iFishes of Terengganu
East coast of Malay Peninsula, Malaysia

ii

iii

Edited by Mizuki Matsunuma, Hiroyuki Motomura, Keiichi Matsuura,
Noor Azhar M. Shazili and Mohd Azmi Ambak
Photographed by Masatoshi Meguro and Mizuki Matsunuma

iv

Copy Right © 2011 by the National Museum of Nature and Science, Universiti Malaysia Terengganu and Kagoshima
University Museum
All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means without
prior written permission from the publisher. Copyrights of the specimen photographs are held by the Kagoshima University Museum.
For bibliographic purposes this book should be cited as follows:
Matsunuma, M., H. Motomura, K. Matsuura, N. A. M. Shazili and M. A. Ambak (eds.). 2011 (Nov.). Fishes ofTerengganu – east coast of Malay Peninsula, Malaysia. National Museum of Nature and Science, Universiti Malaysia
Terengganu and Kagoshima University Museum, ix + 251 pages.
ISBN 978-4-87803-036-9
Corresponding editor: Hiroyuki Motomura (e-mail: motomura@kaum.kagoshima-u.ac.jp)

v

Preface
Tropical seas in Southeast Asian countries are well known for their rich fish diversity
found in various environments such as beautiful coral reefs, mud flats, sandy beaches,mangroves, and estuaries around river mouths. The South China Sea is a major water bodycontaining a large and diverse fish fauna. However, many areas of the South China Sea,
particularly in Malaysia and Vietnam, have been poorly studied in terms of fish taxonomy
and diversity. Local fish scientists and students have frequently faced difficulty when trying to identify fishes in their home countries.
During the International Training Program of the Japan Society for Promotion of Science(ITP of JSPS), two graduate students of Kagoshima University, Mr. Mizuki Matsunuma(the first editor of this book) and Mr. Masatoshi Meguro, were provided with the opportunity to survey marine fishes along the east coast of peninsular Malaysia at the Institute of
Oceanography and Environment (INOS), Universiti Malaysia Terengganu (UMT). Theirfaunal study succeeded through cooperative activities between Malaysian and Japaneseichthyologists, and resulted in this field guide, “Fishes of Terengganu – East coast of Malay Peninsula, Malaysia”, that includes taxonomic accounts with color photographs of 441
species belonging to 276 genera of 108 families. This field guide could not have been completed without the help of Prof. Noor Azhar bin Mohamed Shazili, Deputy Vice-Chancellor(Academic and Internationalization) of UMT, Dr. Zainudin bin Bachok, Deputy Directorof INOS, Mr. Yusri bin Yusuf, Lecturer of INOS and Mrs. Siti Tafzilmeriam bt. Sheikh Abdul Kadir, Research Officer of South China Sea Reference Center & Repository at INOS.
Funds to publish this field guide were provided through grants from UMT and the National
Museum of Nature and Science.

Keiichi Matsuura
Director of Department of Zoology
National Museum of Nature and Sciencevi

PrefaceThis field guide to the fishes of Terengganu was intended to assist local fisheries biologists in the identification of fishes in the coastal zone of Malaysia and the publicationof this book is timely as there has always been a shortage of such guides in this region.
This publication is the culmination of collaborative efforts of scientists from Universiti
Malaysia Terengganu and Kagoshima University, and is another significant milestone in
the long history of collaboration between the two institutions. This effort would not havebeen possible without the support of Kagoshima University and their students who spent
many months in Terengganu, and the support of the Institute of Oceanography and Environment, Universiti Malaysia Terengganu for providing the laboratories, equipment andlogistical support. Our sincere gratitude also goes to scientists from University Malaysia
Sabah and Universiti Kebangsaan Malaysia for their chapter contributions. Universiti Malaysia Terengganu especially appreciates the effort of Prof. Hiroyuki Motomura in leading
the study in Malaysia from the very beginning.
This work shows that taxonomic studies of fishes in this region still have important
contributions to be made, as a number of new species were identified in the coastal zone.
We hope this field guide will be useful in assisting scientists in fish identification in the
Malaysian context.

Noor Azhar bin Mohamed Shazili
Deputy Vice-Chancellor (Academic and International Affairs)
Universiti Malaysia Terengganu

vii

Editors
Mizuki Matsunuma – The United Graduate School of Noor Azhar bin Mohamed Shazili – Academic & InterAgricultural Sciences, Kagoshima University, 1-21-24
Korimoto, Kagoshima 890-0065, Japan (e-mail: k1139853@
kadai.jp)

national Affairs, Universiti Malaysia Terengganu, 21030
Kuala Terengganu, Malaysia (e-mail: nazhar@umt.edu.my)

1-21-30 Korimoto, Kagoshima 890-0065, Japan (e-mail:motomura@kaum.kagoshima-u.ac.jp)

Food Science, Universiti Malaysia Terengganu, 21030
Kuala Terengganu, Malaysia (e-mail: ambak@umt.edu.
my)Mohd Azmi bin Ambak – Department of Fisheries SciHiroyuki Motomura – Kagoshima University Museum, ence and Aquaculture, Faculty of Agrotechnology and

Keiichi Matsuura – Department of Zoology, National

Museum of Nature and Science, 4-1-1 Amakubo, Tsukuba,
Ibaraki 305-0005, Japan (e-mail: matsuura@kahaku.go.jp)

Authors (alphabetical order)
Aziz Arshad – Department of Aquaculture, Faculty of Mizuki Matsunuma – See editors’ details
Agriculture, Universiti Putra Malaysia, 43400 UPM Serdang,Selangor D. E., Malaysia (e-mail: azizar.upm@gmail.com)

Mazlan Abdul Ghaffar – Marine Ecosystem ResearchCentre, School of Environmental and Natural Resource
Sciences, Faculty of Science and Technology, Universiti
Kebangsaan Malaysia, 43600 UKM Bangi, Selangor D.E.,
Malaysia (e-mail: magfish05@yahoo.com; mag@ukm.my)

Keiichi Matsuura – See editors’ details
Hiroyuki Motomura – See editors’ details
Koichi Shibukawa – Nagao Natural Environment Foundation, 3-10-10 Shitaya, Taito-ku, Tokyo 110-0004, Japan(e-mail: shibu@crux.ocn. ne.jp)

Hisashi Imamura – Laboratory of Marine Biology and Siti Tafzilmeriam bt. Sheikh Abdul Kadir – South
Biodiversity (Systematic Ichthyology), Faculty of Fisheries
Sciences, Hokkaido University, 3-1-1 Minato-cho, Hakodate,
Hokkaido 041-8611, Japan (e-mail: imamura@fish.hokudai.
ac.jp)

China Sea Reference Center & Repository, Institute of
Oceanography and Environment, Universiti Malaysia Terengganu, 21030 Kuala Terengganu, Terengganu, Malaysia(e-mail: sititafzil@ umt.edu.my)

Seishi Kimura – Fisheries Research Laboratory, Mie Yusri bin Yusuf – South China Sea Reference Center &University, 4190-172 Wagu, Shima, Mie 517-0703, Japan(e-mail: kimura-s@bio.mie-u.ac.jp)

B. Mabel Manjaji-Matsumoto – Borneo Marine ResearchInstitute, Universiti Malaysia Sabah, 88400 Kota Kinabalu,
Sabah, Malaysia (e-mail: mabel@ums.edu.my)

Repository, Institute of Oceanography and Environment,
Universiti Malaysia Terengganu, 21030 Kuala Terengganu,
Terengganu, Malaysia (e-mail: yusriyusuf@umt.edu.my);current address – Molecular Ecology Laboratory, Australian Rivers Institute, Nathan Campus, Griffith University,
Nathan, Queensland 4111, Australia

Photographers
Masatoshi Meguro – The United Graduate School of Mizuki Matsunuma – See editors’ details
Agricultural Sciences, Kagoshima University, 1-21-24
Korimoto, Kagoshima 890-0065, Japan (e-mail: fisheries_
kago_mm7190@hotmail.co.jp)

viii

Mizuki Matsunuma, PhD student of United Graduate School of Agricultural
Sciences, Kagoshima University

Keiichi Matsuura, Director of Department of
Zoology, National Museum of Nature and Science

Masatoshi Meguro, PhD student of
United Graduate School of Agricultural
Sciences, Kagoshima University (middle
in front row), Siti Tafzilmeriam bt.
Sheikh Abdul Kadir, Research Officerof Institute of Oceanography and Environment (INOS), Universiti Malaysia
Terengganu (on extreme right in frontrow) and Meii bt. Mohamad Norizam,
lecturer of INOS (between Masatoshi andSiti) at Fish Taxonomy Workshop in Setiu
Wetland, 2008

Left to right: Hiroyuki Motomura,
Professor of Kagoshima University
Museum, Noor Azhar bin Mohamed
Shazili, Deputy Vice-Chancellor (Academic and International Affairs) of
Universiti Malaysia Terengganu (UMT),
Aziz bin Deraman, Vice-Chancellor of
UMT, Mohd Azmi bin Ambak, Professor of Department of Fisheries Scienceand Aquaculture, UMT and Yahaya bin
Ibrahim, Deputy Vice-Chancellor(Student Affairs and Alumni) of UMT
Boldfaced names: editors and/or authors of this bookix

Contents
Introduction.............................................................1
Hemiscyllidae...........................................................8
Stegostomatidae.....................................................10
Scyliorhinidae.........................................................11
Carcharhinidae......................................................12
Sphyrnidae.............................................................14
Narkidae.................................................................15
Rhynchobatidae.....................................................16
Dasyatidae..............................................................17
Gymnuridae...........................................................21
Myliobatidae..........................................................22
Elopidae..................................................................24
Megalopidae...........................................................25
Albulidae................................................................26
Ophichthidae..........................................................27
Muraenesocidae.....................................................29
Nettastomatidae.....................................................30
Pristigasteridae......................................................31
Engraulidae............................................................32
Chirocentridae.......................................................36
Clupeidae................................................................37
Chanidae.................................................................40
Plotosidae................................................................41
Ariidae....................................................................42
Synodontidae..........................................................44
Carapidae...............................................................46
Ophidiidae..............................................................47
Antennariidae........................................................48
Mugilidae................................................................49
Atherinidae.............................................................52
Exocoetidae............................................................53
Hemiramphidae.....................................................54
Belonidae................................................................55
Aplocheilidae..........................................................56
Holocentridae.........................................................57
Syngnathidae..........................................................58
Fistulariidae...........................................................59
Centriscidae...........................................................60
Dactylopteridae......................................................61
Apistidae.................................................................63
Scorpaenidae..........................................................64
Tetrarogidae...........................................................66
Aploactinidae.........................................................68
Synanceiidae...........................................................69
Platycephalidae......................................................72
Ambassidae............................................................76
Latidae....................................................................78
Serranidae..............................................................79
Pseudochromidae...................................................85
Priacanthidae.........................................................86
Apogonidae.............................................................87
Sillaginidae.............................................................93
Coryphaenidae.......................................................95
Rachycentridae......................................................96
Echeneidae.............................................................97

Carangidae.............................................................98
Menidae................................................................108
Leiognathidae.......................................................109
Lutjanidae............................................................116
Caesionidae...........................................................119
Gerreidae..............................................................121
Haemulidae..........................................................124
Nemipteridae........................................................127
Lethrinidae...........................................................132
Sparidae................................................................135
Polynemidae.........................................................136
Sciaenidae.............................................................138
Mullidae................................................................142
Pempheridae........................................................145
Monodactylidae...................................................146
Toxotidae..............................................................147
Kyphosidae...........................................................148
Drepaneidae.........................................................149
Chaetodontidae....................................................151
Pomacanthidae.....................................................153
Terapontidae........................................................154
Cepolidae..............................................................156
Pomacentridae.....................................................157
Labridae...............................................................169
Scaridae................................................................176
Champsodontidae................................................179
Pinguipedidae.......................................................180
Uranoscopidae.....................................................181
Tripterygiidae......................................................182
Blenniidae.............................................................183
Callionymidae......................................................189
Eleotridae.............................................................191
Gobiidae................................................................193
Ptereleotridae.......................................................210
Ephippidae...........................................................211
Scatophagidae......................................................213
Siganidae..............................................................214
Acanthuridae.......................................................217
Sphyraenidae.......................................................218
Trichiuridae..........................................................219
Scombridae...........................................................220
Stromateidae........................................................223
Psettodidae...........................................................225
Citharidae.............................................................226
Paralichthyidae....................................................227
Bothidae................................................................230
Samaridae.............................................................232
Soleidae.................................................................233
Cynoglossidae.......................................................235
Triacanthidae.......................................................237
Balistidae..............................................................238
Monacanthidae....................................................240
Tetraodontidae.....................................................243
Diodontidae..........................................................245
Scientific Name Index..........................................247

x1Introduction

By Mizuki Matsunuma and Hiroyuki Motomura
The South China Sea lies in the tropical zone of thewestern Pacific Ocean, off the southeast corner of the
Asian continent, and is known for both its high productivity and the rich diversity of plants and animals. Randall and Lim (2000) listed at least 3,365 species of marine
fishes from the South China Sea. Recently, several pictorial books of fishes in Malaysian waters were published.
Mohsin and Ambak (1996) reported 710 species of marine
fishes from Malaysian waters and adjacent seas. Adrim
et al. (2004) recorded 430 marine fish species from theAnambas and Natuna islands on the Sunda Shelf between
the Malay Peninsula and Borneo in the South China Sea.
More recently, Ambak et al. (2010) estimated 2,243 fish
species occurring in Malaysian waters.
Terengganu, a State of Malaysia, is located on the east
coast of the Malay Peninsula and faced to the South China
Sea. The coastal environment of Terengganu is characterized by having long sandy beaches with only a few rocky
or coral reefs. This is because it is strongly influenced by
the large amount of fresh water discharge from two large
rivers, e.g., the Terengganu and Dungun rivers. These rivers have wide estuaries with mud flats and mangroves. In
particular for coral reefs, these ecosystems are well developed on islands, i.e., Bidong and Redang islands, which
are distant from the influences of river discharge, turbidity, and siltation. These marine and coastal habitats area vital support for a vast variety of marine and estuarine
biota, including fishes.
We carried out field surveys of fishes in Terengganu in
2008–2009 to produce a field guide for marine and estuarine fishes of the area. As a result of the surveys, 441 marine
and estuarine fish species of 108 families were recorded.
They make up around 13 % of over 3,365 fish species recorded by Randall and Lim (2000) from the South China Sea.
Among these, four species including Dasyatis parvonigra
Last & White, 2008 (family Dasyatidae), Larimichthys
crocea (Richardson, 1846) (Sciaenidae), Iniistius trivittatus (Randall & Cornish, 2000) and Leptojulis lambdastigma Randall & Ferraris, 1981 (Labridae) are recorded forthe first time in the east coast of the Malay Peninsula[the latter two species were reported by Matsunuma et al.
(2011)]. This book is produced based on the above surveys
and cover most of the diversity of coastal fishes occurringin marine and estuarine habitats, and commercial fishes
available at fish markets and ports in Terengganu. Species
reported in this book are based on the collected specimens
and are deposited in the South China Sea Natural History
Museum at Institute of Oceanography, Universiti Malaysia Terengganu and the Kagoshima University Museum
to make them available for future scientific studies. We
intend this book to be useful for the study and research ofichthyology and fishery science by researchers, students
and local government administrators.
This project was supported by the International Training Program of the Japan Society for Promotion of Science(ITP of JSPS), Universiti Malaysia Terengganu (UMT),

the National Museum of Nature and Science (NSMT), andthe Kagoshima University Museum (KAUM).

Collection Sites
Surveys of fishes in Terengganu were made during two
periods from 2008 to 2009. The first survey, carried out
from September to November 2008, mainly focused on
fishes inhabiting shallow coastal waters of coral or rocky
reefs, mangroves, and estuaries. The collection of fishes inthe survey was made at several localities in Terengganu,
including Bidong Island, Setiu, Merang, Cendering, Dungun,
and Kemaman. Coral and rocky reef associated fishes
were obtained at Bidong Island. Fishes occurring in mangrove-lined estuary were obtained in Setiu. Cendering,
located just south of Kuala Terengganu, has rocky coasts;
shallow water fishes found in tide pools were collected
here. Fishes were also caught at the beaches and estuaries
near Universiti Malaysia Terengganu and at Merang, Dungun, and Kemaman. Fishes were collected mainly by using hand-net and fence-net while SCUBA diving or snorkeling. Cast-net, small bottom trawl, and gill nets werealso used for collecting in estuaries and mangroves.

Collection Sites

2
The second survey observed from December 2008 to January 2009, highlighted fishes in commercial catches, obviously by trawlers, at the port or those of artisanal fisheries(mainly by traps, hand lines, and gill nets) sold at local fish
markets in Kuala Terengganu and Kampung Raja. A small
number of fishes captured around Redang Island were obtained from a fishing port at Kampung Raja.
The systematic arrangement of families generally follows Nelson (2006). Species in families are arranged inalphabetical order by species name. Each species record
was compiled from voucher specimens. Records of the Zebra Shark (Stegostomatidae), Stegostoma fasciatum (Hermann, 1783), and the Blotched Fantail Stingray (Dasyatidae), Taeniurops meyeni (Müller & Henle, 1841), arebased on photographs; these photographs are cataloged at
the Fish Image Database of Kagoshima University Museum (KAUM–II). On figure legends of photographs of fish
specimens which were obtained at ports or fish markets
during the second survey, sampling localities are shown
as “off Terengganu”; “KT” and “KR” in parentheses indicates the specimens were obtained in Kuala Terengganu
and Kampung Raja, respectively. Specimens obtained
during the surveys are deposited at the South China Sea
Natural History Museum at Institute of Oceanography,
Universiti Malaysia Terengganu (UMTF), Malaysia andthe Kagoshima University Museum (KAUM), Japan.

Bidong Island

Coral reefs around Bidong Island

Fish market in Kuala Terengganu

Mangroves at Setiu

Fish landing port in Kuala Terengganu

Cast netting in estuary at Setiu

3

Methods of Measurements and Counts
Methods of measurements and counts generally follow
Nakabo (2002). For fin formulae, the number of spinousand soft fin rays are described by Roman numerals (I, II,
III, ......) and Arabic numerals (1, 2, 3, ......), respectively.
The unbranched soft rays are sometimes expressed insmall Roman numerals (i, ii, iii, ......). Spinous fin rays are
generally called spines. In the case of the dorsal or anal(sometimes pectoral or pelvic) fins containing spine andsoft rays, the number of spines and soft rays are separated
by a comma. When the dorsal (or anal) fin consists of
two or more fins (i.e. first dorsal fin, second dorsal fin,......), each fin is separated by a “+” sign. Gill rakers on
the first gill arch on the right side of the body are used forcounting. Number of gill rakers on the upper and lower
limbs are separated by a “+” sign. When present, the one
or more gill rakers between the limbs are included in thelower limb counts. Number of vertebrae includes the urostyle. Counts of abdominal and caudal vertebrae are separated by a “+” sign.

A – number of anal-fin rays.
BR – number of branchiostegal rays
D – number of dorsal-fin rays.
DW – disc width: extremities of the left and right pectoral
fins.
DPC – number of dorsal procurrent caudal-fin rays.
FL – fork length: linear distance from most anterior pointof head to bottom of concave margin of caudal fin.

GR – number of gill rakers.
LGR – number of gill rakers on lower limb.
LL – number of lateral line scales: number of scales onlateral line from the scale behind the posttemporal tothe caudal-fin base.
LLp – number of pored scales on lateral line: only the
number of pored scales on lateral line is counted.
LR – number of scales in longitudinal row: number ofscales in longitudinal row from the dorsal end of theopercular membrane to the caudal-fin base.
MP – number of mandibular pores.
P1 – number of pectoral-fin rays.
P2 – number of pelvic-fin rays.
PDS – number of predorsal scales: number of scales on
the dorsal midline from the origin of dorsal fin forward to occipital region.
PLp – preanal lateral-line pores: number of pores along
lateral line behind gill opening to just above the anus.
SL – standard length: linear distance from most anterior
point of upper jaw (or snout) with mouth closed to caudal fin base (posterior end of hypurals, roughly wherefold formed by bending caudal fin).
TL – total length: greatest linear distance between
most anteriorly projecting part of head with mouthclosed and farthest tip of caudal fin when caudal rayssqueezed together. All unspecified lengths are assumed to be total lengths.
V – number of vertebrae.

first dorsal fin

second dorsal fin

anal fintailgill slits
spiracle

pectoral fin

pelvic fin

eye
snout

disc width

Body parts of elasmobranchs

caudal fin

subterminal notch

venomous spine

4

lateral line
spinous portion of dorsal fin
opercle
snout

nape

soft-rayed portion of dorsal fin

eyecaudal peduncle

upper jaw

lower jaw

cheekpreopercle

caudal fin
anal finpectoral fin

pelvic fin

caudal-fin base

Body parts of bony fish

1st dorsal fin

2nd dorsal finscutesadipose eyelid

lateral line

finlets
detached anal-fin spines

Body parts of carangid

upper limb

gill raker

lower limb
gill filament
Gill arches of carangid (left) and sparid (right)

5

Acknowledgements
We express our thanks to Yusuf Shuaib bin Ibrahim,
Meii bt. Mohamad Norizam, Joseph Anak Bidai, Che
Mohd. Kamarul Anuar bin Che Abdullah, Subarjo bin
Merehojono, Nur Fatiha bt. Che Wail, Ahmad Nazila
bin Ali, Harith Nasutieon bin Abdul Rahman, Noratikah
bt. Abd. Manaf, Azri bin Muhamad, Nurhayati bt. Mat
Semawi, Norhayati bt. Hasan, Hazrul bin Ahmad, Johari
bin Mohd. Nor, Mohd. Sharol bin Ali, Zakari bin Ismail,
Asmadi bin Salleh, Mat Alwi bin Omar, Imran bin Mohamad, Abd. Rashid bin Omar, Syed Ahmad Rizal bin
Tn. Nek, Muhamad Shukri bin Shafie, Kasim bin Muda,
Adnan bin Abd. Ghani, Mohd Rusli bin Ngah and Mohd
Kamaruzaman bin Mohd Ariffin (the staff of UMT),
Eiji Suzuki (Kagoshima University), and Shuhei Nishida(Project Leader of JSPS Asian Core Program – Coastal
Ecosystems in Southeast Asia; Atmosphere and Ocean
Research Institute of the University of Tokyo) for their
special support and cooperation. We are grateful to thefollowing persons for providing information on species
identifications: Yukio Iwatsuki (University of Miyazaki),
Tetsuo Yoshino (formerly University of Ryukyus), Kunio
Sasaki and Hiromitsu Endo (Kochi University), Hiroshi
Senou (Kanagawa Prefectural Museum of Natural History), Atsunobu Murase (Tokyo University of Marine Science and Technology), Gota Ogihara, Masahiro Yamashita, Tomohiro Yoshida, Hiroki Iwatsubo, and Harutaka
Hata (KAUM). The first editor and first photographer are
deeply indebted to late Mohamad bin Taib, Tapsilah bt.
Mohd Nordin, Siti Tafsil Raudah bt. Sheikh Abdul Kadirand Siti Tafsir Mazlina bt. Sheikh Abdul Kadir for theirgenerous assistance during their stay in Terengganu. This
work was supported in part by “International Training
Program to Protect Diversity of Bioresources in the Tropical Area” in Kagoshima University, from JSPS.

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6
Gill, A. C. 2004. Revision of the Indo-Pacific dottybackfish subfamily Pseudochrominae (Perciformes: Pseudochromidae). Smithiana Monogr., (1): 1–213, pls. 1–12.
Gon, O. 1993. Revision of the cardinalfish genus
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Hidaka, K., Y. Iwatsuki, and J. E. Randall. 2008. A reviewof the Indo-Pacific bonefishes of the Albula argentea
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Kimura, S., K. Ikejima, and Y. Iwatsuki. 2008b. Eubleekeria
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Kimura, S., H. Motomura, and Y. Iwatsuki. 2008c.
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7
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a new eeltail catfish (Siluriformes: Plotosidae) fromJapan. Bull. Natl. Mus. Nat. Sci., Ser. A, Suppl., 2: 1–11.

8

HEMISCYLLIIDAE

Longtail Carpetsharks (Bamboosharks)
By B. Mabel Manjaji-Matsumoto

Small sharks; maximum size toabout 130 cm; around 70 cm in most
species. Body cylindrical, elongated;precaudal tail somewhat longer thanhead and trunk. Nostrils with barbels; nasoral grooves and circumnarial grooves present. Mouth small,
subterminal (located well anterior toeye); teeth small, not blade-like,monocuspid, similar in both jaws.
Eyes without nictitating eyelids.
Spiracles large, situated behind andbelow eyes. Five small gill slits, theposterior 3 over the pectoral-fin base.
Two dorsal fins, without spines; firstdorsal-fin origin above or posterior
to pelvic-fin base; second dorsal fin
subequal and similar in shape as thefirst, well anterior to anal-fin origin;
anal-fin base connected with caudal
fin, with a deep notch between thefins; caudal fin strongly asymmetrical, with a strong subterminal notch.
Color: adults dark gray to lightbrown, plain; head and body withspiracles large
subterminal
notch
head and trunk

precaudal tailnumerous dark spots and faint bands
in some species. Juveniles more intensely spotted, usually with blackedged saddlemarkings.
Remarks: occurring in tropicaland subtropical coastal areas, from
Madagascar to Japan and Australia.
Bottom dwellers. Feed on polychaetes,
crustaceans and small fishes. Oviparous.
Similar families occurring in thearea: Hemiscylliidae is distinguishedfrom other Indo-Pacific orectolobiform
shark families in having an almost
cylindrical body, a long precaudal tail(longer than head and body), second
dorsal fin located anterior to anal fin,and anal-fin base connected with lower
lobe of caudal fin. Orectolobidae –nasal barbels numerous, branched;head and body with elaborate variegated pattern. Ginglymostomatidae
and Stegostomatidae – no circumnarial
grooves; caudal fin elongate, bladelike; precaudal tail shorter than head
and trunk. Scyliorhinidae (carcharhiniform shark) – eyes with weakly differentiated nictitating lower eyelids;
second dorsal fin located posterior toanal fin; anal-fin base not connectedwith lower lobe of caudal fin; headand body with light spots, dark
blotches, bars, and saddles.

Chiloscyllium griseum
Müller & Henle, 1839
Gray Bambooshark
Body elongated, moderately slender,
without lateral ridges. Nostrils subterminal on snout; mouth closer to eyes
than snout tip. Dorsal fins smaller thanpelvic fins, without free posterior projection; interspace between dorsal finsusually more than 9.3% TL; first dorsalfin height more than 6.6% TL; second
dorsal-fin height usually more than
5.8% TL. Color: pale brown to graybrown dorsally, cream ventrally; with
12 or 13 saddle markings in juveniles,
markings not outlined in black. Size:maximum length 74 cm. Distribution:
Indo-West Pacific, from the Arabian
Sea to New Guinea, north to Japan.
However, not recorded from Borneo.
Remarks: reef associated; inshore
bottom dweller; marketed fresh.
Chiloscyllium griseum, KAUM–I. 16868, 54.0 cm TL
off Terengganu (KT), 5 Dec. 2008

9

Chiloscyllium punctatum
Müller & Henle, 1838
Brownbanded Bambooshark
Body elongated, moderately stout,
without lateral ridges. Nostrils subterminal on snout; mouth closer to eyes
than snout tip. Dorsal fins larger thanpelvic fins, with free posterior projections, posterior margins of dorsal finsstraight or distinctly concave; anal-finbase shorter than caudal-fin lobe
between lower origin and subterminal
notch. Color: plain dark gray or with
faint grayish bands; with dark transverse bands usually with a scatteringof small dark spots in juveniles. Size:maximum length ca. 1 m. Distribution: Indo-West Pacific, from India tonorthern Australia, north to Japan.
Remarks: inshore bottom dweller;
very tenacious of life, can survive outof water for a long period (half a day);marketed fresh, or as fillets in fish
markets of Sabah.

Chiloscyllium punctatum, KAUM–I. 16869, 41.0 cm TL
off Terengganu (KT), 5 Dec. 2008

10

STEGOSTOMATIDAE
Zebra Shark

By Seishi Kimura and B. Mabel Manjaji-MatsumotoThis family is represented by asingle species; see the following species account.
Similar families occurring in
the area: Stegostomatidae is distinguished from other Indo-Pacificorectolobiform shark families in having a very long caudal fin, almost ahalf of total length. Hemiscylliidae –origin of second dorsal fin well anterior to origin of anal fin. Ginglymostomatidae – spiracles much smaller
than eyes; body without lateral ridge.
Rhinco-dontidae – head very wideand flattened; mouth very wide, almost terminal on the head.

spiracles large

very long caudal fin
dermal ridges

Stegostoma fasciatum(Hermann, 1783)
Zebra Shark
A large shark; maximum size toabout 235 cm. Body moderately stout
with prominent ridges on dorsal surface and sides; caudal fin very long,
blade-like. Nostrils close to anterior
margin of snout, with barbels; connected with mouth by nasoral grooves;no circumnarial grooves and folds.
Mouth small, located well anterior toeye; teeth small, tricuspid. Eyes without nictitating eyelids. Spiracle similarin size to eye. Two dorsal fins, without
spines; first dorsal-fin origin anterior
to pelvic-fin origin; second dorsal fin
smaller than the first, located just posterior to the first and anterior to analfin origin; anal fin present, rounded;caudal fin very long, almost as long as
the body, with a deep subterminal
notch but with the lower lobe hardly
developed. Color: adults yellowish
brown with numerous, dark brownspots. Juveniles darker, head and body

Stegostoma fasciatum, KAUM–II. 39, ca. 100 cm TL
off Terengganu (KT), 19 Jan. 2009

dark brown or blackish, with vertical ical and subtropical coastal areas.
yellow bars, spots and reticulations. Feeds usually on mollusks and small
Distribution: Indo-West Pacific, from fishes. Oviparous.
South Africa to New Caledonia, north (S. Kimura & B. M. Manjaji-Matsumoto)to Japan. Remarks: occurring in trop-

11

SCYLIORHINIDAE
Catsharks

By B. Mabel Manjaji-Matsumoto
Small sized sharks; maximum size
to about 70 cm; around 45 cm inmost species. Body slender, elongated; tail very slender. Head small,depressed; snout short to moderately
long; eyes with weakly differentiatednictitating lower eyelids; nostrils
with or without nasoral or circumnarial grooves; mouth moderately
large, subterminal (located beloweye); teeth very small, numerous,
with a single medial cusp and usually
1 or more cusplets on each side near
the center of mouth; 5 small gill slits,
the posterior 2 over the pectoral-finbase. Two dorsal fins, without spines,
both fins of similar shape; first dorsalfin origin above or posterior to pelvicfin base; second dorsal-fin origin posterior to anal-fin origin; caudal fin
strongly asymmetrical, with a strong
subterminal notch. Caudal peduncle
without lateral keels or precaudal
pits. Color: head and body gray,brown, yellowish, or black, often

first dorsal-fin origin behind pelvic-fin origin
caudal fin strongly
heteroceralcaudal peduncle
without pits and keels

mouth located beneath eyeswith light or dark spots and dark
blotches, bars and saddles.
Remarks: occurring in tropical andsubtropical areas. Bottom dwellers,
ranging from shallow coastal waters to
depths greater than 2,000 m. Feed onpolychaetes, crustaceans and small
fishes. Used for fishmeal, oil and lobster bait.
Similar families occurring in thearea: Scyliorhinidae is distinguishedfrom other Indo-Pacific carcharhiniform shark families in having a small

slender body, location of the last 2gill slits behind pectoral-fin origin,
the posterior position of the first dorsal
fin, and the absence of keels or precaudal pits on the caudal peduncle.
Also distinguishable from the superficially similar orectolobiform shark(Hemiscylliidae) by the presence of
weakly differentiated nictitating lower
eyelids; and anal-fin base not connected with lower lobe of caudal fin.

Atelomycterus marmoratus(Bennett, 1830)
Coral Catshark
Body elongated, very slender. Nostrils subterminal on snout; nasoral
grooves present; anterior nasal flaps
greatly expanded, reaching mouth.
Mouth large, located below eyes;
labial furrows very long. Dorsal finslarge, subequal in shape and size,
angled rearwards. Color: adults andadolescents with marbled appearance;head and body with numerous lightgray and white spots; saddles obsolete. Juveniles darker, head and bodydark gray with pale vertical lines,
black spots with pale margins along
dorsal body. Size: maximum length 70
cm. Distribution: tropical Indo-West
Pacific, from Pakistan to New Guinea
and southern China and Taiwan. Remarks: a common, inshore bottom
dweller, often found in crevices andholes on rocky reefs; oviparous; feed
mainly on bony fishes; marketed fresh,or as fillets in fish markets of Sabah.

Atelomycterus marmoratus, KAUM–I. 17214, 46.8 cm TL
off Terengganu (KT), 10 Jan. 2009

12

CARCHARHINIDAE

Whaler Sharks (Requiem Sharks)
By B. Mabel Manjaji-Matsumoto and Seishi Kimura

Small to large sharks; maximum
size 7.5 m; around 1–3 m in most species. Body cylindrical, weak lateralkeels on caudal peduncle in some
genera; precaudal tail much shorter
than head and trunk. Head conical todepressed; snout very short to long;spiracles large. Nostrils withoutbarbels, nasoral grooves or circumnarial grooves. Mouth usually large,
arched, and extending well behind
eyes; teeth small to large, blade-like,with a single cusp and cusplets variably developed; anterior teeth inupper jaw smaller than lateral teeth
and not separated from them bysmaller teeth. Eyes on sides of head,with developed nictitating lower
eyelid. Spiracles usually absent. Fivesmall to medium sized gill slits, thelast 1 to 3 behind pectoral-fin origin.
Two dorsal fins, without spines, thefirst larger than the second or subequal; first dorsal-fin base locatedabove interspace between pectoraland pelvic-fin bases; anal fin moderately large; caudal fin strongly asymmetrical, much less than a half oftotal length, with an undulated dorsal
margin, a strong subterminal notch,

caudal fin
strongly asymmetricaleyes with nictitating
membraneprecaudal pits

and a well-defined lower lobe; precaudal pits well developed. Intestine withscroll valve, lacking spiral valve. Color: head and body gray, yellowish, or
bluish dorsally, white or pale yellow
ventrally, sometimes with prominent
dark or light marking on fins.
Remarks: circumglobal in thetropic and temperate areas of Pacific,
Atlantic, and Indian Oceans. Marine,
occasionally in freshwater lakes orrivers. Typical predator, feed on almost
all aquatic animals. Dominant in tropicalwaters; the larger species are dangerous to people.
Similar families occurring in thearea: Carcharhinidae is distinguishedfrom other Indo-Pacific carcharhiniform shark families in having a character combination of a long snout,spiracles, upper teeth with cusplets,

lower teeth well differentiated fromuppers, long labial furrows, and second dorsal fin subequal to first dorsal
fin, its origin anterior to that of theslightly smaller anal fin. Scyliorhinidae – head and body small, slender,
location of the last 2 gill slits behindthe pectoral-fin origin, the posterior
position of the first dorsal fin, andthe absence of keels or precaudal pits
on the caudal peduncle. Triakidae– dorsal margin of caudal fin not undulated; precaudal pits absent; intestine with spiral valve. Hemigaleidae– intestine with spiral valve; spiracle
present but minute; upper teeth withstrong distal cusplets. Sphyrnidae– head hammer-shaped. Lamnidae(lamniform shark) – caudal peduncle
with large lateral keels.

Carcharhinus dussumieri(Müller & Henle, 1839)
Whitecheek Shark
Body moderately stout. Snout long,broadly parabolic. Upper teeth withstrongly oblique cusp, flanked on one
side by strong serrated cusplets; lower
teeth narrow, upright, no cusplets.
First dorsal fin moderately tall, broad,not falcate. Low interdorsal ridge usually present. Color: grayish; second
dorsal fin tip black (usually covering
about half of fin). Size: maximum
length at least 94 cm. Distribution:
tropical waters of the Indo-West Pacific,
from the Red Sea to northern Australia, New Guinea and Japan. Remarks:
a common demersal inshore species
from close inshore to about 170 m
depth. Viviparous. Used for its finsand meat but of limited value due toits small size.
(B. M. Manjaji-Matsumoto)

Carcharhinus dussumieri, KAUM–I. 17084, 45.4 cm TL
off Terengganu (KT), 28 Dec. 2008

13

Carcharhinus melanopterus(Quoy & Gaimard, 1824)
Blacktip Reef Shark
Body moderately stout. Snout very
short, broadly rounded, preoral length
subequal to internarial space. Upper
teeth with narrow oblique central cusp
and low basal cusplets; lower teeth
narrow, upright to oblique, finely serrated edges. First dorsal fin large, its
apex narrowly rounded, its origin over
inner margins of pectoral fins; second
dorsal fin high, its inner margin lessthan twice the fin height. Interdorsal
ridge absent. Color: yellow brown
dorsally, distinct pale stripe alongeach flank; first dorsal fin with a thickblack tip; caudal fin lower lobe withthick black tip; other fins often withsmaller black tips. Size: maximum
length at least 142 cm. Distribution:
tropical Indo-West Pacific and Central
Pacific, from southeastern Africa to
the Central Pacific Islands, and eastern
Mediterranean. Remarks: a common
reef shark, living in lagoons and nearfringes of reefs. Viviparous.
(B. M. Manjaji-Matsumoto)

Carcharhinus melanopterus, KAUM–I. 17237, 55.8 cm TL
off Terengganu (KT), 14 Jan. 2009

14

SPHYRNIDAE

Hammerhead Sharks
By B. Mabel Manjaji-Matsumoto

Medium- to large-sized sharks;
maximum size to about 600 cm;
around 450 cm in most species. Body
elongate, moderately slender. Head
broad, its anterior portion flattened
dorsoventrally and widely expanded
laterally in "hammer" form, withthe eyes at its outer edges; eyes withwell-developed nictitating lower
eyelids; teeth blade-like, monocuspid. Two dorsal fins, the first highand pointed, its base much shorter
than caudal fin and anterior to pelvichead hammer-shaped;
fin origin; caudal fin asymmetrical, eyes located at tips of lateral bladeswith a strong subterminal notch
and a small, but well-defined lower
lobe. Caudal peduncle not strongly
flattened dorsoventrally or widely
Remarks: cosmopolitan in coastal area: none. Not other shark family
expanded laterally, without longitu- tropical and warm temperate seas. has the characteristic hammer-shaped
dinal ridges but with precaudal pits. Viviparous.
head of the Sphyrnidae.
Color: gray to brassy.
Similar families occurring in the

Sphyrna lewini(Griffith & Smith, 1834)
Scalloped Hammerhead
Body elongate, laterally compressed.
Head broad, hammer-shaped, anterior
margin arched, indented at midline.
Upper teeth triangular, anteriorly upright, posteriorly oblique. First dorsal
fin tall, moderately falcate; second
dorsal fin short with long rear tip andweakly concave posterior margin.
Precaudal pit present, crescentshaped. Color: plain gray to olivaceous; pectoral fins tip gray, black
ventrally. Size: maximum length 350
cm. Distribution: cosmopolitan in all
tropical and warm temperate seas. Remarks: occurs over continental andinsular shelves and adjacent deep
water, to 275 m depth; feed mainly onbony fishes and cephalopods, but also
sharks and rays; potentially dangerous
to humans.

Sphyrna lewini, KAUM–I. 17006, 56.6 cm TL
off Terengganu (KT), 14 Dec. 2008

15

NARKIDAE
Sleeper Rays

Small batoids; maximum size
9–46 cm in total length. Head andgreatly enlarged pectoral fins fully
fused to form a strongly flattenedoval, circular, or pear-shaped disc;trunk short and stout, fully fused withenlarged pelvic fins; tail stout, sharklike, narrower than trunk. Five pairsof gill openings on ventral side ofdisc. Body entirely naked above andbelow, without dermal denticles orthorns. Large kidney-shaped electricorgans present at bases of pectoral
fins (visible through skin). Tail without stinging spine (stings), or electricorgans. Color: brownish or reddish
brown dorsally, white or brownish
ventrally; dorsal surface either plain
or with a few large dark spots or
blotches, paired white spots, andwhite side bands on the tail and pos-

By B. Mabel Manjaji-Matsumoto
no dorsal fin or a single dorsal fin

caudal fin large

terior pelvic bases.
Remarks: a small group of inshore
to deep-water batoids confined to thetemperate and tropical Indo-West Pacificfrom South Africa to Japan and Indonesia. Occur on the intertidal and subtidal zone to offshore on the outer
shelf and upper slope on soft bottom
down to 330 m.
Similar families occurring in thearea: Narcinidae – 2 dorsal fins; snout
supported by a broad, trough-shaped

rostral cartilage, nasal curtain broader
and shorter; snout longer, deep labial
groove present. Torpedinidae – 2 dorsal fins; snout truncate; pectoral disctransversely elliptical; mouth strongly
arched, without labial folds or groove.
Hypnidae – 2 dorsal fins; pectoral
disc longitudinally pear-shaped;
mouth strongly arched, without labial
folds or groove; teeth tricuspidate;tail, dorsal, and caudal fins verysmall.

Temera hardwickii
Gray, 1831
Finless Sleeper Ray
Disc greatly enlarged, its length(measured to cloaca) much longer
than tail. Eyes bulging, not embedded
in skin. Spiracle posterior to eye. Nostrils circular. Mouth small, straight.
No dorsal fins. Caudal fins about aslong as deep. Color: dorsally plainbrown; ventral surfaces of disc andpelvic fins with broad brownish margins. Size: maximum size dubiously
reported to about 46 cm, but mostly
less than 15 cm. Distribution: tropical Indo-West Pacific, from the Andaman Sea off Myanmar to Singapore,
and Vietnam. Remarks: demersal on
the continental shelf. Paired electricorgans in the disc are capable of delivering a moderate shock on direct contact.

Temera hardwickii, KAUM–I. 17198, 6.7 cm DW
off Terengganu (KR), 5 Jan. 2009

16

RHYNCHOBATIDAE
Wedgefishes

Large shark-like batoids; maximum size 270 cm. Head and greatly
enlarged pectoral fins fully fused
to form a small pectoral disc; trunk
thick, shark-like, tail long and stout.
Head narrow and depressed; snout
moderately elongated, angular andsupported by a stout rostral cartilage;eyes and spiracles dorsolateral onhead. Five pairs of gill openings onventral side of disc. Body covered
with small dermal denticles; enlarged
denticles present on dorsal surface on
orbits, and midline of trunk and tail.
Color: yellowish, brownish, graybrown, or greenish dorsally, whiteventrally; dorsal surface with small
to large white spots and sometimes
dark spots on pectoral fins.
Remarks: a small group of inshore

By B. Mabel Manjaji-Matsumoto
pectoral fin ends anterior to pelvic fin

snoutwedge-shapedorsal fins large, falcate

tropical batoids, from tropical West
Africa, and the Indo-West Pacific.
Feed on polychaetes, crustaceans andsmall fishes. Ovoviviparous.
Similar families occurring in thearea: Rhinidae – snout broadly rounded;no spiracular folds; mouth with prominent knobs and depressions; enlarged
thorns present on dorsal surface on

snout, orbits, shoulders, and midlineof trunk. Rhinobatidae – pectoral finswith origins in front of nostrils andwith free rear tips posterior to pelvicfin origin; lower lobe of caudal fin
short. Pristidae – snout with a rostral
saw; no thorns around eyes, on back,or on tail; dorsal surface plain, without spots or ocelli.

Rhynchobatus australiae
Whitley, 1939
Whitespotted Wedgefish
Snout bottle-shaped, constricted neartip; spiracles with two skin folds on
posterior margin; first dorsal fin lessthan 1.5 times height of second (inadults), its origin slightly posterior topelvic-fin origin. Color: disc gray or
brown with inconspicuous white margin; adults with a faint (sometimes
absent) black spot on each pectoral fin,spot diffused-edged in juveniles (lessthan 60 cm), diagonal row of threeequidistant white spots usually above
black pectoral spot. Size: maximum
size 270 cm. Distribution: tropicalIndo-West Pacific, from Taiwan tonorthern Australia, including the Philippines. Remarks: demersal on soft
bottoms near the coast and sand patcheson coral reefs, inshore to depths of atleast 60 m; feeds on bottom curstaceans and molluscs; marketed fresh,
fins prized for the sharkfin trade. Species listing in Rhynchobatidae follows
Last and Stevens (2009) and Compagno
and Last (2010). Listed in Rhinobatidae
by Nelson (1994), and in Rhinidae by
Compagno and Last (1999).
Rhynchobatus australiae, KAUM–I. 16870, 41.0 cm TL
off Terengganu (KT), 5 Dec. 2008

17

DASYATIDAE
Stingrays

Small to very large batoids; maximum size 5 m in total length, 2 m indisc width. Head, trunk, and greatly
enlarged pectoral fins fully fused toform a strongly flattened oval, circular,
or rhomboidal disc; tail moderately
depressed and or cylindrical, whiplike. Head not projecting from disc;eyes and spiracles dorsolateral onhead. Five pairs of gill openings onventral side of disc. Body usuallywith denticles, thorns, and tubercles
on the dorsal surface of disc and tail.
In most species, tail with 1–4 stinging spines (stings). Color: grayish
brown or brownish green dorsally,
paler ventrally; a few species withbluish, whitish, or blackish spots,
rings, or reticulations.
Remarks: occur in marine, estuarine, and fresh-water habitat in temperate and tropical areas worldwide.

By B. Mabel Manjaji-Matsumoto
head not projectingfrom disctail long, slender, whip-like,
without dorsal or caudal finsventral finfold

stinging spine (s) usually present

Similar families occurring in thearea: Plesiobatidae – tail stout, witha long leaf-like caudal fin. Rajidae –tail not whip-like, with small posterior
dorsal fins, and/or caudal fins; pelvic
fins subdivided. Narcinidae – stout
tail with dorsal and caudal fins. Gym-nuridae – disc extremely broad (morethan 1.5 times as broad as long),
tail very short (shorter than ½ discwidth). Myliobatidae – head distinct
from disc; pectoral apices angular,
acute.

Dasyatis parvonigra
Last & White, 2008
Dwarf Black Stingray
Disc rhomboidal with somewhat
angular apices; tail moderately long,
becoming much more slender beyond
sting; short, low dorsal skin fold on tailbeyond sting; long, low ventral skinfold on tail; thorns confined to central
disc (single row on midline with shorter row either side) dorsally (in adults);
no thorns on tail before sting. Color:
adults with disc dark brown to olivaceous dorsally, young disc brownish;disc dark edged (not yellowish) ventrally in young and adult; tail beyond
sting paler than anterior tail in adults,
a broad white segment near tail tip,
dorsally and ventrally in young. Size:maximum size 51 cm disc width. Distribution: known only from northwestern Australia, Indonesia and Malaysia. Remarks: demersal on insularand continental shelves in 60–125 m;
marketed fresh.
Dasyatis parvonigra, KAUM–I. 16871, 19.5 cm DW
off Terengganu (KT), 5 Dec. 2008

18

Dasyatis zugei(Müller & Henle, 1841)
Sharpnose Stingray
Disc with broadly rounded apices;tail relatively short, tapering in thickness beyond sting; weak dorsal skinfold on tail beyond sting; long, lowventral skin fold on tail; a few small
thorns along midline of disc dorsally;
row of small thorns on tail before stingin adults; snout very elongate; no oral
papillae in mouth. Color: disc brownish dorsally; disc pale or dark edged
ventrally. Size: maximum size 290 cm
disc width. Distribution: Indo-West
Pacific, from India to eastern Indonesia (Bali) and northwards to southern
Japan. Remarks: demersal on insularand continental shelves, to depths of atleast 40 m. viviparous, gives birth tolitters of 1–10 pups; feeds on bottom
crustaceans; marketed fresh.

Dasyatis zugei, KAUM–I. 16872, 15.7 cm DW
off Terengganu (KT), 5 Dec. 2008

Himantura gerrardi(Gray, 1851)
Whitespotted Whipray
Disc quadrangular; tail long, whiplike, no skin folds on tail; widely
spaced, granular denticles on central
disc (absent in small juveniles); central disc usually with 1–5 small thorns;
tail lacking thorns; sting situated anteriorly on tail. Color: disc with numerous white spots (rarely plain grayish
brown) dorsally; white ventrally; tailwith alternating light and dark bands(rarely faint). Size: maximum size 85
cm disc width. Distribution: widespread in the Indo-West Pacific, from
India to eastern Indonesia, north to
Taiwan. Remarks: demersal on soft
bottoms, from inshore to depths of atleast 60 m; feeds primarily on smallbottom crustaceans, but also small
fishes; marketed fresh, the skin isvaluable as leather.

Himantura gerrardi, KAUM–I. 16982, 23.8 cm DW
off Terengganu (KT), 12 Dec. 2008

19

Himantura walga(Müller & Henle, 1841)
Dwarf Whipray
Disc almost oval; tail short, notwhip-like (end bulbous in adult females); no skin folds on tail; narrow
band of flat denticles on central disc in
adults; mid-disc thorns absent or rudimentary; tail thorns very elongate,bases nearly half eye diameter inlength; sting situated anteriorly on tail;snout broadly triangular. Color: grayish or brownish dorsally (without apattern); whitish ventrally, sometimes
with a yellowish gray posterior margin. Size: maximum size 240 cm discwidth. Distribution: off Thailand,
Malaysia and Indonesia. Remarks:
demersal on insular and continental
shelves, and occasionally in coastal
embayments. Used for its meat, freshand dried, and is in demand locally.

Himantura walga, KAUM–I. 16922, 17.7 cm DW
off Terengganu (KT), 10 Dec. 2008

Taeniura lymma(Forsskål, 1775)
Bluespotted Fantail Ray
Disc oval; tail base broad, tapering
beyond sting; ventral skin fold on tailrelatively deep, extending to tail tip;
dorsal surface almost smooth to granular (denticles very small); 2 stings,
located posteriorly on tail. Color: discand tail brownish or orangish dorsally,
disc with numerous bright blue spots,
tail with blue stripe on each side before sting. Size: maximum size 350
cm disc width. Distribution: widespread in Indo-West Pacific, fromsouthern Africa to the Solomon Islands, south to tropical Australia, andnorth to the Philippines and Viet Nam.
Remarks: dominant ray in coral reef
habitats; occurs in inshore to depths ofat least 20 m; feeds on molluscs, polychaetes and crustaceans; marketed
fresh.

Taeniura lymma, KAUM–I. 17283, 22.9 cm DW (tail cut off)
Redang Island, 18 Jan. 2009

20

Taeniurops meyeni(Müller & Henle, 1841)
Blotched Fantail Stingray
Disc circular; tail relatively short,becoming much more slender beyond
sting; long, deep ventral skin fold on
tail, extending to tail tip; dorsal surface almost smooth (denticles verysmall); a large sting, located posteriorly on tail. Color: disc dark with white
blotches and mottling (often faint); tailuniform black posterior to sting. Size:maximum size 180 cm disc width.
Distribution: widespread in IndoWest Pacific, from South Africa toJapan, south to Micronesia, including
northern Australia. Remarks: demersal on soft bottoms near coral reefs,
to offshore depths exceeding 400 m;
feeds on bottom bivalves, crustaceans
and small fishes.

Taeniurops meyeni, KAUM–II. 41, ca. 130 cm DW
off Terengganu (KT), 17 Dec. 2008

21

GYMNURIDAE
Butterfly Rays

Small to moderately large batoids; maximum size 2.6 m in total
length, 4 m disc width. Head, trunk,and greatly enlarged pectoral fins fully fused to form a laterally expanded,rounded rhomboidal or diamondshaped disc; precaudal tail moderately depressed or cylindrical; tail short,
whip-like. Five pairs of gill openings on ventral side of disc. Body
entirely naked, except a few large
species with fine denticles dorsally.
In some species, tail with 1 stinging spine (sting), located just behindpelvic fins, absent in others. Color:dorsal surface of disc and pelvic finsbrown, gray or greenish, whitish ventrally; dorsal surface either unspottedor variegated with light or dark spots
and reticulations, with a pair of large
white spots in some species; tail withblack and white stripes.

By B. Mabel Manjaji-Matsumotorounded-rhomboidal
pectoral disc, about
twice as wide as longtail very thin and shorter
than ½ disc width

Remarks: occurring in all tropical and subtropical continental seas.
Range from the intertidal to the ofshore shelf on soft bottom, to depthsof 110 m. Feed on crustaceans andsmall fishes. Ovoviviparous.
Similar families occurring in

the area: no other batoids in the area
combine large rounded-rhomboidal
pectoral discs nearly or quite twice aswide as long and short slender whiplike tails without caudal fins.

Gymnura poecilura(Shaw, 1804)
Longtail Butterfly Ray
Tail length about equal to snout-cloaca length; no dorsal fin; no sting on
tail. Color: disc with faint whitish
spots (sometimes plain) dorsally; discpale ventrally. Size: maximum size 99
cm disc width. Distribution: tropicalIndo-West Pacific, from the Red Sea
eastward to southern Japan and eastern Indonesia. Remarks: demersal on
the continental shelf, to depths of atleast 30 m; appears to prefer sandy
habitats, often in shallow inshore waters and on sand banks; marketed
fresh.

Gymnura poecilura, KAUM–I. 17014, 26.3 cm DW
off Terengganu (KT), 15 Dec. 2008

22

MYLIOBATIDAE
Eagle Rays

Medium to large sized batoids;
maximum size 330 cm disc width.
Head with rostral or cephalic fins;eyes and spiracles present laterally
on head. Five pairs of gill openings
on ventral side of disc. Trunk fusedwith broad, wing-like pectoral fins,forming a large flattened rhomboidal disc. Body naked or with small
denticles covering the dorsal surface.
Tail slender, long, whip-like, without caudal fin. A single, small dorsal
fin on tail base; one or more stings
in most species. Color: plain green,brown, gray or yellowish to blackish dorsally, most species white ventrally; dorsal surface either unspotted or variegated with light spots,
light transverse stripes, dark stripes
and reticulations, or small ocelli, nopaired ocelli on pectoral bases.

By B. Mabel Manjaji-Matsumoto
rostral fin

tail slender, whip-like,
without caudal finseyes and spiracles
lateraly on headstinging spine (s)usually present

dorsal fin

Remarks: circumglobal in temperate to tropical marine, coastal andoffshore areas in Atlantic, Indian andPacific Oceans. Viviparous.
Similar families occurring in thearea: Rhinopteridae – snout notched
medially, formed as 2 rounded lobes;

3 rows of laterally expanded medialplate-like teeth on jaws. Mobulidae– snout formed into paired elongatedcephalic fins; teeth very small, notplate-like; internal gill arches withlarge and complex filter plates.

Aetobatus ocellatus(Kuhl, 1823)
Whitespotted Eagle Ray
Snout moderately long, broadly
rounded; spiracles dorsolateral onhead; nasal curtain V-shaped; teeth ina single row in both jaws, chevronshaped. Color: disc dark brown dorsally, with numerous white spots; paleventrally. Size: maximum size 330 cm
disc width. Distribution: tropical andwarm temperate seas of the Indo-West
Pacific, from the Red Sea and South
Africa to Hawaii, north to Japan, andsouth to Australia. Remarks: inshoreand benthopelagic on continental
shelves, in estuaries, near atolls andoffshore in open water. Viviparous.

Aetobatus ocellatus, KAUM–I. 17339, 43.6 cm DW
off Terengganu (KT), 21 Jan. 2009

23

Aetomylaeus nichofii(Bloch & Schneider, 1801)
Banded Eagle Ray
Fleshy ridge on side of head not connected to edge of disc; spiracles lateralon head; posterior margin of dorsal fin
strongly angled, its origin slightly anterior to or opposite pelvic-fin insertions; edge of nasal curtain near mouth
almost straight. Color: disc yellowish
dorsally, with 5 bluish bands (often
faint); disc pale ventrally, posterior
margin of pectoral fins and pelvic finsdark. Size: maximum size 72 cm discwidth. Distribution: Indo-West Pacific, from India through Indonesia
and northward to Japan. Remarks:bottom intertidal, and offshore todepths of at least 70 m; viviparous.

Aetomylaeus nichofii, KAUM–I. 17338, 28.6 cm DW
off Terengganu (KT), 21 Jan. 2009

24

ELOPIDAE
Ladyfishes

Medium to large sized (maximum 100 cm, commonly 50 cm)
fishes, slender and moderately compressed. Mouth large, terminal; jaws
subequal; posterior tip of upper jaw
extending posterior margin of eye; agular plate present between arms oflower jaw. Teeth small and granular.
Branchiostegal rays numerous, approximately 27–35. Fins without
spines; a dorsal fin begins slightly
behind midbody, with 20–25 rays,
the last ray not elongate; anal fin
short with 13–18 rays, situated wellbehind base of dorsal fin; pectoral
fins with 17–18 rays, set on side ofbody, near ventral outline; pelvic finsabdominal with 12–16 rays, below
origin of dorsal fin; caudal fin deeply
forked. Scales very small; usually
95–120 in lateral line. Vertebrae 63–
79. Color: body silver, blue or greenish gray dorsally; fins sometimes
with a faint yellow tinge.

By Mizuki Matsunuma
last dorsal-fin ray not elongate

lateral line present
mouth terminal

Remarks: one genus (Elops) with
about six species, two of which distributed in the Pacific Ocean. Coastal fishes found in lagoons, bays, and estuaries; commonly travelling in schools inopen water; feeds on various fishes andcrustaceans. Leptocephalus larva having developed caudal fin. Food fishes
used as fresh, dried or salted one; also
known as sport fishes.
Similar families occurring in thearea: Elopidae differs from other similar families in having gular plate, noscutes along midline of ventral, large

number of lateral line scales, muchnumber of branchiostegal rays. Clupeidae – lateral line absent; gularplate absent; most species have scutes
along midline of ventral. Megalopidae (Megalops cyprinoides) – scales
much larger, about 30 to 40 in lateralline; last ray of dorsal fin elongate
and filamentous. Albulidae (Albulaspp.) – mouth inferior. Chanidae(Chanos chanos) – mouth smaller,
gape not extending behind eye; gularplate absent; branchiostegal rays few(4 or 5).

Elops hawaiiensis
Regan, 1909
Hawaiian Ladyfish
D 23–27; A 15–17; P1 17–18; P2
14–15; LL 93–100; BR 27–35; GR
7–8 + 13–15 = 20–23; V 68–70. Body
elongate and slender. Mouth large andterminal; upper jaw extending beyonda vertical through posterior margin ofeye; a gular plate present between
arms of lower jaw. Scales very small.
Dorsal fin large; without elongatedlobe. Caudal fin deeply forked. Color:silvery sides, darker dorsally; fins yellowish. Size: commonly 50 cm. Distribution: widely distributed in the
Indo-Pacific including the Hawaiian
Islands. Remarks: found mainly incoastal waters; larvae to juveniles occurs estuaries, and mouth of river.
Marketed fresh, dried, or salted.

Elops hawaiiensis, UMTF 1046 (KAUM–I. 16452), 12.8 cm SL
estuary near UMT, 28 Sept. 2008

Elops hawaiiensis, KAUM–I. 16460, 26.9 cm SL
estuary near UMT, 28 Sept. 2008

25

MEGALOPIDAE
Tarpons

Medium to large sized (up to 2.4m) fishes. Body moderately deep,
compressed, deepest in middle, tapering toward both ends. Eye large.
Mouth large, gape ending at level ofposterior margin of eye; lower jaw
projects beyond snout; a gular plate
present between arms of lower jaw.
Teeth small, granular. Branchiostegal rays numerous, 23–27. Gill rakers long and slender. Fins without
spines; a dorsal fin at midbody with
13–21 rays, last ray elongate andfilamentous; anal fin relatively longwith 22–29 rays; pectoral fins set low
on side of body near ventral margin;
pelvic fins abdominal with 10 or 11
rays; caudal fin deeply forked. Scales
large, about 40–50 in lateral line.
Color: body silvery, bluish green
dorsally.
Remarks: one genus (Megalops),
with two species are known fromworldwide, one of which (M. cyprinoides) distributed in the Indo-Pa-

By Mizuki Matsunuma
last ray of dorsal
fin elongate and filamentous
mouth terminal
lateral line present

cific. Coastal fishes found in lagoons,bays, and estuaries; commonly travelling in schools in open water; feeds on
various fishes and crustaceans. Leptocephalus larva having forked caudal
fin. Food fishes used as fresh, dried orsalted one; also known as sport fishes.
Similar families occurring in thearea: Megalopidae differs from othersimilar families by having a gularplate, no scutes on ventral, fewer number of lateral line scales, much number of branchiostegal rays, last dorsal-fin ray elongate and filamentous.
Elopidae (Elops spp.) – scales muchsmaller, about 100 on lateral line;
last dorsal-fin ray not elongate andfilamentous. Albulidae (Albula spp.)– mouth inferior; last dorsal-fin ray
not elongate and filamentous; scales
smaller. Clupeidae – lateral line absent; gular plate absent; most species have scutes on midline of belly.
Chanidae (Chanos chanos) – mouthsmaller, gape not reaching beyond
anterior part of eye; scales smaller;
last dorsal-fin ray not elongate andfilamentous; gular plate absent; fewer
branchiostegal rays (4).

Megalops cyprinoides(Broussonet, 1782)
Indo-Pacific Tarpon
D 16–20; A 23–31; P1 10–11; P2 14–
15; LL 30–40; BR 23–27; GR 7–8 +13–15 = 20–23; V 66–70. Body moderately deep and compressed. Mouth
large and terminal; lower jaw projects
beyond snout; a gular plate present between arms of lower jaw. Scales large.
Dorsal fin large; with elongated lobe.
Caudal fin deeply forked. Color: bodysilvery, darker dorsally; dorsal andcaudal fins dark yellowish. Size: 80
cm SL. Distribution: Indo-Pacific, including the Red Sea, from the east
coast of Africa to the Society Islands,
north to southern Japan. Remarks:
marketed fresh.

Megalops cyprinoides, KAUM–I. 16463, 10.5 cm SL
estuary near UMT, 29 Sept. 2008

Megalops cyprinoides, KAUM–I. 17054, 26.4 cm SL
off Terengganu (KT), 18 Dec. 2008

26

ALBULIDAE
Bonefishes

Medium to large sized fishes (upto 80 cm) with body slender, moderately elongate, slightly compressed.
Eye large. Snout conical, projecting well beyond lower jaw. Mouth
small, inferior, gape not reachinganterior margin of eye; a small gularplate present between arms of lower
jaw, but small and inconspicuous.
Teeth small, granular, in patches onjaws and on roof and floor of mouth.
Branchiostegal rays about 10–15.
Gill rakers rudimentary, consistingof small patches of minute, villiform teeth. All fins without spines; adorsal fin situated at midbody, with
16–21 rays, last ray filamentous inAlbula nemoptera; anal fin short with
7–9 rays, located well behind dorsal
fin; pectoral fins set low on side ofbody, near ventral outline; pelvic finsabdominal, located under posterior
part of dorsal fin; caudal fin deeply

By Mizuki Matsunuma

lateral line present

mouth inferior

forked. Scales small, 66–84 in lateralline. Color: bluish green dorsally,
often with several faint saddles ofslightly darker color; silvery on sides,with several faint, narrow, longitudinal lines; belly white.
Remarks: occurring usually inshallow coastal waters on sandy or
muddy bottoms, rarely found in brackish and freshwater. One genus, Albula,with about six species including several undescribed species. Food fish marketed fresh; also known as sport fishes.

Similar families occurring in
the area: Elopidae – mouth terminal;
gular plate well developed; branchiostegal rays 27–35; upper jaw extending beyond mid-eye. Megalopidae –mouth supraterminal; gular plate welldeveloped; branchiostegal rays 23–
27; upper jaw extending beyond mideye; scales larger. Sillaginidae – two
dorsal fins, first with IX–XII spines;
anal fin base long with II spines and
15–27 soft rays.

Albula argentea(Forster, 1801)
Pacific Bonefish
D 16–17; A 7–8; P1 16–18; P2 9–10;
LL 62–72; BR 12–15; GR 7–11 + 10–
12 = 17–22; V 65–73. Body slender.
Snout long, conical; mouth inferior,
lower jaw in ventral view pointed;
posterior tip of upper jaw not reaching
to front margin of eye. Tip of pelvic finreaching anterior edge of anus. Villiform teeth in small patches on premaxilla and dentary; molariform teethin broad bands on parasphenoid, basibranchials and mesopterygoid; anterior end of tooth bands on mesopterygoids almost corresponds to that of
tooth band on parasphenoid. Scales
cycloid. Color: head and body brightsilver; posterior margin of caudal fin
black; posterior tip of lower caudal

Albula argentea, KAUM–I. 16854, 30.0 cm SL
off Terengganu (KT), 5 Dec. 2008

lobe white; first pelvic-fin ray white;anal fin white; base of pectoral andpelvic fins slightly yellow; a vivid yellow line on preopercle from end of
maxilla; anterior margin of snout
black. Size: 80 cm. Distribution:
west and south Pacific, from the Indo-

Malayan region east to the Society Islands and Marquesas. Remarks: according to Hidaka et al. (2008), Albula
forsteri Valenciennes, 1847 is juniorsynonym of A. argentea. Found incoastal waters, bay, and mouth of river. Marketed fresh, dried, or salted.

27

OPHICHTHIDAE

Snake Eels and Worm Eels
By Mizuki Matsunuma and Hisashi Imamura

Marine fishes, some species in or
occasionally entering freshwater.
Body snake-like or worm-like, moreor less rounded in cross section.
Throat swollen. Eye usually small
and just above mouth. Snout pointed.
Anterior nostril tubular, near tip ofsnout. Posterior nostril low on head,on lip, or opening inside of mouth.
Mouth moderate to large, terminal orinferior. Teeth variable in form, fromfang-like to conical to molariform tovilliform. Gill opening midlateral tocompletely ventral, round or slit-like.
Dorsal, anal, pectoral and caudal finspresent or absent; when caudal fin
present, confluent with dorsal andanal fins, when absent, tip of tail often hard and pointed. Lateral line
complete, usually with well developed pores on head and body. Scales
absent. Color: highly variable, from
uniform light or dark to various patterns of spots, stripes, or bars; usually darker dorsally than ventrally.

dorsal fin present or absent

throat swollen
anal fin present
or absent

pectoral fin
present or absent

Remarks: ophichthyids occur in awide variety of habitats from muddy
estuaries to coral reefs to the midwater
realm, from the shore to depths of
700–800 m or more, but most occur inless than 200 m.
Similar families occurring in thearea: Muraenidae – Posterior nostril
high on head, above or before eye, a
simple pore or in a tube. Muraenesocidae – throat not swollen.

anterior nostril tubular, near tip
of snout

posterior nostril low on head,on lip, or opening inside of mouth

Ophichthus lithinus(Jordan & Richardson, 1908)
Evermann's Snake Eel
P1 14; PLp 68–74; V 151. Body extremely elongate; tip tail finless, pointed. Mouth moderately large; teeth ofupper and lower jaws small and sharp,
those on maxilla arranged in 1–2 rows;
vomer teeth present. Dorsal fin begins
just behind tip of pectoral fin; pectoral
fin well developed. Color: head andbody cream; white ventrally; about 30
poorly defined brown saddles along
head and body; sensory pores on headspotted brown. Size: maximum length
about 90 cm. Distribution: West Pacific, from Australia north to southern
Japan. Remarks: according to McCosker
et al. (2006), Ophichthus evermanni
Jordan & Richardson, 1909 is juniorsynonym of O. lithinus. Inhabits muddy bottom of coastal waters and mouthof river.
(M. Matsunuma)

Ophichthus lithinus, KAUM–I. 17256, 79.0 cm TL
off Terengganu (KT), 17 Jan. 2009

28

Pisodonophis cancrivorus(Richardson, 1848)
Longfin Snake Eel
P1 13–14; PLp 55–60; V 153–162.
Body extremely elongate. Mouth
moderately large; teeth of upper andlower jaws, and vomer villiform,
forming wide bands. Dorsal fin begins above pectoral fin; tip tail finless,pointed; pectoral fin well developed.
Color: head and body yellowish
brown, darker dorsally; dorsal andanal fins dark brown. Size: maximum
length about 100 cm. Distribution:
Indo-West Pacific, including the Red
Sea, from the east coast of Africa east
to French Polynesia, Australia northto southern Japan. Remarks: inhabits
coastal waters of muddy bottom.
(M. Matsunuma)

Pisodonophis cancrivorus, KAUM–I. 17126, 51.8 cm TL
off Terengganu (KT), 31 Dec. 2008

29

MURAENESOCIDAE
Pike Congers

By Mizuki Matsunuma
Median to large sized (up to morethan 1 m) marine fishes. Body moderately elongate, compressed alongtail. Head moderate to moderately
elongate. Eye well developed. Snout
moderate to acute, projects somewhat beyond tip of lower jaw. Mouth
large, gape ends behind posterior
margin of eye; lips without fleshy
flanges; tips of lower jaw with enlarged teeth that fit into a notch inunderside of snout when mouthclosed. Teeth large, prominent, sharp;
multiserial on jaws; typically in 3
rows on vomer, with a median row ofcanines flanked on each side by a rowof much smaller teeth. Anterior nostril tubular, on side of snout just behind tip; posterior nostril a simple
opening in front of eye at approximately mid-eye level. Gill opening alarge, oblique slit in front of and below pectoral fins; gill openings of the2 sides nearly meet on ventral midline, interspace much smaller thanlength of gill opening. Dorsal andanal fins well developed. Lateral line
complete, but usually opening throughmouth terminal

mouth extends
beyond eyelarge gill opening

a complex or branching system ofmultiple pores rather than a single
pore per segment. Scales absent. Color: brown or silver gray to black,lighter ventrally; vertical fins usually
edged in black; no distinct patterns ormarkings.
Remarks: muraenesocids are foundin tropical and subtropical coastal waters of worldwide, inhabit primarilyon sand or mud bottoms. Commonly
collected with bottom trawls.
Similar families occurring in thearea: Congridae – underside of snout
without a conspicuous notch; teeth onvomer typically inconspicuous; gill
openings not meeting across midline.
Anguillidae – body covered with embedded scales; lower jaw projecting
slightly. Ophichthidae – no caudal fin
but tail tip pointed in most genera;
throat swollen; a median frontal poreon head. Muraenidae – no pectoral
fin; gill opening a small hole.
Chlopsidae – gill opening a small
hole; vomer teeth in 2 divergent rows;
lateral line reduced; pectoral finspresent or absent.

Muraenesox cinereus(Forsskål 1775)
Daggertooth Pike Conger
D before anus 66–78; PLp 40–47; V
145–159. Body moderately elongate;head broader, interorbital width about
8 times in head length; anus before
mid-body. Snout relatively blunt, notwell pointed; mouth large, end of
mouth slit behind posterior margin ofeye; teeth almost canine like; vomer
with a median series of compressed,
strong canine teeth with 1 or 2 basal

Muraenesox cinereus, KAUM–I. 16898, 49.2 cm TL
off Terengganu (KT), 6 Dec. 2008

cusps. Pectoral and caudal fins developed. Color: head and body uniformly brown; lighter ventrally; medianfins blackish marginally; pectoral fin
black; pores in lateral line and on head
white. Size: maximum length about

80 cm. Distribution: Indo-West Pacific, from the Red Sea to Indonesia,
northern Australia north to Japan. Remarks: inhabits coastal waters to
depths of about 100 m. Feeds on benthic fishes and crustaceans.

30

NETTASTOMATIDAE
Duckbill Eels

By Mizuki Matsunuma
Medium to large sized (up to 1m) snake-like marine fishes. Body
very elongate; anus before mid-body;tail slender, attenuate. Head slender.
Eye well developed. Snout and jaws
elongate, snout projects a variable
distance beyond tip of lower jaw.
Mouth large, gape extends to about
rear margin of eye; no fleshy flange
on upper or lower lip; some teeth visible when mouth closed; tip of lower
jaw fits into depression behind intermaxillary tooth patch. Teeth generally small, conical, multiserial on jaws
and vomer, some vomerine teeth enlarged but no species in the area haslarge fangs. Anterior nostril tubular,
near tip of snout; posterior nostril
variable in position, either in front ofeye, on lip, on top of head, or on topof body behind head. Dorsal and anal
fins present, confluent with caudal
fin; dorsal fin begins over or slightly
behind gill opening. Pectoral fins absent in all species occurring in theelongate jaws

pectoral fins absent

area. Lateral line complete. Scales absent. Color: brown, lighter ventrally,
without markings; dorsal and anal finsoften edged in black, especially posteriorly.
Remarks: six genera with about
38 species. Found in moderate to deep
water. Collected with bottom trawls,
but no commercial importance.
Similar families occurring in thearea: Congridae – posterior nostril located far forward on snout, closer toanterior nostril than to eye; inner row
of teeth on upper jaw separated fromouter rows by a longitudinal toothlessgroove. Muraenesocidae – pectoral
fins well developed. Ophichthidae –pectoral fins well developed; caudal
fin absent with pointed tail tip, orpresent. Serrivomeridae – pectoral
fins present. Nemichthyidae – pectoral fins present; anus located under or
shortly behind pectoral fins.

Saurenchelys cancrivora
Peters, 1864
Slender Sorcerer
PLp 29–39; V 211. Body extremely
elongated; head slender; tail very long.
Posterior nostril front of eye at mideye level. Mouth large, end of maxilla
extending beyond posterior margin ofeye. Teeth small conical on both jaw,
premaxilloethmoid, vomer and pterygoid. Dorsal fin behind gill opening.
Color: body semi-translucent; a narrow silver band on head from snout to
opercle; tail brackish. Size: maximum
length about 70 cm. Distribution:
West Pacific. Remarks: according toKlausewitz and Zajoz (2000), Chlopsis
fierasfer Jordan & Snyder, 1901 is junior synonym of S. cancrivora. Collected with bottom trawls.

Saurenchelys cancrivora, KAUM–I. 17036, 24.5 cm TL
off Terengganu (KT), 17 Dec. 2008

31

PRISTIGASTERIDAE
Longfin Herrings

By Seishi Kimura and Mizuki Matsunuma
Moderate to large fishes (maximum length 55 cm SL, usually 20–25
cm SL), with compressed, elongatedor deep body. Mouth usually superior
or terminal in some species; upper
jaw with 2 supramaxillae; lower jaw
articulation under eye; teeth on jaws
small or minute, canines only in awestern Atlantic species, Chirocentrodon bleekerianus. Eyelids withbroad vertical opening in middle.
Branchiostegals 6. Fins without
spines; dorsal fin single, short (absent in Indo-West Pacific species,
Raconda russeliana), situated near
midpoint of body; anal fin long, 30–
92 rays; pectoral fins moderate orlarge; pelvic fins with 6 or 7 rays,
situated before dorsal fin origin, butabsent in some genera; caudal fin
forked. Body covered with cycloid
scales; a complete series of scutes
along abdomen; lateral line absent.
Color: bluish green dorsally, blightsilver laterally and ventrally.
Remarks: schooling fishes
found in coastal areas of tropical andsubtropical seas; some species enter-

lateral line absent
mouth usually
superior
pelvic-fin insertion
before dorsal-fin origin
lower jaw
articulation under eye

more than 30 anal-fin rays

complete series of scutes along abdomening estuaries, and a few species restricted to freshwater. Food fishes
used as fresh, dried, or salted ones.
Similar families occurring in thearea: Pristigasteridae differs fromother similar families in having lower
jaw articulation under eye, pelvic finsinserting before dorsal-fin origin, longanal fin with more than 30 rays, acomplete series of scutes along abdomen, and no lateral line. Elopidae –lateral line present; no abdominal
scutes; gular plate present; numerous
branchiostegal rays (more than 20).
Megalopidae – lateral line present, no

abdominal scutes; gular plate present;
numerous branchiostegal rays (morethan 20). Albulidae – lateral line present; no abdominal scutes; gular plate
present. Chirocentridae – body highly compressed and elongate; no abdominal scutes; jaws with developed
fanglike canine teeth. Clupeidae –pelvic fins inserting just below dorsal
fin origin; anal fin short with less than
28 rays. Engraulidae – snout pig-likeand projecting; jaw articulation wellbehind eye. Chanidae – lateral line
present; no abdominal scutes; branchiostegal rays few (4 or 5).

Opisthopterus tardoore(Cuvier, 1829)
Tardoore
A 51–60; P1 12–14; LGR 22–28.
Body elongate and strongly compressed, moderately deep (body depth
27–33% SL); abdomen concave infront, with 29 to 35 sharply-keeledscutes. Mouth directed obliquely upward; upper jaw short, not reaching
posteriorly to vertical through mideye; no toothed hypomaxilla between
posterior tip of premaxilla and bladeof maxilla. Pectoral fins relatively
long (length 21–26% SL; usually
about equal with or greater than headlength); dorsal fin very small; anal fin

Opisthopterus tardoore, KAUM–I. 16996, 12.4 cm SL
off Terengganu (KT), 14 Dec. 2008

long; pelvic fin absent. Color: bodysilver, yellowish brown dorsally; dorsal and anal fins translucent, with yellow margin; caudal fin pale yellow.
Size: maximum length 18 cm. Distri-bution: tropical Indo-West Pacific,
from the Gulf of Oman to Indonesia.
Remarks: found in coastal waters andestuaries. Marketed fresh, dried, orboiled.
(M. Matsunuma)

32

ENGRAULIDAE
Anchovies

By Mizuki Matsunuma and Seishi Kimura
Small to medium-sized fishes(mostly less than 20 cm, some larger)
with silver-colored, fusiform or slender body. Mouth inferior; snout projecting beyond tip of lower jaw; jaw
articulation well behind eye; upper
jaw with 2 supra-maxillae; teeth onjaws absent or minute, rarely with canines. Fins without spines; dorsal fin
short and single, situated near midpoint of body, far forward in Coilia;
anal fin mostly short, less than 25
rays, but long in some (to over 100rays); pectoral fins set low on body;
pelvic fins abdominal usually with 7
rays; caudal fin deeply forked, but
small and pointed in some. Body
covered with cycloid scales; scutes
often present along abdomen; lateralline absent.
Remarks: schooling fishes foundin marine coastal areas, mostly feeding on small planktonic animals andplants. Small but important food fish-snout projecting

lateral line absent
6–7 pelvic-fin rays

mouth inferior
upper jaw extending beyond hind margin of eye

es used as fresh, dried, or salted ones,
taken by beach seine and other varietyof nets. Regionally some species havegreat contribution to local catches.
Similar families occurring in thearea: Engraulidae differs from otherclupeoid families in having projecting
and pig-like snout, slender lower jaw,
and jaw articulation well behind eye.
Chirocentridae – body highly compressed and elongate; no abdominal

scutes; jaw articulation below eye;jaws with developed fang-like canine
teeth. Clupeidae – snout not pig-likeand projecting; jaw articulation anterior to below eye. Pristigasteridae –snout not pig-like and projecting;lower jaw projecting; jaw articulation
anterior to vertical below eye.

Coilia dussumieri
Valenciennes, 1848
Gold-spotted Grenadier Anchovy
A 80 or more; P1 9–11; P2 7; LGR
23–26. Body tapering, abdomenrounded anterior to pelvic fins; abdomen with 5 or 6 prepelvic and 7–9
postpelvic keeled scutes, 12–15 in total number; a small predorsal spinelike scute present. Maxilla short; two
Coilia dussumieri, KAUM–I. 16993, 14.1 cm SL
supramaxillae present. Teeth in jaws
off Terengganu (KT), 14 Dec. 2008
small. Dorsal fin far forward, beginning in first third of body length; anal
fin long; posteriormost fin ray joined
to caudal fin; caudal fin very small, extremely pointed; pectoral fins long andfilamentous, reaching posteriorly atleast to anal-fin origin. Light organs
present along flanks and abdomen.
Color: body yellowish silver, brownish dorsally; flanks and abdomen withlongitudinal rows of golden or pearly
Coilia dussumieri, UMTF 1870, 13.3 cm SL
off Terengganu (KT), 24 Dec. 2008
spots (light organs) Size: commonly
13–17 cm, maximum 20 cm. Distribution: eastern Indian Ocean andwestern Pacific, from India to the Gulf Marketed fresh, dried, dried-salted orof Thailand and Indonesia. Remarks: made into fish sauce or fish balls.
(M. Matsunuma)
occurs inshore waters and estuaries.

33

Encrasicholina devisi(Whitley, 1940)
Devis' anchovy
D 15; A 18–20; P1 15; P2 7; LGR
20–27. Body elongate; abdomenslightly rounded, with 3–6 (usually 5)needle-like prepelvic scutes; no postpelvic scutes and no predorsal scutes;
posterior tip of maxilla pointed, reaching to subopercle; isthmus muscle
short, preceded by a small bony plateon urohyal between gill membrane;
initial 3 dorsal and anal fin rays unbranched; anal-fin origin slightly posterior to a vertical through end of dorsal-fin base. Color: in life, body withgolden hue; a broad bright silvery tobrassy stripe on side, with a thin blueline above; dorsal part of body bluegray. Size: maximum length 9.5 cm.
Distribution: Indo-West Pacific from
Gulf of Aden east to Samoa; northern
Australia north to Taiwan. Remarks:
schooling fish found in coastal waters.
Marketed fresh or dried; also used as abaitfish for tuna.
(M. Matsunuma)

Encrasicholina devisi, KAUM–I. 16932, 7.8 cm SL
off Terengganu (KT), 11 Dec. 2008

Setipinna taty, KAUM–I. 16994, 12.5 cm SL
off Terengganu (KT), 14 Dec. 2008

Setipinna taty
(Valenciennes, 1848)
Scaly Hairfin Anchovy
A 48–60; P2 7; LGR 17–21 (usually
18–20). Body deep and strongly compressed; abdomen with complete series of keeled scutes from isthmus toanus; 20–29 (usually 22–27) prepelvic
and 9–14 (usually 11 or 12) postpelvic
scutes, 32–40 (usually 33–39) in total
number; a predorsal spine-like scutepresent. Mouth nearly horizontal, jaws
slender; first supramaxilla absent; second supramaxilla relatively small,rounded. Dorsal-fin origin anterior to
mid-body midpoint, slightly anterior
to vertical through anal-fin origin; anal
fin long; pectoral fin with first fin ray
filamentous, its tip reaching posteriorly to base of 23rd to posteriormost
anal-fin ray; caudal fin well forked,
upper lobe truncated. Scales cycloid,
present on dorsal and anal fins; longitudinal scale rows about 40–48. Color: body yellowish silver, brown dorsally; fins pale to deep yellow, caudal
fin with black margin. Size: commonly 10 cm, maximum length 14 cm.
Distribution: Western Central Pacific, from Gulf of Thailand south to In-

Stolephorus andhraensis, KAUM–I. 17284, 8.0 cm SL
off Terengganu (KT), 19 Jan. 2009

donesia. Remarks: schooling fish
found in coastal waters, often entering
estuaries. Marketed fresh or dried.
(M. Matsunuma)

Stolephorus andhraensis
Babu Rao, 1966
Andhra Anchovy
D 13–14; A 19–20; LGR 20–21.
Body slender, somewhat compressed;
abdomen with 6–7 (usually 6) small
needlelike prepelvic scutes, but nopostpelvic scutes; pelvic scute without
spine; no predorsal spine-like scute;pelvic scute without spine. Posterior
tip of maxilla pointed, extending to or

beyond posterior border of preopercle;
posterior border of preopercle concave. Isthmus muscle reaching to andbeyond gill membrane. Anal fin short,
its origin situated below middle ofdorsal fin base; caudal fin large andforked. Scales cycloid; longitudinalscale rows about 37–39. Color: bodysemi-translucent, with a broad silver
stripe midlaterally; caudal fin dusky.
Size: maximum ca. 5 cm SL. Distribution: eastern coast of India, Andaman Sea coast of Thailand, Singapore,
Gulf of Papua, and northern and northeastern Australia. Remarks: schooling fish found in coastal waters.
(S. Kimura)

34

Stolephorus baganensis
Hardenberg, 1933
Bagan Anchovy
A 21–22; LGR 20–23. Body slender,
somewhat compressed; abdomen with
6–7 (usually 6) small needle-like prepelvic scutes, but no postpelvic scutes;
pelvic scute with spine; a small predorsal spine-like scute present. Posterior tip of maxilla pointed, extending
to or beyond posterior border of preopercle; posterior border of preopercle
rounded, indented near maxilla tip.
Isthmus muscle reaching to and beyond gill membrane. A few small teeth
somewhat present on upper edge ofhyoid bones. Anal fin short, its origin
situated below middle of dorsal finbase; caudal fin large and forked.
Scales cycloid; longitudinal scalerows about 34–36. Color: body semitranslucent, with a broad silver stripe
midlaterally; a double pigment line on
dorsum posterior to dorsal fin. Size:maximum length about 9 cm. Distribution: eastern Indian Ocean andwestern Pacific. Remarks: schoolingfish found in coastal waters.
(M. Matsunuma)

Stolephorus baganensis, KAUM–I. 17116, 9.3 cm SL
off Terengganu (KT), 31 Dec. 2008

Stolephorus indicus, KAUM–I. 16822, 14.0 cm SL
off Terengganu (KT), 5 Dec. 2008

Stolephorus indicus(van Hasselt, 1823)
Indian Anchovy
D 15–17; A 19–21; P1 15–17; P2 7;
LGR 20–28. Body slender, round incross-section; abdomen with 3–5
small needle-like prepelvic scutes, but
no postpelvic scutes; pelvic scutewithout spine; no predorsal spine-like
scute; posterior tip of maxilla pointed,
extending to or slightly beyond anterior border of preopercle; posteriorborder of preopercle convex androunded. Isthmus muscle tapering andreaching to gill membrane. Pelvic finsinserting well before vertical throughdorsal fin origin; anal fin short, its origin situated at vertical through middleof dorsal fin base; caudal fin large andforked. Scales cycloid; longitudinalscale rows about 40. Color: bodytranslucent, yellowish dorsally, with abroad silver stripe midlaterally; headsilver. Size: commonly 12 cm, maximum length 15 cm. Distribution:
widely distributed in the Indo-Pacific,
including the Red Sea, from the east
coast of Africa to Samoa and Tahiti.
Remarks: schooling fish found inThryssa dussumieri, KAUM–I. 17020, 8.8 cm SL
off Terengganu (KT), 16 Dec. 2008

coastal waters, often entering estuar- base in adults. Anal fin long, its origin
ies.
(M. Matsunuma) situated posterior to dorsal-fin base;upper pectoral-fin ray not extended asa filament. Scales cycloid; longitudiThryssa dussumieri
nal scale rows 36–40. Color: body(Valenciennes, 1848)
silvery white, grayish brown dorsally;
Dussumier's Thryssa
a dark blotch posterior to upper part ofA 32–40 (usually 34–39); P2 7; LGR gill opening, usually joined to a dark
17–19. Body somewhat deep, com- saddle on nape; dorsal and caudal finspressed; abdomen with 15–16 prepel- yellowish; caudal fin with broad black
vic and 6–9 (usually 7–8) postpelvic margin. Size: maximum length 11 cm.
keeled scutes, 21–24 (usually 22–23) Distribution: Indo-West Pacific, fromin total number; a small predorsal the coast of Pakistan to Indonesia,
spine-like scute present. Teeth in jaws north to Taiwan. Remarks: schoolingsmall or minute, not canine-like. Max- fish found in inshore waters and estuilla very long, reaching posteriorly to a aries. Marketed fresh, dried and dried(M. Matsunuma)
point at least halfway along length of salted.
pectoral fin and nearly to pelvic-fin

35

Thryssa mystax(Bloch & Schneider, 1801)
Moustached Thryssa
A 32–40 (usually 35–38); P2 7; LGR
14–16. Body somewhat deep, compressed; abdomen with 17–19 prepelvic and 8–13 (usually 11) postpelvic
keeled scutes, 24–32 (usually 28–30)
in total number; a small predorsal
spine-like scute present. Teeth in jaws
small or minute, not canine-like. Maxilla long, reaching or almost reaching
posteriorly to base of first pectoral-fin
ray. Anal fin long, its origin posterior
to vertical through end of dorsal-finbase; upper pectoral-fin ray not extended as a filament. Scales cycloid;
longitudinal scale rows 41–46. Color:body silvery white, brownish dorsally;a dark blotch of wavy lines posterior
to upper part of gill opening; dorsalfin tip black; upper, lower, and posterior margins of caudal fin black. Size:maximum length 16 cm. Distribution: Indo-West Pacific, from the west
coast of India to Indonesia, north toGulf of Thailand. Remarks: schooling fish found in inshore waters andestuaries. Marketed fresh, dried anddried-salted.
(M. Matsunuma)

Thryssa mystax, KAUM–I. 16988, 13.0 cm SL
off Terengganu (KT), 14 Dec. 2008

Thryssa mystax, UMTF 1865, 11.9 cm SL
off Terengganu (KT), 24 Dec. 2008

Thryssa setirostris(Broussonet, 1782)
Longjaw Thryssa
D 12; A 32–39 (usually 35–38); P2 7;
LGR 10–12. Body somewhat deep,
compressed; abdomen with 16–18
prepelvic and 9–10 postpelvic keeledscutes, 25–28 in total number; a small
predorsal spine-like scute present.
Teeth in jaws small or minute, not canine-like. Maxilla very long, reaching
posteriorly at least to tip of pectoral
fins, usually to pelvic-fin base, or evento anal-fin origin. Anal fin long, its
origin at or posterior to verticalthrough end of dorsal-fin base; upper
pectoral-fin ray not extended as a filament. Scales cycloid; longitudinalscale rows 41–46 (usually 41–44).
Color: body silvery white, brownish
dorsally; a diffuse patch of horizontal
wavy gray or charcoal lines on shoul-Thryssa setirostris, KAUM–I. 17021, 10.6 cm SL
off Terengganu (KT), 16 Dec. 2008

der posterior to upper part of gill
opening; dorsal and caudal fins dusky
to deep yellow, margins charcoal; anal
fin deep yellow or white; pectoral andpelvic fins pale yellow. Size: maximum length 18 cm. Distribution: Indo-West Pacific, from the Gulf ofOman east to Vanuatu, northern Australia north to Taiwan. Remarks:
schooling fish mostly found in closeinshore, entering bays and estuaries.
Marketed fresh, dried and dried-salted.
(M. Matsunuma)

36

CHIROCENTRIDAE
Wolf Herrings

By Seishi Kimura
Medium to large-sized silvery
fishes; maximum size about 1 m SL.
Body elongate and strongly compressed; no pelvic scutes. Two fanglike teeth on upper jaw; a series ofcanine teeth on lower jaw. Eyes relatively small, with eyelids completely
covering eyes. Gill rakers 17–22;
branchiostegal rays 8. No spinous fin
rays; dorsal-fin with 16–19 rays, its
origin just above anal-fin origin, itsbase shorter than anal-fin base; anal
fin with 30–35 rays; caudal fin deeply forked; pelvic fins small, abdominal, with 6–7 rays. Scales small cycloid; lateral line absent. No pyloric
caeca; spiral valve present in intes-

2 fang-like teeth on
upper jaw
lateral line absent

no scutes on belly

tine. Total vertebrae 69–75. Color:dark bluish green dorsally, blight silver laterally and ventrally.
Remarks: occurring in coastal
area of Indo-West Pacific, from South
Africa to Japan and Australia. Feeds on
fishes. A single genus, Chirocentrus,
with two species.

