Carpenter 1999b - TheLivingMarineResourcesOfTheWCP Vol4 FAO PDF

FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES
ISSN 1020-6868
THE LIVING MARINE RESOURCES OF THE
WESTERN CENTRAL
PACIFIC
Volume 4 Bony fishes part 2 (Mugilidae to Carangidae)
FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES
THE LIVING MARINE RESOURCES OF THE
WESTERN CENTRAL PACIFIC
VOLUME 4
Bony fishes part 2 (Mugilidae to Carangidae)
edited by
Kent E. Carpenter
Department of Biological Sciences
Old Dominion University
Norfolk, Virginia, USA
and
Volker H. Niem
Marine Resources Service
Species Identification and Data Programme
FAO Fisheries Department
with the support of the
South Pacific Forum Fisheries Agency (FFA)

and the
Norwegian Agency for International Development (NORAD)
FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS

Rome, 1999
ii
The designations employed and the presentation of material in

this publication do not imply the expression of any opinion
whatsoever on the part of the Food and Agriculture
Organization of the United Nations concerning the legal status
of any country, territory, city or area or of its authorities, or
concerning the delimitation of its frontiers and boundaries.
M-40
ISBN 92-5-104301-9
All rights reserved. No part of this publication may be reproduced by

any means without the prior written permission of the copyright owner.
Applications for such permissions, with a statement of the purpose
and extent of the reproduction, should be addressed to the Director,
Publications Division, Food and Agriculture Organization of the United
Nations, via delle Terme di Caracalla, 00100 Rome, Italy.
© FAO 1999
iii
Carpenter, K.E.; Niem, V.H. (eds)

FAO species identification guide for fishery purposes. The living marine resources of
the Western Central Pacific. Volume 4. Bony fishes part 2 (Mugilidae to
Carangidae).
Rome, FAO. 1999. pp. 2069-2790.
SUMMARY
This multivolume field guide covers the species of interest to fisheries of the major marine
resource groups exploited in the Western Central Pacific. The area of coverage includes
FAO Fishing Area 71 and the southwestern portion of Fishing Area 77 corresponding to
the South Pacific Commission mandate area. The marine resource groups included are
seaweeds, corals, bivalves, gastropods, cephalopods, stomatopods, shrimps, lobsters,
crabs, holothurians, sharks, batoid fishes, chimaeras, bony fishes, estuarine crocodiles,
sea turtles, sea snakes, and marine mammals. The introductory chapter outlines the en-
vironmental, ecological, and biogeographical factors influencing the marine biota, and
the basic components of the fisheries in the Western Central Pacific. Within the field
guide, the sections on the resource groups are arranged phylogenetically according to
higher taxonomic levels such as class, order, and family. Each resource group is intro-
duced by general remarks on the group, an illustrated section on technical terms and
measurements, and a key or guide to orders or families. Each family generally has an ac-
count summarizing family diagnostic characters, biological and fisheries information,
notes on similar families occurring in the area, a key to species, a checklist of species,
and a short list of relevant literature. Families that are less important to fisheries include
an abbreviated family account and no detailed species information. Species in the im-
portant families are treated in detail (arranged alphabetically by genus and species) and
include the species name, frequent synonyms and names of similar species, an illustra-
tion, FAO common name(s), diagnostic characters, biology and fisheries information,
notes on geographical distribution, and a distribution map. For less important species,
abbreviated accounts are used. Generally, this includes the species name, FAO com-
mon name(s), an illustration, a distribution map, and notes on biology, fisheries, and dis-
tribution. Each volume concludes with its own index of scientific and common names.
Production staff, Species Identification and Data Programme, Marine Resources

Service, Fishery Resources Division, Fisheries Department, FAO:
Project manager: P. Oliver.

Editorial assistance: N. DeAngelis and M. Kautenberger-Longo.
Desktop publishers: M. Kautenberger-Longo, N. DeAngelis, and G. Sciarappa-Demuro.
Scientific illustrator: E. D’Antoni.
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iv
Table of Contents
Page
BONY FISHES
Order MUGILIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2069
Mugilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2069
Order ATHERINIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2109
Pseudomugilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2109
Atherinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2113
Isonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2140
Telmatherinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2144
Dentatherinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2145
Phallostethidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2146
Order BELONIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2149
Adrianichthyidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2149
Belonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2151
Exocoetidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2162
Hemiramphidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2180
Order CYPRINODONTIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2197
Aplocheilidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2197
Poeciliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2199
Order STEPHANOBERYCIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2201
Melamphaidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2201
Gibberichthyidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2203
Rondeletiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2204
Barbourisiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2205
Cetomimidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2206
Mirapinnidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2208
Megalomycteridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2209
Order BERYCIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2210
Anoplogastridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2210
Diretmidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2211
Anomalopidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2212
Monocentrididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2214
Trachichthyidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2215
Berycidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2218
Holocentridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2225
Order ZEIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2257
Parazenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2257
Macrurocyttidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2258
Zeidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2259
Grammicolepididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2260
Caproidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2261
Order GASTEROSTEIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2262
Pegasidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2262
Solenostomidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2263
Syngnathidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2264
Aulostomidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2277
Fistulariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2278
Macrorhamphosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2280
Centriscidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2281
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Order SCORPAENIFORMES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2283
Dactylopteridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2283
Scorpaenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2291
Caracanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2353
Aploactinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2354
Triglidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2359
Bembridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2383
Platycephalidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2385
Hoplichthyidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2422
Cottidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2425
Psychrolutidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2427
Order PERCIFORMES: PERCOIDEI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2429
Centropomidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2429
Ambassidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2433
Acropomatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2436
Symphysanodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2438
Serranidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2442
Centrogeniidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2549
Ostracoberycidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2551
Callanthiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2553
Pseudochromidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2557
Plesiopidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2578
Notograptidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2586
Opisthognathidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2588
Priacanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2590
Apogonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2602
Epigonidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2611
Sillaginidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2614
Branchiostegidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2630
Lactariidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2649
Pomatomidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2650
Scombropidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2651
Echeneidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2652
Rachycentridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2655
Coryphaenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2656
Carangidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2659
INDEX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2757
LIST OF COLOUR PLATES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2790
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Mugiliformes: Mugilidae 2069
Mugiliformes: Mugilidae Order MUGILIFORMES
MUGILIDAE
Mullets
by I.J. Harrison and H. Senou
D iagnostic characters: Medium- to large-sized fishes; elongate with subcylindrical body. Head often
broad and flattened dorsally (except Aldrichetta and Cestraeus). Eyes often partly covered by
adipose “eyefold” tissue. Mouth small or moderate in size, terminal or inferior; premaxillae protractile;
teeth relatively small, hidden or absent. Two short dorsal fins, well separated; first with IV slender
spines; second dorsal fin usually with 9 or 10 soft rays; anal fin short with II or III spines and 7 to 12 soft
rays in adults; caudal fin emarginate or forked; pectoral fins inserted high on body; dorsal ray of pectoral
fins developed as a short spur or ‘spine’ (not a true spine); pelvic fins with I spine and 5 soft rays, inserted
about equidistant between origins of pectoral fins and first dorsal fin. Lateral line absent. Scales
large to moderate size, cycloid or ctenoid on head and body, with 1 or more longitudinal rows of striae
(grooves); with membranous hind margin in Valamugil; 24 to 64 scales in longitudinal series on midline,
counted from just behind head (behind operculum), above pectoral fins, to point of caudal flexure (i.e. not
including scales on caudal fin); 18 to 16 transverse scales, counted from origin of pelvic fins to origin of
first dorsal fin; 15 to 27 scales in transverse series entirely around caudal peduncle; large, modified scales
may be present above pectoral and pelvic fins (axillary scales) and below first dorsal fin. Oral and
branchial filter-feeding mechanism involving gill rakers and a specialized “pharyngobranchial organ”
comprising large, denticulate “pharyngeal pad” and pharyngeal “sulcus” on each side of pharyn-
gobranchial chamber (less evident in Aldrichetta and Cestraeus). Pharyngobranchial organ may be seen
by lifting operculum and pulling first 3 gill arches forward from fourth arch. The pharygeal pad is a large,
rounded structure with numerous, fine denticulate teeth giving an apparently “furry” surface. The sulcus is
the deep groove anterior to the denticulate pad. Anterior wall of sulcus may bear 1 or 2 “valves” which can
be difficult to see but, when present, are small to moderate flaps of tissue on lower or midpart of sulcus
and lying back against sulcus wall. Sulcus and valves best seen by pulling first 2 gill arches forward from
third and fourth arches. Stomach with muscular gizzard (except in Aldrichetta and Cestraeus) and pyloric
caeca positioned ventrally. Stomach and caeca can be seen by cutting the fish along the abdomen and
removing the liver, lying ventral to the alimentary tract. Intestine elongate and elaborately coiled (less
so in Aldrichetta, Cestraeus, and Myxus). Vertebrae 24 to 26. Colour: dark blue, dark olive, greenish, or
greyish dorsally; flanks silvery, often with more or less distinct dark stripes (about 3 to 9) following rows
of scales; ventral parts of body also silvery, or pale/yellowish; fins dusky or pale yellowish (particularly
pelvic fins) perhaps with margins dusky; dark bar or spot sometimes dorsally at base of pectoral fins.
pectoral modified
1st scale in axillary transverse scale
lateral count scale scales
adipose last scale in
eyefold lateral count
mouth
small
preorbital
bone pectoral fins
placed high pelvic axillary II-III valve
scale spines sulcus
pyloric
caecum
maxilla
“gizzard” denticulate
“crop” preorbital pharyngeal pad
gill arches 1-4
(these are cut, the dashed lines indicate the position
of the arches relative to pharyngobranchial organ)
scales stomach head with pharyngobranchial organ exposed
2070 Bony Fishes
Habitat, biology, and fisheries: Most species are euryhaline; inhabiting coastal marine waters, brackish
water lagoons, estuaries, and may enter fresh waters (particularly at young stages); usually to depths of
20 m, but will go deeper. Some species more typically inhabit fresh water but can be found in brackish
waters. Coastal species usually spawn offshore; fresh-water species spawn in brackish waters. Feeding:
browsing on submerged surfaces and filtering large quantities of benthic detritus; ingesting microalgae,
detritus, small invertebrates, micro-organisms, and particulate organic material. Several species are of
moderate to major importance to fisheries and are caught with diverse net types. From 1990 to 1995, the
FAO Yearbook of Fishery Statistics reports a range of yearly catch of Mugilidae of around 41 900 to 55 100 t
from the Western Central Pacific. Small-scale and subsistence fisheries are probably also relatively large.
The hardiness, simple diet, and rapid growth of mullets has made some species the object of aquaculture.
Remarks: Mugilidae are most speciose in the Indo-West Pacific region; many species are morphologically
very similar (e.g. Liza and Valamugil). Several nominal species appear to be conspecific and others show
a degree of morphological or meristic variability which makes straightforward diagnoses almost impossible.
The taxonomy of Mugilidae in this region is therefore very confused and parts of the following key and
diagnoses are provisional pending forthcoming revision.
Similar families occurring in the area

Atherinidae: body more slender, a prominent
silvery stripe along flanks; eyes larger; anal-fin
rays usually more than 10 (usually, but not
always, less than 10 in Mugilidae).
Atherinidae
Key to the species of Mugilidae occurring in the area
1a. Lower jaw bordered ventrally by a ridge of numerous lamellae (Fig. 1a); upper and lower
jaws ending posteriorly in fleshy lobes (Figs 1a and 3) . . . . . . . . . . . . . . . (Cestraeus) ® 2
1b. No such ridges of lamellae or fleshy lobes associated with jaws (Fig. 2) . . . . . . . . . . . . . ® 4
fleshy lobe
at end of
upper jaw
ridge of medial lobes

numerous of lower jaw
lamellae
a) ventrolateral view of head b) upper jaw teeth ventrolateral view of head
Fig. 1 Cestraeus oxyrhynchus Fig. 2 Liza
2a. Medial lobes on lower jaw not reaching corner of mouth (Figs 1a and 3a); teeth in upper
jaw multicuspid and close-packed (Fig. 1b) . . . . . . . . . . . . . . . . . . Cestraeus oxyrhynchus
2b. Medial lobes on lower jaw reaching corner of mouth (Fig. 3b and c); teeth in upper jaw
bicuspid or unicuspid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Pectoral fins as long as or medial
longer than head; usually lobes
distinct tooth patches on
vomer; interorbit strongly
convex (Fig. 4) and den-
tary symphysis pointed
(Fig. 3b) . . . . . . . Cestraeus goldiei
3b. Pectoral fins shorter than
head; vomer usually
edentate; interorbit only
moderately convex and
dentary symphysis more
a) Cestraeus oxyrhynchus b) Cestraeus goldiei c) Cestraeus plicatilis
ro un d ed th an p ointed
(Fig. 3c) . . . . . . Cestraeus plicatilis Fig. 3 ventral view of head
4a. Scales in longitudinal series 55 or more; several rows of teeth directly attached to jaw
bones (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aldrichetta forsteri
4b. Scales in longitudinal series less than 50; teeth borne on edges of lips (Fig. 6), often
minute or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
upper lip
teeth attached
directly to teeth borne
bones of on edge of tip
upper and
lower jaws
frontal view of head

lateral view of head lateral view of head
Fig. 4 Cestraeus
goldiei Fig. 5 Aldrichetta forsteri Fig. 6 Liza sp.
5a. Posteroventral edge of preorbital very strongly serrate, forming large spurs (Fig. 7a, b);
opercle with short spine at dorsoposterior corner, above pectoral-fin base (Fig. 7a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sicamugil hamiltoni
(fresh-water and brackish estuaries of Myanmar; not yet recorded from the area)
5b. Posteroventral edge of preorbital serrate, but not distinctly so, and not forming large
spurs (Fig. 8a, b); opercle lacking short spine at dorsoposterior corner (Fig. 8a) . . . . . . . . . ® 6
spine
preorbital preorbital
b) posteroventral b) posteroventral
a) lateral view of head edge of preorbital a) lateral view of head edge of preorbital
Fig. 7 Sicamugil hamiltoni Fig. 8 Liza sp.
6a. Head very distinctly flattened and concave between eyes (Fig. 9a); anterior nostril
positioned low on snout and eyes dorsolateral, high on head; snout projecting beyond
inferior, upper lip (Fig. 9b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinomugil nasutus
6b. Head moderately flattened and more or less convex between eyes (Fig. 10a); nostrils
positioned high on snout and eyes lateral on head; upper lip not inferior to snout
(Fig. 10b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
nostrils
nostrils
a) frontal view of head b) lateral view of head a) frontal view of head b) lateral view of head
Fig. 9 Rhinomugil nasutus Fig. 10 Liza sp.

2072 Bony Fishes
7a. Upper lip usually distinctly thickened, its depth at point of snout 5 to 11 times in head
length (usually less than 10 times in head length); lower part of upper lip with rugose
fringe or bearing distinct crenulations or papillae (Fig. 11a-c); (thickening of lip and
papillae may be less distinct in small specimens) . . . . . . . . . . (Crenimugil, Oedalechilus) ® 8
7b. Upper lip not distinctly thickened, its depth at point of snout greater than 12 times in
head length (usually about 15 to 20 times in head length); upper lip not rugose nor
bearing distinct crenulations or papillae (Fig. 11d) . . . . . . . . . . . . . . . . . . . . . . . . ® 10
papillae united into ridges upper lip with several preorbital deeply
2-5 rows on lateral part of lip rows of papillae notched
of papillae
on medial
part of lip
lower lip without lower lip turned down lip with rugose
crenate fringe with fine crenate fringe fringe
a) Crenimugil heterocheilus b) Crenimugil crenilabis c) Oedalechilus labiosus d) Liza sp.
Fig. 11 mouth
8a. Preorbital deeply notched; lips with distinct rugose fringe (Fig. 11c); upper lip thickened,
split into longitudinal lobes (Fig. 12); lips deeply folded into preorbital notch at corner of
mouth (Fig. 11c); 3 or 4 pyloric caeca . . . . . . . . . . . . . . . . . . . . . Oedalechilus labiosus
8b. Preorbital only slightly or not notched; upper lip not split into longitudinal lobes but
thickened with lower part bearing rows of papillae; lips not deeply folded into corners
of mouth (Fig. 11a, b); 6 to 10 pyloric caeca . . . . . . . . . . . . . . . . . . . . (Crenimugil) ® 9
9a. Upper lip with 1 to 10 rows of papillae on medial and lateral parts; upper lip thickness
5.2 to 9.5 times in head length; anterior edge of lower lip turned slightly out and down,
with fine crenate fringe on inner part (Fig. 11b) . . . . . . . . . . . . . . . . . Crenimugil crenilabis
9b. Upper lip with 2 to 5 rows of papillae, these rows united into file-like ridges on lateral
parts of lip; upper lip thickness 6 to 11 times in head length; lower lip thin, without crenate
fringe (Fig. 11a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Crenimugil heterocheilus
10a. Lower lip thick-edged, deflected downwards; teeth on lower lip directed down, with trifid
tips, and in 2 or more rows (Fig. 13a and b) . . . . . . . . . . . . . . . . . . . Neomyxus leuciscus
10b. Lower lip thin, directed forwards; teeth on lower lip directed forwards or upwards, without
trifid tips, or teeth absent (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
upper lip
lower
lip
lower
lower lip
lip teeth
frontal view of head a) lateral view of head b) tooth lateral view of head
Fig. 12 Oedalechilus labiosus Fig. 13 Neomyxus leuciscus Fig. 14 Liza sp.

11a. Maxilla straight, posterior tip not curved down (Fig. 15a); preorbital slender with straight
anteroventral edge and a pointed posteroventral end (Fig. 15a); adipose eyefold always
extensive in adults (Fig. 16a); dentary symphysis pointed, usually less than 90°
(Fig. 17a); in adults, anal fin with III spines and 8 or 9 soft rays (if 8, then this character
in combination with longitudinal series scale count of 36 to 44); (anal fin with II spines
and 9 or 10 soft rays in juveniles about 30 mm standard length or less) . . . . . . . . (Mugil) ® 12
11b. Maxilla with posterior tip slightly curved down (less distinct in Myxus) or more distinctly
sigmoidally curved (Fig. 15b, c); preorbital with a slightly concave or distinctly kinked
anteroventral edge and a blunt (Valamugil; Fig. 15b) or broad squarish (Liza, Myxus)
posteroventral end (Figs 15c and 20); adipose eyefold often marginal or absent in adults
(Fig. 16b, c), rarely as extensive as in Mugil (never extensive in species with more than
43 scales in longitudinal series); dentary symphysis usually blunt, more than 90°
(Fig. 17b); in adults, anal fins usually with III spines and 9 (rarely 8) soft rays (corre-
spondingly II spines and 10 or 9 soft rays in juveniles about 30 mm standard length or
less), except typically III spines and 8 soft rays in adult Liza vaigiensis with less than
30 scales in longitudinal series (cf. Mugil) . . . . . . . . . . . . . . (Myxus, Liza, Valamugil) ® 13
adipose eyefold adipose eyefold adipose eyefold

preorbital preorbital preorbital
maxilla
maxilla maxilla
a) Mugil b) Valamugil c) Liza a) Mugil b) Valamugil c) Liza
Fig. 15 lateral view of mouth (maxilla shaded) Fig. 16 lateral view of head
12a. Anal fin usually with 8 (rarely 9) soft

rays in adults; second dorsal and
anal fins with scales only on anterior
and basal parts of fins . . . . . . Mugil cephalus
12b. Anal fin with 9 soft rays in adults;
second dorsal and anal fins with
a) Mugil b) Liza
scales on all parts of fins . . Mugil broussonetti
Fig. 17 ventral view of head
13a. Scales in longitudinal series 43 or more; predorsal scales never keeled, and ventral
valve of pharyngobranchial organ appears as papilla covered pad (Fig. 18a); (2 pyloric
caeca - specimen must be dissected to see this) . . . . . . . . . . . . . . . . . . . . (Myxus) ® 14
13b. Scales in longitudinal series 43 or less; predorsal scales with or without keel; in species
with more than 40 scales in longitudinal series the pharyngobranchial organ either lacks
valves (Fig. 18b) and predorsal keel, or if valves present, then they are not papillose
and predorsal keel also present (Fig. 19); (4 or more pyloric caeca, except in Liza
argentea with 2 caeca but also less than 40 scales in longitudinal series, cf. Myxus)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Liza, Valamugil) ® 15
sulcus fat body

sulcus
valve
papillate denticulate area

valve pad denticulate area keels
a) Myxus petardi b) Valamugil sp. c) Liza vaigiensis
Fig. 18 lateral view of pharyngobranchial organs Fig. 19 Liza affinis (dorsal view)
2074 Bony Fishes
14a. Scales in longitudinal series 47 to 52; teeth in upper lip small and ciliiform or absent;
gill rakers on first arch 1/2 length of longest gill filaments . . . . . . . . . . . . . . . Myxus petardi
14b. Scales in longitudinal series 43 to 46; teeth in upper lip spatulate with constricted tips
(Fig. 20a, b); gill rakers on first gill arch as long as longest gill filaments . . . . . . Myxus elongatus
15a. Preorbital with a kink (often distinct) on the serrate anteroventral edge and a broad,
squarish, posteroventral tip (Figs 15c and 16c); maxilla stocky and curved sigmoidally
near posterior tip, which is often visible posteroventral to corner of closed mouth
(Figs 15c and 16c); scales ctenoid but lacking a membranous, digitated hind margin
(Fig. 21a); pharyngobranchial organ with 1 or usually 2 valves (Fig. 18c) . . . . . . . . (Liza) ® 16
15b. Preorbital with a more or less concave (rather than kinked) serrate anteroventral edge
and a narrow posteroventral tip (Figs 15b and 16b); maxilla slender and weakly curved
down at posterior tip, which is usually not clearly visible posteroventral to corner of
closed mouth (exposed when mouth opened) (Figs 15b and 16b); scales with a
membranous, digitated hind margin (Fig. 21b) or appearing cycloid where membrane
poorly developed (Valamugil georgii); pharyngobranchial organ with a broad sulcus
and lacking valves (Fig. 18b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Valamugil) ® 25
preorbital
teeth b) spatulate teeth

maxilla membranous
digitated hind
a) lateral view of mouth a) Liza b) Valamugil margin
Fig. 20 Myxus elongatus Fig. 21 scales
16a. Anal fin with 8 soft rays in adults; caudal fin

squarish, its posterior margin nearly straight;
pectoral fins black (lower section yellowish in
adults) (Fig. 22); (10 or more pyloric caeca -
specimen must be dissected to see this) . . Liza vaigiensis
16b. Anal fin with 9, or less frequently 8, soft rays in
adults; caudal fin clearly emarginate or forked;
pectoral fins perhaps dusky but never black;
(less than 10 pyloric caeca - specimen must be Fig. 22 Liza vaigiensis
dissected to see this) . . . . . . . . . . . . . . . . . . ® 17
17a. Predorsal scales with keel (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liza affinis

17b. Predorsal scales lacking keel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
18a. Scales in transverse series 13 to 15 ½ (Fig. 23); usually 10 or more anal-fin rays in
adults; gill rakers on lower limb of first gill arch 100 or more and very long (equal to or
longer than longest gill filaments) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
18b. Scales in transverse series less than 13 (Fig. 24); usually less than 10 anal-fin rays in
adults; gill rakers on lower limb of first gill arch less than 100 and of moderate or short
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
14 transverse scales 10 transverse scales
Fig. 23 Liza argentea Fig. 24 Liza alata

19a. Adults usually with 10 anal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liza argentea

19b. Adults with 11 anal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liza ramsayi
20a. All fins long and falcate; pelvic and pectoral fins longer than length of head minus snout
(Fig. 24); scale margins dark, giving reticulate appearance to flank pigmentation . . . . . Liza alata
20b. Fins less distinctly falcate and shorter, particularly pelvic, second dorsal, and anal fins
(Figs 25 and 26); pelvic fins shorter than length of head minus snout, and pectoral fins
usually shorter than or equal to head minus snout; flank pigmentation less obviously
reticulate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 21
21a. Body slender and elongate (body depth at origin of first dorsal fin 22% or less of standard
length, and at anal-fin origin 20% or less of standard length); head depressed and
pointed (Fig. 26); eye diameter usually less than 20% of head length; 5 to 7 indistict
stripes sometimes present on upper flanks of larger specimens . . . . . . . . . . . . Liza planiceps
21b. Body more robust (body depth at origin of first dorsal fin 22% or more of standard length,
and at anal-fin origin greater than 20% of standard length); head not markedly de-
pressed or pointed (Fig. 25); eye diameter 21% or more of head length; stripes on flanks
absent (except in Liza subviridis with 3 to 6 indistinct stripes) . . . . . . . . . . . . . . . . . . ® 22
11 transverse scales
Fig. 25 Liza macrolepis Fig. 26 Liza planiceps
22a. Upper lip with an outer row of short, yet distinct, peg-like setiform teeth which are very
close-set in a fine “comb”, and 1 or 2 inner rows of finer, more wide-set teeth (Fig. 27a);
31 to 35 (commonly 32 or 33) scales in longitudinal series, and 10 or 11 in transverse
series (Fig. 25); adipose eyefold marginal (Fig. 28a), or absent . . . . . . . . . . . Liza macrolepis
22b. Upper lip with an outer row of short, fine teeth which can be wide-set or moderately
close-set, but do not appear peg-like or form a distinct fine-set “comb” (cf. Liza
macrolepis); 1 or 2 inner rows of smaller ciliform or setiform teeth may be present, but
these are small and irregularly spaced (Fig. 27b) (cf. more regular arrangement in Liza
macrolepis); 25 to 32 (rarely 33; usually 30 or less) scales in longitudinal series, and 9
to 11 (rarely 12) in transverse series; adipose eyefold either marginal or partly devel-
oped over eye and side of head (Fig. 28b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
adipose eyefold adipose eyefold
outer row of teeth preorbital preorbital
maxilla maxilla
inner row of teeth
a) marginal eyefold b) partly developed eyefold
a) Liza macrolepis b) Liza subviridis e.g. Liza macrolepis e.g. Liza subviridis
Fig. 27 inner surface of upper lip Fig. 28 Liza spp.
23a. Lower limb of first gill arch with 30 to 45 gill rakers; body depth at origin of anal fin 29
to 33% of standard length; pectoral fins usually extend to ninth or tenth (rarely eighth
or eleventh) scale in longitudinal series . . . . . . . . . . . . . . . . . . . . . . . . . Liza parmata
23b. Lower limb of first gill arch usually with 45 to 68 or more gill rakers (rarely 41 to 44 in
Liza subviridis); body depth at origin of anal fin 21 to 29% of standard length; pectoral
fins extend to seventh or eighth (rarely ninth) scale in longitudinal series . . . . . . . . . . . . ® 24
2076 Bony Fishes
24a. Adipose eyefold covering about 1/3 to 1/2 of iris (in specimens about 10 cm standard
length or larger) (Fig. 28b; body appears fusiform or not distinctly deep in profile
(Fig. 29a), body depth at origin of first dorsal fin usually 22 to 26% of standard length
(sometimes attaining 27 to 30%); caudal fin bluish with dark pigmentation restricted to
margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liza subviridis
24b. Adipose eyefold marginal, not extending over iris (Fig. 28a); body appears more distinctly
“robust” or deep in profile (Fig. 29b), body depth at origin of first dorsal fin 27 to 31% (rarely
to 34%) of standard length; dark pigmentation over entire caudal fin . . . . . . . . . . Liza melinoptera
a) Liza subviridis b) Liza melinoptera

Fig. 29 lateral view
25a. Eighteen or more (usually 19 or 20) scales in transverse series entirely around caudal
peduncle (i.e. inclusive of both sides of body); origin of fully erected second dorsal fin
on vertical through anal-fin origin, or only just behind it (Fig. 30a, b) . . . . . . . . . . . . . . ® 26
25b. Eighteen or less (usually 16) scales in transverse series entirely around caudal pedun-
cle (i.e. inclusive of both sides of body); origin of fully erected second dorsal fin on
vertical through about third soft ray of anal fin, i.e. behind anterior quarter or more of
anal fin (Fig. 31a, b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .® 27
25th scale in 22nd scale in
longitudinal series longitudinal series
13th scale in 11th scale in falcate fin
longitudinal series longitudinal series
18-20 scales in transverse 19-20 scales in

series around caudal peduncle transverse series
around caudal
a) Valamugil seheli b) Valamugil buchanani
Fig. 30 lateral view peduncle
26a. Scales in longitudinal series 36 or more; second dorsal and anal fins not strongly falcate;
at least 23 (usually 24) scales in longitudinal series anterior to origin of second dorsal
fin (Fig. 30a); pectoral fins 84 to 104% of head length, but not often longer than head;
snout usually 18% or more of head length (rarely 17%) . . . . . . . . . . . . . . . Valamugil seheli
26b. Scales in longitudinal series 32 to 36 (rarely 37); second dorsal and anal fins falcate;
usually less than 24 scales in longitudinal series anterior to origin of second dorsal fin
(Fig. 30b); pectoral fins 90 to 139% of head length, often longer than head; snout 14 to
19% of head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Valamugil buchanani
16 scales in transverse 16 scales in transverse

10.5-12.5 transverse series around caudal 10.5-11.5 series around caudal
scales peduncle transverse scales peduncle
a) Valamugil engeli b) Valamugil perusii
Fig. 31 lateral view

27a Scales in longitudinal series 37 to 43 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 28
27b. Scales in longitudinal series 30 to 36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 29
adipose eyefold
28a. Second dorsal and anal fins weakly preorbital
scaled, with scales only on anterior preorbital
and basal parts; 62 to 75 gill rakers
on lower limb of first gill arch; maxilla
slightly curved downwards and ante-
rior edge of preorbital weakly con-
cave (Fig. 32a) . . . . . . . Valamugil cunnesius
28b. Second dorsal and anal fins well
scaled on all parts; 35 to 45 gill rakers maxilla maxilla
on lower limb of first gill arch; maxilla a) Valamugil cunnesius b) Valamugil speigleri
and anterior edge of preorbital al-
most straight (Fig. 32b) . . . .Valamugil speigleri Fig. 32
29a. Tip of pectoral fins distinctly anterior to level of first dorsal fin (Fig. 33); 12 or 13 (11 ½)
scales in transverse series; pyloric caeca extensively branched, up to 22 in total
(Fig. 34a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Valamugil georgii
29b. Pectoral fins just reaching level of origin of first dorsal fin or extending beyond this
(Fig. 31); 10 or 11 (rarely 12) scales in transverse series; 5 to 7 pyloric caeca (Fig. 34b) . . . . ® 30
pyloric pyloric
caeca caeca
12-13
transverse lateral view a) Valamugil georgii b) Valamugil perusii
scales Fig. 33 Valamugil georgii Fig. 34 stomach
anterior limit of
30a. Second dorsal and anal fins with scales only predorsal scales
on anterior and basal parts; adipose eyefold
rudimentary; origin of first dorsal fin usually
nearer base of caudal fin than anterior tip of
snout (Fig. 31); predorsal scales extending to
tip of snout (Fig. 35a) . . . . . . . . . . Valamugil engeli
30b. Second dorsal and anal fins moderately or
well scaled on all parts; adipose eyefold rea-
sonably developed, covering up to 1/2 of iris;
origin of first dorsal fin usually equidistant a) Valamugil engeli b) Valamugil perusii
between anterior tip of snout and base of Fig. 35 dorsal view of heads
caudal fin (Fig. 31); predorsal scales extend-
ing to posterior nostril (Fig. 35b) . . . . Valamugil perusii
List of species occurring in the area

The symbol 0 is given when species accounts are included.
0 Aldrichetta forsteri (Valenciennes, 1836)
0 Cestraeus goldiei Macleay, 1883
0 Cestraeus oxyrhynchus Valenciennes, 1836
0 Cestraeus plicatilis Valenciennes, 1836
0 Crenimugil crenilabis (Forsskål, 1775)
0 Crenimugil heterocheilus (Bleeker, 1855)
0 Liza affinis (Günther, 1861)
0
0 Liza argentea (Quoy and Gaimard, 1825)
Liza alata (Steindachner, 1892)
2078 Bony Fishes
0 Liza macrolepis (Smith, 1846)

0
0 Liza parmata (Cantor, 1850)
Liza melinoptera (Valenciennes, 1836)
0 Liza planiceps (Valenciennes, 1836)

0 Liza ramsayi (Macleay, 1883)
0 Liza subviridis (Valenciennes, 1836)
0 Liza vaigiensis (Quoy and Gaimard, 1825)
0 Mugil broussonneti Valenciennes, 1836
0 Mugil cephalus Linnaeus, 1758
0 Myxus elongatus Günther, 1861
0 Myxus petardi (Castelnau, 1875)
0 Neomyxus leuciscus (Günther, 1871)
0 Oedalechilus labiosus (Valenciennes, 1836)
0 Rhinomugil nasutus (DeVis, 1883)
1/
? Sicamugil hamiltoni (Day, 1869)
0 Valamugil buchanani (Bleeker, 1853)
0 Valamugil cunnesius (Valenciennes, 1836)
0 Valamugil engeli (Bleeker, 1858)
0 Valamugil georgii (Ogilby, 1897)
0 Valamugil perusii (Valenciennes, 1836)
0 Valamugil seheli (Forsskål, 1775)
0 Valamugil speigleri (Bleeker, 1859)
References
Ingham, S.E. 1952. The biology and taxonomy of the Mugilidae. Unpublished manuscript in the Library of the Natural
History Museum, London.
Schultz, L.P. 1946. A revision of the genera of mullets, fishes of the family Mugilidae, with descriptions of three new
genera. Proc. U.S. Natl. Mus., 96(3204):377-395.
Smith, J.L.B. 1935. The fishes of the family Mugilidae in South Africa. Ann. South African Mus., 30(5):587-644.
Smith, M.M. and J.L.B. Smith. 1986. Family Mugilidae. In Smith’s Sea Fishes, edited by M.M. Smith and P.C. Heemstra.
Johannesburg, Macmillan South Africa, pp. 714-720.
Thomson, J.M. 1954. The Mugilidae of Australia and adjacent seas. Austr. J. Mar. Freshw. Res., 5(1):70-131.
Thomson, J.M. 1984. Mugilidae. In FAO species identification sheets for fishery purposes. Western Indian Ocean
(Fishing Area 51), edited by W. Fischer and G. Bianchi. Vol. 3. Rome, FAO (unpaginated).
Thomson, J.M. 1997. The Mugilidae of the world. Mem. Qld. Mus., 41(3):457-562.
Yoshino T. and H. Senou. 1984. Family Mugilidae. In The fishes of the Japanese Archipelago, edited by H. Masuda, K.
amaoka, C. Araga, T. Uyeno, and T. Yoshino. Tokyo, Tokai University Press, pp. 119-120.
Weber, M. and L.F. de Beaufort. 1922. Fishes of the Indo Australian Archipelago . Vol. 4. Leiden, E.J. Brill, 448 p.
1/ Known from Sittang and Irrawaddy rivers and estuaries of Myanmar. Not yet recorded from the area, but might occur
along the Malay Peninsula.
Aldrichetta forsteri (Valenciennes, 1836)

Frequent synonyms / misidentifications: Agonostomus forsteri (Valenciennes, 1836); Agonostoma
diemensis (Richardson, 1840) / None.
FAO names: En - Yelloweye mullet.
Diagnostic characters: A small to medium-sized species; body slender. Interorbit not strongly convex.
Snout longer than eye diameter and moderately pointed. Dentary symphysis acutely pointed. Lips
thin, without papillae; lower lip directed forwards. No lobes associated with mouth. Teeth small, attached
directly to jaw bones, in 2 or 3 rows in each jaw, perhaps indistinct in large specimens. Sparse, movable,
ciliiform teeth attached on lower lip. Vomer well toothed. Maxilla and serrate anteroventral edge of preorbital
straight; posteroventral tip of preorbital broad and squarish. Distance between anterior and posterior nostril
less than maximum diameter of posterior nostril. Adipose eyefold absent or very rudimentary anteriorly.
Gill rakers on lower limb of first gill arch 24 to 48. Origin of first dorsal fin midway between tip of snout and
base of caudal fin, or slightly nearer latter. Origin of fully erected second dorsal fin on vertical through
anterior half of anal fin; both fins scaled only anterobasally. Anal fin with III spines and 12 (rarely 13)
soft rays in adults (usually II spines and 13 soft rays in juveniles about 30 mm standard length or less).
Caudal fin emarginate. Pectoral fins with I ‘spine’ and 15 soft rays, not reaching level of origin of first dorsal
fin; pectoral fins 19 to 23% standard length, 73 to 93% head length. Scales strongly ctenoid on abdomen,
weakly ctenoid or cycloid on other parts of body; 53 to 64 scales in longitudinal series; 14 or 15 in
transverse series; 22 or more (perhaps up to 27) scales in transverse series entirely around caudal
peduncle. Pharyngobranchial organ not well developed; sulcus V-shaped, shallow dorsally, a deep
furrow ventrally; tissue anterior to sulcus papillose; anterior and medial pharyngobranchial teeth
stout, directly attached to bones; posterior and lateral teeth loosely attached to denticulate pads.
Pyloric caeca 2. Colour: olive-brown dorsally, silvery to yellowish white on flanks and abdomen; eyes with
yellow iris; fins with brownish margin.
Size: Maximum reported total length 40 cm; commonly between 15 and 20 cm total length, rarely above
32 cm total length.
Habitat, biology, and fisheries: Inhabits coastal waters, bays, estuaries, and may ascend rivers into fresh
water. Usually found from the surface to a depth of 10 m, over sandy-muddy bottoms and seagrass
meadows. Frequently schooling. Probably spawns near the mouths of estuaries. Omnivorous, feeding on
benthic detritus, algae, and
worms. Constitutes an impor-
tant fishery; caught in beach
and seine nets and can be
taken by hook.
Distribution: Generally re-
stricted to temperate waters
of Australia and New Zealand.
Specimens might be found at
the southern extreme of the
area, in northern New South
Wales.
2080 Bony Fishes
Cestraeus goldiei (Macleay, 1883)

Frequent synonyms / misidentifications: Aeschrichthys goldiei Macleay, 1883 / None.
FAO names: En - Goldie river mullet.
Diagnostic characters: A small to medium-sized species; body relatively deep and robust. Eye diameter
14 to 22% head length. Interorbit convex. Snout longer than eye diameter but rather blunt in profile.
Dentary symphysis acutely pointed. Small fleshy lobes at ends of upper and lower jaws, in each corner
of mouth. Lower jaw also with lobes positioned medially in the gular region and extending
posteriorly as far as or beyond level of corner of mouth. Upper lip moderately thick, not papillate.
Lower lip thick-edged, directed forwards, and bordered ventrally by a ridge of numerous lamellae.
Teeth attached directly to jaw bones. Upper jaw with outer row of bicuspid or unicuspid teeth which
are not very close-packed; inner row of smaller, unicuspid teeth present or absent. Small teeth scarce
on lower jaw or absent in large specimens (about 30 cm standard length). Vomer usually with distinct
tooth patches on each lateral part. Maxilla and serrate anteroventral edge of preorbital straight;
posteroventral tip of preorbital broad and squarish. Distance between anterior and posterior nostril less
than maximum diameter of posterior nostril. Adipose eyefold absent or very rudimentary. Gill rakers on
lower limb of first gill arch 50 to 65. Origin of first dorsal fin midway between tip of snout and base of caudal
fin, or slightly closer to snout. Origin of fully erected second dorsal fin on vertical posterior to origin of anal
fin; both fins scaled only anteriorly and basally. Anal fin with III spines and 9 soft rays in adults (II spines
and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin emarginate. Pectoral fins
with I ‘spine’ dorsally, 13 or more soft rays enclosed in fin membrane and, ventrally, about 12 or more
soft rays free from fin membrane; pectoral fins reaching level of origin of first dorsal fin or beyond;
pectoral fins almost always as long as head or longer, 23 to 30% standard length. Ctenoid scales
in longitudinal series 36 to 43; 12 to 14 in transverse series; about 23 to 26 scales in longitudinal series
anterior to origin of second dorsal fin; 22 in transverse series entirely around caudal peduncle. Rudimen-
tary pharyngobranchial organ with narrow, shallow sulcus and 2 small denticulate pads bearing
stout, sessile teeth. Pyloric caeca 2. Colour: greyish dorsally, silvery on flanks and white ventrally; dorsal
fins and caudal fin dark with paler, speckled distal margins; dark spot or bar at base of caudal fin; anal fin
and pectoral fins darkish, but may have basal or distal margins paler; pelvic fins pale but with some dark
pigment on midportion of fin.
Size: Maximum reported fork length 40 cm, perhaps commonly reaching about 20 cm total length.
Habitat, biology, and fisheries: Fresh and brackish waters; adults ascending some way up rivers,
reported to 350 m above sea-
level. Probably taken inciden-
tally or as part of subsistence
fisheries in rivers.
Distribution: Indo-West Pa-
cific from Philippines to New
Caledonia. New Guinean re-
ports are commonly from Port
Moresby district.
Cestraeus oxyrhynchus Valenciennes, 1836

Frequent synonyms / misidentifications: None / None.
FAO names: En - Sharpnosed river mullet.
22 to 29% head length. Interorbit convex. Snout longer than eye diameter and pointed, in profile, in
adults. Dentary symphysis acutely pointed. Small fleshy lobes at ends of upper and lower jaws, in
each corner of mouth. Lower jaw also with lobes positioned medially in the gular region and not
extending posteriorly to level of corner of mouth. Upper lip moderately thick, not papillate. Lower lip
thick-edged, directed forwards, and bordered ventrally by a ridge of numerous lamellae. Teeth
attached directly to jaw bones. Upper jaw with outer row of close-packed, multicuspid teeth, and 1
to 3 inner rows of close-packed, spade-like teeth. Caniniform teeth sparse or absent on lower jaw.
Vomer with sparse teeth, in small patches on lateral parts of bone. Maxilla and serrate anteroventral edge
of preorbital straight; posteroventral tip of preorbital broad and squarish. Distance between anterior and
posterior nostril less than maximum diameter of posterior nostril. Adipose eyefold absent or very rudimen-
tary. Gill rakers on lower limb of first gill arch 36 to 54. Origin of first dorsal fin midway between tip of snout
and base of caudal fin. Origin of fully erected second dorsal fin on vertical posterior to origin of anal fin;
both fins scaled only anteriorly and basally. Anal fin with III spines and 9 (rarely 10) soft rays in adults
(usually II spines and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin emarginate.
Pectoral fins with I ‘spine’ and about 18 to 20 soft rays, ventral 4 or 5 soft rays free from fin membrane;
pectoral fins not reaching level of origin of first dorsal fin; pectoral fins 18 to 20% standard length,
75 to 85% head length. Ctenoid scales in longitudinal series 41 to 47; 12 to 14 in transverse series;
26 to 29 scales in longitudinal series anterior to origin of second dorsal fin; 22 in transverse series entirely
around caudal peduncle. Rudimentary pharyngobranchial organ with narrow, shallow sulcus and 2
small denticulate pads bearing stout, sessile teeth. Pyloric caeca 2. Colour: olivaceous dorsally, silvery
ventrally; dorsal fins and caudal fin uniformly speckled brown or black; vertical dark bar at base of caudal
fin; anal fin paler, speckled brown distally; pectoral fins weakly speckled and pelvic fins pale.
Size: Maximum reported total length 39 cm, perhaps commonly reaching about 20 cm total length.
Habitat, biology, and fisheries: Very little available data. Probably found in both fresh and brackish
waters, possibly ascending some way up rivers. Probably taken incidentally or as part of subsistence
fisheries in rivers.
cific from Indonesia to Fiji;
north to Philippines, south to
New Caledonia.
2082 Bony Fishes
Cestraeus plicatilis Valenciennes, 1836

FAO names: En - Lobed river mullet.
21 to 25% head length. Interorbit only moderately convex. Snout longer than eye diameter but
moderately blunt in profile. Dentary symphysis more rounded than pointed. Small fleshy lobes at
ends of upper and lower jaws, in each corner of mouth. Lower jaw also with lobes positioned
medially in the gular region and extending posteriorly as far as level of corner of mouth. Upper lip
moderately thick, not papillate. Lower lip thick-edged, directed forwards, and bordered ventrally by a
ridge of numerous lamellae. Teeth attached directly to jaw bones. Upper jaw with outer row of
bicuspid or unicuspid teeth which are not very close-packed; inner row of smaller, unicuspid teeth
present or absent. Small teeth scarce or absent on lower jaw. Vomer may be toothed, but usually
edentate. Maxilla and serrate anteroventral edge of preorbital straight; posteroventral tip of preorbital broad
and squarish. Distance between anterior and posterior nostril less than maximum diameter of posterior
nostril. Adipose eyefold absent or very rudimentary. Gill rakers on lower limb of first gill arch 52 to 66.
Origin of first dorsal fin midway between tip of snout and base of caudal fin, or slightly closer to snout.
Origin of fully erected second dorsal fin on vertical posterior to origin of anal fin; both fins scaled only
anteriorly and basally. Anal fin with III spines and 9 soft rays in adults (II spines and 10 soft rays in juveniles
about 30 mm standard length or less). Caudal fin emarginate. Pectoral fins with I ‘spine’ and about 18 or
19 soft rays, ventral 4 or 5 soft rays free from fin membrane; pectoral fins not reaching level of origin
of first dorsal fin; pectoral fins 19 to 21% standard length, shorter than head (83 to 95% head length).
Ctenoid scales in longitudinal series 40 to 45; 13 or 14 in transverse series; about 25 or 26 scales in
longitudinal series anterior to origin of second dorsal fin; 22 or 23 in transverse series entirely around
caudal peduncle. Rudimentary pharyngobranchial organ with narrow, shallow sulcus and 2 small
denticulate pads bearing stout, sessile teeth. Pyloric caeca 2. Colour: dark greenish dorsally, flanks
greyish, white ventrally; all fins well speckled brown or black except anterobasal part of anal fin and basal
part of pelvic fins, which are less speckled; dark spot or vertical bar at base of caudal fin.
Size: Maximum reported total length 32.5 cm, perhaps commonly reaching about 20 cm total length.
Habitat, biology, and fisheries: Very little available data. Probably found in both fresh and brackish
waters, possibly ascending
some way up rivers. Probably
taken incidentally or as part of
subsistence fisheries in riv-
ers.
Distribution: Celebes, New
Caledonia, Vanuatu, and Fiji.
Crenimugil crenilabis (Forsskål, 1775)

Frequent synonyms / misidentifications: Mugil cirrhostomus Valenciennes, 1836; M. macrocheilos
Bleeker, 1854 / None.
FAO names: En - Fringelip mullet; Fr - Mulet boxeur; Sp - Lisa labride.
(from Senou, 1988)
Diagnostic characters: A medium-sized species; body moderately deep. Head relatively flattened
dorsally. Snout shorter than eye diameter and blunt in profile. Dentary symphysis obtuse (blunt). Upper
lip very thick; lip thickness at point of snout 5.2 to 9.5 times in head length. Ventral quarter to half
of upper lip with 1 to 10 rows of papillae medially and laterally; papillae in lower rows are larger
than those in upper rows, flask-shaped, and may have bifid tips; papillae first appear in fish of about
6 cm standard length. Lower lip thin, with anterior margin turned out and downwards; 1 or 2 rows
of papillae on inner part of lower lip, giving a fine crenate fringe. Lips and vomer edentate. Maxilla
curved down, weakly sigmoid near posterior tip. Preorbital weakly concave (not kinked) on serrate
anteroventral edge and slightly expanded into squarish, posteroventral tip. Distance between anterior and
posterior nostril less than maximum diameter of posterior nostril. Adipose eyefold forms narrow rim around
eye. Gill rakers on lower limb of first gill arch 50 to 78. Origin of first dorsal fin midway between tip of snout
and base of caudal fin, or slightly closer to latter. Origin of fully erected second dorsal fin just posterior to
vertical level of origin of anal fin; both fins at least moderately scaled (on anterior and basal parts) or well
scaled on all parts. Anal fin with III spines and 9 (rarely 8 or 10) soft rays in adults (usually II spines and
10 soft rays in juveniles about 30 mm standard length or less). Caudal fin forked. Pectoral fins with I ‘spine’
and 15 to 18 (usually 16) soft rays, falcate, reaching origin of first dorsal fin; pectoral fins 20 to 27%
standard length, 76 to 109% head length (usually more than 90%). Scales cycloid or very weakly ctenoid;
36 to 42 (usually 37 to 40) scales in longitudinal series; 12 to 14 in transverse series; 24 to 26 scales
in longitudinal series anterior to origin of second dorsal fin; 19 or 20 scales in transverse series entirely
around caudal peduncle. Pharyngobranchial organ with broad sulcus, small papillose anterior valve and
longer-based, low posterior valve. Pyloric caeca 7 to 10. Colour: olive-green dorsally, flanks silvery and
abdomen whitish; fins greyish except pectoral fins which is yellowish and has distinct dark purplish spot
at upper part of base.
Size: Maximum reported standard length 50 cm; commonly to 26 cm standard length.
Habitat, biology, and fisheries: Found in coastal waters, in sandy or muddy areas of lagoons, reef flats
and tidepools; entering harbours. Forms large schools before spawning, at beginning of ebb tide (in June),
over shallow, open areas of the lagoon slope. Important food-fish in Polynesia; caught in gill nets and taken
as a bycatch with seines.
Marketed fresh. Also used as
live bait in pole-and-line tuna
fishing.
Distribution: W i d e s p r e a d
throughout tropical Indo-Pa-
cific, fr om R ed Sea a nd
Madagascar to Tuamotu Is-
lands; south to Lord Howe Is-
land and north to southern
Japan and, perhaps, Hawaii.
click for next page

2084 Bony Fishes
Crenimugil heterocheilus (Bleeker, 1855)

Frequent synonyms / misidentifications: None / Crenimugil crenilabis (Forsskål, 1755).
FAO names: En - Half fringelip mullet.
(from Senou, 1988)
Diagnostic characters: Gross morphology very similar to C. crenilabis. A medium-sized species; body
moderately deep. Head relatively flattened dorsally. Snout equal to or shorter than eye diameter, blunt in
profile. Dentary symphysis obtuse (blunt). Upper lip thick; lip thickness at point of snout 6 to 11 times
in head length. Ventral part of upper lip with 2 to 5 rows of papillae medially; papillae rows united
into file-like ridges on lateral parts of lip. Papillae not distinct in fish less than about 10 cm standard
length. Lower lip thin, not papillate or crenate. Lips edentate. Maxilla curved down, weakly sigmoid
near posterior tip. Preorbital weakly concave (not kinked) on serrate anteroventral edge and slightly
expanded into squarish, posteroventral tip. Anterior and posterior nostrils close together. Adipose eyefold
forms narrow rim around eye. Gill rakers on lower limb of first gill arch 37 to 57. Origin of first dorsal fin
midway between tip of snout and base of caudal fin, or slightly closer to latter. Origin of fully erected second
dorsal fin just posterior to vertical level of origin of anal fin; both fins lightly to moderately scaled. Anal fin
with III spines and 9 (rarely 8) soft rays in adults (usually II spines and 10 soft rays in juveniles about
30 mm standard length or less). Caudal fin forked. Pectoral fins with I ‘spine’ and 15 to 17 (usually 16) soft
rays, falcate, reaching origin of first dorsal fin; 21 to 27% standard length, 91 to 100% head length.
Scales cycloid or very weakly ctenoid; 35 to 41 (rarely 32) scales in longitudinal series; 11 or 12
(perhaps up to 14) in transverse series; 24 or 25 scales in longitudinal series anterior to origin of second
dorsal fin. Pyloric caeca 6 to 8. Colour: olivaceous dorsally, flanks and abdomen silvery or white; dorsal
fins blackish, second dorsal with orange tip; anal fin whitish with yellow tip; caudal fin pale bluish with tips
of lobes orange-yellow; pectoral fins bluish black with orange-yellow tip and dark spot at upper part of base.
Size: Maximum reported standard length 50 cm, perhaps commonly to 23 cm standard length.
Habitat, biology, and fisheries: Found in coastal waters and ascending rivers into fresh water, penetrating
far upstream from river mouth. Found in turbid channels and over gravel-bottoms. No fisheries data, but
recorded as a food fish.
Distribution: Tropical Indo-
Pacific from Indonesia and
Philippines to Vanuatu. North
to southern Japanese islands.
Liza affinis (Günther, 1861)

Frequent synonyms / misidentifications: None / Liza carinata (Valenciennes, 1836); L. klunzingeri
(Day, 1888).
FAO names: En - Eastern keelback mullet.
Diagnostic characters: A medium-sized species. Body rather elongate; depth at first dorsal fin 19 to 27%
standard length; depth at anal fin 17 to 22% standard length. Mid-dorsal line more or less sharply keeled
on scales anterior to first dorsal fin (keel weaker in small specimens); keel between first and second
dorsal fins weak or absent. Head deeper than wide, only weakly convex interorbitally. Eye diameter 22
to 32% head length. Snout shorter than or equal to eye diameter and slightly pointed. Dentary symphysis
90° or more obtuse. Upper lip thin or slightly thickened; outer row of close-set, fine, recurved, unicuspid
teeth; inner row of smaller teeth (sometimes indistinct). Lower lip thin, directed forwards, no teeth
present. Vomer edentate. Maxilla more or less sigmoidally curved near posterior tip, which is visible
posteroventral to corner of closed mouth. Serrate anteroventral edge of preorbital with distinct
kink; posteroventral tip broad and squarish. Distance between anterior and posterior nostril less than
maximum diameter of posterior nostril. Adipose eyefold relatively well developed, covering about 1
and 2 of iris. Gill rakers on lower limb of first gill arch 43 to 60, shorter than longest filaments. Origin of
first dorsal fin nearer tip of snout than base of caudal fin. Origin of fully erected second dorsal fin on vertical
through anterior third of anal fin; both fins moderately scaled (i.e. basally but not distally) or well
scaled (on all parts). Anal fin with III spines and 9 (rarely 8 or 10) soft rays in adults (usually II spines
and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin emarginate. Pectoral fins
with I ‘spine’ and 15 to 18 soft rays; short, not reaching origin of first dorsal fin; pectoral fins 14 to 18%
standard length, 67 to 76% head length; pectoral axillary scale rudimentary. Pectoral and pelvic fins shorter
than head minus snout. Scales weakly ctenoid on flanks, strongly ctenoid ventrally; 33 to 43 (usually 35
to 40) scales in longitudinal series, 11 to 13 in transverse series; 7 to 10 scales in longitudinal series
anterior to tip of pectoral fins and 23 or 24 anterior to origin of second dorsal fin; 17 to 20 in transverse
series entirely around caudal peduncle. Pharyngobranchial organ with wide sulcus and 2 large,
flap-like valves, the lower (anterior) one fimbriate. Pyloric caeca 4 to 6 (usually 5). Colour: bluish or
greenish dark dorsally, silver laterally and ventrally; several longitudinal dark stripes along scale rows on
middle part of trunk; upper part of iris orange; dorsal fins darkish; caudal fin with broad, dark margin and
dark at origin, but pale in middle part; anal fin partially darkish; pectoral fins darkish, basal part of uppermost
soft ray black but not forming distinctive spot; pelvic fins white.
Size: Maximum reported standard length 30 cm.
Habitat, biology, and fisheries: Inhabits inlet waters and estuaries of rivers. Used as a food fish in Japan
but perhaps not commercially
fished in the Western Central
Pacific.
Distribution: From Japan
(except northern Hokkaido)
through to Ryukyu Islands
and Taiwan Province of
China, and coasts from
Shanghai to Peihar and Hai-
nan Island. Might occur as a
very rare visitor to the north-
ern extreme of the area.
2086 Bony Fishes
Liza alata (Steindachner, 1892)

Frequent synonyms / misidentifications: Mugil diadema Gilchrist and Thompson, 1911; Pteromugil
diadema (Gilchrist and Thompson, 1911); Moolgarda (Planiliza) ordensis Whitley, 1945 / ?Liza
melinoptera (Valenciennes, 1836).
FAO names: En - Diamond mullet; Fr - Mulet diamant; Sp - Lisa diamante.
Diagnostic characters: A medium to large species. Body moderately deep, but elongate, and com-
pressed at caudal peduncle; body depth at first dorsal fin 20 to 25% standard length; depth at anal fin
21 to 22% standard length. Head usually as wide as deep, dorsally flattened. Eye diameter 16 to 20% head
length. Snout longer than eye diameter and more or less pointed. Dentary symphysis obtuse (blunt). Lips
thin. Upper lip with outer row of very small, close-set, unicuspid teeth and an inner band of up to 5
irregular rows of smaller teeth; inner band well spaced from outer row. Lower lip directed forwards
with minute ciliiform teeth present or absent. Vomer edentate. Maxilla sigmoidally curved near
posterior tip, which is visible posteroventral to corner of closed mouth. Serrate anteroventral edge
of preorbital with weak kink; posteroventral tip broad and squarish. Distance between anterior and
posterior nostril less than maximum diameter of posterior nostril. Adipose eyefold poorly developed as rim
around eye. Gill rakers on lower limb of first gill arch 64 to 79, shorter than longest filaments. Origin of first dorsal
fin more or less midway between tip of snout and base of caudal fin, or slightly nearer snout. Origin of fully
erected second dorsal fin on vertical through anterior one-third to two-thirds of anal fin; both fins well scaled
on all parts and falcate. Anal fin with III spines and 9 soft rays in adults (II spines and 10 soft rays
in juveniles about 30 mm standard length or less). Caudal fin deeply forked. Pectoral fins with I ‘spine’
and 16 soft rays; not reaching origin of first dorsal fin; pectoral fins 18 to 20% standard length, 78 to 95%
head length, falcate, greater than length of head minus snout; pectoral axillary scale absent. Pelvic
fins almost equal to pectoral-fin length, longer than head minus snout, falcate. Ctenoid scales in
longitudinal series 29 to 32; 10 to 12 in transverse series; 7 scales in longitudinal series anterior to tip
of pectoral fins and 19 to 22 (usually 19 or 20) anterior to origin of second dorsal fin; 16 scales in transverse
series entirely around caudal peduncle. Pharyngobranchial organ with anterior valve a fleshy
swelling, posterior valve a shallow fold. Pyloric caeca 5. Colour: light brown dorsally, flanks silvery,
abdomen whitish; scales on flanks, especially upper part of body, with dark margins giving reticulate
appearance, and with dark horizontal streak; first dorsal fin hyaline with olive spines; second dorsal and
pectoral fins grey; anal and caudal fins dusky with yellow or orange margin; pelvic fins bright yellow or orange.
Size: Maximum reported total length 75 cm; commonly to 35 cm total length.
Habitat, biology, and fisheries: Inhabits coastal waters, estuaries; sometimes ascending rivers into fresh
water. Preferring slow moving waters or still
lagoons. Water often turbid, over muddy
substrate, and with well-substantiated
aquatic vegetation; water temperature in
range 25° to 35°C. Feeds on microalgae,
detritus, terrestrial plant material, and
aquatic insects. Perhaps catadromous.
Distribution: Reported from South and
East Africa, Madagascar, northern Australia
(Timor Sea), New Guinea, and Tonga.
Liza argentea (Quoy and Gaimard, 1825)

Frequent synonyms / misidentifications: Mugil australis Steindachner, 1879; M. ferrandi Cuvier, 1929;
M. peronii Valenciennes, 1836 / Liza ramsayi (Macleay, 1883); Aldrichetta forsteri (Valenciennes, 1836).
FAO names: En - Flat-tail mullet.
Diagnostic characters: A small to medium-sized species. Body rather slender but with deep, moderately
compressed caudal peduncle. Head deeper than wide, somewhat flattened dorsally. Eye diameter 21 to
31% head length. Snout length approximately equals eye diameter and is more or less pointed. Dentary
symphysis 90° or more obtuse. Lips thin, each with a single row of minute setiform teeth (almost
invisible to naked eye). Vomer toothed. Maxilla sigmoidally curved near posterior tip, which is visible
posteroventral to corner of closed mouth. Preorbital relatively large; serrate anteroventral edge
slightly kinked; posteroventral tip broad and squarish. Anterior and posterior nostrils close together.
Adipose eyefold rudimentary, forming narrow rim around eye. Gill rakers on lower limb of first gill arch
100 to 150, very long, equal to or longer than longest filament. Origin of first dorsal fin usually nearer
base of caudal fin than tip of snout. Origin of fully erected second dorsal fin on vertical through anterior
quarter to half of anal fin; both fins moderately scaled on anterior and basal parts. Second dorsal fin
with 9 or 10 soft rays. Anal fin with III spines and 10 soft rays in adults (II spines and 11 soft rays in
juveniles about 30 mm standard length or less). Caudal fin forked. Pectoral fins with I ‘spine’ and 14 to 16
soft rays; not reaching origin of first dorsal fin; 19 to 23% standard length, 71 to 78% head length; pectoral
axillary scale absent. Scales weakly ctenoid; 34 to 38 in longitudinal series, 13 to 15 ½ in transverse
series; 10 to 12 scales in longitudinal series anterior to tip of pectoral fins and 23 to 25 scales anterior to
origin of second dorsal fin; 18 to 20 scales in transverse series entirely around caudal peduncle.
Pharyngobranchial organ with broad, deep, sulcus and 2 valves. Pyloric caeca 2. Colour: dorsally
steely-blue or light brown; flanks silvery with dark longitudinal stripes formed by alternating silver and dark
on scales; abdomen silvery; gold patch on posteroventral corner of operculum; iris purple with gold flecks;
dorsal, caudal and, to a lesser extent, anal fins dusky, but may have golden-yellow margins; origin of
pectoral fins dark.
Size: Maximum total length perhaps 45 cm; commonly between 15 and 22 cm.
Habitat, biology, and fisheries: Adults inhabit brackish waters. Abundant in estuaries; also found in
shallows, bogs, and saline lagoons. Fish younger than one year can be found in fresh waters, but not when
older. Forms schools and often jumps at surface. Spawning probably occurs at sea. Feeds on benthic
microscopic plants and ani-
mals. Fished from shallow,
coastal waters and estuaries
using gill nets and beach
seines. Marketed fresh and fil-
leted.
Distribution: Southern Aus-
tralia from Geraldton (Western
Australia) to Cooktown
(Queensland).
2088 Bony Fishes
Liza macrolepis (Smith, 1846) LZM

Frequent synonyms / misidentifications: Mugil borneensis Bleeker, 1851; M. troscheli Bleeker, 1858 /
Liza alata (Steindachner, 1892); L. melinoptera (Valenciennes, 1836); L. parsia (Hamilton Buchanan,
1822); L.subviridis (Valenciennes, 1836).
FAO names: En - Largescale mullet; Fr - Mulet à grandes écailles; Sp - Lisa godeya.
(from Senou, 1988)
Diagnostic characters: A medium to large species; body moderately robust; depth at first dorsal fin 26
to 31% standard length; depth at anal fin 23 to 25% standard length. Head depth equal to or greater than
width; head flattened dorsally. Eye diameter 21 to 30% head length. Snout length less than or equal to eye
diameter, moderately pointed. Dentary symphysis obtuse. Lips thin. Upper lip with outer row of very
close-set, small, peg-like unicuspid teeth, forming fine comb; 1 or 2 irregular rows of smaller, more
wide-set teeth; inner rows well spaced from outer row. Lower lip directed forwards with small ciliiform
teeth present or absent. Vomer toothed (sometimes not evident). Maxilla stocky and sigmoidally curved
near posterior tip, which is visible posteroventral to corner of closed mouth. Serrate anteroventral
edge of preorbital with moderate to distinct kink; posteroventral tip broad and squarish. Distance
between anterior and posterior nostril less than or equal to maximum diameter of posterior nostril. Adipose
eyefold poorly developed as rim around eye or absent. Gill rakers on lower limb of first gill arch 35 to 78,
shorter than longest filament. Origin of first dorsal fin closer to base of caudal fin than tip of snout. Origin
of fully erected second dorsal fin on vertical through anterior quarter to half of anal fin; both fins scaled
on anterobasal parts and often well scaled on all parts. Anal fin with III spines and 9 (rarely 8 or
10) soft rays in adults (usually II spines and 10 soft rays in juveniles about 30 mm standard length or
less). Caudal fin emarginate. Pectoral fins with I spine and 15 to 18 (usually 16, rarely 14) soft rays; not
reaching origin of first dorsal fin; pectoral fins 17 to 23% (usually 20%) standard length, 65 to 85% (usually
75 to 80%) head length; less than length of head minus snout; pectoral axillary scale rudimentary or absent.
Ctenoid scales in longitudinal series 31 to 35 (usually 32 or 33); 10 or 11 in transverse series; 7 to
9 scales in longitudinal series anterior to tip of pectoral fins and 19 to 23 (usually 21 or 22) anterior to origin
of second dorsal fin; 16 scales in transverse series entirely around caudal peduncle. Pharyngobran-
chial organ with large denticulate area; broad sulcus with teeth on anterior border slightly larger than other
teeth; anterior and posterior valves are large flaps. Pyloric caeca 4 to 6. Colour: greenish grey dorsally;
flanks and abdomen silvery; fins grey or bluish with dusky margins and may appear yellowish at base;
pectoral fins with golden base, and perhaps a dark spot.
Size: Maximum reported total length about 60 cm; commonly to 26 cm total length.
Habitat, biology, and fisheries: Schools in shallow coastal waters, estuaries, and backwaters. Can survive
in ponds at 87% salinity; also sometimes enters fresh water. Juveniles might be found in shallow inundated
areas along coast. Forms large schools during spawning at sea. Feeds on small algae, diatoms, foraminifera,
benthic polychaets, crustacea, molluscs, organic matter and detritus; fry take copepods and floating algae.
Fished throughout its range; taken with gill nets, cast nets,
stake nets, barrier nets, lift nets, beach seines, and pouch
nets during spawning. Marketed fresh; roe also mar-
keted.
Distribution: Common throughout most of Indo-Pacific
from East Africa and Red Sea to Marquesas and Tuamoto
islands; north to Japan and Marianas Islands.
Liza melinoptera (Valenciennes, 1836)

Frequent synonyms / misidentifications: Mugil ceramensis Bleeker, 1852; M. oligolepis (of Smith,
1935) / Liza alata (Steindachner, 1892); L. macrolepis (Smith, 1846); L. parmata (Cantor, 1850);
L. subviridis (Valenciennes, 1836); L. vaigiensis (Quoy and Gaimard, 1825).
FAO names: En - Otomebora mullet; Fr - Mulet otomebora; Sp - Lisa otomebora.
(from Senou, 1988)
Diagnostic characters: A small to medium-sized species; body distinctly robust or deep in profile; caudal
peduncle compressed; body depth at origin of first dorsal fin 27 to 31% (perhaps up to 34%)
standard length; depth at origin of anal fin 23 to 29% standard length. Head wider than deep and
depressed. Eye diameter 24 to 28% head length. Snout shorter than eye diameter, blunt in profile. Dentary
symphysis obtuse. Lips thin. Upper lip with outer row of close-set, small setiform teeth and inner row
of more sparse teeth, well embedded in lip tissue. Lower lip directed forwards with small ciliiform
teeth present or absent. Vomer edentate. Maxilla stocky and sigmoidally curved near posterior tip,
which is often visible posteroventral to corner of closed mouth. Serrate anteroventral edge of
preorbital with moderate to distinct kink; posteroventral tip broad and squarish. Distance between
anterior and posterior nostril less than maximum diameter of posterior nostril. Adipose eyefold usually
only a narrow rim around eye. Gill rakers on lower limb of first gill arch 45 to 59 (perhaps more),
shorter than longest filaments. Origin of first dorsal fin distinctly closer to base of caudal fin than tip of
snout. Origin of fully erected second dorsal fin on vertical through anterior half to three-quarters of anal
fin; both fins moderately scaled (i.e. on all but distal parts) or well scaled on all parts. Anal fin with
III spines and 9 (rarely 8 or 10) soft rays in adults (usually II spines and 10 soft rays in juveniles about
30 mm standard length or less). Caudal fin emarginate. Pectoral fins with I ‘spine’ and 14 or 15 (infrequently
16) soft rays; not reaching origin of first dorsal fin; pectoral fins 18 to 23% standard length, 67 to 80%
(infrequently up to 89%) head length, less than or equal to length of head minus snout; pectoral axillary
scale rudimentary or absent. Scales ctenoid, usually 26 to 29 (perhaps up to 31) in longitudinal series;
9 to 11 in transverse series; 7 scales in longitudinal series anterior to tip of pectoral fins and 18 to
20 anterior to origin of second dorsal fin; 16 in transverse series entirely around caudal peduncle.
Pharyngobranchial organ with large denticulate area; anterior valve a small finger-like process,
posterior valve a large fold with papillate margin. Pyloric caeca 5 or 6. Colour: olive-brown dorsally,
silvery or white ventrally; distal margins of dorsal fins and entire caudal fin dusky.
Size: Maximum reported standard length 30 cm; commonly to 18 cm total length.
Habitat, biology, and fisheries: Schools in shallow coastal waters; enters lagoons, estuaries and rivers,
to feed; fry may be found in shallower areas of these regions. Feeds on plant detritus, microalgae, minute
bottom-living organisms, and organic mat-
ter contained in sand and mud; juveniles
take planktonic algae. Fished from shallow
coastal waters, estuaries, backwaters, and
lagoons using cast nets, stake nets, beach
seines, and gill nets. Marketed fresh.
Distribution: Indo-Pacific from East Africa
to the Marquesas Islands; north to South
China Sea and south to tropical Australia
and Tonga.
2090 Bony Fishes
Liza parmata (Cantor, 1850)

Frequent synonyms / misidentifications: Liza oligolepis (Bleeker, 1859) / Liza macrolepis (Smith,
1846); L. melinoptera (Valenciennes, 1836); L. subviridis (Valenciennes, 1836).
FAO names: En - Broadmouthed mullet.
(after Day, 1876)
Diagnostic characters: A medium-sized species. Deep bodied and compressed at caudal peduncle;
body depth at first dorsal fin 30 to 37% standard length; depth at anal fin 29 to 33% standard length.
Head deeper than wide and flat dorsally. Eye diameter 24 to 29% head length. Snout length less than or
equal to eye diameter, and moderately blunt in profile. Dentary symphysis very obtuse (blunt); mouth
appearing transverse in ventral view. Lips thin. Upper lip with 1 row of small, setiform teeth. Lower
lip directed forwards and lacking teeth. Vomer edentate. Maxilla sigmoidally curved near posterior tip,
which is visible posteroventral to corner of closed mouth. Serrate anteroventral edge of preorbital
with distinct kink; posteroventral tip broad and squarish. Distance between anterior and posterior
nostril usually less than or equal to maximum diameter of posterior nostril. Adipose eyefold poorly
developed as rim around eye or absent. Gill rakers on lower limb of first gill arch 30 to 45, shorter than
longest filaments. Origin of first dorsal fin distinctly closer to base of caudal fin than tip of snout. Origin of
fully erected second dorsal fin on vertical through anterior quarter to half of anal fin; both fins scaled on
anterobasal parts and may be scaled distally. Anal fin with III spines and 9 soft rays in adults (II
spines and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin emarginate. Pectoral
fins with I spine and 14 or 15 (rarely 13) soft rays; almost reaching origin of first dorsal fin; pectoral fins 20
to 26% standard length, 83 to 91% head length; pectoral axillary scale rudimentary or absent. Scales
cycloid or ctenoid (thorasic and abdominal scales ctenoid); 23 to 32 (usually 25 to 28) scales in
longitudinal series, 9 or 10 (rarely 11) in transverse series; 9 or 10 (rarely 8 or 11) scales in
longitudinal series anterior to tip of pectoral fins and 19 to 22 anterior to origin of second dorsal fin;
15 or 16 in transverse series entirely around caudal peduncle. Pharyngobranchial organ with
anterior valve a long, low fold with a small ventral finger-like process, and posterior valve a
moderate size, more or less fimbriate flap. Pyloric caeca 5 to 9. Colour: greenish dorsally, silvery
ventrally.
Size: Maximum reported total
length 30 cm.
Habitat, biology, and fisher-
ies: Found in seas, estuaries,
and rivers. No other available
data.
Distribution: Malaysia, Indo-
nesia, New Guinea, Philip-
pines, and South China Sea.
Liza planiceps (Valenciennes, 1836)

Frequent synonyms / misidentifications: Mugil belanak Bleeker, 1857; M. tade Forsskål, 1775; Liza
tade (Forsskål, 1775) / Liza parsia (Hamilton Buchanan, 1822); L. subviridis (Valenciennes, 1836); L.
vaigiensis (Quoy and Garmard, 1825).
FAO names: En - Tade mullet; Fr - Mulet tade; Sp - Lisa tade.
Diagnostic characters: A medium to large species. Body elongate; depth at first dorsal fin 20 to 22%
standard length; depth at anal fin 19 to 20% standard length. Head broad, depressed, pointed; head
width greater than or equal to depth; head may appear somewhat bulging laterally. Eye diameter 14 to
33% head length, usually less than 20%. Snout length greater than or equal to eye diameter; pointed in
profile. Dentary symphysis distinctly obtuse (blunt). Lips thin. Upper lip with outer row of fine, unicuspid,
setiform teeth; inner band of 2 to 4 (perhaps 8) rows of smaller, finer, unicuspid setiform teeth.
Lower lip directed forwards with 1 row of sparse ciliiform teeth present or absent. Vomer toothed.
Maxilla sigmoidally curved near posterior tip, which is often visible posteroventral to corner of
closed mouth. Serrate anteroventral edge of preorbital with distinct kink; posteroventral tip broad
and squarish. Distance between anterior and posterior nostril less than maximum diameter of posterior
nostril. Adipose eyefold more or less developed, covering up to 1 and 2 of iris. Gill rakers on lower
limb of first gill arch 48 to 70. Origin of first dorsal fin midway between tip of snout and base of caudal fin
or slightly nearer the latter. Origin of fully erected second dorsal fin on vertical through anterior third to half
of anal fin; both fins well scaled basally but distal parts are sometimes weakly scaled. Anal fin with
III spines and 9 soft rays in adults (II spines and 10 soft rays in juveniles about 30 mm standard length
or less). Caudal fin forked. Pectoral fins with I ‘spine’ and 15 to 16 soft rays; short, not reaching origin of
first dorsal fin; pectoral fins 16 to 20% standard length, 70 to 80% head length, less than length of head
minus snout; pectoral axillary scale rudimentary or absent. Pelvic fins shorter than head minus snout.
Scales ctenoid, sometimes very weakly so; 30 to 35 in longitudinal series; 10 or 11 in transverse series;
8 or 9 scales in longitudinal series anterior to tip of pectoral fins and 21 to 23 anterior to origin of second
dorsal fin; 16 scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ
with reduced denticulate area; broad sulcus; anterior valve a low fold and posterior valve a small, thin
flap. Pyloric caeca 5. Colour: greenish brown dorsally, whitish ventrally; flanks with 5 to 7 indistinct
longitudinal lines along scale rows.
Size: Maximum reported total length 70 cm; commonly to 33 cm standard length.
Habitat, biology, and fisheries: Usually marine, found in schools in shallow coastal waters and lagoons.
Also enters estuaries and rivers to feed. Juveniles might be found in rice fields and mangrove swamps.
Spawning occurs at sea. Feeds on floating algae, minute benthic organisms, and organic material in sand
and mud. Fished in shallow coastal waters,
estuaries, and backwaters using gill nets,
stake nets, beach seines, and cast nets.
Marketed fresh, salted; perhaps also boiled
(Thailand), canned or frozen (Australia).
Roe marketed salted.
Distribution: Indo-Pacific; from Red Sea in
west to China and Marianas Islands in north,
and Vanuatu in south. Reports from tropical
Australia might be misidentifications of
other species.
2092 Bony Fishes
Liza ramsayi (Macleay, 1883)

Frequent synonyms / misidentifications: Gracimugil ramsayi (Macleay, 1883); Trachystoma ramsayi
(Macleay, 1883) / Liza argentea (Quoy and Gairmard, 1825).
FAO names: En - Ramsay’s mullet.
Diagnostic characters: A small to medium-sized species; morphologically very similar to Liza argentea,
hence considered by some biologists as conspecific. Body moderately deep. Eye diameter 25% head
length. Snout length more or less equals eye diameter and is pointed. Dentary symphysis 90° or more
obtuse. Upper lip perhaps slightly thickened; lower lip thin. Each lip with a single row of minute setiform
teeth (almost invisible to naked eye). Vomer toothed. Maxilla sigmoidally curved near posterior tip,
which is visible posteroventral to corner of closed mouth. Preorbital relatively large; serrate an-
teroventral edge slightly kinked; posteroventral tip broad and squarish. Anterior and posterior nostrils
close together. Adipose eyefold rudimentary, forming narrow rim around eye. Gill rakers on lower limb
of first gill arch extremely numerous and as long as gill filaments. Origin of first dorsal fin nearer base
of caudal fin than tip of snout. Origin of fully erected second dorsal fin on vertical through anterior half of
anal fin; both fins moderately scaled. Second dorsal fin with 9 or 10 soft rays. Anal fin with III spines
and 11 (rarely 10) soft rays in adults (usually II, 12 in juveniles about 30 mm standard length or less).
Caudal fin forked. Pectoral fins with I ‘spine’ and 15 soft rays, not reaching origin of first dorsal fin; pectoral
fins 21 to 25% standard length, 77 to 91% head length; pectoral axillary scale very small or absent. Scales
cycloid dorsally, weakly ctenoid laterally and more distinctly ctenoid ventrally; 36 or 37 in longitudinal
series, about 12 to 15 in transverse series; 10 scales in longitudinal series anterior to tip of pectoral fins
and 23 anterior to origin of second dorsal fin. Colour: bluish dorsally, silvery on flanks and abdomen; dark
spot at origin of pectoral fins.
Size: Maximum reported fork length 26.1 cm.
Habitat, biology, and fisheries: Brackish waters. No fisheries data.
Distribution: Tropical Aus-
tralia; all records coming from
Queensland.
Liza subviridis (Valenciennes, 1836)

Frequent synonyms / misidentifications: Mugil alcocki Ogilby, 1908; M. dussumieri Valenciennes,
1836; M. javanicus Bleeker, 1852; M. jerdoni Day, 1876; M. sudanensis Bleeker, 1853 / Liza macrolepis
(Smith, 1846); L. parmata (Cantor, 1850); L . parsia (Hamilton Buchanan, 1822); L . planiceps
(Valenciennes, 1836); L. vaigiensis (Quoy and Gaimard, 1825).
FAO names: En - Greenback mullet; Fr - Mulet dos vert; Sp - Lisa lomo verde.
Diagnostic characters: A medium-sized species; body fusiform or not distinctly deep; depth at first
dorsal fin usually 22 to 26% standard length (but may reach 27 to 30%); depth at anal fin 21 to 24%
standard length (perhaps up to 26%). Head broad, but head depth usually equal to or greater than width;
head dorsally flattened. Eye diameter 21 to 30% head length. Snout short and blunt, usually shorter than
eye diameter. Dentary symphysis obtuse (blunt). Lips thin. Upper lip with an outer row of very small,
close-set unicuspid teeth and 1 or 2 irregular, inner rows of smaller, ciliiform teeth. Lower lip directed
forwards with 1 row of fine ciliiform teeth present or absent. Vomer edentate. Maxilla sigmoidally
curved near posterior tip, which is often visible posteroventral to corner of closed mouth. Serrate
anteroventral edge of preorbital with moderate to distinct kink; posteroventral tip broad and
squarish. Distance between anterior and posterior nostril less than maximum diameter of posterior nostril.
Adipose eyefold moderately developed, covering up to 1 and 2 of iris (less distinct in specimens
smaller than about 10 cm standard length). Gill rakers on lower limb of first gill arch 40 to 68 (rarely
41 to 44), shorter than longest filaments. Origin of first dorsal fin midway between tip of snout and base of
caudal fin, or just closer to latter. Origin of fully erected second dorsal fin on vertical through anterior one
to two-thirds of anal fin; both fins well scaled on all parts. Anal fin with III spines and 9 (rarely 8) soft
rays in adults (usually II spines and 10 soft rays in juveniles about 30 mm standard length or less). Caudal
fin emarginate. Pectoral fins with I ‘spine’ and 14 to 16 (usually 14 or 15) soft rays; short, not reaching
origin of first dorsal fin; pectoral fins 16 to 22% standard length, 70 to 78% head length, less than length
of head minus snout; pectoral axillary scale rudimentary or absent. Ctenoid scales in longitudinal series
28 to 32 (rarely 26, 27, or 33); 9 to 11 (rarely 12) in transverse series; 7 or 8 (rarely 9) scales in
longitudinal series anterior to tip of pectoral fins and 18 to 21 (usually 19 or 20) anterior to origin of
second dorsal fin; 16 or 17 (rarely 15) in transverse series entirely around caudal peduncle.
Pharyngobranchial organ usually with anterior valve as one or more small, finger-like processes,
and posterior valve a small fold; but some putatively conspecific specimens have both valves
developed as moderate size flaps. Pyloric caeca 4 to 6. Colour: dark greenish dorsally, brownish over
head, white ventrally; 3 to 6 indistinct, dark stripes along upper rows of scales; first dorsal fin greyish;
second dorsal fin yellowish grey, with dusky margin; anal fin also with dusky margin; caudal fin bluish
with black margin; pectoral fins yellowish and may have a blue spot at fin origin.
Size: Maximum reported standard length 40 cm; commonly to 25 cm total length.
Habitat, biology, and fisheries: Schools in shallow coastal waters and enters lagoons, estuaries, and
fresh waters to feed. Juveniles might be found in rice fields and mangroves. Adults feed on microalgae,
filamentous algae, diatoms, and benthic detrital material taken in with sand and mud; fry take zooplankton,
diatoms, detrital material, and inorganic sediment.
Fished using gill nets, cast nets, stake nets, lift nets, and
beach seines. Marketed fresh and salted; perhaps also
boiled (Thailand), canned or frozen (Australia). Roe mar-
keted salted. Also used as a bait-fish.
Distribution: Indo-Pacific from Red Sea to Samoa; north
to Japan, where it is rare.
2094 Bony Fishes
Liza vaigiensis (Quoy and Gaimard, 1825)

Frequent synonyms / misidentifications: Liza waigiensis (Quoy and Gairmard, 1825); Ellochelon
vaigiensis (Quoy and Gairmard, 1824) / ?Liza melinoptera (Valenciennes, 1836); L. parmata (Cantor,1850).
FAO names: En - Squaretail mullet (= Diamond-scale mullet, Fishing Areas 57 and 71); Fr - Mulet mopiro;
Sp - Lisa mopiro.
(from Senou, 1988)
Diagnostic characters: A medium to large species. Body robust; depth at first dorsal fin 20 to 27%
standard length; depth at anal fin 23 to 25% standard length. Caudal peduncle deep. Head broad, wider
than deep (except in young) and dorsally flattened. Eye diameter 17 to 33% head length. Snout longer
than eye diameter in adults but shorter in juveniles. Dentary symphysis obtuse (blunt). Lips thin. Upper lip
with 1 or 2 rows of very short, unicuspid, setiform teeth (absent in specimens over about 25 cm
standard length). Lower lip directed forwards with 1 row of fine ciliiform teeth (absent in specimens
over 10 cm standard length). Vomer edentate. Maxilla slightly sigmoidally curved near posterior tip,
which is visible posteroventral to corner of closed mouth. Serrate anteroventral edge of preorbital
with weak kink; posteroventral tip broad and squarish. Distance between anterior and posterior nostril
less than or equal to maximum diameter of posterior nostril. Adipose eyefold poorly developed as rim
around eye. Gill rakers on lower limb of first gill arch 40 to 67, shorter than longest filaments. Origin of first
dorsal fin distinctly closer to base of caudal fin than tip of snout. Origin of fully erected second dorsal fin
on vertical through anterior third to half of anal fin in advance of second dorsal fin; both fins always well
scaled basally, and more or less heavily scaled on distal parts. Anal fin with III spines and 8 soft
rays (rarely 7 or 9) in adults (usually II spines and 9 soft rays in juveniles about 30 mm standard length
or less). Caudal fin square. Pectoral fins with I ‘spine’ and 15 to 18 (usually 15) soft rays; short, just
reaching origin of first dorsal fin; pectoral fins 19 to 26% standard length, 77 to 92% head length; greater
than length of head minus snout; pectoral axillary scale absent. Scales weakly ctenoid (cycloid in juveniles),
25 to 29 (commonly 26) in longitudinal series, 8 to 10 in transverse series; 7 to 9 scales in longitudinal
series anterior to tip of pectoral fins and 17 to 20 (usually 17 or 18) anterior to origin of second dorsal fin;
16 scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ with broad
sulcus; anterior valve a large fleshy lobe and posterior valve a small papilla. Pyloric caeca much
divided; up to 23 in about 2 bunches. Colour: olive-brown dorsally, flanks silvery, abdomen white or
suffused pale yellow; about 6 longitudinal stripes on flanks formed by longitudinal marks on scales;
scales also with darkened margins giving slight chequered appearance to flanks; iris with yellow patches;
margins of fins dusky, otherwise yellowish white; pelvic fins pale except for darker margins; caudal fin
distinctly yellow; pectoral fins completely black in small fish; lower section yellowish in adults.
Size: Maximum reported total length about 60 cm; commonly to 35 cm total length.
Habitat, biology, and fisheries: Common along shallow coastal areas and protected sandy shores in
lagoons, reeflats, estuaries, and coastal creeks. Usually within tidal influence, but may enter fresh water,
ascending up to about 10 km into rivers. Juveniles might be found in rice fields and mangroves. Forms shoals.
Spawning probably occurs at sea. Feeds on small algae, diatoms, benthic polychaets, molluscs, crustaceans,
as well as living and detrital organic matter; fry take copepods and floating algae. Caught in cast nets, stake
nets, beach seines, and gill nets. Marketed fresh and salted;
perhaps also boiled (Thailand), canned or frozen (Austra-
lia). Roe marketed salted. Juveniles used as bait fish.
Distribution: Throughout the Indo-Pacific from East Af-
rica to Tuamotu Islands; north to southern Japan, south
to southern Great Barrier Reef and New Caledonia.
Mugil broussonneti Valenciennes, 1836

Frequent synonyms / misidentifications: Mugil macrolepidotus of Richardson, 1846; ?Myxus
malayanus Herre, 1936 / ?Mugil cephalus Linnaeus, 1758; ?Valamugil speigleri (Bleeker, 1859).
FAO names: En - Broussonnet’s mullet.
Diagnostic characters: Apparently a small species, but may grow larger than reported. Head dorsally
broad and flat, but deeper than wide. Snout length equal to or less than eye diameter; snout more or less
blunt in profile. Dentary symphysis acute (pointed). Lips thin, lower lip directed forwards. Upper lip with
2 rows of small, close-set, unicuspid, setiform teeth; tips of teeth in outer row recurved. Lower lip
with 1 row of small, unicuspid, setiform teeth. Vomer edentate. Maxilla and serrate anteroventral
edge of preorbital straight. Preorbital posteroventrally slender with pointed tip. Distance between
anterior and posterior nostril equal to or greater than maximum diameter of posterior nostril. Adipose
eyefold extensive over iris (except in specimens about 30 mm standard length or less). Gill rakers
on lower limb of first gill arch 32 to 68. Origin of fully erected second dorsal fin on vertical through anterior
third of anal fin; both fins moderately or well scaled on all parts. Second dorsal fin with 9 or 10 soft
rays. Anal fin with III spines and 9 soft rays in adults (II spines and 10 soft rays in juveniles about
30 mm standard length or less). Caudal fin emarginate. Pectoral fins with I ‘spine’ and 16 soft rays, not
reaching origin of first dorsal fin; pectoral fins 18 to 21% standard length, 68 to 80% head length.
Moderately long pectoral axillary scale. Scales ctenoid, not covered by minute secondary, cycloid
scales. Scales in longitudinal series 37 to 39; about 12 or 13 in transverse series; 24 or 25 scales in
longitudinal series anterior to origin of second dorsal fin; 18 scales in transverse series entirely around
caudal peduncle. Pharyngobranchial organ with large denticulate area; broad, deep, sulcus; single, large
valve, often shorter than depth of its base. Pyloric caeca 2. Colour: dark dorsally, flanks silvery, whitish
ventrally.
Size: Maximum reported standard length 19.4 cm.
Habitat, biology, and fisheries: Apparently a rare species with little or no available data.
Distribution: Very difficult to determine due to paucity of records. Perhaps ranging from south China
through south Pacific. Possi-
bly from Celebes, but no con-
firmed reports from Indonesia.
Remarks: C o n s i d e r e d b y
some biologists as conspecific
with Mugil cephalus because
of the close morphological
similarity.
2096 Bony Fishes
Mugil cephalus Linnaeus, 1758

Frequent synonyms / misidentifications: Mugil cephalotus Valenciennes, 1836; M. chaptalii Eydoux
and Souleyet, 1841; M. dobula Günther, 1861; M. japonicus Temminck and Schlegel, 1845; M. oeur
Forsskål, 1775; M. öur Forsskål, 1775 / None.
FAO names: En - Flathead mullet; Fr - Mulet à grosse tête; Sp - Lisa pardete.
(from Senou, 1988)
Diagnostic characters: Medium to large species. Head dorsally broad and flat, but deeper than wide.
Snout length equal to or less than eye diameter; snout more or less blunt in profile. Dentary symphysis
90° or more acute. Lips thin, lower lip directed forwards. Upper lip with 1 or 2 outer rows of moderately
close-set, unicuspid teeth and 1 to 5 inner rows of smaller, bicuspid teeth; inner, bicuspid teeth
well spaced from outer, unicuspid teeth. Lower lip with outer row of moderately close-set unicuspid
teeth; 1 or more inner rows of smaller, bicuspid teeth present or absent. Vomer edentate. Maxilla
and serrate anteroventral edge of preorbital straight. Preorbital posteroventrally slender with
pointed tip. Distance between anterior and posterior nostril equal to or greater than maximum diameter
of posterior nostril. Adipose eyefold extensive over iris (except in specimens about 30 mm standard
length or less). Gill rakers on lower limb of first gill arch 50 to 90. Origin of fully erected second dorsal fin
just posterior to vertical level of origin of anal fin; both fins scaled only anterobasally. Second dorsal fin
with 9 soft rays. Anal fin with III spines and 8 (rarely 9) soft rays in adults (usually II spines and 9 soft
rays in juveniles about 30 mm standard length or less). Caudal fin forked. Pectoral fins with I ‘spine’ and
15 to 17 soft rays, not reaching origin of first dorsal fin; pectoral fins 16 to 21% standard length, 61 to 83%
head length. Long pectoral axillary scale. Scales very weakly ctenoid; minute cycloid scales super-
imposed on main body scales, particularly on occipital region. Longitudinal series scale count can
vary from 36 to 44 in Mugil cephalus, but usually in range 38 to 42 for specimens from the area; 13 to
15 (rarely 16) in transverse series; 25 or 26 in longitudinal series anterior to origin of second dorsal fin;
19 to 21 scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ with
large denticulate area; broad, deep, sulcus; single, large valve, often shorter than depth of its base. Pyloric
caeca 2. Colour: specimens caught from sea olive-green dorsally, flanks silvery and abdomen off-white;
about 7 longitudinal dark stripes along flanks, following rows of scales; these stripes less con-
spicuous ventrally; fish from estuarine waters have duller flanks and suffused deep blue or dirty brown
dorsally; fins dusky; dorsal fins perhaps greyish blue; anal and caudal fin may be yellowish green; pectoral
fins have whitish margin and distinct, dark purple spot at origin; pelvic fins pale yellow.
Size: Maximum reported total length 91 cm; commonly to 35 cm total length.
Habitat, biology, and fisheries: Inhabits inshore marine waters, estuaries, lagoons, and rivers; tolerant
of water temperatures from 12° to 25°C and salinities from hypersaline to fresh water. Adults roaming;
forming schools and sometimes jumping; but fish forage singly. First spawning in third year; mature fish
group in estuaries, form shoals, and move out to sea to spawn in surface waters. Spent fish return to
brackish and fresh waters of estuaries, rivers, or lakes. Newly hatched fry migrate to estuaries and swim
upstream. Juveniles found in sheltered lagoons and bays. Immature “hardgut” fish make preliminary
migrations out to sea but do not spawn, and return without feeding (hence “hardgut”). Adults feed on fine
particulate material, detritus, microscopic algae, and animals. Major commercial fishery; represents 1/2 to
the entire mullet landings in eastern Australia and Tonga. Caught from beaches and estuaries using gill
nets, cast nets, stake nets (barrier nets), lift nets, beach
seines, fish corrals. Also used in aquaculture. Marketed
fresh, salted, and frozen; roe sold fresh or smoked.
Distribution: Worldwide in tropical, subtropical, and
warm temperate waters, although less abundant in trop-
ics and apparently rare in Indonesia.
Myxus elongatus Günther, 1861

FAO names: En - Sand mullet.
Diagnostic characters: A small to medium-sized species; body moderately slender. Head relatively
flattened dorsally. Snout long, longer than eye diameter, pointed in profile. Dentary symphysis obtuse
(blunt) in large specimens (less so in small specimens). Lips thin. Upper lip with 1 row of spatulate teeth
with constricted tips. Lower lip with 1 row of fine ciliiform teeth at edge of lip; a few, scattered teeth
sometimes present behind these. Vomer toothed. Maxilla stocky, only slightly sigmoidally curved near
posterior tip. Preorbital large; serrate anteroventral edge weakly concave; posteroventral tip blunt
or squarish. Anterior and posterior nostrils close. Adipose eyefold absent. Gill rakers on first gill arch
as long as longest gill filaments. Origin of first dorsal fin approximately midway between tip of snout and
base of caudal fin, or nearer the latter. Origin of fully erected second dorsal fin on vertical through anterior
third of anal fin; both fins scaled only anterobasally. Anal fin with III spines and 9 soft rays in adults (II
spines and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin forked. Pectoral fins
with I ‘spine’ and 15 or 16 soft rays, short, not reaching origin of first dorsal fin; pectoral fins 17 to 19%
standard length, 67 to 71% head length; pectoral axillary scale absent. Scales cycloid except for
those on breast, which are ctenoid. Scales in longitudinal series 43 to 46; 13 ½ to 16 in transverse
series; 29 or 30 scales in longitudinal series anterior to origin of second dorsal fin; 19 or 20 scales
in transverse series entirely around caudal peduncle. Pharyngobranchial organ with large denticulate
area; wide sulcus; valve as ventrally placed, double papillose pad. Pyloric caeca 2. Colour: dark
olive-green or reddish brown dorsally, greenish wash on upper flanks, lower flanks silvery; gold blotch on
operculum; golden-yellow iris around dark eye; fins greenish brown, except anal fin with white margin;
black spot at base of pectoral fins.
Size: Maximum reported total length 40 cm, but this is unusual; certainly commonly to 20 cm total length,
and specimens up to 28 cm total length might not be rare.
Habitat, biology, and fisheries: Inhabits open coastline and bays. Common in brackish water, over
shallow-water sand bars, river mouths, ocean inlets, offshore island lagoons, and seagrass meadows;
preferring cleaner conditions than other estuarine mullets. Found at depths of 1 to 10 m, forming schools.
Perhaps spawning near estuary mouths or in sea. First year fish found in fresh water, but seldom thereafter.
Feeds during the day, sifting sand and ingesting small crustacea, molluscs, and microscopic algae. Fished
in shallow coastal waters and
estuaries; caught using gill
nets, seines, and on lines.
Marketed fresh or filleted, with
other rare mullets.
Distribution: Generally re-
stricted to temperate waters of
Australia, Lord Howe Island,
and Norfolk Island. Specimens
might be found at the southern
extreme of the area, in coastal
waters of southern Queens-
land.
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2098 Bony Fishes
Myxus petardi (Castelnau, 1875)

Frequent synonyms / misidentifications: Trachystoma petardi (Castelnau, 1875); Mugil breviceps
Steindachner, 1866 / None.
FAO names: En - Freshwater mullet.
Diagnostic characters: A medium to large species; moderately deep-bodied. Snout long, longer than
eye diameter, pointed in profile. Dentary symphysis more or less obtuse (blunt). Upper lip very slightly
thickened, lower lip thin. Lips with 1 or more rows of fine, ciliiform teeth, or teeth absent. Vomer
toothed. Maxilla stocky, only slightly sigmoidally curved near posterior tip. Preorbital large; serrate
anteroventral edge straight or only weakly concave; posteroventral tip broad and squarish. Distance
between anterior and posterior nostril more or less equal to maximum diameter of posterior nostril. Adipose
eyefold absent. Gill rakers on first gill arch 1/2 length of longest gill filaments. Origin of first dorsal
fin nearer base of caudal fin than tip of snout. Origin of fully erected second dorsal fin on vertical through
anterior half of anal fin; both fins scaled only on anterobasal parts. Anal fin with III spines and 9 (rarely 8)
soft rays in adults (usually II ‘spines’ and 10 soft rays in juveniles about 30 mm standard length or less).
Caudal fin forked. Pectoral fins with I spine and 14 or 15 soft rays, short, not reaching origin of first dorsal
fin; pectoral fins 16 to 20% standard length, 74 to 85% head length; pectoral axillary scale rudimentary
or absent. Ctenoid scales in longitudinal series 47 to 52; 15 or 16 in transverse series; 32 to 34
scales in longitudinal series anterior to origin of second dorsal fin; 22 scales in transverse series
entirely around caudal peduncle. Pharyngobranchial organ with large denticulate area; wide sulcus;
valve as ventrally placed, double papillose pad. Pyloric caeca 2. Colour: greenish black dorsally, lighter
on flanks and silvery white ventrally; eye yellow-orange to pink; upper edge of operculum golden; fins pale
yellow.
Size: Maximum reported total length 81 cm; commonly to 40 cm total length or less.
Habitat, biology, and fisheries: In fresh and brackish waters of streams and lagoons at 9° to 27°C; may
be found in sea after heavy floods. Active surface and midwater fish; small groups may be found in deep
pools where stream flow is slow. Mature after about 4 years and perhaps spawning in estuaries at low
salinities, during late summer and early autumn. Feeds on filamentous algae, microscopic plants and
animals, and detritus. Com-
mercially netted after heavy
rain, but not an important spe-
cies; caught by anglers. Flesh
edible but may have muddy
taste.
Distribution: East coast of
Australia, in New South Wales
and extending north to Burnett
River, southern Queensland.
Neomyxus leuciscus (Günther, 1871)

Frequent synonyms / misidentifications: Chaenomugil nauticus Bryon and Herre, 1903; C. leuciscus
(Günther, 1871); Neomyxus sclateri (Steindachner, 1878) / Neomyxus chaptalii Eydoux and Souleyet,
1841.
FAO names: En - Acute-jawed mullet.
(from Senou, 1988)
Diagnostic characters: A medium-sized species; body elongate. Head relatively flattened dorsally. Snout
long, longer than eye diameter, pointed in profile. Dentary symphysis acute (pointed). Both lips
thick, but upper lip not rugose nor bearing crenulations or papillae, and upper lip thickness at point
of snout usually 12 or more times in head length. Lips bearing 2 or 3 irregular rows of long teeth
which are slightly curved and tricuspid at their tips; lower lip with outer edge folded downwards.
Lower jaw fits snugly under upper jaw. Vomer edentate. Maxilla more or less straight. Serrate
anteroventral edge of preorbital straight; posteroventral tip broad and squarish. Distance between anterior
and posterior nostril more or less equal to maximum diameter of posterior nostril. Adipose eyefold marginal.
Gill rakers on lower limb of first gill arch 33 to 64, (usually 50 or more). Origin of first dorsal fin nearer base
of caudal fin than tip of snout. Origin of fully erected second dorsal fin on vertical through anterior half to
three-quarters of anal fin; both fins scaled only anterobasally. Anal fin with II spines and 10 or 11 soft
rays in adults (I spine and 11 or 12 soft rays in juveniles about 30 mm standard length or less). Caudal
fin emarginate. Pectoral fins with I spine and 15 or 16 soft rays, not reaching origin of first dorsal fin; pectoral
fins 20 to 23% standard length, 80 to 95% head length. Cycloid scales in longitudinal series 43 to 49;
13 or 14 in transverse series; 28 or 29 scales in longitudinal series anterior to origin of second dorsal fin;
21 or 22 scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ with
deep sulcus; denticulate cushion with long setiform teeth; anterior valve tripartite; small digitate fold
posterior to anterior valve. Pyloric caeca 2. Colour: grey dorsally, silvery flanks, white ventrally; pectoral
fins darkish but with bright yellow spot at origin.
Size: Maximum reported standard length 46 cm.
Habitat, biology, and fisheries: In shallow coastal waters (up to a depth of about 4 m), around reef flats,
tidepools, lagoons, drainage ditches, and docks. At night tending to move inshore to surface waters and
shallows close to beaches; at-
tracted to lights. Forms
schools. Can be caught at
night by dazzling with strong
light and scooping up with
hand nets.
Distribution: Central Pacific,
from southern Japanese and
Hawaiian islands in north;
south to Tubai and Ducie Is-
lands; rare in Marianas Is-
lands.
2100 Bony Fishes
Oedalechilus labiosus (Valenciennes, 1836)

Frequent synonyms / misidentifications: Crenimugil labiosus (Valenciennes, 1836); Plicomugil
labiosus (Valenciennes, 1836) / None.
FAO names: En - Hornlip mullet; Fr - Mulet labéon; Sp - Lisa morruda.
(from Senou, 1988)
Diagnostic characters: A small to medium-sized species; body moderately deep. Head relatively flattened
dorsally. Snout shorter than eye diameter and blunt in profile. Dentary symphysis very obtuse; mouth may
appear transverse in ventral view. Upper lip very thick, lip thickness at point of snout about 7 to 10
times in head length. Upper lip with deep longitudinal fold near its ventral edge, splitting it into
upper and lower lobes; these lobes fringed with ridges of horny epidermis; ridges small and appear
as papillae on upper lobe (but different from papillae of Crenimugil); ridges on lower lobe better
developed. Lower lip thin, directed forwards, also fringed with ridges of horny epidermis except at
medial part. Both lips form folds tucked under preorbital at corners of mouth. Lips and vomer
edentate. Maxilla curved down posteriorly. Preorbital deeply notched midway along anteroventral
edge and expanded into a broad, squarish, posteroventral tip. Distance between anterior and posterior
nostril less than maximum diameter of posterior nostril. Adipose eyefold absent. Gill rakers on lower limb
of first gill arch 30 to 41, shorter than gill filaments. Origin of first dorsal fin slightly nearer base of caudal
fin than tip of snout. Origin of fully erected second dorsal fin on vertical through anterior half to
three-quarters of anal fin; both fins usually well scaled on all parts. Anal fin with III spines and 9 soft rays
in adults (II spines and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin
emarginate. Pectoral fins with I ‘spine’ and 15 to 18 soft rays, ventral 1 or 2 soft rays more or less free
from fin membrane; pectoral fins not reaching origin of first dorsal fin; pectoral fins 23 to 28%
standard length, 94 to 112% head length. Scales cycloid or weakly ctenoid; 32 to 37 scales in longitudinal
series; 11 or 12 in transverse series; 23 to 25 scales in longitudinal series anterior to origin of second
dorsal fin; 16 scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ
with broad sulcus; small anterior valve and longer-based posterior valve. Pyloric caeca 3 to 4. Colour:
olive dorsally, silvery ventrally.
Size: Maximum standard length 40 cm or more; commonly to 20 cm total length.
ies: Found in shallow coastal
waters, around coral reefs,
and in harbours. Caught with
gill nets, lift nets, and seines;
also used as live bait in pole-
and-line tuna fishing.
throughout tropical Indo-Pa-
cific, from the Red Sea and
Madagascar to Samoa; north
to southern Japan and south to
the Great Barrier Reef.
Rhinomugil nasutus (DeVis, 1883)

Frequent synonyms / misidentifications: Squalomugil nasutus (DeVis, 1883) / None.
FAO names: En - Shark mullet.
Diagnostic characters: A medium-sized species; body elongate. Head noticeably wider than deep,
flattened dorsally and concave between eyes; eyes positioned dorsolaterally high on head. Eye
diameter 14 to 20% head length. Snout short, less than or equal to eye diameter, but projecting
beyond upper lip. Dentary symphysis about 90° or only slightly obtuse (blunt). Lips thin; lower lip directed
forwards. Teeth small, fine and spatulate on both lips, often not easily discernable. Vomer edentate.
Maxilla straight. Serrate anteroventral edge of preorbital very weakly concave; posteroventral tip narrow.
Nostrils set low on snout; distance between anterior and posterior nostril greater than maximum diameter
of posterior nostril. Adipose eyefold developed posteroventral to eye and not overlying iris. Gill rakers
on lower limb of first gill arch 45 to 55 (perhaps more). Origin of first dorsal fin nearer base of caudal fin
than tip of snout. Origin of fully erected second dorsal fin on vertical through posterior half of anal
fin, or behind anal fin; both fins scaled basally but not distally. Anal fin with III spines and 8 soft rays
in adults (II spines and 9 soft rays in juveniles about 30 mm standard length or less). Caudal fin truncate.
Pectoral fins with I ‘spine’ and 13 soft rays, long, just reaching origin of first dorsal fin; pectoral fins 25 to
26% standard length, 95 to 100% head length. Ctenoid scales in longitudinal series 28 to 30; 9 ½ or
10 in transverse series; 21 or 22 scales in longitudinal series anterior to origin of second dorsal fin; 16
scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ with broad sulcus,
small valve ventrally and finger like processes in sulcal groove. Pyloric caeca 2. Colour: dark slaty-grey
dorsally, silvery on flanks and abdomen; fins yellowish.
Size: Maximum fork length at least 32 cm.
Habitat, biology, and fisheries: Usually found in small schools in muddy fresh waters and coastal waters
(e.g. mangroves). Swimming at the surface with eyes and snout exposed. Perhaps feeding on surface
algae and insects which alight on surface; also feeding on muddy banks and bottoms. Capable of breathing
air and wriggling over mudbanks for short distances. No reported fisheries data.
Distribution: Tropical Austra-
lia and New Guinea.
2102 Bony Fishes
Valamugil buchanani (Bleeker, 1853)

