of vomerine tooth plate nearly square. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, thickness of all first dorsal-fin spine bases similar; second dorsal fin with I spine
and 11 to 13 (mode 12) soft rays; anal fin with III spines and 12 to 14 (mode 13) soft rays, anal-fin base slightly greater thansecond dorsal-fin base; pectoral fin with 14 to 16 (mode 15) rays (all rays unbranched), its length 20 to 24% (mean 22%) of
standard length, posterior tip not reaching to level of posterior tip of pelvic fin; pectoral filaments 9 (rarely 8 or asymmetrically
8 and 9), first filament shortest, not reaching to or extending slightly beyond level of posterior tip of pelvic fin; lengths of

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FAO Species Catalogue for Fishery Purposes No. 3

second to ninth pectoral filaments similar, its length 30 to 41% (mean 36%) of standard length, extending beyond level ofposterior tip of pelvic fin or level of anal-fin origin, sometimes not reaching to level of posterior tip of pelvic fin; entire
well-developed membranes present on dorsal margins of pectoral filaments; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 31 to 38% (mean 35%) and lower lobe 30 to 36% (mean 33%) of
standard length. Pored lateral-line scales 66 to 74 (mode 70); lateral line simple, extending from upper end of gill opening toupper end of lower caudal-fin lobe; scale rows above lateral line 7 to 10 (mode 8), below 12 to 15 (mode 13). Gillrakers 13 to
18 (mode 16) on upper limb, 17 to 20 (mode 19) on lower limb, 31 to 37 (mode 34) total. Vertebrae 10 precaudal and 14
caudal; supraneural bones 2. Swimbladder absent. Colour: Head and body pale green dorsally, silver ventrally; posterior
margins of first and second dorsal fins dark yellow, remaining parts dusky; pectoral fin dusky; base of pectoral filaments
white, becoming yellow on posterior tips; posterior tip of pelvic fin yellow, remaining parts white; base of anal fin white,remaining parts vivid yellow; lower caudal-fin lobe dark yellow, upper lobe dusky.
Geographical Distribution: Currently known from theeastern Pacific Ocean (Fig. 113), where it ranges from
northern Los Angeles Harbor, California, USA to off
Paita, Peru, being relatively rare north of Baja
California, Mexico. The record from off Paita, Peru wasbased on a single juvenile specimen (12 mm total
length; Hildebrand, 1946).
Habitat and Biology: Generally inhabits muddy or
sandy bottoms in coastal waters and estuaries, but also
occurs along sandy beaches (Allen and Robertson,1994). Most of the collection data available indicates it
occurs in depths less than 46 m.
Size: Maximum total length at least 45 cm (Allen andRobertson, 1994).

200 m

Interest to Fisheries: Esteemed as a food fish
throughout the tropical eastern Pacific.

Fig. 113 Polydactylus opercularis
Known distribution

Local Names: COLOMBIA: Barbeta; MEXICO:
Barbudo amarillo.
Literature: Motomura, Kimura and Iwatsuki (2002).
Remarks: P. melanopoma, originally described by Günther (1864) from San José, Guatemala on the basis of a single
specimen (BMNH 1864.1.26.321, 263 mm standard length), was synonymized under P. opercularis, originally described by
Gill (1863) (as Trichidion opercularis) on the basis of a single specimen (apparently lost) (Motomura, Kimura and Iwatsuki,2002).

Polydactylus opercularis is easily distinguished from other congeners by having usually 9 pectoral filaments (usually 4 to 8
filaments in the latter). The number of pectoral filaments in an Atlantic west coast species, P. octonemus, although usually
8, infrequently number 9, thereby overlapping with P. opercularis which has 8 pectoral filaments on rare occasions.
However, P. octonemus differs from the latter in having lower pored lateral-line scale counts [56 to 64 (mode 59) versus 66 to
74 (mode 70) in P. opercularis], well-developed lower-jaw lips throughout life (lips on anterior parts of lower jaws poorly
developed in larger specimens of P. opercularis) and a swimbladder present (absent in P. opercularis). The condition of thelower jaw lips and teeth (anterior parts of lower jaw with villiform teeth extending onto lateral surface, adjacent portion of lip
poorly developed) found in larger P. opercularis is unique among the genus, although the condition is also found in all stagesof Leptomelanosoma indicum and larger Parapolynemus verekeri.
Polydactylus opercularis lacks a swimbladder, and 3 Indo-West Pacific Polydactylus species, P. multiradiatus,
P. nigripinnis and P. siamensis also lack a swimbladder. Polydactylus opercularis can be easily distinguished from theothers by having 9 pectoral filaments (5 in P. siamensis, 6 in P. nigripinnis and 7 in P. multiradiatus). Comparisons of
P. opercularis with a eastern Pacific species, P. approximans are given in the account of the latter.

Polydactylus persicus Motomura and Iwatsuki, 2001

Fig. 114; Plate IVc

Polydactylus persicus Motomura and Iwatsuki, 2001b: 347, figs. 5, 7D [type locality: Kuwait Bay, Kuwait, Persian Gulf
(29°30’N, 47°50’E); holotype (MCZ 60001, 121 mm standard length); 12 paratypes (AMS I. 40432-001, 103 mm standard
length; BMNH 2000.9.25.1, 93 mm standard length; KU 10528, 112 mm standard length; MCZ 59251 (5 specimens,
including 1, 63 mm standard length, cleared and stained), 48-80 mm standard length; MCZ 158350 (2), 92-121 mm standard
length; MUFS 20410, 124 mm standard length; USNM 363075, 96 mm standard length)].
Synonyms: None.

Threadfins of the World

67

FAO Names: En - Persian blackspot threadfin; Fr - Barbure à tâche noire de Perse; Sp - Barbudo de mancha negra persa.

Fig. 114 Polydactylus persicus
Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 33 to 35% (mean 34%) of
standard length; head length 31 to 34% (mean 33%) of standard length. Snout pointed; occipital profile nearly straight.
Posterior margin of maxilla not reaching to level of posterior margin of adipose eyelid; upper-jaw length 13 to 15% (mean14%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentaryteeth restricted to dorsal surface; teeth villiform in broad bands on palatines and ectopterygoids; vomerine tooth plate
covered with skin and teeth absent; palatines straight anteriorly. Posterior margin of preopercle serrated. First dorsal fin with
VIII spines, second spine slightly more robust than others; second dorsal fin with I spine and 12 or 13 (mode 13) soft rays;
anal fin with III spines and 12 soft rays, anal-fin base approximately equal to second dorsal-fin base; pectoral fin with 12 to 14(mode 12) rays (all rays unbranched, except uppermost 1 or 2), its length 18 to 20% (mean 19%) of standard length,
posterior tip not reaching to level of posterior tip of pelvic fin; pectoral filaments 6, first filament shortest; sixth pectoral
filament longest, its length 26 to 36% (mean 32%) of standard length, extending well beyond level of posterior tip of pectoral
fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 35 to 40% (mean 38%)
and lower lobe 34 to 44% (mean 37%) of standard length. Pored lateral-line scales 46 to 49 (mode 48); lateral line simple,
extending from upper end of gill opening to upper end oflower caudal-fin lobe; scale rows above lateral line 5 or 6
(mode 6), below 8 or 9 (mode 9). Gillrakers 12 to 16(mode 13) on upper limb, 17 to 20 (mode 19) on lower
limb, 29 to 35 (mode 32) total. Vertebrae 10 precaudal
and 14 caudal; supraneural bones 3. Swimbladderpresent, well developed. Colour: Head and upper sidesof trunk with slightly darkish silver tinge, becoming lighter
on lower sides; snout semi-translucent; posterior marginof first and second dorsal fins and caudal fin indistinctly
blackish, other parts translucent; pectoral-fin membrane
white with scattered melanophores; pectoral filaments
whitish, becoming blackish on posterior tips; pelvic fin
whitish; a large black spot anteriorly on lateral line.
Geographical Distribution: Currently known only fromthe Persian Gulf (Fig. 115).
Habitat and Biology: Occurs on sandy and muddy
bottoms in shallow waters (less than 10 m). No other data
are available.
Size: Maximum standard length at least 16 cm.
Interest to Fisheries: An important fishery species in
Kuwait, caught mainly by trawl and gill net.
Local Names: ISLAMIC REPUBLIC OF IRAN: Taer;KUWAIT: Ghazal.

200 m

Fig. 115 Polydactylus persicus
Known distribution

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FAO Species Catalogue for Fishery Purposes No. 3

Literature: Motomura and Iwatsuki (2001b); Motomura (2002).
Remarks: P. persicus, previously identified as P. sextarius (e.g. Kuronuma and Abe, 1972; Randall, 1995; Carpenter et al.,1997), was recently described as a new species on the basis of 13 specimens (Motomura and Iwatsuki, 2001b).
Five Polydactylus species: P. malagasyensis, P. microstomus, P. mullani, P. persicus and P. sextarius, are characterizedby having all pectoral-fin rays branched, except the uppermost 1 or 2, the vomer without teeth and a large black spot
anteriorly on the lateral line. Polydactylus persicus can be easily distinguished from P. microstomus and P. mullani by the
number of pectoral filaments [6 versus 5 (rarely asymmetrically 5 and 6) and 7 (rarely asymmetrically 6 and 7), respectively,
in P. microstomus and P. mullani].

Polydactylus persicus differs from P. sextarius in having a well-developed swimbladder (length about 40 to 45% of standard
length versus an atrophied, string-like swimbladder, about 20% of standard length in the latter; see Motomura and Iwatsuki,
2001b: fig. 7), higher counts of gillrakers [29 to 35 (mode 32) versus 25 to 30 (mode 28) in P. sextarius]. Furthermore, thesecond dorsal-fin spine length in P. persicus [6 to 9% (mean 7%) of standard length] is significantly longer than that of
P. sextarius [5 to 8% (mean 6%) of standard length]. Comparisons of P. persicus with P. malagasyensis are given in theaccount of the latter.

Polydactylus plebeius (Broussonet, 1782)

Fig. 116; Plate IVd-h

Polynemus plebeius Broussonet, 1782: described on twentyseventh page from the table of contents, eighth plate (seventh
species) (no pagination) [original locality: Tahiti, Society Islands and Tanna Island, New Hebrides Islands, Vanuatu; type
locality: Tahiti, Society Islands, based on a neotype (FMNH 108655, 88 mm standard length) designated by Motomura,
Iwatsuki and Yoshino, 2001].
Synonyms: Polynemus emoi Lacepède, 1803: 410, 412 (type locality: Tahiti, Society Islands, replacement name forP. plebeius Broussonet; no types known, see Motomura, Iwatsuki and Yoshino, 2001). Polynemus lineatus Lacepède,
1803: 410, pl. 13, fig. 2 [type locality: Réunion Island, Mascarene Islands; holotype (MNHN A. 5440, stuffed specimen,
255 mm standard length)]. Polynemus niloticus Shaw, 1804:151 [type locality: Nile River, Africa (but probably erroneous,see Motomura, Iwatsuki and Yoshino, 2001), based on a figure and description by J. Bruce; no types known]. Polynemus
commersonii Shaw, 1804: 156 (type locality: Indian seas, based on a figure by P. Commerson; replacement name forP. lineatus Lacepède; no types known, see Motomura, Iwatsuki and Yoshino, 2001). Polynemus lineatus Günther, 1860
(not of Lacepède): 327 (primary homonym of P. lineatus Lacepède). Polynemus taeniatus Günther, 1860: 526 [type locality:
Ambon, Indonesia and Guadalcanal, Solomon Islands; replacement name for P. lineatus Günther; 2 syntypes (BMNH
1855.11.7.35, 139 mm standard length; BMNH 1858.4.21.85, 147 mm standard length)]. Polydactylus agonasi Jordan and
McGregor, 1906: 814, unnumbered figure on page 815 [type locality: Tokyo, Japan; holotype (USNM 55608, 155 mm
standard length); 2 paratypes (CAS 109879, 105 to 148 mm standard length)]. Polynemus lydiae Curtiss, 1938: 43 (typelocality: Tahiti, Society Islands; no types known, see Motomura, Iwatsuki and Yoshino, 2001).
FAO Names: En - Striped threadfin; Fr - Barbure rayé; Sp - Barbudo rayado.

Fig. 116 Polydactylus plebeius

Threadfins of the World

69

Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 25 to 32% (mean 28%) of standard
length; head length 25 to 34% (mean 31%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla reaching to or extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw length 13 to
16% (mean 15%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw welldeveloped, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids.
Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal
fin with I spine and 12 or 13 (mode 13) soft rays; anal fin with III spines and 11 or 12 (mode 11) soft rays, anal-fin baseapproximately equal to second dorsal-fin base; pectoral fin with 16 to 18 (mode 17, rarely 15) rays (all rays unbranched), its
length 17 to 28% (mean 20%) of standard length, posterior tip not reaching to level of posterior tip of pelvic fin; pectoral
filaments 5; first pectoral filament shortest, just short of or reaching to level of pelvic-fin origin; second to fourth pectoral
filaments not reaching to level of posterior tip of pelvic fin; fifth pectoral filament longest, its length 22 to 40% (mean 32%) of
standard length, reaching to or extending beyond level of posterior tip of pelvic fin; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 33 to 44% (mean 39%) and lower lobe 30 to 41% (mean 36%) of
standard length. Pored lateral-line scales 60 to 68 (mode 63); lateral line simple, extending from upper end of gill opening toupper end of lower caudal-fin lobe; scale rows above lateral line 8 or 9 (mode 8), below 12 or 13 (mode 12). Gillrakers 9 to 14(mode 11) on upper limb, 13 to 18 (mode 15) on lower limb, 24 to 32 (mode 26) total. Vertebrae 10 precaudal and 14 caudal;
supraneural bones 3. Swimbladder present. Colour: Head and upper sides of trunk tinged slightly blackish silver, becoming
lighter silver on lower sides; abdominal region white; snout semi-transparent; first and second dorsal fins and caudal fin palewith blackish posterior margins; pectoral-fin membrane blackish; pectoral filaments white; anterior margins and origins ofpelvic and anal fins white, other parts dusky yellowish white; 7 or 8 prominent dark stripes along scale rows above lateral line,
7 to 9 faint stripes below.
Geographical Distribution:
Widely distributed in the
Indo-Pacific ranging from South
Africa to French Polynesia(Fig. 117). In the Pacific Ocean,
the northernmost and southernmost recorded ranges of thespecies are Korea and Port
Stephens, New South Wales,
Australia, respectively. There
are no records of P. plebeius
having been collected from the
Red Sea or Persian Gulf.
Habitat and Biology: Occurs
along shallow, sandy or muddy
200 m
coastal beaches to relatively
deep waters (less than 122 m),as well as in estuaries. The
species also occurs offshore in
Fig. 117 Polydactylus plebeius
Miyazaki, southern Japan
Known distribution
during the rainy (June and July)and typhoon seasons (September and October) (Motomura, Iwatsuki and Yoshino, 2001). Apparently, P. plebeius usually
inhabits estuaries and shallow coastal waters at Miyazaki, but migrates during periods of high rainfall. Large numbers ofjuveniles are taken from river mouths. The species usually forms loosely-associated schools throughout life.
Hida (1967) reported the sex composition of 10 specimens (180 to 264 mm standard length) of P. plebeius from western
Pakistan as: 1 hermaphrodite, 1 immature male and 8 immature females. No other biological studies, including maturation,
size at maturity, spawning, spawning grounds or age, have been published.
Venkataramam (1960) examined the stomachs of 42 specimens (99 to 185 mm standard length) from inshore waters off
Kozhikode, southwestern India, finding 15 specimens to be empty and the remainder gorged with a crab, Emerita asiatica.
In addition, a congener, P. mullani, was found in the stomach of a single individual P. plebeius from Pakistan (Hida, 1967).
Feeding behaviour of the species in an aquarium was reported by Motomura, Sado and Kimura (2002).
Size: Maximum standard length at least 45 cm (Feltes in Carpenter and Niem, 2001).
Interest to Fisheries: An important fishery species for South Asia, Southeast Asia, and (especially) Melanesia and
Polynesia, caught by trawls, gill nets, handlines and beach seines.
Local Names: FIJI: Uculuka; INDONESIA: Kuro, Lajan, Lausan, Mamangi, Manangi, Sumbal, Tapasan; JAPAN:
Tsubame-konoshiro; MALAYSIA: Senangin, Senangin buih; MARIANAS: Pááwánér; MAURITIUS: Mulet bâtard, Thread-fish;
MOZAMBIQUE: Barudo raiado; MYANMAR: Za yaw; NEW CALEDONIA: Hwa-n le kale-n, Noxan; PAPUA NEW GUINEA:
Common threadfin; SAMOA; I’ausi, Umi’umia; SOMALIA: Samaduul; SOUTH AFRICA: Gestreepe, Striped threadfin; SRI
LANKA: Barmeen, Polekala; TAHITI: Moi; UNITED REPUBLIC OF TANZANIA: Kupe, Mkizi komo maji; TUAMOTO: Moi.
Literature: Motomura, Iwatsuki and Yoshino (2001); Motomura (2002).

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FAO Species Catalogue for Fishery Purposes No. 3

Remarks: P. plebeius, which is the oldest available name for Indo-Pacific species in that genus, was originally described byBroussonet (1782) from Tahiti, Society Islands and Tanna Island, New Hebrides Islands, Vanuatu (syntypes apparently lost).
Although the original description of P. plebeius is very poor, the figure clearly shows 16 pectoral fin rays, 5 pectoral filaments
and several longitudinal dark stripes along each scale row on the lateral body surface.
The holotype (a dried specimen) of Polynemus lineatus, 2 syntypes of P. taeniatus and the holotype and 2 paratypes of
Polydactylus agonasi were all found to be conspecific with P. plebeius (see Motomura, Iwatsuki and Yoshino, 2001).
Günther (1860) described Polynemus lineatus as a new species on the basis of 2 specimens from Ambon, Indonesia andGuadalcanal, Solomon Islands. Because that name was preoccupied by P. lineatus of Lacepède (1803), Günther (1860)
proposed a replacement name, P. taeniatus, in an addenda to the original description of the species. In addition, Günther(1860) described P. plebejus (sic) (= plebeius) as being characterized by 5 pectoral filaments and a black blotch on thelateral line near its origin. Because the characters for P. plebejus given by Günther (1860) agree with those diagnostic of
Polydactylus microstomus, it is considered that he had misidentified examples of the latter.
Lacepède (1803) and Shaw (1804) proposed new names, Polynemus emoi, for P. plebeius and P. commersonii forP. lineatus, respectively, but did not list any specimens. The new names have no standing in nomenclature, because theprinciple of priority was not followed, contrary to the requirements of Article 23 (ICZN-1999). Polynemus niloticus was
described by Shaw (1804) as a new species on the basis of a description and figure by J. Bruce. According to Shaw (1804),
the species had a unique character, a reddish snout. However, the snout of polynemid fishes is semi-transparent andcartilagenous and easily damaged, causing a reddish coloration. Polynemus niloticus is also considered conspecific with
Polydactylus plebeius (Cantor, 1849; Daget and Njock in Daget et al., 1986; Motomura, Iwatsuki and Yoshino, 2001).

Polynemus lydiae was described by Curtiss (1938) as a new species from Tahiti, Society Islands (no types known).
According to the original description, the species was characterized by 5 pectoral filaments and longitudinal black stripes on
the lateral body surface, characteristics being consistent with those of Polydactylus plebeius. Therefore, Polynemus lydiae
is also regarded as a junior synonym of the latter (Motomura, Iwatsuki and Yoshino, 2001).
Fricke (1999) was synonymized Sciaena pentadactyla Lacepède, 1802 with P. plebeius. However, Motomura, Iwatsuki and
Yoshino (2001) recognized that S. pentadactyla is not a junior synonym of P. plebeius, although the taxonomic status of theformer is still unknown (treated as nomen dubium).
To date, only P. plebeius has been recognized as having 5 pectoral filaments and several prominent dark stripes along thelongitudinal scale rows above and below the lateral line. Two other species, P. bifurcus and P. siamensis, with thecharacters has recently been described from Indonesia and Thailand, respectively. Comparisons of P. plebeius with
P. bifurcus are given in the account of the latter.
Although P. plebeius is very similar to P. siamensis in having the above characters, the former differs in having higher countsof pectoral-fin rays [16 to 18 (mode 17; rarely 15, 1 of 100 specimens examined by Motomura, Iwatsuki and Yoshino, 2001)
versus 15 in the latter], scales above and below the lateral line [8 or 9 (mode 8) and 12 or 13 (mode 12), respectively versus 7
and 10 or 11 (mode 11), respectively in P. siamensis], pored lateral-line scales [60 to 66 (mode 63; rarely 68, 1 of 96
specimens) versus 54 to 58 (mode 54) in P. siamensis] and gillrakers [9 to 14 (mode 11) upper series, 13 to 18 (mode 15)
lower and 24 to 30 (mode 26; rarely 32, 1 of 96 specimens) total versus 9 or 10 (mode 10), 13 or 14 (mode 13) and 22 to 24(mode 23), respectively in P. siamensis] (see Motomura, Iwatsuki and Yoshino, 2001: table 3). Furthermore, P. plebeius
tends to have a slightly shorter upper jaw [13 to 16% (mean 15%) of standard length] than P. siamensis [16 to 17% (mean17%) of standard length], although the proportional measurements for upper-jaw length overlapped between the 2 species(Motomura, Iwatsuki and Yoshino, 2001: fig. 3).