Similar families occurring in
the area: Elopidae and Chanidae –no fanglike teeth; lateral line present.
Engraulidae, Pristigasteridae, andClupeidae – no fang-like teeth; scutes
usually present on belly.

Chirocentrus dorab(Forsskål, 1775)
Dorab Wolf-herring
D 16–18; A 29–36; P1 13–16; P2 6;
GR 1–4 + 10–16. Body elongate,strongly compressed, belly sharp; pectoral fin short, 11–13% SL. Mouth
large, directed upward; both jaws withlarge canine teeth anterioly. Scales cycloid, very small. Color: dark bluegreen dorsally, silvery ventrally; black
marking on the upper part of dorsal
fin; caudal fin blackish. Size: maximum 1 m SL, commonly 30–50 cm
SL. Distribution: widely distributed

Chirocentrus dorab, KAUM–I. 17023, 25.0 cm SL
off Terengganu (KT), 16 Dec. 2008

in Indo-West Pacific from Red Sea andSouth Africa to Solomon Is., north tosouthern Japan. Remarks: occurringin inshore waters. Feeds on small fishes, crustaceans, squids, etc. Marketed
fresh.

37

CLUPEIDAE

Herrings (Sardines)
By Seishi Kimura and Mizuki Matsunuma
Small to medium-sized fishes(mostly 10 to 20 cm, some exceeding
50 cm) with silver-colored fusiform
body, compressed or oval in crosssection. Mouth terminal or somewhat
superior except for members of Dolateral line absent
rosomatinae with mouth fully inferior; upper jaw mostly with 2 supramaxillae; jaw articulation always
anterior to vertical below eye; teeth
on jaws mostly absent or minute, branchiostegal rays usually 5–10
8–9 pelvic-fin rays
sometimes with canines. Fins without spines; dorsal fin single, situatednear midpoint of body; anal fin short plants. Small but important food fishes no abdominal scutes; gular plate presless than 28 rays; pectoral fins set used as fresh, dried, or salted ones, ent; numerous branchiostegal rays
low on body; pelvic fins abdominal taken by beach seine and other variety (more than 20). Albulidae – lateralwith 8 or 9 rays; caudal fin deeply of nets. Regionally some species have line present; no abdominal scutes; guforked. Body covered with cycloid great contribution to local catches.
lar plate present. Chirocentridae –scales; a series of scutes along abdoSimilar families occurring in the body highly compressed and elonmen; lateral line absent. Color: typi- area: Clupeidae differs from other gate; no abdominal scutes; jaws withcally blue-green dorsally, silvery on similar families in having jaw articula- developed fang-like canine teeth.
flanks; with variable darker markings tion before vertical through eye, mostly Pristigasteridae – pelvic fins insertincluding spot behind gill cover, 2 supramaxillae, pelvic fins inserting ing anterior to vertical through dorsal
spots along flanks, spot at dorsal-fin below short dorsal fin, short anal fin fin origin; anal fin long more than 30
origin, and dark pigmentation on part less than 28 rays, a series of scutes rays. Engraulidae – snout pig-likeof dorsal, pectoral, anal, and caudal along abdomen, and no lateral line. and projecting; jaw articulation well
fins.
Elopidae – lateral line present; no ab- behind eye. Chanidae – lateral line
Remarks: schooling fishes found dominal scutes; gular plate present; nu- present; no abdominal scutes; branin marine coastal areas, mostly feed- merous branchiostegal rays (more than chiostegal rays few (4 or 5).
ing on small planktonic animals and 20). Megalopidae – lateral line present,

Amblygaster clupeoides
Bleeker, 1849
Bleeker's Smoothbelly Sardinella
D 18; A 18; P1 17; P2 8; LGR 26–31.
Body moderately slender, somewhat
cylindrical; belly rounded with 16–19(usually 17) prepelvic and 11–14 (usually 13) postpelvic somewhat weakscutes. Upper jaw rounded without distinct median notch or cleft; 2 supramaxillae present; second (posterior)supramaxilla symmetrical; posterior tipof maxilla not reaching posteriorly tovertical through anterior margin of eye.
Opercle smooth, without bony radiating striae. Shoulder girdle (cleithrum)
margin with 2 outgrowths. Scales cycloid; a median series of predorsal
scales present. Color: dark blue-green

Amblygaster clupeoides, KAUM–I. 17040, 17.6 cm SL
off Terengganu (KT), 17 Dec. 2008

dorsally, silvery ventrally; no blackish
spots laterally on trunk; a black blotch
behind gill opening; dorsal fin blackish. Size: commonly to 15 cm, maximum length 18 cm. Distribution: Indo-West Pacific, from southern coasts

of India to Indonesia, and also Fiji.
Remarks: schooling fish found incoastal waters. Marketed fresh, dried,or dried-salted.
(M. Matsunuma)

38

Amblygaster sirm
(Walbaum, 1792)
Spotted Sardinella
D 17–19; A 15–19; P1 16–17; P2 8;
LGR 33–43; V 41–43. Body slender,
somewhat cylindrical; belly roundedwith 16–17 prepelvic and 13–15 postpelvic somewhat weak scutes (29–32
in total). Upper jaw with 2 supramaxillae; the second (posterior) supramaxillae symmetrical. Opercle smooth,
without bony radiating striae. Shoulder girdle (cleithrum) margin with 2
outgrowths. Scales cycloid; a medianseries of predorsal scales present. Color: dark blue-green dorsally, silvery
ventrally; a series of 10–20 blackish
spots laterally on trunk. Size: commonly to 20 cm, maximum 23 cm SL.
Distribution: widely distributed in
Indo-West Pacific from the Red Sea
and Madagascar to Kiribati and Fiji,north to Japan. Remarks: schoolingfish found in coastal waters, feeds on
copepods and other zoo- and phytoplanktons. Marketed fresh, dried, or
dried-salted.
(S. Kimura)

Amblygaster sirm, KAUM–I. 17041, 16.5 cm SL
off Terengganu (KT), 17 Dec. 2008

Anodontostoma chacunda, KAUM–I. 17057, 11.8 cm SL
off Terengganu (KT), 24 Dec. 2008

Anodontostoma chacunda(Hamilton, 1822)
Chacunda Gizzard Shad
D 17–19; A 19–20; P1 15–16; P2 8;
LGR 54–96; V 41–43. Body very
deep, almost oval, strongly compressed (40–50% SL in specimens >10 cm SL); belly keeled with scutes(ca. 28). Mouth inferior; upper jaw
with a distinct median notch. Opercle
smooth, without bony striae. Longest
lower gill rakers shorter than corresponding gill filaments. Shoulder girdle (cleithrum) margin without fleshy
outgrowths. Last dorsal-fin ray not filamentous. Scales cycloid with denticulation along posterior margin; a median series of predorsal scales present.
Color: body silvery, darker dorsally;
occiput to nape yellowish or gold; alarge black spot behind gill opening.
Size: maximum ca. 18 cm SL. Distribution: widely distributed in IndoWest Pacific from the Persian Gulf to
New Caledonia. Remarks: found ininshore waters and estuaries, feeds onzoo- and phytoplanktons. Marketed
fresh, dried, or dried-salted.
(S. Kimura)

Dussumieria elopsoides, KAUM–I. 17130, 15.0 cm SL
off Terengganu (KT), 1 Jan. 2009

Dussumieria elopsoides, KAUM–I. 17201, 13.8 cm SL
off Terengganu (KT), 10 Jan. 2009

Dussumieria elopsoides
Bleeker, 1849
Slender Rainbow Sardine
D 18–21; A 15–17; P1 14–15; P2 8;
LGR 21–32. Body elongated, cylindrical, slender (16–22% SL); belly
rounded without prepelvic or postpelvic scutes. Premaxillae rectangular;
branchiostegal rays numerous (13–18). Scales cycloid with numerous

tiny radiating striae posteriorly. Color: dark blue-green dorsally, silvery
ventrally, with a narrow lateral band ofsilvery gray and gold extending fromupper operculum to caudal-fin base.
Size: maximum ca. 20 cm SL. Distribution: Indo-West Pacific. Remarks:
pelagic, inshore fish. Feeds on planktonic crustaceans and small fishes.
Marketed fresh, dried, or dried-salted.
(S. Kimura)

39

Escualosa thoracata(Valenciennes, 1847)
White Sardine
D 16; A 18–19; P1 12–13; P2 7; LGR
27–40. Body somewhat deep (27–37%
SL), compressed; belly strongly
keeled with 17–19 (mostly 18) prepelvic and 10–12 (mostly 11) postpelvic
scutes (28–30, mostly 29, in total).
Upper jaw rounded, without distinct
median notch or cleft; 2 supramaxilla
present. Opercle smooth, withoutbony radiating striae. Shoulder girdle(cleithrum) margin without fleshy outgrowths. Color: body semi-translucent
with broad silvery midlateral stripe;
inner edges of caudal fin broadly darkish. Size: maximum ca. 10 cm SL.
Distribution: widely distributed in
Indo-West Pacific from the Persian
Gulf to northeastern Australia, northto Vietnam and Philippines. Remarks:
schooling fish found in coastal, inshore waters.
(S. Kimura)

Escualosa thoracata, KAUM–I. 17291, 7.3 cm SL
off Terengganu (KT), 20 Jan. 2009

Hilsa kelee(Cuvier, 1829)
Kelee Shad
D 18; A 20–22; P2 8; LGR 100–175.
Body very deep and strongly compressed; abdomen with sharply-keeled
15–17 prepelvic and 12–14 postpelvic
scutes, 27–31 (usually 28–30) in total
number. Mouth terminal; upper jaw
with a distinct median notch. Opercle
smooth, without bony striae. Gill rakers fine and numerous, those on innerarches curled outward; Shoulder girdle(cleithrum) margin without fleshy outgrowths. Frontoparietal striae (on topof head) numerous, 8–14. Last dorsalfin ray not filamentous. Scales cycloid
with denticulation along posterior margin; vertical striae overlapping at center of scale present. Color: body silvery, blue-green dorsally; occiput tonape yellowish or gold; a black spot
behind gill opening. Size: commonly
15–18 cm, maximum 25 cm SL. Distribution: Indo-West Pacific. Remarks: found in inshore waters andestuaries, feeds on zoo- and phytoplanktons. Marketed fresh, dried, or
dried-salted.
(M. Matsunuma)

Hilsa kelee, KAUM–I. 17297, 15.0 cm SL
off Terengganu (KT), 20 Jan. 2009

Sardinella albella, KAUM–I. 16995, 11.6 cm SL
off Terengganu (KT), 14 Dec. 2008

LGR 41–68. Body somewhat compressed, but variable from slender tomoderately deep (body depth 25–40%SL); abdomen keeled with prepelvic
and postpelvic scutes; total number ofscutes 29–33 (usually 30–32); pelvic
scute with ascending arms. Second supramaxilla symmetrical and paddleshaped; opercle smooth without radiating bony striae; posterior border ofSardinella albella
gill opening with 2 fleshy outgrowths.
(Valenciennes, 1847)
Anal fin short with 2 posteriormost
White Sardinellarays enlarged. Scales cycloid; vertical
D 17–19; A 18–19; P1 14–15; P2 8; striae discontinuous, not meeting at

center; posterior part with a few perforations and somewhat produced posteriorly; longitudinal scale rows usually
41–43. Color: body silvery, bluegreen dorsally; a dark spot at dorsalfin origin; dorsal fin pale yellow; tipsof dorsal and caudal fins blackish.
Size: commonly 10 cm, maximum 14
cm SL. Distribution: Indo-West Pacific. Remarks: schooling fish foundin reef-associated coastal waters. Marketed fresh, dried and dried-salted.
(M. Matsunuma)

40Sardinella gibbosa(Bleeker, 1849)
Goldstripe Sardinella
D 17–20; A 17–21; P1 14–17; P2 8;
LGR 45–60. Body moderately slender(body depth 25–40% SL); abdomen
keeled with prepelvic and postpelvic
scutes; total number of scutes 32–34;
pelvic scute with ascending arms. Second supramaxilla symmetrical andpaddle-shaped; opercle smooth without radiating bony striae; posteriorborder of gill opening with 2 fleshy
outgrowths. Anal fin short with 2 posteriormost rays enlarged. Scales cycloid; vertical striae discontinuous,
not meeting at center; posterior partwith numerous small perforations;longitudinal scale rows 43–47. Color:

Sardinella gibbosa, KAUM–I. 17174, 13.2 cm SL
off Terengganu (KR), 5 Jan. 2009body silvery, blue-green dorsally, witha thin golden midlateral line; a dark
spot at dorsal-fin origin; margins of
dorsal and caudal fins blackish. Size:
commonly 15 cm, maximum 17 cm
SL. Distribution: widely distributedin Indo-West Pacific from East African coasts to eastern Australia, northto Taiwan. Remarks: schooling fish
found in coastal waters. Marketed
fresh, dried, dried-salted and/or boiled.
(M. Matsunuma)

CHANIDAE
Milkfish

By Mizuki Matsunuma
This family represented by a single
species; see the following species account.
Similar families occurring in thearea: Megalopidae – a bony gularplate present between arms of lower
jaw; last dorsal-fin ray filamentous;
scales large, 30–40 in lateral line.
Elopidae – mouth much larger; abony gular plate present between
arms of lower jaw. Mugilidae – 2
dorsal fins; pectoral fins set high on
body; no lateral line. Clupeidae – size
much smaller; usually 6 or 7 bran-mouth terminal
no teeth on jaws
lateral line present
4 branchiostegal rays

chiostegal rays (only 4 in Chanos); toral fins in 2 parts, lower part withno lateral line; scutes usually present long unattached rays.
along belly. Polynemidae – mouthlarge, subterminal; 2 dorsal fins; pec-

Chanos chanos(Forsskål, 1775)
Milkfish
D 13–17; A 9–11; P2 9–11; LL 75–91;
GR 147–160 + 107–165. Body elongate, moderately compressed; no scutes
along velly. Eye covered by adipose tissue. Mouth small, terminal, without
teeth; lower jaw with small tubercle attip, fitting into a notch in upper jaw; nobony gular plate between arms of lower
jaw. Branchiostegal rays 4. Dorsal situated on about midpoint of body; anal
fin short, close to caudal fin; caudal finlarge and deeply forked, with scale
flaps at base in adults; pectoral and pelvic fins with axillary scales. Scales
small and smooth. Color: silver later-

Chanos chanos, KAUM–I. 16451, 13.5 cm SL
estuary near UMT, 28 Sept. 2008

ally and ventrally, olive-green or blue
dorsally; dorsal, anal, and caudal finswith dark margin. Size: commonly
70–100 cm SL, maximum 1.5 m SL.
Distribution: widely distributed intropical and subtropical Indo-Pacific.
Remarks: occurring in marine andbrackish waters. Important food fish;
marketed fresh, dried, canned, andsmoked.

41

PLOTOSIDAE

Eeltail Catfishes (Eel Catfishes)
By Mizuki Matsunuma
Small to medium-sized (commonly less than 40cm, some up to
100 cm) catfishes. Body elongate,
tapering posteriorly; without scales
or bony plates. Lateral line complete,running along middle of body andextending nearly to caudal-fin base.
A dendritic organ, located along ventral midline of body just anterior toanal-fin origin and posterior to anus,
found in all marine, and some freshwater species. Mouth surroundedwith barbels in 4 pairs; 1 pair between widely separated anterior andposterior nostrils; 1 pair at corner ofmouth and 2 pairs on lower jaw.
Teeth present on jaws, except on upper jaw of some fresh-water species.
Palate with teeth, often molariform;
molariform teeth sometimes found inlower jaw. Dorsal and pectoral finslocated just posterior of head, eachwith a pungent serrated spine at leading edge; fin spines venomous andcapable of producing painful sting;anal fin long and continuous withrounded or pointed caudal fin; dorsal

second dorsal, caudal and anal fins confluent

no scales

4 pairs of berbels

series of procurrent caudal-fin raysconsists of elongate rays which, in marine species, extend the fin anteriorly
to at least vertical line at anal-fin origin, thus forming a second dorsal finthat is continuous with caudal fin; dorsal adipose fin absent. Color: bodytan, brown, or black; lighter ventrally;sometimes with light stripes alongside of body, extending onto head.
Remarks: found in fresh, brackish, and marine waters of tropical andsubtropical regions of the Indo-Pacific; freshwater species restricted toAustralia and New Guinea. Venomous, but marketed mostly fresh and locally may be important.

Similar families occurring in
the area: Plotosidae differs from allother catfish families in having adendritic organ (marine species), andcaudal fin confluent with dorsal procurrent caudal, and anal fins. Ariidae– caudal fin forked; adipose fin present; anal fin short and not confluent
with caudal fin; nasal barbel absent.
Other freshwater catfish families(Siluridae, Clariidae and Heteropneustidae) – dorsal and pectoral finswith no pungent serrated spine at anterior edge; anal fin not confluent
with caudal fin.

Plotosus lineatus(Thunberg, 1787)
Striped Eel Catfish
D I, 4; DPC 89–111; A 68–80; P1 I,
10–12; P2 10–13; GR 6–8 + 20–23 =27–31. Body somewhat elongate;
body depth more than 15% SL; anterior nostril situated dorsal to upper lip,
Plotosus lineatus, UMTF 1355 (KAUM–I. 16658), 26.5 cm SL
opening directed anteriorly; gill memBidong Island, 13 Oct. 2008branes narrowly attached across isthmus; nasal and maxillary barbels
short, reaching to or slightly beyond
posterior margin of eye; origin of dorsal procurrent caudal fin inserting vertical above pelvic-fin base. Color:head and body dark brown, whitish
ventrally; 2 or 3narrow pale-yellow
longitudinal stripes on body, of which
2 extending onto head; barbels darkbrown; dendritic organ yellowish
Plotosus lineatus, KAUM–I. 17010, 20.4 cm SL
brown. Size: commonly 20 cm, maxioff Terengganu (KT), 15 Dec. 2008
mum 30 cm. Distribution: widelydistributed in the Indo-West Pacific,
from the Red Sea and east coast of Af- found in coastal waters, often entering occurring in small schools. Each serrica to Samoa, northern Australia estuaries. Juveniles forming dense rate spine of dorsal and pectoral finsnorth to southern Japan. Remarks: ballshaped schools, adults solitary or venomous and highly dangerous.

42

ARIIDAE

Sea Catfishes
By Seishi Kimura
Medium to large sized marine
and freshwater catfishes; maximum
size about 1.8 m. Body elongated,robust; head depressed but tail compressed. Bony shield dorsally onhead usually visible, with a dorsomedian grove, extending posteriorly
to nuchal bone; snout pointed orrounded anterior and posterior nostrils close together; mouth terminal
or inferior; one, two, or three pairsof barbells present around the mouth,
no nasal barbels; jaw teeth small, arranged into narrow or broad bands;
palatal teeth on small or large patches; gill membranes joined together
and attached isthmus anteriorly;
branchiostegal rays 5–7. Dorsal fin
with a long, robust, usually serrated
spine followed by 7 branched rays; avery small spinelet or buckler present prior to dorsal spine; anal fin with
14–36 soft rays; caudal fin deeply
forked with 15 (7 + 8) principal rays;
pectoral fins low on the sides of bodywith a usually long and serrated
spine and 7–13 branched rays; pelvic fins abdominal with 6 branched
rays; adipose fin present just above
anal fin. Body naked; lateral line welldorsal-fin spine
usually serrated
adipose fin just above anal fin

no scales

short anal-fin basecaudal fin
deeply forked

1–3 pairs of berbels

developed. Color: body dark, brownish dorsally, silvery ventrally; adipose
fin with a black spots in some species.
Remarks: occurring in tropicalto temperate marine, estuarine, andfreshwater areas of the world, usuallyfound in coastal and estuarine habitats,abundant in mangrove areas and large
river estuaries. Omnivorous, usually
feed on crustaceans, mollusks, andfishes. Males incubating the fertilized
eggs in its mouth cavity. Food fish.
Similar families occurring in thearea: Ariidae is distinguished fromother Indo-Pacific catfish families inhaving a long, usually serrated dorsal
fin spine, deeply forked caudal fin,short-based anal fin, adipose fin justabove anal fin, naked body, and nonasal barbells. Plotosidae – caudal
fin confluent with long-based anal
fin; no adipose fin; nasal barbells
present. Bagridae (freshwater) – nasal barbells present. Siluridae andClariidae (both freshwater) – nasal
barbells present; no dorsal fin spine;
long-based anal fin. Pangasidae(freshwater) – long-based anal fin.

Arius maculatus(Thunberg, 1792)
Spotted Catfish
D I, 7; A 19–23; P1 I, 9–10; P2 6; GR
5–7 + 11–14; free vertebrae 41–44.
Body elongated, compressed; head depressed; head shield and supraoccipital process weakly striated and granulated; dorsomedian head groove
narrower and deeper posteriorly; snout
rounded; palatal teeth small conicalwith blunt tip or molariform, on a single pair of oval patches, located wellposterior to upper oral valve; inner gillrakers present on all gill arches; lateralline bifurcates behind caudal fin base.
Color: body silvery, brownish green
dorsally; all fins somewhat brownish;adipose fin pale with a prominent darkmarking. Size: maximum length 50
cm. Distribution: Vietnam, Gulf ofArius maculatus, KAUM–I. 16925, 15.3 cm SL
off Terengganu (KT), 10 Dec. 2008

Thailand, Pacific coasts of PeninsularThailand and Malaysia, Borneo, Sumatra, and Java. Remarks: inhabits
coastal waters and estuaries.

43

Hexanematichthys sagor(Hamilton, 1822)
Sagor Sea Catfish
D I, 7; A 16–19; P1 I, 11; P2 6; GR
12–18; V 53–55. Body elongated,compressed; head broad, depressed;
head shield striated; supraoccipital
hemispherical in adults, its apex convex; dorsomedian head groove short;
palatal teeth conical, tips sharp, on 2
pairs of patches; inner patches smaller
than outer; barbels flattened, straplike; inner gill rakers absent on the anterior 2 gill arches; pectoral-fin spine
longer than dorsal-fin spine. Color:body silvery, blue-brownish dorsally;
several golden green vertical stripes
on body; adipose fin pale. Size: maximum length 45 cm SL. Distribution:
Indo-West Pacific, from Pakistan to
Borneo and Java. Remarks: inhabits
coastal waters and estuaries.

Netuma thalassina(Rüppell, 1837)

Hexanematichthys sagor, KAUM–I. 17013, 40.0 cm SL
off Terengganu (KT), 15 Dec. 2008

Hexanematichtys sagor, UMTF 1071 (KAUM–I. 16679), 10.7 cm SL
Kemaman, 21 Oct. 2008

Giant Sea Catfish
D I, 7–8; A 14–19; P1 I, 11–13; P2 6;
GR 4–5 + 8–10; free vertebrae 38–45.
Body elongated, compressed; head depressed; head shield and supraoccipital process finely granulated; dorsomedian head groove short and shallow,
V-shaped posteriorly; palatal teeth
minute, tips sharp, on 3 pairs of patches, located just posterior to upper oral
valve; inner patches on both sides
separated; inner gill rakers absent on
the anterior 2 gill arches; caudal fin
lobes somewhat long, slender; adipose
fin small; lateral line bifurcates behindcaudal fin base. Color: body silvery,
brownish dorsally; adipose fin with ablack marking. Size: maximum length
130 cm SL. Distribution: widely in
Indo-West Pacific, from East Africa toeastern Australia, north to Philippines
and southern China. Remarks: inhabits coastal waters.

Plicofollis dussumieri(Valenciennes, 1840)
Blacktip Sea Catfish
D I, 7; A 14–18; P1 I, 10; P2 6; GR
14–16. Body elongated, compressed.
Head depressed; head shield and supraoccipital process weakly granulated posteriorly; dorsomedian groove
distinctly deeper anterior to supraoccipital process; supraoccipital process

Netuma thalassina, KAUM–I. 16969, 19.0 cm SL
off Terengganu (KT), 11 Dec. 2008

Plicofollis dussumieri, KAUM–I. 17072, 14.9 cm SL
off Terengganu (KT), 27 Dec. 2008

triangular with sharp median keel;
palatal teeth granular or conical withblunt tips, on 2 pairs of patches longitudinally arranged; anterior patch
small and oval; posterior patch elongate, slender; inner gill rakers absent
on the anterior 2 gill arches. Color:body silvery, brownish dorsally; allfins dusky; adipose fin distally blackish or with a black marking. Size:maximum length 75 cm. Distribution: Indian Ocean, Pacific coast ofPeninsular Thailand and Malaysia,
Borneo, and Sumatra. Remarks: inhabits coastal waters and estuaries.

44

SYNODONTIDAE
Lizardfishes

By Mizuki Matsunuma and Seishi Kimura
Small to medium-sized marine
fishes; maximum size about 70 cm,
commonly 20–40 cm. Body slender,
cylindrical, and moderately elongated. Mouth large; hind tip of upper
jaw extending well beyond hind margin of eye; a single supramaxilla
small or absent; numerous needlelike teeth on jaws and palatines;
branchiostegal rays 12–18. No spinous fin rays; dorsal fin located about
midpoint of body, 9–15 rays; anal fin
posterior to dorsal fin base, 8–17
rays; caudal fin forked, 19 principal
rays; pelvic fins close together, 8–9
rays; adipose fin present above analfin. Cycloid scales present at least on
the posterior half of body and along
lateral line. Color: body usually

9–18 dorsal-fin rays
mouth large

adipose fin present

numerous needle-liketeeth on jaws andpalatines

7–18 anal-fin rays

brown or reddish with variable markings. Peritoneum either pale with several black spots, or black.
Remarks: occurring in tropical totemperate marine and estuarine areas,usually found on rocky, coral, sandy,or muddy bottoms in shallow waters.
Feed on small fishes and crustaceans.
Similar families occurring in thearea: Aulopidae – two supramaxillae

present; hind tip of upper jaw not orslightly extending beyond hind marginof eye. Bathysauridae – head depressed
conspicuously; branchiostegal rays
8–13; adipose fin present or absent;
pelvic fin rays 8. Chlorophthalmidae– a single elongate supramaxilla
present; hind tip of upper jaw not extending beyond center of eye.

Harpadon nehereus(Hamilton, 1822)
Bombay Duck
D 10–13; A 13–15; P1 10–12; P2 9.
Body elongate, well compressed;
mouth large; lower jaw longer thanupper jaw. Scales restricted to posterior half of body; lateral line extending
as a median lobe of caudal fin. Pectoral fin very long, its posterior tip extending beyond origin of dorsal fin;dorsal and pelvic fins large. Color:body cream, with black pigments dorsally; fins blackish. Size: maximum
40 cm SL. Distribution: widely distributed in Indo-West Pacific from India to Solomon Islands, north tosouthern Japan. Remarks: found incoastal waters of sandy mud bottom.
Marketed extensively as a relish withcurry.
(M. Matsunuma)

Harpadon nehereus, KAUM–I. 17293, 7.9 cm SL
off Terengganu (KT), 20 Jan. 2009

Saurida tumbil(Bloch, 1795)

Saurida tumbil, KAUM–I. 17241, 27.8 cm SL
off Terengganu (KT), 15 Jan. 2009

Greater Lizardfish
D 11–13; A 10–11; P1 13–16; P2 9;
LL 59–65; PDS 16–19; V 56–61.
Body subcylindrical, elongate. Headand caudal peduncle somewhat depressed; mouth large; numerous small
teeth exposed on side of jaws whenmouth closed; palatine teeth arranged
in two bands on each side of roof ofmouth; palatine teeth on the outerbands in 3 or 4 rows anteriorly. Outermost ray of pelvic fin subequal to innermost ray. Pectoral fin short, its posterior tip not or only just reaching toinsertion of pelvic fin. Color: bodylight brown dorsally, silvery whiteventrally; pectoral fin and lower lobeof caudal fin dusky; upper margin of
caudal fin without any marking; adi-pose fin with a black marking. Size:maximum length about 25 cm SL.
Distribution: widely distributed in
the Indo-West Pacific from east coastof Africa, Red Sea to eastern Australia. Remarks: found in shallow coastal sandy or muddy bottoms. Feeds on
fishes, crustaceans, and squids. Marketed fresh.
(S. Kimura)

45

Saurida undosquamis(Richardson, 1848)
Brushtooth Lizardfish
D 10–13; A 10–11; P1 13–14; P2 9;
LL 49–55; PDS 18–21; V 50–53.
Body subcylindrical, elongate. Headand caudal peduncle somewhat depressed; mouth large; numerous small
teeth exposed on side of jaws whenmouth closed; palatine teeth arranged
in two bands on each side of roof ofmouth; palatine teeth on the outerbands in only 2 rows anteriorly. Outermost ray of pelvic fin subequal to innermost ray. Pectoral fin short, its posterior tip not reaching a line between
origins of pelvic and dorsal fins. Color: body light brown dorsally, silvery
white ventrally; pectoral fin and lower
lobe of caudal fin dusky; upper marginof caudal fin with 6–16 black dots;
adipose fin with a black marking.
Size: maximum length about 28 cm
SL. Distribution: eastern Indian
Ocean and West Pacific from Arabian
Gulf to northern and southwestern
Australia, north to Gulf of Thailand.
Remarks: found in shallow coastal
sandy or muddy bottoms.
(M. Matsunuma)

Saurida undosquamis, KAUM–I. 17242, 24.7 cm SL
off Terengganu (KT), 15 Jan. 2009

Synodus tectus, KAUM–I. 17094, 14.4 cm SL
off Terengganu (KT), 28 Dec. 2008

Synodus tectus(Cressey, 1981)
Tectus Lizardfish
D 13–14; A 9–10; P1 12–13; P2 8; PLS
55–57; PDS 13–15; V 54–56. Body fusiform, elongate, caudal peduncle
slightly compressed. Head somewhat
depressed; mouth large; anteriormost
palatine teeth longer than more posterior teeth. Outermost ray of pelvic fin
distinctly shorter than innermost ray;anal fin base shorter than dorsal finbase; pectoral fin short, its posterior tip
not reaching a line between origins ofpelvic and dorsal fins. Color: bodylight brown dorsally, silvery white ventrally; with about 8 brown saddle-likepatches extending laterally, forming
diamond-shaped patches at level of thelateral line; a black spot on upper edgeof opercle, divided dorsally into 3 or 4
separate finger-like branches; 9–11
peritoneal spots. Size: maximum
length 17 cm SL. Distribution: West
Pacific from northern Australia to Taiwan. Remarks: found in shallow
coastal sandy or muddy bottoms.
(M. Matsunuma)

Synodus tectus, UMTF 1891, 16.7 cm SL
off Terengganu (KT), 11 Jan. 2009

Trachinocephalus myops, UMTF 1360 (KAUM–I. 16572), 16.6 cm SL
Bidong Island, 13 Oct. 2008

Trachinocephalus myops(Forster, 1801)
Bluntnose Lizardfish
D 11–14; A 13–18; P1 11–13; P2 8;
LL 51–61; V 54–55. Body fusiform,
elongate; snout very short; mouthlarge; palatine with a row of teeth.
Outermost ray of pelvic fin distinctly
shorter than innermost ray; anal finbase longer than dorsal fin base; pectoral fin short. Color: body with alter-

nating narrow dark-edged pale blue
and yellow stripes; a large black spot
at upper end of gill opening; about 5
yellow stripe on dorsal fin; adipose
and anal fins with yellow margin; pelvic fin with a yellow stripe basally;
caudal fin yellow. Size: maximum
length 25 cm SL. Distribution: circumglobal in tropical and subtropicalseas except for Eastern Pacific. Remarks: found in shallows to 400 m on
sandy bottom.
(M. Matsunuma)

46

CARAPIDAE
Pearlfishes

By Hiroyuki Motomura and Mizuki Matsunuma
Small to medium-sized fishes,
rarely reaching 50 cm TL. Body
elongate and compressed or subcylindrical. Eye diameter subequal tosnout length in adults. Supramaxilla
absent. Mouth terminal, often withstrong teeth. No spines on opercle.
Scapula and coracoid fused; upperdistal radial of pectoral fins enlarged;
hyomandibula with large foramen.
Dorsal and anal fins long, connected
to caudal fin; dorsal fin rays shorter
than opposing anal fin rays; highly
modified first dorsal fin ray in larvae.
Pelvic fin with a soft ray, or without
ray. Scales absent. Anus located below pectoral fins. Color: semitransparent to brownish.
Remarks: occurs in shallow tomoderately deep waters. Usually pe-

dorsal fin long, connected to caudal finmouth terminal
no scales

anal fin long, connected to caudal fin
anus below pectoral finslagic in larvae and benthic in adults;
free-living adults in some species;
commensals in body cavity of invertebrates, primarily holothurians (sea cucumbers), in most species when adults.
A bycatch in some sea cucumber fish-eries, but no importance to fisheries.
Similar families occurring in
the area: Ophidiidae – body scales
present; anus located behind posterior tip of pectoral fin.

Encheliophis homei(Richardson, 1846)
Silver Pearlfish
P1 17–21; P2 0; V 116–128; Body
elongate, compressed; head depth
slightly greater than body depth.
Mouth moderate; maxilla free fromcheek, movable; villiform teeth onjaws, without fangs. Dorsal fin origin
posterior to anal fin origin; pectoral
fin relatively long, its length subequal
to upper jaw length. Anus located anterior to vertical through at pectoral
fin base. Swimbladder without ridges
an intrinsic constriction. Color: bodytranslucent; cheek and abdomen silvery; side of jaws with some blackish
pigment. Size: maximum length 19
cm. Distribution: widely distributed

Encheliophis homei, KAUM–I. 17186, 10.5 cm SL
off Terengganu (KR), 5 Jan. 2009

Encheliophis homei, KAUM–I. 17187, 10.6 cm SL
off Terengganu (KR), 5 Jan. 2009

in the Indo-Pacific, from Madagascar
east to Society Islands, and northern
Australia north to southern Japan. Remarks: commensal in sea cucumbers.
(M. Matsunuma)

47

OPHIDIIDAE
Cusk Eels

By Mizuki Matsunuma
Median to large-sized marine
fishes, reaching 200 cm. Body moderately elongate. Anterior nostril
placed midway between upper lip
and posterior nostril. Teeth usuallysmall, densely distributed, and blunttipped. Supramaxilla present. Very
seldom less than 7 long gill rakers on
anterior gill arch. A well developed
spine on opercle usually present.
Dorsal and anal fins long, joined tocaudal fin. Dorsal fin rays normallylonger than opposing anal fin rays.
Pelvic fin rays 0–2. Scales present;
lateral line present or absent. Anusplaced posterior to pectoral fins except in species with prolonged pectoral fins. Color: very variable, somewith horizontal or vertical bars andeye-spots.
Remarks: most of species occurs pelagically at great depths; thefew shallow water species are cryp-spine on opercle

pelvic fin rays 0–2
anus usually posterior to pectoral finstic. Usually no commercial importance.
Similar families occurring in thearea: Carapidae – scales absent; analfin rays longer than opposing dorsal-finrays; anus placed below pectoral fins.
Bythitidae – anterior nostril placed immediately above upper lip; very seldom

more than 7 long gill rakers on anterior gill arch. Macrouridae – pelvic
fins well separated from each other,with more than 2 rays. Gadidae andMoridae – pelvic fins well separated
from each other; dorsal and anal fins
not joined to caudal fin.

Sirembo jerdoni(Day, 1888)
Brownbanded Cuskeel
D 89–95; A 64–68; P1 22–24; P2 1.
Body relatively elongate, compressed,
but not attenuate; snout bluntly rounded; eyes well developed. Dorsal andanal fins long, joined to caudal fin;pelvic fins closed to each other, located below level of posterior margin ofeye. No spines on preopercle; spine on
opercle short, not reaching posterior
margin of opercle. Scale cycloid. Color: body silvery white, with 3–4
broad, oblique bands on head and anterior body, connecting over the predorsal and head to the other side; several brown blotch on dorsal and caudal
fins; anal fin with a brown stripe basally, blackish marginally. Size: commonly 10 cm SL. Distribution: IndoWest Pacific, including the Red Sea,
Bay of Bengal,Western Australia, Gulfof Thailand, Philippines, and Southand East China Sea. Remarks: foundin moderate to deep waters at depthsof 57–99 m.

Sirembo jerdoni, KAUM–I. 17170, 14.2 cm SL
off Terengganu (KR), 5 Jan. 2009

Sirembo jerdoni, KAUM–I. 17028, 10.8 cm SL
off Terengganu (KT), 17 Dec. 2008

48

ANTENNARIIDAE
Frogfishes

By Hiroyuki Motomura
Small to moderate fishes, largest
species attaining to over 50 cm; most
species less than 20 cm. Body short,
deep, globose, slightly compressed.
Mouth large, strongly oblique to vertical; villiform teeth on jaws andpalatines. Gill opening, below or behind pectoral-fin base, with a small
pore. Dorsal fin with III spines and
10–16 soft rays; spinous portion of
fin widely separated from soft-rayed
portion; first spine free from rest offin, nearly always with a well-developed terminal bait (esca); second andthird spines covered by thick skin.
Anal fin with 6–10 soft rays. Caudal
fin rounded. Pectoral fin with 6–14
rays, elongate, leg-like. Pelvic fin
with I spine and 5 soft rays. Body
covered with loose skin, naked orwith denticles. Color: extremely
variable, often closely matching thatof surroundings; changing color

dorsal fin III, 10–16

esca

mouth large, strongly
oblique to vertical

6–14 pectoral-fin rays

6–10 anal-fin rays
pelvic fin I, 5

within a few weeks in some species.
Remarks: most species inhabiting
on bottom in shallow waters; one species pelagic, clinging in floating algae.
Feeds on fishes, sometimes large fishes longer than themselves; also canni-

balistic. Released eggs embedded ina single, large, buoyant gelatinous
mass; floating on water surface.
Similar families occurring in thearea: families with an esca not occurring in shore waters in the area.

Antennarius nummifer(Cuvier, 1817)
Spotfin Frogfish
D I + I + I, 12–13; A 7–8; P1 10–11;
P2 I, 5; C 9. Body globose, compressed. Gill opening below pectoralfin base. First dorsal-fin spine with afleshy tentacle; base of the spine behind upper jaw symphysis; second
spine without membrane posteriorly;
illicium short, subequal in length tosecond dorsalfin spine. Posterior endof dorsal fin not broadly connected bymembrane to caudal peduncle. Pectoral-fin rays unbranched. Color: bodyred, pink, orange, yellow, tan, or
brown, with a large black spot at dorsal-fin base. Size: maximum length 13
cm. Distribution: widely distributedin the Indo-Pacific, from east coast of
Africa east to the Hawaiian and Marquesas Islands, and also from islandsof the eastern Atlantic. Remarks: usually found on reefs in shallow waters
to a depth of 293 m.

Antennarius nummifer, KAUM–I. 16629, 1.4 cm SL
Bidong Island, 13 Oct. 2008

49

MUGILIDAE
Mullets

By Koichi Shibukawa, Siti Tafzilmeriam S. A. K. and Seishi Kimura
Medium to large sized (up to 100cm) moderately slender fishes. Body
subcylindrical, compressed posteriorly; lateral line absent. Head subcylindrical, flattened dorsally in many
species; eyes often partly coveredby well-developed adipose eyelid;mouth moderately small, terminal orinferior; teeth minute or absent. Two
dorsal fins well separated from one
another; first dorsal fin with IV pungent spines, the anterior three close
together at base; second dorsal fin
with I spine and usually 9–10 softrays; anal fins with II–III spines and
7–12 soft rays; pectoral fins high onside of body; pelvic fins subabdominal, with I spine and 5 rays; caudal
fin lunate, emarginate or nearlytruncate. Scales cycloid or ctenoid,
moderately large; enlarged axillary
scales usually present at base of pectoral and pelvic fins. Color: head andIV first dorsal fin spines; the anteriror
3 close together at baselateral line absent

II–III anal-fin spines

body more or less silvery, darkened
dorsally; fins transparent, dusky or
yellowish.
Remarks: fast swimming fishes;
inhabits coastal waters (usually todepths of 20 m) including estuaries;
often enter freshwater areas and lagoons. Frequently schooling. Feed on
detritus or small organisms. Many species are important to fisheries, caught
by various nets (e.g., seines, cast nets,

and gill nets) and usually marketed
fresh, frozen, salted or boiled; roe
may be also marketed; often used aslive bait in tuna fishing. Generic assignment follows Senou (2002) andGhasemzadeh et al. (2004).
Similar families occurring in thearea: Atherinidae – first dorsal fin
with III–VIII flexible spines, anterior
three spines separate at base; a single
anal-fin spine.

Chelon subviridis(Valenciennes, 1836)
Greenback Mullet
D IV + 8–10; A III, 8–9; P1 14–17;
LR 27–33. Body moderately robust(depth usually 22–26% of SL), compressed posteriorly; no elongate scale(axillary scale) above pectoral-finbase. Head width subequal to depth,
flattened dorsally; adipose eyelid
weakly developed posteriorly, notcovering iris; hind tip of maxilla exposed when mouth closed. Second
dorsal and anal fins pointed but not
falcate; caudal fin emarginate. Scales
ctenoid. Pyloric caeca 4–6. Color:head and body silvery, darkened dorsally; 3–6 indistinct, dusky longitudinal stripes on body; caudal fin bluishwith dusky narrow margin. Size: 40
cm SL, commonly to 25 cm TL. Distribution: Indo-Pacific. Remarks:
found in shallow coastal waters; frequently enters lower reaches of
streams, estuaries, and mangroveswamps. Marketed fresh and salted.
(K. Shibukawa & S. Tafzilmeriam)

Chelon subviridis, KAUM–I. 17064, 19.0 cm SL
off Terengganu (KT), 26 Dec. 2008

Chelon subviridis, UMTF 1025 (KAUM–I. 16788), 11.9 cm SL
Merang, 2 Nov. 2008

50

Ellochelon vaigiensis(Quoy & Gaimard, 1825)
Squaretail Mullet
D IV + 7–9; A III, 8; P1 15–18; LR
25–29. Body moderately robust (depth
usually 20–27% of SL), compressed
posteriorly; no elongate scale (axillary
scale) above pectoral-fin base. Headwider than deep, flattened dorsally;
adipose eyelid poorly developed; hind
tip of maxilla exposed when mouthclosed. Second dorsal and anal finspointed but not falcate; caudal fintruncate. Scales ctenoid. Pyloric caeca
much divided, up to 23 in about 2bunches. Color: head and body silvery, darkened dorsally; 5–6 indistinct, dusky longitudinal stripes onbody; pectoral fin entirely black; analand caudal fins yellowish. Size: 60 cm
SL, commonly to 35 cm TL. Distribution: Indo-Pacific. Remarks: foundin shallow coastal waters, and enters
estuaries. Marketed fresh and salted.
(K. Shibukawa & S. Tafzilmeriam)

Moolgarda delicata(Alleyne & Macleay, 1877)
Delicate Mullet
D IV + 8; A III, 9; P1 15–17; LR
34–36. Body elongate; mouth small
and terminal; snout rounded; posterior
end of maxilla concealed when mouthclosed; adipose eyelid covering noneof iris; scales cycloid with crenulated
membranous edge; pyloric caeca notbranching; caudal fin forked. Color:greenish or brownish gray dorsally,shading silvery on side and ventrally;
posterior margin of caudal fin black; adark blue spot at origin of pectoral fin.
Size: about 13 cm SL. Distribution:
western Pacific. Remarks: occurringin beaches and estuaries. AlthoughThomson (1997) treated this nominal
name as a synonym of Valamugil seheli(Forsskål, 1775), the former is considered as valid species (H. Senou pers.
com.).
(S. Kimura)

Ellochelon vaigiensis, KAUM–I. 17314, 24.8 cm SL
off Terengganu (KT), 20 Jan. 2009

Ellochelon vaigiensis, UMTF 1314 (KAUM–I. 16684), 4.1 cm SL
Kemaman, 21 Oct. 2008

Moolgarda delicata, UMTF 1293 (KAUM–I. 16454), 8.6 cm SL
estuary near UMT, 28 Sept. 2008

Moolgarda delicata, UMTF 1229 (KAUM–I. 16789), 7.1 cm SL
Merang, 2 Nov. 2008

51

Moolgarda perusii(Valenciennes, 1836)
Longfinned Mullet
D IV + 8–10; A III, 9; P1 15–17; LR
31–34. Body moderately robust (depth
24–35 % SL), compressed posteriorly.
Head deeper than wide and dorsally
flattened; adipose eyefold developed,
extending over iris; hind tip of maxillaconcealed when mouth closed. Second
dorsal and anal fins long, but not falcate; caudal fin emarginate. Scaleswith a digitated membranous posterior
margin; thoracic and abdominal scales
more distinctly ctenoid. Pyloric caeca
5–7. Color: greenish dorsally, flanks
and abdomen silvery; a small gold
spot on opercle; pectoral fins withdark spot dorsally at origin. Size:maximum 25 cm TL, commonly to 15
cm TL. Distribution: Indo-West Pacific. Remarks: found in coastal waters and estuaries. (S. Tafzilmeriam)

Moolgarda perusii, KAUM–I. 17106, 15.0 cm SL
off Terengganu, 28 Dec. 2008

Oedalechilus labiosus(Valenciennes, 1836)
Hornlip Mullet
D IV + 8–9; A III, 9; P1 15–18; LR
33–37. Body moderately deep. Head
relatively flattened dorsally; adipose
eyefold absent; upper lip with deep
longitudinal fold near its ventral edge,
splitting it into upper and lower lobes;
these lobes fringed with ridges ofhorny epidermis; ridges small and appear as papillae on upper lobe; ridges
on lower lobe better developed; upperand lower lips form folds tucked underpreorbital at corners of mouth. Caudal
fin emarginate. Scales cycloid or
weakly ctenoid. Pyloric caeca 3–4.
Color: body olive dorsally, silvery
ventrally; fins greenish or yellowish.
Size: maximum 40 cm SL, commonly
to 20 cm TL. Distribution: Indo-Pacific. Remarks: found in shallow
coastal waters, coral reefs and in harbors. Caught with gill nets, lift nets,
and seines.
(S. Tafzilmeriam)

Moolgarda perusii, UMTF 1217 (KAUM–I. 16669), 7.7 cm SL
Kemaman, 20 Oct. 2008

Oedalechilus labiosus, UMTF 1230 (KAUM–I. 16623), 4.0 cm SL
Bidong Island, 12 Oct. 2008

52

ATHERINIDAE
Silversides

By Seishi Kimura
Small to medium sized marine
and freshwater fishes; maximum size
about 20 cm in the area, commonly
5–10 cm. Body slender cylindrical or somewhat compressed withround belly. Eyes large; interorbital
area flat; mouth small and terminal;hind tip of upper jaw not extending
to posterior margin of eye. Two dorsal fins well separated, the first with
III to VIII flexible spines separated
each other at base; the second dorsal and anal fins with a single spine
followed by soft rays (D2 I, 7–13; AI, 8–21); caudal fin forked; pectoral
fins high on the sides of body with
10–19 rays; pelvic fins abdominal tosubabdominal with I, 5. Scales cycloid, relatively large; lateral line absent. Color: body greenish or greenish brown dorsally, silvery ventrally;a broad silvery (black in preserved
specimens) stripe on the side of body.

III–VIII first dorsal-fin spines, their bases
separated

lateral line absent

mouth terminal

I anal-fin spine

Remarks: occurring in tropicalto temperate marine, estuarine, andfreshwater areas; usually found inbeaches and seagrass areas. Eggs withentangling filaments on chorion.
Similar families occurring in
the area: Atherinidae is distinguishedfrom the following similar families inhaving round belly, anus located behind pelvic fin base, 2 dorsal fins, firstdorsal-fin spines separated each otherat base, and a single anal fin spine.
Engraulidae – hind tip of upper jaw
extending beyond posterior marginof eye; a single dorsal fin; no spines
in fins. Isonidae – body compressedwith keeled belly. Mugilidae – firstdorsal fin with IV spines, anterior 3
spines close together at base; II or
III anal fin spines. Phallostethidae –anus anterior, located under head.

Atherinomorus lacunosus(Forster, 1801)
Wide-banded Hardyhead Silverside
D IV–VII + I, 9–11; A I, 12–16; P1
15–18; LR 40–44; LGR 18–24; V 20–
25 + 18–22 = 41–45. Body rather
stout, deep; anus near or usually beyond pelvic-fin tips; mouth large, upper jaw reaching to or slightly beyonda vertical through front margin of pupil; ascending process of premaxilla
short and broad; upper margin of thedentary almost flat distally, without adistinct tubercle at the posterior end;
posterior margin of lateral scales usually entire, but sometime crenulated.
Color: body greenish tan dorsally; abroad (ca. 1.5 scale width) silvery longitudinal band on side. Size: maximum length about 14 cm SL. Distribution: almost entire Indo-Pacific,
from East Africa to Tonga. Remarks:
occurring in beaches, sometimes entering estuaries. The genus Atherinomorus is characterized by having a

Atherinomorus lacunosus, KAUM–I. 16376, 7.3 cm SL
Dungun, 23 Sept. 2008

short and blunt ascending process anda low and wide lateral process of thepremaxilla. Seven species of Atherinomorus (A. aetholpis, A. crenolepis,
A. duodecimalis, A. endrachtensis, A.
lacunosus, A. pinguis, and A. regina)
are distributed in the West Pacificcoasts of Southeast Asian countries.
Of these, A. endrachtensis, A. lacunosus, and A. pinguis differ from the remaining four species by lacking a tubercle at the posterior end of dentary.
Although A. lacunosus closely resem-

bles A. pinguis in general morphology,
the former is distinguished from thelater in having a wider midlateral band(the lower margin reaching to almost
the center of the fourth scale row atlevel of the anal fin origin vs. the lower margin reaching to the ventral endof the third scale row in the later) andmore numerous midlateral scales (40–
44 vs. 38–41). Atherinomorus endrachtensis has 33–35 midlateral scales.

53

Atherinomorus pinguis(Lacepède, 1803)
Narrow-banded Hardyhead Silverside
D IV–VI + I, 9–11; A I, 12–16; P1
14–17; LR 38–41; LGR 19–23 V 20–
23 + 19–20 = 39–43. Body rather
stout, deep; anus usually just anterior
to pelvic-fin tips; mouth large, upper
jaw reaching to or slightly beyond a
vertical through front margin of pupil;
ascending process of premaxilla shortand broad; upper margin of dentaryalmost flat, without a distinct tubercle
at the posterior end; preopercle
notched; posterior margin of lateralscales usually entire, but sometime

Atherinomorus pinguis, UMTF 1070 (KAUM–I. 16523), 9.9 cm SL
Cendering, 6 Oct. 2008

crenulated. Color: body greenish tan
dorsally; a broad (1 scale width orless) silvery longitudinal band on side.
Size: maximum length about 14 cm
SL. Distribution: widely in Indo-West Pacific, from East Africa tonorthern Australia. Remarks: occurring in beaches, sometimes entering
estuaries.

EXOCOETIDAE
Flyingfishes

By Mizuki Matsunuma
Medium sized (up to about 40 cm)
marine fishes. Body elongate, cylindrical; flattened ventrally in somespecies. Head short; snout blunt,
shorter than eye in all Western Central Pacific species. Mouth small;jaws of equal size; teeth on jaws absent or very small. Gill rakers welldeveloped; upper pharyngeal bonesof third gill arches close together,but not fused into a single plate. Fin
spines absent; dorsal and anal fins set
equally far back on body, their bases
short and opposed; pectoral fins highon sides, strikingly long, always extending beyond dorsal-fin origin;pelvic fins abdominal in position,and greatly enlarged in many, but notall, species; caudal fin deeply forked,
its lower lobe longer than the upper.
Scale large, cycloid; lateral line pres-pectoral fins extremely longmouth small
lateral line low

ent on lower body. Young stages (toabout 10 cm) quite different in appearance from adults, with pectoral finsshorter, dorsal fin often higher thanin adults, color patterns variable, andspots and bars often developed; single
or paired chin barbels conspicuous inmany species. Color: dark dorsally,
pale laterally and ventrally; pectoral
fins in some species with dark spots or
pale stripes; dorsal fin in some species
with black pigment.

Remarks: inhabits surface watersof open ocean as well as neritic andinshore areas. Important food fish.
Similar families occurring in thearea: Hemiramphidae – pectoral
fins short to medium length, neverreaching dorsal-fin origin; lower jaw
much longer than upper jaw, except
in some genera; body more elongate; upper pharyngeals of third archfused, forming a single plate.

Cheilopogon katoptron(Bleeker, 1866)
Indonesian Flyingfish
D 13–14; A 9–10; P1 12–14; P2 6; GR
6–8 + 19–21 = 25–28; PDS 16–20; LL
38–40. Body elongate, cylindrical.
Mouth small and terminal; upper jaw
protrusible. Lateral line with pectoral
branch. Dorsal fin large, its origin justabove anal fin origin; pectoral fin relatively short, its posterior tip extending
beyond pelvic fin origin, but notreaching end of anal fin base; caudal
fin well forked, length of lower lobe
longer than upper lobe. Color: body

Cheilopogon katoptron, KAUM–I. 17095, 16.1 cm SL
off Terengganu (KT), 28 Dec. 2008

silver, green to blue dorsally; dorsal northern Australia north to Viet Nam.
fin entirely dark to black; pectoral fin Remarks: pelagic in nearshore surwith large black area; lower caudal fin face waters; never spread to open sea.
lobe dusky. Size: maximum 18 cm SL.
Distribution: western Pacific from

54

Parexocoetus mento(Valenciennes, 1847)
African Sailfin Flyingfish
D 9–11; A 10–12; P1 12–14; P2 6;
PDS 23–26. Body elongate, cylindrical. Mouth small and terminal; bothjaws of equal length; teeth on jaws noticeable (conspicuous to the touch).
Lateral line with pectoral branch. Dorsal fin large, its origin well before anal
fin origin; pectoral fin long, its posterior tip reaching end of anal fin base,with an uppermost ray unbranched;caudal fin well forked, length of lower
lobe much longer than upper lobe.
Color: body silver, green to blue dorsally; dorsal fin uniformly pale; pecto-

Parexocoetus mento, KAUM–I. 17080, 9.7 cm SL
off Terengganu (KT), 27 Dec. 2008

ral fins with pale oblique cross band,no dark spots; no black spot on pelvic
fin Size: maximum 10 cm SL. Distribution: widely distributed in IndoWest Pacific, from Red Sea and East

Africa to Marshall, Fiji, andQueensland, north to southern Japan.
Remarks: pelagic in neritic surface
waters, never spread to the open sea.

HEMIRAMPHIDAE
Halfbeaks

By Koichi Shibukawa and Mizuki Matsunuma
Small to medium sized (up to 45cm) elongate fishes with long, javelin-like lower jaw, projecting far
beyond upper jaw. Body elongate
and compressed; lateral line runningalong ventral margin of body witha branch to origin of pectoral fin.
Lower jaw usually long and javelinlike, whereas upper jaw short andtriangular; eyes large; several rowsof small teeth on jaws; gill rakers
0–78. Dorsal and anal fins positioned
at posterior part of body; dorsal fin
with 8–25 soft rays; anal fin with
8–19 soft rays; no isolated finlets
behind dorsal and anal fins; pectoral
fin usually short, with 7–14 soft rays;
pelvic fins abdominal, with 6 rays;

upper jaw shortpectoral fin shortlarge scales

lower jaw extremely longcaudal fin rounded, truncate, emarginated or deeply forked. Scales moderately large, cycloid. Color: head andbody silvery, darkened dorsally; tip oflower jaw tinged with red or orange inmany species; fins hyaline, often withblackish markings.
Remarks: surface-swimming fishes,
chiefly found in marine and brackish
waters, whereas some in freshwaters.
Omnivorous, feed on floating sea-

grass pieces, insects, crustaceans,and small fishes. Large species commonly esteemed as food fish, caught
by seine and gill nets; some freshwater species often treated as aquarium
fish.
Similar families occurring in thearea: Belonidae – usually both jaws
elongate; scale small. Exocoetidae– pectoral fin very large; jaws short.

Hemiramphus archipelagicus
Collette & Parin, 1978
Jumping Halfbeak
D 12–15; A 10–13; P1 10–13; GR 6–8+ 19–24 = 25–32. Body elongate, subcyrindrical. Lower jaw greatly prolonged, beak-like (broken in photographed specimen); upper jaw shortand triangular, without scale; preorbital ridge (bony ridge behind nostril)
absent. Dorsal fin without developed
anterior lobe; pectoral fins relatively
short, not reaching posterior nasal pit
when folded forward; caudal fin
rounded forked, lower lobe longer

Hemiramphus archipelagicus, KAUM–I. 16924, 16.1 cm SL (upper jaw broken)
off Terengganu (KT), 10 Dec. 2008

than upper lobe. Color: body silvery
white, dark bluish dorsally; a dark
stripe with black upper margin on laterally; no vertical bars on body side;
beak dark with a bright red fleshy tip;
caudal fin dusky yellow, tip of dorsal-

fin lobe with some yellow. Size: maximum about 23 cm SL. Distribution:
western Indian Ocean and western
Central Pacific. Remarks: found incoastal waters. Marketed fresh anddried salted.
(M. Matsunuma)

55

Zenarchopterus dunckeri
Mohr, 1926
Duncker's River Garfish
D 10–12; A 10–13; P1 10; V 37–42.
Body elongate, subcyrindrical. Lower
Zenarchopterus dunckeri, UMTF 1491 (KAUM–I. 16750), male, 6.7 cm SL
jaw much prolonged, javelin-like; upSetiu, 30 Oct. 2008
per jaw wider than long; nasal papillaelongate and pointed. Base of dorsal
fin longer than anal-fin base; 6th anal
ray of males greatly thickened andelongated, reaching beyond caudal
base; 4th and/or 5th dorsal ray(s) elongated; caudal fin nearly rounded. ColZenarchopterus dunckeri, UMTF 1492 (KAUM–I. 16751), female, 4.9 cm SL
or: light brown dorsally, becoming
Setiu, 30 Oct. 2008
silvery white ventrally; no longitudinal black line on dorsal midline of upper jaw. Size: 12.5 cm SL. Distribu- cially mangrove area. Surface swimtion: Andaman Sea and West Pacific. ming fish, feeds on small terrestrial
Remarks: found in estuaries, espe- insects.
(K. Shibukawa)

BELONIDAE
Needlefishes

By Mizuki Matsunuma
Medium to large sized (up to 2 m)elongate fishes. Body elongate, withboth jaws extended into long beaks
filled with sharp teeth; nostrils in a
pit anterior to eyes. No spines in fins;dorsal fin single with 11–43 rays;
anal fin with 12–39 rays; dorsal andanal fins posterior in position; pelvic fins located in abdominal position, with 6 rays; pectoral fins shortwith usually 10–20 rays; caudal fin
variable, rounded to forked. Scales
small, cycloid; lateral line running
down from pectoral-fin origin andthen along ventral margin of body.
Color: these fishes live at the surface and are protectively colored
for this mode of life by being green

11–43 dorsal-fin soft rays

both jaws extremely longsmall scales

pectoral fin shortor blue on back and silvery white on
lower sides and belly; usually, a dusky
or dark blue stripe along sides; tip oflower jaw frequently red or orange.
Remarks: mostly found in marine,
but some species occur in freshwater.
Carnivorous, feeding largely on small
fishes. Many species are important for
fisheries; caught by casting or trollingsurface or near-surface lures.

12–39 anal-fin soft rays

Similar families occurring in thearea: Hemiramphidae – only the lower jaw prolonged or neither jaw prolonged and lacking the needle-sharp
teeth that stud the upper and lower
jaws of needlefishes. Sphyraenidae –jaws pointed but not prolonged intoa beak; 2 dorsal fins, the first spiny;
pelvic fins in thoracic position.

Ablennes hians(Valenciennes, 1846)
Flat Needlefish
D 23–26; A 24–28; P1 13–15; PDS
340–430; V 87–93. Body elongate,
greatly compressed laterally; no caudal peduncle keel; lateral line without
pectoral branch; jaws greatly elongate,
studded with small sharp teeth; gillrakers absent. Dorsal and anal finswith high falcate lobes anteriorly; origin of dorsal fin posterior to anal fin
origin; pectoral fin falcate; caudal fin

Ablennes hians, KAUM–I. 16880, 43.6 cm SL
off Terengganu (KT), 5 Dec. 2008

deeply forked, lower lobe much longer than upper lobe. Color: body silvery white, bluish green dorsally; anindistinct dark blue stripe along sides;
about 12–14 prominent dark verticalbars on sides; tip of lower jaw red;scales and bones green. Size: maximum 120 cm TL. Distribution: widely distributed in tropical to warmtemperate waters in worldwide. Remarks:
found in offshore surface waters. Marketed fresh.

56

Strongylura strongylura(van Hasselt, 1823)
Spottail Needlefish
D 12–15; A 15–18; P1 10–12; PDS
100–130; V 59–65. Body elongate, belly rounded; no caudal peduncle keel;lateral line with pectoral branch; jaws
greatly elongate, studded with sharp
teeth; gill rakers absent. Dorsal and anal
fins with moderate lobes anteriorly; origin of dorsal fin posterior to anal fin
origin; pectoral fin not falcate; caudal
fin rounded or truncate. Color: bodysilvery white, grayish to greenish dorsally; no markings on body sides; caudal fin with a prominent round black
spot basally. Size: 40 cm SL. Distribution: Indo-West Pacific. Remarks:
found in coastal areas and mangrove.

Strongylura strongylura, KAUM–I. 16792, 30.8 cm SL
Merang, 2 Nov. 2008

Tylosurus crocodilus crocodilus, KAUM–I. 17261, 71.0 cm SL
off Terengganu (KT), 17 Jan. 2009

271–340; V 82–86. Body elongate,
belly rounded; a distinct lateral keel
on caudal peduncle; lateral line with
pectoral branch; jaws greatly elongate,
studded with sharp teeth; gill rakers
Tylosurus crocodilus crocodilus absent. Dorsal and anal fins with relatively high lobes anteriorly; origin of(Péron & Lesueur, 1821)
dorsal fin posterior to anal fin origin;
Hound Needlefish
pectoral fin rounded; caudal fin well
D 21–25; A 19–22; P1 13–15; PDS forked, lower lobe longer than upper

lobe. Color: body silver, dark bluishgreen dorsally; a dark blue stripe along
body sides; anterior lobes of dorsal
and anal fins black; caudal fin black;scales and bones green. Size: 120 cm
SL. Distribution: widely distributed
in tropical to warm temperate waters
worldwide, except for the eastern Pacific. Remarks: found in coastal andoffshore waters areas. Marketed fresh.

APLOCHEILIDAE
Asian Rivulines

By Koichi Shibukawa
Small sized (up to 10 cm SL),
slender and compressed fishes. Anus
just before origin of anal fin; lateralline absent. Eyes large; mouth terminal, protrusible. Single dorsal fin,located at posterior part of body, with
7–20 soft rays; anal fins with 15–19
soft rays; length of anal-fin base longer than dorsal-fin base; pectoral finsrounded, with 14–21 soft rays; pelvic
fins abdominal, with 6 soft rays; caudal fin rounded. Scales cycloid, small
to moderately large. Color: highlyvariable, appearing characteristic

mouth protrusible

single dorsal fin

caudal fin roundedlong anal-fin basepattern in each species.
Remarks: surface-swimming schooling fish, found in fresh- or brackishwaters. Oviparous.
Similar families occurring in thearea: Adrianichthyidae – pectoral fin

falcate (rather than rounded); mouth
non-protrusible. Phallostethidae –small first dorsal fin present in many
species; pelvic fins absent; caudal fin
emarginate. Poeciliidae – anal-finbase rather short.

Aplocheilus panchax(Hamilton, 1822)
Blue Panchax
D 7–8; A 15–16; P1 14; LR 30–34.
Body moderately slender and compressed. Eyes large; mouth small, protrusible; lower jaw projecting beyond
upper jaw. Base of anal fin much longer
Aplocheilus panchax, UMTF 1106 (KAUM–I. 16383), 3.9 cm SL
estuary near UMT, 23 Sept. 2008
than dorsal-fin base. Caudal fin rounded. Color: light grayish brown dorsally, whitish ventrally; edge of scale Distribution: South and Southeast estuaries and around wetland, ponds,
pockets dusky; a distinct jet-black spot Asia from Pakistan to the Indo-Malay- ditches and creeks with dense waterat base of dorsal fin. Size: 6 cm SL. sian Archipelago. Remarks: found in plants.

57

HOLOCENTRIDAE
Soldierfishes

By Mizuki Matsunuma
Medium sized (up to 45 cm SL)
marine fishes. Body ovate to moderately elongate, compressed. Headwith ridges and mucous channels
dorsally; edges of external bonesof head serrate or with spines; eyevery large; mouth moderately large,
terminal or with lower jaw projecting; 2 supramaxillae; small villiform
teeth in bands in jaws and on roof ofmouth (on vomer, palatines, and forsome species on ectopterygoids).
Dorsal fin base long, with IX–XIII
stout spines and 12–17 soft rays;
deeply notched between spinous andsoft portions or between last 2 dorsal
fin spines; anal fin with IV spines,
the third stoutest and often longest,and 7–16 soft rays; caudal fin forked
with 18 or 19 principal caudal rays;
pelvic fins with I spine and 7 soft
rays. Large and very rough ctenoid
scales; lateral line complete with
25–56 pored scales. Color: usually
reddish or pink; scale on body sides
often lighter or silvery white, thus

deeply notchedlarge eyelarge ctenoid scales

pelvic fin with
I spine and 7 soft ray

forming longitudinal bands; black pigment somewhat present on opercularmembrane or as markings in fins; finswith yellow or white markings.
Remarks: mostly found in shallow
coastal waters. Feeds on large zooplankton, benthic invertebrates, andsmall fishes. Some species of Sargocentron having the preopercular spinewith venomous.
Similar families occurring in theanal fin with IV spinesarea: none. The serrate bony edges
and spines on head, in combination
with the large eyes, the very longbase on spinous portion of dorsal fin,
the presence of IV anal spines and 7
pelvic soft rays, readily distinguishes
the squirrelfishes and soldierfishes
from other fish families occurring in
the area.

Sargocentron rubrum(Forsskål, 1775)
Redcoat
D XI, 12–14; A IV, 8–10; P1 13–15;
LL 34–38; GR 6–8 + 9–12. Body
moderately deep; dorsal profile ofhead convex; snout short and blunt;corner of preopercle with a sharp
spine; nasal fossa without spinules;
first suborbital bone with 1 or 2 shortlateral spines near upper margin. Last
dorsal fin spine shortest; second anal
fin spine very strong; caudal fin forked
with slightly rounded tips. Scale rows
above lateral line to middle of spinous
portion of dorsal fin 2.5; cheek with 5
oblique rows of scales. Color: bodywith alternate stripes of silvery whiteand brownish red; a triangular streakof brownish red on cheek; often a concentration of pigment forming anelongate brown spot beneath soft portion of dorsal fin and a roundish blotch
above base of soft portion of anal fin;pelvic fin spine white with red membrane; upper and lower edges of cau-

Sargocentron rubrum, UMTF 1362 (KAUM–I. 16575), 15.2 cm SL
Bidong Island, 14 Oct. 2008

dal fin brownish red; rest of caudal fin
and soft portions of dorsal and anal
fins yellowish; no dark spot on pectoral fin axil. Size: maximum length 32
cm SL. Distribution: Indo-West Pacific, from the Red Sea to New Cale-donia and New South Wales of Australia, north to southern Japan. Remarks:
found in protected habitats of shallow
waters.

58

SYNGNATHIDAE
Pipefishes (Seahorses)
By Mizuki Matsunuma
Usually small fishes (4–50 cm
TL). Body typically slender and elongate, without scales, encased in a series of bony rings. Head either alongsame axis as rest of body (subfamily
Syngnathinae), or bent in ventral direction from main body axis (subfamily Hippocampinae). Snout generally long and tubular; mouth small,
toothless, located at tip of snout. Gill
openings reduced to a pore in the
opercular membrane. Branchiostegalrays 1–3. No spines in fins; pelvic fin
absent, other fins variously present
or absent; a single dorsal fin, usuallywith 15–60 soft rays; anal fin small,with 2–6 rays; caudal fin, if present,with 8–10 rays; pectoral fins usually with 10–23 rays. Some species
develop dermal appendages along
body, head, and snout. No lateralline. Color: variable with the species, generally adapted to the preferred habitat; species living on seagrass, sand, and coral rubble usually
have gray, green, brown, or black

tube-like snout

1 dorsal fin with 15–60 soft rays

pelvic fin absent

caudal fin present
or absent

small anal fin

ground color with various patterns;
coral-reef species sometimes colorfulwith white, yellow, orange, blue, red,and black stripes and bands; fins usually transparent; caudal fin sometimes
with colorful patterns.
Remarks: usually live in coastal
marine waters; some are found in estuarine waters, and only a few in fresh
water; marine species found in various
habitats. Feed on minute benthic andplanktonic fauna, preferably microcrustaceans, by sucking into a tubular
snout. Males have a brood pouch inwhich the eggs are laid and where they
are fertilized and incubated. Mostly noor minor importance as food, but somespecies are for the aquarium fishes orused as medicine in Asian markets.
Similar families occurring in
the area: Aulostomidae – larger,
reaching 80 cm TL; body compressed and scaly; distinct separate dorsal fin spines, followed bya normal second dorsal fin; caudal
fin well developed; lateral line present. Fistulariidae – larger, over 100
cm TL; body depressed rather thancompressed; with minute prickles
and linear row of scutes; caudal fin
forked with a distinct elongate filament. Solenostomidae – body short,compressed, with large stellate bonyplates; 2 separate dorsal fins.

Halicampus brocki(Herald, 1953)
Brock's Pipefish
D 21–23; A 4; P1 11–14. Trunk rings
14; tail rings 33–37. Body slender andelongate; snot relatively long (2.4–3.0
in head length); superior trunk and tailridges discontinuous; inferior trunk
and tail ridges discontinuous, lateraltrunk ridge confluent with inferior tailridge; median dorsal snout ridge discontinuous, not essentially concave inlateral profile, with 2–3 short ridges;
lateral snout ridge with 1–2 spines;
with simple or branched dermal flaps
on head. Caudal fin small, typicallywith 10 rays. Color: pale to tan, headblotched or mottled; snout and suborbital often with 3 brown bars; usuallywith 10–11 diffuse pale bars crossing
dorsum and upper part of body; oftenwith diffuse dark bars on lower part ofside and ventral of trunk rings. Size:maximum ca. 12 cm. Distribution:eastern Indian Ocean and west Pacific,
from Western Australia to the Marianaand Marshall Islands, north to south-

Halicampus brocki, UMTF 1456 (KAUM–I. 16630), 5.3 cm SL
Bidong Island, 13 Oct. 2008

Hippichthys (Hippichthys) heptagonus, KAUM–I. 16548, 12.3 cm SL
estuary near UMT, 30 Sept. 2008

ern Japan. Remarks: inhabits rock or present on some or most scutella. Caudal fin small, typically with 10 rays.
coral habitats.
Color: variably tan to dark brown, often
Hippichthys (Hippichthys) heptagonus without distinctive markings; ventral oftrunk usually dusky brown, shading to
Bleeker, 1849
dark brown near median ridge, infreBelly Pipefish
quently with faint transverse bars; dorsal
D 23–30; A 2–3; P1 13–16. Trunk rings fin usually hyaline. Size: maximum
14–15; tail rings 36–42. Body slender length about 15 cm SL. Distribution:and elongate; superior trunk and tail Indo-West Pacific, from east coast of
ridges discontinuous, posterior end of Africa east to Solomon Islands, easternlateral trunk ridge deflected downward; Australia north to southern Japan. Reinferior trunk and tail ridges continuous; marks: found in lower reach of river,dorsal-fin origin on first tail ring; keels estuary, and mangroves.

59

Trachyrhamphus bicoarctatus(Bleeker, 1857)
Double-ended Pipefish
D 24–32; A 4; P1 15–19. Trunk rings
21–24; tail rings 55–63. Body slenderand very elongate; snout relatively
long (1.5–2.0 in head length), smoothdorsally; superior trunk and tail ridges
discontinuous; inferior trunk and tailridges discontinuous, lateral trunk
ridge confluent with inferior tail ridge;
dorsal-fin origin on trunk rings. Caudal fin small, typically with 9 rays.
Color: body variably near white to Trachyrhamphus bicoarctatus, UMTF 1178 (KAUM–I. 16577), ca. 27 cm SL (tail tip broken)
Bidong Island, 30 Sept. 2008
almost black, spotted or mottled;
somewhat with 12–13 pale bars cross- distributed in Indo-West Pacific, from pan. Remarks: found in shallow
ing dorsum and side of tail. Size: max- the Red Sea and East Africa to New coastal of sandy, reef or grass bottoms
imum ca. 40 cm. Distribution: widely Caledonia, Australia to southern Ja- in depths of 1.6–42 m.

FISTULARIIDAE
Cornetfishes

By Mizuki Matsunuma
Large elongate marine fishes,reaching 2 m TL. Body extremely
elongate and slightly depressed.
Mouth small, with a long tubular
snout; teeth in jaws small. Dorsal
and anal fins short based and opposite, with 14–17 soft rays; pectoral fins with 13–17 soft rays; pelvic
fins small and abdominal, with 6 soft
rays. Lateral line arched, running
anteriorly along back, then bending
downward on side and continuingposteriorly onto an elongate filament
produced by the middle 2 caudal-finrays; lateral line composed of tubeshaped ossifications that graduallytake the form of long bony shields
sometimes bearing sharp spines.
Body covered with rows of smallslightly depressed bodytube-like snout
no barbel

forked caudal fin
with a long filament

spinules in some species; a row ofelongate bony plates may be present
along dorsal and ventral midlines ofbody just anterior to dorsal and / oranal fin. Color: red to orange-brownabove and silvery below or brownish
olive above, lighter below, with a series of blue spots on back and snout.
Remarks: no importance in commercial fisheries, but usually caught

with trawls.
Similar families occurring in
the area: Aulostomidae – no caudal
fin filament; barbel present on lower
jaw; distinct separate spines anterior to soft dorsal fin. Syngnathidae– smaller; body covered with armor;
anal fin reduced or absent.

Fistularia petimba
Lacepède, 1803
Red Cornetfish
D 14–16; A 14–15; P1 15–17. Body
extremely elongate, slightly depressed;
snout prolonged and tubular, posttemporal ridge with large well developed
antrorse serrations. A row of elongate
bony plates embedded in skin along
midline of back anterior to dorsal fin;posterior lateral line ossifications ending in a sharp spine. Caudal fin forked
with long whip-like tail filament. Color: head and body red to orange-

Fistularia petimba, KAUM–I. 16899, 39.0 cm SL
off Terengganu (KT), 6 Dec. 2008

brown dorsally, silver laterally andventrally; median fins orangish. Size:
commonly 100 cm, maximum 200 cm
TL. Distribution: widely distributedin Atlantic, Indian, and western Pacific oceans, including Hawaii. Re-

marks: found in coastal waters oversoft bottoms, usually at depths greaterthan 10 m. Marketed fresh, dried salted, or smoked; also reduced to fishmeal.

60

CENTRISCIDAE

Shrimpfishes (Snipefishes)
By Mazlan Abdul Ghaffar
Small fishes (14–30 cm TL);
body elongate, strongly compressed,and blade-like. Head elongate; snout
long, slender, and tubular; mouthsmall, toothless, located at tip ofsnout. Two dorsal fins; first dorsalfin spine long and sharp, located at
hindmost end of body; all other spinous and soft portions of dorsal finson ventral surface of body; caudal fin
small, on ventral surface; pelvic finssmall, with 4 soft rays, originating ator behind midbody. Body enclosed
in a flattened, transparent, bony casing with sharp ventral edge. Lateralline absent. Color: variable with thespecies; either silvery or yellowish
brown to pale green on back, silvery
on sides; dusky to conspicuous lateral streak running length of body andtube-like snout

thin, translucent bony casing

1st dorsal-fin spineat end of bodysoft dorsal fin
sharp ventral edge

through eye.
Remarks: two genera with 4 species. Found in coastal waters; feed
mainly on small animals of the demersal zooplankton, especially crustaceans. No importance in commercial fisheries, but usually caught withtrawls.
Similar families occurring in
the area: Aulostomidae – no caudal
fin filament; barbel present on lower

anal fincaudal fin

jaw; body compressed rather than depressed; distinct separate spines anterior to soft dorsal fin. Syngnathidae– smaller; body covered with armor;
anal fin reduced or absent; no caudal
fin filament. Belonidae – no caudal
fin filament; mouth large, both jaws
produced into slender beak, lower
jaw longer, with single series of distinct canines.

Centriscus scutatus
Linnaeus, 1758
Serrate Razorfish
D III–V-10–12; A 11–13; P1 10–11;
P2 I, 3–5. Body encased in an armor ofthin, translucent plates. First dorsalfin spine fused with body armor plate,
without a movable spinous ray at its
end. Interorbital space with a groovecontinued to crown of head; sutures oflateral plates serrated; postorbital partof head 1 / 2 or more distance of operculum from base of pectoral fins. Color:body silvery, with a dusky lateral
streak; 7 or 8 silvery crossbars on ventral plates. Size: maximum total length
about 14 cm. Distribution: Indo-West
Pacific. Remarks: found in seagrassbeds and coral reefs; feed on minutecrustaceans in the zooplankton. No
commercial importance, but caught
with trawls.

Centriscus scutatus, KAUM–I. 17308, 14.9 cm SL
off Terengganu (KT), 20 Jan. 2009

Centriscus scutatus, KAUM–I. 17272, 8.8 cm SL
off Terengganu (KT), 18 Jan. 2009

61

DACTYLOPTERIDAE
Flying Gurnards

By Hiroyuki Motomura
Moderate sized fishes. Body elongate, not strongly compressed; headlarge, heavily armored; eyes large;
interorbital remarkably wide. Preopercle with a prominent elongate
spine. Gill opening restricted, fused
to isthmus. Spinous and soft-rayed
portions of dorsal fin separated by a
deep notch; first or anterior 2 spines
separated from remainder of fin; dorsal fin with VII or VIII total spinesand 8 or 9 soft rays. Anal fin without spines and with 6 or 7 soft rays.
Caudal fin emarginate. Pectoral finsremarkably large, horizontal, divided
into 2 portions; anterior portion with
5–7 short rays, posterior portion with
25–31 long rays, tip reaching to caudal fin base in adults. Pelvic fins with
I spine and 4 soft rays. Body scales
scute-like, forming prominent keels.

first 2 dorsal-fin spines detached from fin

pectoral fin very
long, wing-likehead armored

Swimbladder divided into 2 halves.
Remarks: found on sandy bottom
Color: variable, from brownish to in tropical and subtropical waters.
reddish, usually with spots or blotchSimilar families occurring in thees. Upper surface of pectoral fin usu- area: Triglidae – lower pectoral finally bluish, with some markings char- rays free.
acterized for each species.

Dactyloptena gilberti
Snyder, 1909
Flateared Helmet Gurnard
D I + I + V + I, 8; A 6–7; P1 28–32;
P2 I, 4. Body moderately elongate.
Second dorsal fin spine separated
from third to seventh spines. Interorbital space extremely wide, its width
18–23% of standard length. Granular
projections on snout arranged in arow; snout wide and rounded. Posttemporal spine well developed, butflat against body. Color: pectoral fin
with numerous dark spots and a large
black blotch on middle of ray; blue
spots and lines inside the middle black
blotch and posterior margin of fin.
Size: maximum total length 22 cm.
Distribution: distributed in the IndoWest Pacific, from the Arabian Peninsula to Japan, but not southern Southeast Asia or Australia. Remarks:
generally found on mud-sand bottom
at depths of 20–71 m.

Dactyloptena gilberti, KAUM–I. 17179, 11.1 cm SL
off Terengganu (KR), 5 Jan. 2009

62

Dactyloptena orientalis(Cuvier, 1829)
Oriental Helmet Gurnard
D I + I + V + I, 8; A 6–7; P1 27–30;
P2 I, 4–5. Body moderately elongate.
Second dorsal fin spine separated
from third to seventh spines. Interorbital space wide, its width 13–15% of
standard length. Snout wide andsomewhat pointed. Preopercular spine
not extending further posteriorly thanposttemporal spine. Color: pectoral
fin with numerous dark spots scatteredon entire fin, except for white ray tips;
margin of middle portion of ray blackwith numerous short blue lines inadults. A large dark ocellus on pectoral
fin in juveniles. Size: maximum total
length 40 cm. Distribution: widelydistributed in the Indo-Pacific, fromthe east coast of Africa and the Red
Sea to the Hawaii and Tuamotu Archipelago. Remarks: generally found on
sand bottom in shallow coastal waters.

Dactyloptena orientalis, KAUM–I. 17140, 15.6 cm SL
off Terengganu (KT), 1 Jan. 2009

63

APISTIDAE

Bearded Waspfishes
By Hiroyuki Motomura
Moderate sized fishes. Body
elongate, moderately compressed.
Dorsal fin with VIII–XV spines and7–10 soft rays; spines strong, venomous. Anal fin with III spines and
6–8 soft rays. Caudal fin rounded.
Pectoral fins long, greater than headlength; lower 1–3 rays detached or
separated from upper part of fin. Pelvic fins with I spine and 5 soft rays.
Lacrimal bone highly mobile. Depthof second suborbital bone subequal
to length of the bone; second suborbital bone broadly connected topreopercle. Preopercular spines welldeveloped. Color: head and bodysilver, yellowish, brownish or grayish, not strongly mottled with spots
or blotches.
Remarks: found on sandy or

lacrimal bonehighly mobile

1–3 free pectoral-fin rays

muddy bottom in river mouth and in- area: families with free pectoral fin
shore waters; body partially buried un- rays and the mobile lacrimal bone
der the soft substrate. Dorsal, anal, and not occurring in the area.
pelvic fin spines venomous.
Similar families occurring in the

Apistus carinatus(Bloch & Schneider, 1801)
Ocellated Waspfish
D XIV–XV, 8–10; A III, 7–8; P1 11–
12; P2 I, 5. Body moderately elongate,
compressed. Lateral surface of headdensely covered with numerous denticulations. Interorbital space 10.0–
16.6 in head length. Ventralmost pectoral-fin ray separate from rest of fin.
Color: body brownish dorsally, whitish ventrally. Spinous portion of dorsal fin with a large black blotch posteriorly; soft-rayed portion grayish,
reticulated, with white margin. Insideand outside of pectoral fin yellowish
and blackish respectively. Pelvic fin
whitish to grayish. Anal fin grayish,
with black band submarginally, whiteband marginally. Color pattern of caudal fin similar to that of soft-rayed
portion of dorsal fin. Size: maximum
standard length 12.5 cm. Distribution: widely distributed in the IndoWest Pacific, from South Africa to Japan and Australia. Remarks: takenover fine sandy bottom at depths ofless than 60 m.