Frequent synonyms / misidentifications: Mugil ceylonensis Günther, 1861 / Valamugil cunnesius
(Valenciennes, 1836); V. engeli (Bleeker, 1859); V. seheli (Forsskål, 1775).
FAO names: En - Bluetail mullet; Fr - Mulet à queue bleue; Sp - Lisa rabo azul.
(from Senou, 1988)
Diagnostic characters: A medium-sized species; body moderately robust. Head dorsally flattened. Snout
short and blunt, 14 to 18% head length; shorter than eye diameter. Dentary symphysis obtuse (blunt).
Lips thin, lower directed forwards. Teeth on both lips minute and ciliiform in 1 row or absent. Vomer
edentate. Maxilla slender and weakly curved down at posterior tip, which is only just posteroventral
to corner of mouth and partially or completely concealed. Serrate anteroventral edge of preorbital
very weakly concave, not kinked; posteroventral tip narrow but not pointed. Distance between
anterior and posterior nostril less than or equal to maximum diameter of posterior nostril. Adipose eyefold
poorly developed as rim around eye. Gill rakers on lower limb of first gill arch 37 to 67. Relative position
of origin of first dorsal fin may vary during ontogeny, but is usually midway between tip of snout and base
of caudal fin, or slightly nearer latter. Origin of fully erected second dorsal fin on vertical through
anal-fin origin, or only just posterior to it; both fins always well scaled basally and medially, and
more or less scaled on distal parts; both fins falcate (more noticeable in specimens over 12 cm
standard length). Anal fin with III spines and 9 soft rays in adults (usually II spines and 10 soft rays in
juveniles about 30 mm standard length or less). Caudal fin deeply forked. Pectoral fins with I ‘spine’ and
17 to 19 (rarely 16) soft rays; long and falcate, reaching level of origin of first dorsal fin; pectoral fins
22 to 27% standard length, 90 to 139% head length; greater than length of head minus snout, and
often greater than head length; pectoral axillary scale moderately long. Scales with a membranous,
digitated hind margin; 32 to 36 (rarely 37) in longitudinal series; 11 to 13 in transverse series; 10 to
14 scales in longitudinal series anterior to tip of pectoral fins and 19 to 24 scales anterior to origin of
second dorsal fin; 19 to 20 (rarely 18) in transverse series entirely around caudal peduncle.
Pharyngobranchial organ with large denticulate area; broad sulcus; no valves. Pyloric caeca 6 or 7.
Colour: greenish dorsally; flanks and abdomen silvery; small gold patch on upper operculum and on iris;
caudal fin distinctive bright blue (dusky when preserved); dorsal fins dusky; anal fin dusky at margins;
pectoral fins yellow with dark blue spot dorsally at origin.
Size: Usually reported to a maximum standard length of 50 cm (100 cm reported for South African
specimens); commonly to 20 to 30 cm standard length.
Habitat, biology, and fisheries: Inhabits shallow coastal waters. Young enter estuaries, lagoons, and
backwaters. Feeds on algae, diatoms, de-
tritus, and crustacea. Caught in coastal wa-
ters using gill nets, cast nets, stake nets,
barrier nets, lift nets, beach seines, and
pouch nets during the spawning run. Mar-
keted fresh.
Distribution: Indo-Pacific from South Af-
rica through parts of Indonesia to parts of
Melanesia and Micronesia; north to the
Marianas Islands and southern Japan.
Valamugil cunnesius (Valenciennes, 1836)

Frequent synonyms / misidentifications: Mugil ophuysenii Bleeker, 1859 / See Remarks.
FAO names: En - Longarm mullet (= Longfin mullet, Fishing Area 57/71); Fr - Mulet longue aile; Sp - Lisa
de aleta larga.
(from Senou, 1988)
Diagnostic characters: A small species; body somewhat fusiform. Head deeper than wide and dorsally
flattened. Snout moderately long and more or less pointed in profile, 20 to 25% head length. Dentary
symphysis obtuse (blunt). Lips thin, lower directed forwards. Teeth minute and ciliiform on both lips.
Vomer edentate. Maxilla slender and weakly curved down at posterior tip, which is only just
posteroventral to corner of mouth and partially or completely concealed. Serrate anteroventral edge
of preorbital very weakly concave, not kinked; posteroventral tip narrow but not pointed. Distance
eyefold poorly developed, usually as rim around eye. Gill rakers on lower limb of first gill arch 62
to 75. Relative position of origin of first dorsal fin variable, usually midway between tip of snout and base
of caudal fin, or slightly nearer former. Origin of fully erected second dorsal fin on vertical through
anterior quarter to half of anal fin; both fins scaled only on anterior and basal parts. Anal fin with III
spines and 9 soft rays in adults (usually II spines and 10 soft rays in juveniles about 30 mm standard length
or less). Caudal fin forked. Pectoral fins with I ‘spine’ and 14 to 16 soft rays; reaching level of origin of first
dorsal fin; pectoral fins 23 to 25% standard length, 86 to 97% head length; greater than length of head
minus snout; pectoral axillary scale moderately long. Scales with a membranous, digitated hind margin;
37 to 43 (usually 38 or 39) in longitudinal series, 11 or 12 in transverse series; 12 or 13 scales in
longitudinal series anterior to tip of pectoral fins and 23 to 26 anterior to origin of second dorsal fin; 17 or
18 (rarely 15 or 16) scales in transverse series entirely around caudal peduncle. Pharyngobranchial
organ with large denticulate area; broad sulcus; no valves. Pyloric caeca 5 to 7. Colour: greenish grey
dorsally; flanks and abdomen silvery; second dorsal and anal fins with black margins.
Size: Largest observed specimen 14 cm standard length, but probably grows larger.
Habitat, biology, and fisheries: Very little reliable data due to earlier taxonomic confusion for this species
(see below). Probably similar to other Valamugil species, inhabiting shallow coastal waters and browsing
on benthic substrates.
Distribution: Reliable reports
are very rare due to earlier
taxonomic confusion. Per-
haps widespread from Red
Sea to West Pacific.
Remarks: The type-series
specimens of V. cunnesius are
non-conspecific and the puta-
tive holotype has frequently
been overlooked. This has re-
sulted in frequent, earlier,
taxonomic confusion of other
species with V. cunnesius.
2104 Bony Fishes
Valamugil engeli (Bleeker, 1859)

Frequent synonyms / misidentifications: Mugil kandavensis Günther, 1877 / Valamugil cunnesius
(Valenciennes, 1836); V. perusii (Valenciennes, 1836).
FAO names: En - Kanda; Fr - Mulet oeil de perdrix; Sp - Lisa ojo de perdiz.
(from Senou, 1988)
Diagnostic characters: A small to medium-sized species; body more or less fusiform; depth at origin of
first dorsal fin 25 to 29% standard length. Head deeper than wide and dorsally flattened. Snout short and
blunt, 17 to 21% head length; shorter than eye diameter. Dentary symphysis about 90° or slightly obtuse.
Lips thin, lower directed forwards. Teeth on both lips minute and ciliiform or absent. Vomer toothed or
edentate. Maxilla slender and weakly curved down at posterior tip which is only just posteroventral
to corner of mouth and partially or completely concealed. Serrate anteroventral edge of preorbital
very weakly concave, not kinked; posteroventral tip narrow but not pointed. Distance between
anterior and posterior nostril more or less equal to maximum diameter of posterior nostril. Adipose eyefold
weakly developed, only slightly overlying iris. Gill rakers on lower limb of first gill arch 44 to 52. Origin
of first dorsal fin nearer base of caudal fin than tip of snout. Origin of fully erected second dorsal
fin on vertical through about third soft ray of anal fin, i.e. behind anterior quarter or more of anal
fin; anal fin scaled basally, second dorsal fin only scaled anterobasally. Anal fin with III spines and
9 (rarely 8) soft rays in adults (usually II spines and 10 soft rays in juveniles about 30 mm standard length
or less). Caudal fin emarginate. Pectoral fins with I ‘spine’ and 14 to 17 (commonly 14 or 15) soft rays;
moderately long, just reaching level of origin of first dorsal fin or extending beyond this; pectoral fins
23 to 26% standard length, 80 to 95% head length; greater than length of head minus snout; pectoral
axillary scale long. Scales with a membranous, digitated hind margin; thoracic and abdominal scales
more distinctly ctenoid; 32 to 36 in longitudinal series; 10 ½ to 12 ½ in transverse series; 10 to 12
scales in longitudinal series anterior to tip of pectoral fins and 19 to 23 (commonly 21 or 22) anterior to
origin of second dorsal fin; 16 scales in transverse series entirely around caudal peduncle; predorsal
scales to tip of snout. Pharyngobranchial organ with large denticulate area; broad sulcus; no valves.
Pyloric caeca 5 or 6. Colour: olive dorsally; flanks and abdomen silvery; fins hyaline; pectoral fins with
dark spot dorsally at origin.
Size: Maximum reported total length 30 cm; commonly to about 15 cm total length.
Habitat, biology, and fisheries: Inhabits coastal waters, shallow lagoons, protected inlets, and over sandy
to muddy areas of reef flats.
Juveniles may enter rivers
and have been found in tide-
pools at 40°C. Caught with
seines and lift nets. Marketed
fresh; also used as live bait in
pole-and-line fishing for tuna.
Distribution: Widespread
across Indo-Pacific, from Af-
rica to Marquesas and
Tuamotu Islands; north to
southern Japan; introduced to
Hawaii.
Valamugil georgii (Ogilby, 1897)

Frequent synonyms / misidentifications: Mugil nortoni Ogilby, 1908 / ?Valamugil cunnesius
(Valenciennes, 1836); ?V. engeli (Bleeker, 1859); ?V. perusii (Valenciennes, 1836).
FAO names: En - Fantail mullet.
Diagnostic characters: A small to medium-sized species; body moderately robust; depth at origin of first
dorsal fin about 30% standard length (ranges include 25 to 39%). Snout short and blunt, 19 to 24% head
length; shorter than eye diameter. Upper lip perhaps slightly thickened, with fine, ciliiform teeth; lower
lip thin, directed forwards, without teeth. Vomer edentate. Maxilla slender and very weakly curved down
at posterior tip, which is only just posteroventral to corner of mouth and partially or completely
concealed. Serrate anteroventral edge of preorbital more or less straight; posteroventral part of
preorbital broadened into a squarish end. Anterior and posterior nostrils close-set. Adipose eyefold
usually well developed, covering most of iris posteriorly and partly anteriorly. Gill rakers on lower
limb of first gill arch 55 or 56. Origin of first dorsal fin nearer base of caudal fin than tip of snout. Origin of
fully erected second dorsal fin on vertical through anterior third to half of anal fin; both fins scaled
only on anterior or basal parts. Anal fin with III spines and 9 soft rays in adults (usually II spines and 10
soft rays in juveniles about 30 mm standard length or less). Caudal fin emarginate. Pectoral fins with I
‘spine’ and 14 or 15 soft rays; relatively short, tip distinctly anterior to level of origin of first dorsal fin;
pectoral fins about 21% standard length, 92% head length; greater than length of head minus snout;
pectoral axillary scale long. Scales appear cycloid or only very weakly ctenoid (membranous,
digitated hind margin of scale not distinct; thoracic and abdominal scales more distinctly ctenoid;
31 or 32 (rarely 30) in longitudinal series, (11 ½) 12 or 13 in transverse series; (9) 10 or 11 scales in
longitudinal series anterior to tip of pectoral fins and 23 anterior to origin of second dorsal fin; about 16
scales in transverse series entirely around caudal peduncle. Pharyngobranchial organ with large
denticulate area; broad sulcus; no distinct valves but small valve-like swelling or fold of tissue at
anteroventral part of sulcus. Pyloric caeca extensively branched, about 22 in total. Colour: greenish
brown dorsally; flanks silvery; abdomen off-white; gold spots on dorsal and ventral parts of iris; second
dorsal, anal, and caudal fins with dusky margins and second dorsal speckled brown; pectoral fins with
purplish spot dorsally at origin.
Size: Maximum reported total length 30 cm; rarely above 25 cm total length.
Habitat, biology, and fisheries: Inhabits coastal waters, estuaries and bays; young may enter fresh water.
Forms schools, shoaling over shallow-water sand flats, particularly on rising tide at dusk. Fish tend to roll
on their sides and flick a col-
umn of water into air. Taken
as a bycatch; used as baitfish
for crabs and snapper.
Distribution: Tropical Austra-
lia, along coasts of Queens-
land and New South Wales.
Remarks: Very similar to V.
engeli and V. perusii, with
which it has previously been
synonymized.
2106 Bony Fishes
Valamugil perusii (Valenciennes, 1836)

Frequent synonyms / misidentifications: ?Mugil amarulus Valenciennes, 1836; M. kelaartii Günther,
1861; M. longimanus Günther, 1861; M. strongylocephalus Richardson, 1846 / Valamugil cunnesius
(Valenciennes, 1836); V. engeli (Bleeker, 1859).
FAO names: En - Longfinned mullet (from Mugil strongylocephalus).
(from Senou, 1988)
Diagnostic characters: A small species; body moderately robust; depth at origin of first dorsal fin 24 to
35% (often less than 30%) standard length. Head deeper than wide and dorsally flattened. Snout short
and blunt, 17 to 21% head length; shorter than eye diameter. Dentary symphysis about 90° or slightly
obtuse. Lips thin, lower directed forwards. Teeth on both lips minute and ciliiform or absent. Vomer
usually edentate. Maxilla slender and weakly curved down at posterior tip, which is only just
posteroventral to corner of mouth and partially or completely concealed. Serrate anteroventral edge
of preorbital very weakly concave, not kinked; posteroventral tip narrow but not pointed. Distance
eyefold reasonably developed, covering most of iris posteriorly and partly anteriorly. Gill rakers on
lower limb of first gill arch 40 to 51. Origin of first dorsal fin midway between tip of snout and base of
caudal fin, or slightly nearer latter. Origin of fully erected second dorsal fin on vertical through
about third soft ray of anal fin, i.e. behind anterior quarter or more of anal fin; both fins moderately
to well scaled on all parts. Anal fin with III spines and 9 (rarely 8 or 10) soft rays in adults (usually II
spines and 10 soft rays in juveniles about 30 mm standard length or less). Caudal fin emarginate. Pectoral
fins with I ‘spine’ and 14 or 15 (rarely 16) soft rays; long, just reaching level of origin of first dorsal fin
or extending beyond this; pectoral fins 21 to 27% standard length, 90 to 110% head length; greater than
length of head minus snout; pectoral axillary scale long. Scales with a membranous, digitated hind
margin; thoracic and abdominal scales more distinctly ctenoid; 31 to 34 (rarely 35) in longitudinal
series, 10 ½ to 11 ½ in transverse series; 11 to 14 (rarely 10) scales in longitudinal series anterior to tip
of pectoral fins and 19 to 22 anterior to origin of second dorsal fin; 16 scales in transverse series entirely
around caudal peduncle; predorsal scales to level of posterior nostril. Pharyngobranchial organ with
large denticulate area; broad sulcus; no valves. Pyloric caeca 5 to 7. Colour: greenish dorsally; flanks
and abdomen silvery; gold opercular spot; fins dusky, particularly at margins; pectoral fins with dark spot
dorsally at origin.
Size: Maximum reported total
length 25 cm; perhaps com-
monly to 15 cm total length.
ies: Inhabits coastal waters
and estuaries. Shoaling over
mudflats. Netted for bait.
cific from Africa to Marianas
Islands.
Valamugil seheli (Forsskål, 1775)

Frequent synonyms / misidentifications: Mugil axillaris Valenciennes, 1836; M. borbonicus Cantor,
1850; M. caeruleomaculatus Lacepède, 1803; M. cylindricus Valenciennes, 1836 / Valamugil buchanani
(Bleeker, 1853); V. cunnesius (Valenciennes, 1836); V. speigleri (Bleeker, 1859).
FAO names: En - Bluespot mullet; Fr - Mulet à tache bleue; Sp - Lisa de mancha azul.
(from Senou, 1988)
Diagnostic characters: A medium-sized species; body moderately robust. Head as wide as deep or slightly
wider and dorsally flattened. Snout short and blunt, 18 to 22% (rarely 17%) head length; shorter than eye
diameter. Dentary symphysis obtuse (blunt). Upper lip thin or slightly thickened; lower lip thin and directed
forwards. Teeth on both lips minute and ciliiform in 1 row or absent. Vomerine teeth present or absent.
Maxilla slender and weakly curved down at posterior tip, which is only just posteroventral to corner
of mouth and partially or completely concealed. Serrate anteroventral edge of preorbital very weakly
concave, not kinked; posteroventral tip narrow but not pointed. Distance between anterior and posterior
nostril less than or equal to maximum diameter of posterior nostril. Adipose eyefold poorly developed as rim
around eye. Gill rakers on lower limb of first gill arch 39 to 73, usually more than 50. Relative position of origin
of first dorsal fin may vary during ontogeny, but is usually midway between tip of snout and base of caudal
fin, or slightly nearer snout. Origin of fully erected second dorsal fin on vertical through anal-fin origin,
or only just posterior to it; both fins always well scaled basally and medially, and more or less scaled
on distal parts; both fins mildly falcate but not as strongly as pectoral fins or fins of Valamugil buchanani.
Anal fin with III spines and 9 (rarely 8 or 10) soft rays in adults (usually II spines and 10 soft rays in juveniles
about 30 mm standard length or less). Caudal fin forked. Pectoral fins with I ‘spine’ and 16 to 18 (rarely 15 or
19) soft rays; long and falcate, usually reaching level of origin of first dorsal fin; pectoral fins 18 to 25%
standard length, 84 to 104% head length (perhaps more, although this would be uncommon); often greater
than length of head minus snout, not usually greater than length of head; pectoral axillary scale moderately
long. Scales with a membranous, digitated hind margin; 36 to 42 (usually 38 to 40) in longitudinalseries;
12 to 14 in transverse series; 10 to 14 scales in longitudinal series anterior to tip of pectoral fins and 23 to
26 (usually 24 or 25) anterior to origin of second dorsal fin; 18 to (commonly) 20 scales in transverse
series entirely around caudal peduncle. Pharyngobranchial organ with large denticulate area; broad
sulcus; no valves. Pyloric caeca 6 to 9. Colour: bluish brown or green dorsally; flanks and abdomen silvery;
dusky spots on upper rows of scales, giving indistinct longitudinal stripes; dorsal and caudal fins bluish, also
dusky in second dorsal and caudal; anal, pelvic, and particularly pectoral fins yellowish; pectoral fins
also with dark blue spot dorsally at origin.
Size: Maximum reported standard length 50 cm; commonly between 20 and 30 cm standard length.
Habitat, biology, and fisheries: Inhabits shallow coastal waters; enters lagoons, estuaries, brackish tidal
creeks and fresh-water parts of rivers, where it feeds. Juveniles might be found in rice fields and mangrove
swamps. Forms schools; larger shoals occur during spawning, which takes place at sea. Feeds on
microalgae, filamentous algae, diatoms, foraminifera, and detritus associated with sand and mud. Caught
using gill nets, cast nets, stake nets, barrier nets, lift nets, beach seines, and pouch nets during the
spawning run; also taken as a bycatch with
other fishes. Marketed fresh and perhaps
salted, boiled (Thailand), canned or frozen
(Australia). Roe marketed salted.
Distribution: Widespread throughout
Indo-Pacific, from East Africa and Red Sea
to the Marquesas Islands; north to Japan and
Hawaii, and south to southern Queensland
and New Caledonia.
2108 Bony Fishes
Valamugil speigleri (Bleeker, 1859)

Frequent synonyms / misidentifications: Mugil suppositus of Day, 1865 / Valamugil cunnesius
(Valenciennes, 1836).
FAO names: En - Speigler’s mullet; Fr - Mulet chiraya; Sp - Lisa chiraya.
Diagnostic characters: A small to medium-sized species; body fusiform. Head deeper than wide and
dorsally flattened. Snout short and blunt, 17 to 22% head length. Dentary symphysis more or less obtuse
(blunt). Lips thin, lower directed forwards. Teeth on upper lip minute and ciliiform; those on lower lip
also ciliiform but slightly larger and more numerous. Vomer edentate. Maxilla slender and more or
less straight; posterior tip only just posteroventral to corner of mouth and partially or completely
concealed. Serrate anteroventral edge of preorbital straight; posteroventral tip narrow but not
pointed. Distance between anterior and posterior nostril more or less equal to maximum diameter of
posterior nostril. Adipose eyefold varying from a narrow rim around eye to a well-developed structure
covering iris posteriorly and partly anteriorly. Gill rakers on lower limb of first gill arch 35 to 45.
Origin of first dorsal fin usually nearer tip of snout than base of caudal fin. Origin of fully erected second
dorsal fin on vertical through anterior third to half of anal fin; both fins well scaled on all parts. Anal
fin with III spines and 9 soft rays in adults (usually II spines and 10 soft rays in juveniles about 30 mm
standard length or less). Caudal fin forked. Pectoral fins with I ‘spine’ and 15 or 16 (rarely 17) soft rays;
reaching level of origin of first dorsal fin; pectoral fins 20 to 24% standard length, 82 to 100% head length;
greater than length of head minus snout; pectoral axillary scale long. Scales with a membranous,
digitated hind margin; 37 to 41 in longitudinal series; 11 or 12 in transverse series; 12 or 13 scales
in longitudinal series anterior to tip of pectoral fins and 23 to 26 (usually 24 or 25) anterior to origin of
second dorsal fin; 16 (less commonly 17) scales in transverse series entirely around caudal peduncle.
Pharyngobranchial organ with large denticulate area; broad sulcus; no valves. Pyloric caeca 4. Colour:
greenish dorsally; flanks and abdomen silvery; first dorsal fin with black margin, other fins dusky; pectoral
fins with dark spot dorsally at origin.
Size: Maximum reported total length 35 cm; commonly between 15 and 20 cm total length.
Habitat, biology, and fisheries: Schools in shallow coastal waters, estuaries, and backwaters; found in
shallow, inundated areas along coast after heavy rains; enters fresh water. Spawns in sea. Juveniles might
be found in rice fields and mangrove swamps. Feeds on small algae, diatoms, and other organic matter
taken in with sand and mud; fry feed on copepods and floating algae. Caught using gill nets, cast nets,
stake nets, barrier nets, lift
nets, beach seines, pouch
nets, and trawls. Marketed
fresh and perhaps salted,
boiled (Thailand), canned or
frozen (Australia). Roe mar-
keted salted.
cific, from Baluchistan to Bor-
neo and New Guinea; north up
to the Chinese coast, perhaps
not reaching south to Australia.
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Atheriniformes: Pseudomugilidae 2109
Order ATHERINIFORMES
PSEUDOMUGILIDAE
Blue eyes
by W. Ivantsoff
D iagnostic characters: Body small, elongate, moderately compressed. Mouth terminal, oblique.
Maxilla extending posteriorly to about vertical through anterior border of orbit or slightly beyond. Upper
jaw strongly curved and slightly protrusible. Villiform teeth in upper jaw; teeth on lower jaw small, curved,
and either restricted to first third or half, or present on all of dentary toothed surface. Gill rakers small to
very small and widely spaced. Two separate dorsal fins; spines usually absent in second dorsal fin and
pectoral fin. Pectoral fins always set high on body above midlateral line. Body scales cycloid,
dorsoventrally elongated, with 22 to 34 (usually less than 31) scales in midlateral line. Scales on head and
cheeks varying from few large to many small scales. Species of this family often exhibiting sexual
dimorphism with males characterized by extremely elongated fin rays and different coloration.
Habitat, biology, and fisheries: A mostly fresh-water family, predominantly restricted to coastal regions
of Australia and Papua New Guinea. Three species are considered to be marine or estuarine and are found
in coastal regions.
Similar families occurring the area

Pseudomugilids are distinct from other atheriniform families in the area by the absence of the mesethmoid
and by having the single infraorbital (lacrimal) articular as high as the dentary (always lower than dentary
in other atherniform fishes).
Juveniles of the family Mugilidae (mullets) could be mistaken for pseudomugilids.
Key to the estuarine species of Pseudomugilidae
1a. Gill rakers on lower limb of first gill arch 14 . . . . . . . . . . . . . . . . . Pseudomugil majusculus
1b. Gill rakers on lower limb of first gill arch 12 or less . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Premaxilla without postmaxillary process, metapterygoid fused to symplectic

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudomugil cyanodorsalis
2b. Premaxilla with pungent postmaxillary process; metapterygoid not fused to symplectic
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudomugil inconspicuus
The symbol 0
is given when species accounts are included.
0
0
Pseudomugil cyanodorsalis Allen and Sarti, 1983
0
Pseudomugil inconspicuus Roberts, 1978
Pseudomugil majusculus Ivantsoff and Allen, 1984
Reference
Saeed, B. and W. Ivantsoff. 1989. Systematic revision of the blue-eyes (family Pseudomugilidae). Aust. J. Mar.
Freshwater Res., 40:719-87.
2110 Bony Fishes
Pseudomugil cyanodorsalis Allen and Sarti, 1983

Frequent synonyms / misidentifications: None / Pseudomugil inconspicuus Roberts, 1978.
FAO names: En - Blueback blue-eye.
Diagnostic characters: Body very small, elongate, moderately compressed. Mouth small, subvertical.
Teeth in jaws small anteriorly, enlarged, pungent on posterior part of premaxilla and exposed when mouth
closed. Premaxilla wide, its ascending process short, lateral process absent. Vomer wide and
slightly curved. Mesethmoid absent. Gill rakers on lower limb of first gill arch 7 to 10. First dorsal fin with
III to V weak spines, its origin at or near vertical through tips of pelvic fins. Second dorsal fin without
spines and 4 to 7 soft rays. Anal fin with I spine and 9 to 13 soft rays. Pectoral fins without spines and
9 to 11 soft rays. Body scales large; 22 to 26 midlateral scales; 5 scales in transverse rows along side
of body. Predorsal scales 10 to 14; interdorsal scales 3 or 4. Ventromedian scales between cleithrum and
origin of pelvic fins 7 to 10 (usually about 8). Colour: semitransparent dorsal half of males metallic blue,
lower half yellowish; first dorsal fin translucent, its posterior border blackish with small yellowish patch in
basal part near last spine; elongated rays and base of second dorsal fin black, remainder of fin creamy;
pectorals translucent or yellowish; body of females semitransparent, with translucent fins and white
abdomen.
Size: Maximum length 3.5 cm.
Habitat, biology, and fisheries: Inhabits mangrove swamps and highly saline creeks flowing into
estuaries, usually swimming over muddy bottom. Often congregate in large numbers. May be found in
hypersaline waters (range 28 to 40‰) and in relatively high temperatures (22 to 31°C). This species is
small but attractive and may have appeal to aquarium trade.
Distribution: Northwestern
Australia near Broome and
between Darwin and Wynd-
ham in the Northern Territory,
Australia.
Atheriniformes: Pseudomugilidae 2111
Pseudomugil inconspicuus Roberts, 1978

Frequent synonyms / misidentifications: None / Pseudomugil cyanodorsalis Allen and Sarti, 1983.
FAO names: En - Inconspicuous blue-eye.
Diagnostic characters: Body elongate, moderately compressed. Mouth fairly narrow with partially
restricted gape. Posterior premaxillary teeth enlarged and exposed when mouth closed; posterior two-
thirds of dentary toothless. Premaxilla with pungent narrow ascending process; postmaxillary process
also long, 1/2 length of ascending process, and pungent. Vomer narrow and slightly curved. Meseth-
moid absent. Gill rakers on lower limb of first gill arch 10 to 12. First dorsal fin with III or IV spines, its origin
behind vertical through tips of pelvic fins. Second dorsal fin without spine and with 5 soft rays. Anal fin
with I spine and 10 to 12 soft rays. Pectoral fins with 11 or 12 soft rays. Midlateral scales 26 to 28; 5 scales
in transverse rows along side of body. Predorsal scales 14 to 17; interdorsal scales 4 to 7. Ventromedian
scales between cleithrum and origin of pelvic fins 6 to 8. Colour: semitransparent in life; both males and
females yellowish with some scattered melanophores on body; dark midlateral line along side of body; fins
clear their bases dark; sexual dimorphism not apparent in this species.
Size: Maximum length not exceeding 3.5 cm.
Habitat, biology, and fisheries: Found in mangrove swamps and saline creeks. A planktivorous species
which adapts well to fresh-water environment. May occur in large schools, frequently in muddy waters.
Sympatric with Pseudomugil cyanodorsalis over part of its range. Of no commercial value.
Distribution: Known from
several sites but probably ex-
tends from Toro Pass near the
mouth of the Fly River (Papua
New Guinea) to Vogelkop
Peninsula (Irian Jaya) and
from Woods Inlet in Darwin
Harbour (Australia).
2112 Bony Fishes
Pseudomugil majusculus Ivantsoff and Allen, 1984

Frequent synonyms / misidentifications: None / Pseudomugil signifer (Kner, 1866).
FAO names: En - Cape blue-eye.
Diagnostic characters: Body elongate, moderately compressed. Mouth subvertical and small, gape
unrestricted by labial ligament. Posterior premaxillary teeth enlarged and exposed when mouth closed;
teeth on lower jaw strong, though small, villiform, pointing backwards and extending at least half-way along
dentary. Premaxilla broad, with short ascending process and no postmaxillary process. Gill rakers on
lower limb of first gill arch 14 or 15, less than 1/2 diameter of pupil but longer than in other species
of this genus. First dorsal fin with IV or V spines, its origin about 2 scales in front of vertical through tips
of pelvic fins. Second dorsal fin without spine and 7 to 9 soft rays. Anal fin without spine and 12 or 13 soft
rays. Pectoral fins without spine and about 10 to 12 soft rays. Body scales dorsoventrally elongated; 26 to
28 midlateral scales; 5 or 6 scales in transverse rows along side of body. Predorsal scales 12; interdorsal
scales 4 or 5. Ventromedian scales between cleithrum and origin of pelvic fins 9 or 10. Colour: preserved
specimen yellow-green with upper part of head and eyes dark; sides of snout and opercular bones with
scattered melanophores; edges of caudal-fin rays pigmented with scattered melanophores; no reticulate
pattern apparent on upper body scales above thin midlateral band.
Size: Maximum length about 5 cm.
Habitat, biology, and fisheries: Found in mangrove swamps and may provide food for young of
commercially important species. Nothing is known of the biology of this species of which only 2 specimens
are presently known.
C a pe W ar d H un t ( n or t h-
eastern Papua New Guinea)
and from Fergusson Island in
the D’Entrecasteaux Group.
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Atheriniformes: Atherinidae 2113
ATHERINIDAE
Silversides (or hardyheads)
by W. Ivantsoff and L.E.L.M. Crowley
D iagnostic characters: Elongate and somewhat compressed silvery fishes (rarely exceeding 12 cm
total length). Mouth generally small, oblique, and terminally placed; premaxilla with ascending
process of variable length, with lateral process present or absent; ramus of dentary bone elevated
posteriorly or indistinct from anterior part of lower jaw, protrusibility of jaws variable; teeth present or absent
in jaws, palatines, pterygoids (roof of mouth) or on outside of mouth; gill rakers variable, from short and
stubby to long and slender, ranging in number from few to numerous (10 to 26) on lower arm of first
gill arch. Dorsal fins widely separated, first with a variable number of small, flexible spines, originating
in front of, or behind, vertical through tips of pelvic fins; second dorsal fin and anal fin with I weak spine,
1 unbranched soft ray and a variable number of branched soft rays; anal fin always originating slightly in
advance of second dorsal fin; pectoral fins set high on body, directly behind posterior rim of gill cover,
with spine greatly reduced and first ray much thicker than those following. Pelvic fins with I spine and 5
soft rays; caudal fin moderately forked. Anus in front or behind tips of pelvic fins. Lateral line absent. Scales
cycloid (smooth to touch), moderately large; midlateral scales usually with pore or pit; axillary pelvic scales
often present. Colour: in life, blue-green, green, or olive on back, translucent with scales delineated by
small chromatophores above midlateral band; sides of head and body, as well as abdomen silvery;
midlateral band usually distinct and running from upper margin of pectoral fin to base of caudal
fin, but sometimes blending with silvery abdomen anteriorly; fins clear or dusky; blotch on pectoral fin
present in some species. st
1 dorsal fin with 2nd dorsal fin with
flexible spines I weak spine
midlateral stripe
pelvic fin with I anal-fin spine

I spine and 5 soft rays (weak)
Habitat, biology, and fisheries: Mostly close inshore, near the surface, to a depth of about 1 to 2 m. Some
species schooling in large numbers; some preferring to stay in small schools. Breeding either annual or
opportunistic. Eggs moderately large (up to about 1 mm) with filaments present. Only 2 Old World species
(Atherina breviceps, Atherinomorus lacunosus) are large enough to be valuable as human food; other
species are important as forage for commercial fishes and used as bait and dried cat food.

Mugilidae: head broader and flattened, snout blunt; first dorsal fin with IV slender spines, anal fin with up
to III spines (I in Atherinidae); eyes often covered with adipose lids; scales frequently ctenoid (rough to
touch) on sides of head; no midlateral band on body.
Engraulidae: a single dorsal fin; no spines in fins; pectoral fins low on body; mouth very large; snout blunt
and projecting.
no midlateral
often an adipose stripe
eyelid 1 dorsal fin
III anal-fin spines

Mugilidae Engraulidae
2114 Bony Fishes
Identification note
In the keys and throughout the following descriptions all soft rays, including the first unbranched ray, have
been included in the ray count.
Key to the genera of Atherinidae occurring in the area

Note: there has been much confusion with atherinid species, possibly due to their small size, with lack of
commercial importance leading to insuffcient attention to this group. They are small silvery fishes which
are superficially similar in size, coloration, and external morphology. Distinct differences, however, do occur
in some measurements, counts, and in osteology. Mouthparts are frequently diagnostic of species.
1a. Preopercular notch present

(Fig. 1a) . . . . . . . . . . . . . . . ® 2 notch on anterior edge
premaxilla of preopercle
1b. Preopercular notch absent
(Fig. 1b) . . . . . . . . . . . . . . . . ® 3
2a. Ascending process of pre-
maxilla short and broad,
(Fig. 2a) its length about 1/3
diameter of eye; lateral proc-
ess of premaxilla broad and a) b) no notch
flat; mandibular ramus not here
Fig. 1 lateral view of head
elevated (Fig. 3a); slight ele-
vation at distal end of den-
tary present or absent . . Atherinomorus
2b. Ascending process of premaxilla moderately long and narrow, its length about 1/3 to
1/2 diameter of eye; lateral process of premaxilla short and conical (Fig. 2b); lower jaw
distinctly elevated posteriorly (Fig. 3b) . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoatherina
2c. Ascending process of premaxilla long, almost equal to diameter of eye; lateral process
narrow and pungent (Fig. 2c); lower jaw elevated; whole of mandibular ramus elevated
posteriorly (Fig. 3c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stenatherina
(a single species, S. panatela, in the area)
ascending ascending
process process
ascending
process
lateral lateral
process process pungent
lateral
process
free edge with
upper jaw teeth distal
(hind)
end
a) Atherinomorus b) Hypoatherina c) Stenatherina
Fig. 2 premaxilla
posterior posterior posterior

ramus ramus ramus
a) Atherinomorus b) Hypoatherina c) Stenatherina
Fig. 3 dentary
no
3a. Shagreen denticles pre- shagreen denticles
sent outs ide of mouth denticles outside
(Fig. 4a); origin of first dor- mouth
sal fin always behind verti-
cal through tips of pelvic
fins; anal-fin rays always
12 or more; midlateral b) Craterocephalus
a) Atherion
scale count always 40 or
more . . . . . . . . . . . . . . Atherion Fig. 4 lateral view of head
(a single species. A. elymus, in the area)
3b. Shagreen denticles absent (Fig. 4b); origin of first dorsal fin either in front or behind tips
of pelvic fins; anal-fin rays 12 or less, never more; midlateral scale count always less
than 40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Craterocephalus
Key to the species of Atherinomorus occurring in the area

1a. Anus always behind tips of pelvic fins; premaxilla very long, extending well past vertical
through anterior border of eye . . . . . . . . . . . . . . . . . . . . . Atherinomorus capricornensis
1b. Anus always at or in front of tips of pelvic fins; premaxilla long, but not usually extending
well past vertical through anterior border of eye . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Midlateral scales never more than 39, with means from 34 to 37 . . . . . . . . . . . . . . . . . ® 6
2b. Midlateral scales may be less than 39, but with mean always greater than 39 . . . . . . . . . . ® 3
3a. Midlateral scales 44 to 47; 2 anteriormost predorsal scales on either side of midline and
in line with vertical through anterior border of eye . . . . . . . . . . . . . Atherinomorus insularum
3b. Midlateral scales 38 to 44; anteriormost predorsal scales single in midline and almost
in front of vertical through anterior border of eye . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Midlateral band narrow, restricted to third transverse scale row; origin of first dorsal fin
from 1 scale in front to 4 scales behind vertical through tips of pelvic fins . . Atherinomorus ogilbyi
4b. Midlateral band not restricted to and always wider than third scale row; origin of first
dorsal fin never in front of vertical through tips of pelvic fins . . . . . . . . . . . . . . . . . . . . ® 5
5a. Midlateral scales 39 to 41; midlateral band wider than single scale but not as wide as
in following species; predorsal scales (18 or 19) with well defined point posteriorly;
dentary slightly elevated posteriorly with tubercle at distal end; ramus of premaxilla just
reaching past vertical through anterior border of eye . . . . . . . . . . . Atherinomorus cylindricus
5b. Midlateral scales 39 to 44; midlateral band very wide; predorsal scales (17 to 22)
rounded posteriorly; dentary without tubercle and not elevated; ramus of premaxilla
reaching well past vertical through anterior border of eye . . . . . . . . . Atherinomorus lacunosus
6a. Anal fin with 10 to 12 soft rays; midlateral scales 33 to 37; series of dots on scales
extending as lines below midlateral band . . . . . . . . . . . . . . . . . . . Atherinomorus lineatus
6b. Anal fin always with 11 or more soft rays; midlateral scales always 34 or more; series
of dots not extending as lines below midlateral band . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Gill rakers never less than 21 and up to 25 on lower limb of first gill arch; tubercle-like
elevation present or absent at distal end of dentary; dentary may be slightly elevated . . . . . . ® 8
7b. Gill rakers on lower limb of first gill arch 18 to 22 (usually about 20); small tubercle-like
elevation often present at distal end of dentary; dentary never elevated . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atherinomorus endrachtensis
8a. Lower jaw protruding, forming anteriormost part of head; origin of second dorsal fin only
slightly behind vertical through origin of anal fin . . . . . . . . . . . . . Atherinomorus balabacensis
8b. Lower jaw not protruding to form anteriormost part of head; origin of second dorsal fin
well behind vertical through origin of anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
2116 Bony Fishes
9a. Position of anus never more than 2 scales in front of tips of pelvic fins; origin of first
dorsal fin always at or up to 2 scales in front of vertical through tips of pelvic fins, never
behind; predorsal scales 14 to 18; width of midlateral band 2.9 to 3.6 times in body depth
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atherinomorus reginae
9b. Position of anus 1.5 to 3 scales in front of tips of pelvic fins; origin of first dorsal fin from
1 scale behind to 1.5 scales in front of vertical through tips of pelvic fins; predorsal scales
17 to 19; width of midlateral band 4.8 to 6.6 times in body depth . . . . Atherinomorus duodecimalis
Key to the species of Craterocephalus occurring in the area

1a. Midlateral scales always 33 or more; 12 to 17 gill rakers on lower limb of first gill arch;
gill rakers moderately long and slender, about 1/2 diameter of pupil; body depth 4.5 to
7.2 times in standard length, with mean never less than 5.2 . . . . . . . . . . . . . . . . . . . . ® 2
1b. Midlateral scales always 33 or less; gill rakers never more than 13 on lower limb of first
gill arch; gill rakers always less than 1/2 diameter of pupil; body depth never more than
5.4 times in standard length, with mean never more than 4.9 . . . . . . . . . . . . . . . . . . . ® 3
2a. Single black spot at base of pectoral fin; gill rakers equal to diameter of pupil; dentary
lateral fossa for insertion of labial ligament immediately adjacent to dentary symphysis
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Craterocephalus mugiloides
2b. No black spot at base of pectoral fin; gill rakers long but slightly less than diameter of
pupil; dentary lateral fossa for insertion of labial ligament never adjacent to dentary
symphysis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Craterocephalus honoriae
3a. Transverse scales 6 to 8; anus always at or behind tips of pelvic fins; midlateral scales
26 to 29 (mean 27); needle-like teeth in 2 rows in upper jaw, in single row in lower jaw
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Craterocephalus munroi
3b. Transverse scales 5 to 5 ½; anus always in front of tips of pelvic fins; midlateral scales
29 to 33 (mean 30.8); small teeth in both jaws in 2 rows . . . . . . . . . . Craterocephalus capreoli
Key to the species of Hypoatherina occurring in the area

1a. Body scales distinctly crenulated on posterior edge; body robust, its depth never more
than 5.9 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Body scales not crenulated; body slender, its depth never less than 5.5 times in standard
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Midlateral scales 39 to 45; anal-fin rays 9 to 12 (mean 11.4); body depth 4.8 to 5.9 (mean
5.6) times in standard length . . . . . . . . . . . . . . . . . . . . . . . Hypoatherina valenciennesi
2b. Midlateral scales 33 to 35; anal-fin rays 8 or 9; body depth 4.1 to 5.2 (mean 4.5) times
in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoatherina crenolepis
3a. Midlateral band wide, its width 2.3 to 3.2 (mean 2.7) times in body depth; pectoral-fin
rays 11 to 14; 15 to 18 (mean 16.7) gill rakers on lower limb of first gill arch; anus always
behind tips of pelvic fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoatherina barnesi
3b. Midlateral band narrow, its width 2.8 to 7.1 times in body depth; pectoral-fin rays 14 to
18; 18 to 25 gill rakers on lower limb of first gill arch; anus in front or behind tips of
pelvic fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Dorsal process of premaxilla long and narrow, 1.3 to 1.8 times in diameter of eye; anal
fin with 8 to 11 (mean 9) soft rays; midlateral band very narrow, its width 4 to 7 times
in body depth; anus always in front of tips of pelvic fins . . . . . . . . . . . Hypoatherina ovalaua
4b. Dorsal process of premaxilla short or moderately long, but never less than 1.7 times in
diameter of eye; anal fin always with 10 or more soft rays; lateral band moderately wide,
its width never more than 5.1 times in body depth; anus in front or behind tips of pelvic
fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Gill rakers on lower limb of first gill arch 21 to 25 (mean 22.9); predorsal scales 16 to
18; dorsal process of premaxilla of moderate length, 1.7 to 2.5 (mean 2) times in
diameter of eye; origin of pelvic fins either behind or in front of tips of pectoral fins . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoatherina temminckii
5b. Gill rakers on lower limb of first gill arch 18 to 22 (mean 20.3); predorsal scales 17 to
22; dorsal process of premaxilla short, 2.2 to 3.7 (mean 2.9) times in diameter of eye;
origin of pelvic fins always in front of tips of pectoral fins . . . . . . . . . . Hypoatherina tropicalis

0 Atherion elymus Jordan and Starks, 1901
0
0
0
Atherinomorus balabacensis (Seale, 1910)
Atherinomorus capricornensis (Woodland, 1961)
0
Atherinomorus cylindricus (Valenciennes, 1835)
0
Atherinomorus duodecimalis (Valenciennes, 1835)
0
Atherinomorus endrachtensis (Quoy and Gaimard, 1824)
0
Atherinomorus insularum (Jordan and Evermann, 1903)
0
Atherinomorus lacunosus (Forster and Schneider, 1801)
0
Atherinomorus lineatus (Günther, 1872)
0
Atherinomorus ogilbyi (Whitley, 1930)
Atherinomorus reginae (Seale, 1909)
0
0
Craterocephalus capreoli Rendahl, 1922
0
Craterocephalus honoriae (Ogilby, 1912)
0
Craterocephalus mugiloides (McCulloch, 1913)
Craterocephalus munroi Crowley and Ivantsoff, 1988
0
0
Hypoatherina barnesi Schultz, 1953
0
Hypoatherina crenolepis Schultz, 1953
0
Hypoatherina ovalaua (Herre, 1935)
0
Hypoatherina temminckii (Bleeker, 1853)
0
Hypoatherina tropicalis (Whitley, 1948)
Hypoatherina valenciennei (Bleeker, 1853)
0 Stenatherina panatela (Jordan and Richardson, 1908)
References
Gomon, M., C.J.M. Glover, and R. Kuiter. 1994. Fishes of Australia’s south coast. Adelaide, State Press, 992 p.
Ivantsoff, W. 1984. Atherinidae. In FAO species identification sheets for fishery purposes. Western Indian Ocean
Ivantsoff, W. and L.E.L.M. Crowley. 1991. Review of the Australian silverside fishes of the genus Atherinomorus
(Atherninidae). Aust. J. Mar. Freshwater. Res., 46(6):479-505.
Ivantsoff, W. and L.E.L.M. Crowley. 1992. Redefinition of the freshwater genus Craterocephalus (Teleostei:
Atherinidae) of Australia and New Guinea, with an analysis of three species. Ichthyol. Explor., Freshwaters,
3(3):1-15.
2118 Bony Fishes
Atherion elymus Jordan and Starks, 1901

Frequent synonyms / misidentifications: Atherion elymus freyi Schultz, 1953; A. e. asper Schultz, 1953;
A. e. aphrozoicus Schultz, 1953 / Atherion maccullochi (Ogilby, 1915).
FAO names: En - Bearded silverside.
Diagnostic characters: Body slender. Mouth smallish, strongly oblique to horizontal, barely protractile, with
thin lips. No notch present on anterior border of preopercle. Shagreen denticles present outside mouth.
Upper jaw well toothed, dentary with patches of small teeth anteriorly. Rest of dentition variable, with teeth
present or absent on vomer, rarely present on palatines, but usually found on basihyal and mesopterygoids.
Free edge of premaxilla rounded and heavily toothed with notch well behind ascending process, then
becoming straight or slightly concave towards corner of mouth. Ascending process of premaxilla blunt, short,
and wide; no lateral processes of premaxilla present. Distal end of maxilla blunt and truncate. Gill rakers
on lower limb of first gill arch 10 to 14, moderately slender and short, less than diameter of pupil. First dorsal
fin with III to V spines, its origin about 6 to 10 scales behind vertical through tips of pectoral fins. Second
dorsal fin with I spine and 8 to 13 soft rays, its origin behind vertical through anal-fin origin. Anal fin with I spine
and 13 to 15 soft rays. Pectoral fins with I spine and 11 to 14 soft rays. Body scales moderately large,
dorsoventrally elongated with circuli restricted to anterior half of scale; 40 to 44 midlateral scales; 5 or 5 ½
scales in transverse rows along side of body. Predorsal scales 16 to 19; interdorsal scales 5 to 8. Anus well
behind tips of pelvic fins. Colour: variable in preserved specimens, from light brown with upper half of
midlateral band extremely dark; heavy pigmentation around snout, dorsum of head, and lower jaw; cheeks
lighter with fewer chromatophores; in live specimens, abdomen, lower half of midlateral band, and opercle
silvery; spots on middorsal line large and silvery blue.
Habitat, biology, and fisheries: Can be collected in tide pools, surge channels, and coralline ridges of outer
reefs. Eggs large. No known commercial value.
Distribution: Wide-ranging in
the Pacific, extending from
about 18° south along the
eastern Australian coast to
about 30° north in Japan.
Seems to be restricted to the
western Pacific Ocean and
has been collected close to the
islands in Micronesia, Melane-
sia, Fiji, and New Guinea.
Atherinomorus balabacensis (Seale, 1910)

Frequent synonyms / misidentifications: None / Atherinomorus endrachtensis (Quoy and Gaimard, 1824);
A. duodecimalis (Valenciennes, 1835).
FAO names: En - Balabac Island silverside.
Diagnostic characters: Body elongate and deep, slightly compressed. Head conical, with depth greater than
width. Mouth large and oblique, just reaching past anterior margin of orbit. Lower jaw protruding, terminal.
Teeth in jaws small; teeth also present on vomer, mesopterygoid, ectopterygoid, and on back of tongue but
absent on palatines. Gill rakers on lower limb of first gill arch 21 to 24, long and slender. First dorsal fin with
IV to VI flexible spines, its origin 1 or 2 scales behind tips of pectoral fins. Second dorsal fin with I spine and
8 to 11 soft rays, its origin slightly behind vertical through anal-fin origin. Anal fin with I spine and 12 to 14 soft
rays. Pectoral fins with I spine and 14 to 18 soft rays. Body scales large with smooth margins; scales apparently
deciduous on head; 36 to 38 midlateral scales; 5 or 5 ½ scales in transverse rows along side of body.
Predorsal scales 16 to 18; interdorsal scales 6 or 7. Anus 1 ½ to 4 scales in front of tips of pelvic fins. Colour:
preserved specimens uniformly yellow-green with thin silvery midlateral band; midlateral band with very
fine, blue-black edge above; scales above midlateral band with sprinkling of fine melanophores; row of scales
below with small black dot on each, forming continuous line along body; scale rows below with indistinct,
incomplete line of dots; dark line along middorsal contour from back of head to base of caudal fin; fins mostly
uniformly yellowish with dusky patch at base of pectoral fins; caudal fin with slight dusky tinge; eye with darker
markings on top.
Size: Maximum length about 6 to 7 cm.
Habitat, biology, and fisheries: Generally found close inshore sometimes close to mouth of streams. Nothing
is known of the biology of this species. Of no commercial value but may be taken as food by larger commercial
species.
Distribution: Philippines (Bala-
bac, Samar, Cebu, Siquijor,
Mindanao, Palawan, Culion,
Samar Islands); also Trobriand
Islands.
2120 Bony Fishes
Atherinomorus capricornensis (Woodland, 1961)

Frequent synonyms / misidentifications: Pranesus capricornensis Woodland, 1961 / Pranesus ogilbyi Whitley,
1930; P. pinguis (Lacepède, 1803).
FAO names: En - Capricorn silverside.
Diagnostic characters: Body small and slender, somewhat compressed. Small shagreen denticles present
on lips of upper and lower jaws. Teeth in both jaws small, villiform, and in more than 1 row on premaxilla, often
extending to outer surface. Teeth on other bones variable in size, often long and pungent on vomer and
either forming a strong ridge or scattered over mesopterygoid; teeth also present on palatines, ectopterygoids,
and tongue; free edge of lower jaw sloping backwards and upwards without distinctly elevating at posterior
end. Ramus of premaxilla long, extending past vertical through anterior border of pupil, anterior process
short, lateral process absent. Gill rakers on lower limb of first gill arch 20 to 24 (usually 22 or 23). First dorsal
fin with III to VII spines; second dorsal fin with I spine and 9 or 10 soft rays; anal fin with I spine and 12 to 15
soft rays; pectoral fins with 14 to 17 soft rays. Midlateral scales 41 to 45 (usually 43 to 45); always 5 scales
in transverse rows along side of body. Predorsal scales 19 to 22; interdorsal scales 7 to 9. Anus up to 3 scales
behind tips of pelvic fins. Colour: blue-green, light or dark grey and silvery below; midlateral band wide,
covered by third and part of fourth row of midlateral scales; flesh translucent above wide midlateral band, its
upper edge iridescent blue; green chromatophores forming middorsal line. Pectoral blotch present or absent;
ventral surface often iridescent; snout and opercle silvery; fins dusky but pectoral fins sometimes yellowish;
posterior half of scales above midlateral band heavily pigmented, often with C-shaped or diamond-shaped
mark on each scale; eyes iridescent blue-green.
Habitat, biology, and fisheries: An island species of the Great Barrier Reef. Schools in numbers ranging
from several hundred to schools of more than 100 m long and 20 m wide, lying close inshore in depths of
about 1 m, generally about from 15 cm below surface to bottom. Feeding occurs mostly at night when schools
disperse. Food comprises a variety of planktonic crustaceans. Subject to predation by sharks, tunas, long
toms, and amberjacks, also by terns, gannets, seagulls, and herons. May be caught with seines in large
numbers but seem to be of little commercial value.
Distribution: From the Capri-
corn Group (e.g. Heron Island
(23°27’S, 151°55’E) and some
islands closer to the mainland,
as far north as Borth Barnard
Island, 17°40’S, 146°10’E). It
is potentially sympatric with 3
other species of Atherino-
morus: A. lacunosus, A. en-
drachtensis a nd A. ogilbyi.
Species of Hypoatherina also
co-occur.
Atherinomorus cylindricus (Valenciennes, 1835)

Frequent synonyms / misidentifications: None / Atherinomorus duodecimalis (Valenciennes, 1835);
A. endrachtensis (Quoy and Gaimard, 1824).
FAO names: En - Waigeo silverside.
Diagnostic characters: Body slender, slightly compressed. Mouth moderately large. Teeth in lower jaw
minute, sparse, but evenly spaced; in upper jaw minute and with some fine shagreen denticles on lips. Teeth
also present on vomer, tongue, mesopterygoids, and ectopterygoids, but may be difficult to see in last 3.
Ascending process of premaxilla short and moderately broad; lateral process broadly rounded. Dentary slightly
elevated posteriorly. Gill rakers on lower limb of first gill arch 20 to 22, long and slender, about equal to
diameter of pupil, with fine spinules along length. First dorsal fin with V or VI weak spines, its origin 2 to 6
scales behind vertical through tips of pectoral fins. Second dorsal fin with I spine and 10 or 11 soft rays; anal
fin with I spine and 12 to 14 soft rays. Pectoral fins with I small spine and 15 or 16 soft rays. Body scales
dorsoventrally elongated and rounded posteriorly; 39 to 41 midlateral scales; 5 scales in transverse rows
along side of body. Predorsal scales 18 or 19, forming well defined point posteriorly; interdorsal scales
6 to 8. Anus 2 to 4 scales in front of pelvic-fin tips. Colour: preserved specimens yellowish with
silvery-brown midlateral band; dark, thick reticulate pattern present on dorsal scales; single row of melano-
phores present (1 on each scale) below midlateral band from origin of pectoral fin as far as vertical through
anal-fin origin; eye and opercle silvery; snout and dorsum of head dark.
Size: Maximum length about 6.5 cm.
Habitat, biology, and fisheries: Like other members of this genus found close inshore around islands.
Nothing is known of its biology. Of no commercial value but probably preyed upon by larger species.
Distribution: Waigeo Island;
Utupua Island, Santa Cruz
Group.
2122 Bony Fishes
Atherinomorus duodecimalis (Valenciennes, 1835)

Frequent synonyms / misidentifications: Pranesus duodecimalis (Valenciennes, 1835); Allanetta
duodecimalis (Valenciennes, 1835) / Atherinomorus endrachtensis (Quoy and Gaimard, 1824); A. cylindricus
(Valenciennes, 1835).
FAO names: En - Tropical silverside; Fr - Athérine tropicale; Sp - Pejerrey tropical.
Diagnostic characters: Body small, robust, subcylindrical, and laterally compressed. Head and eyes
moderately large. Posterior edge of preopercle more or less straight and not projected into a lobe. Distal end
of upper jaw extending slightly backward slightly beyond vertical through anterior border of orbit, its
ascending process moderately short and wide. Outer surface of premaxilla frequently covered with fine
denticles scarcely visible to naked eye, but no shagreen denticles elsewhere on head. Teeth in several rows
in both jaws. On roof of mouth, teeth usually distinctly visible (but may be weak or lacking) on palatines,
ectopterygoids, or mesopterygoids; teeth also present around edge of tongue. Dentary bone sloping
backward and upward, with distinct tubercle-like elevation at its distal end. Gill rakers moderately long,
slightly less than diameter of pupil, 21 to 25 on lower limb of first gill arch. First dorsal fin with IV to VI spines;
second dorsal fin with I spine and 9 or 10 soft rays; anal fin with I spine and 12 or 13 soft rays; pectoral fins
with I spine and 14 to 17 soft rays. Body scales large; 33 to 38 midlateral scales; about 5 scales in transverse
rows along side of body. Predorsal scales 17 to 19; interdorsal scales 6 or 7. Anus 2 to 4 (rarely more than
3) scales in front of pelvic-fin tips. Colour: greenish above, whitish below when alive, with silvery midlateral
band; preserved specimens brown-green with black midlateral band; edges of scales on back with numerous
small chromatophores; rows of spots on third and fourth row of scales extending from about tips of pectoral
fins to origin of anal fin or slightly beyond.
Habitat, biology, and fisheries: Nothing is known of the biology of this species except that it inhabits shallow
coastal waters. It can be caught with seines but is probably only of use as a forage fish for larger commercial
species. It may also be used as bait.
Distribution: This species ex-
tends well beyond the area into
the Indian Ocean. Found off
the coast of Java, Thailand,
Viet Nam, Papua New Guinea,
New Britain, Solomon Islands,
New Caledonia, and Utupa Is-
land (Santa Cruz Group).
Atherinomorus endrachtensis (Quoy and Gaimard, 1824)

Frequent synonyms / misidentifications: Pranesus endrachtensis (Valenciennes, 1835); Atherinomorus
eendrachtensis (Quoy and Gaimard, 1824) (misspelling) / Atherinomorus ogilbyi (Whitley, 1930);
A. duodecimalis (Valenciennes, 1835); A. lineatus (Günther, 1872).
FAO names: En - Endracht land silverside.
Diagnostic characters: Body moderately small, subcylindrical. Mouth barely protractile, large, and moder-
ately oblique. Teeth in both jaws small, premaxilla covered with fine shagreen denticles, occasionally also
on snout; teeth always present on vomer, but present or absent on palatines and ectopterygoids; strong ridge
of teeth always present on mesopterygoids. Free edge of premaxilla moderately short, not extending
past vertical through anterior border of pupil; dorsal process of premaxilla broad, lateral process absent.
Small tubercular elevation often present at distal end of dentary, but dentaries never elevated. Gill rakers
on lower limb of first gill arch 18 to 22 (usually about 20), relatively long, equal to or greater than diameter
of pupil. First dorsal fin with IV to VI weak spines; second dorsal fin with I spine and 8 to 10 soft rays; anal
fin with I spine and 11 to 14 soft rays; pectoral fins with 12 to 16 soft rays. Midlateral scales 34 to 39 (usually
36 to 37); 5 or 5 ½ scales in transverse rows along side of body. Predorsal scales 16 to 19; interdorsal scales
5 or 7. Anus 2 to 4 scales in front of pelvic-fin tips, usually close to origin of pelvic fins. Colour: live specimens
from Western Australia brown-yellow with distinct chromatophores on edges of scales; eye, preopercle, and
midlateral band silvery; snout and all fins slightly dusky; pectoral blotch, if present, distinct but not intense.
Silvery peritoneum and posterior end of swimbladder visible through translucent flesh; preserved specimens
light yellow, yellow-brown, or yellow-green; upper half of body darker; midlateral silvery black band edged by
narrow black line above; blotch on pectoral fins present or absent.
Size: Maximum standard length about 8.4 cm.
Habitat, biology, and fisheries: Inhabits shallow coastal waters. Nothing is known of the biology of this
species other than that it schools together in shallow waters with Atherinomorus lacunosus, A. ogilbyi,
Craterocephalus mugiloides, and Hypoatherina temminckii in coastal areas of northern Australia. Potentially
sympatric with Atherinomorus duodecimalis in New Guinea, New Britain, and the Solomon Islands. May be
collected with seines but has no commercial value.
Distribution: Northern waters
of Australia; Vanuatu, Papua
New Guinea, Admiralty Is-
lands, New Britain, Solomon
Islands, and New Caledonia.
Sold in small quantities at
some markets in the Philip-
pines.
2124 Bony Fishes
Atherinomorus insularum (Jordan and Evermann, 1903)

Frequent synonyms / misidentifications: Atherina insularum (Jordan and Evermann, 1903) /
Atherinomorus lacunosus (Forster and Schneider, 1801); Pranesus pinguis (Lacepède, 1803).
FAO names: En - Hawaiian silverside.
Diagnostic characters: Body slender, moderately compressed. Head and eyes not particularly large
(when compared with Atherinomorus duodecimalis). Outer surface of mouth frequently covered by fine
shagreen denticles; labial ligament long and fleshy, reaching to 2/3 way along lower jaw. Teeth in jaws small,
needle-like, and usually in single row. Teeth also present on posterior part but absent from front of basihyal;
single ridge of teeth on mesopterygoid and ectopterygoid; teeth on vomer very small and difficult to see.
Free edge of premaxilla reaching past vertical through anterior border of orbit, its ascending process short
and blunt and lateral process broadly rounded. Dentary gently sloping upward and backward. Gill rakers with
spinules, slender and longer than diameter of pupil, 19 to 23 on lower limb of first gill arch. First dorsal fin
with V to VII weak spines, its origin 4 to 7 scales behind tips of pectoral fins. Second dorsal fin with I spine
and 10 or 11 soft rays, its origin behind vertical through origin of anal fin. Anal fin with I spine and 15 to 18
soft rays. Pectoral fins with I small spine and 14 to 17 soft rays. Midlateral body scales 44 to 47, moderately
large, dorsoventrally elongated; 5 or 5 ½ scales in transverse rows along side of body. Predorsal scales 20
to 23; 2 predorsal scales on dorsum of head, on either side of centre line and almost in front of vertical through
orbit; interdorsal scales 6 to 9. Anus at, or only slightly (1 scale) in front of tips of pelvic fins. Colour: preserved
specimens yellowish with fine blue-black line above diffuse but broad silvery midlateral band; opercle silvery;
eye mostly silvery with a dark section at the top; dorsum of head dark; snout dark above lips; scales above
midlateral band outlined with melanophores forming a reticulate pattern; fins clear.
Habitat, biology, and fisheries: The species lays eggs of about 1 to 1.5 mm diameter which have long tendrils
attached to chorion. Larvae hatch at 4 mm. Fully mature at about 15 mm. This silverside swims in small schools
during daylight hours close to the shore line and disperses at night spreading over the reef. A nocturnal
plankton eater, its gut content containing mostly crustaceans and foraminiferans. Like other species of this
family, it is of no commercial
value but may be taken by
larger species as food.
Distribution: Found in the wa-
ters of Hawaii. A population
identified as its subspecies,
occurs off the south coast of
Molokai Island, 22°N 3°08’E.
Saipan, Mariana Islands.
Atherinomorus lacunosus (Forster and Schneider 1801)

Frequent synonyms / misidentifications: Pranesus pinguis (Lacepède, 1803); P. vaigiensis (Quoy and
Gaimard, 1825); P. maculatus Taylor, 1964); Atherina forskalii (Rüppell, 1838); A. morrisi Jordan and Starks,
1906 / Atherinomorus ogilbyi (Whitley, 1930), A. capricornensis (Woodland, 1961).
FAO names: En - Hardyhead silverside; Fr - Atherin têtu; Sp - Pejerrey cabezón.
Diagnostic characters: Body subcylindrical, laterally compressed. Head and eyes moderately large. Distal
end of upper jaw extending backward to vertical through anterior border of pupil, its ascending process
short and blunt, lateral process low and broad. Outer surface of premaxilla frequently covered with fine
denticles, scarcely visible to naked eye; no shagreen denticles elsewhere on head. Teeth in jaws small,
villiform, usually in more than 1 row on premaxilla. Teeth also present on roof of mouth (palatines, vomer,
sometimes also ectopterygoids). Free edge of dentary bone almost flat, with no posterior elevation. Gill
rakers long and slender, equal to or longer than diameter of pupil, 18 to 25 on lower limb of first gill arch. First
dorsal fin with IV to VII spines; second dorsal fin with I spine and 8 to 11 soft rays; anal fin with I spine and
12 to 17 soft rays; pectoral fins with 14 to 18 soft rays. Body scales large; 39 to 44 midlateral scales; 5 or
5 ½ transverse scales rows along side of body. Predorsal scales 17 to 22; interdorsal scales 6 to 9. Anus at,
or slightly in front of, tips of pelvic fins. Colour: blue-green and translucent with silvery parts of abdomen and
head frequently iridescent; midlateral band wider than 1 scale row along side, silvery and often merging with
silvery abdomen on anterior part of body; opercle and iris silvery; upper surface of head heavily pigmented;
scales on back usually outlined with small chromatophores; fins clear to dusky, pectoral fins frequently with
blackish blotch.
Habitat, biology, and fisheries: During the day, this species aggregates in large, relatively inactive schools,
close inshore. The fish disperse over reefs at sunset, keeping about 2 to 4 m apart. Feeding occurs at night
and food includes planktonic eggs, crustaceans, foraminiferans, amphipods, ostracods, calenoid copepods,
and caridean shrimp larvae (most plankton less than 0.5 mm in size). Small fish such as Clupea and Sardinella
species are also taken as food. The schools reform in the early morning and move back closer to shore. Small
individuals, less than 3.5 cm standard length, form small schools also relatively close to the shore and feed
actively during daylight hours. Atherinomorus lacunosus is one of the largest and most robust of silversides
occurring in the area. It is known to be sold at fish markets in various parts of the world.
Distribution: Ambon Island
(Indonesia), Waigeo Island
(West Irian), New Guinea,
Vanuatu; northern Australia,
Fiji, New Caledonia, and
Tonga. Outside the area also
found in the eastern Mediterra-
nean, Indian Ocean, and in the
Pacific as far north as Japan.
2126 Bony Fishes
Atherinomorus lineatus (Günther, 1872)

Frequent synonyms / misidentifications: Atherina lineata Günther, 1872 / Atherinomorus endrachtensis
(Quoy and Gaimard, 1824).
FAO names: En - Lined silverside.
Diagnostic characters: Body slender. Eyes large, their diameter equal to interorbital space. Snout very
short. Lips thin, fusing at junction of premaxilla and maxilla. Teeth in upper and lower jaws in several rows;
shagreen denticles present on lips; teeth also present on palatines, vomer, and mesopterygoids. Premaxilla
extending just past vertical through anterior margin of orbit; ascending process of premaxilla blunt, short, and
broad based. Gill rakers on lower limb of first gill arch 20 to 23. First dorsal fin with IV or V weak spines, its
origin at, or up to 2 ½ scales behind, vertical through tips of pelvic fins. Second dorsal fin with I spine and 7
to 9 soft rays. Anal fin with I spine and 10 to 12 soft rays. Pectoral fins long and pointed. Body scales
dorsoventrally elongated; 33 to 37 midlateral scales; 5 scales in transverse rows along side of body.
Predorsal scales 17 to 19; interdorsal scales 6 or 7. Anus 2 ½ to 5 scales in front of pelvic-fin tips. Colour: in
preserved specimens, only narrow upper edge of midlateral band visible as lighter yellow-gold or black;
series of black dots on scale rows extending as lines below midlateral band; eye silvery black.
Habitat, biology, and fisheries: Appears to live in shallow waters in cays or close to shore. Nothing is known
of the biology. The species is not well known and is unlikely to have any commercial value.
Distribution: Amboyna Cay
(Spratly Islands), Halmahera
Island (Indonesia), and Cebu
(Philippines).
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Atherinomorus ogilbyi (Whitley, 1930)

Frequent synonyms / misidentifications: Pranesus ogilbyi Whitley, 1930 / Pranesus endrachtensis (Quoy
and Gaimard, 1824); Atherinomorus lacunosus (Forster and Schneider, 1801); Pranesus pinguis (Lacepède,
1803).
FAO names: En - Ogilby’s silverside.
Diagnostic characters: Body large, robust, slightly compressed. Free edge of premaxilla extending to just
beyond anterior margin of orbit. Mouth relatively large and oblique; lips thin, with teeth in both jaws small
and barely visible; anterior surface of premaxilla covered by fine shagreen denticles, often extending onto
surface of snout. Teeth always present on vomer and palatines, present or absent on ectopterygoids and
forming strong ridge on mesopterygoids. Ascending process of premaxilla short, lateral process low and broad.
Free edge of dentary flat but distal end slightly elevated. Gill rakers on lower limb of first gill arch 22 to 28
(usually 24 to 26), long, usually equal to or longer than diameter of pupil. First dorsal fin with IV to VII weak
spines; second dorsal fin with I spine and 8 to 11 soft rays; anal fin with I spine and 11 to 16 soft rays; pectoral
fins with 15 to 18 soft rays. Midlateral scales 38 to 43; 5 or 5 ½ scales in transverse rows along side of body.
Predorsal scales 16 to 21; interdorsal scales 6 to 9. Anus 1/2 to 4 scales in front of tips of pelvic fins. Colour:
live specimens bluish green and translucent; midlateraI band narrow, restricted to third transverse scale;
abdomen, opercle, and preopercle usually with iridescent sheen; upper half of midlateral band bright blue; iris
silvery and with black spot on its dorsal half; iris also frequently with red spots (superficial capillaries) on ventral
half.
Size: Maximum standard length about 12.4 cm.
Habitat, biology, and fisheries: Found mostly in marine or estuarine habitats and can survive in water with
salinity as low as 3‰. The life history of this species has been extensively studied in its southern ranges. Its
diet comprises amphipods, copepods, isopods, crab and barnacle larvae, gastropods, polychaetes, chiro-
nomid midges, hymenopterans ants, and plant matter, but the main diet appears to consist of crustaceans
and polychaete worms. Gut content varies from season to season indicating that they are opportunistic
feeders. Life span is 2 years with sexual maturity and breeding occurring during the second year during the
southern hemisphere late winter and spring, from August to October. Breeding may take place both in marine
and estuarine environments. They can be collected with seines and are taken by commercial fishermen in
eastern Australia, probably for bait or fishmeal.
Distribution: Common silver-
side in southeastern and
southwestern Australia, with
range extending into the area
to Rockhampton on the east-
ern coast.
2128 Bony Fishes
Atherinomorus reginae (Seale, 1910)

Frequent synonyms / misidentifications: Atherina regina (Seale, 1910); Allanetta regina (Seale, 1910) /
Atherinomorus duodecimalis (Valenciennes, 1835); A. balabacensis (Seale, 1910).
FAO names: En - Culion silverside.
Diagnostic characters: Body slender. Preopercular notch present. Mouth small. Teeth in both jaws minute
with shagreen denticles present on outside of mouth. Teeth on vomer also minute; no teeth present on
palatines. Ascending process of premaxilla short and broad. Gill rakers on lower limb of first gill arch (22 to
24) long and slender, about 3/4 diameter of pupil. First dorsal fin with V to VII slender spines; first dorsal-fin
origin at, or up to 2 scales in front of, vertical through pelvic-fin tips. Second dorsal fin with I spine and
9 or 10 soft rays, its origin behind vertical through origin of anal fin. Anal fin with I spine and 10 to 12 soft rays.
Body scales large, posterior margin smooth; 36 to 38 midlateral scales; 5 or 6 scales in transverse rows along
side of body. Predorsal scales 14 to 18; interdorsal scales 6 to 8. Anus 1 ½ or 2 scales in front of pelvic-fin
tips. Colour: preserved specimens yellowish with posterior margins of dorsal rows of scales; sides of dentary,
snout, dorsal part of gill cover, and upper surface of eye with dark pigmentation; silvery midlateral band
covering midlateral scale row; line of small dark dots above midlateral band and minute black dots on row of
scales below silvery band; snout and lower jaw dusky; upper part of eye also dark; fins clear to yellowish, but
caudal fin dusky, with 2 or more distinct blotches on base.
Habitat, biology, and fisheries: Like other atherinids, this species is usually found in close inshore habitats.
Nothing is known of its biology and it has no known commercial value. Probably taken by commercial fish
species as food.
Distribution: K n o wn fr om
Culion and Busuanga islands
in the Philippines.
Craterocephalus capreoli Rendahl, 1922

Frequent synonyms / misidentifications: Craterocephalus anticanus Whitley, 1955 / Craterocephalus
pauciradiatus (Günther, 1861).
FAO names: En - Rendahl’s hardyhead.
Diagnostic characters: Body small, robust, subcylindrical. Teeth in both jaws small and conspicuous, all
other bones in mouth without teeth. Premaxilla with long ascending process, just reaching into interorbital
space; lateral process strongly reduced. Free edge of premaxilla reaching vertical through anterior margin of
orbit. Coronoid of dentary highly elevated. Gill rakers short, with spinules, 10 to 13 on lower limb of first
gill arch. First dorsal fin with IV to VI spines, its origin 0 to 2 ½ scales in front of vertical through pelvic-fin
tips. Second dorsal fin with I spine and 6 to 8 soft rays; anal fin with I spine and 7 to 10 soft rays;
pectoral fins with I spine and 11 to 14 soft rays. Body scales dorsoventrally elongated, posterior edge irregular
in larger fish; 29 to 33 midlateral scales; 5 or 6 scales in transverse rows along side of body. Predorsal scales
12 to 15; interdorsal scales 5 to 7. Anus 3 to 5 scales in front of tips of pelvic fins, very close to pelvic-fin
origin. Colour: live specimens pale green or yellow, with silvery abdomen and midlateral band; opercles and
preopercles iridescent; body often with silver iridescent sheen; dorsum of head dark; dorsal fins dusky, caudal
fin dusky yellow, and pelvic fins creamy.
Habitat, biology, and fisheries: Essentially an estuarine and coastal species, co-occurring with Cratero-
cephalus pauciradiatus, C. mugiloides, and Atherinomorus endrachtensis. Nothing is known of its biology
except that this species is capable of withstanding the very high salinities found in Shark Bay, Western
Australia. Can be collected with seines but has no commercial value. It is possibly taken as food by larger
commercial species.
Distribution: Fro m the
Abrolhos Islands in Western
Australia northwards to the
Gulf of Carpentaria (Northern
Territory, Australia).
2130 Bony Fishes
Craterocephalus honoriae (Ogilby, 1912)

Frequent synonyms / misidentifications: Atherina honoriae Ogilby, 1912; Stenatherina honoriae (Ogilby,
1912) / Atherinosoma microstoma (Günther, 1861).
FAO names: En - Estuarine hardyhead.
Diagnostic characters: Body slender, subcylindrical. Mouth small, with gape restricted by labial ligament.
Teeth in both jaws well developed, always present on mesopterygoid and frequently on vomer; edentulous
otherwise. Premaxilla with long ascending process, just reaching into interorbital space; lateral process
reduced. Free edge of premaxilla reaching vertical through anterior edge of orbit. Coronoid of dentary highly
elevated. Gill rakers on lower limb of first gill arch 12 to 15, moderately long, but less than 1/2 diameter
of pupil. First dorsal fin with V or VI spines; second dorsal fin with I spine and 7 or 8 soft rays, its origin 1 ½
to 5 scales in front of tips of pelvic fins; anal fin with I spine and 9 to 12 soft rays; pectoral fins with I spine and
12 or 13 soft rays. Body scales ovate; 33 to 36 midlateral scales; 4 ½ to 6 scales in transverse rows along
side of body. Predorsal scales 12 to 15; interdorsal scales 6 to 8. Anus at or up to 2 scales in front of tips of
pelvic fins. Colour: live specimens dark yellow to greenish brown; edges of scales above midlateral band
outlined by chromatophores; bases of unpaired fins dusky; opercle, eye, and abdomen often silvery; midlateral
band silvery or gold; silvery peritoneum over abdominal cavity visible through skin; silvery sheen lost soon
after death.
Size: Maximum about 7 cm.
Habitat, biology, and fisheries: Estuarine or enclosed marine habitat. Craterocephalus honoriae breeds in
midsummer (January) in southeastern waters of Australia and has a 1 year life cycle. Like many other
atherinids, it is predominantly carnivorous. Frequently caught in estuaries and enclosed coastal lakes, but of
no known commercial value. Likely to be forage for larger juvenile commercial species which occur in estuarine
nursery habitats.
Distribution: Mostly south-
eastern Australia as far north
as Tewantin in southeastern
Queensland.
Craterocephalus mugiloides (McCulloch, 1913)

Frequent synonyms / misidentifications: Atherinichthys punctatus De Vis, 1885; Allanetta mugiloides
(McCulloch, 1913) / Craterocephalus honoriae (Ogilby, 1912); C. pauciradiatus (Günther, 1861); C. capreoli
Rendahl, 1922.
FAO names: En - Spotted hardyhead.
Diagnostic characters: Body robust, subcylindrical. Mouth moderately large, labial ligament inserting
adjacent to symphysis of dentary. Teeth in both jaws of medium size, in 2 rows. Teeth usually present on
mesopterygoid and vomer, otherwise edentulous. Premaxilla with long ascending process, just reaching into
interorbital space; lateral process pungent. Free edge of premaxilla reaching vertical through anterior edge
of orbit or just beyond. Coronoid of dentary highly elevated. Gill rakers on lower limb of first gill arch 13 to
17, their length about 1/2 diameter of pupil, relatively longer than in other species of Craterocephalus. First
dorsal fin with V to VII spines, its origin 1/2 to 5 scales in front of vertical through tips of pelvic fins. Second
dorsal fin with I spine and 7 to 10 soft rays; anal fin with I spine and 9 to 13 soft rays; pectoral fins with I spine
and 12 to 15 soft rays. Body scales on large specimens usually crenulated; 33 to 37 midlateral scales;
4 ½ to 6 scales in transverse rows along side of body. Predorsal scales 11 to 13; interdorsal scales 6 to 9.
Anus 1 to 4 scales in front of tips of pelvic fins. Colour: live specimens almost translucent, dorsal surface
dusky to greenish yellow with chromatophores at margins of scales; distinct concentration of pigment along
middorsal line, from dorsum of head to caudal fin; snout, eye, opercle, preopercle, and abdomen iridescent
or silvery; midlateral band wide, either silvery or olive to golden with narrow orange stripe above it, extending
from origin of pectoral fin to base of caudal fin; black spot often present at base of pectoral fins.
Habitat, biology, and fisheries: A mainly coastal and estuarine hardyhead, which often schools together
with Craterocephalus capreoli, Atherinomorus endrachtensis, and A. ogilbyi. Diet includes crustaceans and
diatoms. It has an extended breeding period throughout southern hemisphere summer months (December to
March). Its life span appears to be 1 year. May be collected with seines but has no known commercial value.
May be useful as food for larger commercially important fish.
Distribution: The range of this
species extends from the
southwest of Western Austra-
lia to Fraser Island on the east-
ern coast. Also found around
the islands in the Gulf of Car-
pentaria and northeastern
Queensland.
2132 Bony Fishes
Craterocephalus munroi Crowley and Ivantsoff, 1988

Frequent synonyms / misidentifications: None / Craterocephalus marjoriae Whitley, 1948.
FAO names: En - Munro’s hardyhead.
Diagnostic characters: Body robust. Mouth moderately large, protractile. Lips somewhat fleshy and gape
restricted by ligament about half-way along free edge of premaxilla. Needle-like teeth in 2 rows in both jaws,
otherwise edentulous. Ascending process of premaxilla moderately long but barely reaching interorbital space;
lateral process small, conical, lying on posterior expansion of ramus. Coronoid of dentary highly elevated. Gill
rakers on lower limb of first gill arch 10 to 12, with small spinules, short, stumpy, less than 1/2 diameter
of pupil. First dorsal fin with V to VII spines; origin of first dorsal fin 4 to 6 scales in front of vertical
through tips of pelvic fins. Second dorsal fin with I spine and 5 to 7 soft rays. Anal fin with I spine and
6 or 7 soft rays. Pectoral fins with I spine and 13 to 15 soft rays. Body scales rounded, not crenulated; 26 to
29 midlateral scales; 6 to 8 scales in transverse rows along side of body. Predorsal scales 10 to 12; interdorsal
scales 5 to 7. Anus from 0 to 2 scales behind tips of pelvic fins. Colour: variable in live specimens, with
abdomen, opercle, and eye silvery to dull brown; midlateral band either distinct or merging with silvery
abdomen; intensity of chromarophores over dorsal half of the body variable; fins clear.
Habitat, biology, and fisheries: Found in fluviatile and estuarine habitats, but nothing is known of the biology
of this species. Unlike many of the marine and estuarine species of atherinids, it does not appear to school
in large numbers nor does it seem to be abundant. Of no known commercial value.
Distribution: E s t u a r i ne
reaches of the Bynoe, Nichol-
son, and Norman rivers in the
lower Gulf of Carpentaria.
Hypoatherina barnesi Schultz, 1953

Frequent synonyms / misidentifications: Allanetta klunzingeri Smith, 1965 / Hypoatherina uislla (Jordan
and Seale, 1906); H. temminckii (Bleeker, 1853).
FAO names: En - Barnes’ silverside; Fr - Atherine de Barnes; Sp - Pejerry de Barnes.
Diagnostic characters: Body small, slender, subcylindrical, and somewhat compressed. Preopercular notch
present (as in all Hypoatherina species). No shagreen denticles on outer surfaces of jaws or other parts of
the head. Fine teeth in both jaws; teeth usually also present on roof of mouth (palatine and pterygoids) and
on tongue. Premaxilla extending backwards to, or just past, vertical through anterior margin of orbit, its
ascending process long and slender (its length, at least twice width); lateral process well defined, although
short and wide at base; second lateral process also present but not visible without dissection. Ramus of
dentary highly elevated posteriorly. Gill rakers moderately long, but less than diameter of pupil, 15 to 19
on lower limb of first gill arch. First dorsal fin with V to VII spines, its origin from 1/2 scales in front to 3
scales behind vertical through tips of pelvic fins. Second dorsal fin with I spine and 8 to 12 soft rays. Anal fin
with I spine and 12 to 17 soft rays. Pectoral fins with 12 to 15 soft rays. Body scales large; 40 to 45 midlateral
scales; 5 scales in transverse rows along side of body. Predorsal scales 16 to 19; interdorsaI scales 6 to 9.
Anus 1 ½ to 3 scales behind pelvic-fin tips. Colour: live specimens green and translucent, midlateral band
and abdomen silvery; preserved specimens brown, green-brown or grey; edges of scales above midlateral
band usually outlined by chromatophores; black middorsal line running from head to end of caudal peduncle,
midlateral band wide, broadest between anus and anal fin and extending onto caudal fin as triangular
blotch on upper half of fin base; upper fifth of band distinctly darker than rest; rows of spots on lower sides
occasionally present, usually merging with strip above origin of anal fin; bases of all fins pigmented; all fins
dusky, especially caudal fin; upper surface of head dark; opercles and preopercles silvery or dark brown.
Size: Maximum length 6.5 cm.
Habitat, biology, and fisheries: Commonly found in large numbers in lagoons. During daylight, schooling
fish may be periodically seen to jump out of the water to avoid predators. They are also easily attracted to
light. The fish are gravid at about 4 to 4.5 cm standard length by July in the northern hemisphere. Like other
species of atherinids, they may be caught by seines but have no known commercial value. Because the
species occurs in larger numbers, it is probably important as forage food for larger fishes.
Distribution: This species
has a very wide distribution
throughout the Pacific Ocean:
Praton Reef (west of Philip-
pines), Marshall Islands, Tami
lsland (New Guinea), Lizard Is-
land (Australia), Vila (Vanu-
a tu), Emirau Island (St.
Mathias Group). Outside the
area, also known from the
western Indian Ocean.
2134 Bony Fishes
Hypoatherina crenolepis (Schultz, 1953)

Frequent synonyms / misidentifications: Allanetta crenolepis Schultz, 1953 / Hypoatherina valenciennei
(Bleeker, 1851); H. woodwardi (Jordan and Starks, 1961).
FAO names: En - Crenulated silverside.
Diagnostic characters: Body small but more robust than in other members of Hypoatherina, slightly
compressed. Eyes large. Mouth oblique to horizontal. Teeth in upper and lower jaws minute; shagreen
denticles along narrow outer margin of premaxilla. A few minute teeth present on vomer and mesoptery-
goid, but none on palatines. Ascending dorsal process of premaxilla short, broad and not entering
interorbital space; lateral process scarcely differentiated. Gill rakers on lower limb of first gill arch 19 to 21,
moderately long and slender, a little shorter than pupil of eye. First dorsal fin with V or VI spines; origin of
first dorsal fin 1/2 to 3 scales in front of vertical through tips of pelvic fins. Second dorsal fin with I spine
and 7 to 9 soft rays. Anal fin with I spine and 9 or 10 soft rays. Pectoral fins longer than in other species
of Hypoatherina, 3.4 to 3.9 times in standard length (c.f. 5 to 6), with I spine and 14 to 16 soft rays. Body
scales with very crenulate posterior edge; 33 to 35 midlateral scales; 5 or 6 scales in transverse rows
along side of body. Predorsal scales 15 to 17; interdorsal scales 6 or 7. Anus 1/2 to 2 scales in front of pelvic-fin
tips. Colour: preserved specimens with darkish pigment at margins of dorsal scales; 2 distinct rows of spots
below silvery lateral band; snout, lower jaw, and dorsal surface of head with dark pigmentation; eye, opercle,
and preopercle silvery.
Habitat, biology, and fisheries: Inhabit close inshore waters around islands. Nothing is known of their
biology. They have no known commercial value but may be taken as forage fish by larger commercial species;
they may also be used as dried pet food or as bait.
Distribution: Tara and Papa-
tog islands (Philippines).
Hypoatherina ovalaua (Herre, 1935)

Frequent synonyms / misidentifications: Allanetta ovalaua (Herre, 1935); Pranesus ovalauus (Herre,
1935) / None.
FAO names: En - Fijian silverside.
Diagnostic characters: Body slender, slightly compressed. Head large, conical. Mouth large, protractile, and
oblique to horizontal. No shagreen denticle on outside of mouth. Lips thin with loose flap of skin lying along
side of dentary. Teeth sparse in lower jaw in many specimens, free edge of premaxilla covered by fine teeth.
Teeth present on vomer and may extend backwards along midline of roof of mouth. Palatine teeth large and
prominent. Teeth always present on mesopterygoids, usually present on tongue present or absent on
ectopterygoids. Premaxilla with long ascending process and 2 lateral processes. Gill rakers on lower
limb of first gill arch 22 to 25. First dorsal fin with IV to VIII spines, its origin 0 to 1 scales in front of vertical
through tips of pelvic fins. Second dorsal fin with I spine and 8 to 10 soft rays. Anal fin with I spine and 9 to
12 soft rays. Pectoral fins with I spine and 15 to 17 soft rays. Body scales dorsoventrally elongated and rounded
posteriorly; 38 to 42 midlateral scales; 5 scales in transverse rows along side of body. Predorsal scales 15 to
19; interdorsal scales 7 to 10. Anus from 0 to 2 scales in front of tips of pelvic fins. Colour: brownish white
with dark scales on back, a silvery midlateral band with dots below it; abdomen pale; dorsum of head very
dark; large dark spot on opercle.
Size: Maximum total length about 7 cm.
Habitat, biology, and fisheries: Inhabits close inshore waters; nothing is known of its biology. Attracted to
light at night in lagoons. Not found in the rougher waters over reefs. Can be collected with seines but has no
commercial value.
Distribution: Ovalau Island,
Fiji, Canton, Hull, Phoenix, Gil-
bert, Marshall, Marianas, and
Hermit islands; New Britain,
and New Ireland.
2136 Bony Fishes
Hypoatherina temminckii (Bleeker, 1853)

Frequent synonyms / misidentifications: Stenatherina temminckii (Bleeker, 1853); Hypoatherina uisila
(Jordan and Seale, 1906); H. gobio (Klunzinger, 1884); Allanetta afra (Peter, 1855) / None.
FAO names: En - Samoan silverside; Fr - Athérine samoan; Sp - Pejerrey samoano.
Diagnostic characters: Body fusiform. No shagreen denticles on external surfaces of jaws. Fine teeth on
dentary and premaxilla. Teeth present on palatines and mesopterygoids; tongue sometimes covered with
small tooth-like papillae. Premaxilla extending to vertical through anterior margin of orbit; ascending process
moderately long and narrow (always longer than twice width); lateral process short and wide. Ramus of
dentary highly elevated posteriorly. Gill rakers on lower limb of first gill arch 21 to 25, moderately long,
but usually less than diameter of pupil. First dorsal fin with V to VII spines, its origin from 2 scales in front to
3 scales behind tips of pelvic fins. Second dorsal fin with I spine and 8 to 10 soft rays. Anal fin with I spine
and 11 to 14 soft rays. Pectoral fins with 15 to 18 soft rays. Body scales large; 38 to 44 midlateral scales;
5 scales in transverse rows along side of body. Predorsal scales 16 to 18, interdorsal scales 7 to 9. Anus from
1 scale in front to 3 scales behind pelvic-fin tips. Colour: live specimens blue-green and translucent; preserved
specimens green-brown, brown, or yellow-green; scales above midlateral band heavily pigmented with edges
outlined with chromatophores; chromatophores often forming pattern resembling incomplete circles along
middorsal line; midlateral band narrow, extending onto caudal fin and forming triangular blotch on upper half
of fin base; body below midlateral band pale; rows of spots often present along third and fourth scale rows
along side of body; fin bases pigmented; fins clear or dusky, pectoral fins frequently with dark blotch.
Size: Maximum length 11 cm.
Habitat, biology, and fisheries: Usually found in coastal waters and harbours within its range. Caught with
seines. Could be of value at subsistence level although not known to be abundant. As with other silversides
and small fishes in general, it can be dried before eating. Can be used as bait fish and is important as forage
species for larger commercial fish.
Distribution: A very widely
distributed species occurring
throughout the Red Sea, In-
dian Ocean, and the South and
Central Pacific Ocean.
Hypoatherina tropicalis (Whitley, 1948)

Frequent synonyms / misidentifications: Atherinosoma tropicalis (Whitley, 1948); Taeniomembras
tropicalis (Whitley, 1948) / Atherinomorus lacunosus (Forster, 1801).
FAO names: En - Whitley’s silverside.
Diagnostic characters: Body slender, slightly compressed. Head almost conical. Mouth large, moderately
oblique to horizontal. Loose flap of skin extending from anterolateral edge of dentary to corner of mouth; apart
from flap of skin, lips absent. Fine teeth present in both jaws, no shagreen denticles on outer surface of jaws.
Teeth usually present on palatines but frequently absent on ectopterygoids; vomerine teeth small but clearly
visible; strong ridge of teeth on mesopterygoids. Tongue with teeth-like papillae in Australian specimens, those
from Madang and Maron islands with distinct teeth; neither teeth nor papillae in specimens from Numa Numa.
Ascending process of premaxilla short and broad, its length usually less than twice its width; lateral process
not well defined. Rami of dentary highly elevated posteriorly, sloping obliquely towards symphysis. Gill rakers
on lower limb of first gill arch 18 to 22, moderately long and slender, but always slightly shorter than diameter
of pupil. First dorsal fin with V to VIII spines, its origin from 1 scale in front to 4 scales behind vertical through
tips of pelvic fins. Second dorsal fin with I spine and 8 to 11 soft rays. Anal fin with I spine and 11 to 14 soft
rays. Pectoral fins with I spine and 15 to 18 soft rays. Body scales dorsoventrally elongated, rounded
posteriorly; 39 to 46 midlateral scales; 5 scales in transverse rows along side of body. Predorsal scales 17
to 22; interdorsal scales 7 to 9. Anus from 1/2 scale in front, to 3 scales behind, tips of pelvic fins. Colour:
live fish translucent with silvery blue sheen; scales above midlateral band frequently heavily pigmented with
inverted C-shaped or diamond-shaped mark present on each predorsal scale; midlateral band series of large
chromatophores with upper fifth darker continuous line; 2 rows of spots present below midlateral band; bases
of fins pigmented, especially pectoral, anal, and second dorsal fins; fins either clear or dusky; caudal fin usually
heavily pigmented.
Habitat, biology, and fisheries: Inhabits waters close inshore. Often seen to jump out the water, especially
after sunset when they appear as fluorescent streaks. At night it appears to aggregate with Atherinomorus
capricornensis around One Tree Island (eastern Australia). Probably eaten by larger commercial species.
Distribution: New Guinea,
n o r t he aster n Australia,
Madang, Maron, and Bougan-
ville islands. Possibly present
over a wide range in Melane-
sian waters. Also found around
Lord Howe Island.
2138 Bony Fishes
Hypoatherina valenciennei (Bleeker, 1853)

Frequent synonyms / misidentifications: Atherina valenciennei Bleeker, 1853; A. bleekeri Günther, 1861;
Allanetta valenciennei (Bleeker, 1853); Pranesus valenciennei (Bleeker, 1853) / Hypoatherina woodwardi
(Jordan and Starks, 1901); Haplocheilus argyrotaenia (Tirant, 1883).
FAO names: En - Sumatran silverside.
Diagnostic characters: Body moderately robust, subcylindrical, and slightly compressed. Free edge of
premaxilla densely covered with minute shagreen denticles; teeth present on vomer, palatines, ectop-
terygoids; a strong ridge of teeth on mesopterygoids; tongue often with teeth around edge. Premaxilla long
and obliquely directed, extending past vertical through anterior margin of orbit. Ascending process of
premaxilla moderately wide and long, lateral process short and wide at base. Coronoid process of dentary
highly elevated but not rounded as in other Hypoatherina species. Gill rakers long and slender, about
diameter of pupil, 20 to 26 on lower limb of first gill arch. First dorsal fin with IV to VII spines, its origin 2 to 5
scales behind vertical through tips of pelvic fins. Second dorsal fin with I spine and 8 to 10 soft rays. Anal fin
with I spine and 10 to 13 soft rays. Pectoral fins with I small spine and 14 to 16 soft rays. Body scales large
and dorsoventrally elongated; 40 to 46 midlateral scales; 5 or 6 scales in transverse rows along side of body.
Predorsal scales 17 to 23; interdorsal scales 6 to 8. Anus 1 to 4 scales in front of tips of pelvic fins. Colour:
live specimens blue-green above, whitish below, and with dark tip to snout; preserved specimens usually
brown with golden sheen; scales probably covered with fine melanophores in life; scales above midlateral
band edged with black; fins clear but with small traces of pigment; head, eye, opercle, and preopercle silvery
gold.
Habitat, biology, and fisheries: Probably important as forage fish. Nothing is known about the biology of
this species.
Distribution: Wide distribu-
tion throughout the southwest-
ern Pacific including Java,
Borneo, Sumatra, Sulawesi,
Viet Nam, Singapore, New
Britain, Hermit Island Group,
Papua New Guinea, Solomon
Islands, Hong Kong, and as far
north as Japan.
Stenatherina panatela (Jordan and Richardson, 1908)

Frequent synonyms / misidentifications: Atherina panatela (Jordan and Richardson, 1908); Hypoatherina
panatela (Jordan and Richardson, 1908) / Stenatherina temminckii (Bleeker, 1853).
FAO names: En - Panatela silverside.
Diagnostic characters: Body slender, slightly compressed. Lips thin, mouth large and extremely protractile
with lips fusing near corners of mouth. Teeth villiform; those on vomer forming characteristic T-pattern.
Palatines and mesopterygoids also with teeth. Free edge of premaxilla convex anteriorly, becoming more or
less straight towards angle of mouth. Ascending process of premaxilla long, extending past orbit into
interorbital space; lateral process very long and pungent. Gill rakers on lower limb of first gill arch 21 to
25, long and slender, longer than 1/2 diameter of pupil. First dorsal fin with IV to VII spines, its origin from
1 scale behind to 2 scales in front of vertical through tips of pelvic fins. Second dorsal fin with I spine and 8 to
10 soft rays. Anal fin with I spine and 9 to 12 soft rays. Pectoral fins with I spine and 16 to 18 soft rays. Body
scales dorsoventrally elongated with ridges restricted to anterior part of scale; 38 to 45 midlateral scales; 5
scales in transverse rows along side of body. Interdorsal scales 8 to 10; predorsal scales 17 to 20. Colour:
live specimens blue-green above and silvery below; distinct crescentic mark present on snout immedi-
ately in front of eye; upper part of midlateral band terminating as half crescent on upper lobe of caudal
fin; preserved specimens yellow below wide and dark midlateral band; upper edge of midlateral band edged
by narrower bluish stripe; scales above midlateral band heavily pigmented, with C-shaped pattern at edge of
each scale; middorsal scales with concentration of pigment forming narrow band from nape and extending
through dorsal fins to caudal fin; bases of pectoral and anal fins dark; caudal fin dusky; opercle and dorsum
of head almost black; distinct black ridge extending from lateral margin of snout to upper edge of orbit.
Habitat, biology, and fisheries: One of the largest and most frequently encountered atherinids in the deeper
lagoon waters of the Marshall Islands; also common in coastal waters, lagoons, atolls, and harbours. May
gather in schools at night when attracted to light but generally not found in very large numbers. Preferred
habitat appears to be deeper enclosed atoll lagoons, reefs, coastal waters, and harbours. May be taken as
food by larger commercial species.
Distribution: W i d e r a n ge
throughout the tropical west-
ern Pacific Ocean. It is known
from the Malaita, Malekula,
and Calayan islands; also Mar-
shall and Phoenix islands,
Philippines, Solomon Islands,
Vanuatu, New Guinea, and
n orthwestern and north-
eastern Australia.
click for next page

2140 Bony Fishes
ISONIDAE
Surf sardines
by W. Ivantsoff
D iagnostic characters: Characterized by highly compressed body, deepest at vertical through of

pectoral-fin origin, tapering rapidly towards caudal fin. Ventral edge of abdomen reduced to
sharp keel. Head small, truncated posteriorly. Snout rounded. Mouth strongly oblique to horizontal, with
premaxilla extending beyond vertical through anterior margin of orbit. Teeth in jaws small but well defined
and curving backward into mouth, those on premaxilla extending outward onto free surface but diminishing
in size. Teeth also present on vomer and palatines. Gill rakers moderately long and well developed, greater
than diameter of pupil. Two dorsal fins present, with first dorsal fin originating about middle of body, second
dorsal fin always originating behind vertical through anal-fin origin. Pectoral fins short and wide, always
set high on body. Body scales cycloid, small, thin, and highly deciduous; area between head and
first dorsal fin, sides of head, and anterior part of abdomen naked. Anus always close to anal fin.
Colour: usually slightly translucent, very bright silvery in life; midlateral band broad and also silvery but
distinctly visible.
pectoral fins high on body
abdomen naked, with sharp keel

Habitat, biology, and fisheries: Frequently found in relatively rough surf along beaches or around rocky
headlands; occasionally also found in tidal inlets and river mouths. Seemingly delicate fishes, unable to
survive where oxygen concentrations are low. There is some evidence that Iso spp. hybridize in some
parts of their range. They have no commercial value but are probably taken as forage by larger commercial
species.
The Isonidae is distinct by the combination of characters in boldface (see above).
Key to the species of Isonidae occurring in the area
1a. Midlateral scales always 42 or more; gill rakers on lower limb of first gill arch 9 to 14
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iso rhothophilus
1b. Midlateral scales always less than 41; gill rakers on lower limb of first gill arch 12 or less . . . . . ® 2
2a. Midlateral scales 34 to 39 (mean 37.4); opercular notch absent on upper part of
operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iso hawaiiensis
2b. Midlateral scales 35 to 41 (mean 38.2); opercular notch present on upper part of
operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Iso nesiotes

The symbol 0
is given when species acounts are included.
0
0
Iso hawaiiensis (Gosline, 1952)
0 Iso nesiotes Saeed, Ivantsoff, and Crowley, 1993
Iso rhothophilus (Ogilby, 1895)
Reference
Ivantsoff, W. 1984. Notocheiridae. In FAO species identification sheets for fishery purposes. Western Indian Ocean
(Fishing Area 51), edited by W. Fischer and G. Bianchi. Vol.3. Rome, FAO (unpaginated).
Atheriniformes: Isonidae 2141
Iso rhothophilus (Ogilby, 1895)

Frequent synonyms / misidentifications: Tropidostethus rhothophilus Ogilby, 1895; Tropidostethops
rhothophilus (Ogilby, 1895) / Iso flosmaris (Jordan and Starks, 1901).
FAO names: En - Surf sardine; Fr - Surfette commune; Sp - Rompeolas.
Diagnostic characters: Body small, laterally compressed. Head short. Dorsal edge of opercle with notch
and small process above it. Mouth small, oblique. Teeth in jaws in single row, covering about half free
edge of premaxilla. Teeth present or absent on vomer, always present on basihyal, but absent on
palatines. Ascending process of premaxilla short and pointed, 2 postmaxillary processes scarcely
developed. Dentary highly elevated. Gill slit present behind last gill arch; gill rakers on lower limb of first
gill arch 9 to 14. First dorsal fin with III to VI small, weak spines, its origin well behind vertical through
tips of pelvic fins. Second dorsal fin with I spine and 10 to 17 soft rays. Anal fin with I spine and 20 to 28
soft rays. Pectoral fins with I spine and 12 to 15 soft rays. Body scales small, cycloid; area between head
and first dorsal fin, sides of head, and abdomen naked. Midlateral scales 42 to 55. Predorsal scales
absent, interdorsal scales 8 to 11. Colour: live fish translucent with side of body silvery; broad silvery
midlateral band terminating in small oval spot before caudal-fin base.
Habitat, biology, and fisheries: Most commonly found in surf around rocky headlands and along the
shore line, occasionally also in the more still waters of river mouths. The fish are very delicate and do not
survive handling. Appears to be able to hybridize with congeners, Iso flosmaris and I. hawaiiensis. Nothing
else is known about this species. Unlikely to have any commercial value.
Distribution: Around the
coasts of Australia, probably
throughout the Pacific Ocean
as it has been collected as far
north as Taiwan Province of
China and Japan.
2142 Bony Fishes
Iso hawaiiensis Gosline, 1952

FAO names: En - Hawaiian surf sardine.
Diagnostic characters: Body small, highly compressed laterally. Dorsal edge of opercle without notch
and process. Mouth small, oblique. Teeth in jaws restricted to first third along free edge of premaxilla;
teeth absent on vomer, palatine, and basihyal. Ascending process of premaxilla short and pointed, dorsal
surface of ramus with slight elevation towards middle. Dentary highly elevated. Gill rakers on lower limb
of first gill arch 9 to 12. First dorsal fin with IV to VI weak spines; second dorsal fin with I spine and 14 to
16 soft rays; anal fin with I spine and 20 to 25 soft rays; pectoral fins with I spine and 12 to 14 soft rays.
Body scales small, deciduous. Midlateral scale count 34 to 39. Predorsal scales absent; interdorsal scales
about 8. Colour: silvery translucent with broad silver midlateral band terminating in oval spot before origin
of caudal fin.
Habitat, biology, and fisheries: Inhabits rough surf and waves around rocky headlands and reefs. Nothing
is known of the biology of this species. It has no commercial value.
Distribution: Presently known
from Hawaii, and the Marshall
and Rapa islands in the Pacific
Ocean.
Atheriniformes: Isonidae 2143
Iso nesiotes Saeed, Ivantsoff, and Crowley, 1993

Frequent synonyms / misidentifications: None / Iso hawaiiensis (Gosline, 1952).
FAO names: En - Samoan surf sardine.
Diagnostic characters: Body highly compressed. Head small, with rounded snout. Dorsal edge of
opercle with notch but no process above it. Mouth small, oblique to horizontal; upper jaw not protractile,
lower jaw deeply elevated posteriorly. Teeth on jaws small, curving backwards into mouth. Teeth restricted
to anterior fifth of free edge of premaxilla. Teeth absent on vomer and palatines, but present on basihyal.
Ascending process of premaxila short and pointed; its ramus with slightly rounded elevation repre-
senting postmaxillary process. Dentary highly elevated. Gill rakers on lower limb of first gill arch 9 to 12,
moderately long, equal to diameter of pupil. First dorsal fin with IV to VI weak spines, its origin about in
line with vertical through tips of pelvic fins. Second dorsal fin with I spine and 13 to 17 soft rays. Anal fin
with I spine and 20 to 25 soft rays; pectoral fins with I spine and 12 to 14 soft rays. Body scales small,
cycloid, and highly deciduous; sides of head and anterior part of abdomen naked. Midlateral scales 35 to
41. Colour: preserved specimens whitish with silvery midlateral band or brownish with darker brown
midlateral band; band ending about half-way along caudal peduncle with no oval spot near base of
caudal fin.
Habitat, biology, and fisheries: Inhabits surf and waves around rocky headlands and reefs. Nothing is
known of the biology of this species. It has no commercial value.
Distribution: Presently known
only from American Samoa
and Pitcairn Island.
2144 Bony Fishes
TELMATHERINIDAE
Sailfin silversides
by W. Ivantsoff
A single species occurring in the area.
Kalyptatherina helodes (Ivantsoff and Allen, 1984)

Frequent syonyms / misidentifications: Pseudomugil helodes Ivantsoff and Allen, 1984 / None.
FAO names: En - Marine sailfin silverside.
Diagnostic characters: Body small, laterally compressed. Eyes large. Mouth slightly protrusible and
oblique, with free edge of premaxilla reaching just past anterior border of orbit. Teeth in upper jaw large,
distribution extending almost to distal end of premaxilla. Teeth in lower jaw curved, villiform, and restricted
to medial third, smaller than those in upper jaw. Teeth frequently present on vomer, palatine, mesoptery-
goid, and basihyal. Ascending process of premaxilla relatively short and broad, lateral process broadly
rounded, about 1/2 height of ascending process. Coronoid process of dentary highly elevated. Gill rakers
on lower limb of first gill arch 16 to 18, moderately long and slender, but less than 1/2 diameter of pupil.
First dorsal fin with III to V weak spines, its origin 1 to 3 scales in front of tips of pelvic fins. Second dorsal
fin lacking spine, with 7 to 9 soft rays. Anal fin with I spine and 12 to 14 soft rays. Pectoral fins with I spine
and 11 or 12 soft rays. Body scales dorsoventrally elongated; 28 to 30 midlateral scales; 5 or 6 scales in
transverse rows along side of body. Predorsal scales 11 to 13; interdorsal scales 4 or 5. Colour: body
translucent with posterior end of swimbladder clearly visible through body wall; preserved specimens
yellow-green; eye black; head dark dorsally; lower edge of eye outlined with melanophores; sides of snout,
chin, and opercle heavily peppered with large and small melanophores; narrow but well defined middorsal
band originating on dorsum of head and extending through bases of dorsal fins to origin of caudal fin; scale
pockets on side of body outlined with melanophores; midlateral band originating as single narrow dark line
at upper edge of pectoral fin and continuing as 2 narrow bands to caudal-fin base.
Pseudomugilidae: Pseudomugil majusculus looks superficially similar to Kalyptatherina helodes, but lacks
a spine in the anal and pectoral fins.
Size: Maximum total length not exceeding 4.5 cm.
Habitat, biology, and fisheries: Inhabits
mangrove swamps close inshore. While
the biology of some telmatherinids may be
known, nothing is known of the biology of
the monotypic Kalyptatherina helodes.
This species is too small to have commer-
cial value but may be taken as food by
young of commercial fish species.
Distribution: Misool and Bantana islands
northwest Irian Jaya.
Reference
Aarn, W. Ivantsoff, and M. Kottelat. (in press). A new telmatherinid genus from Sulawesi, and a phylogenetic analysis
of Telmatherinidae (Teleostei: Atherinomorpha). Ichthyol. Explor. Freshwaters.
Atheriniformes: Dentatherinidae 2145
DENTATHERINIDAE
Tusked silversides
by W. Ivantsoff
Dentatherina merceri Patten and Ivantsoff, 1983

FAO names: En - Mercer’s tusked silverside.
Diagnostic characters: Body very small, slender, subcylindrical; caudal peduncle very slender. Eye
and orbit large. Mouth small. Labial ligament to about half-way along premaxilla forming cylindrical
pouch lateral to ramus of dentary and just below edge of premaxilla. Premaxilla broad and long,
extending just past vertical through anterior border of eye but with anterior two-thirds obscured by anterior
process of maxilla and labial pouch. Premaxilla with ventrally directed tusk-like process. Gill rakers on
lower limb of first gill arch 10 to 12, slender, their length less than 1/2 diameter of pupil. First dorsal fin with
V to VIII spines, its origin 1 to 4 scales in front of vertical through tips of pelvic fins. Second dorsal fin with
I spine and 12 to 14 soft rays; anal fin with I spine and 14 to 16 soft rays, pectoral fins with I spine and 12
to 14 soft rays. Body scales small, thin, and highly deciduous; lateral body scales ellipsoid or irregular; 40
to 43 midlateral scales; 7 to 9 scales in transverse rows along side of body. Predorsal scales 14 to
17; interdorsal scales 8 to 12. Colour: preserved specimens yellow-brown to green-brown; middorsal
band extending over width of 1 scale from head to origin of first dorsal fin, wider at this point with this part
of line frequently broken into 2 or 3 rows of melanophores, then extending as thin irregular line to origin of
caudal fin; caudal fin usually dusky, with 2 triangular marks at its base; midlateral band originating above
upper edge of opercle.
Dentatherina merceri is distinguished from other atheriniform families in the area by the combination of
the characters in boldface (see above).
Size: Maximum length not exceeding 5 cm, but usually much smaller.
i e s : Found close inshore
around islands and over coral
reefs. Except for larval biol-
ogy, little is known about this
species. It has no commercial
value but may be taken by
commercial fishes as food.
D i s t r i b u t i o n : Philippines,
New Guinea, Moluccas, and
other parts of Indonesia; also
around islands of the Barrier
Reef in northeastern Australia.
Reference
Patten, J.M and W. Ivantsoff. 1983. A new genus and species of atherinid fish from western Pacific. Japan. J. Ichthyol.,
29(4):329-339.
2146 Bony Fishes
PHALLOSTETHIDAE
Priapium fishes
by L.R. Parenti
D iagnostic characters: Small (to 3.7 cm standard length), laterally compressed, atherinomorph fishes.
Eyes large. Mouth supraterminal, moderate gape, slightly to highly protrusible. Dentition variously
reduced; small, unicuspid teeth in single row in outer jaw; enlarged teeth on distal portion of premaxilla in
some species. Single row of small, unicuspid teeth on paradentary bone in some species. Pectoral
fins falcate. Caudal fin emarginate, forming incipient lobes. Anus and urogenital openings ante-
rior, under head, may be covered by fleshy hood or papilla in females. Fleshy, rayless keel on
ventral surface of body from point perpendicular to pectoral-fin base to beginning of anal fin. Males
bilaterally asymmetric; elaborate bony, externalized intromittent organ, priapium, under head with
seminal papilla offset to one (aproctal) side of body and anus offset to opposite (proctal) side. First
dorsal fin with I or II short spines or thickened rays, or fin absent; second dorsal fin with 5 to 10 soft
rays; anal fin with 14 to 28 soft rays; pectoral fins with 9 to 13 soft rays; pelvic-fin rays modified into
portion of priapium in males, vestigial or absent in females. Scales cycloid, small to moderate,
deciduous; no lateral line. Colour: nearly translucent in life with orange blotch on caudal peduncle in some
species; scattered minute dark brown to black melanophores on head and body that may be concentrated
at base of priapium.
1st dorsal fin
mouth supraterminal, present or absent 2nd dorsal fin
membranous dome always present
slightly to highly in some species
protrusible
caudal fin
fleshy, rayless ventral keel emarginate
males with elaborate, externalized (after Roberts, 1971a)
intromittent organ under head derived
from pelvic fins, and part of pectoral fins
head and
anterior
portion of
body of
females of Neostethus lankesteri
2 phallostethid anus
fishes urogenital opening
Neostethus bicornis
urogenital opening
anus
(after Parenti, 1989)
Habitat, biology, and fisheries: Fresh to brackish water, species in the area in tidal portions of rivers,
including estuaries, and mangroves. Small, surface-feeding, schooling fishes, may be seen in groups near
shore. Omnivorous, but feed principally on copepods and mollusc larvae. Oviparous, internal fertilization
followed by female laying fertilized eggs.
Atheriniformes: Phallostethidae 2147
Similar families occurring in area

2nd dorsal fin
Adrianichthyidae: readily distinguished from st always present
male priapium fishes by the absence of a bony 1 dorsal fin absent
externalized intromittent organ, urogenital
opening just anterior to anal fin (not under
head), caudal fin truncate, and first dorsal fin
absent; in addition, the mouth in adrianichthy-
ids is small, terminal, and non-protrusible,
whereas it may be highly protrusible in
priapium fishes.
caudal
Aplocheilidae: readily distinguished from male fin
priapium fishes by absence of a bony, exter- no fleshy ventral keel urogenital opening truncate
anterior to anal fin
nalized intromittent organ, with urogenital
Adrianichthyidae
opening just anterior to anal fin, not under
head, caudal fin rounded, no first dorsal fin, (after Iwamatsu et al., 1982)
and black blotch on middle portions of anterior
dorsal-fin rays.
Poeciliidae: livebearers of this family in the area may be confused with priapium fishes because of a
superficially similar intromittent organ which, in male priapium fishes, is under the head and derived from
pectoral and pelvic fins; in male livebearers, the anal-fin rays are modified into an intromittent organ, the
gonopodium; livebearing poeciliids bear live young, while priapium fishes lay fertilized eggs; also,
livebearing poeciliid species may be more brightly coloured and densely pigmented than priapium fishes
which are nearly transparent in life and have sparse pigmentation; finally, livebearers have a rounded or
truncate caudal fin and lack the first dorsal fin.
body and fins may be

densely pigmented 2nd dorsal fin
2nd dorsal fin always present always present caudal fin
truncate or
1st dorsal fin absent st
1 dorsal fin absent lanceolate
no fleshy
ventral keel urogenital opening caudal fin rounded,
anterior to anal fin middle rays males with intromittent
no fleshy organ at midbody
slightly elongate
Aplocheilidae ventral keel Poeciliidae derived from anterior
(after Smith, 1945) (after Rosen, 1973) portion of anal fin
Key to the genera of Phallostethidae

1a. No pulvinulus; outer jaws somewhat protrusible; perforated gular flap of skin through
which anterior end of first ctenactinium may pass; aproctal axial bone projects beyond
ventral body profile (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gulaphallus
1b. Shield-like pulvinulus present (Fig. 2); outer jaws highly protrusible; no perforation of
skin in gular region; aproctal axial bone does not project beyond ventral body profile . . . . . . → 2
2148 Bony Fishes
2a. Main externalized bone of priapium a toxactinium that curves to right or to left under
head of male (Fig. 2); large, fleshy, smooth, or ruffled seminal papilla (Fig. 2); translu-
cent, membranous dome on dorsal surface of head . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Main externalized bone of priapium a ctenactinium; seminal papilla small; no translu-
cent, membranous dome on dorsal surface of head . . . . . . . . . . . . . . . . . . . . Neostethus
large, ruffled
seminal papilla
toxactinium
perforated
gular skin
flap
ctenactinium small, smooth, shield-like
seminal papilla pulvinulus
aproctal axial bone projects
beyond ventral body profile
Fig. 1 Gulaphallus male (lateral view) Fig. 2 Phenacostethus male (ventral view)
(after Parenti, 1989) (after Roberts, 1971a)
3a. Anal-fin rays 26 to 28, vertebrae 40; second ctenactinium serrated; outer jaws equal;
no first dorsal fin; second dorsal-fin rays 8 to 10 . . . . . . . . . . . . . . . . . . . . . Phallostethus
3b. Anal-fin rays 14 or 15; vertebrae 33 to 35; second ctenactinium greatly reduced and not
serrated; lower jaw projects beyond upper jaw; first dorsal fin present, a single ray;
second dorsal-fin rays 5 to 7 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phenacostethus

Gulaphallus panayensis (Herre, 1942)
Neostethus amaricola (Villadolid and Manacop, 1935)
Neostethus bicornis Regan, 1916
Neostethus borneensis Herre, 1939
Neostethus ctenophorus (Aurich, 1937)
Noestethus djajaorum Parenti and Louie, 1998
Neostethus lankesteri Regan, 1916
Noestethus palawanensis (Myers, 1935)
Neostethus robertsi Parenti, 1989
Neostethus thessa (Aurich, 1937)
Neostethus villadolidi Herre, 1942
Neostethus zamboangae Herre, 1942
Phallostethus lehi Parenti, 1996
Phenacostethus smithi Myers, 1928
Phenacostethus trewavasae Parenti, 1986
References
Parenti, L.R. 1989. A phylogenetic revision of the phallostethid fishes (Atherinomorpha, Phallostethidae). Proc. Calif.
Acad. Sci., 46:243-277.
Parenti, L.R. 1996. Phylogenetic systematics and biogeography of phallostethid fishes (Atherinomorpha,
Phallostethidae) of northwestern Borneo, with description of a new species. Copeia, 1996(3):703-712.
Roberts, T.R. 1971a. The fishes of the Malaysian family Phallostethidae (Atheriniformes). Breviora, 374:1-27.
Roberts, T.R. 1971b. Osteology of the Malaysian phallostethid fish Ceratostethus bicornis, with a discussion of the
evolution of remarkable structural novelties, in its jaws and external genitalia. Bull. Mus. Comp. Zool.,
142:393-418.
click for next page

Beloniformes: Adrianichthyidae 2149
Order BELONIFORMES
ADRIANICHTHYIDAE
Ricefishes
by L.R. Parenti
D iagnostic characters: Small to medium-sized, laterally compressed beloniform fishes (to about 18 cm
standard length, but most species mature at under 3 cm standard length). Eyes large. Mouth
terminal, small to large, not protrusible. Small, unicuspid teeth in single or multiple rows on premaxilla
and dentary. Males, and also females of several species in the area may have several enlarged teeth on
distal portion of premaxilla and dentary. Pectoral fins falcate, set relatively high on side of body. Caudal
fin truncate or lanceolate, forming incipient lobes. Single dorsal fin set posteriorly, with 5 to 17 soft rays;
anal fin with 16 to 32 soft rays; caudal fin asymmetrical, with 4 or 5 upper and 5 or 6 lower branched rays;
pectoral fins with 6 to 16 soft rays; pelvic fins with 5 to 7 soft rays. Scales moderate to minute, cycloid, 24
to 85 in lateral series; no lateral line. Dorsal- and anal-fin rays of male longer and thicker than those
of female and often with bony contact organs on distal segments of anal-fin rays. Colour: smaller
ricefish species nearly translucent in life, a black line from dorsal surface of head to dorsal-fin origin, a
midlateral black line from head to base of caudal fin, a black line along anal-fin base, a black submarginal
line on dorsal and on ventral portion of caudal fin, body may be covered with scattered, tiny melanophores
concentrated at pectoral-fin base; males may be more darkly coloured than females.
a single dorsal fin,

set far back on body
caudal fin
mouth small, pectoral fins falcate truncate
non-protrusible
female
no fleshy ventral keel

urogenital opening
(after Iwamatsu et al., 1982)
male
anal-fin rays elongate and may

have bony contact organs
Habitat, biology, and fisheries: Fresh to brackish water, the species listed in the area principally in
brackish water, estuaries, and mangroves. Small to moderate, surface-feeding, schooling fishes. Om-
nivorous, but reported to feed principally on copepods, insects, and mollusc and crustacean larvae.
Oviparous, females may carry a cluster of fertilized eggs near urogenital opening prior to hatching or to
depositing eggs on plants.
2150 Bony Fishes

Aplocheilidae: superficially similar to Oryzias spe-
mouth large, pectoral fins
cies with which it is often confused, but differing most protrusible rounded
notably by having a protrusible mouth, teeth on
vomer, and silvery, “pineal” spot on dorsal surface of
head; a large, black spot on middle portions of dor-
sal-fin rays; and may have thin, black horizontal line
from tip of lower jaw through eye.
Phallostethidae: male phallostethids readily distin-
guished from ricefishes by the presence of a bony, caudal fin rounded,
externalized intromittent organ that may also have a middle rays slightly
large, fleshy seminal papilla. Female phallostethids elongate
superficially similar to ricefishes but with highly pro- Aplocheilidae
trusible mouths and urogenital openings under the (after Smith, 1945)
head.
Poeciliidae: superficially similar to ricefishes, but
with a protrusible rather than non-protrusible mouth
and males with an intromittent organ derived from the
anterior anal-fin rays.
mouth densely pigmented
supraterminal, membranous 1st dorsal fin caudal fin
slightly to dome in present or truncate or
highly some species absent 2nd dorsal fin lanceolate
protrusible always present mouth small,
protrusible
fleshy, rayless
ventral keel
caudal fin
males with elaborate, externalized emarginate
intromittent organ under head derived males with intromittent
from pelvic fins, and part of pectoral fins organ at midbody
derived from anterior
Phallostethidae Poeciliidae portion of anal fin
(after Roberts, 1971a) (after Rosen, 1973)

Oryzias celebensis (Weber, 1894)
Oryzias dancena (Hamilton, 1822)
Oryzias haugiangensis Roberts, 1998
Oryzias hubbsi Roberts, 1998
Oryzias javanicus (Bleeker, 1854)
References
Iwamatsu, T., A. Imaki, A. Kawamoto, and A. Inden. 1982. On Oryzias javanicus collected at Jakarta, Singapore and
West Kalimantan. Ann.Zool. Japon., 55(3):190-198.
Kottelat, M., A.J. Whitten, S.N. Kartikasari, and S. Wirjoatmodjo. 1993. Freshwater fishes of Western Indonesia and
Sulawesi. Periplus Editions (HK) Ltd.
Roberts, T.R. 1998. Systematic observations on tropical Asian medakas or ricefishes of the genus Oryzias, with
descriptions of four new species. Ichthyol.Res., 45(3):213-224.
Rosen, D.E. and L.R. Parenti. 1981. Relationships of Oryzias, and the groups of atherinomorph fishes. Amer. Mus.
Novitat., (2719):25 p.
Beloniformes: Belonidae 2151
BELONIDAE
Needlefishes
by B.B. Collette
D iagnostic characters: Elongate fishes with both upper and lower jaws extended into long beaks
filled with sharp teeth; nostrils in a pit anterior to eyes. No spines in fins; dorsal and anal fins posterior
in position; pelvic fins located in abdominal position and with 6 soft rays; pectoral fins short. Lateral line
running down from pectoral-fin origin and then along ventral margin of body. Scales small, cycloid (smooth),
easily detached. Colour: these fishes live at the surface and are protectively coloured for this mode of life
by being green or blue on back and silvery white on lower sides and belly; usually, a dusky or dark blue
stripe along sides; tip of lower jaw frequently red or orange.
head length caudal peduncle

keel
elongate tooth-studded pelvic fin

jaws
body length
Habitat, biology, and fisheries: Most species are marine, but some occur in fresh water. Carnivorous,
feeding largely on small fishes which they catch sideways in their beaks. Needlefishes tend to leap and
skitter at the surface and some people have been injured when accidentally struck by them, particularly at
night when the fishes are attracted by lights. Caught by casting or trolling surface or near-surface lures.
Flesh excellent in flavour although some people have misgivings about eating it due to the green colour
of the bones. From 1990 to 1995, FAO’s Yearbook of Fishery Statistics reports a range of yearly catch of
needlefishes of around 32 900 to 39 400 t from the Western Central Pacific (Philippines, Indonesia). Some
fresh-water needlefishes reach only 6 to 7 cm in total length while some marine species may attain 2 m.