Polydactylus quadrifilis (Cuvier, 1829)

Fig. 118; Plate Va

Polynemus quadrifilis Cuvier in Cuvier and Valenciennes, 1829: 390, pl. 68 [type locality: Senegal; holotype (MNHN 756,
lacking caudal fin and half of caudal peduncle, 380 mm from anterior tip of snout to posterior end of remaining peduncle)].
Synonyms: None.

Threadfins of the World

71

FAO Names: En - Giant African threadfin; Fr - Gros capitaine; Sp - Barbudo gigante africano.

Fig. 118 Polydactylus quadrifilis
Diagnostic Features: A large species. Body depth at first dorsal-fin origin 24 to 27% (mean 25%) of standard length; headlength 30 to 34% (mean 32%) of standard length. Snout pointed; occipital profile nearly straight. Posterior margin of maxilla
not reaching to or just reaching to level of posterior margin of adipose eyelid; upper-jaw length 13 to 14% (mean 14%) of
standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth
restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of
preopercle serrated. First dorsal fin with VIII spines, second spine more robust than others; second dorsal fin with I spine
and 13 soft rays; anal fin with III spines and 11 soft rays, anal-fin base approximately equal to second dorsal-fin base;pectoral fin with 12 or 13 (mode 13) rays (all rays unbranched), its length 20 to 24% (mean 22%) of standard length, posterior
tip not reaching to level of posterior tip of pelvic fin; pectoral filaments 4; first (shortest) to third pectoral filaments, extending
beyond level of pelvic-fin origin, but not reaching to level of posterior tip of pelvic fin; fourth pectoral filament longest, its
length 27 to 39% (mean 33%) of standard length, just short of or extending slightly beyond level of posterior tip of pelvic fin;caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 34 to 41% (mean 38%) andlower lobe 33 to 40% (mean 37%) of standard length. Pored lateral-line scales 70 or 71 (mode 70); lateral line simple,
extending from upper end of gill opening to upper end of lower caudal-fin lobe; scale rows above lateral line 8 or 9 (mode 9),
below 11 to 13 (mode 11). Gillrakers 8 or 9 (mode 9) on upper limb, 12 to 14 (mode 14) on lower limb, 21 to 23 (mode 23)
total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 2. Swimbladder present, well developed. Colour: Headand upper sides of trunk tinged slightly blackish silver, becoming lighter silver on lower sides; abdominal region white; snout
semi-transparent; first and second dorsal fins and caudal fin pale with blackish posterior margins; pectoral fin vivid yellow;
pectoral filaments white; anterior margins and origins of pelvic and anal fins white, other parts dusky.
Geographical Distribution: On the west coast of Africa
ranging from Senegal to the Congo (Fig. 119).
Habitat and Biology: Occurs on sandy and muddy
bottoms in shallow waters (less than 55 m), sometimes
also in brackish waters. Feeds mainly on crabs(Longhurst, 1960: table 1) and fishes.
Size: Maximum total length 2 m (Daget and Njock in
Daget et al., 1986; Njock in Quéro et al., 1990), common
to 1.5 m (Allen in Fischer et al., 1981).
Interest to Fisheries: One of the most importantfishery (see Table 3; mainly caught by trawl, gill net andbeach seine) and sport species on the west coast of
Africa.
Local Names: ANGOLA: Barbudo; BENIN: Chikoué,
Gainfio; CAPE VERDE: Barbo, Barbudo-gigante,
Capitão, Peixe-barba; GUINEA: Sori; SENEGAL:
Capitaine, Njaane, Njaane jaara.
Literature: Allen in Fischer et al. (1981); Daget andNjock in Daget et al. (1986); Njock in Quéro et al. (1990).

200 m

Fig. 119 Polydactylus quadrifilis
Known distribution

Remarks: P. quadrifilis is easily distinguished from other congeners by having 4 pectoral filaments (5 to 9 in all othercongeners).

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Polydactylus sexfilis (Valenciennes, 1831)

Fig. 120; Plate Vb-d

Polynemus sexfilis Valenciennes in Cuvier and Valenciennes, 1831: 515 [type locality: Mauritius; lectotype (MNHN 9731,
265 mm standard length) designated by Motomura, Iwatsuki and Kimura, 2001a; 2 paralectotypes (MNHN A. 3027, 96 mm
standard length; MNHN 9728, 92 mm standard length)].
Synonyms: Polynemus kuru Bleeker, 1853b: 600 [type locality: Jakarta, Java, Indonesia; 2 syntypes (RMNH 6006, 87 to
198 mm standard length) determined from 5 Bleeker specimens by Motomura, Iwatsuki and Kimura, 2001a].
FAO Names: En - Sixfinger threadfin; Fr - Barbure à six doigts; Sp - Barbudo de seis dedos.

Fig. 120 Polydactylus sexfilis

Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 26 to 32% (mean 29%) of standard
length; head length 27 to 34% (mean 32%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla reaching to level of posterior margin of adipose eyelid; upper-jaw length 13 to 15% (mean 15%) of standard
length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentary teeth restricted todorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 12 or 13(mode 13) soft rays; anal fin with III spines and 11 or 12 (mode 11) soft rays, anal-fin base approximately equal to second
dorsal-fin base; pectoral fin with 15 or 16 (mode 16) rays, its length 20 to 23% (mean 21%) of standard length, posterior tip
not reaching to level of posterior tip of pelvic fin; all pectoral-fin rays unbranched except larger specimens (in larger
specimen, third pectoral-fin ray bifurcated at tip, fourth and fifth pectoral-fin rays divided into 3 at tip, and remaining
pectoral-fin rays unbranched); pectoral filaments 6; first pectoral filament shortest, not reaching to level of pelvic-fin origin;second to fifth pectoral filaments extending beyond level of pelvic-fin origin; sixth pectoral filament longest, its length 28 to41% (mean 35%) of standard length, not reaching to or extending beyond level of posterior tip of pelvic fin; caudal fin deeply
forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 39 to 48% (mean 44%) and lower lobe 37 to
46% (mean 41%) of standard length. Pored lateral-line scales 60 to 67 (mode 64); lateral line simple, extending from upper
end of gill opening to upper end of lower caudal-fin lobe; scale rows above lateral line 8 to 10 (mode 9), below 12 to 14 (mode13). Gillrakers 11 to 14 (mode 13) on upper limb, 15 to 18 (mode 17) on lower limb, 27 to 31 (mode 30) total. Vertebrae 10
precaudal and 14 caudal; supraneural bones 3. Swimbladder present. Colour: Upper sides of head and trunk tinged silvery
brown, becoming more silver on lower sides; first dorsal fin black; posteriormost ray of second dorsal fin white, remainder of
fin greyish black; base of pectoral fin greyish black, becoming dense black posteriorly; bases and tips of pectoral filaments
white, middle parts of filaments greyish black; anterior part of pelvic fin greyish black, remainder of fin white; posteriormost
ray of anal fin white, remainder of fin greyish black; caudal fin uniformly greyish black; 7 to 9 prominent dark stripes along
longitudinal scale rows above lateral line, 1 to 12 faint stripes below (sometimes no stripes, especially in smaller specimens).

Threadfins of the World

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Geographical Distribution: Widelydistributed in the Indo-Pacific region,
generally in the vicinity of oceanicislands (Fig. 121). Specimens nevercollected from the coast of theAustralian continent. The species is
considered to be less dependent uponlarge fresh-water rivers (based on
known locality data) than other,heavily-dependent polynemid species.
Distributional implications of thespecies in Japanese waters werediscussed by Motomura, Burhanuddin
and Iwatsuki (2000).
Habitat and Biology: Occurs along
shallow (less than 50 m), sandy androcky coastal beaches, in lagoons andnear reef areas in the vicinity ofoceanic islands, frequently in zones ofturbulence. Feeds mainly on
crustaceans and teleosts.

200 m

Fig. 121 Polydactylus sexfilis
Known distribution

Matures first as males at between 20 and 25 cm fork length, subsequently transforming into functional females at between
30 and 40 cm fork length following a hermaphroditic stage (Santerre and May, 1977; May, Akiyama and Santerre, 1979).
Displays a lunar spawning rhythm. Observations of spawning behaviour given by May, Akiyama and Santerre (1979). The
potential of the species for aquaculture has also been studied (e.g. May, 1976; Ostrowski et al., 1996; Leber, Brennan andArce, 1998; Masuda and Ziemann, 2000).
Size: Maximum standard length at least 45 cm (Motomura and Senou, 2002).
Interest to Fisheries: One of the most important commercial and sport fisheries’ species in Hawaii, where it is actively
cultivated.
Local Names: AUSTRALIA: Six-fingered threadfin; BANGLADESH: Dagi, Golden sixthread tesselfish, Topshi; FRENCHPOLYNESIA: Moi; HAWAII: Moi, Moili’I, Pacific thread-fin fish, Pacific threadfin; INDONESIA: Mulut tikus; JAPAN:
Nan’you-agonashi; MARSHALL ISLANDS: Atkaru, Becadulce; PAPUA NEW GUINEA: Six-fingered threadfin; PITCAIRN:
Moi; SAMOA: I’ausi, Umiumia, Umi’umia; SEYCHELLES: Mulet barbe; SOLOMON ISLANDS: Bou na pana, Six-fingered
threadfin; SRI LANKA: Gatha, Kala; TAHITI: Moi; TUAMOTO ISLANDS: Moi;
Literature: Motomura, Burhanuddin and Iwatsuki (2000); Motomura, Iwatsuki and Kimura (2001a); Motomura and Senou(2002); Motomura (2002).
Remarks: P. sexfilis has usually been regarded as a valid species (e.g. Sauvage, 1891; Munro, 1955; Jones and Kumaran,
1980). Although P. kuru has also been regarded as a valid species (e.g. Kendall and Goldsborough, 1911; Weber and deBeaufort, 1922; Mishra and Krishnan, 1993), type materials of the 2 have never been directly compared. Recent
examination of 2 syntypes of Polynemus kuru showed them to correspond closely with lectotype and 2 paralectotypes ofP. sexfilis. Therefore, P. kuru is regarded as a junior synonym (Motomura, Iwatsuki and Kimura, 2001a).
Fricke (1999) synonymized Polydactylus sexfilis with P. sextarius, but his opinion was clearly erroneous. Polydactylus
sexfilis can be easily distinguished from P. sextarius in lacking a large black spot anteriorly on the lateral line (present in thelatter) and possessing villiform teeth in broad bands on the vomer (teeth absent).
Although Munro (1955, 1967) and Menon and Babu Rao in Fischer and Bianchi (1984) described the pectoral fin of
P. sexfilis as being black, the fin membrane pigmentation was found to be quite variable (from no pigment to entirely black).
The pectoral-fin rays of P. sexfilis were thought to remain unbranched throughout life. However, the pectoral fin of a large
specimen (KPM-NI 8127, 456 mm standard length, from the Izu Islands, Japan) was found to have 3 branched rays.
Although the first, second, and sixth to sixteenth pectoral-fin rays of the specimen were unbranched, the third, and fourth andfifth rays were bifurcated and divided into 3, respectively, near the tips. This change from unbranched to branched fin rays isapparently related to fish growth. Five similar Polydactylus species: P. malagasyensis, P. microstomus, P. mullani,
P. persicus and P. sextarius, characterized by having a large black spot anteriorly on the lateral line, are also known to havebranched pectoral-fin rays. However, the pectoral-fin ray condition in these species differs from that of P. sexfilis in that all of
the rays, except the uppermost 1 or 2, are branched throughout life.
Like P. sexfilis and 2 other Indo-Pacific Polydactylus species, such as P. longipes and P. nigripinnis, are characterized byhaving 6 pectoral filaments and lacking a large black spot anteriorly on the lateral line. Comparisons of P. sexfilis with
P. longipes and P. nigripinnis are given in the accounts of the latter 2 species.

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FAO Species Catalogue for Fishery Purposes No. 3

Polydactylus sexfilis is most similar to P. plebeius in overall body appearance, but differs from the latter in having higher
counts of pectoral filaments (6 versus 5 in the latter), lower counts of pectoral-fin rays [15 or 16 (mode 16) versus 16 to 18(mode 17; rarely 15)] and higher counts of gillrakers [27 to 31 (mode 30) versus 24 to 32 (mode 26)]. Furthermore, P. sexfilis
tends to have a longer second dorsal-fin ray than P. plebeius, although the proportional length measurements for suchoverlapped between the 2 species [21 to 30% (mean 26%) of standard length versus 19 to 28% (mean 22%) of standard
length; see Motomura, Iwatsuki and Kimura, 2001a: fig. 3]. Both anal- and caudal-fin rays of P. sexfilis are also slightly
longer than those of P. plebeius [anal fin: 20 to 26% (mean 23%) of standard length versus 17 to 23% (mean 19%); upper
caudal fin: 39 to 48% (mean 44%) versus 33 to 44% (mean 39%); lower caudal fin: 37 to 46% (mean 41%) versus 30 to 41%(mean 36%)].

Polydactylus sextarius(Bloch and Schneider, 1801)

Fig. 122; Plate Ve-f

Polynemus sextarius Bloch and Schneider, 1801: 18, pl. 4 [type locality: Tranquebar, Tamil Nadu, India; holotype (ZMB 565,
125 mm standard length)].
Synonyms: None.
FAO Names: En - Blackspot threadfin; Fr - Barbure à tâche noire; Sp - Barbudo de mancha negra.

Fig. 122 Polydactylus sextarius
Diagnostic Features: A small- to medium-sized species. Body depth at first dorsal-fin origin 29 to 36% (mean 32%) of
standard length; head length 30 to 35% (mean 33%) of standard length. Snout pointed; occipital profile nearly straight.
Posterior margin of maxilla not reaching to or extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw
length 12 to 14% (mean 13%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw
well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on palatines and ectopterygoids;
vomerine tooth plate covered with skin and teeth absent; palatines inwardly turned anteriorly. Posterior margin of preopercle
serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 12 or 13(mode 13) soft rays; anal fin with III spines and 11 to 13 (mode 12) soft rays, anal-fin base approximately equal to second
dorsal-fin base; pectoral fin with 13 to 15 (mode 14) rays (all rays unbranched, except uppermost 1 or 2), its length 19 to 25%(mean 22%) of standard length, posterior tip not reaching to level of posterior tip of pelvic fin; pectoral filaments 6, first
filament shortest; sixth pectoral filament longest, its length 20 to 29% (mean 25%) of standard length, not reaching to level ofposterior tip of pectoral fin; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe
33 to 41% (mean 36%) and lower lobe 31 to 41% (mean 34%) of standard length. Pored lateral-line scales 45 to 51 (mode46); lateral line simple, extending from upper end of gill opening to upper end of lower caudal-fin lobe; scale rows above
lateral line 5 or 6 (mode 6), below 8 to 11 (mode 11). Gillrakers 10 to 14 (mode 12) on upper limb, 14 to 18 (mode 16) on lower
limb, 25 to 30 (mode 28) total. Vertebrae 10 precaudal and 14 caudal; supraneural bones 3. Swimbladder atrophied, like afine string. Colour: Head and upper sides of trunk with slightly darkish silver tinge, becoming lighter on lower sides; snout

Threadfins of the World75

semi-translucent; posterior margin of first and second dorsal fins and caudal fin slightly blackish, other parts translucent;posterior tip of anal fin whitish, other parts blackish; pectoral-fin membrane white with scattered melanophores; pectoral
filaments whitish; anterior parts of pelvic fin slightly yellowish white, other parts whitish; a large black spot anteriorly onlateral line.
Geographical Distribution: Known from theeastern Indian to western Pacific oceans where it
ranges from southwestern India to Papua New
Guinea and Miyazaki, Kyushu Island, Japan(Fig. 123). Although Munro (1967) reported asingle example (CSIRO C. 1002, 157 mm standard
length) collected from Papua New Guinea,
P. sextarius is considered to occur rarely west of
Huxley’s line (sensu George in Whitmore, 1981).
Therefore, it may occur rarely in the Philippines,
eastern Indonesia, including the Lesser Sunda
Islands, Sulawesi Island, Moluccas and Irian Jaya,
and Papua New Guinea.
Habitat and Biology: Occurs on sandy andmuddy bottoms, being taken from depths of 16 to
73 m. The species feeds mainly on shrimps (lessthan 20 mm) and amphipods, other importantdietary items (in 50% or more of the stomachs)
being crabs, mysids, fishes and polychaetes(Hida, 1967).

200 m

Fig. 123 Polydactylus sextarius
Known distribution

Hida (1967) reported the sex composition of 140
specimens of P. sextarius from the Bay of Bengal, India as: 112 hermaphrodites ranging from 50 to 149 mm standard length,
27 mature females ranging from 100 to 105 mm standard length and 1 sex undetermined. The hermaphrodites withrelatively large testes were considered as mature males (90 to 134 mm standard length) and those with granulated ova about
0.3 to 0.5 mm diameter as mature female (105 to 149 mm standard length). The species undergoes a protandrous change
through a juvenile-hermaphrodite-female progression, first reaching sexual maturity as a functional male hermaphrodite.
There is no evidence that females develop directly from juveniles.
Size: Maximum standard length at least 17 cm (Motomura and Iwatsuki, 2001b).
Interest to Fisheries: An important fishery species, generally caught by trawl in the Bay of Bengal and Thailand.
Local Names: INDIA: Blackspot threadfin; JAPAN: Kataguro-agonashi; MALAYSIA: Benong, Gubal, Kurau, Lelaoh tanah,
Melong, Mancong, Senangin, Senangin buis, Senohong; MYANMAR: Za yaw; PAPUA NEW GUINEA: Black-spot threadfin;
SRI LANKA: Kutlikala.
Literature: Motomura et al. (1999); Motomura and Iwatsuki (2001b); Motomura (2002).
Remarks: Fricke (1999) synonymized Polynemus sexfilis and P. astrolabi under Polydactylus sextarius. However,
Polydactylus (= Polynemus) sexfilis is presently recognized as a valid species (Motomura, Iwatsuki and Kimura, 2001a)and P. astrolabi was synonymized with Galeoides decadactylus (Motomura, Iwatsuki and Kimura, 2001b).
Five Polydactylus species: P. malagasyensis, P. microstomus, P. mullani, P. persicus and P. sextarius, are characterizedby having all pectoral-fin rays branched, except the uppermost 1 or 2, the vomer without teeth and a large black spot
anteriorly on the lateral line. Polydactylus sextarius can be easily distinguished from P. microstomus and P. mullani by the
number of pectoral filaments [6 versus 5 (rarely asymmetrically 5 and 6) and 7 (rarely asymmetrically 6 and 7), respectively,in the latter]. Comparisons of P. sextarius with P. malagasyensis and P. persicus are given in the accounts of the latter 2
species.

Polydactylus siamensis Motomura, Iwatsuki and Yoshino, 2001

Fig. 124; Plate Vg

Polydactylus siamensis Motomura, Iwatsuki and Yoshino, 2001: 122, fig. 1B [type locality: northern Gulf of Thailand(purchased at Samyan market, Bangkok); holotype (URM-P 14050, 252 mm standard length); paratype (MUFS 18280,
137 mm standard length)].
Synonyms: None.

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FAO Names: En - Largemouth striped threadfin; Fr - Barbure à grande bouche; Sp - Barbudo rayado de boca grande.