Apistus carinatus, KAUM–I. 17184, 8.9 cm SL
off Terengganu (KR), 5 Jan. 2009

Apistus carinatus, KAUM–I. 17274, 4.3 cm SL
off Terengganu (KT), 18 Jan. 2009

64

SCORPAENIDAE
Scorpionfishes

By Hiroyuki Motomura and Mizuki Matsunuma
Small to relatively large fishes.
Body usually weakly compressed;
head moderate to large; eyes andmouth small to large. Numerous conical or villiform teeth on jaws. Branchiostegal rays 7, rarely 6. Gill rakers
usually short to moderate, with spinous points on each raker. All species
with a suborbital ridge, extending
backward across cheek and usuallyfirmly bound to preopercle; the ridgewith spines in some species. Numerous spines on head in most species;
numerous tentacles on head andbody in some species. Dorsal fin with
VIII–XVIII spines and 3–14 soft rays;
spines strong, venomous in most species. Anal fin with II–IV spines and
3–15 soft rays. Caudal fin rounded or
truncate. Pectoral fins usually large.
Pelvic fins with I spine and 4 or 5
soft rays. Lateral line with 4–54 pored
scales. Color: highly variable, fromblackish to reddish, usually with spots
or blotches; barred or mottled color
patterns in most species.

dorsal fin usually single with
VIII–XVIII, 3–14
caudal fin roundedor truncate

anal fin with II–IV, 3–15

suborbital ridge, extending
backward across cheek

Remarks: found on or near bottom from inshore shallow waters to
depths of about 1200 m; entering into
freshwater in some species; pelagicor semipelagic occurring offshore in
depths of 200–800 m in some species.
Well camouflaged and ambush predators, feeding mainly on arthropods
and fishes. Dorsal, anal, and pelvic
fin spines venomous in most species.
Ovoviviparous or viviparous.

Similar families occurring in
the area: Platycephalidae – headhighly depressed; ventral margin of
lacrimal smooth, without distinct
spines; two dorsal fins well separated. Aploactinidae – fin rays usually
unbranched; head with comparatively blunt spines. Serranidae – no suborbital ridge attaching to preopercle;
large canine teeth in jaws in many
species.

Neomerinthe procurva
Chen, 1981
Curvedspine Scorpionfish
D XII, 10; A III, 5; P1 19; P2 I, 5.
Body oblong, moderately compressed.
Interorbital region scaled. Posterior
lacrimal spine well developed, directed posteroventrally. Lateral lacrimal
spine present. Three suborbital spines.
No median ridge on lateral surface ofmaxilla. Color: body color variable,
blackish to reddish, and variously
blotched dorsally. Size: maximum
standard length 14 cm. Distribution:
distributed in the eastern Indian andwestern Pacific Oceans, from India toJapan. Remarks: usually taken by
trawl; no depth data available.
(H. Motomura)

Neomerinthe procurva, KAUM–I. 17168, 6.5 cm SL
off Terengganu (KR), 5 Jan. 2009

65

Parascorpaena picta(Cuvier, 1829)
Painted Scorpionfish
D XII, 9; A III, 5; P1 16–19; P2 I, 5;
LR 43–49. Body oblong, moderately
compressed. Posterior lacrimal spine
directed anteriorly in adults, ventrally
or anteroventraly in young. First andsecond suborbital ridges fused, forming a single ridge with two spines behind level of orbit, spines absent below eye. Interorbital ridges poorly
developed, not encircling a depression. Occipital pit very weakly developed, nearly flat. Color: body usuallybrownish, mottled with poorly defined
blackish blotches. Size: maximum
standard length 12.3 cm. Disribution:
distributed in Southeast Asia and northern Australia. Remarks: common innearshore waters on reefs and overrocky bottoms.
(H. Motomura)Parascorpaena picta, KAUM–I. 16506, 8.1 cm SL
Cendering, 6 Oct. 2008

Pterois russelii
Bennett, 1831
Plaintail Firefish
D XIII, 10–11; A III, 7–8; P1 12–14;
P2 I, 5; LR 69–86. Body oblong, compressed; eye high set, broadly separated from suborbital ridge. Coronal andtympanic spines absent. Longest dorsal-fin spines subequal to body depth;
membranes of spinous portion incised
nearly to base of fin. All pectoral-fin
rays unbranched; membranes of upperrays strongly incised nearly to base ofray; each membrane reaching ray tip.
Scales cycloid. Color: body red with
about 12 dark reddish bars; ventral
surface of mandible and chest without
markings; soft-rayed portion of dorsal
and anal fins, and caudal fin pale red,
without markings; pectoral fin rays
with narrow black to reddish brownbands; numerous small white spots on
inner surface of pectoral fin and pelvic
fin membrane. Size: maximum length
about 20 cm SL. Distribution: IndoWest Pacific. Remarks: found on
sandy bottoms among corals in shallow waters.
(M. Matsunuma)

Scorpaenopsis neglecta
Heckel, 1837
Yellowfin Scorpionfish
D XII, 9; A III, 5; P1 16–18; P2 I, 5;
LLp 20–22; LR 42–45; GR 4–5 +8–10. Nape and anterior body highly
arched, giving a humpback appear-

Pterois russelii, KAUM–I. 17127, 13.2 cm SL
off Terengganu (KT), 1 Jan. 2009

Scorpaenopsis neglecta, KAUM–I. 16507, 4.0 cm SL
Cendering, 6 Oct. 2008

ance. Interorbital space broad, its
width 3.8–4.4 in head length. Orbit
diameter 1.7–2.0 in snout length. Supraorbital and suborbital ridges serrated. Upper posttemporal spine simple. Upper opercular spine divided
into two or more spines. Color: bodycolor highly variable, strongly mottledwith yellowish to reddish and grayish

blotches. Axil of pectoral fins withblack spots. Size: maximum standard
length 18.3 cm. Distribution: eastern
Indian Ocean and the western Pacific
Ocean, ranging from India east to Japan, Indonesia and Australia. Remarks: occurs in shallow rocky andcoral reefs.
(H. Motomura)

66

TETRAROGIDAE
Waspfishes

By Hiroyuki Motomura
Small to medium-sized fishes.
Body usually strongly compressed;
head moderate to large; eyes andmouth small to large. Numerous
conical or villiform teeth on jaws.
Ventralmost pectoral fin rays not detached or separated. Lacrimal bonehighly mobile. Skinny sensory canal between pterotic and preopercle
absent. Suborbital stay usually notforming wide connection to preopercle. Anterior dorsal proximal pterygiophores sutured to neurocranium
and supracarinalis anterior absent.
Color: highly variable, from blackish to reddish, usually with spots or
blotches; barred or mottled color patterns in most species.
Remarks: found on bottom fromdorsal fin origin
usually above eyebody naked or with cycloid scales

inshore shallow waters to deep waters;
one species lives in freshwater rivers. Dorsal, anal, and pelvic fin spines
venomous in most species. This family was formerly included in the family Scorpaenidae.

Similar families occurring in
the area: Aploactinidae – fin rays
usually unbranched. Body covered
with modified, prickly scales. Synanceiidae – skin at gill opening broadly
connected to isthmus.

Cottapistus cottoides(Linnaeus, 1758)
Yellow Waspfish
D XIV–XV, 5–6; A III, 5–6; P2 I, 4;
LR 64–68; GR 18–21. Body oblong,strongly compressed; body depth 34–
42% of standard length. Posterior
margin of maxilla extending a verticalthrough middle of eye. No scales onanterior part of trunk above opercle.
Interorbital space wide, exceeding orbit diameter. Caudal fin rounded. Color: usually brownish dorsally, whitish
ventrally, with numerous dark spots
on head, body and fins, except for pelvic fin. Size: maximum standard
length 9.2 cm. Distribution: widelydistributed in the western Pacific
Ocean, from China to Australia. Remarks: occurs on sandy bottoms inshallow waters to 24 m depth.

Cottapistus cottoides, KAUM–I. 17161, 5.5 cm SL
off Terengganu (KR), 5 Jan. 2009

67

Richardsonichthys leucogaster(Richardson, 1848)
Whitebellied Rougefish
D XII–XIV, 5–9; A III, 6; P1 14–16.
Body oblong, not strongly compressed. Head profile rounded. Lacrimal with two spines. Uppermost preopercular spine strongly developed.
Dorsal fin origin over posterior onethird of eye; first spine more than half
length of second spine. Body entirely
without scales, except for lateral line.
Caudal fin rounded, without elongate
outer rays. Color: highly variable, but
ventral body, and pectoral, pelvic, analand caudal fins usually reddish. Size:maximum standard length 6.6 cm.
Distribution: widely distributed in
the Indo-West Pacific, from Zanzibar
and Seychelles east to New Caledonia.
Remarks: occurs on sandy bottoms oramong corals in shallow waters to 90
m depth.

Richardsonichthys leucogaster, KAUM–I. 16949, 3.8 cm SL
off Terengganu (KT), 11 Dec. 2008

Vespicula trachinoides(Cuvier, 1829)
Goblinfish
D XIII–XVI, 3–5; A III, 3–4. Body
oblong, compressed. Head profile
pointed. Lacrimal with two spines.
Anterior three spines forming a nearly
separate fin; dorsal fin origin well behind eye. Eye small, orbit diameter
6–9% of standard length. Caudal fin
rounded. Color: body mottled withblack and greenish yellow blotches.
Caudal fin translucent with an indistinct vertical dark band. Soft-rayed
portions of dorsal and anal fins with abroad black band. Size: maximum
standard length 5.8 cm. Distribution:
distributed in the eastern Indian andwestern Pacific Oceans, from Myanmar to China and Indonesia. Remarks: found on sandy and broken
shell bottoms.

Vespicula trachinoides, KAUM–I. 16812, 2.0 cm SL
Merang, 2 Nov. 2008

Vespicula trachinoides, KAUM–I. 16767, 2.5 cm SL
Setiu, 31 Oct. 2008

68

APLOACTINIDAE
Velvetfishes

By Hisashi Imamura
Small marine fishes usually under 5 cm. Body usually compressed.
Head usually with knoblike lumps.
Eye small to relatively large. Snout
often prominent. Mouth moderate
to large. Upper and lower jaws withsmall conical teeth. Palatine always
lacking tooth. Vomerine teeth present
or absent. Gill rakers usually small
or moderate; 0–6 on upper limb offirst gill arch and 3–11 on lower.
Anterior isthmus with fleshy extension in most species. All fin rays
unbranched. Dorsal fin origin above
eye or almost so, except in Adventor
and Peristrominous. Dorsal fin with
9–16 spines and 7–16 soft rays; first
3–5 spines usually appear segregated, either elevated or largely without
membrane connection with the restof the spinous dorsal. Anal fin usu-

dorsal-fin origin
usually above eye

prickly body scales

lateral like with 9–16
tube-like scales

all fin rays unbranchedally with 0–5 weak or blunt spines and
5–16 soft rays. Pelvic fin with 1 spine
and usually 2 or 3 soft rays (rarely 1).
Lateral line with 9–16 tube-like scales.
Body scales modified to prickly scales(but some without scales). Color:body usually brown, reddish brown,
or greenish.
Remarks: aploactinids usuallyknown from the shallow sea bottom,

often found between or under vegetation or rocks. Most species wellcamouflaged. With no commercial
importance.
Similar family occurring in
the area: Scorpaenidae – pectoral
fin with 1 spine and 5 soft rays.
Tetrarogidae – anterior isthmus lacking fleshy extension.

Acanthosphex leurynnis(Jordan & Seale, 1905)
Wasp-spine Velvetfish
D III, IX–X, 7–9; A I–II, 6–8; P1
9–10; P2 I, 2; LLp 9 (10 in total, 1 onbase of caudal fin). Body notably compressed. Lacrimal with 2 strong spines.
Preopercle having 4 spines; lowermost
small. Vomerine teeth present. Gillmembrane broadly united to isthmus.
Dorsal fin origin above near middle ofeye. First dorsal spine stout and strong.
Membrane between 3rd and 4th dorsal
spines deeply concaved. Head andbody lacking scales, except for tubelike lateral line scales. Color: body andhead brownish or creamish, having orlacking brown spots; posterior marginof caudal fin clear; pectoral fin withdark band posteriorly. Size: maximum
standard length about 2.1 cm. Distribution: Indo-West Pacific, including
southeast India, South China Sea, Gulfof Thailand and Australia. Remarks:
benthic species, inhabiting inshore,seagrass and soft bottom.

Acanthosphex leurynnis, KAUM–I. 16628, 1.2 cm SL
Bidong Island, 13 Oct. 2008

pressed. Lacrimal with 2 blunt spines.
Preopercle having 5 spines; lowermost small. Vomerine teeth present.
Gill membrane free from isthmus.
Dusky Velvetfish
Dorsal fin origin above posterior marD XIII, 13; A I, 13; P1 13; P2 I, 2; gin of eye. Head and body with prickLLp 14. Body elongate and com- les covered by skin. Color: body pale

Aploactis aspera(Richardson, 1845)

Aploactis aspera, KAUM–I. 17189, 4.2 cm SL
off Terengganu (KR), 5 Jan. 2009

brown to dark brown, with small whiteand dark spots. Size: maximum length
about 10 cm. Distribution: West Pacific, from southern Japan to South
China Sea, New Caledonia and Australia. Remarks: occurring in shallow
sandy bottoms.

69

SYNANCEIIDAE
Stonefishes

By Hiroyuki Motomura
Small to large-sized fishes. Body
shape variable; head moderate tolarge; eyes and mouth small to large.
Skin at gill openings broadly connected to isthmus. Suborbital stay
becoming wider posteriorly. Lower
pectoral fin rays free in some species.
Head and body, except for lateralline, without exposed scales. Color:
highly variable, from blackish to reddish, usually with spots or blotches;barred or mottled color patterns inmost species. Inside of pectoral fin
with various color markings.
Remarks: found on bottom frominshore shallow waters to relatively
deep waters. Dorsal, anal, and pelvic

dorsal, anal, and pelvic fin
spines usually venomoushead and body
apparently naked

pectoral fin baseusually longfin spines highly venomous in most opening not broadly connected to
species.
isthmus. Aploactinidae – body covSimilar families occurring in ered with modified, prickly scales.
the area: Scorpaenidae – skin at gill

Choridactylus multibarbus
Richardson, 1848
Threefinger Scorpionfish
D XII–XIV, 8–9; A II, 8–9; P1 12;
GR 9–11. Body oblong, moderately
compressed. Head relatively small. Interorbital space wide, its width equal
to 1.5 times orbit diameter. No teethon vomer and palatines. Three lower
pectoral fin rays free and detachedfrom remainder of fin; uppermost pectoral fin ray not filamentous in adults.
Color: body yellowish to brownish,with a yellow to reddish marking on
shoulder. Inside of pectoral fin with
about five oblong white areas, surrounded by black or brown pigment
and usually with white specks in axil.
Size: maximum standard length 10.3
cm. Distribution: widely distributedin the Indo-West Pacific, from the Red
Sea east to China and the Philippines.
Remarks: occurs in coastal waters. A
highly venomous species.

Choridactylus multibarbus
KAUM–I. 17196, 8.6 cm SL
off Terengganu (KT), 5 Jan. 2009

70

Inimicus cuvieri(Gray, 1835)
Longsnout Stinger
D XVII–XVIII, 8–9; A II, 11–13; GR
8–11. Body oblong, moderately compressed. Head relatively small. Orbit
only slightly elevated. Snout length
usually equal to or longer than postorbital length. Interspinous membranefrom fourth rewards less than one
fourth spine height. Lower pectoral fin
rays free and detached from remainderof fin; no filamentous upper pectoral
fin rays in adults. Color: body brownor grayish brown dorsally, paler ventrally. Head usually with minute blackspots. Inner surface of pectoral fin
without distinctive pattern, mostly
solid brown, without spots. Size: maximum standard length 19.5 cm. Distribution: distributed in the Andaman
Sea and the western Pacific, from thePhilippines south to northern Australia. Remarks: occurs on sandy or
mud-sand bottom less than 50 m
depth. A highly venomous species.

Inimicus cuvieri
KAUM–I. 17233, 9.6 cm SL
Setiu, 5 Jan. 2009

Inimicus didactylus(Pallas, 1769)
Bearded Ghoulfish
D XV–XVII, 7–9; A II, 10–12; GR
9–11. Body oblong, moderately compressed. Head relatively small. Orbit
only slightly elevated. Snout length
longer than postorbital length. Interspinous membrane from fourth rewards less than one-third spine height.
Lower pectoral fin rays free and detached from remainder of fin; no filamentous upper pectoral fin rays inadults. Color: body color variable. Inner surface of pectoral fin with a broadband medially, remaining parts of fin
blackish with numerous whitish spots
and lines. Size: maximum standard
length 14.3 cm. Distribution: distributed in the western Pacific Ocean,from Japan to Australia and Vanuatu.
Remarks: occurs on sandy or sandymud bottom less than 80 m depth. A
highly venomous species.

Inimicus didactylus
UMTF 1344 (KAUM–I. 16574), 14.7 cm SL
Bidong Island, 12 Oct. 2008

71

Inimicus sinensis(Valenciennes, 1833)
Spotted Stonefish
D XVII–XVIII, 7–9; A II, 11–13; GR
7–10. Body oblong, moderately compressed. Head relatively small. Orbit
only slightly elevated. Snout length
equal to or longer than postorbital
length. Interspinous membrane fromfourth rewards less than one-fourth
spine height. Lower pectoral fin rays
free and detached from remainder offin; no filamentous upper pectoral fin
rays in adults. Color: body brown orgrayish brown dorsally, paler ventrally. Inner surface of pectoral fin darkwith various sized, white or yellow
spots, without bands. Size: maximum
standard length 15 cm. Distribution:
distributed in the eastern Indian andwestern Pacific Oceans, from Sri Lanka to China and Australia. Remarks:
occurs on sandy or mud-sand bottom.
A highly venomous species.

Inimicus sinensis
KAUM–I. 17236, 8.1 cm SL
Setiu, 5 Jan. 2009

Minous pictus
Günther, 1880
Painted Stinger
D X–XII, 11–13; A II, 9–11; GR
13–17. Body oblong, moderately
compressed. Posterior lacrimal spine
much longer than anterior lacrimal
spine. Posterior tip of pectoral finreaching a vertical through middle ofanal fin base. Dorsal fin spines moderately strong; first dorsal fin spine
shorter than second spine. Color:body reddish brown without distinct
bands or spots. Inner surface of pectoral fin reddish brown with yellowish
irregular bands between rays, the band
becoming wider posteriorly, forming abroad marginal band. Caudal fin pale,
without dark markings. Size: maximum standard length 12 cm. Distribution: widely distributed in the tropical western Pacific Ocean. Remarks:
occurs in depths of 27–90 m.

Minous pictus
KAUM–I. 17160, 7.8 cm SL
off Terengganu (KR), 5 Jan. 2009

72

PLATYCEPHALIDAE
Flatheads

by Hisashi Imamura
Small to large marine fishes,
usually ranging from 7 to 50 cm, a
few attaining to 70cm or greater.
Body elongate. Head moderately orstrongly depressed, usually armed bymany spines and ridges. Eye small
to relatively large. Snout prominent.
Mouth large, lower jaw longer thanupper. Teeth present on upper andlower jaws, vomer and palatine. Gillrakers few, relatively short or mere
stubs. Gill membrane free from isthmus. Two dorsal fins well separated;
1st dorsal fin with VI–X spines, firstshort, isolated or scarcely connected
to second; second dorsal fin with
10–15 soft rays. Anal fin with 10–15
soft rays; no spines present. Pelvic
fin with 1 spine and 5 soft rays. Lateral line complete, with about 30 tomore than 100 scales. Upper surface

1st dorsal-fin with VI–X spines
1st dorsal-fin
spine short

2nd dorsal fin with
10–15 soft rays

head depressed

lower jaw longer
anal fin with10–15 soft raysof body covered by ctenoid scales andlower by cycloid. Color: usually dark
above and pale below; the dark colors
with various shades of brown, gray or
black.
Remarks: platycephalids known
from less than 300 m, most found atless than 100 m and inhabiting on themud, sand, rocky shore and coral reef.
Many species excellent for eating.

Similar family occurring in thearea: Plectrogeniidae and Bembridae – lower jaw shorter than or equal
to upper jaw. Parabembridae – anal
fin with 3 spine. Holpichthyidae –head and body extremely depressed
and lateral line with bony plates.
Percophidae – head without spinesand ridges.

Elates ransonnettii(Steindachner, 1876)
Dwarf FlatheadD VI + 12–14 (usually 13); A 12–14(usually 13); P1 19–22; P2 I, 5; LLp
83–107. Body slender and elongate.
Head moderately depressed. Preopercle with 1 long spine. Eye without ocular flaps. Iris lappet absent. Teeth onvomer in 2 separate patches. Interopercular flap absent. Cheek region lacking skinny sensory tubes. Caudal fin
concaved posteriorly; upper lobe with
an elongate ray. Lateral line scaleswith single opening to exterior. Color:body and head pale brown, with a series of small brownish spots along
body sides; 1st and 2nd dorsal, andcaudal fins with dark spots; anal fin
pale. Size: maximum length about 19
cm. Distribution: widespread in West
Pacific, including Philippines, Indonesia, Malaysia and Australia. Remarks:
taken from sandy and muddy bottoms
in 5–53 m depths.

Elates ransonnettii, KAUM–I. 17277, 14.3 cm SL
off Terengganu (KT), 18 Jan. 2009

Elates ransonnettii, KAUM–I. 16902, 13.2 cm SL
off Terengganu (KT), 6 Dec. 2008

73

Grammoplites scaber(Linnaeus, 1758)
Rough Flathead
D I-VIII or I-VII-I + 11–13 (usually12); A 11–12 (usually 12); P1 19–22;
P2 I, 5; LLp 51–55. Body elongate.
Head moderately depressed. Suborbital ridge with 4–5 spines. Eye without ocular flaps. Iris lappet simple andbroad. Teeth on vomer in 2 separate
patches. Interopercular flap absent.
Cheek region lacking skinny sensory
tubes. Lateral line scales with spineand single opening to exterior. Color:body and head brownish above, whitish below, with about 6 dark bands
crossing dorsal surface in some, obscure or absent in others; 1st dorsal
and pelvic fins dusky; 2nd dorsal fin
with small dark spots; anal fin whiteor with a submarginal row of dark
spots; caudal fin dusky, some with arow of dark spots along upper edge;
pectoral fin spotted on upper half,
dusky on lower. Size: maximum
length about 30 cm, commonly to 20
cm. Distribution: Indo-West Pacific,
including southern Arabian Sea, Bay
of Bengal, Malaysia, Indonesia, Gulfof Thailand, Hong Kong, Taiwan andsouthern Philippines. Remarks: taken
by trawls over mud and sand in about
55 m depth.

Grammoplites scaber, KAUM–I. 17074, 17.8 cm SL
off Terengganu (KT), 27 Dec. 2008

Inegocia japonica, KAUM–I. 16966, 16.8 cm SL
off Terengganu (KT), 11 Dec. 2008

Inegocia japonica, KAUM–I. 17221, 14.4 cm SL
off Terengganu (KT), 11 Jan. 2009

Inegocia japonica(Cuvier, 1829)
Japanese Flathead
D I-VIII or IX + 11–13; A 11–13; P1
19–21; P2 I, 5; LLp 51–55. Body elongate. Head moderately depressed. Suborbital ridge with 2 spines. Eye without
ocular flaps. Iris lappet cirrose. Teethon vomer in 2 separate patches. Fingerlike interopercular flap present. Cheek
region lacking skinny sensory tubes.
Lateral line scales with 2 openings toexterior. Color: body and head grayish
or brown above, whitish below, with 6
obscure dark bands crossing dorsal surface; 1st and 2nd dorsal, pectoral, pelvic and caudal fins with dark spots;
anal fin dusky posteriorly in smaller
specimens or entirely in larger specimens. Size: maximum length about 25
cm. Distribution: widespread in IndoWest Pacific, ranging from Sri Lanka tonorthern Australia and to southern Japan. Remarks: occurring in muddy or
sandy bottom to depths of 85 m.

Kumococius rodericensis, KAUM–I. 17222, 17.7 cm SL
off Terengganu (KT), 11 Jan. 2009

Kumococius rodericensis(Cuvier, 1829)
Spiny Flathead
D I-VIII or IX + 11–12 (usually 11);A 11–13 (usually 12); P1 19–22; P2 I,
5; LLp 50–54. Body elongate. Headmoderately depressed. Suborbital
ridge with many spines. Preopercle
with 3 spines; uppermost longest,
reaching nearly to or just past opercular margin. Eye without ocular flaps.
Iris lappet simple or slightly bilobed.
Teeth on vomer in 2 separate patches.
Finger-like interopercular flap pres-ent. Cheek region lacking skinny sensory tubes. Pectoral fin concaved posteriorly. Lateral line scales with 1
opening to exterior. Color: body andhead brownish above, whitish below,
with about 5 dark bands crossing dorsal surface; pectoral fin dark brown,with a clear or pale area in center.
Size: maximum length about 25 cm.
Distribution: widespread in IndoWest Pacific, including Gulf of Oman,
southern Japan and Australia. Remarks: occurring in muddy or sandy
bottom at depths of 18–130 m.

74

Platycephalus cultellatus
Richardson, 1846
D I-VII + 13; A 13; P1 20; P2 I, 5;
LLp 67. Body elongate. Head strongly
depressed. Spines on dorsal surface ofhead obscure. Preopercle with 2
spines. Eye without ocular flaps. Iris
lappet simple elongated lobe. Teeth onvomer in single band. Finger-like interopercular flap present. Cheek region lacking skinny sensory tubes.
Single short spine between first andsecond dorsal fins present or absent.
Lateral line scales with single opening
to exterior. Color: body and head lightbrown above, whitish below; dorsal
fins with small dark spots; anal fin
pale; pectoral and pelvic fins lightbrown with brown spots; caudal fin
with 4 horizontal blackish bands, including that on middle of the fin. Size:attaining to at least about 45 cm SL.
Distribution: South China Sea, including Canton, Vietnam and Malaysia.

Platycephalus cultellatus, KAUM–I. 16999, 25.0 cm SL
off Terengganu (KT), 14 Dec. 2008

Platycephalus indicus, KAUM–I. 16733, 12.3 cm SL
Setiu, 28 Oct. 2008

Platycephalus indicus(Linnaeus, 1758)
Bartail Flathead
D I-VII–VIII + 13; A 13; P1 18–20;
P2 I, 5; LLp 68–82. Body elongate.
Head strongly depressed. Spines on
dorsal surface of head obscure inadults. Preopercle with 2 spines. Eye
without ocular flaps. Iris lappet simple
elongated lobe. Teeth on vomer insingle band. Finger-like interopercular
flap present. Cheek region lacking
skinny sensory tubes. Single shortspine between first and second dorsal
fins present or absent. Lateral line
scales with single opening to exterior.
Color: body and head covered bymany small spots, whitish below; 1stand 2nd dorsal, pectoral and pelvic
fins with dark spots; anal fin pale; caudal fin with 2 horizontal dark bands,
single yellow blotch near middle offin. Size: maximum length about 50
cm. Distribution: widespread in Indo-West Pacific. Population in theeastern Mediterranean is migrant. Remarks: occurring in muddy or sandy
shallow coastal waters. Records fromJapan and Korea are doubtful.

Rogadius tuberculatus, KAUM–I. 16832, 6.9 cm SL
off Terengganu (KT), 5 Dec. 2008

Rogadius tuberculatus(Cuvier, 1829)
D I-VIII or IX + 10–12; A 10–12; P1
19–22; P2 I, 5; LLp 47–54. Body elongate. Head moderately depressed.
Dorsal surface of head with spines andtubercles. Two to 6 preocular spines
present. Suborbital ridge with fine serrations. Lower half of preopercle
roughly serrated, lacking the antrorse
spine. Eye without ocular flaps. Iris
lappet scalloped. Teeth on vomer in 2
separate patches. Interopercular flap
absent. Cheek region with well developed skinny sensory tubes. Lateralline scales with two opening to exterior. Scales on anterior portion of bodywith one or more spines. Color: bodyand head light brown above, whitish
below; body with several indistinct
brown bands dorsally; dorsal fins withmany brownish spots; pectoral fin
with many irregular brownish bands,
posterior portion of pectoral fin blackish except for upper; pelvic fin withirregular brownish spots; caudal fin
dusky. Size: maximum length about 15
cm. Distribution: widespread in IndoWest Pacific. Remarks: occurring in
mud and sand to a depth of ca. 80 m.

75

Suggrundus macracanthus(Bleeker, 1869)
Largespined Flathead
D I-VIII + 11–12 (usually 12); A
12–13 (usually 12); P1 20–23; P2 I, 5;
LLp 50–55. Body elongate. Headmoderately depressed. Suborbital
ridge with many spines. Eye without
ocular flaps. Iris lappet bilobed in
adults, crenate in juveniles. Teeth onvomer in 2 separate patches. Interopercular flap present. Cheek region partially covered by skinny sensory tubes.
Uppermost preopercular spine beyond
the posterior margin of opercle. Lateral line scales with 2 openings to exterior. Color: body and head brownabove, whitish below, with about 7
obscure dark bands crossing dorsal
surface in some; 1st dorsal fin dusky,
with black blotches; 2nd dorsal fin
with small brown spots; anal fin pale,a few dark streaks on posterior rays;
caudal fin with yellowish marking;pectoral fin grayish below, with dark
spots above; pelvic grayish. Size:maximum length about 26 cm, commonly to 18 cm. Distribution: widespread in Indo-West Pacific, ranging
from southern India and Sri Lanka to
Taiwan and to Coral Sea. Remarks:
occurring in muddy or sandy bottom
to depths to 132 m.

Thysanophrys chiltonae
Schultz, 1966
Longsnout Flathead
D I-VII–VIII + 11–12 (usually 11); A
12; P1 19–22; P2 I, 5; LLp 50–54.
Body elongate. Head moderately depressed. Suborbital ridge with manyspines. Eye without ocular flaps. Iris
lappet with short branches. Teeth onvomer in 2 separate patches. Interoperclar margin incised, forming a broadlobe. Cheek region completely covered by skinny sensory tubes. Lateralline scales with 2 openings to exterior.
Color: body light tan mottled with numerous white spots, white below, with
about 5–7 brown bands crossing back;Suggrundus macracanthus, KAUM–I. 17220, 15.1 cm SL
off Terengganu (KT), 11 Jan. 2009

Thysanophrys chiltonae, UMTF 1890, 21.1 cm SL
off Terengganu (KT), 11 Jan. 2009

Thysanophrys chiltonae, KAUM–I. 17219, 15.8 cm SL
off Terengganu (KT), 11 Jan. 2009

1st dorsal fin with a large black submarginal blotch; 2nd dorsal fin withalternating white and brown spots onrays; anal fin pale; caudal fin withrows of small white spots separating
2–3 dark bands; pectoral fin white below, with several small brown blotches in middle and a series of blackishblotches above; pelvic fin with aprominent blackish blotch near base.

Size: maximum length about 22 cm,
commonly to 16 cm. Distribution:
widespread in Indo-West Pacific,
ranging from Red Sea to southern Japan and to Marquesas Islands, including Malaysia, Indonesia, northern
Australia, and Mariana and MarshallIslands. Remarks: inhabits sand areaadjacent to coral reefs to a depth of ca.
38 m.

76

AMBASSIDAE

Perchlets (Glassfishes)
By Koichi Shibukawa
Small to moderate sized (up to 12cm), oblong and compressed fishes.
Lateral line continuous or interrupted
at midway. Head compressed; eyes
large; lower jaw subequal or projecting beyond upper jaw; spines orserrations on various parts of head(e.g., round of eye, snout, and margins of preopercle and interopercle);no opercular spine; villiform teeth
on jaws and roof of buccal cavity;
branchiostegals 6. Dorsal fin single,
deeply notched before last spine,with VII or VIII spines and 7–11 softrays; anal fin with III spines and 8–11
soft rays; pectoral fin with 11–17 softrays; pelvic fin I, 5; caudal fin forked.
Scales cycloid, moderately large andthin, and usually easy to be deciduous. Vertebratae 24–25. Color: subtranslucent or opaque, with silvery

single dorsal fin with VII–VIII, 7–11
no opercular spines

forked caudal fin

cycloid scales

anal fin with III, 7–11

head and belly; fins hyaline, usuallywith black streaks and/or blotches on
dorsal and caudal fins, tinged with yellow in some species.
Remarks: found in freshwaters,
brackish estuaries, mangroves, andshallow coastal water; usually forming aggregations, and feed on small

crustaceans, insects, and fishes. Minor importance in fisheries, often
marketed as dried and salted; some
freshwater species treated as popularaquarium fish.
Similar families occurring in
the area: Apogonidae – two separate
dorsal fins; II anal-fin spines.

Ambassis interrupta
Bleeker, 1853
Longspined Perchlet
D VII + I, 9–10; A III, 9–10; P1 14–
16; LL 11–13 + 9–13; LR 27–28.
Body deep, compressed; lateral line
interrupted at midway, anterior andposterior series with 11-13 and 9-13
scales, respectively. Single supraorbital spine (posteriorly-directed spine
at dorsoposterior margin of orbit);hind margin of preopercle smooth; 2
scale rows on cheek. Color: body
semitransparent, beige dorsally; headand belly silvery; dorsal fins hyaline,
exclusive of 2nd spine and membrane
between 2nd and 3rd spines of firstdorsal fin blackish; caudal fin tinged
with yellow or yellowish orange, narrowly margined dusky posteriorly.
Size: 9 cm SL. Distribution: IndoWest Pacific. Remarks: found inbrackish estuaries and tidal creeks, especially mangroves.

Ambassis interrupta, UMTF 1513 (KAUM–I. 16705), 4.5 cm SL
Setiu, 27 Oct. 2008

77

Ambassis nalua(Hamilton, 1822)
Scalloped Perchlet
D VII + I, 10–11; A III, 9; P1 15–17;
LL 27–29; LR 27–29. Body deep,
compressed; lateral line continuous.
Single supraorbital spine (posteriorlydirected spine at dorsoposterior margin of orbit); hind margin of preopercle smooth; 2 scale rows on cheek.
Color: body semitransparent, headand belly silvery; dorsal fins hyaline,
exclusive of 2nd spine and membrane
between 2nd and 3rd spines of firstdorsal fin blackish. Size: 9.5 cm SL.
Distribution: Indo-West Pacific. Remarks: found in bays, brackish estuaries and tidal creeks, especially mangroves.

Ambassis nalua, UMTF 1511 (KAUM–I. 16701), 4.4 cm SL
Setiu, 27 Oct. 2008

Ambassis urotaenia
Bleeker, 1852
Bleeker's Perchlet
D VII + I, 10; A III, 9–10; P1 16; LR
26–27. Body oblong, compressed; lateral line continuous with 26–27 tubed
scales. A single supraorbital spine(posteriorly-directed spine at dorsoposterior margin of orbit); hind andventral margins of preopercle serrate;
single scale row on cheek. Color:body semitransparent, head and belly
silvery; dorsal fins hyaline, exclusiveof membrane between 2nd and 3rd
spines of first dorsal fin blackish;dusky longitudinal streak on each caudal-fin lobe. Size: 5 cm SL. Distribution: Eastern Indian Ocean and West
Pacific. Remarks: found in brackish
estuaries and tidal creeks, especiallymangroves.

Ambassis urotaenia, UMTF 1126 (KAUM–I. 16503), 6.3 cm SL
Cendering, 6 Oct. 2008

Ambassis vachellii
Richardson, 1846
Vachell's Glass Perchlet
D VII + I, 9; A III, 9; P1 14–16; LL
10–13 + 12–14; LR 27–28. Body oblong, compressed; lateral line interrupted at midway, anterior and posterior series with 10–13 and 12–14
scales, respectively. Some or more(most frequently 3–5) supraorbital
spines (posteriorly-directed spines at
dorsoposterior margin of orbit); hindAmbassis vachellii, KAUM–I. 16445, 4.2 cm SL
and ventral margins of preopercle withestuary near UMT, 28 Sept. 2008
serrate; 2 or more scale rows on cheek.
Color: body semitransparent, head and 3rd spines of first dorsal fin black- Indo-West Pacific. Remarks: foundand belly silvery; dorsal fins hyaline, ish; caudal fin often tinged with yel- in bays, brackish estuaries and tidal
exclusive of membrane between 2nd low. Size: 5.5 cm SL. Distribution: creeks, especially mangroves.

78

LATIDAE
Lates

By Mizuki Matsunuma
Medium to large sized (up to 2 mTL) percoid fishes. Body elongate,
compressed; dorsal profile behind
eyes concave or convex. Eyes medium sized, relatively close to tip ofsnout and dorsal profile. Preopercle
with serrated posterior or ventral margins and a stout flat spine at angle;
opercle with small flat spine; serrated
supracleithrum exposed, near beginning of lateral line. Snout rounded.
Mouth large, almost horizontal. Teeth
small, in villiform bands on upperand lower jaws, vomer, and palatines(may be present on tongue). Dorsal fin deeply incised between spiny
and soft portions; but not completely
separated or, if separated, with one or
two isolated spines present between
them; with VII–IX spines and 10–14
soft rays. Anal fin with III spines and
7–9 soft rays. Pelvic fin with axillarydorsal fins not completely separated

lateral line extending
onto caudal fin

preopercle serrated, astout flat spine at angle

scale, and I spine and 5 soft rays. Pectoral fin with 16 or 17 rays and spiny flap
just above fin base. Caudal fin rounded.
Scales large, ctenoid; covering bases of
caudal, soft dorsal, and anal fins; lateral-line scales 45–50, extending onto
caudal fin in 1 or 3 series. Vertebrae
11 + 14 = 25. Branchiostegal rays 7.
Color: adults greenish or silvery grayto brown, juveniles with stripes; eyeswith characteristic red reflection.
Remarks: occurring in coastal
marine and estuarine to fresh-water
habitats. Important and popular foodfish.
Similar families occurring in thearea: Serranidae – preopercle usually
without large flat spines or serrations;
opercle with 3 flat spines; lateral line
not extending onto caudal fin.

Lates calcarifer(Bloch, 1790)
Barramundi
D VII–IX, 10–11; A III, 7–8; P1 17–
18; LLp 54–57; LGR 16–17; V 25.
Body moderately deep, elongate, andcompressed; dorsal profile of headconcave anteriorly; snout and jaws
pointed; lower edge of preopercle with
3 or 4 (rarely more) large flat triangular spines; anterior and posterior nostrils close together near eye; mouthlarge, posterior tip of maxilla extending beyond eye. Dorsal fin deeply incised before last dorsal-fin spine; 3rdanal fin spine longest; caudal fin
rounded. Scales firmly fixed, ctenoid;
lateral line extending onto caudal fin.
Color: silver with olive-gray or grayblue bakes; eyes brown to golden; juveniles brown to grayish brown with 3
white stripes on head and scattered
white patches on body sides. Size:maximum over 2 m TL. Distribution:
widely distributed in the Indo-West
Pacific, from the Persian Gulf to Papua New Guinea and northern Australia, north to China. Remarks: foundin coastal marine and estuarine waters.
Feeds on fishes and some crustaceans.

Lates calcarifer, UMTF 1898, 23.5 cm SL
off Terengganu (KT), 12 Jan. 2009

Lates calcarifer, UMTF 1518 (KAUM–I. 16749), 11.7 cm SL
Setiu, 30 Oct. 2008

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SERRANIDAE

Groupers (Sea Basses, Fairy Basslets)
By Yusri Yusuf and Hiroyuki Motomura
Body variable in shape, fromdeep-bodied to elongate and little
compressed to notably compressed.
Opercle bearing 3 flat spines; posterior margin of preopercle nearly
always serrate or with 1–4 spines.
Mouth large, terminal; maxilla exposed when mouth closed; lower jaw
usually projecting; bands of small,
slender teeth in jaws; canines usuallypresent at front of jaw and sometimes
at side; small teeth present on vomer
and palatines of most species; nomolar or incisiform teeth. Opercular membrane separate, with 7 branchiostegal rays. Dorsal fin single or
may be notched, with IV–XIII spinesand 9–25 soft rays; anal fin with III(rarely II) spines and 6–24 soft rays;
caudal fin rounded to lunate in shapewith 12–15 branched rays; pelvic
fins with I spine and 5 soft rays, no
scaly axillary pelvic process. Scales
small to moderate, adherent, ctenoid(or secondarily cycloid). A single
complete lateral line, extending on
caudal fin less than ½ length of middle caudal fin rays. Color: variablewith patterns of light or dark stripes,
spots, vertical or diagonal bars, or

single dorsal fin with VII–XIII spines
caudal fin usuallyrounded, truncate,or lunate

3 opercular spines

usually ctenoid
scalesanal fin with III spines

nearly plain.
Remarks: benthic or bottom-orientated fishes, usually found on coral
reefs or rocky substrata; most species
occur on continental or insular shelvesin depths less than 200 m. All are predaceous, the larger species feeding
mainly on fishes, crustaceans, andcephalopods, while the smaller ones
feed on zooplankton. Most species areprotogynous hermaphrodites.
Similar families occurring in
the area: Latidae – lateral line extends to rear margin of caudal fin;
opercle with a single flat spine.
Lutjanidae – no spines on opercle; scaly
axillary process at base of pelvic finsusually well developed. Haemulidae– no teeth on vomer or palatines; nospines on opercle. Sparidae – jaws
with incisiform and/or molariform
teeth; no spines on opercle; edge ofpreopercle smooth. Lobotidae – nospines on opercle; no teeth on vomer or palatines. Kuhliidae – rearedge of opercle froming only 2 flat
points; branchiostegal rays 6; scaly
sheath at bases of dorsal and anal
fins. Acropomatidae – rear edge of
opercle forming 2 flat points, or thelowest point developed as a cluster of
sharp spines; several distinct, spaced
canines along lower jaw.

Cephalopholis boenak(Bloch, 1790)
Chocolate Hind
D IX, 15–17; A III, 8; P1 16–17; P2 I,
5; LLp 46–51; GR 7–9 + 14–17. Body
moderately elongate; body depth lessthan head length, 2.6–3.0 in SL. Head
length 2.3–2.7 in SL. Teeth on palatines. Dorsal-fin membranes distinctly
incised between spines. Caudal fin
rounded. Pectoral fin middle rays longest, its length 1.3–1.6 in head length.
Longest pelvic-fin soft ray length 1.7–
2.2 in head length. No sclaes on maxilla; no axillary scales on body. Color:body dark brown, often dark reddish
brown, usually with 7 or 8 dark bars
on body. Black spot between upperand middle opercular spines. Size:maximum length 26 cm. Distribution: widely distributed in the IndoWest Pacific, from east coast of Africa

Cephalopholis boenak, KAUM–I. 16987, 14.2 cm SL
off Terengganu (KT), 13 Dec. 2008

east to New Caledonia, including to 64 m. Feeds mainly on crustaceans.
southern Japan and northern Australia. Common in the area.
Remarks: usually found on silty dead(H. Motomura & Y. Yusuf)
reefs in protected waters in depths of 4

80

Cephalopholis formosa(Shaw, 1812)
Bluelined Grouper
D IX, 15–17; A III, 7–8; P1 16–18 ;LLp 47–51; GR 7–10 + 15–18. Body
moderate; its depth 2.5–2.9 in SL. Caudal fin rounded; pectoral fins short,
1.6–1.8 in head length. Snout largely
scaled; maxilla partially scaled; scales
on abdomen ctenoid. Color: darkbrown to yellowish brown, with irregular dark blue lines on head, body andfins; snout, lips, and ventral part ofhead and chest with small dark blue
spots. Size: maximum length about 34
cm. Distribution: Indo-West Pacific.
Remarks: found in shallow coastal
waters with dead coral, rocky or silty
reefs; depth range 10–30 m.
(Y. Yusuf)

Cephalopholis formosa, UMTF 1357 (KAUM–I. 16646), 20.7 cm SL
Bidong Island, 13 Oct. 2008

Cephalopholis microprion(Bleeker, 1852)
Freckled Hind
D IX, 14–16; A III, 8; P1 14–16; LLp
45–51; GR 7–9 + 14–17. Body depth
2.5–2.8 in SL; head length 2.3–2.5 in
SL; interorbital area flat or slightly concave; preopercle rounded, finely serrate, lower edge fleshy; subopercle andinteropercle smooth or finely serrate;
upper edge of operculum very convex;
maxilla naked, extending past eye. Pectoral fin usually reaching past verticalat anus, its length 1.4–1.6 in headlength; pelvic fin length 1.9–2.1 inhead length; caudal fin well rounded.
Color: dark brown; proximal part ofeach scale darker than posterior part;somewhat with indistinct dark bars on
body; head including jaws and anterior
part of body with numerous small darkedged blue spots; somewhat blue spots
extending onto fins; soft dorsal, anal,and caudal fins usually with a grayish
blue margin. Size: maximum length
about 24 cm. Distribution: tropicaleastern Indian Ocean and western Pacific. Remarks: found in shallow
coastal waters with silty bottoms; depth
range 2–20 m. Often confused with
Cephalopholis boenak, but have small
blue spots on anterior part of body.
(Y. Yusuf)

Cephalopholis microprion, KAUM–I. 17050, 13.2 cm SL
off Terengganu (KT), 18 Dec. 2008

Cephalopholis sonnerati, KAUM–I. 17073, 15.7 cm SL
off Terengganu (KT), 27 Dec. 2008

5; LLp 66–80; GR 7–9 + 14–16. Body
relatively deep; body depth greater thanor subequal to head length, 2.3–2.7 in
SL. Head length 2.5–2.7 in SL. Dorsalfin membranes distinctly incised between spines. Caudal fin rounded. PecCephalopholis sonnerati
toral fin middle rays longest, its length,(Valenciennes, 1828)
subequal to longest pelvic-fin ray, 1.5–
Tomato Hind
1.7 in head length for specimens of
D IX, 14–16; A III, 9; P1 18–20; P2 I, 15–40 cm length. Color: body gener-

ally pale reddish to yellowish brown,with small brownish red or dark brownspots on head and fainter spots on bodyand fins. Size: maximum length 57 cm.
Distribution: widely distributed in the
Indo-Pacific. Remarks: usually foundon coral reefs in lagoon and outer reef,
depths of 10 to 150 m.
(H. Motomura & Y. Yusuf)

81

Diploprion bifasciatum
Cuvier, 1828
Barred Soapfish
D VIII, 13–16; A II, 12–13; P1 17–18;
P2 I, 5; LLp 71–76: GR 9–10 + 20–22.
Body deep, compressed; body depth
greater than head length, 2.0–2.4 in SL.
Preopercular, subopercular and interopercular margins serrated. Caudal
fin rounded. Posterior tip of depressed
pelvic fin extending beyond anal fin
origin. Scales mainly ctenoid, not deeply embedded. Color: body yellow witha black bar through eye and a broad one
in middle of body continuing onto posterior two thirds of spinous portion of
dorsal fin. Size: maximum length 25
cm. Distribution: Indo-West Pacific.
Remarks: found in coral reefs and adjacent habitats in depths of 5–50 m. Secrete skin toxin under stress.
(H. Motomura & Y. Yusuf)

Diploprion bifasciatum, KAUM–I. 16842, 12.3 cm SL
off Terengganu (KT), 5 Dec. 2008

Epinephelus amblycephalus(Bleeker, 1857)
Banded Grouper
D XI, 15–16; A III, 8; P1 18–19; P2 I,
5; LLp 47–52; GR 8 + 14–16. Body
moderately elongate; body depth lessthan head length, 2.5–3.0 in SL. Head
length 2.1–2.4 in SL. Posteroventral
preopercular margin with 3–6 enlarged
serrae. Dorsal-fin membranes slightly
incised between spines. Caudal fin
rounded. Pectoral fin middle rays longest, its length equal to longest pelvicfin ray, 1.7–2.3 in head length. Lateralbody scales ctenoid, with a few axillary
scales in adults. Color: body pale gray,with 5 broad dark brown bars, first 2
extending into spinous portion of dorsal fin, third and fourth bars extending
into soft-rayed portions of dorsal andanal fins, fifth dark bar at base of caudal fin. Small black spots on dorsal
parts of body bars, mainly along theedges of the bars. Size: maximum
length 50 cm. Distribution: eastern Indian Ocean and central western Pacific.
Remarks: usually found on offshore
coral and rocky reefs in depths of 80–
130 m.
(H. Motomura & Y. Yusuf)

Epinephelus amblycephalus, KAUM–I. 17230, 34.5 cm SL
off Terengganu (KT), 12 Jan. 2009

Epinephelus areolatus, KAUM–I. 16889, 10.7 cm SL
off Terengganu (KT), 6 Dec. 2008

SL. Caudal fin slightly convex in juvenile, truncate or emarginate in adults;
Epinephelus areolatus
pectoral fins relatively long, 1.5–1.8 in(Forsskål, 1775)
head length, longer than pelvic fins.
Areolate Grouper
Scales on body ctenoid, except for thoD XI, 15–17; A III, 7–8; P1 17–19; rax and ventrally on abdomen; maxilla
LLp 49–53; GR 8–10 + 14–16. Body with very small scales. Color: head,moderately elongate; depth 2.8–3.3 in body and fins pale gray, with numerous

close-set roundish to polygonal brownto yellowish brown spots; pectoral fin
pale, with small dark spots; caudal fin
with distinct white straight margin.
Size: 40 cm. Distribution: Indo-West
Pacific. Remarks: usually found inrocky reefs, dead coral, or alcyonarians;
depth range 6–200m.
(Y. Yusuf)

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Epinephelus bleekeri(Vaillant, 1878)
Duskytail Grouper
D XI, 16–18; A III, 8–9; P1 17–19;
LLp 49–53; GR 9–11 + 15–18. Body
elongate; its depth 3.0–3.5 in SL. Preopercle with 2–9 enlarged serrate at
the angle. Caudal fin truncate toslightly convex. Scales on body ctenoid except for nape, thorax, and ventrally on abdomen; maxilla with verysmall scales. Color: brownish to purplish gray, with numerous small orange-yellow spots on head, body, dorsal fin, and upper third of caudal fin;lower two-thirds of caudal fin dark
purplish gray, with white margin.
Size: maximum length 76 cm. Distribution: Indo-West Pacific. Remarks:
occurs on silty coastal reef areas, but
is not known from well developed
coral reefs; depth to 50m. Marketed
fresh.
(Y. Yusuf)

Epinephelus bleekeri, KAUM–I. 16985, 14.4 cm SL
off Terengganu (KT), 13 Dec. 2008

Epinephelus coeruleopunctatus(Bloch, 1790)
White-spotted Grouper
D XI, 15–17; A III, 8; P1 17–19; LLp
Epinephelus coeruleopunctatus, KAUM–I. 16511, 7.4 cm SL
51–61; GR 7–10 + 14–17. Body modCendering, 6 Oct. 2008
erately elongate; its depth 3.0–3.4 in
SL; head pointed, dorsal profile nearlystraight; preopercle rounded, finely
serrate; opercular spines inconspicuous. Caudal fin rounded; pectoral finslarge and fleshy, its length 1.5–2.1 inhead length. Scales on body of adults
in a broad ctenoid zone along side ofbody, cycloid elsewhere; a patch ofsmall scales on maxilla. Color: adults
brownish gray; body, posterior head,and dorsal fin scattered with large
whitish spots, and numerous palespots; dorsal, caudal and pectoral finswith a very narrow white margins; aEpinephelus coioides, UMTF 1218 (KAUM–I. 16512), 11.8 cm SL
series of indistinct dark blotches at
Cendering, 6 Oct. 2008base of dorsal fin, and one on caudal
peduncle; prominent black streak on
body gray-brownish dorsally, shading
maxillary groove. Juveniles dark grayEpinephelus coioides
to whitish ventrally, with numerous
to black, covered with prominent pu(Hamilton, 1822)
small brownish orange or reddish
pil- size white spots and smaller white
Orange-spotted Grouper
brown spots on head, body, and medidots. Size: maximum length 60 cm.
D XI, 13–16; A III, 8; P1 18–20; LLp an fins; body with 5 faint, irregular,
Distribution: Indo-West Pacific. Remarks: found in rocky or coral areas; 58–65; GR 8–10 + 14–17. Body elon- oblique, dark bars which bifurcate
depth range 2–65 m.
(Y. Yusuf) gate; its depth 2.9–3.7 in SL; preoper- ventrally. Size: maximum length 95
cle with enlarged serrae at angle. Cau- cm. Distribution: Indo-West Pacific.
dal fin rounded. Scales on body Remarks: found in coral or rocky
ctenoid; anterior lateral line scales of reefs. Important commercial fish andadults branched. Color: head and reared in cage culture in Malaysia.
(Y. Yusuf)

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Epinephelus erythrurus(Valenciennes, 1828)
Cloudy Grouper
D XI, 15–17; A III, 8; P1 17–19; P2 I,
5; LLp 53–62; GR 8–9 + 14–17. Body
moderately elongate; body depth lessthan head length, 2.8–3.2 in SL. Head
length 2.4–2.7 in SL. Preopercularmargin finely serrated. Dorsal-fin
membranes not incised or moderately
incised between spines. Caudal fin
rounded. Pectoral fin middle rays longest. Lateral body scales distinctly
ctenoid, with axillary scales. Color:body olive to reddish brown, usuallywith irregular pale spots and blotches.
Three dark streaks across opercle. Median and pelvic fins mottled. Pectoral
fin uniform. Size: maximum length
about 43 cm. Distribution: Indo-West
Pacific. Remarks: inhabits in coral
reef area, harbors and estuaries; depth
range 1–20 m.
(H. Motomura & Y. Yusuf)

Epinephelus erythrurus, KAUM–I. 16333, 8.0 cm SL
Cendering, 16 Sept. 2008

Epinephelus fasciatus(Forsskål, 1775)
Blacktip Grouper
D XI, 15–17; A III, 8; P1 18–20; P2 I,
5; LLp 49–75; GR 6–8 + 15–17. Body
moderately elongate; body depth lessthan head length, 2.8–3.3 in SL. Head
length 2.3–2.6 in SL. Posteroventral
preopercular margin serrated. Dorsalfin membranes distinctly incised between spines. Caudal fin slightly tomoderately rounded. Pectoral fin middle rays longest, its length longer thanlongest pelvicfin ray length, 1.5–2.0 inhead length. Lateral body scales distinctly ctenoid, with numerous axillary
scales; nape and dorsoposterior part ofhead densely covered with minute axillary scales. Color: body greenish gray,pale reddish yellow to scarlet, oftenwith 5 or 6 faint dark bars. Fins reddish
orange, pale yellowish green or greenish brown; outer triangular part of interspinous membranes of dorsal fin
black. Size: maximum length about 40
cm. Distribution: Indo-West Pacific.
Remarks: inhabits in shallow coraland rocky reefs; depths range 3–160 m.
(H. Motomura & Y. Yusuf)

Epinephelus fasciatus, KAUM–I. 17151, 19.1 cm SL
off Terengganu (KR), 4 Jan. 2009

Epinephelus fuscoguttatus, KAUM–I. 17053, 18.5 cm SL
off Terengganu (KT), 18 Dec. 2008

5; LLp 52–58; GR 10–12 + 18–21.
Body depth 2.6–2.9 in SL; head length
2.3–2.5 in SL; interorbital margin flator slightly concave; preopercle rounded, finely serrated; upper edge of operculum distinctly convex; anterior edgeof preorbital deeply indented below
nostrils; maxilla extends well posteriEpinephelus fuscoguttatusor to eye; midlateral part of lower jaw(Forsskål, 1775)
with 3 or 4 rows of teeth, inner teeth
Brown-marbled Grouper
about twice longer than outer teeth;
D XI, 14–15; A III, 8; P1 18–20; P2 I, canines inconspicuous. Color: paleyellowish brown, with irregular 5 vertical series of dark brown blotches;
head, body, and fins with numerous
small brown spots; small black saddle
spot on rear half of caudal peduncle; 2or 3 faint, dark bars at side of jaws.
Size: maximum length 95 cm. Distribution: Indo-Pacific. Remarks: foundin shallow coral or rocky reefs to depthsof 60 m. Important commercial fish andreared in cage culture in Malaysia.
(Y. Yusuf)

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Epinephelus heniochus
Fowler, 1904
Bridled Grouper
D IX, 14–15; A III, 8; P1 16–18; P2 I,
5; LLp 54–60; GR 7–9 + 14–16. Body
depth 2.7–3.2 in SL; interorbital areaslightly convex; dorsal head profile distinctly convex; preopercle angular,
with 2–4 large spines; upper edge of
operculum approximately straight;
maxilla usually reaching to or slightly
past a vertical at rear edge of eye; lower
edge of maxilla with a step-like bend in
adults; canines at front of jaws well developed; midlateral part of lower jaw
with 2 rows of teeth. Color: head andbody pale brown dorsally, shading towhitish or pale pink ventrally; faintdark brown stripe from eve to end of
operculum, another darker stripe fromlower edge of eye to subopercle and athird from edge of preorbital to interopercle; dorsal, pectoral, and caudal
fins hyaline grayish yellow. Size: maximum length 43 cm. Distribution:
western Pacific. Remarks: often foundon soft bottom, rather than rocky areas;
depth range 40–235 m.
(Y. Yusuf)

Epinephelus quoyanus(Valenciennes, 1830)

Epinephelus heniochus, KAUM–I. 17144, 17.0 cm SL
off Terengganu (KT), 1 Jan. 2009

Epinephelus quoyanus, KAUM–I. 17153, 15.0 cm SL
off Terengganu (KR), 4 Jan. 2009

Longfin Grouper
D XI, 16–18; A III, 8; P1 16–19; LLp
48–52; GR 6–8 + 14–16. Body depth
2.8–3.2 in SL; serrae at corner of preopercle enlarged. Caudal fin rounded;pelvic fins not reaching or just reaching anus. Scales on body ctenoid;
maxilla naked or with a few very small
scales. Color: body whitish, with numerous large dark brown to blackspots on head, body and fins; chestwith 2 dark brown bands joining below pectoral fin bases; ventral edge ofanal and caudal fins and leading edgeof pelvic fins with white line andblackish submarginal band. Size:maximum length about 40 cm. Distribution: the Andaman Sea and western
Pacific. Remarks: usually found on
inshore silty reefs; less than 50 m. The
enlarged fleshy pectoral fins appear to
be related to its habit of sitting on thesubstrate, with its pectoral fins in contact with the bottom.
(Y. Yusuf)

Epinephelus sexfasciatus(Valenciennes, 1828)
Sixbar Grouper

Epinephelus sexfasciatus, KAUM–I. 16888, 9.6 cm SL
off Terengganu (KT), 6 Dec. 2008

D XI, 14–16; A III, 8–9; P1 17–19;
LLp 46–51; GR 7–8 + 13–15. Body
depth 2.7–3.2 in SL; anterodorsal headprofile convex; preopercle with 2–4
greatly enlarged serrae at angle. Caudal
fin rounded; pectoral fins not fleshy;
pelvic fins not reaching to anus. Scales
on body ctenoid except anterodorsally
above lateral line, on thorax, and abdomen; maxilla naked. Color: head andbody pale brownish gray, with 5 darkbrown bars; scattered with pale spots

may be present on body, and some faintsmall brown spots are often visible on
the edges of the dark bars; soft dorsal,
caudal, and anal fins with dark brownspots, pectoral fins grayish or duskyorange-red; jaws and ventral parts of
the head sometimes pale reddish
brown. Size: maximum length about
28 cm. Distribution: the Andaman Sea
and western Pacific. Remarks: usuallyfound on silty sand or muddy bottoms
at depths of 10–80 m.
(Y. Yusuf)

85

PSEUDOCHROMIDAE
Dottybacks

By Mizuki Matsunuma and Hiroyuki Motomura
Small to moderate fishes, up to
45 cm. Body moderately elongate
and compressed. Mouth moderate,
weakly protrusible; maxilla not extending beyond posterior margin oforbit. Dorsal fin with IV or fewer
spines and 21–74 soft rays; base of
fin long; pelvic fin (absent in somespecies) with one spine and 3–5 softrays, inserted below or in front ofpectoral fin. Branchiostegal rays 6,
with a single ray articulating withposterior ceratohyal. Lateral line
modified, disjunct, posteriorly truncated or multiple. Color: highlyvariable with stripes, spots, blotchesor nearly plain in each species; colorful in most species; sexually dichro-dorsal fin withIV or fewer spines

maxilla not extending
beyond posterior margin of orbit

lateral line modefied, disjunct,posteriorly truncated or multiple

matic in some species.
Remarks: occurs in shallow intertidal areas to depths of about 100 m;
preferring coral and rocky reefs. Feeds
on fishes, crustaceans, molluscs, andworms. Sequental hermaphrodite insome species. Of commercial importance in aquarium trade in colorful

species.
Similar families occurring in
the area: Opistognathidae and Plesiopidae – dorsal fin with IX or morespines. Serranidae – opercle usuallywith 3 spines; dorsal fin with VII or
more spines.

Pseudochromis fuscus
Müller & Troschel, 1849
Dusky Dottyback
D III, 25–28; A III, 13–15; P1 17–20;
P2 I, 5; LLp 23–36 + 4–13. Body elongate, relatively deep; caudal peduncle
short; posterior margin of maxilla below pupil; lower lip interrupted atsymphysis. Caudal fin rounded totruncate or emarginate. Lateral line
disjunct; circumpeduncular scales
usually 20; dorsal and anal fins withweakly to strongly developed scaly
sheaths. Color: bright yellow to darkgray, usually with blue spots on nape
and at least anterior part of body; upper part of body and dorsal fin sometimes abruptly yellow; caudal peduncle and fin sometimes abruptly pale.
Size: 4 cm SL. Distribution: eastern
Indian Ocean and West Pacific. Remarks: found on reef habitats at
depths of 1–30 m.
(M. Matsunuma)

Pseudochromis fuscus, KAUM–I. 16571, 6.2 cm SL
Bidong Island, 14 Oct. 2008

Pseudochromis ransonneti
Steindachner, 1870
Yellowbelly Dottyback
D III, 23–24; A III, 13–14; P1 17–19;
P2 I, 5; LLp 25–32 + 6–14. Body elongate, moderately compressed; caudal
peduncle short; posterior margin of
maxilla below pupil; lower lip incomplete. Caudal fin rounded to emarginate. Lateral line disjunct; circumpeduncular scales 19–20; dorsal and anal

Pseudochromis ransonneti, UMTF 1336 (KAUM–I. 16564), 3.4 cm SL
Bidong Island, 14 Oct. 2008

fins without scaly sheaths. Color: bluish gray dorsally, pinkish or yellow
ventrally; a dark gray to dark blue
stripe extending from snout tip to anterior edge of eye; dorsal fin bluishgray with irregular pale blue mark-ings; anal fin bluish gray with 3 or 4
indistinct yellow stripes. Size: 6 cm
SL. Distribution: western Pacific.
Remarks: found in coral and rocky
reefs at depths of 1–20 m.
(M. Matsunuma)

86

PRIACANTHIDAE
Bigeyes

By Mizuki Matsunuma
Medium sized (up to 36 cm) marine fishes. Body deep, laterally compressed. Eye extremely large (about
1/2 head length); upturned mouth.
Weak spine on posterior opercle andprominent to remnant spine at angleof preopercle. Branchiostegal rays 6.
Total gill rakers on first gill arch 17–
32. Dorsal fin single, with X spinesand 11–15 soft rays; spinous andsoft-rayed portions of fin continuous,
relatively short to long, soft-rayed
portion broadly rounded to broadly
pointed. Anal fin with III spines and
10–16 soft rays; soft-rayed portionof fin relatively short to long andbroadly rounded to broadly pointed.
Caudal fin rounded, emarginate, or
lunate, with 16 principal rays. Pectoral fin with 17–21 rays. Pelvic finswith I spine and 5 soft rays, broadly
attached to belly by membrane andpositioned in advance of pectoral
fins. Head and body mostly covered
with extremely adherent, rough,spiny scales. Color: head and body
generally reddish, sometimes withsilvery blotches or, in some species,
occasionally a pattern of red and sil-single dorsal fin with X, 11–15

eye very large

caudal fin not forked
pelvic fin
connected to belly
by membraneanal fin with III, 10–16

ver-white bars; fins reddish to duskyor black, occasionally yellowish insome species; some species with dark
spots or speckling on fin membranes.
Remarks: occurring near coral
reefs or rock formations but occasionally in more open areas at depths of
5–400 m, or deeper. Not important inmost fisheries but some species occasionally common in trawl catches ofsoutheast Asian waters.
Similar families occurring in
the area: Holocentridae – opercularmargin with spines; spinous and soft

portions of dorsal fin separated; anal
fin with IV spines; pelvic fin witha spine and usually 7 soft rays andnot attached to belly by membrane;
caudal fin deeply forked with 18–19
rays. Berycidae – dorsal fin withshort base, only IV–VII spines; anal
fin with IV spines; pelvic fin with
7–13 rays; caudal fin deeply forked.
Pempheridae – dorsal-fin with shortbase, IV or V spines and 8 or 9 softrays; anal fin with very long base, III
spines and 22 or more soft rays.

Priacanthus macracanthus
Cuvier, 1829
Brownspot Bigeye
D X, 12–14; A III, 13–14; P1 18–19;
LLp 72–82. Body relatively deep,
ovate and compressed; anterior profileslightly asymmetrical, protruding
lower jaw tip above midline of body;
One strong spine at angle of preopercle. Caudal fin generally truncate.
Scales modified; with posterior field
elevated as a separate flange with spinules both on surface and posterioredge. Color: pinkish silver, reddish
dorsally; fins pinkish; dorsal, anal andpelvic fins with numerous rusty yellow or yellowish brown spots, being
equal to 1/ 2 of pupil diameter. Size:maximum length 35 cm. Distribution: widespread in the Indo-West Pacific. Remarks: found in around the
sandy and rocky shore or open sandy
bottom. Marketed fresh, dried, salted,
and as fish balls.

Priacanthus macracanthus, KAUM–I. 17011, 13.8 cm SL
off Terengganu (KT), 15 Dec. 2008

87

Priacanthus tayenus
Richardson, 1846
Purplespot
D X, 12–14; A III, 13–14; P1 18–19;
LLp 72–82. Body moderately deep,
laterally compressed. A strong spine atangle of preopercle. Caudal fin truncate but becoming very lunate in somemales. Scales modified; scales ofmidlateral region with elevated posterior
field reduced and lacking spinules inlarger specimens. Color: pinkish silver, reddish dorsally; fins pinkish; pelPriacanthus tayenus, KAUM–I. 16841, 14.8 cm SL
vic fins with numerous mall deep puroff Terengganu (KT), 5 Dec. 2008
ple to inky black spots in membrane.
Size: maximum length 29 cm. Distri- Pacific. Remarks: found in around sandy bottom.
bution: the Andaman Sea and West the sandy and rocky shore or open

APOGONIDAE
Cardinalfishes

By Mizuki Matsunuma and Koichi Shibukawa
Small sized (up to about 20 cm)
compressed and oblong or moderately
elongate fishes. Eyes large; mouthlarge, oblique with variable dentition;a single opercular spine. Scales usually ctenoid, but cycloid in some groups
and absent in Gymnapogon and Paxton; lateral line continuous, incomplete or absent in some groups. Two
separated dorsal fins, the first with
VI–VIII spines and the second with
one spine and 8–14 soft rays (except
for Paxton having a single dorsal fin);anal fin with II spines (except for Paxton with a single spine) and 8–18 softrays; pelvic fin I, 5; caudal fin forked,
emarginate, truncate, or rounded. Col-

2 dorsal finslarge eyeusually ctenoid
scaleslarge mouth
anal fin with II spines

or: variable with patterns of colored
stripes, reticulation, spots, vertical or
diagonal bars, or nearly plain.
Remarks: chiefly inhabit marine
coastal waters, but also found in freshand brackish water areas in somegroups.
Similar families occurring in thearea: Ambassidae – a single dorsal fin
but deeply notched, III anal-fin spines.

Apogon cavitensis(Jordan & Seale, 1907)
Caviti Cardinalfish
D VII + I, 9; A II, 8; LL 24. Body oblong, moderately deep and compressed;
lateral line well developed. Snout moderately pointed. Posterior margin of
preopercle serrated. No enlarged caninoid teeth on jaws. Base of anal fin subequal to base of second dorsal fin in
length; caudal fin emarginate. Color:head and body pinkish, with narrow
yellow stripe on back; yellow to bronze
stripe with silvery white margins on
Apogon cavitensis, UMTF 1222 (KAUM–I. 16633), 4.2 cm SL
mid-laterally; small black spot (muchBidong Island, 13 Oct. 2008
smaller than pupil) on front middle ofcaudal fin, which is smaller than pupil cm SL. Distribution: western Pacific marks: found in silty coastal reefs to(M. Matsunuma)
diameter; fins subtranslucent. Size: 8 from Australia to Philippines. Re- 20 m.

88

Apogon endekataenia
Bleeker, 1852
Candystripe Cardinalfish
D VII + I, 9; A II, 8; P1 14–15; LL
27–28; GR 2 + 10. Body oblong, moderately deep and compressed; lateralline well developed, extending posteriorly to caudal-fin base. Snout moderately pointed. Posterior margin of
preopercle weakly serrated. No enlarged caninoid teeth on jaws. Base ofanal fin subequal to base of second
dorsal fin in length; caudal fin emarginate. Color: body pale white, with
about 7 longitudinal yellowish brown
stripes on body sides; fifth stripe (middle stripe) posteriorly reaching to caudal-fin base; a large black spot (about
size of eye) at caudal-fin base; finssubtranslucent with pinkish rays. Size:
11 cm SL. Distribution: western Pacific from Australia to southern Japan.
Remarks: found in coral and rocky
reefs.
(M. Matsunuma)

Apogon endekataenia, KAUM–I. 16634, 3.1 cm SL
Bidong Island, 13 Oct. 2008

Apogon fasciatus(White, 1790)
Broadbanded Cardinalfish
D VII + I, 9; A II, 8; P1 15–16 (usually 16); LL 24–25; GR 2–4 + 10–14.
Body oblong, moderately deep andcompressed; lateral line well developed, extending posteriorly to caudalfin base. Snout moderately pointed.
Posterior margin of preopercle serrated. No enlarged caninoid teeth on
jaws. Base of anal fin subequal to baseof second dorsal fin in length; caudal
fin emarginate. Color: body pinkish,with two black stripes on body sides,
the upper stripe extending from snout
to the dorsal portion of the caudal-finbase, the lower stripe from snout extending onto caudal-fin margin; lower
stripe (middle stripe) without extensional bar on lower margin; fins semitranslucent to pinkish; second dorsal
and anal fins with an indistinct reddish
brown stripe basally. Size: 5 cm SL.
Distribution: widely distributed in
the Indo-West Pacific, from the east
coast of Africa to eastern Australia
north to southern Japan. Remarks:
found in coastal waters at depths of
2–128 m.
(M. Matsunuma)

Apogon endekataenia, KAUM–I. 16637, 5.9 cm SL
Bidong Island, 13 Oct. 2008

Apogon fasciatus, KAUM–I. 17223, 6.0 cm SL
off Terengganu (KT), 11 Jan. 2009

89

Apogon hyalosoma
Bleeker, 1852
Humpbacked Cardinalfish
D VI + I, 9; A II, 8; P1 14; LL 24–25;
1–3 + 6–7. Body deep, compressed;
lateral line well developed, extending
posteriorly to caudal-fin base. Snout
moderately pointed; dorsal profile ofhead distinctly concaved, forming
hump-back appearance. Posterior
margin of preopercle serrated. No enlarged caninoid teeth on jaws. Base ofanal fin subequal to base of second
dorsal fin in length; caudal fin emarginate. Color: head and body graydorsally, becoming silvery ventrally; adistinct, large (about size of eye) black
spot at caudal-fin base; fins transparent, exclusive of anterodorsal marginof first dorsal fin black. Size: 12 cm
SL. Distribution: Western Central Pacific. Remarks: inhabits estuaries, especially mangrove areas.
(K. Shibukawa)

Apogon hyalosoma, UMTF 1501 (KAUM–I. 16696), 8.2 cm SL
Setiu, 27 Oct. 2008

Apogon lineatus
Temminck & Schlegel, 1842
Indian Perch
D VII + I, 9; A II, 8; P1 14–16 (usually 15); LL 27; GR 2–3 + 11–12.
Body oblong, moderately deep; lateralline well developed, extending posteriorly to caudal-fin base. Snout blunt.
Posterior margin of preopercle serrated. No enlarged caninoid teeth on
jaws. Base of anal fin subequal to baseof second dorsal fin in length; caudal
fin rounded, or slightly emarginate.
Color: body pinkish to yellowish
white, darker dorsally; 7–11 narrowdark brown bars on body sides; a dark
bar on cheek; distal third to half offirst dorsal fin black; second dorsal fin
and caudal fins dusky with darker distal margin. Size: 8 cm SL. Distribution: Western Central Pacific. Remarks: found in sandy mud bottom ofcoastal waters. Belonging to subgenus
Jaydia.
(M.Matsunuma)

Apogon nigrocincta(Smith & Radcliffe, 1912)
D VII + I, 9; A II, 8; P1 14; LL 24;
GR 3 + 14. Body oblong, moderately
deep and compressed; lateral line welldeveloped, extending posteriorly tocaudal-fin base. Snout moderately
pointed. Posterior margin of preopercle serrated. No enlarged caninoid

Apogon lineatus, KAUM–I. 16881, 4.8 cm SL
off Terengganu (KT), 6 Dec. 2008

Apogon nigrocincta, KAUM–I. 17267, 7.4 cm SL
off Terengganu (KT), 18 Jan. 2009

teeth on jaws. Base of anal fin subequal to base of second dorsal fin in
length; caudal fin emarginate. Color:body plain light purplish brown;
brown to black stripe on snout, fromtip of lower jaw extending upper jaw
to anterior margin of eye; a verticaldark bar on caudal fin base, with small
black spot (much smaller than pupil)on middle; anterior distal margin of
first dorsal fin reddish brown to black;second dorsal and anal fins with narrow reddish brown to black stripe basally; caudal fin reddish. Size: 9 cm
SL. Distribution: Western Central
Pacific. Remarks: found in coral androcky reefs.
(M. Matsunuma)

90

Apogon pleuron
Fraser, 2005
D VII + I, 9; A II, 8; P1 15–16 (usually 15); LL 24–25; GR 3–5 + 13–17.
Body oblong, moderately deep andcompressed; lateral line well developed, extending posteriorly to caudalfin base. Snout moderately pointed.
Posterior margin of preopercle serrated. No enlarged caninoid teeth on
jaws. Base of anal fin subequal to baseof second dorsal fin in length; caudal
fin emarginate. Color: body pinkish,with two black stripes on body sides,
the upper stripe extending from snout
to dorsal portion of caudal-fin base,the lower stripe from snout extending
onto caudal-fin margin; lower stripe(middle stripe) with about 5–9 narrowvertical bars on lower edge. Size: 5 cm
SL. Distribution: eastern Indian Ocean
and West Central Pacific. Remarks:
found in coastal waters at depths of 3–91m.
(M. Matsunuma)

Apogon pleuron, KAUM–I. 16928, 6.8 cm SL
off Terengganu (KT), 11 Dec. 2008

Apogon poecilopterus
Cuvier, 1828
Pearly-finned Cardinalfish
D VII + I, 9; A II, 8; P1 14-17; LL
Apogon poecilopterus, KAUM–I. 17121, 8.0 cm SL
27–29. Body oblong, moderately deep
off Terengganu (KT), 31 Dec. 2008
and compressed; lateral line well developed, extending posteriorly to caudal-fin base. Snout moderately pointed. Posterior margin of preopercle
smooth. No enlarged caninoid teeth on
jaws. Base of anal fin subequal to baseof second dorsal fin in length; caudal
fin truncate. Color: head and bodygray, darkened dorsally, shading to silvery gray ventrally; several faint horizontal wavy lines on side of body; gillchamber and first gill arch dusky; distal part of first dorsal fin dusky; second dorsal fin with some series of
Apogon seminigracaudus, UMTF 1413 (KAUM–I. 16560), 1.7 cm SL
small blackish gray spots; distal edge
Bidong Island, 13 Oct. 2008of caudal fin dusky. Size: 11 cm SL.
Distribution: Indo-West Pacific. Reeral line well developed, extending red gill filaments visual through opermarks: found in inshore areas withposteriorly to caudal-fin base; one cle; blackish pigment on caudal pesandy or muddy bottom. Belonging toscale (plus 1/2 sometimes) between duncle increasing posteriorly, becomsubgenus Jaydia.
(K. Shibukawa)lateral line and base of third spine of ing black at caudal-fin base and onto
first dorsal fin. Snout moderately lower caudal-fin lobe; upper caudalpointed; anterior nasal pore without fin lobe and the other fins pink to red.
Apogon seminigracaudus
posterior flap. Posterior margin of pre- Size: 5 cm SL. Distribution: West
Greenfield, 2007
opercle serrated. No enlarged caninoid Pacific, from the Gulf of Thailand east
D VII + I, 9; A II, 8; P1 14; LL 24; teeth on jaws. Base of anal fin sub- to Fiji and Tonga, north to southern
GR 4–5 + 13–15. Body oblong, mod- equal to base of second dorsal fin in Japan. Remarks: found in reefs on sand,erately deep and compressed; caudal length; caudal fin forked. Color: body silt, and algae.
(M. Matsunuma)
peduncle relatively long and low; lat- semitransparent, pinkish to reddish;

91

Apogon truncatus
Bleeker, 1854
Flagfin Cardinalfish
D VII + I, 9; A II, 8; P1 16–17; LL 25;
GR 1 + 10. Body oblong, moderately
deep; lateral line well developed, extending posteriorly to caudal-fin base.
Snout blunt. Posterior margin of preopercle serrated. No enlarged caninoid
teeth on jaws. Base of anal fin subequal to base of second dorsal fin in
length; caudal fin rounded. Color:body silvery white, darkish dorsally;
4–6 indistinct dark bars on body sides;
a dark bar on cheek; distal half of 1st
dorsal fin black; 2nd dorsal fin white,with black stripe marginally, a black
stripe on middle; anal fin white with ablack stripe on basally; caudal fin withblack margin, white submarginally,yellowish basally. Size: 12 cm SL.
Distribution: Indo-West Pacific. Remarks: found in sandy mud bottoms
at depths of 50–80 m. Belonging tosubgenus Jaydia.
(M. Matsunuma)

Apogon truncatus, KAUM–I. 17203, 12.4 cm SL
off Terengganu (KT), 10 Jan. 2009

Apogonichthyoides niger(Döderlein, 1883)
D VII + I, 9; A II, 8; P1 15; LLp 26;
GR 2 + 8. Body oblong, deep andcompressed; lateral line well developed, extending posteriorly to caudal
fin base. Snout blunt. Posterior marginof preopercle serrated. No enlarged
caninoid teeth on jaws. Base of anal
fin subequal to base of second dorsal
fin in length; posterior tip of pelvic fin
extending beyond anal fin origin when
depressed; caudal fin slightly emarginate. Color: head and body dusky
gray, darker dorsally; eye pale yellow.
Size: 10 cm SL. Distribution: West
Central Pacific. Remarks: Fraser andAllen (2010) placed this species in genus Apogonichthyoides, previously
treated as synonym of Apogon.
(M. Matsunuma)

Apogonichthyoides niger, KAUM–I. 17296, 6.3 cm SL
off Terengganu (KT), 20 Jan. 2009

Archamia fucata(Canton, 1849)

Archamia fucata, KAUM–I. 16647, 5.7 cm SL
Bidong Island, 13 Oct. 2008

Orangelined Cardinalfish
D VII, 9; A II, 15–17; P1 13–15; LLp
25. Body oblong, moderately deep andcompressed; lateral line well developed, extending posteriorly to caudal
fin base. Snout moderately pointed.
Posterior margin (at least posteroventral corner) of preopercle serrated. No

enlarged caninoid teeth on jaws. Baseof anal fin longer than base of second
dorsal fin in length; caudal fin emarginate. Scaly sheath developed aroundbase of anal fin. Color: body translucent gray with a large black spot (neareye-sized) at caudal fin base; 20–23
narrow, curved oblique orange lines
on body; fins largely transparent. Size:
7 cm SL. Distribution: Indo-Pacific.
Remarks: inhabits coral and rocky
reefs.
(K. Shibukawa)

92

Cheilodipterus quinquelineatus
Cuvier, 1828
Five-lined Cardinalfish
D VI + I, 9; A II, 8; P1 12–13; LL
26–28. Body relatively elongate, compressed; lateral line well developed,
extending posteriorly to caudal-finbase. Snout moderately pointed. Posterior margin of preopercle serrated(maybe near smooth in large specimen). Some enlarged, fang-like canines on jaws, but canines absent
around lower-jaw symphysis. Base ofanal fin subequal to base of second
dorsal fin in length; caudal fin emarginate. Color: body with 4–5 broad
dusky longitudinal stripes (distinctly
narrower than paler interspaces); asmall black spot (smaller than pupil)
encircled by yellow at caudal-fin base;anterodorsal part of first dorsal fintinged with black. Size: 10 cm SL.
Distribution: Indo-West Pacific. Remarks: inhabits coral reefs.
(K. Shibukawa)

Cheilodipterus quinquelineatus, UMTF 1012 (KAUM–I. 16596), 4.8 cm SL
Bidong Island, 12 Oct. 2008

Ostorhinchus cooki
Macleay, 1881
Cook's Cardinalfish
D VII + I, 9; A II, 8; P1 15; LL 24.
Body oblong, moderately deep andcompressed; lateral line well developed, extending posteriorly to caudalfin base. Snout moderately pointed.
Posterior margin of preopercle serrated. No enlarged caninoid teeth on
jaws. Base of anal fin subequal to baseof second dorsal fin in length; caudal
fin emarginate. Color: head and body
pale with 6 longitudinal dark brown
stripes with various widths; 3rd stripe
short and extends from dorsal marginof eye to, or slightly beyond, a verticalthrough origin of second dorsal fin; adusky rounded spot (larger than pupil)at caudal-fin base; fins subtranslucent
or a little tinged with red. Size: 8 cm
SL. Distribution: Indo-Pacific. Remarks: inhabits rocky or coral reefs.
Randall (2005) placed this species ingenus Ostorhinchus, previously treated as subgenus of Apogon.
(K. Shibukawa)

Ostorhinchus cooki, UMTF 1057 (KAUM–I. 16410), 4.7 cm SL
Dungun, 24 Sept. 2008

Ostorhinchus cyanosoma, KAUM–I. 16632, 4.3 cm SL
Bidong Island, 13 Oct. 2008

erately deep and compressed; lateralline well developed, extending posteriorly to caudal-fin base. Snout moderately pointed. Posterior margin of preopercle serrated. No enlarged caninoid
teeth on jaws. Base of anal fin subequal
Ostorhinchus cyanosoma
to base of second dorsal fin in length;(Bleeker, 1853)
caudal fin emarginate. Color: head
Yellow-striped Cardinalfish
and body pale with 6 longitudinal yelD VII + I, 9; A II, 8; P1 14; LL 28; low stripes with various; 3rd stripe
GR 6–7 + 15–16. Body oblong, mod- short and extends from dorsal marginof eye to, or slightly beyond, a verticalthrough origin of second dorsal fin;
4th (middle stripe) stripe reaching
onto caudal fin margin; a narrow stripe
on whitish interspace on cheek andflank. Size: 8 cm SL. Distribution:
Indo-Pacific. Remarks: inhabits
rocky or coral reefs. Randall (2005)
placed this species in genus Ostorhinchus, previously treated as subgenus
of Apogon.
(M. Matsunuma)

93

Pristicon trimaculatus(Cuvier, 1828)
Cheekbar Cardinalfish
D VI + I, 9; A II, 8; P1 14; LL 25–26;
GR 2 + 10. Body deep, compressed;
lateral line well developed, extending
posteriorly to caudal-fin base. Snout
moderately pointed; dorsal profile ofhead almost straight. Posterior marginof preopercle serrated. No enlarged
caninoid teeth on jaws. Base of anal
fin subequal to base of second dorsal
fin in length; caudal fin emarginate.
Color: body dark reddish brown with
3 dark saddle bands below dorsal fins;
Pristicon trimaculatus, KAUM–I. 17046, 10.5 cm SLa small black spot on opercle; a small
off Terengganu (KT), 18 Dec. 2008
black spot at caudal-fin base. Size: 12
cm SL. Distribution: West Central Pa- Randall (2005) placed this species in subgenus of Apogon.
(M. Matsunuma)
cific. Remarks: found in coastal reefs. genus Pristicon, previously treated as

SILLAGINIDAE

Sillagos (Smelts, Whitings)
By Mizuki Matsunuma
Moderate-sized (up to 70 cm)
elongate perciform fishes. Opercle
with small sharp spine; lower partof preopercle horizontal. Snout longand conical. Mouth small, terminal;end of upper jaw sliding below preorbital bone. Teeth villiform, in broad
bands; small teeth on roof of mouth
restricted to anterior part of vomer,
none on palatines. Two separate dorsal fins, the first with 9–12 slenderspines; the second with I spine and
16–27 soft rays; anal fin long, with
II weak spines and 14–27 soft rays;
caudal fin emarginate. Scales small,ctenoid; lateral line slightly arched.
Swimbladder absent or vestigial tohighly complex with various exten-

2 dorsal fins
1 opercular spine

ctenoid scales
small terminal
mouth

sions. Color: silvery to sandy gray or
green, sometimes with black spots onbody and pectoral fin base.
Remarks:
bottom-dwelling,
schooling fishes; found in inshore
brackish estuaries and shallow coastal
waters.
Similar families occurring in theanal fin with II, 14–27

area: Branchiostegidae – a single
continuous dorsal fin; mouth large
with fleshy lips. Pinguipedidae –dorsal fin spines short; spinous dorsal fin sometimes joined to soft dorsal fin; pelvic fin base in advance of
pectoral fin base.

Sillago aeolus
Jordan & Evermann, 1902
Oriental Sillago
D XI + I, 18–20; A II, 17–19; LL
67–72. Body elongate, subcylindrical,
covered with ctenoid scales. First pelvic-fin ray normal, not thickened clublike structure; caudal fin slightly
forked with rounded tips. Swimbladder with 3 rudimentary anterolateral
Sillago aeolus, KAUM–I. 17060, 15.3 cm SL
extensions instead of 4; no divided
off Terengganu (KT), 26 Dec. 2008
posterior extension. Color: body silvery, with scattered dark brown elon- cm. Distribution: Indo-West Pacific.
gate blotches on body sides; pelvic fin Remarks: found in coastal inshore
yellowish; caudal fin dusky. Size: 30 waters.

94

Sillago asiatica
McKay, 1983
Asian Sillago
D XI + I, 20–21; A II, 21–23; LL
67–70. Body elongate, subcylindrical,
covered with ctenoid scales. First pelvic-fin ray normal, not thickened clublike structure; caudal fin relatively
small, slightly forked with rounded
tips. Swimbladder with 3 anterior extensions, the middle one projecting
forwards and the anterolateral onesrecurved backwards up to 1/2 lengthof swimbladder; a single posterior extension. Color: body silvery, brownish dorsally; somewhat with an indistinct pale midlateral band; upper andlower margins of caudal fin darkbrown to almost black. Size: 16 cm.
Distribution: West Central Pacific.
Remarks: found in inshore waters.

Sillago chondropus
Bleeker, 1849

Sillago asiatica, KAUM–I. 17062, 15.7 cm SL
off Terengganu (KT), 26 Dec. 2008

Sillago chondropus, UMTF 1346 (KAUM–I. 16545), 19.1 cm SL
beach near UMT, 8 Oct. 2008

Clubfoot Sillago
D XI–XII + I, 20–21; A II, 22–23; LL
66–73. Body elongate, subcylindrical,
covered with ctenoid scales. First pelvic-fin ray modified into a laterallycompressed thickened club-like structure; caudal fin truncate. Swimbladder
reduced in size, no duct-like processfrom the ventral surface to the urogenital aperture. Color: body silvery,head yellowish; pectoral and pelvic
fins yellowish; distal margin of anal
fin yellowish; caudal fin dusky. Size:
35 cm. Distribution: Indo-West Pacific. Remarks: found in shallow
coastal waters. Marketed fresh.