Hemiramphidae (halfbeaks): 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 (barracudas): jaws pointed but not prolonged into a beak; 2 dorsal fins, the first spiny; pelvic
fins in thoracic position.
.
upper jaw 2 dorsal fins
never prolonged
Hemiramphidae Sphyraenidae
2152 Bony Fishes
Key to the species of Belonidae occurring in the area

Note: out of 10 genera and about 32 species in the family, only 10 marine species belonging to 4 genera
occur in the Western Central Pacific.
1a. Body strongly laterally compressed and marked with a series of vertical bars (Fig. 1);
anal-fin rays 24 to 28 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ablennes hians
1b. Body rounded or squarish in cross-section; no vertical bars present; anal-fin rays 13
to 23 (rarely 24) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Caudal peduncle strongly depressed (flattened dorsoventrally) and with well-devel-
oped lateral keels, least depth of caudal peduncle about 1/2 the width (Fig. 2); gill rakers
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Platybelone argalus platyura
2b. Caudal peduncle not strongly depressed, a small lateral keel on caudal peduncle or no
keel at all, caudal peduncle deeper than wide; gill rakers absent . . . . . . . . . . . . . . . . . ® 3
dorsal fin
lateral
vertical bars keel
depth of
caudal
peduncle
width of
caudal
peduncle
anal fin
cross-section through caudal peduncle
Fig. 1 Ablennes hians Fig. 2 Platybelone argalus platyura
3a. Caudal fin rounded or truncate; no keels on caudal peduncle (Fig. 3); no posterior black
dorsal-fin lobe at any size; dorsal-fin rays 12 to 21 . . . . . . . . . . . . . . . . (Strongylura) ® 4
3b. Caudal fin distinctly forked, with lower lobe longer than upper lobe; narrow raised dark
lateral keel on each side of caudal peduncle (Fig. 4); juveniles with an expanded black
lobe in the posterior part of the dorsal fin (Fig. 5); dorsal-fin rays 19 to 27 . . . . . (Tylosurus) ® 7
caudal fin
no lateral caudal fin forked
not forked lateral
keel keel
Fig. 3 Strongylura Fig. 4 Tylosurus
expanded black lobe
Fig. 5 Tylosurus (juvenile)

4a. Dorsal-fin rays 12 to 15; anal-fin rays 15 to 18; bases of dorsal and anal fins covered
with scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .®5
4b. Dorsal-fin rays 17 to 21; anal-fin rays 21 to 27; bases of dorsal and anal fins without
scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .®6
5a. Prominent black spot at base of caudal fin; predorsal scales 100 to 130 . . Strongylura strongylura
5b. No black spot at base of caudal fin; predorsal scales about 80 . . . . . . . . . Strongylura urvillii
6a. Predorsal scales 100 to 125; dorsal-fin origin over anal-fin rays 4 to 6 . . . . . . Strongylura incisa
6b. Predorsal scales 130 to 180; dorsal-fin origin over anal-fin rays 7 to 10 . . . . . Strongylura leiura
7a. Lower jaw slightly curved upward anteriorly, ending in a thick fleshy formation (Fig. 6);
predorsal scales 165 to 225; no teeth on tongue . . . . . . . . . . . . . . . Tylosurus punctulatus
7b. Lower jaw not curved upward anteriorly, not ending in a thick fleshy formation;
predorsal scales 270 to 340; teeth present on tongue . . . . . . . . . . . . . . . . . . . . . . . ® 8
fleshy formation
Fig. 6 Tylosurus punctulatus
8a. Dorsal-fin rays 24 to 27; anal-fin rays 22 to 24; upper jaw curved producing a gap
between upper and lower jaws . . . . . . . . . . . . . . . . . . . . . . . Tylosurus acus melanotus
8b. Dorsal-fin rays 20 to 25; anal-fin rays 19 to 22; upper jaw straight so there is no
appreciable gap between the jaws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Dorsal-fin rays 21 to 25 (usually 22 or 23); predorsal scales 271 to 340; vertebrae 82
to 86 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tylosurus crocodilus crocodilus
9b. Dorsal-fin rays 20 to 22; predorsal scales 185 to 252; vertebrae 78 to 81 . . Tylosurus gavialoides

0 Ablennes hians (Valenciennes, 1846)
0 Platybelone argalus platyura (Bennett, 1832)
0 Strongylura incisa (Valenciennes, 1846)
0 Strongylura leiura (Bleeker, 1850)
0 Strongylura strongylura (van Hasselt, 1823)
0 Strongylura urvillii (Valenciennes, 1846)
0 Tylosurus acus melanotus (Bleeker, 1850)
0 Tylosurus crocodilus crocodilus (Peron and LeSueur, 1821)
0 Tylosurus gavialoides (Castelnau, 1873)
0 Tylosurus punctulatus (Günther, 1872)
Reference
Parin, N.V. 1967. Review of the marine Belonidae of the western Pacific and Indian oceans. Trudy Inst. Okean., 84:3-83.
[In Russian, English translation in Natl. Mar. Fish. Serv. Syst. Lab. Trans. No.68]
2154 Bony Fishes
Ablennes hians (Valenciennes, 1846)

FAO names: En - Flat needlefish; Fr - Orphie plate; Sp - Agujón sable.
Gulf of Guinea, 41.8 cm body length
(from Collette and Parin, 1970)
Diagnostic characters: Body elongate and greatly compressed laterally. Upper and lower jaws greatly
elongate and studded with small sharp teeth. Gill rakers absent. Anterior parts of dorsal and anal fins
with high falcate lobes; dorsal-fin rays numerous, 23 to 26 (usually 24 or 25); posterior part of dorsal fin
with a prominent dark lobe; anal-fin rays numerous, 24 to 28 (usually 26 or 27); pectoral fins falcate;
pectoral-fin rays 13 to 15; caudal peduncle without lateral keels, caudal fin deeply forked, lower lobe much
longer than upper. Right gonad absent in females; absent or greatly reduced in males. Total number of
vertebrae 87 to 93. Colour: bluish green above, silvery white below; a broad dark blue stripe along sides
and about 12 to 14 prominent dark vertical bars on body; tip of lower jaw red; juveniles and adults
with elevated black lobe in posterior part of dorsal fin; scales and bones green.
Size: Maximum total length at least 120 cm; maximum body length (without head and caudal fin) 90 cm,
commonly to 70 cm.
Habitat, biology, and fisheries: A pelagic species inhabiting offshore surface waters; inshore occurrences
seem to be more frequent around islands than along the mainland coast. Carnivorous, feeding mainly on
small fishes. Caught mainly by casting or trolling surface or near surface lures; also with seines and drift
nets. Marketed mostly fresh.
Distribution: A w o r ld w i d e
species of tropical and warm-
temperate waters. Found off
coasts and islands through-
out the area from southern Ja-
pan through Indonesia to
Australia.
Platybelone argalus platyura (Bennett, 1832)

Frequent synonyms / misidentifications: Belone platyura Bennett, 1832 / None.
FAO names: En - Keeltail needlefish; Fr - Orphie carène; Sp - Agujón de quilla.
Belize, 16.8 cm body length (figure of Platybelone argalus argalus)
(from Collette, 1977)
Diagnostic characters: Body elongate, rounded in cross-section. Upper and especially lower jaws greatly
elongate and studded with fine teeth. Gill rakers present. Anterior parts of dorsal and anal fins not forming
prominent lobes; dorsal-fin rays few, 12 to 15; anal-fin rays 15 to 20; pectoral fin not falcate, pectoral-fin
rays 10 to 12 (usually 11); caudal peduncle greatly depressed with very large lateral keels; caudal fin
forked, upper and lower lobes of about equal length. Predorsal scales (in front of dorsal fin) comparatively
few and large, 108 to 120. Both right and left gonads present, right longer than left. Total vertebrae 62 to
75. Colour: bluish green above, silvery below; a dark blue stripe along sides; fins unpigmented; scales
and bones green.
Size: Maximum standard length (without caudal fin) at least 38.2 cm, commonly to 30 cm; maximum body
length 25.6 cm (without head and caudal fin).
Habitat, biology, and fisheries: A pelagic species inhabiting offshore surface waters but particularly
abundant about islands. Carnivorous, feeding mainly on small fishes. Caught mainly by casting or trolling
surface or near-surface lures; also with seines and trammel nets, often using lights. Marketed fresh.
Distribution: In the Western
Pacific Ocean, known from
many island groups extend-
ing into the Central and South
Pacific. Other subspecies are
found worldwide in tropical
and warm temperate waters.
2156 Bony Fishes
Strongylura incisa (Valenciennes, 1846)

Frequent synonyms / misidentifications: Rhaphiobelone robusta Schultz, 1953; Belone leiuroides
FAO names: En - Reef needlefish.
Australia, 21 cm body length
Diagnostic characters: Body elongate, slightly compressed, almost rectangular in cross-section. Head
large, about 2.5 times in body length; upper and lower jaws greatly elongate and studded with sharp teeth;
upper surface of head with prominent ridges; gill rakers absent. Anterior parts of dorsal and anal fins
forming well-developed lobes; origin of dorsal fin over fourth to sixth ray of anal fin; dorsal-fin rays 18
to 20; anal-fin rays 21 to 24; pectoral fins not falcate; pectoral-fin rays 11 to 13. Caudal peduncle without
lateral keels; caudal fin emarginate, not deeply forked. Predorsal scales (in front of dorsal fin) moderately
large, 100 to 125, crowded just posterior to head; bases of dorsal and anal fins without scales. Colour:
body dark grey with narrow silver lateral stripe, ventral surface white; prominent elongate spot between
opercle and preopercle above level with pectoral-fin origin; cheeks with clumped patches of melanophores;
fins clear with scattered melanophores, pectoral fin with a slight yellow tinge.
Size: Maximum standard length (without caudal fin) 75 cm, commonly to 60 cm; maximum body length
50 cm (without head and caudal fin).
Habitat, biology, and fisheries: Inhabits coastal areas, particularly near coral reefs. Carnivorous, feeding
mainly on small fishes. Taken with drift nets and shore seines. Marketed fresh.
cific from the Maldive and Ni-
cobar islands in the Indian
Ocean through Indonesia,
New Guinea, and northern
Australia; north to the Philip-
pines and South China Sea
and east into Oceania (Sa-
moa, Fiji, Marianas, Marshall,
and Caroline islands).
Strongylura leiura (Bleeker, 1850)

Frequent synonyms / misidentifications: Belone ciconia Richardson, 1846; B. leiurus Bleeker, 1850 /
None.
FAO names: En - Yellowfin needlefish.
Diagnostic characters: Body elongate, laterally compressed, almost rectangular in cross-section. Upper
and lower jaws greatly elongate and studded with sharp teeth; gill rakers absent. Anterior parts of dorsal
and anal fins forming distinct lobes; origin of dorsal fin over seventh to tenth ray of anal fin; dorsal-fin
rays 17 to 21; anal-fin rays 23 to 25; pectoral fins not falcate; pectoral-fin rays 10 or 11. Caudal peduncle
without lateral keels; caudal fin emarginate, not deeply forked. Predorsal scales (in front of dorsal fin)
moderately small, 130 to 180; bases of dorsal and anal fins covered with scales. Colour: top of head and
back greenish; a silver stripe along sides widening posteriorly, lower sides and ventral surface white; pelvic
fins whitish; pectoral fins with a distal dark spot, tip of fins yellow when fresh; tips of dorsal and anal-fin
lobes yellowish, some black pigmentation along middle of fins; caudal fin dark with a yellowish tinge to
upper lobe.
Size: Maximum total length 73 cm; maximum body length (without head and caudal fin) 46 cm, commonly
to 35 cm.
Habitat, biology, and fisheries: Inhabits coastal areas and estuaries. Carnivorous, feeding mainly on
small fishes. Taken with drift nets, shore seines, and purse seines. Marketed fresh.
within the Indo-West Pacific
from South Africa east along
the coasts of Africa, Pakistan,
India, and Sri Lanka to the
western Central Pacific. In the
area, from southeast Asia to
New Guinea, Australia, and
the Philippines.
2158 Bony Fishes
Strongylura strongylura (van Hasselt, 1823)

Frequent synonyms / misidentifications: Tylosurus strongylura van Hasselt, 1823 / None.
FAO names: En - Spottail needlefish; Fr - Aiguillette ocelée; Sp - Agujón ocelado.
Diagnostic characters: Body elongate, rounded in cross-section. Upper and lower jaws greatly
elongate and studded with sharp teeth; gill rakers absent. No spines in fins; anterior parts of dorsal and
anal fins forming moderate lobes; dorsal-fin rays 12 to 15; anal-fin rays 15 to 18; pectoral fins not falcate;
pectoral-fin rays 10 to 12 (usually 11). Caudal peduncle without lateral keels; caudal fin rounded or
truncate, not emarginate or forked. Predorsal scales (in front of dorsal fin) few and relatively large, 100
to 130; bases of dorsal and anal fins covered with scales. Total number of vertebrae 59 to 65. Colour:
greenish above, silvery laterally, white ventrally; pectoral, pelvic, and anal fins light; dorsal and anal fins
with some pigmentation along middle of the rays; caudal fin light with a prominent round black spot
near its base; dorsal-fin lobe and distal margin of caudal fin yellow in live adults, anterior margin of anal
fin orange.
Size: Maximum standard length (without caudal fin) 40 cm, commonly to 22 cm; maximum body length
26 cm (without head and caudal fin).
Habitat, biology, and fisheries: Inhabits coastal areas and mangrove-lined lagoons, where it is common
(especially small specimens) and also enters fresh water. Carnivorous, feeding mainly on small fishes,
especially clupeoids. Caught by casting or trolling surface or near-surface lures; also with seines, often
using lights and with cast nets in mangrove-lined lagoons. Marketed fresh.
cific from the Persian Gulf
along the coasts of Pakistan,
India, and Sri Lanka to the
w e s t er n C e nt r al P a c if i c .
Within the area, north to the
Philippines and southern
China, south to northern Aus-
tralia.
Tylosurus acus melanotus (Bleeker, 1850)

Frequent synonyms / misidentifications: Belone appendiculatus Klunzinger, 1871 / None.
FAO names: En - Agujon needlefish; Fr - Aiguille voyeuse; Sp - Marao ojón (= Aguja imperial).
Puerto Rico, 38.2 cm body length (figure of Tylosurus acus acus)
bony appendage of some

specimens
lateral view of head
Diagnostic characters: Body elongate, rounded in cross-section. Upper and lower jaws greatly elongate
and studded with sharp teeth; a conspicuous appendage sometimes present at tip of lower jaw; gill rakers
absent. Anterior part of dorsal fin with a low lobe, contained 10.5 to 13.3 times in body length;
dorsal-fin rays numerous, 24 to 27; anal-fin lobe low, contained 9.7 to 11.7 times in body length; anal-fin
rays numerous, 22 to 24 (usually 23); pectoral and pelvic fins relatively short, 8 to 12.4 and 10 to 14.1
times in body length, respectively; pectoral-fin rays 13 or 14; a distinct black lateral keel on caudal
peduncle; caudal fin deeply forked, lower lobe much longer than upper. Predorsal scales (in front of
dorsal fin) very numerous and tiny, 280 to 340. Left gonad absent or greatly reduced in both sexes. Total
number of vertebrae 90 to 95. Colour: dark bluish above, silvery white below; juveniles with elevated
black lobe in posterior part of dorsal fin which is lost with growth.
Size: Maximum total length 90 cm; maximum body length (without head and caudal fin) 60 cm, commonly
to 50 cm.
Habitat, biology, and fisheries: Pelagic, inhabiting more offshore waters than Tylosurus crocodilus, but
also found in coastal waters. Carnivorous, feeding mainly on small fishes. Caught by casting or trolling
surface or near-surface lures; also with seines and gill nets, often using lights. Marketed mostly fresh. The
flesh is of good quality.
Distribution: Worldwide in
tropical and warm temperate
waters. This subspecies ex-
t e nd s fr o m S o ut h A f r i c a
t hr o ug h t he C en tral and
South Pacific. Other subspe-
cies are found in the Atlantic
and eastern Pacific.
2160 Bony Fishes
Tylosurus crocodilus crocodilus (Peron and LeSueur, 1821)

Frequent synonyms / misidentifications: Tylosurus raphidoma (Ranzani, 1842) / None.
FAO names: En - Hound needlefish; Fr - Aiguille crocodile; Sp - Marao lisero.
Jamaica, 48 cm body length (from Collette, 1977)
teeth point forward
head of juvenile (32 cm body length)
Diagnostic characters: Body elongate, rounded in cross-section. Upper and lower jaws greatly elongate
and studded with sharp teeth jaw teeth point anteriorly in juveniles (to 40 cm body length) but are straight
at all sizes in other species of Tylosurus; gill rakers absent. Anterior part of dorsal and anal fins with
relatively high lobes, contained 5.4 to 10.6 and 5.5 to 8 times in body length, respectively; dorsal-fin rays
21 to 25 (usually 22 or 23); anal-fin rays 19 to 22 (usually 21 or 22); pectoral and pelvic fins long, contained
6.6 to 8.3 and 7.3 to 10.6 times in body length, respectively; pectoral-fin rays 13 to 15 (usually 14 or 15);
a distinct black lateral keel on caudal peduncle; caudal fin deeply forked, lower lobe much longer
than upper. Predorsal scales numerous and tiny, 271 to 340. Both right and left gonads present, right longer
than left. Total number of vertebrae 82 to 86. Colour: dark bluish green above, silvery below; a dark blue
stripe along sides; juveniles (to 20 cm body length) with elevated black lobe in posterior part of dorsal
fin which is lost with growth; scales and bones green.
Size: Maximum standard length (without caudal fin) at least 124 cm, commonly to 90 cm; maximum body
length (without head and caudal fin) 82.5 cm; unverified reports give up to 150 cm total length.
Habitat, biology, and fisheries: A pelagic species inhabiting more coastal waters than Tylosurus acus,
but also found in offshore waters. Carnivorous, feeding mainly on small fishes. Large individuals may be
dangerous when leaping out of the water. Caught by casting or trolling surface or near-surface lures; also
with purse seines and drift nets. Marketed mostly fresh.
Distribution: A w o r ld w i d e
species in tropical and warm-
temperate waters (replaced
by T. crocodilus fodiator in
the eastern Pacific). Through-
out the area, from southern
Japan through the Philippines
and Indonesia to northern
Australia.
Strongylura urvillii (Valenciennes, 1846)

En - Urville’s needlefish.
Maximum body length 26.3 cm; maximum standard length 43.8 cm, commonly to 35 cm. A coastal
species, sometimes entering brackish water. Found only in the waters of Indonesia, Philippines,
and New Guinea, and east to the Solomon and Santa Cruz islands.
(after Bleeker, 1866-72)
Tylosurus gavialoides (Castelnau, 1873)

En - Stour longtom.
Maximum body length 75 cm; maximum standard length 10.5 cm, commonly to 80 cm. A coastal
species. Restricted to the northern three-quarters of Australia.
Australia, 45 cm body length
Tylosurus punctulatus (Günther, 1872)

En - Spongyjawed needlefish.
Maximum body length 39.5 cm; maximum standard length 58.3 cm, commonly to 50 cm. A coastal
species. Restricted to the western Central Pacific, north to the Philippines and east to the Solomon
islands.
Philippines, 34 cm body length
click for next page

2162 Bony fishes
EXOCOETIDAE
Flyingfishes
by N.V. Parin
D iagnostic characters: Elongate fishes, their bodies broadly cylindrical (round or elliptical in cross-section),
flattened ventrally in some species. Head short. Snout blunt, shorter than eye in all Western Central Pacific
species. Mouth small. Jaws of equal size. Jaw teeth absent or very small. Gill rakers well developed. Upper
pharyngeal bones of third gill arches close together, but not fused into a single plate. No spines in fins. Dorsal
and anal fins set equally far back on body, their bases short and opposed. Pectoral fins high on sides,
strikingly long, always extending beyond dorsal-fin origin. Pelvic fins abdominal in position, and greatly
enlarged in many, but not all, species. Caudal fin deeply forked, its lower lobe longer than the upper. Lateral
line low on body. Scales large, cycloid (smooth to touch), easily shed. Swimbladder large, extending posteriorly
beyond body cavity. Young stages (to about 10 cm) quite different in appearance from adults, with pectoral fins
shorter, dorsal fin often higher than in adults, colour patterns variable, and spots and bars often developed;
single or paired chin barbels conspicuous in many species. Colour: dark above, pale below; dark colours
usually iridescent blue or green in life; pectoral fins in some species with dark spots or pale stripes; dorsal fin
in some species with black pigment.
pectoral fins high on

sides, greatly enlarged
lower lobe of
caudal fin
longer than
upper lobe
pelvic fins abdominal,

lateral line low enlarged in some species
Habitat, biology, and fisheries: Inhabit surface waters of the open ocean as well as neritic and inshore areas.
Well known for their habit of leaping out of the water and gliding over long distances. Schooling, do not undertake
extensive migrations. Feed on zooplankton, larger species also on small fishes. Very abundant in most tropical
seas but are objects of only small-scale fisheries in Philippines, Indonesia, Viet Nam, Thailand, Kiribati, and
some other regions of the world. From 1990 to 1995, FAO’s Yearbook of Fishery Statistics reports a range of
yearly catch of Exocoetidae (and Hemiramphidae) of around 25 900 to 67 200 t from the Western Central Pacific.
Main fishing gears are traps, gill nets, and purse seines. Marketed mostly fresh and dried salted, also as
fermented fish paste.

Hemiramphidae: pectoral fins short to medium length, never reaching dorsal-fin origin; lower jaw much
longer than upper jaw, except in adult Oxyporhamphus, Melapedalion, and Arrhamphus; body more
elongate, except in Oxyporhamphus ; upper pharyngeals of third arch fused, forming a single plate;
swimbladder not extending posteriorly beyond body cavity.
Hemiramphus Oxyporhamphus
(a typical representative) (a representative without prolonged lower jaw)
Hemiramphidae
Beloniformes: Exocoetidae 2163
Key to the genera and species of Exocoetidae occurring in the area

1a. Pectoral fins moderately long, not reaching beyond posterior part of anal-fin base;
pectoral branch of lateral line present (Fig. 1); upper jaw protrusible (Fig. 2) . . (Parexocoetus) ® 2
1b. Pectoral fins very long, reaching beyond anal-fin base to, or almost to, caudal-fin base;
pectoral branch of lateral line absent; upper jaw not protrusible . . . . . . . . . . . . . . . . . . ® 3
protrusible
pectoral branch
mouth opened mouth closed

Fig. 1 Parexocoetus Fig. 2 Parexocoetus
2a. Anal-fin rays 12 to14; longest dorsal-fin rays reaching beyond origin of upper caudal-fin
lobe (Fig. 3a); predorsal scales 20 to 24 . . . . . . . . . . . . . . . . . Parexocoetus brachypterus
2b. Anal-fin rays 10 to 12; longest dorsal-fin rays scarcely reaching origin of upper caudal-fin
lobe (Fig. 3b); predorsal scales 16 to 21 . . . . . . . . . . . . . . . . . . . . . Parexocoetus mento
a) Parexocoetus brachypterus b) Parexocoetus mento

Fig. 3
3a. Pelvic fins short, not reaching anal-fin origin (barely reaching in juveniles), inserted
nearer to pectoral-fin insertion than to anal-fin origin (Fig. 4) . . . . . . . . . . . . (Exocoetus) ® 4
3b. Pelvic fins long, reaching well beyond anal-fin origin, inserted nearer to anal-fin origin
than to pectoral-fin insertion (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
4a. Gill rakers on first gill arch 29 to 37 (usually 32 to 34); as a rule 6 transverse rows of
scales between dorsal-fin origin and lateral line (Fig. 4b) . . . . . . . . . . . . . Exocoetus volitans
4b. Gill rakers on first gill arch 21 to 29 (usually 24 to 27); as a rule 7 transverse rows of
scales between dorsal-fin origin and lateral line (Fig. 4a) . . . . . . . . . . . . . . . . . . . . .®5
7 scale rows 6 scale rows

a) Exocoetus monocirrhus b) Exocoetus volitans
Fig. 4
2164 Bony fishes
5a. Juveniles less than 10 cm standard length with a single chin barbel (Fig. 5a); distributed
north of 10°S latitude . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Exocoetus monocirrhus
5b. Juveniles unbarbelled (Fig. 5b); distributed south of 10°S latitude . . . . . Exocoetus obtusirostris
a) Exocoetus monocirrhus b) Exocoetus obtusirostris

Fig. 5 juveniles
6a. Origin of anal fin slightly before, under or not more than 2 rays behind origin of dorsal
fin (Fig. 6b); dorsal fin usually with less, or equal number of rays than anal fin; juveniles
not barbelled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Hirundichthys) ® 29
6b. Origin of anal fin 3 rays or more behind origin of dorsal fin (Fig. 6a); dorsal fin usually
with 2 to 5 rays more than anal fin; juveniles barbelled or not barbelled . . . . . . . . . . . . . ® 7
a) Prognichthys, Cypselurus, Cheilopogon b) Hirundichthys

Fig. 6
7a. First 3 or 4 pectoral-fin rays unbranched; juveniles not barbelled . . . . . . . . (Prognichthys) ® 8

7b. First 1 or 2 pectoral-fin rays unbranched (Fig. 7a, b); juveniles barbelled or not barbelled . . . . . ® 9
8a. First 3 pectoral-fin rays unbranched (Fig. 7c) . . . . . . . . . . . . . . . . Prognichthys brevipinnis

8b. First 4 pectoral-fin rays unbranched (Fig. 7d) . . . . . . . . . . . . . . . . . . . Prognichthys sealei
a) Cheilopogon rapanouiensis b) Cypselurus, Cheilopogon c) Prognichthys brevipinnis d) Prognichthys sealei
Fig. 7
9a. Lower jaw a little shorter than upper jaw and included beneath the latter (Fig. 8a);
juveniles with a single barbel or not barbelled . . . . . . . . . . . . . . . . . . . (Cypselurus) ® 10
9b. Both jaws of equal length, or lower jaw a little longer than upper jaw (Fig. 8b, c); juveniles
with paired barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Cheilopogon) ® 16
a) Cypselurus b) Cheilopogon c) Cheilopogon

10a. Jaw teeth not tricuspid (Fig. 9a);

juveniles not barbelled, with dark
cross bars on body . . . . Cypselurus hexazona
10b. At least some of jaw teeth tricuspid
(Fig. 9b); juveniles barbelled or not a) Cypselurus hexazona b) other Cypselurus spp.
barbelled, without dark cross bars on
Fig. 9 jaw teeth
body . . . . . . . . . . . . . . . . . . . . ® 11
11a. Pelvic fins inserted much nearer to head than origin of lower caudal-fin lobe (distance
between pelvic fins and caudal fin 0.7 to 0.9 times in distance between head and pelvic
fins) (Fig. 10a); pectoral fins spotted or not spotted; juveniles without barbels . . . . . . . . . ® 12
11b. Pelvic fins inserted about midway between head and origin of lower caudal-fin lobe, or
nearer to the latter (distance between pelvic fins and caudal fin 0.85 to 1.3 times in
distance between head and pelvic fins) (Fig. 10 b, c); pectoral fins not spotted; juveniles
with single barbel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
a) Cypselurus poecilopterus b) Cypselurus naresii c) Cypselurus opisthopus

Fig. 10
12a. Pectoral fins without dark spots, their tip light (Fig. 11a) . . . . . . . . . . . . Cypselurus oligolepis
12b. Pectoral fins with dark spots, their tip not light (Fig. 11b, c) . . . . . . . . . . . . . . . . . . . ® 13
13a. Predorsal scales 28 to 34; pectoral fins with varying number of dark spots not arranged
in regular transverse bands (Fig. 11b) . . . . . . . . . . . . . . . . . . . . . . . . Cypselurus simus
13b. Predorsal scales 24 to 28; pectoral fins with numerous dark spots arranged in regular
transverse bands (Fig. 11c) . . . . . . . . . . . . . . . . . . . . . . . . . . Cypselurus poecilopterus
a) Cypselurus oligolepis b) Cypselurus simus c) Cypselurus poecilopterus

Fig. 11
2166 Bony fishes
14a. Pelvic fins inserted much nearer origin of lower caudal-fin lobe than head (distance
between pelvic fins and caudal fin 1.1 to 1.3 times in distance between head and pelvic
fins) (Fig. 10c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cypselurus opisthopus
14b. Pelvic fins inserted about midway between head and origin of lower caudal-fin lobe
(distance between pelvic fins and caudal fin 0.85 to 1.1 times in distance between head
and pelvic fins) (Fig. 10b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
15a. Predorsal scales 28 to 32; dorsal-fin rays 10 to 12; head 0.95 to 1.1 times in origin of
dorsal fin to origin of upper caudal-fin lobe distance . . . . . . . . . . . . . . . .Cypselurus naresii
15b. Predorsal scales 24 to 30; dorsal-fin rays 12 to 14; head 1.1 to 1.25 times in origin of
dorsal fin to origin of upper caudal-fin lobe distance . . . . . . . . . . . . . Cypselurus angusticeps
16a. Dorsal fin moderately high (longest

ray usually less than 10 times in
standard length), often bearing grey
or black spot (Fig. 12a); jaw teeth
noticeable (conspicuous to the
touch); juveniles with a pair of short
or long, rounded or tapering barbels . . . . . ® 17
16b. Dorsal fin moderately low (longest
ray usually more than 10 times in
standard length), without black or
grey spot (Fig. 12b) (may be present a) Cheilopogon b) Cheilopogon
i n Cheilopogon intermedius); jaw spilonotopterus furcatus
teeth minute (not conspicuous to the
touch); juveniles with a pair of short
(long in Cheilopogon antoncichi) ta- Fig. 12 dorsal fin
pering barbels . . . . . . . . . . . . . . . . ® 25
17a. First 2 pectoral-fin rays unbranched (Fig. 7a); juveniles with a pair of short and flattened
rounded barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon rapanouiensis
17b. Only the first pectoral-fin ray unbranched (Fig. 7b) . . . . . . . . . . . . . . . . . . . . . . . . ® 18
18a. Pectoral fins without dark spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19

18b. Pectoral fins with varying number of small dark spots (Fig. 13a); juveniles with a pair of
elongated and flattened rounded barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
19a. Pectoral fins without pale cross band (Fig. 13c); predorsal scales 28 to 41; juveniles
with a pair of very long tapering barbels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
19b. Pectoral fins with pale oblique cross band (Fig. 13b); predorsal scales 23 to 29; juveniles
with a pair of short tapering barbels bearing proximal flap-like appendage . . . . . . . . . . . ® 21
a) Cheilopogon suttoni b) Cheilopogon dorsomacula c) Cheilopogon cyanopterus

Fig. 13 pectoral fin
20a. Predorsal scales 28 to 35; pectoral fins dark brown . . . . . . . . . . Cheilopogon spilonotopterus
20b. Predorsal scales 33 to 41; pectoral fins dark blue . . . . . . . . . . . . . Cheilopogon cyanopterus
21a. Pelvic fins with a prominent black spot; cross band of pectoral fins usually yellow . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon abei
21b. Pelvic fins without black spot; cross band of pectoral fins not yellow . . . . . . . . . . . . . . . ® 22
22a. Dorsal fin usually with a greyish or blackish spot; predorsal scales 25 to 29 . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon dorsomacula
22b. Dorsal fin uniformly pale; predorsal scales 23 to 26 . . . . . . . . . . . . . . Cheilopogon katoptron
23a. Predorsal scales 28 to 34; dorsal fin usually without black spot . . . . . . . Cheilopogon spilopterus
23b. Predorsal scales 33 to 42; dorsal fin with black spot . . . . . . . . . . . . . . . . . . . . . . . ® 24
24a. Pelvic fins inserted nearer head than origin of lower caudal-fin lobe; head shorter than
distance from dorsal-fin origin to origin of upper caudal-fin lobe (Fig. 14a) . . . Cheilopogon atrisignis
24b. Pelvic fins inserted about midway between head and origin of lower caudal-fin lobe; head
subequal to distance from dorsal-fin origin to origin of upper caudal-fin lobe (Fig. 4b) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cheilopogon suttoni
a) Cheilopogon atrisignis b) Cheilopogon suttoni

Fig. 14
25a. Jaws subequal (Fig. 8b); palatine teeth usually present; pelvic fins inserted about
midway between head and origin of lower caudal-fin lobe . . . . . . . . . Cheilopogon intermedius
25b. Lower jaw a little longer than upper jaw (Fig. 8c); no palatine teeth; pelvic fins inserted
nearer head than origin of lower caudal-fin lobe . . . . . . . . . . . . . . . . . . . . . . . . . . ® 26
26a. Pectoral fins usually with varying number of scattered dark dots; 8 or 9 transverse rows
of scales between dorsal-fin origin and lateral line . . . . . . . . . . . . Cheilopogon pitcairnensis
26b. Pectoral fins without dark dots; 7 or 8 transverse rows of scales between dorsal-fin origin
and lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 27
27a. Anal-fin rays 8 or 9; predorsal scales 26 to 29 . . . . . . . . . . . . . . . . . Cheilopogon arcticeps

27b. Anal-fin rays 9 to 11; predorsal scales usually 29 to 33 . . . . . . . . . . . . . . . . . . . . . ® 28
28a. Pectoral fins light or dark grey

with pale oblique cross band
(Fig. 15a) . . . . . . Cheilopogon furcatus
28b. P e c t o r a l f i n s t r a n s p a r e n t
(Fig. 15b) . . . . . . Cheilopogon unicolor
a) Cheilopogon furcatus b) Cheilopogon unicolor

2168 Bony fishes
29a. First 2 pectoral-fin rays unbranched . . . . . . . . . . . . . . . . . . Hirundichthys albimaculatus

29b. Only the first pectoral-fin ray unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 30
30a. Palatine teeth present; pecto-

ral fin with distinct pale oblique
cross band (Fig. 16a) . . . . .
. . . . . . . . . . Hirundichthys speculiger
30b. No palatine teeth; pectoral fin
without distinct cross band
(Fig. 16b) . . . Hirundichthys oxycephalus
a) Hirundichthys speculiger b) Hirundichthys oxycephalus

0
0
0
Cheilopogon abei Parin, 1996
Cheilopogon arcticeps (Günther, 1866)
0 Cheilopogon atrisignis (Jenkins, 1904)
0
Cheilopogon cyanopterus (Valenciennes, 1846)
0
Cheilopogon dorsomacula (Fowler, 1944)
0
Cheilopogon furcatus (Mitchill, 1815)
0
Cheilopogon intermedius Parin, 1961
0
0
Cheilopogon katoptron (Bleeker, 1866)
Cheilopogon pitcairnensis (Nichols and Breder, 1935)
0
Cheilopogon rapanouiensis Parin, 1961
0
Cheilopogon spilonotopterus (Bleeker, 1866)
0
Cheilopogon spilopterus (Valenciennes, 1846)
0
Cheilopogon suttoni (Whitley and Colefax, 1938)
Cheilopogon unicolor (Valenciennes, 1846)
0
0
Cypselurus angusticeps Nichols and Breder, 1935
0
Cypselurus hexazona (Bleeker, 1853)
0
Cypselurus naresii (Günther, 1889)
0
Cypselurus oligolepis (Bleeker, 1866)
0
Cypselurus opisthopus (Bleeker, 1866)
0
Cypselurus poecilopterus (Valenciennes, 1846)
Cypselurus simus (Valenciennes, 1846)
0
0
Exocoetus monocirrhus Richardson, 1846
0
Exocoetus obtusirostris Günther, 1866
Exocoetus volitans Linnaeus, 1758
0
0
Hirundichthys albimaculatus (Fowler, 1934)
0
Hirundichthys oxycephalus (Bleeker, 1852)
Hirundichthys speculiger (Valenciennes, 1846)
0
0
Parexocoetus brachypterus (Richardson, 1846)
Parexocoetus mento (Valenciennes, 1846)
0
0
Prognichthys brevipinnis (Valenciennes, 1846)
Prognichthys sealei Abe, 1955
Reference
Parin, N.V. 1996. On the flying fish species (Exocoetidae) in the western and central pacific Ocean. Voprosy Ikhtiologii,
36(3):300-307 (English transl. in J. Ichthyol., 36(5):357-364).
Cheilopogon abei Parin, 1996

En - Abe’s flyingfish.
Maximum standard length about 22 cm. Pelagic in neritic surface waters. No importance in fisheries.
Distributed in the northern Indian and western Pacific oceans from East Africa to Solomon Islands.
(after Parin, 1996)
Cheilopogon arcticeps (Günther, 1866)

En - Bearhead flyingfish.
Maximum standard length about 21 cm. Pelagic in nearshore and neritic surface waters, never
spread to open sea. No importance in fisheries. Distributed in the western Pacific Ocean (southern
China, Viet Nam, Thailand, Indonesia, New Guinea, Solomon Islands).
(after Parin, 1961)
Cheilopogon atrisignis (Jenkins, 1904)

En - Glider flyingfish; Fr - Exocet planeur; Sp - Volador planeador.
Maximum standard length about 33 cm. Pelagic in open ocean and neritic surface waters. Minor
importance to fisheries in Polynesia. Distributed in the Indian and Pacific oceans from Somalia and
Madagascar to Revillagigedos and Galapagos islands in the eastern Pacific.
(after Jordan and Evermann, 1905)

2170 Bony fishes
Cheilopogon cyanopterus (Valenciennes, 1846)

En - Margined flyingfish; Fr - Exocet codène; Sp - Volador bordiblanco.
Maximum standard length about 34 cm. Pelagic in open ocean and neritic surface waters. No
importance in fisheries. Distributed in the Atlantic, Indian, and western Pacific oceans eastward to
southern Japan, Taiwan Province of China, Queensland (Australia), and the Mariana, Caroline, and
Solomon islands.
(after Bruun, 1935)
Cheilopogon dorsomacula (Fowler, 1944)

En - Backspot flyingfish; Fr - Exocet à dos tacheté; Sp - Volador de dorso manchado.
Maximum standard length about 23 cm. Pelagic in open ocean surface waters. No importance in
fisheries. Distributed in the tropical Pacific Ocean, not found in inland seas of Southeast Asia.
(drawn by G. Pokhilskaya)
Cheilopogon furcatus (Mitchill, 1815)

En - Spotfin flyingfish; Fr - Exocet tacheté; Sp - Volador manchado.
fisheries. Widespread in tropical zone of all oceans, absent in inland seas of Southeast Asia.
(after Bruun, 1935)

Cheilopogo intermedius Parin, 1961

En - Intermediate flyingfish.
Maximum standard length about 22 cm. Pelagic in nearshore and neritic surface waters. Minor
importance in fisheries (Thailand). Distributed in the Indian and western Pacific oceans from western
India to Palau and Solomon islands.
(after Parin, 1961)
Cheilopogon katoptron (Bleeker, 1866)

En - Indonesian flyingfish.
Maximum standard length about 18 cm. Pelagic in nearshore surface waters, never spread to open
sea. Minor importance in fisheries in Viet Nam and Philippines. Distributed in the western Pacific
Ocean (Viet Nam, Thailand, Indonesia, Philippines, and northern Australia).
(after Parin and Besednov, 1965)
Cheilopogon pitcairnensis (Nichols and Breder, 1935)

En - Pitcairn flyingfish.
importance to fisheries in Polynesia. Distributed in southern tropical Pacific from Tonga to Pitcairn
Islands.
(after Nichols and Breder, 1935)

2172 Bony fishes
Cheilopogon rapanouiensis Parin, 1962

En - Rapanoui flyingfish.
Maximum standard length about 30 cm. Pelagic in neritic surface waters. No importance in fisheries.
Distributed in southern tropical Pacific Ocean from New Caledonia, Fiji, and Easter Island; a single
stray collected in eastern Pacific north of the equator.
(after Parin, 1961)
Cheilopogon spilonotopterus (Bleeker, 1866)

En - Stained flyingfish; Fr - Exocet marbré; Sp - Volador jaspeado.
importance to fisheries in Polynesia. Distributed in the Indian and Pacific oceans eastward to
Revillagigedos, Malpelo, and Galapagos islands.
(after Parin, 1960)
Cheilopogon spilopterus (Valenciennes, 1846)

En - Manyspotted flyingfish.
Maximum standard length to about 25 cm. Pelagic in nearshore and neritic surface waters. Minor
importance in fisheries (Thailand). Distributed in the eastern Indian and western Pacific oceans from
Andaman Sea to Samoa.
(from Bleeker, 1866-72)

Cheilopogon suttoni (Whitley and Colefax, 1938)

En - Sutton’s flyingfish; Fr - Exocet de Sutton; Sp - Volador de Sutton.
importance to fisheries in Polynesia. Distributed in the Indian and western Pacific oceans from Gulf
of Aden to 140°E.
(after Parin, 1960)
Cheilopogon unicolor (Valenciennes, 1846)

En - Limpidwing flyingfish.
importance in fisheries (Polynesia). Distributed in the Pacific Ocean from southern Japan, Philip-
pines, and Queensland (Australia) to Hawaii and Tahiti.
(after Abe, 1953-56)
Cypselurus angusticeps Nichols and Breder, 1935

En - Narrowhead flyingfish; Fr - Exocet bécune; Sp - Volador picudo.
Maximum standard length about 24 cm. Pelagic in open ocean and neritic surface waters, most
abundant off oceanic islands. Minor object of fisheries in Polynesia, caught by dip net and lights at
night. Widespread in the Indian and Pacific oceans, from East Africa to Hawaii and Tuamotu Islands;
separate population in the eastern tropical Pacific at Soccora Island (Mexico).
(after Parin, 1961)
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2174 Bony fishes
Cypselurus hexazona (Bleeker, 1853)

En - Darkbar flyingfish.
sea. No importance to fisheries. Distributed in the Indian and western Pacific oceans from the Red
Sea to Philippines, New Guinea, and Queensland (Australia).
Cypselurus naresii (Günther, 1889)

En - Pharao flyingfish; Fr - Exocet pharaon; Sp - Volador farsón.
importance to fisheries (Viet Nam), caught by gill nets. Distributed in the Indian and western Pacific
oceans from Gulf of Aden and Madagascar to southern Japan, New South Wales (Australia), and
Fiji.
(after Parin, 1961)
Cypselurus oligolepis (Bleeker, 1866)

En - Largescale flyingfish; Fr - Exocet à grandes écailles; Sp - Volador escamudo.
sea. Limited importance to fisheries in Thailand, Indonesia, and Philippines, caught by purse seines,
traps, and gill nets. Widespread in the tropical Indian and western Pacific oceans from East Africa
to southern China, Solomon islands, and Queensland (Australia).

Cypselurus opisthopus (Bleeker, 1866)

En - Rearfin flyingfish.
Maximum standard length about 18 cm. Pelagic in nearshore surface waters, juveniles spread to
open sea with floating seaweed. Minor importance to fisheries in Philippines and, probably,
Indonesia. Distributed in the eastern Indian and western Pacific oceans from Bay of Bengal to
Okinawa (Japan), and Solomon Islands.
Cypselurus poecilopterus (Valenciennes, 1846)

En - Yellowing flyingfish; Fr - Exocet aile jaune; Sp - Volador de ala amarilla.
Maximum standard length about 21 cm. Pelagic, mostly in neritic surface waters. Minor importance
to fisheries in Viet Nam, Thailand, and Indonesia. Widespread in the tropical Indian and western
Pacific oceans from East Africa to southern Japan and Queensland (Australia).
Cypselurus simus (Valenciennes, 1846)

En - Ficklespotted flyingfish.
Maximum standard length about 26 cm. Pelagic in neritic surface waters, most abundant off oceanic
islands. Minor importance to fisheries in Polynesia. Distributed in central Pacific Ocean (Hawaii,
Johnston Island, Tonga, Samoa, Rarotonga, Tahiti, Marquesas and Easter islands). Southern
“population” may represent a closely related undescribed species.
(after Parin, 1961)

2176 Bony fishes
Exocoetus monocirrhus (Richardson, 1846)

En - Barbel flyingfish; Fr - Exocet barbu; Sp - Volador barbudo.
importance to fisheries, caught by gill nets (Viet Nam). Widespread in the Indian and Pacific oceans
from East Africa to Central America.
Exocoetus obtusirostris (Günther, 1866)

En - Oceanic two-wing flyingfish; Fr - Exocet bouledogue; Sp - Volador ñato.
Maximum standard length about 20 cm. Pelagic in open ocean surface waters. No importance to
fisheries. Distributed in the Atlantic Ocean and southern subtropical Pacific, between 10° and 30°S
latitude, from Australia to Peru.
(after Bruun, 1935)
Exocoetus volitans Linnaeus, 1758

En - Tropical two-wing flyingfish; Fr - Exocet volant; Sp - Volador.
Maximum standard length about 18 cm. Pelagic in open ocean surface waters. The most abundant
flying fish of the open sea but has no importance to fisheries. Widespread in tropical and subtropical
zone of all oceans, probably absent in the inland seas of Southeast Asia.
(after Bruun, 1935)

Hirundichthys albimaculatus (Fowler, 1934)

En - Whitespot flyingfish; Fr - Exocet à tache blanche; Sp - Volador de mancha blanca.
fisheries. Distributed in central Pacific Ocean.
Hirundichthys oxycephalus (Bleeker, 1852)

En - Bony flyingfish; Fr - Exocet casque; Sp - Volador casquete.
importance to fisheries in Viet Nam, Indonesia, and the Philippines. Distributed in the Indian and western
Pacific oceans from the Arabian Sea to southern Solomon Islands, and New South Wales (Australia).
(after Abe, 1953-1956)
Hirundichthys speculiger (Valenciennes, 1846)

En - Mirrorwing flyingfish; Fr - Exocet miroir; Sp - Volador espejo.
fisheries. Widespread in tropical zone of all oceans, probably absent in the inland seas of Southeast
Asia.

2178 Bony fishes
Parexocoetus brachypterus (Richardson, 1846)

En - Sailfin flyingfish; Fr - Exocet voilier; Sp - Volador aletón.
Maximum standard length about 14 cm. Pelagic in neritic surface waters, rare in the open ocean.
Not important as food fish, but occasionally taken by fisheries. Widespread in the Indian and Pacific
oceans from East Africa to southern Japan, Hawaii, Queensland (Australia), and Marquesas
Islands; separate population in the eastern tropical Pacific (Bay of Panama).
Parexocoetus mento (Valenciennes, 1846)

En - African sailfin flyingfish; Fr - Exocet voiler africain; Sp - Volador aletón africano.
Maximum standard length 10 cm. Pelagic in neritic surface waters, never spread to the open sea.
Minor importance in coastal fisheries. Widespread in the Indian and Pacific oceans from East Africa,
including the Red Sea and the Persian Gulf, to southern Japan, Marshall Islands, Fiji, and
Queensland (Australia).
Prognichthys brevipinnis (Valenciennes, 1846)

En - Shortfin flyingfish; Fr - Exocet alle courte; Sp - Volador de ala corta.
Maximum standard length about 19 cm. Pelagic, usually in neritic surface waters. Minor importance
to coastal fisheries in Viet Nam. Widespread in the Indian and Pacific oceans from East Africa to
Okinawa, and Fiji.
Prognichthys sealei Abe, 1955

En - Sailor flyingfish; Fr - Exocet matelot; Sp - Volador marinero.
fisheries. Widespread in the Indian and Pacific oceans from East Africa to Central America, probably
absent in inland seas of Southeast Asia.
(after Abe, 1955)
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2180 Bony Fishes
HEMIRAMPHIDAE
Halfbeaks
by B.B. Collette
D iagnostic characters: Elongate fishes with prolonged lower jaw (except in Oxyporhamphus,
Arrhamphus, and Melapedalion) and short triangular upper jaw (except in Oxyporhamphus).
Nostrils in pit anterior to eyes. No spines in fins; dorsal and anal fins posterior in position; pelvic fins
abdominal in position, with 6 soft rays; pectoral fins usually short. Lateral line running down from pectoral-fin
origin and then backwards along ventral margin of body. Scales moderately large, cycloid (smooth), easily
detached. Colour: these fishes live near the surface and are protectively coloured for this mode of life by
being green or blue on the back and silvery white on the sides and ventrally; tip of lower jaw bright red or
orange in most species.
lower jaw length head length
lower jaw pelvic fin

prolonged in
most species
standard length
Habitat, biology, and fisheries: Most species are marine, but some inhabit fresh waters. Omnivorous,
feeding on floating sea grass, crustaceans, and small fishes. They are prone to leap and skitter at the
surface and 3 offshore species, Euleptorhamphus viridis and Oxyporhamphus spp. leap out of the water
and glide like a flyingfish. Although at present these fishes are not of great commercial importance, many
species are regularly found in local markets. The flesh is excellent and halfbeaks are utilized as food in
many parts of the world. From 1990 to 1995, FAO’s Yearbook of Fishery Statistics reports a range of yearly
catch of Hemiramphidae (and Exocoetidae) of around 25 900 to 67 200 t from the Western Central Pacific.
They are mainly caught with seines and pelagic trawls, and dipnetted under lights at night. They are utilized
fresh, dried salted, or smoked.

Belonidae (needlefishes): both upper and lower jaws elongate and armed with needle-sharp teeth.
Exocoetidae (flyingfishes): lack the prolonged lower jaw characteristic of most halfbeaks; pectoral fin or
both pectoral and pelvic fins enlarged and used for aerial gliding.
Belonidae Exocoetidae
Beloniformes: Hemiramphidae 2181
Key to the species of Hemiramphidae occurring in the area

Note: out of about 12 genera and 100 species in the family, 20 species belonging to 7 genera, occur in the
marine waters of the Western Central Pacific. The small species of the fresh-water and estuarine genera
Dermogenys, Nomorhamphus, Hemirhamphodon, and Zenarchopterus are not included in the following key.
1a. Lower jaw not noticeably elongate in adults (Fig.1); anterior margin of upper jaw straight,
not forming a prominent triangular anterior projection; pectoral fins long, 30 to 35% of
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Oxyporhamphus) → 2
1b. Lower jaw elongate (Fig. 2) or not; anterior margin of upper jaw forming a prominent
triangular anterior projection; pectoral fins not more than 28% of standard length . . . . . . . . → 3
lower jaw short lower jaw prolonged in most
Fig. 1 Oxyporhamphus Fig. 2 lateral view of head
2a. Swimbladder cellular (multi-chambered) in specimens longer than 12 cm; pelvic fins in
juveniles (3 to 9 cm standard length) intensely pigmented on distal margins; distance
from pelvic-fin origin to caudal-fin base contained 1.05 to 1.35 times in distance from
pectoral-fin origin to pelvic-fin origin . . . . . . . . . . . . . . . Oxyporhamphus convexus convexus
2b. Swimbladder simple (single-chambered) at all sizes; pelvic fins in juvenilies unpigmented or
with slight pigmentation on outer rays; distance from pelvic-fin origin to caudal-fin base
contained 0.9 to 1.2 times in distance from pectoral-fin origin to pelvic-fin origin . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oxyporhamphus micropterus micropterus
3a. Nasal papilla rounded, fan-shaped, or fimbriate; not projecting far beyond nasal fossa
(Fig. 3a, b); caudal fin emarginate or forked, frequently with an elongate lower lobe; anal
fin of males not different from those of females . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4
3b. Nasal papilla elongate and pointed, not fimbriate; projecting well beyond nasal fossa
(Fig. 3c); caudal fin rounded or truncate, with the longest rays in middle of fin; anal fin
of males modified, some rays widened and elongate . . . . . . . freshwater and estuarine genera
nasal fossa
a) rounded b) fimbriate c) elongate, pointed
Fig. 3 nasal papilla

2182 Bony Fishes
4a. Body compressed and ribbon-shaped; dorsal-fin rays 20 to 25; anal-fin rays 20 to
25 (Fig. 4a); pectoral fins long, 25 to 28% of standard length; pectoral-fin rays
usually 7 to 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Euleptorhamphus viridis
4b. Body not ribbon-shaped; dorsal-fin rays 12 to 18; anal-fin rays 10 to 19 (Fig. 4b);
pectoral fins short, less than 20% of standard length; pectoral-fin rays 10 to 14 . . . . . . . . . → 5
long dorsal fin with 20-25 rays short dorsal fin with 12-18 rays
long anal fin with 20-25 rays short anal fin with 10-19 rays
a) Euleptorhamphus b) Hemiramphus
Fig. 4 posterior part of body
5a. Scales absent on upper jaw; preorbital ridge absent (Fig. 5a) . . . . . . . . . . (Hemiramphus) → 6
5b. Scales present on upper jaw; preorbital ridge well developed (Fig. 5b) . . . . . . . . . . . . . . → 9
no preorbital ridge
preorbital ridge
no scales scales scales
preorbital canal preorbital canal

lateral view dorsal view lateral view dorsal view
a) Hemiramphus b) Hyporhamphus
Fig. 5 detail of head
6a. Pectoral fins relatively long, 4.5 to 5.4 times in standard length (length greater than
distance from their origin to anterior margin of nasal fossa); gill rakers on first arch 33
to 46, usually 36 or more; predorsal scales 35 to 43, usually more than 37 . . . Hemiramphus lutkei
6b. Pectoral fins short, 5.2 to 6.8 times in standard length (length less than distance from
their origin to anterior margin of nasal fossa); gill rakers on first arch 25 to 36, usually
fewer than 34; predorsal scales 29 to 39, usually fewer than 37 . . . . . . . . . . . . . . . . . . → 7
7a. Dorsal fin without well-developed anterior lobe, pigmented on margins; body width
contained 1.8 to 2 times in its depth; adults without spots on side of body . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiramphus archipelagicus
7b. Dorsal fin with well-developed anterior lobe, pigmented on anterior part; body width
contained 1.3 to 1.8 times in its depth; adults usually with short vertical bars on side of
body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8
8a. Adults usually with 3 to 9 (usually 4 to 6), short dark vertical bars on sides of body;
anal-fin rays 9 to 12, usually 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiramphus far
8b. Adults usually with 1 dark spot on sides of body; anal-fin rays 11 to 14, usually 12 or 13
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemiramphus robustus
9a. Nasal papillae fimbriate; upper jaw arched; gill rakers on first arch 47 to 78; lateral line
with 2 branches ascending behind opercle and pectoral-fin origin (Fig. 6a) . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Rhynchorhamphus) → 10
9b. Nasal papillae not fimbriate; upper jaw flat or nearly flat; gill rakers on first arch 19 to 47;
lateral line with 1 branch ascending toward pectoral-fin origin (Fig. 6b) . . . . . . . . . . . . . . . → 11
2 ascending branches of nasal fossa not

upper jaw lateral line fimbriate 1 ascending branch of
nasal papilla arched upper jaw lateral line
fimbriate flattened
a) Rhynchorhamphus b) Hemiramphus
Fig. 6 anterior part of body
10a. Gill rakers on first arch 47 to 67, usually 56 to 63; on second arch 41 to 63 . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhynchorhamphus georgii
10b. Gill rakers on first arch 47 to 59, usually 50 to 55; on second arch 39 to 53 . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhynchorhamphus naga
11a. Lower jaw slightly elongate, less than 1/5 head length . . . . . . . . . . . . . . . . . . . . . . → 12
11b. Lower jaw longer, at least 1/2 head length . . . . . . . . . . . . . . . . . . (Hyporhamphus) → 13
12a. Preorbital canal without posterior branch (Fig. 7a); caudal fin emarginate or slightly
forked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Arrhamphus sclerolepis
12b. Preorbital canal T-shaped, with posterior branch (Fig. 7b); caudal fin deeply forked
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Melapedalion breve
13a. Preorbital canal simple, without posterior branch (Fig. 7a); caudal fin emarginate or
slightly forked . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus (Hyporhamphus) →14
13b. Preorbital canal T-shaped, with posterior branch (Fig. 7b); caudal fin distinctly forked
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus (Reporhamphus) → 17
preorbital distance posterior branch
preorbital canal simple preorbital canal

a) T-shaped b)

(scale removed in front of eye to expose preorbital canal)
14a. Distinct black spots on tips of dorsal fin and upper and lower lobes of caudal fin; caudal
fin forked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus melanopterus
14b. No distinct black spots on dorsal fin or caudal fin; caudal fin emarginate, not forked . . . . . . →15
2184 Bony Fishes
15a. Lower jaw usually longer than head length in adults, its length contained 0.7 to 1.3 times
in head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus limbatus
15b. Lower jaw usually not as long as head length in adults, its length contained 0.9 to 1.8
times in head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 16
16a. Preorbital distance (distance from the anterior margin of the orbit to the posterior vertical
margin of the upper jaw; see Fig. 7b) contained 1.4 to 2 times in orbit diameter and 0.8
to 1.35 times in upper-jaw length . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus neglectus
16b. Preorbital distance contained 1.05 to 1.35 times in orbit diameter and 0.6 to 0.8 times
in upper-jaw length . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus neglectissimus
17a. Width of triangular portion of upper jaw about equal to its length (0.8 to 1.05 times in its
length); preorbital canal narrow, not enlarged ventrally; anterior branch on a straight line
with posterior branch (Fig. 8a) . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus acutus
17b. Width of triangular portion of upper jaw usually wider than its length (0.5 to 0.85 times
in its length); preorbital canal enlarged ventrally; anterior branch at an angle with
posterior branch (Fig. 8b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 18
anterior branch on straight anterior branch at angle with

line with posterior branch posterior branch
ventral pore narrow ventral pore enlarged

(scale removed in front of eye to expose preorbital canal)
18a. Preorbital bone contained 1 to 1.2 times in orbit diameter and 0.65 to 1 times in upper-
jaw length; lower jaw long, its length contained 0.7 to 0.9 times in head length . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hyporhamphus balinensis
18b. Preorbital bone contained 1.35 to 2.15 times in orbit diameter and 0.85 to 1.4 times in
upper-jaw length; lower jaw shorter, its length contained 0.8 to 2 times in head length . . . . . → 19
19a. Preorbital bone contained 1.35 to 1.9 times in orbit diameter; pectoral-fin length con-
tained 7.7 to 10 times in standard length . . . . . . . . . . . . . . . . . . . Hyporhamphus affinis
19b. Preorbital bone contained 1.7 to 2.15 times in orbit diameter; pectoral-fin length con-
tained 6.8 to 8.2 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 20
20a. Upper-jaw tip pointed; gill rakers on first gill arch 36 to 47, as few as 33 in eastern
Australian populations; distance from pelvic-fin origin to caudal-fin base contained 1 to
1.3 times in distance from pectoral-fin origin to pelvic-fin origin; lower jaw length
moderate, in large fishes contained up to 1.4 times in head length . . . Hyporhamphus dussumieri
20b. Upper-jaw tip blunt and rounded; gill rakers on first gill arch 26 to 34, up to 36 to 39 in
Australian populations; distance from pelvic-fin origin to caudal-fin base contained 0.9
to 1.05 times in distance from pectoral-fin origin to pelvic-fin origin; lower jaw short, in
large fishes contained up to 2 times in head length . . . . . . . . . . . . . . Hyporhamphus quoyi

0 Arrhamphus sclerolepis sclerolepis (Günther 1866)
0 Euleptorhamphus viridis (van Hasselt, 1823)
0 Hemiramphus archipelagicus Collette and Parin, 1978
0 Hemiramphus far (Forsskål, 1775)
0 Hemiramphus lutkei (Valenciennes, 1846)
0 Hemiramphus robustus Günther, 1866
0 Hyporhamphus (Reporhamphus) acutus acutus (Günther, 1871)
0 Hyporhamphus (R.) affinis (Günther, 1866)
0 Hyporhamphus (R.) balinensis (Bleeker, 1858)
0 Hyporhamphus (R.) dussumieri (Valenciennes, 1846)
0 Hyporhamphus (Hyporhamphus) limbatus (Valenciennes, 1846)
0 Hyporhamphus (H.) melanopterus Collette and Parin, 1978
0 Hyporhamphus (H.) neglectissimus Parin, Collette, and Schcherbachev, 1980
0 Hyporhamphus (H.) neglectus (Bleeker, 1866)
0 Hyporhamphus (Reporhamphus) quoyi (Valenciennes, 1846)
0 Melapedalion breve (Seale, 1909)
0 Oxyporhamphus convexus convexus (Weber and de Beaufort, 1922)
0 Oxyporhamphus micropterus micropterus (Valenciennes,1846)
0 Rhynchorhamphus georgii (Valenciennes, 1846)
0 Rhynchorhamphus naga Collette, 1976
References
Collette, B.B. 1974. The garfishes (Hemiramphidae) of Australia and New Zealand. Rec. Australian Mus.,
29(2):11-105.
Collette, B.B. and J. Su. 1986. The halfbeaks (Pisces, Beloniformes, Hemiramphidae) of the Far East. Proc. Acad.
Nat. Sci. Philad. , 138(1):250-301.
Parin, N.V., B.B. Collette, and Y.N. Shcherbachev. 1980. Preliminary review of the marine halfbeaks
(Hemiramphidae, Beloniformes) of the tropical Indo-West Pacific. Trudy Inst. Okean., 97:7-173 [In Russian
with English abstract].
2186 Bony Fishes
Hemiramphus archipelagicus Collette and Parin, 1978

Frequent synonyms / misidentifications: None / Hemiramphus marginatus Forsskål, 1775.
FAO names: En - Jumping halfbeak; Fr - Demi-bec saltou; Sp - Agujeta saltona.
8° 21’N, 104° 38’E, 19 cm standard length
Diagnostic characters: An elongate fish with greatly prolonged, beak-like lower jaw; upper jaw short,
triangular and scaleless; preorbital ridge (bony ridge behind nostril) absent. Total number of gill
rakers on first gill arch 25 to 32; 6 to 8 on upper, and 19 to 24 on lower limb of arch. Dorsal fin without
well-developed anterior lobe; dorsal-fin rays 12 to 15; anal-fin rays 10 to 13; pectoral fins short, not
reaching past nasal pit when folded forward and with 11 to 13 rays (usually 12); caudal fin deeply forked,
lower lobe much longer than upper. Predorsal scales 32 to 37 (usually 34 to 36). Colour: dark bluish
above, silvery white below, without any vertical bars on sides; beak dark with a bright red fleshy tip;
upper lobe of caudal fin yellow, tip of dorsal-fin lobe with some yellow.
Size: Maximum total length about 34 cm; maximum standard length 23 cm (from tip of upper jaw to base
of caudal fin).
Habitat, biology, and fisheries: Inhabits the immediate vicinity of coasts, but juveniles of this species
may sometimes be found with floating plants carried out to sea. Taken with purse seines at Karwar on the
west coast of India and by dol nets at Bombay. Marketed fresh and dried salted.
Distribution: An Indo-West
Pacific species known from
the western part of the Indian
Ocean and western Central
P acific. Within the area,
known from the Gulf of Thai-
land, Philippines, and Indone-
sia eastward to New Guinea
and western Polynesia.
Hemiramphus far (Forsskål, 1775)

Frequent synonyms / misidentifications: Hemiramphus commersonii Cuvier, 1829 / None.
FAO names: En - Black-barred halfbeak; Fr - Demi-bec bagnard; Sp - Agujete manchada.
Philippines, 26.3 cm standard length
rakers on first gill arch 25 to 36 (usually 29 to 33); 6 to 10 on upper and 19 to 26 on lower limb of arch.
Dorsal fin with well-developed anterior lobe; dorsal-fin rays 12 to 14; anal-fin rays 10 to 12 (usually 11);
pectoral fins short, not reaching past nasal pit when folded forward and with 11 to 13 rays (usually
12); caudal fin deeply forked, lower lobe much longer than upper. Predorsal scales 32 to 39 (usually 34
to 37). Colour: dark bluish above, silvery white below, with 3 to 9 (usually 4 to 6) vertical bars on sides;
beak dark, with a bright red fleshy tip.
of caudal fin), commonly to 27 cm.
Habitat, biology, and fisheries: Found in proximity of continental coasts and islands, chiefly in areas of
rich submerged vegetation. Adults feed mainly on sea grasses, to a lesser extent on green algae and
diatoms. Taken with gill nets, shore seines, drag nets, or by drifting a fine line with tiny hooks baited with
shrimp. Marketed mostly fresh and dried salted.
Pacific species found in tropi-
cal waters of the Indian and
western parts of the Pacific
oceans. In the area, extends
eastward to the Philippines,
Palau, Fiji, Samoa, and
Tonga south to New Guinea,
New Caledonia, and northern
Australia; north to the Izu
Peninsula of Japan.
2188 Bony Fishes
Hemiramphus lutkei (Valenciennes, 1846)

Frequent synonyms / misidentifications: None / Hemiramphus marginatus Forsskål, 1775.
FAO names: En - Lutke’s halfbeak; Fr - Demi-bec de Lutke; Sp - Agujeta de Lutke.
Sri Lanka, 22.1 cm standard length
(from Parin et al., 1980)
rakers on first gill arch 33 to 46 (usually 36 to 41); 9 to 14 on upper, and 24 to 32 on lower limb of arch.
Dorsal-fin rays 12 to 15 (usually 13 or 14); anal-fin rays 10 to 13 (usually 12); pectoral fins long, reaching
beyond anterior margin of nasal pit when folded forward and contained 4.8 to 5.4 times in standard
length, with 10 to 12 rays (usually 11); caudal fin deeply forked, lower lobe much longer than upper.
Predorsal scales 35 to 43 (usually 37 to 41). Colour: dark bluish above, silvery white below, with no spots
or vertical bars on sides; beak dark with a bright red fleshy tip; upper lobe of caudal fin bluish.
of caudal fin), commonly to 25 cm.
Habitat, biology, and fisheries: Found more offshore than other species of Hemiramphus in the area;
juveniles and immature individuals are relatively common far from shore, usually among floating plants.
Marketed fresh and dried salted.
Distribution: Known from In-
dian and western Central Pa-
cific oceans. Within the area,
known from Indonesia, Philip-
pines, and New Guinea; north
to southern Japan, and east
to Marcus Islands, the Gilbert
Islands, and Samoa.
Hyporhamphus (Reporhamphus) dussumieri (Valenciennes, 1846)

FAO names: En - Dussumier’s halfbeak; Fr - Demi-bec de Dussumier; Sp - Agujeta de Dussumier.
Marshall Is., 20.8 cm standard length
Diagnostic characters: An elongate fish with a prolonged, beak-like lower jaw, contained 4.2 to 5.9
times in standard length and 0.95 to 1.4 times in head length, decreasing with growth; upper jaw short,
triangular, and scaly, its width 0.6 to 0.9 times in its length; length of preorbital bone contained 1.7
to 2.2 times in diameter of orbit and 1 to 1.4 times in length of upper jaw; preorbital ridge (bony ridge
between nasal opening and eye) present; posterior branch to preorbital lateral-line canal present. Total
number of gill rakers on first gill arch 33 to 47, usually 38 to 43 (except in Queensland, Australia, usually
35 to 37), 10 to 14 on upper and 26 to 32 on lower limb of arch. Dorsal and anal fin rays 14 to 16, dorsal
rays usually 15 or 16, anal rays usually 14 or 15; pectoral fins short with 11 or 12 rays; pelvic fins located
closer to caudal-fin base than to origin of pectoral fins; distance from pelvic-fin origin to caudal-fin base
contained 1.05 to 1.3 times in distance from pectoral-fin origin to pelvic-fin origin; caudal fin forked, with
lower lobe longer than upper. Anterior part of dorsal fin and all of anal fin covered with scales; predorsal
scales 37 to 44 (usually 38 to 41). Colour: greenish above, silvery white below; fleshy tip of beak red.
Size: Maximum standard length about 29.8 cm (tip of upper jaw to base of caudal fin) plus 5.3 cm beak,
commonly to 19.0 cm.
Habitat, biology, and fisheries: Most common around islands and coral reefs. Taken mostly with shore
seines. Marketed fresh and dried salted.
Distribution: I n h a b i t s t h e
tropical Indian and western
Pacific oceans. Within the
area, known from Indonesia,
Borneo, Philippines, New
Guinea and eastwards as far
as Tuamotu Islands; north to
Hong Kong and Okinawa.
2190 Bony Fishes
Hyporhamphus (Reporhamphus) quoyi (Valenciennes, 1846)

FAO names: En - Quoy’s halfbeak; Fr - Demi-bec de Quoy; Sp - Agujeta de Quoy.
Nagasaki, Japan, 19.5 cm standard length
Diagnostic characters: An elongate fish with a prolonged, beak-like lower jaw, shorter than head
length, its length contained 4.7 to 8.6 times in standard length and 1.1 to 2 times in head length,
decreasing with growth; upper jaw short, scaly, blunt and rounded, its width contained 0.5 to 0.6 times
in its length; preorbital bone 1.75 to 2.15 times in diameter of orbit and 0.9 to 1.15 times in length
of upper jaw; preorbital ridge (bony ridge between nasal opening and eye) present; posterior branch
to preorbital lateral-line canal present. Total number of gill rakers on first gill arch 26 to 39, usually
27 to 31 (except in Australia, usually 31 to 35), 6 to 14 on upper and 18 to 25 on lower limb of arch.
Dorsal-fin rays 14 to 17 (usually 16), anal-fin rays 13 to 17 (usually 15 or 16); pectoral fins short with
11 to 13 rays; pelvic fins located approximately half way between origin of pectoral fin and base of caudal
fin, distance from pelvic-fin origin to caudal-fin base contained 0.9 to 1.05 times in distance from pectoral-fin
origin to pelvic-fin origin; caudal fin forked, with lower lobe longer than upper. Anterior part of dorsal
fin and all of anal fin covered with scales; predorsal scales 36 to 43 (usually 37 to 40). Colour: greenish
above, silvery white below; fleshy tip of beak red.
Size: Maximum standard length 31.2 cm (tip of upper jaw to base of caudal fin).
Habitat, biology, and fisheries: Inhabits more turbid and estuarine situations than does Hyporhamphus
dussumieri and is not found around oceanic islands. Taken mostly with shore seines. Marketed fresh and
dried salted.
Distribution: Inhabits tropi-
cal western Indian and west-
ern Pacific oceans. Within
the area, known from Thai-
land, Indonesia, Borneo, and
Philippines; north to China
and Nagasaki, Japan, south
to New Guinea and the north-
ern half of Australia.
Hyporhamphus (Hyporhamphus) limbatus (Valenciennes, 1846)

Frequent synonyms / misidentifications: N o n e / Hemiramphus gaimardi Valenciennes, 1846;
Hyporhamphus unifasciatus (Ranzani, 1847).
FAO names: En - Congaturi halfbeak; Fr - Demi-bec congaturi; Sp - Agujeta congaturí.
Diagnostic characters: An elongate fish with greatly prolonged, beak-like lower jaw, equal to, or
longer than head length; upper jaw short, triangular and scaly, its width 0.6 to 0.8 times in its length.
Preorbital distance 1.3 to 2.1 times in diameter of orbit and 0.75 to 1.2 times in length of upper jaw;
preorbital ridge (bony ridge between nasal opening and eye) present; posterior branch to preorbital
lateral-line canal absent. Total number of gill rakers on first gill arch 23 to 37 (usually 25 to 31), 6 to
11 on upper and 19 to 23 on lower limb of arch. Dorsal-fin rays 13 to 16, usually 13 or 14; anal-fin rays
13 to 16 (usually 14 or 15); pectoral fins short, with 10 to 12 rays; caudal fin emarginate, not strongly
forked. Anterior part of dorsal and anal fins covered with scales; predorsal scales (in front of dorsal fin)
30 to 38 (usually 32 to 35). Colour: greenish above, the silvery lateral stripe widening posteriorly, white
ventrally; fleshy tip of beak reddish.
Size: Maximum total length 22 cm; maximum standard length 17 cm (tip of upper jaw to base of caudal
fin), commonly to 13 cm.
Habitat, biology, and fisheries: A coastal species, frequently enters estuaries and even strictly fresh-
water. Taken with cast nets on coasts of India. Marketed fresh and dried salted.
Distribution: Extends from
the Persian Gulf to China
along the mainland coast of
Asia. Within the area, found
from Thailand northward to
China. Replaced in Indone-
sia, Borneo, and the Philip-
pines by the closely related
H yp o r h a m p h u s neglectus,
and around southern New
Guinea and northern Austra-
lia by H. neglectissimus.
2192 Bony Fishes
Oxyporhamphus convexus convexus (Weber and de Beaufort, 1922)

Frequent synonyms / misidentifications: Oxyporhamphus meristocystis Parin, 1961 / Oxyporhamphus
micropterus (Valenciennes, 1846).
FAO names: En - Flying halfbeak.
Diagnostic characters: An elongate fish; adults without the prolonged lower jaw characteristic of
most halfbeaks, but lower jaw up to 4 times in standard length in young (specimens between 3.5 and 4.5 cm
standard length); anterior margin of upper jaw straight, not forming a prominent triangular anterior
projection as in other halfbeaks; preorbital ridge (bony ridge behind nostril) absent. Total number of gill
rakers on first gill arch 26 to 33 (usually 27 to 31). Dorsal fin without well-developed anterior lobe; dorsal-fin
rays 12 to 15; anal-fin rays 13 to 17; pectoral fins elongate, 2.8 to 3.3 times in standard length, barely
reaching origin of pelvic fins, with 11 to 13 rays; pelvic fins short, 7.5 to 11.1 times in standard length,
located about midway between origin of pectoral fins and caudal-fin base; distance from pelvic-fin origin
to caudal-fin base contained 1.05 to 1.35 times in distance from pectoral-fin origin to pelvic-fin
origin; caudal fin forked, with lower lobe much longer than upper. Fins not covered with scales; predorsal
scales large, 28 to 34. Swimbladder in adults (greater than 12 cm standard length) alveolar, composed of
many small cells. Colour: pelvic fins intensively pigmented, with exception of innermost ray; pelvic fins
always intensively pigmented distally in young at 2.5 to 3 cm standard length and larger; skin folds along
sides of lower jaw black in juveniles, with expanded lobes anteriorly (noticeably only in undamaged fish).
Size: Maximum standard length 17.6 cm, commonly to 13 cm.
Habitat, biology, and fisheries: More coastal than Oxyporhamphus micropterus and frequently found
near islands. Marketed fresh with flyingfishes in the markets of Cebu (Philippines) and elsewhere.
Distribution: Widespread in
the Indo-West Pacific in some-
what warmer waters than
Oxyporhamphus micropterus
and extending east only to the
Philippines, New Guinea, New
Britain, and New Ireland. Re-
placed by O. convexus bruuni
Parin, Collette, and Scher-
bachv, 1980, in the Red Sea,
Arabian Sea, and coastal wa-
ters in between.
Arrhamphus sclerolepis sclerolepis Günther, 1866

En - Northern snubnose garfish.
Maximum standard length 21.8 cm. A coastal species that extends into fresh water. Considered
an excellent food fish and taken by commercial and recreational fishermen in Queensland.
Confined to northern Australia and southern Papua New Guinea. Known south along the coast of
Western Australia to the Gascoyne River and along the coast of eastern Australia to Lindeman
Island and Bowen, Queensland. Replaced south of Rockhampton, Queensland, by a southern
subspecies, Arrhamphus s. krefftii (Steindachner, 1867) which extends to Sydney.

Clarence River, North South Wales, 22 cm standard length
(figure of A. s. krefftii)
Euleptorhamphus viridis (van Hasselt, 1823)

En - Longfinned halfbeak.
Maximum standard length 40.5 cm. An oceanic species found throughout the tropical and subtropi-
cal waters of the Indo-Pacific. Not known to be of importance to fisheries.
juvenile
Hemiramphus robustus Günther, 1866

En - Three-by-two garfish.
Maximum standard length 31.5 cm. A coastal species mostly found in more turbid inshore waters
replacing Hemiramphus far, which is more common in more oceanic waters. A high quality food
fish well known in markets of Brisbane, Queensland. Its common name is an allusion to its similarity
to a 3-inch by 2-inch plank of wood. An Australian endemic extending south to Sydney on the east
coast and to Perth in Western Australia.
Perth, Western Australia, 26.5 cm standard length

2194 Bony Fishes
Hyporhamphus (Reporhamphus) acutus acutus (Günther, 1871)

En - Sharpnose halfbeak.
Maximum standard length 18.5 cm. Of no known interest to fisheries. An oceanic species confined
to areas around islands on the Pacific Plate of central Oceania, from Marshall Islands to Easter
Island. Replaced in Hawaii by Hyporhamphus a. pacificus (Steindachner, 1900).
Easter Island, 15.6 cm standard length
Hyporhamphus (Reporhamphus) affinis (Günther, 1866)

En - Coral reef halfbeak.
Maximum standard length 24.3 cm. Found chiefly around coral reefs and islands but extends a little
further from shore than Hyporhamphus (R.) dussumieri which has a similar range. Widely distrib-
uted in tropical Indo-West Pacific from Madagascar and Red Sea through western Central Pacific
east to Tuamotu Archipelago.
Marshall Islands, 24.4 cm standard length
Hyporhamphus (Reporhamphus) balinensis (Bleeker, 1859)

En - Bali halfbeak.
Maximum standard length 16.5 cm. Widely distributed in the tropical Indo-West Pacific from
Mozambique Channel and southern part of the Red Sea through the western Central Pacific east
to Samoa.
New Guinea, 13.8 cm standard length

Hyporhamphus (Hyporhamphus) melanopterus Collette and Parin, 1978

En - Blackfinned halfbeak.
Maximum standard length 17 cm. Confined to the Gulf of Thailand, Java Sea, and Sulu Sea.
Gulf of Thailand, 13.4 cm standard length

Hyporhamphus (Hyporhamphus) neglectissimus Parin, Collette, and Schcherbachev, 1980

En - Australian neglected halfbeak.
Maximum standard length 14.4 cm. A coastal species that replaces Hyporhamphus
(Hyporhamphus) neglectus in waters around New Guinea and northern Australia.
New Guinea, 13.7 cm standard length
Hyporhamphus (Hyporhamphus) neglectus (Bleeker, 1866)

En - Neglected halfbeak.
Maximum known size 16.5 cm standard length. Replaces the coastal Hyporhamphus
(Hyporhamphus) limbatus in waters around Java, Sumatra, Borneo, Sulawesi (Celebes), and the
Philippines.
Kalimantan, 13.9 cm standard length

Melapedalion breve (Seale, 1909)

En - Philippine snubnose halfbeak.
Maximum standard length 22.7 cm. A coastal species limited to waters of South China and Sulu
seas.
(from Seale, 1909)

2196 Bony Fishes
Oxyporhamphus micropterus micropterus (Valenciennes, 1846)

En - Oceanic flying halfbeak.
Maximum standard length 18.5 cm. A true oceanic fish, not bound to coastal waters at any period
of its life. Widespread throughout the Indo-Pacific from 20°N to 20°S and extending north to
southern Japan in the western Pacific. Replaced in the Atlantic by Oxyporhamphus m. similis
Bruun, 1935.
Gulf of Mexico, 14.6 cm standard length

(figure of O. m. similis)
Rhynchorhamphus georgii (Valenciennes, 1846)

En - George’s halfbeak.
Maximum standard length 23.1 cm. The most widespread of the 4 species of Rhynchorhamphus,
found from the Persian Gulf through the Arabian Sea and Bay of Bengal through the western Central
Pacific north to Taiwan Province of China and Hong Kong and east to New Guinea and northern
Australia.

N. Borneo, 16.2 cm standard length
Rhynchorhamphus naga Collette, 1976

En - Naga halfbeak.
Maximum standard length 17.7 cm. Restricted to the Gulf of Thailand, South China Sea, and
western Java Sea
Gulf of Thailand, 16.5 cm standard length

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Cyprinodontiformes: Aplocheilidae 2197
Order CYPRINODONTIFORMES
APLOCHEILIDAE
Egg-laying toothcarps
by L.R. Parenti
D iagnostic characters: Small, fusiform, cyprinodontiform fishes (average size at maturity 6 cm

standard length). Dorsal surface of head flat. Eyes large. Mouth terminal, large, protrusible. Small,
unicuspid teeth in pavement dentition in outer jaw. Pectoral fins rounded. Caudal fin rounded or middle
rays of caudal fin elongate. Dorsal fin with 7 to 20 soft rays; anal fin with 15 to 19 soft rays; caudal fin with
14 to 21 branched rays; pectoral fins with 14 to 21 soft rays; pelvic fins with 6 soft rays. Scales moderate,
cycloid, 25 to 42 in lateral series; no lateral line. Small patch of teeth on vomer. Colour: (of Aplocheilus
panchax): highly variable; somewhat translucent in life, with a silvery, “pineal” spot on dorsal surface
of head; head and body greyish yellow to medium brown dorsally, lighter brown to yellow ventrally; thin,
black horizontal line from tip of lower jaw through eye variably present; large, black spot on middle
portions of dorsal-fin rays; pectoral fins hyaline; pelvic, anal, and caudal fins with submarginal
brownish yellow band.
2nd dorsal fin

always present
1st dorsal fin absent
pectoral fins
mouth large, rounded
protrusible
no fleshy ventral keel

caudal fin
rounded, middle
rays slightly
urogenital opening elongate
(after Smith, 1945)
Habitat, biology, and fisheries: Fresh to brackish water; a single species (Aplocheilus panchax) in the
area found in brackish water, estuaries, and mangroves. Small, surface-feeding, schooling fishes.
Omnivorous, but feeds principally on insects and mosquito larvae. Oviparous.
Remarks: The family Aplocheilidae is distributed broadly throughout tropical Africa, Madagascar, and
from the Indian subcontinent to the Indo-Australian Archipelago. Meristic data are for the family, with
diagnostic colour of A. panchax.

pectoral fins
Adrianichthyidae: superficially similar to egg-laying mouth small, falcate
toothcarps, but readily distinguished by a non-pro- non-protrusible
trusible mouth, no teeth on vomer, dorsal- and anal-
fin rays of male longer and thicker than those of
females and with bony contact organs on distal
segments of anal-fin rays, and pectoral fins falcate.
Adrianichthyids are nearly transparent in life, with
sparse pigmentation.
caudal fin
truncate
female
Adrianichthyidae
(after Iwamatsu et al., 1982)
2198 Bony Fishes
Phallostethidae: readily distinguished from egg-laying toothcarps by the presence, in males, of a bony,
externalized intromittent organ, and urogenital openings under head; nearly transparent in life, with sparse
pigmentation.
mouth membranous dome 1st dorsal fin 2nd dorsal fin

supraterminal, in some species present or absent always present
slightly to highly
protrusible
caudal fin
emarginate
fleshy, rayless ventral keel

males with elaborate, externalized (after Roberts, 1971a)
intromittent organ under head derived
from pelvic fins, and part of pectoral fins
Phallostethidae
Poeciliidae: readily distinguished from egg-laying toothcarps by males with an intromittent organ derived
from anterior anal-fin rays, and a small mouth.
2nd dorsal fin always

present
1st dorsal fin body and fins may be
absent densely pigmented
mouth small,
protrusible
caudal fin
truncate or
lanceolate
no fleshy ventral keel males with intromittent organ at midbody

derived from anterior portion of anal fin
(after Rosen, 1973)
Poeciliidae
A single species occurring in the area

Aplocheilus panchax (Hamilton, 1822)
References
Parenti, L. R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha).
Bull. Amer. Mus. Nat. Hist. 168(4):335-557.
Smith, H.M. 1945. The freshwater fishes of Siam, or Thailand. Bull. U.S. Natn. Mus., 188(11):1-622.
Cyprinodontiformes: Poeciliidae 2199
POECILIIDAE
Livebearing toothcarps
by L.R. Parenti
D iagnostic characters: Small to moderate, laterally compressed, cyprinodontiform fishes; females

usually larger than males. Eyes moderately large. Mouth supraterminal, small, and protrusible.
Small, unicuspid teeth in single or multiple rows on premaxilla and dentary; no tooth patch on vomer.
Pectoral fins falcate, may be set relatively high on side of body. Caudal fin truncate or slightly lanceolate.
Single, soft-rayed dorsal fin at midbody or set posteriorly. Dorsal fin with 4 to 16 soft rays; anal fin with 8
to 10 soft rays; pectoral fins with 9 to 16 soft rays; pelvic fins with 6 soft rays; caudal fin with 11 to 15
branched rays. Species in the area with anal fin of males modified into a gonopodium used to
transfer sperm bundles to females. Anal-fin rays 3 through 5 of males thickened and elongate;
haemal arches expanded. Scales moderate, cycloid, 30 to 34 in lateral series; no lateral line. Colour:
some species highly sexually dichromatic and dimorphic; males may be more brightly coloured and smaller
than females; body nearly transparent in life in females and juveniles.
a single dorsal fin
densely pigmented
mouth small, caudal fin

protrusible truncate
fleshy, rayless ventral keel intromittent organ at midbody derived

from anterior portion of anal fin
Poecilia sphenops (male)
females less brightly anal fin unmodified

coloured than males
(after Rosen, 1973) Poecilia sphenops (female)

Habitat, biology, and fisheries: Fresh to brackish water, species in the area in tidal portions of rivers,
estuaries, or in mangroves. Small, surface-feeding, abundant, schooling fishes. Omnivorous. Tolerant of
extremes of temperature and salinity. Species in the area viviparous, males pass sperm bundles to females
through a modified anal fin or gonopodium. Poeciliids are not native to the area; species found here were
either introduced for mosquito control or released, presumably accidentally, from aquarium stocks.
2200 Bony Fishes
Remarks: The family Poeciliidae as diagnosed by Parenti (1981) comprises 3 subfamilies: Poeciliinae,
Fluviphylacinae, and Aplocheilichthyinae. Parenti’s Poeciliinae is equivalent to the Poeciliidae of previous
authors, such as Rosen and Bailey (1963). The only species in the family Poeciliidae that have been
introduced into the Western Central Pacific are in the subfamily Poeciliinae. The characters listed above
refer only to that subfamily.
single dorsal fin set
mouth non- far back on body
Similar families occurring in area protrusible
Adrianichthyidae: superficially similar to livebearers
but with a non-protrusible rather than protrusible
mouth and lacking an intromittent organ.
Aplocheilidae: differing most notably from livebearers
by having a large mouth and lacking an intromittent
organ.
Phallostethidae: superficially similar to livebearers, caudal fin
but with an intromittent organ in males modified from truncate
no fleshy, ventral keel urogenital opening
pectoral- and pelvic-fin rays and under the head, not
a modified anal fin at midbody; in addition, phal-
lostethids are nearly transparent in life, with sparse Adrianichthyidae
pigmentation. (after Iwamatsu et al., 1982)
single dorsal fin set far mouth

supraterminal, 1st dorsal fin
back on body
mouth large, slightly to highly may be present 2nd dorsal
protrusible protrusible membranous or absent fin always
dome in some present
caudal fin anal-fin rays

no fleshy ventral keel rounded, unmodified caudal fin
middle elaborate, externalized
anal-fin rays rays intromittent organ under head fleshy, rayless emarginate
urogenital opening slightly derived from pelvic fins, and
anterior to anal fin unmodified ventral keel
elongate part of pectoral fins
Aplocheilidae Phallostethidae
(after Smith, 1945) (after Roberts, 1971)

Gambusia affinis (Baird and Girard, 1853)
Poecilia latipinna (LeSueur, 1821)
Poecilia reticulata Peters, 1859
Poecilia sphenops Valenciennes in Cuvier and Valenciennes, 1846
References
Parenti, L.R. 1981. A phylogenetic and biogeographic analysis of cyprinodontiform fishes (Teleostei, Atherinomorpha).
Bull. Amer. Mus. Nat. Hist., 168(4):335-557.
Rosen, D.E. 1973. Suborder Cyprinodontoidei. Superfamily Cyprinodontoidea. Families Cyprinodontidae, Poeciliidae,
Anablenidae. Fishes of the Western North Atlantic, part 6. Sears Foundation for Marine Research.
Rosen, D.E. and R.M. Bailey. 1963. The poeciliid fishes (Cyprinodontiformes), their structure, zoogeography, and
systematics. Bull. Amer. Mus. Nat. Hist., 126(1):1-176.
Stephanoberyciformes: Melamphaidae 2201
Order STEPHANOBERYCIFORMES
MELAMPHAIDAE
Bigscales (ridgeheads)
by J.A. Moore and J.R. Paxton
D iagnostic characters: Small (to about 16 cm total length) stephanoberyciform fishes with
subcylindrical body. Head large; large mucous cavities separated by thin ridges. Eyes small in
most species. Snout short and steep. Jaws long, extending to or beyond posterior margin of eye. Teeth
minute, in bands or uniserial row in jaws; palate toothless. One dorsal fin with I to III weak spines and
9 to 18 soft rays; anal fin with I weak spine and 7 to 10 soft rays; caudal fin with 3 or 4 procurrent spines
preceding upper and lower lobes; pelvic fins thoracic, with I spine and 6 to 8 soft rays; pectoral fins
with 13 to 16 rays. Scales thin and cycloid, usually deciduous, moderate-sized to very large, 12 to 40 in
longitudinal series. Lateral line reduced to 1 or 2 pored scales behind upper edge of operculum. No
photophores or light organs. Colour: body and head dark brown or black.
Habitat, biology, and fisheries: Occurring in meso- and bathypelagic waters in depths of 200 to 2 000 m.
Many species widely distributed in Atlantic and Indo-Pacific. Feed primarily on gelatinous organisms and
small crustaceans. Little is known of behaviour. In Western Central Pacific, Scopeloberyx opisthopterus,
Melamphaes simus, and M. danae are dwarf species maturing at sizes less than 3 cm. Frequently taken
in deep-water trawls, but of no commercial importance.
Remarks: A total of 33 species in 5 genera is recognized, found in the midwaters of all oceans except the
Arctic Ocean and Mediterranean Sea. The genera Poromitra and Scopeloberyx need revision. Further
deep-sea trawling should add more species to the area.

Gibberichthyidae: dorsal fin preceded by V to VII
stout, wide-based spines; anal fin preceded by III to
V stout, wide-based spines; 5 to 7 procurrent spines
in upper and lower lobes of caudal fin; pelvic fins
with I spine and 5 soft rays; lateral line fully devel-
oped.
Gibberichthyidae
2202 Bony Fishes

Melamphaes danae Ebeling, 1962
Melamphaes eulepis Ebeling, 1962
Melamphaes janae Ebeling, 1962
Melamphaes longivelis Parr, 1933
Melamphaes polylepis Ebeling, 1962
Melamphaes simus Ebeling, 1962
Poromitra crassa Parin and Ebeling, 1980
Poromitra crassiceps (Günther, 1878)
Poromitra megalops (Lütken, 1877)
Poromitra oscitans Ebeling, 1975
Scopeloberyx opisthopterus (Parr, 1933)
Scopeloberyx robustus (Günther, 1887)
Scopelogadus mizolepis mizolepis (Günther, 1878)
Scopelogadus unispinis Ebeling and Weed, 1963
Sio nordenskjoldii (Lönnberg, 1905)
Reference
Ebeling, A.W. 1986. Family Melamphaidae. In Smiths’ sea fishes, edited by M.M. Smith and P.C. Heemstra. Johan-
nesburg, Macmillan South Africa, pp. 427-431.
Stephanoberyciformes: Gibberichthyidae 2203
GIBBERICHTHYIDAE
Gibberfishes
by J.R. Paxton
D iagnostic characters: Small (to 13 cm) stephanoberyciform fishes; body deep (young) to moderately
robust. Head moderate; large mucous cavities on top of head separated by very thin ridges and
covered by thin skin in undamaged (rarely) specimens. Eyes very large in juveniles, becoming smaller
than snout length in larger specimens. Mouth moderate, jaws extending to posterior margin of eye in larger
specimens. Teeth villiform in bands on jaws; no teeth on vomer or palatine. Gill rakers lath-like. Separate,
rigid, wide-based fin spines precede dorsal and anal fins; origin of soft dorsal and anal-fin rays posterior
and opposite; dorsal fin with V to VII spines and 8 or 9 soft rays; anal fin with IV or V spines and 7
to 9 soft rays; pelvic fins abdominal, with no spines and 5 or 6 soft rays; elaborate pelvic-fin appendage
from elongated third pelvic-fin ray in larvae and prejuveniles; pectoral fins with 13 to 15 rays. Scales
cycloid, deciduous. Lateral line as vertical rows of papillae overlying scales. No photophores or
luminous tissue. No cavernous tissue. Total vertebrae 28 to 31. Colour: brown-black in largest specimens.
Habitat, biology, and fisheries: Meso-, bathy-, and benthopelagic, mostly over continental and island
slopes. Distinctive kasidoron larval/prejuvenile stage in epipelagic waters. Feed as zooplankton pickers,
with small crustaceans in stomachs of both prejuveniles and adults. Rare deep-sea fishes of no
commercial importance.
Remarks: Two species in a single genus, one each in the tropical West Atlantic and tropical Indo-West
Pacific.