Fig. 124 Polydactylus siamensis
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 30 to 33% (mean 32%) of standard
length; head length 31 to 34% (mean 32%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla extending slightly beyond level of posterior margin of adipose eyelid; upper-jaw length 16 to 17% (mean17%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed, dentaryteeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of
preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and
12 or 13 (mode 13) soft rays; anal fin with III spines and 11 soft rays, anal-fin base less than second dorsal-fin base; pectoral
fin with 15 rays (all rays unbranched), its length 20 to 22% (mean 21%) of standard length, posterior tip not reaching to levelof posterior tip of pelvic fin; pectoral filaments 5; first pectoral filament shortest, just short of or reaching to level of pelvic-fin
origin; second to fourth pectoral filaments not reaching to level of posterior tip of pelvic fin; fifth pectoral filament longest, its
length 31 to 43% (mean 36%) of standard length, reaching to or extending beyond level of posterior tip of pelvic fin; caudal fin
deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 38 to 41% (mean 40%) and lower lobe
34 to 37% (mean 35%) of standard length. Pored lateral-line scales 54 to 58 (mode 54); lateral line simple, extending fromupper end of gill opening to upper end of lower caudal-fin lobe; scale rows above lateral line 7, below 10 or 11 (mode 11).
Gillrakers 9 or 10 (mode 10) on upper limb, 13 or 14 (mode 13) on lower limb, 22 to 24 (mode 23) total. Vertebrae 10
precaudal and 14 caudal; supraneural bones 3. Swimbladder absent. Colour: 7 or 8 prominent dark stripes along scalerows above lateral line, 7 to 9 faint stripes below.
Geographical Distribution: Currently known only from
Bangkok, Songkhla and Phuket, Thailand (Fig.125).
According to local fishermen at Bangkok, SamutPrakan and Prachuap Khirikhan, Thailand, P. siamensis
rarely occurs in the Gulf of Thailand and the record fromthe Gulf of Thailand is based only on 3 specimens. The
species is mainly distributed in the Andaman Sea.
Habitat and Biology: No data are available.
Size: Maximum standard length at least 25 cm(Motomura, Iwatsuki and Yoshino, 2001).
Interest to Fisheries: None.
Local Names: MYANMAR: Za yaw.
Literature: Motomura, Iwatsuki and Yoshino (2001);
Motomura (2002).

200 m

Remarks: P. siamensis was recently described as a
Fig. 125 Polydactylus siamensis
new species by Motomura, Iwatsuki and Yoshino (2001)
and characterized by having 5 pectoral filaments,
Known distribution
several longitudinal dark stripes along each scale row
on the lateral body surface, and lower counts of pectoral-fin rays (15), scales above and below the lateral line [7 and 10 or 11(mode 11), respectively], pored lateral-line scales [54 to 58 (mode 54)] and gillrakers [9 or 10 (mode 10) upper series, 13 or
14 (mode 13) lower and 22 to 24 (mode 23) total]. Comparisons of P. siamensis with related species, P. bifurcus andP. plebeius, are given in the accounts of the latter 2 species.

Threadfins of the World

77

Polydactylus virginicus (Linnaeus, 1758)

Fig. 126; Plate Vh

Polynemus virginicus Linnaeus, 1758: 317 (type locality: America; no types known).
Synonyms: Polynemus naso Walbaum, 1793: 108 (type locality: probably Brazil; no types known). Polynemus mango
Lacepède, 1803: 411, 413 (type locality: America; no types known). Polydactylus plumierii Lacepède, 1803: 419, 420, pl.
14, fig. 3 (type locality: unknown; no types known). Polynemus americanus Cuvier 1829a: 155 [type locality: Americanseas; 3 syntypes (MNHN 5504, 141 mm standard length; MNHN A. 3028, 157 mm standard length; MNHN A. 3031, 121 mmstandard length)]. Polynemus antillarum Perugia, 1896: 16 [type locality: St. Pierre, Martinique Island, West Indies;
holotype (MSNG 6695, 84 mm standard length)].
FAO Names: En - Barbu; Fr - Barbure de flaque; Sp - Barbudo de charco.

Fig. 126 Polydactylus virginicus
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 29 to 32% (mean 30%) of standard
length; head length 29 to 34% (mean 32%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla just short of or reaching to level of posterior margin of adipose eyelid; upper-jaw length 13 to 16% (mean14%) of standard length; depth of posterior margin of maxilla less than eye diameter; lip on lower jaw well developed,
dentary teeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior
margin of preopercle serrated. First dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I
spine and 11 or 12 (mode 12) soft rays; anal fin with III spines and 11 to 14 (mode 13) soft rays, anal-fin base greater thansecond dorsal-fin base; pectoral fin with 14 to 16 (mode 15) rays (all rays unbranched), its length 23 to 28% (mean 26%) of
standard length, posterior tip not reaching to, just reaching to or extending slightly beyond level of posterior tip of pelvic fin;pectoral filaments 7; first pectoral filament shortest, not reaching to or extending slightly beyond level of pelvic-fin origin;second to fifth pectoral filaments extending beyond level of pelvic-fin origin; sixth pectoral filament just short of or
extending beyond level of posterior tip of pelvic fin; seventh pectoral filament longest, its length 36 to 47% (mean 42%) of
standard length, extending beyond level of anal-fin origin,sometimes extending beyond level of midpoint of anal-finbase; caudal fin deeply forked, upper and lower caudal-fin
lobes not filamentous, upper caudal-fin lobe 35 to 41%(mean 38%) and lower lobe 32 to 41% (mean 36%) of
standard length. Pored lateral-line scales 54 to 63 (mode58); lateral line bifurcated on caudal fin, extending toposterior margins of upper and lower caudal-fin lobes; scalerows above lateral line 6 to 8 (mode 7), below 10 to 12 (mode11). Gillrakers 26 to 30 (mode 28). Vertebrae 10 precaudal
and 14 caudal; supraneural bones 3. Swimbladder present,
well developed. Colour: Upper sides of head and trunk withslightly darkish silver tinge, becoming lighter on lower sides;
snout semi-translucent; margins of second dorsal andcaudal fins blackish, other parts translucent; intensity ofpigmentation of pectoral and upper parts of first dorsal finvery variable; pectoral filaments, pelvic and anal fins white.
200 m

Geographical Distribution: In the western Atlantic Ocean,
ranging from New Jersey, USA to Salvador, Brazil (Fig. 127).
However, the species does not occur in the northern andwestern Gulf of Mexico.

Fig. 127 Polydactylus virginicus
Known distribution

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FAO Species Catalogue for Fishery Purposes No. 3

Habitat and Biology: Inhabits sandy and muddy bottoms in coastal waters, estuaries and mangroves, as well as
sometimes occurring in the surf. A large number of young specimens aggregate in river mouths. The species feeds primarily
at night (Austin and Austin, 1971), taking mainly crustaceans, followed by chaetognaths, plant material and polychaetes. It
appears to have a prolonged spawning season because juveniles are commonly found throughout the year.
Size: Maximum total length 33 cm (Feltes in Carpenter, 2002).
Interest to Fisheries: Caught incidentally by seines and trawls and is of little commercial importance.
Local Names: BARBADOS: Beard fish; BELIZE: Barbú; CUBA: Barbi, Barbu, Barbudo, Catfish; DOMINICA: Barbudo;
DOMINICAN REP.: Barbú; GUADELOUPE: Barbi, Barbu argenté, Threadfin; GUYANA: Chinese nose, Threadfin,
Threadfish; HAITI: Sevenfingered threadfin; MARTINIQUE: Barbi, Barbu argenté, Threadfin; NICARAGUA: Barbudo decharco; PUERTO RICO: Barbudo, Threadfin; ST. LUCIA: Barbu; SURINAME: Barbu; Sardijntje.
Literature: Feltes in Carpenter (2003).
Remarks: Polydactylus virginicus, the most common species in the western Atlantic, has been regarded as a seniorsynonym of Polydactylus plumierii, Polynemus americanus and P. mango (see Eschmeyer, 1998). The holotype (MSNG
6695, 84 mm standard length) of Polynemus antillarum is also conspecific with specimens considered here as
Polydactylus virginicus, the former herein being regarded as a junior synonym of the latter.

Polynemus naso was originally described by Walbaum (1793) as a species locally named “Piracoada”, which occurred off
the Atlantic coast of Brazil. Although Eschmeyer (1998) believed that the species does not belong to the family
Polynemidae, Walbaum (1793) stated that P. naso is characterized by having 6 pectoral filaments and a cartilaginous snout,
which are included in the diagnostic characteristics of the family. Therefore, P. naso clearly belongs to the family
Polynemidae. Two polynemids, Polydactylus oligodon and P. virginicus, are also known to occur in Brazil (type locality of
Polynemus naso). Because Polydactylus virginicus is more common than P. oligodon and occasionally has 6 pectoral
filaments (as has P. plumierii), Polynemus naso is herein treated as a likely junior synonym of Polydactylus virginicus.
However, Polynemus naso has at no time been reported during the 200 years following its original description. Therefore, if
P. naso is a different species from Polydactylus virginicus and an earlier name of some other polynemid, the former cannot
become a valid taxon under the Reversal of Precedence (treated as nomen oblitum; Article 23.9, ICZN-1999).
Polydactylus virginicus and 2 other species, P. octonemus and P. oligodon, occur in the western Atlantic, being very similar
to each other in overall body appearance. Comparisons of P. virginicus with P. octonemus and P. oligodon are given in theaccounts of the latter 2 species.

Polynemus Linnaeus, 1758
Polynemus Linnaeus, 1758: 317 (type species: Polynemus paradiseus Linnaeus, 1758, see Opinion 93, ICZN-1926).
Synonyms: Polistonemus Gill, 1861: 277 (type species: Polynemus multifilis Temminck and Schlegel, 1843).
Diagnostic Features: Body and head elongate. Adipose eyelid poorly developed; eye diameter less than snout length. Lip
on lower jaw well developed, dentary teeth restricted to dorsal surface; width of tooth band on upper and lower jaws greaterthan space (on symphysis) separating tooth bands on opposing premaxillae; teeth villiform in broad bands on jaws, palatinesand ectopterygoids, vomerine tooth plate without teeth in some species. Posterior margin of maxilla extending well beyond
level of posterior margin of adipose eyelid. Posterior margin of preopercle serrated. Basisphenoid absent; sphenotics not
visible dorsally between anterior margins of parietal and pterotic. First dorsal fin with VII or VIII spines; second dorsal fin
with I spine and 13 to 18 soft rays; anal fin with II or III spines and 10 to 14 soft rays; anal-fin base less than head length;
pectoral fin with 14 to 19 rays, all unbranched; pectoral-fin insertion near midline of body; pectoral-fin base (including base ofpectoral filaments) less than upper-jaw length; pectoral filaments 7 to 16, extending beyond level of posterior tip of caudal
fin; caudal fin deeply forked, but upper and lower caudal-fin lobes not filamentous. Pored lateral-line scales 66 to 109; scalerows above lateral line 6 to 12, below 10 to 22. Total gillrakers 23 to 34. Vertebrae 10 precaudal and 15 caudal; supraneuralbones 2. Swimbladder present or absent.
Habitat and Biology: Polynemus species are generally found on sandy or muddy bottoms in fresh-water rivers, estuaries
and coastal waters, being more heavily dependent upon a fresh-water habitat than other genera. With the exception of
P. paradiseus, no biological information for members of the genus has been reported, owing to the confusion surrounding
specific identifications. Information on P. paradiseus is given below under the species’ account.
Geographic Distribution: Indo-West Pacific, ranging over continental shelves from western India to southern Viet Nam,
including Southeast Asian countries, but has at no time been recorded from the Philippines or Papua New Guinea.

Threadfins of the World

79

Interest to Fisheries: Important commercial fishes in South Asia and Southeast Asia, especially in India, Thailand and Viet
Nam. Some species have been exported to Japan as aquarium fishes.
Species: The genus comprises 8 species:

Polynemus aquilonaris: Thailand, Cambodia, Viet Nam and Lao People's Democratic Republic.
Polynemus dubius: Malaysia and Indonesia.
Polynemus hornadayi: Sarawak, Malaysia.
Polynemus kapuasensis: Kapuas River, Kalimantan
Polynemus melanochir dulcis: Lake Tonle Sap, Cambodia.
Polynemus melanochir melanochir: Lower Mekong River and related rivers, and Kalimantan.
Polynemus multifilis: Thailand and Indonesia.
Polynemus paradiseus: India to Thailand.
Remarks: Polynemus, which is the oldest available generic name for the family Polynemidae, was originally described by
Linnaeus (1758), who simultaneously included 3 species, P. paradiseus, P. quinquarius and P. virginicus, in the genus.
Polynemus paradiseus was later established as the type species of Polynemus by ICZN (1926). Polynemus quinquarius
and P. virginicus are presently regarded as members of Pentanemus and Polydactylus, respectively.
Polistonemus was originally proposed for Polynemus multifilis by Gill (1861), but characters of the former are identical with
Polynemus in all respects, except for an increased number of pectoral filaments. Accordingly, Polistonemus is considereda junior synonym of Polynemus (e.g. Roberts, 1989).
Polynemus and Parapolynemus are characterized by the following characters: eye diameter 1.3 or more in snout length (1.3 orless in other genera); pectoral-fin insertion near midline of body (well below midline of body); pectoral filaments longer thanstandard length (shorter than standard length in other genera, except in Pentanemus and Polydactylus macrophthalmus).
Polynemus differs from Parapolynemus in having the posterior margin of the preopercle serrated (largely not serrated in thelatter), higher counts of caudal procurrent rays (15 to 19 versus 12 to 14) and pectoral-filament length greater than 1.5 total
length (less than 1.5 total length). Further detailed comparisons of the 2 genera were given by Feltes (1993).

Key to the Species of Polynemus
1a. Pectoral filaments 13 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Pectoral filaments 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Pectoral filaments usually 15 on each side of body; pored lateral-line scales 100 to 110 (mode 103);
scale rows above lateral line 9 to 11 (mode 10), below 17 to 20 (mode 18); posterior tip of pectoral fin notextending beyond level of anal-fin origin in adults (Fig. 128, Plate VId) . . . . . . . . . . Polynemus kapuasensis(Kapuas river, Kalimantan)
2b. Pectoral filaments usually 14 of each side of body; pored lateral-line scales 83 to 99 (mode 86); scalerows above lateral line 7 or 8 (mode 8), below 14 to 18 (mode 15); posterior tip of pectoral fin reaching or
extending beyond level of anal-fin origin throughout life (Fig. 129, Plate VIg) . . . . . . . . Polynemus multifilis(Thailand and Indonesia)
3a. First dorsal-fin spines VII; anal-fin spines II or III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3b. First dorsal-fin spines VIII (first spine minute); anal-fin spines III . . . . . . . . . . . . . . . . . . . . . . . ® 5
100-110 pored lateral-line scales
9-11 scale rows
above lateral line

7-8 scale rows
above lateral line

15 pectoral
filaments

Fig. 128 Polynemus kapuasensis

83-99 pored lateral-line scales

14 pectoral filaments

Fig. 129 Polynemus multifilis

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4a. Pored lateral-line scales 90 to 103 (mode 94); scale rows above lateral line 8 to 12 (mode 11), below 16
to 21 (mode 18); gillrakers 26 to 28 (mode 26); occipital profile strongly concave; swimbladder present,
well developed (Fig. 130, Plate VIc) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynemus hornadayi(Sarawak, Malaysia)
4b. Pored lateral-line scales 66 to 71 (mode 70); scale rows above lateral line 6 or 7 (mode 7), below 10 to
12 (mode 11); gillrakers 30 to 34 (mode 32 or 33); occipital profile nearly straight; swimbladder absent(Fig. 131, Plate VIh) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynemus paradiseus(India to Thailand)
8-12 scale rows above lateral line

66-71 pored lateral-line scales

90-103 pored lateral-line scales
6-7 scale rows
above lateral line

16-21 scale rows
below lateral line

30-34
gillrakers

26-28
gillrakers

swimbladder absent

10-12 scale rows
below lateral line

swimbladder present

Fig. 130 Polynemus hornadayi

Fig. 131 Polynemus paradiseus

5a. Vomer with teeth; pectoral fin usually not black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 65b. Vomer without teeth; pectoral fin usually black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6a. Pored lateral-line scales 80 to 86 (mode 81); scale rows below lateral line 14 to 16 (mode 14, rarely 13 or17); gillrakers 25 to 29 (mode 27) (Fig. 132, Plate VIa) . . . . . . . . . . . . . . . . . . Polynemus aquilonaris(Thailand, Cambodia, Viet Nam and Lao People’s Democratic Republic)
6b. Pored lateral-line scales 69 to 79 (mode 78); scale rows below lateral line 13 (rarely 12); gillrakers 29 to
33 (mode 30) (Fig. 133, Plate VIb) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynemus dubius(Malaysia and Indonesia)
80-86 pored lateral-line scales
69-79 pored lateral-line scales

25-29
gillrakers

14-16 (rarely 13 or 17) scale rows below lateral line

29-33
gillrakers

Fig. 132 Polynemus aquilonaris

13 (rarely 12) scale rows
below lateral line

Fig. 133 Polynemus dubius

7a. Snout length 7% of standard length; upper caudal-fin lobe length 34 to 35% (mean 35%) of standard
length; seventh pectoral filament longest (Fig. 134; Plate VIe) . . . . . . . . . . . Polynemus melanochir dulcis(Lake Tonle Sap, Cambodia)
7b. Snout length 5 to 6% (mean 6%) of standard length; upper caudal-fin lobe length 35 to 44% (mean 39%) of
standard length; sixth pectoral filament usually longest (Fig. 135; Plate VIf) . . Polynemus melanochir melanochir(lower Mekong River and related rivers, and Kalimantan)
snout length
7% ofstandard
lengthupper caudal-fin lobe length
35% of standard length

7th pectoral filament longest

Fig. 134 Polynemus melanochir dulcis

snout length 6%(mean) ofstandard length

upper caudal-fin lobe 39%(mean) of standard length

6th pectoral filament longest

Fig. 135 Polynemus melanochir melanochirThreadfins of the World

81

Polynemus aquilonaris Motomura, 2003

Fig. 136; Plate VIa

Polynemus aquilonaris Motomura, 2003a: 155, figs 1a-b, 3 [type locality: Chao Phraya River, Thailand; holotype (URM-P
13930, 136 mm standard length); 12 paratypes (ANSP 177982, 108 mm standard length; ANSP 177984, 2 specimens, 88 to
96 mm standard length; CAS 92821, 114 mm standard length; NRM 24267, 104 mm standard length; UMMZ 181145, 3
specimens, 132 to 139 mm standard length; UMMZ 195407, 105 mm standard length; UMMZ 224815, 64 mm standard
length; UMMZ 232331, 2 specimens, 138 to 145 mm standard length)].
Synonyms: None.
FAO Names: En - Northern paradise fish; Fr - Barbure paradis du nord; Sp - Barbudo paraíso norteño.

Fig. 136 Polynemus aquilonaris
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 23 to 26% (mean 25%) of standard
length; head length 24 to 29% (mean 26%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 10 to 12% (mean 11%)of standard length, approximately equal to caudal-peduncle depth [10 to 12% (mean 11%) of standard length]; depth ofposterior margin of maxilla [2 to 3% (mean 3%) of standard length] slightly less than eye diameter [3 to 4% (mean 3%) of
standard length]; lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands onvomer, palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine
bases of similar thickness; second dorsal fin with I spine and 15 to 19 (mode 16) soft rays; anal fin with III spines and 11 to 13(mode 12) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 15 to 17 (mode 16) rays (all rays
unbranched), its length 34 to 41% (mean 37%) of standard length, posterior tip just short of or extending slightly beyond levelof anal-fin origin; pectoral filaments 7; first pectoral filament shortest, not reaching to level of posterior tip of pelvic fin; second
pectoral filament extending slightly beyond or not reaching to level of posterior tip of pelvic fin; third pectoral filament
extending beyond (rarely just short of) level of posterior tip of pelvic fin; fourth pectoral filament extending beyond level ofanal-fin origin or level of posterior base of anal fin; fifth pectoral filament extending beyond level of posterior tips of caudal-fin
lobes; sixth, usually longest, its length 260 to 371% (mean 308%) of standard length, and seventh pectoral filaments longer
than other filaments, extending well beyond posterior tips of caudal-fin lobes; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 39 to 44% (mean 43%) and lower lobe 33 to 41% (mean 38%) of
standard length. Pored lateral-line scales 80 to 86 (mode 81); lateral line simple, extending from upper end of gill opening to
mid-distal margin of caudal-fin membrane; scale rows above lateral line 6 to 8 (mode 7), below 13 to 17 (mode 14). Gillrakers
9 to 11 (mode 10) on upper limb, 16 to 18 (mode 17) on lower limb, 25 to 29 (mode 27) total. Vertebrae 10 precaudal and 15
caudal; supraneural bones 2. Swimbladder not apparent or present (varying in size less than about 23% of standard length).
Colour: Head and body greyish silver dorsally, silver ventrally; anterior margin of first dorsal fin and posterior margin of
second dorsal fin blackish, remaining parts translucent; pectoral fin translucent; base of pectoral filaments white, becoming
blackish on posterior tips; base and posterior margin of pelvic fin white, remaining parts translucent; posterior margin of anal
fin translucent, remaining parts white; posterior margin of caudal fin translucent, remaining parts white.