Sillago ingenuua, KAUM–I. 17061, 16.7 cm SL
off Terengganu (KT), 26 Dec. 2008

Sillago ingenuua
McKay, 1985
Bay Sillago
D XI + I, 17; A II, 17; LL 66–70.
Body elongate, subcylindrical; cheekcovered with ctenoid scales. First pelvic-fin ray normal, not thickened clublike structure; caudal fin relatively
large, moderately emarginate. Swimbladder with a short median anterior
extension and about 5 small pointed
anterolateral projections. Color: bodysilvery, brownish dorsally; no markings on body sides; dorsal fins brownish distally. Size: 20 cm. Distribution: eastern Indian Ocean and West
Pacific. Remarks: found in inshore
coastal waters. Marketed fresh.

Sillago sihama, KAUM–I. 16678, 19.8 cm SL
Kemaman, 21 Oct. 2008

Sillago sihama(Forsskål, 1775)
Silver Sillago
D XI + I, 20–23; A II, 21–23; LL
66–72. Body elongate, subcylindrical.
First pelvic-fin ray normal; caudal fintruncate to almost rounded. Swimbladder with 2 anterior extensions extending forward and diverging to terminate on each side of the basioccipital;

2 lateral extensions commence anteriorly; 2 posterior tapering extensions of
the swimbladder project into caudal
region. Color: body silvery, brownish
dorsally; no markings on body sides;
pelvic fin yellow; distal margin of anal
fin broadly yellow; caudal fin brownish. Size: 30 cm. Distribution: IndoWest Pacific. Remarks: found in inshore coastal waters.

95

CORYPHAENIDAE

Dolphinfishes (“dolphins”)
By Mizuki Matsunuma
Large sized (up to 2.1 m) marine
fishes. Body elongate, compressed.
Mouth large, with many fine teeth inbands; adult males develop a bony
crest on front of head. Lateral line
curved upward above pectoral fins.
Dorsal and anal fins very long, continuing almost to caudal fin, without
sharp spines, or finlets; dorsal-fin
origin on nape; anal-fin origin at or
before midpoint of body; caudal fin
deeply forked, without any keels on
fin or caudal peduncle; pelvic finsfitting into a groove on body. Scales
small and cycloid. Color: in life veryvariable, sides with golden hues andback brilliant metallic greens andblues; numerous small, black spots
on head and body; specimens lessdorsal fin begins
on napepelvic fin developed

than 15 cm have dark vertical bars.
Remarks: pelagic fishes. Feed on
small fishes and crustaceans associated with floating weed. Taken by bothcommercial and recreational fisheries
wherever they occur.
Similar families occurring in
the area: none. No other fishes havedorsal and anal finsalmost to caudal fin

a combination of characters such as
dorsal fin from nape almost to caudal
fin; anal fin from about midpoint ofbody almost to caudal fin; no sharp
spines in dorsal and anal fins; caudal fin deeply forked; and pelvic finswell developed.

Coryphaena hippurus
Linnaeus, 1758
Common Dolphinfish
D 55–67; A 25–30; P1 17–20; LL
200–300. Body elongate and compressed, greatest body depth in adults
less than 25% SL; head profile becoming vertical with development of abony crest with growth, in males.
Tooth patch on tongue small and oval.
Dorsal fin long, extending from aboveeyes almost to caudal fin; anal fin concave, extending from anus almost to
caudal fin; pectoral and pelvic finslarge; caudal fin deeply forked. Color:back brilliant metallic blue-green inlife, after death fading to gray with agreen tinge; sides silvery with a golden sheen, and 1 row of dark spots orgolden blotches running below dorsal

Coryphaena hippurus, KAUM–I. 17213, 45.0 cm SL
off Terengganu (KT), 10 Jan. 2009

fin and 1, 2, or more rows on and below lateral line, some scattered irregularly; dorsal and anal fins black, thelatter with a white edge; pectoral fins
pale; caudal fin silvery with a golden
sheen. Size: commonly 1 m, maxi-

mum about 2 m. Distribution: worldwide in tropical and subtropical seas.
Remarks: pelagic, inhabiting open
waters, but also approaching the coast.
Marketed fresh.

96

RACHYCENTRIDAE
Cobia

By Hiroyuki Motomura
This family is represented by asingle species; see the following species account.
Similar families occurring in
the area: Echeneidae – dorsal fin
spines absent. Sucking disc present.
Carangidae – II detached spines infront of anal fin.

dorsal-fin spines isolated
dorsal-fin soft rays 26–33

head broad anddepressed

small scales embeddedin thick skinanal-fin soft rays 22–28

Rachycentron canadum(Linnaeus, 1766)
Cobia
D VII–IX, 26–33; A II–III, 22–28; P1
21–22; P2 I, 5. Body elongate, subcylindrical, body depth 5.6–8.0 in standard length. Head broad and depressed; sucking disc absent. Dorsal
fin spines short, not connected by amembrane; longest dorsal fin soft rays
shorter than longest anal fin soft rays.
Caudal fin truncate in young, progressively more emarginate with growth.
Pectoral fins pointed, becoming more
falcate with growth. Scales small, embedded in thick skin; lateral line
slightly wavy anteriorly. Color: body

Rachycentron canadum, KAUM–I. 16991, 27.0 cm SL
off Terengganu (KT), 14 Dec. 2008

grayish to blackish dorsally, whitish tograyish ventrally, with a pale graystripe from front of snout through eye
to upper caudal peduncle and a faintdark stripe on lower side of trunk.
Size: maximum total length about 2
m. Distribution: circumglobal intropical to warm temperate seas, except for the eastern Pacific. Remarks:
semipelagic, but also found over shallow coral and rocky reefs, occasionally in estuaries. Occurs more in continental than insular waters.

97

ECHENEIDAE
RemorasBy Hiroyuki Motomura
Medium-sized (up to 90 cm),
elongate and fusiform fishes. Headwide, depressed; a transversely
laminated, oval-shaped cephalicdisc. Opercle without spines. 8–11
branchiostegal; rays. Dorsal fin with
18–45 soft rays. Anal fin with 18–41;
dorsal and anal fin bases long, lacking spines. Caudal fin slightly forked,
emarginate, or slightly rounded in
adults; with elongate median caudal
fin filament in juveniles of some species. Pectoral fin with 18–32 rays.
Pelvic fin with I spine and 5 soft
rays. Scales small, cycloid, usuallyembedded in skin. No swimbladder.
Color: body brown, grayish to black,oval-shaped cephalic discwith transverse laminae
small cycloid scales
dorsal and anal fins without spines

sometimes whitish, with light anddark longitudinal stripes on trunk.
Remarks: attaching to otherfishes, cetaceans, sea turtles, whales,or dolphins with a sucking disc onhead; a great preference or specificity
toward certain hosts in some species.
Feeds on parasitic copepods attached
to a host and food scraps dropped by

the host. No commercial importance,but some species are taken in coastal
fisheries along with other fishes andsold in local markets.
Similar families occurring in
the area: no other family of fishes
has a sucking disc on dorsally on thehead.

Echeneis naucrates
Linnaeus, 1758
Sharksucker
D 34–42; A 31–41; P1 21–24; P2 I, 5;
LGR 11–16. Body elongate, body
depth 8–14 in standard length. Sucking disc large, with 18–28 laminae.
Caudal fin lanceolate in young, middle rays elongate and filamentous; almost truncate in adults, with upper andlower lobes longer than middle rays.
Pectoral fins pointed. Color: bodygray to blackish, with a white-edged
black stripe from tip of lower jaw tocaudal fin base. Upper and lower mar-Echeneis naucrates, KAUM–I. 17004, 23.2 cm SL
off Terengganu (KT), 14 Dec. 2008

gins of fins whitish in juveniles. Pectoral fins blackish. Pelvic fins grayish
to whitish. Size: maximum standard
length about 1 m. Distribution: circumglobal in tropical to warm temperate seas, except for Pacific coast of

America. Remarks: attaching temporarily to a wide variety of hosts such as
sharks, sea turtles and ships. Often
found free-swimming and occurringin inshore waters.

98

CARANGIDAE

Jacks (Scads, Trevallies)
By Seishi Kimura
Medium to large sized marine
fishes; maximum size about 2 m.
Body shape extremely variable,
ranging from elongate and fusiform
to deep and strongly compressed.
Adipose eyelid developed in somespecies; jaw teeth small, villiform,or absent in some species. Two dorsal fins, the first with IV–VIII spines(obsolete or embedded in adult ofsome species) and the second with
one spine and 17–44 soft rays; anal
fin usually with III spines, the anterior two (rarely only one) detachedfrom the rest of the fin (becoming
embedded in adults of some species), and 15–39 soft rays; caudal
fin forked; pectoral fin usually long,extending beyond a vertical throughsecond dorsal-fin origin except forsome species; pelvic fin I, 5. Scales
small cycloid in most species; scutespresent and prominent on the lateralline in most species, but reduced insome species and absent in some
genera. Vertebral counts 24–27. Color: body silvery, darker (brownish or
greenish) dorsally, paler ventrally;

2 dorsal fins
IV–VIIII, 17–44
caudal fin forked

cycloid scales

anal fin with III, 15–39
anterior 2 anal-fin
spines detacheddark bars or spots on body, and/or yellowish fins in some species.
Remarks: occurring in tropicalto temperate marine, estuarine, andsometimes freshwater areas. Almost
all species having commercial importance.
Similar families occurring in thearea: Carangidae is distinguished fromthe following similar families in having
2 detached anal fin spines (sometimes
completely embedded in adults inseveral genera), and enlarged scutes
usually along at least posterior lateral line. Centrolophidae and Lactariidae – no detached anal fin spine; noscutes in the lateral line. Scombridae– dorsal-fin spines IX–XXVII; no detached anal fin spine; no scutes in thelateral line. Stromateid genus Pampus
very similar to Parastromateus niger
in body shape, but the former lacking
a lateral keel of scutes at caudal peduncle.

Alectis ciliaris(Bloch, 1787)
African Pompano
D IV–VII (embedded) + I, 18–20; A
II (embedded) + I, 15–17; GR 4–6 +12–17 = 18–22; V 10 + 14. Body deep,strongly compressed, diamond shapedin juvenile, becoming more elongatedwith growth; profile of forehead tonape rounded; anterior soft rays of
dorsal and anal fins very long, filamentous in juveniles; pelvic fins elongated in juveniles; body superficiallynaked, scales minute and embedded
where present; straight part of lateralline with scutes (8–30) only posteriAlectis ciliaris, KAUM–I. 17210, 16.8 cm SL
orly. Color: body silvery bluish or
off Terengganu (KT), 10 Jan. 2009
grayish dorsally, silvery white ventrally; 5 chevron-shaped blackish
bands on body in juveniles; a black
botch at base of anterior soft portion wide in tropical seas. Remarks: ocof dorsal fin. Size: maximum length curring in coastal waters; feeds mainly
about 150 cm. Distribution: world- on fishes and crustaceans.

99

Alectis indicus(Rüppell, 1830)
Indian Threadfish
D V–VI (embedded) + I, 18–20; A II
(embedded) + I, 15–17; GR 7–11 +21–26 = 29–37; V 10 + 14. Body deep,strongly compressed, diamond shapedin juvenile, becoming more elongatedwith growth; profile of forehead tonape somewhat angular; anterior softrays of dorsal and anal fins very long,
filamentous in juveniles; pelvic finselongated in juveniles; body superficially naked, scales minute and embedded where present; straight part oflateral line with scutes (6–13) only
posteriorly. Color: body silvery greenish or grayish dorsally, silvery whiteventrally; dark bands on body in juveniles; anal fin yellowish. Size: maximum length about 160 cm. Distribution: widely in Indo-West Pacific,
from East Africa to New Guinea, northto southern Japan. Remarks: occurring in coastal waters; feeds mainly on
fishes, squids, and crustaceans.

Alectis indicus, KAUM–I. 17215, 9.1 cm SL
off Terengganu (KT), 11 Jan. 2009

Alepes djedaba(Forsskål, 1775)
Shrimp Scad
D VIII + I, 23–26; A II + I, 18–21;
Alepes djedaba, KAUM–I. 17077, 9.0 cm SL
GR 10–14 + 25–33 = 35–47; V 10 +off Terengganu (KT), 27 Dec. 2008
14. Body oblong, compressed; dorsal
and ventral profiles almost equallyconvex; snout pointed; adipose eyelid
well developed on posterior half ofeye; a single row of small slender
teeth on both jaws; straight part of lateral line almost entirely with scutes(39–51 following 0–3 cycloid scales),longer than curved part (31–36 cycloid scales + 0–3 scutes). Color:body silvery blue dorsally, silvery
white ventrally; a distinct black spot
on upper margin of opercle. Size:maximum length about 40 cm. Distribution: widely in Indo-Pacific, from
Alepes melanoptera, KAUM–I. 16906, 14.3 cm SL
off Terengganu (KT), 6 Dec. 2008
East Africa to Hawaii, north to southern Japan, south to Australia. Remarks: occurring in coastal waters; Body deep, compressed; dorsal and or: body silvery blue or green dorsalfeeds mainly on crustaceans and small ventral profiles almost equally con- ly, silvery white ventrally; a large
fishes.
vex; snout rounded; adipose eyelid black spot on upper margin of opercle;
well developed on posterior half of first dorsal fin black. Size: maximum
eye; a single row of small slender length about 25 cm. Distribution: InAlepes melanoptera
curved teeth on both jaws; straight do-West Pacific, from the Persian Gulf
(Swainson, 1839)
part of lateral line almost entirely with to western Indonesia, north to southBlackfin Scadscutes (49–69 following 0–4 cycloid ern China. Remarks: occurring in inD VIII + I, 23–26; A II + I, 18–21; scales), longer than curved part (31– shore waters; feeds mainly on shrimps
GR 7–9 + 17–24 = 24–30; V 10 + 14. 50 cycloid scales + 0–2 scutes). Col- and copepods.

100

Alepes vari
(Cuvier, 1833)
Herring Scad
D VIII + I, 23–27; A II + I, 20–23;
GR 9–12 + 23–27 = 32–38; V 10 + 14.
Body oblong, compressed; dorsal andventral profiles almost equally convex; snout pointed; adipose eyelid
well developed on posterior half ofeye; a single row of small slendercurved teeth on both jaws; straightpart of lateral line almost entirely withscutes (48–69 following 0–7 cycloid
scales), longer than curved part (42–50 cycloid scales + 0–2 scutes). Color: body silvery blue or green dorsally, silvery white ventrally; a diffuse
dark spot on upper margin of opercle.
Size: maximum length about 50 cm.
Distribution: widely in Indo-West
Pacific, from Red Sea to northern Australia, north to southern Japan. Remarks: occurring in shallow waters;
feeds mainly on shrimps and fishes.

Alepes vari, KAUM–I. 17287, 32.0 cm SL
off Terengganu (KT), 19 Jan. 2009

Atropus atropos(Bloch & Schneider, 1801)
Cleftbelly Trevally
D VIII + I, 19–22; A II + I, 17–18;
GR 8–11 + 19–23 = 29–34; V 10 + 14.
Body deep, ovate, strongly compressed;
nape strongly convex; a deep midventral groove on belly, accommodating pelvic fins and anal-fin spines;
adipose eyelid not developed; upper
jaw with a narrow band of small teeth,
lower jaw with a single series of small
teeth; central rays of second dorsal
and anal fins produced into filaments
in adult male; pelvic fins long; straightpart of lateral line entirely with scutes(31–37), longer than curved part. Color: body bluish green dorsally, silvery
ventrally; pelvic fins black. Size:maximum length about 50 cm. Distribution: Indo-West Pacific, from Arabian Gulf to Philippines, north tosouthern Japan. Remarks: occurringin shallow waters; feeds mainly on
shrimps and copepods.

Atule mate
(Cuvier, 1833)
Yellowtail Scad
D VIII + I, 22–25; A II + I, 18–21;
GR 10–13 + 24–31 = 36–44; V 10 +14. Body elongate oval, moderately
compressed; dorsal and ventral pro-Atropus atropos, KAUM–I. 17202, 10.5 cm SL
off Terengganu (KT), 10 Jan. 2009

Atule mate, KAUM–I. 17003, 11.1 cm SL
off Terengganu (KT), 14 Dec. 2008

files almost equally convex; snout
pointed; adipose eyelid well developed and completely covering eye except for a vertical slit centered on pupil; shoulder girdle (cleithrum) marginsmooth; last rays of dorsal and anal
fins about twice as long as penultimate
rays and more separated; straight partof lateral line almost entirely withscutes (36–49), shorter than curved
part. Color: body silvery olive-green
dorsally, silvery white ventrally; ablack spot posteriorly on opercle; caudal fin yellow. Size: maximum length
about 30 cm. Distribution: widely in
Indo-Pacific, from East Africa to Hawaii. Remarks: occurring in schools
at depths to about 50 m.

101Carangoides coeruleopinnatus(Rüppell, 1830)
Coastal Trevally
D VIII + I, 20–23; A II + I, 16–20;
GR 5–8 + 15–19 = 21–27; V 10 + 14.
Body ovate, strongly compressed;
dorsal profile more strongly convex
than ventral profile; breast widely naked to pectoral-fin base and behindinsertion of pelvic fins, but naked area
not extending above pectoral-fin base;rarely scaled area present between
pectoral-fin base and breast; anterior
second dorsal- and anal-fin rays elongated, filamentous in juveniles, becoming shorter with growth: straightpart of lateral line shorter than curvedpart, with 16–20 week scutes posteriorly. Color: body silvery blue-graydorsally, silvery white ventrally, withnumerous small yellow spots laterally;
dark blotch usually on upper marginof opercle. Size: maximum length
about 40 cm. Distribution: widely in
Indo-Pacific, from East Africa to Fiji,north to southern Japan. Remarks:
occurring in deep coastal waters.

Carangoides coeruleopinnatus, KAUM–I. 17251, 11.8 cm SL
off Terengganu (KT), 17 Jan. 2009

Carangoides gymnostethus(Cuvier, 1833)
Bludger
D VIII + I, 28–32; A II + I, 24–27;
GR 7–9 + 19–22 = 27–31; V 10 + 14–
15. Body ovate, deep, compressed inyoung, becoming elongate and subcylindrical with growth; dorsal profile ofhead and nape gently convex; breast
naked ventrally beyond insertion ofpelvic fins; lateral naked area extending to pectoral fin base; lobes of second dorsal and anal fins falcate; straightpart of lateral line shorter than curvedpart, with 21–37 week scutes posteriorly. Color: body silvery greenish tan
dorsally, silvery white ventrally; a few
yellow spots sometimes laterally on
body. Size: maximum length about 80
cm. Distribution: widely in IndoWest Pacific, from East Africa to New
Caledonia, north to southern Japan.
Remarks: occurring in offshore reefs;
feeds mainly on crustaceans and small
fishes.

Carangoides gymnostethus, KAUM–I. 17105, 16.6 cm SL
off Terengganu (KT), 28 Dec. 2008

Carangoides gymnostethus, KAUM–I. 16907, 16.4 cm SL
off Terengganu (KT), 6 Dec. 2008

102

Carangoides hedlandensis(Whitley, 1934)
Bumpnose Trevally
D VIII + I, 19–22; A II + I, 16–18;
GR 6–11 + 14–17 = 20–27; V 10 + 14.
Body deep, strongly compressed; dorsal contour of forehead convex, with“bump” on interorbital space; breast
widely naked to pectoral fin base andbehind insertion of pelvic fins, but naked area not extending above pectoral
fin base; second dorsal- and anal-fin
soft rays produced into filaments inadult males; straight part of lateral line
shorter than curved part, with 17–29
week scutes posteriorly. Color: bodysilvery blue-gray or green-gray dorsally, silvery white ventrally; a black
spot on upper margin of opercle; caudal fin yellowish. Size: maximum
length about 30 cm. Distribution:
widely in Indo-Pacific, from South
Africa to Samoa, north to southern Japan. Remarks: occurring in coastal
waters.

Carangoides malabaricus(Bloch & Schneider, 1801)
Malabar Trevally
D VIII + I, 20–23; A II + I, 17–19;
GR 7–12 + 21–27 = 32–38; V 10 + 14.
Body ovate, strongly compressed;
dorsal profile more strongly convex
than ventral profile; naked area ofbreast very wide, extending dorsallybeyond pectoral fin base and posteriorly beyond insertion of pelvic fins,usually extending to anal-fin origin;straight part of lateral line shorter thancurved part, with 19–36 week scutes
posteriorly. Color: body silvery bluegray dorsally, silvery white ventrally,
dark or black blotch on upper marginof opercle. Size: maximum length
about 60 cm. Distribution: widely in
Indo-West Pacific, from South Africa
to northern Australia, north to southern Japan. Remarks: occurring incoastal waters.

Carangoides hedlandensis, KAUM–I. 16910, 24.0 cm SL
off Terengganu (KT), 6 Dec. 2008

Carangoides malabaricus, KAUM–I. 17250, 11.9 cm SL
off Terengganu (KT), 17 Jan. 2009

103

Caranx sexfasciatus
Quoy & Gaimard, 1825
Bigeye Trevally
D VIII + I, 19–22; A II + I, 14–17;
GR 6–8 + 15–19 = 21–25; V 10 + 15.
Body oblong, compressed; dorsal profile moderately convex anteriorly,
ventral profile slightly convex; adipose eyelid developed on posterior
half of eye; posterior end of upper jaw
extending to or beyond posterior margin of eye; breast completely scaled;straight part of lateral line almost entirely with scutes (27–36). Color:body silvery yellowish-green dorsally,
silvery white ventrally; small black
opercular spot; anal and caudal fins
yellowish; scutes dark or black. Size:maximum length about 120 cm. Distribution: widely in Indo-Pacific andtropical eastern Pacific Ocean. Remarks: found in coastal waters; juveniles occurring in brackish waters;
feeds mainly on fishes and crustaceans.

Caranx sexfasciatus, KAUM–I. 17286, 24.5 cm SL
off Terengganu (KT), 19 Jan. 2009

Decapterus macrosoma
Bleeker, 1851
Shortfin Scad

Decapterus macrosoma, KAUM–I. 17103, 14.3 cm SL

D VIII + I, 32–38 +1; A II + I, 26–30
off Terengganu (KT), 28 Dec. 2008
+ 1; GR 10–12 + 34–38 = 43–50; V 10
+ 14. Body elongate, slender, cylindrical; adipose eyelid well developed andcompletely covering eye except for a
vertical slit centered on pupil; posterior end of maxilla slightly concave
above, rounded and produced below;
predorsal scaled area not reaching tolevel of center of eye; shoulder girdle(cleithrum) margin with 2 small papillae; a single detached finlet in dorsal
and anal fins; straight part of lateralline with scutes (24–40) posteriorly.
Decapterus russelli, KAUM–I. 17001, 13.5 cm SL
Color: body blue-gray dorsally, siloff Terengganu (KT), 14 Dec. 2008very ventrally; a black spot posteriorly
on opercle. Size: maximum length(cleithrum) margin with 2 small papilabout 35 cm. Distribution: widely in
Decapterus russellilae; a single detached finlet in dorsal
Indo-Pacific, from East Africa to Ha(Rüppell, 1830)
and anal fins; straight part of lateralwaii, and eastern Pacific Ocean, from
Indian Scadline usually entirely with scutes (30–Gulf of California to Peru. Remarks:
pelagic species, feeds mainly on plank- D VIII + I, 27–32 + 1; A II + I, 24–28 40), but rarely a few cycloid scales
tonic crustaceans and larval fishes.
+ 1; GR 10–14 + 30–39 = 41–53; V 10 anteriorly. Color: body bluish gray+ 14. Body elongate, slender, cylindri- dorsally, silvery ventrally; a black spot
cal; adipose eyelid well developed and posteriorly on opercle. Size: maximum
completely covering eye except for a length about 30 cm SL. Distribution:
vertical slit centered on pupil; posteri- widely in Indo-Wesxt Pacific, fromor end of maxilla almost truncate; pre- East Africa to eastern Australia, northdorsal scaled area not reaching to level to southern Japan. Remarks: inhabitsof center of eye; shoulder girdle inshore waters up to 100 m depth.

104

Elagatis bipinnulata(Quoy & Gaimard, 1825)
Rainbow Runner
D V–VI + I, 23–28+2; A (I) + I, 15–
20 + 2; GR 9–11 + 25–26; V 10 + 14.
Body elongated and almost fusiform;
snout pointed; mouth small, upper jaw
not extending to anterior margin ofeye in adult; villiform teeth bands on
jaws; caudal peduncle grooves present; a detached 2-rayed finlet in dorsal
and anal fins; a single isolated anal-fin
spine covered by skin in adults; noscutes. Color: dark olive-green dorsally, yellow laterally, white ventrally;
2 narrow light blue stripes along body;
caudal and pelvic fins yellowish. Size:maximum length 180 cm. Distribution: widespread circumtropical species. Remarks: pelagic species; feeds
on crustaceans and fishes.

Elagatis bipinnulata, KAUM–I. 16973, 16.4 cm SL
off Terengganu (KT), 12 Dec. 2008

Gnathanodon speciosus(Forsskål, 1775)
Golden Trevally
D VII–VIII + I, 18–21; A II + I, 15–
17; GR 7–9 + 19–22 = 27–30; V 10 +14. Body oblong, somewhat deep;
compressed; both jaws without teethin adults, a few feeble teeth on lower
jaw in juveniles; lips fleshy and remarkably thick; breast completely
scaled; straight part of lateral line withscutes (17–26) posteriorly, shorter
than curved part. Color: body silvery
yellow with 7–11 black vertical bands,
usually alternating broad and narrowin juveniles and young adults; the firstband oblique through eye; all fins yellow, caudal fin tips black. Size: maximum length about 120 m. Distribution: widely in Indo-Pacific from East
Africa to Tuamotu Islands, including
Hawaii, and tropical eastern Pacific
Ocean, from Mexico to Peru. Remarks: occurs in deep lagoon andseaward reefs; feeds mainly on crustaceans and small fishes.

Gnathanodon speciosus, KAUM–I. 17052, 16.6 cm SL
off Terengganu (KT), 18 Dec. 2008

Megalaspis cordyla, KAUM–I. 16846, 17.2 cm SL
off Terengganu (KT), 5 Dec. 2008

Megalaspis cordyla(Linnaeus, 1758)
Torpedo Scad
D VIII + I, 9–11 + 7–9; A II + I, 8–10
+ 6–8; GR 8–11 + 18–22 = 26–32; V
10 + 14. Body elongate, somewhat compressed; caudal peduncle depressed;
adipose eyelid well developed andcompletely covering eye except for a
vertical slit centered on pupil; shoulder girdle (cleithrum) margin smooth
without papillae; several detached fin-lets in dorsal and anal fins; straightpart of lateral line entirely with large
scutes (51–59), much longer thancurved part; scutes on caudal peduncle
forming a lateral keel. Color: bodysilvery brownish-green dorsally, silvery white ventrally; a large black spot
posteriorly on opercle. Size: maximum length about 80 cm. Distribution: widely in Indo-West Pacific,
from East Africa to Fiji, north tosouthern Japan. Remarks: pelagic,
schooling fish; feeds mainly on fishes.

105

Parastromateus niger(Bloch, 1795)
Black Pomfret
D IV–V + I, 40–45; A II + I, 35–39;
GR 5–7 + 13–16; V 10 + 14. Body
deep, rhomboidal, strongly compressed;
dorsal and anal profiles equally muchconvex; mouth small, terminal; a single row of small conical teeth on both
jaws; spinous portion of dorsal fin anddetached anal-fin spines embedded,
not seen in adults; pelvic fins absent inyoung and adults (> 10 cm FL), present at jugular anterior to pectoral-finbase in juvenile; straight part of lateralline much shorter than curved part,with 8–19 week scutes. Color: bodydark brown or black uniformly. Size:maximum length about 75 cm. Distribution: widely in Indo-West Pacific,
from East Africa to northern Australia.
Remarks: inhabits coastal waters
with muddy bottom; feeds on zooplankton.

Parastromateus niger, KAUM–I. 16909, 13.8 cm SL
off Terengganu (KT), 6 Dec. 2008

Scomberoides commersonnianus
Lacepède, 1801
Talang Queenfish
D VI–VII + I, 19–21; A II + I, 16–19;
GR 0–3 + 7–12 = 8–15; V 10 + 16.
Body oblong, somewhat deep, strongly compressed; no caudal peduncle
groove; posterior tip of upper jaw extending well beyond a vertical throughposterior margin of eye in adults; tworows of small conical teeth and 1–2
symphyseal canines on lower jaw;
body covered with broadly lanceolate
scales; no scutes along lateral line.
Color: body blue-greenish gray dorsally, silvery white ventrally, with aseries of 6–8 dusky roundish blotches
dorsolaterally in adults; dorsal andcaudal fins uniformly dusky; anal andpelvic fins whitish in young. Size:maximum length about 120 cm. Distribution: widely in Indo-West Pacific, from East Africa to Australia. Remarks: inhabit shallow coastal waters
to offshore areas; feeds on mainly
fishes.

Scomberoides tol(Cuvier, 1832)
Barred Queenfish
D VI–VII + I, 19–21; A II + I, 18–22;
GR 4–7 + 17–20 = 21–26; V 10 + 16.
Body slender, strongly compressed;
posterior tip of upper jaw not reaching

Scomberoides commersonnianus, KAUM–I. 16877, 26.0 cm SL
off Terengganu (KT), 5 Dec. 2008

Scomberoides tol, KAUM–I. 17076, 11.9 cm SL
off Terengganu (KT), 27 Dec. 2008

to a vertical through posterior marginof eye in adults; dentary with 2 teeth
rows separated by a shallow groove in
adults, teeth in both rows almost same
sized; body covered with slender, needle-like scales; no scutes along lateralline. Color: body blue-greenish graydorsally, silvery white ventrally, witha series of 5–8 oblong dark blotches
laterally in adults; lobe of second dorsal fin black. Size: maximum length
about 60 cm. Distribution: widely in
Indo- Pacific, from East Africa to Fiji,north to southern Japan, south tonorthern Australia. Remarks: found incoastal waters; feeds on mainly fishes.

106

Selar boops(Cuvier, 1833)
Oxeye Scad
D VIII + I, 23–25; A II + I, 19–21;
GR 8–12 + 25–29; V 10 + 14. Body
elongated and compressed; eye verylarge; adipose eyelid well developed
and completely covering eye except
for a vertical slit centered on pupil;shoulder girdle (cleithrum) margin
with upper (small) and lower (large)
papillae; dorsal and anal fins without
detached finlets; straight part of lateralline almost entirely covered by relatively large scutes (37–46), longer
than curved part. Color: body bluegreen dorsally, silvery white ventrally,
with a narrow yellow longitudinal
stripe from border of opercle to caudal
peduncle. Size: maximum length
about 25 cm FL. Distribution: IndoWest Pacific, from Pakistan to Solomon Islands; East Atlantic (Portugal).
Remarks: feeds mainly on planktonic
or benthic crustaceans.

Selar boops, KAUM–I. 16843, 13.1 cm SL
off Terengganu (KT), 5 Dec. 2008

Selar crumenophthalmus(Bloch, 1793)
Bieye Scad

Selar crumenophthalmus, KAUM–I. 16893, 11.7 cm SL

off Terengganu (KT), 6 Dec. 2008
D VIII + I, 24–27; A II + I, 21–23;
GR 9–12 + 27–31; V 10 + 14. Body
elongated and compressed; eye verylarge; adipose eyelid well developed,
completely covering eye except for a
vertical slit centered on pupil; shoulder girdle (cleithrum) margin with upper (small) and lower (large) papillae;
dorsal and anal fins without detachedfinlets; straight part of lateral line almost entirely covered by relatively
small scutes (29–42), almost the samelength as curved part. Color: bodyblue-green dorsally, silvery white ventrally, with a narrow yellow longitudinal stripe from border of opercle toSelaroides leptolepis, KAUM–I. 16948, 11.6 cm SL
caudal peduncle. Size: maximum
off Terengganu (KT), 11 Dec. 2008
length about 70 cm. Distribution:worldwide in tropical and subtropicalwaters. Remarks: found mainly inshallow inshore waters; feeds on pose eyelid well developed on poste- trally, with a broad longitudinal yelplanktonic and benthic invertebrates. rior half of eye; no teeth on upper jaw, low stripe from upper margin of eye toa single row of minute teeth on lower caudal peduncle; upper opercle with ajaw; shoulder girdle (cleithrum) mar- prominent black spot. Size: maximum
Selaroides leptolepis
gin smooth without papillae; dorsal length about 22 cm. Distribution: In(Cuvier, 1833)
and anal fins without finlets; straight do-West Pacific, from Persian Gulf toYellowstripes Scadpart of lateral line with 13–25 cycloid eastern Australia. Remarks: inhabits
D VIII + I, 24–26; A II + I, 20–23; scales followed by 24–29 relatively soft bottom area at depths shallower
GR 10–14 + 27–32 = 40–46; V 10 + small scutes. Color: body metallic than 50 m.
14. Body oblong, compressed; adi- blue-green dorsally, silvery white ven-

107

Seriolina nigrofasciata(Rüppell, 1829)
Blackbanded Trevally
D VII–VIII + I, 30–37; A II + I, 15–
18; GR 4–10; V 11 + 13. Body elongate, slightly compressed; posterior
tip of upper jaw broadly rounded,reaching to a vertical through posterior margin of eye; teeth on both jaws
minute, forming a broad band; gillrakers on first gill arch consisting
mostly of rudiments; caudal peduncle
with fleshy lateral keel and dorsal andventral grooves; no scutes. Color:body dark brown, paler below, with
5–7 black oblique bands dorsolaterally; tips of dorsal and anal fin lobes
whitish. Size: maximum length about
70 cm. Distribution: widely in IndoWest Pacific, from East Africa to Australia, north to southern Japan. Remarks: found mainly in offshore
reefs; feeds on benthic fishes, crustaceans and cephalopods.

Seriolina nigrofasciata, KAUM–I. 17157, 20.1 cm SL
off Terengganu (KR), 4 Jan. 2009

Ulua mentalis(Cuvier, 1833)
Whitemouth JackD VIII + I, 21–22; A II + I, 17–18;
GR 23–27 + 51–61 = 74–86; V 10 +14. Body deep, strongly compressed;
dorsal profile strongly convex inadults; breast widely naked to pectoral
fin base and insertion of pelvic fins;teeth on both jaws small, pointed in asingle row in adults, or in a irregularrow or a narrow band in juveniles;
posterior 2 or 3 spines in first dorsal
fin and detached anal fin spines usually embedded; straight part of lateralline almost entirely with 24–39 scutes.
Color: body silvery blue-green dorsally, silvery white ventrally; juveniles
with 7–8 dark bands laterally. Size:maximum length 1 m. Distribution:
widely in Indo-West Pacific, fromSouth Africa to northeastern Australia,
north to southern Japan. Remarks:
occurring in shallow coastal waters.

Seriolina nigrofasciata, UMTF 1889, 27.6 cm SL
off Terengganu (KT), 11 Jan. 2009

Ulua mentalis, KAUM–I. 17114, 8.6 cm SL
off Terengganu (KT), 31 Dec. 2008

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MENIDAE
Moonfish

By Mizuki Matsunuma
This family is represented by asingle species; see the following species account.
Similar families occurring in
the area: none. No other fishes havea combination of characters such as
body very deep, almost triangular,
extremely compressed; long dorsal and anal fin bases; thin, flexible
and rudimental dorsal fin spines;
extremely long pelvic fin, the first 2
soft rays fused.

body very deepand compressedcaudal fin forked

pelvic fin long

Mene maculata(Bloch & Schneider, 1801)
Moonfish
D III–IV + 40–45; A 30–33; P1 15; P2
I, 5; GR 6–8 + 23–25. Body very deep,
almost triangular, extremely compressed; breast sharp-edged; covered
with minute scales, invisible to the naked eye. Mouth small, almost vertical,
protrusible; villiform bands in jaws.
Dorsal fin long-based, and low posteriorly; spines thin and flexible. Anal
fin long-based, commencing at pointof insertion of pelvic fins, uniformly
very low. Caudal fin deeply forked.
Pectoral fins shorter than head length.
Pelvic fins inserted slightly in front ofpectoral fins, first 2 soft rays fused andgreatly elongated. Color: upper sides
deep metallic blue, remainder silvery,
with a row of round to ovoid, darkslaty-blue spots above and below lateral line, sometimes a few additionalspots above and below these rows;
pelvic fins with a trace of blue, otherfins hyaline or slightly dusky. Size:
commonly 20 cm, maximum 30 cm.
Distribution: Indo-West Pacific fromthe east coast of Africa east to Melanesia, Australia north to Japan. Remarks: found in coastal waters to
depths of 200 m. Marketed fresh anddried.

Mene maculata, KAUM–I. 16946, 10.1 cm SL
off Terengganu (KT), 11 Dec. 2008

109

LEIOGNATHIDAE

Slipmouths (Ponyfishes)
By Seishi Kimura
Small to medium-sized marine
and brackish fishes; maximum size
about 25 cm. Body slimy and deep,
ovate, strongly compressed, except
a few species. Head usually naked(cheek scales present in some species) with bony ridges on dorsal
surface, a median bony ridge (crest)
on nape; mouth small, highly protrusible; no pseudobranchiae; gillmembranes united with isthmus.
Unique light organ encircling theposterior part of esophagus. A single
dorsal fin, lacking a distinct notch,with VII–IX spines (usually VIII)
and 15–17 (usually 16) soft rays;
anal fin usually with III spines and
13–15 (usually 14) soft rays; caudal fin forked; pelvic fin I, 5. Scales
small cycloid easily shed. Vertebral
counts 22–24. Color: body generally
silver; dark vertical bars or yellow
spots on body, a black blotch on topof dorsal fin, yellow vertical fin margins in some species.

single dorsal fin with VII–IX, 14–17
head usually naked

forked caudal finmouth highly
protrusible
anal fin with III, 13–14

Remarks: occurring in tropical totemperate marine and estuarine areas.
The generic names follow Kimura et al.
(2008a, b, c).
Similar families occurring in thearea: Leiognathidae is distinguishedfrom the following similar families
in having strongly protrusible mouth,

naked head (except cheek in somespecies), and small cycloid scales
on body. Carangidae – 2 dorsal fins;
mouth not highly protrusible. Gerreidae – scales large; head scaled; scaly
sheath along the bases of dorsal andanal fins well developed.

Equulites laterofenestra(Sparks & Chakrabarty, 2007)
D VIII, 16; A III, 14; P1 17–19; LL
69–77; GR 5–6 + 13–16 = 18–22; V
10 + 14. Body somewhat elongate, compressed; body depth 31–39% SL;
mouth protruding downwards; ventral
profile of lower jaw slightly concave
or almost straight; cheek naked; breast
naked below a line from lower tip of
opercle to pelvic fin insertion; second
dorsal fin spine somewhat elongated.
Color: body brownish silver dorsally,
brilliant silvery-white ventrally; peculiar dark irregular ring marks andoblique wavy dark lines dorsolaterally
on body. Size: maximum length about
12 cm SL. Distribution: West Pacificfrom Philippines to Indonesia. Remarks: this species is distinguishable
from very similar congeners, E. leuciscus and E. stercorarius by having a
naked area on lower breast and peculiar ring marks dorsolaterally on body.

Equulites laterofenestra, KAUM–I. 17110, 8.4 cm SL
off Terengganu (KT), 31 Dec. 2008

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Equulites leuciscus(Günther, 1860)
Whipfin Ponyfish
D VIII, 15–16 (usually 16); A III, 14;
P1 16–19; LL 55–68; GR 5–7 + 12–15= 17–22; V 10 + 14. Body oblong,somewhat elongate, compressed; body
depth 35–47% SL; dorsal and ventral
profiles similarly convex; mouth protruding downwards; ventral profile oflower jaw slightly concave or almost
straight; small slender teeth on both
jaws; cheek naked; breast completely
scaled laterally with narrow naked
area ventrally; second dorsal fin spinedistinctly elongated, filamentous; second anal fin spine also elongated in
adults. Color: body brownish silver
dorsally, brilliant silvery-white ventrally; irregular oblique dark lines dorsolaterally on body; a few yellow
spots sometimes laterally on body.
Size: maximum length about 12 cm
SL. Distribution: widely in IndoWest Pacific, from East Africa to Australia, northward to southern Japan.
Remarks: usually found in shallow
waters; feeds on small crustaceans andpolychaetes.

Equulites leuciscus, KAUM–I. 17266, 11.7 cm SL
off Terengganu (KT), 18 Jan. 2009

Equulites oblongus(Valenciennes, 1835)
Oblong Ponyfish
D VIII, 16; A III, 13–14 (usually 14);
P1 16–19; LL 52–63; GR 4–6 + 12–15= 17–20; V 10 + 14. Body oblong,compressed; body depth 35–46% SL;
dorsal profile more strongly convex
than ventral profile; mouth protruding
downwards; ventral profile of lower
jaw slightly concave; small slender
teeth on both jaws; cheek naked;
breast completely scaled laterally withnarrow naked area ventrally; second
dorsal- and anal-fin spines not distinctly elongated. Color: body brownish silver dorsally, brilliant silverywhite ventrally; irregular oblique dark

Equulites oblongus, KAUM–I. 16861, 6.7 cm SL
off Terengganu (KT), 5 Dec. 2008lines dorsolaterally on body; dorsal, Remarks: usually found in shallow
anal and caudal fins yellowish. Size: waters; feeds on small crustaceans andmaximum length about 10 cm SL. polychaetes.
Distribution: widely in Indo-West
Pacific, from Mauritius to Australia.

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Equulites stercorarius(Evermann & Seale, 1907)
Slender Ponyfish
D VIII, 16; A III, 14; P1 16–18; LL
58– 61; GR 5–6 + 11–14 = 16–20.
Body elongate, moderately compressed;
body depth 30–36% SL; mouth protruding downwards; profile of lower
jaw almost straight; a single row ofsmall slender teeth on jaws; retrorse
spines of dorsal- and anal-fin pterygiophores embedded; cheek, breast, andbelly completely scaled. Color: bodybrownish dorsally, silvery-white ventrally; black blotch on snout; dark irregular oblique lines indistinctly on
dorsolateral body surface; dorsal andanal fins with yellow bands. Size:maximum length about 12 cm. Distribution: found in Philippines, Vietnam, Thailand, Malaysia, and Indonesia. Remarks: inhabits soft bottom
area at depths shallower than 50 m.

Equulites stercorarius, KAUM–I. 16860, 9.9 cm SL
off Terengganu (KT), 5 Dec. 2008

Eubleekeria jonesi(James, 1971)
Yellowlined Ponyfish
D VIII, 16; A III, 14–15 (usually 14);
P1 16–19; LL 50–61; GR 5–7 + 21–24= 24–31; V 10 + 14. Body deep, ovate,strongly compressed; dorsal and ventral profiles similarly convex; lower
margin of eye located above level ofmouth gape; mouth protruding downwards; ventral profile of lower jaw almost straight; small slender teeth on
both jaws; cheek naked; breast almost
completely scaled; semicircular naked
area on nape. Color: head and bodyalmost uniformly silvery-white; tip ofsnout dark; lateral line scales prominent yellow but easily fade out; distal
half of fin membranes between secondand fifth spines of dorsal fin with asomewhat pale black blotch. Size:maximum length about 14 cm SL.
Distribution: Indo-West Pacific, from
Mauritius to Indonesia, northward to
Hainan Island, China, south to Australia. Remarks: usually found in muddy
bottoms in shallow waters.

Eubleekeria jonesi, KAUM–I. 16864, 8.8 cm SL
off Terengganu (KT), 5 Dec. 2008

Eubleekeria jonesi, KAUM–I. 16865, 7.3 cm SL
off Terengganu (KT), 5 Dec. 2008

112

Gazza minuta(Bloch, 1795)
Toothpony
D VIII, 16; A III, 13–14 (usually 14);
P1 14–19; LL 52–69; LGR 13–18; V
10 + 14. Body oval, compressed, somewhat elongated; mouth protruding forwards, not downwards, with distinct
canine teeth on both jaws; a long narrow anterodorsal extension of subocular silvery region, proximal contact
only with orbit; pelvic-fin tips notreaching to origin of anal fin when appressed; scaled area of anterior dorsolateral surface of body extending anteriorly beyond a vertical throughposterior tip of sensory canal on temporal; breast completely naked. Color: body bluish silver dorsally, brilliant silvery-white ventrally; anal fin
yellowish distally; caudal fin yellowish with faintly black margin. Size:maximum length about 16 cm. Distribution: widely in Indo-West Pacific,
from East Africa to northern Australia,
north to southern Japan. Remarks:
feeds on benthic animals and small
fishes.

Gazza minuta, KAUM–I. 16858, 8.8 cm SL
off Terengganu (KT), 5 Dec. 2008

Gazza rhombea
Kimura, Yamashita & Iwatsuki, 2000
Rhomboid Toothpony
D VIII, 16; A III, 14; P1 15–18; LL
56–66; LGR 13–17; V 10 + 14. Body
deep, rhomboid, strongly compressedin adults, oblong in juveniles; dorsal
and ventral profiles similarly convex;
mouth protruding forwards, not downwards, with distinct canine teeth on
both jaws; a long narrow anterodorsal
extension of subocular silvery region,
proximal contact only with orbit; pelvic-fin tips not reaching to origin of
anal fin when appressed; scaled areaof anterior dorsolateral surface ofGazza rhombea, KAUM–I. 17163, 11.5 cm SL (preserved specimen)
off Terengganu (KR), 5 Jan. 2009 (photo by Seishi Kimura)

body not reaching to a vertical throughposterior tip of sensory canal on temporal; breast completely naked. Color: body bluish silver dorsally, brilliant silvery-white ventrally; anal fin
yellowish distally. Size: maximum

length about 18 cm SL. Distribution:
Indo-West Pacific, from Pakistan to
Vanuatu, northward to southern Japan.
Remarks: usually found in sandy or
muddy bottoms in coastal and brackish waters.

113

Leiognathus equulus(Forsskål, 1775)
Common Ponyfish
D VII–VIII, 16–17 (usually VIII,
16); A III, 13–14 (usually 14); P1 19–
22; LL 54–68; GR 4–7 + 13–18 = 19–
24; V 10 + 14. Body extremely deep,
compressed, hump-back shaped; mouth
protruding downwards; small slender
teeth on both jaws; profile of lower
jaw strongly concave; supraorbital
ridge serrated; cheek, breast, and belly
completely naked; second dorsal- andanal-fin spines slightly elongated, but
not filamentous; retrorse spines of
dorsal and anal-fin pterygiophores
strong and exposed. Color: bodyblue-grayish silver dorsally, brilliant
silvery white ventrally; dorsolateralbody with 17–22 narrow dark verticallines. Size: maximum length about 30
cm. Distribution: widely in IndoWest Pacific, from East Africa to Fiji,north to southern Japan and south tonorthern Australia. Remarks: inhabits
shallow coastal areas, usually entering
into estuaries; feeds chiefly on polychaetes, small crustaceans, and small
fishes.

Leiognathus equulus, KAUM–I. 16862, 11.3 cm SL
off Terengganu (KT), 5 Dec. 2008

Nuchequula longicornis
Kimura, Kimura & Ikejima, 2008
Longspine Ponyfish
D VIII, 16–17 (usually 16); A III, 14;
P1 18–19; LL 54–58; GR 5–6 + 16–17= 22–23; V 10 + 14. Body deep, oval,
compressed; dorsal and ventral profiles equally convex; mouth protruding downward; a narrow band ofsmall, slender, villiform teeth in both
jaws; minute, sparse, slender projections on the upper oral valve; cheeknaked; no naked area on nape; breast
naked; second dorsal-fin spine elongated, extending beyond the base of
fifth soft ray when fin appressed; second anal-fin spine not elongated, notreaching to the base of fifth soft ray
when fin appressed. Color: body almost uniformly silver; a distinct dark
blotch on the nape; a yellow marking
on the abdomen between base of pectoral fin and origin of anal fin. Size:maximum length about 80 cm SL.
Distribution: West Pacific, from Gulfof Thailand to Java.

Leiognathus equulus, UMTF 1497 (KAUM–I. 16729), 8.4 cm SL
Setiu, 28 Oct. 2008

Nuchequula longicornis, KAUM–I. 16920, 9.9 cm SL
off Terengganu (KT), 10 Dec. 2008

114

Photopectoralis bindus(Valenciennes, 1835)
Orangefin Ponyfish
D VIII, 15–16 (usually 16); A III, 14;
P1 15–19; LL 51–88; GR 4–7 + 16–20= 21–27; V 10 + 14. Body rounded,disc-like, strongly compressed; ventral profile similarly convex as, ormore strongly convex than ventral
profile; mouth protruding forwards; asingle row of small conical teeth inboth jaws; ventral profile of lower jaw
almost straight or slightly concave;cheek naked; breast almost completely scaled laterally with narrow naked
area ventrally. Color: body brownish
silver dorsally, brilliant silvery-whiteventrally; irregular oblique dark lines
and/or dark vermiculation dorsolaterally on body; spinous portion of dorsal
fin dark orange to yellow distally, anterior anal fin yellowish distally. Size:maximum length about 10 cm SL.
Distribution: Indo-West Pacific, fromRed Sea to New Caledonia, north tosouthern Japan. Remarks: usuallyfound in muddy bottoms of coastal inshore waters.

Photopectoralis bindus, KAUM–I. 17263, 7.5 cm SL
off Terengganu (KT), 18 Jan. 2009

Secutor hanedai
Mochizuki & Hayashi, 1989
Haneda’s Ponyfish
D VIII, 16; A III, 14; P1 15–18; GR
5–7 + 17–20 = 22–26; V 10 + 14. Body
oblong, compressed; ventral profile
more strongly convex than dorsal profile; mouth protruding upwards; minute teeth on both jaws; ventral profile
of lower jaw slightly concave; cheekand breast naked; lateral line incomplete, number of scale pockets along
lateral line to caudal-fin base 60–70;
scale rows above and below lateralline 16–22 and 42–53, respectively.
Color: body silvery brownish green
dorsally, brilliant silvery-white ventrally; 9 or 10 vertical series of darkbars or blotches dorsolaterally on
body. Size: maximum length about 10
cm. Distribution: western and eastern
coasts of Malay Peninsula, Gulf ofThailand, Sumatra, Java, and Borneo.
Remarks: usually found in muddy
bottoms in shallow waters.

Secutor hanedai, KAUM–I. 17276, 5.1 cm SL
off Terengganu (KT), 18 Jan. 2009

Secutor hanedai, KAUM–I. 17275, 3.4 cm SL
off Terengganu (KT), 18 Jan. 2009

115

Secutor indicius
Monkolprasit, 1973
Dots-and-dashes Ponyfish
D VIII, 16; A III,14; P1 16–18; LGR
20–21; V 10 + 14. Body oblong,somewhat elongated, compressed;
ventral profile more strongly convex
than dorsal profile; mouth protruding
upwards; minute teeth on both jaws;
ventral profile of lower jaw concave;cheek naked; breast naked ventrally;
lateral line incomplete, number ofscale pockets along lateral line to caudal-fin base 87–111; scale rows aboveand below lateral line 18–22 and 39–
48, respectively. Color: body silvery
brownish dorsally, brilliant silverywhite ventrally; about 15 irregularvertical dark markings consisted ofdots and dashes dorsolaterally on
body. Size: maximum length about 10
cm. Distribution: West Pacific Ocean,from Gulf of Thailand to New Guinea,
north to southern Japan. Remarks:
usually found in muddy bottoms inshallow waters.

Secutor indicius, KAUM–I. 16857, 8.7 cm SL
off Terengganu (KT), 5 Dec. 2008

Secutor indicius, KAUM–I. 17265, 8.4 cm SL
off Terengganu (KT), 18 Jan. 2009

116

LUTJANIDAE
Snappers

By Siti Tafzilmeriam S. A. K.
Typical perch-like fishes, bodysymmetrical and moderately compressed, oblong in shape (size to 160cm). Eye usually moderate; premaxilla usually moderately protrusible;
mouth terminal and fairly large;
supramaxilla absent. Jaws usuallywith more or less distinct canines;
vomer and palatines usually withteeth; pterygoids usually toothless.
Cheek and operculum scaly; snout,lacrimal, and lower jaw naked. Dorsal fin single, continuous or slightly
notched, with X–XII spines and 10–
19 soft rays; anal fin with III spinesand 7–11 soft rays; pectoral fin rays
14–19; pelvic fins with I spine and 5
soft rays; caudal fin shape variablefrequently truncate, emarginated or
lunate. Scales moderate to rather
small, ctenoid. Lateral line complete.
Pelvic axillary process usually welldeveloped. Branchiostegal rays 7.
Vertebrae 24 (10 + 14). Color: highly variable; mainly from red throughyellow to blue; often with blotches,lines, or other patterns.
Remarks: mostly bottom-associated fishes, occurring from shallow
inshore areas to depths of about 500

single dorsal fin with X–XII, 10–19

preopercle scaled

ctenoid scales

snout andlacrimal naked

anal fin with III, 7–11

m, mainly over reefs or rocky outcrops. Active predators, mostly nocturnal, feeding on crustaceans, mollusks, and fishes.
Similar families occurring in
the area: Caesionidae – mouth small;premaxillae extremely protrusible;
teeth small and minute. Haemulidae –scales present on snout and lacrimal;
preoperculomandibular canal lateralsystem opening under chin; usuallyno teeth on vomer and palatines.
Lethrinidae – opercular membranes

broadly united to each other; preopercular margin typically smooth;branchiostegal rays 6; soft rays of
dorsal fin 9 or 10; no teeth on vomer
or palatines. Nemipteridae – preopercular margin typically smooth; softrays of dorsal fin 9; soft rays of anal
fin usually 7; branchiostegal rays 6.
Sparidae – preopercular margin typically smooth; branchiostegal rays 6;teeth in jaws variable-conical, incisiform, or molariform.

Lutjanus argentimaculatus(Forsskål, 1775)
Mangrove Red Snapper
D X, 13–14; A III, 8; P1 16–17; P2 I,
5; LLp 44–48; GR 6–8 + 9–12. Body
moderately deep, its depth 2.5–3.1 instandard length. Preorbital bone relatively broad, wider than eye diameter;
preopercular notch and knob poorly
developed. Posterior profile of dorsal
and anal fins rounded; caudal fin
emarginated to nearly truncate. Scalerows on back more or less parallel tolateral line, or parallel below spinouspart of dorsal fin and sometimes risingobliquely posteriorly, or rarely withentirely oblique rows. Color: generally greenish brown on back, grading
to reddish on sides and ventral parts;
scales usually with dark centers andwhite margins, giving a reticulated appearance; pectoral fin hyaline; pelvic
and anal fins dark brown becoming

Lutjanus argentimaculatus, UMTF 1519 (KAUM–I. 16722), 16.9 cm SL
Setiu, 28 Oct. 2008

paler towards base; specimens fromdeep water frequently overall reddish;juveniles with a series of about 8 whitish bars crossing sides, and 1 or 2 bluelines across cheek. Size: 120 cm. Dis-

tribution: Indo-West Pacific. Remarks: a euryhaline species. Juveniles and young adults occur inmangrove estuaries and in the lower
reaches of freshwater streams.

117

Lutjanus fulviflamma(Forsskål, 1775)
Blackspot Snapper
D X, 12–14; A III, 8; P1 16–17; P2 I,
5; LLp 46–49; GR 6–7 + 9–12. Body
moderately deep to somewhat slender;
its depth 2.6 to 2.9 times in standard
length. Preopercular notch and knob
poorly developed. Caudal fin truncate
or slightly emarginate. Scale rows onback rising obliquely above lateralline. Color: back and upper sides reddish brown; lower sides whitish or
light brown; whitish to yellow on belly; 6 or 7 yellow stripes on sides; aprominent black spot at level of lateralline below base of anterior part ofsoft-rayed portion of dorsal fin; fins
yellowish. Size: 35 cm. Distribution:
Indo-Pacific. Remarks: inhabits coastal waters around coral reef areas. Juveniles sometimes found in estuary.

Lutjanus fulviflamma, KAUM–I. 16504, 7.5 cm SL
Cendering, 6 Oct. 2008

Lutjanus lutjanus(Bloch, 1790)
Bigeye Snapper
D X–XII, 12; A III, 8; P1 16–17; P2 I,
5; LLp 48–50; GR 6–8 + 17–19. Body
fusiform, slender; its depth 2.9 to 3.3
times in standard length. Preorbitalbone very narrow, much less than eye
diameter. Vomerine tooth patch triangular, with a medial posterior extension. Caudal fin truncate or slightly
emarginated. Scale rows on back rising obliquely above lateral line. Color: upper back golden brown; a broad
yellow to brownish stripe from eye tocaudal-fin base; yellow horizontal
lines (1 per scale row) on lower half ofbody, and similar lines runningobliquely above lateral line; fins yellowish. Size: 30 cm. Distribution: Indo-West Pacific. Remarks: inhabits
offshore coral reefs.

Lutjanus lutjanus, UMTF 1010 (KAUM–I. 16639), 12.4 cm SL
Bidong Island, 12 Oct. 2008

Lutjanus malabaricus(Bloch & Schneider, 1801)
Malabar Blood Snapper
D XI, 12–14; A III, 8–9; P1 16–17; P2
I, 5; LLp 46–50; GR 4–7 + 12–14.
Body relatively deep; its depth 2.2 to
2.8 times in standard length. Preopercular notch and knob poorly developed. Vomerine tooth patch crescentic
or triangular, without a medial posterior extension. Caudal fin truncate.
Scale rows on back rising obliquely

Lutjanus malabaricus, KAUM–I. 17019, 11.5 cm SL
off Terengganu (KT), 16 Dec. 2008

above lateral line. Color: back andsides red or red-orange, lighter onlower parts; fins reddish; juveniles
with a broad, oblique band of brownor black from upper jaw to beginningof dorsal fin, and a prominent blackband across caudal peduncle with apearly white anterior border. Size: 100
cm. Distribution: Indo-West Pacific.
Remarks: inhabits both coastal andoffshore reefs; depth range 12–100 m.

118

Lutjanus quinquelineatus(Bloch, 1790)
Five-lined Snapper
D X, 13–15; A III, 8; P1 16–17; P2 I,
5; LLp 47–50; GR 7–8 + 13–15. Body
moderately deep, its depth 2.3–2.9 instandard length. Preorbital width usually less than eye diameter; preopercular notch and knob well developed.
Caudal fin truncate or slightly emarginate. Scale rows on back risingobliquely above lateral line. Color:body generally bright yellow, with 5
blue stripes on side; 4 originating frombehind eye and the other originatingfrom cheek below eye; a round black
spot, about the size of eye or larger, onside below soft-rayed portion of dorsal
fin; fins yellowish. Size: 38 cm. Distribution: Indo-Pacific. Remarks:
inhabits sheltered lagoons and exposed outer slope coral reefs at depth
range of 2–40 m.

Lutjanus quinquelineatus, KAUM–I. 16974, 12.5 cm SL
off Terengganu (KT), 12 Dec. 2008

Lutjanus russellii(Bleeker, 1849)
Russell's Snapper
D X, 14; A III, 8; P1 16–17; P2 I, 5;
LLp 47–50; GR 6–7 + 7–11. Body
moderately deep to somewhat slender;
its depth 2.6-2.8 in standard length.
Anterodorsal profile of head steeply tomoderately sloped; preorbital widthabout equal to, or slightly less than eye
diameter. Caudal fin truncate or slightly emarginate. Scale rows on back rising obliquely above lateral line. Color: body brownish dorsally, pink towhitish ventrally with a silvery sheen;a large black spot, sometimes faint, on
lateral line below the anterior portionof soft-rayed portion of dorsal fin;adults usually with 6–8 narrow goldenbrown stripes on sides; juveniles,
whitish with about 4 black stripes onsides, and a round black spot on upper
back. Size: 45 cm. Distribution: Indo-West Pacific.Remarks: juveniles
are often found in mangrove areas.
Marketed fresh.

Lutjanus vitta(Quoy & Gaimard, 1824)

Lutjanus russellii, UMTF 1043 (KAUM–I. 16796), 7.8 cm SL
Merang, 2 Nov. 2008

standard length. Preorbital widthabout equal to eye diameter; preoperBrownstripe Snapper
cular notch and knob poorly develD X, 12–13; A III, 8–9; P1 15–16; P2 oped. Caudal fin emarginated. Scales
I, 5; LLp 49–51; GR 6–7 + 9–12. rows on back rising obliquely above
Body moderately deep to relatively lateral line. Color: generally whitish
slender; its depth 2.6 to 3 times in or pink, with a dark brown to blackish

Lutjanus vitta, KAUM–I. 16941, 10.9 cm SL
off Terengganu (KT), 11 Dec. 2008

stripe on side from eye to upper half ofcaudal peduncle; narrow brown lines,
1 per scale row, on sides; fins yellowish. Size: 40 cm. Distribution: IndoWest Pacific. Remarks: inhabits thevicinity of coral reefs.

119

CAESIONIDAE
Fusiliers

By Mizuki Matsunuma
Medium sized (up to 50 cm) marine fishes. Body oblong to fusiform,
moderately compressed; longitudinal
axis from tip of snout to middle ofcaudal fin passing through center ofeye. Eye moderately large, its diameter longer than snout length. Mouth
small and highly protrusible. Dentition variously reduced; small or minute conical teeth; premaxilla, vomer,
and palatines with or without teeth.
Dorsal fin with X–XV slender weakspines and 8–22 soft rays; anal fin
with III spines and 9–13 soft rays;
pelvic fin with I spine and 5 soft rays;
pectoral fin with 16–24 rays; caudal
fin distinctly forked, with pointed
lobes. Branchiostegal rays 7. Scales
moderate to small, weakly ctenoid;
lateral line scales 45–88; scale rows
on body running horizontally; dorsal and anal fins with scales in most
species. Color: sides with or without
longitudinal stripes; caudal fin either without markings, with blackishblotch on tips of lobes, or with a longitudinal blackish streak in middle ofeach lobe; axil of pectoral fins black.
Remarks: found mostly oncoral reefs; occur near the surface tosingle dorsal fin with X–XV, 8–22

forked caudal fin

weakly ctenoid scales

anal fin with III, 9–13

depths of 60 m. Feed by picking zooplankton. Major importance in coralreef fisheries.
Similar families occurring in
the area: Lutjanidae – closely allied
to Caesionidae and difficult to distinguish from Caesionidae on the basisof any single external character, but
most members of Lutjanidae having
a deeper body, the eye well above thehorizontal axis of the body, and lacking a strongly forked caudal fin; thoselutjanid genera with the horizontal
passing near the center of eye either
having scale rows running obliquely
upward, fewer than 9 anal fin rays, noscales on dorsal and anal fins, or thecaudal fin much less deeply forked.
Nemipteridae – eye above horizontal
axis in most species; 9 dorsal and 7
or 8 anal fins soft rays. Lethrinidae– eye always above horizontal axisof body; base of soft part of dorsal
fin generally shorter than base ofspinous part; 8–10 anal fin soft rays;
usually enlarged canines in frontof jaws, sometimes lateral molars.
Emmelichthyidae – superficially similar but caught in deep water; maxilla
fully scaly; dorsal fin with IX spines
and usually 12 soft rays.

Caesio cuning(Bloch, 1791)
Redbelly Yellowtail Fusilier
D X, 14–16; A III, 10–12; P1 17–20;
LL 45–51. Body fairly deep and compressed. A single postmaxillary process; small conical teeth in jaws, vomer and palatines. Scale rows above
lateral line 7–9; supratemporal band ofscales confluent at dorsal midline; 4–5
scales on cheek; predorsal scales 20–
26; dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest
spinous height covered with scales;supratemporal band of scales confluent at dorsal midline. Color: head andbody bluish white, pinkish ventrally;
posterior portion of back, upper caudal peduncle, and caudal fin yellow;
axil and upper base of pectoral fin
black; dorsal fin yellow posteriorly
and grayish blue anteriorly; pelvic andCaesio cuning, KAUM–I. 16984, 10.5 cm SL
off Terengganu (KT), 13 Dec. 2008

anal fins reddish. Size: maximum
length 50 cm. Distribution: Indo-West
Pacific. Remarks: inhabits coastal areas to the depth of 60 m.

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Dipterygonotus balteatus(Valenciennes, 1830)
Mottled Fusilier
D XII–XV, 8–11; A III, 9–11; P1
17–19; LL 68–80. Body slender, fusiform, elongate, and moderately compressed. Two postmaxillary processes;
small conical teeth on dentary and vomer; premaxilla and palatines without
teeth. Scales above lateral line to origin of dorsal fin 9–11; scales below
lateral line to origin of anal fin 15–18;
usually 6–9 scale rows on cheek; predorsal scales usually 29–34; dorsal
and anal fins without scales. Color:body brownish bronze dorsally, silvery white ventrally; with a thin,straight, tan stripe about 1 scale widefrom orbit to caudal fin, above lateralline; above and parallel to this stripe 2
thin, irregular, and usually interrupted
stripes of same color; caudal fin tan topinkish. Size: maximum length 14 cm.
Distribution: Indo-West Pacific. Remarks: feeds on zooplankton. Marketed fresh.

Dipterygonotus balteatus, KAUM–I. 17169, 7.9 cm SL
off Terengganu (KR), 5 Jan. 2009

Pterocaesio chrysozona, KAUM–I. 16989, 12.8 cm SL
off Terengganu (KT), 14 Dec. 2008

Pterocaesio chrysozona(Cuvier, 1830)
Goldband Fusilier
D X–XI, 14–16; A III, 11–12; P1
17–20; LL 64–69. Body fusiform,
elongate, and moderately compressed.
Two postmaxillary processes; small
conical teeth in jaws, vomer, and palatines. Scales above lateral line to origin of dorsal fin 7–9; scales below
lateral line to origin of anal fin 14–16;
predorsal scales usually 23–26; dorsal
and anal fins scaly, dorsal fin with
about 1/2 of its greatest spinous heightcovered with scales. Color: body light
blue to brownish dorsally, white topinkish ventrally; a bright yellow banddirectly below lateral line for most of
its length, from behind eye to base of
caudal fin, 2 to 3 scales wide anteriorly, tapering to 1 scale in width on caudal peduncle where it is above lateralline; a less conspicuous yellow stripe
along dorsal midline; dorsal fin slightly dusky distally; tips of caudal-fin
lobes black. Size: maximum length 21
cm. Distribution: Indo-West Pacific.
Remarks: schooling fish found incoral reefs.

Pterocaesio digramma, KAUM–I. 17226, 14.9 cm SL
off Terengganu (KT), 12 Jan. 2009

Pterocaesio digramma(Bleeker, 1864)
Doublelined Fusilier
D X, 14–16 (usually 15); A III, 11–
12 (usually 12); P1 20–22; LL 66–75.
Body fusiform, elongate, and moderately compressed. Two postmaxillary
processes; small conical teeth in jaws,
vomer, and palatines. Scales above
lateral line to origin of dorsal fin 9–11;
scales below lateral line to origin of
anal fin 17–18; cheek with 4–5 scalerow; upper peduncular scales usually
12–13; dorsal and anal fins scaly. Color: body blue to greenish dorsally,

white ventrally; with 2 yellow longitudinal stripes laterally, the lower running from behind upper orbit about 1
scale below the lateral line to a verticalat about middle of soft portion of dorsal fin where it crosses and usually
passes above lateral line for the lengthof caudal peduncle; upper stripe 1–2
scales below dorsal profile foremostof length of body, ending dorsally on
caudal peduncle; caudal fin lobes withblack tips. Size: maximum length 30
cm. Distribution: western Pacific. Remarks: schooling fish found in coastal
areas, primarily around coral reefs. Marketed fresh and dried salted.

121

GERREIDAE
Mojarras

By Seishi Kimura and Siti Tafzilmeriam S. A. K.
Small to medium sized marine
and brackish fishes; maximum size
about 40 cm. Body compressed,
depth variable, oblong to rhomboidal. Head scaled; mouth small,
highly protrusible; teeth on jaws
small, brush-like; gill membrane freefrom isthmus. A single dorsal fin,lacking a distinct notch, with IX–X
spines and 9–17 soft rays; anal fin
usually with II–IV spines and 6–18
soft rays; scaly sheath along bases of
dorsal and anal fins; pelvic fin I, 5;
caudal fin deeply forked. Scales thin,
cycloid, moderately large, but deciduous. Vertebral counts 24. Color:body generally brilliant silver, withdark vertical bars or spots on body insome species.
Remarks: occurring in tropicalto temperate marine and estuarine areas. Feed mainly on benthic animals.

scaly sheathcaudal fin deeply forked

head scaled
large cycloid scales

mouth highly
protrusible

scaly sheath

Similar families occurring in
the area: Gerreidae is distinguishedfrom the following similar families
in having strongly protrusible mouth,
scaled head, moderately large cycloid

scales on body, and scaly sheathalong the bases of dorsal and anal
fins. Haemulidae, Lutjanidae, andSparidae – mouth not highly protrusible.

Gerres erythrourus(Bloch, 1791)
Deepbody Silverbiddy
D IX, 10; A III, 7; P1 15–16; LL 35–
38; GR 4–5 + 8–9. Body deep, compressed; anterodorsal profile almost
straight. Dorsal- and anal-fin spines
strong; second dorsal-fin spine longer
than the third; second anal-fin spine
robust, nearly the same in length ofanal-fin base; caudal fin short, deeply
forked, tips broadly rounded. Color:body brownish silver dorsally, brilliant silver ventrally, with indistinct
dark longitudinal stripes along scalerows above and 4–6 rows immediately
below lateral line; often 4–11 indistinct vertical narrow dark bars on sideof body; anal and pelvic fins yellow.
Size: maximum length about 37 cm.
Distribution: Indo-Pacific, fromwestern India to Micronesia. Remarks: inhabits soft bottom. Juveniles often observed in estuarine waters. Caught by bottom set nets andbottom trawls. Marketed fresh or driedand salted.
(S. Kimura & S. Tafzilmeriam)

Gerres erythrourus, KAUM–I. 16879, 11.8 cm SL
off Terengganu (KT), 5 Dec. 2008

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Gerres filamentosus
Cuvier, 1829
Whipfin Mojarra
D IX, 10; A III, 7; P1 15–16; LL 43–
46 (usually 44–45); GR 4–6 + 8. Body
deep, compressed, anterodorsal profile somewhat convex. Second dorsalfin spine very long and filamentous,
reaching slightly beyond base of lastdorsal-fin ray; anal-fin spines generally robust, third longest; caudal fin
deeply forked, lobes pointed. Preopercle with 3 scale rows; 4½–5½ scales
between 5th dorsal fin spine base andlateral line. Color: body brownish silver dorsally, brilliant silver ventrally,
with vertical rows of indistinct dark
ovoid spots laterally in adults and subadults; all fins tinged with tan, caudal
fin with a blackish margin. Size: maximum length about 32 cm. Distribution: widely distributed in tropicalIndo-Pacific, from eastern coast of
Africa to Micronesia. Remarks: inhabits sandy or muddy bottoms incoastal areas to depths of at least 50m.
Marketed fresh or dried, sometimes
used in making fish crackers. Caught
mainly by beach seines and bottom
trawls. (S. Kimura & S. Tafzilmeriam)

Gerres filamentosus, KAUM–I. 16878, 17.8 cm SL
off Terengganu (KT), 5 Dec. 2008

Gerres limbatus
Cuvier, 1830
Saddleback Silverbiddy
D IX, 10; A III, 7; P1 15; LLp 34–36.
Body relatively deep and then, becoming slender with growth. Usually 2 ½
scale rows between 5th dorsal-fin
spine base and lateral line; usually 4 ½
scales above lateral line, 7 ½–9 ½
scales below; 2nd dorsal-fin spine
slightly shorter than 3rd dorsal-fin
spine; 2nd anal-fin spine more robust
than 3rd. Color: body silvery with 4
or 5 diffuse, dark saddle patches mainly along back in life, extending downsides to midline. Pelvic and anal finspale yellow. Size: maximum length
about 12 cm SL; commonly less than
10 cm. Distribution: eastern Indian
Ocean and West Central Pacific. Remarks: found in estuaries, lower
reaches of coastal rivers.
(S.Tafzilmeriam)

Gerres limbatus, UMTF 1527 (KAUM–I. 16732), 8.0 cm SL
Setiu, 28 Oct. 2008

Gerres limbatus, UMTF 1175 (KAUM–I. 16801), 3.7 cm SL
Merang, 2 Nov. 2008

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Gerres macracanthus
Bleeker, 1854
Longspined Silverybiddy
D IX, 10; A III, 7; P1 15; LLp 41–44.
Body relatively slender and compressed. Usually 4 ½ scale rows between 5th dorsal-fin spine base andlateral line; 5 ½ or 6 ½ scales above
lateral line, 9 ½ or 10 ½ scales below.
Second dorsal-fin spine elongated andfilament; 2nd and 3rd anal-fin spines
short. Caudal fin deeply forked. Color: body silvery with usually 6–10
(rarely up to 14 as faint bands in larger
specimens) indistinct vertical darkbands on body; pectoral, pelvic, andanal fins yellowish. Size: maximum
total length 20 cm. Distribution: Indo-West Pacific. Remarks: found incoastal waters to depths of at least 30
m.
(S. Tafzilmeriam)

Gerres macracanthus, KAUM–I. 16963, 11.5 cm SL
off Terengganu (KT), 11 Dec. 2008

Gerres oyena(Forsskål, 1775)
Common Silverbiddy
D IX, 10; A III, 7; P1 15–17; LL
35–40; GR 3–6 + 8–9 (usually 8).
Body oblong, compressed, anterodorsal profile convex. Posterior tip ofmaxilla extending beyond anterior
margin of eye. Second dorsal-fin spine
not elongated; pectoral fins short, notreaching to anal-fin origin. Scales
mostly absent on premaxillary groove;
scale rows above lateral line 4. Color:body silvery, sometime with faint darkvertical bands laterally; tip of dorsal
and caudal fins with black margins; arow of small dusky spots on dorsal-fin
membrane near base; anal and pelvic
fins yellowish. Size: maximum length
about 25 cm. Distribution: Indo-Pacific.
Remarks: inhabits inshore, especially
soft bottom area at depths shallower
than 30 m.
(S. Kimura & S. Tafzilmeriam)

Gerres oyena, KAUM–I. 16509, 6.4 cm SL
Cendering, 6 Oct. 2008

Pentaprion longimanus(Cantor, 1850)
Longfin Silverbiddy
D IX–X, 12–15; A V–VI, 12–14; P1
17; LLp 44–48. Body compressed,
ovoid, moderately deep, its depth 2.5–
2.8 times in standard length. Anal-finbase longer than base of soft-rayed
portion of dorsal fin; pectoral fins
long; caudal fin deeply forked withpointed lobes. Color: body silvery

Pentaprion longimanus, KAUM–I. 17087, 9.1 cm SL
off Terengganu (KT), 28 Dec. 2008

with a mirror-like stripe from snout to inhabits inshore areas on muddy-sandcaudal peduncle; fins dusky yellow. bottoms to depths of 70 m. Marketed
Size: maximum length 20 cm. Distri- fresh; commonly used for fishmeal.
bution: Indo-West Pacific. Remarks:(S. Tafzilmeriam)

124

HAEMULIDAE
Sweetlips (Grunts)

By Hiroyuki Motomura, Koichi Shibukawa and Mizuki Matsunuma
Medium-sized (up to 120 cm),
oblong and compressed fishes. Lateral line continuous. Head almost
entirely scaled, exclusive of lips,
chin and tip of snout; mouth small
or moderate in size, subterminal;
lips thick in Plectorhinchus; chinwith distinct pores; hind margin of
lacrimal not exposed; opercle with asingle spine; teeth conical, forming
narrow band in each jaw; teeth onoutermost row of jaws enlarged, but
not canine-like; palatine toothless;branchiostegals 7. A single dorsal fin
with IX–XV strong spines (second
spine usually very strong) and 11–26
soft rays; anal fin with III spines and6–18 soft rays; pelvic fin below baseof pectoral fin, with I spine and 5 softrays; caudal fin truncate or emarginated in adults, rounded in juveniles.
Scales small and ctenoid. Vertebrae
26–27. Color: highly variable, appearing characteristic pattern (e.g.,banded and spotted) in each species;
in many species, juveniles strikingly
differ from adult in color.

single dorsal fin with IX–XV, 11–26

head scaled

ctenoid scales
thick lips
anal fin with III, 7–9
2nd anal-fin
spine strong

chin withpores

Remarks: inhabit coastal waters
including reefs, bays, and estuaries,
down to about 80 m. Carnivorous,
feed on small benthic invertebrates or
fishes. Esteemed as food fish, caught
by spear, line, and various nets; marketed fresh or salted.
Similar families occurring in thearea: Lutjanidae – suborbital area na-ked; palatine usually toothed. Lethrinidae – no scales on preopercle;
dorsal fin with 9–10 soft rays. Nemipteridae – no pores on chin; hindmargin of lacrimal exposed. Sparidae– suborbital area naked; preopercularmargin not serrated.

Diagramma picta picta(Thunberg, 1792)
Yellow-spotted Slatey
D IX–X, 20–26; A III, 7; P1 16–18;
LLp 55–74; GR 17–23. Body deepand compressed. Profile of snout
steep; lips fleshy; chin with six pores;
lower jaw without longitudinal grooveat midline. Color: varying much bysize; large adult silvery gray, with orwithout dusky spots; subadult pale
gray with numerous bright yellow or
dusky spots or lines on head, body andfins; juvenile with broad longitudinal
black bands on body. Size: commonly
45 cm, maximum 100 cm. Distribution: Indo-West Pacific. Remarks:
inhabits rocky and coral reefs andsandy bottoms. Nocturnal, forming
small aggregations by day and dispersing at night for feeding; feeds on
small benthic invertebrates. Johnson
et al. (2001) regarded it as a valid subspecies.
(K. Shibukawa)Diagramma picta picta, KAUM–I. 16901, 12.3 cm SL
off Terengganu (KT), 6 Dec. 2008

125

Plectorhinchus gibbosus(Lacepède, 1802)
Harlequin Sweetlip
D XIII–XIV, 15–17; A III, 7–8; P1
16–18; P2 I, 5; LLp 46–55; GR 8–10 +18–20. Body deep, compressed. Lip
fleshy, greatly swollen with age. Chinwith 6 pores; no median pit; not covering barbels or papillae. Caudal fin
rounded in juveniles, rounded to truncate in adults. Color: body dark gray.
Centers of scales paler than edges. No
distinct spots or blotches on fins. Size:maximum about 75 cm. Distribution:
widely distributed in the Indo-West
Pacific, from South Africa east to the
Samoa Islands, and southern Japan
south to New South Wales, Australia.
Remarks: generally found in silty
reef areas, young penetrating estuarine
habitats.
(H. Motomura)

Plectorhinchus gibbosus, KAUM–I. 17058, 38.7 cm SL
off Terengganu (KT), 24 Dec. 2008

Pomadasys argenteus(Forsskål, 1775)
Silver Grunt
D XII, 13–14; A III, 7; P1 16–18; P2 I,
5; LLp 47–49. Body deep, compressed; head blunt. Lip fleshy, greatly
swollen with age. Chin with 2 pores
and a median pit. Circumpeduncularscales 21 or 22; 5 scales between lateral line and dorsal-fin origin. Caudal
fin emarginate. Color: in adults, bodysilvery with dark blotches on dorsal
fin and numerous scattered darkbrown to blackish spots on body side.
Juveniles with body pale brownish,lighter below, back with irregular longitudinal streaks on alternate scalerows; dorsal fin with dusky membranes; a dark spot on gill cover. Size:
commonly 40 cm, maximum 60 cm.
Distribution: Indo-West Pacific fromthe Red Sea to Melanesia, northern
Australia north to southern Japan. Remarks: found in coastal inshore waters in open bays and estuaries. Marketed fresh.
(M. Matsunuma)

Plectorhinchus gibbosus, UMTF 1011 (KAUM–I. 16672), 10.5 cm SL
Kemaman, 21 Oct. 2008

Pomadasys argenteus, KAUM–I. 17252, 13.1 cm SL
off Terengganu (KT), 17 Jan. 2009

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Pomadasys kaakan(Cuvier, 1830)
Javelin Grunt
D XII, 13–15; A III, 7–8; P1 17–18; P2
I, 5; LLp 43–50; GR 5–6 + 13–14.
Body moderately elongate, compressed. Chin with two pores and amedian pit. Caudal fin emarginate.
Color: body silver with six to ten vertical series of blackish spots or double
spots in four rows, third row along lateral line; spots less distinct withgrowth. No blotches or spots on fins.
Size: maximum length about 80 cm.
Distribution: Indo-West Pacific. Remarks: inhabits turbid inshore waters
with sandy to muddy bottoms todepths of 75 m.
(H. Motomura)

Pomadasys kaakan, KAUM–I. 17000, 11.2 cm SL
off Terengganu (KT), 14 Dec. 2008

Pomadasys maculatus(Bloch, 1793)
Saddle Grunt
D XII, 13–14; A III, 7; P1 17; P2 I, 5;
LLp 50–52; GR 5–6 + 13–15. Body
moderately elongate, compressed. Chinwith two pores and a median pit. Posterior portion of maxilla slightly covered or not covered by lacrimal laterally in western Pacific population;
largely covered by lacrimal in Indian
Ocean population. Caudal fin slightly
emarginate. Color: body silver with alarge black blotch across nape, extending downward to below lateral line;
three broken black blotches on upper
half of trunk. A large dark blotch on
membrane centrally between secondand seventh dorsal-fin spines. Size:maximum length 59.3 cm. Distribution: Indo-West Pacific. Remarks:
occurs in coastal inshore waters inopen bay and estuaries to depths ofless than 110 m.
(H. Motomura)

Pomadasys maculatus, KAUM–I. 16835, 15.7 cm SL
off Terengganu (KT), 5 Dec. 2008

Pomadasys unimaculatus
Tian, 1982
Red Patched Grunter
D XII, 13–14; A III, 7; P1 16–17; P2 I,
5; LLp 47–50; GR 3–4 + 12–13. Body
moderately elongate, compressed. Chinwith two pores and a median pit. Middle and posterior portion of maxilla
largely covered by lacrimal laterally.
Caudal fin slightly emarginate. Color:body silver with a large dark brown
blotch across nape, extending downward to lateral line; four small faintly
dark semicircular patches on upper halfof trunk. A large dark red blotch on

Pomadasys unimaculatus, KAUM–I. 17085, 16.1 cm SL
off Terengganu (KT), 28 Dec. 2008

membrane distally between third and man Sea. Remarks: occurs in coastal
sixth to seventh dorsal-fin spines. Size: inshore waters on sandy and muddy
maximum length about 25 cm. Distri- bottoms.
(H. Motomura)bution: the South China Sea and Anda-

127

NEMIPTERIDAE

Threadfin Breams and Monocle Bream
By Koichi Shibukawa, Hiroyuki Motomura and Mizuki Matsunuma
Medium sized (up to 35 cm),
moderately elongate fishes. Headcompressed; eyes moderately large;
jaws subequal, or upper jaw slightly
beyond lower jaw; hind margin ofsuborbital bone exposed; branchiostegal rays 6. Single dorsal fin withX spines and 9 soft rays; anal fin with
III spines and 7–8soft rays; pectoral
fin more or less falcate, with 14–19
soft rays (uppermost 2 rays unbranched); pelvic fin with I spine and5 soft rays; caudal fin emarginated,
lunate or forked, with filamentous
tip on each or both lobes in many
species. Scales ctenoid; cheek andoperculum scaled; a pair of elongate,
pointed scales at base of pelvic fin.
Color: pinkish, yellowish, grayish,or brownish dorsally, turned to silvery ventrally, with various-coloredvertical bands or longitudinal stripes
on head and body in many species.
Remarks: found in coral reefs
and coastal to offshore shelf waters

single dorsal fin with X, 9ctenoid scales

anal fin with III, 7–8

elongate, pointed
scale at pelvic-fin basewith sandy or muddy bottoms, down
to about 300 m depth. Carnivorous,
feed on small fishes, crustaceans,
cephalopods, and polychaetes. Esteemed as food fish in many species;
marketed fresh, dried and salted, or
minced for fish balls or fish cakes.
Similar families occurring in thearea: Caesionidae – dorsal fin with
X–XV spines and 8–22 soft rays; anal

fin with III spines and 9–13 soft rays;
no elongated, pointed scales at baseof pelvic fin. Haemulidae – dorsal fin
with IX–XIV spines and 11–26 soft
rays. Lethrinidae – anal fin with III
spines and 8–10 soft rays; no scales
on preopercle in Lethrinus (scaly inother genera). Lutjanidae – no elongate and pointed scales at base ofpelvic fin.