Melamphaidae: dorsal fin with I to III weak spines;
anal fin with I weak spine; pelvic fins with I spine and
6 to 8 soft rays; lateral line reduced to 1 or 2 scales
behind operculum, no arborescent, elongate rays in
larvae or prejuveniles.
Gibberichthys latifrons (Thorpe, 1969) Melamphaidae
References
de Sylva, D.P. and W.N. Eschmeyer. 1977. Systematics and biology of the deep-sea fish family Gibberichthyidae, a
senior synonym of the family Kasidoroidae. Proc. Cal. Acad. Sci., 41(6):215-231.
Kotlyar, A.N. 1996. Beryciform fishes of the world ocean. Moscow, VNIRO Publishing, 368 p. [In Russian]
2204 Bony Fishes
RONDELETIIDAE
Redmouth whalefishes
by J.R. Paxton
D iagnostic characters: Small (to 11 cm) stephanoberyciform fishes, body flabby, somewhat whale-
shaped with median fins opposite and far posterior. Head large; mucous cavities on top of head
indistinct, covered by thick skin. Eyes small. Snout very long; nasal organ moderately developed. Mouth
large, jaws not extending beyond posterior margin of eye. Teeth small and closely set, on jaws and
vomer; palatine, ectopterygoid, and copula lacking teeth. Gill rakers well developed, lath-like. Fins without
spines; 1 dorsal fin with 13 to 16 soft rays; anal fin with 13 to 16 soft rays; pectoral fins with 9 to 11 rays;
pelvic fins abdominal with 5 or 6 soft rays; caudal fin with 19 principal rays. Lateral line as vertical
rows of papillae without supporting internal scales. No external body scales. No photophores or
luminous tissue. No cavernous tissue. Pleural ribs present. Total vertebrae 24 to 27. Colour: in life,
orange-brown, inside mouth and gill cavities red-orange; in preservative, brown.
Habitat, biology, and fisheries: Meso- and bathypelagic. Feeding mode as predator on amphipods and
crustaceans. Uncommon deep-sea fishes of no commercial importance.
Remarks: One genus with 2 species throughout the world ocean in tropical and temperate latitudes.
Rondeletia bicolor is now known from 3 Pacific records, including 1 in the area.
Barbourisiidae: mouth very large with jaws extending far behind eye; head and body covered with tiny
scales with central spine giving velvet-like texture; live colour bright red-orange.
Cetomimidae: no pelvic fins; mouth extremely large with jaws extending far behind eye; most with lateral
line as broad tube pierced by large pores, a single species with vertical rows of papillae.
Barbourisiidae Cetomimidae

Rondeletia bicolor Goode and Bean, 1895
Rondeletia loricata Abe and Hotta, 1963
Reference
Kotlyar, A.N. 1996. The osteology, intraspecific structure, and distribution of Rondeletia loricata (Rondeletiidae). Vopr.
Ikhtiol., 36(2):154-168. [In Russian, English transl. J. Ichthyol., 26(3)].
Stephanoberyciformes: Barbourisiidae 2205
BARBOURISIIDAE
Redvelvet whalefish
by J.R. Paxton
D iagnostic characters: Moderate-sized (to 33 cm) stephanoberyciform fishes; body somewhat

flabby, moderately robust, whale-shaped with median fins opposite and far posterior. Head large;
mucous cavities on top of head separated by thick ridges and covered by skin. Eyes small. Snout very
long; nasal organ moderately developed. Mouth very large, jaws extending far behind eye. Teeth small
and closely set, on jaws, vomer, and ectopterygoid; palatine and copula lacking teeth. Gill rakers well
developed, lath-like. Fins without spines; 1 dorsal fin with 19 to 22 soft rays; anal fin with 15 to 18 soft
rays; pectoral fins with 13 or 14 rays; pelvic fins abdominal with 6 soft rays; caudal fin with 19 principal
rays. Lateral line well developed as a broad tube pierced by small pores and supported by internal
scales. Body and head covered by small, adherent, non-imbricate scales with a single, central spine
giving a velvet-like texture. No photophores or luminous tissue. No cavernous tissue. Pleural ribs
present. Total vertebrae 40 to 43. Colour: in life, bright red-orange; in preservative, white.
Habitat, biology, and fisheries: Mesopelagic as juveniles, benthopelagic as adults. Feeding mode
unknown, presumably as predator on crustaceans. Rare deep-sea fishes of no commercial importance.
Remarks: One genus and species throughout the world ocean in tropical and temperate latitudes.
Rondeletiidae: no external body scales; lateral line as vertical rows of papillae; mouth large, but jaws not
extending beyond posterior margin of eye; live colour orange-brown, in preservative brown.
Cetomimidae: no external body scales; no pelvic fins; colour brown or black.
Rondeletiidae Cetomimidae

Barbourisia rufa Parr, 1945
References
Kotlyar, A.N. 1995. Osteology and distribution of Barbourisia rufa (Barbourisiidae). Vopr. Ikhtiol., 35(3):282-289. [In
Russian, English transl. J. Ichthyol., 35(6)]
Paxton, J.R. and D.J. Bray. 1986. Family Barbourisiidae. In Smiths’ sea fishes, edited by M.M. Smith and P.C. Heemstra.
Johannesburg, Macmillan, p. 434.
2206 Bony Fishes
CETOMIMIDAE
Whalefishes
by J.R. Paxton
D iagnostic characters: Small to moderate-sized (to 40 cm) stephanoberyciform fishes; body soft and
flabby, slender to robust, whale-shaped with median fins opposite and far posterior. Head very
large; mucous cavities on top of head indistinct and covered by skin in undamaged specimens. Eyes tiny.
Snout very long; nasal organ poorly developed (well developed in Procetichthys). Mouth enormous, jaws
extending far behind eye. Teeth tiny and closely set, small and widely spaced, or elongate in well defined,
closely set rows; jaws, pharyngobranchials, and copula always with teeth; vomer, palatine, and ectop-
terygoid usually with teeth. Gill rakers club-shaped, or as tooth patches, tooth plates, or individual
teeth, never lath-like. Fins without spines; 1 dorsal fin with 13 to 37 soft rays; anal fin with 11 to 34 soft
rays; pectoral fins with 15 to 24 rays; no pelvic fins; caudal fin with 10 to 19 principal rays. Lateral
line very well developed, supported with internal scales; in most as a broad tube pierced by large
pores (Procetichthys with vertical rows of lateral-line papillae). No external body scales. No photopho-
res or apparent luminous tissue. Unique cavernous tissue around anus in most, over base of anal fin
and other areas in many species. No pleural ribs. Total vertebrae 38 to 59. Colour: brown or black;
in fresh specimens fin rays and inside mouth reddish orange.
Habitat, biology, and fisheries: Bathypelagic as adults, some may be benthopelagic. Feeds on crusta-
ceans. Rare deep-sea fishes of no commercial importance.
Remarks: Nine genera with about 35 species (including 15 undescribed) throughout the world ocean from
boreal latitudes to the Antarctic. When the genus Cetomimus is revised, additional species will be recorded
from the area.

Rondeletiidae: pelvic fins present; lateral line as vertical rows of papillae; jaws not extending beyond
posterior margin of eye; pleural ribs present.
Barbourisiidae: pelvic fins present; head and body covered with tiny scales with central spine giving
velvet-like texture; live colour bright red-orange.
Rondeletiidae Barbourisiidae
Stephanoberyciformes: Cetomimidae 2207
Megalomycteridae: nasal organ enormous; jaws not extending behind eye; pelvic fins in some; body scales
in some.
Mirapinnidae: jaws not extending behind eye; pelvic fins present.
Megalomycteridae Mirapinnidae

Cetomimus gillii Goode and Bean, 1895
Cetostoma regani Zugmayer, 1914
Danacetichthys galathenus Paxton, 1989
Ditropichthys storeri (Goode and Bean, 1895)
Gyrinomimus bruuni Rofen, 1959 (unpublished record, Paxton ms)
Gyrinomimus sp. nov. b (unpublished record, Paxton ms)
Gyrinomimus sp. nov. c (unpublished record, Paxton ms)
Rhamphocetichthys savagei Paxton, 1989
Reference
Paxton, J.R. 1989. Synopsis of the whalefishes (family Cetomimidae) with descriptions of four new genera. Rec. Aust.
Mus., 41(2):135-206.
2208 Bony Fishes
MIRAPINNIDAE
Tapetails (hairyfish)
by J.R. Paxton
D iagnostic characters: Small (to 5 cm) stephanoberyciform fishes; body slender to very elongate,
with median fins opposite and far posterior. Head moderate. Eyes small to moderate. Snout
moderate to large; nasal organ poorly developed. Mouth moderate, jaws not extending behind eye.
Teeth small and closely set in 1 row on premaxillary and few rows on dentary; no teeth on vomer or palatine.
Gill rakers lath-like. Fins without spines; 1 dorsal fin with 16 to 33 soft rays; anal fin with 14 to 29 soft rays;
pectoral fins with 13 to 24 rays; pelvic fins jugular with 4 to 10 soft rays; caudal fin with 19 principal
rays, caudal streamer longer than body in larvae. No lateral line. No body scales. Body of 1 genus
(Mirapinna) covered with dense, hair-like papillae; others with naked skin. No photophores or luminous
tissue. No cavernous tissue. No pleural ribs. Total vertebrae 42 to 55. Colour: brown, black, or
transparent.
Habitat, biology, and fisheries: Larvae and juveniles epipelagic; mature adults unknown, perhaps meso-
pelagic. Feeding mode zooplankton pickers on copepods. Rare oceanic fishes of no commercial importance.
Remarks: Three genera with 6 species (1 undescribed) throughout the world ocean in tropical and
subtropical latitudes. The most recent review is that of Bertelsen (1986), while that of Bertelsen and
Marshall (1984) illustrates the most species. A revision of the family is needed, but mature specimens have
yet to be collected. More species in the area are expected.

Megalomycteridae: nasal organ enormous; pelvic fins thoracic with 1 to 3 rays, or absent; scales present
in some; no skin papillae or caudal streamer.
Cetomimidae: pelvic fins absent; mouth enormous, jaws extending far behind eye.
Megalomycteridae Cetomimidae

Eutaeniophorus festivus (Bertelsen and Marshall, 1956)
Parataeniophorus sp.
References
Bertelsen, E. 1986. Family Mirapinnidae. In Smiths’ sea fishes, edited by M.M. Smith and P.C. Heemstra.
Johannesburg, Macmillan, pp. 406-407.
Bertelsen, E. and N.B. Marshall. 1984. Mirapinnatoidei: development and relationships. In Ontogeny and systematics
of fishes, edited by H.G. Moser et al. Amer. Soc. Ichthyol. Herpetol. Spec. Publ., (1):380-383.
Stephanoberyciformes: Megalomycteridae 2209
MEGALOMYCTERIDAE
Bignose fishes
by J.R. Paxton
D iagnostic characters: Small (to 7 cm) stephanoberyciform fishes; body elongate, with median fins
opposite and far posterior. Head moderate. Eyes small to moderate. Snout elongate to very large;
nasal organ enormous, covering much of snout. Mouth moderate, jaws not extending behind eye.
Teeth small and closely set in 1 to few rows on premaxillary and dentary; teeth present or absent on vomer,
absent on palatine and copula. Gill rakers poorly developed, few in number. Fins without spines; 1 dorsal
fin with 15 to 31 soft rays; anal fin 13 to 29 soft rays; pectoral fins with 18 to 23 rays; pelvic fins thoracic
with 1 to 3 rays, or absent; caudal fin with 16 principal rays. Lateral line poorly developed or unknown.
Body scales small to moderate, non-imbricate, cycloid, or absent. No photophores or luminous tissue.
No cavernous tissue. No pleural ribs. Total vertebrae 41 to 55. Colour: brown, black, or unknown.
Habitat, biology, and fisheries: Meso- and bathypelagic. Feed as zooplankton pickers on copepods. All
specimens histologically examined (more than 20) are males. Very rare deep-sea fishes of no commercial
importance.
Remarks: Four genera with 7 or 8 species (3 undescribed) throughout the world ocean in tropical and
subtropical latitudes. A family revision is necessary. More species are expected with further deep-sea
collecting in the area.

Mirapinnidae: nasal organ poorly developed; pelvic fins jugular with 4 to 10 rays.
Cetomimidae: jaws extending far behind eye; lateral line well developed; all but a single species with poorly
developed nasal organs.
Mirapinnidae Cetomimidae

Ataxolepis apus Myers and Freihofer, 1966
Cetomimoides parri Koefoed, 1955 (unpublished record from the Philippines)
Reference
Becker, V.E. 1981. On the first record of a rare bathypelagic fish from the genus Ataxolepis (Megalomycteridae) in the
southern hemisphere. Vopr. Ikhtiol., 21(3):558-561. [In Russian, English transl. J.Ichthyol., 21(3)]
click for next page

2210 Bony Fishes
Order BERYCIFORMES
ANOPLOGASTRIDAE
Fangtooths
by J.R. Paxton
D iagnostic characters: Small (to 16 cm)

beryciform fishes, body short, deep, and
compressed. Head large, steep; deep mu-
cous cavities on top of head separated by
serrated crests; very large temporal and pre-
opercular spines and smaller orbital (frontal)
spine in juveniles of one species, all disap-
pearing with age. Eyes smaller than snout
length in adults (but larger than snout
length in juveniles). Mouth very large, jaws
extending far behind eye in adults; one
supramaxilla. Teeth as large fangs in pre-
maxilla and dentary; vomer and palatine
toothless. Gill rakers as gill teeth in adults
(elongate, lath-like in juveniles). No fin spines; dorsal fin long based, roughly in middle of body, with
16 to 20 rays; anal fin short-based, far posterior, with 7 to 9 rays; pelvic fin abdominal in juveniles,
becoming subthoracic with age, with 7 rays; pectoral fin with 13 to 16 rays. Scales small, non-overlap-
ping, spinose, cup-shaped in adults; lateral line an open groove partly covered by scales. No light
organs. Total vertebrae 25 to 28. Colour: brown-black in adults.
Habitat, biology, and fisheries: Meso- and bathypelagic. Distinctive caulolepis juvenile stage, with greatly
enlarged head spines in one species. Feeding mode as carnivores on crustaceans as juveniles and on
fishes as adults. Rare deepsea fishes of no commercial importance.
Remarks: One genus with 2 species throughout the world ocean in tropical and temperate latitudes. The
family was revised by Kotlyar (1986).

Diretmidae: No fangs, jaw teeth small, in bands;
anal fin with 18 to 24 rays.
Trachichthyidae: No fangs, jaw teeth small, in
bands; dorsal and anal fins with III to VIII and II or
III spines respectively, in addition to soft rays.
Anomalopidae: No fangs, jaw teeth small, in bands;
dorsal and anal fins with II to IV and I or II spines
respectively, in addition to soft rays.
Trachichthyidae
Diretmidae Anomalopidae
Anoplogaster brachycera Kotlyar, 1986
Anoplogaster cornuta (Valenciennes, 1833)
Reference
Kotlyar, A. N., 1986. Classification and distribution of fishes of the family Anoplogasteridae. Vopr. Ikhtiol., 26(4):531-551.
[in Russian, English transl. J. Ichthyol., 26(4)]
Beryciformes: Diretmidae 2211
DIRETMIDAE
Spinyfins
by J.R. Paxton
D iagnostic characters: Moderate-

sized (to 37 cm) beryciform fishes;
body round to oval, strongly
compressed. Head moderate to large,
deep; deep mucous cavities on top of
head separated by thin, serrated ridges;
no spine on preopercle (except in young
juveniles of some). Eyes very large,
much longer than snout length. Mouth
large, very oblique, jaws not extending
behind eye; 1 supramaxilla. Teeth
small, in villiform bands, on premaxilla
a nd de nt ar y ; v om er and palatine
toothless. Gill rakers lath-like. No true
fin spines; dorsal fin very long based,
origin just behind pectoral-fin origin, with 24 to 30 soft rays; anal fin posterior, with 18 to 24 soft
rays; pelvic fins subthoracic to thoracic, with I laminar, serrate spine and 6 soft rays; pectoral fins with 16
to 20 soft rays. Scales small, spinose; present on dorsal and anal-fin rays; no lateral line; enlarged
midventral scutes present. No light organs. Total vertebrae 26 to 32. Colour: silver and grey back.
Habitat, biology, and fisheries: Meso- and bathypelagic; largest adults of some species benthopelagic.
Feeding mode as planktivores. Uncommon deep-sea fishes, sometimes locally more abundant in deep
water, of no present commercial importance.
Remarks: Three genera with 4 species throughout the world ocean in tropical and temperate latitudes,
except eastern North Pacific and Mediterranean Sea.
Anoplogastridae: jaws with very large fangs; anal fin with 7 to 9
soft rays; pelvic fins without laminar spine.
Trachichthyidae: mucous cavities of head extensive; spine pre-
sent on preopercle; dorsal and anal fins with spines; lateral-line
scales distinct and usually enlarged.
Anomalopidae: large light organ present under eye; dorsal and
anal fins withspines;lateral-linescalesdistinct and oftenenlarged.
Anoplogastridae
Anomalopidae
Trachichthyidae

Diretmichthys parini (Post and Quero, 1981)
Diretmoides pauciradiatus (Woods, 1973)
Diretmoides veriginae Kotlyar, 1987
Diretmus argenteus Johnson, 1864
Reference
Kotlyar, A.N. 1990. Diretmichthys, a new genus of Diretmidae (Beryciformes). Vopr. Ikhtiol., 30(1):144-151. [in
Russian, English transl. J. Ichthyol., 30(2)]
2212 Bony Fishes
ANOMALOPIDAE
Flashlight fishes
by J.R. Paxton and G.D. Johnson
D iagnostic characters: Small to moderate-sized beryciform fishes (to 26 cm), oblong to moderately
deep. Head moderate; snout blunt; moderate mucous cavities on top of head separated by broad
ridges and covered by skin. Eyes large, their diameter longer than snout length; large light organ
below eyes, sometimes obscured by rotation downward or by curtain of dark skin. Mouth moderate,
oblique, jaws not extending behind eyes; 1 (Anomalops) or 2 supramaxillae. Teeth very small, conical,
on jaws and palatines, vomer toothless. Snout blunt. One dorsal fin or spinous dorsal fin separate from
second dorsal; dorsal fin with II to VI spines and 14 to 19 soft rays; anal fin with I or II spines and 9
to 14 soft rays; caudal fin deeply forked with 10+9 principal rays; pelvic fins with 0 to I spine and 5 to 7 soft
rays; pectoral fins with 16 or 17 soft rays. Branchiostegal rays 8, ventral margins spiny or smooth. Scales
small, strongly ctenoid; body scale rows 50 to 150, lateral-line scales 30 to 69; midventral scutes 5 to 19,
reduced and discontinuous or moderate and continuous. Total vertebrae 30 (13-15+15-17). Colour: head
and body brown to black; external surface of light organ white; lateral-line scales reflective or not.
Habitat, biology, and fisheries: Mostly around coral reefs, from shallow water to a depth of 350 m, with
some species migrating to the surface on moonless nights. Light organ colonized by luminous bacteria.
Feeding mode as zooplankton pickers. At least 1 species schooling in large groups. Popular species in
public aquaria, and a target for some local fishers as bait.
Remarks: Seven species in 6 genera are recognised, distributed in the Caribbean, Indian eastern Pacific
and western Pacific from Tahiti to Japan, Australia, the Red Sea, and Comoro Islands. Parmops coruscans
is known from only 3 specimens (Tahiti, Fiji) from 350 m, while Protoblepharon rosenblatti is known only
from the holotype (Cook Islands) from 275 m.

Trachichthyidae: no light organ under eyes in any
species; midventral scutes of most species fewer
and larger than those of anomalopids.
Trachichthyidae
Beryciformes: Anomalopidae 2213
Key to species of Anomalopidae in the area

1a. One dorsal fin with II spines and 18 or
19 soft rays; light organ not rotatable,
obscured by raising black elastic shut-
ter to cover lateral surface; midventral
scutes reduced and discontinuous
(Fig. 1) . . . . . . . . Photoblepharon palpebratus
1b. Two dorsal fins, the first with V or VI
spines and the second with I spine and
14 to 16 soft rays; light organ rotatable
downwards; midventral scutes well
developed and continuous . . . . . . . . . . . → 2 Fig. 1 Photoblepharon palpebratus
2a. Second dorsal fin with 16 soft rays; anal fin with II spines and 12 soft rays; 2 supramax-
illa; lateral-line scales 30 (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . Parmops coruscans
2b. Second dorsal fin with 14 or 15 soft rays; anal fin with II spines and 10 or 11 soft rays; 1
supramaxillae; lateral-line scales 57 to 60 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3
3a. First dorsal fin with IV or V spines; gill rakers on first arch 28 to 34; light organ length
35.4% head length (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anomalops katoptron
3b. First dorsal fin with VI spines; gill rakers on first arch 21, including bony plates; light organ
length 14.5% head length . . . . . . . . . . . . . . . . . . . . . . . . . Protoblepharon rosenblatti
Fig. 2 Parmops coruscans Fig. 3 Anomalops katoptron

Anomalops katoptron (Bleeker, 1856)
Parmops coruscans Rosenblatt and Johnson, 1991
Parmops sp. nov. (Fiji)
Photoblepharon palpebratus (Boddaert, 1781)
Protoblepharon rosenblatti Baldwin, Johnson, and Paxton, 1997
References
Baldwin, C.C., G.P. Johnson, and J.R. Paxton. 1997. Protoblepharon rosenblatti, a new genus and species of flashlight
fish (Beryciformes: Anomalopidae) from the tropical South Pacific, with comments on anomalopid phylogeny.
Proc. Biol. Soc. Wash., 110(3):373-383.
Rosenblatt, R. H. and G.D. Johnson. 1991. Parmops coruscans, a new genus and species of flashlight fish
(Beryciformes: Anomalopidae) from the south Pacific. Proc. Biol. Soc. Wash., 104(2):328-334.
2214 Bony Fishes
MONOCENTRIDIDAE
Pineapple fishes
by J.R. Paxton
D iagnostic characters: Moderate-sized

(to 26 cm) beryciform fishes; body
short, deep, and compressed. Head large
and deep; deep mucous cavities on top of
head separated by thin, spinous ridges and
covered by skin. Eyes moderate to large,
somewhat larger than snout length. Snout
protruding slightly beyond mouth. Mouth
large, jaws extending to or somewhat be-
hind eyes; 1 supramaxilla. Teeth small, in
bands on jaws and palatine, present or
absent on vomer. Gill rakers lath-like. Two
dorsal fins, first with IV to VII strong free
spines alternating side to side, second
with 9 to 12 soft rays; anal fin with no
spines and 10 to 12 soft rays; pelvic fins
subthoracic with I very large spine and 2
to 4 small soft rays: pectoral fins with 13 to
15 rays. Scales large, non-overlapping,
plates with central keel and posterior spine; ventral scales forming keel between pelvic and anal
fins with lateral groove for pelvic-fin spine. No lateral line. Light organ on lower jaw, either on anterior
just below tip or on side under eye. Total vertebrae 26 to 28. Colour: yellow with black between scale
plates, lateral light organ (Cleidopus) red.
Habitat, biology, and fisheries: Demersal from 3 m to about 300 m. Feeding mode as plankton pickers or
carnivores on crustaceans. Uncommon shelf fishes popular in aquaria, but of no other commercial importance.
Remarks: Two genera and 3 species known only in the Indian and Pacific Oceans, except the Northeast
Pacific, in tropical and temperate latitudes.
Berycidae: no greatly enlarged free spines in dorsal or pelvic fins; scales not greatly enlarged and plate-like;
body colour not yellow.
Holocentridae: no greatly enlarged free spines in dorsal or pelvic fins; anal fin with spines; body colour not yellow.
Berycidae Holocentridae

Cleidopus gloriamaris DeVis, 1882
Monocentris japonica (Houttuyn, 1782)
Reference
Kotlyar, A.N. 1985. Taxonomy and distribution range of the family Monocentridae (Beryciformes). Vopr. Ikhtiol.
25(4):531-545. [in Russian, English transl. J. Ichthyol. 25(4)]
Beryciformes: Trachichthyidae 2215
TRACHICHTHYIDAE
Slimeheads (roughies)
by J.A. Moore and J.R. Paxton
D iagnostic characters: Medium-sized (to about 60 cm) beryciform fishes; body oval, laterally com-
pressed. Head large, extensive mucous cavities separated by spinous ridges and covered with
membranous skin; flat, triangular spine on preopercle. Eyes moderate to large in diameter, longer or
shorter than snout length. Mouth large, obliquely angled when closed; 1 supramaxilla. Teeth small, in
villiform bands on jaws, present or absent on vomer, present on palatine. One dorsal fin with III to
VIII striated spines and 10 to 19 soft rays; anal fin with II or III spines and 8 to 12 soft rays; caudal fin
forked, with 4 to 6 procurrent spines in upper and lower lobes; pelvic fins with I spine and 6 soft rays;
pectoral fins with 11 to 20 rays. Scales thick and ctenoid or thin and cycloid, adherent to very deciduous.
Lateral-line scales usually more or less enlarged. Enlarged scales along midventral part of belly
between pelvic and anal fins, forming row of well-developed scutes. Light organs in a few species.
Colour: generally reddish orange, pinkish, or dusky silver.
Habitat, biology, and fisheries: Occurring near bottom in depths of 2 to 1 500 m (most commonly between
150 and 650 m, but abundant aggregations of certain species occur between 700 and 1 200 m). Can be
very abundant locally. Feed primarily on small crustaceans. Extensive fisheries for orange roughy
(Hoplostethus atlanticus) in New Zealand, Australia, Namibia, west of Britain, and the mid-Atlantic Ridge,
where marketed fresh, exported frozen, or made into fishmeal. Useful oils and waxes are also extracted
from this species. Local abundance and recent trends towards development of deep-water fisheries make
other species of potential commercial importance.
Remarks: At least 40 species in 8 genera are currently recognized, with a number of undescribed species
awaiting description. The family occurs in slope waters of all oceans and shallow waters of Australia and
New Zealand. The most comprehensive reviews are that of Kotlyar (1980 and 1996), but some of the
Pacific species still require revision, and more species in the area are expected.

Anomalopidae: large light organ under eye in all
species (may be obscured by curtain of dark skin or
rotation inward); midventral scutes more numerous
and smaller than those of trachichthyids.
Anomalopidae
2216 Bony Fishes
Berycidae: no spine on preopercle; 2 supramaxillae (1 in Trachichthyidae); anal-fin spines IV (II or III in

Trachichthyidae); pelvic fin with 7 to 12 soft rays (6 in Trachichthyidae).
Diretmidae: no spine on preopercle; no dorsal-or anal-fin spines; no lateral line.
Berycidae Diretmidae
Key to the genera of Trachichthyidae occurring in the area

1a. Anus with dark ring between pelvic fins, anterior to abdominal scutes; striated area
before pectoral-fin base and over abdominal scutes and anal fin (Fig. 1) . . . . . . Aulotrachichthys
1b. Anus just anterior to anal fin, posterior to abdominal scutes; no striated areas on body
(except for Hoplostethus metallicus) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2
2a. Body depth less than 40% of standard length (Fig. 2) . . . . . . . . . . . . . . . . . . . . . Optivus
2b. Body depth greater than 40% of standard length . . . . . . . . . . . . . . . . . . . . . . . . . → 3
Fig. 1 Aulotrachichtys Fig. 2 Optivus
3a. Dorsal fin with III spines; dorsal and anal fins sickle-shaped with elongate anterior soft
rays (Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachichthys
3b. Dorsal fin with IV to VIII spines; dorsal and anal fins rounded to emarginate, anterior
soft rays not elongated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4
4a. Lateral-line scales enlarged and diamond-shaped (Fig. 4) . . . . . . . . . . . . . . . Hoplostethus

4b. Lateral-line scales not enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5
Fig. 3 Trachichthys Fig. 4 Hoplostethus

Beryciformes: Trachichthyidae 2217
5a. Dorsal fin with VIII (very rarely VII) spines; body scales with flat, triangular, posteriorly
projecting spines (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gephyroberyx
5b. Dorsal fin with V spines; body scales with small, round, laterally projecting spines
(Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parinoberyx
Fig. 5 Gephyroberyx Fig. 6 Parinoberyx
List of species occuring in the area

Aulotrachichthys latus (Fowler, 1938)
Aulotrachichthys sp.
Gephyroberyx darwini (Johnson, 1866)
Gephyroberyx philippinus Fowler, 1938
Hoplostethus crassispinus Kotlyar, 1980
Hoplostethus japonicus Hilgendorf, 1879
Hoplostethus mediterraneus Cuvier, 1829
Hoplostethus melanopterus Fowler, 1938
Hoplostethus melanopus (Weber, 1913)
Hoplostethus metallicus Fowler, 1938
Hoplostethus sp.
Optivus sp.
Parinoberyx horridus Kotlyar, 1984
Trachichthys australis Shaw, 1799
References
Kotlyar, A.N. 1980. Systematics and distribution of trachichthyid fishes (Trachichthyidae, Beryciformes) of the Indian
Ocean. Trudy Inst. Okeanol. Akad. Nauk S.S.S.R. 110:177-224. [in Russian]
Kotlyar, A.N. 1996. Beryciform fishes of the world ocean. Moscow, VNIRO Publishing, 368 p. [in Russian]
2218 Bony Fishes
BERYCIDAE
Alfonsinos
by J.R. Paxton
D iagnostic characters: Moderate-sized (to 60 cm) beryciform fishes, body moderately deep to very
deep, compressed. Head moderately large; large, deep mucous cavities on top of head separated
by thin ridges and covered by skin; small spines present or absent on snout, over eyes and behind end
of jaw; anterior edge of nasal bone on snout dentate (Centroberyx) or smooth (Beryx); cheeks and
opercles with scales; opercle crenate or smooth, with 1 or 2 flat keels, no opercular spines; no spine on
preopercle. Eyes very large, its diameter greater than snout length. Mouth large, oblique, jaws not
reaching posterior margin of eyes; maxilla expanded posteriorly; lower jaw protrudes anteriorly
beyond upper; 2 supramaxillae. Teeth small, in villiform bands on jaws, vomer and palatine. One dorsal
fin with III to VII spines and 11 to 20 soft rays; anal fin with III or IV spines and 12 to 30 soft rays;
caudal fin distinctly forked, with 17 or 18 principal rays; pelvic fins with I spine and 7 to 13 soft rays;
pectoral fins with 13 to 19 rays. Branchiostegal rays 7 or 8. Scales spinose, very rough to touch; lateral
line with 36 to 82 scales, extending onto caudal fin (Beryx) or not (Centroberyx); enlarged scales between
pelvic and anal fins present (Centroberyx) or absent (Beryx). Gill rakers lath-like, total of 22 to 34 on first
gill arch. No light organs present. Total vertebrae 24 or 25. Colour: basic colour pinkish red on head,
body and fins, with silvery, golden, yellow, or white in smaller proportions.
Habitat, biology, and fisheries: Benthic or benthopelagic fishes of shelf and slope to depths of at least
1 300 m; often found over seamounts; vertical migration into shallower water at night for some species.
Feeding mode as carnivores on mesopelagic crustaceans, squids, and fishes. The species of Beryx are
commercially important in many areas of the world, while at least 1 species of Centroberyx is commercially
important, primarily in eastern Australia. They are usually taken in bottom trawls, or on longlines.
Remarks: Two genera and 9 or 10 species throughout the world ocean in temperate and tropical latitudes,
except the Northeast Pacific.
Beryciformes: Berycidae 2219

Trachichthyidae: pelvic fins with I spine and 6 soft rays; anal fin with II or III spines and 8 to 12 soft rays;
1 supramaxilla.
Pempheridae: pelvic fins with I spine and 5 soft rays; very long anal fin; narrow caudal peduncle; colour
bronze, tan, or cream, never pink or red.
Trachichthyidae Pempheridae
Key to the species of Berycidae occurring in the area

1a. Dorsal fin with III to V spines; soft anal-fin rays 25 to 30; pelvic fins with I spine and 9 to
13 soft rays; lateral-line scales 60 to 82, extending onto caudal fin; 1 or 2 small spines
present on either side of snout, most pronounced in juveniles . . . . . . . . . . . . . . . . . . . ® 2
1b. Dorsal fin with VI or VII spines; soft anal-fin rays 12 to 16; pelvic fins with I spine and 7
soft rays; lateral-line scales 36 to 62, not extending onto caudal fin; no spine present on
snout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Dorsal fin with 12 to 15 soft rays; head with only 1 spine on snout, most pronounced in
juveniles; body moderately deep, its depth about equal to head length and 2.4 to 2.8
times in standard length to base of caudal fin (Fig. 1) . . . . . . . . . . . . . . . . . Beryx splendens
2b. Dorsal fin with 16 to 20 soft rays; head with 4 spines, 2 on snout, 1 over eye, 1 behind
end of jaw, most pronounced in juveniles; body very deep, its depth much greater than
head length and 1.9 to 2.5 tim es in standard length to base of caudal (Fig. 2) . . Beryx decadactylus
Fig. 1 Beryx splendens Fig. 2 Beryx decadactylus

2220 Bony Fishes
3a. Anal fin with 12 soft rays; lateral-line scales 40 to 44; 3 rows of scales on cheek (Fig. 3)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centroberyx affinis
3b. Anal fin with 15 to 17 soft rays; lateral-line scales 53 to 62; 5 rows of scales on cheek
(Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centroberyx druzhinini
Fig. 3 Centroberyx affinis Fig. 4 Centroberyx druzhinini

0
0
Beryx decadactylus Cuvier, 1829
Beryx splendens Lowe, 1834
0
0
Centroberyx affinis (Günther, 1859)
Centroberyx druzhinini (Busakhin, 1981)
References
Busakhin, S.V. 1982. Systematics and distribution of the family Berycidae (Osteichthyes) in the World Ocean. Vopr.
Ikhtiol., 22(6):904-921. [in Russian, English transl. J. Ichthyol., 22(6)]
Kotlyar, A.N. 1996. Beryciform fishes of the world ocean. Moscow, VNIRO Publishing, 368 p. [in Russian]
Beryx decadactylus Cuvier, 1829

Frequent synonyms / misidentifications: None / Beryx splendens Lowe, 1834.
FAO names: En - Alfonsino; Fr - Beryx commun; Sp - Alfonsino palometón.
Diagnostic characters: Body very deep, greatest depth 1.9

to 2.5 times in standard length and significantly more than
head length. Two small spines on snout, 1 over eye and 1
behind end of jaw, less distinct in larger fish. Dorsal fin with
III to V spines and 16 to 20 soft rays, first 3 dorsal-fin rays
may be elongate in smaller specimens; anal fin with III or IV
spines and 25 to 30 soft rays; pelvic fins with I spine and 9 or 10
soft rays, first pelvic-fin ray may be very elongate in smaller
specimens, all rays very elongate in some specimens; pec-
toral fins with 14 to 18 rays. Lateral line with 59 to 73 scales,
extending onto caudal fin. Pyloric caecae 61 to 98. Colour:
scale free portion
upper head and body, all fins and iris of eye bright red, becoming
silvery-pink below.
Size: Maximum total length about 60 cm, commonly to about 35 cm; weight to 2.5 kg.
Habitat, biology, and fisheries: Benthopelagic fishes of outer shelf (180 m) and slope to at least 1 000 m,
moving further from the bottom at night; often found over seamounts; juveniles pelagic. Commercially
important in some areas (Madeira, Indian Ocean), where it is caught by both trawlers and longliners and
marketed mostly fresh. In most areas it is both less common and less desirable than Beryx splendens.
Distribution: Circumglobal,
except Northeast Pacific; the
limited number of records in
the area is doubtless the re-
sult of limited fishing effort be-
low 200 m.
2222 Bony Fishes
Beryx splendens Lowe, 1834

Frequent synonyms / misidentifications: None / Beryx decadactylus Cuvier, 1829; B. mollis Abe, 1959.
FAO names: En - Slender alfonsino; Fr - Beryx long; Sp - Alfonsino besugo.
Diagnostic characters: Body depth moderate, greatest depth 2.4 to 2.8 times in standard length and
equal to or slightly more than head length. One small spine on snout, less distinct in larger
specimens, none elsewhere on head. Dorsal fin with III to V spines and 12 to 15 soft rays, first
dorsal-fin rays may be elongate in smaller specimens; anal fin with IV spines and 25 to 30 soft rays;
pelvic fins with I spine and 9 to 13 soft rays; pectoral fins with 15 to 19 rays. Lateral line with 69 to 82
scales, extending onto caudal fin. Pyloric caecae 23 to 34. Colour: upper head and body, all fins and
iris of eye bright red, becoming silvery-pink below.
Size: Maximum total length 60 cm.
Habitat, biology, and fisheries: Benthopelagic fishes of outer shelf (180 m) and slope to at least 1 300 m,
probably moving further from the bottom at night; often found over seamounts; juveniles pelagic. Com-
mercially important in many areas (Madeira, Japan, New Caledonia), where it is caught by both trawlers
and longliners and marketed mostly fresh. In most areas, this is both the more common and the preferred,
of the 3 species of Beryx.
Distribution: Circumglobal,
except Northeast Pacific and
Mediterranean Sea; the lim-
ited number of records in the
area is doubtless the result of
limited fishing effort below
200 m.
Centroberyx affinis (Günther, 1859)

Frequent synonyms / misidentifications: None / Centroberyx gerrardi (Günther, 1887).
FAO names: En - Nannygai.
Diagnostic characters: Cheek with 3 rows of scales. Pelvic-fin origin before dorsal-fin origin. Dorsal
fin with VI or VII spines and 11 to 13 soft rays; anal fin with IV spines and 12 soft rays; pelvic fins with
I spine and 7 soft rays; pectoral fins with 13 or 14 rays. Lateral line with 40 to 44 scales, not extending
onto caudal fin. Colour: upper head and body, caudal fin, and iris of eye orange-red, becoming
silvery-pink below, with horizontal golden-silver lines sometimes present; other fins pink.
Size: Maximum total length 50 cm (angling record 2 kg); present catches average less than 20 cm fork
length.
Habitat, biology, and fisheries: Juveniles in estuaries, adults on shelf and upper slope, rarely to a depth
of 700 m; adults benthic, moving into water column at night; largest trawl catches between 100 and 300 m.
Commercially important off southeastern Australia, where it is caught by bottom trawlers and marketed
fresh. Previously more than 2 000 t/year were caught, but present catches are less than 1 000 t/year.
Distribution: Eastern Austra-
lia from the western edge of
Bass Strait (previous records
from southern and south-
western Australia based on
misidentifications) and mid-
way down the east coast of
Tasmania north to just north
of Brisbane, Queensland;
New Zealand around the
North Island, rare south of
Cook Strait; recently recorded
from the Chesterfield Islands
and New Caledonia.
2224 Bony Fishes
Centroberyx druzhinini (Busakhin, 1981)

Frequent synonyms / misidentifications: None / Centroberyx lineatus (Cuvier, 1829).
FAO names: En - Druzhinin’s alfonsino.
Diagnostic characters: Cheek with 5 rows of scales. Pelvic-fin origin under dorsal-fin origin. Dorsal
fin with V to VII spines and 12 to 15 soft rays; anal fin with IV spines and 15 to 17 soft rays; pelvic fins
with I spine and 7 soft rays; pectoral fins with 13 rays. Lateral line with 53 to 62 scales, not extending
onto caudal fin. Colour: head and upper body and all fins except pectoral fins bright red, becoming silvery
white below; pectoral fins pink; iris of eye yellow.
Habitat, biology, and fisheries: A benthic or benthopelagic fish trawled in 100 to 300 m. Only a few
records have been published, nothing is known of its biology, and the few specimens captured do not
indicate a potential commercial fishery.
Distribution: Described from
the western Indian Ocean,
and recorded from Japan, the
Sulu Sea, and more recently
New Caledonia.
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Beryciformes: Holocentridae 2225
HOLOCENTRIDAE
Squirrelfishes (soldierfishes)
by J.E. Randall and D.W. Greenfield
D iagnostic characters: Body ovate to moderately elongate; body compressed; caudal peduncle
slender; ridges and mucous channels dorsally on head; edges of external bones of head serrate
or with spines. Eyes very large. Mouth terminal or with lower jaw projecting; gape slightly to strongly
oblique; mouth moderately large, the maxilla extending posterior at least to a vertical at front edge of pupil
(often beyond middle of eye); upper jaw protractile; 2 supramaxillae present; small villiform teeth in bands
in jaws and on roof of mouth (on vomer, palatines, and for some species on ectopterygoids). Branchiostegal
rays 8. Dorsal-fin base long, base of spinous portion 2 to 4 times that of soft portion; dorsal fin with XI to
XIII (rarely XIII) stout spines and 12 to 17 soft rays, deeply or completely notched between spinous and
soft portions or between last 2 dorsal-fin spines; anal fin with IV spines, the third stoutest and often
longest, and 7 to 16 soft rays; pelvic fins with I spine and 7 soft rays; caudal fin forked with 17
branched rays. Lateral-line complete, the pored scales 25 to 56; scales of body strongly ctenoid;
surface of scales smooth or with ridges. Colour: usually red or pink, scale centres of body often lighter
(may be silvery white), thus forming longitudinal bands; black pigment may be present on opercular
membrane or as markings in fins; fins may also have yellow or white markings.
spiny portion of dorsal fin

much longer than soft portion
eyes large
nasal fossa
external bones
of head serrate
or with spines IV spines
in anal fin
pelvic fins with I
spine and 7 soft rays
Habitat, biology, and fisheries: Most of the squirrelfishes (subfamily Holocentrinae) and soldierfishes
(subfamily Myripristinae) live in relatively shallow water on coral reefs or rocky bottoms, but a few
(particularly the genus Ostichthys) occur in depths of 200 m or more. The holocentrid fishes are believed
to be derived from deep-dwelling berycoid ancestors. Their large eyes suit them well for their nocturnal
habits. The family is also well known for sound production. The squirrelfishes feed mainly on crustaceans
living on or near the bottom, whereas the soldierfishes of the genus Myripristis feed on the larger elements
of the zooplankton. The preopercular spine of at least some species of Sargocentron is venomous.
Although wounds from these spines may be very painful, they are not as serious as those from the dorsal-fin
spines of most scorpionfishes. Many of the holocentrids are too small to be of any commercial value; the
largest are frequently seen in local markets, but rarely in abundance.

None. The serrate bony edges and spines on the head, in combination with the large eyes, the very long
base on the spinous portion (as compared to soft portion) of the dorsal fin, the presence of IV spines in
the anal fin and of 7 soft rays in the pelvic fins, readily distinguishes the squirrelfishes and soldierfishes
from other fish families occurring in the area.
2226 Bony Fishes
Key to the genera of Holocentridae occurring in the area

1a. Corner of preopercle either rounded or without
a sharp spine (at most with a short, very broad
spine) (Fig. 1a); front of swimbladder
bifurcate, each anterior projection connected
to an auditory bulla of skull; anal-fin rays 10 to
16 (rarely 10, except for Ostichthys delta) . .
. . . . . . . . . . . . . . (subfamily Myripristinae) → 2
1b. Corner of preopercle with a sharp stout spine,
l o n g e r than broad ( Fig. 1b ); front o f
swimbladder blunt, with no connection to
auditory bullae of skull; anal-fin rays 7 to 10 sharp spine
(rarely 10, except in Sargocentron spiniferum a) Myripristinae b) Holocentrinae
and S. ensifer) . . . . . . (subfamily Holocentrinae) → 5 Fig. 1 lateral view of head
2a. Dorsal-fin spines XI; first gill arch with 19 to 33 gill rakers on lower limb; scales finely to
moderately ctenoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis
2b. Dorsal-fin spines XII (except Ostichthys delta with XI); first gill arch with 11 to 18 gill rakers
on lower limb; scales coarsely ctenoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3
3a. Premaxillary groove long and narrow, nasal bone margins parallel (Fig. 2a); vertebrae 29 . . Pristilepis
(a single species, P. oligolepis (Whitley), in the genus; known from Western Australia, Lord Howe Island,
Japan, and Easter Island, but not yet recorded from the area)
3b. Premaxillary groove either broadly V-shaped or rhomboidal (Fig. 2b, c); vertebrae 26 or
27 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4
premaxillary premaxillary premaxillary
groove long and groove groove
narrow V-shaped rhomboidal
a) Pristilepis b) Ostichthys c) Plectrypops

Fig. 2 dorsal view of head
4a. Premaxillary groove broadly V-shaped; longest dorsal-fin spine 1.9 to 2.6 times in head
length; lateral-line scales 28 to 38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostichthys
4b. Premaxillary groove narrow and rhomboidal; longest dorsal-fin spine 2.5 to 2.9 times in
head length; lateral-line scales 40 to 42 (in Indo-Pacific species) . . . . . . . . . . . . . . Plectrypops
(a single species, P. lima, occurring in the area)
5a. Last dorsal-fin spine shortest, equidistant between penultimate spine and first soft ray
1/
(Fig. 3a); lower jaw not projecting (except in S. spiniferum); ground colour red . . . . Sargocentron
5b. Last dorsal-fin spine longer than penultimate spine, much closer to first soft ray than to
2/
penultimate spine (Fig. 3b); lower jaw projecting; ground colour silvery to silvery pink . . Neoniphon
last spine shortest
a) Sargocentron b) Neoniphon
Fig. 3 dorsal fin

1/ The genus Adioryx is a junior synonym.
2/ The genus Flammeo is a junior synonym.
Key to the species of Myripristis occurring in the area

1a. No scales in axil of pectoral fins (except occasionally in M. numerous small scales
amaena, then only a few small scales ventrally in axil);
lateral-line scales 27 to 44 . . . . . . . . . . . . . . . . . . . . . . → 2
1b. Numerous small scales (Fig. 4a) or 1 or 2 large scales
(Fig. 4b) in axil of pectoral fins (except Marquesan
specimens of M. berndti which lack scales in axil of pectoral
fins, or have only a few); lateral-line scales 25 to 32 . . . . . . . . → 12 a) Myripristis hexagona
2a. Two pairs of symphyseal tooth patches, one above the

other, at tip of lower jaw just outside gape (Fig. 5) (lower pair
usually absent in specimens smaller than about 9 cm
standard length); peritoneum black; lateral-line scales 27 to
29; total gill rakers on first gill arch 30 to 38 . . . . . . . . . . . . . → 3
b) Myripristis adusta
2b. A single pair of symphyseal tooth patches at tip of lower jaw
just outside gape (occasional large specimens of M.
Fig. 4 axil of pectoral fin
chryseres from Hawaii, and specimens of M. trachyacron,
with a second smaller lower pair); peritoneum translucent (after Randall and Greenfield, 1996)
whitish; lateral-line scales 28 to 44; total gill rakers on first
gill arch 29 to 48 . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4
3a. Interorbital not very narrow, its width 4.5 to 5.65 times in head length; caudal-peduncle
length 6.2 to 7.25 times in standard length; first gill arch with 10 to 13 gill rakers on upper
limb (total gill rakers on first gill arch 32 to 38); a prominent black spot distally on caudal-fin
lobes and elevated portions of soft dorsal and anal fins . . . . . . . . . . . . . . . Myripristis botche
(western Pacific and Indian oceans)
3b. Interorbital very narrow, its width 5.75 to 5.95 times in head length; caudal-peduncle
length 8.4 to 9.1 times in standard length; first gill arch with 10 gill rakers on upper limb
(total gill rakers on first gill arch 30 to 32); no distinct black spot distally on caudal-fin lobes
and elevated portions of soft dorsal and anal fins (but some black pigment may be present
on rays at these locations) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis robusta
(Philippines and Papua New Guinea)
4a. A broad dark reddish brown bar from upper end of gill opening to base and axil of pectoral
fins; outer part of spinous portion of dorsal fin yellow to yellow-orange; dorsal-fin rays 15
to 17 (modally 16, rarely 15); anal-fin rays 14 to 16 (modally 15); lateral-line scales 37 to
44 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis kuntee
(Indo-Pacific)
4b. No continuous broad dark reddish brown bar from upper end of gill opening to base and
axil of pectoral fins; outer part of spinous portion of dorsal fin red or red with white spine
tips; dorsal-fin rays 12 to 16 (modally 14 or 15); anal-fin rays 10 to 15 (modally 12 to 14);
lateral-line scales 28 to 40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5
5a. Lateral edge of frontal bone (in dorsal view) above anterior half of orbit slightly convex
(Fig. 6a); longest anal-fin spine 1.3 to 1.55 times in head length; no dark pigment on
opercular membrane or axil of pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6
5b. Lateral edge of frontal bone (in dorsal view) above anterior half of orbit slightly concave
(Fig. 6b); longest anal-fin spine 1.55 to 2.8 times in head length; dark pigment present on
opercular membrane and axil of pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7
slightly concave
convex
2 pairs of tooth patches a) Myripristis trachyacron b) Myripristis formosa
Fig. 5 frontal view of head Fig. 6 dorsal view of head
2228 Bony Fishes
6a. Lateral-line scales 31 to 33; total gill rakers on first gill arch 38 to 41; third anal-fin spine
very long, 1.3 to 1.55 times in head length; dorsal surface of frontal bone next to edge of
orbit obliquely striated anteriorly and rugose (from small nodules) posteriorly; leading
edge of soft dorsal, anal, caudal, and pelvic fins not white; dorsal-fin spines not tipped with
white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis trachyacron
(Indo-Malayan region)
6b. Lateral-line scales 35 to 40; total gill rakers on first gill arch 33 to 38; third anal-fin spine
1.55 to 1.8 times in head length; dorsal surface of frontal bone next to edge of orbit smooth
to slightly rugose; leading edge of soft dorsal, anal, caudal, and pelvic fins white; dorsal-fin
spines tipped with white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis vittata
(Indo-Pacific)
7a. Third dorsal-fin spine clearly longer than fourth spine (both measured from base of scaly
sheath), its tip usually extending beyond fourth spine when depressed . . . . . . . . . . . . . . . → 8
7b. Third dorsal-fin spine shorter than or about equal to fourth spine (both measured from
base of scaly sheath), its tip not extending beyond fourth spine when depressed . . . . . . . . . → 9
8a. Lateral-line scales 32 to 38; total gill rakers on first gill arch 33 to 38; pair of symphyseal
tooth patches outside gape in lower jaw separated by less than a tooth-patch diameter,
the narrow anterior part of lower jaw bearing these teeth fitting into a distinct notch in upper
jaw; median and pelvic fins largely bright yellow in life . . . . . . . . . . . . . . Myripristis chryseres
(Indo-Pacific)
8b. Lateral-line scales 28 to 31; total gill rakers on first gill arch 41 to 45; pair of symphyseal
tooth patches outside gape in lower jaw separated by a space equal to or greater than a
tooth-patch diameter, the anterior part of jaw bearing these teeth not fitting into a distinct
notch in upper jaw; median and pelvic fins mainly red with white leading edges . . Myripristis randalli
(Pitcairn, Austral Islands, Samoa, and Tonga)
9a. Total gill rakers on first gill arch 39 to 48 (modally 44); dorsal-fin rays 13 or 14 (modally
14); anal-fin rays 12 or 13 (modally 12); lateral-line scales 30 to 34 (modally 32); a small
white spot in life at top of pectoral-fin base, adjacent to black or deep red of axil of pectoral
fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis woodsi
(French Polynesia to Micronesia)
9b. Total gill rakers on first gill arch 34 to 42 (modally 37 or 38); dorsal-fin rays 13 to 17
(modally 15); anal-fin rays 12 to 15 (modally 13 or 14); lateral-line scales 32 to 40; no white
spot at top of pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10
10a. Anal-fin rays modally 14; posterior edge of nasal fossa strongly serrate; dark pigment of
opercular membrane not extending ventral to notch below opercular spine; leading edges
of soft dorsal, anal, and caudal fins white . . . . . . . . . . . . . . . . . . . . . . Myripristis pralinia
(Indo-Pacific)
10b. Anal-fin rays modally 13; posterior edge of nasal fossa smooth or with a few small serrae;
dark pigment of opercular membrane extending ventral to notch below opercular spine;
leading edges of soft dorsal, anal, and caudal fins not white or very narrowly white . . . . . . . . → 11
11a. Lateral-line scales 35 to 40 (modally 37); cteni on scale edges 27 to 47; body elongate for
the genus, its depth 2.5 to 3 times in standard length; a distinct striped pattern on body in
life and in preservative . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis tiki
(Easter Island, Pitcairn Group, and Rapa)
11b. Lateral-line scales 32 to 36 (modally 34); cteni on scale edges 45 to 94; body less elongate,
its depth 2.3 to 2.7 times in standard length; body not distinctly striped . . . . . . . . Myripristis amaena
(islands of Oceania)
12a. A single large scale (rarely 2) in lower half of axil of pectoral fins; outer third of caudal-fin
lobes and elevated part of soft dorsal and anal fins black; spinous dorsal fin reddish black
with a median whitish band; scales rimmed with black dorsally on body, and salmon pink
on sides and ventrally; largest species of the genus (to 25 cm standard length) . . . Myripristis adusta
(Indo-Pacific)
12b. Numerous small scales in ventral part of axil of pectoral fins; outer part of median fins not
black; scales rimmed with brownish red, red, or pink (except dorsally on M. violacea
where edged in black) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 13
13a. Two pairs of tooth patches, one above the other, at front of lower jaw outside gape in
specimens larger than about 9 cm standard length; lateral-line scales 25 to 29 (modally
27); leading edges of caudal-fin lobes, soft dorsal, anal, and pelvic fins not white . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Myripristis hexagona
(Samoa to East Africa)
13b. A single pair of tooth patches at front of lower jaw just outside gape; lateral-line scales 27
to 32 (modally 28 to 30); leading edge of soft dorsal, anal, caudal, and pelvic fins white . . . . . → 14
14a. Scales dorsally on body with broad dark brown to black edges, those on nape almost
completely black; modally 14 gill rakers on upper limb of first gill arch . . . . . . . Myripristis violacea
(Indo-Pacific)
14b. Scales dorsally on body without dark brown to black edges; modally 10 to 13 gill rakers
on upper limb of first gill arch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →15
mucous channel deep
15a. Mucous channel on each side of anterior interorbital
space deep (Fig. 7); first gill arch with 10 gill rakers on
upper limb (total gill rakers on first gill arch 32); scales of
body strongly silvery in life . . . . . . . . . . . Myripristis aulacodes
(known from a single specimen taken off Lombok, Indonesia)
15b. Mucous channel on each side of anterior interorbital
space not deep; first gill arch with 11 to 15 gill rakers on
dorsal view of head
upper limb (total gill rakers on first gill arch 35 to 44);
scales of body not strongly silvery in life . . . . . . . . . . . . → 17 Fig. 7 Myripristis aulacodes
16a. Interorbital space narrow, its width 4.5 to 5.2 times in head length; lower jaw of adults
strongly projecting when mouth is closed; modally 12 gill rakers on upper limb of first gill
arch; outer part of spinous dorsal fin yellow or yellow-orange in life . . . . . . . . . Myripristis berndti
(Indo-Pacific and tropical eastern Pacific)
16b. Interorbital space broad, its width 3.65 to 4.4 times in head length; lower jaw of adults
slightly projecting when mouth closed; modally 13 gill rakers on upper limb of first gill arch;
outer part of spinous dorsal fin red in life . . . . . . . . . . . . . . . . . . . . . . Myripristis murdjan
(Samoa and Marshall islands to East Africa and Red Sea)
Key to the species of Neoniphon occurring in the area

1a. Scales above lateral line to base of middle dorsal-fin spines 3 ½; lateral-line scales 42 to
47; silvery pink with yellow stripes on body . . . . . . . . . . . . . . . . . . Neoniphon aurolineatus
(Indo-Pacific)
1b. Scales above lateral line to base of middle dorsal-fin spines 2 ½; lateral-line scales 36 to
43; no yellow stripes on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 2
2a. First dorsal fin without dark markings; pectoral-fin rays usually 13 . . . . . . . . Neoniphon argenteus
(Indo-Pacific)
2b. Spinous portion of dorsal fin with a conspicuous black spot or large black area; pectoral-fin
rays usually 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 3
2230 Bony Fishes
3a. A black spot about as large as eye between first and third dorsal-fin spines; anal-fin rays 8
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoniphon sammara
(Indo-Pacific)
3b. A broad black median zone across entire spinous portion of dorsal fin; anal-fin rays
usually 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neoniphon opercularis
(Indo-Pacific)
Key to the species of Ostichthys occurring in the area

1a. Scales above lateral line to middle of spinous portion of dorsal fin 3 ½ . . . . . . . . . . . . . . . → 2
1b. Scales above lateral line to middle of spinous portion of dorsal fin 2 ½ . . . . . . . . . . . . . . . → 5
2a. Anterior end of each nasal bone with a sharp forward-directed spine projecting to or
beyond median part of upper lip; a short stout spine at corner of preopercle; gill rakers on
first gill arch 10 to 12 on upper limb and 16 to 18 on lower limb . . . . . . . Ostichthys acanthorhinus
(Gulf of Oman to southern Indonesia)
2b. Anterior end of nasal bones without a forward-directed spine (except juveniles); no spine
at corner of preopercle notably larger than spinules along entire margin; gill rakers on first
gill arch 6 to 11 on upper limb and 12 to 17 on lower limb . . . . . . . . . . . . . . . . . . . . . . → 3
3a. Lateral-line scales 38; dorsal fin with XI spines and 16 rays; pectoral-fin rays 15 . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostichthys ovaloculus
(known from a single specimen taken off Tahiti at a depth of about 300 m)
3b. Lateral-line scales 28 to 30; dorsal fin with XII spines and 12 to 14 rays; pectoral-fin rays
16 or 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 4
4a. Last dorsal-fin spine 2 to 3 times longer than penultimate spine; depth of second suborbital
bone (measured vertically below eye centre) about 1/2 orbit diameter; dorsal profile of
head convex; pectoral-fin rays usually 17; first gill arch with 12 to 14 gill rakers on lower
limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostichthys japonicus
(western Pacific and Andaman Sea)
4b. Last 2 dorsal-fin spines subequal; depth of second suborbital bone about 1/3 to 1/4 orbit
diameter; dorsal profile of head to above rear edge of eye nearly straight; pectoral-fin rays
usually 16; first gill arch with 15 or 16 gill rakers on lower limb . . . . . . . . . . . . Ostichthys sandix
(Society Islands and Hawaii)
5a. Anal-fin origin below anterior soft portion of dorsal fin; anal-fin rays 9 to 11 (usually 10);
lateral-line scales 27 or 28 (usually 27); upper procurrent spiniform caudal-fin rays 5, and
lower procurrent spiniform caudal-fin rays 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6
5b. Anal-fin origin below posterior spinous portion of dorsal fin; anal-fin rays 11 or 12 (usually
11); lateral-line scales 28 to 30; upper procurrent spiniform caudal-fin rays 4, and lower
procurrent spiniform caudal-fin rays 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7
6a. Dorsal fin with XI spines and 13 or 14 rays; vomerine teeth in a near-triangular patch; head
length 2.5 to 2.65 times in standard length; longest dorsal-fin spine 2.05 to 2.3 times in head
length; first gill arch with 7 or 8 gill rakers on upper limb, 13 or 14 on lower limb . . . . . Ostichthys delta
(Réunion and Samoa)
6b. Dorsal fin with XII spines and 12 rays; vomerine teeth in a V-shaped patch; head length
2.45 times in standard length; longest dorsal-fin spine 2.6 times in head length; first gill
arch with 8 gill rakers on upper limb, 15 on lower limb . . . . . . . . . . . . Ostichthys brachygnathus
(known from a single specimen taken off Guam at a depth of 230 m)
7a. A half scale directly anterior to upper half of first pored scale of lateral line; pectoral-fin
rays 15; dorsal profile of head straight to above posterior edge of preopercle; snout length
3.7 to 4.3 times in head length . . . . . . . . . . . . . . . . . . . . . . . . Ostichthys archiepiscopus
(Réunion, Tahiti, southern Japan, and Hawaii)
7b. No half scale anterior to upper half of first pored scale of lateral line; pectoral-fin rays 15
to 17 (strongly modal as 16); dorsal profile of head slightly convex, especially region of
nape; snout length 4.2 to 4.9 times in head length . . . . . . . . . . . . . . . . . Ostichthys kaianus
(Réunion, northwestern Australia, and Indonesia to southern Japan)
Key to the species of Sargocentron occurring in the area

1a. Scales above lateral line to base of middle dorsal-fin spines 3 ½ . . . . . . . . . . . . . . . . . . → 2
1b. Scales above lateral line to base of middle dorsal-fin spines 2 ½ . . . . . . . . . . . . . . . . . . → 7
2a. Lateral-line scales 57 to 58; eyes very large, 2.6 times in head length . . . . . Sargocentron megalops
(known from a single 7.9 cm specimen, Henderson Island, Pitcairn Group)
2b. Lateral-line scales 38 to 48; eyes not very large, the orbit diameter 2.85 to 4.8 times in
head length (in specimens more than 7 cm standard length) . . . . . . . . . . . . . . . . . . . . → 3
3a. Anal-fin rays 10 (rarely 9); membranes of spinous portion of dorsal fin not or only slightly
incised; maximum standard length greater than 20 cm . . . . . . . . . . . . . . . . . . . . . . . → 4
3b. Anal-fin rays 9 (rarely 8); membranes of spinous portion of dorsal fin distinctly incised;
maximum standard length about 8.5 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 5
4a. No small spines at edge of nasal fossae; dorsal-fin rays usually 15; preopercular spine of
adults usually greater than orbit diameter; first dorsal-fin spine about 3/4 length of second
spine; body very deep, its depth 2.4 to 2.6 times in standard length; a large oval dark red
spot on preopercle behind eye; spinous portion of dorsal fin uniform dark red . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron spiniferum
(Indo-Pacific)
4b. One or more small spines at edge of nasal fossae (Fig. 8a); dorsal-fin rays usually 14;
preopercular spine of adults less than orbit diameter; first and second dorsal-fin spines
subequal; body not very deep, its depth 2.5 to 2.8 times in standard length; no large dark
red spot behind eye; spinous portion of dorsal fin yellow with a red margin . . . Sargocentron ensifer
(Hawaii, Japan, New Caledonia, Pitcairn, and Paracel Islands in South China Sea)
1 or more small retrorse

small spines recumbent spine
spine
slightly
retrorse,
a) b) c) blunt spine d)
5a. No spine on nasal bone between edge of nasal fossa and premaxillary groove; lateral-line
scales 42; pectoral-fin rays 14; total gill rakers on first gill arch 20 or 21 . . . . Sargocentron shimizui
(2 specimens: Sulawesi, Indonesia)
5b. A small recumbent spine on nasal bone between edge of nasal fossa and premaxillary
groove (Fig. 8b); lateral-line scales 43 to 49; pectoral-fin rays usually 15; total gill rakers
on first gill arch 17 to 20 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6
2232 Bony Fishes
6a. A retrorse spine at edge of premaxillary groove (Fig. 8c); body very deep, its depth 2.3 to
2.6 times in standard length; length of upper jaw 2.6 to 2.9 times in head length . . Sargocentron iota
(Hawaii, Fiji, New Caledonia, Chesterfield Bank, Palau, Papua New Guinea, and Indonesia)
6b. No spine at edge of premaxillary groove; body not very deep, its depth 2.65 to 2.9 times
in standard length; length of upper jaw 2.35 to 2.6 times in head length . . . . . Sargocentron lepros
(Pitcairn, Cook Islands, Samoa, Rotuma, and Christmas Island in Indian Ocean)
7a. A retrorse spine on nasal bone between nasal fossa and premaxillary groove (Fig. 8c);
edge of premaxillary groove with a small retrorse spine (occasionally 2 spines); largest
specimen, 11.3 cm standard length . . . . . . . . . . . . . . . . . . . . . . Sargocentron inaequalis
(Comoro Islands, Chagos Archipelago, Seychelles, and Line Islands, Central Pacific)
7b. No spine on nasal bone between nasal fossa and premaxillary groove; no spine at edge
of premaxillary groove (except S. microstoma); maximum standard length 13 cm or more . . . . . → 8
8a. Lateral-line scales 32 to 38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9

8b. Lateral-line scales 38 to 55 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 14
9a. A serrate ridge on upper edge of suborbital bones, preceded by a laterally projecting,
slightly retrorse, blunt spine on suborbital below or slightly posterior to anterior edge of
orbit (Fig. 8d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10
9b. No serrate ridge along upper edge of suborbital bones (or only a few small serrae below
posterior part of orbit), and without a laterally projecting blunt spine on suborbital below
anterior edge of orbit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 13
10a. A large black blotch on first 2 membranes of dorsal fin; tips of caudal-fin lobes broadly
rounded; body light orange-red with narrow darker red stripes following scale edges; no
blackish spot at base of soft dorsal, anal, and caudal fins; lateral-line scales modally 33
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron dorsomaculatum
(Ryukyu Islands, Palau, and Caroline Islands)
10b. No blackish blotch on first 2 membranes of dorsal fin (or if blackish pigment is present on
these membranes, it is also present on remaining spinous membranes of fin); tips of
caudal-fin lobes pointed to slightly rounded; body alternately striped with red and silvery
white; a blackish spot at base of soft dorsal, anal, and caudal fins (may be faint at base of
anal and caudal fins); lateral-line scales modally 34 to 36 . . . . . . . . . . . . . . . . . . . . . → 11
11a. Spinous portion of dorsal fin largely blackish red in life (dark brown in preservative);
upper-jaw length 2.75 to 2.95 times in head length; body depth 2.7 to 3 times in standard
length; spinules present on edge of nasal fossae . . . . . . . . . . . . . . . Sargocentron cornutum
(East Indies and Great Barrier Reef)
11b. Spinous portion of dorsal fin largely red in life (not dark brown in preservative); upper-jaw
length 2.35 to 2.75 times in head length; body depth 2.4 to 2.8 times in standard length;
no spinules present on edge of nasal fossae (or rarely a single spinule in large S. praslin) . . . . → 12
12a. Oblique rows of scales on cheek 4; lateral-line scales modally 34; least interorbital width
3.9 to 4.4 times in head length; upper-jaw length 2.35 to 2.55 times in head length; dark
pigment on pelvic fins mainly on first soft ray . . . . . . . . . . . . . . . . . . Sargocentron praslin
(East Africa to Society Islands)
12b. Oblique rows of scales on cheek 5; lateral-line scales modally 35; least interorbital width
4.5 to 5.2 times in head length; upper-jaw length 2.5 to 2.75 times in head length; dark
pigment on pelvic fins confined to distal part of second to sixth (or fewer) membranes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron rubrum
(Red Sea and Gulf of Oman to western Pacific)
13a. Dorsal-fin rays modally 14; membranes of spinous portion of dorsal fin not incised; body
depth 2.3 to 2.6 times in standard length; no blackish spot at base of median fins; body
dark brownish to purplish red with a silvery white vertical line on each scale; a black spot
on upper opercular membrane . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron violaceum
(islands of western Indian Ocean to Society Islands)
13b. Dorsal-fin rays modally 13; membranes of spinous portion of dorsal fin incised; body depth
2.6 to 3 times in standard length; a large black oval spot at base of soft portion of dorsal
fin, a second smaller spot at base of caudal fin, and usually a third spot at base of soft
portion of anal fin; body orange-red to orange-yellow with silvery white stripes; no black
spot on opercular membrane . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron melanospilos
(East Africa to Marshall Islands and Samoa)
14a. Lateral-line scales 38 to 43; preopercular-spine length usually more than 2/3 orbit
diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 15
14b. Lateral-line scales 41 to 55; preopercular-spine length less than 2/3 orbit diameter (except
S. tiere) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 16
15a. Premaxillary groove not reaching

or just reaching a vertical at
anterior edge of orbit (Fig. 9a);
dorsal-fin rays usually 14; oblique
rows of scales on cheek 5; edge of
nasal fossa usually with 1 or more
spinules; body red without stripes,
the edges of scales narrowly
a) Sargocentron b) Sargocentron
silver; a prominent silvery white caudimaculatum tiereoides
spot anterodorsally on caudal
peduncle (posterior third of body
silvery white on some individuals) Fig. 9 dorsal view of head
. . . . . . . . Sargocentron caudimaculatum
(Indo-Pacific)
15b. Premaxillary groove extending distinctly posterior to a vertical at anterior edge of orbit
(Fig. 9b); dorsal-fin rays usually 13; oblique rows of scales on cheek 4; edge of nasal fossa
usually without spinules; body with alternating stripes of red and silvery white; no silvery
white spot on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron tiereoides
(Indo-Pacific)
16a. Dorsal-fin spines short, the longest 2.6 to 3.5 times in head length; preopercular spine of
adults about equal to orbit diameter; head and body primarily red (faint silvery stripes on
body, those on lower part with blue iridescence); maximum standard length 27 cm . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron tiere
(islands of western Indian Ocean to Hawaii and Pitcairn)
16b. Dorsal-fin spines not short, the longest 1.5 to 2.6 times in head length; preopercular spine
of adults less than 2/3 orbit diameter; body distinctly striped with red and silvery white;
maximum standard length 17.5 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 17
17a. Lateral-line scales 41 to 49 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 18

17b. Lateral-line scales 48 to 56 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 20
18a. Pectoral-fin rays usually 14; membranes of spinous portion of dorsal fin reddish black
(black in preservative) with a disjunct longitudinal whitish band (lower in anterior part of fin
and higher in posterior part) and whitish tips . . . . . . . . . . . . . . . . . . Sargocentron diadema
(Indo-Pacific)
18b. Pectoral-fin rays usually 15; membranes of spinous portion of dorsal fin not reddish black
(if a whitish longitudinal band is present, it is not interrupted) . . . . . . . . . . . . . . . . . . . → 19
2234 Bony Fishes
19a. Snout very short, its length less than interorbital width; body usually peppered with small
blackish dots; outer part of membranes of spinous portion of dorsal fin red; no blackish
blotch on first 1 or 2 interspinous membranes of dorsal fin; lateral-line scales 41 to 47
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron punctatissimum
(Indo-Pacific)
19b. Snout not short, its length equal to or greater than interorbital space; body without
numerous small blackish dots; tips of membranes of spinous portion of dorsal fin white
(except posteriorly); preserved specimens often with a large blackish blotch on first 1 or 2
interspinous membranes of dorsal fin; lateral-line scales 43 to 49 . . . . . . . . Sargocentron ittodai
(Indo-Pacific)
20a. Third anal-fin spine very long, its length 1 to 1.2 times in head length; longest dorsal-fin
spine 1.7 to 1.95 times in head length; spinous portion of dorsal fin completely separate
from soft portion; silvery white stripes on body broader than red stripes (except for narrow
white stripe following lateral line) . . . . . . . . . . . . . . . . . . . . . . . Sargocentron microstoma
(islands of western Indian Ocean to French Polynesia)
20b. Third anal-fin spine not very long, 1.2 to 1.45 times in head length; longest dorsal-fin spine
1.95 to 2.2 times in head length; spinous portion of dorsal fin linked by last membrane to
base of exposed part of first dorsal-fin ray; red stripes on body much broader than silvery
white stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sargocentron hormion
(Pitcairn Group, Rapa, and Rarotonga in Cook Islands)

Subfamily MYRIPRISTINAE
0Myripristis adusta Bleeker, 1853
0Myripristis amaena (Castelnau, 1873)
0Myripristis berndti Jordan and Evermann, 1903
Myripristis aulacodes Randall and Greenfield, 1996
0Myripristis botche Cuvier, 1829

0Myripristis chryseres Jordan and Evermann, 1903
0Myripristis hexagona (Lacepède, 1802)
0Myripristis kuntee Valenciennes, 1831
0Myripristis murdjan (Forsskål, 1775)
0Myripristis pralinia Cuvier, 1829
0Myripristis randalli Greenfield, 1974
0Myripristis tiki Greenfield, 1974
Myripristis robusta Randall and Greenfield, 1996
0Myripristis trachyacron Bleeker, 1863

0Myripristis violacea Bleeker, 1851
0Myripristis vittata Cuvier, 1831
Myripristis woodsi Greenfield, 1974
0
Ostichthys acanthorhinus Randall, Shimizu, and Yamakawa, 1982
Ostichthys archiepiscopus (Valenciennes, 1862)
Ostichthys brachygnathus Randall and Myers, 1993
0
Ostichthys delta Randall, Shimizu, and Yamakawa, 1982
0
Ostichthys japonicus (Cuvier, 1829)
Ostichthys kaianus (Günther, 1880)
Ostichthys ovaloculus Randall and Wrobel, 1988
Ostichthys sandix Randall, Shimizu, and Yamakawa, 1982
Plectrypops lima (Valenciennes, 1831)
Subfamily HOLOCENTRINAE
Neoniphon argenteus (Valenciennes, 1831)
0Neoniphon opercularis (Valenciennes, 1831)

Neoniphon aurolineatus (Liénard, 1839)
0Neoniphon sammara (Forsskål, 1775)

0Sargocentron caudimaculatum (Rüppell, 1835)
Sargocentron cornutum (Bleeker, 1853)
Sargocentron diadema (Lacepède, 1802)
Sargocentron dorsomaculatum (Shimizu and Yamakawa, 1979)
Sargocentron ensifer (Jordan and Evermann, 1904)
Sargocentron hormion Randall, 1998
Sargocentron inaequalis Randall and Heemstra, 1985
Sargocentron iota Randall, 1998
Sargocentron ittodai (Jordan and Fowler, 1903)
Sargocentron lepros (Allen and Cross, 1983)
Sargocentron melanospilos (Bleeker, 1858)
Sargocentron megalops Randall, 1996
0
Sargocentron microstoma (Günther, 1859)
Sargocentron praslin (Lacepède, 1802)
0
Sargocentron punctatissimum (Cuvier, 1829)
0
Sargocentron rubrum (Forsskål, 1775)
0
Sargocentron spiniferum (Forsskål, 1775)
Sargocentron tiere (Cuvier, 1829)
Sargocentron tiereoides (Bleeker, 1853)
Sargocentron violaceum (Bleeker, 1853)
References
Greenfield, D.W. 1978. A revision of the squirrelfish genus Myripristis Cuvier (Pisces: Holocentridae). Sci. Bull. Nat.
Hist. Mus. Los Angeles County, (19):54 p.
Randall, J.E. 1998. Revision of the Indo-Pacific squirrelfishes (Beryciformes: Holocentridae: Holocentrinae) of the genus
Sargocentron, with descriptions of four new species. Indo-Pac. Fishes, (27):105 p.
Randall, J.E. and D.W. Greenfield. 1996. Revision of the Indo-Pacific holocentrid fishes of the genus Myripristis , with
descriptions of three new species. Indo-Pac. Fishes, (25):61 p.
Randall, J.E. and P. Guézé. 1981. The holocentrid fishes of the genus Myripristis of the Red Sea, with clarification of
the murdjan and hexagonus complexes. Contr. Sci. Nat. Hist. Mus. Los Angeles County, (334):16 p.
Randall, J.E. and P.C. Heemstra. 1985. A review of the squirrelfishes of the subfamily Holocentrinae from the western
Indian Ocean and Red Sea. Ichthyol. Bull., (49):27 p.
Randall, J.E., T. Shimizu, and T. Yamakawa. 1982. A revision of the holocentrid fish genus Ostichthys, with descriptions
of four new species and a related genus. Japan. J. Ichthyol., 29(1):1-26.
Shimizu, T. and T. Yamakawa. 1979. Review of the squirrelfishes (subfamily Holocentrinae: order Beryciformes) of
Japan, with description of a new species. Japan. J. Ichthyol., 26(2):109-147.
2236 Bony Fishes
Myripristis adusta Bleeker, 1853

FAO names: En - Shadowfin soldierfish; Fr - Marignan ombré; Sp - Candil sombreado.
Diagnostic characters: Body oblong, its depth 2.1 to 2.55 times in standard length. Usually a single pair
of tooth patches at front of lower jaw just outside nape (rarely a smaller second pair below). Dorsal
fin with XI spines and 14 to 16 soft rays, notched to back between last 2 spines; eleventh spine more than
twice length of tenth; anal fin with IV spines (third spines strongest, but fourth spine longest) and 12 to 14
soft rays. Lateral-line scales 27 to 29 (usually 28); a single scale (occasionally 2) on lower part of
pectoral-fin axil. Total gill rakers on first gill arch 35 to 40. Colour: scale centres pale bluish dorsally,
pale pink on sides and ventrally, the scale edges dark brown to black dorsally and orangish brown to
salmon pink on sides and ventrally; a large blackish spot posteriorly on gill cover; distal third of soft
dorsal fin, anal fin, and caudal-fin lobes black; spinous dorsal fin reddish black with a broad median
whitish band.
Size: The largest of the soldierfishes; maximum total length about 32 cm, commonly to 25 cm.
Habitat, biology, and fisheries: Occurs on coral reefs and rocky areas in relatively shallow water; seeks
refuge in caves and beneath ledges by day, often in aggregations. Usually caught at night with gill nets or
hook-and-line; sometimes
speared during daylight
hours.
Distribution: Indo-Pacific ex-
cept the Red Sea, Arabian
Sea, and Persian Gulf in the
Indian Ocean region and Ha-
waii and Pitcairn Group in the
Pacific.
Myripristis amaena (Castelnau, 1873)

Frequent synonyms / misidentifications: Myripristis argyromus Jordan and Evermann, 1904 / None.
FAO names: En - Brick soldierfish.
(after Randall and Greenfield, 1996)
Diagnostic characters: Body depth 2.3 to 2.7 times in standard length; interorbital width 3.4 to 4 times in
head length. Lower jaw not projecting when mouth closed. A single pair of tooth patches at front of
lower jaw outside mouth. Dorsal fin with XI spines and 13 to 16 (usually 15) soft rays, notched to back
between last 2 spines; eleventh spine more than twice length of tenth; anal fin with IV spines (third and
fourth spines subequal, the third most stout) and 12 to 15 (usually 13) soft rays. Lateral-line scales 32 to
36 (modally 34). Axil of pectoral fins naked. Total gill rakers on first gill arch 34 to 41. Colour: red,
the centres of scales pale red dorsally and silvery white on sides and ventrally, thus forming a longitudinal
banding; black on opercular membrane extending well below opercular spine; median fins red without
conspicuous white leading edges.
Size: Largest specimen 26 cm, commonly to 18 cm.
Habitat, biology, and fisheries: Nocturnal, like others of the genus, hiding in caves and beneath ledges
by day. Occurs in shallower water, in general, than other species. Usually caught by gill nets or by spearing.
Distribution: Islands of
Oceania, including Hawaii
and Pitcairn Group.
2238 Bony Fishes
Myripristis berndti Jordan and Evermann, 1903

Frequent synonyms / misidentifications: None / Myripristis murdjan (Forsskål, 1775).
FAO names: En - Blotcheye soldierfish; Fr - Marignan à oeillères; Sp - Candil ojo manchado.
Diagnostic characters: Body depth 2.3 to 2.65 times in standard length; interorbital narrow, its width 4.5
to 5.2 times in head length. Lower jaw of adults strongly projecting when mouth closed. A single pair
of tooth patches at front of lower jaw outside mouth; vomerine teeth in a triangular patch with rounded
corners. Dorsal fin with XI spines and 13 to 15 (usually 14) soft rays, notched to back between last 2 spines;
eleventh spine more than twice length of tenth; anal fin with IV spines (third and fourth spines subequal,
the third strongest) and 11 to 13 (usually 12) soft rays. Lateral-line scales 28 to 31 (modally 29). Small
scales on lower half to three-quarters of pectoral-fin axil. Total gill rakers on first gill arch 35 to 42.
Colour: red, the scale centres pale red dorsally and silvery white on sides and ventrally; opercular
membrane black to well below opercular spine; axil of pectoral fins black; outer third to half of spinous
dorsal fin yellow to orange-yellow; leading edges of caudal-fin lobes, soft dorsal, anal, and pelvic fins
white.
Size: Maximum total length 29 cm, commonly to 22 cm.
Habitat, biology, and fisheries: Among the most common of the larger species of the genus; generally
found at moderate depths of 10 to 30 m; a cave resident by day. Caught mainly by gill nets, hook-and-line,
and by spear.
has the broadest distribution
of the genus, from the coast of
East Africa (but not the Red
Sea or Persian Gulf) to is-
lands of the tropical eastern
Pacific; in the western Pacific
it ranges from southern Japan
to Lord Howe Island.
Myripristis botche Cuvier, 1829

Frequent synonyms / misidentifications: Myripristis melanostictus Bleeker, 1863 / Myripristis
hexagona (non Lacepède, 1802).
FAO names: En - Blacktip soldierfish; Fr - Marignan potelé; Sp - Candil rechoncho.
Diagnostic characters: Body robust, its depth 2.1 to 2.45 times in standard length; interorbital width 4.5
to 5.65 times in head length. Lower jaw of adults slightly projecting when mouth closed. Two pairs of tooth
patches at front of lower jaw outside mouth; vomerine teeth in a triangular patch with rounded corners.
Dorsal fin with XI spines and 13 or 15 (usually 14) soft rays, notched to back between last 2 spines; eleventh
spine more than twice length of tenth; anal fin with IV spines (third spine stoutest, but fourth usually slightly
longer) and 11 or 12 (usually 12) soft rays. Lateral-line scales 27 to 29 (modally 28). No small scales
in pectoral-fin axil. Total gill rakers on first gill arch 32 to 38. Colour: silvery white, the scales rimmed
with brownish red dorsally and red on sides and ventrally; black pigment of opercular membrane extending
to about level of lower edge of eye; outer part of spinous dorsal fin and most of soft dorsal and anal fins
red; tips of soft dorsal and anal fins and caudal-fin lobes with a black spot, the leading edges of
these fins white.
ies: A species of moderate to
relatively deep water (gener-
ally more than 30 m). Not
common, but of importance
as a food fish due to its larger
size than most species of
Myripristis.
Distribution: Coast of East
Africa to the western Pacific,
east to New Caledonia.
2240 Bony Fishes
Myripristis chryseres Jordan and Evermann, 1903

FAO names: En - Yellowfin soldierfish.
Diagnostic characters: Body depth 2.3 to 2.6 times in standard length; interorbital narrow, its width 4.55
to 5.2 times in head length. Lower jaw of adults slightly projecting when mouth closed. A single pair of
tooth patches on knob-like front of lower jaw outside mouth, the knob fitting into a distinct notch
in upper jaw when mouth closed. Dorsal fin with XI spines and 14 or 15 (usually 14) soft rays, notched
to back between last 2 spines; eleventh spine more than twice length of tenth; anal fin with IV spines (third
anal-fin spine longer and stronger than fourth) and 11 to 13 (usually 12) soft rays. Lateral-line scales
32 to 38 (modally 34). No small scales in pectoral-fin axil. Total gill rakers on first gill arch 33 to 39.
Colour: red, the scale centres lighter; opercular membrane black to well below opercular spine; axil of
pectoral fins black; soft dorsal, anal, and caudal fins largely bright yellow.
Habitat, biology, and fisheries: Generally found at depths greater than 30 m; seeks refuge in caves or
crevices by day.
Distribution: East Africa to
the Samoa Islands and Ha-
waii, but not reported from
many intervening localities.
Its occurrence in deeper
water is no doubt the reason
for the relatively few records.
Myripristis kuntee Valenciennes, 1831

Frequent synonyms / misidentifications: Myripristis borbonicus Valenciennes, 1831; M. multiradiatus
Günther, 1874 / None.
FAO names: En - Shoulderbar soldierfish; Fr - Marignan ardoisé; Sp - Candil de lomo manchado.
Diagnostic characters: Body depth 2.2 to 2.6 times in standard length; interorbital width 3 to 5 times in
head length. Lower jaw of adults slightly projecting when mouth closed. A single pair of tooth patches
at front of lower jaw outside mouth. Dorsal fin with XI spines and 15 to 17 (usually 16) soft rays,
notched to back between last 2 spines; eleventh spine more than twice length of tenth; anal fin with IV
spines (third spine stoutest, but fourth slightly longer) and 14 to 16 (usually 15) soft rays.
Lateral-line scales 37 to 44. No small scales in pectoral-fin axil. Total gill rakers on first gill arch 33
to 41. Colour: light red, the edges of the scales darker than centres; a continuous reddish brown bar
from upper end of gill opening to axil of pectoral fins; outer part of spinous dorsal fin orange-yellow;
red pigment in soft dorsal and anal fins concentrated in a large spot at tip of fins.
Habitat, biology, and fisheries: A coral-reef species generally found in relatively shallow water. Often
seen in aggregations in and at the entrance to caves. Although one of the most common of soldierfishes,
its small size reduces its value as a food fish.
Distribution: A widespread
Indo-Pacific species (though
absent from the Red Sea and
Persian Gulf).
Remarks: Myripristis kuntee
is similar in counts to another
common small species, M.
pralinia ( s e e a b b r e v i a t ed
species account below), but
the dark pigment on the oper-
cular membrane of the latter
extends only slightly below
the opercular spine.
2242 Bony Fishes
Myripristis murdjan (Forsskål, 1775)

Frequent synonyms / misidentifications: Myripristis parvidens Cuvier, 1829; M. axillaris Valenciennes,
1831 / None.
FAO names: En - Pinecone soldierfish; Fr - Marignan pomme de pin; Sp - Candil piñón.
Diagnostic characters: Body depth 2.3 to 2.5 times in standard length; interorbital width narrow, 3.65 to
4.4 times in head length. Lower jaw of adults slightly projecting when mouth closed. A single pair of tooth
patches at front of lower jaw outside mouth; vomerine teeth in a triangular patch with rounded corners.
Dorsal fin with XI spines and 13 to 15 (usually 14) soft rays, notched to back between last 2 spines; eleventh
spine more than twice length of tenth; anal fin with IV spines (third and fourth spines subequal, the third
stoutest) and 11 to 14 (usually 12 or 13) soft rays. Lateral-line scales usually 28 to 30 (modally 29).
Small scales on lower one-fourth to three-fourths of pectoral axil. Total gill rakers on first gill arch
36 to 44. Colour: light red, the edges of scales reddish brown; opercular membrane dark brown to black
to about level of middle of eye; axil of pectoral fins dark brown to black; spinous dorsal fin light red on
basal two-thirds, bright red on outer third; leading edges of soft dorsal, anal, caudal, and pelvic fins
white, sometimes with blackish pigment submarginal to the white on soft dorsal and anal fins; a blackish
blotch on iris above pupil, and a lesser blotch below.
Habitat, biology, and fisheries: A common coral-reef species, usually found in less than 10 m, but known
to depths of at least 50 m. Nocturnal like others of the genus, feeding mainly on larger animals of the
zooplankton.
Distribution: Red Sea and
coast of East Africa to the
Marshall and Samoa islands;
in the western Pacific from
southern Japan to New South
Wales.
Myripristis violacea Bleeker, 1851

Frequent synonyms / misidentifications: Myripristis microphthalmus Bleeker, 1852 / None.
FAO names: En - Violet soldierfish; Fr - Marignan violacé; Sp - Candil guru-guru.
Diagnostic characters: Body moderately deep, its depth 2.15 to 2.45 times in standard length; interorbital
width 3.6 to 4.1 times in head length. Lower jaw of adults slightly projecting when mouth closed. A single
pair of tooth patches at front of lower jaw outside mouth. Dorsal fin with XI spines and 13 to 16 (usually
15) soft rays, notched to back between last 2 spines; eleventh spine more than twice length of tenth; anal
fin with IV spines (third and fourth spines subequal, the third strongest) and 12 to 14 (usually 13) soft rays.
Lateral-line scales 27 to 29 (modally 28). Small scales usually present on lower half of pectoral-fin
axil. Total gill rakers on first gill arch 38 to 48. Colour: scale centres bluish silver (sometimes pinkish
silver), the edges of scales above lateral line dark brown to black (broadest on nape); lateral-line
scales rimmed with reddish brown, and scales below lateral line edged with brownish red; opercular
membrane dark reddish brown, the darker pigment ending at about level of middle of eye; axil of pectoral
fins black; spinous dorsal fin largely bright red; caudal fin and outer part of soft dorsal and anal fins bright
red; leading edges of caudal-fin lobes, soft dorsal, anal, and pelvic fins white (narrow on median fins).
Habitat, biology, and fisheries: A common coral-reef species generally found in shallow protected waters
such as lagoons.
Distribution: I n d o - P a c i f i c
(except the Red Sea, Persian
Gulf, Hawaii, and Pitcairn
Group); in the western Pacific
from the Ryukyu Islands to the
Great Barrier Reef (Australia).
2244 Bony Fishes
Neoniphon opercularis (Valenciennes, 1831)

Frequent synonyms / misidentifications: Flammeo opercularis (Valenciennes, 1831); Holocentrus
opercularis Smith and Smith, 1963; Kutaflammeo opercularis Carcasson, 1977 / None.
FAO names: En - Blackfin squirrelfish; Fr - Marignan aile noire; Sp - Candil de aleta negra.
Diagnostic characters: Body moderately elongate, its depth 3.1 to 3.3 times in standard length. Dorsal
profile of head slightly convex; snout pointed; lower jaw strongly projecting when mouth closed;
preopercular spine short, less than 1/2 eye diameter. Nasal fossa large, without spinules. Dorsal fin with
XI spines, the last longer than the penultimate and close to first soft ray, the soft rays 13; anal fin
with IV spines and 9 soft rays; pectoral-fin rays 13 or 14 (usually 14). Lateral-line scales 38 to 40; scales
above lateral line to middle of spinous portion of dorsal fin 2 ½. Colour: silvery pink (scale centres light
red, the edges broadly silver); snout and top of head light red; spinous dorsal fin black, the membranes
tipped with white, with a diagonal white spot near base of each membrane; lobes of caudal fin broadly
light red, the rest of fin yellowish; soft dorsal and anal fins yellow, tinged with pink; pectoral fins light red;
pelvic fins pinkish white.
Size: Maximum total length about 32 cm, commonly to 23 cm.
Habitat, biology, and fisheries: A reef species which may be found in shallow to moderate depths. The
largest member of the genus Neoniphon. Taken by hook-and-line and in traps. Marketed fresh.
Distribution: Occurs through-
out the Indo-Pacific region (ex-
cept the Red Sea and Hawaii);
in the western Pacific from the
Ryukyu Islands to the Great
Barrier Reef.
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Neoniphon sammara (Forsskål, 1775)

Frequent synonyms / misidentifications: Flammeo sammara (Forsskål, 1775); Holocentrus sammara
Smith and Smith, 1963; Kutaflammeo sammara Carcasson, 1977 / None.
FAO names: En - Sammara squirrelfish; Fr - Marignan tacheté; Sp - Candil samara.
Diagnostic characters: Body moderately elongate, its depth 3 to 3.6 times in standard length. Dorsal
profile of head slightly convex; snout pointed; lower jaw projecting when mouth closed; preopercular
spine short, about 1/3 eye diameter. Nasal fossa large, without spinules. Dorsal fin with XI spines, the
last longer than the penultimate and close to first soft ray, the soft rays 11 or 12 (usually 12); anal
fin with IV spines and 7 or 8 (mostly 8) soft rays; pectoral-fin rays 13 or 14. Lateral-line scales 39 to
43; scales above lateral line to middle of spinous portion of dorsal fin 2 ½. Colour: silvery, each scale with
a horizontally elongate dark reddish brown spot, thus forming longitudinal dark lines on body; a reddish
stripe following lateral line; snout and top of head light red; spinous portion of dorsal fin with a large
black spot tinged with red on membranes between first and fourth spines; caudal fin yellowish with
a broad red band at margins of lobes; a reddish streak anteriorly in soft dorsal and anal fins.
Habitat, biology, and fisheries: Associated with coral reefs; most often seen in shallow water of protected
bays and lagoons. Among the first to leave shelter with the advent of darkness for feeding. About 67% of
the diet consists of crabs and about 15% small fishes. Taken by hook-and-line and in traps. Marketed fresh.
throughout the Indo-Pacific
region, including the Red Sea
and Hawaii; in the western
Pacific from Japan (the young
ranging as far north as
s o uth er n Honshu) to the
southern Great Barrier Reef.
2246 Bony Fishes
Ostichthys archiepiscopus (Valenciennes, 1862)

Frequent synonyms / misidentifications: Ostichthys pillwaxii (Steindachner, 1893) / None.
FAO names: En - Straighthead soldierfish; Fr - Marignan cuirassé; Sp - Candil acorazado.
Diagnostic characters: Body oblong, its depth 2.1 to 2.35 times in standard length; caudal peduncle
slender, its least depth 3.8 to 4.7 times in head length. Dorsal profile of head straight; head length 2.45
to 2.6 times in standard length; snout 3.7 to 4.3 times in head length; no anteriorly directed spine on
nasal bone of adults and no large spine at corner of preopercle. Premaxillary groove broadly V-shaped;
vomerine teeth in a V-shaped patch. First gill arch with 7 to 9 gill rakers on upper limb, 13 to 15 on lower
limb. Dorsal fin continuous though deeply notched between spinous and soft portions; dorsal fin with XII
spines (last spine shortest and close to soft portion of fin) and 13 or 14 soft rays; anal fin with IV
spines (the third much stronger and larger) and 11 soft fin rays; pectoral fins with 15 rays. Lateral-line
scales 28 to 30 (usually 29); rows of scales above lateral line to middle of spinous portion of dorsal
fin 2 ½; a half scale directly anterior to upper half of first lateral-line scale. Colour: light red, the
centres of the scales paler, thus forming faint longitudinal bands.
ies: A benthic species, prob-
ably occurring mainly over
hard substratum; known from
its depth range of 146 to 400
m. Usually caught by hook-
and-line, rarely by trawling.
Distribution: Reported only
from Réunion in the south-
western Indian Ocean (type
locality), the Ryukyu Islands,
Hawaii, and Society Islands,
t h u s s u g g e s t i n g a n a n-
tiequatorial distribution.
Ostichthys japonicus (Cuvier, 1829)

FAO names: En - Japanese soldierfish.
slender, its least depth 4 to 4.5 times in head length. Dorsal profile of head convex; head length 2.35 to
2.5 times in standard length; snout short, 4.65 to 5.6 times in head length; height of second suborbital
bone (measured vertically below eye centre) about 1/2 orbit diameter. No anteriorly directed spine
on nasal bone of adults or subadults; no spine at corner of preopercle of adults (subadults may have a
small spine at angle). Premaxillary groove broadly V-shaped. Vomerine teeth in a V-shaped patch. First
gill arch with 7 to 10 gill rakers on upper limb, 12 to 14 on lower limb. Dorsal fin continuous, though deeply
notched between spinous and soft portions; dorsal fin with XII spines, the last distinctly longer than
penultimate spine, and 12 to 14 (usually 13) soft rays; space between last spine and first ray of dorsal
fin much less than space between last 2 dorsal-fin spines; anal fin with IV spines (the third longest
and stoutest) and 10 to 12 (usually 11) soft rays; pectoral fins with 16 or 17 (usually 17) rays. Lateral-line
scales 28 to 30 (modally 28); rows of scales above lateral line to middle of spinous portion of dorsal
fin 3 ½; no half scale anterior to first lateral-line scale. Colour: edges of scales red, the centres silvery
pink; spinous portion of dorsal fin mottled light red and whitish; remaining fins with light red rays and pale
membranes; iris red.
Habitat, biology, and fisheries: Most specimens have come from fish markets with little or no information
on type of bottom or depth of capture. Those for which the depth is known were taken in 90 to 194 m.
Caught mostly by hook-and-line and marketed fresh.
Distribution: Reported in
Japan (type locality) from the
Ryukyu Islands to Honshu at
31°31’N; Korea; China off
Shanghai and Hong Kong;
South China Sea; Lesser
Sunda Islands, Indonesia;
northwestern Australia; New
S o u th W a l e s a t 3 4 ° 26 ’S
(where named Holotrachys
major W h i t l e y ) , a n d t h e
Andaman Sea.
2248 Bony Fishes
Ostichthys kaianus (Günther, 1880)

FAO names: En - Kai soldierfish.
slender, its least depth 3.8 to 4.6 times in head length. Dorsal profile of head slightly convex; head
length 2.15 to 2.4 times in standard length; snout moderately short, 4.2 to 4.9 times in head length. No
anteriorly directed spine on nasal bone of adults or subadults; no spine at corner of preopercle of
adults (subadults may have a small spine at angle). Premaxillary groove broadly V-shaped. Vomerine teeth
in a V-shaped patch. First gill arch with 7 to 9 gill rakers on upper limb, 14 to 16 on lower limb. Dorsal fin
continuous, though deeply notched between spinous and soft portions; dorsal fin with XII spines, the
last 2 spines subequal, and 12 or 13 (usually 13) soft rays; space between last spine and first ray of
dorsal fin about 1/2 as broad as space between last 2 dorsal-fin spines; anal fin with IV spines (the
third longest and stoutest) and 11 soft rays; pectoral fins with 15 to 17 (usually 16) rays. Lateral-line
scales 28 to 30 (modally 28); rows of scales above lateral line to middle of spinous portion of dorsal
fin 2 ½; no half scale anterior to first lateral-line scale. Colour: red with a series of silvery white dashes
(1 on each scale for nearly its full exposed width) forming longitudinal bands on body; fins pale red; iris
red.
ies: Obtained by hook-and-
line and by trawling in the
depth range of 310 to 640 m.
Distribution: Reported from
Kai Island, eastern Banda
Sea (type locality), South
China Sea, the Ryukyu Is-
lands, northwestern Australia,
a nd R é un i on ( l oc a lity of
Myripristis (Holotrachys)
guezei Postel).
Sargocentron caudimaculatum (Rüppell, 1838)

Frequent synonyms / misidentifications: Adioryx caudimaculatus (Rüppell, 1838); Holocentrus
caudimaculatus Smith and Smith, 1963; Holocentrum andamanense Day, 1870 / None.
FAO names: En - Silverspot squirrelfish; Fr - Marignan rouge et argent; Sp - Candil platero.
Diagnostic characters: Body depth 2.3 to 2.7 times in standard length. Dorsal profile of head slightly
convex; preopercular spine long, its length about 1 eye diameter. Nasal fossa with 1 or 2 spinules on
anterior margin. Dorsal fin with XI spines (last spine shortest) and 13 to 15 (usually 14) soft rays; anal
fin with IV spines and 8 to 10 (usually 9) soft rays; pectoral-fin rays 13 to 15 (usually 14). Lateral-line
scales 38 to 43; scale rows above lateral line to middle of spinous portion of dorsal fin 2 ½. Colour: red,
the scales rimmed posteriorly with silvery white; a saddle-like silvery white spot dorsally on caudal
peduncle; spinous dorsal fin light red to whitish, the outer triangular part of each membrane deep red; no
blackish markings; in life, posterior third of body may be silvery white.
Habitat, biology, and fisheries: A reef species which may be locally abundant. Caught by hook-and-line
and in traps. Marketed fresh.
Distribution: Wide-ranging
in the Indo-Pacific, from the
Red Sea south to Transkei,
South Africa, east to the Mar-
shall Islands and Society Is-
lands; in the western Pacific
from the Ryukyu Islands to the
southern Great Barrier Reef.
2250 Bony Fishes
Sargocentron praslin (Lacepède, 1802)

FAO names: En - Brownspot squirrelfish; Fr - Marignan chocolat; Sp - Candil chocolate.
Diagnostic characters: Body deep, its depth 2.5 to 2.8 times in standard length. Dorsal profile of head
convex; preopercular spine about 3/4 eye diameter; first suborbital bone with 1 or 2 short lateral spines
near upper margin. Nasal fossa without spinules. Dorsal fin with XI spines (last spine shortest) and 12 or
13 (usually 13) soft rays; anal fin with IV spines and 8 or 9 (rarely 8) soft rays; pectoral-fin rays 13 to 15
(usually 14). Lateral-line scales 33 to 36; scale rows above lateral line to middle of spinous portion
of dorsal fin 2 ½; cheek with 4 diagonal rows of scales. Colour: body with alternate stripes of silvery
white and reddish brown; a triangular streak of brownish red on cheek, from eye to corner of preopercle;
often a concentration of pigment forming an elongate brown spot beneath soft portion of dorsal fin
and a roundish blotch above base of soft portion of anal fin, spinous portion of dorsal fin red with a
median band of cojoined whitish spots, the membranes tipped with white; a large dark brown spot in axil
of pectoral fins; front edge of pelvic fins dark.
Habitat, biology, and fisheries: A shallow-water species, more often found in a coral reef environment
than the closely related Sargocentron rubrum.
Distribution: Indo-Pacific,
but exact distribution not
known due to confusion with
S. rubrum. Definite records
from Kenya, Mozambique
(15°S), Aldabra, Comoro Is-
lands, Chesterfield Islands
(Madagascar), Indonesia,
New Guinea, New Britain
(type locality), Solomon Is-
lands (locality of neotype),
Fiji, Vanuatu, Palau, Philip-
pines, Ryukyu Islands, Mari-
ana Islands, Samoa Islands,
and Society Islands.
Sargocentron rubrum (Forsskål, 1775)

Frequent synonyms / misidentifications: Adioryx ruber (Forsskål, 1775); Holocentrus ruber Smith and
Smith, 1963 / None.
FAO names: En - Redcoat; Fr - Marignan rouget; Sp - Candil rubí.
convex, preopercular spine about 3/4 eye diameter; first suborbital bone with 1 or 2 short lateral spines
near upper margin. Nasal fossa without spinules. Dorsal fin with XI spines (last spine shortest) and
12 to 14 (usually 13) soft rays; anal fin with IV spines and 8 to 10 (rarely 10) soft rays; pectoral-fin rays 13
to 15 (usually 14). Lateral-line scales 34 to 38; scale rows above lateral line to middle of spinous
portion of dorsal fin 2 ½; cheek with 5 diagonal rows of scales. Colour: body with alternate stripes of
silvery white and brownish red; a triangular streak of brownish red on cheek from eye to corner of
preopercle; often a concentration of pigment forming an elongate brown spot beneath soft portion of
dorsal fin and a roundish blotch above base of soft portion of anal fin; spinous portion of dorsal fin
red with a median band of cojoined whitish spots, the membranes tipped with white; no dark spot in
pectoral-fin axil; tips of second to fourth soft pelvic-fin rays darker; upper and lower edges of caudal
fin brownish red; rest of caudal fin and soft portions of dorsal and anal fins yellowish.
Habitat, biology, and fisheries: A shallow-water species usually found in protected habitats such as bays
and lagoons, but less often around coral reefs than the closely related Sargocentron praslin.
Distribution: Known from the
Red Sea (type locality), Medi-
terranean Sea (an immigrant
via the Suez Canal), Gulf of
Oman, India, Sri Lanka, An-
daman Sea (off Thailand),
and northwestern Australia,
but remains unreported from
the coast of East Africa or any
islands of the western Indian
Ocean; definite records in the
Pacific include Indonesia, Ma-
laysia, Great Barrier Reef,
New Caledonia, and southern
Japan.
2252 Bony Fishes
Sargocentron spiniferum (Forsskål, 1775)

Frequent synonyms / misidentifications: Holocentrus spiniferus: Smith and Smith, 1963 / None.
FAO names: En - Sabre squirrelfish; Fr - Marignan sabre; Sp - Candil sable.
straight; lower jaw protruding when mouth closed; preopercular spine long, slightly longer than eye
diameter. Nasal fossa without spinules. Dorsal fin with XI spines (last spine shortest) and 14 to 16
soft rays, the membranes of spinous portion only slightly incised; anal fin with IV spines and 9 or 10 (usually
10) soft rays; pectoral-fin rays 14 to 16 (usually 15). Lateral-line scales 41 to 46; scale rows above
lateral line to middle of soft portion of dorsal fin 3 ½. Colour: head and body red, darker dorsally than
ventrally, the scales rimmed with silvery white; a large vertically elongate deep red spot behind eye;
pectoral-fin axil and region above pectoral-fin base deep red; spinous portion of dorsal fin solid deep
red; remaining fins yellowish.
Habitat, biology, and fisheries: Largest of holocentrid fishes. Associated with coral reefs. Like other
squirrelfishes, it tends to hide in caves by day and emerges for foraging with the onset of darkness. Feeds
mainly on crustaceans, especially crabs, but occasionally ingests small fishes. Caught by hook-and-line,
in traps and by spearing. Marketed fresh. Has been implicated in ciguatera poisoning.
the Indo-Pacific from the Red
Sea (type locality) and coast
of East Africa south to Natal
and east to Hawaii and Pit-
cairn Group in Oceania; in the
western Pacific from the
Ruykyu Islands to the south-
ern Great Barrier Reef.
Sargocentron tiere (Cuvier, 1829)

FAO names: En - Tahitian squirrelfish.
Diagnostic characters: Body moderately deep, its depth 2.6 to 2.95 times in standard length; dorsal profile
of head straight to slightly convex; mouth terminal; preopercular spine nearly as long as orbit diameter in
adults; no lateral spinules on first suborbital bone; nasal fossa without spinules at edges. First gill arch
with 7 to 9 gill rakers on upper limb, 13 to 16 on lower limb. Dorsal fin with XI spines and 13 to 15 (usually
14) soft rays; anal fin with IV spines and 9 (rarely 10) soft rays; dorsal-fin spines short, the longest 2.6
to 3.5 times in head length; membranes of spinous portion of dorsal fin slightly incised; pectoral-fin rays
13 to 15. Lateral-line scales 45 to 52; scale rows above lateral line to middle of spinous portion of
dorsal fin 2 ½. Colour: red, the sides and ventral part of body with faint silvery red stripes overlaid with
blue iridescence (particularly the ventral stripes); fins red except white tips of dorsal-fin spines and a white
spot in middle of each interspinous membrane of dorsal fin except first 2 membranes where near the fin
base; leading edge of anal and pelvic fins white.
Habitat, biology, and fisheries: A coral-reef species occurring more often on exposed than sheltered
reefs. It may be seen in as little as 1 to 2 m, but is found at least as deep as 20 m. Feeds at night, mainly
on crabs. At night it displays 2 prominent white bars on its otherwise bright red body.
Distribution: Ranges from
the Pitcairn Group and Hawaii
to islands of the western In-
dian Ocean. It is primarily a
species of oceanic islands,
rather than continental waters.
It remains unknown from the
Red Sea, the coasts of Africa
and Asia, and the islands of
Indonesia. In the western Pa-
cific it is reported from the
Great Barrier Reef, Solomon
Islands, Palau Islands, and the
Ryukyu Islands.
2254 Bony Fishes
Myripristis hexagona (Lacepède, 1802)

En - Doubletooth solderfish.
Maximum standard length about 16 cm. Distributed from Samoa to East Africa; in the western
Pacific it ranges from the Ryukyu Islands in the north to the Great Barrier Reef and New Caledonia
in the south.
Myripristis pralinia Cuvier, 1829

En - Scarlet soldierfish.
Maximum standard length about 16 cm. Generally found in moderately shallow water on coral reefs
in depths of 1 to 30 m. Distributed from the Tuamotu Archipelago and Line Islands to East Africa;
in the western Pacific it ranges from the Ryukyu Islands in the north to the southern Great Barrier
Reef in the south.
Myripristis randalli Greenfield, 1974

En - Randall’s soldierfish.
Maximum standard length about 15 cm. Found at depths of 15 to 53 m. Known from Pitcairn Island,
Austral Islands, American Samoa, and Tonga.