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FAO Species Catalogue for Fishery Purposes No. 3

Geographical Distribution: Currently known from the
Chao Phraya River system (Thailand), Mekong Riversystem below the Khone waterfalls of Lao People's
Democratic Republic (Cambodia, southernmost of Lao
People's Democratic Republic and southern Viet Nam),and Lake Tonle Sap and related rivers (Cambodia)(Fig. 137). Two specimens (NSMT-P 21772, 21776, 133
to 138 mm standard length), collected from the Chao
Phraya River at Nakhon Sawan, Thailand, represent thenorthernmost reliable record of the species.

Polynemus aquilonaris is currently known from
Indochina, whereas a related species, P. dubius, isdistributed in rivers on the Malay Peninsula, Sumatra
and Kalimantan. The distributions of both species aremost likely to be as relics, and are consistent with thelocation of Sundaland during the last Pleistocene glacial
period. According to Bornbusch and Lundberg (1989),
200 m
Sundaland was drained by several major river systems,
at least 2 of which may have participated in faunal
mixing. These included the South Indo-China River andNorth Sunda River which are now restricted to drainages
Fig. 137 Polynemus aquilonaris
on Indochina, and Malay Peninsula, Sumatra andKnown distribution
Kalimantan, respectively, following submergence of
Sundaland owing to increased sea levels. The present distributional ranges of P. aquilonaris and P. dubius indicate thatthey evidently originated in the South Indo-China and North Sunda rivers, respectively.
Habitat and Biology: Occurs on sandy or muddy bottoms in fresh-water rivers and estuaries. The species feeds oncrustaceans, small fishes and benthic organisms. Gonad examinations (based on 12 specimens) of the species by theauthor revealed that the species should be considered as probably having separate sexes (hermaphroditism not present).
Size: Maximum standard length at least 16 cm (Motomura, 2003a).
Interest to Fisheries: Esteemed as a food fish at least along the Chao Phraya and Mekong rivers, and Lake Tonle Sap. The
species has been often exported to Japan as an aquarium fish.
Local Names: CAMBODIA: Trey pream sor; LAO PEOPLE’S DEMOCRATIC REPUBLIC: Jin.
Literature: Motomura (2003a).
Remarks: Polynemus aquilonaris previously identified as P. dubius or P. longipectoralis (e.g. Rainboth, 1996; Kottelat,2001), has recently been described as a new species by Motomura (2003a).
Larger specimens (over about 110 mm standard length) of Polynemus aquilonaris collected from Lake Tonle Sap had afleshy lip on the upper jaw, smaller specimens (less than about 100 mm standard length) from the lake and all those fromother localities having a relatively thin lip. Apart from the lip condition in larger specimens, however, the 2 forms were difficult
to distinguish between because their meristic characters and proportional measurements fully overlapped. Therefore, thedifference in lip condition between the 2 forms is considered to represent geographical variation. More studies of the 2 formsof P. aquilonaris are needed to assess whether the 2 represent separate subspecies like P. melanochir dulcis and P. m.
melanochir or not.
Although P. aquilonaris is very similar to P. dubius, the former can be clearly distinguished from the latter by having higher
counts of pored lateral-line scales [80 to 86 (mode 81) versus 69 to 79 (mode 78) in P. dubius] and scale rows below thelateral line [14 to 17 (mode 14, rarely 13 or 17) versus 13 (rarely 12) in P. dubius], and lower counts of gillrakers [9 to 11(mode 11) in upper series, 16 to 18 (mode 17) in lower and 25 to 29 (mode 27) total versus 11 to 13 (mode 12), 18 to 21(mode 18) and 29 to 33 (mode 30), respectively]. Furthermore, P. aquilonaris differs from P. dubius in having a slightly
higher second dorsal-fin soft ray counts (15 to 19 versus 14 to 16 in P. dubius) and lower pectoral-fin ray counts (15 to 17
versus 16 to 18).

Polynemus dubius Bleeker, 1854

Fig. 138; Plate VIb

Polynemus dubius Bleeker, 1854: 92 [type locality: Banjarmasin, Sampit or Palembang, Indonesia; lectotype (RMNH 6014,
126 mm standard length) designated by Motomura, 2003a; 7 paralectotypes (RMNH 34452, 77 to 132 mm standard length)].
Synonyms: Polynemus longipectoralis Weber and de Beaufort, 1922: 213 [type locality: Banjarmasin, Kalimantan,
Indonesia; holotype (ZMA 112570, 133 mm standard length)].

Threadfins of the World83

FAO Names: En - Eastern paradise fish; Fr - Barbure paradis d'Orient; Sp - Barbudo paraíso oriental.

Fig. 138 Polynemus dubius
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 23 to 28% (mean 24%) of standard
length; head length 25 to 27% (mean 26%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 11 to 13% (mean 11%)of standard length, approximately equal to caudal-peduncle depth [11 to 13% (mean 11%) of standard length]; depth ofposterior margin of maxilla [3 to 4% (mean 3%) of standard length] slightly greater than eye diameter [2 to 3% (mean 3%) of
standard length]; lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands onvomer, palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine
bases of similar thickness; second dorsal fin with I spine and 14 to 16 (mode 16) soft rays; anal fin with III spines and 12 softrays, anal-fin base less than second dorsal-fin base; pectoral fin with 16 to 18 (mode 16) rays (all rays unbranched), its
length 30 to 40% (mean 37%) of standard length, posterior tip just short of or extending slightly beyond level of anal-fin origin;pectoral filaments 7; first pectoral filament shortest, not reaching to level of posterior tip of pelvic fin; second pectoral filament
extending slightly beyond or not reaching to level of posterior tip of pelvic fin; third pectoral filament extending beyond (rarely
just short of) level of posterior tip of pelvic fin; fourth pectoral filament extending beyond level of anal-fin origin or level ofposterior base of anal fin; fifth pectoral filament extending beyond level of posterior tips of caudal-fin lobes; sixth, usually
longest, its length 264 to 312% (mean 293%) of standard length, and seventh pectoral filaments longer than other filaments,
extending well beyond posterior tips of caudal-fin lobes; caudal fin deeply forked, upper and lower caudal-fin lobes notfilamentous, upper caudal-fin lobe 35 to 50% (mean 42%) and lower lobe 33 to 47% (mean 39%) of standard length. Poredlateral-line scales 69 to 79 (mode 78); lateral line simple, extending from upper end of gill opening to mid-distal margin of
caudal-fin membrane; scale rows above lateral line 6 to 8 (mode 7), below 12 or 13 (mode 13). Gillrakers 11 to 13 (mode 12)
on upper limb, 18 to 21 (mode 18) on lower limb, 29 to
33 (mode 30) total. Vertebrae 10 precaudal and 15
caudal; supraneural bones 2. Swimbladder notapparent. Colour: (preserved specimens) Head andbody grey dorsally, pale silver ventrally; anterior marginand posterior tip of first dorsal fin blackish, remaining
parts yellowish silver; posterior margins of second
dorsal, anal and caudal fins translucent, remaining
parts pale yellow; pectoral fin and filaments translucent
or white without melanophores; pelvic fin pale yellow.
Geographical Distribution: Currently known from theKangsar and Muar rivers (western Malaysia in Malay
Peninsula), Musi and Batanghari rivers (southeastern
Sumatra, Indonesia), and Sampit and Barito rivers(southern Kalimantan, Indonesia) (Fig. 139).
Habitat and Biology: Occurs on sandy or muddy
bottoms in fresh-water rivers and estuaries. Feeds oncrustaceans, small fishes and benthic organisms.
Size: Maximum standard length at least 18 cm(Motomura, 2003a).
Interest to Fisheries: Unknown.

200 m

Fig. 139 Polynemus dubius
Known distribution

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FAO Species Catalogue for Fishery Purposes No. 3

Local Names: None known.
Literature: Motomura (2003a).
Remarks: P. dubius was described by Bleeker (1854) on the basis of Bleeker’s (1851a, 1852) descriptions of a species
originally believed by him to represent P. longifilis (true P. longifilis Cuvier in Cuvier and Valenciennes, 1829b has recently
been regarded as a junior synonym of P. paradiseus; see Motomura et al., 2002b). Although P. dubius has been regarded asa valid species (e.g. Kottelat et al., 1993; Randall and Lim, 2000), P. longipectoralis has also been regarded as a valid
species by many researchers (e.g. Chevey, 1932; Kottelat, 1989, 2001; Talwar and Jhingran, 1991; Mishra and Krishnan,1993), but without comparisons of each nominal species. Weber and de Beaufort (1922), Myers (1936) and Rainboth (1996)
regarded both species as valid, the first-mentioned believing P. dubius and their new species, P. longipectoralis, to be
distinguished by the number of lateral-line scales [stated as 67 in P. dubius (based on a single specimen) versus 84 in
P. longipectoralis (based on the holotype), but in fact 79, according to examination of that specimen by the author]. The
diagnostic characters (seven pectoral filaments, 8 spines in the first dorsal fin, 79 pored lateral-line scales, 13 scale rows
below lateral line, vomer with villiform teeth and posterior portion of the maxilla less than orbit diameter) found in the holotypeof P. longipectoralis are consistent with those of specimens of P. dubius. Therefore, P. longipectoralis is regarded as a
junior synonym.
Rainboth (1996) also distinguished between the 2 nominal species by the number of lateral-line scales (65-67 in P. dubius
versus 80-85 in P. longipectoralis). However, his P. longipectoralis is in fact P. aquilonaris (see account of P. aquilonaris).
Myers (1936) distinguished the 2 nominal species by the number of first dorsal-fin spines (7 in P. dubius versus 8 in
P. longipectoralis). However, P. dubius, including the holotype of P. longipectoralis, in fact has 8 spines in the first dorsal
fin. Polynemus hornadayi and P. paradiseus are the only polynemid species with 7 spines in the first dorsal fin.

Polynemus dubius is easily distinguished from P. kapuasensis and P. multifilis by having 7 pectoral filaments (13 to 16 in
the latter). Polynemus melanochir dulcis and P. m. melanochir lack villiform teeth on the vomer and have black
pigmentation on (usually) more than half of the posterior margin of the pectoral fins, whereas P. dubius has villiform
vomerine teeth and lacks black pigmentation on the pectoral fins. Furthermore, P. dubius differs from both of the above
subspecies in having longer pectoral-fin rays [34 to 40% (mean 37%) of standard length in P. dubius versus 31 to 35% (mean33%) in P. m. dulcis and 30 to 35% (mean 32%) in P. m. melanochir] and pectoral filaments [longest filament 264 to 312%
(mean 293%) of standard length in P. dubius versus 128 to 153% (mean 141%) and 141 to 193% (mean 159%),
respectively]. Comparisons of P. dubius with P. aquilonaris are given in the account of the latter.

Polynemus hornadayi Myers, 1936

Fig. 140; Plate VIc

Polynemus hornadayi Myers, 1936: 376, fig. 1 [type locality: Ensengi River (emptying into the Sadong River), Sarawak,
Kalimantan, Malaysia; holotype (USNM 100632, 193 mm standard length); 9 paratypes presently known (BMNH
1935.8.29.31, 142 mm standard length; USNM 35719, 8 specimens, 109 to 146 mm standard length)].
Synonyms: None.
FAO Names: En - Hornaday’s paradise fish; Fr - Barbure paradis de Hornaday; Sp - Barbudo paraíso de Hornaday.

Fig. 140 Polynemus hornadayi

Threadfins of the World

85

Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 24 to 30% (mean 26%) of standard
length; head length 26 to 29% (mean 28%) of standard length. Snout pointed; occipital profile strongly concave. Posterior
margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 13 to 16% (mean 14%)of standard length, greater than caudal-peduncle depth [10 to 11% (mean 11%) of standard length]; depth of posterior
margin of maxilla [4 to 5% (mean 5%) of standard length] greater than eye diameter [1 to 2% (mean 1%) of standard length];
lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatinesand ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VII spines, all spine bases of similar
thickness; second dorsal fin with I spine and 14 to 16 (mode 15) soft rays; anal fin with II or III (mode III; first of 3 spines
vestigial, confirmed by radiograph) spines and 11 or 12 (mode 11) soft rays, anal-fin base less than second dorsal-fin base;pectoral fin with 16 to 19 (mode 18) rays (all rays unbranched), its length 39 to 47% (mean 42%) of standard length, posterior
tip reaching to level of midpoint of anal-fin base; pectoral filaments 7; first pectoral filament shortest, not reaching to level ofposterior tip of pelvic fin; second pectoral filament not reaching to level of anal-fin origin; third pectoral filament reaching tonear level of posterior end of anal-fin base or reaching between levels of origin and posterior end of anal-fin base; fourth
pectoral filament slightly longer or slightly shorter than standard length; fifth to seventh pectoral filaments longer than total
length; fifth pectoral filament longest, its length 323 to 339% (mean 331%) of standard length; caudal fin deeply forked,
upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 41 to 46% (mean 44%) and lower lobe 37 to 46%(mean 41%) of standard length. Pored lateral-line scales 90 to 103 (mode 94); lateral line simple, extending from upper endof gill opening to mid-distal margin of caudal-fin membrane; scale rows above lateral line 8 to 12 (mode 11), below 16 to 21(mode 18). Gillrakers 10 or 11 (mode 10) on upper limb, 16 or 17 on lower limb, 26 to 28 (mode 26) total. Vertebrae 10
precaudal and 15 caudal; supraneural bones 2. Swimbladder present, well developed. Colour: (preserved specimens)
Head and body greyish black dorsally, pale yellowish silver ventrally; membranes of first dorsal fin translucent, spines paleyellow; bases of second dorsal, pectoral, pelvic, anal and caudal fins pale yellow, posterior margins of these fins translucent;pectoral filaments uniformly whitish yellow.
Geographical Distribution: Currently known only from
3 rivers, Ensengi, Rajang and Sungai rivers, in western
Sarawak, Kalimantan, Malaysia (Fig. 141).
Habitat and Biology: Occurs in muddy, fast flowing
rivers, 12 m wide and 3 m deep (Feltes in Carpenter andNiem, 2001). Inhabits only fresh-water regions, having
never been recorded from the estuarine areas behind theriver mouths.
Size: Maximum standard length at least 20 cm(Motomura et al., 2002b).
Interest to Fisheries: Esteemed as a food fish inwestern Sarawak, Kalimantan, Malaysia.
Local Names: None known.
Literature: Motomura et al. (2002b).

200 m

Remarks: P. hornadayi and P. paradiseus, both have
VII first dorsal-fin spines, whereas all other polynemid
Fig. 141 Polynemus hornadayi
species have VIII spines. However, radiographs of
P. hornadayi indicate that 4 of 26 specimens examined
Known distribution
by the author have a grain of oval bone (diameter lessthan about 1 mm) buried under the subcutaneous tissue in front of the first dorsal-fin spine and associated with the firstpterygiophore (see Motomura et al., 2002b: fig. 3B). While the anteriormost dorsal-fin spine of other polynemid species isalso small, the spine is conventionally “spine-shaped” (not oval) with a dorsally exposed tip (see Motomura, Iwatsuki andKimura, 2001b: fig. 4, Motomura et al., 2002b: fig. 3C). The oval bone associated with the first pterygiophore in some
P. hornadayi is considered to represent the final stages of degeneration from an original spine precursor.
At a glance, P. hornadayi and P. paradiseus can be easily distinguished from each other by the shape of the occipital profile,that of P. hornadayi protruding strongly whereas that of P. paradiseus is nearly straight throughout life. In meristic
characters, P. hornadayi differs from P. paradiseus in having lower counts of anal-fin soft rays [11 or 12 (mode 11) versus 12(rarely 11 or 13) in the latter] and gillrakers [26 to 28 (mode 26) versus 30 to 34 (mode 32 or 33) in P. paradiseus], and higher
counts of pectoral-fin rays [16 to 19 (mode 18) versus 15 to 18 (mode 17) in P. paradiseus], scales above and below thelateral line [8 to 12 (mode 11) and 16 to 21 (mode 18) versus 6 or 7 (mode 7) and 10 to 12 (mode 11), respectively in
P. paradiseus] and pored lateral-line scales [90 to 103 (mode 94) versus 66 to 71 (mode 70) in P. paradiseus]. While
P. hornadayi is similar to P. paradiseus in having 7 pectoral filaments, in the former the fifth pectoral filament is the longest[323 to 339% (mean 331%) of standard length] and the fourth filament extends well beyond the posterior central margin of
the caudal fin, whereas in the latter the sixth filament is the longest [181 to 248% (mean 208%) of standard length] and thefourth filament fails to reach the posterior central margin of the caudal fin. The pectoral-fin ray of P. hornadayi is also
relatively longer than that of P. paradiseus [posterior tip of pectoral fin reaching to midpoint of anal-fin base, 39 to 47% (mean42%) of standard length versus not reaching, 30 to 35% (mean 33%) of standard length, respectively]. Furthermore, theposterior margin of the premaxilla of P. hornadayi is deeper than that of P. paradiseus [4 to 5% (mean 5%) of standard length

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FAO Species Catalogue for Fishery Purposes No. 3

versus 3 to 4% (mean 4%) of standard length, respectively], P. hornadayi also tending to have a slightly longer head [26 to
29% (mean 28%) of standard length], postorbital [18 to 21% (mean 20%) of standard length], and pectoral-fin base [14 to
15% (mean 14%) of standard length], and greater pelvic-fin ray lengths [19 to 22% (mean 20%) of standard length] thanP. paradiseus [24 to 27% (mean 27%), 17 to 19% (mean 18%), 13 to 14% (mean 13%) and 15 to 18% (mean 16%) of
standard length, respectively], although the proportional length measurements overlapped between the 2 species (seeMotomura et al., 2002b: fig. 8). Internally, P. hornadayi differs from P. paradiseus in having a well-developed swimbladder(absent in the latter). Furthermore, the former frequently has a vestigial anal fin spine buried under subcutaneous tissue(62%, 16 of 26 specimens), such being absent in P. paradiseus (see Motomura et al., 2002b: table 3).

Polynemus kapuasensis Motomura and van Oijen, 2003

Fig. 142; Plate VId

Polynemus kapuasensis Motomura and van Oijen, 2003: 394, figs 1, 3a, 6 [type locality: Kapuas River Basin, fish market at
Sintang, Kalimantan, Indonesia; holotype (CAS 47198, 151 mm standard length); 11 paratypes (CAS 217348, 3 specimens,
133 to 172 mm standard length; CAS 49454, 5 specimens, 33 to 76 mm standard length; CAS 217161, 3 specimens, 83 to
108 mm standard length)].
Synonyms: None.
FAO Names: En - Kapuas elegant paradise fish; Fr - Barbure paradis de Kapuas; Sp - Barbudo paraíso de Kapuas.