Nemipterus bathybius
Snyder, 1911
Yellowbelly Threadfin BreamD X, 9; A III, 7; P1 15–17; P2 I, 5; GR
13–16. Body moderately elongate,
compressed; body depth 2.9–3.6 in
SL. Posterior margin of suborbital andlower preopercular margin not serrated; suborbital spine absent. Ventral
margin of orbit just reaching or notreaching a horizontal line throughsnout tip and upper base of pectoral
fin. First 2 dorsal-fin spine separated
by a membrane; membranes between
dorsal-fin spines not strongly incised.
Upper lobe of caudal fin falcate or
ribbon-like extension (broken in thephoto specimen). Pectoral fin notreaching a vertical through anal fin
origin. Three transverse scale rows on
preopercle. Color: body pinkish dorsally, silvery ventrally; belly yellow.
Two yellow stripes, first below lateralline from behind opercle to upper caudal-fin base; second from behind upper of pectoral fin base to middle ofNemipterus bathybius, KAUM–I. 17133, 11.4 cm SL
off Terengganu (KT), 1 Jan. 2009caudal-fin base. Caudal fin pink, upper lobe and filament yellow. Size:maximum 20 cm SL. Distribution:
western Pacific, including southern
Japan, South China Sea, Southeast
Asian countries and northwestern
Australia. Remarks: usually occurs
on sandy or muddy bottoms at depthsof 35 to 300 m, but most common between 45 and 90 m. Feeds mainly on
crustaceans, small fishes, and cephalopods in adults and copepods, ostracods, and amphipods in young.
(H. Motomura)

128

Nemipterus furcosus(Valenciennes, 1830)
Forktailed Threadfin BreamD X, 9; A III, 7; P1 16–18; P2 I, 5; GR
7–10. Body moderately elongate, compressed; body depth 3–3.9 in SL. Snout
length equal to or greater than eye diameter. Suborbital spine absent; 4–6
transverse scale rows on preopercle.
Ventral margin of orbit tangent to or
above just reaching or not reaching ahorizontal line through snout tip andupper base of pectoral fin. Pectoral finreaching to or just short of a verticalthrough anus; pelvic fins short, reaching to or short of a vertical throughanus; caudal fin deeply forked; upper
lobe of the fin pointed, not produced.
Color: body pinkish dorsally, silvery
ventrally. Nine indistinct reddish saddle
bars on back; forming a reddish spot behind origin of lateral line. Dorsal fin
pale rosy, sometimes with yellowish
tinge, outer margin darker pink; a yellow stripe submarginally. Anal fin bluish white, with row of transparent or
faint yellowish spots near base. Caudal
fin pale rosy, with yellow tinge; lower
margin of fin white; posterior tip of upper lobe red. Size: maximum length 20
cm SL. Distribution: eastern Indian
Ocean and West Pacific. Remarks: occurs on sand and mud bottoms in depths
of 8–110 m.
(M. Matsunuma)

Nemipterus furcosus, UMTF 1080 (KAUM–I. 16573), 13.0 cm SL
Bidong Island, 13 Oct. 2008

Nemipterus hexodon, KAUM–I. 17017, 8.6 cm SL
off Terengganu (KT), 16 Dec. 2008

Nemipterus hexodon(Quoy & Gaimard, 1824)
Ornate Threadfin BreamD X, 9; A III, 7; P1 15–18; P2 I, 5; GR
11–17. Body moderately elongate,
compressed. Posterior margin of suborbital and lower preopercular margin notserrated; suborbital spine absent. Ventral margin of orbit not reaching a horizontal line through snout tip and upper
base of pectoral fin. Enlarged canines
anteriorly in upper and lower jaws.
First 2 dorsal-fin spines separated by amembrane. Upper lobe of caudal fin
slightly rounded. Pectoral fin not extending beyond a vertical through anal
fin origin. Three transverse scale rows
on preopercle. Color: body pinkishdorsally, silvery ventrally, with 6–8
pale yellow stripes below lateral line. A
red, ovoid spot below lateral line origin, bordered below by bright yellow.
Size: maximum length 21 cm SL. Distribution: the Andaman Sea and West

Nemipterus marginatus, KAUM–I. 16821, 8.7 cm SL
off Terengganu (KT), 5 Dec. 2008

Pacific. Remarks: usually occurs on produced into a short filament. Threesandy or muddy bottoms at depths of transverse scale rows on preopercle.
10 to 80 m.
(H. Motomura) Color: body pinkish dorsally, silvery
ventrally; with a broad yellow stripe,
divided above pectoral fin, from beNemipterus marginatus
low lateral line origin to upper part of(Valenciennes, 1830)
caudal peduncle; a second yellowishRed Filament Threadfin Breamorange stripe from above base of pecD X, 9; A III, 7; P1 16; P2 I, 5; GR 12. toral fin to lower part of caudal peBody moderately elongate, com- duncle; a reddish spot below and justpressed. Posterior margin of suborbit- behind lateral line origin; caudal fin
al and lower preopercular margin not including filament, reddish, its medianserrated; suborbital spine absent. Ven- rays yellowish. Size: maximum lengthtral margin of orbit above a horizontal 15 cm SL. Distribution: West Pacific.
line through snout tip and upper base Remarks: found on sand or mud botof pectoral fin. Pectoral and pelvic fins toms in depths of 12 to 70 m.
long; caudal fin forked, upper lobe
(M. Matsunuma)

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Nemipterus tambuloides(Bleeker, 1853)
Fivelined Threadfin BreamD X, 9; A III, 7; P1 16; P2 I, 5; GR
6–8. Body moderately elongate, compressed; body depth 3.2 to 3.6 in SL.
Posterior margin of suborbital andlower preopercular margin not serrated; suborbital spine absent. Ventral
margin of orbit tangent to or just above
a horizontal line through snout tip andupper base of pectoral fin. Pectoraland pelvic fins long; caudal fin deeply
forked, tip of upper lobe pointed.
Three transverse scale rows on preopercle. Color: body pinkish dorsally,
silvery ventrally, 5 well-defined yellow stripes along body. Dorsal fintranslucent pink, with yellow margin;
a narrow yellow stripe just above basefin. Anal fin translucent bluish whitewith pale yellow stripe near base offin. Caudal fin bright rosy, upper tip
yellow. Size: maximum length 23 cm
SL. Distribution: the Andaman Sea
and West Central Pacific. Remarks:
usually occurs on sand or mud bottoms in depths of 50 to 70 m.
(M. Matsunuma)

Nemipterus tambuloides, KAUM–I. 17132, 11.0 cm SL
off Terengganu (KT), 1 Jan. 2009

Nemipterus zysron, KAUM–I. 17134, 12.0 cm SL
off Terengganu (KT), 1 Jan. 2009

Nemipterus zysron(Bleeker, 1857)
Slender Threadfin BreamD X, 9; A III, 7; P1 15–17; P2 I, 5; GR
10. Body moderately elongate, compressed; body depth 3.8–4.6 in SL.
Posterior margin of suborbital andlower preopercular margin not serrated; suborbital spine absent. Ventral
margin of orbit just reaching or notreaching a horizontal line throughsnout tip and upper base of pectoral
fin. First two dorsal-fin spine separated
by a membrane; membranes between
dorsal-fin spines not strongly incised.
Upper lobe of caudal fin falcate or ribbon-like extension. Pectoral and pelvic
fins short, not reaching to a verticalthrough anal fin origin. Color: body
pinkish dorsally, silvery ventrally; belly white. Two yellow stripe on snout,
upper extending onto eye posteriorly.
Dorsal fin bluish translucent, with yellow margin, broad pale yellow stripe
on center of fin; anal fin bluish translucent with pale yellow stripe near baseof fin ; caudal fin pink, with reddish
posterior margin; upper margin andfilament yellow. Size: maximum

Pentapodus setosus, KAUM–I. 16819, 10.1 cm SL
off Terengganu (KT), 5 Dec. 2008

length 18 cm SL. Distribution: IndoWest Pacific. Remarks: found on sandbottoms near rocks in depths between
10 to 125 m.
(K. Shibukawa)Pentapodus setosus(Valenciennes, 1830)
Butterfly Whiptail
D X, 9; A III, 7; P1 15; P2 I, 5; LL
46–48; GR 16–17. Body moderately
elongate, fusiform; snout length greater than eye diameter. Scales on top ofhead reaching forward to or in front of
middle of eyes; six transverse scalerows on preopercle; lower limb of preopercle naked. Pelvic fins short, notreaching level of anus; caudal fin
forked, upper lobe produced into avery long trailing filament. Color:body whitish, pale brownish dorsally;a yellow stripe from behind eye, gradually arching on back and terminating
in a black spot on upper caudal peduncle; a narrow blue line running throughyellow stripe, this line convergent
with a blue line from origin of anal fin,both lines meeting at an acute angle
behind black spot on caudal peduncle;
2 bluish stripes across snout, first frommiddle of eye to tip of snout, second
from upper lip to lower margin of eye.
Dorsal fin pale blue, with yellow margin; caudal fin pinkish, filament pinkish brown. Size: maximum length17.5 cm SL. Distribution: West Central Pacific. Remarks: found in offshore waters.
(M. Matsunuma)

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Scolopsis margaritifera(Cuvier, 1830)
Pearly Monocle BreamD X, 9; A III, 7; P1 16–17; P2 I, 5; LL
36–39. Body relatively deep, compressed; body depth 2.4–3.8 in SL.
Posterior margin of suborbital with alarge backwardly pointing spine and aseries of smaller spines or serrations; asmall forward pointing spine below
eye absent. Preopercular margin serrated. Caudal fin forked, upper lobe
not produced into a filament. Scales on
dorsal head reaching forward to or in
front of level of anterior margin of eye;temporal region covered with scales.
Color: body olive dorsally, whitish
ventrally. Center of scales with pearlyor yellowish spots, forming longitudinal and transverse lines. In juveniles,
upper half of body white, lower yellow, with two dark stripes; a large dark
blotch on dorsal fin anterioly. Size:maximum length 19.5 cm SL. Distribution: the Andaman Sea and thewestern central Pacific. Remarks: occurs in shallow waters on sandy or
muddy bottoms close to reefs, todepths of 20 m.
(H. Motomura)

Scolopsis margaritifera, UMTF 1318 (KAUM–I. 16595), 4.6 cm SL
Bidong Island, 12 Oct. 2008

Scolopsis monogramma(Cuvier, 1830)
Monogrammed Monocle BreamD X, 9; A III, 7; P1 16–17; P2 I, 5; LL
36–39. Body relatively deep, compressed; body depth 2.4–3.8 in SL.
Suborbital margin with a large backwardly pointing spine; no antrorse
spine just below eye; lower limb of
preopercle scaled. Pelvic fin reaching
to anus when deppressed; caudal fin
forked or lunate, upper lobe a little
longer than lower lobe or produced
into a short filament. Color: head andbody grayish dorsally, silvery ventrally; dusky midlateral stripe on body;some sky blue streaks on head. Size:
commonly 18 cm SL. Distribution:
the Andaman Sea and West Pacific.
Remarks: found in coastal waters
with sandy or muddy bottom aroundcoral reefs in depths to 50 m.
(K. Shibukawa)

Scolopsis monogramma, KAUM–I. 16848, 18.9 cm SL
off Terengganu (KT), 5 Dec. 2008

Scolopsis monogramma, UMTF 1358 (KAUM–I. 16576), 17.0 cm SL
Bidong Island, 13 Oct. 2008

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Scolopsis taenioptera(Cuvier, 1830)
Lattice Monocle BreamD X, 9; A III, 7; P1 17–18; P2 I, 5; LL
45–48. Body relatively deep, compressed; body depth 2.7–3.1 in SL.
Suborbital with a large backwardly
pointing spine; no antrorse spine justbelow eye; exterminal edge of maxilla
smooth; lower limb of preopercular
naked. Pelvic fin reaching to anus
when deprresed; caudal fin forked or
lunate, upper lobe not produced into afilament. Color: head and body grayish dorsally, silvery ventrally; orangeor red spot at upper part of pectoral finbase. Size: commonly 15 cm, maximum 20 cm SL. Distribution: the Andaman Sea and West Pacific. Remarks: found in offshore waters withsandy or muddy bottom in depths to
50 m.
(K. Shibukawa)

Scolopsis taenioptera, KAUM–I. 16970, 19.8 cm SL
off Terengganu (KT), 11 Dec. 2008

Scolopsis vosmeri(Bloch, 1792)
Whitecheek Monocle BreamD X, 8–9; A III, 7; P1 18–19; P2 I, 5;
LL 41–44; GR 11–12. Body deep,
compressed; depth 2–2.6 in SL. Suborbital with a large backwardly-pointed spine; no antrorse spine just below
eye; external edge of maxilla smooth.
Pectoral fin short, not reaching to levelof anus; pelvic fin reaching to or beyond anus when depressed; caudal fin
forked. Scales on top of head extending forward to between level of snout
and anterior nostril; lower limb of pre-Scolopsis vosmeri, KAUM–I. 16853, 16.7 cm SL
off Terengganu (KT), 5 Dec. 2008

opercle scaled. Color: head and bodybrownish, paler ventrally; distinct,
broad white vertical bar on head. Size:
16 cm SL, commonly 15 cm SL. Distribution: Indo-West Pacific. Re-

marks: found in offshore or coastal
waters with sandy or muddy bottom
around coral reefs. Caught by traps
and trawls; marketed fresh.
(K. Shibukawa)

132

LETHRINIDAE
Emperors

By Koichi Shibukawa, Mizuki Matsunuma and Hiroyuki Motomura
Medium to large sized (up to 100cm), oblong and compressed fishes.
Body compressed, covered with ctenoid scales; lateral line continuous.
Cheek scales absent in Lethrinus,
whereas present in the other genera;
mouth terminal with relatively thick
lips; strong canines at front of jaws;
either conical or molariform teeth
on sides of jaws; no teeth on roof ofmouth; branchiostegals 6. A single
dorsal fin with X spines and 9–10
soft rays; anal fin with III spinesand 8–10 soft rays; pectoral fin with
13–15 soft rays; pelvic fin below
base of pectoral fin, with I spine and5 soft rays; caudal fin emarginated orforked. Scales ctenoid, moderate insize. Vertebrae 24. Color: head andbody silvery, gray, and light or reddish brown, frequently mottled, spot-

single dorsal fin with X, 9–10

strong canines atfront of jaws

mouth terminal
with thick lips
anal fin with III, 8–10

ted or striped with dusky, blue, yellow
or red.
Remarks: commonly found incoastal waters around coral and rocky
reefs, seagrass beds and mangroves.
Carnivorous, feeds on various organisms, e.g., fishes, mollusks, crustaceans, polychaetes, and sea urchins.
Esteemed as food fish, caught by han-dline, traps and trawls.
Similar families occurring in
the area: Haemulidae – chin with
distinct pores; scales on cheek.
Lutjanidae – teeth on roof of mouth;
scales on cheek. Sparidae – usuallymore spines in dorsal fin; scales oncheek.

Gymnocranius elongatus
Senta, 1973
Forktail Large-eye BreamD X, 10; A III, 10; P1 14; P2 I, 5; LLp
47–48. Body oblong, compressed; its
depth 2.2–2.4 in SL. Outer surface ofmaxilla smooth. Ventral margin of orbit intersected by line from tip of snout
to middle of caudal-fin fork. Sides ofjaws with canines and villiform teeth.
Cheek with 4–6 transverse scale rows;
inner surface of pectoral fin axil notcovered with scales. Caudal fin
strongly forked, with pointed tips; median rays distinctly shorter than eye
diameter. Color: body silvery, sometimes slightly brownish dorsally;
about 8 transverse brown bars onsides; first bar through eye and acrosscheek. Fins mainly clear to yellow-orange; a narrow brown bar often acrosspectoral-fin base; caudal fin margins
and tips often deep red. Size: maximum length about 35 cm. Distribution: eastern Indian Ocean and western Pacific Ocean. Remarks: occurs
in coastal and shelf waters at depths
between about 50 and 100 m. Caught
mostly with bottom trawls.
(M. Matsunuma)

Gymnocranius elongatus, KAUM–I. 17069, 13.9 cm SL
off Terengganu (KT), 27 Dec. 2008

Gymnocranius elongatus, KAUM–I. 17109, 13.5 cm SL
off Terengganu (KT), 31 Dec. 2008

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Gymnocranius griseus(Temminck & Schlegel, 1843)
Gray Large-eye BreamD X, 10; A III, 10; P1 14; P2 I, 5; LLp
46–48. Body oblong, compressed; its
depth comparatively deep, about 1.9–
2.2 in SL. Outer surface of maxilla
smooth. Ventral margin of orbit abovehorizontal line through snout tip andmiddle of caudal fin fork. Sides of jaws
with canines and villiform teeth. Cheek
with 4 transverse scale rows; inner surface of pectoral fin axil not covered
with scales. Caudal fin moderately
forked, with pointed tips; median rays
slightly greater than eye diameter. Color: body silvery; frequently with a diffuse to vivid pattern of 5 to 8 narrowdark bars on side; first bar through eye
and across cheek. Fins mainly clear toyellowish; a narrow brown bar often
across pectoral-fin base. Size: maximum length about 35 cm. Distribution: distributed in the eastern Indian
Ocean and the western Pacific fromsouthern Japan to western Australia.
Remarks: occurs in continental shelfand coastal inshore waters at depths between about 20 and 80 m. Caught mostly with bottom trawls. (H. Motomura)Gymnocranius griseus, KAUM–I. 17316, 16.5 cm SL
off Terengganu (KT), 21 Jan. 2009

Lethrinus genivittatus
Valenciennes, 1830
Longspine Emperor
Lethrinus genivittatus, KAUM–I. 16980, 12.4 cm SL
D X, 9; A III, 8; P1 13; P2 I, 5; LLp
off Terengganu (KT), 12 Dec. 2008
46–47. Body moderately elongate andslender; its depth 2.9–3.5 in SL. Headprofile near eye slightly convex. Outersurface of maxilla with a distinct knob.
Second dorsal-fin spine the longest,much longer than other dorsal-fin
spines. First or second anal-fin soft ray
longest. Cheek not covered with scales;
inner surface of pectoral fin axil covered with small scales; 4 ½ scale rows
between lateral line and base of middle
dorsal fin spines; 15 or 16 scale rows inlower series around caudal peduncle.
Color: body tan or brown on uppersides, lower sides white with 3 brownor tan stripes, sides often with scatteredirregular black oblique bars and asquare black blotch above pectoral fins
Lethrinus genivittatus, KAUM–I. 17240, 11.6 cm SL
and bordering below lateral line; head
off Terengganu (KT), 15 Jan. 2009
brown or tan sometimes with several
broad, somewhat indistinct vertical andoblique bands; fins pale, speckled with length about 25 cm. Distribution: east- marks: found in shallow coastal wasmall white blotches. Size: maximum ern Indian Ocean and West Pacific. Re- ters. Marketed fresh. (M. Matsunuma)

134

Lethrinus lentjan(Lacepède, 1802)
Pink Ear Emperor
D X, 9; A III, 8; P1 13; P2 I, 5; LLp
46–47. Body moderately deep; its
depth 2.5–2.8 in SL. Dorsal profile ofsnout around straight. No scales oncheek; inner surface of pectoral-finbase scaled or naked; 6 longitudinalscale rows (including small scales beneath dorsal-fin base) between lateralline and base of middle dorsal-fin
spine. Color: body grayish dorsally,
silvery ventrally; whitish spot at center of each scale on body; posterior
margin of opercle red; base of pectoral
fin sometimes with red marking; caudal fin mottled. Size: 52 cm. Distribution: Indo-West Pacific Remarks:
found in lagoons, coral reefs and seagrass beds; juveniles enter estuaries
and mangrove swamps. Carnivorous,
feeds on crustaceans, mollusks, polychaetes and fishes. Esteemed as foodfish.
(K.Shibukawa)

Lethrinus lentjan, KAUM–I. 16923, 16.1 cm SL
off Terengganu (KT), 10 Dec. 2008

Lethrinus ornatus
Valenciennes, 1830
Ornate Emperor
D X, 9; A III, 8; P1 13; P2 I, 5; LLp
46–47. Body moderately deep; its
Lethrinus ornatus, UMTF 1061 (KAUM–I. 16334), 7.4 cm SL
depth 2.3–2.6 in SL. Dorsal profile ofCendering, 16 Sept. 2008
snout around eye convex. No scales oncheek; inner surface of pectoral finbase scaled; 6 longitudinal scale rows
(including small scale beneath dorsal
fin base) between lateral line and baseof middle spine of dorsal fin. Color:body grayish dorsally, silvery ventrally, with 5–6 longitudinal yellow
stripes; posterior margin of preopercle
and opercle red. Size: 45 cm. Distribution: Indo-West Pacific. Remarks:
found in coastal waters, e.g., lagoons,seagrass beds, and areas adjacent to
reef. Carnivorous, feeds on crustaLethrinus variegatus, KAUM–I. 17048, 12.8 cm SL
ceans, mollusks, polychaetes and fishoff Terengganu (KT), 18 Dec. 2008
es. Utilized as food fish, caught bytraps, hand-lines, trawls, and seines.
(K. Shibukawa) to anterior nostril than orbit. Outer irregular dark spots; often 2 dark
surface of maxilla smooth. Fourth dor- bands below eye; a dark band acrosssal-fin spine usually longest. Cheek interorbital space; dorsal, anal, pectoLethrinus variegatus
naked; inner surface of pectoral fin ral, and pelvic fins light or translucent;
Valenciennes, 1830
axil not covered with scales; 4 ½ scale caudal fin light and dark striped. Size:
Slender Emperorrows between lateral line and base of 20 cm. Distribution: Indo-West PaD X, 9; A III, 8; P1 13; P2 I, 5; LLp middle dorsal fin spines; 13 or 14 cific. Remarks: inhabits sandy bot46–48. Body slender, its depth 3.2–3.9 scale rows in lower series around cau- toms near coral reefs.
in SL. Head profile near eye convex or dal peduncle. Color: body brown and(M. Matsunuma)
nearly straight; posterior nostril closer gray, lighter ventrally, with scattered

135

SPARIDAE

Seabreams or Porgies
By Mizuki Matsunuma
Medium to large sized (up to
100 cm) fishes, with body oblong,moderately deep and compressed.
Head large, often with a steep upper profile; mouth subhorizontal
and slightly protrusible, upper jaw
never reaching reaching a verticalline through middle of eye; hind tipof premaxilla overlapping maxilla;
jaw teeth well developed, differentiated into either conical, or flattened,
and often rounded; vomer and palatines toothless. Gill rakers variable,
7–20 on lower limb of first gill arch.
Dorsal fin single, with X–XIII spinesand 9–17 soft rays, the spiny and soft
portions not separated by a notch;
anal fin with III spines and 7–15 softrays; pectoral fins usually long andpointed; pelvic fin with I spine and5 soft rays, and an axillary scale atbase; caudal fin moderately deeply
emarginate or forked. Scales cycloid
or weakly ctenoid. Color: overallcolor highly variable, from pinkish
or reddish to yellowish or grayish,

single dorsal fin with X–XIII, 9–17

anal fin with III, 7–15

often with silvery or golden reflections, often with dark or colored spots,
stripes, or bars.
Remarks: inhabits tropical andtemperate coastal waters; occasionallyfound in estuaries. Most seabreams
are excellent food fish and are of notable commercial importance.
Similar families occurring in thearea: Haemulidae – edge of preopercle serrate; suborbital space scaled;; no molar teeth. Lutjanidae – edgeof preopercle usually serrated, andoften excavated to accommodate abony knob; no molar or incisor-liketeeth. Lobotidae – edge of preopercle
strongly toothed; no molar teeth;
dorsal, anal and caudal fins rounded.
Lethrinidae – posterior tip of maxilla
overlapping premaxilla; 8–11 soft
dorsal-fin rays (9–17 in Sparidae);incisor-like teeth never present.

Acanthopagrus pacificus
Iwatsuki, Kume & Yoshino, 2010
Pacific Seabream
D XI, 11; A III, 8; P1 15; LLp 42–46.
Body deep and moderately compressed; mouth slightly oblique; lips
thick; 6 canines anteriorly in upper
jaw, 6 in lower jaw; ventral edge ofanterior two infraorbitals straight injuveniles, but a moderately curved
concavity in adults. Second anal-fin
spine robust, clearly longer than 3rd;caudal fin slightly forked, with rounded lobes. Scale rows between 5th dorsal fin spine base and lateral line 3½;
front edge of predorsal fin scaly area
on head slightly convex, without small
scales. Color: head and body slivery
gray, whitish ventrally; pectoral finhyaline or somewhat dusky dorsally;other fins dusky. Size: 45 cm SL. Distribution: West Central Pacific, fromthe Ryukyu Islands south to northern
Australia. Remarks: found in shallow
coastal waters; juvenile entering lower
reaches of rivers.

Acanthopagrus pacificus, KAUM–I. 17141, 18.0 cm SL
off Terengganu (KT), 1 Jan. 2009

136

POLYNEMIDAE
Threadfins

By Hiroyuki Motomura
Body elongate to moderately
deep, compressed; size from about
10 to 200 cm. Snout obtusely conical, overhanging. Adipose eyelid
covering eye. Mouth ventral, nearhorizontal 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 onjaws, vomer, palatines and ectopterygoids (vomerine teeth absent in somespecies); canine, molariform or incisiform teeth absent. 7 branchiostegal rays, one ray present on epihyal.
Two well-separated dorsal fins; firstdorsal fin with 7 or 8 spines; second
dorsal fin with a spine and 11 to 18
soft rays. Anal fin with 2 or 3 spinesand 10 to 18 soft rays. Pectoral finsdivided into an upper part with 12 to
19 rays joined by membrane and alower part with 3 to 16 separate softrays (pectoral filaments). Pelvic finswith a spine and 5 soft rays. Caudal
fin deeply forked. Scales weakly

2 dorsal fins well separated
adipose eyelid

lower pectoral-fin rays
forming pectoral filaments

ctenoid, extending onto head; scales
covering most of dorsal, pectoral, analand caudal fins; lateral line extending onto posterior margin of caudal
fin; trisegmental pterygiophores absent. Vertebrae 10 precaudal and 14
or 15 caudal; supraneural bones 0 to
3. Color: silvery, golden or brown. A
large black spot at anterior lateral line
or several longitudinal dark stripes insome species.

Remarks: taken over sandy or
muddy bottoms in tropical and subtropical coastal, brackish and frshwaters. Feeds mainly on crustaceans
and small fishes. Some species hermaphrodites.
Similar families occurring in
the area: pectoral fin and filamentcharacters (see above description)
distinguish Polynemidae from allother families.

Eleutheronema tetradactylum(Shaw, 1804)
Fourfinger Threadfin
D VIII + I, 13–15; A III, 14–16; P1
15–19 + 4 filaments; P2 I, 5; LLp 71–
80; GR 6–18. Body elongate, moderately compressed. Posterior margin of
maxilla extending well beyond a vertical through posterior margin of adipose eyelid. Upper jaw length 5.9–7.1
in standard length. Anterior parts oflower jaw with small teeth extending
onto lateral surface, adjacent portionof lip absent; tooth plate extension
onto lateral surface of lower jaw 11.1–
14.3 in standard length; vomer withdeciduous tooth plates on both sides,
except in juveniles. Longest pectoral
filament not reaching origin of analfin. Swimbladder absent. Color: upper sides of head and trunk with slightdarkish silver tinge, becoming lighter
on lower sides. Anterior margins of

Eleutheronema tetradactylum, KAUM–I. 17039, 23.8 cm SL
off Terengganu (KT), 17 Dec. 2008

first and second dorsal fins blackish,remaining parts translucent and slightly blackish, respectively; pectoral fin
vivid yellow (dusky yellow in large
specimens over ca. 350 mm standard
length). Size: maximum total length
about 200 cm. Distribution: distributed in the Indo-West Pacific, from thePersian Gulf east to Papua New Guin-ea and northern Australia. Remarks:
generally occurs on continental shelves
on muddy and sandy substrata, andfrequently enters brackish waters, especially as juveniles. Sex changing
from male to female with growth.

137

Leptomelanosoma indicum(Shaw, 1804)
Indian Threadfin
D VIII + I, 12–13; A III, 10–11; P1
12–14 + 5 filaments; P2 I, 5; LLp 69–
72; GR 18–21. Body elongate, moderately compressed. Posterior margin of
maxilla extending well beyond a vertical through posterior margin of adipose eyelid. Anterior one-third oflower jaw with small teeth extending
onto lateral surface, adjacent portionof lip poorly developed; vomer withteeth. Longest pectoral filament notreaching origin of anal fin. Swimbladder with many appendages inserted
into lateral walls of abdominal cavity.
Color: head and upper sides of trunk
tinged slightly blackish brown, becoming deep black on lower sides;
snout and abdominal regions blackish.
Origin of pectoral filaments dusky
yellowish, grading to blackish posteriorly; pectoral fin membrane deep
black. Size: maximum total length 1.4
m. Distribution: distributed in the Indo-West Pacific, from Pakistan to
Papua New Guinea. Remarks: mainly
occurs near estuaries, sometimes entering rivers, and is occasionally takenfrom depths to 100 m.

Polydactylus sextarius(Bloch & Schneider, 1801)
Blackspot Threadfin
D VIII + I, 12–13; A III, 11–13; P1
13–15 + 6 filaments; P2 I, 5; LLp 45–
51; GR 25–30. Body moderately deep,
compressed. Posterior margin of maxilla reaching to or slightly short of avertical through posterior margin of
adipose eyelid. Lip on lower jaw well-

Leptomelanosoma indicum, KAUM–I. 17104, 20.9 cm SL
off Terengganu (KT), 28 Dec. 2008

Polydactylus sextarius, KAUM–I. 16905, 12.3 cm SL
off Terengganu (KT), 6 Dec. 2008

developed, dentary teeth restricted todorsal surface; teeth villiform in broad
bands on palatines and ectopterygoids;
vomerine tooth plate covered withskin and teeth absent. All pectoral fin
rays, except uppermost 1 or 2, unbranched. Longest pectoral filament
not reaching to posterior tip of pectoral fin. Swimbladder atrophied, like afine string. Color: head and body silver. A large black spot anteriorly onlateral line. Posterior margins of firstand second dorsal fins and caudal fin
slightly blackish, remaining partstranslucent white. Pectoral fin membrane dark yellow; pectoral filaments
faintly white. Size: maximum standard length 17 cm. Distribution: distributed in the eastern Indian andwestern Pacific Oceans, from India
east to Japan and Indonesia. Not occur
in Australian waters. Remarks: occurs on sandy and muddy bottoms, being taken from depths of 16 to 73 m.

138

SCIAENIDAE
Croakers

By Aziz Arshad and Hiroyuki Motomura
Small to large fishes, reaching
200 cm standard length. Body moderately elongate, compressed. Body,
except snout tip, completely covered
with scales. Enlarged teeth always
form outer series in upper jaw, inner series in lower jaw; vomer andpalatines without teeth. Dorsal fin
continuous, with deep notch between
anterior and posterior portions; with
VIII to X spines anteriorly and I
spines and 21 to 44 soft rays posteriorly; base of posterior portion much
longer than anal fin base. Anal fin
with II spines and 6 to 12 soft rays.
Caudal fin emarginate to pointed,
never deeply forked. Pelvic fins with
I spine and 5 soft rays. Lateral line
scales extending to posterior marginof caudal fin. Swimbladder well developed with thick wall, with horn-

dorsal fin continuous with deep notch
lateral line scales
extending onto caudal fin

anal fin with 2 spines

like, tube-like, or arborescent appendages. Color: variable from silvery to
black; with spots and dark bands insome species.
Remarks: occurs in coastal waters, estuaries and rivers. No records
from oceanic island groups distantly
separated from a continental shelf.
Usually found over muddy or sandy

bottoms.
Similar families occurring in
the area: the following combinationof characters distinguish Sciaenidae
from all other families: anal fin with
II spines and lateral line scales extending to posterior margin of caudal
fin.

Chrysochir aureus(Richardson, 1846)
Reeve's Croaker
D XI, 25–28; A II, 6–7; P2 I, 5; LGR
8–9. Body slender. Mouth nearly horizontal; posterior margin of maxilla
extending beyond a vertical throughposterior margin of orbit. No barbel onchin; upper rostral pores 3, marginal
rostral pores 5; mental pores in 3 pairs,
the first slit-like on either side of symphysis. Swimbladder carrot-shaped,with 27–30 pairs of fan-like appendages along its sides, none entering
head. Color: body metallic blue dorsally, shading to silvery laterally;
oblique streaks along scale rows on
back; pectoral fins yellow, other fins

Chrysochir aureus, KAUM–I. 17063, 19.7 cm SL
off Terengganu (KT), 26 Dec. 2008

gray suffused with orange. Size: max- Bengal east to southern China and Inimum standard length about 30 cm, donesia. Remarks: inhabits shallow
commonly to 25 cm. Distribution: coastal waters.
(A. Arshad)Indo-West Pacific, from the Bay of139

Dendrophysa russelli(Cuvier, 1829)
Goatee Croaker
D XI, 25–28; A II, 7; P2 I, 5; LGR
8–10. Body relatively deep. Mouth inferior; posterior margin of maxilla notextending beyond a vertical throughposterior margin of orbit. A pointed
barbel on chin. Swimbladder carrotshaped, with about 14–17 pairs of fanlike appendages along its sides; firstpair entering head beyond transverse
septum. Color: body gray dorsally,
white ventrally. A dark brown broadband on nape. Opercle with a deep
blue blotch. Spinous portion of dorsal
fin faintly edged with black. Size:maximum standard length 25 cm. Distribution: distributed in the eastern
Indian Ocean and the western central
Pacific Ocean, from the Bay of Bengal
east to southern China, the Philippinesand eastern Indonesia. Remarks: occurs in coastal waters and estuaries.
(H. Motomura)

Dendrophysa russelli, KAUM–I. 16997, 11.9 cm SL
off Terengganu (KT), 14 Dec. 2008

Johnius amblycephalus(Bleeker, 1855)
Bearded Croaker
D XI, 23–26; A II, 7; P2 I, 5; LGR
6–9. Body moderately elongate. A
stiff, blunt barbel on chin. Teeth differentiated into large and small in upper jaw, villiform teeth only in lower
jaw. Second to fifth dorsal fin spines
prolonged. Scales on head and body
cycloid; small scales covering softrayed portions of dorsal and anal fins.
Swimbladder hammershaped, with 14
or 15 pairs of arborescent appendages
along its sides, first pair entering headbeyond transverse septum and sending
a palmate branch to the front of pectoral arch. Color: body black or darkbrown dorsally, whitish or cream yellow ventrally; prolonged spinous portion of dorsal fin black distally. Size:maximum standard length about 25
cm. Distribution: Indo-West Pacific,
from Pakistan east to southern China
and northeastern Australia. Remarks:
occurs in shallow coastal waters andestuaries.
(H. Motomura)Johnius amblycephalus, KAUM–I. 17255, 8.6 cm SL
off Terengganu (KT), 17 Jan. 2009

Johnius macrorhynus, KAUM–I. 16873, 15.6 cm SL
off Terengganu (KT), 5 Dec. 2008

deep. Mouth inferior; posterior marginof maxilla extending beyond below
middle of eye, but not reaching below
posterior margin of orbit. No barbel onchin; upper rostral pores 5, marginal
rostral pores 5; mental pores in 3 pairs,
Johnius macrorhynus
the first open close behind symphysis(Lal Mohan, 1976)
in a common pit. Swimbladder hamBigsnout Croaker
mer-shaped, with 13 or 14 pairs of arD XI, 27–30; A II, 7; P2 I, 5; LGR borescent appendages along its sides,
5–9. Body elongate and relatively the first pair entering head beyond

transverse septum and sending a palmate branch to the front of pectoralarch. Color: head and body silver,
lower part of body with a golden tinge;
pectoral, pelvic, and anal fins yellow;
a faint steel blue blotch on gill cover.
Size: maximum length about 30 cm.
Distribution: the South China Sea
west to India and Pakistan. Remarks:
inhabits shallow coastal waters.
(A. Arshad)140

Larimichthys crocea(Richardson, 1846)
Large Yellow Croaker
D X, 31–33; A II, 8; P2 I, 5; LGR
18–20. Body elongate. Mouth terminal; posterior margin of maxilla extending beyond a vertical through posterior margin of orbit. No barbel on
chin. Small cycloid scales coveringsoft-rayed portions of dorsal and anal
fins. Swimbladder carrot-shaped, with
31–33 pairs of arborescent appendages along its sides, each appendage
with a well-developed dorsal and ventral limb, the first pair entering headbeyond transverse septum. Color:body slivery, darker dorsally, yellowish laterally; fins yellowish. Size:maximum length about 60 cm. Distribution: the South China Sea and East
China Sea. Remarks: Generic assignment follows Nakabo (2002). Inhabits
shallow coastal waters. (A. Arshad)

Larimichthys crocea, KAUM–I. 16977, 20.0 cm SL
off Terengganu (KT), 12 Dec. 2008

Nibea semifasciata
Chu, Lo & Wu, 1963
Sharpnose Croaker
D XI, 26–29; A II, 7; P2 I, 5; LLp
50–52; LGR 9–10. Body deep; napehighly arched. Snout acutely pointed,
projecting in front of upper jaw.
Mouth large, slightly inferior; posterior margin of maxilla slightly extending beyond a vertical through posterior margin of orbit. No barbel on chin;upper rostral pores 3, marginal rostral
pores 5; mental pores in 3 pairs, thefirst close together, united by a crescent-shaped groove just behind symphysis. Scales on head cycloid, thoseon body ctenoid; small scales coveringsoft-rayed portions of dorsal and anal
fins. Swimbladder carrot-shaped, with
17–20 pairs of aborescent appendages
along its sides, the first pair entering
head beyond transverse septum. Color: body brownish gray, dark obliquelines on dorsally; margin of spinous
portion of dorsal fin darker; pectoraland pelvic fins yellowish. Size: maximum standard length 24 cm. Distribution: the East China Sea and the
Gulf of Thailand. Remarks: inhabits
coastal waters.
(A. Arshad)Nibea semifasciata, KAUM–I. 16876, 15.9 cm SL
off Terengganu (KT), 5 Dec. 2008

48–51; LGR 7–12. Body relatively
deep; nape highly arched. Snout not
swollen or projecting. Posterior margin of maxilla reaching a verticalthrough posterior margin of orbit; upper jaw length 2.3–2.6 in head length,
lower jaw length 1.8–2.3 in headlength. No barbel on chin. Scales on
Nibea soldado
head cycloid, those on body ctenoid;(Lacepède, 1802)
small scales covering soft-rayed porSoldier Croakertions of dorsal and anal fins. SwimD X–XI, 27–33; A II, 7; P2 I, 5; LLp bladder carrot-shaped, with 18 to 22

Nibea soldado, KAUM–I. 16998, 17.2 cm SL
off Terengganu (KT), 14 Dec. 2008

pairs of aborescent appendages along
its sides, first pair entering head beyond transverse septum. Color: bodysilvery with faint series of oblique
stripes along scale rows. Soft-rayed
portion of dorsal fin dark distally. Pectoral and pelvic fins yellowish. Size:maximum standard length 60 cm. Distribution: Indo-West Pacific, from
India east to Queensland, Australia.
Remarks: occurs in shallow coastal
waters.
(H. Motomura)

141

Otolithes ruber(Bloch & Schneider, 1801)
Tigertooth Croaker
D X–XI, 26–30; A II, 7; P2 I, 5; LGR
8–11. Body slender, cylindrical. Snoutnot swollen or projecting, its dorsal
profile rising evenly to dorsal fin origin. Posterior margin of maxilla reaching a vertical through posterior marginof orbit. No barbels on chin. Teeth differentiated into large and small in both
Otolithes ruber, KAUM–I. 16874, 17.7 cm SL
jaws, with 1 or 2 pairs of strong caoff Terengganu (KT), 5 Dec. 2008
nines at front of one or both jaws.
Scales cycloid, but a few ctenoid on
lower part of caudal peduncle. Swimbladder carrot-shaped with 32–36
pairs of fan-like appendages alongsides, each appendage lodged beside
bladder and none widely lapping dorsal surface of bladder wall, first pair
not entering head. Color: body brownish dorsally, silvery with a golden
sheen on middle and ventrally, oftenwith oblique dark streaks dorsally.
Pectoral, pelvic, and anal fins reddish
or light brown. Size: maximum standard length 70 cm. Distribution: widely distributed in the Indo-West Pacific,
Pennahia anea, KAUM–I. 16875, 13.3 cm SL
off Terengganu (KT), 5 Dec. 2008
from South Africa east to southern
China and northeastern Australia. Remarks: occurs in shallow coastal waters to depths of 40 m. Feeds mainly on orbit. No barbels on chin. Teeth well reflection to whitish abdomen. Gill
prawns and fishes.
(H. Motomura) differentiated into large and small in cover with a diffused dark blotch. Upboth jaws, but none canine-like. Gill per two-thirds of spinous portion ofrakers slender, as long as gill filaments dorsal fin dusky. Size: maximum stanPennahia anea
at angle of arch. Caudal fin truncate. dard length 30 cm. Distribution:(Bloch, 1793)
Scales cycloid on head and flanks, widely distributed in the Indo-West
Donkey Croakerelsewhere ctenoid. Swimbladder car- Pacific, from the Persian Gulf east to
D X–XI, 22–24; A II, 7; P2 I, 5; LGR rot-shaped, with 17–22 pairs of fan- western Indonesia. Remarks: occurs
9–12. Body moderately deep. Snout like appendages along its sides, first in shallow coastal waters to depths ofpointed, but not swollen or projecting. not entering head. Color: body gray- 60 m.
(H. Motomura)
Posterior margin of maxilla reaching a ish light brown dorsally, becoming
vertical through posterior margin of paler ventrally, with increasing silvery

142

MULLIDAE
Goatfishes

By Siti Tafzilmeriam S. A. K., Hisashi Imamura and Seishi Kimura
Body moderately elongate andsomewhat compressed (size to 50cm). Mouth low on head; lower jaw
inferior; two long unbranched barbels on chin; a single flat spine posteriorly on opercle; margin of preopercle smooth. Two well-separated
dorsal fins, the first with VII–VIII
slender spines; the second fin with 9
soft rays (first unbranched); anal fin
with I spine and 6–7 soft rays; caudal fin deeply forked; pectoral finswith 13–18 soft rays; pelvic fins with
I spine and 5 soft rays. Scales finely
ctenoid; lateral line complete, following the contour of back, with 27–
38 pored scales. Color: ground color
often whitish to light red; most species with distinctive black, brown,
red, or yellow markings; median finsoften with stripes or oblique bands.
Remarks: inhabit shallow seas,on open sand or mud bottoms. They
are carnivorous, feeding on a wide

2 separate dorsal fins

2 barbels on chinvariety of small animals, especiallycrustaceans and worms. The barbels,
with chemosensory receptors, are actively moved over or into the sedimentto find food organisms.
Similar families occurring in thearea: Polymixiidae is the only otherfamily with a pair of long barbels onchin; the species occur in deep water,

anal fin with a single spine

about 200–400 m. They are distinguished from the Mullidae by thefollowing combination of characters:a single, unnotched dorsal fin with
IV– VI spines; anal fin with III–IV
spines; barbels inserted well behind
tip of lower jaw.

Mulloidichthys flavolineatus(Lacepède, 1801)
Yellowstripe Goatfish
D VIII + 9; A I, 7; P1 15–19; P2 I, 5;
LLp 33–36; GR 8–9 + 18–22. Body
elongate and moderately compressed.
Chin with 2 slender barbels, usually
not reaching a vertical through posterior margin of preopercle when depressed. No teeth on vomer and palatine; 2 rows of small villiform teeth on
jaws. Spinous and soft-rayed portionsof dorsal fin separated by 5 rows ofscales. Caudal fin forked. Color: body
Mulloidichthys flavolineatus, KAUM–I. 16855, 13.4 cm SL
off Terengganu (KT), 5 Dec. 2008
silvery white with a yellow mid lateral
stripe, usually containing a blackish
spot below first dorsal fin; pectoral,
anal and pelvic fins whitish to paleyellowish; barbels yellow; peritoneum in lagoons and bays; feeds on small
blackish. Size: maximum total length invertebrates and small fishes. Mar40 cm, commonly to 28 cm. Distribu- keted fresh or dried. (S. Tafzilmeriam)tion: Indo-Pacific. Remarks: found143

Parupeneus heptacanthus(Lacepède, 1802)
Cinnabar Goatfish
D VIII + 9; A I, 6; P1 15–17; LLp
27–28; GR 6–7 + 20–23. Body slightly elongate and moderately compressed. Chin with 2 moderately longbarbels, extending beyond a verticalthrough posterior margin of preopercle when depressed. Anterodorsal profile of head convex. No teeth on vomer and palatines. Spinous andsoft-rayed portions of dorsal fin separated by 3 rows of scales. Caudal fin
forked. Color: body brownish yellow
to light red, silvery white ventrally; asmall reddish brown spot beneath thelateral line under the rear of the firstdorsal fin; faint iridescent blue lines extending dorsoposteriorly and ventroanteriorly from eye. Size: maximum total
length about 37 cm. Distribution: Indo-Pacific. Remarks: occurs overmuddy, sandy, rubble, or seagrass bottoms of lagoon and seaward reefs.
(S.Tafzilmeriam)

Parupeneus heptacanthus, KAUM–I. 16856, 14.5 cm SL
off Terengganu (KT), 5 Dec. 2008

Upeneus guttatus(Day, 1868)

Upeneus guttatus, KAUM–I. 16964, 11.0 cm SL
off Terengganu (KT), 11 Dec. 2008

Two-tone Goatfish
D VII, 9; A I, 7; P1 13–14 (usually13); GR 5–8 + 15–18 = 21–25; V 10 +14. Body elongate, cylindrical. Chinwith 2 slender barbels, usually justreaching to posterior margin of preopercle. Mouth small; small conicalblunt teeth in a narrow band on jaws,
palatines, ectopterigoids, and in 2
small patches on vomer. Scales present on side of snout; 6 scales between
dorsal fins; 10–11 scales along dorsaledge of caudal peduncle. Color: bodysilvery pink with bright vermilion
spots or blotches; barbels yellow;
bright red bars on both dorsal fins;
faint reddish bars on pelvic fins; upper
lobe of caudal fin with 5 red obliquebars, the lower lobe with a red wideband from center of caudal fin base to
inner hind margin and 5 red obliquebars. Size: 11 cm SL. Distribution:
Indo-West Pacific. Remarks: similar
to Upeneus japonicus Houttuyn, 1782,
but distinguished from it by having
red oblique bars on lower lobe of caudal fin.
(S. Kimura)

Upeneus sulphureus, KAUM–I. 17097, 13.0 cm SL
off Terengganu (KT), 29 Dec. 2008

Upeneus sulphureus
Cuvier, 1829
Sulpher Goatfish
D VIII + 9; A I, 7–8; P1 14–17; LL
33–36; GR 8–9 + 19–21. Body moderately elongate. Chin with 2 short barbels, 1.4–1.9 in head length. Lacrimal
region lacking preorbital scales. Bothjaws, vomer and palatines with teeth.
Second dorsal and anal fins withscaled area basally. Color: body sil-very greenish or pink dorsally, shading to silver on side and ventrally, with
2 narrow, brassy yellow stripes on sideof body; 1st dorsal fin broadly tippedwith black; caudal fin lacking darkcross bands. Size: reaching to ca. 20
cm. Distribution: Indo-West Pacific,
including east coast of Africa, IndoMalayan region, Japan, Australia andFiji. Remarks: generally found on
mud substrata at depths of 20–60 m.
(H. Imamura)

144

Upeneus sundaicus(Bleeker, 1855)
Ochreband Goatfish
D VIII + 8; A I, 6; P1 14; LLp 31–32;
GR 4–6 + 19–22. Body moderately
elongate and slender, its depth 3.6–4.0
in standard length. Chin with 2 barbels, reaching to a vertical throughposterior margin of preopercle. Mouth
small; small conical blunt teeth in anarrow band on jaws, palatines, and in
2 small patches on vomer. No scales
on side of snout; 4 ½ scales between
dorsal fins; 10–11 scales along dorsaledge of caudal peduncle. Caudal fin
forked. Color: body light reddishbrown dorsally, silvery white ventrally; a yellow stripe on body side fromeye to caudal peduncle; caudal fin
brownish-yellow with dusky lower
lobe; pectoral fin pale yellow; pelvic
and anal fins yellowish; barbels yellow; peritoneum pale. Size: commonly to 13 cm. Distribution: Indo-West
Pacific. Remarks: found in mud or
salty sand substrata at depth of 3–20
m.
(S. Tafzilmeriam)

Upeneus tragula
Richardson, 1846
Freckled Goatfish
D VIII + 9; A I, 7; P1 13–14; LLp
28–30; GR 5–7 + 15–18. Body elongate; its depth 3.9–4.3 in standard
length. Chin with 2 slender barbels,
usually not reaching a vertical throughposterior margin of preopercle when
depressed. Mouth small with villiform
teeth in a narrow band in jaws, on palatines, and in 2 small patches on vomer. Second dorsal and anal fins withscaled area basally. Caudal fin forked.
Color: highly variable in color, from
red, to irregular dots and blotches onbody; a dark reddish brown to blackish stripe from front of snout to base of
caudal fin; head and body above stripe
brownish to greenish gray, fleckedwith small dark reddish brown or
blackish spots; upper lobe of caudal
fin with 4–6 blackish cross bands, andlower lobe with 5–7; outer third to halfof first dorsal fin with a large dark red

Upeneus sundaicus, KAUM–I. 17098, 12.2 cm SL
off Terengganu (KT), 29 Dec. 2008

Upeneus tragula, KAUM–I. 17150, 18.0 cm SL
off Terengganu (KR), 4 Jan. 2009

Upeneus tragula, UMTF 1062 (KAUM–I. 16513), 7.8 cm SL
Cendering, 6 Oct. 2008

to black blotch containing 2 to several West Pacific Remarks: found oversmall yellow spots; second dorsal fin sand and mud bottoms near coralwith 3 dark reddish or blackish bands; reefs, to the depth of 40 m.
barbels usually yellow. Size: maxi(S. Tafzilmeriam)
mum total length about 33 cm, commonly to 25 cm. Distribution: Indo-

145

PEMPHERIDAE
Sweepers

By Hisashi Imamura and Mizuki Matsunuma
Small to moderate sized (to 30cm) marine fishes. Body compressed,very deep (Pempheris) or slender
single dorsal fin with IV–VII, 7–12
(Parapriacanthus). Eye very large.
Snout short. Maxilla broad, reaching to below pupil. Teeth small, in eye very large
band in jaws, on palatine, and in
V-shaped patch on vomer. Gill rakers long, usually 25–31. Dorsal finsingle, with IV–VII graduated spinesand 7–12 soft rays. Anal fin with
III (rarely II) spines and 17–45 soft
rays. Pelvic fin with I spine and 5
soft rays. Lateral line scales usually
anal fin with III (rarely II), 17–45
40–82. Scales vary from ctenoid andadherent to cycloid and deciduous.
Color: variable (e.g., silvery, pinkish, yellowish and brownish).
ledges, dispersing at night to feed in- area. Berycidae – pelvic fin with
Remarks. usually seen by day dependently on zooplankton.
7–13 soft rays.
in aggregations in caves or beneath
Similar family occurring in thePempheris oualensis
Cuvier, 1831
Copper Sweeper
D VI, 9–10 (usually 9); A III, 36–45;
P1 16–18 (usually 17); LL 54–79; GR
7–10 + 19–26. Body very deep at origin of dorsal fin, strongly tapering tonarrow caudal peduncle; body depth
about 1.9–2.4 in standard length; prepelvic area narrow and keeled. Base ofanal fin very long; caudal fin slightly
forked. Body covered by ctenoid
scales except for chest with cycloid;
5–6 scale rows between lateral line
and dorsal fin base. Color: bodybronze, darker dorsally; dorsal fin
Pempheris oualensis, KAUM–I. 17178, 11.6 cm SL
off Terengganu (KR), 5 Jan. 2009
with a dark leading edge that expands
distally to produce dark tips on thefirst 2 or 3 soft rays; base and axil of cific, from northern Australia to thepectoral fin dark to black. Size: maxi- Philippines, east to Micronesia. Remum length 18 cm. Distribution: marks: found in coastal rocky reefs.
(M. Matsunuma)
eastern Indian Ocean and western Pa-

146

MONODACTYLIDAE
Moonyfishes (Fingerfishes)
By Mizuki Matsunuma
Small compressed fishes (up to
25 cm). Body oval, deep, strongly
compressed. Eye moderately large,
its diameter longer than snout length.
Mouth small and oblique. Jaws withbands of small conical teeth; vomer,
palatines, and tongue with granularteeth. Dorsal fin single, with V–VIII
slender spines and 26–31 soft rays;
anterior soft dorsal-fin rays elongated. Anal fin with III spines and
26–31 soft rays; anterior soft anal
fin rays elongated. Pelvic fin shortor rudimental. Caudal fin truncate toforked. Body, head, and unpaired finscovered by small, deciduous scales.
Color: head and body silvery, mostly with 1 or more black vertical bar
through eye, in front of pectoral-finbase (usually continuous with black
margin along anterior edge of analfin) and on body; tip of dorsal andanal fins usually black.
Remarks: occurs in shoals in estuaries and over shallow coral reefs,

single dorsal fin with V–VIII, 26–31

gill membranes
united mediallypelvic fins shortor rudimental

occasionally entering freshwater. Minor commercial importance; but someare caught for the aquarium fish trade.
Similar families occurring in
the area: Drepaneidae and Ephippidae – pelvic fins well developed;
gill membranes attached to isthmus.
Chaetodonti-dae and Scatophagidae– pelvic fins well-developed; dorsal

anal fin with III, 26–31

fin with X–XVII spines; spinous portion of dorsal fin well separated fromsoft-rayed portion. Carangidae – caudal fin forked; anal fin with II spines.
Menidae – ventral profile conspicuously more convex than dorsal profile; first 2 pelvic fin rays elongated.

Monodactylus argenteus(Linnaeus, 1758)
Silver Moony
D VII–VIII, 26–31; A III, 26–31.
Body oval, deep and strongly compressed; its depth 1.2–1.6 in SL; distance between tips of dorsal and anal
fins 0.8–0.9 in SL. Eye moderately
large; mouth small and oblique. Anterior soft fin rays of both dorsal andanal fins elongated; posterior edge of
dorsal and anal fins distinctly concave; caudal fin slightly emarginate;pelvic fin rudimentary or absent in
adults. Color: adults bright silver, tipof dorsal and anal fins dusky; juveniles more colorful with yellow overmost of the dorsal fin; tip of dorsal andanal fins black; anterior edge of anal
fin with broad black margin; two vertical black bars over the head, one
through the eye and the other in frontof pectoral-fin base. Size: maximum
length about 25 cm. Distribution: Indo-West Pacific. Remarks: commonly found in mangrove estuaries, sometimes found in silty coastal reefs.

Monodactylus argenteus, UMTF 1525 (KAUM–I. 16725), 8.4 cm SL
Setiu, 28 Oct. 2008

147

TOXOTIDAE
Archerfishes

By Mazlan Abdul Ghaffar
Small to medium sized fishes (toabout 50 cm). Body oval or rhomboidal-shaped and moderately compressed. Eye large, diameter nearly
equal to snout length. Mouth moderately large, protractile, with lower
jaw protruding; angle of jaw oblique,maxilla slender, scaly and without asupplemental bone. Fine villiform
teeth on jaws, vomer, and palatines;
a deep longitudinal groove on roofof mouth, which is converted to a
tube when tongue is pressed against
it. First gill arch with 2–8 gill rakers
on lower limb. Dorsal fin single with
IV–VI spines and 11–14 soft rays;
anal fin with III spines and 15–18
soft rays; pelvic fins with I spine and5 soft rays; pectoral fins with 11–15
rays; caudal fin truncate to slightly
emarginate. Scales moderate to relatively large and ctenoid, coveredhead and median fins. Color: most
species silvery white with a patternsingle dorsal fin with IV–V, 15–18

a deep longitudinalgroove on roof of mouth

anal fin with III, 15–18

of dark bars, large spots, or irregular
stripes on sides.
Remarks: inhabit mangrove
shores, estuaries, and fresh waters,always in shallow depths. Exhibit impressive hunting techniques in which
they use jets of water to knock aerial
insects into the water where they canbe eaten.
Similar families occurring in
the area: Pempheridae – snout blunt,
dorsal-fin base short, positioned
above pectoral fins; anal fin long,usually with more than 30 soft rays;
occur in marine rather than estuarine
and fresh-water habitats.

Toxotes jaculatrix(Pallas, 1767)
Banded Archerfish
D IV–V, 11–13; A III, 15–17; P1 13;
LGR 5–7. Body shape rhomboid,
moderately compressed. Eye large.
Mouth moderately large, protractile,
with lower jaw protruding; longitudinal axis from tip of snout to caudal fin
passing through center of eye. Dorsalfin spines usually IV, third spine longest. Scales in lateral line usually 26–
30; horizontal scale rows above lateralline 3 or 4, below lateral line 8 or 9.
Color: body silvery white, brownish
green dorsally, with 4 or 5 black
blotches or bars along the sides ofbody; caudal fin pale yellowish; anal
fin pale basally, outer half of the fin
blackish. Size: maximum total length
about 30 cm, commonly to about 20
cm. Distribution: eastern Indian Ocean
and western Pacific. Remarks: inhab-

Toxotes jaculatrix, UMTF 1008 (KAUM–I. 16787), 7.5 cm SL
Merang, 2 Nov. 2008

its mangrove shores and brackish estuaries, always in shallow depths. Feedson terrestrial insects. Marketed mostly
fresh.

148

KYPHOSIDAE

Sea Chubs (Rudder Fishes)
By Mizuki Matsunuma
Medium-sized fishes (to 75 cm);
body oblong or elliptical, moderately
compressed. Head small. Snout short.
Posteroventral corner of preopercle
serrate. Opercle with 2 small, weak
retrorse spines. Eye moderately small,its diameter shorter than snout length.
Mouth small and terminal, single rowof lanceolate incisor-like teeth on both
jaws; minute canine-like teeth in 2 or
3 rows medial to incisor-like teeth;
palatines, vomer, and tongue witha band of villiform teeth. Preorbital
narrow, covering little of maxilla;
maxilla barely reaching eye. Dorsal
fin continuous, beginning above origin of pelvic fins, with XI spines and
12 to 15 soft rays; anal fin beginningslightly behind middle of body, with
III spines and 11 to 14 soft rays; pectoral fins bluntly pointed posteriorly,
with 16 to 20 rays, slightly longer thanpelvic fins; pelvic fins beginning alittle behind pectoral-fin base; caudal
fin more or less forked, with pointed

single dorsal fin with XI, 12–15

single row of lanceolate
incisor-like teeth on both jaws

anal fin with III, 11–14

lobes. Scales ctenoid and not deciduous, extending onto most of soft portions of dorsal and anal fins and proximal part of caudal fin. Color: dull olive
or silvery black, lighter below; distinct
longitudinal lines on sides.
Remarks: occurs on rocky andcoral reefs in tropical and temperate waters. Herbivorous, feeding pri-marily on benthic algae. Schooling,sometimes in a group of mixed species of other kyphosids.
Similar families occurring in
the area: Sparidae – molar teeth
often present; pectoral fins long. Lethrinidae – head much larger; lateral
teeth conical or molar-like; pectoral
fins long.

Kyphosus cinerascens(Forsskål, 1775)
Blue Sea Chub
D XI, 11–13 (usually 12); A III, 10–
12 (usually 11); P1 18; LLp 49–52; GR
7–9 + 19–22 = 26–31. Body ovate,
compressed. Snout short, subequal toor shorter than eye diameter; dorsal
contour of snout slightly steep. Mouth
terminal; teeth incisor-like. Base ofspinous portion of dorsal fin longer
than base of soft rayed portion; softrayed portion of fin well elevated, anterior soft rays clearly longer than longest dorsal fin spine; anterior softrayed portion of anal fin well elevated,
similar to soft rayed portion of dorsal
fin; caudal fin slightly forked withpointed lobes. Nine to twelve (usually
10 or 11) scales between lateral line
and dorsal fin origin; 17–21 (usually
18–20) scales between lateral line toanal fin origin. Color: body bluishdorsally, dusky grayish ventrally, withseveral bluish brown longitudinal

Kyphosus cinerascens, KAUM–I. 17044, 31.0 cm SL
off Terengganu (KT), 17 Dec. 2008lines on flank, large specimens (about
40 cm SL) uniformly dusky or dark
blue dorsally; head with 2 obliquebrown or bluish brown bands; onethrough eye, the other one below eye;dorsal and anal fins dark blue, marginof soft portion with darker band; pectoral fins silver blue near base, distal
half slightly darker. Size: commonly

40 cm; maximum 50 cm TL. Distribution: widespread in the Indo-West
Pacific, including the Red Sea, fromSouth Africa east to Australia; northto southern Japan. Remarks: inhabits
coastal areas, primarily coral androcky reefs, often occurring on inner
reefs. Herbivorous.

149

DREPANEIDAE
Sicklefishes

By Mizuki Matsunuma
Body oval and strongly compressed (up to 50 cm). Interorbital,
preorbital, and broad preopercularflange naked; opercle scaly dorsally;
ventral edge of preopercle serrate injuveniles. Mouth highly protrusible,
forming a downward-pointing tube
when protruded; jaws with bands ofsetiform teeth; no teeth on roof ofmouth; maxilla exposed posteriorly,
no supramaxilla. Branchiostegal
membranes joined to isthmus; branchiostegal rays 6. A single dorsal
fin, with VIII–X spines and 19–22
soft rays; anal fin with III spines and
16–19 soft rays; caudal fin roundedor bluntly wedge-shaped (almost
truncate in large adults); pectoral finselongate, falciform, reaching caudal
peduncle, with 16–18 rays; pelvic
fins with I spine, 5 soft rays, and afleshy axillary process. Scales small,finely ctenoid, extending onto top ofhead and base of median fins; lateralline complete, strongly curved over
pectoral fin; lateral-line scales 48–
55. Vertebrae 10 + 14 = 24. Color:

single dorsal fin with VIII–X, 19–22
body very deep andstrongly compressedcaudal fin rounded or
bluntly wedge-shaped

pectoral fins elongate,
falcate, reaching caudal
peduncleanal fin with III, 16–19

silvery gray above, silverywhite below, with dusky spots or gray verticalbars.
Remarks: inhabits a variety ofhabitats including sand or mud bottoms, coral reefs, estuaries and harbours. Mostly caught with trawls.
Similar families occurring in
the area: Chaetodontidae and Pentacerotidae – mouth not highly pro-trusible; no notch between spinousand soft-rayed portions of dorsal fin.
Ephippidae – pectoral fins short, notreaching past anal-fin base; mouth
not protrusible. Scatophagidae – IVanal-fin spines; head profile concave.
Monodactylidae – mouth not highly
protrusible; pectoral fins shorter thanhead; eye centered on horizontal axis
through mouth.

Drepane longimana(Bloch & Schneider, 1801)
Sicklefish
D VIII–IX, 19–23; A III, 17–19; P1
16–18; LLp 46–55; GR 6–8 + 10–12.
Body oval and strongly compressed,
its depth 1.2–1.3 in SL. Mouth highly
protrusible, forming a downwardpointing tube when protruded. Caudal
fin rounded or bluntly wedgeshaped(almost truncate in large adults); pectoral fins elongate, falciform, reachingcaudal peduncle. Large adults with abump or bony knob on interorbital region, a result of hyperostosis of thefrontal bones. Color: head and bodysilvery with 4–10 vertical dark bars
usually visible on dorsal part of bodyfrom head to caudal-fin base. Size:maximum length about 50 cm. Distribution: temperate and tropical IndoWest Pacific, from South Africa and
Red Sea to New Guinea and northern
Australia, north to southern Japan. Remarks: occurs in inshore waters withsand or mud bottoms.

Drepane longimana, KAUM–I. 16903, 12.2 cm SL
off Terengganu (KT), 6 Dec. 2008

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Drepane punctata(Linnaeus, 1758)
Spotted Sicklefish
D VIII–X, 20–22; A III, 16–19; P1
16–18; LLp 46–50; GR 5 + 10–11.
Body oval and strongly compressed,
its depth 1.2–1.3 in SL. Mouth highly
protrusible, forming a downwardpointing tube when protruded. Caudal
fin rounded or bluntly wedgeshaped(almost truncate in large adults); pectoral fins elongate, falciform, reachingcaudal peduncle. Large adults with abump or bony knob on interorbital region, a result of hyperostosis of frontalbones. Color: generally silvery with agreenish tinge on upper half of body;
5–10 series of black spots arranged invertical lines on dorsal part of bodyfrom below dorsal fin to caudal peduncle. Size: maximum length about
40 cm. Distribution: temperate andtropical Indo-West Pacific, from Red
Sea and east coast of Africa to New
Guinea, Samoa, and northern Australia, north to southern Japan. Remarks:
occurs in inshore habitats, such as
sand or mud bottoms, reefs and estuaries. Feeds on benthic invertebrates.

Drepane punctata, KAUM–I. 17067, 25.7 cm SL
off Terengganu (KT), 26 Dec. 2008

Drepane punctata, KAUM–I. 17118, 2.2 cm SL
off Terengganu (KT), 31 Dec. 2008

151

CHAETODONTIDAE
Butterflyfishes
By Yusri Yusuf
Small sized (up to 30 cm) marine
fishes, with body deep and highly
compressed. Head short; preopercle
smooth. Eye moderately small, nearlocated on longitudinal axis fromtip of snout to middle of caudal fin.
Snout highly variable, very short toextremely elongate. Mouth small,terminal, protractile; teeth bristlelike, curved, arranged in rows orbands across jaws; vomer and palatines without teeth. Branchiostegalrays 6 or 7. Gill rakers short. A single dorsal fin, usually with X–XIV
strong, stout spines and 15–30 softrays; no notch between spinous andsoft rayed portions of dorsal fin. Anal
fin with III–V strong, stout spinesand 14–23 soft rays: interspinous
membranes deeply incised; marginusually rounded but sometimes angular. Pectoral fin with 13–15 rays.
Pelvic fin with I stout spine and 5 soft
rays. Caudal fin rounded to slightly
emarginate. Scales ctenoid, covering
head, body, and median fins; scaly
axillary process at upper base of pelvic fins; number of lateral line scalesvariable, 20–90. Vertebrae 11 + 13.

single dorsal fin with
VI–XVI, 15–30

scales ctenoid
mouth verysmall

anal fin with III–V, 14–23

Color: most species very brightly colored with complex and varied color
patterns; many species with a dark
occellate “false-eye” spot on posterior
portion of body.
Remarks: generally found oncoral reefs, usually in shallow water
above depths of 20–30 m. Feed oncoral polyps, colonial sea anemones,
tentacles of tubeworms, small crustaceans, zooplankton, and algae. No importance in commercial fisheries; but

commonly treated as an aquarium
fish.
Similar families occurring in
the area: Pomacanthidae – a strong
spine at angle of preopercle; most ofconspicuous species larger and morecolorful. Scatophagidae – dorsal fin
with a deep notch; IV anal fin spines;
mouth not protractile. Zanclidae –strongly produced snout, bony supraocular projections, and only VII
dorsal fin spines.

Chaetodon octofasciatus
Bloch, 1787
Eightbanded Butterflyfish
D X–XII, 17–20; A III, 14–17; P1
12–14; LLp 27–38; LR 36–42. Body
rounded, almost circular; snout short,blunt; predorsal contour straight. Lateral line incomplete. Dorsal and anal
fins strongly rounded, caudal fin truncate to slightly rounded. Color: overall
creamy yellow with eight black vertical
stripes, including an eye band, 5 central
body stripes, and the other 2 stripes
crossing dorsal and anal fins; pelvic
fins yellow. Size: maximum length
about 10 cm. Distribution: eastern Indian Ocean and western Pacific, from
Maldives and Sri Lanka to New Guinea
and Great Barrier Reef. Remarks: inhabits coral reefs at depths of 3–20 m.
Forms pairs or small groups. The mostcommon butterflyfish found in coral
reef of Malay Peninsula; juvenile usually found hiding in branching coral.

Chaetodon octofasciatus, UMTF 1235 (KAUM–I. 16601), 4.1 cm SL
Bidong Island, 12 Oct. 2008

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Coradion chrysozonus(Cuvier, 1831)
Orangebanded Coralfish
D IX, 28–30; A III, 19–21; P1 15–16;
LR 48–52. Body rounded, almost circular; its depth 1.3–1.5 in SL; snout
short and blunt. Dorsal and anal finsstrongly rounded and elevated; caudal
fin truncate to slightly rounded. Lateral line complete. Color: overall
whitish, with 2 dark yellow-orange
bars at the level of pelvic fin; darkbrown bars through eye, and caudal
fin base; pelvic fin dark brown to
black. Juveniles with a white-edged
ocellus in dorsal fin that reduces to a
black spot in adults. Size: maximum
length about 15 cm. Distribution:eastern Indian Ocean and western Pacific: from the Andaman Sea eastward
throughout Indo-Malayan region, and
the Great Barrier Reef northward to
southern Japan. Remarks: found incoastal to outer reef drop-offs; depth
range 3–60 m. Feeds on sponges.

Coradion chrysozonus, UMTF 1361 (KAUM–I. 16640), 11.2 cm SL
Bidong Island, 12 Oct. 2008

Parachaetodon ocellatus(Cuvier, 1831)
Ocellate Coralfish
D VI–VII, 28–30; A III, 18–20; P1
14–15; LR 39–46. Body rounded, almost circular; its depth 1.3–1.4 in SL.
Dorsal fin triangular in shape and first
soft ray at apex. Color: head and body
pearly white, with five dusky to orange bands; the first band through theeye orange, with black edges; an ocellated black spot in forth body band;
band on caudal peduncle with a silver
anterior margin; upper portion of dorsal fin mostly yellow. Size: maximum
length about 17 cm. Distribution:eastern Indian Ocean and western Pacific, from Sri Lanka eastward
throughout the Indo-Malayan region,and the Great Barrier Reef northward
to the Ryukyu Islands, Japan. Remarks: inhabits lagoons and coral
reefs at depths of 3 to 50 m. Occurs inpairs on flat sand or silty bottoms oncoastal reefs. Adults school over open
muddy substrates in deep water; juveniles sometimes in large lagoons withseagrasses in depths of about 5 m or
more. Rare in coral reefs area of Malay Peninsula.

Parachaetodon ocellatus, KAUM–I. 17217, 10.5 cm SL
off Terengganu (KT), 11 Jan. 2009

153

POMACANTHIDAE
Angelfishes

By Yusri Yusuf
Small to medium sized (up to 50cm) marine fishes, with deep body
oblong to oval-shaped, highly compressed. Head short; 1 or more prominent spines at angle of preopercle.
Mouth small, terminal, protractile;teeth bristlelike, usually tricuspid, arranged in rows or bands across jaws;
vomer and palatines without teeth. A
single dorsal fin with IX–XV strong,stout spines and 15–33 soft rays; nonotch between spinous and soft-rayed
portions. Anal fin with III strong, stout
spines and 14–25 soft rays. Caudal fin
variable, rounded to slightly emarginate, or forked with long filamentous
extensions. Pectoral fin with 16–21
rays. Pelvic fin with a single stout
spine and 5 soft rays. Scales ctenoid,
covering head, body, and median fins;
number of lateral line scales variable,
30–90. Vertebrae 10 + 14. Color: almost all very brightly colored withcomplex and varied color patterns. Juveniles are often strikingly different incolor from adults. Color patterns are

single dorsal fin with IX–XV, 15–33

one or more
prominent spines atangle of preopercle

most diagnostic for species identification.
Remarks: generally found on coral reefs at depths of 5–40 m. Frequently
utilized as aquarium fish.
Similar families occurring in thearea: Chaetodontidae – lack a strong
spine at angle of preopercle; an axil-

anal fin with III, 14–25

lary scaly process present at bases ofpelvic fins. Ephippidae – lack a strong
spine at angle of preopercle; generally
less colorful as adults; dorsal and anal
fins often very elongate. Scatophagidae – dorsal fin with a deep notch,
lacking spines at angle of preopercle,
and having IV anal-fin spines.

Pomacanthus annularis(Bloch, 1787)
Blueringed Angelfish
XIII, 20–22; A III, 20–21; P1 19; GR
4–5 + 12. Body oblong, very deep andcompressed; a long spine at corner of
preopercle. Soft portions of dorsal andanal fins round; caudal fin rounded.
Color: body yellowish brown, with
5–8 diagonal brilliant blue stripes onsides; a blue ring above upper edge ofgill opening, at the origin of lateralline; 2 horizontal blue stripes on headacross nape before eye onto edge of
opercle, the upper stripe at the level of
mid eye, the lower at the level just beneath lower edge of eye; base of pectoral fins with blue band; caudal fin
white. Juveniles bluish-black with aseries of white and blue narrow bars
on sides, anterior bars slightly curved;
Pomacanthus annularis, KAUM–I. 17159, 25.0 cm SL
no white ring before caudal peduncle;
Redang Island, 4 Jan. 2009no marking on caudal fin. Size: maximum length about 30 cm. Distribution: Indo-Pacific, from East Africa to turbid water, around caves and wrecks cates. The most common anglefish in
New Caledonia. Remarks: inhabits at depths of 10–30 m. Found solitary coral reef areas of Malay Peninsula.
lagoons and coral reefs, frequently in or in pairs. Feeds on sponges and tuni-

154

TERAPONTIDAE

Grunters (Tigerperches)
By Mizuki Matsunuma
Small to medium-sized (to 35cm) perch-like fishes. Body oblongand slightly to moderately compressed. Mouth moderate, protractile, upper jaw not extending beyond
center of orbit; jaw teeth usually invilliform bands, shape of teeth conical, flattened, or tricuspidate; vomer
and palatines toothless in adults
of most species. Opercle bearing
2 spines, the lower one larger andstronger; posttemporal bone exposedand expanded posteriorly, with posterior margin serrate in some species.
Dorsal fin single, notched, with XI–
XIV spines and 8–14 soft rays; anal
fin with III and 7–12 soft rays; pelvic
fins inserted behind base of pectorals,with I spine and 5 soft rays; caudal
fin usually emarginate. Scales adherent, finely ctenoid; lateral line single
and complete, reaching on caudal
fin. Color: body tan or light gray, often silvery in life with various dark

single dorsal fin with XI–XIV, 8–14

2 opercular spines

anal fin with III, 7–12

markings; most marine species with 3
or more dark, straight or downwardly
curved longitudinal stripes on body;some with dark transverse bands on
lobes of caudal fin.
Remarks: marine terapontids inhabiting inshore marine and brackish
waters, with some species also enter-ing hypersaline and fresh waters.
Similar families occurring in
the area: Serranidae – mouth large,
with upper jaw typically reaching
to below vertical through posterior
margin of eye; caudal fin typicallyrounded; 3 strong spines on opercle.
Kuhliidae – dorsal fin with X spines.

Pelates quadrilineatus(Bloch, 1790)
Fourlined Terapon
D XII–XIII, 9–11; A III, 9–10; LLp
66–75; GR 16–18 + 22–27. Body oblong and compressed laterally. Preopercle serrate; lower opercular spine
stronger and longer, but not extending
beyond margin of opercular lobe;
posttemporal bone not expanded or
exposed posteriorly, covered with skinand scales. Color: body silvery white,grayish dorsally; 4–6 narrow darkbrown or black horizontal stripes onbody; spinous portion of dorsal fin
with a black blotch dorsally on membranes between third to seventh dorsal-fin spines; a large black blotch onside of body posterior to nape; lobes
of caudal fin without prominent transverse black stripes. Size: maximum
length about 30 cm, commonly to 20
cm. Distribution: Indo-West Pacific:

Pelates quadrilineatus, KAUM–I. 17033, 13.8 cm SL
off Terengganu (KT), 17 Dec. 2008

Red Sea and East Africa to New Guinea, northern Australia north to Japan.
Remarks: often found in brackish waters. Feeds on small fishes and invertebrates.

155

Terapon jarbua(Forsskål, 1775)
Jarbus Terapon
D XI–XII, 9–11; A III, 7–10; LLp
75–100; GR 6–8 + 12–15. Body oblong and moderately compressed. Preopercle strongly serrate, particularly atangle; lower opercular spine very longand strong, extending distinctly beyond
margin of opercular lobe; posttemporalbone expanded, exposed and serrate
posteriorly. Caudal fin emarginate.
Color: body silvery white, grayish dorsally; 3 or 4 dark brown or black downwardly curved longitudinal stripes onbody; spinous portion of dorsal fin witha blackish blotch dorsally on membranes between third and sixth spines;
both caudal-fin lobes with dark tips anda transverse band. Size: maximum
length about 35 cm. Distribution: Indo-Pacific: East Africa to Samoa, northto southern Japan, south to Australia.
Remarks: found over shallow sandy
bottoms, in the vicinity of river mouths;
enters estuaries and rivers.

Terapon jarbua, KAUM–I. 16986, 19.5 cm SL
off Terengganu (KT), 13 Dec. 2008

Terapon theraps
Cuvier, 1829
Largescaled Terapon
D XI–XII, 9–11; A III, 7–9; P1 14–
15; LLp 46–56; GR 6–8 + 14–17.
Body oblong, moderately compressed.
Preopercle serrate, the serrations largest at angle; lower opercular spine
very long and strong, extending distinctly beyond margin of opercular
lobe; posttemporal bone expanded,
exposed and serrate posteriorly. Spinous part of dorsal fin strongly archedand deeply notched; penultimate spine
about half length of ultimate spine.
Caudal fin emarginate with rounded
lobes. Color: body dusky dorsally, silvery white ventrally; 4 dark brownhorizontal stripes on body; spinouspart of dorsal fin with a black blotch
dorsally on fin membranes between
third and seventh spine; caudal fin
with medial rays pigmented; upper
lobe of caudal fin with dark tip; both
lobes of caudal fin with a dark, transverse band. Size: maximum length
about 30 cm. Distribution: Indo-Pacific, from East Africa and Red Sea to
Solomon Islands, northern Australia
north to Japan. Remarks: found in inshore waters, sometimes entering
brackish waters.

Terapon jarbua, UMTF 1068 (KAUM–I. 16484), 7.0 cm SL
estuary near UMT, 30 Sept. 2008

Terapon theraps, KAUM–I. 16892, 11.7 cm SL
off Terengganu (KT), 6 Dec. 2008

156

CEPOLIDAE
Bandfishes

By Mizuki Matsunuma
Small to medium (up to 40 cm)
elongate marine fishes. Body moderately to noticeably elongate fisheswith compressed, tapering body andlanceolate caudal fin. Head short,with blunt snout. Eyes relatively
large and high on head. Mouth large,
oblique; upper jaw broad at end,
without supramaxilla, and extending
to below posterior margin of eye; asingle row of slender, slightly curved
teeth in jaws with an inner clusterof teeth at symphysis in some species. Dorsal fin continuous, with 0
to IV flexible spines and 21 to 89
segmented rays; anal fin with 0 or I
spine and 13 to 102 segmented rays;
caudal fin lanceolate, middle 9 to 15
rays branched; pelvic fins positioned
below or slightly anterior to pectoral
fins, with I spine and 5 segmented
rays; outermost segmented pelvic-fin
ray unbranched or weakly branched,
4 inner rays distinctly branched. Lat-dorsal fin with 0–IV spines and 21–89 soft rays

pelvic fins below
pectoral finscaudal fin
lanceolate

anal fin with 0 or I spine and 13–102 soft rayseral line high on body, close to dorsalfin base, terminating posteriorly nearend of fin; lateral-line tubes or canals
on scales not embedded in skin. Scales
cycloid (smooth) or with crenulate
margins, relatively large to minute.
Color: in life red or pink; most species have a distinctive dark stripe on
the membrane (usually hidden) connecting the premaxillary and maxillary bones of the upper jaw.
Remarks: relatively uncommon
fishes taken by trawls in shallow to

deep depths (to at least 475 m). Occurring on level bottom, sand or mud
substrates.
Similar families occurring in
the area: the combination of a lanceolate tail, large oblique mouth, andthe arrangement of the pelvic-finrays, consisting of I spine and 5 segmented rays (the outermost ray unbranched or weakly branched and theinner 4 branched), distinguishes the
bandfishes from all other families.

Acanthocepola abbreviata(Valenciennes, 1835)
Yellowspotted Bandfish
D 64–75; A 63–76; P1 16; P2 I, 5; V
12 + 44–48. Body well elongate andcompressed. Head short, with bluntsnout; eyes large and high on head;mouth large, oblique; preopercularmargin with spines; cheeks scaly. Pelvic fin origin situated slightly anterior
to pectoral fin; caudal fin pointed,
joined to dorsal and anal fins; 8–10
rays branched. Color: head and body
pale pink, reddish dorsally; body with
about 4–5 orange spots (same as pupilsize) on dorsoanteriorly and about
11–13 indistinct orange vertical bars
posteriorly, from mid-point of body tocaudal fin base. Dorsal and anal finsreddish yellow, whitish basally; no
black markings on dorsal fin; pectoral
fin translucent tinged with reddish;pelvic fin white; posterior tip of cau-

Acanthocepola abbreviata, KAUM–I. 17135, 28.4 cm SL
off Terengganu (KT), 1 Jan. 2009

Acanthocepola abbreviata, KAUM–I. 17136, 21.2 cm SL
off Terengganu (KT), 1 Jan. 2009

dal fin dark red to black. Size: maxi- Philippines. Remarks: inhabits mud
mum length about 30 cm. Distribu- or sand bottoms of coastal waters.
tion: eastern Indian Ocean and West Caught mostly with bottom trawls.
Pacific from the Gulf of Oman tonorthwestern Australia, north to the

157

POMACENTRIDAE

Damselfishes and Anemonefishes
By Yusri Yusuf and Koichi Shibukawa
Small to medium-sized fishes,
usually less than 20 cm. Body compressed, ovate to almost circular,
covered with ctenoid scales usuallyextending onto fins; dorsal fin continuous with VIII–XVII spines andno notch; anal fin with II spines and
10–16 soft rays; caudal fin emarginated to forked or lunate. Lateral line
interrupted; anterior part of pored
scales more or less parallel with dorsal profile and usually ending below
soft portion of dorsal fin; posterior
part comprising several pits or obscure pores along mid-lateral caudal
peduncle; snout usually short andblunt; mouth small, slightly protractile; teeth conical or compressed,
uniserial or in 2 or more rows; noteeth on palatine. Color: highly variable; mainly from pale through yellowish, orange, reddish or bluish to
blackish; sometimes with blotches,bands, spots, or other patterns.

single dorsal fin with VIII–XVII spines

lateral line interrupted

mouth small

anal fin with II, 10–16

Remarks: found in coastal waters
such as bays, coral reefs and rocky
shores, and often enter brackish estuaries or freshwater streams. Frequently utilized as aquarium fish.

Similar families occurring in
the area: Pomacanthidae – a long,strong spine at the corner of preopercle. Chaetodontidae – elongate,somewhat tube-like snout.