Myripristis tiki Greenfield, 1974

En - Tiki soldierfish.
Maximum standard length about 21 cm. Found at depths of 1.5 to 15 m. Known only from the Easter
Islands, Pitcairn Group, and Rapa.
Myripristis trachyacron Bleeker, 1863

En - Roughskull soldierfish.
Maximum standard length about 12 cm. Apparently occurs in deep water, but was also reported
from shallower waters. Known only from the Indo-Australian Archipelago (Philippines, Indonesia,
and Solomon Islands).
(after Greenfield, 1974)
Myripristis vittata Cuvier, 1831

En - Whitetip soldierfish.
Maximum standard length about 16 cm. Generally found in depths of more than 20 m, but a single
specimen was collected at 3 m; most likely will be found in other locations. Distributed from the
Marquesas Islands, and Hawaii westward to the Mascarene Islands and Seychelles.
2256 Bony Fishes
Myripristis woodsi Greenfield, 1974

En - Wood’s soldierfish.
Maximum standard length about 21 cm. A shallow-water species typically found on exposed
outer-reef areas, more often at low islands and atolls than high islands. Western Central Pacific.
Confirmed records from the Marquesas Islands, Line Islands, Phoenix Islands, Kiribati, Marshall
Islands, Minami tori shima (Marcus Island), and Mariana Islands.
(after Greenfield, 1974)
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Zeiformes: Parazenidae 2257
Order ZEIFORMES
PARAZENIDAE
Parazen
by P.C. Heemstra
D iagnostic characters: Body oblong and compressed (size to 30 cm total length), its depth about
equal to head length, contained 0.9 to 2.1 times in standard length. Eye large, its diameter greater
than snout length and contained 2.8 to 5.2 times in head length. Mouth large, terminal, the upper jaw
highly protrusible; jaws with 1 or 2 rows of short, slender, conical teeth; similar teeth on vomer; maxilla
exposed posteriorly; no supramaxilla. Gill rakers short and flattened, 1 on upper limb and 4 to 6 on lower
limb of first arch (not counting rudiments). Gills 3 ½ (no slit behind last hemibranch). Branchiostegal
membranes separate from isthmus; branchiostegal rays 7. Two dorsal fins, the first with VI to VIII
slender spines, the second dorsal fin with 26 to 30 soft rays; anal fin with I small spine and 28 to
34 soft rays; caudal fin forked, with 11 segmented rays and 9 branched rays; pectoral fins much shorter
than head, with 12 to 16 rays; pelvic fins with 1 unbranched soft ray and 6 branched soft rays (no
spine). Two lateral lines anteriorly, merging to a single line on the caudal peduncle. Head naked; body
scales moderate in size, deciduous and weakly ctenoid. Vertebrae 34. Colour: reddish silver.
Habitat, biology, and fisheries: Caught with trawls in depths of 150 to 600 m.
Remarks: This family comprises a single genus and species. Not reported yet from the WCP area but very
likely to occur there. Common off Japan in the western Pacific and also known from specimens collected
off Tanzania in the western Indian Ocean and Cuba in the western Atlantic.
Macrurocyttidae: eye diameter distinctly longer than snout; pelvic-fin spine well developed.
Zeidae: body depth distinctly greater than head length, contained 0.9 to 2.1 times in standard length.
Macrurocyttidae Zeidae

? Parazen pacificus Kamohara, 1935
Reference
Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino (eds). 1988. The fishes of the Japanese Archipelago.
Second edition. Tokyo, Tokai University Press, 456 p.
2258 Bony Fishes
MACRUROCYTTIDAE
Dwarf dories
by P.C. Heemstra
D iagnostic characters: Body oval or oblong and compressed, its depth contained 2.2 to 2.8 times
in standard length; head length 2 to 2.8 times in standard length. Eye huge, its diameter about
twice snout length, and 30 to 52% head length. Mouth slightly or greatly protrusible; jaws with a row of
minute conical teeth; no teeth on vomer; maxilla exposed posteriorly; no supramaxilla. Gills 3½ (no slit
behind last hemibranch); gill rakers rudimentary low knobs. One dorsal fin, with IV to VII spines and
25 to 30 soft rays, the second spine longest and serrate along its front margin; anal-fin spine
rudimentary or absent, anal-fin rays 25 to 32; caudal fin rounded or almost truncate, with 12 to 16 rays;
pectoral fins much shorter than head, with 12 to 17 rays; pelvic fins with I strong spine and 2, 3 or 6
soft rays. Lateral line complete or replaced by fleshy tubercles. Body scales small, rudimentary, or absent;
head naked. Vertebrae 25 to 34. Colour: Macrurocyttus acanthopodus has the head and front part of body
blackish brown, the rear part of body and fins pale dusky or brownish. Zenion hololepis is dusky silver,
with a reddish sheen; black spot distally on first dorsal fin; caudal fin reddish distally.
Habitat, biology, and fisheries: Taken with trawls over sand or mud bottoms in depths of 330 to 1 140 m.
Biology unknown. These small, rare fishes are of no importance to fisheries.
Remarks: This poorly defined family comprises 2 or 3 genera and 4 or 5 species. The relationship of
Macrurocyttus to Zenion may be more superficial than real.

Parazenidae: eye shorter than snout; pelvic fin with 1
unbranched soft ray, 6 branched soft rays, and no
spine.

Macrurocyttus acanthopodus Fowler, 1934
Zenion hololepis Goode and Bean, 1896 Parazenidae
(= Zenion japonicum Kamohara, 1934)
(= Cyttula macropus Weber, 1913)
Reference
Nyako, C.O. and K. Amaoka. 1996. First records of the zeiform fish, Macrurocyttus acanthopodus Fowler, 1933, from
the Coral Sea. Ichthyol. Res., 43:93-96.
Zeiformes: Zeidae 2259
ZEIDAE
Dories
by P.C. Heemstra
D iagnostic characters: Body oval and strongly compressed; head also strongly compressed;
body depth greater than head length, contained 0.9 to 2.1 times in standard length. Mouth highly
protrusible, forming a downward-pointing tube when protruded; jaws with bands of villiform granular
teeth or minute conical teeth, or rows of small canines; similar teeth on vomer; maxilla exposed posteriorly,
no supramaxilla. Gills 3 ½ (no slit behind the last hemibranch); gill rakers rudimentary low knobs.
Branchiostegal membranes separate from isthmus; branchiostegal rays 7. A single dorsal fin, with VII
to X spines and 20 to 37 soft rays; anal fin with I to IV spines and 20 to 39 soft rays; caudal fin
rounded or almost truncate, the upper lobe with 6 segmented rays and 5 branched rays, the lower lobe
with 7 segmented rays and 6 branched rays; pectoral fins much shorter than head, with 11 to 18 rays;
pelvic fins with I spine and 6 or 7 rays, or lacking the spine and with 6 or 7, or 9 or 10 soft rays.
Lateral line complete, strongly curved over pectoral fin. Body scales small, rudimentary or absent; head
naked (cheek and operculum scaly in Cyttomimus). Swimbladder present. Vertebrae 29 to 42. Colour:
Zenopsis: silvery with faint dusky blotches; Zeus: bronzy copper, with prominent black ocellus at midside;
Cyttopsis: reddish silvery, Cyttomimus: dusky silver.
Cyttopsis rosea
Habitat, biology, and fisheries: Most zeids are caught with trawls over sand or mud bottom in depths of
35 to 600 m. They usually occur near the bottom and feed on fishes and benthic invertebrates (mainly
crustaceans and worms).
Remarks: This poorly defined family comprises 5 or 6 genera and 11 or 12 species. Zeids are rare in
tropical and subtropical waters, but 5 species have been reported from the Western Central Pacific.

Drepanidae: pectoral fins falciform, longer than head, reaching cau-
dal peduncle; pelvic fins with I spine and 5 soft rays.

Cyttomimus affinis Weber, 1913
Cyttopsis cypho (Fowler, 1934)
Cyttopsis rosea (Lowe, 1843)
Zenopsis nebulosa (Temminck and Schlegel, 1847)
Zeus faber Linnaeus, 1758 Drepanidae
Reference
Heemstra, P.C. 1986. Zeidae. In Smith’s sea fishes, edited by M.M. Smith and P.C. Heemstra. Johannesburg, Macmillan
South Africa, pp. 435-438.
2260 Bony Fishes
GRAMMICOLEPIDIDAE
Tinselfishes
by P.C. Heemstra
D iagnostic characters: Body oval, oblong, or diamond-shaped, and strongly compressed, its
depth much greater than head length, contained 0.8 to 2.3 times in standard length; head length
2.9 to 4.4 times in standard length; body, cheeks, and operculum covered with vertically elongated
scales; a row of small spines along each side of dorsal- and anal-fin bases. Mouth small; maxilla
ridged, bound to ascending processes of premaxillae, loosely connected to palatines; jaws with 1 or 2 rows
of small, slender teeth; vomer with or without teeth. Gills 3 ½, no slit behind last hemibranch; gill rakers
rudimentary flat tooth plates attached to skin covering the first gill arch. Branchiostegal rays 7, with
membranes joined to front of isthmus. Two dorsal fins, the first with V to VII slender spines, the second
dorsal fin with 17 to 34 unbranched soft rays; anal fin with II spines and 27 to 39 unbranched soft
rays; juveniles with first anal-fin spine and second dorsal-fin spines greatly elongated; caudal fin
with 13 branched rays; pelvic fins with I spine and 6 branched soft rays. Vertebrae 37 to 46. Colour:
adults silvery (with black spots in Xenolepidichthys).
adult, 60 cm total length

Grammicolepis brachiusculus juvenile, 17 cm total length
Habitat, biology, and fisheries: Apparently rare,

no information available on biology. Caught with
trawls, but of no importance for fisheries.
Remarks: This family comprises 2 genera, each
with a single species. Tinselfishes occur worldwide
(but not in polar seas) at depths of 100 to 800 m.

Caproidae: have small, oval ctenoid scales; branched dorsal-
and anal-fin rays; pelvic fins with I spine and 5 soft rays; caudal
fin with 10 branched rays; branchiostegal rays 6; vertebrae
9+12.
The transparent, compressed, deep-bodied acronurus post-
larva of surgeonfishes (family Acanthuridae) superficially re-
sembles a small tinselfish, but acanthurids have a restricted gill
opening, and the postlarva lacks elongate fin spines.

Grammicolepis brachiusculus Poey, 1873
Caproidae
Xenolepidichthys dalgleishi Gilchrist, 1922
Reference
Karrer, C. and P.C. Heemstra. 1986. Grammicolepididae. In Smith’s Sea Fishes, edited by M.M. Smith and P.C.
Heemstra. Johannesburg, Macmillan South Africa, pp. 440-441.
Zeiformes: Caproidae 2261
CAPROIDAE
Boarfishes
by P.C. Heemstra
D iagnostic characters: Body disc-

shaped, strongly compressed; dorsal
profile angular, ventral profile semicircu-
lar; body depth more than twice head
length and contained 0.8 to 1.4 times in
standard length; head length 2.4 to 3 times
in standard length. Mouth small, oblique; up-
per jaw protrusible; bands of small teeth on
jaws, none on vomer or palatines. Gills 4, a
small slit behind last gill; gill rakers on first gill
arch 15 to 23. Branchiostegal rays 6, the mem-
branes joined far forward. Single dorsal fin
with VII to IX spines and 26 to 38 soft rays;
anal fin with III spines and 24 to 34 soft
rays; dorsal- and anal-fin rays branched;
caudal fin truncate, with 10 branched rays;
pectoral-fin rays 12 to 14; pelvic fins with I
strong spine and 5 branched soft rays.
Scales small, round or slightly oval, with en-
larged ctenii. Vertebrae 9+12. Colour: reddish
orange.
Habitat, biology, and fisheries: Tropical and
temperate waters of all oceans; adults usually
found near the bottom in depths of 50 to
600 m; the larvae are pelagic.
Remarks: Two genera are presently recog-
nized. The monotypic Capros aper (Linnaeus,
1758) is common in the North Atlantic and
Mediterranean. Antigonia comprises 10 spe-
cies, of which 4 are known from the Western
Central Pacific. The placement of this family in
the order Zeiformes is controversial and it is
often considered more closely aligned with the
Perciformes.

Grammicolepididae: scales greatly elongated verti-
cally; dorsal- and anal-fin rays unbranched; pelvic fins
with I spine and 6 branched soft rays; caudal fin with
13 branched rays; brachiostegal rays 7; vertebrae 37
to 46.
juvenile

Antigonia capros Lowe, 1843
Antigonia malayana Weber, 1913
Antigonia rubescens Günther, 1860
Antigonia rubicunda Ogilby, 1910 Grammicolepididae
References
Parin, N.V. and O.C. Borodulina. 1986. Preliminary review of the benthopelagic fish genus Antigonia Lowe (Zeiformes,
Caproidae). Trans. P.P. Shirshov Inst. Oceanol., 121:141-172.
Zehren, S.J. 1987. Osteology and evolutionary relationships of the boarfish genus Antigonia (Teleostei: Caproidae).
Copeia, 1987(3):564-592.
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2262 Bony Fishes
Order GASTEROSTEIFORMES
PEGASIDAE
Seamoths (seadragons)
by T.W. Pietsch and W.A. Palsson
D iagnostic characters: Small fishes (to 18 cm total length); body depressed, completely encased in
fused dermal plates; tail encircled by 8 to 14 laterally articulating, or fused, bony rings. Nasal bones
elongate, fused, forming a rostrum; mouth inferior. Gill opening restricted to a small hole on dorsolat-
eral surface behind head. Spinous dorsal fin absent; soft dorsal and anal fins each with 5 rays, placed
posteriorly on body. Caudal fin with 8 unbranched rays. Pectoral fins large, wing-like, inserted horizon-
tally, composed of 9 to 19 unbranched, soft or spinous-soft rays; pectoral-fin rays interconnected by broad,
transparent membranes. Pelvic fins thoracic, tentacle-like, with I spine and 2 or 3 unbranched soft rays.
Colour: in life highly variable, apparently capable of rapid colour change to match substrata; head and
body light to dark brown, olive-brown, reddish brown, or almost black, with dorsal and lateral surfaces
usually darker than ventral surface; dorsal and lateral body surface often with fine, dark brown reticulations
or mottled lines, sometimes with irregular white or yellow blotches; tail rings often encircled with dark brown
bands; pectoral fins with broad white outer margin and small brown spots forming irregular, longitudinal
bands; unpaired fins with small brown spots in irregular rows.
dorsal view lateral view
Habitat, biology, and fisheries: Benthic, found on sand, gravel, shell-rubble, or muddy bottoms. Collected
incidentally by seine, trawl, dredge, or shrimp nets; postlarvae have been taken at surface lights at night.
They apparently “walk” over the bottom with the aid of the tentacular pelvic fins. In Hong Kong and other
southeast Asian localities, specimens are dried and sold for medicinal purposes; the dried bodies are
subsequently boiled in water and the resulting broth is sipped as a cure for various throat irritations; dried
specimens have also been collected and maintained as curiosities in Chinese insect boxes. Otherwise
these fishes are of no commercial value.
Dactylopteridae: head large, snout blunt, body covered with
scute-like scales, not encased in fused bony plates; tail not
encased in bony rings. Spinous dorsal fin well developed;
pectoral fins extremely large, with 28 to 37 rays; pelvic fins with
I spine and 4 soft rays.
Eurypegasus draconis (Linnaeus, 1766)
Pegasus laternarius Cuvier, 1816
Pegasus volitans Linnaeus, 1758 Dactylopteridae
References
Orr, J.W. and T.W. Pietsch. 1994. Pipefishes and their allies. In Encyclopedia of fishes, edited by J. R. Paxton and
W.N. Eschmeyer. Sydney, University of New South Wales Press, pp. 168-172.
Palsson, W.A. and T.W. Pietsch. 1989. Revision of the acanthopterygian fish family Pegasidae (order
Gasterosteiformes). Indo-Pac. Fishes, (18):38 p.
Pietsch, T.W. 1978. Evolutionary relationships of the sea moths (Teleostei: Pegasidae) with a classification of
gasterosteiform families. Copeia, 1978:517-529.
Gasterosteiformes: Solenostomidae 2263
Gasterosteiformes: Solenostomidae SOLENOSTOMIDAE

Ghost pipefishes
by R.A. Fritzsche and K.G. Thiesfeld
D iagnostic characters: Small fishes (12 to 16 cm total length); body elongate, laterally compressed,
depth greatest between origin of spinous dorsal and pelvic fins, strongly constricted between
pelvic-fin insertion and origin of soft dorsal and anal fins. Head elongate, approximately 44% of
standard length; snout elongate, tubular, with nasal lamellae (number of lamellae sexually dimorphic);
mouth small, vertically oriented, toothless, with single mandibular barbel. Two dorsal fins, the first with
V weak spines and the second with 17 to 22 unbranched soft rays; spinous and soft dorsal fins widely
separated, each on a raised base; anal fin rounded, situated opposite soft dorsal fin and on a raised
base, composed of 17 to 22 unbranched soft rays; caudal fin truncate, rounded, or lanceolate; pectoral
fins small, rounded; pelvic fins large, situated opposite spinous dorsal fin. Lateral line absent. Body
covered with stellate plates that bear spines, variously with cutaneous papillae. Colour: overall
colour highly variable, ranging between light and dark phases: light colour phase with light background
often overlain with spots and/or reticulations; dark colour phase with overall dusky to black coloration or
strongly contrasting reticulating pattern of deep red-brown with yellow to orange spots, lines, or blotches.
Similar familes occurring in the area

Aulostomidae: larger; distinct separate spines anterior to soft
dorsal fin; lateral line present; never cutaneous papillae.
Syngnathidae: body encased in bony rings; no spines in fins;
pelvic fins absent; anal fin reduced or absent; male with
specialized ventral brooding surface or pouch.
Fistulariidae: body depressed rather than compressed; spi-
nous dorsal fin absent; a distinct caudal-fin filament present. Fistulariidae
Aulostomidae Syngnathidae
Habitat, biology, and fisheries: Ghost pipefishes are found over sand or mud bottoms, in seagrass beds,
marine algae, or in association with reef invertebrates. They feed on small benthic and pelagic inverte-
brates, mainly crustaceans. Reproduction involves the brooding of eggs in a ventral brood pouch formed
by the pelvic fins of the female. Ghost pipefish are not utilized as food. Sought by aquarium hobbyists.
Taken in bottom trawls and by hand. Occur throughout the Indo-West Pacific.

Solenostomus armatus Weber, 1913
Solenostomus cyanopterus Bleeker, 1854
Solenostomus paradoxus (Pallas, 1770)
Reference
Orr, J.W. and R.A. Fritzsche. 1993. Revision of the ghost pipefishes, family Solenostomidae (Teleostei: Syngnathoidei).
Copeia, 1993(1):168-182.
2264 Bony Fishes
SYNGNATHIDAE
Pipefishes and seahorses
by T. Paulus pectoral fin
D iagnostic characters: Usually small fishes (4 to 55 cm total length).

Body typically slender and elongate, without scales, encased in a
series of bony rings; with or without a prehensile tail. Head either more or
less along same axis as rest of body (subfamily Syngnathinae), or clearly
bent in ventral direction from main axis of body (subfamily Hippocampinae).
Snout generally long and tubular; mouth small, toothless, located at tip
of snout. Branchiostegal rays 1 to 3; gill opening reduced to a pore in the
opercular membrane. Spinous dorsal and pelvic fins absent; other fins
variously present or absent; a single dorsal fin, usually with 15 to 60 soft
rays; anal fin small, with 2 to 6 soft rays; caudal fin, if present, with 8 to 10
rays; pectoral fins usually with 10 to 23 rays. Some species develop dermal
appendages along body, head, and snout. No lateral line. Colour: variable anal fin
with the species, generally adapted to the preferred habitat; species living
on seagrass, sand, and coral rubble usually have grey, green, brown, or dorsal
fin
black ground colour with various patterns; coral-reef species sometimes
colourful with white, yellow, orange, blue, red, and black stripes and bands;
fins usually transparent; caudal fin sometimes with colourful patterns.
subfamily Hippocampinae (seahorses)
anal fin subfamily Syngnathinae (pipefishes)
G DFL
MDSR STR
LTR
ITR
LSR MVTR
OPR PR TR S cross-section
lateral view of head (after Dawson, 1985)
STR
STAR
LTR
ITR ITAR
MVTR
AR S LTAR TAR cross-section
posterior part of body (after Dawson, 1985)
morphological features used in the identification key

(adapted from Dawson, 1985)
AR - anal ring; DFL - dermal flap; G - gill opening; ITAR - inferior tail ridge; ITR - inferior trunk ridge;
LSR - lateral snout ridge; LTAR - lateral tail ridge; LTR - lateral trunk ridge; MDSR - median dorsal snout
ridge; MVTR - median ventral trunk ridge; OPR - opercular ridge; PR - pectoral ring (1st trunk ring);
S - scutellum; STAR - superior tail ridge; STR - superior trunk ridge; TAR - tail ring; TR - trunk ring.
Gasterosteiformes: Syngnathidae 2265
Habitat, biology, and fisheries: Syngnathidae usually live in coastal marine waters; some are found in
estuarine waters, and only a few in fresh water. The marine species live in a wide variety of habitats, such
as on sand and rubble substrate, seagrass beds, in caves and crevices of coral reefs and on steep drop-offs
of the reef. Juveniles often with a planktonic stage. Some species are associated with floating objects such
as Sargassum seaweeds. Generally found in shallow waters at depths of 1 to more than 100 m, planktonic
juveniles are sometimes caught in trawls from greater depths in the open sea. The majority of the known
species feed on minute benthic and planktonic fauna, preferably microcrustaceans. Perhaps the most
peculiar behaviour displayed by the seahorses and pipefishes is their habit of male egg incubation. The
female deposits the eggs on the ventral surface of the male body, where they are fertilized. The eggs are
kept in a pouch or on a specially vascularized surface of the trunk or tail. The brood is carried by the male
until the young hatch. Most of the relatively small species are of no or minor commercial importance.
However, some colourful species regularly appear in the aquarium trade, e.g. Corythoichthys spp.,
Doryrhamphus spp., Halicampus spp., and Hippocampus spp. In addition, seahorses and pipefishes are
sold primarily for use on Asian markets as medicine and aphrodisiacs, but also as curios and food.

The body encased in bony rings, typical body shape, and the lack of pelvic and second dorsal fins easily
distinguishes the seahorses and pipefishes from other families. Members of the Aulostomidae, Fistularii-
dae, and Solenostomidae may superficially resemble syngnathids.
Aulostomidae: larger, reaching a maximum length of 80 cm; body compressed, elongate, and scaly; series
of 8 to 12 isolated dorsal-fin spines, followed by a normal dorsal fin; well-developed lateral line; caudal fin
well developed and rounded.
Fistulariidae: generally larger, with maximum length of 1.8 m, but usually less than 1 m; body depressed,
elongate, and naked, or with minute prickles and linear row of scutes; caudal fin forked, with elongate
filament produced by the median 2 caudal-fin rays.
Solenostomidae: body short, compressed, with large stellate bony plates; 2 separate dorsal fins, the first
with 5 long feeble spines; pelvic fins relatively large; females with brood pouch formed by the pelvic fins;
maximum length 16 cm.
Aulostomidae
Fistulariidae Solenostomidae
Identification note
The identification of seahorses and pipefishes is based on specific morphological features such as the
typical bony ridges on the head, trunk, and tail (see figure on previous page) as well as the number of trunk
and tail rings. An additional important morphological character is the position and type of the male brood
area. A revision of the subfamily Hippocampinae is urgently needed.
Key to the genera and subgenera of Syngnathidae occurring in the area

(Key adapted from Dawson, 1985; figures after Dawson, 1985)
1a. Distal part of tail clearly prehensile and tapered . . . . . . . . . . . . . . . . . . . . . . . . . . → 2
1b. Distal part of tail not prehensile or tapered, its tip with a distinct caudal fin . . . . . . . . . . . . → 8
2a. Head clearly bent in ventral direction from longitudinal axis of body, usually more than
70° . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hippocampus
2b. Head essentially in line with longitudinal axis of body or bent very little in ventral direction . . . . . → 3
2266 Bony Fishes
3a. Principal body ridges with enlarged spines; bony

snout long, head 1.6 to 1.7 times as long as platelets
snout . . . . . . . . . . . . . . . . . . . . . . Haliichthys
3b. Principal body ridges without enlarged spines . . . . . → 4
4a. With bony platelets on sclera of eye (Fig. 1) . . . . . . → 5

4b. Without bony platelets on sclera of eye . . . . . . . . → 6 Fig. 1 lateral view of head
5a. Superior trunk and tail ridge continuous (Fig. 2a) . . . . . . . . . . . . . . Solegnathus (Runcinatus)
5b. Superior trunk and tail ridge discontinuous (Fig. 2b) . . . . . . . . . . . Solegnathus (Solegnathus)
6a. Lateral trunk ridge bent dorsally, ending just below superior ridge near rear of dorsal-fin
base (Fig. 2c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Syngnathoides
6b. Lateral trunk ridge not bent dorsally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 7
7a. Superior trunk and tail ridge continuous (Fig. 2d) . . . . . . . . . . . . . Acentronura (Acentronura)
7b. Superior trunk and tail ridge discontinuous (Fig. 2e) . . . . . . . . . . . Acentronura (Idiotropiscis)
8a. Superior trunk and tail ridges continuous (Fig. 2d) . . . . . . . . . . . . . . . . . . . . . . . . . → 9

8b. Superior trunk and tail ridges discontinuous (Fig. 2e) . . . . . . . . . . . . . . . . . . . . . . . → 19
9a. Inferior trunk and tail ridges continuous, lateral trunk ridge not confluent with inferior tail
ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 10
9b. Inferior trunk and tail ridges discontinuous, lateral trunk ridge confluent with inferior tail
ridge. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 16
10a. Anal fin present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 11

10b. Anal fin absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 14
11a. Lateral trunk ridge deflected ventrally near rear anal ring, dorsal-fin origin on tail (Fig. 2f)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ichthyocampus
11b. Lateral trunk ridge not deflected ventrally, dorsal-fin origin on trunk . . . . . . . . . . . . . . . . → 12
superior tail discontinuous

superior trunk ridge
ridge
a) b) c)
discontinuous dorsal-fin
origin
d) e)
trunk tail
f)
Fig. 2 lateral view of midbody
12a. Principal tail ridges produced laterally, the

posterior angles of tail rings usually hook-
like (Fig.3) . . . . . . . . . . . . . . . . Phoxocampus
12b. Principal tail ridges essentially straight,
the posterior angles of tail rings not pro-
duced and hook-like . . . . . . . . . . . . . . . . → 13
Fig. 3 end of tail
13a. Median dorsal snout ridge low, essentially entire and concave in lateral profile; snout 2.5
to 6.5 times as long as deep; total number of rings 46 to 59, dorsal-fin rays 18 to 28,
pectoral-fin rays 10 to 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Festucalex
13b. Median dorsal snout ridge elevated or with semi-isolated dorsal projections; snout 1.3 to
2.3 times as long as deep; total number of rings 39 to 52, dorsal-fin rays 16 to 19,
pectoral-fin rays 7 to 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Campichthys
14a. Mouth inferior, not on projecting snout; dorsal fin absent in subadults and adults (Fig. 4a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bulbonaricus
14b. Mouth terminal on projecting snout; dorsal fin present (Fig. 4b) . . . . . . . . . . . . . . . . . → 15
mouth
terminal
mouth
inferior
a) b)
15a. Pectoral fins present . . . . . . . . . . . . . . . . . . . . . . . . . . Nannocampus (Mannarichthys)
15b. Pectoral fins absent . . . . . . . . . . . . . . . . . . . . . . . . . . Nannocampus (Nannocampus)
16a. Pectoral fins present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . →17

16b. Pectoral fins and dorsal fin absent in subadults and adults . . . . . . . . . . . . . . . . . . . . →18
17a. Dorsal-fin origin on trunk; caudal-fin rays typically 10; anal fin present; total number of
rings 31 to 44, pectoral-fin rays 18 to 23; anal fin present . . . . . . . . . . . . . . . . Choeroichthys
17b. Dorsal-fin origin on tail, usually between first and sixth tail ring; total number of rings 57
to 73, pectoral-fin rays 9 to 13; anal fin absent . . . . . . . . . . . . . . . . . . . . . . Siokunichthys
18a. Trunk rings 10 to 12; brood pouch without bilateral folds, a closed sac-like structure
opening through an anteromesial pore (Fig. 5a) . . . . . . . . . . . . . . . . . . . Apterygocampus
18b. Trunk rings 17 to 19; brood pouch with bilateral membranous folds and semi-pouch
closure (Fig. 5b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Penetopteryx
19a. Inferior trunk and tail ridges continuous, lateral trunk ridge not confluent with lateral and
superior tail ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 20
19b. Inferior trunk and tail ridges discontinuous, the lateral trunk ridge confluent with the
inferior tail ridge (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 25
eggs
pouch
a) b) fold
Fig. 5 cross-section of body and brood pouch Fig. 6 lateral view of midbody
2268 Bony Fishes
20a. S nout with dorsolateral spines and

spinules, adults with bony platelets in gill
membranes (Fig. 7) . . . . . . . . . . . . . . . Bhanotia
20b. Snout without dorsolateral spines and
bony
spinules, without bony platelets in gill mem- plates
branes . . . . . . . . . . . . . . . . . . . . . . . . → 21
21a. Caudal-fin rays typically 9; trunk rings 15 to 26; male brood area under trunk . . . . . . Doryichthys
21b. Caudal-fin rays typically 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 22
22a. With lateral snout ridge and/or dermal flaps . . . . . . . . . . . . . . . . . . . . . . . Cosmocampus

22b. Without lateral snout ridge or dermal flaps . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 23
23a. Anal-fin rays 4; pouch plates absent, semi-pouch closure (Fig. 8a) . . . . . . . . . . Corythoichthys
23b. Anal-fin rays 2 or 3; pouch plates present, everted pouch closure (Fig. 8b) . . . . . . . . . . . .→ 24
24a. Lateral trunk ridge deflected ventrally at anal ring (Fig. 9a) . . . . . . . . . Hippichthys (Hippichthys)
24b. Lateral trunk ridge not deflected at anal ring (Fig. 9b) . . . . . . . . . Hippichthys (Parasyngnathus)
everted
pouch
a) closure b)
a) b)
Fig. 8 cross-section of body and brood pouch Fig. 9 lateral view of midbody
25a. Caudal-fin rays typically 8 or 9 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 26

26a. Caudal-fin rays typically 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 27

27a. Median dorsal snout ridge with 1 to 3 semi-isolated dorsal projections; pectoral-fin rays
8 to 13; anal-fin rays 2 or 3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minyichthys
27b. Median dorsal snout ridge entire; pectoral-fin rays 14 to 16; anal-fin rays 4 . . . . . . . . Filicampus
28a. Caudal fin often stubby (Fig. 10); trunk

rings 21 to 24, the first clearly longer
than second; male brood area under
tail . . . . . . . . . . . . . . . . Trachyrhamphus
28b. Caudal fin well developed; trunk rings
15 to 21, the first not much longer than Fig. 10 end of tail
second; male brood area under trunk . . . . →29
29a. Opercular ridge distinct, typically complete; pectoral-fin base with 1 or 2 distinct ridges;
lateral and inferior trunk ridges distinct . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 30
29b. Opercular ridge vestigial or obsolete; pectoral-fin base without distinct ridges in
subadults and adults; lateral and inferior trunk ridges indistinct . . . . . . . . Microphis (Coelonotus)
30a. Without supplemental ridges below longitudinal opercular ridge (Fig. 11a) . . . . . . . . . . . . ® 31
30b. With 1 or more supplemental ridges below longitudinal opercular ridge (Fig. 11b) . . . . . . . . ® 32
31a. Dorsal-fin rays 57 to 74; subdorsal trunk rings 5.5 to 12; usually with some keeled
scutella in subadults-adults (Fig. 12a) . . . . . . . . . . . . . . . . . . . . Microphis (Belonichthys)
31b. Dorsal-fin rays 28 to 56; subdorsal trunk rings 0.25 to 3.5; scutella without longitudinal
keels (Fig.12b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microphis (Microphis)
ridges
a) b)
a) keeled b) unkeeled
Fig. 11 lateral view of head Fig. 12 detail of body (lateral view)
32a. Snout relatively long and slender, on average 4.3 to 10.6 times as long as deep; scutella
without keels . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Microphis (Oostethus)
32b. Snout relatively short and deep, its depth averages 3.5 to 4.1 in snout length; usually
with some keeled scutella . . . . . . . . . . . . . . . . . . . . . . . . . . Microphis (Lophocampus)
33a. First trunk ring much longer than second (Fig. 13a); pectoral fins typically emarginate;
caudal fin large; dermal flaps absent; male brood area under trunk . . . . . . . . . . . . . . . ® 34
33b. First trunk ring not much longer than second (Fig. 13b); pectoral fins rounded; caudal fin
not large; dermal flaps usually present; male brood area under tail . . . . . . . . . . . . . . . . ® 35
34a. Snout with 1 to 5 rows of dorsolateral spinules; without banded colour pattern; membra-
nous pouch folds present in brooding males (Fig. 14a) . . . . . . . Doryrhamphus (Doryrhamphus)
34b. Snout without rows of dorsolateral spinules; mostly with banded colour pattern; membra-
nous pouch folds absent in brooding males (Fig.14b) . . . . . . Doryrhamphus (Dunckerocampus)
1st trunk ring not longer than 2nd
1st trunk ring longer than 2nd
a) b)
Fig. 13 head and anterior part of trunk (lateral view) no pouch folds
pouch folds a) b)
35a. Median dorsal snout ridge low, entire, essentially Fig. 14 cross-section of body and
concave in lateral profile; lateral snout ridge ab- brood pouch
sent; without dermal flaps on eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . Micrognathus
35b. Median dorsal snout ridge not essentially concave in lateral profile, often elevated or
spiny; lateral snout ridge or spine usually present; usually with dermal
flaps on eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Halicampus

Note: species that occur in brackish or fresh water are marked in the following list by an asterisk (*). A plus
mark (+) indicates commercial species (i.e. used in the aquarium trade, as curios, medicine, or food).
The symbol 0is given when species accounts are included.
Subfamily HIPPOCAMPINAE
0
Hippocampus angustus Günther, 1870
0
Hippocampus histrix Kaup, 1856 +
Hippocampus kuda Bleeker, 1852 +
0
Hippocampus planifrons Peters, 1877
Hippocampus whitei Bleeker, 1855 +
2270 Bony Fishes
Subfamily SYNGNATHINAE
Acentronura (Acentronura) gracilissima (Temminck and Schlegel, 1850)
Acentronura (Acentronura) tentaculata Günther, 1870
Acentronura (Idiotropiscis) larsonae Dawson, 1984
Apterygocampus epinnulatus Weber, 1913
Bhanotia fasciolata (Duméril, 1870)
Bhanotia nuda Dawson, 1978
Bulbonaricus brauni (Dawson and Allen, 1978)
Bulbonaricus davaoensis (Herald, 1953)
Campichthys tricarinatus Dawson, 1977
Campichthys tryoni (Ogilby, 1890)
Choeroichthys brachysoma (Bleeker, 1855)
Choeroichthys cinctus Dawson, 1976
Choeroichthys sculpatus (Günther, 1870)
Choeroichthys suillus Whitley, 1951
0
0
Corythoichthys amplexus Dawson and Randall, 1975 +
0
Corythoichthys flavofasciatus (Rüppell, 1838) +
Corythoichthys haematopterus (Bleeker, 1851) +
0
Corythoichthys intestinalis (Ramsay, 1881) +
Corythoichthys nigripectus Herald, 1953 +
Corythoichthys ocellatus Herald, 1953
Corythoichthys paxtoni Dawson, 1977
0
Corythoichthys polynotatus Dawson, 1977
Corythoichthys schultzi Herald, 1953 +
Cosmocampus banneri (Herald and Randall, 1972)
Cosmocampus darrosanus (Dawson and Randall, 1975)
Cosmocampus howensis (Whitley, 1948)
Cosmocampus investigatoris (Hora, 1925)
Cosmocampus maxweberi (Whitley, 1933)
Doryichthys boaja (Bleeker, 1851) *
Doryichthys deokhatoides (Bleeker, 1853) *
Doryichthys heterosoma (Bleeker, 1851) *
Doryichthys martensii (Peters, 1869) *
0 Doryrhamphus (Doryrhamphus) excisus excisus Kaup, 1856 +
Doryrhamphus (Doryrhamphus) janssi (Herald and Randall, 1972) +
Doryrhamphus (Doryrhamphus) negrosensis malus (Whitley, 1954)
Doryrhamphus (Doryrhamphus) negrosensis negrosensis Herre, 1934
0
Doryrhamphus (Dunckerocampus) chapmani (Herald, 1953)
0
Doryrhamphus (Dunckerocampus) dactyliophorus (Bleeker, 1853) +
Doryrhamphus (Dunckerocampus) pessuliferus (Fowler, 1938) +
Festucalex cinctus (Ramsay, 1882)
Festucalex erythraeus (Gilbert, 1905)
Festucalex gibbsi Dawson, 1977
Festucalex prolixus Dawson, 1984
Festucalex wassi Dawson, 1977
Filicampus tigris (Castelnau, 1879)
Halicampus boothae (Whitley, 1964)
0
Halicampus brocki (Herald, 1953) +
Halicampus dunckeri (Chaubanaud, 1929) +
Halicampus grayi Kaup, 1856 +
Halicampus macrorhynchus Bamber, 1915 +
Halicampus marquesensis Dawson, 1984
Halicampus mataafae (Jordan and Seale, 1906)
Halicampus nitidus (Günther, 1873) +
Haliichthys taeniophorus Gray, 1859
Hippichthys (Hippychthys) cyanospilus (Bleeker, 1854)

Hippichthys (Hippychthys) heptogonus Bleeker, 1849
Hippichthys (Hippychthys) spezifer (Rüppell, 1838)
Hippichthys (Parasyngnathus) parvicarinatus (Dawson, 1978) *
Hippichthys (Parasyngnathus) penicillus (Cantor, 1849) *
Ichthyocampus carce (Hamilton Buchanan, 1822) *
Micrognathus andersonii (Bleeker, 1858)
Micrognathus brevirostris pygmaeus Fritzsche, 1981
Micrognathus micronopterus (Fowler, 1936)
Micrognathus natans Dawson, 1982
Microphis (Belonichthys) mento (Bleeker, 1856)
Microphis (Belonichthys) spinachioides (Duncker, 1915)
Microphis (Coelonotus) angulus (Peters, 1855) *
Microphis (Coelonotus) leiaspis (Bleeker, 1853) *
Microphis (Lophocampus) brevidorsalis (de Beaufort, 1913) *
Microphis (Lophocampus) caudocarinatus (Weber, 1908) *
Microphis (Lophocampus) ocellatus (Duncker, 1910) *
Microphis (Lophocampus) retzii (Bleeker, 1856) *
Microphis (Microphis) cruentus Dawson and Fourmanoir, 1981 *
Microphis (Oestethus) brachyurus brachyurus Bleeker, 1853
Microphis (Oestethus) jagorii Peters, 1869 *
Microphis (Oestethus) manadensis (Bleeker, 1856) *
Microphis (Oestethus) pleurostictus (Peters, 1869 *
Minyichthys brachyrhinus (Herald, 1953)
Minyichthys myersi (Herald and Randall, 1972)
Nannocampus (Mannarichthys) pictus (Duncker, 1915)
Nannocampus (Mannarichthys) weberi Duncker, 1915
Penetopteryx taeniocephalus Lunel, 1881
Phoxocampus blecheri (Kaup, 1856)
Phoxocampus diacanthus (Schultz, 1943)
Phoxocampus tetrophthalmus (Bleeker, 1858)
Siokunichthys breviceps Smith, 1963 +
Siokunichthys herri Herald, 1953
Siokunichthys nigrolineatus Dawson, 1983 +
Siokunichthys southwelli (Duncker, 1910)
Solegnathus (Runcinatus) dunckeri Whitley, 1927
Solegnathus (Solegnathus) hardwickii (Gray, 1830)
Solegnathus (Solegnathus) lettiensis Bleeker, 1860
Solegnathus (Solegnathus) spinosissimus Günther, 1870
0 Syngnathoides biaculeatus (Bloch, 1785) +
0 Trachyrhamphus bicoarctatus (Bleeker, 1857) +
Trachyrhamphus longirostris Kaup, 1856 +
References
Dawson, C.E. 1985. Indo-Pacific pipefishes (Red Sea to the Americas). Ocean Springs, Mississippi, Gulf coast Res.
Lab., 230 p.
Dawson, C.E. and Vari, R.P. 1982. Fishes of the western north Atlantic, Part 8: order Gasterosteiformes. Mem. Sears.
Found. Mar. Res., 1(8):189 p.
Fritzsche, R.A. 1980. Revision of eastern Pacific Syngnathidae (Pisces: Syngnathiformes) including both recent and
fossil forms. Proc. Calif. Acad. Sci., 42:181-227.
Herald, E.S. 1959. From pipefish to seahorse. - a study of phylogenetic relationship. Proc. Calif. Acad. Sci., 4. Ser.,
29(13):465-473.
Vincent, A.C.J. 1996. The international trade in seahorses (species in danger). TRAFFIC International report.
2272 Bony Fishes
Hippocampus histrix Kaup, 1856

En - Thorny seahorse.
Maximum total length 15 cm (stretched). Colour varying. In coastal
areas among clumps of algae or in seagrass beds. Feeds primarly
on crustacean zooplankton. Sometimes sold as medicine and aph-
rodisiac, but also as aquarium fish, curios, and food. Widespread in
the western Indian Ocean, including the Red Sea, and the western
and eastern Central Pacific.
Hippocampus kuda Bleeker, 1852

En - Yellow seahorse.
Maximum total length 30 cm (stretched). Colour ranges from dirty
yellow over reddish brown to black, and may be blotched or banded.
In coastal areas on seagrass, also in sandy and coral areas, to a depth
of about 30 m. Often sold as curios and aphrodisiacs on Asian
markets. Feeds on zooplankton. Widespread in the Indian Ocean and
the western and eastern Central Pacific.
Hippocampus whitei Bleeker, 1855

En - White’s seahorse.
Maximum total length 15 cm (stretched). Colour varying from grey to
brown and black, sometimes with saddle-like markings. Along coastal
areas among seaweeds. Feeds on microcrustaceans. Regularly im-
ported by the aquarium trade and sold as curios and medicine.
Common along the coasts of Australia, Madagascar, Mozambique,
and South Africa.
Corythoichthys amplexus Dawson and Randall, 1975

En - Brownbanded pipefish.
Maximum total length about 10 cm. Body with broad dark bands crossing side and dorsum. Occurs
on shallow reefs and deep walls in depths to about 30 m. Feeds on microcrustaceans by sucking
in through the tube snout. Sometimes imported by the aquarium trade. Widespread in the eastern
Indian Ocean and Pacific.
Corythoichthys flavofasciatus (Rüppell, 1838)

En - Network pipefish.
Maximum total length about 14 cm. Body with about 13 to 25 reticulate bands crossing side and
dorsum; males during courtship with intensive dark blue or black blotch on the ventral of the anal
ring. Occurs in small schools along the coastline on sand and seagrass beds near coral reefs. Feeds
on live zooplankton. Couples seem to be monogamous. Rarely in the aquarium trade. Widespread
in the Indian Ocean, including the Red Sea and the Persian Gulf, the northwestern and western
Central Pacific, except the Philippines and Indonesia.
Corythoichthys haematopterus (Bleeker, 1851)

En - Reeftop pipefish.
Maximum total length about 20 cm. Feeds on zooplankton. Occurs in groups or schools along the
coastline in depths to 20 m, but mostly from 0 to 3 m. Sold regularly in the aquarium trade in Europe
and America. Widespread in the Indian Ocean from the African coast to the western Central and
northeastern Pacific.
2274 Bony Fishes
Corythoichthys nigripectus Herald, 1953

En - Blackbreasted pipefish.
Maximum total length about 11 cm. Coloration variable without distinct bars or stripes, opercle
mostly orange-red, breast dark brown to black. Not a common species, occurs alone or in pairs in
lagoons and seaweed reefs in caves and crevices in depths from 4 to about 30 m. Feeds on
zooplankton. Very rarely in the aquarium trade. Found in the northern Red Sea, throughout
Micronesia, and the Society and Caroline islands.
Corythoichthys schultzi Herald, 1953

En - Schultz’ pipefish.
Maximum total length about 17 cm. Coloration varying in geographic regions; usually with dark
stripes and spots on the side of the snout, trunk with 10 to 15 bars. A very common species in large
groups on reef crests, sandy areas with coral patches and rubble flats near seagrass beds down to
a depth of about 30 m. Feeds on small crustaceans, sucked in from the sandy substratum or from
between the branches of soft corals. Very rarely in the aquarium trade. Widespread throughout the
Indo-Pacific including the Red Sea and the northwestern and western Central Pacific.
Doryrhamphus (Doryrhamphus) dactyliophorus (Bleeker, 1853)

En - Banded pipefish.
Maximum total length about 18 cm. Specimens regularly banded with white and red or black bands
along trunk and tail. Occurs in pairs or large groups in protected coastal reefs and lagoons, mostly
in caves. Feeds mostly on planktonic amphipods. Very common in the aquarium trade, exported
from Indonesia and the Philippines in large numbers. Widespread in the Indo-Pacific including the
Red Sea, western and eastern Central Pacific, and northwestern Pacific, from South Africa to Japan
and Australia.
Doryrhamphus (Doryrhamphus) excisus excisus Kaup, 1856

En - Bluestripe pipefish.
Maximum total length about 7 cm. A colourful species with a blue longitudinal stripe along the trunk
and tail; yellowish dorsally and ventrally; fan-like tail fin. Several subspecies are reported from
different geographic regions. Lives secretive in crevices and caves along the coastline in depths of
usually more than 35 m. Sometimes hides between the spines of sea urchins. Feeds on floating
planktonic organisms. Hovers in pairs over the substratum. Regularly seen in large numbers in the
aquarium trade, and used as curios and medicine in Asia. Widespread in the Indian Ocean, including
the Red Sea, and throughout the Pacific.
Doryrhamphus (Doryrhamphus) pessuliferus (Fowler, 1938)

En - Yellowbar pipefish.
Maximum total length about 14 cm. Similar to Doryrhamphus (Doryrhamphus) multiannulatus, but
differs in coloration and tail and trunk rings. Snout bright yellow, trunk and tail regularly with red and
yellow bands; red tail fin with yellow dot in centre. Occurs secretively in caves and crevices along
coral reefs in depths of 30 m and more. No data on food preferences available. Frequently imported
to Europe by the aquarium trade. A single reliable record from the Sulu Archipelago in the western
Central Pacific.
Halicampus dunckeri (Chaubanaud, 1929)

En - Duncker’s pipefish.
Maximum total length about 12 cm. Colour variable from pale to brown and black with black, brown,
and white bands. Different morphological types have been reported from various geographical
regions. Occurs on sand and rubble along the coastline of coral reefs in a depth of more than 20 m.
Feeds on small planktonic organisms. Sometimes used as curios and sold in the aquarium trade.
Widespread in the western and eastern Indian Ocean, including the Red Sea, northwestern and
western Central Pacific.
2276 Bony Fishes
Syngnathoides biaculeatus (Bloch, 1785)

En - Alligator pipefish.
Maximum total length about 30 cm. Colour varying from green and brown with irregular white
markings. Tail fin absent in adults, tail prehensile like a seahorse; dermal flaps over much of trunk
and tail. Occurs in shallow waters in seagrass beds or close to floating weeds. Juveniles are found
in offshore samples. Feeds on small planktonic organisms. Regularly enters the aquarium trade
and primarily used as curios and medicine. Widespread in the Indian Ocean including the Red Sea,
South Africa, and Mozambique, as well as the northwestern, eastern, and western Central Pacific.
Trachyrhamphus bicoarctatus (Bleeker, 1857)

En - Double-ended pipefish.
Maximum total length about 42 cm. Colour varying from near white, green, grey, or brown to black;
body plain, spotted, or mottled, adapted to the substratum. Tip of snout often pale. Planktonic
juveniles have elongate appendages on the dorsum of trunk and tail. Adults loose their tail fin, which
cannot be regenerated. Occurs on sand and rubble along coastlines in depths from 1 to more than
45 m, where it imitates drifting seagrass. Feeds on crustaceans, such as Gammarida, Mysidacea,
Amphipoda, and Caprellidae. Sometimes sold for use as medicine, curios, and in the aquarium
trade. Widespread in the western and eastern Indian Ocean, including the Red Sea, the Persian
Gulf, and South Africa; also in the northwestern, eastern, and western Central Pacific to China,
Japan, and southern Australia.
Gasterosteiformes: Aulostomidae 2277
Gasterosteiformes: Aulostomidae AULOSTOMIDAE

Trumpetfishes

Aulostomus chinensis (Linnaeus, 1766)
Frequent synonyms / misidentifications: Aulostomus valentini (Bleeker, 1853) / None.
FAO names: En - Chinese trumpetfish; Fr - Trompette chinoise; Sp - Trompetero chino.
Diagnostic characters: Body elongate and compressed. Mouth located at tip of elongate snout;
single barbel on chin. First dorsal fin with VIII to XII isolated spines; second dorsal fin and anal
fin opposite to each other and similarly shaped; second dorsal fin with 24 to 27, and anal fin with 26
to 29 segmented soft rays; pelvic fins small, abdominal, with 6 soft rays. Lateral line continuous. Body
covered with small ctenoid (rough) scales, except for the head and anterior part of the back, which are
scaleless. Vertebrae 62 to 64, the first 4 elongate and fused. Colour: overall colour variable; body most
commonly brownish with irregular light vertical bar, or with several white horizontal stripes, or uniform
yellow colour; a black maxillary stripe usually present, but may be reduced; dorsal and anal fins light, but
with a dark basal bar; caudal fin usually with 2 round black spots; a black spot at base of each pelvic fin.
Fistulariidae: a distinct caudal-fin filament present; body depressed rather than compressed; no spines
before soft dorsal fin.
Solenostomidae: smaller; dorsal-fin spines not isolated; no lateral line; body variously covered with
cutaneous papillae.
Fistulariidae Solenostomidae
Size: Maximum total length 80 cm; commonly to 40 cm.

Habitat, biology, and fisheries: Occurs in shallow, clear water, most frequently observed hanging
vertically in the water with the head down and associated with gorgonians. Also will conceal itself by hiding
among schools of other fish. Capable of colour changes depending on surroundings. Feeds on small fishes
and crustaceans. Taken occasionally as bycatch in artisanal fisheries. Of no commercial importance.
Occasionally consumed by the local population.
Distribution: T r o p i c a l In d o -
Pacific, just reaching offshore
is la nd s o f e astern Pacific,
including southern Japan,
Hawaii, and Lord Howe and
Easter islands.
Reference
Wheeler, A.C. 1955. A preliminary
revision of the fishes of the
genus Aulostomus. Ann.
M a g . Nat, H ist., Ser.,
12(8):613-623.
2278 Bony Fishes
Gasterosteiformes: Fistulariidae FISTULARIIDAE

Cornetfishes (flute mouths)
D iagnostic characters: Large fishes (to about 2 m total length); body elongate and depressed. Mouth
small, at end of a long tubular snout, hexagonal in cross-section; teeth in jaws small. Dorsal and anal
fins short based and opposite, with 14 to 17 segmented soft rays; pectoral fins with 13 to 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 continuing posteriorly onto an elongate filament produced by the middle 2 caudal-fin
rays; lateral line composed of tube-shaped ossifications that gradually take the form of long bony shields
sometimes bearing sharp spines. Body of juveniles covered with rows of small spinules (retained in adults of
only 1 of the Indo-Pacific species, Fistularia petimba); a row of elongate bony plates may be present along
dorsal and ventral midlines of body just anterior to dorsal and/or anal fin. Total number of vertebrae 76 to 85,
with the first 4 elongate and fused. Colour: variable with the species; either red to orange-brown above and
silvery below, or brownish olive above, lighter below, with a series of blue spots on back and snout.
Habitat, biology, and fisheries: Fistularia petimba is typically found in coastal areas over soft bottoms,
usually at depths greater than 10 m. F. commersonii is most often seen in seagrass beds and coral reefs.
Cornetfishes feed on small fishes and crustaceans. Although not important in commercial fisheries of the
area, they are frequently taken in trawls and by various types of artisanal gear and may appear in local
fish markets. Although edible, they are most often used for fishmeal.

Aulostomidae: no caudal-fin filament present; barbel pre- Aulostomidae
sent on lower jaw; body compressed rather than de-
pressed; distinct separate spines anterior to soft dorsal fin.
Syngnathidae: smaller; body covered with armor; anal fin
reduced or absent; no caudal-fin filament present. Belonidae
Belonidae: no caudal-fin filament present; mouth large,
both jaws produced into slender beak, lower jaw longer,
with single series of distinct canines.
Syngnathidae
Key to the species of Fistulariidae occurring in the area elongate body plates dorsal fin
1a. A row of elongate bony plates embedded in skin along
midline of back anterior to dorsal fin (Fig. 1); posterior
lateral-line ossifications ending in a sharp spine; im-
maculate red or brown above . . . . . . . . . . . Fistularia petimba
1b. No elongate bony plates along midline of back; poste-
rior lateral-line ossifications without a spine; rows of dorsal view
blue spots on back, sides and snout . . . . Fistularia commersonii
Fig. 1 Fistularia petimba
Gasterosteiformes: Fistulariidae 2279

0
0
Fistularia commersonii Rüppell, 1835
Fistularia petimba Lacepède, 1803
Reference
Fritzsche, R.A. 1976. A review of the cornetfishes, genus Fistularia (Fistulariidae), with a discussion of intrageneric
relationships and zoogeography. Bull. Mar. Sci., 26(2):196-204.
Fistularia commersonii Rüppell, 1835

En - Bluespotted cornetfish; Fr - Cornette à taches bleues; Sp - Corneta pintada.
Maximum total length about 1.6 m; commonly to 1 m. Brownish to olive above, becoming lighter to
silvery below; a pair of blue stripes or a row of blue spots along back; dorsal and anal fins with an
orange cast becoming transparent at base; caudal-fin filament white. Juveniles may have dark
barred pattern with a dusky caudal-fin filament. Most common in seagrass beds and coral reefs in
shallow water. Feeds on small fishes and crustaceans. No special fishery, but taken frequently in
trawls and artisanal fisheries. Caught with bottom trawls, gill nets, and line gear. Utilized fresh, dried
salted, or smoked; also reduced to fishmeal. Indo-Pacific: Red Sea to Panama, north to southern
Japan and Hawaii, south to Lord Howe and Easter islands, throughout Micronesia.
Fistularia petimba Lacepède, 1803

En - Red cornetfish; Fr - Cornette rouge; Sp - Corneta colorada.
Maximum total length about 2 m; commonly to 1 m. Colour in life red to orange-brown above, silvery
below; vertical fins also have an orange cast. Found in coastal areas over soft bottoms, usually at
depths greater than 10 m. Feeds on small fishes and crustaceans. No special fishery, but caught
frequently in bottom trawls and in artisanal fisheries. Caught with bottom trawls, gill nets, and line
gear. Utilized fresh, dried salted, or smoked; also reduced to fishmeal. Atlantic, Indian, and western
Pacific oceans, including Hawaii.
2280 Bony Fishes
Gasterosteiformes: Macrorhamphosidae MACRORHAMPHOSIDAE

Snipefishes

Macrorhamphosus scolopax (Linnaeus, 1758)
Frequent synoyms / misidentifications: Macrorhamphosus gracilis (Lowe, 1839); M. velitaris (Pallas,
1776) / None.
FAO names: En - Longspine snipefish; Fr - Bécasse de mer; Sp - Trompetero.
Diagnostic characters:
Body compressed, deep,
widest point at or posterior
to midbody at origin of first
dorsal fin, narrowing at
caudal peduncle. Head
elongate; snout long, tu-
bular; mouth small, tooth-
less, located at tip of snout;
eyes lateral, large, con-
tained about 4.5 times in
snout length. First dorsal
fin originating at or behind
midbody; second spine greatly enlarged, posterior edge serrated, other spines stout but short; pelvic fins
small, originating at or behind midbody. Lateral line absent. Body covered with small but distinct scales, 2
series of bony plates embedded in the skin on the back between head and dorsal fin, each series
consisting of 3 well-developed plates and a fourth much smaller plate. There are 2 forms, slender and
deep bodied, often treated as distinct species. Colour: orange, pink, or red on back; paler and silvery on sides.
Centriscidae: body blade-like and enclosed in transpar-
ent, bony casing with sharp ventral edge; first dorsal-fin
spine greatly enlarged, first dorsal-fin spine at hindmost
point of body. Centriscidae
Triacanthidae: mouth much wider than snout; gill opening
small, less than or equal to width of pectoral-fin base; first
dorsal-fin origin anterior to midbody, dorsal-fin spines not
enlarged.
Solenostomidae: body strongly constricted between pel-
vic-fin insertion and origin of soft dorsal and anal fins,
covered with stellate plates bearing spines; mouth with
singular mandibular barbel; spinous and soft dorsal fin
widely separated and on a raised base; pelvic fins large. Triacanthidae
Habitat, biology, and fisheries: Snipefishes are benthic
or benthopelagic fishes commonly found in schools over
mud or sand bottoms in deep water to 600 m. Juveniles
are epipelagic in oceanic waters. They feed on zooplank-
ton and benthic invertebrates, mainly crustaceans. Not a
commercially important species. Not considered a food Solenostomidae
fish. Caught in abundance by bottom trawl.
Distribution: Ranges across tropical and temperate Atlan-
tic, Pacific, and Indian oceans, probably worldwide.
Reference
Heemstra, P.C. 1986. Family Macrorhamphosidae. In Smiths’
sea fishes, edited by M.M. Smith and P.C. Heemstra.
Gasterosteiformes: Centriscidae 2281
Gasterosteiformes: Centriscidae CENTRISCIDAE

Shrimpfishes (razorfishes) 1st dorsal
fin
by R.A. Fritzsche and K.G. Thiesfeld 2nd dorsal
fin
D iagnostic characters: Small fishes (14 to 30 cm total length); body elon-
gate, strongly compressed, and blade-like. Head elongate; snout long,
slender, and tubular; mouth small, toothless, located at tip of snout. Two
caudal
fin
short-based dorsal fins; first dorsal-fin spine greatly enlarged, originating at
hindmost end of body; all other spinous and soft portions of dorsal fins on
ventral surface of body; caudal fin small, on ventral surface, nearly at right anal
fin
angle to body axis; pelvic fins small, with 4 short soft rays, originating at or
behind midbody. Lateral line absent. Body enclosed in a flattened, transparent,
bony casing with sharp ventral edge. Colour: variable with the species; either
silvery or yellowish brown to pale green on back, silvery on sides; dusky to
conspicuous lateral streak running length of body and through eye.
Habitat, biology, and fisheries: Razorfishes are found on muddy bottoms near
mangroves to inshore coral reefs, frequently seeking refuge among coral branches
or the spines of long-spined sea urchins. They swim in small groups, each fish in
a vertical position, with its snout pointing downwards. They feed on small benthic
invertebrates, mainly crustaceans. Razorfishes are not utilized as food, have no
commercial importance, but are sought by aquarium hobbyists. Taken in bottom
trawls and by hand.
Macrorhamphosidae: body not blade-like, lacking bony casing, with small but
distinct scales; first dorsal fin originating near midbody; second dorsal-fin spine
greatly enlarged, serrated on posterior edge; caudal fin not at right angle to body
axis on ventral surface.
Solenostomidae: body not blade-like, covered with stellate plates that bear spines;
pelvic fins enlarged; caudal fin not at right angle to body axis on ventral surface. natural orientation in life
Macroramphosidae Solenostomidae
Key to the species of Centriscidae occurring in the area
1a. First dorsal-fin spine hinged at its base, slightly movable, with a movable spinous ray at
its end; interorbital space striated, convex, without a longitudinal groove . . . . . . Aeoliscus strigatus
1b. First dorsal-fin spine fused with body armour plate, without a movable spinous ray at its
end; interorbital space convex or with a groove continued to crown of head, which is
striated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ®2
2a. Interorbital space with a groove continued to crown of head; sutures of lateral plates
serrated; postorbital part of head 1/2 or more than 1/2 distance of operculum from base
of pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centriscus scutatus
2b. Interorbital space convex, without groove; sutures of lateral plates smooth; postorbital part
of head 3 times the distance of operculum from base of pectoral fins . . . . . . . Centriscus cristatus
0 Aeoliscus strigatus (Günther, 1860)
0
0
Centriscus cristatus (De Vis, 1885)
Centriscus scutatus Linnaeus, 1758
Reference
Mohr, E.W. 1937. Revision der Centriscidae (Acanthopterygii, Centrisciformes). Dana Rept., 13:1-69.
2282 Bony Fishes
Aeoliscus strigatus (Günther, 1860)

En - Shrimpfish.
Maximum total length about 14 cm. Colour yellowish brown to pale green on back, silvery on sides;
conspicuous black stripe running from the snout, through the eye, to the base of spinous dorsal fin.
Occurs in small schools that frequently seek refuge among the spines of the long-spined sea urchin
Diadema setosum, or coral branches. Feed primarily on crustacean zooplankton. Indo-West Pacific,
but not in the Red Sea.
(after Randall, Allen, and Steene, 1990)
Centriscus cristatus (De Vis, 1885)

En - Smooth razorfish.
Maximum total length 30 cm. Colour silvery, with a pronounced black band running through the eye
to the base of spinous dorsal fin. Western Pacific, northern Australia.
Centriscus scutatus Linnaeus, 1758

En - Serrate razorfish.
Maximum total length 14 cm. Colour silvery, with a dusky lateral streak running length of body; 7 or
8 silvery crossbars on ventral plates. Indo-West Pacific.
(after Sainsbury, Kailola, and Leyland, 1984)
click for next page

Scorpaeniformes: Dactylopteridae 2283
Order SCORPAENIFORMES
DACTYLOPTERIDAE
Flying gurnards (helmet gurnards)
by S.G. Poss and W.N. Eschmeyer
D iagnostic characters: Moderately elongate scorpaeniform fishes (size to 40 cm). Head large, heavily
armored. Eyes large. Interorbit wide, concave. Mouth small, subterminal, and protractile; teeth minute,
nodular or absent. A small movable bone (pontinal) between plate-like bones around eye and
preopercle. Preopercle with a prominent elongate spine. Gill openings restricted and fused to
isthmus. Spinous and soft parts of dorsal fin separated by a deep notch and I short spine (reduced to a
short spinous point); total number of dorsal-fin spines VII or VIII, with first or first 2 anterior spines
separated from remainder of fin; soft-rayed part of dorsal fin with 8 or 9 rays. Anal fin without spines and
with 6 or 7 soft rays. Caudal fin emarginate. Base of pectoral fins horizontal, the fins divided into 2
sections: a short anterior part, with 5 to 7 rays, and an elongate posterior part reaching to caudal-fin
base in adults, with 25 to 31 rays. Pelvic fins with I spine and 4 soft rays. Lower side of posterior part of
trunk with 2 to 4 enlarged, keel-like scales. Scales scute-like, forming prominent keels. Lateral line present
or absent (obscured by scute-like scales), if present, extending to near base of caudal fin. Swimbladder
divided at midline into 2 halves; swimbladder musculature present. Colour: usually, dusky red or purple
with brown spots above, pinkish or whitish below, changing somewhat with growth; upper surface of
pectoral-fin rays brightly coloured, usually bluish, with dark brown and often whitish spots, and young of
some species with prominent ocellus.
first 2 dorsal-fin spines

detached from fin
pectoral fins very

long, wing-like
head
armoured
scales
scute-like
Habitat, biology, and fisheries: Small to moderate-sized bottom-living

fishes. Most are taken inshore with seines over primarily sandy bottoms.
Adults forage on the bottom using the anterior part of the pectoral fins to
search for food. The young are pelagic and are often taken offshore with anterior part
lights at night. Species posses well-developed swimbladder musculature separate
and probably produce sounds. Found only in tropical or warm-temperate pectoral fin
waters. These fishes are usually of little commercial importance, but are
often taken as bycatch in nearshore fisheries. The flesh is edible and
consumed in some localities.

Triglidae: also with large, heavily armored head, but addi-
tionally with rostral projections; bony scutes present; pecto-
ral fins smaller, with lowermost rays free; lack the small
movable bone (pontinal) at the angle of the preopercle; all lower pectoral-fin rays free
dorsal fin-spines united by a fin membrane.
Triglidae
2284 Bony Fishes
Key to the species of Dactylopteridae occurring in the area

1a. A single elongate spine anterior to continuous part of spinous dorsal fin (Fig. 1a) . . . . . . . . . ® 2
1b. A spine about midway between elongate anteriormost spine and those in continuous part
of spinous dorsal fin (Fig. 1b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
a) Dactyloptena peterseni, b) other species

D. tiltoni
Fig. 1 first dorsal fin (after Eschmeyer, 1997)
2a. Snout rounded (Fig. 2a); scales on sides with a single, strong, transverse knife-like ridge
at middle of each scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactyloptena peterseni
2b. Snout pointed (Fig. 2b); scales on sides with multiple transverse ridges on each scale,
with middle ridges slightly stronger . . . . . . . . . . . . . . . . . . . . . . . . Dactyloptenta tiltoni
snout pointed snout somewhat pointed
a) Dactyloptena b) Dactyloptena c) Dactyloptena d) Dactyloptena

peterseni tiltoni gilberti orientalis
Fig. 2 dorsal view of head (after Eschmeyer, 1997)
3a. Interorbit very wide, 16 to 23% of standard length (Fig. 2c, d) . . . . . . . . . . . . . . . . . . . ® 4
3b. Interorbit moderate, 12 to 16% of standard length . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
4a. Posttemporal spine lying flat against body; granular projections on snout arranged in
rows; snout somewhat rounded and wide (Fig. 2c) . . . . . . . . . . . . . . . . Dactyloptena gilberti
4b. Posttemporal spine elevated at rear; granular projections on snout scattered uniformly;
snout somewhat pointed and narrow (Fig. 2d) . . . . . . . . . . . . . . . . . . Dactyloptena papilio
5a. Preopercular spine not extending further posteriorly than posttemporal spine; many dark
spots on pectoral fins, upper flank, and top of head in specimens over 15 cm standard
length; 1 dark ocellus about 2/3 from tip of fin in specimens between about 5 and 6.5 cm
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactyloptena orientalis
5b. Preopercular spine usually extending further posteriorly than posttemporal spine; usually
1 dark blotch, which often contains small pale spots on pectoral fins, situated about 1/2
distance from tip of fin . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactyloptena macracantha
Dactyloptena cheirophthalmus (Bleeker, 1854) 1/
0
?
0
Dactyloptena gilberti Snyder, 1909
Dactyloptena macracantha (Bleeker, 1854)
0
0
Dactyloptena orientalis (Cuvier, 1829)
0
Dactyloptena papilio Ogilby, 1910
Dactyloptena peterseni (Nyström, 1887)
0 Dactyloptena tiltoni Eschmeyer, 1997
Reference
Eschmeyer, W.N. 1997. A new species of Dactylopteridae (Pisces) from the Philippines and Australia, with a brief
synopsis of the family. Bull. Mar. Sci., 60(3):727-738.
1/ May be a synonym of Dactyloptena macracantha. Not included in the identification key.
Dactyloptena gilberti Snyder, 1909

Frequent synonyms / misidentifications: None / Dactyloptena jordani Franz, 1910.
FAO names: En - Flateared helmet gurnard.
(after Masuda et al., 1984)
Diagnostic characters: Body moderately elongate, squarish in cross-section. Head very broad, blunt,
depressed anteriorly, with a prominent keeled spine extending posteriorly from nape to below second spine
of continuous spinous part of dorsal fin. Posttemporal spine strong, but not markedly elevated above
rest of cranium. Granular projections on snout arranged in rows. Angle of preopercle with a long,
prominent spine. Eyes large. Interorbit extremely wide (width 18 to 23% of standard length) and
strongly concave, with depth 12 to 17% of head length. Mouth small, subterminal, and protractile. Upper
jaw largely obscured by bones surrounding eye. Minute teeth present on jaws, vomer, and palatines.
Anterior 2 dorsal-fin spines separated from remainder of spinous dorsal fin, the first spine elongate, the
second short; continuous part of spinous dorsal fin with V spines, followed by I short spinous point and the
soft dorsal fin with 8 rays (dorsal-fin formula: I, I, V+I, 8). Anal fin with 6 or 7 soft rays. Caudal fin elongate.
Pectoral fins with 28 to 32 rays. Pelvic fins with I spine and 4 soft rays. Scales strong, each with a distinct
ridge and no secondary ridges. Large scute-like scales posteriorly on caudal peduncle. Lateral line
present, extending to caudal fin. Colour: pectoral fins dusky, with large dark spots arranged in row
and especially dark over middle fin rays; separated anterior portion of pectoral fins paler, with no
large ocellus or spot; pectoral fins mostly black in juveniles, but without black spot or ocellus.
Habitat, biology, and fisheries: Little is known of the biology of this species. Taken nearshore with seines
or trawls and occasionally marketed fresh, it forms a minor component to the fisheries of the area. Captured
over sand or mud-sand bot-
toms at depths of 20 to 71 m.
Distribution: Ranges from
southern Japan and Gulf of
Thailand to India and the Ara-
bian Peninsula.
2286 Bony Fishes
Dactyloptena macracantha (Bleeker, 1854)

FAO names: En - Spotwing helmet gurnard; Fr - Grondin volant cocarde; Sp - Alón buho.
Diagnostic characters: Body moderately elongate, squarish in

cross-section. Head broad, blunt, depressed anteriorly, with a
prominent keeled spine extending posteriorly from nape to below
second spine of continuous spinous part of dorsal fin. Angle of
preopercle with a long, prominent spine. Eyes large. Interorbital
width 12 to 15% of standard length; interorbit weakly concave,
with depth 6 to 8% of head length. Mouth small, subterminal, and
protractile. Upper jaw largely obscured by bones surrounding eye. pectoral fin
Minute teeth present on jaws, vomer, and palatines. Anterior 2
dorsal-fin spines separated from remainder of spinous dorsal fin, the first spine elongate, the second short
(rarely missing); continuous part of spinous dorsal fin with V spines, followed by I short spinous point and
the soft dorsal fin with 8 or 9 rays (dorsal-fin formula: I, I, V+I, 8-9; rarely I, 0, V+I, 8). Anal fin with 6 soft
rays. Caudal fin elongate. Pectoral fins with 31 to 34 rays. Pelvic fins with I spine and 4 or 5 soft rays.
Scales strong, each with a distinct ridge. Large scute-like scales posteriorly on caudal peduncle. Scales
in about 45 to 47 rows. Lateral line present, extending to caudal fin. Colour: dusky violet above and
pinkish below; an oblong black blotch over middle of pectoral fins.
Size: Maximum total length 16.5 cm.
Habitat, biology, and fisheries: Typically, this species has been reported from depths of 45 to 125 m.
However, a single record from 128 to 177 m is known. Usually taken near shore with trawls or handlines
and occasionally marketed
fresh.
ranges eastward in the Indian
Ocean to India, Sri Lanka,
and western Pacific Ocean.
Within the area found in Java,
the Philippines, the Malay
Peninsula, Sumatra, Indone-
sia, and the Northern Terri-
tory (Australia).
Dactyloptena orientalis (Cuvier, 1829)

FAO names: En - Oriental helmet gurnard; Fr - Grondin volant oriental; Sp - Alón oriental.

cross-section. Head broad, blunt, depressed anteriorly, with a promi-
nent keeled spine (posttemporal) extending posteriorly from the nape
to below second spine of continuous spinous part of dorsal fin. Angle pectoral fin
of preopercle with a long, prominent spine. Eyes large. Interorbital
width 13 to 15% of standard length; interorbit moderately concave, with depth 8 to 11% of head
length. Mouth small, subterminal, and protractile. Upper jaw largely obscured by bones surrounding eye.
Minute teeth present on jaws, vomer, and palatines. Anterior 2 dorsal-fin spines separated from
remainder of spinous dorsal fin, the first spine elongate, the second short; continuous part of spinous
dorsal fin with V spines, followed by I short spinous point and the soft dorsal fin with 8 rays (dorsal-fin
formula: I, I, V+I, 8). Anal fin with 6 or 7 soft rays. Caudal fin elongate in adults (much shorter in young).
Base of pectoral fins horizontal, the fins divided into 2 sections, a short anterior part with 5 rays, and a long
posterior section with 27 to 30 rays. Pelvic fins with I spine and 4 or 5 soft rays. Scales strong, each with
a distinct ridge. Large scute-like scales posteriorly on caudal peduncle. Scales in about 45 to 47 rows.
Lateral line absent or greatly obscured. Colour: variable, usually yellowish brown above, lighter brown
below; small orange spots over top of head and back; dusky, golden spots on pectoral fins (larger distally);
4 golden bands on caudal fin; a yellow band along upper part of spinous dorsal fin.
Habitat, biology, and fisheries: This is a benthic species primarily inhabiting sandy bottoms in shallow
coastal waters. It is capable of “walking” on the bottom by alternately moving the pelvic fins and short
pectoral-fin rays. Feeds on benthic crustaceans, clams, and small fishes. The enlarged pectoral-fin rays
are spread as a defensive be-
havior. Not commercially
fished, but taken incidentally
throughout its range.
Distribution: This species is
the most widely ranging dac-
tylopterid. It has been cap-
tured from the Red Sea and
South Africa to the Tuamoto
Archipelago and Hawaii.
2288 Bony Fishes
Dactyloptena papilio Ogilby, 1910

FAO names: En - Sharpeared helmet gurnard.
Diagnostic characters: Body moderately elongate, squarish in cross-section. Head very broad, blunt,
depressed anteriorly, with a prominent keeled spine extending posteriorly from the nape to below second
spine of continuous spinous part of dorsal fin. Posttemporal spine strong and elevated above rest of
cranium. Angle of preopercle with a long, prominent spine. Eyes large. Interorbit extremely wide (width
16 to 20% of standard length) and strongly concave, with depth 12 to 17% of head length. Granular
projections on snout scattered uniformly. Mouth small, subterminal, and protractile. Upper jaw largely
obscured by bones surrounding eye. Minute teeth present on jaws, vomer, and palatines. Anterior 2
dorsal-fin spines separated from remainder of spinous dorsal fin, the first spine elongate, the second short;
continuous part of spinous dorsal fin with V spines, followed by I short spinous point and the soft dorsal
fin with 8 rays (dorsal-fin formula: I, I, V+I, 8). Anal fin with 6 or 7 soft rays. Caudal fin elongate. Pectoral
fins with 28 to 32 rays. Pelvic fins with I spine and 4 soft rays. Scales strong, each with a single median
ridge. Large scute-like scales posteriorly on caudal peduncle. Lateral line present, extending to caudal fin.
Colour: pectoral fins with group of dark spots and an irregular dark area extending 1/3 length of fin
near base; faint dark saddles on dorsum in specimens between 8 and 12 cm standard length.
Habitat, biology, and fisheries: This species is poorly known and no fishery exists for it.
Distribution: Dactyloptena
papilio is found in western
and northern Australia and
the Arafura Sea in depths of
13 to 137 m.
Dactyloptena peterseni (Nyström, 1887)

Frequent synonyms / misidentifications: Daicocus peterseni (Nyström, 1887) / None.
FAO names: En - Starry helmet gurnard; Fr - Grondin volant étoilé; Sp - Alón estrellado.

cross-section. Head broad, blunt, depressed anteriorly, with a promi-
nent keeled spine (posttemporal) extending posteriorly from the nape
to below second spine of continuous spinous part of dorsal fin. Angle
of preopercle with a long, prominent spine. Spinous and soft dorsal pectoral fin
fins separated by a deep notch. Eyes large. Interorbital width 13 to
14% of standard length; interorbit region weakly concave, its depth a midline about 2/3 width of
pupil (5 to 7% of head length). Mouth small, subterminal, and protractile. Upper jaw largely obscured by
bones surrounding eye. Nodular teeth present on jaws. Small teeth on vomer and palatines. A single
elongate filamentous spine widely separated from remainder of spinous dorsal fin, continuous part
of spinous dorsal fin with VII spines, soft dorsal fin with 8 rays (dorsal-fin formula: I, VII, 8). Anal fin
with 6 soft rays. Caudal fin emarginate. Bases of pectoral fins horizontal, the fins divided into 2 sections,
a short anterior part with 5 rays and a long posterior part, with 25 or 26 rays that reach to caudal-fin base
in adults. Lower side of posterior part of trunk with 3 enlarged keel-like scales, the first above middle
of anal fin. Scales each with a strong median ridge. Lateral line absent or greatly obscured. Colour:
dusky violet above, pinkish below; an oblong black blotch over middle of pectoral fins.
Habitat, biology, and fisheries: Benthic, found on sandy bottoms in coastal waters at depths of 20 to
71 m. It feeds primarily on benthic crustaceans. It is not fished commercially and is rarely used for food.
Most adults are taken with trawls.
occurs from southern Japan
and the South China Sea to
South Africa. The extent of its
range in the West Pacific is
uncertain as it is sometimes
confused with Dactyloptena
orientalis.
2290 Bony Fishes
Dactyloptena tiltoni Eschmeyer, 1997

FAO names: En - Plain helmet gurnard.
(after Eschmeyer, 1997)
Diagnostic characters: Body moderately elongate, squarish in cross-section. Head broad, blunt, de-
pressed anteriorly, with a prominent keeled spine (posttemporal) extending posteriorly from the nape to
below second spine of continuous spinous part of dorsal fin. Angle of preopercle with a long, prominent
spine. Eyes large. Interorbital width 13 to 14% of standard length; interorbit region weakly concave,
its depth at midline about 2/3 width of pupil (5 to 8% of head length). Mouth small, subterminal, and
protractile. Upper jaw largely obscured by bones surrounding eye. Nodular teeth present on jaws. Small
teeth on vomer and palatines. Spinous and soft dorsal fins separated by a deep notch. A single elongate
filamentous spine widely separated from remainder of spinous dorsal fin; continuous part of
spinous dorsal fin with V spines, followed by I short spinous point and the soft dorsal fin with 8
rays (dorsal-fin formula: I, V, I, 8). Anal fin with 6 soft rays. Caudal fin emarginate. Bases of pectoral fins
horizontal; pectoral fins with 29 to 32 rays, the fins divided into 2 sections, a short anterior part and a long
posterior part that reaches to caudal-fin base. Lower side of posterior part of trunk with 3 enlarged
keel-like scales, the first above middle of anal fin. Scales scute-like, with those on upper flank with
several wavy keel-like ridges, the middle ridge strongest. Lateral line absent or greatly obscured.
Colour: unknown in life, but without spots when preserved, with mostly dusky pectoral fins and faint
bands on the soft dorsal-fin rays, and with pale pelvic, anal, and caudal fins.
Size: Maximum standard length at least 9.4 cm.
ies: Nothing is known about
biology of this recently de-
scribed species other than it
appears to be the deepest liv-
ing dactylopterid; the known
specimens were trawled at
depths of 119 to 565 m.
Distribution: Known from the
Philippines and Western Aus-
tralia.
click for next page

Scorpaeniformes: Scorpaenidae 2291
SCORPAENIDAE
Scorpionfishes (also, lionfishes, rockfishes, stingfishes, stonefishes, and waspfishes)
by S.G. Poss
D iagnostic characters: Body usually weakly, rather than strongly, compressed; body depth 21 to 50%
standard length. Head moderate to large, 37 to 50% standard length, often notably depressed with
cirri, particularly above eye. Eye small to relatively large, 4 to 14% of standard length. Snout short to long,
often prominent, 6 to 20% of standard length. Mouth often large and upturned, upper jaw 9 to 23% standard
length. Numerous small conical teeth present on upper and lower jaws, with those on vomer and palatine
present or absent. Branchiostegal rays typically 7 (rarely 6). Gill rakers usually small or moderate, 1 to 9
in upper arch and 4 to 20 in lower arch. All species with suborbital stay (or ridge), an extension of the
third infraorbital bone (second suborbital) extending backward across the cheek and usually firmly
bound to preopercle. Most species with numerous head spines, with those on lacrimal bone (or first
infraorbital bone), those above orbital margin and those behind occiput most prominent. Dorsal fin with
strong venomous spinous part bearing VIII to XVIII spines connected to soft-rayed part posteriorly,
with 3 ½ to 14 soft rays, the last typically split to its base and counted as 1 ½. Anal fin with II to IV, but
usually III strong, sharp spines, the second usually longest, followed by 3 ½ to 15 soft rays, the last
usually split to its base and counted as 1 ½. Caudal fin typically rounded or truncate, never forked, 15 to
40% standard length, usually about 27 to 35% standard length. Pectoral fins usually large, with 11 to 24
rays; with rays of larger individuals of most species branched. Pelvic fins with I strong spine and 5, or less
often, 4 branched rays. Scales in most species relatively small and either ctenoid or pseudocycloid, entirely
absent in others, or present only as deeply embedded scale rudiments. Lateral-line scales present, with 4
to 54 pored or tubed scales (lateral-line scales trough-like in the subfamily Setarchinae). When present,
scales above lateral line 4 to 8; scales below lateral line 10 to 19. All species possess striated
swimbladder musculature that is extrinsic in nearly all species, with musculature present even in those
without swimbladders. Pyloric caecae 1 to 16. Vertebrae 24 to 29. Colour: most species strongly
camouflaged and red, reddish brown, or brown in colour, and usually have barred or mottled colour patterns
that are typically darker dorsally than ventrally.
VIII-XVIII
dorsal-fin spines
supraocular cirrus dorsal fin
(some species) continuous, notched
caudal fin usually
rounded
suborbital stay preopercle

(or ridge) spiny III (II-IV) anal-fin
spines lacrimal bone (shape and
spines often used as specific
diagnostic characters)
Habitat, biology, and fisheries: Scorpionfishes and their near relatives
are typically found on or near the bottom, which they often strongly
resemble. Most species in the area are found on relatively nearshore hard
bottoms and reefs or associated with coral rubble, from the surface to a
depth of 150 m. Some species in the area range, into deeper waters (to
800 m), although outside the area captures to 1 113 m have been re-
ported. A few species, such as those of the genera Setarches, Lioscor-
pius, and Ectreposebastes, are pelagic or semipelagic occurring offshore
in depths of 200 to 800 m. Many species are relatively small, typically
under 20 cm standard length, and their biology poorly studied. Nonethe-
less, most are known to lead solitary lives, and evidently aggregate only
for reproduction. The young of most species are planktonic, with many detail of head
2292 Bony Fishes
settling out of the plankton relatively quickly. postorbital spine lower posttemporal spine
Most feed primarily on arthropods and many
feed on small fishes as they attain larger
sizes. Most species are extremely well upper
camouflaged and excellent ambush preda- posttemporal
spine
tors. The vividly (aposematically) coloured
lionfishes or turkeyfishes are notable excep-
tions, cornering their prey with their elongate
pectoral fins. Most scorpionfishes are ovovi-
viparous, producing between a few hundred
and a few thousand eggs, although some
are viviparous. Nearly all possess well-de-
veloped venom glands and should be han-
dled with extreme caution, lest painful and
potentially fatal wounds be inflicted by their
sharp fin and head spines. Although all are nasal supraocular
edible, most species in the Western Central spine tympanic upper
spine posttemporal
Pacific are small and dangerous to handle postocular spine
spine spine
and thus do not form the basis of large lower
fisheries. However, a few species in the area posttemporal
spine
are relatively large, occur in considerable
number, and are marketed fresh. Numerous
species outside the area for important fish-
eries.
Remarks: Scorpaenoid fishes form a large
(approximately 500 species) and hetero- opercular
genous assemblage of fishes. The limits of spines
the Scorpaenidae, included subfamilies,
and associated families are not well estab-
lished nor is there agreement on what family postorbital
spine
or subfamily names should be used. Some
phylogenetically derived taxa, such as the
Synanceiinae and Tetraroginae, are often
treated as distinct families by many authors,
whereas such authors usually regard more lacrimal
spines
phylogenetically distant relatives as belong-
suborbital ridge (and spines) preopercular spines
ing to the Scorpaenidae. Other derived taxa,
such as Caracanthidae and Aploactinidae, diagram of head spines used in the identification key
are almost universally regarded as separate (after Eschmeyer, 1969)
families, although they too are more closely
related than are some genera invariably in-
cluded in the Scorpaenidae. For purposes of this general treatment a broad definition of the Scorpaenidae
is adopted. With a few commonly accepted exceptions, this avoids use of an unfamiliar and highly split
classification.
Fishes of several other bony-fish families are superficially similar to scorpaenids in general appearance.
Like all near relatives (Triglidae, Platycephalidae, Caracanthidae, and Aploactinidae), scorpaenids pos-
sess a bony suborbital stay below and behind the eye that attaches to the preopercle. In addition, most
scorpionfishes bear numerous head spines not seen in species otherwise of similar colour or body shape
(see above diagram of head spines).
Triglidae: possess a very broad suborbital stay; all
species have heavily armoured heads and free, highly
mobile lowermost pectoral-fin rays (only few scor-
paenids possess heavily armoured heads or free pec-
toral-fin rays, and none have these characteristics in
combination as do sea robins, except for the Apisti-
nae, which can be distinguished from triglids by their
movable lacrimal bone). lowermost pectoral-fin rays free
Triglidae
Platycephalidae: possess similar head spines, but are distinguished by their highly depressed head (head
depth less than about 20% standard length), and by having the ventral margin of the lacrimal bone smooth
(ventral margin of lacrimal often bearing numerous large spines in scorpaenids; see diagram of head spines
on previous page).
Caracanthidae: also have a suborbital stay and similar head spines anterior and posterior to the eye, but
are distinguished by their notably compressed heads and bodies with rounded profiles that are covered
with numerous fleshy papillae-like projections not seen in scorpaenids (which usually have ctenoid or
pseudocycloid scales; a few species lack scales).
Platycephalidae Caracanthidae
Aploactinidae: closely related to the subfamily Synanceiinae and also have a suborbital stay and head
spines, but most aploactinids are distinguished by the possession of highly modified tack-like scales (but
many are without scales); all aploactinids lack branched fin rays (as seen in nearly all scorpaenids, except
subfamilies Synanceiinae and Minoinae), and most have comparatively blunt spines (instead of strongly
pungent spines seen in scorpaenids).
Serranidae: similar to some scorpaenids in head and body shape, but often having concave, lunate, or
forked caudal fins; no suborbital stay under the eye that attaches to preopercle; often 3 opercular spines
(typically only 2 in scorpaenoids); many species possess large canine teeth common in anterior end of
upper and lower jaws (in contrast to the comparatively uniform small teeth of scorpaenoids).
Aploactinidae Serranidae
Centrogeniidae: the only member of this family,
Centrogenys vaigiensis (false scorpionfish, some-
times placed in the Serranidae), sufficiently resem-
bles scorpaenids in coloration and size and shape
of its fins, perhaps for purposes of mimicry, that it
was originally described in the genus Scorpaena.
However, C. vaigiensis does not possess a subor-
bital stay that extends back toward the eye and
firmly attaches to the preopercle, nor does it have
venom glands associated with the fin spines.
Identification note Centrogeniidae
Many scorpaenoid species, but not all, have the last fin ray of the dorsal and anal fins split to their base
yet borne on a single pterygiophore, as revealed by radiography. Workers have differed in how such fin
rays are counted, some counting this condition as 1 ray, others as 2. A convention has developed among
scorpaenoid workers to count this last double ray as 1 ½. This method is used here as it has the advantage
of permitting the split-rayed condition to be distinguished from the single-rayed condition, while eliminating
the ambiguity inherent in counting the split ray as either 1 or 2 rays. Vertical scale row counts are taken
above the lateral line from the uppermost posttemporal spine to the base of the caudal fin. Gill raker counts
are presented as formulas, with the first range indicating the rakers in the upper arm of the first gill arch,
the middle number indicating a gill raker at the angle between the upper and lower arms and the last range
referring to the rakers on the lower arm (e.g. 3-5+1+7-11=12+16) with the range after the equal sign
indicating the total number of gill rakers.
2294 Bony Fishes
Key to the species of Scorpaenidae occurring in the area

Note: in the anticipation that they may be recorded in the future from the area, some species known from
adjoining seas are included in the key, with their distribution indicated in parenthesis.
1a. Skin at gill openings broadly connected to isthmus . . . . . . . . . .(subfamily Syanceiinae) ® 2

1b. Skin at gill openings connected to each other or connected to isthmus only narrowly
anteriorly, not broadly connected to isthmus (partially united in Taenianotus triacanthus) . . . . . ® 26
2a. Pectoral fins with lowermost 1 to 3 rays not free from other rays . . . . . . . . . . . . . . . . . ® 3
2b. Pectoral fins with lowermost 1 to 3 rays free, and independently mobile . . . . . . . . . . . . . ® 9
3a. Mouth terminal, slightly oblique; eyes lateral on head . . . . . . . . . . . . . . . . . . (Erosa) ® 4

3b. Mouth strongly upturned; eyes on dorsal surface of head . . . . . . . . . . . . . . . . . . . . ® 5
4a. Pectoral-fin rays 14 to 16, usually 15; dorsal-fin spines XIII to XV, usually XV; dorsal-fin
rays 5 ½ or 6 ½, usually 6 ½; anal-fin spines III; anal-fin rays 5 ½ or 6 ½ . . . . . . . . Erosa erosa
4b. Pectoral-fin rays 12 or 13, usually 12; dorsal-fin spines XII or XIII; dorsal-fin rays 8 ½
or 9 ½ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erosa daruma
(Western Australia; not yet recorded from the area)
5a. Dorsal-fin spines XVI or more . . . . . . . . . . . . . . . . . . . . . . Leptosynanceia asteroblepa
5b. Dorsal-fin spines XI to XIV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Anal fin with II spines and 12 to 15 soft rays, the last split to base . . Trachicephalus uranoscopus
6b. Anal fin with III spines and 4 ½, 5 to 6 ½, or 7 soft rays . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Pectoral-fin rays 10 to 15, usually 11 . . . . . . . . . . . . . . . . . . . . . . . . . Synanceia alula

7b. Pectoral-fin rays 14 to 19 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Orbital margin heavily ossified, forming a “crest” above eye; pectoral-fin rays 15 to 17
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synanceia horrida
8b. Orbital margin not heavily ossified, not forming a “crest” above eye; pectoral-fin rays
17 to 19 (usually 18 or 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Synanceia verrucosa
9a. Only lowermost pectoral-fin ray free and relatively mobile . . . . . . . . . . . . . . (Minous) ® 11
9b. Lowermost 2 or 3 pectoral-fin rays free and relatively mobile . . . . . . . . . . . . . . . . . . ® 10
10a. Dorsal-fin spines XIII to XV; lowermost 3 pectoral-fin rays free; eyes small to moderate
(about 7 to 11% head length), positioned laterally on head . . . . . . . Choridactylus multibarbus
10b. Dorsal-fin spines XV to XVIII; lowermost 2 pectoral-fin rays free; eyes small (about 4
to 7% head length), positioned on top of head . . . . . . . . . . . . . . . . . . . . (Inimicus) ® 17
11a. First dorsal-fin spine equal to or longer than second dorsal-fin spine; first spine widely
separated from base of second spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
11b. First dorsal-fin spine notably shorter than second dorsal-fin spine (first spine typically
less than 1/2 length of second spine); first dorsal-fin spine close to base of second
dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
12a. Caudal fin without transverse dark bars; medial (axillary) surface of pectoral fin with
dark brown stripes; posterior spine of lacrimal bone about twice length of anterior spine
and not bayonet-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minous quincarinatus
12b. Caudal fin with transverse dark bars; medial (axillary) surface of pectoral fin either
uniformly coloured or spotted; posterior spine of lacrimal bone long and bayonet-
shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Soft portion of dorsal fin with wavy bands; dorsal-fin spines usually IX; caudal fin usually
with 3 or 4 transverse wavy bands . . . . . . . . . . . . . . . . . . . . . . . . . Minous versicolor
13b. Soft portion of dorsal fin black anteriorly; dorsal-fin spines usually X or more; caudal
fin with 2 broad, dark vertical bars . . . . . . . . . . . . . . . . . . . . . . . Minous monodactylus
14a. Dorsal-fin spines weak and hair-like . . . . . . . . . . . . . . . . . . . . . . . . . Minous pusillus

14b. Dorsal-fin spines strong or thin, but not hair-like . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
15a. Posterior lacrimal spine about equal in length to anterior lacrimal spine; soft portion of
dorsal fin with 8 to 11 rays, usually 10; anal-fin rays plus anal-fin spines together 9 to
11; caudal-fin rays usually with alternating dark and pale markings . . . . . Minous trachycephalus
15b. Posterior lacrimal spine much longer than anterior lacrimal spine; soft portion of dorsal
fin with 11 to 13 rays; anal-fin rays plus anal-fin spines together 11 to 13; caudal fin
pale, without dark markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
16a. Medial (axillary) surface of pectoral-fin stripes radiating distally along course of fin rays
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minous pictus
16b. Medial (axillary) surface of pectoral fin with irregular black spots on a pale background
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Minous coccineus
17a. Snout shorter than postorbital length (0.7 to 0.9 in postorbital length); fin membranes
of dorsal-fin spines subsequent to third spine incised to about midlength of spines . . . . . . ® 18
17b. Snout about equal to, or usually, longer than postorbital length (0.9 to 1.7 in postorbital
length); fin membranes of dorsal-fin spines subsequent to third spine incised for nearly
entire length of spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
18a. Medial (axillary) surface of pectoral fins dusky or blackish, often with black spots or
streaks, but without a transverse white band in middle of fin . . . . . . . Inimicus brachyrhynchus
18b. Medial (axillary) surface of pectoral fins dusky, with black spots, and a transverse band
in middle of fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
19a. Third pectoral-fin ray from bottom not free, except at its tip . . . . . . . . . . . Inimicus japonicus
19b. Third pectoral-fin ray from bottom free for about 1/3 of its length . . . . . . . . . . Inimicus joubini
20a. Orbits extremely elevated and closely spaced at their bases, which are broadly joined;
interorbit narrow, about equal to orbit diameter; uppermost 2 pectoral-fin rays filamen-
tous in juveniles and adults . . . . . . . . . . . . . . . . . . . . . . . . . . . Inimicus filamentosus
20b. Orbits slightly elevated and widely spaced at their bases, which are joined only by a
low ridge; interorbit wide, usually 1.5 times orbit diameter; uppermost 2 pectoral-fin
rays not filamentous in specimens longer than about 5 cm standard length . . . . . . . . . . ® 21
21a. Medial surface of pectoral fins nearly uniform grey, without markings, except with
sometimes a few scattered dark spots . . . . . . . . . . . . . . . . . . . . . . . . Inimicus cuvieri
21b. Medial surface of pectoral fins not uniformly coloured in adults . . . . . . . . . . . . . . . . . ® 22
22a. Medial surface of pectoral fins with a broad terminal dark band in adults . . . . . . . . . . . . ® 23
22b. Medial surface of pectoral fins without a broad terminal dark band in adults, or with a
thin dusky band terminally in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 24
23a. Medial surface of pectoral fins with broad dark transverse bar near proximal end of fin
in adults, which may be streaked with white lines extending along fin rays and a white
area separating it from terminal dark band . . . . . . . . . . . . . . . . . . . . Inimicus didactylus
23b. Medial surface of pectoral fins without a broad dark transverse bar near proximal end
of fin in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .® 25
2296 Bony Fishes
24a. Medial surface of pectoral fins either with irregular dusky spots proximally or entirely
pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Inimicus gruzovi
24b. Medial surface of pectoral fins with irregular dusky spots proximally and 2 large dark
spots on an otherwise pale background . . . . . . . . . . . . . . . . . . . . . . Inimicus smirnovi
25a. Medial surface of pectoral fins with irregularly-sized pale spots on a dark grey back-
ground . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Inimicus sinensis
25b. Medial surface of pectoral fins with 2 dark regions separated by a pale area . Inimicus caledonicus
26a. Lacrimal bone (infraorbital 1) highly mobile, hinged to lateral ethmoid dorsally and
abutting but not firmly bound to first suborbital bone (infraorbital 2) posteriorly . . . . . . . . . ® 27
26b. Lacrimal bone (infraorbital 1) relatively immobile; strongly bound to the lateral-ethmoid
dorsally and to first suborbital bone (infraorbital 2) posteriorly . . . . . . . . . . . . . . . . . ® 51
27a. Ventralmost 1 or 3 pectoral-fin rays detached or separated from rest of fin so that
independent movement is possible; second suborbital bone (third infraorbital) very
deep (about as deep as long) covering nearly entire cheek and very broadly connected
to preopercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Subfamily Apistinae) ® 28
27b. Ventralmost pectoral-fin rays not detached or separated from more dorsal-fin rays;
second suborbital (third infraorbital) notably longer than deep, not covering entire
cheek, and usually not forming wide connection to preopercle . . .(Subfamily Tetraroginae) ® 30
28a. Ventralmost 3 pectoral-fin rays not detached or separate from more dorsal-fin rays; no
barbels on lower jaw; no ocellus in dorsal fin . . . . . . . . . . . . . . . Cheroscorpaena tridactyla
28b. Ventralmost pectoral-fin ray not detached or separate from more dorsal-fin rays;
barbels on lower jaw; a pronounced ocellus at rear of spinous dorsal fin . . . . . . . . . . . . ® 29
29a. Orbit 8 to 10% head length, interorbit 13 to 17% of head length and 18 to 24% of body
depth in specimens greater than about 4 cm standard length; head bones of larger
specimens relatively smooth, not densely covered with numerous denticulations . Apistops caloundra
29b. Orbit 3 to 6% head length, interorbit 6 to 10% of head length and 6 to 13% of body
depth in specimens greater than about 4 cm standard length; head bones of larger
specimens densely covered with numerous denticulations, often closely set and
arranged in rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apistus carinatus
30a. Numerous cycloid scales present on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 31

30b. Scales absent on body or few that are present form weakly developed spinous points . . . . ® 46
31a. Dorsal-fin origin distinctly posterior to posterior margin of orbit . . . . . . . . . . . . . . . . . ® 32

31b. Dorsal-fin origin distinctly anterior to posterior margin of orbit . . . . . . . . . . . . . . . . . ® 34
32a. Occipital region large, relatively elongate . . . . . . . . . . . . . . . . . . . . . . Notesthes robusta

32b. Occipital region greatly foreshortened . . . . . . . . . . . . . . . . . . . . . . (Centropogon) ® 33
33a. A broad naked area below anterior part of spinous dorsal fin; jaw short, 10 to 15%
standard length; body with strong dark saddles . . . . . . . . . . . . . . . . Centropogon australis
33b. No broad naked area below anterior part of spinous dorsal fin; jaw moderate, 15 to 19%
standard length; interorbital ridges relatively weak; body marmorated . . . Centropogon marmoratus
34b. Dorsal fin with anterior 3 dorsal-fin spines forming a nearly separate fin, with fin
membrane deeply incised posterior to third spine . . . . . . . . . . . . . . . . . (Vespicula) ® 43
34a. Dorsal fin continuous with anterior 3 dorsal-fin spines not forming a nearly separate fin . . . . ® 35
35a. Pelvic fins with I spine and 4 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .® 36

35b. Pelvic fins with I spine and 5 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .® 38
36a. Pectoral fins with 13 to 15 rays, usually 14; body depth 39 to 46% standard length;
upper jaw 17 to 19% standard length; scales in about 40 to 45 vertical rows; gill rakers
3-5+1+7-11=12-16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liocranium praepositum
36b. Pectoral fins with 10 or 11 rays; body depth 31 to 38% standard length; upper jaw 12
to 15% standard length; scales in 44 to 66 vertical rows; gill rakers 2-3+1+4-7=7-11
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Paracentropogon) ® 37
37a. Lateral-line scales 19 to 24; dorsal-fin spines XII to XV (usually XIII or XIV, rarely XV);
pectoral-fin rays 10 or 11 (rarely 11); total gill rakers on first gill arch (including
rudiments) typically 7 to 10, rarely 11 . . . . . . . . . . . . . . . . . Paracentropogon longispinis
37b. Lateral-line scales 16 to 18; dorsal-fin spines XV; pectoral-fin rays 11; total gill rakers
on first gill arch 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracentropogon zonatus
38a. Dorsal fin with XVII or XVIII spines; upper jaw small, about 12 to 14% standard length
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Ablabys) ® 39
38b. Dorsal fin with XIV to XVI spines; upper jaw moderate or large, about 16 to 25%
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 41
39a. Dorsal fin with XVII or XVIII spines (usually XVII) and 6 ½ or (usually) 7 ½ soft rays;
anal fin with III spines and 4 ½ or (usually) 5 ½ soft rays . . . . . . . . . . . Ablabys taenianotus
39b. Dorsal fin XV or XVI spines and 8 ½ to 10 ½ soft rays; anal fin with III spines and 7 or
8 ½ soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ablabys macracanthus
40a. Gill rakers elongate, rakers 4-6+1+12-14=18-21; body depth 34 to 42% standard
length; scales in 64 to 68 vertical rows . . . . . . . . . . . . . . . . . . . . . . Cottapistus cottoides
40b. Gill rakers stout, rakers 2-5+1+6-12=9-17; body depth 29 to 36% standard length;
scales in 74 to 95 vertical rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 41
41a. Palatine teeth absent; body depth 34 to 42% standard length; upper jaw 14 to 17%
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Snyderina yamanokami
(based on a single unverified record from Indonesia)
41b. Palatine teeth present; body depth 29 to 33% standard length; upper jaw 17 to 19%
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Neocentropogon) ® 42
42a. Body with a large blotch behind head above pectoral fins, but no other distinct dark
markings elsewhere; scales in about 85 to 95 vertical rows on body . . . . Neocentropogon affinis
42b. Body with a blotch behind head above pectoral fins, but with 2 dark blotches on dorsal
fin and numerous spots over body; scales in about 74 to 85 vertical rows . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neocentropogon trimaculatus
43a. Dorsal fin with XIII to XVI spines and 3 ½ to 5 ½ soft rays; anal fin with III spines and
3 ½ or 4 ½ soft rays; orbit 6 to 9% standard length . . . . . . . . . . . . . . Vespicula trachinoides
43b. Dorsal fin with XIII or XIV spines and 6 ½ or 7 ½ soft rays; anal fin with III spines and
4 ½ or (usually) 5 ½ soft rays; orbit 9 to 12% standard length . . . . . . . . . . . . . . . . . . ® 44
44a. Pectoral-fin rays 11; orbit 9 to 10% standard length; body notably compressed . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Vespicula depressifrons
44b. Pectoral-fin rays 12; orbit 11 to 13% standard length; body compressed but not greatly so . . . . ® 45
2298 Bony Fishes
45a. Dorsal fin with XIII or XIV spines and 6 ½ soft rays; body depth about 31 to 37%
standard length; body nearly uniform brown . . . . . . . . . . . . . . . . . . . Vespicula zollingeri
45b. Dorsal fin with XIII spines and 7 ½ soft rays; body depth about 37% standard length;
body with spotted or reticulate pattern . . . . . . . . . . . . . . . . . . . . . . . . Vespicula cypho
46a. Body entirely without scales, except for lateral line . . . . . . . . . . . . . . . . . . . . . . . . ® 47

46b. Body with a few scattered tack-like or rudimentary scales (most evident above lateral
line behind head, but must be closely examined), as well as on lateral-line scales . . . . . . . ® 50
47a. Body robust, not strongly compressed; head profile rounded; pectoral-fin rays 14 to 16
(rarely 13); dorsal fin with XII to XIV spines and 5 to 9 segmented rays . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Richardsonichthys leucogaster
47b. Body notably compressed; head profile angular; pectoral-fin rays 11 to 13 (usually 12);
dorsal fin with XIV to XVII spines and 7 to 9 segmented rays . . . . . . . . . . . . (Ocosia) ® 48
48a. Second and third dorsal-fin spines notably elongate relative to succeeding spines; no
small spine on lower margin of second infraorbital bone; no small spine on lateral face
of lacrimal bone; body with distinct brown spots . . . . . . . . . . . . . . . . . . . Ocosia zaspilota
48b. Second dorsal-fin spine notably elongate relative to succeeding spines; a small spine
on lower margin of second infraorbital bone; a small spine usually present on lateral
face of lacrimal bone; small flecks of brown on sides, but no distinct spots . . . . . . . . . . . ® 49
49a. Ratio of length of second dorsal-fin spine to length of third spine 1.6 to 1.7; pectoral-fin
rays 12 or 13 (usually 12); gill rakers 2-5+1+8-15=12-21 . . . . . . . . . . . . . . . . . Ocosia apia
49b. Ratio of length of second dorsal-fin spine to length of third spine about 1.15, with both
notably longer than succeeding spines; gill rakers about 4+1+9=15 . . . . . . . . . Ocosia spinosa
(Taiwan Province of China; not yet recorded from the area)
50a. Anal fin with III spines and 4 ½ or 5 ½ soft rays; gill rakers 2-3+1+5-7=8-10 . . . Tetraroge barbata
50b. Anal fin with III spines and 6 ½ or 7 ½ soft rays; gill rakers 3-4+1+6-7=10-11. . . . Tetraroge niger
51a. Pectoral fins strongly notched or notably bilobate with upper rays longest. . . . . . . . . . . . ® 52
51b. Pectoral fins rounded or elongate, but not strongly notched or bilobate (somewhat
bilobate in some species of Neosebastes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 53
52a. Pectoral-fin rays 22 or 23; dorsal-fin spine X extremely short, with membrane between
it and spines IX and XI absent or nearly so . . . . . . . . . . . . . . . . . . Plectrogenium nanum
52b. Pectoral-fin rays 20 to 22; dorsal-fin spine X short, but not extremely so, with membrane
between it and spines IX and XI normally developed as between other spines . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Trachyscorpia capensis
(New Zealand; not yet recorded from the area)
53a. Opercle usually with a single weak ridge that typically ends in a small spine; lacrimal
and suborbital bones (infraorbital bones 1 to 3) relatively broad, flat, and thin; postor-
bital always absent; dorsal-fin membranes incised nearly entire length of all spines;
dorsal-fin spines greatly elongate in most species . . . . . . . . . . . (subfamily Pteroinae) ® 54
53b. Opercle with 2 distinct ridges that diverge at an acute angle, both usually strongly
developed and bearing spines; lacrimal and suborbital bones (infraorbital bones 1 to
3) relatively narrow and usually somewhat convex laterally and usually strongly
ossified; lacrimal bone with strong spines along ventral margin; dorsal-fin spines short,
usually less than 1/2 body depth; dorsal-fin membranes not incised nearly entire length
of all spines; dorsal-fin spines not greatly elongate in most species . . . . . . . . . . . . . . ® 67
54a. Pectoral-fin rays unbranched in juveniles and adults . . . . . . . . . . . . . . . . . (Pterois) ® 61

54b. Pectoral-fin rays branched in juveniles and adults . . . . . . . . . . . . . . . . . . . . . . . . ® 55
55a. Dorsal-fin spines relatively short, less than 1/2 body depth and nearly equal to length
of dorsal-fin rays; mandible scaled, with serrated ridges . . . . . . . . . . Brachypterois serrulatus
55b. Dorsal-fin spines long, usually longer than body depth and much longer than dorsal-fin
rays; mandible without serrate ridges or scales . . . . . . . . . . . . . . . . . . . . . . . . . ® 56
56a. Anal fin with II spines (III spines in juveniles) and 7 ½ or 8 ½ soft rays; lateral surface
of lacrimal bone with numerous ventrally directed spines; caudal fin with outer rays
elongate and becoming filamentous in larger individuals . . . . . . . . . . . Parapterois heterurus
56b. Anal fin with III spines and 5 ½ to 8 ½ soft rays; lateral surface of lacrimal bone without
ventrally directed spines; caudal fin with outer rays never elongate . . . . . . . . . . . . . . . ® 57
57a. Preopercle with IV spines, with third from dorsalmost notably enlarged and often with
multiple points; males bearing a thin bony crest above orbit; pectoral-fin rays 15 to 18
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ebosia bleekeri
(Taiwan Province of China; not yet recorded from the area)
57b. Preopercle with III spines, none notably enlarged; males without a bony crest above
orbit; pectoral-fin rays 17 to 21 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 58
58a. Two (rarely 1 or 3) large ocelli on soft part of dorsal fin; 2 large cirri on snout; pectoral-fin
rays 20 or 21 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrochirus biocellatus
58b. No ocelli on soft part of dorsal fin; nasal without cirrus; pectoral-fin rays 17 or 18 . . . . . . . ® 59
59a. Body with broad red vertical bars on a pale background; vertical scale rows about 34;
snout 8 to 10% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrochirus bellus
59b. Body with red or reddish brown saddles or with broad red vertical bars that alternate
with thinner red or reddish brown bars; vertical scale rows 45 to 54; snout 11 to 15%
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 60
60a. Dorsal fin with XII or XIII spines and 9 ½ or 10 ½ soft rays (usually 9 ½); anal fin with
III spines and 5 ½ soft rays; supraocular cirrus absent or shorter than orbit diameter
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dendrochirus brachypterus
60b. Dorsal fin with XIII spines and 10 ½ or 11 ½ soft rays (usually 10 ½); anal fin with III
spines and 6 ½ or 7 ½ soft rays; supraoccular cirrus usually well developed and
typically longer than orbit diameter . . . . . . . . . . . . . . . . . . . . . . . . Dendrochirus zebra
61a. Vertical scale rows more than 65 (65 to 80 in Pterois russellii) . . . . . . . . . . . . . . . . . ® 62