Fig. 142 Polynemus kapuasensis
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 23 to 25% (mean 24%) of standard
length; head length 22 to 23% (mean 23%) of standard length in adults (more than about 130 mm standard length), 24 to
29% (mean 27%) of standard length in juveniles (less than about 110 mm standard length). Snout pointed; occipital profile
nearly straight or slightly protruding. Posterior margin of maxilla extending well beyond level of posterior margin of adipose
eyelid; upper-jaw length 10 to 11% (mean 10%) of standard length, approximately equal to caudal-peduncle depth [10 to
11% (mean 11%) of standard length]; depth of posterior margin of maxilla (2% of standard length) slightly less than eye
diameter [3 to 4% (mean 3%) of standard length]; lip on lower jaw well developed, dentary teeth restricted to dorsal surface;teeth villiform in broad bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal
fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 15 or 16 (mode 15) soft rays; anal
fin with III spines and 11 or 12 (mode 12) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 15 to 17(mode 15) rays (all rays unbranched), its length 29 to 33% (mean 31%) of standard length, posterior tip not extending
beyond level of anal-fin origin in adults (more than about 70 mm standard length) or extending beyond level of anal-fin origin
in juveniles (less than about 50 mm standard length); pectoral filaments usually 15 on each side of body, rarely 16 on eachside or asymmetrically 15 and 16; first pectoral filament shortest, extending beyond level of pelvic-fin origin; second toseventh pectoral filaments not reaching to level of posterior tip of pelvic fin; eighth and ninth pectoral filaments extending
beyond level of posterior tip of pelvic fin, but not reaching to level of anal-fin origin; tenth pectoral filament extending slightly
beyond level of anal-fin origin; eleventh pectoral filament extending slightly beyond level of posterior end of anal-fin base;twelfth pectoral filament extending beyond level of posterior margin of central caudal fin, but not reaching to level of posteriorThreadfins of the World87

tips of caudal-fin lobes; thirteenth to fifteenth (or sixteenth if present) pectoral filaments longer than total length; thirteenth
pectoral filament probably longest, 332 to 397% (mean 365%) of standard length; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 41 to 51% (mean 46%) and lower lobe 35 to 50% (mean 40%) of
standard length. Pored lateral-line scales 100 to 110 (mode 103); lateral line simple, extending from upper end of gill
opening to mid-distal margin of caudal-fin membrane; scale rows above lateral line 9 to 11 (mode 10), below 17 to 20 (mode18); caudal-peduncle scales 35 to 40 (mode 38). Gillrakers 9 to 11 (mode 11) on upper limb, 16 to 18 (mode 16 and 18) onlower limb, 25 to 29 (mode 28) total. Vertebrae 10 precaudal and 15 caudal; supraneural bones 2. Swimbladder present.
Colour: Head and body bluish gray dorsally, silver ventrally; bases of first and second dorsal, anal and caudal fins reddish
gray; first and second dorsal and caudal fins grayish white; posterior part of anal fin bluish white, remaining parts of anal andpelvic fins and pectoral filaments white; pectoral fin translucent.
Geographical Distribution: Currently known only
from the Kapuas River system, western Kalimantan,
Indonesia (Fig. 143).
Habitat and Biology: No data available.
Size: Maximum standard length at least 17 cm(Motomura and van Oijen, 2003).
Interest to Fisheries: Esteemed as an importantfood fish along the Kapuas River.
Local Names: None known.
Literature: Motomura and van Oijen (2003).
Remarks: Polynemus kapuasensis , previously
identified as P. multifilis (e.g. Myers, 1936; Roberts,1989), was recently described as a new species on
the basis of 12 specimens from the Kapuas River(Motomura and van Oijen, 2003).

200 m

Fig. 143 Polynemus kapuasensis
Polynemus kapuasensis and P. multifilis are easily
distinguished from all other congeners by having
Known distribution
higher counts of pectoral filaments (13 to 16 versus 7
in the latter). Polynemus kapuasensis differs from P. multifilis in having higher counts of pectoral filaments (usually 15 oneach side of body, rarely asymmetrically 15 and 16, or 16 on each side versus usually 14 on each side, rarely 13 on eachside, asymmetrically 13 and 14, or 14 and 15 in the latter), pored lateral-line scales [100 to 110 (mode 103) versus 83 to 99(mode 86)], scale rows above and below lateral line [9 to 11 (mode 10) and 17 to 20 (mode 18), respectively versus 7 or 8
(mode 8) and 14 to 18 (mode 15), respectively] and caudal-peduncle scales [35 to 40 (mode 38) versus 28 to 37 (mode 32)]
(see Motomura and van Oijen, 2003: tables 1-4). Furthermore, P. kapuasensis tends to have a slightly shorter head andanal-fin base lengths [22 to 23% (mean 23%) of standard length in adults more than about 130 mm standard length and 12 to
14% (mean 14%) of standard length, respectively] than P. multifilis [23 to 29% (mean 25%) of standard length and 15 to
17% (mean 15% of standard length, respectively], although the proportional measurements for the lengths overlapped
between the two species, especially in young stages (see Motomura and van Oijen, 2003: fig. 6a-b). Moreover, the former
has a short pectoral fin [29 to 33% (mean 31%) of standard length], its posterior tip not extending beyond the level of theanal-fin origin in adults (more than about 70 mm standard length), whereas the latter has a long pectoral fin [33 to 38%(mean 36%) of standard length], its posterior tip reaching or extending beyond the level of the anal-fin origin throughout life
(see Motomura and van Oijen, 2003: fig. 6c).

Polynemus melanochir dulcis Motomura and Sabaj, 2002

Fig. 144; Plate VIe

Polynemus melanochir dulcis Motomura and Sabaj, 2002: 182, fig. 1 [type locality: Lake Tonle Sap, Cambodia; holotype(ANSP 178011, 135 mm standard length); 2 paratypes (AMS I. 40968-001, 126 mm standard length; BSKU 14850, 128 mmstandard length)].
Synonyms: None.

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FAO Names: En - Lake blackhand paradise fish; Fr - Barbure paradis à doigts noirs de lac; Sp - Barbudo paraíso de manonegra lacustre.

Fig. 144 Polynemus melanochir dulcis
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 23% of standard length; head length 27
to 28% (mean 28%) of standard length. Snout strongly pointed; occipital profile nearly straight. Posterior margin of maxilla
extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 14 to 15% (mean 14%) of standard
length, greater than caudal-peduncle depth [10 to 11% (mean 10%) of standard length]; depth of posterior margin of maxilla[3 to 4% (mean 4%) of standard length] greater than eye diameter (3% of standard length); lip on lower jaw well developed,
dentary teeth restricted to dorsal surface; teeth villiform in broad bands on palatines and ectopterygoids; vomerine toothplate covered with skin and teeth absent. Posterior margin of preopercle serrated. First dorsal fin with VIII spines, all spine
bases of similar thickness; second dorsal fin with I spine and 16 soft rays; anal fin with III spines and 11 or 12 (mode 12) softrays, anal-fin base less than second dorsal-fin base; pectoral fin with 16 or 17 (mode 16) rays (all rays unbranched), its
length 34 to 35% (mean 33%) of standard length, posterior tip not reaching to or just reaching to level of anal-fin origin;pectoral filaments 7; first (shortest) and second pectoral filaments extending beyond level of pelvic-fin origin but not reaching
to level of posterior tip of pelvic fin; third pectoral filament reaching near anal-fin origin; fourth pectoral filament extending
slightly beyond level of middle of anal-fin base or posterior base of anal fin; fifth pectoral filament extending slightly beyond
caudal-fin base or posterior mid-distal margin of caudal fin; sixth and seventh [longest, its length 128 to 153% (mean 141%)of standard length] pectoral filaments extending well beyond posterior tips of caudal fin; caudal fin deeply forked, upper andlower caudal-fin lobes not filamentous, upper caudal-fin lobe 34 to 35% (mean 35%) and lower lobe 34 to 38% (mean 36%)of standard length. Pored lateral-line scales 71 or 72 (mode 71); lateral line simple, extending from upper end of gill opening
to mid-distal margin of caudal-fin membrane; scale rows above lateral line 7, below 12 or 13 (mode 13). Gillrakers 12 on
upper limb, 17 to 19 (mode 17) on lower limb, 29 to 31 (mode 29) total. Vertebrae 10 precaudal and 15 caudal; supraneuralbones 2. Swimbladder absent. Colour: (preserved specimens) Head and body greyish black dorsally, pale whitish yellow
ventrally; posterior tip of first dorsal fin intense black, other parts whitish yellow; posterior margins of second dorsal, pelvic,
anal and caudal fins translucent (posterior margin of second dorsal fin blackish in a single specimen, BSKU 14850), anterior
margin of second dorsal fin blackish, other parts whitish
yellow; pectoral fin, except posterior tip and base,intense black (translucent in a single specimen, AMS I.
40968-001); base of pectoral filaments whitish yellow,
becoming blackish posteriorly.
Geographical Distribution: Currently known only
from Lake Tonle Sap, upper end of the lower MekongRiver floodplain, Cambodia (Fig. 145).
Habitat and Biology: No data available.
Size: Maximum standard length at least 135 mm(Motomura and Sabaj, 2002).
Interest to Fisheries: Unknown.
Local Names: CAMBODIA: Trey pream.
Literature: Motomura and Sabaj (2002).
Remarks: P. melanochir dulcis and P. m. melanochir
are uniquely characterized by the absence of vomerine
teeth (present in all congeners). Likewise, bothsubspecies have a very black pectoral fin. Although thepectoral fin of P. paradiseus is occasionally tinged with

200 m

Fig. 145 Polynemus melanochir dulcis
Known distribution

Threadfins of the World89

black, that species has villiform vomerine teeth. The vomer and pectoral fin characters of P. m. dulcis and P. m. melanochir
are very similar. Furthermore, the meristic characters of both subspecies completely overlap (see Motomura and Sabaj,
2002: table 3). However, P. m. dulcis differs from the latter in having a greater snout length [7% of standard length versus 5
to 6% (mean 6%) of standard length in the latter; see Motomura and Sabaj, 2002: tables 1-2, figs. 4-5] and shorter upper
caudal-fin lobe length [34 to 35% (mean 35%) of standard length versus 34 to 44% (mean 39%) of standard length; seeMotomura and Sabaj, 2002: tables 1-2, fig. 5].
In addition, the head length [27 to 28% (mean 28%) of standard length], eye diameter (3%), orbit diameter [3 to 4% (mean3%)], upper-jaw length [14 to 15% (mean 14%)], second dorsal-fin base length [20 to 21% (mean 21%)] and longest
pelvic-fin ray length [17 to 18% (mean 18%)] in P. m. dulcis are greater than those in P. m. melanochir [24 to 27% (mean26%), 1 to 2% (mean 2%), 2 to 3% (mean 2%), 12 to 14% (mean 13%), 18 to 21% (mean 20%) and 14 to 17% (mean 16%),
respectively; see Motomura and Sabaj, 2002: tables 1-2].

Polynemus m. dulcis appears to be one of the rarest polynemids and only 3 type specimens are known from museum
collections.

Polynemus melanochir melanochir Valenciennes, 1831

Fig. 146; Plates VIf

Polynemus melanochir Valenciennes in Cuvier and Valenciennes, 1831: 513 [mistakenly reported from India (seeMotomura and Sabaj, 2002), based solely on a drawing sent from Sumatra, Indonesia (see Feltes, 1991: fig. 1); type locality:
Kuching Bay, Sarawak, Kalimantan, Malaysia, based on a neotype (ZRC 37829, 179 mm standard length) designated byMotomura and Sabaj, 2002].
Synonyms: Polynemus macronema Bleeker, 1852 (not of Pel): 419 (primary homonym of Polynemus macronemus Pel).
Polynemus borneënsis Bleeker, 1857b: 3 (replacement name for P. macronema Bleeker, but incorrect original spelling).
Polynemus borneensis Bleeker, 1858a: 2 [type locality: Banjarmasin, Pontianak and Sampit, Kalimantan, Indonesia;
justified emendation of P. borneënsis Bleeker; 3 syntypes (RMNH 6013, 148 to 190 mm standard length) determined from 5
Bleeker specimens by Motomura and Sabaj, 2002]. Galeoides microps Steindachner, 1869a: 126 [type locality: China (butprobably erroneous, see Motomura and Sabaj, 2002); holotype (NMW 77569, 171 mm standard length)]. Polynemus
melanopus Sauvage, 1881: 101 [type locality: Saigon (= Ho-chi-minh), Viet Nam; 4 syntypes (MNHN A. 3048, 2 specimens,
139 to 160 mm standard length; MNHN A. 3049, 2 specimens, 130 to 160 mm standard length)]. Trichidion hilleri Fowler,
1905: 502, fig. 11 [type locality: Baram River, Sarawak, Kalimantan, Malaysia; holotype (ANSP 114895, 166 mm standard
length)].
FAO Names: En - Blackhand paradise fish; Fr - Barbure paradis à doigts noirs; Sp - Barbudo paraíso de mano negra.

Fig. 146 Polynemus melanochir melanochir
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 22 to 25% (mean 23%) of standard
length; head length 25 to 27% (mean 26%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 12 to 14% (mean 13%)of standard length, greater than caudal-peduncle depth [10 to 13% (mean 11%) of standard length]; depth of posterior
margin of maxilla [3 to 5% (mean 4%) of standard length] greater than eye diameter [1 to 2% (mean 2%) of standard length];
lip on lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on palatines andectopterygoids; vomerine tooth plate covered with skin and teeth absent. Posterior margin of preopercle serrated. First
dorsal fin with VIII spines, all spine bases of similar thickness; second dorsal fin with I spine and 15 or 16 (mode 16; rarely

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18, 1 of 40 specimens) soft rays; anal fin with III spines and 11 or 12 (mode 12; rarely 13, 1 of 40 specimens) soft rays,
anal-fin base less than second dorsal-fin base; pectoral fin with 15 to 18 (mode 17) rays (all rays unbranched), its length 30
to 35% (mean 32%) of standard length, posterior tip not reaching to, just reaching to or extending slightly beyond level ofanal-fin origin; pectoral filaments 7; first pectoral filament shortest, extending beyond level of pelvic-fin origin but notreaching to level of posterior tip of pelvic fin; second pectoral filament extending slightly beyond level of posterior tip of pelvic
fin; third pectoral filament reaching near anal-fin origin or extending slightly beyond level of posterior tip of pelvic fin; fourth
pectoral filament reaching to or extending slightly beyond level of posterior base of anal fin; fifth pectoral filament extending
well beyond caudal-fin base, but not reaching posterior tips of caudal fin, or extending slightly beyond caudal-fin base, notreaching to mid-distal margin of caudal-fin; sixth and seventh [longest, its length 134 to 193% (mean 159%) of standard
length] pectoral filaments extending well beyond posterior tips of caudal fin; caudal fin deeply forked, upper and lower
caudal-fin lobes not filamentous, upper caudal-fin lobe 35 to 44% (mean 39%) and lower lobe 29 to 42% (mean 36%) of
standard length. Pored lateral-line scales 68 to 74 (mode 70); lateral line simple, extending from upper end of gill opening to
mid-distal margin of caudal-fin membrane; scale rows above lateral line 6 to 8 (mode 7), below 11 to 14 (mode 12). Gillrakers
11 to 14 (mode 13) on upper limb, 16 to 20 (mode 18) on lower limb, 27 to 33 (mode 31) total. Vertebrae 10 precaudal and 15
caudal; supraneural bones 2. Swimbladder absent. Colour: Head and body greyish black dorsally, yellow ventrally;
posterior tip of first dorsal fin intense black, other parts yellow; posterior margin and tip of second dorsal fin black, other parts
yellow; pelvic fin uniformly yellow; posterior margins of anal and caudal fin translucent, other parts yellow; pectoral fin intense
black, except for vivid yellow base; base of pectoral filaments vivid yellow, becoming blackish posteriorly.
Geographical Distribution: Currently known from thelower Mekong River and related rivers (Cambodia andsouthern Viet Nam), and Kalimantan (Malaysia andIndonesia) (Fig. 147).
Habitat and Biology: No data available.
Size: Maximum standard length at least 20 cm(Motomura and Sabaj, 2002).
Interest to Fisheries: Esteemed as an important foodfish at least along the lower Mekong River and related
rivers.
Local Names: CAMBODIA: Trey pream loeung;
INDONESIA: Kurau.
Literature: Motomura and Sabaj (2002).
200 m
Remarks: The name, Polynemus melanochir, hasbeen used by many researchers [e.g., Bleeker, 1849;
Myers, 1936 (as Filimanus melanochir ) ;Fig. 147 Polynemus melanochir melanochir
Gloerfelt-Tarp and Kailola, 1984 (as F. melanochir);Bleeker, 1983 (as Trichidion melanochir)]. However,Known distribution
Feltes (1991) recognized that the polynemid species
commonly identified as P. melanochir (F. melanochir or T. melanochir) was not true P. melanochir, but in fact represented
a separate undescribed species (described as F. perplexa Feltes, 1991).

Although the original description of P. melanochir was brief, the drawing clearly shows the very black pectoral-fin rays (seeFeltes, 1991: fig. 1), such being a characteristic of the specimens considered here as P. melanochir melanochir.

Polynemus borneensis has been regarded as a valid species by many researchers (e.g. Weber and de Beaufort, 1922;
Kottelat et al., 1993; Rainboth, 1996). However, recent examination of the 3 syntypes of P. borneensis, the neotype of
P. m. melanochir, and other specimens showed that they represented a single species (see Motomura and Sabaj, 2002).
Therefore, P. borneensis is regarded as a junior synonym of the latter.
Galeoides microps was described by Steindachner (1869a) on the basis of a single specimen and later described in moredetail (Steindachner, 1869b). Although there is no indication in the literature that the holotype of G. microps has been
re-examined since Steindachner’s (1869a-b) descriptions (Springer, 1982), the species has been regarded as valid,
belonging to the genus Galeoides (e.g. Fowler, 1935; Myers, 1936; Springer, 1982; Hureau in Whitehead et al., 1986).
Recently, Motomura, Mikschi and Iwatsuki (2001) and Motomura and Sabaj (2002) examined the holotype of G. microps andregarded it as a junior synonym of P. m. melanochir.
Polynemus melanopus was described by Sauvage (1881) on the basis of 4 specimens from Ho-chi-minh, Viet Nam. The
characters, black pectoral fin, vomer without teeth and short snout (5 to 6% of standard length), of the 4 syntypes were foundto be consistent with those of the specimens considered here as P. m. melanochir (see Motomura and Sabaj, 2002).
Therefore, P. melanopus is also regarded as a junior synonym of the latter.
Trichidion hilleri was described by Fowler (1905) on the basis of a single specimen from Baram River, Sarawak,
Kalimantan, Malaysia. Although the species has been regarded as valid (as Polynemus hilleri) by many researchersThreadfins of the World91(e.g. Kottelat et al., 1993; Kottelat and Lim, 1995), examination of the holotype of T. hilleri and the specimens consideredhere as P. m. melanochir showed that they represented a single species (see Motomura and Sabaj, 2002). Therefore,
T. hilleri is also regarded as a junior synonym of the latter.

Polynemus m. melanochir and P. m. dulcis are uniquely characterized by the absence of vomerine teeth (present in allcongeners). Comparisons of P. m. melanochir with P. m. dulcis are given in the account of the latter.

Polynemus multifilis Temminck and Schlegel, 1843

Fig. 148; Plate VIg

Polynemus multifilis Temminck and Schlegel, 1843: 29 [type locality: near Banjarmasin, south Kalimantan; holotype(RMNH 436, 136 mm standard length)].
Synonyms: Polynemus quatordecimfilis Pel, 1851: 10 [type locality: near Banjarmasin, south Kalimantan; objectivesynonym of P. multifilis Temminck and Schlegel; holotype (RMNH 436, 136 mm standard length)].
FAO Names: En - Elegant paradise fish; Fr - Barbure paradis élégante; Sp - Barbudo paraíso elegante.

Fig. 148 Polynemus multifilis
Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 23 to 27% (mean 24%) of standard
length; head length 23 to 29% (mean 25%) of standard length. Snout pointed; occipital profile nearly straight or slightly
protruding. Posterior margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 9
to 11% (mean 10%) of standard length, approximately equal to caudal-peduncle depth [11 to 12% (mean 11%) of standard
length]; depth of posterior margin of maxilla [2 to 3% (mean 2%) of standard length] slightly less than eye diameter [3 to 4%(mean 3%) of standard length]; lip on lower jaw well-developed, dentary teeth restricted to dorsal surface; teeth villiform inbroad bands on vomer, palatines and ectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VIII
spines, all spine bases of similar thickness; second dorsal fin with I spine and 14 to 16 (mode 15) soft rays; anal fin with III
spines and 11 to 13 (mode 12) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 14 to 16 (mode 15)rays (all rays unbranched), its length 33 to 38% (mean 36%) of standard length, posterior tip reaching or extending beyond
level of anal-fin origin throughout life; pectoral filaments usually 14 on each side of body, rarely 13 on each side orasymmetrically 13 and 14, or 14 and 15; first pectoral filament shortest, extending beyond level of pelvic-fin origin; second tosixth pectoral filaments not reaching to level of posterior tip of pelvic fin; seventh and eighth pectoral filaments not reachingor extending beyond level of posterior tip of pelvic fin; ninth pectoral filament extending well beyond level of posterior tip ofpelvic fin, sometimes extending beyond level of anal-fin origin; tenth pectoral filament extending beyond level of anal-fin
origin, rarely extending beyond level of posterior end of anal-fin base; eleventh pectoral filament extending beyond level ofposterior end of hypural plate, but not reaching to level of posterior tips of caudal-fin lobes; twelfth to fourteenth (or fifteenth if
present) pectoral filaments longer than total length; twelfth pectoral filament probably longest, 182 to 376% (mean 295%) of
standard length; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 38 to 46%(mean 43%) and lower lobe 32 to 41% (mean 38%) of standard length. Pored lateral-line scales 83 to 99 (mode 86); lateralline simple, extending from upper end of gill opening to mid-distal margin of caudal-fin membrane; scale rows above lateralline 7 or 8 (mode 8), below 14 to 18 (mode 15); caudal-peduncle scales 28 to 37 (mode 32). Gillrakers 9 to 11 (mode 10) on

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upper limb, 15 to 17 (mode 16) on lower limb, 24 to 27 (mode 26) total. Vertebrae 10 precaudal and 15 caudal; supraneuralbones 2. Swimbladder present. Colour: Head and body dark purplish blue dorsally, silver ventrally; bases of first and second
dorsal, anal and caudal fins grayish white, anterior margin of pelvic fin white, remaining parts of those fins semitransparent;
pectoral fin semitransparent; pectoral filaments white.
Geographical Distribution: Currently known from the
Chao Phraya River system (Thailand), Musi and
Batanghari rivers (southeastern Sumatra, Indonesia),
and Sampit and Barito rivers (southern Kalimantan,
Indonesia). The species is not known from the Kapuas
River (where it is replaced by the endemic
P. kapuasensis) (Fig. 149).
Habitat and Biology: Occurs on sandy or muddy
bottoms in fresh-water rivers. Feeds on crustaceans,small fishes and benthic organisms.
Size: Maximum standard length at least 28 cm (Feltes
in Carpenter and Niem, 2001).
Interest to Fisheries: Esteemed as a food fish at least
along the Chao Phraya River. The species has been
often exported to Japan as an aquarium fish.
Local Names: None known.