Abudefduf bengalensis(Bloch, 1787)
Bengal Sergeant
D XIII, 13–15; A II, 13–14; P1 16–
20; LLp 19–23. Body deep and compressed. Posterior margin of preopercle smooth. Suborbital scaled, withentire ventral margin. Teeth on jaws
uniserial, incisor-like. Caudal fin
forked with rounded tail lobes; noconspicuous projecting spine-like rays
at upper and lower edges of caudal-finbase. Color: ground color pale or lightgray, gradually darkened dorsally; 7narrow blackish bars (narrower thanpaler interspaces) from nape to caudal
fin base; dorsal end of pectoral-finbase with a distinct black spot. Size:
14 cm SL. Distribution: Indo-West
Pacific: from Pakistan to Great Barrier
Reef, north to southern Japan. Remarks: found in shallow coral reefs
and lagoons to depths of 6 m.
(K. Shibukawa & Y. Yusuf)

Abudefduf bengalensis, UMTF 1053 (KAUM–I. 16401), 6.2 cm SL
Dungun, 24 Sept. 2008

158

Abudefduf sexfasciatus(Lacepède, 1801)
Scissortail Sergeant
D XIII, 11–14; A II, 11–13; P1 17–
20; LLp 18–22; GR 23–30. Body
ovate, compressed; margins of suborbital and preopercle smooth; teeth onanterior part of jaws flattened, uniserial; suborbital scaled. Caudal fin
forked; no conspicuous projecting
spine-like rays at upper and lower
edges of caudal-fin base. Color: bodywhite with 5 black bars on side; ablack stripe on each caudal fin lobe;
fins whitish but a short black stripe on
soft dorsal and anal fins. Size: 17 cm
SL. Distribution: Indo-West Pacific.
Remarks: inhabits inshore and offshore coral or rocky reefs; depth range
1–20 m. One of the most common
damselfish in coral reef areas of Malay Peninsula.
(Y. Yusuf)

Abudefduf sexfasciatus, KAUM–I. 17047, 9.7 cm SL
off Terengganu (KT), 18 Dec. 2008

Abudefduf sordidus(Forsskål, 1775)
Black-spot Sergeant
D XIII, 14–16; A II, 13–15; P1 18–
20; LLp 21–23. Body deep and compressed. Posterior margin of preopercle smooth. Suborbital scaled, withentire ventral margin. Teeth on jaws
uniserial, incisor-like. Caudal fin
forked; no conspicuous projecting
spine-like rays at upper and lower edges of caudal-fin base. Color: groundcolor gray; 6 dark brown bars (broader
than pale interspaces) from nape tocaudal-fin base; axil and dorsal end of
pectoral-fin base black; a distinct,
small saddle-like black blotch on anterior part of caudal peduncle. Size: 17
cm SL. Distribution: Indo-West Pacific. Remarks: found in shallow
rocky shore, reef flat and piers exposed
to waves; depth range 0.2–3 m. Feedon algae, crustaceans and other invertebrates. (K. Shibukawa & Y. Yusuf)

Abudefduf sordidus, UMTF 1486 (KAUM–I. 16335), 2.2 cm SL
Cendering, 16 Sept. 2008

Amblypomacentrus breviceps(Schlegel & Müller, 1839)
Black-banded Demoiselle
D XIII, 11–12; A II, 12–13; P1 16–
17; LLp 16–17. Body slender, its depth
2.1–2.3 in SL; caudal fin forked, withlobes filamentous in adults. Color:
pearly white, with 3 black bars, thefirst running across the head throughthe eye, and other 2 from the back to

Amblypomacentrus breviceps, UMTF 1254 (KAUM–I. 16585), 3.8 cm SL
Bidong Island, 13 Oct. 2008

about mid portion of the side; usuallywith longitudinal rows of blue spots onsides. Size: 8.5 cm TL. Distribution:
Western Central Pacific. Remarks: in-habits lagoon and coastal areas in sandy or silty areas around outcrops of
rubble on sand, mud, sponge or rocks;
depth range 2–35 m.
(Y. Yusuf)

159

Amphiprion clarkii(Bennett, 1830)
Clark’s Anemonefish
D X–XI, 14–17; A II, 12–15; P1
18–21; LLp 34–45. Body moderately
deep and compressed. Opercle, subopercle, preopercle and interopercle
serrate. Teeth on jaws uniserial. Caudal fin weakly emarginate; no conspicuous projecting spine-like rays at
upper and lower edges of caudal-finbase. Color: ground color variablefrom reddish yellow to blackish; threewhite bars on head and body. Tailwhite or yellow; other fins variable
black to yellow-orange. Size: 8.5 cm
SL. Distribution: Indo-West Pacific:
Persian Gulf east to Vanuatu and Marshall Islands, northern Australia northto southern Japan. Remarks: found incoral reefs to depth of 55 m; symbiotically associated with anemones Heteractis magnifica and Stichodactyla
mertensii.
(K. Shibukawa & Y. Yusuf)

Amphiprion clarkii, UMTF 1238 (KAUM–I. 16578), 8.8 cm SL
Bidong Island, 13 Oct. 2008

Amphiprion frenatus
Brevoort, 1856
Tomato Anemonefish
D IX–X, 16–18; A II, 13–15; P1
18–21; LL 31–44. Body moderately
deep and compressed; its depth 1.7–
2.0 in SL; posterior margins of opercle, interopercle and subopercle
strongly serrated. Caudal fin weakly
emarginate. Color: adults with a single white bar on head; females mainly
blackish on sides with red or orange
snout, breast, belly, and fins; males redor orange overall and lacking blackishcoloration, considerably smaller thanfemales. Juveniles with two or threewhite bars. Size: 14.0 cm TL. Distribution: western Pacific: Gulf of Thailand, South China Sea to southwestern
Palau, north to southern Japan, south
to Java, Indonesia. Remarks: found inlagoon reefs and embayment; depth
range 1–12 m. Monogamous and protandrous hermaphrodite. Associatedwith the anemone Entacmaea quadricolor. Feeds on algae, fish eggs, andpelagic copepods. Similar to A. melanopus, but with a broader white headbar and black pelvic and anal fins.
(Y. Yusuf)

Amphiprion frenatus, KAUM–I. 16579, 6.7 cm SL
Bidong Island, 13 Oct. 2008

Amphiprion frenatus, UMTF 1319 (KAUM–I. 16580), 5.6 cm SL
Bidong Island, 13 Oct. 2008

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Amphiprion ocellaris
Cuvier, 1830
False Crown Anemonefish
D X–XI, 14–17; A II, 12–15; P1
18–21; LLp 34–45. Body moderately
deep and compressed. Opercle, subopercle, preopercle and interopercle
serrate. Teeth on jaws uniserial. Caudal fin weakly emarginate; no conspicuous projecting spine-like rays at
upper and lower edges of caudal-finbase. Color: ground color of head,
body and fins orange, with three broad
white bars; fins with marginal or submarginal narrow black areas. Size: 8
cm SL. Distribution: east part of Indian Ocean, Andaman Sea and West
Pacific. Remarks: found in coral
reefs to depths of 15 m; symbioticallyassociated with anemones Heteractis
magnifica, Stichodactyla gigantea andStichodactyla mertensii. The mostcommon anemonefish found in coral
reef of Malay Peninsula.
(K. Shibukawa & Y. Yusuf)

Amphiprion ocellaris, UMTF 1236 (KAUM–I. 16582), 5.9 cm SL
Bidong Island, 13 Oct. 2008

Amphiprion perideraion
Bleeker, 1855
Pink Anemonefish
D IX–X, 16–17; A II, 12–13; P1
16–18; LL 32–43. Body depth 1.9–2.2
in SL; posterior margins of opercle,
interopercle and subopercle strongly
serrated; caudal fin rounded. Color:
pink to pinkish orange; fins transparent; narrow white head bar and white
stripe on top of head beginning between the eyes and extending alongbase of dorsal fin; adult males havenarrow orange margin on soft dorsal
fin and upper and lower edges of tail.
Size: 10 cm. Distribution: eastern Indian Ocean and western Pacific, fromthe Andaman Sea east to Cocos Islands and Tonga, northern Australia
north to southern Japan. Remarks:
inhabits lagoon and seaward reefs todepth of 38 m. Associated with theanemones Heteractis magnifica (usually), H. crispa, Macrodactyla doreensis, and Stichodactyla gigantea. Similar to A. akallopisos and A. sandaracinos
but lack the white head bar.
(Y. Yusuf)

Amphiprion ocellaris, UMTF 1406 (KAUM–I. 16559), 2.4 cm SL
Bidong Island, 13 Oct. 2008

Amphiprion perideraion, UMTF 1233 (KAUM–I. 16581), 6.2 cm SL
Bidong Island, 13 Oct. 2008

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Chromis atripectoralis
Welander & Schultz, 1951
Black-axil Chromis
D XII, 9–10; A II, 9–10; P1 18–20;
LLp 15–16. Body deep and compressed. Posterior margin of preopercle smooth. Suborbital scaled, without
visible ventral free edge. Teeth onjaws conical, multiserial; teeth of outermost row larger than those of innerrows. Caudal fin forked; three conspicuous projecting spine-like rays at
upper and lower edges of caudal-finbase. Color: head, body and most partof fins bright greenish blue; single orsome minute blue dots or short irregular lines on each scales; axil of pectoral black. Size: 8.5cm SL. Distribution: Indo-Pacific. Remarks: foundaround branching corals on coral reefs
in depths of 2–15 m; forms aggregations; feeds on zooplankton and fish
eggs.
(K. Shibukawa & Y. Yusuf)

Chromis atripectoralis, UMTF 1391 (KAUM–I. 16591), 6.5 cm SL (preserved specimen)
Bidong Island, 12 Oct. 2008

Chrysiptera unimaculata(Cuvier, 1830)
Onespot Demoiselle
D XIII, 13–14; A II, 12–14; P1 18–
19; LLp 16–18. Body moderate toslender, its depth 2.1–2.4 in SL. Posterior margin of preopercle smooth. No
scales on infraorbitals; cheek with 2
rows of scales. No conspicuous projecting spine-like rays at upper andlower edges of caudal-fin base. Color:
adults usually uniformly dark brown,with a black spot at the end of the dorsal fin base; a small orange spot on
gill-plate; pectoral fin yellowish. Juvenile yellowish with blue stripe on
upper head, blue-ring black spot below mid dorsal fin, and dark spot atrear dorsal fin. Size: 10.0 cm TL. Distribution: Indo-West Pacific: Red
Sea and East Africa to Fiji, north to
Ryukyu Islands, south to southern
Great Barrier Reef; Palau in Micronesia. Remarks: found solitarily or insmall groups in shallow water amongcoastal algal reefs, rubble or over open
beach-rock of reef flats exposed tomoderate surge; to depth of 3 m. Feeds
mainly on benthic algae. (Y. Yusuf)

Chrysiptera unimaculata, UMTF 1837 (KAUM–I. 16397), 4.5 cm SL
Dungun, 24 Sept. 2008

Chrysiptera unimaculata, KAUM–I. 16404, 1.7 cm SL
Dungun, 24 Sept. 2008

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Dascyllus reticulatus(Richardson, 1846)
Reticulate Dascyllus
D XII, 14–16; A II, 12–14; P1 19–21;
LLp 20; GR 7 + 19–21. Body deep,
compressed. Margins of preorbital,
suborbital, and preoperculum finely
serrated. Caudal fin emarginate, withrounded lobes; two conspicuous projecting spinelike rays at upper andlower edges of caudal-fin base. Color:various; influenced by ecological andbehavioral conditions. Basically bodywhitish with snout, interorbital andforehead greenish; black bar on anterior through pectoral base and a fainterone on posterior part of the body;
blackish scale margins; spinous dorsal
fin same as body color on basal halfwith broad black margin on outer half;
soft dorsal fin transparent to slightly
bluish pelvic fins mostly black; pectorals transparent with a spot on its base.
Size: 9.0 cm TL. Distribution: eastern Indian Ocean and western Pacific.
Remarks: occurs in reef outer lagoon
and seaward reefs to depths of 50 m.
Inhabits branching coral heads, particularly Pocillopora sp. and Acropora spp. Forms schools. Feeds onplants, benthic and planktonic invertebrates. Common reef-associated fishin the area; replaced by D. carneus in
the Indian Ocean.
(Y. Yusuf)

Dascyllus reticulatus, UMTF 1418 (KAUM–I. 16600), 2.5 cm SL
Bidong Island, 12 Oct. 2008

Dascyllus trimaculatus(Rüppell, 1829)
Three-spot Dascyllus
D XII, 14–16; A II, 14–15; P1 19–21;
LLp 17–19. Body deep, compressed.
Posterior margin of preopercle serrate.
Suborbital scaled, with serrate ventral
free margin. Teeth on jaws conical,
multiserial; teeth of outermost row
larger than those of inner rows. Caudal
fin emarginate, with rounded lobes;
two conspicuous projecting spinelikerays at upper and lower edges of caudal-fin base. Color: head, body andfins black or gray, with a white spot on
dorsal part of side of body, as well as asimilar-colored spot on forehead(these white spots may obscure inlarge adult); body of large individual
light gray, with dark-margined scale

Dascyllus trimaculatus, UMTF 1302 (KAUM–I. 16587), 3.5 cm SL
Bidong Island, 13 Oct. 2008

pockets forming reticulated pattern;
juveniles overall black with lighter
scale centers, prominent white blotch
present on forehead and upper sides.
Size: 11.0 cm SL. Distribution: IndoWest Pacific. Remarks: found in cor-al and rocky reefs; inhabits lagoons,passes, and outer reed to depths of 55m; juveniles frequently symbioticallyassociated with various sea anemones;
common reef-associated fish in thearea.
(K. Shibukawa & Y. Yusuf)

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Dischistodus fasciatus(Cuvier, 1830)
Banded Damsel
D XIII, 13–14; A II, 13–14; P1 17–
18; LLp 16–17. Body depth 2.0–2.1 in
SL. Upper and lower accessory caudal
rays not spinous. Posterior margin of
preopercle serrated. Both jaws withbiserial teeth. Predorsal scales reaching above anterior part of orbit. Color:head and body pale, with a dark bar
through eye; 3 dark bars on body bars,
widest in the middle. Size: 11.5 cm
SL. Distribution: western Pacific, including Indonesia, Philippines, Taiwan and northern Australia. Remarks:
inhabits lagoon and coastal reefs withsilt and sandy bottom, around coraloutcrops and seagrass beds; depth
range 1–8 m. Can be aggressive toward SCUBA diver, defending its territory.
(Y. Yusuf)

Dischistodus fasciatus, UMTF 1337 (KAUM–I. 16621), 4.5 cm SL
Bidong Island, 12 Oct. 2008

Dischistodus prosopotaenia(Bleeker, 1852)
Honey-head Damsel
D XIII, 14–16; A II, 14–15; P1 17;
LLp 16–17. Body depth 2.1–2.2 in SL.
Upper and lower accessory caudal
rays not spinous. Posterior margin of
preopercle serrated. Both jaws withbiserial teeth. Predorsal scales reaching above anterior part of orbit. Color:
adults with body golden brown anteriorly, white posteriorly; three indistinct
brown saddles or bars below soft portion of dorsal fin; axil of pectoral fin
black; light blue dots and vertical lines
on scales. Juveniles brown, with two
white bars on sides; an ocellated spot
on dorsal fin base. Size: 17.0 cm TL.
Distribution: widely distributed in
the Indo-West Pacific from Nicobar
Islands to Vanuatu, north to the
Ryukyu Islands, south to northwest
Australia and the Great Barrier Reef.
Remarks: inhabits lagoons and coastal reefs, usually in sand and silty bottoms; depth range 1–12 m. (Y. Yusuf)

Dischistodus prosopotaenia, UMTF 1207 (KAUM–I. 16611), 9.4 cm SL
Bidong Island, 12 Oct. 2008

Dischistodus prosopotaenia, KAUM–I. 16614, 3.5 cm SL
Bidong Island, 12 Oct. 2008

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Neoglyphidodon melas(Cuvier, 1830)
Bowtie Damselfish
D XIII, 14–15; A II, 13–15; P1 18–
19; LLp 16–17. Body deep; its depth
1.7–2.0 in SL. Posterior margin of preopercle and suborbital smooth. Suborbital scaled. Teeth on jaws flattened
distally with biserial teeth. No conspicuous projecting spine-like rays at upper and lower edges of caudal fin base.
Color: adult bluish-black uniformly,
without marking. Juveniles white tolight gray, dorsum part of head andbody yellow; dorsal fin, and upper andlower margins of caudal fin yellow;
frontal margins of pelvic and anal fins
black. Size: 18.0 cm TL. Distribution:
Indo-West Pacific. Remarks: found incoral-rich areas usually with abundant
soft corals; depth range 1–12 m. Juveniles are encountered around staghorn
corals, Acropora spp.
(Y. Yusuf)

Neoglyphidodon melas, UMTF 1334 (KAUM–I. 16622), 4.3 cm SL
Bidong Island, 13 Oct. 2008

Neopomacentrus anabatoides(Bleeker, 1847)
Silver Demoiselle
D XIII, 10–11; A II, 10–11; P1 17–
18; LLp 16–18. Body relatively slender, compressed. Posterior margin of
preopercle weakly serrate. Ventral
margin of suborbital concealed byscales. Teeth on jaws biserial, incisor
like. Caudal fin forked; no conspicuous projecting spine-like rays at upperand lower edges of fin base. Color:head and body metallic green, whitish
ventrally; scales dark marginally;small black blotches (subequal pupil
sized) on upper end of gill opening,
and upper pectoral fin base. Size: 10.5
cm TL. Distribution: western Pacific.
Remarks: schooling fish inhabiting
soft bottoms around coral or rock outcrops; depth range 2–15 m. (Y. Yusuf)

Neopomacentrus anabatoides, UMTF 1239 (KAUM–I. 16565), 5.5 cm SL
Bidong Island, 14 Oct. 2008

Neopomacentrus cyanomos(Bleeker, 1856)
Regal Damoiselle
D XIII, 11–12; A II, 11–12; P1 17–
18; LLp 17–18. Body relatively slender, compressed. Posterior margin of
preopercle weakly serrate. Ventral
margin of suborbital with scales. Teethon jaws biserial, incisor-like. Caudal
fin forked, with filamentous upper andlower lobes; no conspicuous projecting spine-like rays at upper and lower

Neopomacentrus cyanomos, UMTF 1021 (KAUM–I. 16648), 6.6 cm SL
Bidong Island, 13 Oct. 2008

edges of caudal-fin base. Color: headand body dark brownish gray; a small
black blotch (subequal pupil sized) on
upper end of gill opening; a small
black spot on upper origin of pectoral
fin; posterior part of dorsal fin pale toyellow; caudal fin pale to yellow, withblack upper and lower margins. Size:
7.0 cm SL. Distribution: Indo-West
Pacific. Remarks: found around
patch reefs in protected bays with sandy muddy bottoms in depth of 5–18 m.
Feeds on plankton in aggregation.
(K. Shibukawa & Y. Yusuf)

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Neopomacentrus taeniurus(Bleeker, 1856)
Freshwater Demoiselle
D XII–XIII, 11–12; A II, 10–11; P1
17–18; LL 16–17. Body depth 2.2–2.6
in SL. Posterior margin of preopercle
serrated. Both jaws with biserial teeth.
Predorsal scales reaching above anterior nostril. Caudal fin with filamentous lobes; no conspicuous projectingrays on base. Color: body brown, withscales dark marginally; a small black
spot on upper edge of opercle near origin of lateral line; a large black spot on
upper pectoral fin base; posterior partof dorsal and anal fins pale white toyellow; caudal fin pale white to yellow, with base and both lobes darkbrown. Size: 10 cm TL. Distribution:
Indo-West Pacific. Remarks: inhabitsmangroves, estuaries, lower reaches offreshwater streams, and harbors withfreshwater discharge; to depths of 3 m.
(Y. Yusuf)

Neopomacentrus taeniurus, UMTF 1028 (KAUM–I. 16762), 5.2 cm SL
Setiu, 31 Oct. 2008

Pomacentrus alexanderae
Evermann & Seale, 1907
Alexander's Damsel
D XIII, 14–15; A II, 14–15; P1 16–
17; LLp 15–17. Body ovate, compressed. Posterior margin of preopercle serrate. Suborbital naked, withserrate ventral margin. Teeth on jaws
biserial, incisor-like. Caudal fin emarginate; no conspicuous projecting
spine-like rays at upper and lower edges of caudal-fin base. Color: head andbody pale gray, with a large blotch on
pectoral fin base; tips of dorsal spines
black. Size: 9.0 cm TL. Distribution:
western Pacific. Remarks: occurs inlagoon, inshore, and offshore reefs;
depth range 5–60 m. Feed on algae,
barnacle nauplii, copepods, fish eggs,
and small gastropods.
(Y. Yusuf)

Pomacentrus alexanderae, UMTF 1339 (KAUM–I. 16556), 3.0 cm SL
Bidong Island, 13 Oct. 2008

Pomacentrus chrysurusCuvier, 1830
Whitetail Damsel
D XIII, 14–16; A II, 15–16; P1 18;
LLp 18–19. Body ovate, compressed.
Posterior margin of preopercle serrate.
Suborbital naked, with serrate ventral
margin. Teeth on jaws biserial, incisorlike. Caudal fin emarginate; no conspicuous projecting spine-like rays at
upper and lower edges of caudal-finbase. Color: head and body dusky,

Pomacentrus chrysurus, UMTF 1227 (KAUM–I. 16613), 4.2 cm SL
Bidong Island, 12 Oct. 2008

darkened posteriorly and abruptly turnto white in caudal fin; a small black
spot at dorsalmost part of pectoral finbase; juvenile tinged with blue andwith orange over back and pale-edged
black ocellus at posterior part of dorsal

fin (ocellus may persisting in adult).
Size: 7.0 cm SL. Distribution: IndoWest Pacific. Remarks: found in sandy areas of shallow inshore coral reefs
to depths of 3 m.
(K. Shibukawa & Y. Yusuf)

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Pomacentrus coelestis
Jordan & Starks, 1901
Neon Damselfish
D XIII, 13–15; A II, 14–15; P1 17–
18; LL 17–18. Body depth 2.5–2.6 in
SL. Posterior margin of preopercle serrated; ventral margin of infraorbitalsmooth. Both jaws with biserial teeth.
Predorsal scales reaching above anterior nostril; no scales on infraorbitals.
Caudal fin emarginate; no conspicuous
projecting spine-like rays at upper andlower edges of caudal fin base. Color:body blue, often with iridescent glow;
yellow-orange on ventral part of body,posterior parts of dorsal, anal and on
caudal fins. Size: 10 cm SL. Distribution: eastern Indian Ocean and western Pacific. Remarks: inhabits lagoon
and seaward reefs, close to bottom
among rubble beds; depth range 1–20
m.
(Y. Yusuf)

Pomacentrus coelestis, KAUM–I. 16594, 3.3 cm SL
Bidong Island, 12 Oct. 2008

Pomacentrus cuneatus
Allen, 1991
Wedgespot Damsel
D XIII, 14–15; A II, 14–15; P1 16–
18; LLp 17. Body ovate, compressed.
Body depth 2.0–2.1 in SL; posterior
margin of preopercle serrate. Pre andsuborbital naked, with serrate ventral
margin. Teeth on jaws biserial. No
conspicuous projecting spine-like rays
at upper and lower edges of caudal-finbase. Color: adults gray-brown uniformly, with dark scale margin; ablack wedge-shaped spot on dorsaledge of opercle; a small black spot on
upper pectoral fin base; tips of dorsal
fin spines bluish. Juvenile yellowish,
with several blue stripes from snout toposterior part of body; blue-edged
black ocellus at posterior part of dorsal fin. Size: 7.5 cm SL. Distribution:
Western Central Pacific. Remarks:
inhabits sheltered inshore coral reefs,
usually in turbid silty, degraded reefarea; depth range 1–15 m. Solitary orin small group.
(Y. Yusuf)

Pomacentrus milleri
Taylor, 1964
Miller's Damselfish
D XIII–XIV, 13–14; A II, 13–14; P1
17–18; LLp 18–19. Body ovate, compressed. Posterior margin of preopercle serrate. Suborbital naked, with serrate ventral margin. Teeth on jaws

Pomacentrus cuneatus, UMTF 1341 (KAUM–I. 16567), 4.1 cm SL
Bidong Island, 14 Oct. 2008

Pomacentrus milleri, UMTF 1226 (KAUM–I. 16566), 3.7 cm SL
Bidong Island, 14 Oct. 2008

biserial, incisor-like. Caudal fin emarginate; no conspicuous projecting
spine-like rays at upper and lower
edges of caudal fin-base. Color: adults
gray, with scales dark marginally; finswith blue margins. Juvenile blue, yellow dorsally; neon-blue lines on dorsal
part of head; black spot edged withblue on rear dorsal fin. Size: 7.5 cm
SL. Distribution: distributed in western and northern Australia, and the
Gulf of Thailand. Remarks: inhabits
inshore reef areas mainly on dead coral, rubble or algal area; depth range
1–6 m.
(Y. Yusuf)

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Pomacentrus moluccensis
Bleeker, 1853
Lemon Damsel
D XIII, 14–15; A II, 14–15; P1 17;
LLp 17–18. Body ovate, compressed.
Posterior margin of preopercle serrate.
Suborbital naked, with serrate ventral
margin. Teeth on jaws biserial, incisorlike. Caudal fin emarginate; no conspicuous projecting spine-like rays at
upper and lower edges of caudal-finbase. Color: head, body and fins yellow, darker dorsally; dosal and anal fin
with narrow black margins; a minute
black spot on upper end of gill opening; subequal or slightly larger black
spot at upper pectoral fin origin; noocellus on dorsal fin. Size: 5.5 cm SL.
Distribution: eastern Indian Ocean
and western Pacific. Remarks: foundin coral reefs areas; forming grouparound live coral; to depths of 14 m.
Feeds mainly on zooplankton, smallbenthic invertebrates, and algae.
(K. Shibukawa & Y. Yusuf)

Pomacentrus moluccensis, UMTF 1007 (KAUM–I. 16568), 3.5 cm SL
Bidong Island, 14 Oct. 2008

Pomacentrus taeniometopon
Bleeker, 1852
Brackish Damsel
D XIII, 13–14; A II,13–14; P1 17–
18; LLp 17–19. Body ovate, compressed; both jaws with biserial teeth;
distinct notch between lacrimal andsuborbital; preopercular margin serrated; predorsal scales reaching above
anterior nostril; no scales on infraorbitals and suborbital. No conspicuous
projecting spine-like rays at upper andlower edges of caudal-fin base. Color:
adults light to dark gray or dark brown,
often with vertical blue streaks or
spots on scales. Juvenile dark brown,with neon-blue lines and dot rows onsides; a small black spot with blue
margin on rear part of dorsal fin; posterior portions of dorsal and anal fins,caudal peduncle, and caudal fin yellow. Size: 12.0 cm TL. Distribution:
western Pacific. Remarks: occurs
solitarily or in small groups in lagoons, harbors, coastal reefs and outerreef slopes; also found in mangrove
creeks (sometimes in freshwater),brackish lagoons and shallow reefs exposed to freshwater runoff; depth
range 0–8 m. Feeds primarily on benthic algae.
(Y. Yusuf)

Pomacentrus taeniometopon, UMTF 1027 (KAUM–I. 16677), 7.6 cm SL
Kemaman, 21 Oct. 2008

Pomacentrus taeniometopon, UMTF 1384 (KAUM–I. 16341), 2.1 cm SL
Cendering, 16 Oct. 2008

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Pomacentrus tripunctatus
Cuvier, 1830
Threespot Damsel
D XIII, 14–15; A II, 14–15; P1 17–
18; LLp 17–18. Body ovate, compressed; its depth 1.8–2.0 in SL. Posterior margin of preopercle serrated;vertral margin of infraorbital finely
serrated. Both jaws with biserial teeth.
Predorsal scales reaching above anterior nostril; no scales on infraorbitals.
No conspicuous projecting spine-likerays at upper and lower edges of caudal-fin base. Color: dark gray-brown,with a small blue margined black saddle-spot on upper caudal peduncle.
Juveniles with a blue-edged ocellus on
dorsal fin. Size: 7.5 cm SL. Distribution: Indo-West Pacific, from Sri Lanka, Andaman Sea, and Indo-AustralianArchipelago eastward to New Britain,
Solomon Islands, Santa Cruz Islands,
and Vanuatu, north to southern Japan.
Remarks: inhabits shallow bays, silty
coastal reefs and harbors with sparse
coral and algal growth; depth range
0–3 m. Solitary species; feeds mainly
on benthic algae.
(Y. Yusuf)

Pomacentrus tripunctatus, UMTF 1402 (KAUM–I. 16328), 6.4 cm SL
Cendering, 16 Sept. 2008

Pristotis obtusirostris(Günther, 1862)
Gulf Damselfish
D XIII, 12–13; A II, 12–14; P1 17–
18; LLp 19–20. Body depth 2.5–2.8 in
SL. Posterior margin of preopercle
and subopercle serrated. Both jaws
with uniserial teeth. Caudal fin forked
without filamentous lobes; no conspicuous projecting spine-like rays at
upper and lower edges of caudal-finbase. Color: body pale gray to bluishwith blue spots on each scale; a small
black spot at base of upper pectoral
rays. Size: 14.0 cm TL. Distribution:
Indo-West Pacific: Red Sea and Persian Gulf to New Guinea, northern
Australia north to Ryukyu Islands, Japan. Remarks: inhabits flat sandy or
rubble bottoms around patch reefs oflagoons and trawling grounds; depth
range 2–80 m. Adults often forming
small groups out in the open on sandy
substrate, swimming well-above bottom; juveniles often entering shallow
estuaries; rare in the area. (Y. Yusuf)

Pomacentrus tripunctatus, UMTF 1393 (KAUM–I. 16281), 4.3 cm SL
Cendering, 14 Sept. 2008

Pristotis obtusirostris, KAUM–I. 16981, 6.6 cm SL
off Terengganu (KT), 12 Dec. 2008

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LABRIDAE
Wrasses

By Mizuki Matsunuma
A diverse group of fishes varying
in body shape and size (usually 20cm); body slightly to extremely compressed. Mouth terminal, slightly to
extremely protrusible, usually withprominent lips; maxilla not exposedon cheek; teeth in jaws usually separate and caniniform, the anteriormost
1 or 2 pairs typically enlarged andoften directed forward; pharyngeal
jaws strong with pharyngeal teeth either sharp, conical, or broad and molariform. A single, long-base dorsal
fin in most species, with VIII–XXI
spines (usually less than XV) and
6–21 soft rays; anal fin with II–VI
spines (often III) and 7–18 soft rays.
Scales cycloid and highly variable insize among species; head never fully
scaly; lateral line interrupted or continuous. Color: most species withbright and intricate color patterns, including stripes, bars, spots, blotches,and ocelli. Most species change color

single dorsal fin

canines atfront part of jaws

and sex with growth, from an initial
phase (IP) of females, the latter able
to change sex into an often brilliantly
colored terminal male phase (TP).
Remarks: most common in shallow waters in a variety of habitats
such as coral reefs, rocky reefs, sand,seagrass, and algae, but rarely foundin muddy areas. Wrasses are diurnal
with diverse feeding habits, including

benthic invertebrates, fishes, coralmucus, zooplankton, ectoparasites,
and algae.
Similar families occurring in
the area: Scaridae – mouth not protrusible; teeth in jaws coalesced atbase or fused into a bony, parrot-likebeak, except a few species; lips continuous with facial skin, without an
indentation.

Cheilinus chlorourus(Bloch, 1791)
Floral Wrasse
D X, 9; A III, 8; P1 12; LLp 14–16 +7–9. Body moderately deep; dorsal
profile of head convex in front of dorsal fin, then straight to tip of snout.
Jaws prominent, anterior tip of lower
jaw even with anterior tip of upper
jaw; 2 strong canines situated anteriorly in each jaw. Lateral line interrupted below posterior portion of dorsal-fin base. Scales reaching well onto
bases of dorsal and anal fins; predorsal
scales extending forward to above
center of eye; cheek and operculumwith 2 rows of scales. Caudal fin
rounded in small fishes, in large individuals the upper and lower rays form
elongate lobes. Color: variable fromgreenish brown to deep reddish brown,
usually flecked with white; reddish
brown fins; each scale on sides usuallywith a blue or brown spot, spots extending onto dorsal and anal fins; ablackish spot basally on first 1 or 2

Cheilinus chlorourus, KAUM–I. 17154, 11.1 cm SL
off Terengganu (KR), 4 Jan. 2009

membranes of dorsal fin; head with
red spots, some joining to form lines
radiating from eye; dark brown
blotches on sides, sometimes forming
a diffuse barred pattern. Small individuals with about 4 more prominent
vertical brown bands on sides. Size:maximum length about 36 cm. Distri-bution: Indo-Pacific from the east
coast of Africa to French Polynesia,
north to southern Japan. Remarks:
feeds primarily on benthic, hardshelled invertebrates.

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Cheilinus fasciatus(Bloch, 1791)
Redbreasted Wrasse
D IX, 10; A III, 8; P1 12; LLp 14–16+ 7–9. Body moderately deep; dorsal
profile of head convex; anterior tip ofsnout forming an acute angle. Jaws
prominent; 2 strong canines situatedanteriorly in both jaws. Lateral line interrupted. Predorsal scales extending
forward to above anterior portion ofeye; cheek and operculum with 2 rowsof scales. Caudal fin rounded in juveniles; in adult with elongate upper andlower lobes. Color: body reddish
brown to blackish with 6–7 narrow
white bars; chest to posterior part ofhead orange to reddish; narrow orange-red lines extending anteriorly
and posteriorly to eye; caudal fin
white with a broad black middle bar
and black margin. Size: maximum
length about 40 cm. Distribution: Indo-Pacific. Remarks: feeds primarily
on benthic, hard-shelled invertebrates.

Cheilinus fasciatus, KAUM–I. 17051, 13.2 cm SL
off Terengganu (KT), 18 Dec. 2008

Cheilinus trilobatus
Lacepède, 1801
Tripletail Wrasse
D IX, 10; A III, 8; P1 12; LLp 15–17+ 7–9. Body moderately deep; dorsal
profile of head convex; anterior tip ofsnout forming an acute angle. Jaws
prominent; 2 strong canines situatedanteriorly in both jaws. Lateral line interrupted. Predorsal scales extending
forward to above anterior portion ofeye; cheek and operculum scaled.
Caudal fin rounded, upper and lower
corners prolonged and middle part offin protruding in large male. Color:variably pigmented from green tobrown with mottled purple and redmarkings; 4 vertical dark bars on bodyside; head with numerous small red
spots; red lines radiating from anterior
and posterior of eye; dorsal, anal, andpectoral fins yellow or green with distal red streaks; caudal fin green with ared posterior margin. Size: maximum
length about 40 cm. Distribution: Indo-West Pacific. Remarks: found oncoral reefs at depths of 1 to 20 m.

Cheilinus trilobatus, KAUM–I. 17158, 20.2 cm SL
off Terengganu (KR), 4 Jan. 2009

Epibulus insidiator, KAUM–I. 17143, 17.5 cm SL
off Terengganu (KT), 1 Jan. 200922–23. Body moderately deep. Dorsal
profile of head convex in front of dorsal
fin, a slight concavity above and before
eye. Jaws highly protrusible, lower jaw
extending posteriorly all the way to gillmembrane when mouth closed. Lateral
Epibulus insidiatorline interrupted. Predorsal scales ex(Pallas, 1770)
tending forward above eye; cheek andSlingjaw Wrasse
operculum scaled; lower jaw naked.
D IX, 10–11; A III, 8–9; P1 12; LLp Pectoral fin comparatively short (20.5–

23.3% SL); caudal slightly rounded injuveniles, upper and lower rays forming elongate lobes in large males. Color: overall brown to occasionally yellow; dorsal fin with several horizontal
dark stripes and a black spot between
first 2 spines. Size: maximum length
about 35 cm. Distribution: Indo-Pacific. Remarks: Found on coral reefs todepths of 5 to 40 m.

171

Halichoeres argus(Bloch & Schneider, 1801)
Argus Wrasse
D IX, 11–12; A III, 11–12; P1 14;
LLp 27; GR 17–19. Body slightly
elongate, compressed. Mouth terminaland small; 2 pairs of prominent canines situated anteriorly in each jaw; alarge curved canine present on eachside at rear of upper jaw; pharyngeal
teeth well developed, molariform.
Dorsal-fin origin above posterior tip of
opercular margin. Head naked. Lateralline complete. Pelvic fins relatively
short, not reaching to anus; caudal fin
moderately rounded. Color: initial
phase with dusky orange-brown stripes
following scale rows alternating withwhitish or pale interspaces; most of
scales dorsally above lateral line withblackish centers; edges of scales inabout 5–6 discrete zones on side dusky
to blackish, thus forming diffuse darkbars on body; 2 pale-edged black spots
on dorsal fin, the first on membrane
between first and second spines, and asecond larger spot on soft-rayed portion; a small pale-edged black spot on
upper caudal-fin base. Terminal phaserose red with a black-edged green spoton each scale; rows of similar spots on
dorsal, anal, and caudal fins; head rosewith irregular green bands and large
spots. Size: maximum length about 11
cm. Distribution: eastern Indian
Ocean and western Pacific. Remarks:
found in shallow coastal reefs.

Halichoeres chloropterus(Bloch, 1791)

Halichoeres argus, UMTF 1304 (KAUM–I. 16624), TP, 5.3 cm SL
Bidong Island, 13 Oct. 2008

Halichoeres argus, KAUM–I. 16517, IP, 4.8 cm SL
Cendering, 6 Oct. 2008

Halichoeres chloropterus, UMTF 1366 (KAUM–I. 16609), 10.4 cm SL
Bidong Island, 12 Oct. 2008

Pastel-green Wrasse
D IX, 10–11; A III, 10–11; P1 13–15;
LLp 27; GR 17–20. Body moderately
deep. Dorsal profile of head only
slightly convex. Mouth terminal andsmall; a single pair of prominent canines situated anteriorly in each jaw,
second pair of teeth moderately enlarged and recurved; a large curved
canine present on each side at rear ofupper jaw. Dorsal fin origin above posterior tip of opercular margin. Lateralline complete. Small patches of scales
present high on opercle, behind eye,and high on cheek. Caudal fin slightly
rounded. Color: initial phase palegreen, whitish ventrally; each scale on
upper part of body side with a small
black dot; a narrow black bar at pecto-

Halichoeres chloropterus, UMTF 1367 (KAUM–I. 16619), 4.4 cm SL
Bidong Island, 12 Oct. 2008

ral-fin base. Terminal phase pastel
green with a faint pink spot in center of
each scale in middle and posterior ofbody; head with irregular bands ofgreen and lavender-pink. Size: maxi-

mum total length about 19 cm. Distribution: the Andaman Sea and Western
Central Pacific. Remarks: foundmostly on coral reefs and surrounding
sandy areas at depths of 1–10 m.

172

Halichoeres marginatus
Rüppell, 1835
Dusky Wrasse
D IX, 13–14; A III, 12–13; P1 14–15;
LLp 27–28; GR 17–20. Body moderately deep. Jaws prominent; 2 pairs ofprominent canines situated anteriorly
in each jaw; a large curved canine
present on each side at rear of upper
jaw. Dorsal-fin origin over first to second lateral-line scales. Lateral line
complete. Head naked except for nape.
Caudal fin rounded; pelvic fins of terminal males long, reaching beyond origin of anal fin. Color: initial phasedark brown with faint pale stripes following scale rows; a small yellowedged deep blue spot at front of dorsal
fin and a large one in middle of fin;caudal fin whitish. Terminal males yellowish brown with a dark bluish spoton each scale, narrow oblique blue
bands on head; caudal fin with a green
crescent at base, followed by a large
purple-edged red crescent containing
small blue-edged green spots. Size:maximum length about 17 cm. Distribution: Indo-Pacific. Remarks: foundin coral reefs to depths of at least 30 m.

Halichoeres marginatus, UMTF 1206 (KAUM–I. 16389), 4.3 cm SL
Dungun, 24 Sept. 2008

Halichoeres melanurus, UMTF 1347 (KAUM–I. 16604), 4.4 cm SL
Bidong Island, 12 Oct. 2008

Halichoeres melanurus(Bleeker, 1851)
Tailspot Wrasse
D IX, 12; A III, 12; P1 14; LLp 27.
Body moderately deep. Jaws prominent; a single pair of prominent canines situated anteriorly in each jaw; alarge curved canine present on eachside at rear of upper jaw. Lateral line
complete. Head naked except for nape.
Pelvic fins of terminal phase reaching
to or beyond anus; caudal fin slightly
rounded. Color: initial phase with alternating narrow orange-yellow andblue stripes; a blue-edged black spot at
upper base of caudal fin; a large spot in
middle of dorsal fin, and a small spot at
front of dorsal fin. Terminal phase withblue-green and orange stripes on body;
3–6 narrow green vertical bars on upper side; head with pink and bluegreen bands; caudal fin blue withcurved orange-red bands and a large
vertically elongate dark spot posteriorly in middle of fin; pectoral-fin basewith a large yellow spot with a smaller
black spot at upper fin base. Size:maximum length about 10.5 cm. Distribution: eastern Indian Ocean and

Halichoeres nigrescens, KAUM–I. 16326, 6.7 cm SL
Cendering, 16 Sept. 2008

West Pacific. Remarks: found mostly row on body which may coalesce intoon shallow coral reefs and rocky and indistinct dark stripe; a short vertical black line behind eye preceded by ashores.
blue line and followed by a dusky pinkbar; a small triangular black spot at upHalichoeres nigrescens
per base of pectoral fins. Males light(Bloch & Schneider, 1801)
green, white ventrally, with 7 reticulate
Bubblefin Wrasse
deep pink blotches, the last 4 interconD IX, 12; A III, 11–12; P1 14–15; nected, and a midlateral series of indisLLp 27; GR 18–22. Body slightly tinct small white spot; a black and yelelongate, compressed. Dorsal-fin ori- low spot in dorsal fin between fifth andgin slightly posterior to upper end of sixth spines. Size: maximum length
gill opening. Scales on nape extending about 14 cm. Distribution: Indo-Westto or slightly anterior to a vertical at Pacific. Remarks: found in shallow
rear edge of eyes; rest of head naked. areas of reef flats, rocky shorelines as
Lateral line complete. Caudal fin well as seagrass beds.
rounded. Color: females with irregulargreen and pink stripes and 6 groups of
7–10 small black spots in a midlateral

173

Hemigymnus melapterus(Bloch, 1792)
Blackeye Thicklip
D IX, 11; A III, 11; P1 14; LLp 27–
28. Body deep. Mouth small; a pair ofprotruding canine teeth in jaws; lips
large and fleshy, the lower lips split on
the midline forming 2 lobes; a broadcanine present at each side at rear ofupper jaw. Lateral line abruptly curved
below posterior portion of dorsal finbase; scales barely reaching onto bases of dorsal and anal fins; scales infront of dorsal fin extending forward
to above center of eye; cheek with anarrow band of scales below and behind eye; opercle and lower jaw without scales. Color: white forward of aline between origins of dorsal and anal
fins, dark behind; head light blue
above, greenish below with red spots
and stripes around eyes; dorsal andanal fins with blue lines and with pinkedges; pink edges also on caudal fin.
Size: maximum length about 50 cm.
Distribution: Indo-West Pacific. Remarks: found mostly on coral reefs
and reef flats at depths of 1 to at least
20 m.

Hemigymnus melapterus, KAUM–I. 17142, 22.1 cm SL
off Terengganu (KT), 1 Jan. 2009

Iniistius trivittatus(Randall & Cornish, 2000)

Iniistius trivittatus, KAUM–I. 16945, 12.1 cm SL
off Terengganu (KT), 11 Dec. 2008

Triplebar Razorfish
D IX, 12; A III, 12; P1 12–13; LLp
19–20 +4–5; GR 14–16. Body deep,
very compressed; caudal peduncle
short. Snout long; its dorsal profile
nearly vertical; remaining head profile
strongly convex. Mouth small, horizontal; lips well developed; jaws witha pair of long curved caniniform teeth
anteriorly. Lateral line interrupted;
head naked, except for 2–3 scales on
upper part of opercle; a broad band ofsmall scales below eye. Membrane between 2nd and 3rd dorsal-fin spines
incised; gap between bases of 2nd and3rd spines larger than those of adjacent interspinous spaces; caudal fin
rounded. Color: body yellowishdusky gray, whitish ventrally; 3 vertical blackish bars on body side; an indistinct large blackish blotch on centerof caudal peduncle; a median pale
blue line on forehead. Dorsal, anal andpelvic fins pale yellow; caudal fin paleblue, yellowish basally, blackish distally, red spots forming irregular shortbars on uppermost edge. Size: maxi-

Labroides dimidiatus, UMTF 1241 (KAUM–I. 16617), 5.6 cm SL
Bidong Island, 12 Oct. 2008

mum length about 13 cm. Distribution: Western Central Pacific. Remarks: found in coastal waters ofsandy bottoms.

Labroides dimidiatus(Valenciennes, 1839)
Cleaner Wrasse
D IX, 11; A III, 10; P1 14–15; LLp 26.
Body slender. Dorsal profile of head
convex; mouth terminal, small; lips
thick, the lower strongly bilobed, the 2half separated by a u-shaped notch; asingle pair of canines anteriorly injaws, a canine a corner of mouth, andseveral rows of small teeth on side ofjaws. Head scaly except for ventral portion. Caudal fin truncate to slightly
rounded. Color: juveniles black with abright blue stripe on back and dorsallyon head. Adults light blue shading topale yellow or white anteriorly, with ablack stripe from snout through eye torear edge of caudal fin, this stripe progressively broader posteriorly. Size:maximum length about 11.5 cm. Distribution: Indo-Pacific. Remarks: inhabits coral reefs. Feeds on crustacean
ectoparasites and mucus of fishes.

174

Leptojulis cyanopleura(Bleeker, 1853)
Shoulderspot Wrasse
D IX, 11; A III, 11–12; P1 13; LLp
27; GR 19–22. Body slender; headpointed; dorsal profile of head nearlystraight, nape slightly convex; snout
relatively long; eye small. Mouth terminal, slightly oblique; front of jaws
with 2 pairs of large canines; a prominent canine tooth posteriorly on upper
jaw. Lateral line complete; body covered with large cycloid scales, headand bases of fins except for caudal fin
naked. Caudal fin varying from slightly rounded in initial phase to veryslightly double emarginate in large
terminal males. Color: initial phase
whitish with 2 orange-brown stripes
on upper half of head and body, thefirst from upper lip through eye tomidbase of caudal fin, the second, narrower, from top of head along base of
dorsal fin to tip of caudal peduncle.
Terminal males bluish gray with ablue-edged orange-yellow stripe fromfront of snout through eye to middle ofcaudal fin; a group of blue-edged
black scales forming a spot on stripe
above pectoral fins; a blue-edged
black spot middorsally on nape; caudal fin blue with 3 or 4 oblique orange-yellow bands in upper and lower
half of fin. Size: maximum length
about 13 cm. Distribution: Central
Indian Ocean to western Pacific, fromthe Gulf of Oman to Great Barrier
Reef, north to Philippines. Remarks:
aggregates usually in small group
above reefs at depths of 6–45 m.

Leptojulis lambdastigma
Randall & Ferraris, 1981
D IX, 12–13; A III, 11–12; P1 13;
LLp 27; GR 16–20. Body slender;head pointed; dorsal profile of headnearly straight, nape slightly convex;
snout relatively long; eye small.
Mouth terminal, slightly oblique; frontof jaws with 2 pairs of large canines; aprominent canine tooth posteriorly on
upper jaw. Lateral line complete; bodycovered with large cycloid scales,
head and bases of fins except for cau-

Leptojulis cyanopleura, KAUM–I. 16584, TP, 9.3 cm SL
Bidong Island, 13 Oct. 2008

Leptojulis cyanopleura, UMTF 1313 (KAUM–I. 16586), IP, 6.6 cm SL
Bidong Island, 13 Oct. 2008

Leptojulis lambdastigma, KAUM–I. 17035, 10.7 cm SL
off Terengganu (KT), 17 Dec. 2008

dal fin naked. Caudal fin rounded; firstpelvic fin ray elongate, especially inlarge male. Color: Body and headpale pink, whitish ventrally; yellow
stripe on body side extending from upper lip to caudal-fin base, via eye; alarge V-shaped black marking on
nape. Dorsal fin pale orange; upper
half of soft-rayed portion of fin paleblue, with a broad pale yellow band
submarginally; lower portion of dorsal
fin with a row of yellow blotches witha pale blue margin on membranes anteriorly, forming a zigzag stripe poste-

riorly. Anal fin pale orange, dusky yellow distally, with a thin light blue
margin; a row of yellow spots alongbase. Pectoral and pelvic fins translucent. Caudal fin white, pale blue distally, blackish marginally, numerous
scattered yellow spots. Size: maximum length about 14 cm. Distribution: Western Central Pacific including Taiwan, Philippines, and Gulf ofThailand. Remarks: found in coastal
waters of sand bottoms.

175

Oxycheilinus digramma(Lacepède, 1801)
Cheeklined Maori Wrasse
D IX, 10; A III, 8; P1 12; LLp 14–16+ 7–9. Body moderately slender; dorsal
profile of head nearly straight. Jaws
prominent; 2 strong canines situatedanteriorly in each jaw. Lateral line interrupte; predorsal scales 6, reaching
forward to vertical at center of eye;cheek scales reaching just anterior to
forward extent of orbit; subopercularscales extending anteriorly to verticalat center of eye. Pelvic fins short, notreaching anus; caudal fin slightly
rounded to truncate. Color: variably
colored; body olive-green to graybrown fading to orange red on anteroventral surface, with an orange-red baror spot on each scale; head with irregular orange lines on upper part parallel todorsal profile, radiating from eye, and aseries of about 8 diagonal purple lines
on lower cheek; large central portion of
caudal fin mainly green. Size: maximum length about 40 cm. Distribution: Indo-Pacific. Remarks: found oncoral reefs at depths of 3–30 m.

Oxycheilinus digramma, KAUM–I. 17055, 16.2 cm SL
off Terengganu (KT), 19 Dec. 2008

Thalassoma lunare, UMTF 1243 (KAUM–I. 16644), 9.9 cm SL
Bidong Island, 12 Oct. 2008

Thalassoma lunare(Linnaeus, 1758)
Moon Wrasse
D VIII, 13; A III, 11; P1 15; LLp 25;
GR 18–20. Body moderately slender;
dorsal profile of head slightly convex.
Mouth small, 2 prominent canines anteriorly in each jaw. Lateral line complete; head naked except for a patch ofscales on upper end of opercle. Caudal
fin varying from truncate in young tohighly lunate with upper and lower
corners produced into filaments inlarge adults. Color: initial phase green
with vertical red lines on scales; headgreen with many irregular rose pinkbands; a blue-edged rose pink band ineach lobe of caudal fin, the broad central and posterior part of fin yellow;
pectoral fins blue with a broad pinkband in upper central part. Terminal
males similar but more blue overall.
Juveniles olive green on upper half ofbody, blue-white below; a large black
spot in middle of dorsal fin and a large
diffuse blackish spot at caudal-finbase. Size: maximum length about 25
cm. Distribution: Indo-Pacific. Remarks: found on coral and rocky reefs
at depths of 1–20 m.

Xiphocheilus typus, KAUM–I. 16939, 11.8 cm SL
off Terengganu (KT), 11 Dec. 2008

Xiphocheilus typus
Bleeker, 1856
Bluebanded Wrasse
D XII, 8; A III, 10; P1 15–17; LLp 29.
Body slender; dorsal profile of head
convex, head and snout blunt. Jaws
prominent; upper jaw with 2 widely
separated prominent anterior canines; 1
or 2 pair large curved canines on upper
jaw posteriorly; lower jaw also with 2large anterior canines. Pelvic fin short,reaching anus only in large specimens;caudal fin slightly rounded to double
emarginate in adults. Lateral line complete; bases of dorsal and anal fins naked; 7–10 large predorsal scales, extending in front of dorsal fin to centerof eye; cheek and opercle scaled. Color: head and body olive-green dorsally,orange or pinkish laterally; sides withnumerous narrow oblique blue bands;
head with 4 or 5 narrow blue bands or
stripes outlined with yellow-orange;
dorsal fin blue with 2 or 3 narrow orange or pink stripes; anal fin yelloworange with numerous narrow blue
bands basally; caudal fin yellow-orange with about 5–8 narrow blue
bands; pectoral fins transparent toorangish with a blue band on base.
Size: maximum length about 12 cm.
Distribution: eastern Indian Ocean
and western Pacific. Remarks: inhabits coral reefs at depths of 35 to 85 m.
Caught mostly with bottom trawls.

176

SCARIDAE
Parrotfishes

By Mizuki Matsunuma
Medium to large sized (up to
120 cm) fishes. Body oblong, moderately deep. Head generally bluntly
rounded anteriorly. Teeth in jaws
usually fused to form a pair of beaklike plates in each jaw, but a few
species have free, imbricate, incisorlike teeth; small, isolated, projecting teeth (canines) occur in somespecies on outer sides of upper jaw;
pharyngeal dentition strong consisting of interdigitating paired upper
pharyngeals with rows of elongate
molariform teeth; these teeth bearagainst the elongate molariform teethon the surface of the single lower
pharyngeal bone. Dorsal fin with IX
slender, flexible spines and 10 softrays; anal fin with III flexible spinesand 9 soft rays; pectoral fins with 2
unbranched rays and 11–14 branched
rays; pelvic fins with I spine and 5
soft rays; caudal fin with 11 branchedrays, varying from rounded to lunate with produced caudal fin lobes.
Scales large, cycloid, usually 22–24
on lateral line; 1–4 rows of scales oncheek; 2–8 median predorsal scales.
Color: most species very colorful;

single dorsal finlarge scales on bodybeak-like toothplates on jaws

may exhibit striking sexual dichromatism. Most species are protogynous
hermaphrodites (maturing first as
females, then sexually transforming
into males), this sexual transformation
is usually accompanied by change incolor phase. In species where 2 adultcolor phases are known the first is
termed "initial phase" (IP), the second"terminal phase" (TP).
Remarks: parrotfishes are mostoften found on or in vicinity of coral
reefs, and usually most abundant inshallow waters to a depth of 30 m.
Herbivorous, usually scraping algae

from dead coral substrates. Bits ofrock eaten with the algae are crushed
into sand and ground with the algae
to aid in digestion, making parrotfish
some of the most important producers of sand on coral reefs.
Similar families occurring in
the area: the beak-like plates ofmost scarids, in addition to features
such as large smooth scales and often bright colors, usually preclude
parrotfishes being confused with anyother fish family.

Hipposcarus longiceps(Valenciennes, 1840)
Pacific Longnose Parrotfish
D IX, 10; A III, 9; P1 14–16 (usually15); PDS 4. Head profile pointed with
distinctly angular snout; eye near dorsal profile; dental plates relatively narrow with cement; terminal male with 1
or 2 canine teeth on upper dental plate;
3 rows of small scales on cheek, upperrow 6–8 scales, middle row 5–7 scales,
lower row 3–6 scales, these in small
triangular patch on cheek. Caudal fin
strongly double emarginate with very
short lobes. Color: initial phase paleyellowish gray to nearly white, edgesof scales darker gray; margin of dorsal
and anal fins pale blue; caudal peduncle and fin yellowish. Terminal male
pale blue-green, scales edged with orange to pink; edge of upper lip narrowly bright orange to pink; dental
plates white; dorsal and anal fins

Hipposcarus longiceps, KAUM–I. 17045, 17.4 cm SL
off Terengganu (KT), 17 Dec. 2008

salmon pink with a blue streak on eachmembrane and a blue margin. Juveniles light gray, with an orange lateral
stripe extending from tip of snout to a
black spot at caudal-fin base. Size:maximum length 40 cm; commonly
20–30 cm. Distribution: West Pacificfrom the Gulf of Thailand east to Tuamotu Archipelago, northern Australia
north to southern Japan; northwestern
Australia in the Indian Ocean. Remarks: relatively common over sandnear seaward reefs or in lagoons.

177

Scarus ghobban
Forsskål, 1775
Yellowscale Parrotfish
D IX, 10; A III, 9; P1 15–16 (usually15); PDS 5–7 (6). Teeth fused to form
dental plates; dental plates relatively
smooth; lips covering more than halfof dental plates; large adults with 1–3
canine teeth on side at rear of upper
dental plate; usually 3 rows of scales
on cheek; caudal fin slightly emarginate in small initial-phase fish to lunate
in large terminal males. Color: initial
phase dull orange yellow, centers ofscales bluish; 5 irregular blue bars often present on body; fins yellowish.
Terminal males green dorsally, scales
rimmed with salmon pink, shading topale green ventrally with pale salmon
pink bar on each scale; head green
dorsally, shading to pale salmon on
cheek and chin, with 2 transverse blue
bands on chin and 3 narrow irregulargreen bands extending posteriorly
from eye. Size: maximum length
about 75 cm. Distribution: widespread Indo-Pacific and tropical eastern Pacific. Remarks: found on coral
reefs to at least 30 m depth.

Scarus ghobban, KAUM–I. 16978, 20.3 cm SL
off Terengganu (KT), 12 Dec. 2008

Scarus psittacus, KAUM–I. 17152, 13.6 cm SL
Redang Island, 4 Jan. 2009

Scarus psittacus
Forsskål, 1775
Common Parrotfish
D IX, 10; A III, 9; P1 13–15 (usually14); PDS 3–5 (4). Teeth fused to form
dental plates; dental plates relatively
smooth; lips largely covering dental
plates; usually1 conical tooth on sideof upper dental plate of initial phase,
additionally 1 conical tooth on lower
plate of terminal males; 2 rows ofscales on cheek; caudal fin slightly
emarginate in initial phase fish to
deeply emarginate terminal males.
Color: initial phase reddish brown togray, snout often paler than rest ofhead, reddish ventrally; a dark spot atbase of first membrane of dorsal fin,and a small black and blue spot at upper base of pectoral fin; median finscolored like body. Terminal malegreen posteriorly, edges of scales pink;
abdomen pink with longitudinal seriesof green spots following scale rows;

Scarus psittacus, UMTF 1309 (KAUM–I. 16588), 5.3 cm SL
Bidong Island, 12 Oct. 2008

head with a green band on edge oflips, joining at corner of mouth andcontinuing below eye; 2 additional
green bands extending posterior toeye; snout to a vertical above posterioredge of eye dark purplish to lavender

gray; dental plates white. Size: maximum length about 26 cm; commonly
16–22 cm. Distribution: widespread
Indo-Pacific. Remarks: commonly
found in coral or rocky reefs; often inlarge mixed species groups.

178

Scarus quoyi
Valenciennes, 1840
Quoy's Parrotfish
D IX, 10; A III, 9; P1 14; PDS 6.
Teeth fused to form dental plates; dental plates relatively smooth; lips largely covering dental plates; 3 rows ofscales on cheek, usually 2 scales inlower row; caudal fin of adults truncate to slightly emarginate. Color: initial phase light grayish brown withfive or six faint bars from scales withpaler centers; three longitudinal whitish stripes on abdomen. Terminal
males blue-green dorsally, the scaleswith narrow pink edges, becoming violet-pink or magenta below; head violet-gray dorsally; a large irregular
patch of bright blue green on cheekwhich extends as a broad band onto
upper lip and continues onto chinwhere invaded by irregular pink

Scarus quoyi, KAUM–I. 17139, 18.5 cm SL
off Terengganu (KT), 1 Jan. 2009

bands; operculum orange dorsally,shading to violet-pink ventrally; caudal and pectoral fins largely bright
deep blue. Size: maximum length
about 30 cm, commonly 20–25 cm.

Distribution: eastern Indian Ocean
and West Pacific. Remark: inhabits
coral-rich areas of outer channels andseaward reefs.

179

CHAMPSODONTIDAE
GapersBy Mizuki Matsunuma
Small perciform fishes (up to
about 15 cm). Body elongate, slightly
compressed. Eyes large, usually witha small posterodorsal cirrus. Mouth
large, oblique; maxilla extending
posteriorly to below eye or beyond.
Premaxilla and dentary with at least
2 rows of teeth; inner row of long,depressible, needle-like teeth; outerrow of shorter fixed teeth; teeth onvomer but not on palatines. Lacrimal
with 2 spines projecting anteroventrally over maxilla and premaxilla.
Preopercle with prominent, posteroventral spine. Gill opening large; firstgill arch with 1 or 2 gill rakers on upper limb, 10–14 on lower limb. Branchiostegals 7. Two dorsal fins; firstdorsal fin with IV–VI spines; second
dorsal fin with 18–23 soft rays. Anal
fin spineless, similar in length andshape to second dorsal fin, with 16–
21 soft rays. Caudal fin forked, with
15 principal rays and 14–18 procurrent rays. Pectoral fin small, set highon body, with 12–16 rays. Pelvic fin
much larger than pectoral fin, with I
spine and 5 soft rays. Body covered
with small, rough, non-overlapping
scales, with 2–9 spinules projectingfrom posterior margin of broad plate.

eye cirrus

lateral-line system

lacrimal spines
preopercular spine

Two horizontal lateral lines composedof numerous small papillae, extending from opercular margin onto caudal
fin; numerous (20–25) transverse rowsof these sensory papillae. Vertebrae
10–12 + 19–21 = 29–33. Color: bodybrownish dorsally; lateral and ventral
parts tan or silvery, with dark spots
along midline in some species; distal
third of first dorsal fin dark brown
to black or, speckled ; upper or bothcaudal-fin lobes dark brown in many
species; dark blotch at caudal fin base.
Remarks: captured in depths of
34 to 1100 m. They sometimes occur in great shoals, evidently rising
to the surface at night, and are quite

commonly cast ashore during storms.
None of the species is fished commercially, but they are often taken
along with commercial species; they
may also be important prey species
for scombrids and other pelagic predators.
Similar families occurring in
the area: none. Gapers are easily
distinguished from other fishes in thearea by having rough, non-overlapping scales; large preopercular spine;
2 lacrimal spines projecting anteriorly over maxilla; pelvic fin, longer
than and inserting in front of pectoral fin; large mouth with depressible
teeth; and 2 horizontal lateral lines.

Champsodon vorax
Günther, 1867
D IV–V + 20–21; A 17–19; P1 12–
14; GR 1 + 10–12; V 12 + 19–20 =31–32. Body elongate, slightly compressed; snout short, its length equal toeye diameter; maxilla extending beyond posterior margin of eye; premaxilla notched lateral to symphysis.
Transverse row of 4 sensory papillae
between posterior margins of pteroticridges. Chin and breast scaled; triangular patch of scales between pectoraland pelvic fin bases; belly partially or
fully scaled. Color: body dark brown
dorsally, pale laterally; distal tip offirst dorsal fin speckled thickly withmelanophores; upper caudal fin lobe

Champsodon vorax, KAUM–I. 17089, 4.4 cm SL
off Terengganu (KT), 28 Dec. 2008

occasionally with a few melanophores; other fins pale; peritoneum
pale or silvery, spotted with distinct
melanophores. Size: maximum length
about 8 cm. Distribution: known

from the South China Sea, northwestern Australia, and Maldives. Remarks: usually caught with bottom
trawls in depths of 34–84 m.

180

PINGUIPEDIDAE
Sandperches

By Hisashi Imamura
Small to large-sized fishes. Body
elongate, cylindrical anteriorly (except Prolatilus, having rather deeperbody). Eye moderate. Snout slender.
Teeth small, in band in jaws. Vomer
with V-shaped tooth patch (teeth
absent in Prolatilus). Palatine teeth
present or absent. Gill membranes
united, free from isthmus. Dorsal
fin single, with 4–7 short spines and
19–27 soft rays. Anal fin with 17–25
rays, including 1 or 2 weak spines
anteriorly. Caudal fin usually with
15 branched rays. Pelvic fin with
1 spine and 5 soft rays. Vertebrae
30–37. Several species of Parapercis
have been shown to be protogynous
hermaphrodites (thus females change
to males). Color: variable (e.g., dark,
pale brownish, and reddish), many

single dorsal fin with IV–VII, 19–27

anal fin with 17–25 rays, including I or II spines

species with distinct and characteristic markings; some change in the color
pattern with the sex change is usuallyknown in Parapercis.
Remarks: five genera are known

from the world; only Parapercis is
distributed in the area.
Similar family occurring in thearea: Percophidae – 2 dorsal finspresent.

Parapercis filamentosa(Steindachner, 1878)
Threadfin Sandperch
D V, 22; A I, 18; P1 16; LLp 58–59.
Body cylindrical. Three pairs of canine teeth on lower jaw anteriorly.
Palatine teeth absent. Middle dorsal
spines longest. Membrane from spinous dorsal connected by membrane
to 1st soft ray near base. First few dorsal soft rays greatly elongated. Color:body light brown dorsally with 6 faintV-shaped brown markings, whitish
ventrally; pelvic fin dusky; upper caudal fin base with a dark spot. Size:maximum total length about 18 cm.
Distribution: West Pacific, including
Singapore, Thailand, Hainan, Java andBorneo, Malaysia.
Parapercis filamentosa, KAUM–I. 16936, 11.9 cm SL
off Terengganu (KT), 11 Dec. 2008

181

URANOSCOPIDAE
Stargazers

By Mizuki Matsunuma
Medium sized (up to 65 cm)
marine fishes. Body moderately
elongate, depressed anteriorly orcompressed. Head massive, nearlycube-shaped, flattened dorsally,rounded anteriorly; dorsal and lateral surfaces of head almost entirely
encased in sculptured bones. Eyes
directed dorsally or dorsolaterally,
placed on or near top of head. Infraorbital bones dilated. Interorbital
space noticeably broad, anterodorsal
part of skull deeply scooped backward (interorbital fossa). Mouth
large, protractile, almost vertical; jaws, prevomer, and palatines
toothed; a pair of pockets on anteriorroof of mouth (between premaxillae
and prevomer). A peculiar dermal
appendage present at central tip of
respiratory valve inside lower jaw
in Uranoscopus and young of Genyagnus. Branchiostegals 6. First gillarch with gill teeth instead of typicalgill rakers; pseudobranchiae present.
Membranes of anal, pelvic, and pectoral fins fleshy and thickened. Dorsal fin single with 0–IV rudimentaryspines and 12–20 soft rays or divided
with IV or V weak spines and 13 orlateral line reaches torear end of caudal finmouth almost
verticalpelvic fins thoracic

14 soft rays; anal fin with 13–19 softrays; caudal fin subtruncate, with 10 or
11 branched rays; pectoral fin broad,“squarish”, or knife-shaped, with
13–25 rays; pelvic fin close together,situated on throat and in advance of
pectoral fins, with I non-visible weakspine and 5 segmented rays. Body naked or covered with adherent, cycloid
scales almost embedded under theskin; lateral line complete, reaching
onto caudal fin margin. Color: generally brownish dark with dark or palespots or mottling on body.
Remarks: stargazers are noctur-

nal and during the day bury theirbody into sand or mud, with only theeyes and mouth cleft protruding fromthe substrate. Caught mostly withbottom trawls.
Similar families occurring in
the area: Batrachoididae – gill openings small, restricted to sides of body(gill openings wide in Uranoscopidae); pelvic fins with I distinct spine
and 5 soft rays (with I non-visible
weak spine and 5 soft rays in Uranoscopidae). Champsodontidae – bodysmaller, more compressed; caudal fin
forked.

Uranoscopus cognatus
Cantor, 1849
Twospined Yellowtail Stargazer
D III + 14–15; A 13–14; P1 16–17.
Body moderately elongate; head massive; short cirri on eye; respiratory
valve inside lower jaw with a threadlike appendage. Preopercle with 4
spines on lower edge; supracleithrum
not forming distinct spine at rear end;
2 basipterygial processes. Color: headand body brown dorsally, finely mottled and stippled dark brown; dusky
white laterally, white ventrally. First
dorsal fin black; second dorsal fintranslucent, rays tinged with brownish; pectoral fin translucent; posterior
half of fin yellow with white margin;

Uranoscopus cognatus, KAUM–I. 16887, 6.5 cm SL
off Terengganu (KT), 6 Dec. 2008

pelvic fin white; caudal fin pale yel- south to the Philippines. Remarks:
low, dusky distally. Size: maximum caught mostly with bottom trawls in
length about 22 cm. Distribution: depths of 50–250 m.
eastern Indian Ocean and West Pacificfrom India east to northern Australia,

182

TRIPTERYGIIDAE
Triplefins

By Hisashi Imamura
Small to moderate-sized marine
fishes (maximum size about 26 cm,
3 dorsal fins with III–X + VII–XXVI + 7–17
most species less than 6 cm). Body
slender. Eye large. Jaws with bandsof conical teeth, broadest anteriorly.
Gill membranes broadly attachedacross isthmus. Dorsal fin divided
into 3 distinct segments, 1st with
III–X spines, 2nd with VII–XXVI
spines and 3rd with 7–17 soft rays.
Anal fin with 0–II (rarely IV) spines
anal fin usually 0–II, 14–31
and 14–31 soft rays. Caudal fin with
branched rays. Pelvic fin with I spineand 2–3 soft rays. Single lateral line. ally color pattern from females.
area: Blenniidae – body always naScales usually ctenoid (1 species ofRemarks: most occurring in shal- ked and dorsal fin comprised of sinLepidoblennius with cycloid). Col- low water, often on reef flats or in gle fin. Clinidae – dorsal fin consistor: highly variable, frequently with tidepools.
ing of 1 or 2 fins and caudal-fin rays
bars or a mottled pattern; males usuSimilar family occurring in the unbranched. Gobiidae – 2 dorsal fins.

Enneapterygius tutuilae
Jordan & Seale, 1906
High-hat Triplefin
D III + IX–XIII + 7–10; A I, 15–20;
P1 13–18; LLp 7–13 + 18–25 (interrupted); MP 2–4 + 2 + 2–4. Body slender. Supraorbital region with lobate
tentacle. First dorsal fin high, 1st spine
higher than 1st and 2nd spines of 2nd
dorsal fin. Abdomen lacking scales.
Color: variable, often translucent
greenish with faint, irregular narrowbars from red to brown pigments onscale edges; single black blotch present on middle of 2nd dorsal fin inmales, absent in females; anal-fin basewith 6–7 dark blotches; pelvic fin
black near base in males, lacking
black mark in females. Size: maximum size about 2.8 cm. Distribution:
Indo-West Pacific, ranging from Madagascar to southern Japan and to New
Caledonia. Remarks: known fromtidepools and shallow rocky reefs.

Helcogramma striata
Hansen, 1986
Neon Triplefin
D III + XII–XV + 9–12; A II, 17–23;
P1 15; P2 I, 2; LLp 14–20; MP 3 + 2 +3. Body slender. Supraorbital region
lacking tentacles. Lateral line ending
under 10th–17th dorsal fin elements.
Head, abdomen and pectoral-fin baseEnneapterygius tutuilae, UMTF 1283 (KAUM–I. 16551), 1.6 cm SL
Bidong Island, 13 Oct. 2008

Helcogramma striata, UMTF 1282 (KAUM–I. 16553), 2.3 cm SL
Bidong Island, 13 Oct. 2008

naked. Color: body and head red dorsally, green to blue ventrally; 3 longitudinal blue stripes on each side ofbody running from snout to caudal
peduncle; a row of 6–7 blue spots
present just below to lateral line andon caudal peduncle; dorsal fins usually red, each with blue longitudinal

stripe near base. Size: attaining to 5.7
cm. Distribution: Indo-West Pacific,
including Sri Lanka, Andaman Sea,
southern Japan, Indonesia, northern
Australia and Solomon Islands. Remarks: present species lacks sexual
dimorphism.

183

BLENNIIDAE

Blennies (Combtooth Blennies, Sabertooth Blennies)
By Koichi Shibukawa and Hisashi Imamura
Small, scaleless, often elongate
fishes (most species much less than
15 cm). Head usually blunt, oftenwith cirri on eyes, nasal openings,
nape, and/or cheeks; gill openings
continuous across lower surfaceof head or restricted to small opening on each side of head. Upper jaw
not protractile; teeth incisor-like insingle row in each jaw, often veryfine and loosely attached; canine
teeth occasionally present; no teeth
in palatines; vomerine teeth present or absent. Dorsal fin continuous
or notched, usually with more segmented rays than spines (III–XVII
flexible spines and 9–119 segmented
soft rays); anal fin with II spines,
often indistinguishable; segmented
caudal fin rays 10–14; pelvic fins inadvance of pectoral fins, pelvic fin
spine not visible externally, soft rays
fewer than 5, sometimes deformed
or absent. Color: very variable, dull
to brilliant, full spectrum; often mottled, with irregular vertical bands or
stripes, some species almost uniform.

single dorsal fin

body with no scales

pelvic fins anterior
to pectoral fins

2 reduced spines in anal fin

Remarks: mostly bottom dwelling species in sea and estuaries, usually at depths much less than 20 m,
mostly among rocky, oyster, or coral
reefs, often in tide pools. Feeding on amixed diet of algae and benthic invertebrates; some are planktivores, andsome are specialized to feed on skinor fins of larger fishes, with mimic as
cleaner.
Similar families occurring in
the area: Clinidae – body with fine,embedded cycloid scales; scales withradii in all fields; many more dorsal

fin spines than segmented rays; jaw
teeth in more than 1 row. Tripterygidae – body with ctenoid scales; dorsal fin clearly divided into 3 parts;
many more dorsal fin spines thanrays; jaw teeth in more than 1 row.
Gobiidae and Eleotridae – body usually scaly; pelvic fins usually with I
spine and 5 soft rays; dorsal fin in 2
well-separated sections, the spinous
portion with less than VIII spines, thesegmented ray portion with I spine at
beginning.

Blenniella bilitonensis(Bleeker, 1858)
D XII–XIV, 19–22; A II, 18–22; P1
13–15; P2 I, 3. Body elongate andcompressed. Simple cirrus present oneye; nasal cirri branched; cirri absent
on nape and cheek. Blade-like fleshy
crest on top head in male. Gill opening
wide, continuous across ventral surface of head. Dorsal fin continuous,with a distinct notch between spinousand segmented-ray portions. Caudal
fin usually with 13 segmented rays,
several of them branched. Color: 4–9
faint dusky longitudinal lines on bodyin male, in addition to numerous whitish spots on posterior part of body
(both absent in female); several diagonal dusky lines at basal half of segmented-ray portion of dorsal fin inmale (absent in female); dark spots on
distal half of pectoral fin. Size: 9 cm.
Distribution: West Pacific. Remarks:
inhabits coastal shallow reefs; common
in tidepools.
(K. Shibukawa)

Blenniella bilitonensis, UMTF 1159 (KAUM–I. 16394), female, 4.9 cm SL
Dungun, 24 Sept. 2008

Blenniella bilitonensis, UMTF 1328 (KAUM–I. 16690), male, 4.5 cm SL
Kemaman, 21 Oct. 2008

184

Ecsenius yaeyamaensis(Aoyagi, 1954)
Yaeyama Blenny
D XII–XIII, 13–15; A II, 15–17; P1
12–14; P2 I, 3. Body elongate andcompressed. No cirri on eye, nape andcheek; nasal cirri simple. Fleshy crest
absent on top of head. Gill opening
wide, continuous across ventral surface of head. Dorsal fin continuous,with a distinct notch between spinousand segmented-ray portions. Caudal
fin usually with 13 segmented rays, allof them unbranched. Color: groundcolor of head and body pale yellowish
gray; 1-3 longitudinal series of blackspots and/or dashes behind eye; arclike black line from lower jaw to ventral part of operculum; pectoral-finbase with a forked black line; usually
many faint pale spots on posterior partof body. Size: 5 cm. Distribution: Indo-West Pacific. Remarks: inhabits
coral reefs.
(K. Shibukawa)

Ecsenius yaeyamaensis, UMTF 1330 (KAUM–I. 16598), 3.6 cm SL
Bidong Island, 12 Oct. 2008

Entomacrodus lighti, KAUM–I. 16373, 5.6 cm SL
Dungun, 23 Sept. 2008

Entomacrodus lighti(Herre, 1938)
D XIII, 15–16; A II, 17–18; P1 14; P2
I, 4. Body elongate and compressed.
Cirrus on eye typically simple; nasal
cirri irregular in size and shape; cirrus
on nape occasionally with frayededge; cirri absent on cheek. Fleshy
crest on top head absent in both sexes.
Middle third of hind margin of upper
lip crenulate. Gill opening wide, continuous across ventral surface of head.
Dorsal fin continuous, with a distinct
notch between spinous and segmented-ray portions. Caudal fin usuallywith 13 branched rays, several of thembranched. Color: body with diffused
dusky banded pattern, without finepale speckles; a dark spot between
first two spines of dorsal fin. Size: 10
cm. Distribution: West Pacific. Remarks: found in shallow rocky shore.
(K. Shibukawa)

Entomacrodus striatus, KAUM–I. 16382, 7.8 cm SL
Dungun, 23 Sept. 2008

Entomacrodus striatus, KAUM–I. 16370, 5.6 cm SL
Dungun, 23 Sept. 2008

cirri absent on cheek. Fleshy crest on
top head absent in both sexes. Hind
margin of upper lip entirely crenulate.
Reef Margin Blenny
Gill opening wide, continuous across
D XIII, 14–17; A II, 15–18; P1 14; P2 ventral surface of head. Dorsal fin
I, 4. Body elongate and compressed. continuous, with a distinct notch beCirrus on eye typically branched, with tween spinous and segmented-ray por1–20; nape and nasal cirri present; tions. Caudal fin usually with 13

Entomacrodus striatus(Valenciennes, 1836)

branched rays, several of thembranched. Color: body pale brown,with several patches of small black
dots; head with numerous, dense pale
speckles. Size: 12 cm. Distribution:
Indo-Pacific. Remarks: found in shallow rocky shore; common in intertidal
zone.
(K. Shibukawa)

185

Istiblennius dussumieri(Valenciennes, 1836)
Dussumier's Rockskipper
D XII–XIV, 19–24; A II, 21–25; P1
13–15. Body elongate. Nape lacking
cirri. Supraorbital tentacle variable,with up to 5 branches on each side.
Single blade-like occipital crest present in adult males. Nasal with short
cirri. Canine teeth absent on dentary.
Several caudal-fin rays branched. Lateral line ending below 4th to 10th dorsal spines. Color: body dusky graywith a series of 7 double, brownish
orange bars; small dark spots scatteredon body in females; outer part of median fins broadly dark in males. Size:attaining to about 13 cm. Distribution: Indo- West Pacific, ranging fromeast coast of Africa to Taiwan and toFiji.
(H. Imamura)

Istiblennius edentulus(Forster & Schneider, 1801)
Rippled Rockskipper
D XII–XIV, 18–23; A II, 20–24; P1
13–14. Body elongate. Nape with amoderately long cirrus on each side.
Supraorbital tentacle usually a longsimple filament. Single blade-like occipital crest present in adult males.
Nasal with 3 cirri. Canine teeth absent
on dentary. Several caudal-fin rays
branched. Lateral line without verticalpairs of pores, ending below last dorsal spine and 15th soft ray. Color:body pale grayish with dark gray double bars extending basally into dorsal
fin; irregular orangish lines in pale interspaces on posterior half of body; apale-edged gray bar from eye acrossupper lip; an oblique, pale-edged grayband behind eye; posterior body anddorsal and anal fins with orangish
brown spots in females. Size: attaining to 17 cm. Distribution: widespread in Indo-Pacific, but no records
from Hawaiian Islands. Remarks:
typically found along the edge ofrocky shores where wave action is notsevere.
(H. Imamura)

Istiblennius dussumieri, UMTF 1099 (KAUM–I. 16329), female, 7.3 cm SL
Cendering, 16 Sept. 2008

Istiblennius dussumieri, UMTF 1098 (KAUM–I. 16336), male, 3.4 cm SL
Cendering, 16 Sept. 2008

Istiblennius edentulus, UMTF 1287 (KAUM–I. 16406), female, 10.1 cm SL
Dungun, 24 Sept. 2008

Istiblennius edentulus, KAUM–I. 16408, male, 8.9 cm SL
Dungun, 24 Sept. 2008

186

Istiblennius lineatus(Valenciennes, 1836)
Lined Rockskipper
D XII–XIV, 21–25; A II, 22–25; P1
13–15. Body elongate. Nape lacking
cirri. Supraorbital tentacle usually atriangular filament with short medial
and lateral branches in adults. Single
large, blade-like occipital crest present
in adult males. Nasal cirri short andpalmate. Canine teeth absent on dentary. Several caudal-fin rays branched.
Lateral line without vertical pairs ofpores, ending below space between
6th dorsal spine and 5th soft ray. Color: body pale brown with several longitudinal, dark brown lines; headcrossed with vertical, irregular, orangish brown lines. Size: attaining to 14
cm. Distribution: widespread in Indo-Pacific, including Maldive Islands,
southern Japan and Tuamotu Archipelago. Remarks: found in rocky
shores of coral reefs and tidepools.
(H. Imamura)

Istiblennius lineatus, KAUM–I. 16343, female, 8.5 cm SL
Cendering, 16 Sept. 2008

Istiblennius lineatus, UMTF 1093 (KAUM–I. 16407), male, 7.9 cm SL
Dungun, 24 Sept. 2008

Laiphognathus multimaculatus
Smith, 1955
Many-spotted Blenny
D X–XII, 18–21; A II, 19–22; P1
12–14; P2 I, 3; V 35–39. Body elongate. Nape lacking cirri. Cirri on bothanterior and posterior nostrils. Gill
opening small, gill membranes onboth sides separated. Caudal fin lacking branched rays. Color: body paleyellow with many dusky golden spots,
abdomen pinkish; spinous dorsal fin
with a narrow longitudinal black band;
distal half of soft dorsal fin speckledwith black; anal fin and margin of caudal fin dusky; pectoral and pelvic finmembranes transparent. Size: attaining to 5.5 cm. Distribution: widespread in Indo-Pacific, including east
coast of Africa, Andaman Sea, andSolomon Islands.
(H. Imamura)

Laiphognathus multimaculatus, UMTF 1404 (KAUM–I. 16627), 2.5 cm SL
Bidong Island, 13 Oct. 2008

Omobranchus elongatus, UMTF 1416 (KAUM–I. 16543), female, 3.5 cm SL
Cendering, 6 Oct. 2008

Omobranchus elongatus(Peters, 1855)
Cloister Blenny
D XI–XIII (usually XII), 18–21; A
II, 20–23; P1 13; P2 I, 2. Body elongate. No cirri or tentacles and crest on
head. Single canine tooth present on
both jaws posteriorly. Edge of lips
smooth. Lower lip with a small, posterior, ventral-projecting flap. Caudal
fin lacking branched rays. Color:

Omobranchus elongatus, UMTF 1317 (KAUM–I. 16536), male, 4.0 cm SL
Cendering, 6 Oct. 2008

body reddish gray with curved andoblique gray bars and white light red
elliptical spot on opercle; small dark
spots sometimes present on ventral
part of head and on chest; a dusky spot
in dorsal fin between 9th and 13th softrays in males. Size: attaining to 6.6
cm. Distribution: widespread in Indo-West Pacific, including east coastof Africa, Andaman Sea and Iriomote
Island. Remarks: found in rocky reefs
of estuaries.
(H. Imamura)

187

Omobranchus ferox(Herre, 1927)
D XI–XIII, 20–23; A II, 20–26; P1
13; P2 I, 2. Body elongate and compressed. No cirri on head. Fleshy creston top head absent in both sexes. Gill
opening narrow, restricted on side ofhead. Dorsal fin continuous, without
distinct notch between spinous andsegmented-ray portions. Caudal fin
usually with 13 segmented rays, all ofthem unbranched. Color: head andbody olive green, typically with several vague dusky bars on body; a short,narrow vertical black band just behindeye (preceded by white vertical line
when alive or fresh); a dusky spot frequently found around end of dorsal fin
in male. Size: 6 cm. Distribution: Indo-West Pacific. Remarks: found in
mangrove swamps and estuaries.
(K. Shibukawa)

Omobranchus ferox, UMTF 1430 (KAUM–I. 16808), 3.9 cm SL
Merang, 2 Nov. 2008

Omobranchus punctatus, KAUM–I. 16327, 6.4 cm SL
Cendering, 16 Sept. 2008

Omobranchus punctatus(Valenciennes, 1836)
Muzzled Blenny
D XI–XIII, 19–24; A II, 22–26; P1
13; P2 I, 2. Body elongate and compressed. No cirri on head. Fleshy creston top head absent in both sexes. Gill
opening narrow, restricted on side ofhead. Dorsal fin continuous, without
distinct notch between spinous andsegmented-ray portions. Caudal fin
usually with 13 segmented rays, all ofthem unbranched. Color: ground color of head and body brownish olive;
head with broad dusky vertical bands;
body with several irregular, horizontal
dusky lines anteriorly. Size: 9 cm.
Distribution: Indo-Pacific; also introduced to Western Atlantic. Remarks:
found in mangrove swamps, brackish
estuaries and adjacent waters.
(K. Shibukawa)Parablennius thysanius(Jordan & Seale, 1907)
Tasseled Blenny
D XII, 13–15; A II, 14–17; P1 14; P2
I, 3. Body relatively short, compressed. Cirrus on eye a short fringed
flap; nasal cirrus simple; no cirri onnape and cheek. Fleshy crest on top
head absent in both sexes. Gill open-

Omobranchus punctatus, UMTF 1147 (KAUM–I. 16337), 4.1 cm SL
Cendering, 16 Sept. 2008

Parablennius thysanius, UMTF 1410 (KAUM–I. 16398), 3.2 cm SL
Dungun, 24 Sept. 2008

ing wide, continuous across ventral
surface of head. Dorsal fin continuous,
without distinct notch between spinous and segmented-ray portions.
Caudal fin usually with 13 segmented
rays, several of them branched. Color:body olive gray dorsally, light gray or
yellowish gray ventrally, mottled withvarious-sized dark spots; head olive
gray, with numerous dense small darkbrown spots, forming reticulated pattern. Size: 5 cm. Distribution: IndoWest Pacific; also introduced to
Hawaii. Remarks: found in shallow
coastal waters and brackish estuaries.
(K. Shibukawa)

188

Salarias fasciatus(Bloch, 1786)
Jeweled Blenny
D XII, 18–20; A II, 19–21; P1 14; P2
I, 3; V 10 + 26–27 = 36–37. Body
moderately elongate. Supraorbital andnuchal cirri branched. Occipital crest
absent in both sexes. Gill opening continuous across ventral surface of head.
Dorsal fin shallowly notched. Last
anal-fin ray bounded by membrane tocaudal peduncle. Several caudal-fin
rays branched. Color: color variable,
usually olivaceous to brown with irregular dark bars and many round to
oblong white spots; a few very small,bright blue spots dorsally on posterior

Salarias fasciatus, UMTF 1208 (KAUM–I. 16642), 10.4 cm SL
Bidong Island, 12 Oct. 2008

half of body. Size: attaining to 14 cm. Barrier Reef and New Caledonia. ReDistribution: Indo-West Pacific, in- marks: usually seen on coral reefs.
cluding east coast of Africa, Red Sea,
(H. Imamura)
Andaman Sea, Ryukyu Islands, Great189

CALLIONYMIDAE
Dragonets

By Koichi Shibukawa
Small to moderate-sized (up to
45 cm), depressed benthic fishes.
Body elongate, depressed or subcylindrical. Head usually wide andwell depressed, triangular when seenfrom above in many species; gill
opening restricted to a small dorsal
or sublateral pore; preopercle witha strong spine; no spines on opercle
and subopercle; jaw with villiform
teeth; no teeth on roof of mouth; upper jaw greatly protractile. Two dorsal fins, comprising spinous (with
III–V spines; rarely absent) and soft
(with 7–10 soft rays) portions; anal
fin with 6–10 soft rays; pelvic finsthoracic, with I spine and 5 soft rays.
No scales on head and body. Color:
highly variable, but dull sandy-colored in many species, e.g., head andbody grayish or brownish dorsally,
whitish ventrally, mottled and/or

2 dorsal finspreopercle witha strong spine

body moderately depressed without scales

spotted with paler colored spots; somehave entirely reddish body (especiallyin deep-water species) or several vivid
color with complex patterns on head,
body and fins.
Remarks: found in seagrass beds,
bays, coral reefs, estuaries, and shelfwaters, and commonly found on sandyor muddy bottoms. Locally utilized asfood fish.
Similar families occurring in thearea: Eleotridae – usually no preopercular spine; gill opening wide;
scales on body. Gobiidae – usuallyno preopercular spine; pelvic fins
fused medially one another in many
species; gill opening wide. Platycephalidae – mouth large, lower jaw
projecting beyond upper jaw; shortspines, serrae and/or bony tubercles
on the head; gill opening wide; scales
on body.

Calliurichthys japonicus(Houttuyn, 1782)
Japanese Longtail Dragonet
D IV + 9; A 8; P1 18–21. Head andbody compressed. A pair of small
bony bumps on dorsal surface of headbehind eye. Preopercular spine slenderand straight, with several serrae along
its dorsal margin. Transverse canalbranch connecting opposite lateralline canals present on caudal peduncle. Anterior two spine of first dorsal
fin elongate and filamentous in large
male. All but posteriormost one segmented rays of second dorsal fin unbranched. Caudal fin very long. Color: head and body light grayish brown
dorsally, pale ventrally; a distinct
black spot between third and fourth
spine of first dorsal fin; a broad submarginal black stripe on anal fin. Size:
22 cm SL. Distribution: West Pacific.
Remarks: inhabits sandy-mud bottoms at the depths of 10–200 m. Generic assignment follows Nakabo(1982).

Calliurichthys japonicus, KAUM–I. 17199, 18.1 cm SL
off Terengganu (KR), 5 Jan. 2009

Calliurichthys japonicus, KAUM–I. 17200, 16.7 cm SL
off Terengganu (KR), 5 Jan. 2009

190

Dactylopus dactylopus(Valenciennes, 1837)
Fingered Dragonet
D IV + 8; A 7; P1 19. Body subcylindrical or slightly compressed posteriorly. No transverse canal branch connecting opposite lateral line canals on
caudal peduncle. Dorsal fin spines
elongate and filamentous in male. All
rays of second dorsal fin branched.
Spine and first ray of pelvic fin connected together, free from other rays.
Color: head and body grayish brown
brownish dorsally, pale ventrally; numerous pale and dusky spots on body.
Size: 25 cm SL. Distribution: eastern
Indian Ocean and West Pacific. Remarks: inhabits sandy-mud bottoms
in shallow coastal waters, particularly
seagrass beds.

Dactylopus dactylopus, KAUM–I. 17192, 10.4 cm SL
off Terengganu (KR), 5 Jan. 2009

Repomucenus belcheri recurvispinnis(Li, 1966)
Concave Dragonet
D IV + 9; A 9; P1 17–21. Head andbody compressed. Preopercular spine
straight, with several serrae along its
dorsal margin. Transverse canalbranch connecting opposite lateralline canals present on caudal peduncle. No filamentous spine of first dorsal fin in both sexes. All but posteriormost one segmented rays of second
dorsal fin unbranched. Caudal fin
moderate in size. Color: head andbody light grayish brown dorsally,
pale ventrally; lower side of body witha series of short, wavy vertical duskylines in large male; anal fin with submarginal black stripe in male, a seriesof black blotches in female. Size: 22
cm SL. Distribution: Indo-West Pacific. Remarks: the generic assignment tentatively follows Nakabo(1982; for R. belcheri). Fricke (1983,
2002) treated this as subspecies ofCallionymus belcheri Richardson,
1844.

Repomucenus planus(Ochiai, 1955)
Japanese Darter Dragonet
D IV + 9; A 9; P1 18–22. Head andbody compressed. Preopercular spinewith 3–6 serrae along its dorsal margin and curved posteriormost tip.
Transverse canal branch connecting
opposite lateral line canals present on

Repomucenus belcheri recurvispinnis, KAUM–I. 16894, 7.5 cm SL
off Terengganu (KT), 6 Dec. 2008

Repomucenus planus, KAUM–I. 17113, 7.9 cm SL
off Terengganu (KT), 31 Dec. 2008

Repomucenus planus, KAUM–I. 17270, 7.1 cm SL
off Terengganu (KT), 18 Jan. 2009caudal peduncle. First dorsal fin small,without filamentous spine in both sexes. All but posteriormost one segmented rays of second dorsal fin unbranched. Color: head and body lightyellowish brown dorsally, pale ventrally, with numerous, dense darkbrown dots; first dorsal fin largely
blackish in female; many white spots
on second dorsal fin in male. Size: 10
cm SL. Distribution: West Pacific.
Remarks: inhabits sandy bottoms inshallow coastal waters. The genericassignment follows Nakabo (1982).

191

ELEOTRIDAE
Sleepers

By Koichi Shibukawa
Small to moderate sized fishes(up to ca. 85 cm, commonly to 20cm), bottom-oriented fishes. Body
moderately elongate, more or lesscompressed posteriorly; no poredor tubed lateral-line scales on body.
Teeth usually conical, forming someor more rows in each jaw; sensorypapillae rows well developed on head
(and few on body and caudal fin);branchiostegals 6. Two dorsal fins,
first one with VI–X flexible spines,
second one with I spine and 6–12 softrays; pelvic fins separated, innermostor preceding rays longest; pelvic fin
usually with I spine and 5 soft rays.
Color: highly variable, typically
blackish or grayish brown with dusky
mottles, whereas some species has
brilliant color with e.g., bright yellow
and/or red spots.
Remarks: typically found in

2 dorsal fins with VI–X + I, 6–12
well developed
sensory papillae
on headno pored or tubed
lateral line scales

pelvic fins separated

fresh and brackish waters, whereassome found in coral reefs. Carnivorous. Large species often esteemed asfood fish; some freshwater colorful
species often treated as aquarium fish.
Similar families occurring in thearea: Blenniidae – single dorsal fin ;teeth incisor-like, forming single row ineach jaw. Callionymidae – preopercle
with a strong spine; no scales on body;gill opening restricted to a small dorsal

anal fin with I, 6–12

or dorsolateral pore. Gobiidae – pelvic fins fused medially one anotherin many species; five branchiostegal
rays. Platycephalidae – many shortspines or bony tubercles on head; nospines on second dorsal and anal fins.
Ptereleotridae – bottom-oriented, freeswimming fishes, with elongate (butnot eel-like) and compressed headand body; five branchiostegal rays.
Tripterygiidae – three dorsal fins.