61b. Vertical scale rows less than 60 (50 to 60 in Pterois lunulata) . . . . . . . . . . . . . . . . . ® 64
62a. Ventral surface of mandible with numerous longitudinal dark stripes; pectoral-fin rays
14; caudal fin and soft parts of dorsal and anal fins spotted . . . . . . . . . . . . . . . . . . ® 63
62b. Ventral surface of mandible pallid, without longitudinal dark stripes; pectoral-fin rays
13; caudal fin and soft parts of dorsal and anal fins without spots . . . . . . . . . . Pterois russellii
63a. Dorsal-fin rays 9 ½ to 11 ½ (almost always 10 ½); anal-fin rays 5 ½ or 6 ½ (almost

always 6 ½) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterois miles
63b. Dorsal-fin rays 10 ½ to 12 ½ (almost always 11 ½); anal-fin rays 5 ½ to 7 ½ (almost
always 7 ½) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterois volitans
64a. Pectoral-fin rays 12 to 14 (usually 13); anal fin with III spines and 7 ½ or 8 ½ soft rays;
no spots on soft-rayed part of dorsal, caudal, and anal fins; vertical scale rows 50 to
60; scales on flank mostly cycloid . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterois lunulata
64b. Pectoral-fin rays 16 to 20; anal fin with III spines and 5 ½ or 6 ½ soft rays; spots on
soft-rayed part of dorsal, caudal, and anal fins; vertical scale rows less than 60; scales
on flank mostly ctenoid . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 65
2300 Bony Fishes
65a. Dorsal fin with XIII spines and 10 ½ soft rays; pectoral-fin rays 18 or 19; supraocular
cirrus usually poorly developed or absent . . . . . . . . . . . . . . . . . . . . . Pterois mombasae
65b. Dorsal fin with XII spines and 10 ½ to 12 ½ soft rays; pectoral-fin rays 16 to 18 (usually
16 or 17); supraocular cirrus well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 66
66a. About 5 broad dark bars on body, bordered by distinct white lines; broad horizontal
stripe on caudal peduncle; supraocular cirrus not banded . . . . . . . . . . . . . . . Pterois radiata
66b. Numerous dark bars on body; thin diagonal bars on caudal peduncle; supraocular cirrus
with alternating light and dark bands . . . . . . . . . . . . . . . . . . . . . . . . Pterois antennata
67a. Lateral-line scales weakly ossified and barely roofing an otherwise open trough; bones
of cranium relatively weakly ossified . . . . . . . . . . . . . . . . . (subfamily Setarchinae) ® 68
67b. Lateral-line scales forming relatively complete tubes, that are sometimes buried; bones
of cranium strongly or moderately ossified . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 71
68a. Anal fin with II spines; greatest body depth less than 28% of standard length; 2 or 3
spines immediately anterior to last dorsal-fin spine extremely small, sometimes cov-
ered by scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lioscorpius longiceps
68b. Anal fin with III spines; greatest body depth greater than 30% of standard length; 2 or
3 spines immediately anterior to last dorsal-fin spine small, but at least 1/5 length of
last spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 69
69a. Anteriormost lacrimal spine much shorter than posterior 2; anal fin with usually 6 ½ soft
rays; eye relatively small with orbit diameter roughly 1/2 of interorbital width . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ectreposebastes imus
69b. Anteriormost lacrimal spine about as long as posterior 2; anal fin with usually 5 ½ soft
rays; eye relatively large with orbit diameter roughly equal to interorbital width . . (Setarches) ® 70
70a. Second preopercular spine nearly equal to or longer than uppermost (first) or third;
interorbital width 7 to 9% standard length . . . . . . . . . . . . . . . . . . . . Setarches guentheri
70b. Second preopercular spine reduced or absent, much shorter than uppermost (first) or
third; interorbital width 9 to 12% standard length . . . . . . . . . . . . . . . Setarches longimanus
71a. Lateral line normal, continuing onto or near base of caudal fin, with 20 to 54 scales . . . . . . ® 72
71b. Lateral line incomplete, continuing for short distance onto flank only, with 4 to 8 scales
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Phenacoscorpius megalops
72a. Posterior lacrimal spine hooked forward (not pronounced in small juveniles) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Parascorpaena) ® 73
72b. Posterior lacrimal spine absent or not hooked forward . . . . . . . . . . . . . . . . . . . . . ® 76
73a. Suborbital ridge with 3 spines or spinous points, first ventral to eye, second and third
posterior to eye and close together; pectoral-fin rays usually 15 or 16 . . . . . . . . . . . . . ® 74
73b. Suborbital ridge with 2 spines or spinous points behind eye (no spinous point ventral
to eye); pectoral-fin rays usually 17 or 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 75
74a. Spinous dorsal fin of males with a distinct black blotch, mostly between spines VIII to
XI (females with smaller or no spot); supraocular cirrus usually small or absent; reddish
in life, not strongly mottled with dark brown . . . . . . . . . . . . . . . . Parascorpaena mcadamsi
74b. Spinous dorsal fin without black blotch in either sex; supraocular cirrus, when present,
relatively large; mottled with dark brown; with 2 pale areas on caudal peduncle . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parascorpaena mossambica
75a. Interorbital ridges well developed from middle of eye posteriorly, joining to form a broad
loop at rear of interorbit and enclosing a depression; depression at occiput fairly well
marked; tympanic spines usually closer together than are postocular spines . Parascorpaena aurita
75b. Interorbital ridges weakly or moderately developed, not joining to enclose a depression;
depression at occiput shallow or of moderate depth; tympanic spines usually further
apart than are postocular spines . . . . . . . . . . . . . . . . . . . . . . . . . Parascorpaena picta
76a. Scales on flank ctenoid, particularly above lateral line; head and body sometimes
slightly compressed but not markedly so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 84
76b. Scales on flank cycloid or each reduced to form small spiny point and widely scattered
on body; head and body strongly compressed, at times extremely so . . . . . . . . . . . . . . ® 77
77a. Scales reduced to small widely scattered points; body extremely compressed; dorsal
fin without or with only weak notch anterior to last dorsal-fin spine . . . . . Taenianotus triacanthus
77b. Scales cycloid; body notably compressed but not extremely so; dorsal fin with relatively
pronounced notch immediately anterior to last dorsal-fin spine . . . . . . . . . . . . . . . . . ® 78
78a. Body not notably elevated, with body depth less than about 36% standard length . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(Pteroidichthys) ® 83
78b. Body notably elevated, with body depth 38 to 54% standard length . . . . . . . . . . . . . . . ® 79
78a. Vertical scale rows 30 to 45 . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinopias filamentosa

79b. Vertical scale rows 65 to 80 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 80
80a. Pectoral fins with 15 or 16 rays; an ocellus or black spot on soft part of dorsal fin . . . . . . . ® 81
80b. Pectoral fins with 17 or 18 rays; no ocellus or black spot on soft part of dorsal fin . . . . . . . ® 82
81a. Head, body, and fins with round and oblong spots or blotches, with spots typically with
pale centres and dark margins . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinopias frondosa
81b. Head, body, and fins with dark reticulations or intricate markings on a pale background
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rhinopias aphanes
82a. Third dorsal-fin spine about 16% standard length; mandibular cirri absent . . . Rhinopias argoliba
82b. Third dorsal-fin spine about 24 to 36% standard length; mandibular cirri present . . Rhinopias xenops
83a. Anal fin with II spines and 6 ½ soft rays . . . . . . . . . . . . . . . . . Pteroidichthys amboinensis
83b. Anal fin with III spines and 5 ½ soft rays . . . . . . . . . . . . . . . . . . .Pteroidichthys noronhai
84a. Third infraorbital bone (suborbital 2) inclined ventrally at an angle of about 30° to 45°
and only narrowly connected to preopercle; dorsal-fin rays usually 11 ½, or more;
ventral margin of lacrimal bone typically without spines or with weak spines . . . Sebastiscus tertius
84b. Third infraorbital bone (suborbital 2) extends nearly straight posteriorly and relatively
broadly connected to preopercle; ventral margin of lacrimal bone usually with numerous
spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 85
85a. Dorsal-fin spines XIII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 86

85b. Dorsal-fin spines XII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 100
86a. Teeth on palatines present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 87

86b. Teeth on palatines absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 89
2302 Bony Fishes
87a. Lengths of longest dorsal-fin spines (third or fourth) much shorter than body depth . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Thysanichthys crossotus
(presence in the area uncertain; reported from northeastern Taiwan Province of China)
87b. Lengths of longest dorsal-fin spines (third or fourth) about equal to body depth . . . . . . . . ® 88
88a. Mandibular foramina each leading to a single large opening . . . . . . . . Neosebastes incispinnis
88b. Mandibular foramina each leading to numerous tiny perforations in membrane covering
each formen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neosebastes entaxis
(Taiwan Province of China, Western Australia; not yet recorded from the area)
89a. Anal fin with II spines and 6 ½ or 7 ½ soft rays (occasionally with III spines and 5 ½
rays); first anal-fin spine rudimentary, about 20% length of second spine . . Hoplosebastes armatus
(Japan, Hong Kong, Taiwan Province of China; not yet recorded from the area)
89b. Anal fin with III spines and 5 ½ soft rays; first anal-fin spine about 50% length of second
spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Scorpaenodes) ® 90
90a. Vertical scale row counts 70 to 75 . . . . . . . . . . . . . . . . . . . . . Scorpaenodes muciparus

90b. Vertical scale row counts 30 to 60 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 91
91a. Pectoral-fin rays 14 to 16, middle rays abruptly longer than those immediately above;
nasal spine absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 92
91b. Pectoral-fin rays 17 to 20, middle rays not abruptly longer than those immediately
above, fin wedge-shaped; nasal spine present, although sometimes reduced . . . . . . . . . ® 93
92a. Dorsal-fin soft rays 9 ½; pectoral-fin rays usually 16; vertical scale rows more than 37
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenodes albaiensis
92b. Dorsal-fin soft rays 8 ½; pectoral-fin rays 15 to 16 (usually 15); vertical scale rows less
than 32 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenodes minor
93a. A pronounced dark spot on subopercle; coronal and interorbital spines almost always
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenodes littoralis
93b. No pronounced dark spot on subopercle; coronal and interorbital spines always or often
absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 94
94a. Vertical scale rows 29 to 35; dorsal-fin rays usually 8 ½ . . . . . . . . . . . . . . . . . . . . . ® 95

94b. Vertical scale rows 38 to 50; dorsal-fin rays usually 9 ½ . . . . . . . . . . . . . . . . . . . . . ® 96
95a. Pectoral-fin rays 17 or 18; usually 4 spines along suborbital ridge, a fifth spine on
second infraorbital bone, below that in main line of suborbital ridge . . . . . Scorpaenodes hirsutus
95b. Pectoral-fin rays 18 to 20, usually 18 or 19; usually 3 spines along suborbital ridge,
without small spine on lateral face of lacrimal bone and without small second spine on
second infraorbital bone ventral to those along main suborbital ridge . . . . Scorpaenodes kelloggi
96a. Five to 15 spines on suborbital ridge; numerous cirri on head and body . Scorpaenodes parvipinnis
96b. Usually 3 spines on suborbital ridge; few or no cirri on head and body . . . . . . . . . . . . . ® 97
97a. A dark spot at rear of spinous portion of dorsal fin; a dark triangular mark at base of
pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenodes varipinnis
97b. No dark spot at rear of spinous portion of dorsal fin; no dark triangular mark at base of
pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 98
98a. No large dark spot over opercle . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenodes smithi

98b. A large dark spot (sometimes diffuse) over opercle . . . . . . . . . . . . . . . . . . . . . . . ® 99
99a. Dark spot on opercle usually pronounced; fourth dorsal-fin spine relatively short, about
3 to 16% standard length; spine at posteroventral margin of lacrimal bone relatively
weak, usually with a cirrus; body moderately blotched, particularly on dorsal and caudal
fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenodes guamensis
99b. Dark spot on opercle usually diffuse; fourth dorsal-fin spine relatively long, about 13 to
20% standard length; spine at posteroventral margin of lacrimal bone relatively strong,
usually without a cirrus; body moderately speckeled . . . . . . . . . . . . . . Scorpaenodes scaber
100a.Palatine teeth absent on roof of mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 101

100b.Palatine teeth present on roof of mouth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 114
101a.Black pigment between dorsal-fin spines I to III (or II and III); fourth dorsal-fin spine
notably elongate relative to other spines in specimens larger than about 6 cm . Iracundus signifer
101b.No black pigment between dorsal-fin spines I to III; fourth dorsal-fin spines not notably
elongate relative to other spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 102
102a.Suborbital ridge without spines or with a single spine at end of ridge, near preopercle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis fowleri
102b.Suborbital ridge with 3 to 5 spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 103
103a.Second dorsal-fin spine 7 to 10% standard length; first dorsal-fin spine 4 to 6% standard
length; head profile notably blunt and with orbit 7 to 10% standard length . . Scorpaenopsis brevifrons
103b.Second dorsal-fin spine 11 to 18% standard length; first dorsal-fin spine 6 to 9%
standard length; head profile relatively pointed or with orbit 12 to 15% standard length . . . . ® 104
104a.Upper opercular spine usually ending in 3 or more points; about 30 to 35 vertical scale
rows; orbit large, 12 to 15% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . ® 105
104b.Upper opercular spine ending in 1 or 2 points; more than 35 vertical scale rows; orbit
moderate 7 to 11% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 106
105a.Predorsal scales 4; predorsal length 39 to 43% standard length; third dorsal-fin spine
12 to 16% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis iop
(Japan; not yet recorded from the area)
105b.Predorsal scales 5 or 6; predorsal length 46 or 47% standard length; third dorsal-fin
spine 17 to 21% standard length . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis cotticeps
106a.Body immediately behind head strongly elevated, giving dorsal profile of back a
strongly humped appearance; mouth superior and strongly upturned; interorbit 10 to
12% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 107
106b.Body immediately behind head elevated, but not strongly so; mouth terminal or superior
but not strongly upturned; interorbit 6 to 8% standard length . . . . . . . . . . . . . . . . . ® 110
107a.Supraorbital spine ending in multiple spinous points . . . . . . . . . . . . . Scorpaenopsis neglecta

107b.Supraorbital spine ending in a single spinous point . . . . . . . . . . . . . . . . . . . . . . . ® 108
108a.Medial surface of pectoral fins with a dark subterminal band along entire margin of fin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis macrochir
108b.Medial surface of pectoral fins with a dark subterminal band incomplete or absent
entirely . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 109
2304 Bony Fishes
109a.Ratio of orbit diameter to snout length 0.39 to 0.53; pectoral-fin rays 17 to 19, but usually
18; black subterminal band on medial (axial) surface of pectoral fins present along
upper 1/2 or less of fin margin (absent in Hawaiian populations); pigment over
proximal 1/2 of upper rays not distinct from that on axial, if present . . . . Scorpaenopsis diabolus
109b.Ratio of orbit diameter to snout length 0.62 to 0.92; pectoral-fin rays 16 to 18, but
usually 17; a black subterminal band on medial (axial) surface of pectoral fins
extending slightly more than 1/2 of fin margin; a distinct large spot over proximal 1/3
of upper rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis gibbosa
(Indian Ocean; not yet recorded from the area)
110a.Pectoral-fin rays 19 or 20; vertical scale rows 60 to 68 . . . . . . . . . . Scorpaenopsis oxycephala

110b.Pectoral-fin rays 15 to 19; vertical scale rows 35 to 61 (Scorpaenopsis cirrhosa
occasionally with more than 60 and rarely 19 pectoral-fin rays) . . . . . . . . . . . . . . . . ® 111
111a.Pectoral-fin rays 16 or 17 (usually 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 112

111b.Pectoral-fin rays 17 to 19 (usually 18 or 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 113
112a.Vertical scale rows about 35 . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis furneauxi

112b.Vertical scale rows 44 to 53 . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis venosa
113a.Vertical scale rows 42 to 52; occipital pit relatively deep; snout relatively pointed
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis papuensis
113b.Vertical scale rows 55 to 61; occipital pit relatively shallow; snout relatively rounded
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaenopsis cirrhosa
114a.Occiput depressed, with at least a shallow pit, a deep pit in some species, never flat
or convex; scales on pectoral-fin base reduced or absent; scales on breast cycloid,
often notably reduced, or absent . . . . . . . . . . . . . . . . . . . . . . . . . (Scorpaena) ® 115
114b.Occiput inclined, but flat or slightly convex, never concave; scales on breast ctenoid
or cycloid, small, but never absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 119
115a.Scales on pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 116

115b.Scales absent on pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 118
116a.Coronal spines present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena papillosus

(New Zealand; not yet recorded from the area)
116b.Coronal spines absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 117
117a.Pectoral-fin rays 16; vertical scale rows about 30; lateral-line scales about 33
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena gibbifrons
117b.Pectoral-fin rays 17 or 18; vertical scale rows 64 to 67; lateral-line scales 24 to 26
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scorpaena cookii
118a.Pectoral-fin rays 17; vertical scale rows 48 to 63 . . . . . . . . . . . . . . . . Scorpaena cardinalis

118b.Pectoral-fin rays 19; vertical scale rows about 43 . . . . . . . . . . . . . . Scorpaena hemilepidota
119a.All pectoral-fin rays simple; no slit behind last hemibranch . . . . . . . . . . Pontinus rhodochrous
119b.At least some rays branched (need to observe carefully); a slit behind last hemibranch
(except N. bauchotae, which does not occur in fishing area) . . . . . . . . . . . . . . . . . ® 120
120a.Dorsal fin-rays 8 ½ to 10 ½, the last split to base; occiput convex . . . . . . (Sebastapistes) ® 121
120b.Dorsal fin-rays 9 ½ to 11 ½, the last split to base; occiput flat or nearly so . . . . . . . . . . ® 126
121a.Coronal spines present; pronounced dark blotch at rear of spinous portion of dorsal fin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sebastapistes mauritiana
121b.Coronal spines absent; no dark blotch at rear of spinous portion of dorsal fin . . . . . . . . . ® 122
122a.Lacrimal bone with 2 spines; a ridge anterior to lower opercular spine . . . . Sebastapistes strongia
122b.Lacrimal bone with 3 to 5 spines; no ridge anterior to lower opercular spine . . . . . . . . . ® 123
123a.Body and fins covered with numerous conspicuous small dark spots . . . . . . . . . . . . . ® 124
123b.Body and fins without numerous dark spots; spots, when present, large and pale . . . . . . ® 125
124a.Lacrimal bone with 4 spines; vertical scale rows 50 to 55 . . . . . . . . . . Sebastapistes coniorta

124b.Lacrimal bone with 5 spines; vertical scale rows 44 to 49 . . . . . . . . . . Sebastapistes tinkhami
125a.Cycloid or emarginate scales on flank; lacrimal bone with 2 spines . . . . Sebastapistes galactacma
125b.Ctenoid scales on flank; lacrimal bone with usually 5 spines . . . . . . . Sebastapistes cyanostigma
126a.Vertebrae 25; posterior lacrimal spine relatively weak, pointing directly ventrally; caudal
peduncle at base of caudal fin probably with scattered large melanophores . . Idiastion pacificum
(Kyushu-Palau Ridge; not yet recorded from the area)
126b.Vertebrae 24; posterior lacrimal spine relatively strong, pointing posteroventrally; caudal
peduncle at base of caudal fin without scattered large melanophores . . . . . . (Neomerinthe) ® 127
127a.Pectoral-fin rays usually 18; 3 spines on suborbital ridge; no spine on lateral face of
lacrimal bone; 5 preopercular spines, with third from dorsalmost longest . . . Neomerinthe rotunda
127b.Pectoral-fin rays usually 19; 4 spines on suborbital ridge; anteriormost on lateral face
of lacrimal bone; 4 preopercular spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 128
128a.Fourth dorsal-fin spine longest; tip of lower jaw protruding just beyond tip of upper jaw
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomerinthe megalepis
128b.Third dorsal-fin spine longest; jaws subequal . . . . . . . . . . . . . . . . . . . . . . . . . ® 129
129a.Suborbital spine ventral to midorbit in line with ridges of succeeding spines . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomerinthe amplisquamiceps
129b.Suborbital spine ventral to midorbit slightly below ridges of succeeding spines . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neomerinthe procurva

The symbol 0 is given when species accounts are included. A question mark indicates that presence in
the area is uncertain.
0 Apistops caloundra de Vis, 1885

Subfamily APISTINAE
0 Apistus carinatus (Bloch and Schneider, 1801)

Cheroscorpaena tridactyla Mees, 1964
0
Subfamily PLECTROGENINAE
Plectrogenium nanum Gilbert, 1905
0 Brachypterois serrulatus (Richardson, 1846)

Subfamily PTEROINAE
0 Dendrochirus bellus (Jordan and Hubbs, 1925)

0 Dendrochirus biocellatus (Fowler, 1938)
0 Dendrochirus brachypterus (Cuvier, 1829)
0 Dendrochirus zebra (Quoy and Gaimard, 1824)
2306 Bony Fishes
? Ebosia bleekeri (Döderlein, 1884)

0 Parapterois heterurus (Bleeker, 1856)
0
0
Pterois antennata (Bloch, 1787)
0
Pterois lunulata Temminck and Schlegel, 1842
0
Pterois miles (Bennett, 1828)
0
Pterois mombasae (Smith, 1957)
0
Pterois radiata Cuvier, 1829
0
Pterois russellii Bennett 1831
Pterois volitans (Linnaeus, 1758)
Subfamily SCORPAENINAE
? Idiastion pacificum Ishida and Amaoka, 1992
0 Iracundus signifer Jordan and Evermann, 1903
0 Neomerinthe amplisquamiceps (Fowler, 1938)
0 Neomerinthe megalepis (Fowler, 1938)
0 Neomerinthe procurva Chen, 1981
0 Neomerinthe rotunda Chen, 1981
0 Parascorpaena mcadamsi (Fowler, 1938)
Parascorpaena aurita (Rüppell, 1838)
0 Parascorpaena mossambica (Peters, 1855)

0 Parascorpaena picta (Kuhl and Van Hasselt in Cuvier 1829)
0 Phenacoscorpius megalops Fowler, 1938
Pontinus rhodochrous (Günther, 1871)
Pteroidichthys amboinensis Bleeker, 1856
Pteroidichthys noronhai (Fowler, 1938)
0 Rhinopias aphanes Eschmeyer, 1973
Rhinopias argoliba Eschmeyer, Hirosaki, and Abe, 1973
0
Rhinopias filamentosa (Fowler, 1938)
Rhinopias frondosa (Günther, 1891)
Rhinopias xenops (Gilbert, 1905)
0
0
Scorpaena cardinalis Richardson, 1842
0
? Scorpaena cookii Günther, 1873
0
Scorpaena gibbifrons Fowler, 1938
Scorpaena hemilepidota Fowler, 1938
0
0
Scorpaenodes albaiensis (Evermann and Seale. 1907)
0
Scorpaenodes hirsutus (Smith, 1957)
0
Scorpaenodes guamensis (Quoy and Gaimard, 1824)
0
Scorpaenodes kelloggi (Jenkins, 1903)
0
Scorpaenodes littoralis (Tanaka, 1917)
Scorpaenodes minor (Smith, 1958)
0
Scorpaenodes muciparus (Alcock, 1889)
0
Scorpaenodes parvipinnis (Garrett, 1864)
Scorpaenodes scaber (Ramsey and Ogilby, 1885)
0
Scorpaenodes smithi Eschmeyer and Rama Rao, 1972
Scorpaenodes varipinnis Smith, 1957
Scorpaenopsella armata Fowler, 19381/
0
0
Scorpaenopsis brevifrons Eschmeyer and Randall, 1975
0
Scorpaenopsis cirrhosa (Thunberg, 1793)
0
Scorpaenopsis cotticeps Fowler, 1938
0
Scorpaenopsis diabolus (Cuvier, 1829)
Scorpaenopsis fowleri (Pietchmann, 1934)
1/ Known from a single specimen taken at Sombero Island, between Balayan Bay and Verde Island Passage in the
Philippines. Not included in the identification key.
Scorpaenopsis furneauxi Whitley, 1959
0
? Scorpaenopsis gibbosa (Bloch and Schneider, 1801)
Scorpaenopsis macrochir Ogilby, 1910
0
Scorpaenopsis neglecta (Heckel, 1840)
Scorpaenopsis oxycephala Bleeker, 1849
0
Scorpaenopsis papuensis (Cuvier, 1829)
Scorpaenopsis venosa (Cuvier, 1829)
0
Sebastapistes coniorta Jenkins, 1903
0
Sebastapistes cyanostigma (Bleeker, 1856)
0
Sebastapistes galactacma Jenkins, 1903
0
Sebastapistes mauritiana (Cuvier, 1829)
0
Sebastapistes strongia (Cuvier, 1829)
Sebastapistes tinkhami (Fowler, 1946)
0 Taenianotus triacanthus Lacepède, 1802
Subfamily NEOSEBASTINAE
Neosebastes incispinnis Ogilby, 1910
Subfamily SEBASTINAE
? Sebastes inermis Cuvier, 18292/
? Sebastes joyneri Günther, 1878 2/
? Sebastes pachycephalus Temminck and Schlegel, 1843 2/
? Sebastiscus albofasciatus Lacepéde, 1801 2/
Sebastiscus marmoratus Cuvier, 1829 2/
0
?
Sebastiscus tertius Barsukov and Chen, 1978
0 Ectreposebastes imus Garman, 1899

Subfamily SETARCHINAE
0 Lioscorpius longiceps Günther, 1880

0 Setarches guentheri Johnson, 1862
0 Setarches longimanus (Alcock, 1894)
0 Choridactylus multibarbus Richardson, 1848
Subfamily SYNANCEIINAE
0 Erosa erosa Langsdorf, 1829

? Erosa daruma (Whitley, 1932)
0
Inimicus brachyrhynchus (Bleeker, 1874)
0
Inimicus caledonicus (Sauvage, 1875)
0
Inimicus cuvieri (Grey, 1835)
Inimicus didactylus (Pallas, 1769)
Inimicus filamentosus (Cuvier, 1829)
Inimicus gruzovi Mandritsa, 1991
Inimicus japonicus (Cuvier, 1829)
0
Inimicus joubini (Chevy, 1927)
Inimicus sinensis (Valenciennes, 1833)
Inimicus smirnovi Mandritsa, 1990
0 Leptosynanceia asteroblepa (Richardson, 1848)
0
Minous coccineus Alcock, 1890
Minous monodactylus (Bloch and Schneider, 1801)
0
Minous pictus Günther, 1880
Minous pusillus Temminck and Schlegel, 1843
0
Minous quincarinatus (Fowler, 1943)
0
Minous trachycephalus (Bleeker, 1854)
Minous versicolor Ogilby, 1910
2/ Reported from the Mariana Islands, but based on records that are unconfirmed and likely erroneous. Not included in
the identification key.
2308 Bony Fishes
0 Synanceia horrida (Linnaeus, 1766)

Synanceia alula Eschmeyer and Rama Rao, 1973
0 Synanceia verrucosa Bloch and Schneider, 1801

0 Trachicephalus uranoscopus (Bloch and Schneider, 1801)
Subfamily TETRAROGINAE
0
Ablabys macracanthus Bleeker, 1857
Ablabys taenianotus (Cuvier, 1829)
0 Centropogon australis (White, 1790)
Centropogon marmoratus Günther, 1862
0 Cottapistus cottoides (Linnaeus, 1758)
0 Liocranium praepositum Ogilby, 1904
0 Neocentropogon trimaculatus Chan, 1965
Neocentropogon affinis (Lloyd, 1909)
0 Notesthes robusta (Günther, 1860)

0 Ocosia apia Poss and Eschmeyer, 1975
0 Ocosia zaspilota Poss and Eschmeyer, 1975
? Ocosia spinosa Chen, 1981
0 Paracentropogon longispinis (Cuvier, 1829)

Paracentropogon vespa Ogilby, 19103/
Paracentropogon zonatus (Weber, 1913) 4/
0 Richardsonichthys leucogaster (Richardson, 1848)
? Snyderina yamanokami Jordan and Starks, 19015/
0
0
Tetraroge barbata (Cuvier, 1829)
Tetraroge niger (Cuvier, 1829)
0
0
Vespicula cypho (Fowler, 1938)
0
Vespicula depressifrons Richardson, 1848
Vespicula trachinoides (Cuvier, 1829)
Vespicula zollingeri (Bleeker, 1848)
References
Chen, L.C. 1981. Scorpaenid fishes of Taiwan. Quart. Journ. Taiwan Mus., 34(1,2):60 p.
Eschmeyer, W.N. 1986. Scorpaenidae. In Smiths' sea fishes, edited by M.M. Smith and P.C. Heemstra. Johannesburg,
MacMillan South Africa, pp. 463-478.
Matsubara, K. 1943. Studies on the scorpaenoid fishes of Japan (II). Trans. Sigenkagaku Kenkyusyo, Tokyo, 2:171-486.
Weber, M. and L.F. de Beaufort (eds). 1962. The fishes of the Indo-Australian Archipelago. XI. Scleroparei,
Hypostomides, Pediculati, Plectognathi, Opisthomi, Discocephali, Xenopterygii. Leiden, E.J. Brill, 481 p.
3/ Not included in the identification key. The differences between this species and Paracentropogon longispinnis are
uncertain.
4/ Known only from 2 syntypes from the Sulu or Postillon Island, Philippines.
5/ Based on a single record from Indonesia that may not be this species.
Ablabys taenianotus (Cuvier, 1829)

En - Cockatoo waspfish.
Maximum standard length 10.7 cm. Among seaweeds in rocky intertidal or littoral habitats to depths
of nearly 80 m. Rocks back and forth in response to surge and can be easily caught with small hand
nets. Of no importance to fisheries, but of interest to the aquarium trade because of its intriguing
appearance. Widely distributed in the western Pacific from Japan in the north, to Australia and Fiji
in the south. Found along the Asian mainland, the Philippines, throughout the Indonesian Archipel-
ago, and westward to the Solomons and Fiji, but does not reach more oceanic islands of the area.
Outside the area, known from India, the Andaman Islands, Taiwan Province of China, Japan, and
Western Australia. Originally described from a specimen said to be taken at Mauritius, but this needs
further confirmation.
Apistops caloundra (de Vis, 1885)

En - Shortspined waspfish.
Maximum standard length 8.8 cm. Taken in muddy and silty areas and near river mouths. Known
to partially bury itself in the soft substrate. Caught in shrimp trawls, but of no commercial importance.
Care should be exercised when removing this species from nets. Known from the Gulf of Carpen-
teria, near Groote Eylandt eastward to Caloundra, Queensland and taken from Jayapura, Irian Jaya.

2310 Bony Fishes
Apistus carinatus (Bloch and Schneider, 1801)

En - Ocellated waspfish; Fr - Rascasse ocellée; Sp - Rascacio ocelado.
Maximum standard length 12.5 cm. Taken over fine sand in depths of 21 to 60 m, particularly in
bays. Quite common in trawls and utilized as a food fish in local markets, despite its venomous
nature. Extreme caution should be exercised in handling this species, as wounds from its fin spines
can be painful. A widespread species, occurring from the mouth of the Umblanga River in South
Africa and the Eilat and Massawa in the Red Sea in the west, to Japan, the Philippines, and Australia
in the east. In the area, reported from the Sunda-Mollucas Archipelago, the Malay Archipelago,
Indonesia, the Philippines (Cavite and Luzon), New Guinea (Merauke), Western Australia (Kimberly
Region), Timor and Arafura seas, and Gulf of Carpentaria, to just south of the area at Ballina, New
South Wales; also known from China and Taiwan Province of China.
Brachypterois serrulatus (Richardson, 1846)

En - Sawcheek scorpionfish.
Maximum standard length about 11.5 cm. A little-known species, occasionally found in trawls and
collected to a depth of 82 m. Of no interest to fisheries, because of its small size and venomous
nature. In the north of the area, known from the western coast of Luzon and Thailand; outside the
area, reported from Japan, Taiwan Province of China, southern China, Western Australia, India,
and the Red Sea.
Centropogon australis (White, 1790)

En - Fortesque.
Maximum standard length at least 8.1 cm. Extremely common in estuaries, particularly in the
reproductive season in the austral Spring. Although of no commercial importance, commonly
entrapped in fishing gear, such as trawls, and can be dangerous to fishermen who attempt to remove
them. Just enters the south of the area, reaching Dunwich, Moreton Bay, in southern Queensland;
found south of the area along the Australian coast to southern New South Wales; reports of this
species in Melanesia are erroneous.
Choridactylus multibarbus Richardson, 1848

En - Threefinger scorpionfish.
Maximum standard length at least 10.3 cm. A highly venomous species, occasionally taken in
coastal waters and tidal inlets. Although consumed only in subsistence fisheries and of little
commercial value, it represents a danger to fishermen in areas where it occurs. Widely distributed,
known from the Red Sea, eastward through the Persian Gulf, Pakistan, and India to the Gulf of
Thailand, China, and the Philippines.
(after Eschmeyer et al., 1979)

2312 Bony Fishes
Cottapistus cottoides (Linnaeus, 1758)

En - Yellow waspfish.
Maximum standard length at least 9.2 cm. Over moderately soft bottoms, from less than a metre to
a depth of 24 m. This species can be common in shrimp trawls. Care must be taken when removing
it from nets as it is highly venomous. Commonly taken in China, Viet Nam, Thailand, Singapore,
Java, Flores, Kei Islands, Borneo, the Dampier Archipelago, New Guinea, northwestern Australia,
the Gulf of Carpentaria, and the east coast of Queensland south to Bowen; not yet reported from
the Philippines, except at Jolo.
(after Weber and de Beaufort, 1962)
Dendrochirus bellus (Jordan and Hubbs, 1925)

En - Butterfly scorpionfish.
Maximum standard length 15 cm. A rare food fish; when taken, it is caught on sandy-mud or gravel
bottoms at depths of 15 to 200 m. Common in the rocky littoral and sublittoral zones of northern
Taiwan Province of China. Also reported from New Caledonia, but better known from Taiwanese
and Japanese waters; likely has a broader distribution than presently reported.
Dendrochirus biocellatus (Fowler, 1938)

En - Twinspot lionfish.
Maximum standard length 12 cm. A little-known species, without importance to fisheries. Found in
rocky littoral and sublittoral areas, which may account for its infrequent capture. Although only
occasionally collected, it has a broad distribution, from South Africa and Mauritius in the west to the
Tuamotu Archipelago in the east; reported from Jolo and the Sulu Islands, near the southern
Philippines, New Caledonia, and as rare in Taiwan Province of China and the Ryukyu Islands.
(from Fowler, 1938)
Dendrochirus brachypterus (Cuvier, 1829)

En - Dwarf lionfish.
Maximum standard length at least 15 cm; possibly to 25 cm. Commonly caught at depths of 32 to
80 m. Crepuscular or nocturnal in habit, remaining inactive by day. Highly venomous, but frequently
used as food in subsistence fisheries. A common and widespread species, ranging from the Red
Sea and South Africa to the Philippines, Samoa, and Hawaii. In the south of the area, likely abundant
throughout all of Indonesia, but becoming less common on the Great Barrier Reef and on the eastern
Australian coast.
(after Smith and Heemstra, 1986)

2314 Bony Fishes
Dendrochirus zebra (Cuvier, 1829)

En - Zebra lionfish.
Maximum standard length about 20 cm. Common in lagoons and outer reef slopes among corals,
as well as rocky littoral and sublittoral areas to depths of about 73 m. Very common in the area, and
ranging widely beyond it; known from South Africa, Madagascar, the Red Sea, India, and Sri Lanka
in the west, to the Philippines, Palau, Guam, Apia, Fiji, and Samoa in the east; commonly reported
in Australia and southern Japan.
(after Masuda et al., 1984 )
Ectreposebastes imus Garman, 1899

En - Black scorpionfish.
Maximum standard length 17 cm. Occurs at considerable depths; taken in trawls and midwater
nets from 150 to 800 m. Although catchable by commercial gear, this small species is not sufficiently
abundant to have potential for fisheries. Within the area, reported only from off Bali, Indonesia, and
off New Caledonia; also known from Taiwan Province of China, Hawaii, and nearly circumglobal in
tropical and subtropical waters of the Atlantic and Pacific Oceans; additional records in the area
can be expected.
Erosa erosa (Langsdorf, 1829)

En - Daruma stinger.
Maximum standard length 14 cm. A common food fish in Japan, but seldom eaten elsewhere, either
because it is infrequently encountered, or perhaps due to its grotesque appearance. Occurs in
depths of 60 to 85 m at Chesterfield Bank, but more frequently seen in shallow waters of southern
Japan, Taiwan Province of China, and Australia. Common in Australia, where it is taken as far south
as New South Wales; also reported from New Caledonia and Chesterfield Bank, but records from
the Indonesian Archipelago do not exist.
(after Allen and Swainston, 1988)
Inimicus caledonicus (Sauvage, 1875)

En - Caledonian stinger.
Maximum standard length 17 cm. This species can be found on sandy bottoms in depths of 15 to
60 m. In the area, reported from New Caledonia and Queensland. Known to range westward to the
Nicobar and Andaman Islands.
(after Day, 1875)
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2316 Bony Fishes
Inimicus cuvieri (Grey, 1835)

En - Longsnout stinger.
Maximum standard length 19.5 cm. Common over sandy or muddy-sand bottoms at depths of 1 to
50 m. Occasionally consumed in subsistence fisheries despite its grotesque appearance and
venomosity. Appears to have a restricted range, from the South China Sea, Gulf of Thailand,
Singapore, Java, Bali, and Sarawak.
(after Eschmeyer, Rama-Rao, and Hallacher, 1979)
Inimicus didactylus (Pallas, 1769)

En - Bearded ghoulfish.
Maximum standard length 14.3 cm. Like other species of the genus, Inimicus didactylus occurs
over sandy and sandy-mud bottoms from inshore to a depth of about 80 m, where it can be
numerous. Used for food throughout its range in subsistence fisheries, but a significant fishery for
species of Inimicus only occurs in Japan, where I. japonicus is cultured as well as caught in the
wild. Study of additional materials may show I. gruzovi and I. smirnovi to be synonyms. Widely
distributed in the area, ranging from the Ryukyu Islands southward through China, Viet Nam,
Thailand, Singapore, Indonesia, New Guinea, Northern Territories and Queensland, the Solomon
Islands, Chesterfield Bank, New Caledonia, and Vanuatu.

Inimicus sinensis (Valenciennes, 1833)

En - Spotted stonefish.
Maximum standard length 15 cm. This dangerous species is frequently taken in trawls over soft,
sandy bottoms. Although likely consumed as food in subsistence fisheries, no significant commer-
cial market exists for it. Known from southern China and Viet Nam, southward through Java and
Bali to Shark Bay, Western Australia; some records are known from the west coast of Luzon; outside
the area, taken in Sri Lanka and the southeast coast of India.
Iracundus signifer Jordan and Evermann, 1903

En - Decoy scorpionfish.
Maximum standard length 11 cm. Found on coral reefs or nearby sandy bottoms to depths of about
10 to 70 m. Moves the dorsal fin to a lure and captures small fishes. Although probably good eating,
no fishery exists for this uncommon species. Known within the area from New Caledonia and
Chesterfield Bank; outside the area, reported from Iriomote-jima in the southern Ryukyus, Taiwan
Province of China, Hawaii, Raratonga, and South Africa. Future collecting will undoubtedly docu-
ment its presence at many Indo-Pacific localities.
(from Jordan and Evermann, 1903)

2318 Bony Fishes
Leptosynanceia asteroblepa (Richardson, 1848)

En - Stareyed stonefish.
Maximum standard length reported as 11.7 cm. Occurs in estuaries and may tolerate fresh water.
Dangerous to fishermen and typically thrown overboard, although edible. Distribution relatively
limited; found off Hong Kong, Viet Nam, Sumatra, Indonesia, Borneo, Sarawak, and New Guinea.
(after Fowler, 1905)
Liocranium praepositum Ogilby, 1904

En - Blackspot waspfish.
Maximum standard length 10.7 cm. Little is known of the biology of this species, which is not
infrequently taken in trawls at depths of about 12 to 65 m. It is highly venomous and can inflict a
painful wound. Found along the northern coast of Australia from Exmouth Gulf to Port Curtis,
Queensland; also known from New Guinea, between Misol and Salawatti.

Lioscorpius longiceps Günther, 1880

En - Slender scorpionfish.
Maximum standard length 13.5 cm. Infrequently taken in trawls at considerable depths (256 to
402 m). Apparently does not occur in large quantities and is not of large enough size to be of
importance to fisheries. Known from southern Japan, Saipan, through the South China Sea and into
the Arafura Sea; also recorded in the Philippines from the vicinity of northern Mindanao and the Kei
Islands.
(from Fowler, 1938)
Minous monodactylus (Bloch and Schneider, 1801)

En - Grey goblinfish.
Maximum standard length 8.6 cm. Frequently taken in trawls, but of no interest to fisheries. Caution
should be exercised when removing this species from nets. Ranges broadly from South Africa and
the Red Sea to New Caledonia; occurs in southern Japan and China southward into the Indonesian
Archipelago; apparently absent in Australia and the Philippines.

2320 Bony Fishes
Minous pusillus Temminck and Schlegel, 1843

En - Dwarf stingfish.
Maximum standard length 5.8 cm; matures at about 4.5 cm total length. Over soft bottoms, taken
at depths of 30 to 110 m. Frequently covered with hydroids, like several other species of Minous.
Found in southern Japan, south to Hong Kong, the Philippines, and New Caledonia.
(from Matsubara, 1943)
Minous trachycephalus (Bleeker, 1854)

En - Striped stingfish.
Maximum standard length 5.5 cm. Over soft bottoms, between depths of 11 and 46 m. Occurs from
Taiwan Province of China southward through Viet Nam and the Philippines into the Indonesian
Archipelago; reported from New Caledonia and from northwestern Australia, just outside the area;
listed also from the Red Sea, but this is likely based on a misidentification of Minous coccineus.

Minous versicolor Ogilby, 1910

En - Plumbstriped stingfish.
Maximum standard length 8.4 cm. Commonly taken in shrimp trawls in northern Australia between
depths of 12 to 64 m. Its sting can be painful and it should be removed from nets with caution. Known
only from the northwestern Australian shelf eastward to Cape Glouscester, Queensland.
Neocentropogon trimaculatus Chan, 1965

En - Threespotted waspfish.
Maximum standard length at least 81 cm, but larger records should be expected. A rare but
dangerous species. Of little importance to fisheries, but may appear in deeper trawls (depth range
203 to 225 m). Known from the South China Sea, off the northwestern Shelf of Australia, at
Chesterfield Bank, and New Caledonia.
(after Chan, 1965)

2322 Bony Fishes
Neomerinthe amplisquamiceps (Fowler, 1938)

En - Orange scorpionfish.
Maximum standard length 15.2 cm. Found at depths of 183 to 294 m. A small species of no
commercial importance. Reported from the Philippines and northwestern Australia. Additional study
may prove it to be the senior synonym of Neomerinthe procurva.
(from Fowler, 1938)
Neomerinthe megalepis (Fowler, 1938)

En - Bigscale scorpionfish.
Maximum standard length 9.9 cm. A little-known species, taken at depths of about 62 to 82 m. Of
no importance to fisheries. In the area, known only from the South China Sea at 21°54’N, and
114°46’E, and from the Philippines; also reported from Taiwan Province of China.
(from Fowler, 1938)

Neomerinthe procurva Chen, 1981

En - Curvedspine scorpionfish.
Maximum standard length at least 14.1 cm. A little-known species; data on its depth range are
unavailable. Evidently, it can be common where it occurs, but probably not in sufficient numbers to
support a significant fishery; possibly taken in subsistence fisheries within the area. Occurs off
Taiwan Province of China and off Indonesia at 09°01’S, 116°18’E. Further study may prove it to be
a junior synonym of Neomerinthe amplisquamiceps.
(after Chen, 1981)
Neomerinthe rotunda Chen, 1981

En - Round scorpionfish.
Maximum standard length 9.4 cm. Biology largely unknown; occurs at depths of 225 to 295 m.
Unlikely to form the basis of an important fishery. At present known only from the western margin
of the area, at Chesterfield Bank and New Caledonia, and outside the area in Taiwan Province of
China.
(after Chen, 1981)

2324 Bony Fishes
Notesthes robusta (Günther, 1860)

En - Bullrout.
Maximum standard length 22.9 cm. Primarily restricted to fresh water and upper estuarine localities.
Can be taken by seines or by hook-and-line. Considerable care should be exercised when handling
this species. Although it can be used for food, it is generally avoided due to its formidable
appearance and because of the extremely venomous spines. Common in the coastal rivers and
estuaries of northeast and southeast coastal Queensland and northern New South Wales to the
Shoalhaven River; also reported from southern Papua New Guinea, but unconfirmed.
Ocosia apia Poss and Eschmeyer, 1975

En - Stoutspine waspfish.
Maximum standard length at least 5.3 cm. Taken in trawls at depths of 225 to 350 m. Of no
commercial importance. Within the area, reported at Chesterfield Bank; previously known from the
Kermadec Islands to the southeast (29°15’S, 177°57’W).
(after Poss and Eschmeyer, 1975)

Ocosia zaspilota Poss and Eschmeyer, 1975

En - Polka dot waspfish.
Maximum standard length 7 cm. This small species has been trawled over moderately muddy
bottoms from depths of 192 to 247 m. Of no commercial importance, but potentially dangerous to
fishermen. To date, known only from the vicinity of Balayan Bay in Luzon, Philippines.
(after Poss and Eschmeyer, 1975)
Paracentropogon longispinis (Cuvier, 1829)

En - Whiteface waspfish.
Maximum standard length 8 cm. A venomous species that should be handled with caution, found
inshore down to nearly 70 m, on and around corals and hard bottoms. Of little importance to
fisheries, but often taken in nets and by hand lines. Some colour patterns are characteristic of a
given region. For example, specimens from the Gulf of Thailand almost always have large blotches
over the body, while those from Indonesia and Australia generally have less solid markings.
Although these various colour patterns may suggest that specimens identified here as Paracentro-
pogon longispinis belong to a species complex, limited information on live coloration exists and
some intermediate colour patterns are observed. Further study may show P. vespa Ogilby, 1910 to
be a synonym. This species has been seen to alter its colour from light to dark in captivity. Known
throughout the western Pacific from Taiwan Province of China and southern China southward
through Indonesia, the Philippines, to New Caledonia; occurs west of the area to southern India.
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2326 Bony Fishes
Parapterois heterurus (Bleeker, 1856)

En - Blackfooted firefish.
Maximum standard length 25 cm. Occasionally appears in trawls over soft bottoms at depths of 40
to 300 m. Too infrequently taken to be of importance to fisheries, but can force trawl fishermen to
handle their catch with greater caution. Reported within the area at Amboina; more commonly taken
in southern Japan and Taiwan Province of China and is also known from South Africa.
Parascorpaena mcadamsi (Fowler, 1938)

En - Ocellated scorpionfish.
Maximum standard length 6 cm. A small relatively uncommon, reef-dwelling or shallow rocky-bot-
tom inhabiting species of minor commercial value, but occasionally seen in subsistence fisheries.
Found from near shore in lagoons to outer reef slopes to depths of 37 m. Widely ranging, like other
species of Parascorpaena, and known from South Africa to the southern Philippines (at Jolo), to
Taiwan Province of China and Queensland, Guadalcanal in the Solomon Islands, Chesterfield
Islands, New Caledonia, and Rapa. Scorpaena moultoni is a probable junior synonym.
(from Fowler, 1938)

Parascorpaena mossambica (Peters, 1855)

En - Mozambique scorpionfish.
Maximum standard length 10 cm. Occupies inshore rocky areas and can be abundant. Although
this species makes its way into local markets and is consumed as food, no significant fisheries exists
for it, due to its small size. Widely distributed, occurring from South Africa and the Red Sea to the
Ryukyu and Palau Islands and eastward to the Society Islands.
Parascorpaena picta (Kuhl and Van Hasselt in Cuvier 1829)

En - Painted scorpionfish.
Maximum standard length 17 cm. Quite common in nearshore waters on reefs and over rocky bottoms.
Utilized as food, but does not form the basis of a significant fishery. Distributed from Sri Lanka to Fiji.

2328 Bony Fishes
Phenacoscorpius megalops Fowler, 1938

En - Noline scorpionfish.
Maximum standard length 5.1 cm. Infrequently trawled at considerable depths (66 to 622 m) over
coralline rubble bottoms. Too small and rarely caught to be of commercial importance. Known from
the Philippines, Indonesia, Taiwan Province of China, Hawaii, and New Zealand.
(from Fowler, 1938)
Plectrogenium nanum Gilbert, 1905

En - Dwarf thronyhead.
Maximum standard length 7.7 cm. Only found at depths of 274 to 600 m, with peak abundance off
Hawaii, occurring at depths of 300 to 450 m. This species does not support a fishery, but can be
very common where it occurs. Currently known from Madagascar, southern Japan, Taiwan Province
of China, New Caledonia, the Kermadec Islands, and New Zealand. Southern populations may
represent a distinct species.
(adopted from Gilbert, 1905)

Pterois antennata (Bloch, 1787)

En - Broadbared lionfish.
Maximum standard length 14 cm. Found in lagoons and on outer reef slope habitats down to depths
of 76 m. Like most lionfishes, living in crevices and holes by day and ventures out over the surface
of reefs at night to forage on shrimps and crabs. Common in the area and consumed as food in
subsistence fisheries, but nowhere supports large scale food fisheries due to its dangerous spines
and small size. Commonly collected for the aquarium trade. A common, wide-ranging species;
reported from off South Africa and the Red Sea eastward to through Indonesia and adjoining parts
of the area to Queensland, New Caledonia, Guam, the Solomon Islands, and onward to southern
Japan, the Kermadec Islands, and the Marquesas and Mangareva.
Pterois lunulata Temminck and Schlegel, 1842

En - Dragon’s beard fish.
Maximum standard length 30 cm. This venomous species is utlized as food and the dorsal-fin
spines are removed by fishermen when marketed. May prove to be conspecific with Pterois
russellii . Additional study of the individual variation in these species, particularly of colour
pattern, is needed. Distributed throughout the western Pacific Ocean from southern Hokkaido
and Korea, southward through China, Taiwan Province of China, Indonesia, northern Australia,
Queensland, and Caledonia.
(after Okada, 1966)

2330 Bony Fishes
Pterois miles (Bennett, 1828)

En - Devil firefish; Fr - Poisson volant.
Maximum standard length 25 cm. Frequently misidentified as Pterois volitans. This species will
direct spines toward an intruder if annoyed. Numerous fatalities have resulted from stings of this
species and its close relative P. volitans, and it should be treated with extreme caution. Used for
human consumption in subsistence fisheries and frequently finds its way into the aquarium trade.
Hardy and easily reared, but dangerous and voracious. Primarily ranges outside the area throughout
the Indian Ocean, but also recorded from the westernmost regions of Indonesia and northwestern
Australia.
Pterois mombasae (Smith, 1957)

En - Deepwater firefish.
Maximum standard length 16 cm. Apparently occurs on deeper reefs. Of no commercial importance
but should be handled cautiously when taken to avoid its highly venomous spines. Ranges from
South Africa eastward into the area in Indonesia and northwestern Australia.
Pterois radiata Cuvier, 1829

En - Radial firefish.
Maximum standard length 10 cm. Common in rock areas and coral reefs. Like other species of
Pterois, it is extremely venomous and should be handled with caution. Of importance as food only
in subsistence fisheries, but forms an important component of the aquarium trade. A wide-ranging
species, taken from South African and the Red Sea eastward to the Tuamotou Archipelago,
northward to Japan and southward to southern Queensland.
Pterois russellii Bennett, 1831

En - Plaintail firefish.
Maximum total length 30 cm. Commonly found on reefs. Despite its venomosity it is sometimes
taken for the aquarium trade. Frequently misidentified as Pterois volitans, but distinguished by the
lack of dark spots on its dorsal, anal, and caudal fin, a lower pectoral-fin ray count (typically 12 or
13), its fewer vertical rows (65 to 80), and its shorter supraocular cirrus. In these respects it
approaches and is perhaps conspecific with P. lunulata. However, variation of these features within
and among lionfishes are poorly understood and are in need of additional study. Occurs throughout
the Indian Ocean and is widespread through all but the easternmost part of the area; reported from
the Philippines and off Queensland.
2332 Bony Fishes
Pterois volitans (Linnaeus, 1758)

En - Lionfish.
Maximum standard length 30 cm. Frequently confused with Pterois miles. A well-known reef
species, frequently taken in trawls at depths of about 10 to 175 m. Largely inactive by day but
forages in deeper water at night, where it evidently reproduces. Its spines are exceedingly
dangerous and numerous fatalities have been reported from wounds resulting from its fin-spines.
Nonetheless, with its fin-spines removed it is used for food in subsistence fisheries. Also extremely
popular in the aquarium trade despite its dangerous reputation. Widespread in the western Pacific
from southern Japan to Australia and the Philippines; it ranges to offshore localities in the Marshall
Islands, New Caledonia, the Kermadec Islands, Fiji, to Tuamotu Archipelago.
(after Matsubara, 1943)
Rhinopias aphanes Eschmeyer, 1973

En - Weedy scorpionfish.
Maximum standard length 24 cm. One of many cryptic and rare scorpionfishes about which little is
known. It is reported from depths of about 30 m, perhaps favouring outer reef slopes. Of no
commercial importance, but of interest to aquarists because of its remarkable appearance. At
present reported only from northeastern Australia, New Caledonia, New Guinea, and southern
Japan, but will probably become known from more widely distributed localities within the area with
further collecting.

Rhinopias frondosa (Günther, 1891)

En - Popeyed scorpionfish; Fr - Poisson scorpion feuillu.
Maximum standard length at least 15.2 cm. Appears to inhabit rock and coralline bottoms at depths
of 13 to 90 m. Little else is known about its biology. Reported from local markets; taken in lobster
gill nets and shrimp trawls, but unlikely to be of significant commercial importance. A widely
distributed species, known from South Africa, Mauritius, and the Seychelles eastward to Sri Lanka,
Viet Nam, Japan, and the Caroline Islands.
Richardsonichthys leucogaster (Richardson, 1848)

En - Whitebellied rougefish.
Maximum standard length 6.6 cm standard length. A little-known, dangerous small species, which
has been reported among corals. Not infrequently taken in trawls at depths of 55 to 90 m. Although
associated with corals and widely reported throughout islands of the Indonesian Archipelago, it
appears more commonly near continental margins. Reported from off Zanzibar, Madagascar, and
the Seychelles eastward to northern Queensland, the Chesterfield Islands, New Caledonia, and the
Philippines.
2334 Bony Fishes
Scorpaena cardinalis Richardson, 1842

En - Northern scorpionfish.
Maximum standard length 38 cm. An inshore species. Good eating and commonly taken by divers
and fishermen over rocky reefs. Diet consists mainly of fishes but also includes crabs, shrimps, and
octopus. Can be aggressive toward divers. Enters into the area in the northern part of its range,
reaching southern Queensland; occurs southward to Tasmania.
(after Ayling and Cox, 1982)
Scorpaena cookii Günther, 1860

En - Sandy-Bay cod.
Maximum standard length 26.4 cm. Fished inshore over rocky reefs and consumed locally for food;
exact depths of capture and statistics on the size of this fishery are unavailable. Limited in its range,
known from the southern part of the area at Raoul Island in the Kermadecs, Lord Howe Island,
Norfolk Island, and at Elizabeth and Middleton Reefs. Reported also from coast of New South Wales
in the vicinity of Coffs Harbor.
(after Paulin, 1982)
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Scorpaena gibbifrons Fowler, 1938

En - Bumphead scorpionfish.
Maximum standard length 6.8 cm. Other than where it is captured, very little is known of this species.
Of no importance to fisheries, but may be encountered while trawling for other species. Known only
from the Philippines in the vicinity of Cabugan Grande Island (10°27’30" N, 125°18’E).
(from Fowler, 1938)
Scorpaena hemilepidota Fowler, 1938

En - Halfscaled scorpionfish.
Maximum standard length at least 15 cm. Nothing is known of the biology of this species other than
it occurs to depths of at least 247 m. Of no importance to fisheries. Only collected from the
Philippines in offshore waters between Samar and Masbate (12°12’35"N, 124°2’48"E).
(from Fowler, 1938)

2336 Bony Fishes
Scorpaenodes albaiensis (Evermann and Seale, 1907)

En - Longfingered scorpionfish.
Maximum standard length 8 cm. Of no importance to fisheries, although it does appear incidentally
in nearshore catches. Caught in depths of about 3 to 8 m. A widely-ranging species, that occurs
from Africa eastward to the Chesterfield Islands, New Caledonia, and the Philippines.
Scorpaenodes guamensis (Quoy and Gaimard, 1824)

En - Guam scorpionfish.
Maximum standard length 12 cm. Common in lagoons and outer reef slope habitats at depths
between 3 and 15 m. Sebastes meleagris may be a junior synonym. A wide-ranging species that
occurs from off Africa and the Red Sea easward to Tonga and the Society Islands. Occurs north to
Japan and south to Capricorn Group at the southern end of the Great Barrier Reef.

Scorpaenodes hirsutus (Smith, 1957)

En - Hairy scorpionfish.
Maximum standard length about 5 cm. A small, little-known species of no commercial importance.
Distributed from East Africa and the Red Sea to the Tuamoto Archipelago; found north to Japan
and south to the Great Barrier Reef.
(after Eschmeyer and Randall, 1975)
Scorpaenodes kelloggi (Jenkins, 1903)

En - Dwarf scorpionfish.
Maximum standard length about 5 cm. Lives on coral reefs in shallow depths. Distributed throughout
the western Central Pacific, including Hawaii, Gilbert Islands, Caroline Islands, Palau, Raiatea, and
Japan and Taiwan Province of China.

2338 Bony Fishes
Scorpaenodes littoralis (Tanaka, 1917)

En - Cheekspot scorpionfish.
Maximum standard length 10 cm. Not consumed as food. Females are gravid in summer. No fishery
exists for this small species. Found in South Africa eastward to Japan, Hawaii, and the southern
end of the Great Barrier Reef, Norfolk, and the Kermadec Islands. Not reported at intermediate
latitudes and may have an antitropical distribution.
Scorpaenodes minor (Smith, 1958)

En - Minor scorpionfish.
Maximum standard length about 5 cm. Virtually nothing is known about this small species. Rarely
seen, except in poison stations on coral reefs and in infrequent trawls over hard bottoms. Ranges
broadly across the African coast of the Indian Ocean to the western Pacific Ocean from the Ryukyus
in the north to the Capricorn Group (Great Barrier Reef) in the south; within the area, reported from
Indonesia, the Marshall Islands, the Philippines, and New Caledonia.

Scorpaenodes parvipinnis (Garrett, 1864)

En - Coral scorpionfish.
Maximum standard length 13 cm; one of the larger species of Scorpaenodes. Known from inshore
reef habitats from depths of 3 to 15 m. A tropical species, known from coral reefs of the Seychelles
and the Red Sea eastward to the Solomon and Chesterfield Islands and New Caledonia. Just north
of the area, it reaches Taiwan Province of China and to the south it is known from the southern end
of the Great Barrier Reef.
Scorpaenodes scaber (Ramsey and Ogilby, 1885)

En - Pygmy scorpionfish.
Maximum standard length 8 cm. Biology poorly known. A small species of no commercial impor-
tance. Occurs on reefs at depths to about 8 to 12 m. Found at dispersed localities throughout the
area from the Society Islands westward to the Indian Ocean; records from the Red Sea can not be
confirmed. Frequently misidentified as Scorpaenodes guamensis or S. littoralis.

2340 Bony Fishes
Scorpaenodes varipinnis Smith, 1957

En - Blotchfin scorpionfish.
Maximum standard length 7 cm. This rare species has been collected to depths of about 15 m in
rocky and coral reef sublittoral areas. Of no importance to fisheries. Distributed from the Seychelles
and the Red Sea eastward to Australia, the Chesterfield Islands, and New Caledonia.
Scorpaenopsis brevifrons Eschmeyer and Randall, 1975)

En - Bigmouth scorpionfish.
Maximum standard length about 11 cm. Taken over coral and rock areas inshore to depths of 35 m.
Reported from South Africa to Hawaii; within the area, known to occur in the Philippine and Tobriand
islands, Fiji, and Samoa.

Scorpaenopsis cirrhosa (Thunberg, 1793)

En - Weedy stingfish; Fr - Poisson arme.
Maximum standard length at least 21 cm. Most specimens of Scorpaenopsis reported throughout
the area have been misidentified as this species. These are often based on specimens correctly
identified as S. oxycephala. Because numerous literature reports that purport to document the
widespread occurrence of this species can not presently be confirmed, this species is included here
even though further research may likely show that it is restricted to Japan, China, and Taiwan
Province of China. S. cirrhosa occurs in coralline intertidal areas to depths of 37 to 91 m. Known
to occur in Japanese waters, on the Chinese mainland, and Taiwan Province of China; unconfirmed
reports within the area are numerous and include New Guinea and scattered Indonesian localities.
Scorpaenopsis cotticeps Fowler, 1938

En - Bulbheaded scorpionfish.
Maximum standard length 4.4 cm. Of no commercial importance. Taken at depths of about 18 m.
Restricted range, presently known only from MacClesfield Bank in the South China Sea, the Sulu
Archipelago, and the southern Philippines. Closely related to Scorpaenopsis iop from Japan. Both
species are in need of further study.
(from Fowler, 1938)

2342 Bony Fishes
Scorpaenopsis diabolus (Cuvier, 1829)

En - False stonefish.
Maximum standard length at least 2.2 cm. Occurs in reef flats and outer reef slope habitats among
coral. Like most scorpaenoids, this species is an ambush predator that seldom moves. However,
when alarmed, this fish reveals bright inner coloured pectoral fins as a warning. Distributed from
East African coast eastward to the Society Islands and Hawaii. Often misidentified as Scorpaenopsis
gibbosa.
Scorpaenopsis fowleri (Pietchmann, 1934)

En - Fowler’s scorpionfish.
Maximum standard length 2.8 cm. A diminutive species of no commercial importance, occuring in
depths of 7 to 38 m in association with Seriatopora corals. It matures at about 2.5 cm standard
length. Known from the Maldive Islands to Mangareva, and the Hawaiian Islands.

Scorpaenopsis macrochir Ogilby, 1910

En - Rough humpbacked scorpionfish.
Maximum standard length 10 cm. Biology little known, but reported to occur at Guam on mixed sand
and coral rubble areas of reef flats and shallow lagoons. Originally described from Queensland, and
reported also from the Moluccas, the Philippines, Tonga, the Marianas, and the Caroline Islands.
Ranges northward to the Ryukyus Islands. Sometimes confused with other humpbacked Scor-
paenopsis: S. diabolus, S. gibbosa, and S. neglecta.
Scorpaenopsis oxycephala Bleeker, 1849

En - Smallscale scorpionfish.
Maximum standard length about 30 cm. An inshore species, common on reefs. Taken as food in
subsistence fisheries. Widely distributed and found from the Red Sea to the Philippines, northward
to the Ryukyu Islands and southward to New Caledonia. Often misidentified as Scorpaenopsis
cirrhosa as are many species of Scorpaenopsis.
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2344 Bony Fishes
Scorpaenopsis venosa (Cuvier, 1829)

En - Raggy scorpionfish; Fr - Lappies.
Maximum standard length at least 18 cm standard length. A little-known species. Occurs inshore
on reefs. Occasionally taken by hook-and-line and in trawls, but of no commercial importance.
Reported from widely scattered localities from southern Africa, the Red Sea, India, Indonesia, and
northwestern Australia.
Sebastapistes cyanostigma (Bleeker, 1856)

En - Yellowspotted scorpionfish.
Maximum standard length about 7 cm. Common in shallow water on coral reefs and hard bottoms
to about 9 m. Of no commercial importance, but occasionally makes its way into the aquarium trade,
because of its small size and coloration. Ranges from South Africa and the Red Sea eastward to
Okinawa, Guadalcanal, and New Caledonia.

Sebastapistes galactacma Jenkins, 1903

En - Galactacma scorpionfish.
Maximum standard length 4.9 cm. Taken over coral and coral rubble in depths of 6 to 29 m. Appears
to be confined to the Pacific Plate, and is known from Guam, Pohnpei, and Rapa within the area
and at Hawaii outside the area.
(from Jordan and Evermann, 1903)
Sebastapistes mauritiana (Cuvier, 1829)

En - Spineblotch scorpionfish; Fr - Rascasse de Suez; Sp - Rascacio de Suez.
Maximum standard length about 8 cm. Typically collected in outer intertidal reef and lagoon habitats
at depths of less than 10 m. Commonly taken, but of no commercial importance. Distributed from
the eastern shores of Africa and the Red Sea eastward to Guam, Marshall Islands, Gilbert Islands,
Phoenix Islands, Rapa, and the Marquesas; also reported to have passed through the Suez Canal
and into the Mediterranean. Further study of the Hawaiian Sebastapistes ballieu (Sauvage, 1875)
may prove it to be a synonym of S. mauritiana or only subspecifically distinct.

2346 Bony Fishes
Sebastapistes strongia (Cuvier, 1829)

En - Barchin scorpionfish.
Maximum standard length about 6 cm. Study of the biology of this small reef-dwelling fish originally
described from the Caroline Islands has been stymied by considerable confusion associated with
its taxonomic identity. Further analysis may prove specimens of this variable species may perhaps
belong to 2 species, which have been thoroughly confused under a variety of inconsistently used
names. Often confused with Scorpaena bynoensis Richardson, 1845, which is a synonym of
Parascorpaena picta. Of no commercial importance, but often caught at shallow depths over coral
reefs. A wide-ranging species, commonly taken from East Africa and the Red Sea, eastward to Sri
Lanka and Taiwan Province of China; occurs throughout the Indonesian Archipelago, Melanesia,
Micronesia, and the Philippines to the Tuamoto Archipelago; it has traversed the Suez Canal and
was reported from Cyprus.
Sebastapistes tinkhami (Fowler, 1946)

En - Darkspotted scorpionfish.
Maximum standard length about 8 cm. A small relatively rare species that appears to occur on
deeper reefs. Of no importance to fisheries, but occasionally appears in trawls. Care should be
taken when handling it. Widely distributed. Known from South Africa eastward to the Tuamotu
Archipelago, and also reported in the area from Fiji.

Sebastiscus tertius Barsukov and Chen, 1978

En - Absent-minded scorpionfish.
Maximum standard length 37 cm. Very closely related to Sebastiscus marmoratus and likely has
similar habits, but occurs in deeper waters at depths of 70 to 500 m. A relatively large species and
would make an excellent food fish as does its near relatives. Although presently no major fishery
exists for this species, discovery of suitable hard-bottom habitats over which it may occur in large
numbers, could make a sustainable fishery possible. Best known from southern Japan, Korea, and
Taiwan Province of China, but also reported in the area from Indonesia (08°58’S, 116°34’E).
Setarches guentheri Johnson, 1862

En - Deepwater scorpionfish; Fr - Rascasse serran; Sp - Rascacio serrano.
Maximum standard length 18 cm. No commercial fishery exists for this deep-dwelling scorpionfish,
although it occurs in large numbers at some localities. Taken with bottom trawls and bottom-set
long lines at depths of 170 to 576 m. Wounds from this species can be painful. The most
widely-ranging scorpaenoid fish, found worldwide on continental slopes in tropical and warm-tem-
perate latitudes, as well as insular localities. In the area, reported from off Sumatra, Bali, Ujung
Padang, the Arafura Sea, Fiji, the Philippines, and near the Kei Islands.
2348 Bony Fishes
Setarches longimanus (Alcock, 1894)

En - Redsmooth scorpionfish.
Maximum standard length 18 cm. Reported from trawls fished at depths of 110 to 763 m, with most
specimens taken at intermediate depths between 180 and 550 m. The spines of this species are
venomous and should be avoided. Distributed from Andaman Islands in the Eastern Indian Ocean
to the Philippines, north to southern Japan and south to northern Australia and New Caledonia.
Synanceia horrida (Linnaeus, 1766)

En - Estuarine stonefish.
Maximum standard length 19.6 cm; may weigh up to 2 kg. Most frequently found in lagoons and
reef flats, but can also be encountered in relatively muddy or rubble bottoms associated with
estuaries. Used as food in subsistence fisheries and found in local markets, but extremely venomous
and therefore of limited value. Capable of inflicting fatal wounds and should be handled, if at all,
with extraordinary caution. Found from the eastern Indian Ocean eastward through the Indonesian
Archipelago to Australia and the Philippines.
(after Eschmeyer and Rama-Rao, 1973)

Synanceia verrucosa Bloch and Schneider, 1801

En - Reef stonefish; Fr - Poisson-pierre.
Maximum standard length 18.7 cm. Usually seen at low tide in coral rubble in pools on shallow reef
flats and in lagoons, often under rocks and ledges. Can be abundant at some localities, reaching
depths of about 13 m. Larger individuals appear to favour surge channels. Also capable of burying
in sand. Feeds on Abudefduf leucozona, A. biocellatus, A. glaucus, Pomacentrus littoralis, and
other small fishes and invertebrates. Because it is remarkably well camouflaged and exceedingly
venomous, this species should be fished and handled only with extreme caution; its spines can
inflict fatal wounds. Although typically feared, it is occasionally fished by spear and utilized for human
consumption. A widely distributed species, known from East Africa and the Red Sea eastward to
the Tuamoto Archipelago.
Taenianotus triacanthus Lacepède, 1802

En - Leaf scorpionfish; Fr - Poisson balance.
Maximum standard length 7.9 cm. Typically inhabits coral beds and rocks in sublittoral and intertidal
habitats in areas of strong wave action, to depths of about 78 to 134 m. Individuals will “sway” with
the current, mimicking its surroundings, even in the absence of current. T. triacanthus molts, at
times, shedding its skin in a nearly single piece like some snakes. A variety of strikingly different
colour morphs exist, with yellow-brown and red and pink morphs being the most common. It is
unknown to what extent individuals can change colours. A poor swimmer and easily caught with
hand nets. Of no commercial importance, but has become a frequent but voracious aquarium fish,
due to its unusual appearance and interesting behaviour. Distributed from East Africa eastward to
the Galapagos Islands, where it has been photographed but not yet collected; also known from
southern Japan in the north to the southern end of the Great Barrier Reef in the south.

2350 Bony Fishes
Tetraroge barbata (Cuvier, 1829)

En - Bearded rougefish.
Maximum standard length 9.3 cm. Found in brackish water and lower fresh water reaches of shallow
slow-flowing coastal drainages. Likely mimics leaf-litter. No fishery exists for this dangerous
species. Distributed from Japan southward through Taiwan Province of China and the Philippines,
throughout Indonesia from Southern Malaysia to the Solomon Islands; not known to occur in
Australia or Fiji.
(after Munro, 1967)
Tetraroge niger (Cuvier, 1829)

En - Blacksea wasp.
Maximum standard length 10 cm. A little-known, highly venomous species; enters brackish and
fresh water often considerable distances (16 to 42 km) from the coast. Females can be extremely
fecund bearing tens of thousands of eggs. Found from India eastward, throughout the Indonesia
Archipelago to Fiji; also occurs in the Philippines and northward to Taiwan Province of China and
the southernmost Ryukyu Islands.

Trachicephalus uranoscopus (Bloch and Schneider, 1801)

En - Stargazing stonefish.
Maximum standard length 8 cm. Generally caught in inshore trawls over mud bottoms in estuaries.
Despite its venomosity, this small species does appear in local market catches. Occurs from India
eastward through the Strait of Mallaca and northward along the coasts of Malaysia and Thailand
northward to Hong Kong and southern China; also reported westward to Java, Borneo, and
Amboina.
(after Eschmeyer and Rama-Rao, 1973)
Vespicula cypho (Fowler, 1938)

En - Hunchbacked goblinfish.
Maximum standard length 3.3 cm. A rare venomous species that may become entrapped in nets,
but is of no commercial importance. Known only from a single specimen taken in the Philippines at
Davao, Mindanao.
(from Fowler, 1938)

2352 Bony Fishes
Vespicula depressifrons Richardson, 1848

En - Leaf goblinfish.
Maximum standard length 6 cm. A little-known species, typically found in brackish or semi-fresh
waters. No fishery exists for this species, but it does appear from time to time in the aquarium trade.
Found in the Nicobar Islands through the Indonesia Archipelago and the Philippines westward to
Papua New Guinea, reports of this species from Japan are erroneous.
Vespicula trachinoides (Cuvier, 1829)

En - Goblinfish.
Maximum standard length 5.8 cm. Appears to be found over soft sand and broken shell bottoms.
This small species is of little commercial importance, but does seem to be marketed fresh in local
fisheries throughout the area. Distributed from the Mergui Archipelago (Myanmar) eastward to the
Philippines and Sulawesi; known outside the area from Hainan Island, China; a single record from
the Red Sea (under the synonym Apistus bottae) is likely based on a locality error.
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Scorpaeniformes: Caracanthidae 2353
CARACANTHIDAE
Orbicular velvetfishes (coral crouchers)
by S.G Poss
D iagnostic characters: Small fishes (typically under 4 cm standard length); body rounded, com-
pressed, although not greatly so. Head moderate to large, 37 to 49% of standard length. Eyes small
to moderate, 8 to 12% of standard length. Snout 8 to 13% of standard length. Mouth moderate to large,
upper jaw 12 to 21% of standard length. Numerous small conical teeth present on upper and lower jaws,
none on vomer or palatines. Lacrimal movable, with 2 spines, posteriormost largest directed ven-
trally. All species with a narrow extension of the third infrarobital bone (second suborbital)
extending backward and downward across the cheek and usually firmly bound to preopercle. No
postorbital bones. Branchiostegal rays 7. Skin over gill covers strongly fused to isthmus. Dorsal fin
with VII or VIII (rarely VI) short spines and 12 to 14 branched soft rays. Anal fin usually with II short spines,
followed by 11 or 12 branched soft rays. Caudal fin rounded, never forked. Pelvic fins small, difficult to see,
with I stout spine and 2 or 3 (rarely 1) rays. Pectoral fins with 14 or 15 thickened rays. Scales absent,
except for lateral line, but body densely covered with tubercles. Lateral-line scales present; usually
11 to 19 tubed scales. All species possess extrinsic striated swimbladder musculature. Vertebrae 24.
Colour: orbicular velvetfish are either pale pinkish white with numerous small black spots, or light brown
or greenish and with orange spots or reticulations.
Habitat, biology, and fisheries: Velvetfishes live within the branches of Acropora, Poecillopora, and
Stylophora corals, rarely venturing far from the coral head. Other than their close association with corals,
little is known of their biology.
Gobid fishes of the genus Gobiodon also occur among coral heads and are easily mistaken for caracan-
thids. However, these fishes lack the suborbital stay that extends back to the preopercle, the large
moveable lacrimal spines, and the large preopercular spines.
Key to the species of Caracanthidae occurring in the area
1a. No deep notch between spinous and soft-rayed parts of dorsal fin, fin membrane
relatively continuous; relatively uniformly light brown or grey, without distinct spots or
mottling; lacrimal with 1 large spine and 2 small blunt spines . . . . . . . . . Caracanthus unipinna
1b. A deep notch between spinous and soft-rayed parts of dorsal fin; body with either spots
or with reticulate pattern dorsally; lacrimal with 1 long spine and 1 small blunt knob . . . . . . . ® 2
2a. Body covered with numerous black spots . . . . . . . . . . . . . . . . . . Caracanthus maculatus
2b. Body with reticulate pattern over dorsum . . . . . . . . . . . . . . . Caracanthus madagascariensis
(Indian Ocean; not yet recorded from the area)
Caracanthus maculatus (Gray, 1831)
Caracanthus unipinna (Gray, 1831)
Reference
Smith, J.L.B. 1958. Fishes of the families Tetrarogidae, Caracanthidae, and Synanceiidae from the Western Indian
Ocean with further notes on scorpaenid fishes. Ichthyol. Bull. Rhodes Univ., 12:167-181.
2354 Bony Fishes
APLOACTINIDAE
Velvetfishes
by S.G. Poss
D iagnostic characters: Small fishes (typically under 5 cm standard length); body usually compressed,
although usually not greatly so. Head moderate to large, 26 to 42% of standard length. Eyes small to
relatively large, 5 to 11% of standard length. Snout often prominent, 6 to 14% of standard length. Mouth
moderate to large, upper jaw 7 to 18% of standard length. Numerous small conical teeth present on upper
and lower jaws, with those on vomer usually present and those on palatine always absent. Branchiostegal
rays typically 7 (rarely 6). Gill rakers usually small or moderate, 0 to 6 on upper limb of first gill arch, and
3 to 11 on lower limb (including raker at angle between upper and lower limb). Most species with a fleshy
papillae at end of isthmus, but with skin over gill covers fused to isthmus in some species. Pseudobranch
with 0 to 18 filaments. No branched rays in fins. Dorsal fin with IX to XVI spines, usually blunt at
their tip, and 7 to 16 soft rays. Anal fin usually with 0 to 5 usually weak or blunt spines, followed by 5 to
16 soft rays. Caudal fin moderate to large (21 to 38% standard length) and rounded, never forked. Pectoral
fins usually large, with 9 to 16 rays. Pelvic fins with I strong spine and 2 or 3 (rarely 1) soft rays. Scales
in most species irregular and forming highly modified tack-like structures (some without scales,
except for lateral line). Lateral-line scales present, with usually 9 to 16 tubed scales. All species with an
extension of the third infraorbital bone (second suborbital) extending backward across the cheek and
usually firmly bound to preopercle. Most species with numerous, usually blunt head spines, with those
on movable lacrimal, ususally most prominent. All species possess striated swimbladder musculature
that is extrinsic in nearly all species, with musculature present even in those without swimbladders. Pyloric
caecae 1 to 16. Vertebrae 21 to 33. Colour: most species strongly camouflaged and brown, reddish brown,
or greenish; usually with barred or mottled colour patterns, typically darker dorsally than ventrally.
Habitat, biology, and fisheries: Velvetfishes live in or on the bottom, where they are often found under
or between rocks or vegetation. Although most are taken at shallow depths, a few species in the area range
into deeper waters (to 510 m). Most species are extremely well camouflaged. They are of no importance
to fisheries.

Scorpaenidae: have a suborbital stay but scales, if present, do not form spinous points (except in
Taenianotus); dorsal fin orginates before eye in waspfishes (tetrarogine scorpaenids), but in these species
all spines are sharp and there is no fleshy extension on the anteriormost part of the isthmus; pelvic fins
with I spine and 3 to 5 segmented rays (I spine and 2 or 3 segmented rays in Aploactinidae); caudal-fin
rays always branched (except in Coccotropsis and Minous); other fins, except pectoral fins, usually with
branched rays (all segmented rays unbranched in Aploactinidae); eyes on top of head or lower pectoral-fin
rays detached in some species (never true of Aploactinidae).
Scorpaeniformes: Aploactinidae 2355
Key to the species of Aploactinidae occurring in the area

Note: in the anticipation that they may be recorded in the future from the area, some species known from
adjoining seas are included in the key, with their distribution indicated in parenthesis.
1a. Gill membranes not broadly united to isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Gill membranes broadly united to isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 26
2a. No fleshy papillae at anterior end of isthmus . . . . . . . . . . . . . . . . . . . . . Eschmeyer nexus

(incertae sedis)
2b. A fleshy papillae at anterior end of isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Frontals, parietals, and infraorbital bones heavily armored; with a wide depression in
interorbit; pelvic-fin insertion well behind pectoral-fin base . . . . . . . . . . Peristrominous dolosus
3b. Frontals, parietals, and infraorbital bones not heavily armored; depression in interorbit
narrow or non-existent; pelvic-fin insertion anterior or just behind pectoral-fin base . . . . . . . . ® 4
4a. Anus just behind pelvic fins, far forward of anal-fin origin . . . . . . . . . . . . Prosoproctus pataecus
4b. Anus slightly before anal-fin origin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Three dorsal fins, scales on body form expanded spinous points; each scale forms a
blade-like ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Neaploactis tridorsalis
5b. One or 2 dorsal fins, scales on body absent or form spinous points that are not expanded
or blade-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Anteriormost 3 dorsal-fin spines form small fin that originates over or immediately behind
eye and is widely separated from remainder of fin . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6b. Dorsal fin continuous or nearly so, without small, separate fin over eye . . . . . . . . . . . . . ® 10
7a. Body smooth, covered with numerous elongate cirri, especially anteriorly; lacrimal weakly
ossified, without spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sthenopus mollis
7b. Body covered with numerous scales that form spinous points; lacrimal with 2 strong sharp
spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Xenaploactis)
8a. Interorbit with ridges nearly parallel; body depth less than 1/3 of standard length; dorsal
fin with III spines, followed by IX spines and 8 or 9 soft rays (based on limited material;
some variation to be expected) . . . . . . . . . . . . . . . . . . . . . . . . . . . Xenaploactis cautes
8b. Interorbit with ridges divergent anteriorly, convergent over middle of orbit, divergent
posteriorly; body depth equal to or greater than 1/3 of standard length; dorsal fin with III
spines, followed by X spines and 8 or 9 soft rays (based on limited material; some variation
to be expected) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Second infraorbital bone with 1 or 2 spines; pore of infraorbital lateral-line canal at second
infraorbital bone a simple obscure pore; body depth greater than 1/3 of standard length;
no finger-like cirri above uppermost preopercular lateral-line pores; dorsal fin with III
spines, followed by X spines and 9 soft rays; anal fin with I spine and 10 soft rays (based
on limited material; some variation to be expected) . . . . . . . . . . . . . . Xenaploactis asperrima
9b. Second infraorbital bone without spines; pore of infraorbital lateral-line canal at second
infraorbital bone a prominent elongate slit; body depth equal to 1/3 of standard length;
finger-like cirri present above uppermost preopercular lateral-line pores; dorsal fin with III
spines, followed by X spines and 8 soft rays; anal fin with I spine and 9 soft rays (based
on limited material; some variation to be expected) . . . . . . . . . . . . . . . . Xenaploactis anopta
10a. Pelvic fins with I spine and 3 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11

10b. Pelvic fins with I spine and 2 (1) soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
2356 Bony Fishes
11a. Interorbital ridges run in parallel or converge posteriorly . . . . . . . . . . . . . . (Cocotropus) ® 12

11b. Interorbital ridges diverge strongly posteriorly . . . . . . . . . . . . . . . . . . . (Paraploactis) ® 16
12a. Total dorsal-fin elements 24 (i.e. XIII spines and 11 soft rays); pectoral-fin rays 11 (based
on limited material; some variation to be expected); prickles on body greatly reduced and
widely spaced . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cocotropus echinatus
12b. Total dorsal-fin elements less than 24 (i.e. XII to XIII spines plus 8 to 10 soft rays);
pectoral-fin rays 12 to 14; prickles on body relatively large, densely covering head and
body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Upper jaw relatively short, 10 to 11% of standard length; a stubby fleshy cirrus near distal
end of maxilla; anal fin with I spine and 8 or 9 soft rays . . . . . . . . . . . . . Cocotropus altipinnis
(Lord Howe Island, Kermadec Islands; not yet recorded from the area)
13b. Upper jaw relatively long, 12 to 14% of standard length; no stubby fleshy cirrus near distal
end of maxilla; anal fin with II spines and 7 to 9 soft rays . . . . . . . . . . . . . . . . . . . . . ® 14
14a. First dorsal-fin spine relatively short, 20 to 21% of standard length; second dorsal-fin spine
relatively short, 19 to 20% standard length . . . . . . . . . . . . . . . . . Cocotropus dermacanthus
14b. First dorsal-fin spine relatively long, 22 to 31% of standard length; second dorsal-fin spine
relatively short, 23 to 26% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
15a. Dorsal fin with XIII spines and 9 or 10 soft rays; anal fin with II spines and 8 or 9 soft rays;
snout in lateral profile notably concave; numerous short dark streaks along body, particu-
larly between lateral-line scales . . . . . . . . . . . . . . . . . . . . . . . . . . Cocotropus larvatus
15b. Dorsal fin with XII spines and 9 soft rays; anal fin with II spines and 7 soft rays; snout in
lateral profile notably convex; few, if any, dark streaks on body . . . . . . . . . . Cocotropus masudai
(Hatizyo Island, Izu Islands; not yet recorded from the area)
16a. Ventral surface of lower jaw smooth, not covered with prickles or papillose villi; mandibular
cirri almost confined to outer margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
16b. Ventral surface of lower jaw covered with numerous prickles or papillose villi and with cirri
on both inner and outer margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
17a. Minute cirri on snout, those in larger patch on tip of snout across lacrimal bone about equal
in size to those in smaller patch on nasal bone; fleshy pad on isthmus with its width about
1/4 to 1/3 of its length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraploactis trachyderma
17b. Large, fleshy, and extensively branched cirri on snout; those in larger patches on tip of
snout and across lacrimal bone large, those in smaller patch on nasal bone smaller; fleshy
bad on isthmus with its width about 1/2 to 3/4 of its length . . . . . . . . . . . . Paraploactis pulvinus
18a. No fleshy pad on isthmus . . . . . . . . . . . . . . . . . . . . . . . . . Paraploactis kagoshimensis

18b. Fleshy pad present on isthmus, its width about 1/4 to 1/3 of its length . . . . . . Paraploactis intonsa
19a. Dorsal fin originates over dorsal ramus of preopercle . . . . . . . . . . . . . . . Adventor elongatus
19b. Dorsal fin originates over or before eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
20a. Lacrimal with 2 sharp spines, second large and extends down over maxilla; preopercle
with 4 sharp spines; pelvic fins in front of pectoral-fin base . . . . . . . . . . . . . . (Erisphex) ® 24
20b. Lacrimal spines, if present, blunt, never sharp; preopercle with 4 or 5 blunt spines; pelvic
fins just under or just behind pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 21
21a. Head 28 to 38% of standard length; body elongate; dorsal fin with XIII or XIV spines and
11 to 15 soft rays; anal fin usually with I spine and 11 to 14 soft rays . . . . . . . . . Aploactis aspera
21b. Head 35 to 40% of standard length; body oblong or gibbous; dorsal fin with XII or XIII
spines and 8 to 10 soft rays; anal fin with I or II spines and 8 or 9 soft rays . . . . (Kanekonia) ® 22
Scorpaeniformes: Aploactinidae 2357
22a. Body gibbous; greatest body depth 37 to 40% standard length; body covered with highly
modified scales each of which forms a small spinous point; interorbital ridges thick and
fused in middle of interorbit; dorsal margin of operculum inclined about 35° above
horizontal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kanekonia queenslandica
22b. Body oblong; greatest body depth 29 to 33% standard length; body without modified
scales, except for a few at base of pectoral fins, on cheek, and sometimes on caudal
peduncle; interorbital ridges thin and do not fuse in middle of interorbit; dorsal margin of
operculum inclined about 15° above horizontal . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
23a. Head bones not strongly sculptured; dorsal profile of head sloping at about 40° to 45° from
horizontal, not notably rounded; interorbital ridges converge and just meet posteriorly;
space between ridges not expanded posteriorly; interorbital bones usually with only fine
striations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kanekonia florida
(Japan; not yet recorded from the area)
23b. Head bones, especially infraorbital series, strongly sculptured; dorsal profile of head
notably rounded; interorbital ridges converge posteriorly but do not meet, notably sculp-
tured; space between ridges narrow but expanded anteriorly and posteriorly . . . . . Kanekonia pelta
24a. Dorsal-fin rays 14 to 16; anal-fin rays 13 to 15; vertebrae 30 to 31 . . . . . . . . Erisphex philippinus
24b. Dorsal-fin rays 9 to 13; anal-fin rays 9 to 11; vertebrae 27 to 29 (rarely 30) . . . . . . . . . . . . ®25
25a. Head and body densely covered with modified scales, the largest of which forms a point
that is about 3 to 6 times as long as wide at base; knob on second infraorbital bone almost
always forms 2 blunt spnes, one above the other . . . . . . . . . . . . . . . . . . . . Erisphex potti
(China, Korea, Japan; not yet recorded from the area)
25b. Head and body covered with modified scales, the largest of which forms a point that is
about 1 to 2 times as long as wide at base; knob on second infraorbital bone almost always
forms a single spine (blunt in larger specimens) . . . . . . . . . . . . . . . . . . . . Erisphex aniarus
26a. Body somewhat elongate, not compressed; dorsal fin with II spines, followed by VI spines and
6 soft rays; pelvic fin with I spine and 1 soft ray; vertebrae 21 . . . . . . . . . Matsubarichthys inusitatus
26b. Body notably compressed; dorsal fin with XI to XV spines and 7 to 9 soft rays . . . . . . . . . . ® 27
27a. Anal fin with I or II spines and 6 to 8 soft rays . . . . . . . . . . . . . . . . . Acanthosphex leurynnis
27b. Anal fin with III or IV spines and 5 to 9 soft rays . . . . . . . . . . . . . . . . (Bathyaploactis) ® 28
28a. Anal fin with III spines and 8 or 9 soft rays; pectoral-fin rays 10; 2 pairs of barbels on lower
jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bathyaploactis curtisensis
28b. Anal fin with IV spines (rarely V) and 5 to 7 soft rays; pectoral-fin rays 11 or 12 (rarely 10);
lower jaw without barbels or with a single pair of barbels . . . . . . . . . . Bathyaploactis ornatissima

A question mark indicates that presence in the area is uncertain.
Acanthosphex leurynnis (Jordan and Seale, 1905)
Adventor elongatus (Whitley, 1952)
Aploactis aspera (Richardson, 1844)
Bathyaploactis curtisensis Whitley, 1933
Bathyaploactis ornatissima Whitley 1933
? Cocotropus altipinnis Waite, 1903
Cocotropus dermacanthus (Bleeker, 1852)
Cocotropus echinatus (Cantor, 1850)
Cocotropus larvatus Poss and Allen, 1987
? Cocotropus masudai Matsubara, 1943
2358 Bony Fishes
Erisphex aniarus (Thompson, 1967) 1/

Erisphex philippinus (Fowler, 1938)
? Erisphex pottii (Steindachner, 1896)
Erisphex simplex Chen, 1981
Eschmeyer nexus Poss and Springer, 1983 (insertae sedis)
? Kanekonia florida Tanaka, 1915
Kanekonia pelta Poss, 1982
Kanekonia queenslandica Whitley, 1952
Matsubarichthys inusitatus Poss and Johnson, 1991
Neaploactis tridorsalis Eschmeyer and Allen, 1978
Paraploactis hongkongiensis (Chan, 1966)
Paraploactis intonsa Poss and Eschmeyer, 1978
Paraploactis kagoshimensis (Ishikawa, 1904)2/
Paraploactis obbesi (Weber, 1913)
Paraploactis pulvinus Poss and Eschmeyer, 1978
Paraploactis trachyderma Bleeker, 1865
Peristrominous dolosus Whitley, 1952
Prosoproctus pataecus Poss and Eschmeyer, 1979
Sthenopus mollis Richardson, 1848
Xenaploactis anopta Poss and Eschmeyer, 1980
Xenaploactis asperrima (Günther, 1860)
Xenaploactis cautes Poss and Eschmeyer, 1980
Reference
Poss, S.G. and W.N. Eschmeyer. 1978. Two new Australian velvetfishes, genus Paraploactis (Scorpaeniformes:
Aploactinidae), with a revision of the genus and comments on the genera and species of the Aploactinidae. Proc.
Calif. Acad. Sci., 41(18):401-426.
1/ May be a complex of species including Erisphex simplex (Taiwan Province of China).