200 m

Literature: Motomura and van Oijen (2003).

Fig. 149 Polynemus multifilis
Remarks: The name, Polynemus multifilis, first
Known distribution
appeared in a footnote of a description of specimens of
Polynemus plebejus Broussonet, 1782 (presently regarded as Polydactylus plebeius) by Temminck and Schlegel (1843).
Many researchers (e.g. Myers, 1936; Eschmeyer, 1998; Feltes in Carpenter and Niem, 2001) indicated the authorship ofP. multifilis as Schlegel. However, the authorship of the species should be treated as Temminck and Schlegel (seeMotomura and van Oijen, 2003). Polynemus multifilis was originally described on the basis of a single specimen (RMNH
436, 136 mm standard length), and later described in more detail by Schlegel (1852). Pel (1851) described
P. quatordecimfilis as a new species, but P. quatordecimfilis is a objective synonym of P. multifilis (see Motomura and van
Oijen, 2003). Type material of P. multifilis and P. quatordecimfilis were determined by Motomura and van Oijen (2003).
Bleeker (1852) described Polynemus polydactylus as a new species on the basis of a single specimen (RMNH 6001,
133 mm standard length) from Banjarmasin, southern Kalimantan, Indonesia. Because that name was preoccupied by
P. polydactylus of Vahl (1798) which has presently been regarded as a junior synonym of Galeoides decadactylus (Bloch,1795), Bleeker’s P. polydactylus is permanently invalid, treated as a primary homonym of Vahl’s P. polydactylus.
Subsequently, Bleeker (1860) recognized that his P. polydactylus is the same species as P. multifilis (his opinion concurredwith here).

Polynemus multifilis is easily distinguished from other congeners, except for P. kapuasensis, by having 13 to 15 pectoral
filaments (7 in the latter). Comparisons of P. multifilis with P. kapuasensis are given in the account of the latter.

Polynemus paradiseus Linnaeus, 1758

Fig. 150; Pl. VIh

Polynemus paradiseus Linnaeus, 1758: 317 [original locality: Bengal, India, based on a figure (see Motomura et al., 2002d:
fig. 5) and description of Edwards (1743-1751); type locality: Gariahat, Calcutta, West Bengal, India, based on a neotype(NRM 47529, 198 mm standard length) designated by Motomura et al., 2002b].
Synonyms: Polynemus risua Hamilton, 1822: 228, 381 (type locality: vicinity of Lukhipur, India; no types known).
Polynemus toposui Hamilton, 1822: 232, 381 (type locality: estuary of Ganges River, West Bengal, India; no types known).
Polynemus aureus Hamilton, 1822: 232, 381 (type locality: Calcutta, West Bengal, India; no types known). Polynemus
longifilis Cuvier in Cuvier and Valenciennes, 1829: 365 [type locality: Pondicherry and Ganges River, India; Manila,
Philippines (probably erroneous, see Motomura et al., 2002b); 7 syntypes (MHNG 148.24, 134 mm standard length; MNHN
2200, 3 specimens, 141 to 157 mm standard length; MNHN A. 3045, 115 mm standard length; MNHN A. 4803, dried
specimen, 154 mm standard length; SMF 439, 124 mm standard length)].

Threadfins of the World

93

FAO Names: En - Paradise threadfin; Fr - Barbure paradis; Sp - Barbudo paraíso.

Fig. 150 Polynemus paradiseus

Diagnostic Features: A medium-sized species. Body depth at first dorsal-fin origin 20 to 28% (mean 24%) of standard
length; head length 24 to 27% (mean 26%) of standard length. Snout pointed; occipital profile nearly straight. Posterior
margin of maxilla extending well beyond level of posterior margin of adipose eyelid; upper-jaw length 13 to 15% (mean 14%)of standard length, greater than caudal-peduncle depth [9 to 12% (mean 10%) of standard length]; depth of posterior marginof maxilla [3 to 4% (mean 4%) of standard length] greater than eye diameter [1 to 2% (mean 2%) of standard length]; lip on
lower jaw well developed, dentary teeth restricted to dorsal surface; teeth villiform in broad bands on vomer, palatines andectopterygoids. Posterior margin of preopercle serrated. First dorsal fin with VII spines, all spine bases of similar thickness;second dorsal fin with I spine and 14 or 15 (mode 15) soft rays; anal fin with II spines and 12 (rarely 11 or 13, 2 of 40
specimens) soft rays, anal-fin base less than second dorsal-fin base; pectoral fin with 15 to 18 (mode 17) rays (all rays
unbranched), its length 30 to 35% (mean 33%) of standard length, posterior tip reaching to or just short of level of anal-fin
origin [but in juveniles (less than about 100 mm standard length) extending slightly beyond anal-fin origin]; pectoral filaments
7; first pectoral filament shortest, not reaching to level of posterior tip of pelvic fin; second pectoral filament not reaching to or
extending slightly beyond level of posterior tip of pelvic fin; third pectoral filament just reaching to or not reaching to level ofanal-fin origin; fourth pectoral filament reaching near level of posterior base of anal fin or just reaching to caudal-fin base; fifth
to seventh pectoral filaments longer than total length; sixth pectoral filament longest, its length 181 to 248% (mean 208%) of
standard length; caudal fin deeply forked, upper and lower caudal-fin lobes not filamentous, upper caudal-fin lobe 39 to 49%(mean 44%) and lower lobe 33 to 47% (mean 40%) of standard length. Pored lateral-line scales 66 to 71 (mode 70); lateralline simple, extending from upper end of gill opening to mid-distal margin of caudal-fin membrane; scale rows above lateralline 6 or 7 (mode 7), below 10 to 12 (mode 11). Gillrakers 12 to 14 (mode 13) on upper limb, 17 to 20 (mode 20) on lower limb,
30 to 34 (mode 32 or 33) total. Vertebrae 10 precaudal and 15 caudal; supraneural bones 2. Swimbladder absent. Colour:
Head and body greyish black dorsally, yellow ventrally; anterior parts of first and second dorsal fins greyish black, other parts
pale yellow; pectoral and pelvic fins uniformly vivid
yellow; base of pectoral filament s vivid yellow,
becoming whitish yellow on posterior tips; anal fin
uniformly yellow; posterior margin of caudal fin yellow,
other parts greyish black.
Geographical Distribution: Currently known from theeastern Indian to western Pacific Oceans, where it
ranges over continental shelves from western India to
Thailand (Fig. 151). Two specimens (ANSP 11498,
135 to 147 mm standard length) collected (probably
pre-1900) from Indonesia, lacked detailed locality andother collection data. An Indonesian distribution of thespecies therefore needs reconfirmation.
Habitat and Biology: Occurs in both estuarine andoffshore waters (from depths of less than 27 m), but
has been recorded as regularly entering fresh water for
breeding purposes (David, 1954). The species feeds
on crustaceans, small fishes and benthic organisms.

200 m

Fig. 151 Polynemus paradiseus
Known distribution

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FAO Species Catalogue for Fishery Purposes No. 3

Sexes of the species are separate (no evidence of hermaphroditism; Kagwade, 1970). Mukhopadhyay et al. (1995) inferredthat males mature at 110 mm total length; females at 120 mm. They suggested that the species spawns from April to
September, in the Hooghly River, India, females collected in October being mostly spent.
According to Gupta (1968), who examined ova diameter in Indian examples of the species, the largest diameter of
intraovarian eggs was 1.0 mm, the ova diameter in fully matured specimens being over 0.83 mm. However, Jones andMenon (1953), who had collected fertilized eggs of the same species, had earlier recorded a measurement of 0.7 mm indiameter and possession of an oil globule of 0.4 mm in diameter. They also reported a specimen of 170 mm standard
length having about 42 000 ova. Gupta (1968) noted that the right gonad was invariably more fecund than the left.
Jones and Menon’s (1953) study of the larval history of P. paradiseus from Indian waters recorded the swimbladder as
developing in the larval stage, but atrophying in juveniles. The yolk sac was absorbed by the 4.3 mm stage. With thecompletion of scale formation and the elongation of the pectoral filaments to adult proportions, the larvae enter the juvenilestage. Body scale rudiments were apparent in juveniles of 13 to 14 mm.
Size: Maximum standard length at least 20 cm (Motomura et al., 2002b).
Interest to Fisheries: One of the most important fisheries species for India (especially in West Bengal) and Bangladesh,and Myanmar.
Local Names: BANGLADESH: Tapasi; INDIA: Paradise threadfin; MYANMAR: Nga ponnar; THAILAND: Pla nuad pram.
Literature: Motomura et al. (2002d).
Remarks: P. paradiseus, 1 of 3 of the oldest available name for the family Polynemidae, was described on the basis of a
figure (reproduced in Motomura et al., 2002b: fig. 5) and description given by Edwards (1743-1751), who briefly described
the figured fish (collected from Bengal, India by Dr Mead) as the “Mango-Fish.” The specimen used for Edwards’ figure anddescription apparently no longer exists. However, Edwards’ figure clearly indicates VII spines in the first dorsal fin and 7
pectoral filaments, the uppermost 3 extending beyond the posterior tips of the caudal-fin lobes and fourth pectoral filament
not extending beyond the caudal-fin base. The characters of the specimens considered here as P. paradiseus agree wellwith those shown on the figure.
Subsequently, Hamilton (1822) described 3 new species with 7 pectoral filaments, Polynemus risua, P. toposui and
P. aureus, all from India. According to their original descriptions (Hamilton, 1822), all 3 species were characterized by having
7 long pectoral filaments and VII first dorsal-fin spines, but differed from each other in the numbers of dorsal-fin soft rays (17in P. risua, 16 in P. toposui and 15 in P. aureus), pectoral-fin rays (17, 16 and 17, respectively) and anal-fin rays (including
spines; 15, 16 and 14, respectively). However, these fin-ray numbers are almost wholly within the range of intraspecificvariation (see Motomura et al., 2002b). The diagnostic characters of 7 long pectoral filaments and VII first dorsal-fin spines
given in the original descriptions (supported by the collection locality) of these species are consistent with those of
P. paradiseus. Accordingly, P. risua, P. toposui and P. aureus are regarded as junior synonyms of P. paradiseus (seeMotomura et al., 2002b).
Although Fricke (1999) believed P. longifilis to be a junior synonym of Polydactylus plebeius, which is distributed in the
Indo-Pacific, the syntypes of the former clearly differ from examples of the latter, including the neotype, in having 7 pectoral
filaments (longer than standard length versus 5, shorter than standard length in the latter) and the orbit diameter smaller
than the posterior margin of the maxilla (larger). Examination of the syntypes of P. longifilis showed them to correspondclosely with the specimens considered here as P. paradiseus. Therefore, P. longifilis is also regarded as a junior synonymof P. paradiseus (see Motomura et al., 2002b).

Threadfins of the World95

3. LIST OF NOMINAL SPECIES OF POLYNEMIDAE
The following list gives in order (i) the scientific name as it originally appeared, in alphabetical order according to the specificname; (ii) the author(s) [Cuvier (1829b) = in Cuvier and Valenciennes (1829b); Jordan and McGregor (1907) = in Jordan and
Seale (1907); Valenciennes (1831) = in Cuvier and Valenciennes (1831)]; (iii) date of publication; and (iv) present
identification.

NOMINAL SPECIESPolydactylus agonasi Jordan and McGregor, 1906
Polynemus americanus Cuvier, 1829a
Polynemus antillarum Perugia, 1896
Polynemus approximans Lay and Bennett, 1839
Polynemus aquilonaris Motomura, 2003a
Polynemus artedii Bennett, 1831
Polynemus astrolabi Sauvage, 1881
Polydactylus auratus McKay, 1970
Polynemus aureus Hamilton, 1822
Polydactylus bifurcus Motomura, Kimura and Iwatsuki, 2001
Polynemus borneensis Bleeker, 1857b
Polynemus coecus Macleay, 1878
Polynemus californiensis Thominot, 1886
Polynemus commersonii Shaw, 1804
Polynemus decadactylus Bloch, 1795
Polynemus diagrammicus Bleeker, 1849
Polynemus dubius Bleeker, 1854
Polynemus emoi Lacepède, 1803
Polynemus enneadactylus Cuvier, 1829b
Polynemus gelatinosus McClelland, 1843
Polynemus heptadactylus Cuvier, 1829b
Polynemus hexanemus Cuvier, 1829b
Trichidion hilleri Fowler, 1905
Polynemus hornadayi Myers, 1936
Polynemus indicus Shaw, 1804
Polynemus intermedius Nichols, 1954
Polynemus kapuasensis Motomura and van Oijen, 2003
Polydactylus konadaensis Mishra and Krishnan, 1993
Polynemus kuru Bleeker, 1853b
Polynemus lineatus Lacepède, 1803
Polynemus longifilis Cuvier, 1829b
Polynemus longipectoralis Weber and de Beaufort, 1922
Polydactylus longipes Motomura, Okamoto and Iwatsuki, 2001
Polynemus lydiae Curtiss, 1938
Polynemus macrochir Günther, 1867
Polynemus macronemus Pel, 1851
Polynemus macrophthalmus Bleeker, 1858b
Polydactylus malagasyensis Motomura and Iwatsuki, 2001b
Polynemus mango Lacepède, 1803
Polynemus melanochir Valenciennes, 1831
Polynemus malanochir dulcis Motomura and Sabaj, 2002

PRESENT ALLOCATION

Polydactylus plebeius
Polydactylus virginicus
Polydactylus virginicus
Polydactylus approximans
Polynemus aquilonaris
Pentanemus quinquarius
Galeoides decadactylus
Polydactylus multiradiatus
Polynemus paradiseus
Polydactylus bifurcus
Polynemus melanochir melanochir
Eleutheronema tetradactylum
Polydactylus approximans
Polydactylus plebeius
Galeoides decadactylus
Filimanus xanthonema
Polynemus dubius
Polydactylus plebeius
Galeoides decadactylus
Leptomelanosoma indicum
Filimanus heptadactyla
Filimanus hexanema
Polynemus melanochir melanochir
Polynemus hornadayi
Leptomelanosoma indicum
Parapolynemus verekeri
Polynemus kapuasensis
Filimanus xanthonema
Polydactylus sexfilis
Polydactylus plebeius
Polynemus paradiseus
Polynemus dubius
Polydactylus longipes
Polydactylus plebeius
Polydactylus macrochir
Pentanemus quinquarius
Polydactylus macrophthalmus
Polydactylus malagasyensis
Polydactylus virginicus
Polynemus melanochir melanochir
Polynemus malanochir dulcis

96

FAO Species Catalogue for Fishery Purposes No. 3
NOMINAL SPECIESPolynemus melanopoma Günther, 1864
Polynemus melanopus Sauvage, 1881
Galeoides microps Steindachner, 1869
Polynemus microstoma Bleeker, 1851b
Polynemus multifilis Temminck and Schlegel, 1843
Polynemus multiradiatus Günther, 1860
Polynemus naso Walbaum, 1793
Polydactylus nigripinnis Munro, 1964
Polynemus niloticus Shaw, 1804
Trichidion octofilis Gill, 1861
Polynemus octonemus Girard, 1858
Polynemus oligodon Günther, 1860
Trichidion opercularis Gill, 1863
Polynemus paradiseus Linnaeus, 1758
Filimanus perplexa Feltes, 1991
Polydactylus persicus Motomura and Iwatsuki, 2001b
Polynemus pfeifferi Bleeker, 1853a
Polynemus plebeius Broussonet, 1782
Polydactylus plumierii Lacepède, 1803
Polynemus polydactylus Vahl, 1798
Polynemus quadrifilis Cuvier, 1829b
Polynemus quatordecimfilis Pel, 1851
Polynemus quinquarius Linnaeus, 1758
Polydactylus rhadinus Jordan and Evermann, 1902
Polynemus risua Hamilton, 1822
Polydactylus sealei Jordan and Richardson, 1910
Polynemus sele Hamilton, 1822
Polynemus senarius Gronow in Gray, 1854
Polynemus sexfilis Valenciennes, 1831
Polynemus sextarius Bloch and Schneider, 1801
Polynemus sextarius mullani Hora, 1926
Polynemus sheridani Macleay, 1884
Polydactylus siamensis Motomura, Iwatsuki and Yoshino, 2001
Filimanus similis Feltes, 1991
Polynemus specularis De Vis, 1883
Polynemus taeniatus Günther, 1860
Polynemus teria Hamilton, 1822
Polynemus tetradactylus Shaw, 1804
Polynemus toposui Hamilton, 1822
Polynemus tridactylus Bleeker, 1849
Polynemus uronemus Cuvier, 1829a
Polynemus verekeri Saville-Kent, 1889
Polynemus virginicus Linnaeus, 1758
Polynemus xanthonemus Valenciennes, 1831
Polydactylus zophomus Jordan and McGregor, 1907

PRESENT ALLOCATION

Polydactylus opercularis
Polynemus melanochir melanochir
Polynemus melanochir melanochir
Polydactylus microstomus
Polynemus multifilis
Polydactylus multiradiatus
Polydactylus virginicus
Polydactylus nigripinnis
Polydactylus plebeius
Polydactylus octonemus
Polydactylus octonemus
Polydactylus oligodon
Polydactylus opercularis
Polynemus paradiseus
Filimanus perplexa
Polydactylus persicus
Filimanus xanthonema
Polydactylus plebeius
Polydactylus virginicus
Galeoides decadactylus
Polydactylus quadrifilis
Polynemus multifilis
Pentanemus quinquarius
Eleutheronema rhadinum
Polynemus paradiseus
Filimanus sealei
Leptomelanosoma indicum
Filimanus hexanema
Polydactylus sexfilis
Polydactylus sextarius
Polydactylus mullani
Polydactylus macrochir
Polydactylus siamensis
Filimanus similis
Polydactylus multiradiatus
Polydactylus plebeius
Eleutheronema tetradactylum
Eleutheronema tetradactylum
Polynemus paradiseus
Eleutheronema tridactylum
Leptomelanosoma indicum
Parapolynemus verekeri
Polydactylus virginicus
Filimanus xanthonema
Polydactylus microstomus

Threadfins of the World97

4. LIST OF SPECIES BY MAJOR FISHING AREASGEOGRAPHICAL DISTRIBUTIONSPECIESFRESHWATERS

MAJOR FISHING AREA FOR STATISTICAL PURPOSES
21

31

34

37

41

47

51

57

Eleutheronema rhadinum•••

Eleutheronema tetradactylum
Eleutheronema tridactylum
Filimanus heptadactyla••

Filimanus hexanema
Filimanus perplexa
Filimanus sealei
Filimanus similis
Filimanus xanthonema
Galeoides decadactylus

••

•

•

•

Leptomelanosoma indicum
Parapolynemus verekeri

••
•
••

06

Pentanemus quinquarius

61

•

71

•

Polydactylus longipes•

Polydactylus macrochir
Polydactylus macrophthalmus

04

Polydactylus microstomus
Polydactylus mullani
Polydactylus multiradiatus
Polydactylus nigripinnis
Polydactylus octonemus
Polydactylus oligodon