Butis butis(Hamilton, 1822)
Duckbill Sleeper
D VI + I, 8; A I, 8; P1 18–19; LR
29–31; PDS 25–33. Body elongate,subcylindrical and compressed posteriorly. Head depressed. Snout elongate
and depressed, duck-bill shaped. LowButis butis, UMTF 1104 (KAUM–I. 16492), 5.3 cm SLer jaw prominent. Bony edge with serestuary near UMT, 30 Sept. 2008
rae on interorbital area around dorsal
margin of eye. Pelvic fins widely separated. Cheek, operculum and interorbital with ctenoid scales; scales present between eye and interorbital bonyridge; small axillary scale between
large body scales. Color: head andbody blackish brown, with numerous
black, orange and pale dots; base ofpectoral fin with a distinct large black
spot edged with two vivid red spots
dorsoventrally. Size: 14.5 cm TL. DisButis koilomatodon, UMTF 1433 (KAUM–I. 16756), 2.7 cm SL
tribution: Indo-West Pacific. ReSetiu, 30 Oct. 2008
marks: found in brackish estuaries
and mangrove swamps.
short, subcylindrical and compressed brown, with indistinct 4–5 broad
posteriorly. Head slightly depressed. dusky oblique bands; typically pectoLower jaw prominent. Bony edge with ral-fin base with a distinct black spot.
Butis koilomatodon
serrae on interorbital area around dor- Size: 7.5 cm SL. Distribution: Indo(Bleeker, 1849)
sal margin of eye. Pelvic fins widely West Pacific. Remarks: found in
Mud Sleeper
separated. Cheek and operculum with mangrove swamps, brackish estuaries
D VI + I, 8; A I, 8; P1 21–22; LR ctenoid scales; no scales on interor- and adjacent coastal waters with mud25–28; PDS 11–15. Body relatively bital area. Color: body blackish dy bottoms.

192

Ophiocara porocephala(Valenciennes, 1837)
Northern Mud Sleeper
D VI + I, 8–9; A I, 6–7; P1 14–15; LR
33–40; PDS 18–26. Body moderately
elongate, subcylindrical and compressed posteriorly. Head depressed.
Lower jaw prominent. Pelvic fins
widely separated. Cheek and operculum with cycloid scales; predorsal
scales extending anteriorly to snout;scales on body ctenoid. Color: headand body dark grayish brown dorsally,
becoming much paler ventrally; numerous, dense white speckles on bodyand dorsal and caudal fins in young
and adult; some pale vertical bands on
body in small specimen. Size: 32 cm
TL. Distribution: Indo-West Pacific.
Remarks: found in mangrove swamps
and brackish estuaries.

Oxyeleotris marmorata(Bleeker, 1852)
Marble Goby
D VI + I, 9; A I, 7–8; P1 17–19; LR
80–90; PDS 60–65. Body elongate,subcylindrical and compressed posteriorly. Lower jaw prominent. No ventrally-directed spinous projection alongposterior margin of preopercle. Vomer
edentate. Pelvic fins widely separated.
Scales on body small cycloid anteriorly, weak ctenoid posteriorly. Color:ground color of head and body lightgrayish brown, with some large blackish brown markings. Size: 63 cm TL.
Distribution: West Pacific. Remarks: found in brackish estuaries,
reservoirs and freshwater rivers andponds. Common cultured fish, highly
esteemed as food fish in South Asian
countries.

Ophiocara porocephala, UMTF 1015 (KAUM–I. 16547), 11.4 cm SL
estuary near UMT, 30 Sept. 2008

Ophiocara porocephala, UMTF 1032 (KAUM–I. 16546), 5.6 cm SL
estuary near UMT, 30 Sept. 2008

Oxyeleotris marmorata, UMTF 1311 (KAUM–I. 16482), 16.4 cm SL (preserved specimen)
estuary near UMT, 29 Sept. 2008

Oxyeleotris urophthalmus(Bleeker, 1851)
Sinuous Gudgeon
D VI + I, 9–12; A I, 8–10; P1 15–18;
LR 85–102; PDS 55–60. Body elongate, subcylindrical and compressed
posteriorly. Lower jaw prominent. No
ventrally-directed spinous projection
along posterior margin of preopercle.
Vomer edentate. Pelvic fins widely
separated. Scales on body small cycloid anteriorly, weak ctenoid posteriorly. Color: ground color of head andOxyeleotris urophthalmus, KAUM–I. 17312, 11.5 cm SL
Terengganu River, 20 Jan. 2009body dark grayish brown; typically an
indistinct black ocellated spot at upper
part of caudal fin basally; 4–5 narrowsaddle-like, vertical yellow bands on
body in small specimen. Size: 28 cm

TL. Distribution: West Pacific. Remarks: found in mangrove swamps
and adjacent brackish estuaries. Carnivorous.

193

GOBIIDAE
GobiesBy Koichi Shibukawa and Mazlan Abdul Ghaffar
Small to moderate-sized (upto ca. 60 cm, commonly to 10 cm),
bottom-oriented fishes. Body shapehighly variable, but, typically, moderately elongate and more or lesscompressed posteriorly; no poredor tubed lateral-line scales on body.
Teeth typically conical (flattened andincisor-like in some genera), forming a single to several rows in each
jaw; minute sensory papillae welldeveloped on head (and few on bodyand caudal fin), forming transverse
or longitudinal rows; five branchiostegal rays. Usually two dorsal fins,
first one with 0–X flexible spines,
second one usually with I spine and
5–66 soft rays; anal fin usually with
I flexible spine and 5–65 soft rays;
pectoral fin with 12–65 soft rays;dorsal- and anal-fin spines usually
narrow and flexible; pelvic fins fused
medially by frenum (between spines)
and connecting membrane (between
innermost rays), exclusive of several coral reef genera with separated
ones; pelvic fin usually with I spineand 4–5 soft rays. Scales cycloid or
ctenoid (with peripheral cteni only),usually 2 dorsal fins with 0–X + I, 5–66
well developed
sensory papillae
on headno pored or tubed
lateral line scales

pelvic fins fused
medially in many species

anal fin with I, 5–65

minute to moderately large in size.
Color: highly variable.
Remarks: found in various habitats from torrential freshwater rivers to
shelf waters (to depth of ca. 500 m),
but most common in brackish waters
and shallow coastal waters. Large species may be esteemed as food fish;
several freshwater species may betreated as aquarium fish.
Similar families occurring in
the area: Blenniidae – single dorsal
fin (may be deep notch developed
between spinous and soft portions);
teeth incisor-like, forming single row
in each jaw; pelvic fins separated.

Callionymidae – preopercle with astrong spine; no scales on body; gill
opening restricted to a small dorsal or dorsolateral pore; pelvic finsseparated. Eleotridae – pelvic finsseparated; six branchiostegal rays.
Platycephalidae – many short spines
or bony tubercles on head; pelvic finsseparated. Ptereleotridae – bottomoriented, freeswimming fishes, withelongate (but not eel-like) and compressed body and separated pelvicfins; mouth almost vertical in many
species. Tripterygiidae – three dorsal
fins; pelvic fins separated.

Acentrogobius caninus(Valenciennes, 1837)
Tropical Sand Goby
D VI + I, 9; A I, 9; P1 18; LR 26–27;
PDS 16. Body moderately elongate
and compressed. Head slightly compressed. Jaws subequal. Gill opening
not extending anteriorly to a verticalthrough posterior margin of preopercle. Single or some enlarged caninoid
teeth on each side of lower jaw.
Tongue truncate. Pelvic fins united
medially; frenum present. Scales ctenoid, excluding those on nape, pectoral base and breast cycloid; no scales
on cheek; operculum may be scaled.
Sensory canals and pores present onhead; longitudinal pattern of sensory-

Acentrogobius caninus, KAUM–I. 17111, 9.9 cm SL
off Terengganu (KT), 31 Dec. 2008

papillae rows on cheek, some rows
multiple. Color: large ovoid black
spot just above dorsalmost of gill
opening; four midlateral dusky spots
on body. Size: 13 cm TL. Distribu-

tion: Indo-West Pacific. Remarks:
found in brackish estuaries and adjacent coastal waters with sandy-mud
bottom.
(K. Shibukawa)194

Acentrogobius viridipunctatus(Valenciennes, 1837)
Spotted Green Goby
D VI + I, 10; A I, 9–10; P1 19–20; LR
33–35; PDS 27–33. Body moderately
elongate and compressed. Head slightly compressed. Jaws subequal. Gill
opening extending anteriorly to a vertical through posterior margin of preopercle. Single or some enlarged
caninoid teeth on each side of lower
jaw. Tongue truncate. Pelvic fins united medially; frenum present. Scales
ctenoid, excluding those on head,nape, pectoral base and breast cycloid;
upper part of cheek and operculum
scaled. Sensory canals and pores present on head; distinct transverse sensory-papillae rows on cheek. Color:head and body grayish brown, withnumerous bright light green or blue
spots (faded immediately after death);
L-shaped dusky marking below eye.
Size: 12 cm SL. Distribution: IndoWest Pacific. Remarks: found inbrackish estuaries and adjacent coastal
waters with sandy-mud bottoms.
(K. Shibukawa)Acentrogobius viridipunctatus, KAUM–I. 16802, 6.0 cm SL
Merang, 2 Nov. 2008

Amblygobius phalaena, UMTF 1249 (KAUM–I. 16562), 5.7 cm SL
Bidong Island, 12 Oct. 2008

Amblygobius phalaena(Valenciennes, 1837)
White-barred Goby
D VI + I, 14–15; A I, 14–15; P1 19–
20; LR 54–57; PDS 22–25. Body
moderately elongate and compressed.
Head compressed. Snout rounded,
protruding beyond jaws. Gill opening
not extending anteriorly to a verticalline through posterior margin of preopercle. Tongue rounded or near truncate. Pelvic fins united medially; frenum present. Scales on body cycloid
anteriorly, ctenoid posteriorly; head
naked, except for dorsal part of operculum with cycloid scales. Sensory
canals and pores present on head; aseries of short transverse rows of sensory papillae below eye. Color:ground color of head and body grayish
olive dorsally, pale ventrally; bodywith five blackish or grayish brown
vertical bands, typically edged withwhite anteroposteriorly; some small
black spot on caudal fin, anterior oneof them well apart from caudal-finbase; color of large specimen frequently darkened (probably as nuptial
color). Size: 8 cm SL. Distribution:
West Pacific. Remarks: found in cor-

Amblygobius stethophthalmus, UMTF 1073 (KAUM–I. 16502), 5.9 cm SL
Cendering, 6 Oct. 2008

al reefs, protected bays, lagoons and ent. Scales on body cycloid anteriorly,
estuaries with sandy bottoms and rub- ctenoid posteriorly; head naked, exbles.
(K. Shibukawa) cept for dorsal part of operculum withcycloid scales. Sensory canals andAmblygobius stethophthalmus pores present on head; a series of shorttransverse rows of sensory papillae(Bleeker, 1851)
below eye. Color: ground color of
Head-stripe Goby
head and body olive dorsally, pale
D VI + I, 14–15; A I, 15–16; P1 20; ventrally; broad black stripe from tip
LR 66–74; PDS 28. Body moderately of snout through eye to anterior part ofelongate and compressed. Head com- body; middle of operculum with faintpressed. Snout rounded, protruding broad dusky stripe, meeting posteriorbeyond jaws. Gill opening not extend- ly with black spot on pectoral-fin base.
ing anteriorly to a vertical line through Size: 8 cm SL. Distribution: West Paposterior margin of preopercle. cific and Andaman Sea. Remarks:
Tongue rounded or near truncate. Pel- found in protected bays, lagoons andvic fins united medially; frenum pres- estuaries.
(K. Shibukawa)

195

Asterropteryx semipunctata
Rüppell, 1830
Starry Goby
D VI + I, 10; A I, 9; P1 16–17; LR
23–25; PDS 7–8. Body relatively
short, compressed. Head compressed.
Two to10 spinous projections at posterior margin of preopercle. Gill opening not extending anteriorly to a vertical through posterior margin of
preopercle. Tongue rounded or neartruncate. Third spine of first dorsal fin
elongate and filamentous; pelvic finsalmost separated; frenum absent.
Scales on body ctenoid, exclusive ofpectoral-fin base and breast with cycloid scales; cheek, operculum andbranchiostegal membrane scaled. Sensory canals and pores present on head;
distinct transverse pattern of sensorypapillae rows on cheek. Color: headand body grayish with numerous
bright blue dots; ca. 4 irregular rowsof small dusky blotches on body. Size:
4 cm SL. Distribution: Indo-Pacific.
Remarks: found around rubbles onshallow protected bays, coral reefs, lagoons and brackish estuaries.
(K. Shibukawa)

Asterropteryx semipunctata, UMTF 1372 (KAUM–I. 16386), 2.5 cm SL
Dungun, 24 Sept. 2008

Bathygobius cocosensis, UMTF 1253 (KAUM–I. 16660), 3.6 cm SL
Kemaman, 20 Oct. 2008

Bathygobius cocosensis(Bleeker, 1854)
Cocos Frill Goby
D VI + I, 8–10; A I, 7–8; P1 16–20;
LR 33–39; PDS 6–14. Body moderately elongate and compressed. Headsubcylindrical. Gill opening not extending anteriorly to a vertical throughposterior margin of preopercle. Mental flap well developed, slightly protruding posterolaterally. No flap-likeprojection at tip of anterior nostril.
Anterior margin of tongue notched.
Upper part of pectoral fin with filamentous free rays. Pelvic fins united
medially; frenum present, with no distinct projection at middle of posterior
margin. Scales ctenoid posteriorly, cycloid anteriorly; head naked. Sensory
canals and pores present on head; longitudinal pattern of sensory papillae
on chin. Color: head and body lightgrayish blown dorsally, pale ventrally;a series of small black blotches on
mid-lateral body. Size: 5 cm SL. Distribution: Indo-Pacific. Remarks:
found in shallow rocky shores.
(K. Shibukawa)Bathygobius cyclopterus, UMTF 1181 (KAUM–I. 16290), 3.7 cm SL
Cendering, 14 Sept. 2008

Bathygobius cyclopterus(Valenciennes, 1837)
Spotted Frill Goby
D VI + I, 8–9; A I, 8; P1 19–23; LR
36–40; PDS 12–20. Body moderately
elongate and compressed. Head subcylindrical. Gill opening not extending anteriorly to a vertical throughposterior margin of preopercle. Mental flap well developed, slightly protruding posterolaterally. A flap-likeprojection at tip of anterior nostril.
Anterior margin of tongue notched.
Upper part of pectoral fin with filamentous free rays. Pelvic fins united

medially; frenum present, with a distinct projection at middle of posterior
margin. Scales ctenoid posteriorly, cycloid anteriorly; head naked, except
for upper part of operculum with cycloid scales. Sensory canals and porespresent on head; longitudinal patternof sensory papillae on cheek. Color:head and body dark gray or grayish
blown dorsally, pale ventrally; a seriesof small black blotches on mid-lateralbody. Size: 5 cm SL. Distribution:
Indo-Pacific. Remarks: found in shallow rocky shores.
(K. Shibukawa)

196

Bathygobius fuscus(Rüppell, 1830)
Dusky Frill Goby
D VI + I, 8–10; A I, 7–8; P1 16–20;
LR 31–40; PDS 10–19. Body moderately elongate and compressed. Headsubcylyndrical. Gill opening not extending anteriorly to a vertical throughposterior margin of preopercle. Mental flap well developed, trapezoid, but
not protruding posterolaterally. No
flap-like projection at tip of anterior
nostril. Anterior margin of tongue
notched. Upper part of pectoral fin
with filamentous free rays. Pelvic finsunited medially; frenum present, withno distinct projection at middle of posterior margin. Scales ctenoid posteriorly, cycloid anteriorly; head naked.
Sensory canals and pores present onhead; longitudinal pattern of sensory
papillae on chin. Color: head andbody grayish blown dorsally, paleventrally; a series of black blotches,
each extending ventrally to ventral
half of body, on mid-lateral body; dorsal fin with a broad pale yellow distal
margin. Size: 8 cm SL. Distribution:
Indo-Pacific. Remarks: found inrocky or coral reefs, sandy beacheswith rubbles and brackish estuaries.
(K. Shibukawa)

Cryptocentrus maudae
Fowler, 1937

Bathygobius fuscus, UMTF 1199 (KAUM–I. 16288), 5.3 cm SL
Cendering, 14 Sept. 2008

Cryptocentrus maudae, UMTF 1274 (KAUM–I. 16540), 2.6 cm SL
Cendering, 6 Oct. 2008

Drombus triangularis, KAUM–I. 16758, 2.4 cm SL
Setiu, 30 Oct. 2008

Maude's Shrimpgobyy
D VI + I, 9–10; A I, 9–10; P1 17; LS
120. Body elongate, compressed.
Head slightly compressed. Jaws subequal. Gill opening reaching anteriorly around a vertical through posterior
margin of preopercle. Pelvic fins united medially; frenum present. Scales
minute, cycloid; head, breast and pectoral-fin base naked. Sensory canals
and pores present on head; distinct
transverse sensory-papillae rows on
cheek. Color: ground color of headand body blackish brown; anterodorsal part of eye, interorbital space,snout and anterior tip of lower jaw
pale or pale brown; several pale orpale brown saddles on head and body,in addition to numerous minute palespots; anal fin with 5–6 diagonal blackbars. Size: 5 cm SL. Distribution:
Andaman Sea and West Pacific. Remarks: found in tidal estuaries andprotected bays with sandy-mud bot-

Drombus triangularis, KAUM–I. 16760, 2.0 cm SL
Setiu, 30 Oct. 2008

toms and rubble patches; symbiotical- medially; frenum present. Scales ctely associates with alpheid shrimps.
noid, excluding nape, breast, pectoral(K. Shibukawa) fin base and predorsal area with cycloid scales; head naked. Sensory
canals and pores present on head; disDrombus triangularis
tinct transverse pattern of sensory-pa(Weber, 1909)
pillae rows on cheek. Color: head andBrown Drombus
body dusky with irregular mottles;
D VI + I, 9–10; A I, 8; P1 16–17; LR yellow triangular spot at uppermost of
30–31; PDS 12–19. Body moderately pectoral base. Size: 5 cm. Distribuelongate and compressed. Head slight- tion: Indo-West Pacific. Remarks:
ly depressed. Lower jaw projecting found in mangrove swamps and brackbeyond upper jaw. Gill opening not ish estuaries with rubbles.
reaching anteriorly to a vertical through(K. Shibukawa)preopercular margin. Pelvic fins united

197

Eugnathogobius variegatus(Peters, 1868)
Stripe-face Brackish Goby
D VI + I, 7–9; A I, 7–8; P1 16–17; LR
29–34. Body elongate and compressed. Head slightly depressed.
Jaws subequal. Gill opening not reaching anteriorly to a vertical throughpreopercular margin. Pelvic fins united medially; frenum present. Scales
on body cycloid anteriorly, ctenoid
posteriorly. Sensory canals and porespresent on head; distinct transverse
pattern of sensory-papillae rows on
cheek. Color: head and body grayish
yellow dorsally, becoming paler ventrally; numerous blackish or dark grayspeckles on head and body; obliquedusky streaks from eye to cheek andmiddle of upper jaw. Size: 4 cm. Distribution: West Pacific. Remarks:
found in mangrove swamps.
(K. Shibukawa)

Eviota pellucida
Larson, 1976

Eugnathogobius variegatus, UMTF 1507 (KAUM–I. 16712), 3.6 cm SL
Setiu, 27 Oct. 2008

Eviota pellucida, UMTF 1263 (KAUM–I. 16636), 1.3 cm SL
Bidong Island, 13 Oct. 2008

Pellucida Pygmy Goby
D VI + I, 7–9; A I, 7; P1 15–16; LR
21–22; PDS 0. Body relatively short,compressed. Head slightly compressed. Jaws subequal. Gill opening
not reaching anteriorly to a verticalthrough preopercular margin. Pelvic
fins almost separated; frenum absent.
Scales ctenoid; head and pectoral-finbase naked. Sensory canals and porespresent on head; reduced longitudinalpattern of sensory-papillae rows on
cheek. Color: when alive, head andbody subtranslucent, with two narrow
longitudinal yellow stripe from eye to
middle of body; longitudinal white
stripe on belly; color immediately
turned to pale yellow or pale reddish
yellow after death. Size: 2 cm SL. Distribution: eastern Indian Ocean andWestern Central Pacific. Remarks:
found in coral reefs. (K. Shibukawa)

Eviota storthynx, UMTF 1442 (KAUM–I. 16635), 1.6 cm SL
Bidong Island, 13 Oct. 2008

Eviota storthynx(Rofen, 1959)
Storthynx Pygmy Goby
D VI + I, 8–9; A I, 7; P1 14–16; LR
22–23; PDS 0. Body relatively short,compressed. Head slightly compressed. Jaws subequal. Gill opening
not reaching anteriorly to a verticalthrough preopercular margin. Pelvic
fins almost separated; frenum absent.
Scales ctenoid; head and pectoral-finbase naked. Sensory canals and porespresent on head; reduced longitudinalpattern of sensory-papillae rows on
cheek. Color: when alive, head andbody subtranslucent, with several
white spots; nasal tube dark red; edgesof scale pockets dusky; 7–8 dark grayish brown subcutaneous vertical bars
on ventral half of tail; color immediately turned to pale after death, with aprominent black spot behind eye.
Size: 2.1 cm SL. Distribution: Western Australia and Western Pacific. Remarks: found in shallow rocky andcoral reefs.
(K. Shibukawa)

198

Exyrias puntang(Bleeker, 1851)
Puntang Goby
D VI + I, 10–11; A I, 8–10; P1 15–18;
LR 29–33; PDS 10–12. Body moderately elongate and compressed. Headsubcylindrical. Lower jaw not projecting anteriorly beyond upper jaw. Gill
opening not reaching anteriorly to avertical through posterior margin of
preopercle. Spines of first dorsal fin
typically elongate and filamentous.
Pelvic fins united medially; frenum
present. Scales on body cycloid anteriorly, ctenoid posteriorly; cheek andoperculum scaled. Sensory canals andpores present on head; longitudinalpattern of sensory-papillae rows on
cheek. Color: ground color of headand body dull yellow or light yellowish brown, paler and tinged with yellow ventrally; several vague verticaldusky bars on body; dorsal fins withnumerous dusky spots. Size: 12 cm
SL. Distribution: Indo-West Pacific.
Remarks: found in mangroveswamps, brackish estuaries and adjacent shallow coastal waters with sandy-mud bottoms.
(K. Shibukawa)Exyrias puntang, KAUM–I. 16795, 10.2 cm SL
Merang, 2 Nov. 2008

Favonigobius reichei, UMTF 1508 (KAUM–I. 16716), 4.4 cm SL
Setiu, 28 Oct. 2008

Favonigobius reichei(Bleeker, 1854)
Indo-Pacific Tropical Sand Goby
D VI + I, 8; A I, 8; P1 16; LR 27; PDS
0–3. Body moderately elongate andcompressed. Head subcylindrical orslightly depressed. Lower jaw slightly
projecting beyond upper jaw. Gill
opening not reaching anteriorly to avertical through posterior margin of
preopercle. Tongue near truncate. Second spine of first dorsal fin elongate
and filamentous in male; pelvic finsunited medially; frenum present.
Scales ctenoid, excluding breast andpectoral-fin base with cycloid scales;
head naked. Sensory canals and porespresent on head; sensory papillae rows
on cheek multiple, but not forming
distinct transverse rows. Color: head

Favonigobius reichei, KAUM–I. 16781, 3.5 cm SL
Merang, 2 Nov. 2008and body pale gray with numerous
black dots. Size: 7 cm SL. Distribution: Indo-West Pacific. Remarks:
found in mangroves, brackish estuaries and adjacent shallow coastal waters with sandy-mud bottoms.
(K. Shibukawa)

199

Fusigobius inframaculatus(Randall, 1994)
Longspine Goby
D VI + I, 9; A I, 8; P1 17–19; LR
23–24; PDS 0. Body moderately elongate, compressed posteriorly. Headsubcylindrical. Snout pointed. Lower
jaw slightly projecting beyond upper
jaw. Gill opening not extending anteriorly to a vertical through posterior
margin of preopercle. First spine offirst dorsal fin elongate, filamentous.
Pelvic fins united medially; frenum
present. Scales on body ctenoid, exclusive of side of nape, breast andpectoral-fin base with cycloid scales;
head naked. Sensory canals and porespresent on head; reduced longitudinalpattern of sensory papillae on cheek.
Color: head and body pale (subtranslucent when alive) with numerous orange
yellow spots; a large black blotch at
caudal-fin base. Size: 4 cm SL. Distribution: Indo-West Pacific. Remarks:
found on flat sandy bottoms in coral
reefs.
(K. Shibukawa)

Fusigobius inframaculatus, UMTF 1329 (KAUM–I. 16626), 3.3 cm SL
Bidong Island, 13 Oct. 2008

Glossogobius aureus, KAUM–I. 17208, 12.9 cm SL
off Terengganu (KT), 10 Jan. 2009

Glossogobius aureus
Akihito & Meguro, 1975
Golden Flat-head Goby
D VI + I, 9; A I, 8–9; P1 18–21; LR
31–34; PDS 22–27. Body elongate
and compressed. Head slightly depressed. Lower jaw prominent. Gill
opening reaching anteriorly to, or alittle beyond, a vertical through posterior margin of preopercle. Anterior
margin of tongue notched. Pelvic finsunited medially; frenum present.
Scales ctenoid, excluding predorsal
area, breast, belly and pectoral-finbase and operculum with cycloid
scales. Sensory canals and pores present on head; longitudinal pattern of
sensory papillae on cheek. Color:head and body light yellowish brown,darkened dorsally; body with midlateral series of five dusky blotches, aswell as many faint irregular duskylines and spots dorsally. Size: 20 cm
SL. Distribution: Indo-West Pacific.
Remarks: found in lower reaches oflarge rivers, mangroves, brackish estuaries and adjacent shallow coastal waters with sandy-mud bottoms.
(K. Shibukawa)

Glossogobius bicirrhosus, KAUM–I. 16779, 4.4 cm SL
Merang, 2 Nov. 2008

Glossogobius bicirrhosus, UMTF 1259 (KAUM–I. 16780), 3.2 cm SL
Merang, 2 Nov. 2008

Glossogobius bicirrhosus(Weber, 1894)
D VI + I, 9; A I, 8; P1 18–19; LR
30–32; PDS 13–15. Body elongate andcompressed. Head slightly depressed.
Lower jaw prominent, with a pair ofshort barbells on chin. Gill opening
reaching anteriorly to, or a little beyond, a vertical through posterior margin of preopercle. Anterior margin of
tongue notched. Pelvic fins united medially; frenum present. Scales on bodylargely ctenoid; blest with cycloid
scales. Sensory canals and pores present on head; longitudinal pattern of
sensory papillae on cheek. Color:head and body light grayish brown orbeige, darkened dorsally; body withmidlateral series of dusky blotches, aswell as many faint irregular duskylines and spots dorsally. Size: 9 cm SL.
Distribution: West Pacific. Remarks:
found in lower reaches of streams,
mangroves and brackish estuaries.
(K. Shibukawa)

200

Glossogobius circumspectus(Macleay, 1883)
Circumspect Flat-head Goby
D VI + I, 8–10; A I, 8; P1 18–19; LR
29–33; PDS 15–19. Body elongate
and compressed. Head slightly depressed. Lower jaw prominent. Gill
opening reaching anteriorly to, or alittle beyond, a vertical through posterior margin of preopercle. Anterior
margin of tongue notched. Pelvic finsunited medially; frenum present.
Scales ctenoid, excluding predorsal
area, breast, belly and pectoral-finbase and operculum with cycloid
scales. Sensory canals and pores present on head; numerous transverserows of sensory papillae on cheek.
Color: head and body beige, darkened
dorsally; body with midlateral seriesof five dusky blotches, as well as
many faint irregular dusky lines andspots. Size: 10 cm SL. Distribution:
Indo-West Pacific. Remarks: foundin lower reaches of large rivers, mangroves and brackish estuaries.
(K. Shibukawa)

Glossogobius circumspectus, UMTF 1371 (KAUM–I. 16807), 5.5 cm SL
Merang, 2 Nov. 2008

Gobiodon quinquestrigatus, UMTF 1258 (KAUM–I. 16615), 2.6 cm SL
Bidong Island, 12 Oct. 2008

Gobiodon quinquestrigatus(Valenciennes, 1837)
Fivelined Coralgoby
D VI + I, 10–11; A I, 8–9; P1 18–20;
LR 0; PDS 0. Body ovate, well compressed. Head compressed; snout
rounded. Jaws subequal. Gill opening
Hemigobius hoevenii, KAUM–I. 16738, 4.1 cm SL
narrow, restricted to pectoral-fin base.
Setiu, 29 Oct. 2008
Tongue narrow, with rounded anterior
margin. Pelvic fins small, united medially; frenum present. No scales onhead and body. Sensory canals andpores present on head; reduced longitudinal pattern of sensory papillae on
cheek. Color: head reddish brown;
body and fins dark reddish brown or
blackish; four and one vertical sky
lines on head and pectoral-fin base,respectively. Size: 3 cm SL. DistribuHemigobius hoevenii, UMTF 1278 (KAUM–I. 16715), 3.4 cm SL
tion: West Pacific. Remarks: foundSetiu, 27 Oct. 2008
in coral reefs; inhabits amongst
branches of acroporid corals of the ge- pressed. Snout rounded, sometimes a tudinal pattern of sensory-papillae rows
nus Acropora.
(K. Shibukawa) little projecting beyond jaws anteriorly. on cheek. Color: head and body olive
Jaws subterminal. Interorbital space or grayish brown, darkened dorsally;broad. Gill opening not extending ante- body typically with irregular black
Hemigobius hoevenii
riorly to a vertical through preopercular blotches and oblique bands; fins tinged(Bleeker, 1851)
margin. Pelvic fins united medially; fre- with yellow in large male. Size: 4 cm
Banded Mullet Gobynum present. Scales ctenoid, excepting SL. Distribution: West Pacific and AnD VI + I, 7–8; A I, 7–9; P1 13–16; LR those on predorsal area, pectoral base, daman Sea. Remarks: found in man25–32; PDS 8–10. Body relatively belly and operculum cycloid. Sensory grove swamps and brackish estuaries.
(K. Shibukawa)short, compressed. Head slightly de- canals and pores present on head; longi-

201

Hemigobius mingi(Herre, 1936)
Blue-eyed Mullet Goby
D VI + I, 6–8; A I, 6–7; P1 13–15; LR
27–32; PDS 8–10. Body relatively
short, compressed. Head slightly depressed. Snout rounded, sometimes alittle projecting beyond jaws anteriorly.
Jaws subterminal. Interorbital spacebroad. Gill opening not extending anteriorly to a vertical through preopercularmargin. Pelvic fins united medially;
frenum present. Scales ctenoid, excepting those on predorsal area, pectoral
base, breast, cheek and operculum cycloid. Sensory canals and pores present
on head; longitudinal pattern of sensory-papillae rows on cheek. Color: headand body dark grayish brown dorsally,
becoming paler ventrally; typically 4–5
faint narrow beige saddles on body; adistinct black spot on posterior part offirst dorsal fin. Size: 6 cm SL. Distribution: West Pacific. Remarks: foundin mangrove swamps and brackish estuaries.
(K. Shibukawa)

Hemigobius mingi, UMTF 1504 (KAUM–I. 16709), 3.7 cm SL
Setiu, 27 Oct. 2008

Istigobius decoratus, UMTF 1252 (KAUM–I. 16563), 4.7 cm SL
Bidong Island, 14 Oct. 2008

Istigobius decoratus(Herre, 1927)
Decorative Lagoon Goby
D VI + I, 11; A I, 10; P1 17–19; LR
30–33; PDS 8–10. Body moderately
elongate and compressed. Head slightly depressed. Snout rounded, slightly
projecting beyond jaws. Jaws subequal.
Gill opening not extending anteriorly toa vertical through preopercular margin.
No free, filamentous rays on upper partof pectoral fin. Pelvic fins united medially; frenum present. Scales ctenoid,
excepting breast, pectoral-fin base andoccipital region with cycloid scales;
head naked. Sensory canals and porespresent on head; longitudinal pattern of
sensory-papillae rows on cheek. Color: head and body pale grayish brown
dorsally, becoming paler ventrally;
body with a midlateral series of blackspots, as well as many dark brownsmaller spots and grayish brown reticulation; a distinct black spot between
first and second spines of first dorsal
fin. Size: 10 cm SL. Distribution: Indo-West Pacific. Remarks: found inshallow coral reefs with sandy bottoms.
(K. Shibukawa)

Istigobius diadema, KAUM–I. 16805, 4.9 cm SL
Merang, 2 Nov. 2008

Istigobius diadema, UMTF 1111 (KAUM–I. 16287), 3.3 cm SL
Cendering, 14 Sept. 2008

Istigobius diadema(Steindachner, 1876)
Black-lined Lagoon Goby
D VI + I, 11; A I, 9; P1 19; LR 31–33;
PDS 18. Body moderately elongate
and compressed. Head slightly depressed. Snout rounded, slightly projecting beyond jaws. Jaws subequal.
Gill opening not extending anteriorly
to a vertical through preopercular margin. No free, filamentous rays on upper part of pectoral fin. Pelvic finsunited medially; frenum present. Scales

ctenoid, excepting breast, pectoral-finbase and occipital region with cycloid
scales; head naked. Sensory canals
and pores present on head; longitudinal pattern of sensory-papillae rows
on cheek. Color: head and body palegrayish brown, with a distinct horizontal black band behind eye. Size: 10
cm SL. Distribution: Eastern Indian
Ocean, Andaman Sea and Indonesia.
Remarks: found in mangrove creeks
and adjacent coastal waters with sandyor muddy bottoms. (K. Shibukawa)

202

Istigobius goldmanni(Bleeker, 1852)
Goldman's Goby
D VI + I, 10–11; A I, 9; P1 17–19; LR
26–27; PDS 7–9. Body moderately
elongate, compressed. Head subcylindrical or slightly depressed. Snout
rounded, slightly projecting beyond
jaws. Jaws subterminal. Gill opening
not extending anteriorly to a verticalthrough preopercular margin. No free,filamentous rays on upper part of pectoral fin. Pelvic fins united medially;
frenum present. Scales ctenoid, excepting breast, pectoral-fin base andoccipital region with cycloid scales;
head naked. Sensory canals and porespresent on head; longitudinal patternof sensory-papillae rows on cheek.
Color: head and body pale gray, withnumerous minute black spots. Size: 4
cm SL. Distribution: West Pacific.
Remarks: found in shallow coral
reefs with sandy bottoms.
(K. Shibukawa)

Istigobius goldmanni, UMTF 1479 (KAUM–I. 16541), 2.5 cm SL
Cendering, 6 Oct. 2008

Istigobius nigroocellatus, UMTF 1388 (KAUM–I. 16561), 2.3 cm SL
Bidong Island, 13 Oct. 2008

Istigobius nigroocellatus(Günther, 1873)
Black Spotted Goby
D VI + I, 10–12; A I, 9–10; P1 17; LR
28; PDS 9. Body moderately elongate,
compressed. Head subcylindrical orslightly depressed. Snout rounded,slightly projecting beyond jaws. Jaws
subterminal. Gill opening not extending anteriorly to a vertical throughpreopercular margin. No free, filamentous rays on upper part of pectoral
fin. Pelvic fins united medially; frenum present. Scales ctenoid, excepting breast, pectoral-fin base and occipital region with cycloid scales;
head naked. Sensory canals and porespresent on head; longitudinal patternof sensory-papillae rows on cheek.
Color: head and body pale gray, withnumerous minute black spots; a distinct black spot between fifth and sixth
spines of first dorsal fin. Size: 4 cm
SL. Distribution: West Pacific. Remarks: found in shallow coral reefs
with sandy bottoms.
(K. Shibukawa)

Istigobius ornatus, UMTF 1190 (KAUM–I. 16676), 7.4 cm SL
Kemaman, 21 Oct. 2008

Istigobius ornatus, UMTF 1058 (KAUM–I. 16291), 4.0 cm SL
Cendering, 14 Sept. 2008

LR 27–28; PDS 9–12. Body moderately elongate and compressed. Headsubcylindrical or slightly depressed.
Snout rounded, slightly projecting beyond jaws. Jaws subequal. Gill opening not extending anteriorly to a vertical through preopercular margin.
Upper part of pectoral fin with somefilamentous free rays. Pelvic fins unitIstigobius ornatus
ed medially; frenum present. Scales(Rüppell, 1830)
ctenoid, excepting breast, pectoral-fin
Ornate Gobybase and occipital region with cycloid
D VI + I, 10–11; A I, 8–10; P1 17–20; scales; head naked. Sensory canals

and pores present on head; longitudinal pattern of sensory-papillae rows
on cheek; curved or near transverse
sensory-papillae row just behind chin.
Color: head and body light grayish
yellow dorsally, becoming paler ventrally; body with numerous small
black dots or horizontally-elongate
blotches; distal tip of first dorsal fintinged with yellow. Size: 10 cm SL.
Distribution: Indo-West Pacific. Remarks: found in shallow silty reefs
and mangroves.
(K. Shibukawa)

203“Lophogobius” bleekeri
Popta, 1921
Ornate Goby
D VI + I, 8; A I, 7; P1 16–18; LR
25–28; PDS 0. Body short and compressed. Head compressed. Lower jaw
a little projecting beyond upper jaw.
Gill opening not extending anteriorly
to a vertical through preopercular margin. Pelvic fins united medially; frenum present. Scales on body largely
ctenoid; pectoral-fin base and narrow
area on anterior part of body with cycloid scales; head naked. Sensory canals and pores present on head; distinct transverse rows of sensory
papillae on cheek. Color: head andbody beige, with several irregular,
broad vertical blackish brown bands;
sometimes entirely blackened in particular the large specimen. Size: 4 cm
SL. Distribution: West Pacific. Remarks: found in lower reaches of
streams and brackish estuaries. Hitherto this species has been mis-assigned
to Lophogobius; the author and his
collaborators are now tackling withthis issue.
(K. Shibukawa)“Lophogobius” bleekeri, UMTF 1460 (KAUM–I. 16806), 2.2 cm SL
Merang, 2 Nov. 2008

Macrodontogobius wilburi, UMTF 1331 (KAUM–I. 16618), 3.4 cm SL
Bidong Island, 12 Oct. 2008

Macrodontogobius wilburi
Herre, 1936
Wilbur's Goby
D VI + I, 9–11; A I, 9–10; P1 15–17;
LR 27–31; PDS 7. Body moderately
elongate, compressed. Head subcylindrical or slightly depressed. Snout
rounded, slightly projecting beyond
jaws. Lower jaw not projecting anteriorly beyond upper jaw. Gill opening
not extending anteriorly to a verticalthrough preopercular margin. Pelvic
fins united medially; frenum present.
Scales on body ctenoid, excepting
isthmus with cycloid scales; cheek andoperculum with ctenoid scales. Sensory canals and pores present on head;
many transverse rows of sensory papillae on cheek. Color: head and bodylight grayish brown, with numerous
minute black spots; anal fin with faintdiagonal dusky barred pattern. Size: 5
cm SL. Distribution: Indo-West Pacific. Remarks: found in shallow lagoons and coral reefs with rubbles.
(K. Shibukawa)

Mugilogobius chulae, KAUM–I. 16713, 2.7 cm SL
Setiu, 27 Oct. 2008

Mugilogobius chulae(Smith, 1932)
Two-spot Mangrove Goby
D VI + I, 6–8; A I, 6–8; P1 12–16; LR
25–32; PDS 11–15. Body moderately
elongate and compressed. Head slightly depressed. Interorbital space broad.
Gill opening not extending anteriorly
to a vertical through preopercular margin. Some spines of first dorsal fin
elongate and filamentous. Pelvic finsunited medially; frenum present.
Scales ctenoid, excepting those on predorsal area, pectoral base, belly and up-

per part of operculum with cycloid
scales; other part of head naked. Sensory canals and pores absent on head;longitudinal pattern of sensory-papillae rows on cheek. Color: head andbody light grayish yellow or beige,
with several diagonal black bars andspeckles; two black spot at caudal finbase; black spot at posterior part offirst dorsal fin. Size: 4 cm SL. Distribution: Indo-West Pacific. Remarks:
found in shallow mangrove swamps
and brackish estuaries.allow coral
reefs with sandy bottoms.
(K. Shibukawa)

204

Mugilogobius tigrinus
Larson, 2001
Narrow-barred Mangrove Goby
D VI + I, 6–7; A I, 6–7; P1 14–17; LR
24–28; PDS 9–12. Body moderately
elongate and compressed. Head slightly depressed. Interorbital space broad.
Gill opening not extending anteriorly
to a vertical through preopercular margin. Some spines of first dorsal fin
may be elongate and filamentous. Pelvic fins united medially; frenum present. Scales ctenoid, excepting those on
predorsal area, pectoral base, belly
and upper part of operculum with cycloid scales; other part of head naked.
Sensory canals and pores absent onhead; longitudinal pattern of sensorypapillae rows on cheek. Color: headand body whitish with four vertical
black bands, in addition to three black
saddles and single black spot at caudal-fin base; first dorsal fin with ablack spot. Size: 2.3 cm SL. Distribution: Thailand, Malaysia and Singapore. Remarks: found in shallow mangrove creeks and swamps.
(K. Shibukawa)

Mugilogobius tigrinus, UMTF 1261 (KAUM–I. 16740), 2.1 cm SL
Setiu, 29 Oct. 2008

Paratrypauchen microcephalus, KAUM–I. 17081, 8.9 cm SL
off Terengganu, 27 Dec. 2008

Paratrypauchen microcephalus(Bleeker, 1860)
Comb Goby

Periophthalmus argentilineatus, UMTF 1133 (KAUM–I. 16493), 5.0 cm SL
estuary near UMT, 30 Sept. 2008

D VI, 48–51; A 48–51; P1 16–17; LR
52–65; PDS 0. Body very elongate
and compressed, with prominent midventral keel on belly. Head compressed. A pouch-like cavity justabove operculum. Eye reduced, difficult to be discerned from externalview. Dorsal and anal fins confluent
with caudal fin. Pelvic fins fused medially, but connecting membrane between innermost rays emarginated
Periophthalmus argentilineatus, UMTF 1039 (KAUM–I. 16550), 3.4 cm SL
posteriorly; frenum low. Scales on
estuary near UMT, 30 Sept. 2008
body cycloid; head and belly naked.
Sensory canals and pores absent onhead; numerous sensory papillae scat- Periophthalmus argentilineatus canals and pores on head. Color: headand body grayish brown with indistered on cheek. Color: head and body
Valenciennes, 1837
tinct dusky mottles; first dorsal fin
entirely reddish. Size: 18 cm SL. DisSilver-lined Mudskipper
with many white spots, in addition to atribution: Indo-West Pacific. ReD IX–XVI + I, 9–12; A I, 8–11; P1 broad distal submarginal black band.
marks: found in bays and brackish
12–14; LR 64–100; PDS 22–37. Body Size: 9 cm SL. Distribution: Indoestuaries with muddy bottom.
(K. Shibukawa) moderately elongate, compressed. Pacific. Remarks: found in schools in
Head slightly compressed. Snout intertidal zone of mangrove areas andsteep. Dermal cup-like process devel- brackish estuaries; feeds on algal mats
oped beneath eye. Single row of coni- and small invertebrates on the sand /cal teeth on each jaws. Pelvic fins al- mud flats.
most separated; frenum absent. Scales(M. A. Ghaffar & K. Shibukawa)on head and body cycloid. No sensory

205

Periophthalmus novemradiatus(Hamilton, 1822)
Dusky-gilled Mudskipper
D IX–XI + I, 12–13; A I, 11–13; P1
11–14; LR 61–76; PDS 20–28. Body
moderately elongate, compressed.
Head slightly compressed. Snout
steep. Dermal cup-like process developed beneath eye. Single row of conical teeth on each jaws. Pelvic fins almost separated with serrated margin;
frenum present. Scales on head andbody cycloid. No sensory canals andpores on head. Color: head and bodygrayish brown with indistinct dusky
mottles; first dorsal fin with several
dusky spots tinged with red, in addition to broad distal submarginal blackband. Size: 10 cm SL. Distribution:
Eastern Indian Ocean, Andaman Sea
and West Pacific. Remarks: normallyfound in large schools on the sand flatsof estuaries; feeds on small invertebrates and algal mats.
(M. A. Ghaffar & K. Shibukawa)

Periophthalmus novemradiatus, UMTF 1505 (KAUM–I. 16741), 5.4 cm SL
Setiu, 30 Oct. 2008

Pleurosicya labiata, UMTF 1483 (KAUM–I. 16552), 1.9 cm SL
Bidong Island, 13 Oct. 2008

Pleurosicya labiata(Weber, 1913)
D VI+I, 7–9; A I, 8–9; P1 15–18; LR
24–27; PDS 9–11. Body relatively
moderately elongate, compressed.
Head slightly depressed. Snout pointed. Upper jaw projecting anteriorly
beyond lower jaw; lower jaw triangular. Gill opening not extending anteriorly to a vertical through preopercularmargin. Pelvic fins united medially;
frenum present; skin around pelvic-fin
spine thickened, forming a lobe-likeappearance. Scales on body ctenoid;
Priolepis semidoliata, UMTF 1390 (KAUM–I. 16388), 2.4 cm SL
Dungun, 24 Sept. 2008
head naked. Sensory canals and porespresent on head; reduced longitudinalpattern of sensory-papillae rows ongate, filamentous. Pelvic fins united
Priolepis semidoliata
cheek. Color: head and body submedially; frenum absent. Scales on(Valenciennes, 1837)
translucent, with numerous minutebody ctenoid; head naked. Sensory cabrown speckles. Size: 2.5 cm SL. DisHalfbarred Goby
nals and pores absent on head; reduced
tribution: Andaman Sea and West PaD VI+I, 9–10; A I, 7–8; P1 16–18; LR longitudinal pattern of sensory-papilcific. Remarks: found in coral reefs; 25–28; PDS 0. Body relatively short, lae rows on cheek. Color: groundcommensal with sponges.
compressed posteriorly. Head subcy- color of head and body orange yellow(K. Shibukawa) lindrical or slightly depressed. Lower to dark grayish brown; several vertical
jaw projecting anteriorly a little be- or oblique pale bars on head and body,yond upper jaw. Gill opening extend- two of them around end of head coning slightly beyond a vertical through nected one another at dorsum. Size: 2
preopercularmarginanteriorly. cm SL. Distribution: Indo-Pacific.
Tongue near truncate or rounded. Remarks: found in shallow coral
Some spines of first dorsal fin elon- reefs.
(K. Shibukawa)

206

Psammogobius biocellatus(Valenciennes, 1837)
Sleepy Goby
D VI + I, 9; A I, 8–9; P1 18; LR 29–
30; PDS 14–15. Body moderately
elongate and subcyrindrical, compressed. Head depressed. Lower jaw
prominent. Tongue bilobate. Small
lappet over iris. Gill opening extending anteriorly beyond a verticalthrough preopercular margin; gillmembranes united medially, with freeposterior margin across isthmus. Pelvic fins united medially; frenum present. Scales ctenoid, excepting those on
belly and pectoral base cycloid. Sensory canals and pores present on head;longitudinal pattern of sensory-papillae rows on cheek. Color: head andbody blackish or grayish brown withindistinct black mottles and lines.
Size: 12 cm TL. Distribution: IndoWest Pacific. Remarks: found in
mangrove swamps, brackish estuaries
and adjacent coastal waters with sandy-mud bottoms.
(K. Shibukawa)Psammogobius biocellatus, UMTF 1381 (KAUM–I. 16752), 3.6 cm SL
Setiu, 30 Oct. 2008

Pseudogobius javanicus, KAUM–I. 16294, female, 2.8 cm SL
Cendering, 14 Sept. 2008

Pseudogobius javanicus(Bleeker, 1856)
Java Fat-nose Goby
D VI + I, 7; A I, 7; P1 14–16; LR
24–28; PDS 7–8. Body moderately
elongate, compressed. Head slightly
depressed. Snout rounded, projecting
beyond upper jaw. Gill opening notextending anteriorly to a verticalthrough preopercular margin. Pelvic
fins united medially; frenum present.
Scales ctenoid, excepting those on
predorsal area, pectoral base, belly
and operclum cycloid. Sensory canals
and pores present on head; longitudinal pattern of sensory-papillae rows
on cheek. Color: head and body pale
gray, with numerous dusky spots and a
diagonal dusky bar below first dorsal

Pseudogobius javanicus, UMTF 1467 (KAUM–I. 16494), male, 2.6 cm SL
estuary near UMT, 30 Sept. 2008

fin; black spot at posterior part of firstdorsal fin; rays of second dorsal andcaudal fins with several dusky dots;
two black spots at caudal-fin base.
Size: 3 cm SL. Distribution: West Pa-

cific and Andaman Sea. Remarks:
Found in mangrove swamps andbrackish estuaries with sandy-mud
bottoms.
(K. Shibukawa)

207

Pseudogobius melanostictus(Day, 1876)
Black Spotted Fat-nose Goby
D VI+I, 7; A I, 7; P1 14–15; LR 25–
26; PDS 7–8. Body moderately elongate, compressed. Head slightly depressed. Snout rounded, projecting
beyond upper jaw. Gill opening notextending anteriorly to a verticalthrough preopercular margin. Pelvic
fins united medially; frenum present.
Scales ctenoid, excepting those on
pectoral base and belly cycloid; operculum with ctenoid and cycloid scales.
Sensory canals and pores present onhead; longitudinal pattern of sensorypapillae rows on cheek. Color: headand body pale gray, with numerous
dusky spots; 2 broad, irregular black
stripes on first dorsal fin; rays of second dorsal and caudal fins with several
dusky dots; two black spots at caudalfin base. Size: 4 cm SL. Distribution:
Indo-West Pacific. Remarks: Found
in mangrove swamps and brackish estuaries with sandy-mud bottoms.
(K. Shibukawa)

Pseudogobius melanostictus, KAUM–I. 16817, female, 3.3 cm SL
estuary near UMT, 3 Dec. 2008

Pseudogobius melanostictus, UMTF 1426 (KAUM–I. 16486), male, 2.6 cm SL
estuary near UMT, 30 Sept. 2008

Redigobius balteatus(Herre, 1935)
Rhinohorn Goby
D VI + I, 7–8; A I, 6–7; P1 17–19; LR
27–28; PDS 9–11. Body relatively
deep, compressed. Head subcylindrical or slightly compressed. Gill opening not extending anteriorly to a vertical through preopercular margin.
Pelvic fins united medially; frenum
present. Scales on body cycloid or
weak ctenoid anteriorly, ctenoid posteriorly; breast, belly and pectoral-finbase with cycloid scales; dorsal part of
operculum with ctenoid scales; cheeknaked. Sensory canals and pores present on head; longitudinal pattern of
sensory-papillae rows on cheek. Color: ground color of head and bodygrayish yellow dorsally, pale beige
ventrally; a conspicuous, broad diagonal black bar from first dorsal fin to
belly; a narrow diagonal black bandfrom eye to middle of operculum.
Size: 3 cm SL. Distribution: IndoWest Pacific. Remarks: found in
mangrove swamps and brackish estuaries.
(K. Shibukawa)

Redigobius balteatus, KAUM–I. 16803, 3.0 cm SL
Merang, 2 Nov. 2008

Redigobius balteatus, UMTF 1271 (KAUM–I. 16774), 2.4 cm SL
Merang, 2 Nov. 2008

208

Stigmatogobius pleurostigma(Bleeker, 1849)
Peach Knight Goby
D VI + I, 7; A I, 7–8; P1 17–21; LR
23–28; PDS 7–13. Body relatively
short, compressed. Head slightly compressed. Jaws subequal. Gill opening
not extending anteriorly to a verticalthrough preopercular margin. Anterior
margin of tongue rounded. Middle
some spines of first dorsal fin elongate, forming tall pointed fin; pelvic
fins united medially; frenum present.
Scales ctenoid, excepting those onnape, opercle, breast and pectoral-finbase cycloid. Sensory canals and porespresent on head; transvers rows of
sensory papillae on cheek. Color:head and body grayish brown or beige,
darkened dorsally; a series of blackspots on midlateral body (some additional black spots frequently found inlarge specimen); a small black spot
between first and second spines offirst dorsal fin; many small black spots
on caudal fin. Size: 6 cm SL. Distribution: West Pacific. Remarks:
found in mangrove creeks and brackish estuaries.
(K. Shibukawa)

Stigmatogobius pleurostigma, UMTF 1237 (KAUM–I. 16739), 3.5 cm SL
Setiu, 29 Oct. 2008

Trypauchen vagina, KAUM–I. 17300, 17.9 cm SL
off Terengganu (KT), 20 Jan. 2009

Trypauchen vagina, KAUM–I. 17302, 13.5 cm SL
off Terengganu (KT), 20 Jan. 2009

Trypauchen vagina(Bloch & Schneider, 1801)
Pink Worm Goby
D VI, 40–50; A39–47; P1 15–20; LR
80–115; PDS 0. Body very elongate
and compressed. Head compressed. A
pouch-like cavity just above operculum. Eye reduced, difficult to be discerned from external view. Dorsal andanal fins confluent with caudal fin.
Pelvic fins fused medialy, but conValenciennea muralis, KAUM–I. 16500, 9.1 cm SL
necting membrane between innermost
Cendering, 6 Oct. 2008
rays emarginated posteriorly; frenum
low. Scales on body cycloid; head naon belly and breast, if present, cycloid;ked. Sensory canals and pores absent
Valenciennea muralis
head and pectoral-fin base naked. Senon head; numerous sensory papillae(Valenciennes, 1837)
sory canals and pores present on head;scattered on cheek. Color: head andMural Goby
longitudinal pattern of sensory-papilbody entirely reddish. Size: 15 cm SL.
D VI + I, 11–13; A I, 11–13; P1 18– lae rows on cheek. Color: head andDistribution: Indo-West Pacific. Remarks: found in bays and brackish 21; LR 73–94; PDS 0. Body moder- body pale gray or pale beige, with 4-5
ately elongate, compressed. Head sub- longitudinal narrow pink or dusky
estuaries with muddy bottom.
(K. Shibukawa) cylindrical. Lower jaw not projecting stripes; lips yellowish; black spot atanteriorly beyond upper jaw. Gill distal part of first dorsal fin just beopening relatively narrow, restricted hind third spine. Size: 10 cm SL. Disto pectoral-fin base. Third spine of tribution: Indo-West Pacific. Refirst dorsal fin slightly elongate, form- marks: found on shallow coastal
ing pointed fin. Pelvic fins separated; waters with flat sandy bottoms andfrenum absent. Caudal fin rounded. rubbles.
(K. Shibukawa)
Scales small ctenoid, excepting those209

Valenciennea sexguttata(Valenciennes, 1837)
Sixspot Goby
D VI + I, 11–13; A I, 11–13; P1 19–
21; LR 71–94; PDS 0. Body moderately elongate, compressed. Headslightly compressed. Lower jaw notprojecting anteriorly beyond upper
jaw. Gill opening relatively narrow,
restricted to pectoral-fin base. Third
spine of first dorsal fin slightly elongate, forming pointed fin. Pelvic finsseparated; frenum absent. Caudal fin
rhomboid. Scales small ctenoid, excepting those on belly, prepelvic region and pectoral-fin base, if present,
cycloid; head naked. Sensory canals
and pores present on head; longitudinal pattern of sensory-papillae rows
on cheek. Color: head and body pale
gray or pale beige; several sky spots
on cheek and operculum; narrow longitudinal faint pinkish stripe on body;
black spot on distal part of first dorsal
fin just behind third spine. Size: 15 cm
SL. Distribution: Indo-Pacific. Remarks: found on flat sandy-mud bottoms with rubbles in shallow coastal
waters.
(K. Shibukawa)

Valenciennea sexguttata, UMTF 1304 (KAUM–I. 16616), 4.8 cm SL
Bidong Island, 12 Oct. 2008

Valenciennea wardii, KAUM–I. 17191, 8.5 cm SL
off Terengganu (KR), 5 Jan. 2009

Valenciennea wardii(Playfair, 1867)
Ward's Goby
D VI + I, 11–12; A I, 11–12; P1 19–
22; LR 70–88; PDS 0. Body moderately elongate, compressed. Headslightly compressed. Lower jaw notprojecting anteriorly beyond upper
jaw. Gill opening relatively narrow,
restricted to pectoral-fin base. First
dorsal fin rounded, without elongate
spines. Pelvic fins separated; frenum
absent. Caudal fin rounded. Scales
small ctenoid, excepting those on belly, prepelvic region and pectoral-finbase, if present, cycloid; head naked.
Sensory canals and pores present onhead; longitudinal pattern of sensorypapillae rows on cheek. Color: headand body pale gray or pale beige, withthree broad vertical brown bars andsome small brown saddles; a narrowbright blue oblique stripe throughcheek and operculum; distinct black
ocellated spot on rear part of first dorsal fin; posterior part of caudal fin
brown, with pale distal margin; small
black spot on dorsal part of caudal fin.

Yongeichthys nebulosus, UMTF 1038 (KAUM–I. 16766), 5.9 cm SL
Setiu, 31 Oct. 2008

Size: 10 cm SL. Distribution: IndoWest Pacific. Remarks: found on
sandy-mud or soft mud bottom inbays.
(K. Shibukawa)

Yongeichthys nebulosus(Forsskål, 1775)
Shadow Goby
D VI + I, 9; A I, 9; P1 17–19; LR
25–30; PDS 0. Body moderately elongate, compressed. Head slightly compressed. Jaws subequal. Gill opening
not extending anteriorly to a verticalthrough preopercular margin. Pelvic

fins united medially; frenum present.
Scales ctenoid, excepting those on
breast and pectoral-fin base cycloid;
head naked. Sensory canals and porespresent on head; longitudinal patternof sensory-papillae rows on cheek,some rows multiple. Color: body pale
gray or pale brown, with two dusky
saddles and three large dusky midlateral spots. Size: 10 cm SL. Distribution: Indo-West Pacific. Remarks:
found in mangrove creeks, brackish
estuaries and protected bays with sandy-mud bottoms.
(K. Shibukawa)

210

PTERELEOTRIDAE
Dart Gobies

By Koichi Shibukawa
Small to moderate-sized (up to
ca. 18 cm SL, commonly to 10 cm),
bottom-oriented free swimming fishes. Body elongate, sometimes highly
compressed; head more or less compressed; no pored or tubed lateral-line
scales on body. Teeth conical, forming some or several rows in each jaw;
minute sensory papillae developedon head (and few on body and caudalfin), forming transverse or longitudinal rows; five branchiostegal rays.
Usually two dorsal fins (continuous in
one species of Ptereleotris), first with
IV–VI flexible spines and second
with single flexible spine and 9–39
soft rays; anal fin with single flexible
spine and 14–37 soft rays; pectoral fin
with 10–25 soft rays; pelvic fins separated, with single spine and 3–5 softrays in each; caudal fin rounded, truncate or often emarginate. Scales small
cycloid (except for Nemateleotris

head compressed
scales mostly embedded and non-imbricate
5 branchiostegals
pelvic fins separated

and some species of Ptereleotris with
ctenoid scales), mostly embedded andnonimbricate. Color: variable.
Remarks: typically found in coral
reefs and brackish estuaries. Several
species often treated as aquarium fish.
Similar families occurring in thearea: Blenniidae – teeth incisor-like,forming single row in each jaw. Eleotridae – six branchiostegal rays; headsubcylindrical or rather depressedin many species; scales imbricate inmost species. Gobiidae – pelvic finsusually united medially; scales, if
present, imbricate in most species.

Parioglossus formosus(Smith, 1931)
Beautiful Hover Goby
D V–VI + I, 13–15; A I, 13–15; P1
15–17; LR 66–78; PDS 0. Body elongate, compressed. Head compressed.
Lower jaw slightly projecting anteriorly beyond upper jaw. Gill opening
not extending anteriorly to a verticalline through middle of operculum.
Middle some spine of first dorsal fin
slightly elongate in male. Pelvic finsseparated. Scales on body small cycloid, embedded and non-imbricate;head naked. Sensory canals and porespresent on head; reduced transverse
pattern of sensory-papillae rows on
cheek. Color: head and body pale

Parioglossus formosus, UMTF 1266 (KAUM–I. 16340), 2.0 cm SL
Cendering, 16 Sept. 2008

beige, with a longitudinal black stripe
from snout to posterior margin of caudal fin through eye and ventral half ofbody; additional black stripe along
dorsal profile from snout to base of
caudal fin, continuous to dark brown
stripe on dorsal part of caudal fin.

Size: 3.5 cm SL. Distribution: Andaman Sea, northwestern Australia andWest Pacific. Remarks: found in shallow coastal area of coral or rocky reefs
at high-tide hour; free swimming species, usually forming aggregations.

211

EPHIPPIDAE

Spadefishes (Batfishes)
By Keiichi Matsuura and Seishi Kimura
Medium sized (up to 20 cm)
marine fishes. Body deep, orbicular and strongly compressed. Headshort, its length shorter than half ofbody depth; interorbital region andanterior part of head naked or head
covered with scales. Mouth small,rear end of upper jaw not reachingvertical through anterior edge of eye;upper jaw not protrusible; jaws covered with bands of slender setiform
or compressed teeth. Dorsal fin single
or deeply notched before soft-rayedpart, with V–IX spines and 19–38
soft rays; anal fin with III spines and
15–27 soft rays; pectoral fins rounded
and usually shorter than head; caudal
fin truncate or wedge shaped. Scales
large and smooth, or small and ctenoid. Lateral line complete. Color:
adults silvery, silvery blue-green, or
yellowish silvery; frequently withvertical dark bars.
Remarks: occurring in various
shallow water habitats including es-

single dorsal fin with V–IX, 19–38

body deep,
orbicular

anal fin with III, 15–27

tuaries, harbors, and coral reefs. Marketed fresh but with small commercial
importance.
Similar families occurring in thearea: Drepaenidae – upper jaw greatly protrusible; pectoral fins elongate
reaching posterior part of anal fin base.
Chaetodontidae – dorsal fin with VI–

XVII spines and 14–34 soft rays; anal
fin with III–V spines and 15–24 soft
rays. Monodactylidae – pelvic fins
rudimentary or absent. Scatophagidae– dorsal fin notched or divided to base
before soft rayed part, with XI or XII
spines and 15–18 soft rays; anal fin
with IV spines and 13–17 soft rays.

Ephippus orbis(Bloch, 1787)
Spadefish
D IX, 19–20; A III, 15–17; P1 18–19;
LL 39–43. Body orbicular, strongly
compressed, head length less than halfof body depth. Mouth small, upper
jaw not reaching posteriorly verticalthrough anterior edge of eye; upper
jaw not protrusible; jaws covered withbands of slender incisiform teeth. Preopercle distinctly serrate with a wide
naked margin. Single dorsal fin with adeep notch before soft rayed part; pectoral fins shorter than head. Scales
relatively large and smooth. Lateralline complete. Color: head and bodysilvery blue-green; 4 to 5 vertical darkbars frequently on body; fins dusky.
Size: maximum size 20 cm. Distribution: tropical regions of the Indo-West
Pacific from India to Indonesia, northto southern Japan.
(K. Matsuura)

Ephippus orbis, KAUM–I. 16900, 11.5 cm SL
off Terengganu (KT), 6 Dec. 2008

212

Platax orbicularis(Forsskål, 1775)
Copper Batfish
D V, 34–39; A III, 25–29; P1 16–18;
LL 44–52. Body orbicular, strongly
compressed. Snout short, dorsal contour somewhat concave in large adults.
Five pores on each side of lower jaw.
Tricuspid teeth on jaws; middle cusp
larger than lateral cusps; no teeth onvomer or palatines. Caudal fin truncate. Color: body yellowish silver
with 2 vertical black bands in adults;
juvenile reddish brown with small
white ocellii on body. Size: maximum
length 50 cm. Distribution: IndoWest Pacific. Remarks: found on nearreefs in adults. Juveniles appear inmangrove areas, mimic floating leaves.
(S. Kimura)

Platax orbicularis, KAUM–I. 17059, 20.1 cm SL
off Terengganu (KT), 24 Dec. 2008

213

SCATOPHAGIDAE
Scats

By Keiichi Matsuura and Mazlan Abdul Ghaffar
Body greatly compressed, reaching to 32 cm. Dorsal profile of headsteeply ascending to dorsal fin origin; snout rounded. Mouth small, terminal; jaws not protrusible; villiform
teeth in several rows on jaws. Eyerelatively large, its diameter slightly
shorter than snout length. Dorsal fin
with XI or XII spines and 16–18 softrays; first dorsal fin spine procumbent; a deep notch between spinousand soft portions of dorsal fin. Anal
fin with IV spines and 13–16 soft
rays. Pectoral fin with 16 or 17 soft
rays. Caudal fin rounded in juveniles,
truncate or slightly emarginated in
adults. Head and body covered withsmall cycloid scales. Color: headand body greenish or silvery withmany black spots or bars.
Remarks: occurring in estuaries,
harbors, and lower reaches of rivers.
Marketed fresh but with small com-

dorsal fin with XI–XII, 16–18

mouth small,
not protrusible
body deepand compressedanal fin with IV, 13–16

mercial importance.
Similar families occurring in
the area: Chaetodontidae – lacking a
deep notch between spinous and soft
parts of dorsal fin; jaws protrusible.
Drepaenidae – anal fin with III spines;

jaws protrusible; pectoral fins elongate, reaching beyond anal fin origin.
Pomacanthidae – preopercle with astrong, backwardly pointed spine; nodeep notch between spinous and soft
parts of dorsal fin.

Scatophagus argus(Linnaeus, 1766)
Spotted Scat
D XI, 16–18; A IV, 14–15; P1 16.
Body quadrangular, strongly compressed. Dorsal profile of head steeply
ascending to dorsal fin origin; snout
rounded. Mouth small, terminal; jaws
protrusible; villiform teeth in severalrows on jaws. Teeth villiform, making
several rows on jaws. Eye relatively
large, its diameter slightly shorter thansnout length. Color: ground color ofhead and body greenish; juveniles
with a few large round blotches being
almost equal to eye in size, or with
5–6 wide dark, vertical bars; adults
with less distinct dark blotches ormarkings. Size: maximum size 35 cm.
Distribution: tropical regions of Indo-West Pacific. Remarks: euryhaline fish commonly found in estuarine
ecosystem extended to uppermost
streams freshwater river tributaries.
Named based on their feeding habit(Scat / human feces feeder = Scato +phagus). However, recent researchfinding denoted that they are omnivorous (subsisting mainly on plant material and detritus).
(K. Matsuura & M. A. Ghaffar)

Scatophagus argus, KAUM–I. 17148, 9.3 cm SL
Setiu, 3 Jan. 2009

214

SIGANIDAE
Rabbitfishes

By Keiichi Matsuura, Seishi Kimura and Mizuki Matsunuma
Body oval, strongly compressed,
single dorsal fin with usually XIII, 10
reaching to 55 cm. Mouth small, terminal; jaws not protrusible; a single
row of compressed, incisiform teeth
recumbent
spine
on jaws. Dorsal fin with XIII spinesand 10 soft rays, preceded by embedded, recumbent spine. Anal fin with
VII spines and 9 soft rays. Pelvic finswith an inner and outer spines andseparated 3 soft rays. Fin spines witha pair of grooves containing venom
glands. Body covered with small,
cycloid scales. Color: species inhabiting coral reefs having bright color
with reticulations and markings, andother species with drab mottled patanal fin with VII, 9tern.
Remarks: dwelling on bottom marily herbivorous. Caught by bottom area: Acanthuridae and Chaetodonin shallow coastal waters. Some spe- trawlers, traps, set nets and by spear. tidae – I spine and 5 soft rays in pelvic fins.
cies found in coral reefs and others Marketed fresh.
Similar families occurring in thein mangroves and seagrass beds. Pri-

Siganus fuscescens(Houttuyn, 1782)
Mottled Spinefoot
D XIII, 10; A VII, 9; P1 15–17. Body
oval, relatively slender, its depth 2.3–
2.9 in SL. Last anal-fin spine 1.3–1.5
in longest anal-fin spine. Soft parts of
dorsal and anal fins low, longest dorsal-fin ray 0.5–1 times in longest dorsal-fin spine. Caudal fin slightly concave in individuals smaller than 10 cm
SL, and becoming more concave withgrowth. Color: greatly variable, but
body ground color olive green or lightbrown on sides and back, silvery on
ventral half; many small light blue
spots scattered on head and body.
Size: maximum size 40 cm, commonly to 25 cm. Distribution: western
coast of Malay Peninsula and West Pacific from Thailand eastward to New
Caledonia, northward to the Ryukyu
Islands, and southward to northern
Australia.
(K. Matsuura)

Siganus fuscescens, KAUM–I. 16844, 18.7 cm SL
off Terengganu (KT), 5 Dec. 2008

Siganus fuscescens, UMTF 1104 (KAUM–I. 16375), 9.3 cm SL
Dungun, 23 Sept. 2008

215

Siganus guttatus(Bloch, 1787)
Orangespotted Spinefoot
D XIII, 10; A VII, 9; P1 15–17. Body
oval, strongly compressed, its depth
1.8–2.3 in SL. Snout short, dorsal profile of head steep anteriorly, somewhat
concave above eye. Fifth to eighth dorsal-fin spine longest. Last anal-fin
spine longest, longer than longest dorsal-fin spine. Caudal fin emarginate butmoderately forked in large adults.
Scales minute. Color: many large tomedium-sized orange to bronze spots
scattered laterally on body with a brightyellow spot at base of last few dorsalfin rays. Size: maximum length about
40 cm, commonly to 25 cm. Distribution: Andaman Sea and western Pacific. Remarks: inhabits shallow coastal
waters, often in brackish waters especially sea grass and mangrove areas.
Marketed fresh.
(S. Kimura)

Siganus guttatus, KAUM–I. 17027, 16.7 cm SL
off Terengganu (KT), 16 Dec. 2008

Siganus javus(Linnaeus, 1766)
Streaked Spinefoot
D XIII, 10; A VII, 9; P1 15–17. Body
relatively deep, its depth 2.0–2.3 in
SL. Last dorsal fin spine 1.2–1.6 in
fourth to sixth spine. Except for shortfirst spine, all anal-fin spines similarin length. Soft parts of dorsal and anal
fins moderately high, longest anal-fin
ray subequal to longest anal-fin spine.
Caudal fin slightly concave. Color:dark bronze on dorsal half, becoming
paler ventrally; many gun-metal blue
spots on head and dorsal half of body;silvery blue undulating lines on ventral half of body. Size: maximum size
50 cm, commonly to 30 cm. Distribution: Indo-West Pacific from the Persian Gulf through the Malay Peninsula
eastward to Vanuatu, northward to thePhilippines, and southward to northern Australia.
(K. Matsuura)

Siganus javus, UMTF 1005 (KAUM–I. 16674), 9.5 cm SL
Kemaman, 21 Oct. 2008

Siganus punctatus(Schneider & Forster, 1801)
Goldspotted Spinefoot
D XIII, 10; A VII, 9; P1 16–17. Body
deep and compressed; its depth 1.9–2.3
in SL. Last dorsal fin spine 1.1–1.3 infourth to seventh spine. Last anal-fin
spine typically longest. Soft parts of
dorsal and anal fins moderately high,longest dorsal-fin ray subequal toSiganus punctatus, KAUM–I. 17083, 23.7 cm SL
off Terengganu (KT), 28 Dec. 2008

slightly shorter than longest dorsal-fin
spine. Caudal fin emarginate in young;deeply forked with rounded tips inlarge adults. Scales minute. Color:head and body brown; numerous golden spots scattered on head, body, andcaudal fin; a large deep brown blotch
below origin of lateral line. Size: maximum size 45 cm. Distribution: eastern
Indian Ocean and western Pacific. Remarks: inhabits shallow coastal waters(M. Matsunuma)
at depths of 3–50 m.

216

Siganus virgatus(Valenclennes, 1835)
Doublebarred Spinefoot
D XIII, 10; A VII, 9; P1 16–17. Body
oval, strongly compressed, its depth
1.8–2.3 in SL. Snout moderate, dorsal
profile of head convex. Fifth to seventh dorsal-fin spine longest. Third or
fourth anal-fin spine usually longest.
Caudal fin emarginate. Color: a darkbrown band running from napethrough eye to chin; another fromfourth to fifth dorsal-fin spine bases topectoral-fin base; these dark bands
containing light blue small spots andshort lines; posterior to second band,orange to yellow dorsally with light
blue spots, silver white ventrally. Size:maximum length about 33 cm, commonly 20 cm. Distribution: eastern
Indian Ocean and western Pacific,
from southern India to northern Australia, north to southern Japan. Remarks: inhabits shallow coastal waters, around coral reefs. Juveniles
often enter mangrove areas.
(S. Kimura)

Siganus virgatus, KAUM–I. 17156, 16.3 cm SL
Redang Island, 4 Jan. 2009

Siganus virgatus, KAUM–I. 17026, 13.9 cm SL
off Terengganu (KT), 16 Dec. 2008

217

ACANTHURIDAE
Surgeonfishes

By Hisashi Imamura
High-bodied, compressed marine
fishes (up to 75 cm) with very small
ctenoid scales. Eye high on head.
Mouth small, terminal; premaxilla
not protractile. Small close-set teeth
on jaws; no teeth on roof of mouth.
Dorsal fin continuous with IV–IX
spines and 19–33 soft rays. Anal fin
with II or III spines and 18–28 soft
rays. Caudal fin emarginated to lunate, with 16 principal rays. Pectoral
fin with 14–18 rays. Pelvic fin with
1 spine and 3 or 5 soft rays. Side of
caudal peduncle with sharp spine(s).
Color: usually predominantly brownto gray, but some species very colorful.
Remarks: acanthurids occur oncoral reefs or over rocky substrata,
generally at depths less than 100 m.
It is divisible into 3 subfamilies: theAcanthurinae, Prionurinae and Nasi-

single dorsal fin with X–XI, 19–33

sharp spine on
caudal peduncle

nae.
Similar family occurring in thearea: Siganidae – pelvic fin with 2
spines and 3 soft rays between them;dorsal fin with 13 spines; anal fin with

7 spines. Zanclidae – snout protruding; premaxilla protractile; dorsal fin
with a long filament from 3rd spine;no spine or keels on caudal peduncle.

Acanthurus mata(Cuvier, 1829)
Elongate Surgeonfish
D IX, 24–26; A III, 23–24; P1 16–17;
GR 13–15. Body deep, 2.1–2.5 in SL;
elongate in adults; caudal peduncle
slender. Mouth small; teeth fixed, denticulate on all of margin, with expanded incurved tips. Caudal fin deeply
emarginate, the caudal concavity 5.5–
8.0 in SL. Caudal peduncle with 1
folding spine. Color: body gray tobrown with lengthwise blue lines onhead and body; a small blackish spot
at upper end of gill opening, precededby a yellow band that continues infront of eye as a double band. Capableof changing body color to pale bluishoverall. Size: maximum length 50 cm,
commonly to 35 cm. Distribution:
Indo-West Pacific, including the Red
Sea, Andaman Sea, Ryukyu Islands
and New Caledonia. Remarks: occurring on rocky and coral reefs.

Acanthurus mata, KAUM–I. 17155, 17.7 cm SL
Redang Island, 4 Jan. 2009

218

SPHYRAENIDAE
Barracudas

By Keiichi Matsuura, Seishi Kimura and Mizuki Matsunuma
Body elongate, slightly compressed, reaching to 170 cm. Headlong; snout pointed. Mouth horizontal, large; lower jaw projecting;large, sharp flattened or conical teeth
on jaws; usually 1 or 2 canine teeth
on tip of lower jaw. Gill rakers, if
present, as short spinules, 1 or 2 atangle of gill arch. Two short dorsal
fins, widely separated; the first with
IV spines located above or slightly
behind pelvic fins; the second with I
spine and 9 soft rays. Anal fin with IIspines and 7–9 soft rays. Caudal fin
deeply forked. Color: gray to blue,
or light brown dorsally, with silvery
reflections, paler or white ventrally.
Remarks: carnivorous pike-like,pelagic to demersal fishes in tropical2 dorsal fins separated
snout long and pointed;
mouth large

sharp canines near
tip of lower jaw

and temperate seas. Most members of
Sphyraenidae found in coastal areas
but also in the surface of open seas or
down to depths of 100 m. Caught byhandlines, gill nets, set nets or trawls.
Marketed fresh, also dried-salted.
Similar families occurring in thearea: Atherinidae and Mugilidae –having 2 separate dorsal fins and silver body, but having a short snout andsmall mouth without canine teeth.

Sphyraena forsteri
Cuvier, 1829
Bigeye Barracuda
D V + I, 9; A II, 8; LL 112–133; GR
0. A medium-sized species. Body
elongate, subcylindrical. Maxilla either not reaching anterior margin of
Sphyraena forsteri, KAUM–I. 17079, 26.1 cm SLeye, or to just below anterior margin.
off Terengganu (KT), 27 Dec. 2008
First gill arch with many platelets,
each bearing distinct small spines.
Caudal fin typically forked in all stages, without a pair of lobes at posterior
margin; origin of pelvic fins before
first dorsal-fin origin. Color: head andbody silvery white, dark gray dorsally,
without bars and stripes on body sides;
a dusky blotch present behind axil of
pectoral fins; upper part of second
Sphyraena putnamae, UMTF 1352 (KAUM–I. 16466), 21.4 cm SL
dorsal fin dusky yellow; caudal fin
estuary near UMT, 29 Sept. 2008
darkish. Size: maximum length about
65 cm, commonly to 50 cm. Distriburior to origin of first dorsal fin. Color:tion: widespread in the tropical andSphyraena putnamae
head and body dark brown dorsally,
subtropical Indo-Pacific, from East
Jordan & Seale, 1905
silvery white ventrally. Many typical
Africa to the Marquesas Islands; north
Sawtooth Barracuda
dark chevron markings crossing laterto southern Japan. Remarks: inhabits
D V + I, 9; A II, 7–9; LLp 124–134; al line on body. Caudal fin blackishcoral reefs. Caught mostly by handlines.
(M. Matsunuma) GR 0. Body elongate, subcylindrical. without white tips. Size: maximum
Snout long, pointed; posterior tip of length 90 cm. Distribution: widely
maxilla just reaching to below anterior distributed in Indo-West Pacific, frommargin of eye. First gill arch without East Africa and Red Sea to Vanuatu
gill raker. Last rays of second dorsal and New Caledonia, north to southernand anal fin elongate in comparison Japan. Remarks: inhabits bays andwith penultimate rays. Caudal fin turbid inner lagoons. Marketed fresh.
deeply forked. Pelvic-fin origin ante(S. Kimura)

219

TRICHIURIDAE
Cutlassfishes

By Mizuki Matsunuma
Body remarkably elongate andcompressed, ribbon-like, with a tapered tail or small forked caudal
fin (up to over 2 m). A single nasal
opening on each side of head. Mouth
large, jaws not protractile, lower jaw
extends anterior to upper jaw. Teeth
extremely strong, fang-like in anterior part of upper jaw and sometimes
in anterior part of lower jaw. Dorsal
fin low and long, beginning shortly
behind eye, its anterior spinous partshorter than posterior soft part, 2
parts continous mostly or interrupted
by a shallow notch sometimes. Anal
fin low or reduced to short spinules.
Caudal fin either small and forked orabsent. Pectoral fins short and low
in position. Pelvic fins reduced to ascale-like spine (plus a rudimentary
ray in Benthodesmus) or completely
absent (in Trichiurus and Lepturacanthus). Preanal length less than 1/2

body extremely elongate and compressedsingle nostril

pelvic fins absent or reduced to a scale-like spine
mouth large, withstrong canine teeth

standard length. Lateral line single.
Scales absent. Color: body generally
metallic silvery or more or less brownin Aphanopus and Lepidopus; pectoral fin semi transparent; dorsal andanal fins sometimes tinged with paleyellow. Usually no distinct marks or
blotches on body.
Remarks: benthopelagic on continental shelves and slopes and underwater rises, from the surface to a depthof about 2 000 m, found in tropicalto warm-temperate waters. Excellent
eating, although the flesh is scanty.
Marketed mostly fresh or salted,
sometimes also frozen.
Similar families occurring in
the area: Gempylidae – double nostrils on each side of head; 2 dorsal
fins.

Trichiurus lepturus
Linnaeus, 1758
Largehead Hairtail
D III, 124–138; A II, 105–108; P1
10–12. Body extremely elongate andstrongly compressed, ribbon-like, tapering to a point posteriorly. Eyelarge; mouth large, with a dermal process at tip of each jaw; 2 or 3 pairs ofenlarged fangs with barbs near tip ofupper jaw, and another pair near tip oflower jaw, a single series of sharp,
compressed lateral teeth in both jaws,
minute teeth on palatines. Lower hindmargin of gill cover concave. Dorsal
fin high and long, without a notch between spinous and soft parts; pectoral
fin rather short, but extend beyond lateral line; caudal and pelvic fins absent.
No scales on body. Color: fresh specimens steel blue with silvery reflection,
pectoral fins semi-transparent, otherfins sometimes tinged with pale yellow hyaline in fresh, especially strong
yellow just after death. Size: maxi-

Trichiurus lepturus, KAUM–I. 16971, 51.0 cm TL
off Terengganu (KT), 11 Dec. 2008

Trichiurus lepturus, KAUM–I. 16972, 43.0 cm TL
off Terengganu (KT), 11 Dec. 2008

mum length about 1.8 m. Distribution: considered as a cosmopolitan
species of tropical and temperate waters. Remarks: basically offshore wa-

ters, and occurs in somewhat deeper
coastal area, but often collected evenin mangrove or estuary basin and to adepth of ca. 150 m depth.

220

SCOMBRIDAE

Mackerels and Tunas
By Seishi Kimura and Mizuki Matsunuma
Medium to large sized marine
fishes, maximum size over 5 m.
Body relatively elongate and fusiform, moderately compressed. Snout
pointed, upper jaw not protrusible.
Caudal peduncle slender with 2 ormore lateral keels on each side. Two
dorsal fins, the first usually shortbased with IX–XXVII spines, separated from or apparently contiguous
to the second. Finlets (5–12 each)present behind second dorsal andanal fins. Caudal fin deeply forked.
Pectoral fins inserted high on body.
Pelvic fins relatively small with I
spine and 5 soft rays, located below
pectoral fins. Body covered withsmall to medium cycloid scales ora corselet developed on area behind
head and pectoral fins. Lateral line
present. Color: body dark blue or2 dorsal fins separated
finlets present behinddorsal and anal finslateral keels on
caudal peduncle

dark green dorsally, silvery white ventrally. Usually dark vertical or longitudinal bands and/or spots on sides.
Remarks: typical pelagic fishes;
smaller fishes usually inhabits inshore
waters; large tunas, Thunnus, transoceanic migrants. Important food fishes.
Similar families occurring in thearea: Carangidae – dorsal-fin spines

III–VIII; II detached anal-fin spines;
scutes developed along posterior partof lateral line in most species. Gempylidae – body dark brown to brown
dorsally without distinct marks or
blotches; no keels on caudal peduncle except Lepidocybium.

Auxis rochei rochei(Risso, 1810)
Bullet Tuna
D X–XI + 10–12; A 12–13; P1 22–
23; GR 43–48. Body robust, elongate
and rounded. Teeth small and conical,
in a single series. Second dorsal fin
followed by 8 finlets; anal fin followed by 7 finlets; pectoral fin short,
not reaching vertical line from anterior
margin of scaleless area above corselet; a large, single-pointed flap (interpelvic process) between pelvic fins.
Body naked except for corselet; corselet well developed posteriorly, extending beyond vertical through second
dorsal fin origin. A strong central keel
on each side of caudal-fin base between 2 smaller keels. Color: backbluish, turning to deep purple or almost black on head; scaleless areawith pattern of 15 or more fairly
broad, nearly vertical dark bars; belly
white; pectoral and pelvic fins purple.
Size: maximum 40 cm, commonly to
35 cm FL. Distribution: a cosmopolitan species of tropical and temperatewaters except for the eastern Pacific.
Remarks: caught with seines, lift
nets, pole-and-line, and by trolling.
Marketed fresh.
(M. Matsunuma)

Auxis rochei rochei, KAUM–I. 17227, 24.8 cm SL
off Terengganu (KT), 12 Jan. 2009

Auxis rochei rochei, UMTF 1899, 23.0 cm SL
off Terengganu (KT), 11 Jan. 2009

221

Euthynnus affinis(Cantor, 1849)
Kawakawa
D XI–XIV + 12–13; A 12–14; P1
25–27; GR 31–34; V 39. Body robust
and fusiform, somewhat elongate.
Jaws with small conical teeth in a single series; no vomerine teeth. Caudal
peduncle with a large lateral keel andcaudal-fin base with 2 small keels.
Two dorsal fins separated by only anarrow interspace. Eight to 10 dorsal
and 6–8 anal finlets. Pectoral finsshort; 2 flaps between pelvic fins (interpelvic process). Body naked except
for corselet and lateral line. Color:head and body dark blue dorsally, silvery-white ventrally. Dark blue oblique
stripes dorsolaterally on posterior body.
Several black spots between pectoraland pelvic fins. Size: maximum length
about 100 cm, commonly to 60 cm
FL. Distribution: widely distributed
in warm waters of Indo-Pacific. Remarks: inhabits coastal waters andaround offshore islands. Marketed
fresh, dried, salted, and smoked.
(S. Kimura)

Rastrelliger kanagurta(Cuvier, 1816)

Euthynnus affinis, UMTF 1886, 29.3 cm SL
off Terengganu (KT), 25 Dec. 2008

Euthynnus affinis, KAUM–I. 17068, 28.3 cm SL
off Terengganu (KT), 26 Dec. 2008

Indian Mackerel
D VIII–XI + 11–13; A I, 11–12; P1
19–22; LGR 30–46. Body slightly
deep, compressed. Head longer thanbody depth. Maxilla covered by lacrimal bone. Adipose eyelid well developed. Gill rakers very long, visible
when mouth is opened. Caudal finbase with 2 small lateral keels. Two
dorsal fins widely separated. Anal fin
spine rudimentary. Five or six dorsal
and anal finlets. Pectoral fins short; asmall single flap between pelvic fins(interpelvic process). Body covered
with small scales. Color: head andbody bluish green dorsally, silverywhite ventrally. One or two rows ofsmall, dark spots dorsolaterally on
body. First dorsal, caudal and pectoral
fins yellowish. Size: maximum length
35 cm, commonly to 25 cm FL. Distribution: widely distributed in IndoWest Pacific, from South Africa and
Red Sea to Samoa, north to southern
Japan. Remarks: found in coastal waters. Feeds on macrozooplanktons.
Marketed fresh, dried, salted, andsmoked.
(S. Kimura)

Rastrelliger kanagurta, KAUM–I. 17002, 13.8 cm SL
off Terengganu (KT), 14 Dec. 2008

Rastrelliger kanagurta, UMTF 1885, 15.8 cm SL
off Terengganu (KT), 25 Dec. 2008

222

Scomberomorus commerson(Lacepède, 1800)
Narrowbarred Spanish Mackerel
D XV–XVIII + 15–20; A 16–21; P1
21–24; GR 0–2 + 1–8 = 1–8, V 19–20+ 23–27 = 42–46. Body elongate,strongly compressed. Upper jaw
reaching to or extending slightly beyond posterior margin of eye. Teeth onjaws strong with finely serrated edge.
Caudal peduncle with a large lateralkeel and caudal-fin base with 2 small
keels. Two dorsal fins scarcely separated. Eight to 11 dorsal and 7–12 anal
finlets. Pectoral fins short, pointed.
Body covered with small scales. A
single lateral line abruptly bent downward below posterior end of dorsal-finbase. Color: head and body bluish silver dorsally, silvery-white ventrally.
Many vertical dark bars on body. Size:maximum length 220 cm, commonly
to 90 cm FL. Distribution: widelydistributed in Indo-West Pacific, fromSouth Africa and Red Sea to Fiji, northto Japan. Remarks: found in coastal
waters. Feeds on fishes. Marketed
mainly fresh.
(S. Kimura)

Scomberomorus commerson, KAUM–I. 16943, 15.0 cm SL
off Terengganu (KT), 11 Dec. 2008

Scomberomorus commerson, UMTF 1893, 14.4 cm SL
off Terengganu (KT), 11 Jan. 2009

Thunnus obesus(Lowe, 1839)
Bigeye Tuna
D XIV–XV + 13–15; A 13–15; P1
32–35; GR 26–28. Body robust, fusiThunnus obesus, UMTF 1901, 26.2 cm SL
form, slightly compressed. Two dorsal
off Terengganu (KT), 12 Jan. 2009
fins, separated only by a narrow interspace, with 8–10 finlets; pectoral finsvery long, its tip reaching to extending
beyond vertical through second dorsal
fin origin; 2 flaps (interpelvic process)between pelvic fins; anal fin followedby 7–10 finlets. Very small scales onbody; corselet of larger and thicker
scales developed, but not very distinct.
Caudal peduncle with a strong lateralkeel between 2 smaller keels. Ventral
surface of liver striated. Color: backmetallic dark blue, lower sides andbelly whitish; a lateral iridescent blue
Thunnus obesus, KAUM–I. 17228, 26.0 cm SLband runs along sides in live specioff Terengganu (KT), 12 Jan. 2009
mens; first dorsal fin deep yellow, second dorsal and anal fins light yellow,
finlets bright yellow edged with black. tion: worldwide in tropical and sub- face to depths of 250 m. Marketed(M. Matsunuma)
Size: maximum length over 200 cm, tropical waters. Remarks: a pelagic fresh.
commonly to 180 cm FL. Distribu- oceanic species, taken from the sur-223

STROMATEIDAE
Butterfishes

By Seishi Kimura
Medium sized (up to 60 cm)
marine fishes. Body strongly deepand compressed. Snout short, blunt.
Mouth small, terminal or slightly inferior, not protractile. Teeth on jaws
uniserial, small and flattened with
3 minute cusps. No teeth on vomer
or palatines. Gill membranes joined
to breast (Pampus) or not usuallyconnected to isthmus (Peprilus andStromateus). Branchiostegal rays
5 or 6. Caudal peduncle short andcompressed with no scutes or keels.
Dorsal fin single, anal fin similar inshape to dorsal fin. Caudal fin forked
or emarginate. Pectoral fins long.
Pelvic fins absent in adults. Body
covered with small cycloid scales.
Lateral line single dorsolaterally on
body. Color: body silvery white,dark brownish in some species.
Remarks: coastal fishes, feeds
mainly on zooplanktons. Three genera, Pampus (Indo-West Pacific),
Peprilus (western Atlantic and eastern Pacific), and Stromateus (Atlantic and eastern Pacific). Importantcaudal fin forked or
emarginate

mouth small,
not protractile

no pelvic fins in adults

food fishes.
Similar families occurring in thearea: Carangidae – Parastromateus
niger, very similar in shape to Pampus,
having small scutes along posterior

part of lateral line; gill membrane
not united to isthmus. Centrophidae– pelvic fins present; 7 branchiostegals. Nomeidae – 2 dorsal fins; pelvic fins present.