2/ May be a complex of species including Paraploactis obbesi and P. hongkongiensis.
click for next page

Scorpaeniformes: Triglidae 2359
TRIGLIDAE
Gurnards, sea robins (also, armoured gurnards, armoured sea robins)
by W.J. Richards
D iagnostic characters (subfamily Triglinae): Body elongate. Head large, bony, with a pair of rostral
projections (rostral process), with spines but no scales or skin covering musculature on head.
Very small barbels present on lower jaw in a few species. Teeth present on upper and lower jaws. Two
separate dorsal fins, first spinous; with either bony plates at spinous base or plates and spines along bases
of both dorsal fins; single anal fin; pectoral fins large, with 3 free lower rays; pelvic fins with I spine and
5 soft rays. Trunk and tail covered with scales of varying size; lateral line distinct, composed of
tube-like scales to enlarged bony scales. Colour: brownish, but often red or reddish when caught; body
often with dark spots, blotches or saddles; first dorsal fin often with black or red blotch; pectoral fins dark
but often with bright blue spots on black background and often with greenish or bluish margins.
scales on body and tail

a pair of rostral
projections
(rostral process)
very small barbels,

or barbels absent 3 free lower
pectoral-fin rays subfamily Triglinae
Diagnostic characters (subfamily Peristediinae): Body elongate. Head large, bony, with rostral projec-
tions, with spines but no scales or skin covering musculature on head. Large barbels on lower jaw.
Teeth present or absent on both jaws. Two separate dorsal fins, first spinous; single anal fin; pectoral fins
with 2 free lower rays; pelvic fins with I spine and 5 soft rays. Trunk and tail completely covered with
large bony scutes. Colour: red or reddish; some with dark spots or markings on body; pectoral fins
generally dark without bright colours.
a pair of rostral
projections bony scutes, no scales
subfamily Peristediinae
always large barbels 2 free lower
pectoral-fin rays
Habitat, biology, and fisheries: Members of the subfamily Triglinae are benthic species with several of
commercial importance inhabiting continental shelves and insular areas from shallow water to 500 m. The
Peristediinae comprise deep benthic species of limited commercial value inhabiting warm tropical seas of
all oceans along continental shelf edge and slope and insular areas from 200 to 500 m.
Remarks: A recent study (Inamura, 1996) provides strong evidence that the Triglidae and Peristediidae
should be recognized as separate families based principally on pelvic bone differences.
2360 Bony Fishes

Dactylopteridae: also with large bony head, but lack
rostral projections and have no barbels; bony
scutes always absent; pectoral fins much enlarged
and brightly coloured, lacking free lower soft rays.
Hoplichthyidae
Hoplichthyidae: also have (3 or 4) free lower pecto-
ral-fin rays, but are distinguished by having the
large bony head extremely depressed, lacking ros-
tral projections, and scales or bony scutes absent.
Bembridae: head also bony, but depressed; no free
lower pectoral-fin rays.
Platycephalidae: head also bony, but depressed; no
free lower pectoral-fin rays.
Bembridae
Platycephalidae
Dactylopteridae
Key to the genera of Triglidae occurring in the area

Note: the genus Pterygotrigla includes many species, several of which are undescribed. It is provisionally
divided into subgenera which can be separated as follows:
1. Subgenus Pterygotrigla - nasal spine absent, opercular spine long, nuchal spine long, antrorse
rostral spine absent, cleithral spine long and strong. Includes P. andertoni and P. pauli.
2. Subgenus Otohime - opercular spine long and slender, extending behind cleithrum, nasal spine
absent, antrorse rostral spine absent, cleithral spine small if present (often absent). Includes P.
hemisticta and P. tagala.
3. Subgenus Bovitrigla - nasal spine absent, opercular spine short, cleithral spine long and strong,
rostral spine long, antrorse rostral spine absent. Includes P. acanthomlopate.
4. Subgenus Parapterygotrigla - nasal spine present or absent, nuchal and cleithral spines long and
strong, opercular spine small, rostral spines long in young and strong in large adults, antrorse rostral
spine present in a single species (P. multiocellata). This subgenus may be divided into additional
subgenera or species groups. Provisionally includes P. hoplites, P. macrorhynchus, P. megalops,
P. multiocellata, and P. ryukyuensis.
1a. Pectoral fins with 3 free lower soft rays; body and tail covered with scales; no large
barbels on lower jaw (Fig.1) . . . . . . . . . . . . . . . . . . . . . . . . . (subfamily Triglinae) ® 2
1b. Pectoral fins with 2 free lower soft rays; body and tail covered with large bony scutes;
large barbels present on lower jaw (Fig. 2) . . . . . . . . . . . . . . . (subfamily Peristediinae) ® 5
Fig. 1 Triglinae Fig. 2 Peristediinae

2a. Bony plates with sharp spines posteriorly along entire length of both dorsal fins (Fig. 3a) . . . . . ® 3
2b. Bony plates only along base of first dorsal fin (Fig. 3b) . . . . . . . . . . . . . . . . . . . . . . ® 4
large, smooth plates
small, spiny plates along bases restricted to base of
of both dorsal fins spiny dorsal fin
a) Lepidotrigla, Chelidonichthys b) other genera
Fig. 3 head and anterior part of body (dorsal view)

3a. Trunk and tail scales large, usually less than 60 rows along lateral line; head usually with
a deep occipital groove (Fig. 4a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lepidotrigla
3b. Trunk and tail scales small, usually more than 60 rows along lateral line; head lacking
deep occipital groove (Fig. 4b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelidonichthys
deep occipital groove no deep occipital

groove
a) Lepidotrigla b) Chelidonichthys
Fig. 4 head and anterior part of body (lateral view)

4a. Scales small, more than 60 rows along lateral line . . . . . . . . . . . . . . . . . . . . Pterygotrigla
(see note at beginning of identification key)
4b. Scales large, less than 60 rows along lateral line . . . . . . . . . . . . . . . . . . . . . . . . Uradia
5a. Upper jaw with villiform teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6

5b. Upper jaw lacking teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
6a. Second dorsal fin with 14 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gargariscus

6b. Second dorsal fin with 20 or more soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Rostral process short and triangular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heminodus

7b. Rostral process long and spatulate . . . . . . . . . . . . . . . . . . . . . . . . . . . Paraheminodus
8a. Preopercular spine long well visible in dorsal view . . . . . . . . . . . . . . . . . . . . . Satyrichthys

8b. Preopercular spine small, not visible in dorsal view . . . . . . . . . . . . . . . . . . . . Peristedion

0 Chelidonichthys kumu (Cuvier, 1829)

Subfamily TRIGLINAE
0 Chelidonichthys spinosus (McClelland, 1844)

Lepidotrigla abyssalis Jordan and Starks, 1904
Lepidotrigla alata (Houttuyn, 1782)
Lepidotrigla annamarae Del Cerro and Lloris, 1997
Lepidotrigla argyrosoma Fowler, 1938
Lepidotrigla deasoni Herre and Kauffman, 1952
2362 Bony Fishes
Lepidotrigla eydouxi Sauvage, 1878

Lepidotrigla japonica (Bleeker, 1857)
Lepidotrigla jimjoebob Richards, 1992
Lepidotrigla kanagashira Kamohara, 1936
Lepidotrigla lepidojugulata Li, 1981
Lepidotrigla longimana Li, 1981
Lepidotrigla macrobrachia Fowler, 1938
Lepidotrigla marisinensis Fowler, 1938
Lepidotrigla musorstom Del Cerro and Lloris, 1997
Lepidotrigla nana Del Cerro and Lloris, 1997
Lepidotrigla oglina Fowler, 1938
Lepidotrigla pectoralis Fowler, 1938
Lepidotrigla punctipectoralis Fowler, 1938
0
Lepidotrigla sereti Del Cerro and Lloris, 1997
Lepidotrigla spiloptera Günther, 1880
Lepidotrigla venusta Fowler, 1938
Lepidotrigla spp. (possibly 20 undescribed forms and in need of revision)
0
0
Pterygotrigla acanthomoplate (Fowler, 1938)
0
Pterygotrigla andertoni Waite, 1910
0
Pterygotrigla hemisticta (Temminck and Schlegel, 1843)
0
Pterygotrigla hoplites (Fowler, 1938)
0
Pterygotrigla leptacanthus (Günther, 1880)
0
Pterygotrigla macrorhynchus (Kamohara, 1936)
0
Pterygotrigla megalops (Fowler, 1938)
Pterygotrigla multiocellata (Matsubara, 1937)
Pterygotrigla pauli Hardy, 1982
0
Pterygotrigla robertsi Del Cerro and Lloris, 1997
0
Pterygotrigla ryukyuensis Matsubara and Hiyama, 1932
Pterygotrigla tagala (Herre and Kauffman, 1952)
Pterygotrigla n. sp. (close to Pterygotrigla tagala)
Pterygotrigla n. sp. (close to Pterygotrigla hemisticta)
Pterygotrigla n. sp. (about 2 undescribed species)
Pterygotrigla n. sp. (several undescribed species)
0 Uradia macrolepidota Kamohara, 1938
0 Gargariscus prionocephalus (Dumeril, 1868)

Subfamily PERISTEDEDIINAE
0 Heminodus philippinus Smith, 1917

Heminodus japonicus Kamohara, 1952
0 Paraheminodus murrayi (Günther, 1880)

Peristedion amblygenys Fowler, 1938
Peristedion halyi (Alcock, 1899)
0
Persitedion investigatoris (Alcock, 1898)
Peristedion liorhynchus Günther, 1871
Peristedion moluccense Bleeker, 1850
Peristedion nierstraszi Weber, 1913
Peristedion orientale Temminck and Schlegel, 1843
Peristedion riversandersoni (Alcock, 1899)
Satyrichthys adeni (Lloyd, 1907)
Satyrichthys amiscus (Jordan and Starks, 1904)
Satyrichthys clavilapis Fowler, 1938
Satyrichthys engyceros (Günther, 1871)
Satyrichthys hians (Gilbert and Cramer, 1897)
Satyrichthys isokawae Yatou and Okamura, 1985
Satyrichthys magnus Yatou, 1985
Satyrichthys orientale (Fowler, 1938)
Satyrichthys piercei Fowler, 1938
0 Satyrichthys rieffeli (Kaup, 1859)

Satyrichthys quadratorostratus (Fourmanoir and Rivaton, 1979)
Satyrichthys serrulatus (Alcock, 1898)

Satyrichthys welchi (Herre, 1925)
References
Chen, M.-H and K.-T. Shao. 1988. Fish of Triglidae (Scorpaenoidei) from Taiwan. J. Taiwan Mus., 41(1):127-138.
Del Cerro, L. and D. Lloris. 1997. Gurnard fishes (Scorpaeniformes, Triglidae) from off New Caledonia, with description
of five new species. Résultats des Campagnes MUSORSTOM, Volume 17. Mém. Mus. natn. Hist., 174:91-124.
Fowler, H.W. 1938. Descriptions of new fishes obtained by the United States Bureau of Fisheries Steamer “Albatross”,
chiefly in Philippine Seas and adjacent waters. Proc. U. S. Natl. Mus., 85(3032):31-135.
Gloerfelt-Tarp, T. and P.J. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Australia . Australian
Development Assistance Bureau, Australia; Directorate General of Fisheries, Indonesia; German Agency for
Technical Cooperation, FRG, 406 p.
Inamura, H. 1996. Phylogeny of the family Platycephalidae and related taxa (Pisces: Scorpaeniformes). Species
Diversity, 1:123-233.
Matsubara, K. and Y. Hiyama. 1932. A review of Triglidae, a family of mail-cheeked fishes, found in the waters around
Japan. J. Imp. fish. Inst., Tokyo., 28:3-67.
Miller, G.C. 1974. Fische des Indischen Ozeans. A. Systematischer Teil, XIV. Scorpaeniformes (2) Family Peristediidae.
Meteor Forsch.-Ergebnisse, D(18):61-72.
Ochia, A. and T. Yatou. 1984. Family Triglidae. In The fishes of the Japanese Archipelago , edited by H. Masuda, K.
Amaoka, C. Araga, T. Ueyeno, and T. Yoshino. Tokyo, Tokai Univ. Press, pp. 333-334.
Paxton, J.R., D.F. Hoese, G.R. Allen, and J.E. Hanley. 1989. Pisces. Petromyzontidae to Carangidae. Zoological
catalogue of Australia, (7):591p.
Richards, W.J. 1984. Triglidae. In FAO species identification sheets for fishery purposes. Western Indian Ocean
Richards, W.J. 1992. Comments on the genus Lepidotrigla (Pisces: Triglidae) with descriptions of 2 new species from
the Indian and Pacific Oceans. Bull. Mar. Sci., 51:45-65.
Richards, W.J. and V.P. Saksena. 1974. Fische des Indischen Ozeans. A. Systematischer Teil, XIV. Scorpaeniformes
(1) Family Triglidae. Meteor Forsch.-Ergebnisse., D(18):55-60.
Richards, W.J. and V.P. Saksena. 1977. Systematics of the gurnards, genus Lepidotrigla (Pisces, Triglidae) from the
Indian Ocean. Bull. Mar. Sci., 27:208-222.
Sainsbury, K.J., P.J. Kailola, and G.G. Leyland. 1985. Continental shelf fishes of northern and north-western Australia.
Canberra, Australia, Clouston and Hall and Peter Pownall Fisheries Information Service, 375 p.
Shen, S.-C. 1984. Coastal fishes of Taiwan. Taipei, National Taiwan Univ., 190 p.
2364 Bony Fishes
Chelidonichthys kumu (Cuvier, 1829)

Frequent synonyms / misidentifications: Trigla kumu Lesson, 1830 / Chelidonichthys spinosus
(McClelland, 1844).
FAO names: En - Bluefin gurnard; Fr - Grondin aile bleue; Sp - Testolín de aleta azul.
(after Ochiai and Okada, 1966)
Diagnostic characters: Head large, triangular, with many ridges and spines, but without a fissure (occipital
groove) on top of head. Eye diameter greater than interorbital width. First gill arch with 8 or 9 gill rakers.
Soft dorsal- and anal-fin rays 14 to 17; bases of first and second dorsal fins with small plates bearing strong
spines (often called bucklers and resulting from outgrowths of the pterygiophores). Body scales small, 70
to 80 along lateral line; breast scaleless. Colour: olive or brownish, becoming red when stressed; lower
half of inner part of pectoral fins with large black blotch surrounded by numerous pale spots (bluish
in life, white or pale in preserved specimens).
Size: Maximum standard length 40 cm.
Habitat, biology, and fisheries: A tropical and subtropical species. Found from estuaries to the edge of
continental shelves generally over sandy bottoms. Separate statistics are generally not reported for this
species except for Australia and New Zealand. Taken mainly by trawls off southern Africa, Australia, and
New Zealand. Reported to be an excellent food fish. Dorsal-fin spines reported to be venomous.
Distribution: Known from South Africa and southern Mozambique, southern Australia, and New Zealand.
One record from Hawaii and unconfirmed records from Chile. Because of Hawaiian record the Chilean
record could be authentic.
Remarks: A thorough study
of the taxonomy of this spe-
cies is called for throughout its
range together with Cheli-
donichthys spinosus t o r e-
solve identification questions.
Chelidonichthys spinosus (McClelland, 1844)

Frequent synonyms / misidentifications: Trigla spinosa McClelland, 1844 / Chelidonichthys kumu
(Lesson, 1830).
FAO names: En - Red gurnard.
(after Ochiai and Okada, 1966)
Diagnostic characters: Head large, triangular, with many ridges and spines, but without a fissure (occipital
groove) on top of head. Eye diameter greater than interorbital width. First gill arch with 8 or 9 gill rakers.
Soft dorsal- and anal-fin rays 15 to 17; bases of first and second dorsal fins with small plates bearing strong
spines (often called bucklers and resulting from outgrowths of the pterygiophores). Body scales small, 70
to 80 along lateral line; breast scaleless. Colour: olive or brownish, becoming red when stressed; lower
half of inner part of pectoral fins with scattered numerous pale spots (bluish in life, white or pale
in preserved specimens); black blotch appears occasionally on lower inner part of pectoral fins.
Habitat, biology, and fisheries: A tropical to warm-temperate species. Taken mainly in trawls throughout
its range. Excellent food fish.
Distribution: Found in the
China Sea, Yellow Sea, and
Japan (principally west of
Kyushu).
Remarks: A thorough study
of the taxonomy of this spe-
cies is needed throughout its
range together with Cheli-
donichthys kumu to resolve
identification questions.
2366 Bony Fishes
Gargariscus prionocephalus Dumeril, 1868

Frequent synonyms / misidentifications: Peristedion undulatus W e b e r , 1 9 1 3 ; Gargariscus
semidentatus Smith, 1917 / None.
FAO names: En - Jaggedhead armoured gurnard.
Diagnostic characters: Head large, expanded, and flattened with crenulations. Rostral process very
broad and short. Preopercular spine sharp and strong. Filamentous barbel short, extending below eye.
Teeth on both jaws. Colour: orange reddish in life with black bands in pectoral fins and black edge on
dorsal fins.
Habitat, biology, and fisheries: A deep-water species. No present fishery, but taken incidentally
throughout its range. Bony scutes reduce commercial value.
Distribution: Only known
from the Philippines.
Heminodus philippinus Smith, 1917

Frequent synonyms / misidentifications: None / Other species of the subfamily Peristediinae.
FAO names: En - Philippine armoured gurnard.
Diagnostic characters: Head large. Rostral process short. Filamentous barbel short. Teeth present
on both jaws. Colour: preserved specimens pale, reddish in life.
from the Philippines.
2368 Bony Fishes
Lepidotrigla spiloptera (Günther, 1880)

Frequent synonyms / misidentifications: None / Chelidonichthys kumu (Lesson, 1830).
FAO names: En - Spotwing gurnard; Fr - Grondin aile tachetée; Sp - Cabete aleta manchada.
prominent spines
rostral processes
(dorsal view)
(after Richards and Saksena, 1977)
Diagnostic characters: Head large, triangular, with many ridges and spines, and a fissure on top of head,
behind eyes (occipital groove). Rostral process with several prominent spines. Bases of first and
second dorsal fins with many small plates bearing strong lateral spines. Large lateral-line scales, fewer
than 70 rows; scale rows below lateral line 19 to 21; breast lacking scales; body scale attachment weak.
Colour: mostly red with definite silvery white breast, belly, and lower flank.
Size: Maximum standard length less than 11 cm.
Habitat, biology, and fisheries: A tropical to warm-temperate species. Taken mainly in trawls throughout
its range. Excellent food fish.
Distribution: Found in the
China Sea, Yellow Sea, and
Japan (principally west of
Kyushu). Also found off East
Africa and northern Australia.
Remarks: A s w i t h Cheli-
donichthys kumu and C. spi-
nosus, a thorough taxonomic
study is needed throughout
the range of Lepidotrigla spi-
loptera.
Paraheminodus murrayi (Günther, 1880)

Frequent synonyms / misidentifications: Satyrichthys murrayi (Günther, 1880) / Satyrichthys spp.
FAO names: En - Murray’s armoured gurnard.
(after Günther, 1887)
Diagnostic characters: Head large, broadly expanded laterally. Rostral process long and flattened.
Preopercular spine long. Filamentous barbel long, extending to first dorsal-fin origin. Teeth on both
jaws. Dorsal-fin spines VII; second dorsal-fin rays 20 to 22; anal-fin rays 20 to 22; pectoral-fin rays 14 or
15. Colour: reddish in life, preserved specimens pale; edge of dorsal fins black.
Distribution: Japan, Banda
Sea, New Caledonia, and off
Horn of Africa.
2370 Bony Fishes
Peristedion liorhynchus Günther, 1871

FAO names: En - Smoothnose armoured gurnard.
(after Okamura, 1985)
Diagnostic characters: Head large. Filamentous barbel moderately long, 12% of standard length.
Dorsal-fin spines VIII; second dorsal-fin rays 21; anal-fin rays 20; attached pectoral-fin rays 11. Colour:
mostly brownish red with dark margin on dorsal fins and banded pectoral fins.
Distribution: Western Pacific
and Indian Oceans.
Pterygotrigla acanthomoplate (Fowler, 1938)

Frequent synonyms / misidentifications: Bovitrigla acanthomoplate (Fowler, 1938) / None.
FAO names: En - Bullheaded gurnard.
Diagnostic characters: Head large, triangular. Nasal spine present, opercular spine short, nuchal spine
long, rostral spines moderately long, antrorse rostral spine absent, and cleithral spine long. Base of first
dorsal fin with about 10 flat plates, first plate extends in advance of first dorsal-fin spine. Lateral-line
scales small, about 50 rows. Colour: preserved specimens uniformly tan with no black spots on dorsum.
Habitat, biology, and fisheries: A rare deep-water species. Incidentally caught by deep trawling or
hook-and-line.
from northern Mindanao in the
Philippines.
2372 Bony Fishes
Pterygotrigla andertoni Waite, 1910

Frequent synonyms / misidentifications: None / Pterygotrigla picta (Günther, 1880).
FAO names: En - Spotted gurnard.
Diagnostic characters: Head large, triangular. No spines around orbit; nasal spine absent, opercular
spine long, nuchal spine long, rostral spines strong but short, no antrorse rostral spine, cleithral spine long.
Base of first dorsal fin with 10 flat plates, first plate extends in advance of first dorsal-fin spine. Lateral-line
scales small, more than 70 rows. Colour: mostly red with prominent black spots; head and dorsum
covered with prominent black spots.
Habitat, biology, and fisheries: A tropical to warm-temperate species in generally deep water of 200 to
500 m, but shallower in New Zealand (90 m). Taken mainly in trawls off New Zealand and Australia.
Excellent food fish but catches are rare.
Distribution: Occurs around
both North Island and South
Island of New Zealand, south-
eastern Australia, and north to
New Caledonia (New Caledo-
nia record from Del Cerro and
Lloris, 1997, specimens not
seen).
Remarks: The closely related
Pterygotrigla picta occurs
around islands and guyots off
the coast of Chile and was
considered to be the senior
synonym of P. andertoni until
recent revisionary studies.
Pterygotrigla hemisticta (Temminck and Schlegel, 1843)

Frequent synonyms / misidentifications: Otohime hemisticta (Temminck and Schlegel, 1843); Trigla
arabica Boulenger, 1887; Prionotus alepis Alcock, 1889 / Pterygotrigla picta (Günther, 1880).
FAO names: En - Blackspotted gurnard; Fr - Grondin encré; Sp - Cabete tintero.
Diagnostic characters: Head large, triangular, with many ridges and spines. Opercular spine long and
slender, extending posteriad of cleithrum; nasal spine and antrorse rostral spine absent; cleithral spine
small; rostral spines strong but short. Base of first dorsal fin with 10 flat plates, first plate extends in advance
of first dorsal-fin spine. Pectoral fins with 12 connected rays. Lateral-line scales small, more than 70 rows.
Colour: mostly red with prominent black spots and large black spot in first dorsal fin; head and dorsum
covered with prominent black spots; inner surface of pectoral fins with large dark area and diagonal
band of separate white spots.
Habitat, biology, and fisheries: A wide depth range of 10 to 420 m. No present fishery, but taken
incidentally throughout its range. Taken mainly in trawls. Excellent food fish but catches are rare.
southwards to the China Sea,
Philippines, and northern
Australia.
Remarks: A closely related
species reported from the
western Indian Ocean, Ptery-
gotrigla arabica Boulenger,
is distinguished from P. hem-
isticta by the white markings
on the inner side of the pecto-
ral fins, and other characters.
2374 Bony Fishes
Pterygotrigla hoplites (Fowler, 1938)

Frequent synonyms / misidentifications: Dixiphichthys hoplites Fowler, 1938 / None.
FAO names: En - Swordspine gunard.
long, rostral spines very long, antrorse rostral spine absent, and cleithral spine long. Base of first dorsal
fins with about 7 flat plates, first plate not extending in advance of first dorsal-fin spine. Pectoral fins
long, more than 50% of standard length. Nape, breast, and belly scaly. Lateral-line scales small, more
than 70 rows. Colour: preserved specimens uniformly tan; no spots on dorsum.
Size: Maximum standard length 10.7 cm.
hook-and-line.
Distribution: Philippines and
northwestern Australia.
Pterygotrigla leptacanthus (Günther, 1880)

Frequent synonyms / misidentifications: Trigla leptacanthus Günther, 1880; Bovitrigla leptacanthus
(Günther, 1880) / None.
FAO names: En - Spotted bullheaded gurnard.
long, rostral spines moderately long, antrorse rostral spine absent, and cleithral spine long. Base of first
dorsal fin with about 10 flat plates, first plate extends in advance of first dorsal-fin spine. Pectoral fins
moderate sized, 38 to 41% of standard length. Lateral-line scales small, more than 70 rows. Colour:
preserved specimens uniformly tan with black spots on dorsum.
ies: A rare deep-water spe-
cies. Incidentally caught by
deep trawling or hook-and-
line.
Australia, Arafura Sea.
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2376 Bony Fishes
Pterygotrigla macrorhynchus Kamohara, 1936

Frequent synonyms / misidentifications: Parapterygotyrigla macrorhynchus (Kamohara, 1936);
Dixiphistes macrorhynchus Fowler, 1938; Dixiphichthys ferculum Whitley, 1952 / None.
FAO names: En - Longnose gurnard.
Diagnostic characters: Head large, triangular. Snout long, 17 to 22% of standard length. Nasal spine
present, opercular spine short, nuchal spine long, rostral spines very long, antrorse rostral spine absent,
and cleithral spine long. Base of first dorsal fin with 10 flat plates, first plate extends in advance of first
dorsal-fin spine. Lateral-line scales small, more than 70 rows; breast and nape scaleless. Colour:
preserved specimens with 3 dark bands on pectoral fins; no spots on dorsum.
hook-and-line.
Distribution: Japan south-
ward to the Philippines; also in
western and eastern Australia
and western Indian Ocean
around Madagascar, Mozam-
bique Channel, and Andaman
Sea.
Pterygotrigla megalops (Fowler, 1938)

Frequent synonyms / misidentifications: Parapterygotyrigla megalops (Fowler, 1938); Dixiphistops
megalops Fowler, 1938 / None.
FAO names: En - Bigeye gurnard.
long, rostral spines very long, antrorse rostral spine absent, cleithral spine long. Base of first dorsal fin with
10 flat plates, first plate extends in advance of first dorsal-fin spine. Pectoral fins short, 30 to 35% of
standard length. Nape, breast, and belly scaly. Lateral-line scales small, more than 70 rows. Colour:
preserved specimens uniformly tan; no spots on dorsum.
hook-and-line.
Distribution: P h i l i p p i n e s
and Saipan.
2378 Bony Fishes
Pterygotrigla multiocellata (Matsubara, 1937)

Frequent synonyms / misidentifications: Parapterygotyrigla multiocellata Matsubara, 1937 / None.
FAO names: En - Antrorse spined gurnard.
(after Kamohara, 1952)
Diagnostic characters: Head large, triangular. No spines around orbit; nasal spine present, opercular
spine short, nuchal spine long, rostral spines strong but short, antrorse rostral spine present, and cleithral
spine long. Base of first dorsal fin with 10 flat plates, first plate extends in advance of first dorsal-fin spine.
Lateral-line scales small, more than 70 rows. Colour: mostly red without black spots.
hook-and-line.
ward to Taiwan Province of
China, eastward to Saipan.
Pterygotrigla ryukyuensis Matsubara and Hiyama, 1932)

FAO names: En - Ryukyu gurnard.
(after Matsubara and Hiyama, 1932)
Diagnostic characters: Head large, triangular. Nasal spine absent, opercular spine short, nuchal spine
long, rostral spines very long, antrorse rostral spine absent, and cleithral spine long. Base of first dorsal
fin with about 10 flat plates, first plate extends in advance of first dorsal-fin spine. Nape scaleless,
breast, and belly scaly. Lateral-line scales small, more than 70 rows. Colour: preserved specimens
uniformly tan; no spots on dorsum.
hook-and-line. Some specimens exhibit extreme hyperostosis of the head and dorsal-fin spines.
ward to Taiwan Province of
China, and northwestern Aus-
tralia.
2380 Bony Fishes
Pterygotrigla tagala (Herre and Kaufman, 1952)

Frequent synonyms / misidentifications: Pterygotrigla spinosa Asano and Okamura, 1963 /
Pterygotrigla hemisticta (Temminck and Schlegel, 1843).
FAO names: En - Tagala gurnard.
(after Asano and Okamura, 1963)
Diagnostic characters: Head large, triangular, with many ridges and spines. Opercular spine long and
slender, extending behind cleithrum, nasal spine absent, antrorse rostral spine absent, cleithral spine
very reduced or absent, with base barely visible. Rostral spines strong but short. Base of first dorsal fin
with 10 flat plates, first plate extends in advance of first dorsal-fin spine. Lateral-line scales small, more
than 70 rows. Pectoral fins with 13 connected rays. Colour: probably reddish in life as are congeners;
preserved specimens brownish; dorsum with a few dark spots; pectoral fins with black area bordered
with white or light spots, remainder dark.
Habitat, biology, and fisheries: Taken in depths of 99 to 119 m. No present fishery, but taken incidentally
throughout its range.
Distribution: Known only
from Manila Bay (Philip-
pines).
Satyrichthys riefflei (Kaup, 1859)

FAO names: En - Spotted armoured gurnard.
(after Okamura, 1985)
Diagnostic characters: Head large, with long preopercular spine and long flattened rostral process.
Filamentous barbel moderately long, 15% of standard length. Dorsal-fin spines VII; second dorsal-fin rays
17. Anal-fin rays 17. Colour: mostly reddish with dark spots on head and dorsum.
and Indian oceans.
2382 Bony Fishes
Uradia macrolepidota (Kamohara, 1938)

Frequent synonyms / misidentifications: Pterygotrigla macrolepidota (Kamohara, 1938) / None.
FAO names: En - Largescaled gurnard.
(after Kamohara, 1938)
Diagnostic characters: Head large, triangular, with large orbit. Rostral spines strong but short. Opercular
spine short, nasal spine absent, antrorse rostral spine absent, and cleithral spine moderate. Base of first
dorsal fin with 10 flat plates, first plate extends in advance of first dorsal-fin spine. Pectoral fins with
12 connected rays. Large lateral-line scales, about 60 rows. Colour: mostly reddish head and dorsum;
flanks and belly white.
throughout its range as bycatch.
Distribution: T o s a B a y ,
Japan and recent records
from New Caledonia (Del
Cerro and Lloris, ms;
specimens not seen).
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Scorpaeniformes: Bembridae 2383
BEMBRIDAE
Deepwater flatheads
by S.G. Poss
D iagnostic characters: Small bottom fishes (size to nearly 30 cm standard length), with large and
depressed heads and subcylindrical bodies. Body without large scute-like spines. Lower jaw
superior or slightly subterminal. Eye moderately large, about 24 to 42% of head length. No iris lapit or
ocular flaps. Infraorbital bones forming a strong ridge, confluent with uppermost preopercular spine
and bearing 4 to 6 spines. Spines on head short or moderate, 8 to 11 above eye. Dorsal fin separated
into a spinous and soft-rayed part. None to III spines present in anal fin. Pectoral fins without free rays.
Lateral-line scales about 28 to 85. Scales on body otherwise ctenoid. Colour: usually red, orange, or light
brown; some species with green, dark green, or black spots or blotches on body and fins; anal fins
sometimes black or nearly so.
Habitat, biology, and fisheries: Bottom fishes living on the continental shelf at depths of 80 to 581 m.
The family limits are not well understood and have been variously regarded as including from 5 to 13
species in 6 genera, with the most recent review (not including Brachybembras) restricting the family to
include only species of the genus Bembras, with the monotypic Parabembras curtus being placed in a
closely related family Parabembridae and species of Bembradium being placed in a family Plectrogeniidae.

Platycephalidae: first dorsal-fin spine relatively short, not broadly attached to the second dorsal fin-spine
(long and broadly attached in Bembridae); pelvic fins inserted behind pectoral-fin insertion (immediately
ventral to pectoral-fin insertion in Bembridae).
Hoplichthyidae: head extremely compressed with numerous minute head spines (head less compressed
with fewer somewhat larger spines in Bembridae); spiny scutes along lateral line (absent in Bembridae);
pectoral fins with 3 or 4 detached lower rays (no detached rays in Bembridae).
Platycephalidae Hoplichthyidae
Key to the species of Bembridae occurring in the area

1a. Anal fin with III spines; lower jaws extended anterior to upper jaws; maxillae relatively
narrow posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parabembras curtus
1b. Anal fin without spines; lower jaws terminal, not extending anterior to upper jaws;
maxillae relatively broad posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2384 Bony Fishes
2a. Lateral-line scales 28 to 32; pectoral-fin rays 21 to 30; head larger, 38 to 46% of standard
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Lateral-line scales 52 to 56; pectoral-fin rays 16 to 22; head smaller, 34 to 38% of
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Dorsal fin with VIII spines and 12 soft rays; lateral-line scales 32; pectoral-fin rays 21;
5 suborbital spines on ridge of second and third infraorbital bones . . Brachybembras aschemeieri
3b. Dorsal fin with IX spines and 11 or 12 soft rays; lateral-line scales 28 to 29; pectoral-fin
rays 25 to 30; 9 to 20 suborbital spines on ridge of second and third infraorbital bones
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bembradium furici
4a. Broad subterminal vertical band in caudal fin; 54 to 62 scale rows above lateral line
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bembras macrolepis
4b. Several narrow vertical bands in caudal fin or with a large dark spot, more densely
pigmented in ventral lobe of fin; 77 to 85 scale rows above lateral line . . . . . . . . . . . . . . ® 5
5a. Anal-fin rays 15; first gill arch with 3 or 4 gill rakers on upper limb . . . . . . Bembras megacephala
5b. Anal-fin rays usually 13 or 14 (rarely 15); first gill arch with 1 or 2 gill rakers on upper
limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Pectoral fins with 16 to 18 rays (usually 17); ratio of pectoral-fin length to caudal-fin
length 0.88 to 1.05 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bembras japonica
6b. Pectoral fins with 17 to 19 rays (usually 18); ratio of pectoral-fin length to caudal-fin
length 1.05 to 1.15 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bembras longispinnis

Bembradium furici Fourmanoir and Rivaton, 1979
Bembras japonica Cuvier, 1829
Bembras longipinnis Imamura and Knapp, 1998
Bembras macrolepis Imamura, 1998
Bembras megacephala Imamura and Knapp, 1998
Brachybembras aschemeieri Fowler, 1938
? Parabembras curtus (Temminck and Schlegel, 1843) 1/
Reference
Imamura, H. and L.W. Knapp. 1998. Review of the genus Bembras Cuvier, 1829 (Scorpaeniformes: Bembridae) with
description of three new species collected from Australia and Indonesia. Ichthyol. Res., 45(2):165-178.
1/ Reported from the area by some authors, but without documentation; no museum records have been located to confirm
these reports.
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Scorpaeniformes: Platycephalidae 2385
PLATYCEPHALIDAE
Flatheads
by L.W. Knapp
D iagnostic characters: Body elongate, head moderately to strongly depressed. Mouth large, lower
jaw longer than upper. Eye partly directed upward; orbit diameter subequal to or less than snout
length. Small villiform or caniniform teeth on jaws, vomer, and palatines in most; stout canines present in
a few species. Bony ridges of head usually bearing spines or serrations. Branchiostegal rays 7. Gill
rakers few, relatively short or mere stubs; gill membranes free from isthmus. Two dorsal fins, well
separated; spinous dorsal fin with VI to X spines, first spine short, isolated or scarcely connected
to second spine; second dorsal fin with 10 to 15 soft rays; anal fin with 10 to 15 soft rays (no spines).
Pelvic fins thoracic in position, behind pectoral-fin base, set far apart towards sides of body, with I
spine and 5 soft rays. Vertebrae 27, usually 11+16 or 12+15. Lateral line complete, number of pored scales
varying from as few as 28 in Onigocia to more than 100 in Elates; most species with a small spine or ridge
on first few anterior lateral-line scales, and in a few species all or most lateral-line scales bear a spine.
Colour: usually dark above and pale below, the dark colours with various shades of brown, grey, or black;
brighter hues of reddish, purplish, or greenish in some species.
VI-X spines (usually IX)
st
1 dorsal-fin 10-15 dorsal-fin rays
spine short
lower jaw
longer lateral view (Solitas)
10-15 anal-fin rays
accessory
preopercular
supraorbital ridge spine
(and spines) (upper)
iris lappet lacrimal preopercular
spines preocular spines spine
dorsal view
(Thysanophrys)
nasal
tentacle
ocular
preorbital flap
spine
suborbital ridge lower
(and spines) opercular spines
diagram of head spines used in the identification

key and species accounts
(after Knapp in Smith and Heemstra, 1986)
Habitat, biology, and fisheries: Benthic fishes frequently found on mud or sand bottoms of the continental
shelf at depths to 300 m, most found at less than 100 m; a second group is associated with rocky shores
or coral reefs. Many species are excellent eating, a few attain sizes of 70 cm or greater.
2386 Bony Fishes

Bembridae: head less depressed; pelvic fins set
close together; first dorsal-fin spine broadly at-
tached to second spine.
Hoplichthyidae: superficially resemble platy-
cephalids, but are mostly scaleless, with a row of
large spiny scutes along the lateral line, and with 3
or 4 detached lower pectoral-fin rays. Bembridae
Percophidae: no bony ridges and spines on head;
pelvic fins set close together, in advance of pecto-
ral fins; first dorsal-fin spine broadly attached to
second spine.
Hoplichthyidae Percophidae
Identification note
Counts of dorsal-fin spines are presented as short formulas, with the first small, isolated spine counted
separately (e.g. I,VIII). In a few species, the spiny dorsal fin is followed by a second small, isolated spine
(e.g. I,VIII,I).
Key to the species of Platycephalidae occurring in the area

1a. A single, long preopercular spine; caudal fin forked, upper lobe longest, a single upper
caudal-fin ray ending in elongate filament; dorsal-fin spines VI . . . . . . . . . . . Elates ransonnetii
1b. Two or more preopercular spines; upper lobe of caudal fin not elongate, lacks elongate
caudal-fin filament; dorsal-fin spines VII to X . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Teeth on vomer in 1 trans-

verse patch (Fig. 1a); 13 or
more dorsal-fin rays; 2 preop-
ercular spines, lower spine
usually longest . . . . (Platycephalus) ® 3
2b. Teeth on vomer in 2 distinct
patches (Fig. 1b); 12 or fewer
dorsal-fin rays; 2 or more pre- a) b)
opercular spines, upper spine
longest . . . . . . . . . . . . . . . . . ® 5 Fig. 1 teeth on vomer
3a. Caudal fin with 4 or more horizontal dark streaks, no yellow blotch; first gill arch with 3
or 4 gill rakers on upper limb, 10 to 12 on lower limb . . . . . . . . . . . . . Platycephalus arenarius
3b. Caudal fin with 2 or 3 horizontal dark bars and a large yellow blotch; first gill arch with 1
to 3 gill rakers on upper limb, 6 to 10 on lower limb . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. First gill arch usually with 1 or 2 gill rakers on upper limb, 5 to 7 on lower limb; yellow
blotch located towards central area of caudal fin . . . . . . . . . . . . . . . . Platycephalus indicus
4b. First gill arch usually with 1 to 3 gill rakers on upper limb, 7 to 10 on lower limb; yellow
blotch located along upper margin of caudal fin . . . . . . . . . . . . . Platycephalus endrachtensis
ocular flap
5a. Pored lateral-line scales 28 to 42; side
of head unicarinate (lower edge of
suborbital turned inward, not visible)
(Fig. 2) . . . . . . . . . . . . . . . (Onigocia) ® 6
5b. Pored lateral-line scales 48 to 56 . . . . . . . ® 8
6a. Upper surface of eye without ocular unicarinate (single ridge on side of head)
flap; first gill arch without gill rakers on
upper limb and 4 or 5 gill rakers on Fig. 2 lateral view of head
lower limb; lateral-line scales 29 to 33
. . . . . . . . . . . . . . . . . Onigocia pedimacula
6b. Upper surface of eye bearing ocular flap (Fig. 2); first gill arch with 1 gill raker on upper
limb, and 4 or 5 on lower limb; lateral-line scales 34 to 42 . . . . . . . . . . . . . . . . . . . . . ® 7
7a. A single preocular spine; suborbital ridge with a smooth notch below eye; antorbital
margin of lacrimal bone with 2 antrorse (forward directed) spines . . . . . . . . Onigocia macrolepis
7b. Three or more preocular spines; suborbital ridge lacks notch; antorbital margin with 3
antrorse spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Onigocia spinosa
8a. All lateral-line scales bearing a stout spine . . . . . . . . . . . . . . . . . . . . (Grammoplites) ® 9

8b. Lateral-line scales lack spines on posterior part of body . . . . . . . . . . . . . . . . . . . . . . ® 10
9a. Interorbital width 9 to 13.1% of head length; lateral-line scale spines strong, extending
beyond rear margin of scale on posterior part of body . . . . . . . . . . . . . . Grammoplites scaber
9b. Interorbital width 6.4 to 8.4% of head length; lateral-line scale spines weak, not reaching
beyond rear margin of scale on posterior part of body . . . . . . . . . . . . . . Grammoplites knappi
10a. Diagonal scale rows slanting downward and backward above lateral-line usually exceed
number of lateral-line scales by 5 or more (about equal in Cymbacephalus beauforti but
pit present behind eye and rear edge of maxilla ends well in front of eye) . . . . . . . . . . . . ® 11
10b. Diagonal scale rows slanting downward and backward above lateral line about equal to
number of lateral-line scales, usually differing by only 1 or 2 scales . . . . . . . . . . . . . . . ® 17
11a. Upper preopercular-spine length approximately equal to second, no accessory spine on

base; iris lappet of eye cirrose (Fig. 3a) . . . . . . . . . . . . . . . . . . . . (Cymbacephalus) ® 12
11b. Upper preopercular-spine distinctly longer than second, accessory spine on base; iris
lappet simple or bilobed (Fig. 3b, c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
a) cirrose b) simple c) bilobed d) pointed e) crenate f) short branches

Fig. 3 various types of iris lappets
12a. No pit behind eye; soft dorsal- and anal-fin rays 12; caudal fin with a black streak along
upper and lower edge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymbacephalus bosschei
12b. A pit located behind upper rear edge of eye; soft dorsal- and anal-fin rays 11; caudal fin
with a mottled pattern of dark blotches and light streaks . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Rear edge of maxilla ends well in front of eye; a series of papillae and cirri on upper
surface of eye; number of oblique scale rows slanting backward above lateral line
approximates number of pored lateral-line scales . . . . . . . . . . . . . . Cymbacephalus beauforti
13b. Rear edge of maxilla located below or behind front of eye; number of oblique scale rows
usually exceeds number of pored lateral-line scales by 5 or more scales . . . . . . . . . . . . ® 14
2388 Bony Fishes
14a. Upper surface of eye with several cirri and papillae; suborbital ridge with a single spine;
short dark streaks crossing fin rays in dorsal and caudal fins . . Cymbacephalus nematophthalamus
14b. Upper surface of eye lacks cirri and papillae; suborbital ridge with 2 spines; no dark
streaks crossing fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cymbacephalus staigeri
15a. Caniniform jaw teeth are depressible; suborbital ridge bearing 3 or more spines below
eye; iris lappet pointed (Fig. 3d) . . . . . . . . . . . . . . . . . . . . . . . . . Ratabulus diversidans
15b. Jaw teeth not depressible; suborbital ridge bearing 2 spines under eye; iris lappet simple
or slightly bilobed (Fig. 3b, c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Cociella) ® 16
16a. Suborbital ridge with 2 spines below eye, several spines behind eye; upper preopercular
spine long, reaching nearly to opercular margin; preorbital spine slight or lacking . . . Cociella hutchinsi
16b. Suborbital ridge with 2 spines below eye, no spines behind eye; upper preopercular
spine shorter, reaching about half-way to opercular margin; preorbital spine usually well
developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cociella punctata
17a. Second dorsal-fin spine much shorter

than third; pectoral fins with falcate rear
margin, black with pale central area; scale
pores of lateral line with a single opening
to outside (Fig. 4a) . . . . . Kumococius rodericensis
17b. Second dorsal-fin spine length about
equal to third; pectoral-fin margin not fal- a) b)
cate, colour pattern not as above; scale Fig. 4 detail of lateral-line scale
pores of lateral line with 2 openings to
exterior (Fig. 4b) . . . . . . . . . . . . . . . . . . ® 18
18a. Bony ridges of head bearing many fine serrations; side of head beneath eye unicarinate
(lower edge of suborbital turned inward, not readily visible) (Fig. 5a); iris lappet bilobed
(Fig. 3c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Rogadius) ® 19
18b. Bony ridges of head bearing some larger spines; side of head beneath eye bicarinate
(lower edge of suborbital more vertically positioned, visible on side of head) (Fig. 5b); iris
lappet with crenate or short branches (Fig. 3e, f) . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
19a. No antrorse preopercular

spine; soft dorsal-fin rays
usually 12; caudal fin whit-
ish, usually with 3 horizon-
tal dark bars . . . . . Rogadius patriciae
19b. Antrorse spine (Fig. 5a)
projecting from lower pre- unicarinate antrorse spine bicarinate
opercle (barely visible in a) b)
some); soft dorsal-fin rays
usually 11; caudal fin with 1 Fig. 5 lateral view of head
or more vertical dark bands . . . . ® 20
20a. Preocular spine with small accessory spine(s) along anterior base; usually 9 (8 to 10)
scale rows between soft dorsal fin and lateral line . . . . . . . . . . . . . . . . . . Rogadius serratus
20b. Preocular spine(s) lack basal accessory spines; usually 6 or 7 (5 to 7) scale rows
between soft dorsal fin and lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 21
21a. A single preocular spine; antrorse preopercular spine stout, easily visible; caudal fin with
a broad dark subterminal band . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rogadius pristiger
21b. Two or more preocular spines; antrorse preopercular spine slight, only tip visible; caudal
fin with several narrow dark bands . . . . . . . . . . . . . . . . . . . . . . . . . Rogadius welanderi
22a. Preocular spines 2 to 6; preopercular spines 4 or more; posterior edge of breast scales
elongated to a slight point; iris lappet crenate (Fig. 3e) . . . . . . . . . . . . . Sorsogona tuberculata
22b. Preocular spines usually 1 (may have a cluster of smaller spines around base,
Ambiserrula jugosa); preopercular spines 3 or less; breast scales normal, without
posterior elongation . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
23a. Upper preopercular spine bayonet-like, reaching to or past opercular margin; total gill
rakers on first gill arch usually 8 or more (rarely 7) . . . . . . . . . . . . Suggrundus macracanthus
23b. Upper preopercular spine shorter, not reaching opercular margin; gill rakers on first gill
arch usually 7 or less (rarely 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 24
24a. Upper 2 preopercular spines short, subequal; base of lower opercular spine extends
across opercle as a bony ridge; iris lappet cirrose (Fig. 3a) . . . . . . (Inegocia, Ambiserrula) ® 25
24b. Upper preopercular spine distinctly longer than next spine; base of lower opercular spine
1/
concealed by scales; iris lappet crenate or with short branches (Fig. 3e, f) . . . (Thysanophrys) ® 27
25a. Preocular spines 1 or 2, with several smaller spines around anterior and distal base;
suborbital ridge with many small spines under entire length of eye; base of lower
opercular spine bearing serrations . . . . . . . . . . . . . . . . . . . . . . . . . . . Inegocia jugosa
25b. A single stout preocular spine, no spines around base; suborbital ridge lacking spines
under front of eye; base of lower opercular spine smooth . . . . . . . . . . . . . . . . . . . . . ® 26
26a. Soft dorsal- and anal-fin rays usually 12, rarely 11;
pectoral-fin rays 18 to 21 (usually 20); interopercular
flap present (Fig. 6) . . . . . . . . . . . . . . . . Inegocia japonica
26b. Soft dorsal- and anal-fin rays usually 11; pectoral-fin
rays 22 to 25 (usually 23 or 24); interopercular flap
lacking . . . . . . . . . . . . . . . . . . . . . . . . Inegocia harrisii
Fig. 6 interopercular flap

27a. One or more dermal papillae present on upper surface
of eye; greatest diameter of eye about equal to snout
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 28
27b. No papillae on eye; greatest diameter of eye 1.3 or more times in snout length . . . . . . . . . ® 30
28a. A row of small papilla on upper eye, 1 or 2 elongate; 7 to 10 scale rows between soft
dorsal-fin base and lateral line; iris lappet crenate (Fig. 3e) . . . . . . . . Thysanophrys carbunculus
28b. A single papilla on upper eye (minute in Thysanophrys armatus); 4 or 5 scale rows
between soft dorsal-fin base and lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 29
29a. Soft dorsal-fin rays 12; anal-fin rays 13; 2 or more preorbital spines . . . . . Thysanophrys celebicus
29b. Soft dorsal-fin rays 11; anal-fin rays 12; 1 preorbital spine . . . . . . . . . . . Thysanophrys armatus
30a. Rear edge of maxilla in front of anterior margin of orbit; least interorbital width 3.8 to 7.2
times in greatest diameter of orbit . . . . . . . . . . . . . . . . . . . . . . . Thysanophrys chiltonae
30b. Rear edge of maxilla behind front margin of orbit; least interorbital width 1.5 to 3.5 times
in orbit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 31
31a. Soft dorsal-fin rays usually 11; lips with a row of small fleshy papillae . . . . Thysanophrys otaitensis
31b. Soft dorsal-fin rays usually 12; lips smooth, without papillae . . . . . . . . . Thysanophrys arenicola
Thysanophrys to include
1/ The classification used for this field guide does not follow Imamura (1996), who restricted
T. celebica, T. chiltonae, T. armata , and 2 species not found in the area. His new genus Eurycephalus includes the
WCP species E. arenicola and E. otaitenis, as well as E. carbunculus from outside the area.
2390 Bony Fishes

The symbol0 is given when species accounts are included.
0 Ambiserrula jugosa (McCulloch, 1914)
0 Cociella hutchinsi Knapp, 1996
0 Cociella punctata (Cuvier in Cuvier and Valenciennes, 1829)
0 Cymbacephalus beauforti (Knapp, 1973)
0 Cymbacephalus bosschei (Bleeker, 1860)
0 Cymbacephalus nematophthalamus (Günther, 1860)
0 Cymbacephalus staigeri (Castelnau, 1875)
0 Elates ransonnetii (Steindachner, 1877)
0 Grammoplites knappi Imamura and Amaoka, 1994
0 Grammoplites scaber (Linnaeus, 1758)
0 Inegocia harrisii (McCulloch, 1914)
0 Inegocia japonica (Tilesius, 1812)
0 Kumococius rodericensis (Cuvier in Cuvier and Valenciennes, 1829)
0 Onigocia macrolepis (Bleeker, 1854)
0 Onigocia pedimacula (Regan, 1908)
0 Onigocia spinosa (Temminck and Schlegel, 1843)
0 Platycephalus arenarius Ramsy and Ogilby, 1886
0 Platycephalus endrachtensis Quoy and Gaimard, 1825
0 Platycephalus indicus (Linnaeus, 1758)
2/
Platycephalus fuscus Cuvier in Cuvier and Valenciennes, 1829
0 Ratabulus diversidans (McCulloch, 1914)

0 Rogadius patriciae Knapp, 1987
0 Rogadius pristiger (Cuvier in Cuvier and Valenciennes, 1829)
0 Rogadius serratus (Cuvier in Cuvier and Valenciennes, 1829)
0 Rogadius welanderi (Schultz, 1966)
0 Sorsogona tuberculata (Cuvier in Cuvier and Valenciennes, 1829)
0 Suggrundus macracanthus (Bleeker, 1869)
0 Thysanophrys arenicola Schultz, 1966)
0 Thysanophrys carbunculus (Valenciennes in Cuvier and Valenciennes, 1833)
Thysanophrys armatus (Fowler, 1938)
0 Thysanophrys celebicus (Bleeker, 1854)
0 Thysanophrys chiltonae Schultz, 1966
0 Thysanophrys otaitensis (Parkinson in Cuvier and Valenciennes, 1829)
References
Beaufort, L.F. de and J.C. Briggs. 1962. The fishes of the Indo-Australian archipelago Vol. 11. Leiden, E.J. Brill, 481 p.
Bleeker, P. 1879. Atlas ichthyologique des Indes Orientales Neerlandaises, publie sous les auspices du Gouvernement
colonial neerlandais. Tome IX, Platycephaloidei: pls 418-420.
Bleeker, P. 1879. Revision des espèces insulindiennes du genre Platycephalus . Verh. Akad. Amsterdam, XIX:1-31.
Gloerfelt-Tarp, T. and P.J. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Australia. Jakarta,
Dir. Gen. Fish. (Indonesia), German Tech. Coop., Aust. Dev. Assoc. Bur., 406 p.
Imamura, H. 1996. Phylogeny of the family Platycephalidae and related taxa (Pisces: Scorpaeniformes). Species
Diversity, 1(2):123-233.
Myers, R.F. 1989. Micronensian reef fishes. Guam, Coral Graphics, 298 p.
Sainsbury, K.J., P.J. Kailola, and G.G. Leyland. 1985. Continental shelf fishes of northern and north-western Australia.
Canberra, Glouston and Hall and Peter Pownall Fisheries Information Service, 375 p.
2/ Not included in the identification key. The dusky flathead, Platycephalus fuscus, commonly occurs in southeastern
Australia and only enters the area from Brisbane to near Mackay, Queensland. It usually has 14 soft rays both in the
second dorsal fin and anal fin, which distinguishes it from the other 3 species of Platycephalus treated here. Its caudal
fin is dusky with some dark spots in the upper lobe; horizontal dark bars are never present.
Ambiserrula jugosa (McCulloch, 1914)

FAO names: En - Mud flathead.
(from McCulloch, 1914)
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla reaches to
slightly behind front edge of eye. Preopercular spines usually 3, upper spine short, subequal with next;
small accessory spine present. Supraorbital ridge serrate, with a cluster of small spines posteriorly.
A series of small preorbital spines present; preocular spines often clustered, with 1 larger spine and
several smaller spines around its base; suborbital ridge with numerous small serrations.Teeth on
vomer in 2 separate patches. Lower side of head bicarinate. Iris lappet cirrose. Interopercular flap
truncated, tongue-shaped. Total gill-rakers on first gill arch 6. Dorsal-fin spines IX or I,VIII; dorsal-fin
rays 11 or 12 (usually 11); anal-fin rays 11 or 12 (usually 11); pectoral-fin rays 21 or 22. Oblique scale
rows slanting downward above lateral line about equal to number of lateral-line scales. Lateral-line scales
52 to 55 (usually 52 or 53), anterior 10 to 25 (mean 16.7) scales bearing a small spine or ridge. Scale pores
of lateral line with 2 openings to the exterior. Colour: head and body brown or grey above, pale below; 4
or 5 dark blotches on cheek including a broad dark bar below eye, lips also with dark blotches; back crossed
by about 4 dark bands; lower side with dark blotches and streaks; spinous dorsal fin with a submarginal
dark band, spines with dark spots; anal fin pale anteriorly, rear portion dusky; other fins with a series of
brown spots, forming vertical bands in caudal fin.
Habitat, biology, and fisheries: Found in shallow coastal waters, taken by trawls up to about 20 m.
Australia from Botany Bay
(New South Wales) to vicinity
of Bowens (Queensland).
2392 Bony Fishes
Cociella hutchinsi Knapp, 1996

Frequent synonyms / misidentifications: Suggrundus sp. 1 [Sainsbury, Kailola, and Leyland, 1985] /
None.
FAO names: En - Brownmargined flathead.
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla reaches behind
front edge of eye. Preopercular spines usually 2 (a weak third spine sometimes present), upper preoper-
cular spine long, nearly reaching to rear margin of opercle. Preorbital spine slight, often lacking.
Supraorbital ridge usually smooth over anterior quarter of eye, bearing 7 to 10 small spines posteriorly.
Suborbital ridge bearing 2 spines below eye, and 3 or 4 spines behind eye. Teeth on vomer in 2 separate
patches. Lower side of head bicarinate. Interopercular flap absent. Iris lappet simple, semicircular.
Total gill rakers on first gill arch 6 or 7 (usually 6). Dorsal-fin spines I,VII or I,VIII (usually I,VIII); dorsal-fin
rays 11 or 12 (usually 11); anal-fin rays 11; pectoral-fin rays 20 to 23 (usually 21 or 22). Oblique scale rows
slanting backward above lateral line 59 to 68 (mean 63). Lateral-line scales 51 to 55, anteriormost 2 to 8
scales bearing weak spines. Scale pores of lateral line with a single opening to the exterior. Colour: body
brownish to tannish above; white below with specks of brown sparse on breast, more heavily stippled
posteriorly; spinous dorsal fin dusky, with a broad submarginal brown band; soft dorsal fin pale, with small
brown spots on rays; anal fin white; pectoral fins dusky brown, with traces of vertical dark bands, lower
margin white; pelvic fins with whitish base, stippled with brown, with well-developed submarginal dark band;
caudal fin dusky, usually with a row of dark streaks forming a submarginal dark band.
Habitat, biology, and fisheries: Benthic species inhabiting the continental shelf off western and northern
Australia to depths of about 108 m. Caught in trawls.
Distribution: Restricted to
the Arafura and Timor seas off
Australia.
Cociella punctata (Cuvier in Cuvier and Valenciennes, 1829)

Frequent synonyms / misidentifications: None / Cociella crocodila (Tilesius, 1812).
FAO names: En - Spotted flathead.
behind front edge of eye. Preopercular spines usually 3 (sometimes 2), uppermost preopercular spine
longest, reaching only about half-way to opercular margin. Preorbital spine present. Supraorbital
ridge usually smooth over anterior half of eye, bearing 5 to 8 small spines posteriorly. Suborbital ridge
bearing 1 spine in front of eye, 2 spines below, and no spines behind eye. Teeth on vomer in 2 separate
patches. Lower side of head bicarinate. Interopercular flap present. Iris lappet simple, semicircular.
Dorsal-fin spines I,VIII; dorsal-fin rays 10 to 12 (usually 11); anal-fin rays 11 or 12 (usually 11); pectoral-fin
rays 19 to 22 (usually 20 or 21). Number of oblique scale rows slanting backward above lateral line 56 to
76 (mean 68). Lateral-line scales 50 to 56 (usually 53 or 54), anteriormost 1 to 16 scales bearing a small
spine. Scale pores of lateral line with a single opening to the exterior. Colour: head and body reddish,
greyish or brown above, whitish below, frequently with 5 or 6 dark bands crossing back; numerous small
dark spots on back reaching to below lateral line, more widely scattered posteriorly; spinous dorsal
fin with broad submarginal dark band; soft dorsal fin with dark spots on rays; anal-fin membranes dusky;
pectoral fins dusky on lower half, spotted above; pelvic fins dusky; caudal fin somewhat variable, usually
with a broad dark marginal band or series of dark spots and horizontal streaks, basal area more or less
pale.
Habitat, biology, and fisheries: Frequents shallow coastal areas in depths of 10 m or less. Taken in trawl
fisheries at depths of 23 to 250 m. Reported from a trap in the Gulf of Aqaba at a depth of 300 m. Young
to juvenile stages utilize mangrove habitat.
the Indo-Pacific; Vanuatu to
Papua New Guinea, north to
Taiwan Province of China,
through Indonesia to Thai-
land, Pakistan, the Red Sea,
and South Africa.
2394 Bony Fishes
Cymbacephalus beauforti (Knapp, 1973)

FAO names: En - Crocodile fish.
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla ends well in
front of eye. Prominent pit present behind upper eye. Preopercular spines short, usually 2 subequal,
rarely 3. Supraorbital ridge usually smooth over eye, bearing a few small spines posteriorly. Preorbital
spines lacking; a single preocular spine; suborbital ridge largely smooth, bearing 2 spines below eye.
Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Iris lappet cirrose. Dermal papillae
(10 to 12 in adults) on upper eye, some simple, some branched, longest not reaching above
supraorbital ridge. Interopercular flap usually broader than long, with several subdivisions. Total
gill rakers on first gill arch 5 or 6 (usually 6). Dorsal-fin spines I,VIII or I,IX; dorsal-fin rays 11; anal-fin
rays 11; pectoral-fin rays 19 to 21 (usually 20). Oblique scale rows slanting backward above lateral
line more or less equal to number of lateral-line scales. Lateral-line scales 50 to 55 (usually 52 or 53),
anterior 1 to 3 scales bearing a weak spine. Scale pores of lateral line with a single opening to the exterior.
Colour: body dark brown above, lower sides frequently with irregular dark brown blotches, pale below,
breast with brown streaks in some; all fins mottled with small to large dark blotches.
Habitat, biology, and fisheries: Frequents shallow sandy or coral rubble areas near seagrasses or
mangroves. Found at depths to about 8 m, but usually at 2 to 3 m.
Distribution: K n o w n f r o m
New Caledonia, Papua New
Guinea, Moluccas, Borneo,
Philippines, Palau, and Yap
Island to the Ryukyu Islands.
Cymbacephalus bosschei (Bleeker, 1860)

FAO names: En - Smalleyed flathead.
(after Taylor, 1964)
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla reaches to or
behind front of eye. No pit present behind upper eye. Preopercular spines short, usually 2 subequal,
rarely 3. Supraorbital ridge usually smooth over eye, bearing a few small spines posteriorly. Preorbital
spines lacking; a single preocular spine; suborbital ridge mostly smooth, bearing 1 slight spine just
behind eye, spine often obscure or lacking in adults. Teeth on vomer in 2 separate patches. Lower
side of head bicarinate. Iris lappet cirrose. No dermal papillae on upper eye. Interopercular flap broad, with
shallow incisions. Total gill rakers on first gill arch 6 or 7 (usually 6). Dorsal-fin spines IX or I,VIII; dorsal-fin
rays 12; anal-fin rays 12; pectoral-fin rays 18 to 21 (usually 19 or 20). Oblique scale rows slanting
backward above lateral line 55 to 68, usually about 61 (specimens from Singapore with the lowest counts).
Lateral-line scales 51 to 55 (usually 52 or 53), anterior 0 to 2 scales bearing a weak spine. Scale pores of
lateral line with a single opening to the exterior. Colour: most with body brownish above, light tan below,
back mottled with dark brown blotches; some darker specimens with white spots on head and back; fins
with dark brown spots, forming vertical bands in caudal fin; caudal fin with a large elongate dark blotch
along upper and lower margins; lower lip with about 4 dark brown bars.
Habitat, biology, and fisheries: Shallow coastal areas and reef flats to about 50 m. Usually captured by
spear fishing or with ichthyocide, occasional specimen taken by trawls.
Distribution: Widespread from Brisbane and Darwin to Western Australia, Papua New Guinea, the
Philippines, Borneo, and Sin-
gapore.
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2396 Bony Fishes
Cymbacephalus nematophthalamus (Günther, 1860)

FAO names: En - Fringe-eyed flathead.
behind front of eye. Prominent pit present behind upper eye. Preopercular spines short, upper 2
subequal, a small third spine often present. Supraorbital ridge usually smooth over eye, with a few
small spines posteriorly. Preorbital spines lacking; a single preocular spine; suborbital ridge mostly
smooth, bearing 1 spine below rear of eye. Teeth on vomer in 2 separate patches. Lower side of head
bicarinate. Iris lappet cirrose. Dermal papillae (usually 6 to 9 in adults) on upper surface of eye, longest
branched, reaching well above supraorbital ridge. Interopercular margin bearing several flaps of
varying sizes. Total gill rakers on first gill arch 5 to 7 (usually 6). Dorsal-fin spines IX or I,VIII; dorsal-fin
rays 11; anal-fin rays 11; pectoral-fin rays 19 to 21 (usually 20). Oblique scale rows slanting backward
above lateral line 57 to 70 (often 62 or 63). Lateral-line scales 51 to 55 (frequently 53), anterior 1 to 6 scales
(usually 2 or 3), bearing a weak spine or ridge. Scale pores of lateral line with a single opening to the
exterior. Colour: upper body grey-green or brown, with about 7 dark bands crossing back and reaching
far down sides; lower parts of body pale to white; fins mottled with dark and light blotches, may be yellow
and green in life; short dark streaks usually crossing rays in dorsal and caudal fins; a broad, dark bar
below eye; about 4 dark bands on lower lip.
Habitat, biology, and fisheries: Inhabits shallow rocky coastal areas, frequently associated with weed
beds and mangroves, most commonly at depths of less than 5 m.
Distribution: From the north-
western shelf of Australia to
Brisbane, New Georgia,
Papua New Guinea, Celebes,
Borneo, Singapore, and the
Philippines.
Cymbacephalus staigeri (Castelnau, 1875)

Frequent synonyms / misidentifications: Cymbacephalus parilis (McCulloch, 1914) / None.
FAO names: En - Blotched flathead.
behind front of eye. A prominent pit present behind upper eye. Preopercular spines short, 2 subequal.
Supraorbital ridge smooth over most of eye, with a few small spines posteriorly. Preorbital spines lacking;
a single preocular spine; suborbital ridge mostly smooth, bearing 2 spines below eye. Teeth on vomer
in 2 separate patches. Lower side of head bicarinate. Iris lappet cirrose. No dermal papillae on upper eye.
A prominent lobate interopercular flap present. Total gill rakers on first gill arch usually 6. Dorsal-fin spines
IX or I,VIII; dorsal-fin rays 11; anal-fin rays 11; pectoral-fin rays 20 to 22 (usually 20 or 21). Oblique scale
rows slanting backward above lateral line 59 to 71 (usually about 65). Lateral-line scales 51 to 54 (usually
52 or 53), anterior 1 or 2 scales bearing a weak spine. Scale pores of lateral line with a single opening to
the exterior. Colour: body brownish, with 6 or 7 dark bands crossing back down to lateral line; a series of
dark blotches on lower side; lower body white; usually a row of dark blotches on lower cheek; spinous
dorsal fin with a broad submarginal dark band widening posteriorly; soft dorsal fin dusky; anal fin with a
series of large brown blotches; pectoral and pelvic fins with small brown spots; and caudal fin mottled with
light streaks and dark blotches.
Habitat, biology, and fisheries: Possibly associated with aquatic vegetation. Has been taken by trawls
at depths of 15 to 46 m, frequently found at depths of 10 m or less.
Distribution: From Port
Moresby (Papua New
Guinea) and northern
Queensland, Gulf of Carpen-
teria, Darwin, Shark Bay, to
Christmas Island.
2398 Bony Fishes
Elates ransonnetii (Steindachner, 1877)

FAO names: En - Dwarf flathead.
(after Allen and Swainston, 1988)
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla ends below
anterior nostril, well in front of eye. Preopercular spine single, long, bayonet-like, reaching to or
past rear opercular margin, accessory spine lacking. Supraorbital ridge smooth. Preorbital spine
present; a single preocular spine; suborbital ridge with 2 spines, a short spine under front of eye, and a
long spine under rear of eye. Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Iris
lappet slightly developed as a simple lobe. No dermal papillae on eye. Interopercular flap lacking. Total
gill rakers on first gill arch 15 to 21. Dorsal-fin spines VI; dorsal-fin rays 12 to 14 (usually 13); anal-fin
rays 12 to 14 (usually 13); pectoral-fin rays 19 to 22 (usually 20 or 21). Caudal fin emarginate, upper
lobe longest, bearing an elongate filament. Lateral-line scales 83 to 107 (often about 94), anterior 0 to
2 scales bearing weak spines. Scale pores of lateral line with a single opening to the exterior. Colour:
head and body light tan or cream; a series of small oval dark blotches along sides; dorsal and caudal fins
with scattered black spots.
Habitat, biology, and fisheries: Taken by trawls over sand and mud bottoms at depths of about 5 to 53 m.
Distribution: From the Timor
Sea off Australia to Papua
New Guinea and northern
Queensland, through Indone-
sia, to Singapore, the Gulf of
Thailand, South China Sea,
and the Philippines.
Grammoplites knappi Imamura and Amaoka, 1994

Frequent synonyms / misidentifications: None / Grammoplites scaber (Linnaeus, 1758).
FAO names: En - Smallspined flathead.
(after Imamura and Amaoka, 1994)
behind front edge of eye. Preopercular spines 3; upper longest, not reaching to opercular margin, bearing
a small accessory spine on base. Supraorbital ridge smooth anteriorly, with a few serrations posteriorly.
Preorbital spine absent; a single preocular spine; suborbital ridge bearing 3 or 4 spines. Teeth on vomer
in 2 separate patches. Lower side of head bicarinate. Iris lappet a simple lobe. Interopercular flap absent.
Total gill rakers on first gill arch 5 to 7 (usually 6). Interorbital width less than 9% of head length.
Dorsal-fin spines IX, I,VIII, or I,VII,I (usually I,VIII); dorsal-fin rays 12; anal-fin rays 12; pectoral-fin rays
20 to 22 (usually 21 or 22). Oblique scale rows slanting backward above lateral line closely approximate
number of lateral-line scales. Lateral-line scales 51 to 55 (frequently 53), each scale with a backward
directed spine that does not extend beyond rear margin of scale on posterior part of the body. Scale
pores of lateral line with a single opening to the exterior. Colour: head and body brown, with about
4 dark bands crossing back; spinous dorsal fin dusky or with a broad dark marginal band; soft dorsal fin
with dark spots; pectoral fins with dark spots that form several vertical bands; caudal fin frequently with
2 dark bands posteriorly.
Habitat, biology, and fisheries: Taken by trawls to depths of about 32 m.
Hainan Island (China), the
South China Sea, the Gulf of
Thailand, and Johore Shoals
off Malaysia.
2400 Bony Fishes
Grammoplites scaber (Linnaeus, 1758)

FAO names: En - Rough flathead.
(after Day, 1878)
behind front edge of eye. Preopercular spines 3; upper longest, not reaching to opercular margin, bearing
a small accessory spine on base. Supraorbital ridge smooth anteriorly, with a few serrations posteriorly.
Preorbital spine usually present; a single preocular spine; suborbital ridge bearing about 3 or 4 spines.
Interorbital width 9% or more of head length. Teeth on vomer in 2 separate patches. Lower side of head
bicarinate. Iris lappet a simple lobe. Interopercular flap absent. Total gill rakers on first gill arch 6 or 7
(usually 6). Dorsal-fin spines IX, I,VIII, or I,VII,I; dorsal-fin rays 11 to 13 (usually 12); pectoral-fin rays 19
to 22 (usually 20 or 21). Oblique scale rows slanting backward above lateral line closely approximate
number of lateral-line scales. Lateral-line scales 51 to 55 (often 53), each scale with a backward directed
spine that extends beyond rear margin of scale, especially noticeable along caudal peduncle. Scale
pores of lateral line with a single opening to the exterior. Colour: head and body brownish above,
whitish below; back crossed by about 6 dark bands in some, obscure or absent in others; first dorsal fin
and pelvic fins dusky; soft dorsal fin with dark spots on rays; anal fin white or with a submarginal row of
dark spots; caudal fin dusky, some with a row of dark spots along upper edge.
Habitat, biology, and fisheries: Taken by trawls over mud and sand at depths to about 55 m.
Distribution: From the
southern Arabian Sea and
Bay of Bengal through Malay-
sia, Indonesia to Bali and
Celebes, Gulf of Thailand,
Viet Nam, Hong Kong, and
southern Philippines.
Inegocia harrisii (McCulloch, 1914)

FAO names: En - Harris’ flathead.
(after Gloerfelt-Tarp and Kailola, 1984)
behind front edge of eye. Preopercular spines usually 3, upper preopercular spine short, subequal with
next; accessory spine absent. Supraorbital ridge smooth anteriorly, with serrations over posterior half
of eye. Preorbital spine absent; a single preocular spine; suborbital ridge usually with 2 spines under
eye and several more spines behind eye.Teeth on vomer in 2 separate patches. Lower side of head
bicarinate. Iris lappet cirrose. Interopercular flap absent. Total gill rakers on first gill arch 5 to 7 (usually 6).
Dorsal-fin spines I,VIII; dorsal-fin rays 11; anal-fin rays 11; pectoral-fin-rays 22 to 25 (usually 23 or
24). Oblique scale rows slanting backward above lateral line closely approximate number of lateral-line
scales. Lateral-line scales 51 to 54 (frequently 52 or 53), anterior 6 to 36 scales bearing a small spine.
Scale pores of lateral line with 2 openings to the exterior. Colour: head and body grey or brown above,
with small brown spots, whitish below with dark stippling, several vague dark bands crossing back; dorsal
fins dusky, with small brown spots; anal fin whitish, rays dusky, darkest posteriorly; pectoral fins dusky,
with brown spots forming vertical bands; pelvic fins dusky, with a few dark spots; caudal fin dusky, with
prominent elongate dark blotches.
Habitat, biology, and fisheries: Taken over sand and mud bottoms by trawls at depths of 8 to 64 m.
Young to juvenile stages commonly found in Darwin Harbour.
Distribution: Northern Aus-
tralia from the northwestern
shelf, Napier Broome Bay,
Darwin, to the Gulf of Carpen-
teria and Pine Peak, Queens-
land; also Daru, Papua New
Guinea.
2402 Bony Fishes
Inegocia japonica (Tilesius, 1812)

Frequent synonyms / misidentifications: Inegocia isacanthus (Cuvier in Cuvier and Valenciennes,
1829) / None.
FAO names: En - Japanese flathead; Fr - Platycéphale japonais; Sp - Chato japones.
behind front edge of eye. Preopercular spines usually 3, upper preopercular spine short, subequal with
next; small accessory spine usually present. Supraorbital ridge smooth anteriorly, with serrations over
posterior half of eye. Preorbital spine absent; a single preocular spine; suborbital ridge usually smooth
except for 2 spines under eye (specimens from Indonesia and Australia frequently have a few additional
small serrations). Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Iris lappet cirrose.
Interopercular flap finger-like, pointed. Total gill rakers on first gill arch 5 to 7 (usually 6). Dorsal-fin
spines IX or I,VIII; dorsal-fin rays 11 to 13 (usually 12); anal-fin rays 11 to 13 (usually 12); pectoral-fin
rays 19 to 21 (frequently 19 or 20). Oblique scale rows slanting downward above lateral line closely
approximate number of lateral-line scales. Lateral-line scales 51 to 55 (usually 52 or 53), anterior 2 to 21
scales bearing a small spine or ridge. Scale pores of lateral line with 2 openings to exterior. Colour: head
and body brown or grey above, whitish below, back crossed by about 6 obscure dark bands; dorsal fins
clear, with small brown spots on rays; pectoral fins with somewhat larger brown spots, pelvic fins with pale
base, several vague dark bands near tip; caudal fin pale, with large dark spots, some elongated.
Habitat, biology, and fisheries: Taken by trawls over sand and mud to depths of about 85 m.
Distribution: Widespread,
from Cockburn Sound (West-
ern Australia) to Moreton Bay
(Queensland); also Papua
New Guinea, Indonesia, Phil-
ippines, southern Japan,
South China Sea, Gulf of
Thailand to India and Sri
Lanka.
Kumococius rodericensis (Cuvier in Cuvier and Valenciennes, 1829)

Frequent synonyms / misidentifications: Kumococius detrusus (Jordan and Seale, 1905) / None.
FAO names: En - Spiny flathead; Fr - Platycéphale cocarde; Sp - Chato rodrigo.
behind front edge of eye. Preopercular spines 3, upper much longer, reaching nearly to or just past
opercular margin; accessory spine present. Supraorbital ridge with stout serrations over eye. Preorbital
spine present; a single preocular spine; suborbital ridge bearing numerous strong spines and serrations.
Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Iris lappet simple or slightly bilobed.
Interopercular flap present. Total gill rakers on first gill arch 8 to 12 (usually 10 or 11). Dorsal-fin spines
XI or I,VIII; dorsal-fin rays 11 or 12 (usually 11); second dorsal-fin spine noticeably shorter than third.
Anal-fin rays 11 to 13 (usually 12); pectoral fin-rays 19 to 22 (usually 19 or 20); pectoral fin with a falcate
posterior margin. Oblique scale rows slanting backward above lateral line closely approximate number
of lateral-line scales. Lateral-line scales 50 to 54 (frequently 52 or 53), anterior 2 to 22 scales (mean 10.6)
bearing small spines (2 specimens from the Moluccas had spines on most of the lateral-line scales). Scale
pores of lateral line with a single opening to the exterior. Colour: head and body brown above, white below
with scattered minute black spots, about 5 broad dark bands crossing back in some; spinous dorsal fin
dusky with a black blotch frequently present distally between second and third spines and a second blotch
at edge above middle of fin; anal fin whitish with a dusky margin; pectoral fins dark brown, with a clear
or whitish central area; pelvic fins dark brown with a light edge; caudal fin dusky on posterior half.
Habitat, biology, and fisheries: Taken by trawls over sand and mud bottoms at depths of 18 to 130 m.
D i s t r i b u t i o n : Widespread
from the Gulf of Oman to the
Bay of Bengal, Malaysia, Gulf
of Thailand, Indonesia, north-
ern Australia, Philippines,
South China Sea, and south-
ern Japan.
2404 Bony Fishes
Onigocia macrolepis (Bleeker, 1854)

FAO names: En - Notched flathead.
front edge of eye. Preopercular spines 3, uppermost longest, with an accessory spine on base. Supraor-
bital ridge smooth over front half of eye, serrated posteriorly. Preorbital spines 1 to 4 (usually 2 or 3);
a single preocular spine; suborbital ridge bearing numerous small spines or serrations, with a distinct
smooth notch below middle of eye. Antorbital margin with 2 antrorse spines. Teeth on vomer in 2
separate patches. Lower side of head unicarinate. Iris lappet bearing numerous short branches with
bifurcate tips. Upper surface of eye with a short, branched cirrus. Interopercular flap absent. Total gill
rakers on first gill arch 5 to 7 (usually 5 or 6). Dorsal-fin spines IX or I,VIII; dorsal-fin rays 11 or 12; anal-fin
rays 11 to 13 (usually 12); pectoral-fin rays 21 to 23 (usually 21 or 22). Oblique scale rows slanting backward
above lateral line closely approximate number of lateral-line scales. Lateral-line scales 34 to 41
(frequently 37 or 38), anterior 2 to 10 (usually 2 to 5) scales bearing spines. Scale pores of lateral line
with 2 openings to the exterior. Colour: body light brown above, pale below; back crossed by about 4 dark
bands; anal fin pale, other fins mostly with brown spots or blotches forming bands.
Habitat, biology, and fisheries: Taken by trawls over sand and mud bottoms to 130 m.
Distribution: Sea of Japan,
Yellow Sea, South China Sea,
Philippines to the northwest-
ern shelf of Australia, Indone-
sia, Gulf of Thailand, and
Andaman Sea.
Onigocia pedimacula (Regan, 1908)

Freguent synonyms / misidentifications: None / None.
FAO names: En - Broadband flathead.
behind front edge of eye. No supraocular cirrus present. Preopercular spines 3, upper longest, with an
accessory spine on base. Supraorbital ridge entirely serrated over eye. Preorbital with 2 to 6 serrations;
preocular spines 3 to 5, slanting towards midline; suborbital ridge bearing numerous small spines or
serrations, no smooth notch under eye. Antorbital margin with 2 antrorse spines. Teeth on vomer in
2 separate patches. Lower side of head unicarinate. Iris lappet bilobed or crenate. Upper surface of
eye smooth, no cirrus present. Interopercular flap absent. Total gill rakers on first gill arch 4 or 5
(usually 4). Dorsal-fin spines IX or I,VIII; dorsal-fin rays 11; anal-fin rays 10 or 11 (usually 11); pectoral-fin
rays 19 to 23 (usually 21 or 22). Oblique scale rows slanting backward above lateral-line about equal to
number of lateral-line scales. Lateral-line scales 29 to 33 (frequently 30 or 31), anterior 2 to 6
(frequently 3) scales bearing spines. Scale pores of lateral line with 2 openings to the exterior. Colour:
head and body grey or brown above, whitish below; back crossed by 4 to 6 brown or olive dark bands, the
widest under first dorsal fin; spinous dorsal-fin dusky; soft dorsal, pectoral, and caudal fins more or less
with dark spots; pelvic fins with a prominent dark blotch in middle, a smaller dark blotch near base.
Habitat, biology, and fisheries: Taken by trawls over sand or coral-rubble bottom at 15 to 110 m, also
taken by scuba divers off coral or rocky slopes.
from off Natal, South Africa to
Karachi, Maldives, South
China Sea, Philippines, Irian
Barat (Teluk Berau), north-
western shelf of Australia, Ti-
mor and Arafura seas, Great
Barrier Reef, Guadalcanal,
and Tonga.
2406 Bony Fishes
Onigocia spinosa (Temminck and Schlegel, 1843)

FAO names: En - Largescaled spiny flathead.
front of eye. Preopercular spines 3, upper longest, with an accessory spine on base. Supraorbital ridge
entirely serrated over eye. Preorbital serrations 3 to 6; preocular spines 3 to 5, not noticeably slanting
towards midline; suborbital ridge bearing numerous small spines; no smooth notch under eye.
Antorbital margin with 3 antrorse spines. Teeth on vomer in 2 separate patches. Lower side of head
unicarinate. Iris lappet bearing numerous short branches with bifurcate tips. Upper surface of eye
with a short, branched cirrus. Interopercular flap absent. Total gill rakers on first gill arch 5 or 6
(usually 5). Dorsal-fin spines IX or I,VIII; dorsal-fin rays 11 or 12 (usually 12); anal-fin rays 11 or 12
(usually 12); pectoral-fin rays 20 to 23 (usually 21 or 22). Oblique scale rows slanting backward above
lateral line are about equal to number of lateral-line scales. Lateral-line scales 34 to 42 (frequently 37
or 38), anterior 7 to 27 bearing a small spine or ridge. Scale pores of lateral line with 2 openings to the
exterior. Colour: head and body tan or brown above, pale below, back crossed by 3 or 4 dark bands;
anterior half of spinous dorsal with brown spots on spines, posterior half of fin with dusky blotches forming
about 3 horizontal bands; soft dorsal fin with series of brown spots forming horizontal bands; pectoral fins
with dark spots forming vertical bands; pelvic fins mostly dark, light distally; anal fin pale, with a faint dusky
submarginal band; caudal fin with several vertical dark bands.
Habitat, biology, and fisheries: Taken by trawls over sand and mud bottoms to about 250 m.
Distribution: Southern Ja-
pan, South China Sea, and
t h e P h i l i p p i n e s , t h r o u gh
Arafura and Timor seas to the
northwestern shelf of Austra-
lia.
Platycephalus arenarius Ramsay and Ogilby, 1886

FAO names: En - Sand flathead.
Diagnostic characters: Body elongate, head strongly depressed. Rear edge of maxilla reaches to about
front third of eye. Preopercular spines 2, lower distinctly longer; no accessory spine on base of upper.
Supraorbital ridge smooth. Preorbital spine lacking; a single preocular spine. Suborbital ridge smooth in
adults, bearing a spine just behind eye in juveniles. Teeth on vomer in a single transverse band. Lower
side of head bicarinate. Iris lappet a simple elongated lobe. Interopercular flap present, finger-like in
shape. Total gill rakers on first gill arch 12 to 16. Dorsal-fin spines I,VI, I,VII, VII,I, or VIII;
dorsal-fin rays 13 or 14 (usually 13); anal-fin rays 13 or 14 (usually 13); pectoral-fin rays 19 to 21
(frequently 20). Diagonal scale rows slanting backward above lateral line 82 to 96 (mean 89).
Lateral-line scales 69 to 83 (mean 74), anteriormost scale usually with a small spine or ridge. Scale
pores of lateral line with 1 opening to the outside. Colour: head and body with many small brown blotches
above, whitish below; dorsal fins with small brown spots on rays; pectoral and pelvic fins with small brown
spots forming bands; anal fin whitish; caudal fin white, with 5 or 6 dark horizontal streaks which
become narrower above.
Habitat, biology, and fisheries: Taken by angling along sandy beaches and by trawls to depths of about
55 m, usually less. Juveniles frequently found in shallows.
Distribution: From Jervis Bay
(North South Wales), across
northern Australia to Hamelin
Bay (Western Australia); also
recorded from Bali (Indone-
sia).
2408 Bony Fishes
Platycephalus endrachtensis Quoy and Gaimard, 1825

Frequent synonyms / misidentifications: None / Platycephalus indicus (Linnaeus, 1758).
FAO names: En - Yellowtailed flathead.
(after Taylor, 1964)
below middle of eye. Preopercular spines 2, lower usually longest; accessory spine usually lacking on
base of upper preopercular spine. Supraorbital ridge smooth. Preorbital spine lacking; a single preocular
spine, obscure in large adults. Suborbital ridge smooth in adults, bearing a spine below rear of eye in
juveniles. Teeth on vomer in a single transverse band. Lower side of head bicarinate. Iris lappet a
simple elongated lobe. Interopercular flap present, finger-like in shape. Total gill rakers on first gill
arch 9 to 13 (usually 10 or 11). Dorsal-fin spines I,I,VI,I, I,VII,I, or I,I,VII,I; dorsal-fin rays 13; anal-fin
rays usually 13; pectoral-fin rays 18 to 20 (usually 19 or 20). Diagonal scale rows slanting backward
above lateral line 93 to 109. Lateral-line scales 67 to 77, anteriormost scale usually with a small
spine or ridge. Scale pores of lateral line with 1 opening to the outside. Colour: head and body covered
with small brown flecks, whitish below, several indistinct dark bands crossing back in some; dorsal,
pectoral, and pelvic fins with small brown spots on rays; caudal fin whitish, with 3 or 4 horizontal dark
bars and a prominent yellow blotch along upper margin.
Size: Maximum total length about 40 cm, commonly to 3 m.
Habitat, biology, and fisheries: Taken by handlines and seines in shallow coastal waters; also taken by
trawls over mud and sand at shallow depths. Frequently found in estuaries.
Freemantle (Western Austra-
lia), across northern Australia
to Brisbane. Also appears to
reach the southern coasts of
Java and Papua New Guinea.
Platycephalus indicus (Linnaeus, 1758)

FAO names: En - Bartail flathead; Fr - Platycéphale indian; Sp - Chato índico.
below middle of eye. Preopercular spines 2, lower usually longest; a trace of an accessory spine usually
present on base of upper spine. Supraorbital ridge smooth. Preorbital spine lacking; a single preocular
spine, obscure in large adults. Suborbital ridge smooth in adults, bearing a spine below rear of eye in
juveniles. Teeth on vomer in a single transverse band. Lower side of head bicarinate. Iris lappet a
simple elongated lobe. Interopercular flap present, finger-like in shape. Total gill rakers on first gill
arch 7 to 10 (usually 8 or 9). Dorsal-fin spines I,VII,I, I,I,VII,I or I,VIII; dorsal-fin rays 13; anal-fin
rays usually 13; pectoral-fin rays 17 to 20 (usually 18 or 19). Diagonal scale rows slanting backward
above lateral line 83 to 107. Lateral-line scales 65 to 81, anteriormost scale usually with a small
spine or ridge. Scale pores of lateral line with a single opening to the outside. Colour: head and body
covered with small brown flecks, whitish below, several indistinct dark bands crossing back in some; dorsal,
pectoral, and pelvic fins with small brown spots on rays; caudal fin with 2 or 3 horizontal dark bars, a
prominent yellow blotch near middle of fin, whitish areas in upper lobe and along lower margin.
Habitat, biology, and fisheries: Taken by handlining and seining in shallow coastal waters; taken by
trawls over mud and sand at depths to 30 m, usually less. Frequently found in estuaries, small juveniles
have been taken in fresh water.
from the eastern Mediterra-
nean, Red Sea, to South Af-
rica, northern Indian Ocean to
Indonesia, Korea, southern
Japan, Philippines, and north-
ern and eastern Australia.
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2410 Bony Fishes
Ratabulus diversidans (McCulloch, 1914)

Frequent synonyms / misidentifications: Ratabulus megacephalus (Tanaka, 1917) / None.
FAO names: En - Freespine flathead.
(after McCulloch, 1914)
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla about even
with front margin of eye. Preopercular spines 2 or 3, uppermost longest, accessory spine present on base;
third spine minute, when present. Supraorbital ridge smooth over anterior half of eye, with stout serrations
posteriorly. Preorbital spine present; a single preocular spine; suborbital ridge bearing numerous spines
and serrations, the 2 largest under the eye. Some caniniform teeth on jaws and palate are depressible;
teeth on vomer in 2 separate patches. Lower side of head bicarinate. Iris lappet a small simple lobe.
Interopercular flap absent. Total gill rakers on first gill arch 7 to 9 (usually 7 or 8). Dorsal-fin spines IX,I
or I,VIII,I; dorsal-fin rays 11; anal-fin rays 11 or 12 (usually 12); pectoral-fin rays 18 to 20 (usually 19 or
20). Diagonal scale rows slanting backward above lateral line 64 to 79. Lateral-line scales 52 to 55 (usually
53 or 54), anterior 2 to 8 scales bearing a small spine. Scale pores of lateral line with a single opening to
the exterior. Colour: body grey to light tan above, whitish below; back crossed by 7 or 8 indistinct dark
bands; a series of purplish blotches along midsides; orange or brown spots on head, larger specimens
may have orange or green spots on upper body; spinous dorsal fin with a dark submarginal band, anterior
basal area clear, with scattered dark spots; soft dorsal and pectoral fins with fine brown flecks; anal fin
whitish; pelvic fins with scattered dark spots; caudal fin with submarginal dark band, basal area whitish
with scattered dark spots above.
Habitat, biology, and fisheries: Taken by trawls at depths of 79 to 261 m. Recorded at Galathea Station
number 539 as being dredged over gravel.
Distribution: From East
China Sea, South China Sea,
and northern Philippines to
northwestern shelf of Austra-
lia, Timor Sea, Coral Sea, and
B o t a n y B a y ( Ne w S o u t h
Wales).
Rogadius patriciae Knapp, 1987

Frequent synonyms / misidentifications: Suggrundus sp. 2 [Sainsbury, Kailola, and Leyland, 1985] /
None.
FAO names: En - Blackbanded flathead.
below front edge of eye. Preopercular spines 2 to 5 (usually 3 or 4), uppermost longest, with an accessory
spine on its base; antrorse spine lacking. Supraorbital ridge bearing many fine serrations. Preorbital
spine lacking, several small serrations present; preocular spine 1. Teeth on vomer in 2 separate patches.
Lower side of head unicarinate. Iris lappet bilobed. Interopercular flap absent. Total gill rakers on first
gill arch 5 to 8 (usually 7). Dorsal-fin spines IX or I,VIII; dorsal-fin rays 11 or 12 (usually 12), anal-fin
rays 11 or 12 (usually 11); pectoral-fin rays 20 to 23 (frequently 22). Number of scale rows between
soft dorsal-fin origin and lateral line 6 to 11 (usually 7 or 8). Oblique scale rows slanting backward
above lateral line are about equal to number of lateral-line scales. Lateral-line scales 49 to 55 (frequently
52 or 53), anterior 4 to 11 bearing a spine or ridge. Scale pores of lateral line with 2 openings to the outside.
Colour: body light tan or orange above, pale below; a brown band crossing nape to preopercular spines;
suborbital dark bar present; spinous dorsal fin with marginal dark band; soft dorsal fin with small black
spots on rays; anal fin white, with dusky basal band becoming black posteriorly; pectoral-fin rays bearing
numerous small brown spots on upper fin, lower part black with white lower edge; pelvic fins black with
outer edge and ray tips white; caudal fin white with several dark blotches along upper edge and 2 or
3 elongate horizontal black bars below.
Habitat, biology, and fisheries: Taken by trawls on coastal shelves at depths of 14 to 100 m.
shelf of Australia, northern
Queensland off Cairns, New
Caledonia, Philippines, Tai-
wan Province of China, and
Ryukyu Islands.
2412 Bony Fishes
Rogadius pristiger (Cuvier in Cuvier and Valenciennes, 1829)

Frequent synonyms / misidentifications: None / Rogadius asper (Cuvier, 1829).
FAO names: En - Thorny flathead.
front edge of eye. Preopercular spines 3 to 6 (usually 4 or 5), upper longest, with an accessory spine on
base; lower edge of preopercle projecting downward as a strong antrorse spine. Supraorbital and
suborbital ridges bearing many fine serrations. Preorbital spine lacking, several small serrations
present; preocular spine 1. Teeth on vomer in 2 separate patches. Lower side of head unicarinate. Iris
lappet bilobed. Interopercular flap absent. Total gill rakers on first gill arch 5 to 8 (frequently 7). Dorsal-fin
spines IX or I,VIII; dorsal-fin rays 10 to 12 (usually 11); anal-fin rays 10 to 12 (usually 11); pectoral-fin
rays 20 to 24 (usually 22 or 23). Number of scale rows between soft dorsal-fin origin and lateral line
5 to 7 (usually 6 or 7). Oblique scale rows slanting backward above lateral line about equal to number of
lateral-line scales. Lateral-line scales 49 to 55 (usually 52 or 53), anterior 4 to 14 scales bearing a small
spine or ridge. Scale pores of lateral line with 2 openings to the outside. Colour: body brown above, pale
below, 3 or 4 dusky bands crossing back; lower lip dusky, with 4 to 6 dark bands; spinous dorsal fin with
a black marginal band and a triangular light basal area towards front; soft dorsal fin with brown spots on
rays; pectoral fins with base and lower edge white, upper rays with brown dash marks, middle of fin dusky;
pelvic fins with black blotches; caudal fin with a white basal band, a broad dusky submarginal band
and a narrow white edge.
Habitat, biology, and fisheries: Taken by trawls over mud and sand bottoms at depths to about 80 m.
New Caledonia, Papua New
Guinea, northwestern shelf of
Australia, Indonesia, Philip-
pines, Gulf of Thailand, and
northern Indian Ocean to Gulf
o f O m a n , R e d S e a , a nd
Madagascar.
Rogadius serratus (Cuvier in Cuvier and Valenciennes, 1829)

FAO names: En - Serrated flathead.
front edge of eye. Preopercular spines 4 or 5 (usually 4), uppermost longest, with an accessory spine on
base; tip of antrorse spine sometimes difficult to see. Supraorbital and suborbital ridges bearing many
fine serrations. Preorbital spine absent, several small serrations present; preocular spine 1, with an
accessory spine on anterior base. Teeth on vomer in 2 separate patches. Lower side of head
unicarinate. Iris lappet bilobed. Interopercular flap absent. Total gill rakers on first gill arch 6 to 8 (usually
7). Dorsal-fin spines IX or I,VIII; dorsal-fin rays 11 (rarely 12); anal-fin rays 11; pectoral-fin rays 20 to 23
(usually 21 or 22). Number of scale rows between soft dorsal-fin origin and lateral line usually 9 (8
to 10). Oblique scale rows slanting backward above lateral line about equal to number of lateral-line scales.
Lateral-line scales usually 51 to 54 (rarely 50), anterior 1 to 9 scales bearing a small spine. Scale pores
of lateral line with 2 openings to the outside. Colour: body light tan above, pale below, back crossed by
about 7 or 8 dark bands, side with a series of dark blotches; a broad suborbital dark bar; spinous dorsal
dusky; soft dorsal fin with dark spots on rays; anal fin whitish, with submarginal dark band; pectoral fins
dark below with whitish edge, pale above with dark spots; pelvic fins with white base, black distally; caudal
fin with dark basal band, whitish central band, blackish submarginal band becoming paler above,
upper area usually with a few distinct dark spots.
Habitat, biology, and fisheries: Taken over sand around coral reefs to a depth of 11 m; a single specimen
taken by trawls at 45 m.
Distribution: Mauritius, Sey-
chelles, Somalia, Pakistan,
Sri Lanka, Maldives, Chagos
Archipelago, Indonesia, Phil-
ippines, Port Moresby, north-
ern Queensland, and New
Caledonia.
2414 Bony Fishes
Rogadius welanderi (Schultz, 1966)

FAO names: En - Welander’s flathead.
(after Gloerfelt-Tarp and Kailola, 1984)
front edge of eye. Preopercular spines 4 or 5, uppermost longest, with an accessory spine on base; small
antrorse spine present. Supraorbital and suborbital ridges bearing many fine serrations. Several
small preorbital serrations present; preocular spines 2 or 3. Teeth on vomer in 2 patches. Lower side of
head unicarinate. Iris lappet slightly bilobed. Interopercular flap absent. Total gill rakers on first gill arch
7 or 8. Dorsal-fin spines IX; dorsal-fin rays 11; anal-fin rays 11; pectoral-fin rays 21 or 22. Number of scale
rows between soft dorsal-fin origin and lateral line usually 6 or 7. Oblique scale rows slanting backward
above lateral line about equal to number of lateral-line scales. Lateral-line scales 52 or 53, anterior 5 to 9
scales bearing a small spine. Scale pores of lateral line with 2 openings to the outside. Colour: body light
brown above, pale below; several indistinct dark bands crossing back; dorsal and pelvic fins dusky; anal
fin pale, with a narrow submarginal dark band; pectoral fins with vague dark spots above, dusky below;
caudal fin pale, with about 4 narrow vertical dark bands.
Habitat, biology, and fisheries: Taken over sand near coral reefs to a depth of 40 m.
Distribution: American
Samoa, Marshall Islands, and
Moluccas; to Réunion Island
(Western Indian Ocean).
Sorsogona tuberculata (Cuvier in Cuvier and Valenciennes, 1829)

FAO names: En - Tuberculated flathead; Fr - Platycéphale poignard; Sp - Chato puñalero.
Diagnostic characters: Body elongate, head moderately depressed. Posterior edge of breast scales
elongated to a point. Rear end of maxilla reaches to below front margin of eye. Preopercular spines 4
to 9, decreasing in size ventrally, accessory spine on base of upper. Supraorbital and suborbital ridges
bearing small spines or serrae (some bicuspid). Preorbital spine absent, several small serrations
present; preocular spines 2 to 6. Teeth on vomer in 2 separate patches. Lower side of head bicarinate.
Iris lappet crenate. Interopercular flap absent. Total gill rakers on first gill arch 7 to 9, frequently 8.
Dorsal-fin spines IX or I,VIII; dorsal-fin rays 10 to 12 (mostly 11); anal-fin rays 10 to 12 (mostly 11);
pectoral-fin rays 19 to 22 (frequently 21). Scales in rows above anterior lateral line sometimes bearing
a hook or tubercle. Oblique scale rows slanting backward above lateral line about equal to number of
lateral-line scales. Lateral-line scales 47 to 54 (usually 51 or 52), anterior 12 to 28 bearing a small spine
or ridge. Pores of lateral-line scales with 2 openings to the outside. Colour: body light olive or tan above,
whitish below, with 3 or 4 dark bands crossing back; spinous dorsal fin with a submarginal brown band;
soft dorsal fin with small brown spots on rays; anal fin white, with a few dark streaks on posterior rays;
pectoral and pelvic fins with several black bands, margins whitish; caudal fin whitish with faint brown spots
on rays and several vague vertical brown bands.
Habitat, biology, and fisheries: Taken by trawls over mud and sand to a depth of about 80 m.
Distribution: From Persian
Gulf, northern Indian Ocean,
Malaysia, Gulf of Thailand, In-
donesia, Philippines, north-
Gulf of Carpenteria, Coral
Sea, and Platypus Bay
(Queensland).
2416 Bony Fishes
Suggrundus macracanthus (Bleeker, 1869)

Frequent synonyms / misidentifications: Suggrundus sundaicus (Bleeker, 1878) / None.
FAO names: En - Largespined flathead.
(after Day, 1878)
front edge of eye. Preopercular spines 3, upper long, bayonet-like, reaching to or past opercular
margin, with an accessory spine on base. Supraorbital ridge smooth over anterior third of eye, with stout
spines posteriorly. Preorbital spine present; a single preocular spine. Teeth on vomer in 2 separate patches.
Lower side of head bicarinate. Iris lappet bilobed in adults, crenate in juveniles. Finger-like interop-
ercular flap present. Total gill rakers on first gill arch 7 to 9 (usually 8). Dorsal-fin spines IX or I,VIII;
dorsal-fin rays 11 or 12 (usually 12); anal-fin rays 12 or 13 (usually 12); pectoral-fin rays 20 to 23 (usually
21 or 22). Oblique scale rows slanting downward above lateral line about equal to number of lateral-line
scales. Lateral-line scales 50 to 55 (usually 52 or 53), anterior 13 to 33 scales bearing a small spine or
ridge. Scale pores of lateral line with 2 openings to the outside. Colour: brown above, whitish below, with
about 7 vague dark bands crossing back in some; spinous dorsal fin dusky, with black blotches; soft dorsal
fin clear with small brown spots on rays; anal fin pale, a few dark streaks on posterior rays; pectoral fins
greyish below, with dark spots above; pelvic fins greyish; caudal fin brownish.
Habitat, biology, and fisheries: Taken by trawls over mud and sand at depths to 132 m.
Distribution: Southern India
and Sri Lanka to Malaysia,
Gulf of Thailand, Hong Kong,
Taiwan Province of China, In-
donesia, Philippines, north-
Coral Sea.
Thysanophrys arenicola Schultz, 1966

FAO names: En - Broadhead flathead.
below front edge of eye. Lips smooth, lacking papillae on edges. Least interorbital width usually
3.5 times or less in greatest diameter of orbit. Preopercular spines 2 or 3 (usually 3), upper spine longest,
usually reaching less than half-way to opercular margin, accessory spine lacking. Supraorbital ridge
smooth over anterior half of eye, with about 4 or 5 spines posteriorly. Suborbital ridge smooth anteriorly,
with about 4 or 5 spines posteriorly. Preorbital spine rarely present; preocular spine 1. Teeth on vomer in
2 separate patches. Lower side of head bicarinate. Base of lower opercular spine concealed by scales.
Iris lappet bearing short branches with bifurcate tips. Interopercular flap absent. Total gill rakers on
first gill arch 5 to 7 (usually 6). Dorsal-fin spines IX or I,VIII; dorsal-fin rays 11 to 13 (usually 12); anal-fin
rays 11 to 13 (usually 12); pectoral-fin rays 19 to 22 (usually 21). Oblique scale rows slanting backward
above lateral line about equal to number of lateral-line scales. Lateral-line scales 50 to 55 (usually 52 or
53), anterior 1 to 3 scales (usually 1 or 2), bearing a small spine. Scale pores of lateral line with 2
openings to the outside. Colour: body tan or greyish, back often covered with small dark flecks and 4 or
5 vague cross bands, some specimens with a marbled pattern of small dark and light blotches on dorsum,
undersides whitish; fins flecked with small brown spots.
Habitat, biology, and fisheries: Taken over sand associated with rocks and coral reefs to a depth of about
30 m.
Distribution: Mozambique,
Mauritius, Seychelles, Aden,
Chagos Archipelago, Indone-
s i a , T a i w a n P r o v i nc e of
China, Ryukyu Islands, Mari-
ana and Marshall islands,
Philippines, Torres Strait, and
northern Queensland to New
Caledonia and Fiji.
2418 Bony Fishes
Thysanophrys carbunculus (Valenciennes in Cuvier and Valenciennes, 1833)

FAO names: En - Papillose flathead.
below front edge of eye. Single row of papillae, often 7 to 9, on upper surface of eye, middle one
elongate. Preopercular spines 2 or 3 (usually 3), upper longest, not reaching to opercular margin,
accessory spine present in most. Supraorbital ridge bearing spines throughout its length; suborbital ridge
with 4 to 6 stout spines. Preorbital spine present, preocular spine 1. Teeth on vomer in 2 separate patches.
Lower side of head bicarinate. Base of lower opercular spine concealed by scales. Iris lappet crenate.
Interopercular flap absent. Total gill rakers of first gill arch 6. Dorsal-fin spines IX, I,VIII, or I,VII; dorsal-fin
rays 10 to 12 (usually 11); anal-fin rays 11 to 13 (usually 12); pectoral-fin rays 18 or 19 (frequently 19).
Oblique scale rows slanting backward above lateral line about equal to number of lateral-line scales.
Lateral-line scales 50 to 54 (usually 52 or 53), anterior 2 to 21 lateral-line scales bearing a small spine.
Scale pores of lateral line with 2 openings to the outside. Colour: body greenish above, whitish below,
back crossed by 4 or 5 dark bands, a series of small dark blotches along lower side; spinous dorsal-fin
with a broad submarginal dark band; soft dorsal, pectoral, and pelvic fins with numerous dark spots or
blotches; anal fin white with a faint dark submarginal band; caudal fin pale with rows of dark spots forming
several vertical bands.
Habitat, biology, and fisheries: May be associated with aquatic vegetation at shallow depths.
Distribution: From Bombay
(India) to Singapore, Gulf of
Thailand, North Borneo, and
Indonesia.
Thysanophrys celebicus (Bleeker, 1854)

FAO names: En - Celebes flathead.
below front edge of eye. A single small unbranched papilla on posterior half of upper surface of eye.
Preopercular spines 3, upper longest, not reaching half-way to opercular margin, accessory spine present
on base. Supraorbital ridge with spines throughout; suborbital ridge with 8 to 10 spines. Preorbital spines
2 or 3; preocular spine l. Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Base of
lower opercular spine concealed by scales. Iris lappet with very short branches. Interopercular flap
absent. Total gill rakers on first gill arch 5 to 7 (usually 5 or 6). Dorsal-fin spines IX, VIII, or I,VII (usually
I,VII); dorsal-fin rays 12; anal-fin rays 12 or 13 (usually 13); pectoral-fin rays 18 to 20 (usually 19).
Oblique scale rows slanting downward above lateral line about equal to number of lateral-line scales.
Lateral-line scales 50 to 53 (usually 51 or 52), anterior 2 to 6 lateral-line scales bearing a small spine.
Scale pores of lateral line with 2 openings to the outside. Colour: body brownish or greenish above, whitish
below, back crossed by 4 or 5 dark bands; spinous dorsal-fin with a broad submarginal dark band; soft
dorsal, pectoral, and pelvic fins with numerous dark spots; caudal fin with a basal dark band followed by
a clear band and 4 dark narrow bands on distal part of fin; rear of head with a narrow dark band, a bold
dark bar under eye.
Habitat, biology, and fisheries: Inhabits shallow coastal areas and appears to be taken over sandy bottom
with aquatic vegetation. Taken by trawls to a depth of 43 m, but usually taken during scuba diving at 20 m
or less. Frequently taken near coral rubble and rock.
Distribution: From South
Africa to Persian Gulf, Sri
Lanka, southern India, Gulf of
Thailand, Hong Kong, Taiwan
Province of China, Philip-
pines, Celebes, Irian Barat,
Arafura Sea, northern
Queensland, and the Solo-
mon Islands.
2420 Bony Fishes
Thysanophrys chiltonae (Schultz, 1966)

FAO names: En - Longsnout flathead; Fr - Platycéphale clin d’oeil; Sp - Chato guiñón.
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla is in front of
anterior margin of eye. Preopercular spines 3, upper longest, accessory spine usually absent. Supraorbital
ridge smooth over anterior half of eye, with serrations posteriorly; suborbital ridge with about 6 or 7 spines.
Least interorbital width 3.8 to 7.2 times in greatest eye diameter. Preorbital spines 1 to 3; preocular
spine 1. Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Base of lower opercular
spine concealed by scales. Iris lappet bearing short branches with bifurcate tips. Interopercular
margin incised, forming a broad lobe. Total gill rakers on first gill arch 5 to 7 (usually 6). Dorsal-fin spines
IX, I,VIII, or I,VII; dorsal-fin rays 11 to 12 (usually 11); anal-fin rays 12; pectoral-fin rays 19 to 22 (usually
20 or 21). Oblique scale rows slanting downward above lateral line about equal to number of lateral-line
scales. Lateral-line scales 50 to 54 (usually 52 or 53), anterior 1 to 3 (usually 3) lateral-line scales,
bearing a small spine. Scale pores of lateral line with 2 openings to the outside. Colour: body light tan
mottled with numerous white spots, white below, with about 5 to 7 brown bands crossing back; lips with
alternating white and brown bands; a broad brown suborbital bar present; spinous dorsal fin with a large
black submarginal blotch, a series of small white and brown spots on spines; soft dorsal fin with alternating
white and brown spots on rays; anal fin pale; pectoral fins white below, with several small brown blotches
in middle and a series of blackish blotches above; pelvic fins whitish, with a prominent blackish blotch near
base and several small black blotches distally; caudal fin with rows of small white spots on rays separating
2 or 3 vertical dark bands.
Habitat, biology, and fisheries: Inhabits sand areas adjacent to coral reefs to a depth of about 38 m,
usually found at 18 m or less.
Distribution: Red Sea, In-
dian Ocean, Malaysia, Indo-
nesia, northern Australia;
Ryukyu Islands, Philippines,
and from the Mariana and
Marshall islands to the Mar-
quesa Islands.
Thysanophrys otaitensis (Parkinson, in Cuvier and Valenciennes, 1829)

FAO names: En - Fringelip flathead.
Diagnostic characters: Body elongate, head moderately depressed. Rear edge of maxilla reaches past
front edge of eye. Lips with a row of small fleshy papillae. Preopercular spines 2 or 3, upper longest,
extending about half-way to opercular margin, accessory spine absent. Supraorbital ridge smooth anteri-
orly, with stout serrations posteriorly; suborbital ridge bearing about 5 stout spines. Preorbital spine rarely
present; preocular spine 1. Teeth on vomer in 2 separate patches. Lower side of head bicarinate. Base of
lower opercular spine concealed by scales. Iris lappet bearing short branches with bifurcate tips.
Interopercular flap absent. Total gill rakers on first gill arch 5 to 7 (usually 6). Dorsal-fin spines IX, I,VIII,
or I,VII,I; dorsal-fin rays 11 or 12 (usually 11); anal-fin rays 11 or 12 (usually 12); pectoral-fin rays 19 to
22 (usually 20 or 21). Oblique scale rows slanting backward above lateral line about equal to number of
lateral-line scales. Lateral-line scales 50 to 54 (usually 52 or 53), anterior 1 to 3 with a small spine. Scale
pores of lateral line with 2 openings to the outside. Colour: back tan, mottled with white flecks or spots, 4
or 5 vague cross bands crossing back, undersides whitish; fins flecked with small brown spots, larger
blotches frequently present on pectoral, pelvic, and caudal fins.
Habitat, biology, and fisheries: Commonly taken over sand around coral reefs at depths of 10 m or less;
maximum depth to about 40 m.
D i s t r i b u t i o n : Widespread
from northern Natal, Mada-
gascar, Comoros, Chagos Ar-
chipelago, Indonesia, Taiwan
Province of China, southern
Japan, Philippines, Timor
Sea, Great Barrier Reef, Mar-
shall and Caroline islands to
the Marquesa and Tuamotu
islands.
click for next page

2422 Bony Fishes
HOPLICHTHYIDAE
Ghost flatheads (spiny flatheads)
by M.A. McGrouther
D iagnostic characters: Flattened, small to medium-sized (to about 18 cm standard length) scorpaeni-
form fishes; body elongate; mostly scaleless, with a row of spiny scutes along lateral line covering
much of back and upper half of sides. Head very wide and flat, much wider than deep, with variously
developed ridges and spines. Eyes on dorsal surface, longer than wide (eyes of larvae lateral, moving
dorsally with growth). Snout broad and greatly flattened. Mouth large. Teeth minute to villiform on jaws,
vomer, palatines, and pharyngeals. First dorsal fin short-based, with V or VI spines; second dorsal fin long
based, with 13 to 16 soft rays, some rays may be elongated in adult males; anal fin with I spine and 15 to
17 soft rays, about as long as second dorsal fin; caudal fin rounded to emarginate; pectoral fins with 11
to 14 upper rays and 3 or 4 detached lower rays; pelvic fins thoracic, with I spine and 5 soft rays. Colour:
most species yellow, pinkish, or brown above and on sides, with varying spots and mottling, fading to pink,
white, or silver below; pectoral fins yellowish brown to hyaline with various crossbars, spots, or mottling;
first dorsal fin often dark posteriorly, or with wavy crosslines; caudal fin usually hyaline, although in some
species with a dark distal margin or basal spots.
spiny scutes along lateral line

body mostly 13-16 dorsal-fin rays
scaleless
(after McCulloch, 1914)

3-4 detached pectoral-fin rays
Habitat, biology, and fisheries: Occur on the continental shelf and slope at depths of 60 to 1 500 m.
Larvae and juveniles collected in shallower water than adults. Feed on a range of crustaceans and fishes.
Of no use in commercial fisheries. Flesh supposedly of good quality but even the largest fish have very
little meat. Hoplichthys haswelli which occurs in southern Australian waters (outside the WCP area) grows
to 43 cm.
Remarks: Currently, 11 species of hoplichthyids are recognized worldwide and 8 of these occur within the
Western Central Pacific. The taxonomy of the family still requires some clarification (see footnote to species
list below).

Platycephalidae: superficially resemble hoplichthyids, but fully scaled on body, do not have detached
pectoral-fin rays, and no spiny scutes along lateral line.
Triglidae: fully scaled on body; less spinose on sides of head; head and body not or less flattened.
Platycephalidae Triglidae
Scorpaeniformes: Hoplichthyidae 2423
Callionymidae: also with strong preopercular spine, but lack

lateral head spination, have no body scutes, and no detached
pectoral-fin rays as found in hoplichthyids.
Callionymidae
Key to the species of Hoplichthyidae occurring in the area

1a. Lateral scutes each with 2 large spines of simi-
lar length (Fig. 1a) . . . . . . . . . . Hoplichthys langsdorfii
1b. Lateral scutes with 1 large spine and possibly 2 large
several tiny spines below it (Fig. 1b) . . . . . . . . . . . ® 2 1 large
spines spine
2a. Dorsal surface of head 2/3 to 1 eye length

behind eye with a strong elevated ridge, or
cluster of 2 or more stout spines (Fig. 2); more
than 1 strong spine usually present immediately
behind eye . . . . . . . . . . . . . . . . Hoplichthys ogilbyi a) Hoplichthys
langsdorfii b) other species
2b. Not as above, dorsal surface of head behind
each eye with small spines in rows, or with a Fig. 1 lateral scutes
single large spine, or without spines . . . . . . . . . . . ® 3 (along lateral line)
3a. Interorbital width greater than or equal to 1/3 elevated ridge or cluster of
eye length . . . . . . . . . . . . . . . . . . . . . . . . . ® 4 2 or more spines
3b. Interorbital width much less than 1/3 eye length . . . . . ® 6
4a. Interorbital width 1/3 to 1/2 eye length; pectoral-

fin rays 4 to 6 of juveniles elongated, often
crossed by 2 or 3 wavy bars . . . Hoplichthys filamentosus Fig. 2 Hoplichthys ogilbyi
4b. Interorbital width greater than eye length; pec-
toral-fin rays 4 to 6 not much longer than other
rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Dorsal profile of eyes partly

c o v e r e d b y o r b i t w h en
viewed from above (Fig. 3a);
head length 5 times eye
length . . . . . . . Hoplichthys fasciatus
(only known from juvenile specimens, most
likely a synonym of one of the other species)
5b. Dorsal profile of eyes not
covered by orbit (Fig. 3b);
head length 8 times eye
a) Hoplichthys fasciatus b) Hoplichthys pectoralis
length . . . . . . . Hoplichthys pectoralis
(only known from juvenile specimens, most Fig. 3 dorsal view of head
likely a synonym of one of the other species)
2424 Bony Fishes
6a. Lateral margin of head less spinose, weakly lobed, almost straight, with only a narrow
gap between the spine rows below the eye (Fig. 4a) . . . . . . . . . . . . . . . Hoplichthys gilberti
6b. Lateral margin of head strongly spinose, more strongly lobed, with a distinct gap between
the spine rows below the eye (Fig. 4b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
narrow gap
distinct gap
strong spines only posteriorly strong, curved spines
a) Hoplichthys gilberti b) Hoplichthys citrinus
Fig. 4 dorsal view of head
7a. Free rays of pectoral fins equal in length to longest joined rays; total anal-fin elements
(spines and soft rays) 16; fresh coloration unknown . . . . . . . . . . . . . . . . Hoplichthys regani
7b. Free rays of pectoral fins shorter than longest joined rays; total anal-fin elements 17;
body bright lemon yellow when fresh . . . . . . . . . . . . . . . . . . . . . . . Hoplichthys citrinus

Hoplichthys citrinus Gilbert, 1905
? Hoplichthys fasciatus Matsubara, 1937 1/
Hoplichthys filamentosus Matsubara and Ochiai, 1950
Hoplichthys gilberti Jordan and Richardson, 1908
Hoplichthys langsdorfii Cuvier and Valenciennes, 1829
Hoplichthys ogilbyi McCulloch, 1914
? Hoplichthys pectoralis (Fowler, 1943)1/
Hoplichthys regani Jordan, 1908
References
Matsubara, K. 1971. Fish morphology and hierarchy. Part II. Ishizaki - Shoten. Japan, pp. 791-1605 (relevant pages
for Hoplichthyidae: 1123-1130).
Matsubara, K. 1971. Fish morphology and hierarchy. Part III (Plates). Ishizaki -Shoten. Japan, pls 1-135 (relevant plate
for Hoplichthyidae pl. 125).
Matsubara, K. and A. Ochiai. 1950. Studies on Hoplichthyidae, a family of mail-cheeked fishes, found in Japan and its
adjacent waters. II. Japan. J. Ichthyol., 1(2):82-88.
1/ Described from juveniles (about 7 cm) and show the juvenile character of a very wide interorbit. These species are
quite likely conspecific with one of the other species in the area, but because of this uncertainty both have been included
in the key above.
Scorpaeniformes: Cottidae 2425
COTTIDAE
Sculpins
by J.S. Nelson and K.E. Carpenter
D iagnostic characters (for the 2 species occurring in the area): Small (to about 51 mm standard
length) moderately elongate scorpaeniform fishes; body depth 15.8 to 20.7% of standard length. Head
large, 38.5 to 41.2% of standard length. Eye width much longer than snout length, orbit length around
2 times snout length; snout length 7.25 to 8.3% of standard length. Long supraorbital cirrus present
(possibly absent in Antipodocottus mesembrinus). Preopercle with 4 posteriorly-directed spines, the
uppermost antler-like, with 3 to 5 secondary projections. Villiform teeth in jaws, vomer, and palatines.
Two dorsal fins, the first with VIII spines, the second with 12 to 14 soft rays. Anal fin with 10 to 12 soft
rays. Caudal fin slightly rounded. Pectoral fins with 17 to 20 soft rays. Pelvic fins thoracic, with I spine
and 2 soft rays. Two rows of modified bony scales on sides, the uppermost directly beneath base of
second dorsal fin, the second along the lateral line (careful examination required). Colour: live coloration
unknown; in alcohol, body brownish or reddish brown, lighter ventrally, sides and head with irregularly
spaced darker spots and blotches, sometimes a dark blotch at base of lower caudal-fin rays; dorsal fin with
irregular dark markings, sometimes with distal end distinctly darker on spinous dorsal fin.
2 rows of bony
4 spines on preopercle, the scales on side
uppermost antler-like
(after Fricke and Brunken, 1983)

pelvic fins thoracic, with
I spine and 2 soft rays
Habitat, biology, and fisheries: Deep water demersal, at depths between 150 and 765 m. Very little
known about the biology of the species occurring in the area. No interest to fisheries. Antipodocottus
elegans is known only from southernmost Queensland and Tasmania while A. mesembrinus is known only
from off Indonesian New Guinea.
Remarks: Both the generic placement and specific status of the 2 species recorded from the area are
questionable.

Bembridae (compared to Cottidae in the area): head depressed (not depressed in Cottidae); snout length
greater than or nearly equal to eye diameter (snout length much smaller than eye diameter in Cottidae).
Psychrolutidae (compared to Cottidae in the area): a single dorsal fin (2 dorsal fins in Cottidae); no
preopercular spines (4 in Cottidae, the uppermost prominent and antler-like); no scales along lateral line
(some scales in Cottidae).
Bembridae Psychrolutidae
2426 Bony Fishes
Scorpaenidae (compared to Cottidae in the area): a single dorsal fin (2 dorsal fins in Cottidae).
Callionymidae (compared to Cottidae in the area): the cottids in the area superficially look like callionymids,
however they can be distinguished by very restricted gill openings (broad gill openings in Cottidae); up to
IV dorsal-fin spines (VIII in Cottidae); 1 stout preopercular spine (4 in Cottidae); pelvic fins subjugular
(thoracic in Cottidae) with I spine and 5 soft rays (I spine and 2 soft rays in Cottidae); body scaleless (scales
present in Cottidae).
Scorpaenidae Callionymidae

Antipodocottus elegans Fricke and Brunken, 1984
Antipodocottus mesembrinus (Fricke and Brunken, 1983)
References
Fricke, R. and H. Brunken. 1984. A new cottid fish of the genus Antipodocottus (Teleostei: Scorpaeniformes) from
eastern Australia, with a key to the species of the genera Stlengis and Antipodocottus. J. Nat. Hist., 18:41-46.
Nelson, J.S. 1990. Redescription of Antipodocottus elegans (Scorpaeniformes: Cottidae) from Australia, with comments
on the genus. Copeia, 1990(3):840-846.
Scorpaeniformes: Psychrolutidae 2427
PSYCHROLUTIDAE
Fathead sculpins
by J.S. Nelson
D iagnostic characters: Small to moderate fishes (to about 65 cm standard length, commonly under
35 cm); body moderately elongate, tadpole-shaped, stout and robust in cross-section behind head,
tapering and compressed posteriorly. Head very large, smooth, or with protruding spines or knobs, cirri
sometimes present. Spines absent from preopercle in species in the area. Interorbital space wide,
greater than exposed eye diameter in species in the area. Eye large. Mouth large, terminal, or nearly
so. Teeth conical, generally reduced; premaxillae and dentary with several rows, prevomer with or without
teeth; palatine without teeth. Branchiostegal rays 7. Gill rakers short spiny knobs. Dorsal fin continuous in
species in the area. Dorsal fin with VI to XII slender weak spines and 12 to 21 soft rays; anal fin with 9 to
17 soft rays; caudal fin rounded to truncate; pelvic fins (thoracic, closely spaced, small, and slender)
with I spine and 3 soft rays; pectoral fins with 15 to 26 (usually 19 to 26) rays. Skin loose in most
species, covering dorsal and anal fins in most, making it difficult to count rays. Body naked or variously
with bony plates or prickles (cirri may be present or absent). Lateral line on trunk reduced, with 20
or fewer small pores. Bone covering cranial sensory canals in interorbital and suborbital areas
reduced to well-developed but relatively narrow bony arches. Postorbital bones 1 or 2. Vertebrae about
28 to 38. Colour: uniformly brown to grey or with irregular markings or spotted pattern of black, grey,
brown, or white.
Habitat, biology, and fisheries: Mostly benthic, from inshore shallow water to depths of 2 800 m.
Occurring primarily in cool waters of the North Pacific and North Atlantic and scattered parts of the southern
hemisphere (e.g. off South America, South Africa, Australia, and New Zealand). Little is known of their
biology; some have been found with gastropods in the stomach. Trawled incidentally, of no commercial
value.
Remarks: Fathead sculpins are poorly represented in the area. The status of M. Weber’s Cottunculus
gyrinoides from the Flores Sea (south of Sulawesi) is uncertain. The related family Cottidae is also known
in the area from 2 species of Antipodocottus.