••
•

•

87

•
•
•
•
•
•
•

•
••••
•
• •
• •

Polydactylus malagasyensis

81

•
•
•
•
•
•

Polydactylus approximans
Polydactylus bifurcus

77

•

•

•

98

FAO Species Catalogue for Fishery Purposes No. 3

GEOGRAPHICAL DISTRIBUTIONSPECIESFRESHWATERS

MAJOR FISHING AREA FOR STATISTICAL PURPOSES
21

31

34

37

Polydactylus persicus
Polydactylus plebeius•

Polydactylus sexfilis
Polydactylus sextarius
Polydactylus siamensis••

Polydactylus virginicus
Polynemus aquilonaris

04

Polynemus dubius

04

Polynemus hornadayi

04

Polynemus kapuasensis

04

Polynemus melanochir dulcis

04

Polynemus melanochir melanochir

04

Polynemus multifilis

04

Polynemus paradiseus

04

47

51

57

61

71

77

81

•
•
•
••••••

Polydactylus opercularis

Polydactylus quadrifilis

41

•

•••••
••••
• •

••

•
•
•
•
•
•
•
•

87

Threadfins of the WorldMAJOR FISHING AREAS FOR STATISTICAL PURPOSES

99

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FAO Species Catalogue for Fishery Purposes No. 3

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6. INDEX OF SCIENTIFIC AND VERNACULAR NAMESEXPLANATION OF THE SYSTEM

Italics: Valid scientific names (double entry by genera and species)

Italics: Synonyms and misidentifications (double entry by genera and species)

ROMAN : Family names
ROMAN : Names of orders, class, subclass, superfamilies
Roman : FAO names
Roman: Local names

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A

Barbudo javanés . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Barbudo mancha negra africano . . . . . . . . . . . . . . . . 54
Barbudo ocho dedos. . . . . . . . . . . . . . . . . . . . . . . . . . 27
Barbudo ocho dedos atlántico . . . . . . . . . . . . . . . . . . 62
Barbudo paraíso . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Barbudo paraíso de Hornaday . . . . . . . . . . . . . . . . . . 84
Barbudo paraíso de mano negra . . . . . . . . . . . . . . . . 89
Barbudo paraíso de mano negra lacustre . . . . . . . . . 88
Barbudo paraíso elegante. . . . . . . . . . . . . . . . . . . . . . 91
Barbudo paraíso enano . . . . . . . . . . . . . . . . . . . . . . . 36
Barbudo paraíso de Kapuas . . . . . . . . . . . . . . . . . . . . 86
Barbudo paraíso norteño . . . . . . . . . . . . . . . . . . . . . . 81
Barbudo paraíso oriental . . . . . . . . . . . . . . . . . . . . . . 83
Barbudo rayado . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
Barbudo rayado de boca grande . . . . . . . . . . . . . . . . 76
Barbudo real . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Barbudo seis barbas . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Barbudo-gigante . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Barbudo-real . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Barbudos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Barbure à doigts oranges . . . . . . . . . . . . . . . . . . . . . . 29
Barbure à dos sombre . . . . . . . . . . . . . . . . . . . . . . . . 59
Barbure à grande bouche . . . . . . . . . . . . . . . . . . . . . . 76
Barbure à huit doigts . . . . . . . . . . . . . . . . . . . . . . . . . 27
Barbure à nageoires noires . . . . . . . . . . . . . . . . . . . . 61
Barbure à petite bouche . . . . . . . . . . . . . . . . . . . . . . . 56
Barbure à petites écailles . . . . . . . . . . . . . . . . . . . . . . 64
Barbure à quatre doigts . . . . . . . . . . . . . . . . . . . . . . . 17
Barbure à quatre doigts d'Asie . . . . . . . . . . . . . . . . . 15
Barbure à sept doigts . . . . . . . . . . . . . . . . . . . . . . . . . 23
Barbure à six doigts . . . . . . . . . . . . . . . . . . . . . . . . . . 72
Barbure à tâche noire . . . . . . . . . . . . . . . . . . . . . . . . . 74
Barbure à tâche noire d'Arabie. . . . . . . . . . . . . . . . . . 57
Barbure à tâche noire de Perse . . . . . . . . . . . . . . . . . 67
Barbure à trois doigts . . . . . . . . . . . . . . . . . . . . . . . . . 19
Barbure bleu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Barbure de flaque . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Barbure de rivière . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Barbure gros-doigts . . . . . . . . . . . . . . . . . . . . . . . . . . 50
Barbure huit doigts atlantique . . . . . . . . . . . . . . . . . . 62
Barbure indien . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Barbure jaune . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Barbure javanais . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Barbure longs-doigts . . . . . . . . . . . . . . . . . . . . . . . . . 48
Barbure magnifique. . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Barbure paradis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 93
Barbure paradis à doigts noirs . . . . . . . . . . . . . . . . . 89
Barbure paradis à doigts noirs de lac . . . . . . . . . . . . 88
Barbure paradis de Hornaday . . . . . . . . . . . . . . . . . . 84
Barbure paradis d'Orient . . . . . . . . . . . . . . . . . . . . . . 83
Barbure paradis du nord. . . . . . . . . . . . . . . . . . . . . . . 81
Barbure paradis élégante . . . . . . . . . . . . . . . . . . . . . . 91
Barbure paradis de Kapuas . . . . . . . . . . . . . . . . . . . . 86
Barbure paradis nain. . . . . . . . . . . . . . . . . . . . . . . . . . 36
Barbure rayé . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68

African blackspot threadfin . . . . . . . . . . . . . . . . . . . . 54
agonasi, Polydactylus . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Aguapuro. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Akin-akin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
americanus, Polynemus . . . . . . . . . . . . . . . . . . . . . . 77-78
antillarum, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 77
approximans, Polydactylus . . . . . . . . . . 19, 39, 44-47, 66
approximans, Polynemus . . . . . . . . . . . . . . . . . . . . . . . 44
aquilonaris, Polynemus . . . . . . . . . . . . . 4, 79, 81-82, 84
Arabian blackspot threadfin . . . . . . . . . . . . . . . . . . . . 57
artedii, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . 37-38
astrolabi, Polynemus . . . . . . . . . . . . . . . . . . . . . 31-33, 75
Atkaru . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Atlantic threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
auratus, Polydactylus. . . . . . . . . . . . . . . . . . . . . . . . 59-60
aureus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 92, 94
Australian threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . 59

B
Baling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Barbeta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45, 66
Barbi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Barbinho . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Barbo. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32, 71
Barbu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77-78
Barbú. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 64, 78
Barbu argenté . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Barbudo . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32, 38, 71, 78
Barbudo amarillo . . . . . . . . . . . . . . . . . . . . . . . . . . 65-66
Barbudo azul. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44-45
Barbudo cuatro dedos de Asia. . . . . . . . . . . . . . . . . . 15
Barbudo de aletas negras. . . . . . . . . . . . . . . . . . . . . . 61
Barbudo de boca pequeña . . . . . . . . . . . . . . . . . . . . . 56
Barbudo de charco. . . . . . . . . . . . . . . . . . . . . . . . . 77-78
Barbudo de cuatro dedos . . . . . . . . . . . . . . . . . . . . . . 17
Barbudo de dedos amarillos . . . . . . . . . . . . . . . . . . . 29
Barbudo de dedos largos . . . . . . . . . . . . . . . . . . . . . . 48
Barbudo de escamas pequeñas. . . . . . . . . . . . . . . . . 64
Barbudo de lomo oscuro . . . . . . . . . . . . . . . . . . . . . . 59
Barbudo de mancha negra . . . . . . . . . . . . . . . . . . . . . 74
Barbudo de mancha negra persa. . . . . . . . . . . . . . . . 67
Barbudo de mancha negra árabe. . . . . . . . . . . . . . . . 57
Barbudo de manos grandes . . . . . . . . . . . . . . . . . . . . 50
Barbudo de rio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Barbudo de seis dedos . . . . . . . . . . . . . . . . . . . . . . . . 72
Barbudo de siete dedos . . . . . . . . . . . . . . . . . . . . . . . 23
Barbudo de siete dedos índico . . . . . . . . . . . . . . . . . 28
Barbudo de tres dedos . . . . . . . . . . . . . . . . . . . . . . . . 19
Barbudo enano africano . . . . . . . . . . . . . . . . . . . . . . . 31
Barbudo esbelto . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Barbudo espléndido . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Barbudo gigante africano . . . . . . . . . . . . . . . . . . . . . . 71
Barbudo indio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Threadfins of the World
Barbure sept doigts des indes . . . . . . . . . . . . . . . . . . 28
Barbure svelte . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Barbure tâche noire d'Afrique. . . . . . . . . . . . . . . . . . . 54
Barbures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Barmeen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Barudo de dez barbas . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Barudo de mancha. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Barudo raiado . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Bearbear . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Beard fish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Beardfish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Becadulce . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Benong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
bifurcus, Polydactylus . . . . . . 9, 19, 39, 46-47, 51, 70, 76
Bikau . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Blackfin threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 61
Black-finned threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Blackhand paradise fish . . . . . . . . . . . . . . . . . . . . . . . 89
Blackspot threadfin . . . . . . . . . . . . . . . . . . . . . . . . 74-75
Black-spot threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Blind tassel-fish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Blue bobo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Blue salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Blue threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Bluenose salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Blunt-nosed salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
borneensis, Polynemus . . . . . . . . . . . . . . . . . . . . . . . 89-90
borneënsis, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 89
Bou na pana. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Burnett salmon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51

C
Cá chét. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
californiensis, Polynemus . . . . . . . . . . . . . . . . . . . . . 44-45
Capitaine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38, 71
Capitaine royal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Capitão. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Catfish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Cekéém . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Char-wen-e-kerning . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Chikoué . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Chinese nose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
coecus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 18
Colonial salmon . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
commersonii, Polynemus . . . . . . . . . . . . . . . . . . . . . 68, 70
Common threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Cooktown salmon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

D
Dagi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
decadactylus, Galeoides . . . . . . . . . . . . . . . . . . . 31-33, 75
decadactylus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 31
diagrammicus, Polynemus . . . . . . . . . . . . . . . . . . . . 29-30
dubius, Polynemus . . . . . . . . . . . . . . . . . . . . . . 79, 82, 84
dulcis, Polynemus melanochir . . . . 79, 82, 84, 87-89, 91

111
Dwarf paradise fish . . . . . . . . . . . . . . . . . . . . . . . . . . . 36

E
East Asian fourfinger threadfin . . . . . . . . . . . . . . . . . 15
Eastern paradise fish. . . . . . . . . . . . . . . . . . . . . . . . . . 83
Eightfinger threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Eight-fingered threadfin . . . . . . . . . . . . . . . . . . . . . . . . . 27
Elegant paradise fish . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Eleutheronema . . . . . . . . . . . . . . . . . . . . . 1, 9, 13-14, 40
Eleutheronema rhadinum . . . . . . . . . . . 13, 15-17, 19-20
Eleutheronema tetradactylum . . . . . . . 3-4, 13, 16, 18-20
Eleutheronema tridactylum . . . . . . . . . . . 9, 13, 16, 19-20
emoi, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
enneadactylus, Polynemus . . . . . . . . . . . . . . . . . . . . 31-32

F
Filimanus . . . . . . . . . . . . . . . . . . . . . . . 1, 9, 20-21, 37, 40
Filimanus heptadactyla . . . . . . . 21, 23-24, 28-30, 58, 60
Filimanus hexanema . . . . . . . . . . . . . . . 21, 24-26, 28, 30
Filimanus melanochir . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Filimanus perplexa. . . . . . . . . . . 20-21, 25-26, 28, 30, 90
Filimanus sealei. . . . . . . . . . . . . . . . . . . . . . 21, 27-28, 30
Filimanus similis . . . . . . . . . . . . . . . . . . . . . 21, 24, 28-30
Filimanus xanthonema . . . . . . . . . . . . . . . . 21, 28-29-30
Fourfinger threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . 17

G
Gainfio . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Galeoides . . . . . . . . . . . . . . . . . . . . . . . . . . 1, 9, 31, 40, 90
Galeoides decadactylus . . . . . . . . . . . . . . . . . . . 31-33, 75
Galeoides microps . . . . . . . . . . . . . . . . . . . . . . . 31, 89-90
Galeoides polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . 32
Gatha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Gbalakassa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
gelatinosus, Polynemus . . . . . . . . . . . . . . . . . . . . . . 33, 35
Genohong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Gestreepe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Ghazal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Giant African threadfin . . . . . . . . . . . . . . . . . . . . . . . . 71
Giant threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Gold threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Golden sixthread tesselfish . . . . . . . . . . . . . . . . . . . . . . 73
Gor-ah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Gros capitaine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
Gubal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Günther’s threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 60

Hheptadactyla, Filimanus . . . . . . . 21, 23-24, 28-30, 58, 60
heptadactylus, Polydactylus . . . . . . . . . . . . . . . 24, 58, 60
heptadactylus, Polynemus . . . . . . . . . . . . . . . . . . . . 23, 58
hexanema, Filimanus . . . . . . . . . . . . . . 21, 24-26, 28, 30
hexanema, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . 24
hilleri, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
hilleri, Trichidion. . . . . . . . . . . . . . . . . . . . . . . . . . . 89, 91

112
Hornaday’s paradise fish . . . . . . . . . . . . . . . . . . . . . . 84
hornadayi, Polynemus . . . . . . . . . . . . . . . . . . . 79, 84-85
Hugao . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Hugau . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Hwa-n le kale-n . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

I
I’ausi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69, 73
Ikan salangan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Indian sevenfinger threadfin. . . . . . . . . . . . . . . . . . . . 28
Indian threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . 33, 35
indicum, Leptomelanosoma . . . . . . . . . . 1, 33-35, 50, 66
indicus, Polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . . 34
indicus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 33, 35
intermedius, Polynemus . . . . . . . . . . . . . . . . . . . . . . 35-36

J
Javanese threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Jenohong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Jin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Jubal senohong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

K
Ka ku yan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Kagau . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kakuyan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Kalawa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kalemeen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kapuas elegant paradise fish . . . . . . . . . . . . . . . . . . . 86
kapuasensis, Polynemus . . . . . . . . . . . . . . . 79, 84, 86-87
Kataguro-agonashi . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Kesumbang . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
King salmon. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
King threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 50-51
konadaensis, Polydactylus . . . . . . . . . . . . . . . . . . . . 29-30
Kubal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kugao . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kupe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55, 69
Kurau . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 35, 75, 90
Kurau hitam . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Kurau janggut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Kuro. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Kurow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Kuru janggut . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
kuru, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 72-73
Kutlikala. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Kuwa-kuwa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57

L
Laas. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Lajan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Lake blackhand paradise fish. . . . . . . . . . . . . . . . . . . 88
Lakhua. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Laos. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

FAO Species Catalogue for Fishery Purposes No. 3
Largemouth striped threadfin. . . . . . . . . . . . . . . . . . . 76
Lausan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 69
Lelaoh tanah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
Leptomelanosoma . . . . . . . . . . . . . . . . . . . . . 1, 9, 33, 40
Leptomelanosoma indicum . . . . . . . . . . . 1, 33-35, 50, 66
Lesser African threadfin . . . . . . . . . . . . . . . . . . . . 31-32
lineatus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Littlescale threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Longfinned threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
longifilis, Polynemus . . . . . . . . . . . . . . . . . . . . . 84, 92, 94
longipectoralis, Polynemus . . . . . . . . . . . . . . . . . . . 82, 84
longipes, Polydactylus . . . . . 33, 39, 45, 48-49, 53, 62, 73
Long-limb threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . 48
lydiae, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . 68, 70

Mmacrochir, Polydactylus . . . . . . . . . 39, 47, 49-51, 53, 60
macrochir, Polynemus. . . . . . . . . . . . . . . . . . . . . . . . . . 49
macronema, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 89
macronemus, Polynemus . . . . . . . . . . . . . . . . . . 37-38, 89
macrophthalmus, Polydactylus . 1, 39, 49, 51, 53, 60, 79
macrophthalmus, Polynemus . . . . . . . . . . . . . . . . . . . . 51
malagasyensis, Polydactylus . . . . 1, 9, 39, 45, 49, 54-55,
57-58, 62, 68, 73, 75
Mamali. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Mamaling babai . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Mamaling bato . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Mamangi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Manangi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Mancong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 35, 75
mango, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . 77-78
Many-rayed threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . 60
melanochir dulcis, Polynemus . . . . 79, 82, 84, 87-89, 91
melanochir melanochir, Polynemus . . . . . 31, 79, 82, 84,
88-89-91
melanochir, Filimanus . . . . . . . . . . . . . . . . . . . . . . . . . 90
melanochir, Polynemus . . . . . . . . . . . . . . 20-21, 26, 89-90
melanochir, Polynemus melanochir . . . . . 31, 79, 82, 84,
88-89-91
melanochir, Trichidion . . . . . . . . . . . . . . . . . . . . . . . . . 90
melanopoma, Polynemus . . . . . . . . . . . . . . . . . . . . . 65-66
melanopus, Polynemus. . . . . . . . . . . . . . . . . . . . . . . 89-90
Melong. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
microps, Galeoides . . . . . . . . . . . . . . . . . . . . . . 31, 89-90
microstoma, Polynemus . . . . . . . . . . . . . . . . . . . . . 55, 57
microstomus, Polydactylus . . . . . 1, 39, 47, 51, 55-58, 62,
68, 70, 73, 75
Minami-konoshiro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Mkizi komo maji . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55, 69
Moi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69, 73
Moili’I . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
MUGILIDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Mulet barbe . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Mulet bâtard. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
mullani, Polydactylus . . . . . . . 1, 9, 24, 39, 53, 55, 57-60,
68-69, 73, 75

Threadfins of the World
mullani, Polynemus sextarius . . . . . . . . . . . . . . . . . . 57-58
multifilis, Polynemus . . . . . . . . . . . . . 4, 78-79, 84, 87, 91
multiradiatus, Polydactylus . . . . . . . . 3, 39, 53, 59-60, 66
multiradiatus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . 59
Mulut tikus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

N
Nan’you-agonashi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
naso, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 77-78
Nga ponnar . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Nga-let-kwa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
nigripinnis, Polydactylus . . . 39, 45, 49, 55, 61-62, 66, 73
niloticus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Njaane . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38, 71
Njaane jaara. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38, 71
Northern paradise fish. . . . . . . . . . . . . . . . . . . . . . . . . 81
Noxan. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

Ooctofilis, Trichidion . . . . . . . . . . . . . . . . . . . . . . . . . 62-63
octonemus, Polydactylus . . . . . . 39, 45, 47, 62-63, 66, 78
octonemus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 62
oligodon, Polydactylus . . . . . . . . . . 39, 45, 47, 63, 65, 78
oligodon, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
opercularis, Polydactylus . . . 27-28, 33, 39, 45-47, 65-66
opercularis, Trichidion . . . . . . . . . . . . . . . . . 27-28, 65-66

P
Pááwánér. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Pacific threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Pacific thread-fin fish . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Paradise threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . 93-94
paradiseus, Polynemus . . . . 3-4, 78-79, 84-85, 88, 92, 94
Parapolynemus . . . . . . . . . . . . . 1, 3, 9, 33, 35, 37, 40, 79
Parapolynemus verekeri . . . . . . . . . . . . . . . . . . 35-36, 66
Peixe-barba . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
pentadactyla, Sciaena . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Pentanemus . . . . . . . . . . . . . . . . . . . . . . . . 1, 9, 37, 40, 79
Pentanemus quinquarius . . . . . . . . . . . . . . . . . . . . 37-38
perplexa, Filimanus . . . . . . . . . . 20-21, 25-26, 28, 30, 90
Persian blackspot threadfin . . . . . . . . . . . . . . . . . . . . 67
persicus, Polydactylus . . . . . . . . . 1, 39, 45, 49, 55, 57-58,
62, 66, 68, 73, 75
Petit capitaine. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
pfeifferi, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 29-30
Pla nuad pram . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
plebeius, Polydactylus . . . 9, 34, 39, 47, 51, 68-70, 76, 94
plebeius, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
plebejus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 57, 70
Plexiglass. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
plumierii, Polydactylus . . . . . . . . . . . . . . . . . 39-40, 77-78
Polekala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Polistonemus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78-79
Polydactylus . . . . . . . 1, 8-9, 19, 33, 39-40, 43, 45, 47, 53,
55, 60, 66, 68, 73, 75, 79
Polydactylus (= Polynemus) sexfilis . . . . . . . . . . . . . . . 75