Pampus argenteus(Euphrasen, 1788)
Silver Pomfret
D V–X + 37–43; A V–X + 34–43; P1
24–27; GR 10–13; V 34–37. Body
oval, strongly compressed. Gill opening short, slit-like. Gill rakers verysmall, vestigial. Anterior dorosal andanal fins falcate. Caudal fin deeply
forked, lower lobe often extended.
Sensory canal area on temporal extending backward along lateral line.
Color: head and body dark brownish
silver dorsally, silvery white ventrally.
Anal and caudal fins yellowish. Size:maximum length 60 cm. Distribution: Indo-West Pacific, from Persian
Gulf to Indonesia. Remarks: found incoastal waters over muddy bottom.
Marketed fresh.

Pampus argenteus, KAUM–I. 17066, 15.6 cm SL
off Terengganu (KT), 26 Dec. 2008

224

Pampus chinensis(Euphrasen, 1788)
Chinese Silver Pomfret
D 43–50; A 39–42; P1 24–27; GR
11–14; V 33. Body oval, deep, strongly compressed. Gill opening short,slit-like. Gill rakers slender, needlelike. Anterior dorsal and anal fins not
falcate. Caudal fin emarginate. Sensory canal area on temporal extending
backward along lateral line. Color:body grayish brown, abdomen paler.
Size: maximum length 40 cm SL. Distribution: Indo-West Pacific, from
Persian Gulf to eastern Indonesia. Remarks: found in inshore waters overmuddy bottom. Marketed fresh.

Pampus chinensis, KAUM–I. 17115, 7.1 cm SL
off Terengganu (KT), 31 Dec. 2008

Pampus chinensis, KAUM–I. 17298, 10.1 cm SL
off Terengganu (KT), 20 Jan. 2009

225

PSETTODIDAE
Spiny Turbots

By Hisashi Imamura
Marine flatfishes in middle size,
maximum size ca. 80 cm. Body
oval, caudal peduncle deeper thanits length. Eyes on right or left sideof head. Mouth large, extending wellbeyond posterior margin of lower
eye. Teeth on jaws large canines,
many with barbed tips; vomer andpalatine with teeth. Margin of preopercle distinct, not covered withskin. Dorsal fin origin well posterior
to upper eye. Anterior rays of dorsal
and anal fins spinous. Pelvic fin with
1 spine and 5 soft rays, nearly symmetrically placed on each sides of
midventral line. Caudal fin with 15
branched rays, separated from dorsal
and anal fins. Lateral line on bothsides of body, only slightly curved
above pectoral fin. Weak ctenoid
scales covering both sides of body.
Vertebrae usually 10 + 14=24. Color: ocular side brownish, blind sidepale brown.
Remarks: single genus, Psettodes, with one species (P. erumei)
from Indo-West Pacific and two (P.

anterior dorsal-fin rays spinous
dorsal fin notextending to head

pelvic finswith I spine
anterior anal-fin rays spinous

belcheri Bennett, 1831 and P. bennettii Steindachner, 1870) from western
Africa. Occurring in coastal waters.
Food fish.
Similar family occurring in thearea: Psettodidae differs from allother flatfish families in having dorsal fin origin well posterior to upper
eye, and anterior rays of dorsal andanal fin spinous. Citharidae – lateralline highly arched above pectoral
fin. Paralichthyidae – pelvic fin with-out spine; lateral line highly archedabove pectoral fin. Bothidae – pelvic
fin without spine; origin of pelvic fin
on eyed side anterior to that on blindside; lateral line highly arched above
pectoral fin. Samaridae – pelvic fin
on blind side absent. Soleidae – pelvic fin without spine; origin of pelvic
fin on eyed side anterior to that on
blind side. Cynoglossidae – caudal
fin continuous with dorsal and anal
fins.

Psettodes erumei(Bloch & Schneider, 1801)
Indian Halibut
D IX–XI, 38–45; A I, 33–43; P1 14–
16; P2 I, 5; LL 61–77. Body oval,
strongly compressed. Eyes on right orleft side of head. Mouth large, extending well beyond posterior margin oflower eye. Teeth on jaws large canines; vomer and palatine with small
conical teeth. Anterior rays of dorsal
and anal fin spinous. Caudal fin separated from dorsal and anal fins. Lateralline on both sides of body, only slightly curved above pectoral fin. Scales on
small, weakly ctenoid on both sides ofbody. Color: eyed side generally
brownish or grayish, sometimes with
4 or 5 broad dark cross bands; dorsal,
anal, posterior portion of caudal in

Psettodes erumei, KAUM–I. 16836, 10.6 cm SL
off Terengganu (KT), 5 Dec. 2008

darker; blind side pale brown. Size:maximum total length about 60 cm,
most commonly 20–40 cm. Distribution: Indo-West Pacific, from South

Africa and Red Sea to northern Australia, north to Taiwan. Remarks:
found on sandy and muddy bottoms.
Marketed fresh.

226

CITHARIDAE

Largescale Flounders
By Hisashi Imamura
Marine flatfishes in small size,
maximum size less than 40 cm. Body
elliptical strongly compressed. Eyeson left side of head in some species, on right side in others. Mouth
moderate or somewhat large. Smalland slender teeth on jaws. Marginof preopercle distinct, not covered
with skin. Dorsal fin origin anterior
to eyes. Anterior rays of dorsal andanal fins not spinous. Pelvic fin with
1 spine and 5 soft rays, fin bases onboth sides short. Caudal fin with
13–15 branched rays, separated fromdorsal and anal fins. Lateral line usually present on both sides of body,with high arch above pectoral fin.
Body scales large; ctenoid scales oneyed side, weakly ctenoid or cycloid
scales on blind side. Color: eyed sidewith some spots on body and fins;
blind side whitish.

no dorsal-fin spines

pelvic finswith I spine

no anal-fin spines

Remarks: five genera, Brachypleura, Citharoides, Citharus, Lepidoblepharon and Paracitharus, are
known from Indo-West Pacific. Usually found on sandy or muddy-sandy
bottoms. Food fish.
Similar family occurring in thearea: Psettodidae – dorsal fin origin
well posterior to upper eye; anteriorrays of dorsal and anal fin spinous;

lateral line slightly curved above
pectoral fin. Paralichthyidae – pelvic
fin without spine. Bothidae – pelvic
fin without spine; origin of pelvic
fin on eyed side anterior to that on
blind side. Samaridae – pelvic fin onblind side absent. Soleidae – pelvic
fin without spine. Cynoglossidae –caudal fin continuous with dorsal andanal fins.

Brachypleura novaezeelandiae(Günther, 1862)
Widemouth Largescale Flatfish
D 65–77; A 41–50; P1 11–13 (eyedside), 10–13 (blind side); P2 I, 5; LL
28–33; GR 5–6 + 7–10. Body elliptical, strongly compressed. Both eyes
on right side of body. Head 2.2–2.7 in
SL. Mouth large, arched, reaching toor beyond center of lower eye, but notreaching to posterior margin of lower
eye. Some anterior rays of dorsal fin
elongate in males, not in females.
Caudal fin rounded posteriorly. Lateral line usually present on both sides ofbody, with high arch above pectoral
fin. Body scales large; ctenoid scales
on eyed side, weakly ctenoid or cycloid scales on blind side. Color: eyedside of body yellowish or yellowish
brown; dorsal, anal and caudal finspaler than body, with dark spots; blindside of body whitish. Size: maximum
total length 14 cm. Distribution: Indo-West Pacific, including Maldives,
Andaman Sea, and northern area ofSouth China Sea. Remarks: found on
sandy and muddy bottoms, at depthsof 18 to 73 m. Marketed fresh.

Brachypleura novaezeelandiae, KAUM–I. 16828, male, 8.6 cm SL
off Terengganu (KT), 5 Dec. 2008

Brachypleura novaezeelandiae, KAUM–I. 16830, female, 8.2 cm SL
off Terengganu (KT), 5 Dec. 2008

227

PARALICHTHYIDAESand Flounders

By Hisashi Imamura
Body elliptical to oval, strongly
compressed, reaching to 40 cm. Headlarge, 3.0–4.4 in SL. Mouth rather
large. Teeth uniserial on both jaws.
Eyes on left side of head, separated
by a bony ridge. Margin of preopercle distinct, not covered with skin.
Dorsal fin origin anterior to eyes.
Anterior rays of dorsal and anal
fins not spinous. Pelvic fin without
spines; pelvic fin bases on both sides
short, subequal and subsymmetrical in position. Caudal fin separated
from dorsal and anal fins. Lateral line
equally developed on both sides ofbody, with high arch above pectoral
fin and supratemporal branch, running upward to anterior part of dorsal
fin. Body scales large; ctenoid scales
on eyed side, weakly ctenoid or cycloid scales on blind side. Color:body brownish or light greenish withdark spots or rings.
Remarks: most species known

eyes onleft side
pectoral fin rays on
ocular side branchedpelvic fin bases
short and subsymmetricalfrom shallow sandy and muddy bottoms of continental shelf; some species from brackish waters near rivermouths.
Similar family occurring in thearea: Psettodidae – dorsal fin origin
well posterior to upper eye; anteriorrays of dorsal and anal fin spinous;
pelvic fin with one spine; lateral line

slightly curved above pectoral fin.
Citharidae – pelvic fin with one
spine. Bothidae – origin of pelvic fin
on eyed side anterior to that on blindside. Samaridae – pelvic fin on blindside absent. Soleidae – eyes on left
side of head. Cynoglossidae – caudal
fin continuous with dorsal and anal
fins.

Pseudorhombus arsius(Hamilton, 1822)
Largetooth Flounder
D 71–84; A 53–62; P1 12–13 (eyedside); LL 69–81. Body oval. Head
3.3–3.6 in SL; upper profile of headwith a slight notch anterior to uppereye. Upper jaw extending to below
posterior margin of lower eye. Several
pairs of moderately large canine teeth
in anterior part of both jaws. Scales
ctenoid on eyed side, cycloid on blindside. Color: body greenish or lightbrown; a dark blotch at junction ofstraight and curved parts of lateralline; a smaller blotch halfway to caudal fin base. Size: maximum standard
length 45 cm, commonly to 30 cm.
Distribution: widespread in IndoWest Pacific, from eastern coast of
Africa eastward to Australia and Fiji.
Remarks: found from shallow muddy-sandy or muddy bottoms fromcoastal areas to river mouths.

Pseudorhombus arsius, UMTF 1516 (KAUM–I. 16720), 14.6 cm SL
Setiu, 29 Oct. 2008

228

Pseudorhombus dupliciocellatus
Regan, 1905
Ocellated Flounder
D 72–78; A 59–63; P1 11–12 (eyedside); LL 73–84. Body oval. Head 3.4–
3.7 in SL; upper profile of head with astrong notch in front of upper eye. Upper jaw extending to below or beyond
middle of lower eye. Both jaws withsmall teeth, lacking strong canines.
Scales ctenoid on eyed side, cycloid on
blind side. Color: body brownish, with
1 or 2 pairs of distinct double ocelli
above and below lateral line. Size:maximum standard length 40 cm,
commonly to 20–30 cm. Distribution:
widespread in Indo-West Pacific, from
Nicobar Islands to Japan and to Australia. Remarks: inhabits shallow waters on mud and sand bottoms of continental shelf, at depths of 50–150 m.

Pseudorhombus dupliciocellatus, KAUM–I. 17278, 26.0 cm SL
off Terengganu (KT), 18 Jan. 2009

Pseudorhombus javanicus(Bleeker, 1853)
Javan Flounder
D 67–76; A 51–56; P1 11–12 (eyedside); LL 66–74. Body oval. Head
3.2–3.6 in SL; upper profile of head
without notch anterior to upper eye.
Upper jaw extending to below middleof lower eye. Teeth on both jaws
small, with slightly enlarged teeth anteriorly. Scales ctenoid on anteriorpart, and dorsal and ventral margins ofbody of eyed side; those on other areasof eyed side and blind side cycloid.
Color: body brownish, with a distinct,
large dark blotch at junction of straightand curved parts of lateral line, and asmaller blotch on middle of straightsection of lateral line. Size: maximum
standard length 35 cm, commonly to
20 cm. Distribution: Indo-West Pacific, from eastern coast of India eastward to western New Guinea and tosouthern China. Remarks: foundfrom shallow waters on mud and sandbottoms of continental shelf.

Pseudorhombus malayanus
Bleeker, 1865
Malayan Flounder
D 72–74; A 58–59; P1 12–13 (eyedside); LL 70–78. Body deeply oval.
Head 3.1–3.3 in SL; upper profile ofhead with a slight notch anterior to upper eye. Upper jaw extending to below
posterior margin of lower eye. Teeth

Pseudorhombus javanicus, KAUM–I. 17303, 12.7 cm SL
off Terengganu (KT), 20 Jan. 2009

Pseudorhombus malayanus, KAUM–I. 17024, 16.6 cm SL
off Terengganu (KT), 16 Dec. 2008

on upper jaw small and close-set laterally, and enlarged anteriorly; teeth onlower jaw stronger and more widely
spaced than those on upper jaw. Scales
ctenoid on both sides of body. Color:body brownish, with a small dark
blotch at junction of straight andcurved parts of lateral line. Size: maximum standard length 35 cm, commonly to 20 cm. Distribution: IndoWest Pacific, from Persian Gulf to
Malay Archipelago and to Philippines.
Remarks: inhabits shallow waters on
mud and sand bottoms.

229

Pseudorhombus neglectus
Bleeker, 1865
Neglected Flounder
D 72–75; A 56–59; P1 12–13 (eyedside); LL 77–79. Body elliptical. Head
3.5–3.6 in SL; upper profile of head
without a distinct notch or with aslight notch anterior to upper eye. Upper jaw extending to below or beyond
middle of lower eye. Teeth on upper
jaw small, slightly enlarged anteriorly;
teeth on lower jaw larger and more
widely spaced than those on upper
jaw. Scales ctenoid on eyed side, cycloid on blind side. Color: body pale
brownish, with small ocelli at junctionof straight and curved parts of lateralline, and another one near middle ofstraight section. Size: maximum standard length 25 cm. Distribution:
known from eastern Indian Ocean toWest Pacific. Remarks: found fromshallow waters on mud and sand bottoms of continental shelf, at depths of
30–40 m.

Pseudorhombus neglectus, KAUM–I. 16979, 16.3 cm SL
off Terengganu (KT), 12 Dec. 2008

Pseudorhombus pentophthalmus
Günther, 1862
Fivespot Flounder
D 68–76; A 53–57; P1 11–12 (eyedside); LL 62–75. Body oval. Head
3.2–3.4 in SL; upper profile of headwith a distinct notch anterior to uppereye. Mouth extending to below or alittle beyond middle of lower eye.
Teeth on upper jaw small and closeset; teeth on lower jaw moderately
larger and more widely spaced thanthose on upper jaw. Scales ctenoid on
eyed side, cycloid on blind side. Color: body pale brownish, with 2 ocelli

Pseudorhombus pentophthalmus, KAUM–I. 16952, 8.8 cm SL
off Terengganu (KT), 11 Dec. 2008

above and below lateral line and 1
ocellus on posterior third of straightsection of lateral line. Size: maximum
standard length 18 cm, commonly 12
cm. Distribution: West Pacific, fromJava Sea through Indo-China, north tonorthern Japan. Remarks: found fromfairly deep waters on mud and sandbottoms of continental shelf, at depthsof 50–150 m.

230

BOTHIDAE

Lefteye Flounders
By Hisashi Imamura
Marine flatfishes. Body ellipticalto oval, strongly compressed, reaching to 88 cm. Mouth small to large.
Eyes on left side of head. Interorbital width variable; males of somespecies with wider interorbit thanfemale. Margin of preopercle distinct, not covered with skin. Dorsal
fin origin above or ahead of anterior
margin of upper eye. Anterior raysof dorsal and anal fins not spinous.
Pelvic fin without spines; pelvic finof eyed side on midventral line withorigin anterior to origin of pelvic
fin of blind side; pelvic fin of blindside above midventral line. Caudal
fin separated from dorsal and anal
fins. Lateral line on eyed side highly
arched above pectoral fin; lateral line
absent below lower eye. Color: eyedside of body brownish, usually withspots, blotches, or rings; blind sideof body whitish, but dark colored insome species.
Remarks: usually found on san-eyes on leftside

dorsal and anal
fins not attached
to caudal fin

no lateral line below
lower eyepelvic fin bases
asymmetricaldy and muddy bottoms, but some species on rocky or coral reefs. Feeds on
small fishes, crustaceans, gastropods
and other benthic animals. Twenty
genera. Food fish.
Similar family occurring in thearea: Psettodidae – dorsal fin origin
well posterior to upper eye; anteriorrays of dorsal and anal fin spinous;
pelvic fin with one spine; lateral line

slightly curved above pectoral fin.
Citharidae – pelvic fin with one
spine. Paralichthyidae – pelvic finbases on both sides short, subequaland subsymmetrical in position.
Samaridae – pelvic fin on blind sideabsent. Soleidae – eyes on left sideof head. Cynoglossidae – caudal fin
continuous with dorsal and anal fins.

Arnoglossus macrolophus
Alcock, 1889
Largecrested Lefteye Flounder
D 91–98; A 70–76; P1 12–14 (eyedside); LL 56–62. Body elliptical. Head
3.6–4.2 in SL; upper profile of headwith a distinct notch anterior to uppereye. Upper jaw extending slightly beyond anterior margin of lower eye.
Teeth on both jaws small, closely
spaced. First 4 or 5 dorsal fin rays
greatly elongate in males, only slightly elongate in females. Color: bodylight brownish, with a dark spot atbase of posterior parts of dorsal andanal fins; a dark spot on distal portionof pectoral fin. Size: maximum total
length 13 cm. Distribution: northern
Indian Ocean and West Pacific, including southern Japan, Taiwan, South
China Sea and Indo-Australian Archipelago.

Arnoglossus macrolophus, KAUM–I. 17145, 7.9 cm SL
off Terengganu (KT), 1 Jan. 2009

231

Engyprosopon grandisquama(Temminck & Schlegel, 1846)
Largescale Flounder
D 80–91; A 57–69; P1 10–13 (eyedside); LL 41–48. Body oval. Head
3.5–4.2 in SL; upper profile of headwith a slight concavity anterior to upper margin of lower eye. Single strong
rostral spine near snout in males, absent or feeble in females. Upper jaw
extending to below anterior portion oflower eye. Teeth on upper jaw biserial,
those on lower jaw uniserial. Scales
large, with short ctenii on eyed side,
cycloid on blind side. Color: eyedside light brown, many dark spots andrings irregularly scattered on it; caudal
fin with a pair of large prominent jetblack blotches. Size: maximum standard length about 11 cm, commonly to
8 cm. Distribution: Indo-West Pacific, including East Africa, southern Japan and northern Australia. Remarks:
found from mud and sand bottoms ofcontinental shelf, at depths of 10–
100 m.

Engyprosopon grandisquama, KAUM–I. 16827, 7.9 cm SL
off Terengganu (KT), 5 Dec. 2008

Grammatobothus polyophthalmus(Bleeker, 1865)
Manyeyed Flounder
D 77–86; A 61–71; P1 14–17 (eyedside); LL 75–77. Body oval. Head
3.5–4.0 in SL; upper profile of headwith a deep notch anterior to interorbital region. Upper jaw reaching to orslightly beyond below anterior portionof lower eye. Teeth on both jaws uniserial. 2nd to 10th dorsal fin rays elongate in both sexes, longer in males
than females. Pectoral fin elongate inmales. Lateral line on both sides ofbody. Scales on eyed side ctenoid,
those on blind side cycloid. Color:
eyed side pale brownish, with 3 large
and prominent dark ocelli, 1 eachabove and below pectoral fin, 1 on
middle of strait section of lateral line.
Size: maximum total length about 21
cm, most specimens about 17cm. Distribution: Indo-West Pacific, including India, Ryukyu Islands, Philippinesand northwestern Australia. Remarks:
inhabits on mud, sand and rubble bottoms at depths of 0–90 m.

Engyprosopon grandisquama, KAUM–I. 16885, 7.2 cm SL
off Terengganu (KT), 6 Dec. 2008

Grammatobothus polyophthalmus, KAUM–I. 17304, 11.7 cm SL
off Terengganu (KT), 20 Jan. 2009

232

SAMARIDAE

Crested Flounders
By Hisashi Imamura
Marine flatfishes, maximum size
smaller than 22 cm. Body elliptical tosomewhat oval, strongly compressed.
Mouth small, reaching t almost anterior margin of upper eye. Eyes on rightside of head. Margin of preopercle
distinct, not covered with skin. Dorsal
fin origin anterior to anterior marginof upper eye. Anterior rays of dorsal
and anal fins not spinous. Pectoral fin
present on eyed side but absent on
blind side. Pelvic fins symmetrical,
without spines. Caudal fin separated
from dorsal and anal fins. Lateral line
well developed or rudimental. Color:
eyed side of body brownish, usuallywith spots, blotches, or rings; blindside of body whitish.
Remarks: known from tropicaland subtropical regions in Indo-Pacific, primarily deeper waters. Three
genera, Plagiopsetta, Samaris andSamariscus, with about 20 species.
Similar family occurring in thearea: Psettodidae – dorsal fin origin

dorsal and anal
fins not attached
to caudal fineyes on right side

body flat and somewhat elongate

pelvic fin bases asymmetricalwell posterior to upper eye; anteriorrays of dorsal and anal fin spinous;
pelvic fin with one spine; lateral line
slightly curved above pectoral fin.
Citharidae – pelvic fin with one spine.
Paralichthyidae – eyes on left side ofhead, pelvic fin bases on both sides

short, subequal and subsymmetricalin position. Bothidae – eyes on left
side of head. Soleidae – preopercle
without free margin. Cynoglossidae –eyes on left side of head; preopercle
without free margin; pectoral fins absent.

Samaris cristatus
Gray, 1831
Cockatoo Righteye Founder
D 71–85; A 49–60; P1 4 (eyed side);
LL 60–80. Body elliptical. Head 3.8–
5.3 in SL; upper profile of head with aslight notch anterior to upper eye.
Mouth small; upper jaw extending to
below anterior portion of lower eye.
Both jaws with small villiform teeth inband. Anterior 12 or 13 rays greatly
elongate and filamentous. Pectoral fin
on eyed side somewhat elongate. Pelvic fin on eyed side continuous withanal fin by membrane. Lateral line
present only on eyed side. Scales oneyed side ctenoid, those on blind side
feebly ctenoid or cycloid. Color: eyedside pale brownish, with a series ofobscure dark blotches along dorsal
and ventral margins of body; blindside whitish. Size: maximum length
22 cm. Distribution: Indo-West Pacific, from southeastern Africa to New
Caledonia, north to Japan and south tonorthern Australia. Remarks: foundfrom mud and sand bottoms.

Samaris cristatus, KAUM–I. 17197, 12.8 cm SL
off Terengganu (KR), 5 Jan. 2009

233

SOLEIDAE
Soles

By Hisashi Imamura
Marine flatfishes (excepting one
species entering rivers in Africa),
reaching to 32 cm. Body somewhat
elongate to oval, strongly compressed. Mouth small and asymmetrical, terminal or slightly inferior.
Eyes on right side of head. Preopercle without free margin, embedded
in skin. Dorsal fin origin anterior toupper eye. Anterior rays of dorsal
and anal fins not spinous. Pectoral
fins sometimes absent; when present, eyed side pectoral fin usually
longer than on blind side. Pelvic finswithout spines, sometimes asymmetrical, either free or fused withanal fin. Caudal fin separated from or
fused with dorsal and anal fins. Lateral line single, running straight on
body, sometimes branched on head.
Color: highly variable according tosubstratum, uniformly dull brownto strikingly colored with scattered
black spots or blotches or dark crossbands on eyed side of body; usually

dorsal fin extending
far forward on headeyes onright side
lateral line single, straight

uniformly yellowish or white on blindside of body.
Similar family occurring in thearea: Soleidae differs from all otherflatfish families, except for Cynoglossidae, in lacking free preoperclar
margin. Psettodidae – dorsal fin origin
well posterior to upper eye; anteriorrays of dorsal and anal fin spinous;
pelvic fin with one spine; lateral line

slightly curved above pectoral fin.
Citharidae – pelvic fin with one
spine. Paralichthyidae, Bothidae and
Cynoglossidae – eyes on left side ofhead.
Remarks: inhabits muddy andsandy bottom in coastal areas. About
35 genera with about 130 species.

Brachirus orientalis(Bloch & Schneider, 1801)
Oriental Sole
D 61–65; A 44–48; P1 7–8 (eyedside); LL 70–85. Body oval, 1.8–2.2 in
SL. Head 4.2–5.3 in SL. Mouth small,terminal, slightly curved. Eyes separated by a scaly interorbital space.
Very small teeth on both jaws on blindside. Pectoral fins well developed.
Pelvic fin separated from anal fin.
Caudal fin separated from dorsal andanal fins. Lateral line with high rounded arch on head not directed posteriorly. Scales strongly ctenoid on eyedside; mostly weakly ctenoid on blindside with some cycloid. Color: eyedside gray or brown with numerous,
cloudy and indistinct spots, and several narrow blackish lines; blind side
uniformly light yellow, without sooty
blotches. Size: maximum total length
about 30 cm, commonly 10 to 12 cm.

Brachirus orientalis, UMTF 1487 (KAUM–I. 16719), 9.5 cm SL
Setiu, 29 Oct. 2008

Distribution: Indo-West Pacific, including Gulf of Thailand, Philippines,
southern Japan, New Guinea and Australia. Remarks: found from mud andsand bottoms in coastal waters.

234

Liachirus melanospilos(Bleeker, 1854)
Darkspotted Sole
D 59–62; A 42–47; LL 57–65. Body
oval, 2.1–2.4 in SL. Head 3.9–4.2 in
SL. Mouth small, slightly curved.
Eyes separated by a scaly interorbital
space. Pectoral fins absent. Pelvic fin
separated from anal fin. Caudal fin
separated from dorsal and anal fins.
Scales on both sides small cycloid.
Color: eyed side light to dark brownwith scattered darker dots and blotches; blind side pale. Size: attaining to
7.5 cm SL. Distribution: Indo-West
Pacific, including South China Sea,
southern Japan and Australia. Remarks: inhabits on mud and sand bottoms around 100 m depth.

Liachirus melanospilos, KAUM–I. 16959, 7.1 cm SL
off Terengganu (KT), 11 Dec. 2008

Pardachirus pavoninus(Lacepède, 1802)
Peacock Sole
D 66–70; A 50–53; LL 75–82. Body
oval. Mouth strongly curved, cleftreaching to anterior margin of lower
eye. Eyes separated by a scaly interorbital space. Pectoral fins absent. Pelvic fin separated from anal fin. Caudal
fin separated from dorsal and anal
fins. Lateral line distinct, straight, single. Series of toxin glands with pores
along bases of dorsal and anal fin rays.
Color: eyed side reddish brown,densely spotted on head, body andfins; spots in various sizes and shapes,
bordered by a dark rim and some witha blackish spot in center. Size: maximum total length about 22 cm, commonly 10–15 cm. Distribution: widespread tropical Indo-West Pacific,
including southern Japan, Philippines
and Australia. Remarks: inhabits on
sandy bottoms of coral reef areas, at
depths of about 40 m.

Pardachirus pavoninus, KAUM–I. 17212, 16.0 cm SL
off Terengganu (KT), 10 Jan. 2009

Zebrias quagga(Kaup, 1858)

Zebrias quagga, KAUM–I. 16957, 8.5 cm SL
off Terengganu (KT), 11 Dec. 2008

Zebra Sole
D 60–75; A 50–62. Body elliptical.
Mouth small, terminal, slightly
curved. Eyes contiguous, each with ashort tentacle. Pectoral fins well developed. Pelvic fin separated fromanal fin. Caudal fin continuous withdorsal and anal fins. Scales ctenoid.
Color: eyed side cream or beige with
11–12 brown single or double cross-bands, slightly wider than interspaces;
body pattern continued onto dorsal
and anal fins; caudal fin cream withelongate black blotches. Size: maximum length 15 cm. Distribution: Indo-West Pacific, including Red Sea,

Persian Gulf, Malaysia, Thailand,
China and Australia. Remarks: foundfrom mud and sand bottoms in shallow coastal waters.

235

CYNOGLOSSIDAE
Tonguesoles

By Hisashi Imamura
Marine flatfishes (some species
entering freshwater), reaching to 40
cm. Body tongue-shaped, strongly
compressed. Mouth small, subterminal and asymmetrical, reaching
posteriorly below lower eye. Eyeson left side of head. Posterior margin of preopercle strongly attached toopercle, without free margin and embedded in skin. Dorsal fin reaching
far forward on head. Anterior rays of
dorsal and anal fins not spinous. Pectoral fins absent. Usually only pelvic
fin on blind side present. Caudal fin
separated fused with dorsal and anal
fins. Scales small, ctenoid or cycloid.
Color: variable even in a species,
usually brownish or grayish, often
variably marked with spots, blotches
or cross bands on eyed side of body;usually uniformly yellowish or whitish on blind side of body.
Remarks: commonly found on
muddy bottoms and other substrates,from tide pools to deep waters oneyes onleft side

dorsal and anal fins
joined with caudal fin

body flat and oblongno spines in all finscontinental shelves and slopes. Marketed fish. Includes 3 genera andabout 130 species.
Similar family occurring in thearea: Cynoglossidae differs from allother flatfish families in this area, except for Soleidae, in having caudal fin
fused with dorsal and anal fins, andlacking free preoperclar margin. It isalso separable from most flatfishes

in lacking pectoral fins. Psettodidae– dorsal fin origin well posterior toupper eye; anterior rays of dorsal
and anal fin spinous; pelvic fin with
one spine; lateral line slightly curved
above pectoral fin. Citharidae – pelvic fin with one spine. Samaridae andSoleidae – eyes on right side of head.

Cynoglossus bilineatus(Lacepède, 1802)
Fourlined Tonguesole
D 107–113; A 80–88; C usually 12;
LL 88–96 (midlateral line). Body
elongate, 3.5–4.5 in SL. Mouth moderate, extending beyond lower eye.
Eyes small, slightly larger than interorbital space. Two lateral lines oneach side of body; upper one along
dorsal fin base, lower one midlaterally
on body. Scales ctenoid on eyed side
and cycloid on blind side. Color: eyedside of body pale brown; blind side ofbody whitish. Size: maximum length
40 cm. Distribution: Indo-West Pacific, from Persian Gulf to New Guinea, north to southern Japan, south toAustralia. Remarks: inhabits sandy
and muddy bottoms at depths of 13 toca. 400 m.

Cynoglossus bilineatus, KAUM–I. 17096, 24.9 cm SL
off Terengganu (KT), 28 Dec. 2008

Cynoglossus bilineatus, UMTF 1874, 10.9 cm SL
off Terengganu (KT), 29 Dec. 2008

236

Cynoglossus lingua
Hamilton, 1822
Long Tonguesole
D 126–138; A 97–114; C usually 10;
LL 90–101 (midlateral line). Body
very elongate, 4.5–5.9 in SL. Mouth
moderate, extending well beyond posterior margin of lower eye. Eyes separated by narrow interorbital space.
Two lateral lines present on eyed sideof body; upper one along dorsal finbase, lower one midlaterally on body.
Lateral lines absent on blind side ofbody. Scales comparatively large, ctenoid on eyed side and cycloid on blindside. Color: eyed side of body reddish
brown, sometimes with irregularbrown-black patches, with a large
black blotch on opercular region.
Size: maximum total length about 40
cm. Distribution: Indo-West Pacific,
including Red Sea, Gulf of Thailand,
Philippines and Indonesia. Remarks:
inhabits mainly shallow sandy andmuddy bottoms on inner continental
shelf, often entering estuaries.

Cynoglossus lingua, KAUM–I. 17005, 27.1 cm SL
off Terengganu (KT), 14 Dec. 2008

Cynoglossus monopus, KAUM–I. 17119, 13.0 cm SL
off Terengganu (KT), 31 Dec. 2008

Cynoglossus monopus(Bleeker, 1849)
D 115–120; A 92–96; C 10; LL 108–125 (midlateral line). Body very elongate, 3.8–4.6 in SL. Mouth moderate,
extending to below posterior marginof eye or just behind it. Eyes pedunculate. Two lateral lines present on eyedside of body; upper one along dorsal
fin base, lower one midlaterally on
body. Lateral lines absent on blindside of body. Scales ctenoid on bothsides of body. Color: eyed side ofbody brownish; dorsal, anal and caudal fins darker. Size: maximum length
about 19 cm. Distribution: IndoWest Pacific, from northern part of
Bay of Bengal to Hong Kong and Malay Archipelago.

Paraplagusia blochii, UMTF 1321 (KAUM–I. 16450), 13.7 cm SL
estuary near UMT, 28 Sept. 2008

Paraplagusia blochii(Bleeker, 1851)
Bloch's Tonguesole
D 99–105; A 76–82; C 8; LL 75–83(midlateral line). Body elongate, 3.6–
3.8 in SL. Mouth extending to below
posterior margin of lower eye. Eyes
small, separated by a scaly interorbital
space about equal to eye diameter.
Two lateral lines present on eyed sideof body; lateral lines absent on blind

Paraplagusia blochii, UMTF 1083 (KAUM–I. 16518), 7.4 cm SL
Cendering, 6 Oct. 2008

side of body. Scales ctenoid on eyed mum length 22 cm. Distribution: Inside and weakly ctenoid on blind side. do-West Pacific, including Oman, InColor: eyed side of body brown; scale dia, Indonesia and Philippines.
centers paler than edges. Size: maxi-

237

TRIACANTHIDAE
Triplespines

By Keiichi Matsuura
Small fishes less than 30 cm,
with moderately elongate, strongly
compressed body covered by moderately thick skin with numerous
minute scales. Mouth small andterminal; teeth in an outer series ofabout 10 heavy incisors in each jaw,
internally to which are several molariform teeth, usually 4 in upper
jaw and 2 in lower jaw. Gill opening a moderately short vertical slit infront of pectoral-fin base. Dorsal fin
spine VI (usually only V visible, thesixth rudimentary), dorsal-fin rays
20 to 26; caudal fin deeply forked;
pelvic fins with I large spine and novisible rays; most dorsal-, anal, andpectoral-fin rays branched. Caudal
peduncle distinctly tapering to a narrow transversely indented region justin front of caudal-fin base, where thepeduncle is wider than deep. Lateralline inconspicuous. Color: generally

V visible spines
20–26 fin rays

caudal fin
deeply forked

pelvic fin withno visible rays

silvery, with upper half of body dusky,
with or without darker blotches.
Remarks: benthic, occurring usually on flat, sandy or weed-covered
bottoms. Feed on bottom invertebrates. Marketed but not commercially important.
Similar families occurring in thearea: Triacanthodidae – caudal fin
not deeply forked, rounded to almost truncate; caudal peduncle notdistinctly tapered, deeper than wide;
dorsal-fin rays 12 to 18 (20 to 26 in
Triacanthidae).

Tripodichthys blochi(Bleeker, 1852)
Longtail Tripodfish
D VI + 20−24; A 15−19; P1 13−15.
Scale covered ventral surface of pelvis
distinctly tapered to a point posteriorly. Second dorsal spine much less thanhalf the length of first dorsal spine.
Pelvis between bases of pelvic spines
narrow (1.4−3.0 %SL), the width 7.2−
12.4 (usually about 9 or 10) times inlength of pelvis. Dorsal rays modally
22 and anal rays modally 17. Color:dorsal half of body silvery light brown,
ventral half silvery white; several irregular dark yellow markings on sideof body; spiny dorsal fin membrane
pale; basal part of spiny dorsal fin yellowish orange; soft dorsal, anal andpectoral fins pale; caudal fin light yellow. Size: 15 cm. Distribution:
known from China and countries inSoutheast Asia, a stray found fromsouthern Japan. Remarks: coastal andestuarine shallow waters at depths of
0−40 m; usually found on sandy or
muddy flat; feeds on benthic invertebrates.

Tripodichthys blochi, KAUM–I. 16730, 9.1 cm SL
Setiu, 28 Oct. 2008

238

BALISTIDAE
Triggerfishes

By Keiichi Matsuura
Small to medium-sized fishes, up
to 80 cm in total length, with deep,
moderately compressed body encased in very thick, tough skin withlarge scale plates easily discernible
as individual unites; scale above
pectoral-fin base in many species enlarged, forming a flexible tympanum.
Gill opening a relatively short vertical to oblique slit in front of pectoralfin base; mouth small and usuallymore or less terminal; teeth heavy, 8
in outer series of upper jaw and lower jaw. Three dorsal-fin spines, thesecond spine more than one half ofthe length of the first; the first spine
capable being locked in an uprightposition of erection by the second;most dorsal, anal and pectoral-fin
rays branched; pelvic fins and spines
rudimentary, represented by a seriesof 4 pairs of enlarged scales encasing
the end of pelvis. Lateral line inconspicuous. Color: variable and many
species having bright color markings

III dorsal-fin spines

scales plate-likesingle pelvic fin

on head and body.
Remarks: most species benthic,
occurring in coral and rocky reefs
from the coastline to a depth of 100m; some species (e.g. Canthidermis)
pelagic in open water. Feed on bottom
invertebrates, but also zooplankton(e.g. Melichthys indicus and Odonus
niger). High valued as food in many
handline fisheries.

Similar families occurring in
the area: Monacanthidae – two
dorsal-fin spines, only the first ofwhich is long and prominent; bodymore laterally compressed; fewer
and less massive teeth in jaws; scales
shagreen-like, with individual basal
plates small and not readily distinguishable from one another to theunaided eye.

Abalistes stellatus(Annonymous, 1798)
Starry Triggerfish
D III + 26−27; A 24−26; P1 14−15(usually 14). Scales enlarged above
pectoral-fin base and just behind gill
opening to form a flexible tympanum.
A groove in front of eye. Mouth terminal. Caudal peduncle depressed, wider
than deep. Color: grayish brown toolivaceous on back, with small pale
blue or yellow spots dorsally and large
yellow spots ventrally; 3 large white
blotches on back, disappearing inlarge specimens. Size: 60 cm. Distribution: widespread in Indo-West Pacific, from East Africa to northern
Australia, north to southern Japan. Remarks: occurring on sand, sponge,
and weed bottoms to depths of 100 m.
Similar to Abalistes filamentosus Matsuura & Yoshino, 2004 but distinguished by lacking produced caudal
fin rays. Feeds on marine invertebrates.

Abalistes stellatus, KAUM–I. 17123, 12.5 cm SL
off Terengganu (KT), 31 Dec. 2008

239

Sufflamen fraenatum(Latreille, 1804)
Masked Triggerfish
D III + 26−28; A 23−26; P1 12−14.
Scales enlarged above pectoral-finbase and just behind gill opening toform a flexible tympanum. A groovein front of eye. Mouth terminal. Caudal peduncle compressed, deeper thanwide. Scales of posterior part of bodywith large tubercles forming longitudinal ridges extending forward to levelof soft dorsal-fin origin. Color: headand body dark brown; a light yellow
line running obliquely from corner ofmouth to below a groove before eye in
male, but lacking in female; all finsdark brown; head and body of juveniles white with many longitudinal
black lines below level of eye, dorsalside above level of eye dark brown.
Size: 35 cm. Distribution: widespread in Indo-West Pacific, from East
Africa eastward to Tuamotu Islands,
northward to southern Japan. Remarks: occurring in coral reefs shallower than 50 m.

Sufflamen fraenatum, KAUM–I. 17108, juvenile, 4.2 cm SL
off Terengganu (KT), 28 Dec. 2008

240

MONACANTHIDAE
Filefishes

By Keiichi Matsuura
Small to medium-sized fishes,
usually less than 20 cm (but up to 50
cm for some species of Aluterus), withdeep, highly compressed body covered by thin but rough or shagreenlike skin with minute scales not individually easily discernible to theunaided eye. Mouth small and usually more or less terminal or slightly
supraterminal; teeth only moderately
heavy, 6 in an outer series in upper
jaw and 6 or fewer in the lower. Gill
opening a relatively short, vertical to
oblique slit in front of pectoral-finbase. Two (sometimes 1) dorsal−finspines, second spine not more than
1/3 the length of first; first spine usually capable of being locked in an
upright position of erection by thesecond; dorsal-, anal- and pectoralfin rays unbranched; pelvic fin andspines rudimentary or absent, represented by a series of 3 or fewer pairsof enlarged scales encasing end of
pelvis, or segments of indeterminate
number, or entirely absent. Scales

II dorsal-fin spines(2nd hidden in skin)scales small, shagreen-likesingle pelvic fin

above pectoral-fin base unmodified,
not forming a tympanum. Lateral line
inconspicuous or only slightly apparent. Color: variable, drab brown, gray,or greenish, but often with strikingly
marked and vivid patterns.
Remarks: found in shallow coraland rocky reefs, seagrass beds, andsandy-muddy bottoms in depths of
10 to over 200 m. Feed on coral polyps, epiphytes attached to seagrass, orother marine invertebrates. Marketed
fresh.
Similar families occurring in
the area: Balistidae – III spines infirst dorsal fin; skin tough but not
shagreen-like, individual scales distinct, usually forming prominent,
oblique rows. Triacanthidae – VIspines (usually only V spines visible) in first dorsal fin; pair of strong
pelvic-fin spines.

Aluterus monoceros(Linnaeus, 1758)
Unicorn Leatherjacket
D II + 45−51; A 47−53; P1 14−15.
Body elongate, strongly compressed.
Dorsal profile of head slightly convex.
Gill opening short oblique slit below
eye extending antero-ventrally beyond
anterior margin of eye. Dorsal spine
above eye, very long and fragile; broken frequently in collected specimens.
Soft part of dorsal fin opposite to analfin. Caudal fin shorter than snout. Color: body grayish with many irregulardark lines and dots; dorsal-, anal- andpectoral-fin rays yellow; caudal fin
dark. Size: 76 cm. Distribution: circum tropical; in West Pacific known
from west coast of peninsular Malay-

Aluterus monoceros, KAUM–I. 16897, 21.0 cm SL
off Terengganu (KT), 6 Dec. 2008

sia eastward to the Solomon Islands,
northward to the Ryukyu Islands. Remarks: forming large schools incoastal waters.

241

Chaetodermis penicilligerus(Cuvier, 1816)
Prickly Leatherjacket
D II + 25−26; A 23−24; P1 12−13.
Body deep; dorsal profile of head ascending from mouth to dorsal spine.
Body covered with many fleshy filaments. Pelvic terminus prominent,
composed of 3 segments of encasing
scales, movable dorso-ventrally.
Scales on body relatively large with abackwardly curved spine. Dorsal
spine with many fleshy filaments, located above gill opening. Caudal fin
rhomboidal. Color: light brown withmany longitudinal black lines on sideof body; fin rays of dorsal, anal andcaudal fins with many black spots.
Size: 31 cm. Distribution: tropicaleastern Indian Ocean and West Pacificfrom west coast of Malaysia eastward
to northern Australia, northward to
southern Japan. Remarks: collectedfrom rocky reefs and sandy-muddy
bottom, usually from depths shallower
than 200 m; feeds on marine invertebrates.

Chaetodermis penicilligerus, KAUM–I. 16852, 18.1 cm SL
off Terengganu (KT), 5 Dec. 2008

Monacanthus chinensis(Osbeck, 1765)
Fanbellied Leatherjacket
D II + 28−30; A 27−30; P1 12. Body
deep; dorsal profile of head concave,ascending from mouth through eye tosoft dorsal fin. Pelvic terminus prominent, composed of 3 segments of encasing scales, movable dorso-ventrally; ventral flap very large. Upper
caudal fin rays produced into a filament. Color: body light brown withmany irregular dark brown blotches.
Size: 38 cm. Distribution: west coastof Malay Peninsula and tropical West
Pacific from Thailand eastward to
New Guinea. Remarks: occurring in
sea grass bed and rocky reefs in shallow waters.

Monacanthus chinensis, KAUM–I. 17071, 13.8 cm SL
off Terengganu (KT), 27 Dec. 2008

242

Paramonacanthus pusillus(Rüppell, 1829)
D II + 25−30; A 24−29; P1 11−13.
Body relatively elongate; dorsal profile of head convex in male, withprominent hump immediately in frontof nostrils, slightly convex or straightin female. Pelvic terminus elongate,
composed of 3 segments of encasing
scales, movable dorso-ventrally; ventral flap developed. First dorsal spine
depressed in cross section, armed with
4 rows of downward-directed barbs.
Color: head and body pale grayish
brown to whitish, with many yellowish to dark brown markings tending toform 3−4 irregular curved stripes onbody; soft dorsal fin pale with 2 dark
blotches at its base, anal fin and pectoral fins pale, caudal fin pale with 2
curved dark cross bands. Size: 18 cm.
Distribution: tropical Indo-West Pacific from South Africa eastward tonorthern Australia, northward to
southern Japan. Remarks: this species is the most widespread member ofParamonacanthus.

Paramonacanthus pusillus, KAUM–I. 16937, 7.7 cm SL
off Terengganu (KT), 11 Dec. 2008

Pseudomonacanthus macrurus(Bleeker, 1856)
Strap-weed Filefish
D II + 29−30; A 27−30; P1 11−12.
Body relatively elongate, strongly
compressed. Doral profile of headslightly concave. Pelvic terminus
small, composed of 2 segments of encasing scales, not movable; ventral
flap large. Dorsal spine located above
posterior half of eye. Gill opening
oblique, extending from below middleof eye to below nasal organ. Color:head and body light brown with many
black spots, ventral half of body paler
than the dorsal half. Fin rays of all fins
yellow; caudal fin with 2 transverse
dark brown bands. Size: 20 cm. Distribution: South China Sea eastward
to New Guinea, northward to the
Ryukyu Islands. Remarks: occurringin sea grass bed and on sandy-muddy
bottom in shallow waters.

Pseudomonacanthus macrurus, KAUM–I. 17042, 15.7 cm SL
off Terengganu (KT), 17 Dec. 2008

243

TETRAODONTIDAE
Puffers

By Keiichi Matsuura
Small to moderate-sized fishes,
most species less than 30 cm, witha heavy blunt body capable of rapid
inflation by intake of water (or air).
Head large and blunt; jaws modified
to form a beak of 4 heavy, powerful teeth, 2 above and 2 below. Gill
openings without distinct opercular
cover, appearing as simple slits anterior to the pectoral fin; eyes locatedhigh on head. Dorsal and anal finslocated far posteriorly bearing nospines, but 7 to 15 soft rays; caudal fin usually truncate to slightly
rounded; pelvic fins absent. Typicalscales absent, but most species are
partially covered with tiny prickles
or spinules, and many species havesmall fleshy tabs or lappets on thedorsal and/or lateral surfaces. Color:most species are mottled, variegated,or barred on the upper and lateralsurfaces, often with spots of various
sizes and colors; ventral surfaces arealmost always unpigmented.
Remarks: occurring in tropicalgill opening
beak

deflated
inflated

pelvic fins absent

and temperate seas, most frequently
in shallow inshore waters, sometimes
entering brackish and fresh waters,
but a few species are pelagic. The viscera, skin, and blood of most species
are poisonous; in some species eventhe flesh is poisonous. Laymen arestrongly recommended not to eat puffers, although connoisseurs like to con-sume puffers in licensed restaurants
in some countries (e.g., Japan).
Similar families occurring in
the area: Diodontidae – head andbody covered with strong elongate
spines; 1 tooth plate in each jaw.

Lagocephalus lunaris(Bloch & Schneider, 1801)
Rough Golden Puffer
D 12−13; A 11−12; P1 16−17. A
patch of spinules on back reaching todorsal-fin origin; belly covered with
spinules. Nasal organ with 2 openings.
Caudal fin concave. Color: dorsal halfof body grayish light brown, side silvery white, and belly white; caudal fin
white in the ventral lobe and dark yellow in the dorsal lobe. Size: 35 cm.
Distribution: tropical Indo-West Pacific from East Africa to northern Australia; found recently from southern
Japan. Remarks: strongly toxic andfrequently confused with other speciesof Lagocephalus such as L. gloveri
Abe & Takita, 1983 and L. spadiceus(Richardson, 1845).

Lagocephalus lunaris, KAUM–I. 17207, 11.7 cm SL
off Terengganu (KT), 10 Jan. 2009

244

Lagocephalus spadiceus(Richardson, 1845)
Half-smooth Golden Puffer
D 11−13; A 10−12; P1 14−17. A
patch of spinules on back between
snout and halfway to dorsal fin origin;
belly covered with spinules. Nasal organ with 2 openings. Caudal fin slightly concave. Color: dorsal half of bodygrayish brown, side silver white, andbelly white; caudal fin white in theventral lobe and dark yellow in thedorsal lobe. Size: 30 cm. Distribution: warm areas in eastern Indian
Ocean and West Pacific from Japanthrough Thailand and Indonesia tonorthern Australia. Remarks: frequently confused with other species of
Lagocephalus including a very dangerous toxic species, L. lunaris, having an enlarged patch of spinules onback reaching posteriorly to dorsal-fin
origin.

Lagocephalus suezensis
Clark & Gohar, 1953
Suez Puffer
D 10−12; A 9−10; P1 14−16. A patchof spinules on back from between
snout to dorsal-fin origin, belly covered with spinules. Nasal organ with 2
openings. Caudal fin deeply lunate.
Caudal peduncle depressed, wider
than deep. Color: dorsal half of bodylight brown with many irregularly
shaped brown dots and lines, side silvery white and belly white; dorsal,
anal and pectoral fins pale; caudal fin
white in the ventral lobe, yellow in thedorsal lobe. Size: 20 cm. Distribution: tropical Indo-West Pacific fromRed Sea to northern Australia, northward to southern Japan. Remarks:
frequently confused with Lagocephalus
sceleratus (Gmelin, 1789) but distinguished from it by lacking many small
black spots on back and by having
fewer fin-ray counts (in L. sceleratus,
dorsal-fin rays 12−13, anal-fin rays
10−12, and pectoral-fin rays 16−17).
Probably toxic.

Tetraodon nigroviridis
Marion de Procé, 1822

Lagocephalus spadiceus, KAUM–I. 17175, 16.2 cm SL
off Terengganu (KR), 10 Jan. 2009

Lagocephalus spadiceus, KAUM–I. 16839, 14.0 cm SL
off Terengganu (KT), 5 Dec. 2008

Lagocephalus suezensis, KAUM–I. 17034, 13.1 cm SL
off Terengganu (KT), 17 Dec. 2008

Tetraodon nigroviridis, KAUM–I. 16706, 9.3 cm SL
Setiu, 27 Oct. 2008

organ composed of a tentacle divided
distally into 2 flattened and broadened
Spotted Green Puffer
elements. Color: back and side ofD 12−14; A 10−12; P1 18−23. 17cm. body dark greenish yellow with many
Body rounded in cross section. Head black spots, belly white; dorsal, anal
and body covered with spinules. Nasal and pectoral fins pale; caudal fin palewith several transverse dark lines.
Size: 15 cm. Distribution: freshwaterand brackish water in tropical Asia
from Sri Lanka to Indonesia, northward to Vietnam. Remarks: toxic.

245

Torquigener gloerfelti
Hardy, 1984
Brown Spotted Puffer
D 8−9; A 6−7; P1 15−16. Body relatively elongate, rounded dorsally andflattened ventrally. Chin distinct. Nasal organ a short, erect papilla with 2
openings. Ventro-lateral skin fold running from behind chin to caudal-finbase. Spinules on back from level ofnasal organ to immediately before
dorsal-fin origin; 7−9 spinules acrossback just behind mid-dorsal branch oflateral lines. Color: back and side ofbody pale yellowish white covered

Torquigener gloerfelti, KAUM–I. 17043, 17.2 cm SL
off Terengganu (KT), 17 Dec. 2008

with irregularly shaped brown circles
and spots, belly white; dorsal, anal andpectoral fins pale; caudal-fin rays yellowish with dark brown margin. Size:

20 cm. Distribution: Indonesia, Malaysia and Vietnam. Remarks: caught
by trawl in shallow waters on sandymuddy bottom; probably toxic.

DIODONTIDAE
Porcupinefishes

By Keiichi Matsuura
Small to medium-sized fishes (upto 100 cm). Body capable of great inflations, covered with strong spines
which may be quite long. Mouth
large, wide, and terminal; teeth fused
to form a strong, beak-like plate
without a median suture dividing left
and right halves. Gill opening a relatively small, vertical slit just in frontof pectoral-fin base. Dorsal and anal
fins without spines, placed far backon body; most fin rays branched;no pelvic fins. Lateral line inconspicuous. No normal scales. Color:background color light tan to brown,
but frequently gray; usually overlain
with dark brown to black spots, bars
and/or blotches; belly white, frequently with yellow overtone.

2 beak-like toothplates on jaws

many long, strong spines on bodyno pelvic fins

Remarks: found in shallow coraland rocky reefs, seagrass beds, andsandy-muddy bottoms in depths of
10 to over 200 m. Feed on coral polyps, epiphytes attached to seagrass, orother marine invertebrates. Marketed
fresh.

Similar families occurring in
the area: no other families having
the following combination of characters: large spines on body, no pelvic
fins, inflatable body, teeth fused intoa single beak-like plate.

Cyclichthys orbicularis(Bloch, 1785)
Birdbeak Burrfish
D 11−13; A 10−12; P1 18−21. Body
rounded with many 3 rooted, non-erectile spines; a small movable spine below and behind corner of mouth. Color: head and body brown dorsally,
white ventrally; black spots on backand side; all fins pale. Size: 16 cm.
Distribution: tropical Indo-West Pacific from East Africa eastward to New
Caledonia, southward to northern Australia, northward to southern Japan.
Remarks: found in coral and rocky
reefs and sandy bottom; usually occurring in depths shallower than 40 m.

Cyclichthys orbicularis, KAUM–I. 17031, 7.7 cm SL
off Terengganu (KT), 17 Dec. 2008

246

Diodon holocanthus
Linnaeus, 1758
Longspined Porcupinefish
D 14−16; A 14−16; P1 21−25. Long
erectile spines on head and body; 16–
21 spines from top of snout to dorsal
fin; frontal spines shorter than spines
posterior to pectoral fin; no downward
pointing spine below anterior marginof eye. Nasal organ a short hollow
tube with two opening near tip. Color:dorsal side of body brown to gray,white ventrally; a vertical white-edged
brown band across rear of head andanother across middle of back; a shortvertical white-edged brown bar infront of gill opening; a large oval
white-edged brown blotch above pec-

Diodon holocanthus, KAUM–I. 41684, 7.5 cm SL
off Terengganu (KT), 10 Sept. 2011

toral-fin base, and another around
dorsal-fin base; fins yellowish. Size:maximum total length 50 cm. Distribution: Indo-Pacific, from east Africa

to Society Islands, north to southern
Japan, south to New South Wales,
Australia. Remarks: occurring mainly on coral reefs.

247

Scientific Name Index
Abalistes stellatus 238
abbreviata, Acanthocepola 156
Ablennes hians 55
Abudefduf bengalensis 157
sexfasciatus 158
sordidus 158
Acanthocepola abbreviata 156
Acanthopagrus pacificus 135
Acanthosphex leurynnis 68
Acanthuridae 217
Acanthurus mata 217
Acentrogobius caninus 193
viridipunctatus 194
aeolus, Sillago 93
Aetobatus ocellatus 22
Aetomylaeus nichofii 23
affinis, Euthynnus 221
albella, Sardinella 39
Albula argentea 26
Albulidae 26
Alectis ciliaris 98
indicus 99
Alepes djedaba 99
melanoptera 99
vari 100
alexanderae, Pomacentrus 165
Aluterus monoceros 240
Ambassidae 76
Ambassis interrupta 76
nalua 77
urotaenia 77
vachellii 77
amblycephalus, Epinephelus 81
Johnius 139
Amblygaster clupeoides 37
sirm 38
Amblygobius phalaena 194
stethophthalmus 194
Amblypomacentrus breviceps 158
Amphiprion clarkii 159
frenatus 159
ocellaris 160
perideraion 160
anabatoides, Neopomacentrus 164
andhraensis, Stolephorus 33
anea, Pennahia 141
annularis, Pomacanthus 153
Anodontostoma chacunda 38
Antennariidae 48
Antennarius nummifer 48
Antherinidae 52
Apistidae 63
Apistus carinatus 63
Aploactinidae 68
Aploactis aspera 68
Aplocheilidae 56
Aplocheilus panchax 56
Apogon cavitensis 87
endekataenia 88
fasciatus 88
hyalosoma 89
lineatus 89
nigrocincta 89
pleuron 90
poecilopterus 90
seminigracaudus 90
truncatus 91
Apogonichthyoides niger 91

Apogonidae 87
Archamia fucata 91
archipelagicus, Hemiramphus 54
areolatus, Epinephelus 81
argenteus, Monodactylus 146
Pampus 223
Pomadasys 125
argentia, Albula 26argentilineatus, Periophthalmus 204
argentimaculatus, Lutjanus 116
argus, Halichoeres 171
Scatophagus 213
Ariidae 42
Arius maculatus 42
Arnoglossus macrolophus 230
arsius, Pseudorhombus 227
asiatica, Sillago 94
aspera, Aploactis 68
Asterropteryx semipunctata 195
Atelomycterus marmoratus 11
Atherinidae 52
Atherinomorus lacunosus 52
pinguis 53
atripectoralis, Chromis 161
atropos, Atropus 100
Atropus atropos 100
Atule mate 100
aureus, Chrysochir 138
Glossogobius 199
australiae, Rhynchobatus 16
Auxis rochei rochei 220
baganensis, Stolephorus 34
Balistidae 238
balteatus, Dipterygonotus 120
Redigobius 207
bathybius, Nemipterus 127
Bathygobius cocosensis 195
cyclopterus 195
fuscus 196
belcheri recurvispinnis, Repomucenus 190
Belonidae 55
bengalensis, Abudefduf 157
bicirrhosus, Glossogobius 199
bicoarctatus, Trachyrhamphus 59
bifasciatum, Diploprion 81
bilineatus, Cynoglossus 235
bilitonensis, Blenniella 183
bindus, Photopectoralis 114
biocellatus, Psammogobius 206
bipinnulata, Elagatis 104
bleekeri, Epinephelus 82
Lophogobius 203
Blenniidae 183
Blenniella bilitonensis 183
blochi, Tripodichthys 237
blochii, Paraplagusia 236
boenak, Cephalopholis 79
boops, Selar 106
Bothidae 230
Brachirus orientalis 233
Brachypleura novaezeelandiae 226
breviceps, Amblypomacentrus 158
brocki, Halicampus 58
Butis butis 191
koilomatodon 191
butis, Butis 191
coeruleopinnatus, Carangoides 101
Caesio cuning 119

Caesionidae 119
calcarifer, Lates 78
Callionymidae 189
Calliurichthys japonicus 189
canadum, Rachycentron 96
cancrivora, Saurenchelys 30
cancrivorus, Pisodonophis 28
caninus, Acentrogobius 193
Carangidae 98
Carangoides coeruleopinnatus 101
gymnostethus 101
hedlandensis 102
malabaricus 102
Caranx sexfasciatus 103
Carapidae 46
Carcharhinidae 12
Carcharhinus dussumieri 12
melanopterus 13
carinatus, Apistus 63
cavitensis, Apogon 87
Centriscidae 60
Centriscus scutatus 60
Cephalopholis boenak 79
formosa 80
microprion 80
sonnerati 80
Cepolidae 156
chacunda, Anodontostoma 38
Chaetodermis penicilligerus 241
Chaetodon octofasciatus 151
Chaetodontidae 151
Champsodon vorax 179
Champsodontidae 179
Chanidae 40
Chanos chanos 40
chanos, Chanos 40
Cheilinus chlorourus 169
fasciatus 170
trilobatus 170
Cheilodipterus quinquelineatus 92
Cheilopogon katoptron 53
Chelon subviridis 49
Chiloscyllium griseum 8
punctatum 9
chiltonae, Thysanophrys 75
chinensis, Monacanthus 241
Pampus 224
Chirocentridae 36
Chirocentrus dorab 36
chloropterus, Halichoeres 171
chlorourus, Cheilinus 169
chondropus, Sillago 94
Choridactylus multibarbus 69
Chromis atripectoralis 161
Chrysiptera unimaculata 161
Chrysochir aureus 138
chrysozona, Pterocaesio 120
Coradion 152
chrysurus, Pomacentrus 165
chulae, Mugilogobius 203
ciliaris, Alectis 98
cinerascens, Kyphosus 148
cinereus, Muraenesox 29
circumspectus, Glossogobius 200
Citharidae 226
clarkii, Amphiprion 159
Clupeidae 37
clupeoides, Amblygaster 37

248
cocosensis, Bathygobius 195
coelestis, Pomacentrus 166
coeruleopunctatus, Epinephelus 82
cognatus, Uranoscopus 181
Coilia dussumieri 32
coioides, Epinephelus 82
commerson, Scomberomorus 222
commersonianus, Scomberoides 105
cooki, Ostorhinchus 92
Coradion chrysozonus 152
cordyla, Megalaspis 104
Coryphaena hippurus 95
Coryphaenidae 95
Cottapistus cottoides 66
cottoides, Cottapistus 66
cristatus, Samaris 232
crocea, Larimichthys 140
crocodilus crocodilus, Tylosurus 56
crumenophthalmus, Selar 106
Cryptocentrus maudae 196
cultellatus, Platycephalus 74
cuneatus, Pomacentrus 166
cuning, Caesio 119
cuvieri, Inimicus 70
cyanomos, Neopomacentrus 164
cyanopleura, Leptojulis 174
cyanosoma, Ostorhinchus 92
Cyclichthys orbicularis 245
cyclopterus, Bathygobius 195
Cynoglossidae 235
Cynoglossus bilineatus 235
lingua 236
monopus 236
cyprinoides, Megalops 25
Dactyloptena gilberti 61
orientalis 62
Dactylopteridae 61
Dactylopus dactylopus 190
dactylopus, Dactylopus 190
Dascyllus reticulatus 162
trimaculatus 162
Dasyatidae 17
Dasyatis parvonigra 17
zugei 18
Decapterus macrosoma 103
russelli 103
decoratus, Istigobius 201
delicata, Moolgarda 50
devisi, Encrasicholina 33
Dendrophysa russelli 139
diadema, Istigobius 201
Diagramma picta picta 124
didactylus, Inimicus 70
digramma, Oxycheilinus 175
Pterocaesio 120
dimidiatus, Labroides 173
Diodon holocanthus 246
Diodontidae 245
Diploprion bifasciatum 81
Dipterygonotus balteatus 120
Dischistodus fasciatus 163
prosopotaenia 163
djedaba, Alepes 99
dorab, Chirocentrus 36
Drepane longimana 149
punctata 150
Drepaneidae 149
Drombus triangularis 196
dunckeri, Zenarchopterus 55
dupliciocellatus, Pseudorhombus 228

dussumieri, Carcharhinus 12
Coilia 32
Istiblennius 185
Plicofollis 43
Thryssa 34
Dussumieria elopsoides 38
Echeneidae 97
Echeneis naucrates 97
Ecsenius yaeyamaensis 184
edentulus, Istiblennius 185
Elagatis bipinnulata 104
Elates ransonnettii 72
Eleotridae 191
Eleutheronema tetradactylum 136
Ellochelon vaigiensis 50
elongatus, Gymnocranius 132
Omobranchus 186
Elopidae 24
Elops hawaiiensis 24
elopsoides, Dussumieria 38
Encheliophis homei 46
Encrasicholina devisi 33
endekataenia, Apogon 88
Engraulidae 32
Engyprosopon grandisquama 231
Enneapterygius tutuilae 182
Entomacrodus lighti 184
striatus 184
Ephippidae 211
Ephippus orbis 211
Epibulus insidiator 170
Epinephelus amblycephalus 81
areolatus 81
bleekeri 82
coeruleopunctatus 82
coioides 82
erythrurus 83
fasciatus 83
fuscoguttatus 83
heniochus 84
quoyanus 84
sexfasciatus 84
Equulites laterofenestra 109
leuciscus 110
oblongus 110
stercorarius 111
equulus, Leiognathus 113
erumei, Psettodes 225
erythrourus, Gerres 121
erythrurus, Epinephelus 83
Escualosa thoracata 39
Eubleekeria jonesi 111
Eugnathogobius variegatus 197
Euthynnus affinis 221
Eviota pellucida 197
storthynx 197
Exocoetidae 53
Exyrias puntang 198
fasciatum, Stegostoma 10
fasciatus, Apogon 88
Cheilinus 170
Dischistodus 163
Epinephelus 83
Salarias 188
Favonigobius reichei 198
ferox, Omobranchus 187
filamentosa, Parapercis 180
filamentosus, Gerres 122
Fistularia petimba 59
Fistulariidae 59

flavolineatus, Mulloidichthys 142
formosa, Cephalopholis 80
formosus, Parioglossus 210
forsteri, Sphyraena 218
fraenatum, Sufflamen 239
frenatus, Amphiprion 159
fucata, Archamia 91
fulviflamma, Lutjanus 117
furcosus, Nemipterus 128
fuscescens, Siganus 214
fuscoguttatus, Epinephelus 83
fuscus, Bathygobius 96
Pseudochromis 85
Fusigobius inframaculatus 199
Gazza minuta 112
rhombea 112
genivittatus, Lethrinus 133
gerrardi, Himantura 18
Gerreidae 121
Gerres erythrourus 121
filamentosus 122
limbatus 122
macracanthus 123
oyena 123
ghobban, Scarus 177
gibbosa, Sardinella 40
gibbosus, Plectorhinchus 125
gilberti, Dactyloptena 61
gloerfelti, Torquigener 245
Glossogobius aureus 199
bicirrhosus 199
circumspectus 200
Gnathanodon speciosus 104
Gobiidae 193
Gobiodon quinquestrigatus 200
goldmanni, Istigobius 202
Grammatobothus polyophthalmus 231
Grammoplites scaber 73
grandisquama, Engyprosopon 231
griseum, Chiloscyllium 8
griseus, Gymnocranius 133
guttatus, Siganus 215
Upeneus 143
Gymnocranius elongatus 132
griseus 133
gymnostethus, Carangoides 101
Gymnura poecilura 21
Gymnuridae 21
Haemulidae 124
Halicampus brocki 58
Halichoeres argus 171
chloropterus 171
marginatus 172
melanurus 172
nigrescens 172
hanedai, Secutor 114
hardwickii, Temera 15
Harpadon nehereus 44
hawaiiensis, Elops 24
hedlandensis, Carangoides 102
Helcogramma striata 182
Hemigobius hoevenii 200
mingi 201
Hemigymnus melapterus 173
Hemiramphidae 54
Hemiramphus archipelagicus 54
Hemiscyllidae 8
heniochus, Epinephelus 84
heptacanthus, Parupeneus 143
heptagonus, Hippichthys (Hippichthys) 58

249
Hexanematichthys sagor 43
hexodon, Nemipterus 128
hians, Ablennes 55
Hilsa kelee 39
Himantura gerrardi 18
walga 19
Hippichthys (Hippichthys) heptagonus 58
Hipposcarus longiceps 176
hippurus, Coryphaena 95
hoevenii, Hemigobius 200
holocanthus, Diodon 246
Holocentridae 57
homei, Encheliophis 46
hyalosoma, Apogon 89
indicius, Secutor 115
indicum, Leptomelanosoma 137
indicus, Alectis 99
Platycephalus 74
Stolephorus 34
Inegocia japonica 73
inframaculatus, Fusigobius 199
ingenuua, Sillago 94
Iniistius trivittatus 173
Inimicus cuvieri 70
didactylus 70
sinensis 71
insidiator, Epibulus 170
interrupta, Ambassis 76
Istiblennius dussumieri 185
edentulus 185
lineatus 186
Istigobius decoratus 201
diadema 201
goldmanni 202
nigroocellatus 202
ornatus 202
jaculatrix, Toxotes 147
japonica, Inegocia 73
japonicus, Calliurichthys 189
jarbua, Terapon 155
javanicus, Pseudogobius 206
Pseudorhombus 228
javus, Siganus 215
jerdoni, Sirembo 47
Johnius amblycephalus 139
macrorhynus 139
jonesi, Eubleekeria 111
kaakan, Pomadasys 126
kanagurta, Rastrelliger 221
katoptron, Cheilopogon 53
kelee, Hilsa 39
koilomatodon, Butis 191
Kumococius rodericensis 73
Kyphosidae 148
Kyphosus cinerascens 148
labiata, Pleurosicya 205
labiosus, Oedalechilus 51
Labridae 169
Labroides dimidiatus 173
lacunosus, Atherinomorus 52
Lagocephalus lunaris 243
spadiceus 244
suezensis 244
Laiphognathus multimaculatus 186
lambdastigma, Leptojulis 174
Larimichthys crocea 140
laterofenestra, Equulites 109
Lates calcarifer 78
Latidae 78
Leiognathidae 109

Leiognathus equulus 113
lentjan, Lethrinus 134
Leptojulis cyanopleura 174
lambdastigma 174
leptolepis, Selaroides 106
Leptomelanosoma indicum 137
lepturus, Trichiurus 219
Lethrinidae 132
Lethrinus genivittatus 133
lentjan 134
ornatus 134
variegatus 134
leuciscus, Equulites 110
leucogaster, Richardsonichthys 67
leurynnis, Acanthosphex 68
lewini, Sphyrna 14
Liachirus melanospilos 234
lighti, Entomacrodus 184
limbatus, Gerres 122
lineatus, Apogon 89
Istiblennius 186
Plotosus 41
lingua, Cynoglossus 236
lithinus, Ophichthus 27
longiceps, Hipposcarus 176
longicornis, Nuchequula 113
longimana, Drepane 149
longimanus, Pentaprion 123
Lophogobius bleekeri 203
lunare, Thalassoma 175
lunaris, Lagocephalus 243
Lutjanidae 116
Lutjanus argentimaculatus 116
fulviflamma 117
lutjanus 117
malabaricus 117
quinquelineatus 118
russellii 118
vitta 118
lutjanus, Lutjanus 117
lymma, Taeniura 19
macracanthus, Gerres 123
Priacanthus 86
Suggrundus 75
Macrodontogobius wilburi 203
macrolophus, Arnoglossus 230
macrorhynus, Johnius 139
macrosoma, Decapterus 103
macrurus, Pseudomonacanthus 242
maculata, Mene 108
maculatus, Arius 42
Pomadasys 126
malabaricus, Carangoides 102
Lutjanus 117
malayanus, Pseudorhombus 228
margaritifera, Scolopsis 130
marginatus, Halichoeres 172
Nemipterus 128
marmorata, Oxyeleotris 192
marmoratus, Atelomycterus 11
mata, Acanthurus 217
mate, Atule 100
maudae, Cryptocentrus 196
Megalaspis cordyla 104
Megalopidae 25
Megalops cyprinoides 25
melanoptera, Alepes 99
melanopterus, Carcharhinus 13
melanospilos, Liachirus 234
melanostictus, Pseudogobius 207

melanurus, Halichoeres 172
melapterus, Hemigymnus 173
melas, Neoglyphidodon 164
Mene maculata 108
Menidae 108
mentalis, Ulua 107
mento, Parexocoetus 54
meyeni, Taeniurops 20
microcephalus, Paratrypauchen 204
microprion, Cephalopholis 80
milleri, Pomacentrus 166
mingi, Hemigobius 201
Minous pictus 71
minuta, Gazza 112
moluccensis, Pomacentrus 167
Monacanthidae 240
Monacanthus chinensis 241
monoceros, Aluterus 240
Monodactylidae 146
Monodactylus argenteus 146
monogramma, Scolopsis 130
monopus, Cynoglossus 236
Moolgarda delicata 50
perusii 51
Mugilidae 49
Mugilogobius chulae 203
tigrinus 204
Mullidae 142
Mulloidichthys flavolineatus 142
multibarbus, Choridactylus 69
multimaculatus, Laiphognathus 186
Muraenesocidae 29
Muraenesox cinereus 29
muralis, Valenciennea 208
Myliobatidae 22
myops, Trachinocephalus 45
mystax, Thryssa 35
nalua, Ambassis 77
Narkidae 15
naucrates, Echeneis 97
nebulosus, Yongeichthys 209
neglecta, Scorpaenopsis 65
neglectus, Pseudorhombus 229
nehereus, Harpadon 44
Nemipteridae 127
Nemipterus bathybius 127
furcosus 128
hexodon 128
marginatus 128
tambuloides 129
zysron 129
Neoglyphidodon melas 164
Neomerinthe procurva 64
Neopomacentrus anabatoides 164
cyanomos 164
taeniurus 165
Nettastomatidae 30
Netuma thalassina 43
Nibea semifasciata 140
soldado 140
nichofii, Aetomylaeus 23
niger, Apogonichthyoides 91
Parastromateus 105
nigrescens, Halichoeres 172
nigrocincta, Apogon 89
nigrofasciata, Seriolina 107
nigroocellatus, Istigobius 202
nigroviridis, Tetraodon 244
novaezeelandiae, Brachypleura 226
novemradiatus, Periophthalmus 205

250
Nuchequula longicornis 113
nummifer, Antennarius 48
obesus, Thunnus 222
oblongus, Equulites 110
obtusirostris, Pristotis 168
ocellaris, Amphiprion 160
ocellatus, Aetobatus 22
Parachaetodon 152
octofasciatus, Chaetodon 151
Oedalechilus labiosus 51
Omobranchus elongatus 186
ferox 187
punctatus 187
Ophichthidae 27
Ophichthus lithinus 27
Ophidiidae 47
Ophiocara porocephala 192
Opisthopterus tardoore 31
orbicularis, Cyclichthys 245
Platax 212
orbis, Ephippus 211
orientalis, Brachirus 233
Dactyloptena 62
ornatus, Istigobius 202
Lethrinus 134
Ostorhinchus cooki 92
cyanosoma 92
Otolithes ruber 141
oualensis, Pempheris 145
Oxycheilinus digramma 175
Oxyeleotris marmorata 192
urophthalmus 192
oyena, Gerres 123
pacificus, Acanthopagrus 135
Pampus argenteus 223
chinensis 224
panchax, Aplocheilus 56
Parablennius thysanius 187
Parachaetodon ocellatus 152
Paralichthyidae 227
Paramonacanthus pusillus 242
Parapercis filamentosa 180
Paraplagusia blochii 236
Parascorpaena picta 65
Parastromateus niger 105
Paratrypauchen microcephalus 204
Pardachirus pavoninus 234
Parexocoetus mento 54
Parioglossus formosus 210
Parupeneus heptacanthus 143
parvonigra, Dasyatis 17
pavoninus, Pardachirus 234
Pelates quadrilineatus 154
pellucida, Eviota 197
Pempheridae 145
Pempheris oualensis 145
penicilligerus, Chaetodermis 241
Pennahia anea 141
Pentapodus setosus 129
Pentaprion longimanus 123
pentophthalmus, Pseudorhombus 229
perideraion, Amphiprion 160
Periophthalmus argentilineatus 204
novemradiatus 205
perusii, Moolgarda 51
petimba, Fistularia 59
phalaena, Amblygobius 194
Photopectoralis bindus 114
picta, Parascorpaena 65
picta picta, Diagramma 124

pictus, Minous 71
Pinguipedidae 180
pinguis, Atherinomorus 53
Pisodonophis cancrivorus 28
planus, Repomucenus 190
Platax orbicularis 212
Platycephalidae 72
Platycephalus cultellatus 74
indicus 74
Plectorhinchus gibbosus 125
pleuron, Apogon 90
Pleurosicya labiata 205
pleurostigma, Stigmatogobius 208
Plicofollis dussumieri 43
Plotosidae 41
Plotosus lineatus 41
poecilopterus, Apogon 90
poecilura, Gymnura 21
Polydactylus sextarius 137
Polynemidae 136
polyophthalmus, Grammatobothus 231
Pomacanthidae 153
Pomacanthus annularis 153
Pomacentridae 157
Pomacentrus alexanderae 165
chrysurus 165
coelestis 166
cuneatus 166
milleri 166
moluccensis 167
taeniometopon 167
tripunctatus 168
Pomadasys argenteus 125
kaakan 126
maculatus 126
unimaculatus 126
porocephala, Ophiocara 192
Priacanthidae 86
Priacanthus macracanthus 86
tayenus 87
Priolepis semidoliata 205
Pristicon trimaculatus 93
Pristigasteridae 31
Pristotis obtusirostris 168
procurva, Neomerinthe 64
prosopotaenia, Dischistodus 163
Psammogobius biocellatus 206
Psettodes erumei 225
Psettodidae 225
Pseudochromidae 85
Pseudochromis fuscus 85
ransonneti 85
Pseudogobius javanicus 206
melanostictus 207
Pseudomonacanthus macrurus 242
Pseudorhombus arsius 227
dupliciocellatus 228
javanicus 228
malayanus 228
neglectus 229
pentophthalmus 229
psittacus, Scarus 177
Ptereleotridae 210
Pterocaesio chrysozona 120
digramma 120
Pterois russelii 65
punctata, Drepane 150
punctatum, Chiloscyllium 9punctatus, Omobranchus 187
Siganus 215

puntang, Exyrias 198
pusillus, Paramonacanthus 242
putnamae, Sphyraena 218
quadrilineatus, Pelates 154
quagga, Zebrias 234
quinquelineatus, Cheilodipterus 92
Lutjanus 118
quinquestrigatus, Gobiodon 200
quoyanus, Epinephelus 84
quoyi, Scarus 178
Rachycentridae 96
Rachycentron canadum 96
ransonneti, Pseudochromis 85
ransonnettii, Elates 72
Rastrelliger kanagurta 221
Redigobius balteatus 207
reichei, Favonigobius 198
Repomucenus belcheri recurvispinnis 190
planus 190
reticulatus, Dascyllus 162
rhombea, Gazza 112
Rhynchobatidae 16
Rhynchobatus australiae 16
Richardsonichthys leucogaster 67
rochei rochei, Auxis 220
rodericensis, Kumococius 73
Rogadius tuberculatus 74
ruber, Otolithes 141
rubrum, Sargocentron 57
russelii, Pterois 65
russelli, Decapterus 103
Dendrophysa 139
russellii, Lutjanus 118
sagor, Hexanematichthys 43
Salarias fasciatus 188
Samaridae 232
Samaris cristatus 232
Sardinella albella 39
gibbosa 40
Sargocentron rubrum 57
Saurenchelys cancrivora 30
Saurida tumbil 44
undosquamis 45
scaber, Grammoplites 73
Scaridae 176
Scarus ghobban 177
psittacus 177
quoyi 178
Scatophagidae 213
Scatophagus argus 213
Sciaenidae 138
Scolopsis margaritifera 130
monogramma 130
taenioptera 131
vosmeri 131
Scomberoides commersonianus 105
tol 105
Scomberomorus commerson 222
Scombridae 220
Scorpaenidae 64
Scorpaenopsis neglecta 65
scutatus, Centriscus 60
Scyliorhinidae 11
Secutor hanedai 114
indicius 115
Selar boops 106
crumenophthalmus 106
Selaroides leptolepis 106
semidoliata, Priolepis 205
semifasciata, Nibea 140

251seminigracaudus, Apogon 90
semipunctata, Asterropteryx 195
Seriolina nigrofasciata 107
Serranidae 79
Setipinna taty 33
setirostris, Thryssa 35
setosus, Pentapodus 129
sexfasciatus, Abudefduf 158
Caranx 103
Epinephelus 84
sexguttata, Valenciennea 209
sextarius, Polydactylus 137
Siganidae 214
Siganus fuscescens 214
guttatus 215
javus 215
punctatus 215
virgatus 216
sihama, Sillago 94
Sillaginidae 93
Sillago aeolus 93
asiatica 94
chondropus 94
ingenuua 94
sihama 94
sinensis, Inimicus 71
Sirembo jerdoni 47
sirm, Amblygaster 38
soldado, Nibea 140
Soleidae 233
sonnerati, Cephalopholis 80
sordidus, Abudefduf 158
spadiceus, Lagocephalus 244
Sparidae 135
speciosus, Gnathanodon 104
Sphyraena forsteri 218
putnamae 218
Sphyraenidae 218
Sphyrna lewini 14
Sphyrnidae 14
Stegostoma fasciatum 10
Stegostomatidae 10
stellatus, Abalistes 238
stercorarius, Equulites 111
stethophthalmus, Amblygobius 194
Stigmatogobius pleurostigma 208
Stolephorus andhraensis 33
baganensis 34
indicus 34
storthynx, Eviota 197
striata, Helcogramma 182
striatus, Entomacrodus 184
Stromateidae 223

Strongylura strongylura 56
strongylura, Strongylura 56
subviridis, Chelon 49
suezensis, Lagocephalus 244
Sufflamen fraenatum 239
Suggrundus macracanthus 75
sulphureus, Upeneus 143
sundaicus, Upeneus 144
Synanceiidae 69
Syngnathidae 58
Synodontidae 44
Synodus tectus 45
taeniometopon, Pomacentrus 167
taenioptera, Scolopsis 131
Taeniura lymma 19
Taeniurops meyeni 20
taeniurus, Neopomacentrus 165
tambuloides, Nemipterus 129
tardoore, Opisthopterus 31
taty, Setipinna 33
tayenus, Priacanthus 87
tectus, Synodus 45
Temera hardwickii 15
Terapon jarbua 155
theraps 155
Terapontidae 154
tetradactylum, Eleutheronema 136
Tetraodon nigroviridis 244
Tetraodontidae 243
Tetrarogidae 66
thalassina, Netuma 43
Thalassoma lunare 175
theraps, Terapon 155
thoracata, Escualosa 39
Thryssa dussumieri 34
mystax 35
setirostris 35
Thunnus obesus 222
thysanius, Parablennius 187
Thysanophrys chiltonae 75
tigrinus, Mugilogobius 204
tol, Scomberoides 105
Torquigener gloerfelti 245
Toxotes jaculatrix 147
Toxotidae 147
Trachinocephalus myops 45
trachinoides, Vespicula 67
Trachyrhamphus bicoarctatus 59
tragula, Upeneus 144
Triacanthidae 237
triangularis, Drombus 196
Trichiuridae 219
Trichiurus lepturus 219

trilobatus, Cheilinus 170
trimaculatus, Dascyllus 162
Pristicon 93
Tripodichthys blochi 237
Tripterygiidae 182
tripunctatus, Pomacentrus 168
trivittatus, Iniistius 173
truncatus, Apogon 91
Trypauchen vagina 208
tuberculatus, Rogadius 74
tumbil, Saurida 44
tutuilae, Enneapterygius 182
Tylosurus crocodilus crocodilus 56
typus, Xiphocheilus 175
Ulua mentalis 107
undosquamis, Saurida 45
unimaculata, Chrysiptera 161
unimaculatus, Pomadasys 126
Upeneus guttatus 143
sulphureus 143
sundaicus 144
tragula 144
Uranoscopidae 181
Uranoscopus cognatus 181
urophthalmus, Oxyeleotris 192
urotaenia, Ambassis 77
vachellii, Ambassis 77
vagina, Trypauchen 208
vaigiensis, Ellochelon 50
Valenciennea muralis 208
sexguttata 209
wardii 209
vari, Alepes 100
variegatus, Eugnathogobius 197
Lethrinus 134
Vespicula trachinoides 67
virgatus, Siganus 216
viridipunctatus, Acentrogobius 194
vitta, Lutjanus 118
vorax, Champsodon 179
vosmeri, Scolopsis 131
walga, Himantura 19
wardii, Valenciennea 209
wilburi, Macrodontogobius 203
Xiphocheilus typus 175
yaeyamaensis, Ecsenius 184
Yongeichthys nebulosus 209
Zebrias quagga 234
Zenarchopterus dunckeri 55
zugei, Dasyatis 18
zysron, Nemipterus 129