113
Polydactylus agonasi . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polydactylus approximans . . . . . . . . . . . 19, 39, 44-47, 66
Polydactylus auratus . . . . . . . . . . . . . . . . . . . . . . . . 59-60
Polydactylus bifurcus. . . . . . . 9, 19, 39, 46-47, 51, 70, 76
Polydactylus heptadactylus . . . . . . . . . . . . . . . . 24, 58, 60
Polydactylus indicus . . . . . . . . . . . . . . . . . . . . . . . . . . . 34
Polydactylus konadaensis . . . . . . . . . . . . . . . . . . . . 29-30
Polydactylus longipes. . . . . . 33, 39, 45, 48-49, 53, 62, 73
Polydactylus macrochir . . . . . . . . . . 39, 47, 49-51, 53, 60
Polydactylus macrophthalmus . . 1, 39, 49, 51, 53, 60, 79
Polydactylus malagasyensis . . . . 1, 9, 39, 45, 49, 54-55,
57-58, 62, 68, 73, 75
Polydactylus microstomus . . . . . 1, 39, 47, 51, 55-58, 62,
68, 70, 73, 75
Polydactylus mullani . . . . 1, 9, 24, 39, 53, 55, 57-60, 62,
68-69, 73, 75
Polydactylus multiradiatus . . . . . . . . 3, 39, 53, 59-60, 66
Polydactylus nigripinnis . . . 39, 45, 49, 55, 61-62, 66, 73
Polydactylus octonemus. . . . . . . 39, 45, 47, 62-63, 66, 78
Polydactylus oligodon . . . . . . . . . . . 39, 45, 47, 63, 65, 78
Polydactylus opercularis . . . . 27-28, 33, 39, 45-47, 65-66
Polydactylus persicus . . . 1, 39, 45, 49, 55, 57-58, 62, 66,
68, 73, 75
Polydactylus plebeius . . . . 9, 34, 39, 47, 51, 68-70, 76, 94
Polydactylus plumierii. . . . . . . . . . . . . . . . . . 39-40, 77-78
Polydactylus quadrifilis . . . . . . . . . . . . . 39, 46-47, 70-71
Polydactylus rhadinus . . . . . . . . . . . . . . . . . . . . . . . 15-16
Polydactylus sealei . . . . . . . . . . . . . . . . . . . . . . . . . . 27-28
Polydactylus sexfilis . . . . . . . 3-4, 9, 39, 45, 49, 62, 72-74
Polydactylus sextarius . . . . . . 1, 9, 24, 32, 39, 45, 49, 55,
57-58, 62, 68, 73-74-75
Polydactylus sheridani . . . . . . . . . . . . . . . . . . . . . . . . . 51
Polydactylus siamensis . . . . . 9, 39, 47, 51, 66, 70, 75-76
Polydactylus virginicus . . 19, 39-40, 45-47, 63, 65, 77-78
Polydactylus zophomus . . . . . . . . . . . . . . . . . . . . . . 55, 57
polydactylus, Galeoides . . . . . . . . . . . . . . . . . . . . . . . . . 32
polydactylus, Polynemus . . . . . . . . . . . . . . . . . . . . . 31-32
POLYNEMOIDEA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Polynemus . . . . . . . . . . . 1, 3, 9, 18, 21, 31, 40, 53, 78-79
Polynemus americanus . . . . . . . . . . . . . . . . . . . . . . . 77-78
Polynemus antillarum . . . . . . . . . . . . . . . . . . . . . . . . . . 77
Polynemus approximans . . . . . . . . . . . . . . . . . . . . . . . . 44
Polynemus aquilonaris . . . . . . . . . . . . . . 4, 79, 81-82, 84
Polynemus artedii . . . . . . . . . . . . . . . . . . . . . . . . . . . 37-38
Polynemus astrolabi . . . . . . . . . . . . . . . . . . . . . . 31-33, 75
Polynemus aureus . . . . . . . . . . . . . . . . . . . . . . . . . . 92, 94
Polynemus borneensis . . . . . . . . . . . . . . . . . . . . . . . 89-90
Polynemus borneënsis . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Polynemus californiensis . . . . . . . . . . . . . . . . . . . . . 44-45
Polynemus coecus . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 18
Polynemus commersonii . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus decadactylus . . . . . . . . . . . . . . . . . . . . . . . . 31
Polynemus diagrammicus . . . . . . . . . . . . . . . . . . . . 29-30
Polynemus dubius . . . . . . . . . . . . . . . . . . . . . . . 79, 82, 84
Polynemus emoi . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus enneadactylus . . . . . . . . . . . . . . . . . . . . 31-32
Polynemus gelatinosus . . . . . . . . . . . . . . . . . . . . . . 33, 35

114

FAO Species Catalogue for Fishery Purposes No. 3

Polynemus heptadactylus . . . . . . . . . . . . . . . . . . . . 23, 58
Polynemus hexanema . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Polynemus hilleri . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Polynemus hornadayi . . . . . . . . . . . . . . . . . . . . 79, 84-85
Polynemus indicus . . . . . . . . . . . . . . . . . . . . . . . . . 33, 35
Polynemus intermedius . . . . . . . . . . . . . . . . . . . . . . 35-36
Polynemus kapuasensis . . . . . . . . . . . . . . . 79, 84, 86-87
Polynemus kuru . . . . . . . . . . . . . . . . . . . . . . . . . . . . 72-73
Polynemus lineatus . . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus longifilis . . . . . . . . . . . . . . . . . . . . . 84, 92, 94
Polynemus longipectoralis . . . . . . . . . . . . . . . . . . . 82, 84
Polynemus lydiae . . . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus macrochir . . . . . . . . . . . . . . . . . . . . . . . . . . 49
Polynemus macronema . . . . . . . . . . . . . . . . . . . . . . . . . 89
Polynemus macronemus . . . . . . . . . . . . . . . . . . 37-38, 89
Polynemus macrophthalmus . . . . . . . . . . . . . . . . . . . . 51
Polynemus mango . . . . . . . . . . . . . . . . . . . . . . . . . . 77-78
Polynemus melanochir . . . . . . . . . . . . . . 20-21, 26, 89-90
Polynemus melanochir dulcis . . . . . 79, 82, 84, 87-89, 91
Polynemus melanochir melanochir . . . . . 31, 79, 82, 84,
88-89-91

Polynemus melanopoma . . . . . . . . . . . . . . . . . . . . . 65-66
Polynemus melanopus . . . . . . . . . . . . . . . . . . . . . . . 89-90
Polynemus microstoma . . . . . . . . . . . . . . . . . . . . . . 55, 57
Polynemus multifilis . . . . . . . . . . . . . 4, 78-79, 84, 87, 91
Polynemus multiradiatus . . . . . . . . . . . . . . . . . . . . . . . 59
Polynemus naso . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77-78
Polynemus niloticus . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus octonemus . . . . . . . . . . . . . . . . . . . . . . . . . . 62
Polynemus oligodon . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
Polynemus paradiseus . . . . 3-4, 78-79, 84-85, 88, 92, 94
Polynemus pfeifferi . . . . . . . . . . . . . . . . . . . . . . . . . 29-30
Polynemus plebeius . . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus plebejus . . . . . . . . . . . . . . . . . . . . . . . . . 57, 70
Polynemus polydactylus . . . . . . . . . . . . . . . . . . . . . . 31-32
Polynemus quadrifilis . . . . . . . . . . . . . . . . . . . . . . . . . . 70
Polynemus quatordecimfilis . . . . . . . . . . . . . . . . . . . . . 91
Polynemus quinquarius . . . . . . . . . . . . . . . . . . . . . . 37, 79
Polynemus risua . . . . . . . . . . . . . . . . . . . . . . . . . . . 92, 94
Polynemus salliah . . . . . . . . . . . . . . . . . . . . . . . . . . 18-19
Polynemus sele . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33, 35
Polynemus senarius . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Polynemus sexfilis . . . . . . . . . . . . . . . . . . . . . . . . . . 72, 75
Polynemus sextarius . . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Polynemus sextarius mullani . . . . . . . . . . . . . . . . . . 57-58
Polynemus sheridani . . . . . . . . . . . . . . . . . . . . . . . . 49-50
Polynemus specularis . . . . . . . . . . . . . . . . . . . . . . . 59-60
Polynemus taeniatus . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Polynemus teria . . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 18
Polynemus tetradactylus . . . . . . . . . . . . . . . . . . 13, 16, 18
Polynemus toposui . . . . . . . . . . . . . . . . . . . . . . . . . 92, 94
Polynemus tridactijlu . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Polynemus tridactylus . . . . . . . . . . . . . . . . . . . . . . . 13, 19
Polynemus uronemus . . . . . . . . . . . . . . . . . . . . . . . 33, 35
Polynemus verekeri . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Polynemus virginicus . . . . . . . . . . . . . . . . . . . . . . . 77, 79

Polynemus xanthonemus . . . . . . . . . . . . . . . . . . . . . 29-30
Pozhakkala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Pugao . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

Q
quadrifilis, Polydactylus . . . . . . . . . . . . 39, 46-47, 70-71
quadrifilis, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 70
quatordecimfilis, Polynemus . . . . . . . . . . . . . . . . . . . . . 91
quinquarius, Pentanemus . . . . . . . . . . . . . . . . . . . . 37-38
quinquarius, Polynemus . . . . . . . . . . . . . . . . . . . . . 37, 79

R
Rashgoo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
rhadinum, Eleutheronema . . . . . . . . . . . . . 13, 15-17, 20
rhadinus, Polydactylus. . . . . . . . . . . . . . . . . . . . . . . 15-16
risua, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 92, 94
River threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 52
Rockhampton salmon . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Royal threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37

S
Saccolih. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Salliah . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
salliah, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . 18-19
Samaduul . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Sanis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Sardijntje . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Sciaena pentadactyla . . . . . . . . . . . . . . . . . . . . . . . . . . 70
SCIAENIDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
sealei, Filimanus . . . . . . . . . . . . . . . . . . . . . 21, 27-28, 30
sealei, Polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . 27-28
Selanghi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
sele, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33, 35
Senangin . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 35, 69, 75
Senangin buih . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Senangin buis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
senarius, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Senohong . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 75
Sesvinger-draadvin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
Sevenfinger threadfin . . . . . . . . . . . . . . . . . . . . . . . . . 23
Sevenfingered threadfin. . . . . . . . . . . . . . . . . . . . . . . . . 78
Seven-fingered threadfin . . . . . . . . . . . . . . . . . . . . . . . . 24
sexfilis, Polydactylus . . . . . . 3-4, 9, 39, 45, 49, 62, 72-74
sexfilis, Polydactylus (= Polynemus) . . . . . . . . . . . . . . 75
sexfilis, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 72, 75
sextarius mullani, Polynemus . . . . . . . . . . . . . . . . . 57-58
sextarius, Polydactylus . . . . . 1, 9, 24, 32, 39, 45, 49, 55,
57-58, 62, 68, 73-74-75
sextarius, Polynemus. . . . . . . . . . . . . . . . . . . . . . . . . . . 74
Sheem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Sheridan’s threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
sheridani, Polydactylus . . . . . . . . . . . . . . . . . . . . . . . . . 51
sheridani, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 49-50
Shi-jih-ma-fa-yu . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Shineose . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

Threadfins of the World
siamensis, Polydactylus . . . . . 9, 39, 47, 51, 66, 70, 75-76
Sibet mba. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Sikket mbàw. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Sikor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
similis, Filimanus . . . . . . . . . . . . . . . . . . . . 21, 24, 28-30
Sinanghi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Sixfinger threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . 55, 72
Six-fingered threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . 73
Slender fivefinger threadfin . . . . . . . . . . . . . . . . . . . . 46
Smallmouth threadfin . . . . . . . . . . . . . . . . . . . . . . . . . 56
Small-mouthed threadfin . . . . . . . . . . . . . . . . . . . . . . . . 57
Smallscale threadfin. . . . . . . . . . . . . . . . . . . . . . . . . . . . 64
Sori . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
specularis, Polynemus . . . . . . . . . . . . . . . . . . . . . . . 59-60
SPHYRAENIDAE. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
Splendid threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
Streamered tasselfish. . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Striped threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . 68-69
Sumbal. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 69
Suro . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

Ttaeniatus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . 68, 70
Taer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
Tahlunkala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Tailla. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
Tapasan . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Tapasi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
teria, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 16, 18
tetradactylum, Eleutheronema . . . . . . . 3-4, 13, 16, 18-20
tetradactylus, Polynemus . . . . . . . . . . . . . . . . . . 13, 16, 18
Threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38, 78
Threadfins. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Threadfish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 78
Thread-fish. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Threefinger threadfin . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Tiekem . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
toposui, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 92, 94
Topshi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

115
Trey pream. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 89
Trey pream loeung . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Trey pream sor. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
Trichidion hilleri . . . . . . . . . . . . . . . . . . . . . . . . . . . 89, 91
Trichidion melanochir . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Trichidion octofilis . . . . . . . . . . . . . . . . . . . . . . . . . . 62-63
Trichidion opercularis. . . . . . . . . . . . . . . . . . 27-28, 65-66
tridactijlus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . 20
tridactylum, Eleutheronema . . . . . . . . . . 9, 13, 16, 19-20
tridactylus, Polynemus . . . . . . . . . . . . . . . . . . . . . . 13, 19
Tsubame-konoshiro . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69

UUculuka . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
Umi’umia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69, 73
Umiumia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
uronemus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . 33, 35

Vverekeri, Parapolynemus . . . . . . . . . . . . . . . . . . 35-36, 66
verekeri, Polynemus . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
virginicus, Polydactylus . . 19, 39-40, 45-47, 63, 65, 77-78
virginicus, Polynemus . . . . . . . . . . . . . . . . . . . . . . . 77, 79

Xxanthonema, Filimanus . . . . . . . . . . . . . . . . 21, 28-29-30
xanthonemus, Polynemus . . . . . . . . . . . . . . . . . . . . . 29-30

Y
Yellow bobo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 65
Yellowthread threadfin . . . . . . . . . . . . . . . . . . . . . . . . 29
Yevakala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Z
Za yaw . . . . . . . . . . . . . . . . . . . . . . . . . . . 29-30, 69, 75-76
Za yaw gyi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18
zophomus, Polydactylus . . . . . . . . . . . . . . . . . . . . . 55, 57

Threadfins of the World117

7. LIST OF COLOUR PLATESPLATE Ia)

Eleutheronema rhadinum, 152 mm SLe)

Filimanus hexanema, 82 mm SLb)

Eleutheronema tetradactylum, 121 mm SL

f)

Filimanus perplexa, 110 mm SLc)

Eleutheronema tridactylum, 180 mm SLg)

Filimanus sealei, 124 mm SLd)Filimanus heptadactyla, 90 mm SLh)Filimanus similis, 98 mm SL

PLATE IIa)

Filimanus xanthonema, 62 mm SLe)

Pentanemus quinquarius, 169 mm SLb)

Galeoides decadactylus, 250 mm SL

f)

Polydactylus approximans, 250 mm SL

c)

Leptomelanosoma indicum, 215 mm SL

g)

Polydactylus bifurcus, 144 mm SL

d)Parapolynemus verekeri, 72 mm SL

h)

Polydactylus longipes, 159 mm SL

PLATE IIIa)

Polydactylus macrochir, 200 mm SLe)

Polydactylus mullani, 145 mm SLb)

Polydactylus macrophthalmus, 88 mm SL

f)

Polydactylus multiradiatus, 170 mm SL

c)

Polydactylus malagasyensis, 140 mm SL

g)

Polydactylus nigripinnis, 138 mm SL

d)

Polydactylus microstomus, 101 mm SL

h)

Polydactylus octonemus, 135 mm SL

PLATE IVa)

Polydactylus oligodone)

Polydactylus plebeius, 46 mm SLb)

Polydactylus opercularis, 250 mm SL

f)

Polydactylus plebeius, 56 mm SL

c)

Polydactylus persicus, 155 mm SL

g)

Polydactylus plebeius, 169 mm SL

d)

Polydactylus plebeius, 12 mm SL

h)

Polydactylus plebeius, 248 mm SL

PLATE Va)

Polydactylus quadrifilise)Polydactylus sextarius, 164 mm SLb)

Polydactylus sexfilis, 456 mm SL

f)

Polydactylus sextarius, 120 mm SL

c)

Polydactylus sexfilisg)Polydactylus siamensis, 174 mm SL

d)

Polydactylus sexfilish)Polydactylus virginicus

PLATE VIa)

Polynemus aquilonaris, 86 mm SLe)

Polynemus melanochir dulcis, 135 mm SLb)

Polynemus dubius, 184 mm SL

f)

Polynemus melanochir melanochir, 134 mm SLc)

Polynemus hornadayi, 170 mm SLg)

Polynemus multifilisd)Polynemus kapuasensis, 127 mm SL

h)Polynemus paradiseus, 155 mm SL

PLATE Ia) Eleutheronema rhadinum
152 mm SL, Linkou, Taipei, Taiwan Province of China(K.-T. Shao)b) Eleutheronema tetradactylum
121 mm SL, Kuala Lumpur, Malaysia(H. Ida)c) Eleutheronema tridactylum
180 mm SL, Jakarta, Java, Indonesia(H. Motomura)d) Filimanus heptadactyla
90 mm SL, Sarawak, Malaysia(H. Ida)e) Filimanus hexanema
82 mm SL, Serang, Java, Indonesia(H. Ida)f) Filimanus perplexa
110 mm SL, Indonesia(CSIRO)g) Filimanus sealei
124 mm SL, Mindanao, Philippines(H. Motomura)h) Filimanus similis
98 mm SL, Chennai, India(H. Motomura)PLATE IIa) Filimanus xanthonema
62 mm SL, Lombok, Indonesia(S. Kimura)b) Galeoides decadactylus
250 mm SL, Cape Verde, Sal Island(P.N. Duarte)c) Leptomelanosoma indicum
215 mm SL, Singapore(H. Motomura)d) Parapolynemus verekeri
72 mm SL, South Alligator River, Australia(H. Motomura)e) Pentanemus quinquarius
169 mm SL, Senegal(H. Motomura)f) Polydactylus approximans
250 mm SL, Panama Bay, Panama(G.R. Allen)g) Polydactylus bifurcus
144 mm SL, Lombok, Indonesia(S. Kimura)h) Polydactylus longipes
159 mm SL, Mindanao, Philippines(H. Motomura)PLATE IIIa) Polydactylus macrochir
200 mm SL, Irian Jaya, Indonesia(M. Allen)b) Polydactylus macrophthalmus
88 mm SL, Indonesia(H. Motomura)c) Polydactylus malagasyensis
140 mm SL, east coast of Africa(NRIFSF)d) Polydactylus microstomus
101 mm SL, Ambon, Indonesia(S. Kimura)e) Polydactylus mullani
145 mm SL, Arabian Sea, India(H. Motomura)f) Polydactylus multiradiatus
170 mm SL, Queensland, Australia(J.W. Johnson)g) Polydactylus nigripinnis
138 mm SL, Arafura Sea(S. Kimura)h) Polydactylus octonemus
135 mm SL, off Texas(H. Motomura)PLATE IVa) Polydactylus oligodon
Northern coast of South America(F. Cervigón)b) Polydactylus opercularis
250 mm SL, Panama Bay, Panama(D.R. Robertson)c) Polydactylus persicus
155 mm SL, Kuwait Bay, Kuwait(H. Motomura)d) Polydactylus plebeius
12 mm SL, Ryukyu Islands, Japan(H. Kobayashi)e) Polydactylus plebeius
46 mm SL, Lombok, Indonesia(K. Matsuura)f) Polydactylus plebeius
56 mm SL, Lombok, Indonesia(K. Matsuura)g) Polydactylus plebeius
169 mm SL, Miyazaki, Japan(H. Motomura)h) Polydactylus plebeius
248 mm SL, Malindi, Kenya(J.E. Randall)PLATE Va) Polydactylus quadrifilis
Bakau, Gambia(J. Jensen)b) Polydactylus sexfilis
456 mm SL, Izu Islands, Japan(H. Senou)c) Polydactylus sexfilis
Coral Sea, Australia(G.R. Allen)d) Polydactylus sexfilis
Hawaii(J.E. Randall)e) Polydactylus sextarius
164 mm SL, Miyazaki, Japan(H. Motomura)f) Polydactylus sextarius
120 mm SL, Gulf of Thailand(H. Motomura)g) Polydactylus siamensis
174 mm SL, Songkhla, Thailand(H. Motomura)h) Polydactylus virginicus
Northern coast of South America(F. Cervigón)

PLATE VIa) Polynemus aquilonaris
86 mm SL, Chao Phraya River, Thailand(M.H. Sabaj)b) Polynemus dubius
184 mm SL, Sumatra, Indonesia(H. Motomura)c) Polynemus hornadayi
170 mm SL, Sarawak, Malaysia(H. Motomura)d) Polynemus kapuasensis
127 mm SL, Kapuas River, Indonesia(M. Kottelat)e) Polynemus melanochir dulcis
135 mm SL, Lake Tonle Sap, Cambodia(H. Motomura)f) Polynemus melanochir melanochir
134 mm SL, Sabah, Malaysia(H. Ida)g) Polynemus multifilils
Thailand(E. Schraml)h) Polynemus paradiseus
155 mm SL, Calcutta, India(H. Motomura)