Aphyonidae, Batrachoididae, Bythitidae, and
Ophidiidae: distinguished from Psychrolutidae in
having pelvic fins in front of pectoral fins.
Ophidiidae
2428 Bony Fishes
Liparidae (known only from nearby WCP area): distinguished from Psychrolutidae in having pelvic fins
modified into a sucking disc or in lacking these fins.
Cottidae: the 2 species occurring within and adjacent to area, Antipodocottus mesembrinus from Indonesia
and A. elegans from off eastern Australia, differ from Psychrolutidae in having dorsal fins distinct and
separated, some scales along lateral line, and 4 preopercular spines, uppermost especially prominant and
antler-like.
Liparidae Cottidae
Key to the species of Psychrolutidae occurring in or near the area

1a. Vomerine teeth present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ebinania spp.
1b. Vomerine teeth absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Pectoral-fin rays usually 24 or 25; body uniform light grey; peritoneum jet black . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psychrolutes marcidus
2b. Pectoral-fin rays 23 or fewer; body variously brownish; peritoneum may be darkish but
not jet black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Cirri absent from head and body; total dorsal-fin rays 22 or 23 . . . . . . . Psychrolutes occidentalis
3b. Cirri sparse but present; total dorsal-fin rays usually 24 to 26 . . . . . . . . . . . . . . . . . . . ® 4
4a. Median chin pore single . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Psychrolutes inermis

4b. Median chin pore paired . . . . . . . . . . . . . . . . . . . . . . . . . Psychrolutes macrocephalus

Note: the taxa listed are known or reported from the area or nearby; additional species occur in, for
example, New Zealand.
Ebinania spp.
Psychrolutes inermis (Vaillant, 1888) or P. macrocephalus (Gilchrist, 1904)
Psychrolutes marcidus (McCulloch, 1926)
Psychrolutes occidentalis Fricke, 1990
References
Jackson, K.L. and J.S. Nelson. 1998. Ambophthalmos, a new genus for “Neophrynichthys” angustus an d
“Neophrynichthys” magnicirrus , and the systematic interrelationships of the fathead sculpins (Cottoidei,
Psychrolutidae). Can. J. Zool., 76:in press.
Nelson, J.S. 1995. Psychrolutes microporos, a new species of cottoid (Teleostei: Scorpaeniformes) from New Zealand
and Japan with biogeographical comments. Proc. Zool. Soc., Calcutta, 48(2):67-76.
click for next page

Perciformes: Percoidei: Centropomidae 2429
Order PERCIFORMES
Suborder PERCOIDEI
CENTROPOMIDAE
Sea perches
by H.K. Larson
D iagnostic characters: Elongate, compressed, medium to large (to 2 cm) percoid fishes with dorsal
profile behind eyes concave or convex. Eyes medium sized, relatively close to tip of snout 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, reaching at least to below eyes. Teeth small, in villiform bands
on upper and lower jaws, vomer, and palatines (may be present on tongue). Branchiostegal rays 7. First
gill arch with 3 to 7 gill rakers on upper limb, 9 to 14 on lower limb. Dorsal fin deeply incised before last
dorsal-fin spine, or with distinct gap between spiny and soft portions of fin. Caudal fin rounded. Dorsal
fin with VII to IX strong spines and 10 to 14 soft rays; anal fin with III spines and 7 to 9 soft rays; pelvic
fins with axillary scale, and I spine and 5 soft rays; pectoral fins with 16 or 17 rays and spiny flap exposed
just above fin base. Scales large, ctenoid; scale rows on body running horizontally; bases of caudal, soft
dorsal, and anal fins covered with scales. Lateral-line scales 45 to 50; lateral line extends onto caudal fin,
nearly to rear margin, in 1 or 3 series. Vertebrae 11+14=25. Three predorsal bones present. Five hypurals
(all separate), 2 epurals, and 1 or 2 uroneurals. Swimbladder with strong ligament running anterodorsally
to posttemporal. Supraoccipital extended forward, separating posterior portions of frontals. Colour: adults
greenish or silvery grey to brown, juveniles with stripes; eyes with characteristic red reflection.
dorsal fin deeply incised
lateral line
extending onto
dorsal profile behind caudal fin
eye concave
preopercle serrated, a
stout flat spine at angle
Habitat, biology, and fisheries: Contains 2 subfamilies, the Centropominae, with 9 species, and Latinae,
with 9 species. Coastal marine and estuarine to fresh water, inhabiting mangrove estuaries and rocky to coral
reefs. Feed on crustaceans (mostly prawns) and fishes. Solitary or in small groups. Lates calcarifer move up
into brackish or fresh water during parts of their life cycle; other centropomids remain in the sea or fresh water.
Popular and sought-after fishes, with some species of very considerable economic importance.
Serranidae: preopercle smooth or serrated, usually with-
out large flat spines or serrations; opercle with 3 flat
spines; teeth conical, pointed, in rows; lateral line not
extending onto caudal fin; dorsal fin single and not divided
in most species. Most similar species in the area: Niphon
spinosus.
Moronidae (Lateolabrax japonicus, occurs near the
northern part of the area): usually forked caudal fin; oper-
cle with 2 spines on upper part; preopercle serrated,
several large flat spines along lower edge; pelvic fins Serranidae (Niphon spinosus)
without axillary scale; lateral line does not extend onto
caudal fin.
2430 Bony Fishes
Key to the species of Centropomidae occurring in the area

1a. Jaws reach back to rear of eyes (Fig. 1); lower edge of preopercle with 3 or 4 large flat
spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lates calcarifer
1b. Jaws reach back to under eyes (Fig. 2); lower edge of preopercle smooth, without
spines, only 1 spine present at angle . . . . . . . . . . . . . . . . . . . . . Psammoperca vaigiensis
a single
preopercular
3-4 spine
preopercular
spines
Fig. 1 Lates calcarifer Fig. 2 Psammoperca vaigiensis

0 Lates calcarifer (Bloch, 1790)
0 Psammoperca vaigiensis (Cuvier, 1828)
References
Copland, J.W. and D.L. Grey (eds). 1987. Management of wild and cultured sea bass/barramundi (Lates calcarifer).
Canberra, Australian Centre for International Agricultural Research. ACIAR Proceedings , (20):210 p.
Greenwood, P.H. 1976. A review of the family Centropomidae (Pisces, Percifomes). Bull. Brit. Mus. (Natl. Hist.) Zool.,
29(1):1-81.
Perciformes: Percoidei: Centropomidae 2431
Lates calcarifer (Bloch, 1790) GIP

Frequent synonyms / misidentifications: None / Lateolabrax japonicus (Cuvier 1828) (family Moronidae).
FAO names: En - Barramundi (= giant seaperch, Fishing Areas 57/72); Fr - Barramundi; Sp - Barramundi.
(after Kailola et al., 1993)
Diagnostic characters: Body moderately deep, elongate, and compressed; caudal peduncle distinct. Dorsal
profile concave anteriorly, convex in front of dorsal fin; snout and jaws pointed. Lower edge of preopercle
with 3 or 4 (rarely more) large flat triangular spines. Nostrils on each side of head close together near
eye. Mouth large, jaws reaching past eyes. Lower limb of first gill arch with 16 or 17 gill rakers. Dorsal fin
deeply incised before last (small) dorsal-fin spine, the fin with VII to IX spines and 10 or 11 soft rays. Anal
fin with III spines and 7 or 8 soft rays. Caudal fin rounded. Scales firmly fixed, ctenoid. Lateral line extends
onto caudal fin. Low scaly sheath at base of soft dorsal and anal fins. Colour: adults silver with olive-grey or
grey-blue backs (fish from turbid waters are darker and duller); juveniles brown to greyish brown with 3 white
stripes on head and scattered white patches on sides (white markings can be intensified or turned off at will);
eyes brown to golden, with bright red reflective glow; fins without markings.
Size: Maximum total length over 2 m; commonly to 1.5 m and over 55 kg.
Habitat, biology and fisheries: Inhabits coastal marine and estuarine to fresh-water habitats; adults mainly
in estuaries (mangroves and river mouths), younger fish extending up rivers into fresh water. Adults return to
estuaries to breed, forming spawning aggregations. Barramundi change sex during their life cycle, with most
fish maturing first as functional males then becoming females after 3 to 5 years of age. Feed on fishes and
some crustaceans (mostly prawns). Popular and sought-after fishes of very considerable economic impor-
tance. From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports a range of yearly catch of around
20 200 to 42 800 t of Lates calcarifer from the Western Central Pacific. Gill nets mostly used, but recreational
fishery (rod-and-line) becoming inceasingly important. Farming presently carried out, for example in Thailand,
Indonesia, and Australia. Marketed mostly fresh.
Distribution: Widely distrib-
uted in coastal areas of the
Indo-West Pacific, from the
eastern edge of the Persian
Gulf to China and southern Ja-
pan, and southwards to north-
ern Australia and southern
Papua New Guinea.
2432 Bony Fishes
Psammoperca vaigiensis (Cuvier, 1828)

FAO names: En - Waigieu seaperch; Fr - Brochete de mer; Sp - Perca de mar.
Diagnostic characters: Body moderately deep, elongate, and compressed; caudal peduncle distinct. Dorsal
profile concave anteriorly, convex in front of dorsal fin; snout and jaws pointed, snout tip slightly rounded.
Lower edge of preopercle smooth, 1 large flat spine at rear angle of preopercle. Nostrils on each side
of head widely separated. Mouth moderate, jaws reaching to below middle of eyes. Lower limb of first
gill arch with 12 to 14 gill rakers. Dorsal fin deeply incised before last (small) dorsal-fin spine, the fin with
VII or VIII spines and 12 to 14 soft rays. Anal fin with III spines and 8 or 9 soft rays. Caudal fin rounded.
Scales firmly fixed, ctenoid. Lateral line extends onto caudal fin. Low scaly sheath at base of soft dorsal and
anal fins. Colour: light silvery grey to dark brown, usually coppery brown, paler ventrally, darker brown lines
may follow scale rows along length of body; eyes golden-brown with distinct red reflection; fins unmarked.
Size: Maximum total length about 47 cm; commonly between 20 and 30 cm.
Habitat, biology, and fisheries: Coastal, marine, found on shallow rocky or coral reefs especially near algal
beds (such as Sargassum). Secretive during the day, hiding in crevices or under overhangs. Hunts at night
for small fishes and crustaceans. No fishery statistics available. Fished mostly with handlines, rod-and-line,
or gill nets. Marketed mostly fresh.
Distribution: Distributed in
the tropical east Indo-West
Pacific, from Bay of Bengal,
Indo-Australian Archipelago
a nd n or t he r n A us tr a l i a,
Philippines, Japan, and the
China Sea.
Perciformes: Percoidei: Ambassidae 2433
AMBASSIDAE
(= Chandidae)
Perchlets, glassfishes
by G.R. Allen
D iagnostic characters: Small (to 12 cm) percoid fishes; body oblong to oval-shaped and compressed.
Eyes large, much greater than snout length. Margins and ridges of preorbital, suborbital, supraor-
bital, preopercle, and interopercle frequently serrate. Mouth moderately large, only slightly protractile,
jaws equal or lower one slightly protruding; angle of jaw oblique, about 40° to horizontal. Bands of villiform
teeth on jaws, vomer, and palatines. First gill arch with 16 to 29 gill rakers on lower limb. A deeply notched
dorsal fin, the front portion of fin with VII spines, the rear portion with I spine and 8 to 11 soft rays. Anal
fin with III spines and 8 to 11 soft rays. Caudal fin forked. Pelvic fins with I spine and 5 soft rays. Pectoral
fins with 11 to 17 rays. Branchiostegal rays 6. Scales large and cycloid, extending onto head and base of
median fins; cheeks and operculum scaly; lateral line continuous or interrupted in middle portion; scales
in midlateral series 24 to 34. Colour: semitransparent, usually with dark scale outlines and silvery sheen
on side of head and belly.
single dorsal fin,

deeply notched
often with serrated

ridges on head III anal-fin
spines
Habitat, biology, and fisheries: Mangrove shores, brackish estuaries, and fresh waters, always in shallow
depths. Forms resting aggregations during the day among the roots of mangrove trees, log snags, and
aquatic plants. They disperse at night to feed on micro-crustaceans (cladocerans, ostracods, and cope-
pods), aquatic and terrestrial insects, and occasional fishes. Too small to be commercially important,
although they are sometimes dried and salted; also used as bait fishes.

Apogonidae: dorsal fin consisting of 2 completely
separate parts; only II anal-fin spines.
2 anal-fin spines
Apogonidae
2434 Bony Fishes
Key to marine and estuarine species of Ambassidae occurring in the area

Remarks on key characters: the serrated margins and ridges on various head bones are useful features
for identifying ambassid fishes (Fig. 1).
1a. Supraorbital spines usually 3 to 5 preorbital ridge
(rarely 2); nasal spine well developed . . . . . ® 2
supraorbital ridge
1b. Single suborbital spine; nasal spine nasal
well developed or absent . . . . . . . . . . ® 4 spine
preorbital
edge hind margin of
2a. Hind margin of preopercle (i.e. ver- preopercle
tical limb) with about 6 to 13 small
preopercular
serrae (Fig. 2) . . . . . . . . Ambassis vachellii ridge
(Indo-Australian Archipelago) interopercle
lower edge
2b. Hind margin of preopercle usually of preopercle suborbital
smooth or weakly crenate without
distinct serrae (Figs 3 and 4) . . . . . . . . ® 3 Fig. 1 serrated ridges and edges of head
bones used in the identification key
3a. Soft anal-fin rays usually 10 (rarely 11); predorsal scales 16 to 18; eye relatively small, 10.7
to 12.7% of standard length; caudal peduncle relatively short and deep, its length and depth
16.1 to 20.8% and 14.4 to 16.3% of standard length, respectively (Fig. 3) . . . . . . . Ambassis marianus
(southern Queensland and northern New South Wales, Australia)
3b. Soft anal-fin rays usually 9; predorsal scales 11 to 14; eye larger, 13 to 13.9% of standard
length; caudal peduncle more slender, its length and depth 20.6 to 22.2% and 13 to 14.8%
of standard length, respectively (Fig. 4) . . . . . . . . . . . . . . . . . . . Ambassis gymnocephalus
(Indo-West Pacific)
6-13 small
serrae
Fig. 2 Ambassis vachellii Fig. 3 Ambassis marianus Fig. 4 Ambassis gymnocephalus

4a. Cheek with 1 row of scales (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . Ambassis urotaenia
(Indo-Australian Archipelago)
4b. Cheek with 2 or more scale rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Lateral line continuous from upper edge of gill opening to caudal-fin base . . . . . . . . . . . . . ® 6
5b. Lateral line interrupted in middle portion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
6a. Pectoral-fin rays usually 16 or 17 (rarely 15); nasal spine absent (Fig. 6); body relatively
deep, maximum depth 45.3 to 50.6% of standard length . . . . . . . . . . . . . . . Ambassis nalua
(India to New Guinea and Australia)
6b. Pectoral-fin rays 13 to 15; nasal spine present, but may be blunt and hidden under skin;
body more slender, maximum depth 29.2 to 44.7% of standard length . . . . . . . . . . . . . . . ® 7
1 row of
scales here
Fig. 5 Ambassis urotaenia Fig. 6 Ambassis nalua

Perciformes: Percoidei: Ambassidae 2435
7a. Dorsal-fin spines relatively weak and flexible; soft dorsal-fin rays usually 10 (occasionally
9 or 11); soft anal-fin rays usually 8 or 9; body slender, maximum depth 29.2 to 33.2% of
standard length; head length 32 to 34% of standard length (Fig. 7) . . . . . . Ambassis jacksoniensis
(New South Wales and southern Queensland, Australia)
7b. Dorsal-fin spines relatively strong and stiff; soft dorsal-fin rays usually 9 (rarely 10); soft
anal-fin rays usually 9 or 10; body deeper, maximum depth 33.4 to 44.7% of standard
length; head length 35.5 to 41.4% of standard length . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Predorsal scales 17 to 22; horizontal scale rows from anal-fin origin to base of dorsal fin
12 or 13 (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ambassis macracanthus
(Sumatra to New Guinea)
8b. Predorsal scales 8 to 15; horizontal scale rows from anal-fin origin to base of dorsal fin 9
to 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Predorsal scales 8 to 11; second dorsal-fin spine longer than third spine; horizontal scale
row from anal-fin origin to base of dorsal fin 11 or 12 . . . . . . . . . . . . . . . . . . Ambassis kopsi
(Malay Peninsula, Kalimantan Sabah, and the Philippines)
9b. Predorsal scales 12 to 15; second dorsal-fin spine slightly shorter than third spine;
horizontal scale row from anal-fin origin to base of dorsal fin 9 or 10 (Fig. 9) . . . . . Ambassis miops
(India to Australia and New Guinea)
Fig. 7 Ambassis jacksoniensis Fig. 8 Ambassis macracanthus Fig. 9 Ambassis miops

10a. Margin of interopercle with 2
to 10 small serrae (Fig. 10);
height of spinous dorsal fin
34 to 3 8 % o f s tandard
length; maximum depth of
body 37.2 to 47.8% of stand- 2-10 small
ard length . . . . . Ambassis interruptus serrae
(Andaman Islands to Vanuatu and New Caledonia) Fig. 10 Ambassis interruptus Fig. 11 Ambassis buruensis
10b. Margin of interopercle
smooth (Fig. 11); height of spinous dorsal fin 27.6 to 32.7% of standard length; maximum
depth of body 32.9 to 39.5% of standard length . . . . . . . . . . . . . . . . . . Ambassis buruensis
(Sumatra to New Guinea and the Philippines)
List of marine-estuarine species occurring in the area
Ambassis buruensis Bleeker, 1857
Ambassis gymnocephalus (Lacepède, 1802)
Ambassis interruptus Bleeker, 1852
Ambassis jacksoniensis (Macleay, 1881)
Ambassis kopsi Bleeker, 1858
Ambassis macracanthus Bleeker, 1849)
Ambassis marianus Günther, 1880
Ambassis miops Günther, 1871
Ambassis nalua (Hamilton, 1822)
Ambassis urotaenia Bleeker, 1852
Ambassis vachellii Richardson, 1846
References
Allen, G.R. and W.E. Burgess. 1990. A review of the glassfishes (Chandidae) of Australia and New Guinea. Rec. West.
Aust. Mus. Supplement, 34:139-206.
Fraser-Brunner, A. 1954. A synopsis of the centropomid fishes of the subfamily Chandinae, with descriptions of a new
genus and two new species. Bull. Raffles Mus., 25:185-213.
2436 Bony Fishes
ACROPOMATIDAE
Temperate ocean-basses (lanternbellies, splitfins)
by K.E. Carpenter
D iagnostic characters: Body moderately compressed (size to about 40 cm). Eyes large, their diameter
greater than snout length. Usually 2, sometimes 3 flat spines on opercle. Mouth subterminal, the
lower jaw slightly projecting; small teeth present on jaws and usually on vomer and palatines; usually canine
teeth in jaws; maxillae not covered by preorbitals, not scaly, and broadened distally. Gill membranes free
from isthmus. Branchiostegal rays 7. Separate spiny and soft dorsal fins, or dorsal fin notched to base
before last spine; first dorsal fin with VII to X spines, and second fin with 0 to I spines and 8 to 10
soft rays. Anal fin with II or III spines and 6 to 8 soft rays. Caudal fin usually forked, sometimes deeply
forked or emarginate. Vertebrae with 10 precaudal and 16 caudal elements. Colour: body either pink or
reddish and whitish or silvery ventrally, silvery, or mostly brownish or blackish; the 2 species of Acropoma
have a light organ and their anus situated near pelvic-fin base.
2 dorsal fins, or a single dorsal fin notched nearly to base
before last spine, with VII-X spines and 9-10 soft rays
2-3 flat
opercular spines
lower jaw usually

with distinct,
spaced canines anal fin with II-III spines
and 6-8 soft rays
Habitat, biology, and fisheries: Demersal, generally over soft bottom in deep water, found at depths
between 20 and 700 m. Incidentally caught in deep bottom trawls and generally too small or not abundant
enough to be exploited commercially; occasionally consumed, some are considered food fish in Japan.
Remarks: The relationships and affinities of the Acropomatidae have been very much confused. Members
of this family have variously been placed in the Percichthyidae, Apogonidae, Howellidae, and Polyprioni-
dae. The Percichthyidae is now restricted to temperate fresh-water perches and the marine forms put in
the Acropomatidae. A species included here under Howella is provisionally placed here and is perhaps
better treated as a separate family (although sometimes also included under the Apogonidae). The species
of Polyprion which apparently are restricted to temperate waters outside the WCP area, are also perhaps
better treated in the separate family, Polyprionidae.
Acropomatids are distinguishable from most other similar percoid fishes (including the Serranidae, their
presumed close relatives) by their divided or nearly divided spinous and soft dorsal fins. Other similar-
shaped families with this dorsal-fin configuration include:
Apogonidae: anal fin with 8 to 18 soft rays (6 to 8 in Acropomatidae); a single opercular spine (2 or 3 in
Acropomatidae).
Epigonidae: anal fin with I to III spines (II or III in Acropo-
matidae); opercle with 1 or 3 spines (2 or 3 in Acropomati-
dae); maxilla narrow (tip of maxilla broad in Acropomatidae).
1-3 opercular spines
8-18 anal-fin rays

anal fin with
I-III spines
Epigonidae Apogonidae
Perciformes: Percoidei: Acropomatidae 2437
nd
2 dorsal fin with I spine
Lactariidae: second dorsal fin with I spine and 19 to 23 soft and 19-23 soft rays
rays (0 to I spines and 8 to 10 soft rays in Acropomatidae);
anal fin with 25 to 28 soft rays (6 to 8 soft rays in Acropoma-
tidae).
Scombropidae: second dorsal fin with I spine and 11 to 14
soft rays (0 to I spines and 8 to 10 soft rays in Acropomati-
dae); anal fin with 11 to 13 soft rays (6 to 8 soft rays in
Acropomatidae); maxilla scaly (not scaly in Acropomatidae).
25-28 anal-fin rays
Emmelichthyidae: dorsal fin with XI to XIII spines and 10 to
12 soft rays (VII to X spines and 8 to 10 soft rays in Lactaridae
Acropomatidae); anal fin with 9 or 10 soft rays (6 to 8 soft
rays in Acropomatidae); broad, scaly maxilla (not scaly in
Acropomatidae). nd
2 dorsal fin with I spine dorsal fin with XI-XIII spines
and 11-14 soft rays and 10-12 soft rays
maxilla scaly maxilla scaly

11-13 anal-fin rays 9-10 anal-fin rays
Scombropidae Emmelichthyidae
Acropoma japonica Günther, 1859
Acropoma lecorneti Fourmanoir, 1988
Apogonops anomalus Ogilby, 1896
Bathysphyraenops simplex Parr, 19331/
Doederleinia berycoides (Hilgendorf, 1879)
Doederleinia gracilispinis Fowler, 1943
Howella brodiei Ogilby, 1899
Malakichthys elegans Döderlein, 1883
Malakichthys sp.
Neoscombrops pacificus Mochizuki, 1979
Pseudohowella intermedia Fedoryako, 1976
Synagrops analis Katayama, 1957
Synagrops argyrea (Gilbert and Cramer, 1896)
Synagrops japonicus (Döderlein, 1884)
Synagrops malayanus Weber, 1913
Synagrops philippinensis Günther, (1880)
Synagrops serratospinosa Smith and Radcliffe, 1912
References
Gloerfelt-Tarp, T. and P.J. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Australia. Jakarta,
Dir. Gen. Fish. (Indonesia), German Tech. Coop., Aust. Dev. Assoc. Bur., 406 p.
Heemstra, P.C. 1986. Family Acropomatidae. In Smiths’ sea fishes, edited by M.M. Smith and P.C. Heemstra.
Mochizuki, K. 1984. Family Percichthyidae. In Fishes of the Japanese Archipelago , edited by H. Masuda, K. Amaoka,
C. Araga, T. Uyeno, and T. Yoshino. Tokai Univ. Press., pp. 123-126.
Paxton, J.R. and J.E. Hanley. 1989. Family Percichthyidae. In Zoological catalogue of Australia 7. Pisces.
Petromyzontidae to Carangidae , edited by J.R. Paxton, D.F. Hoese, G.R. Allen, and J.E. Hanley. Canberra,
Australian Government Publishing Service, 665 p.
1/ Listed here provisionally, most recently considered ‘incertae sedis’ in the Percoidei.
2438 Bony Fishes
SYMPHYSANODONTIDAE
Bunquelovelies (also wampeejawed fishes, slopefishes, and shelf beauties)
by W.D. Anderson, Jr.
D iagnostic characters: Body slender to moderately deep, somewhat compressed (size to about
17 cm). Head moderate. Eyes moderate to large, their diameter about equal to snout length to
considerably longer than snout. Suborbital extremely narrow, its height (width) about 1% of standard
length. Opercular spines 2. Snout relatively blunt. Anterior ends of premaxillae incised, forming
conspicuous symphysial notch that receives anterior ends of dentaries. Mouth terminal and oblique;
jaws about equal. Extreme dorsalmost margin of maxilla covered by very narrow suborbital with
mouth closed. Premaxilla with small teeth (usually larger anteriorly); symphysial notch toothless. Dentary
with small teeth usually extending from posterior elevation of the bone almost to symphysis; teeth on and
near posterior elevation usually larger; usually a number of relatively large exserted teeth at anterior
end of dentary, these teeth fitting into symphysial notch in premaxillae when mouth closed. Teeth,
when present, small on vomer, palatines, and pterygoids; no teeth on tongue. Branchiostegal rays 7. First
gill arch with 9 to 14 gill rakers on upper limb and 20 to 29 on lower limb (total 29 to 42). Dorsal fin not
incised at junction of spinous and soft rays. Caudal fin deeply forked. Both lobes of caudal fin and pelvic
fins extremely produced in some individuals. Dorsal fin usually with IX spines and 10 soft rays. Anal
fin with III spines and 7 or 8 soft rays. Principal caudal-fin rays 17 (9 in upper lobe, 8 in lower lobe); branched
caudal-fin rays 15 (8 in upper lobe, 7 in lower lobe). Pectoral-fin rays 15 to 18 (usually 16 or 17). Pelvic
fins with I spine and 5 soft rays. Dorsal and anal fins without scales, but with scaly sheaths at their bases.
Axillary scales of pelvic fins and scaly interpelvic process well developed. Most of head, including
maxillae and dentaries, covered with scales. Scales moderate, ctenoid. Tubed lateral-line scales 42 to
61. Vertebrae 10+15=25. Colour: mainly shades of red or orange.
single dorsal fin, usually with

IX spines and 10 soft rays
2 opercular spines
caudal fin
deeply
forked
(after Anderson, 1970)
Habitat, biology, and fisheries: Bottom-associated fishes, known from depths of 50 to 500 m on the
continental shelf and upper continental slope, around islands, and over reefs and submarine ridges.
Probably planktivorous.
Remarks: Have been considered by various workers to be members of either the family Serranidae or the
family Lutjanidae, but species of Symphysanodontidae possess characters that clearly distinguish them
from serranids and lutjanids and lack characters that would associate them with either of those groups of
fishes.
Perciformes: Percoidei: Symphysanodontidae 2439

Serranidae: 3 opercular spines (2 in Symphysanodontidae); dorsalmost margin of maxilla not covered by
suborbital when mouth closed; vertebrae rarely 25, usually 24 or 26 (25 in Symphysanodontidae).
Lutjanidae: maxilla covered to considerable degree by suborbital when mouth closed; anterior ends of
premaxillae not incised to form conspicuous symphysial notch that receives anterior ends of dentaries
when mouth closed; X to XII dorsal-fin spines (almost always IX dorsal-fin spines in Symphysanodontidae);
vertebrae 24 (25 in Symphysanodontidae).
3 opercular
spines
Serranidae (subfamily Anthiinae) Lutjanidae
Key to the species of Symphysanodontidae occurring in the area

Remarks on key characters: counts of lateral-line scales are of tubed scales along the body, excluding
those posterior to the base of the caudal fin. Counts of gill rakers include rudiments, when present.
1a. Lateral-line scales 42 to 49; first gill arch with 9 to 11 gill rakers on upper limb and 20 to
26 on lower limb (total 29 to 37); length of pelvic fins 22 to more than 80% of standard
length; length of upper caudal-fin lobe 29 to 75% of standard length; length of lower
caudal-fin lobe 29 to 76% of standard length (individuals with well-produced to extremely
produced caudal-fin lobes and pelvic fins are probably males) . . . . . . Symphysanodon maunaloae
1b. Lateral-line scales 52 to 55; first gill arch with 10 to 12 gill rakers on upper limb and 25
to 28 on lower limb (total 36 to 40); length of pelvic fins 22 to 26% of standard length;
both lobes of caudal fin produced slightly, but apparently never extremely produced, as
in some individuals of S. maunaloae . . . . . . . . . . . . . . . . . . . . . . . Symphysanodon typus

The symbol 0
0
0
Symphysanodon maunaloae Anderson, 1970
Symphysanodon typus Bleeker, 1878
Reference
Anderson, W.D., Jr. 1970. Revision of the genus Symphysanodon (Pisces: Lutjanidae) with descriptions of four new
species. Fish. Bull., 68(2):325-346.
2440 Bony Fishes
Symphysanodon maunaloae Anderson, 1970

Frequent synonyms / misidentifications: None / Symphysanodon typus Bleeker, 1878.
FAO names: En - Beautimous wampeejaw.
Diagnostic characters: Body slender to moderately deep, its depth 21 to 33% of standard length (depth
tending to become relatively greater with increase in standard length). First gill arch with 9 to 11 gill rakers
on upper limb and 20 to 26 on lower limb (total 29 to 37). Caudal fin deeply forked, both lobes produced;
length of upper caudal-fin lobe 29 to 75% of standard length; length of lower caudal-fin lobe 29 to
76% of standard length. Length of pelvic fins 22 to more than 80% of standard length. (Individuals
with well-produced to extremely produced caudal-fin lobes and pelvic fins are probably males.) Lateral-line
scales 42 to 49. Colour: upper body and caudal fin light red; iris pink; dark bar between eyes; dark vertical
bar just behind operculum.
Size: Maximum standard length about 16 cm; commonly to 9 cm.
Habitat, biology, and fisheries: Occurs in depths of 150 to 500 m. No other information available.
Distribution: Wide ranging
from the eastern South Pacific
(Sala y Gómez Ridge) and
Hawaii to Indonesia (off Lom-
bok) and the Kyushu-Palau
Ridge.
Perciformes: Percoidei: Symphysanodontidae 2441
Symphysanodon typus Bleeker, 1878

FAO names: En - Aristocratic bunquelovely.
Diagnostic characters: Body slender, its depth 22 to 29% of standard length (depth tending to become
relatively greater with increase in standard length). First gill arch with 10 to 12 gill rakers on upper limb
and 25 to 28 on lower limb (total 36 to 40). Caudal fin deeply forked, both lobes produced, but apparently
never produced into extremely long filaments. Pelvic fins short, not reaching anal fin; length of pelvic
fins 22 to 26% of standard length. Lateral-line scales 52 to 55. Colour: (from literature descriptions)
body rose above, silvery below; caudal fin yellowish; other fins apparently some shade of red.
Size: Maximum standard length about 17 cm.
Habitat, biology, and fisheries: Occurs in depths of 50 to 235 m. No other information available.
Distribution: Wide ranging
from Hawaii to the Philippines
(off Luzon) and Indonesia (off
Lombok and the Kai Islands).
click for next page

2442 Bony Fishes
SERRANIDAE
Groupers and sea basses (also, soapfishes, anthiines, etc.)
by P.C. Heemstra and J.E. Randall
D iagnostic characters: Body variable in shape, from deep-bodied to elongate and little compressed
(at least anteriorly) to notably compressed (size from a few to 250 cm). Opercle with 3 (rarely 2) flat
spines; margin of preopercle nearly always serrate (but serrae rudimentary in adults of a few
species) or with 1 to 4 spines. Mouth large, terminal; maxilla exposed when mouth is closed; lower
jaw usually projecting; bands of small, slender teeth in jaws; canines usually present at front of
jaws and sometimes at side; small teeth present on vomer and palatines of most species; no molars or
incisiform teeth. Gill membranes separate, with 7 branchiostegal rays. Dorsal fin single or divided into
spinous and soft portions, with IV to XIII spines and 9 to 25 soft rays; anal fin with III (rarely II) spines
and 6 to 24 soft rays; last dorsal and anal-fin rays usually split to their base but counted as a single ray;
caudal fin with 12 to 15 branched rays, the fin varying in shape from rounded to lunate; pelvic fins with I
spine and 5 soft rays, inserted below or slightly anterior or posterior to base of pectoral fins; no scaly axillary
process at base of pelvic fins. Scales small to moderate, adherent, ctenoid (or secondarily cycloid). A single
complete lateral line (except Pseudogrammini and some species of Plectranthias), extending on caudal
fin less than 1/2 length of middle caudal-fin rays. Vertebrae 24 to 30. Colour: variable with patterns of light
or dark stripes, spots, vertical or diagonal bars, or nearly plain; many species are capable of rapid colour
changes; xanthic (yellow) phases are known in some species and several species have distinctively
coloured deep- and shallow-water forms; colour patterns are generally the most useful field characters
as the morphometric and meristic characters often overlap to a considerable degree.
dorsal fin single or divided into spinous and soft
3 (rarely 2) flat portions, with IV-XIII spines and 9-25 soft rays
opercular spines
mouth large,
maxilla
exposed when
mouth closed
margin of preopercle nearly anal fin usually with III

always serrate or with 1-4 spines spines and 6-25 soft rays
Habitat, biology, and fisheries: Serranids are benthic or bottom-oriented fishes, usually found on coral
reefs or rocky substrata; the great majority of species occur on continental or insular shelves in depths
less than 200 m. All are predaceous, the larger species feeding mainly on fishes, crustaceans, and
cephalopods, while many of the smaller ones feed on zooplankton. Most serranids represented in the area
are protogynous hermaphrodites, i.e. they first mature as females and, after spawning one or more times,
they will then change sex, spawning thereafter as males. Fishes of the subfamily Serraninae are
synchronous hermaphrodites, with both sexes combined (and functional) at the same time in a single
individual. Although these synchronous hermaphrodites can fertilize their own eggs, they normally spawn
in pairs and alternate the release of eggs or sperm in order to have their eggs fertilized by the other fish.
Some groupers (subfamily Epinephelinae) form large aggregations at specific sites at the time of spawning,
making them vulnerable to over-fishing. These spawning aggregations should be protected. Except for
occasional spawning aggregations, most groupers are solitary fishes, and tagging studies have shown that
they are generally resident on a particular reef for a long time (often years). This site specificity and the
relatively slow growth rate of groupers make them particularly vulnerable to over-fishing. Groupers are
among the most highly priced food fishes and are actively sought by commercial and sport fishermen. They
are caught with hook-and-line, gill nets, spear, traps, and in trawls. For 1995, FAO’s Yearbook of Fishery
Statistics reports a total catch of around 45 600 t of Serranidae from the Western Central Pacific. Separate
statistics per species are not available from the area. Some groupers are important in aquaculture, and a
few species have been spawned in captivity. Several species are used in cage-culture operations. Some
of the smaller serranids, particularly the colourful Anthiinae and Liopropomatini are of value as aquarium
fishes.
Perciformes: Percoidei: Serranidae 2443

Callanthiidae: lateral line running close to dorsal-fin base and terminating on upper surface of caudal
peduncle or below last dorsal-fin ray; nasal organ without lamellae; preopercle edge smooth; opercle with
1 or 2 spines.
Glaucosomatidae: rear edge of opercle with 2 small, flat points; scaly axillary process at base of pelvic fins
well developed; dorsal-fin spines VIII.
lateral line close to

dorsal-fin base
Callanthiidae Glaucosomatidae
Centropomidae: head depressed anteriorly; pectoral fins shorter than pelvic fins; lateral line extends to
rear margin of caudal fin; opercle with a single flat spine.
Lutjanidae: maxilla mostly covered by preorbital bone when mouth is closed; no spines on opercle; scaly
axillary process at base of pelvic fins usually well developed.
head depressed lateral line extends

anteriorly onto caudal fin
Centropomidae Lutjanidae
Haemulidae: maxilla mostly covered by preorbital bone when mouth is closed; no teeth on vomer or
palatines; no spines on opercle.
Sparidae: jaws with incisiform and/or molariform teeth; maxilla mostly covered by preorbital bone when
mouth is closed; no spines on opercle; edge of preopercle smooth.
maxilla mostly
covered by
preorbital bone
Haemulidae Sparidae
2444 Bony Fishes
Lobotidae: no spines on opercle; no teeth on vomer or palatines; soft dorsal and anal fins project well past
caudal-fin base.
Kuhliidae: rear edge of opercle forming only 2 flat points; branchiostegal rays 6; scaly sheath at bases of
dorsal and anal fins.
Lobotidae Kuhliidae
Acropomatidae: rear edge of opercle forming 2 flat

points, or the lower point developed as a cluster of
sharp spines; dorsal fin deeply divided or as 2 sepa-
rate spiny and soft-rayed fins, the first part with VII to
X spines, the second with I spine and 9 or 10 soft
rays; several distinct, spaced canines along lower
jaw.
Moronidae (Lateolabrax japonicus, occurs near the
northern part of the area): opercle ends in 2 flat Acropomatidae
points; vertebrae 34 to 37; swimbladder extends into
hollow of first anal-fin pterygiophore.
lateral scale series; the 1st, 5th, 10th, 15th, 20th, 25th, and 30th series are indicated
Identification note
Lateral scale series are the oblique
series of scales that run above the
lateral line from the upper end of the
gill opening to the base of the caudal
fin (Fig.1).
lateral-line scales Fig. 1 methods of counting scales
Key to the genera of Serranidae occurring in the area

Note: species names are given when a genus contains a single species, or if only a single species of the
genus is known from the area.
dorsal fin with XIII spines
1a. Dorsal fin with XIII spines and 10 or and 10-11 soft rays
11 soft rays; preopercle with en-
larged spine at corner, extending
past subopercle (Fig. 2) . . . . . Niphon spinosus
1b. Dorsal fin with IV to XIII spines and
10 to 25 soft rays, the rays more nu-
merous than the spines; spine at cor- enlarged
ner of preopercle enlarged in some preopercular
species, but not extending past spine
subopercle . . . . . . . . . . . . . . . . . . . ® 2
Fig. 2 Niphon spinosus
2a. Scaly flap of skin joining base of upper pectoral-fin rays to body; oblique scale series
more numerous than lateral-line scales; branched caudal-fin rays 15; dorsal-fin spines
IV, or VI to XI (rarely X) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. No scaly flap of skin joining base of upper pectoral-fin rays to body; oblique scale series
not more numerous than lateral-line scales; branched caudal-fin rays 12 to 15; dorsal-fin
spines VIII to XIII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 21
3a. Less than 1/2 of upper border of opercle joined to body by skin; dorsal-fin spines VII to
XI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (tribe Epinephelini) ® 12
3b. Most of upper border of opercle joined to body by skin; dorsal-fin spines IV, or VII to IX . . . . . ® 4
4a. Scales mainly ctenoid, not deeply embedded; preopercle edge strongly serrate; subop-
ercle and interopercle serrate . . . . . . . . . . . . . . . . . . . . . . . . (tribe Diploprionini) ® 5
4b. Scales cycloid, deeply embedded; preopercle edge smooth, except for a few short blunt
spines dorsally; subopercle and interopercle smooth; pectoral fins longer than pelvic fins
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (tribe Grammistini) ® 7
5a. Dorsal fin with VIII spines and 13 to 16 soft rays; body depth greater than head length
and 2.0 to 2.4 times in standard length . . . . . . . . . . . . . . . . . . . . . Diploprion bifasciatum
5b. Dorsal fin with IX spines and 10 to 12 soft rays; body depth less than or equal to head
length and 2.9 to 3.7 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Dorsal fin cleft to base before last spine, which is more than twice length of eighth spine;
dorsal-fin rays 10; anal fin with II spines and 8 soft rays; caudal fin truncate . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Belonoperca chabanaudi
6b. Dorsal fin indented before soft-rayed part but not to base, the last 2 spines subequal;
dorsal-fin rays 12; anal fin with III spines and 9 soft rays; caudal fin rounded . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aulacocephalus temmincki
7a. Lateral line interrupted; dorsal-fin rays 19 to 25; anal-fin rays 15 to 21 . . . . . . . . . . . . . . . ® 8

7b. Lateral line complete to caudal-fin base; dorsal-fin rays 10 to 16; anal-fin rays 8 to 13 . . . . . ® 10
8a. Lateral line double, upper part ends below middle dorsal-fin rays, lower line runs along
midlateral part of body above anal fin; large spine on preopercle pointing dorsally;
anal-fin rays 19 to 21; body of adults pale brown, with scattered, vertically-elongate dark
brown spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aporops bilinearis
8b. Lateral line single, ending below middle dorsal-fin rays; large spine on rear edge of
preopercle pointing ventrally; anal-fin rays 15 to 18, or 19 to 22 . . . . . . . . . . . . . . . . . . ® 9
9a. Anal-fin rays 19 to 22; dorsal-fin rays 22 to 25; pale mid-dorsal stripe on head . . . . Suttonia lineata
9b. Anal-fin rays 16 to 19; dorsal-fin rays 19 to 23; no pale mid-dorsal stripe on head . . Pseudogramma
10a. No flap; body depth less than head length, 3.1 to 3.7 times in standard length; pectoral-fin
rays 14 or 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grammistops ocellatus
10b. A fleshy flap on chin; body depth about equal to head length, 2.2 to 2.7 times in standard
length; pectoral-fin rays 16 to 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
11a. Chin flap large, about equal to eye diameter; anal fin with III spines and 8 soft rays; head
and body brown, with numerous small dark-edged white spots; 4 triangular black
saddle-blotches on body, the largest on nape, the smallest on front of caudal peduncle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pogonoperca punctata
11b. Chin flap shorter than pupil diameter; anal fin with II spines and 9 soft rays; head and
body dark brown or black with yellow stripes which break up into a series of dashes on
large adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grammistes sexlineatus
2446 Bony Fishes
12a. Dorsal-fin spines VII or VIII; lower edge of preopercle with 1 to 3 enlarged spines
(usually hidden by skin, but these spines can be detected by running a finger or probe
along preopercle edge) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
12b. Dorsal-fin spines IX to XI; lower edge of preopercle smooth (except for a few species of
Epinephelus with 1 to 4 enlarged serrae) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13a. Anal-fin spines strong, all 3 distinct; preorbital depth 1/2 or less of eye diameter; head
length 2.5 to 2.6 times in standard length (Fig. 3) . . . . . . . . . . . . . . . . . . . Saloptia powelli
13b. Anal-fin spines weak, the first and second covered by skin; preorbital depth 0.7 to 2 times
eye diameter; head length 2.8 to 3.1 tim es in standard length (Fig. 4) . . . . . . . . . . Plectropomus
head length 2.5-2.6 times
in standard length head length 2.8-3.1 times in standard length
strong
anal-fin spines anal-fin
Fig. 3 Saloptia Fig. 4 Plectropomus spines weak
14a. Caudal fin deeply lunate or forked; dorsal-fin spines IX (Fig. 5) . . . . . . . . . . . . . . . . . Variola
14b. Caudal fin rounded, truncate, or concave; dorsal-fin spines IX to XI . . . . . . . . . . . . . . . ® 15
15a. No teeth on palatines; body and head elongate and markedly compressed, the greatest
body width 11 to 15% of standard length and more than 3 times in head length (Fig. 6)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Anyperodon leucogrammicus
15b. Palatines with teeth; body compressed in some species, but its width only 1.8 to 3 times
in head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
dorsal fin with IX spines and 13-14 soft rays
no teeth on
palatines
Fig. 5 Variola Fig. 6 Anyperodon
X dorsal-fin spines
profile of
16a. Dorsal profile of head markedly head concave
concave; dorsal-fin spines X; rear
nostrils of adults a long vertical slit
(Fig. 7) . . . . . . . . . . . Cromileptes altivelis
16b. Dorsal profile of head straight,
c o n v e x o r s lightly concave;
dorsal-fin spines IX or XI (rarely
X); rear nostrils round or oblong . . . . . ® 17
rear nostrils a slit
Fig. 7 Cromileptes
17a. Pectoral fins distinctly asymmetric, the fifth or sixth rays longest (Fig. 8a); dorsal fin with
IX spines and 17 or 18 soft rays; caudal fin truncate (Fig. 9) . . . . . . . . . . . Aethaloperca rogaa
17b. Pectoral fins symmetric or nearly so, the middle rays longest (Fig. 8b); dorsal fin with IX
to XI spines and 12 to 21 soft rays; caudal fin rounded, truncate, or emarginate . . . . . . . . . ® 18
dorsal fin with IX spines and 17-18 rays
caudal fin
truncate
pectoral fins asymmetric pectoral fins symmetric

a) b)
Fig. 8
pectoral fins
18a. Dorsal-fin spines IX . . . . . . . . . . . ® 19 asymmetric
18b. Dorsal-fin spines XI . . . . . . . . . . . ® 20
Fig. 9 Aethaloperca
19a. Caudal fin truncate; head small, 2.9 to 3.2 times in standard length; distal part of maxilla
with step or hook-like process on lower edge (hidden by lip); dorsal-fin membranes not
incised between spines (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . Gracila albomarginata
19b. Caudal fin rounded (truncate in Cephalopholis polleni); head length 2.2 to 3.1 times in
standard length; adults with a knob at lower rear corner of maxilla (hidden by upper lip);
dorsal-fin membranes distinctly incised between spines (Fig. 11) . . . . . . . . . . . . Cephalopholis
dorsal-fin membranes not incised dorsal-fin membranes distinctly incised between spines
caudal fin
maxilla with step truncate
or hook-like maxilla with caudal fin
process knob rounded
head length 2.9-3.2 times in standard length head length 2.2-3.1 times in standard length
Fig. 10 Gracila Fig. 11 Cephalopholis
20a. Body depth distinctly greater than head length and 2.4 to 2.7 times in standard length;
dorsal fin with XI spines and 18 to 21 soft rays, the base of soft-rayed part longer than
that of spinous part (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Triso dermopterus
20b. Body depth 2.4 to 4.1 times in standard length, usually less than head length; dorsal fin
with XI spines and 12 to 19 soft rays, the base of soft-rayed part shorter than or equal
to that of spinous part (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus
Fig. 12 Triso Fig. 13 Epinephelus

2448 Bony Fishes
21a. Dorsal fin with VIII spines and 11 to 14 soft rays; nostrils far apart, the anterior nostril
tubular, at front of snout near upper lip; maxilla with well-developed supramaxillary bone
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (tribe Liopropomatini) ® 22
21b. Dorsal fin with IX to XIII spines and 9 to 21 soft rays; nostrils close together on rear half
of snout; supramaxilla rudimentary or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
22a. Body elongate, depth 5.4 to 5.7 times in standard length; caudal peduncle depth about
70% of body depth; pectoral fins short and rounded, about 1/2 head length . . Rainfordia opercularis
22b. Body depth 3 to 4.2 times in standard length; caudal peduncle depth 40 to 55% of body
depth; pectoral fins long and pointed, more than 1/2 head length . . . . . . . . . . . . . Liopropoma
23a. Branched caudal-fin rays 15; dorsal fin with X spines and 9 or 10 soft rays; lateral line
parallel to dorsal contour of body; maxilla naked; vertebrae 24 . . . . . . . . . . . . . . Chelidoperca
23b. Branched caudal-fin rays 12 to 15 (usually 13); dorsal fin with X or XI spines and 13 to
20 soft rays or XIII spines and 15 soft rays; maxilla scaly or naked; vertebrae 25 to 28
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (subfamily Anthiinae) ® 24
24a. Dorsal fin divided to base between spinous and soft portions; lateral-line scales 51 to 78;
body elongate, its depth 3.3 to 5.3 times in standard length . . . . . . . . . . . . . . . Luzonichthys
24b. Dorsal fin continuous or deeply notched between spinous and soft portions; lateral-line
scales 25 to 65 (when lateral line complete); body less elongate, its depth 1.9 to 4.2 times
in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 25
25a. No teeth on vomer or palatines; body depth 3.4 to 4.2 times in standard length; dorsal-fin
spines slender; in adult males, all dorsal-fin spines elongated to form a tall sail-like dorsal
fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Rabaulichthys altipinnis
25b. Teeth present on vomer and palatines; body depth 1.9 to 3.6 times in standard length . . . . . ® 26
26a. Patch of teeth on vomer diamond-shaped; a patch of minute teeth present on tongue . . . . . . ® 27
26b. Patch of teeth on vomer triangular or V-shaped; no teeth on tongue . . . . . . . . . . . . . . . ® 28
27a. Dorsal-fin rays 19 to 21; lateral-line scales 57 to 61; caudal fin rounded to slightly
emarginate; no elongate fin spines or rays . . . . . . . . . . . . . . . . . . . . . Caprodon schlegelii
27b. Dorsal-fin rays 13 to 17; lateral-line scales 30 to 47; caudal fin forked; 1 or more fin spines
or rays prolonged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holanthias
28a. Dorsal-fin spines XIII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthistius ocellatus

28b. Dorsal-fin spines X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 29
29a. Dorsal fin distinctly notched between spinous and soft part; supramaxilla present (may
be rudimentary); branched caudal-fin rays 12 to 15 (usually 15); maxilla scaly or naked . . . . ® 30
29b. Dorsal fin not notched between spinous and soft portions (except slightly notched in
Dactylanthias and Sacura); supramaxilla absent; branched caudal-fin rays 13; maxilla
scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 31
30a. Lateral-line scales 41 to 65; anal-fin rays 7 to 9; maxilla naked . . . . . . . . . . . . Hypoplectrodes

30b. Lateral-line scales 25 to 41; anal-fin rays 6 or 7; maxilla scaly or naked . . . . . . . . . . . . . ® 32
31a. Total gill rakers on first gill arch 28 to 33; ascending process of premaxilla extending
between frontals; anterior canines of lower jaw short, stout, and projecting obliquely
outward . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Selenanthias
31b. Total gill rakers on first gill arch 14 to 25; ascending process of premaxilla not extending
between frontals; anterior canines not projecting obliquely outward . . . . . . . . . . . Plectranthias
32a. Serrae at corner of preopercle distinctly enlarged; posterior soft rays of dorsal and anal
fins distinctly shorter than anterior soft rays; second to fourth dorsal-fin rays elongated
in adults; third dorsal-fin spine greatly elongated in males . . . . . . . . . . . . . . . . . . . . Sacura
32b. Serrae at corner of preopercle not distinctly enlarged; posterior soft dorsal- and anal-fin
rays not shorter than anterior soft rays, the second to fourth dorsal-fin rays not elongated . . . . ® 33
33a. Body deep, its depth 1.9 to 2.2 times in standard length; dorsal-fin rays 18 to 20;
pectoral-fin rays 13 or 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Serranocirrhitus latus
33b. Body less deep, its depth 2.4 to 3.6 times in standard length; dorsal-fin rays 13 to 18;
pectoral-fin rays 13 to 21 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 34
34a. Pectoral-fin rays 16 to 21, middle rays, at least, branched; lateral-line scales 37 to 64 . . . Pseudanthias
34b. Pectoral-fin rays 13 to 18, all unbranched; lateral-line scales 31 to 39 . . . . . . . . . . . . . . ® 35
35a. Pectoral-fin rays 18; lateral-line scales 39; body depth 2.4 to 2.5 times in standard length
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Dactylanthias haplodactylus
35b. Pectoral-fin rays 13 to 17; lateral-line scales 31 to 37; body depth 2.6 to 3.6 times in
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 36
36a. Dorsal-fin rays 13 to 15; anal-fin rays 6 or 7; lateral-line scales 34 to 37; body elongate,
its depth 3 to 3.6 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . Tosana niwae
36b. Dorsal-fin rays 17; anal-fin rays 8; lateral-line scales 31 to 33; body depth 2.6 to 2.7 times
in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tosanoides flavofasciatus
Key to the species of Cephalopholis occurring in the area

1a. Caudal fin rounded; head length 2.2 to 2.7 times in standard length; colour pattern not
of alternating stripes of blue and orange-yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Caudal fin truncate to slightly emarginate; head small, 2.7 to 3.1 times in standard length;
head and body with alternating stripes of blue and orange-yellow; juveniles
golden-yellow, with wavy purple stripes along dorsal part of head and body, black spot
on snout in front of eye and black maxillary streak . . . . . . . . . . . . . . . Cephalopholis polleni
2a. Body deep, its depth 2 to 2.4 times in standard length; body width 2.3 to 2.6 times in body
depth; lateral line strongly arched above pectoral fin; head and body red with yellow bars;
pelvic-fin tips black; juveniles with large black spot in dorsal fin . . . . . Cephalopholis igarashiensis
2b. Body depth 2.3 to 3.2 times in standard length; lateral line not strongly arched above
pectoral fin; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Pelvic fins short, their length 2.3 to 2.5 times in head length; pectoral-fin rays 15 or 16;
dorsal-fin rays 14; anal-fin rays 8; colour reddish brown with indistinct dark blotch basally
on pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis aitha
3b. Pelvic-fin length 1.5 to 2.3 times in head length; pectoral-fin rays 15 to 20; dorsal-fin rays
14 to 17; anal-fin rays 8 to 10; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Anal-fin rays usually 8; colour generally brown to dark brown . . . . . . . . . . . . . . . . . . . ® 5

4b. Anal-fin rays 9 (rarely 10); colour generally red, orange, or yellow (except C. argus and
some C. urodeta or C. sonnerati) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
5a. Small dark spots or dark-edged pale blue spots on head and/or body . . . . . . . . . . . . . . . ® 6
5b. No small dark spots or blue ocelli on head or body . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
2450 Bony Fishes
6a. Dorsal-fin rays 14 to 16; lateral scale series 84 to 98; pectoral-fin length 1.4 to 1.5 times in
head length; dark-edged blue spots only on head and anteriorly on body . . . Cephalopholis microprion
6b. Dorsal-fin rays 15 to 17; lateral scale series 92 to 106; pectoral-fin length 1.5 to 1.8 times
in head length; blue ocelli on head, body, and basally on median fins; juveniles greenish
grey, the median fins yellow . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis cyanostigma
7a. Pectoral fins short, their length 1.5 to 1.8 times in head length; colour generally brown or
yellowish brown, with dark blue lines on head, body, and fins; black spot between upper
2 opercular spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis formosa
7b. Pectoral-fin length 1.3 to 1.6 times in head length; body brown, with 7 or 8 more or less
distinct dark bars; fins dark brown, with a pale blue line at corners of caudal fin . . Cephalopholis boenak
8a. Dorsal-fin rays 15 to 17; lower limb of first gill arch with 17 to 19 gill rakers; auxiliary
scales present on body; colour dark brown, covered with small dark-edged blue ocelli; 5
or 6 pale bars often visible on rear half of body . . . . . . . . . . . . . . . . . . Cephalopholis argus
8b. Dorsal-fin rays usually 14 or 15; lower limb of first gill arch with 13 to 16 gill rakers; no
auxiliary scales on body scales; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Lateral-line scales 66 to 80; lateral scale series 115 to 134; pectoral-fin rays 18 to 20;
body depth 2.3 to 2.8 times in standard length; colour generally red to reddish brown
(juveniles and some adults may be dark purple or brown) with widely scattered whitish
blotches (Indian Ocean) or generally brownish, covered with small dark red to reddish
brown spots and irregular white blotches (Pacific) . . . . . . . . . . . . . . Cephalopholis sonnerati
9b. Lateral-line scales 45 to 68; lateral scale series 79 to 121; pectoral-fin rays 16 to 19; body
depth 2.6 to 3.5 times in standard length; colour not as above . . . . . . . . . . . . . . . . . . ® 10
10a. Lateral-line scales 54 to 68; caudal fin blackish red, the corners broadly red, each set off
by an oblique white stripe; pectoral fins red, shading to orange-yellow distally (Pacific);
or caudal and pectoral fins uniformly blackish (Indian Ocean) . . . . . . . . . Cephalopholis urodeta
10b. Lateral-line scales 45 to 56; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
11a. Lateral scale series 79 to 90; head length 2.2 to 2.4 times in standard length; dark brown
saddle spot on caudal peduncle, followed by a smaller spot; submarginal dark streak at
corners of caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis leopardus
11b. Lateral scale series 90 to 121; head length 2.3 to 2.6 times in standard length; colour not
as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Head, body, and fins covered with small blue ocelli . . . . . . . . . . . . . . . . . . . . . . . . ® 13
12b. No blue spots on head, body, or fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13a. Body with 4 or 5 dark blotches along base of dorsal fin, a faint blotch on nape and 2
smaller ones on peduncle (blotches sometimes merging with or being replaced by dark
red vertical bars); most specimens with dark-edged blue lines radiating from eyes . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis sexmaculata
13b. No dark blotches on body or blue lines radiating from eyes . . . . . . . . . . Cephalopholis miniata
14a. Edge of subopercle and interopercle distinctly serrate; pelvic fins usually reaching anus,
their length 1.6 to 2 times in head length; colour generally orange-yellow to orange-red
or golden, with red to orange dots on head and dorsally on body . . . . . . . Cephalopholis aurantia
14b. Subopercle and interopercle usually smooth (rarely with a few small serrae); pelvic fins
not reaching anus, 1.9 to 2.2 times in head length; colour pale reddish orange, mottled
with dark red or brownish red . . . . . . . . . . . . . . . . . . . . . . . . Cephalopholis spiloparaea
Key to the species of Chelidoperca occurring in the area

1a. Caudal fin truncate; interorbital area naked; body reddish orange, with 5 oblong black
blotches along flanks . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelidoperca pleurospilus
1b. Caudal fin emarginate; interorbital area scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Scale rows from dorsal-fin origin to lateral line 3; no scales on anterior half of interorbital
area . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelidoperca margaritifera
2b. Scale rows from dorsal-fin origin to lateral line 4 or 5; interorbital scales extend to front
edge of eyes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelidoperca hirundinacea
Key to the species of Epinephelus occurring in the area

1a. Caudal fin of adults emarginate to truncate (slightly rounded on some E. bleekeri and
juveniles, and convex if broadly spread in adults) . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Caudal fin rounded (truncate on some E. fasciatus from Oceania) . . . . . . . . . . . . . . . . ® 12
2a. Interspinous membranes of dorsal fin not incised (Fig. 14a) . . . . . . . . . . . . . . . . . . . . ® 3

2b. Interspinous membranes of dorsal fin incised (Fig. 14b) . . . . . . . . . . . . . . . . . . . . . . ® 6
interspinous membranes of dorsal fin not incised interspinous membranes of dorsal fin incised
a) b)
Fig. 14
3a. Gill rakers elongate, no rudiments, 20 to 23 rakers on lower limb of first gill arch;
dorsal-fin rays 17 to 19; colour purplish to brownish grey with yellowish brown dots on
head and longitudinal brown lines on dorsal part of body (lines usually lost on large
adults) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus undulosus
3b. Gill rakers not elongate and rudiments often present, 13 to 18 rakers on lower limb of
first gill arch; dorsal-fin rays 15 to 17; colour not as above . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Second dorsal-fin spine of adults elongated, its length 1.8 to 2.4 times in head length;
total gill rakers on first gill arch 20 to 23; body depth 2.7 to 3.2 times in standard length;
body reddish brown with a white dot on each scale; broad dark red margin on spinous
portion of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus irroratus
4b. Second dorsal-fin spine not elongate (third or fourth spines longest); total gill rakers on
first gill arch 24 to 28; body depth 2.3 to 2.9 tim es in standard length . . . . . . . . . . . . . . . ® 5
5a. Body dark purplish grey with scattered irregular whitish blotches; body depth 2.6 to 2.9
times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus multinotatus
5b. Head, body, and fins bluish grey with numerous blackish dots; large adults with scattered
irregular blackish spots and blotches, most smaller than pupil; body depth 2.4 to 2.7
times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus cyanopodus
6a. Lateral-line scales 48 to 54; head and at least front of body with small spots, either yellow
(pale in preservative) or brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6b. Lateral-line scales 56 to 76; spots on head and body dark brown or absent . . . . . . . . . . . ® 10
2452 Bony Fishes
7a. Caudal fin truncate to slightly rounded; body depth 3.0 to 3.5 times in standard length; head,
body, dorsal fin, and upper third of caudal fin with small orange-yellow spots, the lower
two-thirds of caudal fin dark grey; anal and paired fins dusky, without spots . . . . Epinephelus bleekeri
7b. Caudal fin slightly emarginate (truncate on some E. chlorostigma); body depth 2.7 to 3.4
times in standard length; spots on head, body, and fins yellow or yellowish brown to dark
brown; anal fin with spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Head and front of body with small round well-separated bright yellow spots (pale in
preservative); distal margin of caudal, soft dorsal, and anal fins with close-set, dark,
yellowish brown spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus timorensis
8b. Head, body, and fins covered with small, close-set, yellowish brown to dark brown spots
(dark in preservative) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Dorsal-fin rays 15 to 17; anal fin of adults rounded to slightly angular, the longest soft ray
2.0 to 2.6 times in head length; 14 to 16 gill rakers on lower limb of first gill arch; pyloric
caeca 11 to 17; dark spots on body of adults about equal to pupil . . . . . . . Epinephelus areolatus
9b. Dorsal-fin rays 16 to 18; anal fin of adults angular or pointed, the longest soft ray 1.9 to
2.3 times in head length; 15 to 18 gill rakers on lower limb of first gill arch; pyloric caeca
26 to 52; dark spots on body of adults distinctly smaller than pupil . . . . . Epinephelus chlorostigma
10a. Anal-fin rays 9 or 10 (rarely 8); 1 to 4 small spines often present on ventral edge of
preopercle near corner; dorsal-fin rays 14 or 15; rear nostrils of adults 2 to 4 times larger
than anterior nostrils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus ergastularius
10b. Anal-fin rays 8; no spines on lower edge of preopercle; dorsal-fin rays 14 to 17; rear
nostrils not enlarged (except E. darwinensis) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
11a. Caudal fin truncate to slightly convex; rear nostrils and anterior nostrils subequal;
pectoral-fin rays 19 or 20; depth of caudal peduncle 3.2 to 3.8 times in head length; no
scales on maxilla; margin of spinous dorsal-fin membranes dark red to black . . Epinephelus retouti
11b. Caudal fin slightly emarginate; rear nostrils of adults more than twice diameter of anterior
nostrils; pectoral-fin rays 18; depth of caudal peduncle 3.9 times in head length; maxilla with
small scales; no conspicuous markings on head, body or spinous dorsal fin . . Epinephelus darwinensis
12a. Anal-fin rays 9 (rarely 10); body with 5 dark bars below dorsal fin, the last 2 bars as broad
as preceding bars; 2 pale interspaces below soft dorsal fin . . . . . . . . Epinephelus octofasciatus
12b. Anal-fin rays 8 (rarely 7 or 9); colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Dorsal-fin rays 12 to 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13b. Dorsal-fin rays 14 to 18 (rarely 13 in E. bruneus) . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
14a. Lateral-line scales 52; lateral body scales rough; rear nostrils twice as large as anterior
nostrils; no obvious dark or light markings . . . . . . . . . . . . . . . . . . . Epinephelus perplexus
(1 specimen, 46.5 cm standard length, from Queensland, Australia)
14b. Lateral-line scales 56 to 65; lateral body scales smooth; rear nostrils and anterior nostrils
subequal; juveniles with 2 broad, longitudinal, black-edged whitish bands that disappear
in adults, the dark edges breaking into dashes and spots, which may be lost in large
adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus latifasciatus
15a. Lateral-line scales with branched tubules; eye small, its diameter about 1/8 head length for
specimens of 20 cm length, about 1/9 head length at 35 cm, and 1/13 head length at 145 cm
standard length; interorbital wide, the width more than 1/5 head length for specimens of 23
to 153 cm standard length; maximum length about 270 cm; juveniles yellow, with 3 broad
black bars on body and irregular black bands on head . . . . . . . . . . . . Epinephelus lanceolatus
15b. Lateral-line scales with single tubule (except anterior scales of large E. bruneus, E.
coioides, and E. malabaricus); eye diameter more than 1/7 head length for specimens
of 20 cm length, and more than 1/8 head length at 35 cm; interorbital width less than 1/5
head length for specimen of 23 to 153 cm length; maximum length less than 150 cm
(except E. tukula, which may reach 200 cm) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
16a. Head and body mostly covered with small dark spots (orange, russet, brown, or black in
life and persist in preservative) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
16b. No distinct dark spots over most of head and body (yellow or orange spots may be
present in life, but these usually do not persist in alcohol; or there may be some dark dots
or scattered small dark spots, but not over most of head and body) . . . . . . . . . . . . . . . ® 40
17a. Lateral-line scales 46 to 53 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18

17b. Lateral-line scales 55 to 74 (except E. fuscoguttatus with 52 to 58) . . . . . . . . . . . . . . . . ® 29
18a. Midlateral body scales smooth, at least on adults (may be rough in area covered by
pectoral fins) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
18b. Midlateral body scales rough (mostly smooth on adults of E. socialis) . . . . . . . . . . . . . . ® 22
19a. Dorsal profile of head nearly straight; orange-red to brownish red spots on head, body,
and median and pelvic fins; 3 or 4 prominent black spots or blotches dorsally on body,
the first (and largest, greater than eye diameter) at base of middle of dorsal fin, the last
blotch dorsally on caudal peduncle; no blackish maxillary streak (Pl. VI, 44) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus trimaculatus
(Japan, Korea, China, Taiwan Province of China; not yet recorded from the area)
19b. Dorsal profile of head convex; spots on head, body, and fins dark brown to black; black
blotches on body at base of dorsal fin present or absent; blackish maxillary streak usually
present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
20a. Dorsal-fin rays 16 to 18; dark spots on pectoral fins progressively smaller distally;
pelvic-fin length 2.2 to 2.6 times in head length . . . . . . . . . . . . . . . . . Epinephelus faveatus
20b. Dorsal-fin rays 15 to 17; no dark spots on distal part of pectoral fins; pelvic fins 1.9 to 2.3
times in head length (for specimens of 10 to 25 cm standard length) . . . . . . . . . . . . . . ® 21
21a. Eye-sized blackish blotch at base of last 3 dorsal-fin spines; dark spots on head, body,
and fins numerous, 15 to 20 on soft dorsal fin of specimens 10 cm standard length, and
25 to 30 on soft dorsal fin of 28 cm specimens; total gill rakers on first gill arch 23 to 26
(modally 25); body width 1.7 to 2.2 times in body depth; jaws subequal . . . . Epinephelus howlandi
21b. No large blackish blotch at base of posterior dorsal-fin spines (though 1 to 3 spots at this
location are usually darker than adjacent spots); dark spots on head, body, and fins less
numerous, 5 to 7 on soft dorsal fin of 11 cm specimens, and 20 or 21 on soft dorsal fin
of 29 cm specimens; total gill rakers on first gill arch 21 to 24 (modally 23); body width
1.4 to 1.8 times in body depth; lower jaw strongly projecting . . . . . . . . Epinephelus macrospilos
22a. Dark spots on body elongate, oblique, and more

numerous posteriorly; dark spots on head separated
by more than 2 spot diameters; ventral edge of
maxilla of subadults (33 to 40 cm standard length) ventral edge
with a distinct step that develops into a bony knob in with a “step”
a)
adults (Fig. 15a) . . . . . . . . . . . . . Epinephelus longispinis
22b. Dark spots on body not elongate and not more nu-
merous posteriorly; dark spots on head separated by
less than a spot diameter (except E. bontoides); no
distinct step on ventral edge of maxilla (Fig. 15b) . . . . . . ® 23
no “step” on b)
ventral edge
Fig. 15 left maxilla
2454 Bony Fishes
23a. Pectoral fins large, their length 1.2 to 1.6

times in head length (except specimens
from Northwest Australia); dorsal-fin rays
16 to 18; serrae at corner of preopercle
distinctly enlarged (Fig. 16a); 2 oblique
dark brown bands (or elongate dark
blotches linked by narrow bands) on chest
. . . . . . . . . . . . . . . . . . Epinephelus quoyanus
23b. Pectoral-fin length 1.5 to 2 times in head serrae at corner of preopercle serrae at corner of preopercle
length; dorsal-fin rays 14 to 17; serrae at distinctly enlarged slightly enlarged
preopercle corner slightly enlarged
b)
(Fig. 16b); no oblique dark bands on chest . . . . ® 24 a)
Fig. 16
24a. Lateral scale series 82 to 86; pectoral-fin rays 18 or 19; total gill rakers on first gill arch
20 to 22; head and body with small dark spots separated by spaces greater than 2 spot
diameters; no dark spots on ventral parts of head and body; caudal, soft dorsal, and
pectoral fins blackish brown with a pale yellow to white margin . . . . . . . . Epinephelus bontoides
24b. Lateral scale series 92 to 120; pectoral-fin rays 16 to 19; total gill rakers on first gill arch
23 to 27 (except E. merra with 21 to 25); dark spots on head and body extending
ventrally, and most within a spot diameter of adjacent spots; fins not dark brown to black
with a distinct pale margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 25
25a. Dorsal-fin rays 14 or 15; length of longest dorsal-fin spines 2.7 to 3.3 times in head
length; black saddle-blotch on caudal peduncle; head and body covered by small dark
brown or orange-brown spots; head and dorsal part of body with irregular dark brown
blotches superimposed over the dark spots . . . . . . . . . . . . . . . . Epinephelus polyphekadion
25b. Dorsal-fin rays 15 to 18; longest dorsal-fin spines 2.1 to 2.7 times in head length (except
E. merra, 2.6 to 2.9 times in head length); colour not as above (especially, no black
saddle on caudal peduncle) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 26
26a. Dark spots on head and body about 1/2 size of pupil; dark brown to black spots on fins
(except spinous dorsal fin) much larger than those on body; maxilla not reaching vertical
at rear edge of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus miliaris
26b. Dark spots on body larger than pupil and larger than spots on fins; maxilla usually
reaching to or past vertical at rear edge of eye . . . . . . . . . . . . . . . . . . . . . . . . . . ® 27
27a. Pectoral-fin rays 16 to 18; length of longest dorsal-fin spine 2.4 to 3.2 times in head
length; some dark brown spots on body often joined to form short bands; pectoral fins
with small black spots largely confined to fin rays . . . . . . . . . . . . . . . . . Epinephelus merra
27b. Pectoral-fin rays 17 to 19; longest dorsal-fin spine 2.1 to 2.6 times in head length; no
confluent dark spots on body; dark spots on pectoral fins not uniformly small and not
confined to fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 28
28a. Dorsal-fin rays 17 or 18; lateral scale series 94 to 102; 3 close-set pairs of dark brown
to black spots on body at base of dorsal fin . . . . . . . . . . . . . . . . . . . Epinephelus bilobatus
28b. Dorsal-fin rays 15 to 17; lateral scale series 102 to 120; no pairs of dark brown to black
spots along base of dorsal fin; 2 large dusky to blackish areas on body and dorsal fin,
these separated by a broad whitish (though still faintly spotted) area . . . . . Epinephelus maculatus
29a. Dorsal-fin rays 17; pectoral-fin rays 20; lateral-line scales 73; body broadly marbled with
brown and finely spotted with dark brown . . . . . . . . . . . . . . . . . Epinephelus lebretonianus
(known only from 1 specimen, 24.8 cm standard length; locality unknown, but probably Indo-Pacific)
29b. Dorsal-fin rays 13 to 17 (only E. corallicola rarely with 17); pectoral-fin rays 17 to 20;
lateral-line scales less than 71 (except E. tauvina) . . . . . . . . . . . . . . . . . . . . . . . . ® 30
30a. Most dark spots on body polygonal and close-set, separated only by pale lines (forming
a reticulum) or white dots; midlateral body scales rough . . . . . . . . . . . . . . . . . . . . . ® 31
30b. Most dark spots on body round or oblong and well separated (except posteriorly on E.
socialis where spots may be confluent to form irregular dark stripes); midlateral body
scales rough or smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 33
31a. Second anal-fin spine 2.1 to 2.3 times in head length; longest dorsal-fin spine 2.5 to 2.8
times in head length; dorsal-fin rays 15 to 17; polygonal dark spots on body separated
mainly by white dots at corners of spots; elongate yellow-brown blotch behind eye or on
opercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus hexagonatus
31b. Second anal-fin spine 2.4 to 3.7 times in head length, longest dorsal-fin spine 2.8 to 3.8
times in head length; dorsal-fin rays 14 to 16; dark spots on body separated by network
of pale lines; no yellow-brown blotch behind eye or on opercle . . . . . . . . . . . . . . . . . . ® 32
32a. A single black blotch on body at base of posterior dorsal-fin spines and extending onto
fin about half-way to margin; dark spots at front of upper lip in 1 or 2 irregular horizontal
rows; caudal peduncle depth 3.2 to 3.7 times in head length . . . . . . . Epinephelus melanostigma
32b. Three or 4 black blotches (or groups of spots darker than others) at base of dorsal fin
and 1 on top of peduncle; blotch at base of rear dorsal-fin spines not extending half-way
to fin margin; small dark spots in 3 or 4 irregular rows along front of upper lip; caudal
peduncle depth 3.7 to 4.3 times in head length . . . . . . . . . . . . . . . . Epinephelus spilotoceps
33a. Dorsal-fin rays 13 to 15; total gill rakers on first gill arch 29 to 31; lateral-line scales 53
to 58; body depth 2.6 to 2.9 times in standard length; head and body pale yellowish
brown, with irregular dark brown blotches and numerous small close-set dark brown
spots; black saddle spot on caudal peduncle; midlateral body scales of adults smooth
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus fuscoguttatus
33b. Dorsal-fin rays 14 to 17; total gill rakers on first gill arch 22 to 29; lateral-line scales 53
to 74; body depth 2.7 to 3.7 times in standard length; colour not as above; midlateral
body scales rough (except large E. tauvina which have mainly smooth scales) . . . . . . . . . ® 34
34a. Irregular black spots (less than 1/2 pupil size) on head, body, dorsal fin, and a few on
caudal fin; no spots on ventral parts of head or body or on anal and paired fins; no
auxiliary scales; maxilla not reaching past vertical at rear edge of eye . . . Epinephelus magniscuttis
34b. Colour not as above; auxiliary scales present on body scales of adults; maxilla usually
reaching past vertical at rear edge of eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 35
35a. Dorsal-fin rays 15 to 17; rear nostrils (at standard length greater than 14 cm) vertically
elongate, their length 2 to 3 times diameter of anterior nostrils; pectoral-fin length 1.5 to
1.7 times in head length; lateral scale series 88 to 109; grey, with well-separated black
spots smaller than pupil on head, body and fins; 3 dusky to blackish blotches on body at
base of rear half of dorsal fin, the largest at base of last 2 or 3 spines; dusky to blackish
saddle spot on caudal peduncle; juveniles with black-edged white spots on head and
body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus corallicola
35b. Dorsal-fin rays 13 to 16; rear nostrils not vertically elongate or more than twice diameter
of anterior nostrils; pectoral-fin length 1.6 to 2.5 times in head length; lateral scale series
95 to 130; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 36
36a. Lateral-line scales 54 to 65, the anterior scales of large adults with branched tubules;
numerous small bony platelets on side of first gill arch; body with 5 irregular dark bars
which tend to bifurcate ventrally (bars may be faint or broken into series of 2 or 3 large
blotches) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 37
36b. Lateral-line scales 62 to 74, none with branched tubules; no small bony platelets on side
of first gill arch; dark bars not present on body (or only faintly on E. tauvina) . . . . . . . . . . . ® 38
2456 Bony Fishes
37a. Head and body with numerous small well-separated black spots (largest spots about 2
times larger than rear nostrils); irregular white or pale spots or blotches usually present
on head and body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus malabaricus
37b. Head, body, and usually median fins with numerous orange, brownish orange or reddish
brown spots (orange spots turn brown after death and poorly defined on preserved
specimens; spots about 4 or 5 times larger than rear nostrils); no white or pale spots on
head or body; orange spots become poorly defined and darker with growth; spots on
head often coalesce and become elongated, arranged in irregular rows radiating from
eye; fins brownish, with proximal parts spotted . . . . . . . . . . . . . . . . . Epinephelus coioides
38a. Body with several large round or oval dark brown spots, arranged in 5 or 6 subvertical
series; head and fins with smaller dark brown spots and irregular dark streaks; lateral
scale series 113 to 130; maximum total length 200 cm . . . . . . . . . . . . . . Epinephelus tukula
38b. Colour not as above; lateral scale series 95 to 112; maximum total length 80 cm . . . . . . . . ® 39
39a. Head length 2.1 to 2.4 times in standard length; head, body, and fins with orange-red to
dark brown spots (those on body nearly as large as eye in young, about pupil size in
large specimens; spots usually absent in outer part of pectoral fins of adults); body often
with faint oblique dark bars; blackish blotch often present at base of last 4 dorsal-fin
spines and extending onto lower part of fin (more evident in young) . . . . . . . Epinephelus tauvina
39b. Head length 2.4 to 2.7 times in standard length; head and body with blackish brown spots
much smaller than pupil, those posteriorly on body often coalescing to form irregular
stripes; 4 blackish blotches often present at base of dorsal fin and 1 on caudal peduncle;
caudal, dorsal, and anal fins with small white spots and white margins . . . . . Epinephelus socialis
40a. Membranes of spinous dorsal fin usually not incised (some specimens with membranes
slightly to moderately incised); head and body dark brown or greenish brown, marbled
with irregular pale spots and blotches; 1 or 2 faint dark streaks running posteriorly from
eye; no dark spots on head, body or fins . . . . . . . . . . . . . . . . . . . . Epinephelus erythrurus
40b. Interspinous dorsal-fin membranes moderately to deeply incised; colour not as above . . . . . ® 41
41a. Body pale greyish brown, with 5 dark bars; a narrow pale bar may be present within dark
bars, nearly dividing them in 2; fins greyish, the median fins with several small dark spots;
13 to 15 gill rakers on lower limb of first gill arch . . . . . . . . . . . . . . . Epinephelus sexfasciatus
41b. Colour not as above; 14 to 19 gill rakers on lower limb of first gill arch . . . . . . . . . . . . . . ® 42
42a. Head, body, and median fins uniform dark grey; pectoral fins pale, pelvic fins blackish;
body depth equal to head length, 2.4 times in standard length; lateral-line scales 67 to
69; dorsal-fin rays 16 or 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus trophis
(2 specimens taken off Western Australia in a depth of 130 m)
42b. Colour not as above; body depth less than head length, 2.5 to 3.6 times in standard
length; lateral-line scales 48 to 72; dorsal-fin rays 13 to 18 . . . . . . . . . . . . . . . . . . . . ® 43
43a. Body pale, with close-set wavy, brown or brownish orange stripes, the intervening pale
lines forming a maze-like pattern; head greyish brown with numerous small close-set
brownish orange spots; dorsal-fin rays 15 to 17; body depth 2.5 to 2.9 times in standard
length; interorbital space convex . . . . . . . . . . . . . . . . . . . . . Epinephelus undulatostriatus
43b. Colour not as above; dorsal-fin rays 13 to 18; body depth 2.6 to 3.6 times in standard
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 44
44a. Dorsal-fin rays 15 to 18 (rarely 15); body usually with vertical or slightly oblique broad
dark bars (faint or absent on some species) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 45
44b. Dorsal-fin rays 13 to 16 (rarely 16); body without dark vertical bars (except juvenile E.
bruneus and E. daemelii which have dark oblique bars containing pale spots and
streaks) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 53
45a. Head, body, and fins tan dorsally; belly and rear part of body whitish ventrally; chest and
ventral part of head reddish orange; dark orange-brown to black dots (less than 2 mm
diameter) on head (behind eye) and body (anteriorly); body with 5 faint, oblique, dark
bars (midlateral part of each bar may be darker, representing a series of squarish dark
blotches continued from the dark band running from eye to end of operculum) soft dorsal
and caudal fins with narrow dark brown margin; dark line along base of soft dorsal fin;
lateral-line scales 48 to 51 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus stictus
45b. No dark brown to black dots on body or head (some species with black spots, but these
are larger than 2 mm); 48 to 75 lateral-line scales . . . . . . . . . . . . . . . . . . . . . . . . . ® 46
46a. Margin of interspinous dorsal-fin membranes black (red in specimens from Western
Australia and deep water); edge of orbit (around eye) narrowly black, surrounded by pale
blue line; body usually with 5 faint dark bars often containing irregular pale spots . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus fasciatus
46b. Margin of spinous dorsal fin not black; rim of orbit not black; dark bars on body present
or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 47
47a. Nape and front of body above lateral line with minute cycloid scales and numerous pores;
body scales with a whitish dot (may be lost in alcohol); semicircular dark red to reddish
brown blotch at base of pectoral fin; dark pigment in groove along base of spinous dorsal
fin; usually some violet lines and spots on cheek . . . . . . . . . . . . . . . . Epinephelus rivulatus
47b. Nape and dorsoanterior part of body without minute scales and numerous pores; colour
not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 48
48a. Corner of preopercle with 1 to 6 large

spines (2 or 3 times longer than serrae
above corner) (Fig. 17a); dark bars on
body vertical or nearly vertical . . . . . . . . . . . ® 50
48b. Serrae at corner of preopercle only slightly
to moderately enlarged (Fig. 17b); dark
bars on body oblique (may be faint or ab-
sent) . . . . . . . . . . . . . . . . . . . . . . . . ® 49 corner of preopercle serrae at corner of preopercle
with 1-6 large spines only slightly enlarged
a) b)
49a. Head large, its length 2.1 to 2.4 times in Fig. 17
standard length; adults with 4 dark brown
bars on body and brown bands on head; small black spots on edges of brown bars and
bands; no dark spots on fins; juveniles with black bars on body and small black spots on
fins; 14 to 16 gill rakers on lower limb of first gill arch . . . . . . . . . . Epinephelus amblycephalus
49b. Head length 2.3 to 2.6 times in standard length; 4 dark bars (sometimes faint or absent)
on dorsal part of body; head and body with numerous small yellow spots (but no black
spots); body and median fins also with pale grey spots; rear margin of median fins with
a broad yellow border; 16 to 18 gill rakers on lower limb of first gill arch . . . . . Epinephelus awoara
50a. Head and body brownish grey, covered (except ventrally) with red, orange, or gold spots
(pale in alcohol); dorsal-fin margin yellow or orange; dusky yellow or orange spots (1 per
membrane) along middle of spinous dorsal fin and another row along base of fin; black
blotch at base of last 3 dorsal-fin spines . . . . . . . . . . . . . . . . . . . . . . Epinephelus akaara
50b. Colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 51
51a. Head and body with numerous small dark brown, brownish yellow, or russet spots (dark
in alcohol); body usually with 5 broad dark bars, the first 4 extend onto dorsal fin (first 2
bars darker dorsally, extending to margin of dorsal fin), the last bar on caudal peduncle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus fasciatomaculosus
51b. Colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 52
2458 Bony Fishes
52a. Head, body, and dorsal fin dark brownish grey, with numerous small white spots and
blotches; on adults (larger than 30 cm standard length) most white spots on the body are
coalesced, forming wavy pale lines and mottlings; a few dark blotches usually visible on
body at base of dorsal fin; midlateral part of lower jaw with 3 to 5 rows of small teeth . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus caeruleopunctatus
52b. Body with 7 or 8 horizontal stripes of greenish brown and yellow-orange; dorsal part of
head and snout dark green; cheek pale green (colour from Valenciennes, 1830);
midlateral part of lower jaw with 2 rows of teeth; dorsal-fin rays 17; body depth 3.4 times
in standard length; lateral-line scales 52 to 54; first gill arch with 8 gill rakers on upper
limb, 15 on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus chlorocephalus
(known only from 1 specimen, 17 cm standard length, from Tonga Islands)
53a. Body of juveniles with oblique dark bars and pale spots; body elongate, its depth
distinctly less than head length, 2.9 to 3.6 times in standard length; lateral-line scales 63
to 72 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 54
53b. No dark bars on juveniles or adults; body depth 2.6 to 3.3 times in standard length;
lateral-line scales 48 to 72 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 55
54a. A black saddle-blotch on caudal peduncle; numerous small white spots on head and
body; body scales smooth (except area covered by pectoral fins), with auxiliary scales;
adults often uniformly dark brown or black, the underside of head pale . . . . . Epinephelus daemelii
54b. No black saddle-blotch on peduncle; white streaks and dark bands radiating from eye,
but no small white spots on head or body (pale blotches may be present, especially
within dark body bars); midlateral body scales rough, without auxiliary scales; adults
(larger than 50 cm) dark greyish brown, the bars and bands replaced by faint dorsal
blotches or absent altogether, the body covered with small pale grey spots forming short
horizontal lines and a mottled pattern . . . . . . . . . . . . . . . . . . . . . . . Epinephelus bruneus
55a. Serrae at corner of preopercle slightly to

moderately enlarged (Fig. 18a); midlateral
part of lower jaw with 3 to 5 rows of teeth
in adults; lateral-line scales 48 to 55; body
scales with auxiliary scales; colour pattern
of white dots, spots, or blotches . . . . . . . . . . ® 56
55b. Preopercle corner with 2 to 5 distinctly
enlarged serrae (Fig. 18b); midside of
lower jaw with 2 rows of teeth in adults;
lateral-line scales 52 to 72; no auxiliary serrae at corner of preopercle corner of preopercle with 2-5
scales on body scales; colour pattern not slightly enlarged distinctly enlarged serrae
as above . . . . . . . . . . . . . . . . . . . . . . ® 57 a) b)
Fig. 18
56a. Lateral scale series 81 to 91; snout length 4.8 to 5.2 times in head length; head, body,
and fins dark brown with numerous whitish dots (may not persist in preservative); rear
margin of median and pectoral fins narrowly orange-red . . . . . . . . . . . Epinephelus polystigma
56b. Lateral scale series 95 to 110; snout length 4.0 to 4.8 times in head length; head, body,
and fins dark grey-brown with numerous small whitish spots and scattered large whitish
blotches; small white spots of adults tend to coalesce to form irregular horizontal stripes
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus ongus
57a. Lateral scale series 89 to 100; total gill rakers on first gill arch 21 to 25; no dark or light
markings on body; yellowish brown stripe from eye to opercular flap and 2 similar bands
on cheek, the lower runs posteriorly from end of maxilla . . . . . . . . . . . Epinephelus heniochus
57b. Lateral scale series 102 to 125; total gill rakers on first gill arch 23 to 28; body with dark
brown bands and/or spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 58
58a. Body with long curved dark brown bands or series of spots, the middle of each band or
row of spots more ventral than the ends . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 59
58b. Dark markings not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 60
59a. Large oval dark brown blotch (or group of small spots on larger specimens) on body at
base of middle dorsal-fin spines, this blotch not joined to lower dark bands; broadly-
curved dark bands or rows of spots on head and body paralleling curvature of isolated
dark blotch; dark markings faint or absent on large adults . . . . . . . . . Epinephelus poecilonotus
59b. Dark brown blotch on body at base of middle dorsal-fin spines joined to dark brown band
passing to upper edge of operculum; a second dark band from upper end of gill opening,
bifurcating above pectoral fin, with branches to anterior and posterior dorsal-fin rays; 2
bands from eye, the upper branch to nape, the lower a broad curve or broken line to
upper part of caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus morrhua
60a. Head and body pale brownish or greenish grey, with small, brownish black spots on
dorsolateral part of body and sometimes on rear part of head and median fins; body
depth 3.0 to 3.3 times in standard length . . . . . . . . . . . . . . . . . . . . Epinephelus epistictus
60b. Head and body with intricate pattern of irregular dark lines, spots, and blotches; body
depth 2.6 to 3.0 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 61
61a. Five irregular dark brown bands (with age only the edges remain dark) passing down-
ward and forward from dorsal surface of body, the first from nape to eye, the second band
from middle dorsal-fin spines to upper end of gill opening, third and fourth bands from
dorsal-fin rays, branching as they pass ventrally, the fifth band on caudal peduncle . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epinephelus radiatus
61b. Dark markings on postorbital head and body forming a coarse broken reticulum, none
passing downward and forward from dorsal surface of body . . . . . . . . Epinephelus tuamotuensis
Key to the species of Holanthias occurring in the area

1a. Dorsal-fin rays 13 or 14; lateral-line scales 30 to 39; third dorsal-fin spine elongate in
adult males; rear part of body with small silvery spots . . . . . . . . . . . . . . . . . . . . . . . ® 4
1b. Dorsal-fin rays 15 to 18; lateral-line scales 36 to 43; third dorsal-fin spine not elongate;
no silvery spots on rear part of body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Body depth 2.0 to 2.3 times in standard length; dorsal-fin rays 17 or 18; lateral-line scales
39 to 43; body pink with large yellow to dark brown blotches (most larger than eye) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holanthias borbonius
2b. Body depth 2.2 to 2.45 times in standard length; dorsal-fin rays 15 to 17; lateral-line
scales 36 to 41; no large blotches on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Caudal-fin lobes 1.3 to 1.6 times in standard length; total gill rakers on first gill arch 42
to 47; scales on top of head extending in front of nostrils; dorsal fin lavender, with yellow
submarginal line in spinous part and yellow filamentous soft rays . . . . . . . . . . Holanthias tapui
3b. Caudal-fin lobes 2.1 to 3.0 times in standard length; total gill rakers on first gill arch 39
to 41; no scales in front of nostrils; dorsal fin yellow, except for magenta spines and ray
tips and purple spots in soft portion . . . . . . . . . . . . . . . . . . . . . . Holanthias chrysostictus
4a. Dorsal-fin rays 14; lateral-line scales 35 to 39; no black and white bars at base of caudal
fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holanthias unimaculatus
4b. Dorsal-fin rays 13; lateral-line scales 30 to 32; curved dark brown bar preceded by white
bar at base of caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . Holanthias rhodopeplus
Key to the species of Hypoplectrodes occurring in the area

1a. Body with 6 narrow, dark brown vertical bars, the last 2 on caudal peduncle; 2 similar bands
from eye to eye over the nape; dorsal-fin rays 17 to 19; anal-fin rays 7 . . . . Hypoplectrodes annulatus
1b. No narrow dark bars on body; dorsal-fin rays 17 to 21; anal-fin rays 7 to 9 . . . . . . . . . . . . ® 2
2a. Body covered with close-set dark brown blotches; lateral-line scales 38 to 42; dorsal-fin
rays 17 to 19 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hypoplectrodes jamesoni
2b. Body pale pink to brownish red, with 5 broad dark saddle-blotches at base of dorsal fin;
lateral-line scales 45 to 49; dorsal-fin rays 19 to 21 . . . . . . . . . . . Hypoplectrodes maccullochi
2460 Bony Fishes
Key to the species of Liopropoma occurring in the area

1a. Dorsal fin continuous (though it may be deeply notched, all spines and soft rays are
connected above their bases by membranes) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Dorsal fin divided into separate spinous and soft-rayed fins . . . . . . . . . . . . . . . . . . . . ® 5
2a. First gill arch with 8 gill rakers on upper limb, and on lower limb 19 or 20 (only 2
rudiments); pectoral-fin rays 14; dorsal-fin rays 11 or 12; caudal fin truncate to slightly
rounded; no scales on front of snout or on preorbital; head and body with 6 or 7 broad
dark stripes; prominent dark ocellus in soft dorsal and anal fins . . . . . . . . . Liopropoma swalesi
2b. First gill arch with 5 to 7 gill rakers on upper limb, and on lower limb 11 to 16 (including
9 to 12 rudiments); pectoral-fin rays 14 to 16; dorsal-fin rays 12 to 14; caudal fin slightly
to deeply emarginate; head covered with scales; no dark stripes on body; no ocelli in fins . . . . . ® 3
3a. Last dorsal-fin spine shorter than sixth and seventh spines; caudal fin with broadly
rounded corners; anterior nostril tube not reaching lip; anal-fin rays 9; pectoral-fin rays
15 or 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liopropoma erythraeum
3b. Last (eighth) dorsal-fin spine longer than sixth and seventh spines; caudal fin corners
pointed; anterior nostril tube overlaps upper lip; anal-fin rays 8; pectoral-fin rays 14 or 15 . . . . . ® 4
4a. Body with scattered, small, dark brown spots (often semicircular); caudal concavity 4 or
5 times in head length; pectoral-fin rays 15 . . . . . . . . . . . . . . . . . . Liopropoma lunulatum
4b. No dark spots on body; caudal concavity 12 or 13 times in head length; pectoral-fin rays
14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liopropoma incomptum
5a. Head and body with 4 to 8 dark brown longitudinal stripes . . . . . . . . . . . . . . . . . . . . . ® 6

5b. No dark longitudinal stripes on head and body . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6a. Body with 6 to 8 dark stripes; pectoral-fin rays 15 or 16 . . . . . . . . . . . . . Liopropoma susumi

6b. Body with 4 or 5 dark stripes; pectoral-fin rays 14 or 15 . . . . . . . . . . . . . Liopropoma collettei
7a. Pectoral-fin rays 15 or 16; no enlarged pore in front of rear nostrils; body reddish
grey-brown anteriorly, dusky or brownish orange-red posteriorly, with faint brownish
orange lines along scale rows . . . . . . . . . . . . . . . . . . . . . . . . . . Liopropoma pallidum
7b. Pectoral-fin rays 13 to 15; 1 or 2 enlarged pores in front of each rear nostril; colour not
as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Two to 10 small pores in a group on each side of anterior interorbital space; scales rarely
extend dorsally on snout more than half-way from rear nostrils to upper lip; body red to
brownish red, often with faint lines following scale rows; yellow stripe from front of snout
to eye, another between eyes, and 1 to 3 running posteriorly from eye . . . . Liopropoma mitratum
8b. A pair of large pores, 1 on each side of interorbital space on a vertical through anterior
edge of pupil; scales dorsally on snout extend more than half-way from rear nostrils to
upper lip . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Head, body, and fins immaculate yellow; pelvic fins short, 22% of standard length and
1.75 times in head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liopropoma flavidum
9b. Colour not entirely yellow; pelvic fins 23 to 28% of standard length, 1.4 to 1.7 times in
head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Body with 2 red bands separated by midlateral white stripe from head to caudal fin; body
depth 3.5 to 3.7 times in standard length . . . . . . . . . . . . . . . . . . . Liopropoma tonstrinum
10b. Body dusky yellow with red lines following scale rows; caudal peduncle reddish, with
midlateral white stripe; body depth 3.0 to 3.5 times in standard length . . Liopropoma multilineatum
Key to the species of Luzonichthys occurring in the area

1a. Anal-fin rays with II spines and 9 soft rays; pectoral-fin rays 21 to 23; lateral-line scales
70 to 78; body elongate, its depth 4.2 to 5.3 times in standard length; some scales on
head and dorsoanterior part of body with enlarged ctenii; body lavender-pink with a
dark-edged orange-yellow stripe between lateral line and dorsal-fin base . . . Luzonichthys williamsi
1b. Anal fin with III spines and 7 soft rays; pectoral-fin rays 17 to 22; lateral-line scales 51
to 74; body depth 3.2 to 5.0 times in standard length; scale ctenii not enlarged; no
dark-edged orange-yellow stripe on back . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Lateral-line scales 65 to 74; pectoral-fin rays usually 21 or 22; body depth 4.2 to 5.0 times
in standard length; upper third of body yellow or orange-yellow, the lower two-thirds
lavender-pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Luzonichthys whitleyi
2b. Lateral-line scales 51 to 68; pectoral-fin rays 17 to 20; body depth 3.2 to 4.1 times in
standard length; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Pectoral-fin rays modally 20; lateral-line scales 59 to 68; caudal fin lunate, the caudal
concavity 6.1 to 7.2 times in standard length; scales dorsally on snout not extending in
front of nostrils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Luzonichthys earlei
3b. Pectoral-fin rays modally 19; lateral-line scales 51 to 60; caudal-fin lobes elongate, the
caudal concavity 4.0 to 5.6 times in standard length; scales dorsally on snout extending
in front of anterior nostrils . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Caudal peduncle depth 2.4 to 2.5 times in head length; snout length 3.7 to 4.1 in head
length; pectoral-fin length 3.2 to 3.4 and pelvic-fin length 3.7 to 4.7 times in standard
length; first gill arch with 7 to 10 gill rakers on upper limb, 20 to 22 on lower limb; no linear
colour pattern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Luzonichthys waitei
4b. Caudal peduncle depth 2.6 to 2.7 times in head length; snout length 4.2 to 4.4 in head
length; pectoral-fin length 3.5 to 3.7 and pelvic-fin length 4.6 to 4.9 times in standard
length; first gill arch with 7 gill rakers on upper limb, 10 on lower limb; orange and pale
yellow stripes on front of body . . . . . . . . . . . . . . . . . . . . . . . . . Luzonichthys taeniatus
Key to the species of Plectranthias occurring in the area

1a. Some pectoral-fin rays branched; lateral line complete . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. No pectoral-fin rays branched; lateral line complete or incomplete . . . . . . . . . . . . . . . . ® 15
2a. Head, including maxilla and chin, covered with scales . . . . . . . . . . . . . . . . . . . . . . . ® 3

2b. Head not completely scaly (most of snout, chin, and maxilla naked) . . . . . . . . . . . . . . . . ® 4
3a. Caudal fin rounded or truncate with rounded corners; no dorsal-fin rays elongated; no
canine teeth in lower jaw; 10 to 12 gill rakers on lower limb of first gill arch; head and
body nearly uniform light red . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias japonicus
3b. Caudal fin slightly emarginate, the second upper ray usually elongate; second dorsal-fin
ray elongate; 2 stout canine teeth at front of lower jaw and another pair on side of jaw;
14 to 17 gill rakers on lower limb of first gill arch; dark red bars on body and a small red
spot on upper basal part of caudal fin . . . . . . . . . . . . . . . . . . . . . . Plectranthias kelloggi
4a. Body depth 3.4 times in standard length; eye diameter 2.9 times in head length; no
canine teeth; preopercle smooth; colour mainly yellow . . . . . . . . Plectranthias megalophthalmus
4b. Body depth 2.2 to 3.2 times in standard length; eye diameter 3.0 to 4.8 in head length;
canine teeth present; preopercle serrate and lower margin with 2 antrorse spines; colour
(when known) not mainly yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
2462 Bony Fishes
5a. Dorsal-fin rays 14 or 15; fourth or fifth dorsal-fin spines longest . . . . . . . . . . . . . . . . . . ® 6

5b. Dorsal-fin rays 16 to 18; third or fourth dorsal-fin spine longest . . . . . . . . . . . . . . . . . . ® 8
6a. Dorsal-fin rays 14; preopercle edge smooth; maxilla scaly . . . . . . . . . . . . . Plectranthias lasti
6b. Dorsal-fin rays 15; preopercle edge serrate; maxilla scaly or naked . . . . . . . . . . . . . . . . ® 7
7a. Lateral-line scales 29; maxilla and lower jaw naked; pectoral-fin rays 14; body pink, with
3 orange-red bars below dorsal fin, a Y-shaped orange-red bar on nape, and 7 red spots
on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias rubrifasciatus
7b. Lateral-line scales 32 to 35; maxilla and rear part of lower jaw scaly; pectoral-fin rays 16;
colour tan, becoming silvery white ventrally; first 4 lateral-line scales dusky . . Plectranthias robertsi
8a. Lateral-line scales 40 or 41; oblique rows of large scales on cheek between eye and
corner of preopercle 8 or 9; body red, with no distinctive markings . . . . . . . . Plectranthias taylori
8b. Lateral-line scales 28 to 36; oblique rows of large cheek scales 5 to 7; colour (when
known) not entirely red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Fourth dorsal-fin spine longest, 2.9 times in head length; pectoral fins just reaching
vertical at base of first anal-fin ray; body pale red with 2 rows of large dark red blotches
dorsally; fins yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias whiteheadi
9b. Third dorsal-fin spine longest, 1.8 to 2.9 times in head length; pectoral fins reaching past
vertical at base of first anal-fin ray; colour not as above . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Lateral-line scales 33 to 36; dorsal-fin rays 17 or 18 . . . . . . . . . . . . . . . . . . . . . . . ® 11

10b. Lateral-line scales 28 to 31; dorsal-fin rays 16 . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
11a. Oblique rows of large cheek scales 5; body with 6 faint dark saddle blotches, first from
nape to dorsal-fin origin, second below middle dorsal-fin spines, sixth at base of last 4
dorsal-fin rays; another 2 diffuse saddle blotches on caudal peduncle . . . Plectranthias anthioides
11b. Cheek scale rows 7; body with 4 dark saddle-bars, first from nape to first 2 dorsal-fin
spines, the fourth bar below twelfth to sixteenth dorsal-fin rays, and 2 more dark bars on
rear half of peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias jothyi
12a. Inner teeth on side of lower jaw about twice as long as teeth in outer rows; length of
longest caudal-fin ray about 1.4 times in head length; body pale, with red bar from middle
of dorsal fin to front of anal fin and another on caudal peduncle . . . . . Plectranthias retrofasciatus
12b. Teeth on side of lower jaw equal in size; longest caudal-fin ray 1.6 to 1.9 times in head
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Pectoral-fin rays 14; eye diameter 3.35 times in head length . . . . . . . . . . Plectranthias knappi
13b. Pectoral-fin rays 13; eye diameter 3.8 to 4.4 times in head length . . . . . . . . . . . . . . . . ® 14
14a. Body depth 2.7 to 2.8 times in standard length; third dorsal-fin spine and second anal-fin
spine 2.0 times in head length; life colour unknown . . . . . . . . . . . . . . . Plectranthias pallidus
14b. Body depth 2.6 to 2.65 times in standard length; third dorsal-fin spine 2.4 to 2.5 times in
head length, second anal-fin spine 2.2 to 2.3 times in head length; body whitish, with
large orange-red blotches suffused with yellow . . . . . . . . . . . . . . . . . Plectranthias wheeleri
15a. Lateral-line usually incomplete, 12 to 22 scales in anterior series . . . . . . . . . . . . . . . . ® 16

15b. Lateral-line complete, 25 to 31 scales in series to caudal-fin base . . . . . . . . . . . . . . . . ® 19
16a. Third dorsal-fin spine elongate, with a pennant-like flap near tip; edge of preopercle smooth
or with a few weak serrae; pectoral fins reaching past anal-fin base, the length 2.2 to 2.5 times
in standard length; body pale, with large close-set red blotches . . . . . . . . . . Plectranthias inermis
16b. Fourth dorsal-fin spine longest; upper edge of preopercle strongly serrate and 2 antrorse
spines on lower edge; pectoral-fin length 2.4 to 3.2 times in standard length; colour not
as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
17a. Pectoral-fin rays 12 or 13; lateral-line scales 12 to 15; subopercle with 2 to 7 coarse
serrae and interopercle with 1 to 8 serrae; body pale, with brown blotches forming
irregular bars; dark brown spot at rear end of dorsal-fin base, another at base of upper
caudal-fin rays, a third at base of last anal-fin ray, and a fourth spot between second and
third spots; small dusky spots along bases of dorsal and anal fins . . . . . Plectranthias longimanus
17b. Pectoral-fin rays 14 to 18; lateral-line scales 14 to 22; subopercle serrae 0 to 2 (1 of 46
P. nanus with 4), and none on interopercle (a single P. nanus with 1); colour not exactly
as above (but that of P. nanus similar) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
18a. Dorsal-fin rays 13 to 15; pectoral-fin rays 14 to 16; colour as in 17a except a vertical
brown line on caudal-fin base separated by a pale zone from 2 dark spots on caudal
peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias nanus
18b. Dorsal-fin rays 16 or 17; pectoral-fin rays 16 to 18; body mottled orange anteriorly,
becoming red and white posteriorly, with red spot at dorsal-fin origin and white spot at
base of last soft ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias winniensis
19a. Caudal fin emarginate, with 1 or more filamentous rays . . . . . . . . . . . . . . . . . . . . . . ® 20

19b. Caudal fin truncate or rounded, without elongated rays . . . . . . . . . . . . . . . . . . . . . . ® 21
20a. No antrorse spines on lower edge of preopercle; several rays of caudal-fin lobes
filamentous; dorsal-fin rays 14 or 15, the second to fourth elongated; small dark blotches
on nape, 5 or 6 at base of dorsal fin, and 2 dorsally on caudal peduncle; 2 dark blotches
just below lateral line, 1 below last dorsal-fin spines and the other just behind head; 2
faint dark blotches on side of caudal peduncle (life colour unknown) . . . . . . Plectranthias foresti
20b. Two large antrorse spines on lower edge of preopercle; only the second branched
caudal-fin ray elongated; dorsal-fin rays 15 to 17, none elongated; body pale, mottled
with red and yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectranthias sagamiensis
21a. Preopercle with 18 to 21 serrae; oblique rows of cheek scales 6; lower pectoral-fin rays
not enlarged; caudal fin truncate; body rose, with blackish blotches dorsally; colour in
alcohol essentially as in P. foresti . . . . . . . . . . . . . . . . . . . . . . . Plectranthias megalepis
(1 specimen, 62 mm standard length, from Kai Islands, Indonesia)
21b. Preopercle with 1 to 4 serrae; cheek scale rows 4 or 5; lower pectoral-fin rays enlarged;
caudal fin rounded; dark bars on body and large black spot basally in anal fin . . . . . . . . . . ® 22
22a. Palatine teeth absent; pectoral-fin rays 12 or 13, the long lower rays slightly enlarged;
dorsal-fin rays 16 to 18; lateral-line scales 25; interorbital area without scales; body pale, with
4 or 5 brown bars, first in front of dorsal fin, last at base of caudal fin; 4 prominent black spots
as follows: covering rear third of spinous dorsal fin, below base of last 6 dorsal-fin rays, at
base of last 4 anal-fin rays and midventrally on abdomen . . . . . . . . . . . Plectranthias fourmanoiri
22b. Palatine teeth present; pectoral-fin rays 14, the lower 6 rays about twice as thick as shorter
upper rays; dorsal-fin rays 15; lateral-line scales 29 or 30; top of head scaled anterior to
nostrils; body pale, with 6 broad brown bars, the first on nape, the last on caudal-fin base,
the ones between tending to form double bars (upper ends extending into basal part of dorsal
fin where darker); large dark spot at base of anal fin, another at pectoral-fin base, and 2 dark
spots, one above the other, on caudal-fin base . . . . . . . . . . . . . . . . Plectranthias cirrhitoides
2464 Bony Fishes
Key to the species of Plectropomus occurring in the area

1a. Anterior rays of soft dorsal and anal fins elevated, the fin margin with a distinct lobe
anteriorly; longest dorsal-fin rays 1.6 to 2.1 times in head length; pectoral-fin rays 14 to
16; adults with vertical blue lines anteriorly on side of body and head . . Plectropomus oligacanthus
1b. Anterior rays of dorsal and anal fins not elevated to form a lobe; longest dorsal-fin rays
2.2 to 3.2 times in head length; pectoral-fin rays 15 to 18; adults not coloured as above . . . . . ® 2
2a. Body uniform brown or brown marbled with olive-green, brownish orange, or white;
juveniles brownish with pale horizontally elongate spots and streaks; pectoral-fin rays 16
to 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectropomus punctatus
2b. Body with numerous blue spots or with saddle-like black blotches and a few blue spots;
pectoral-fin rays 15 to 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Caudal fin truncate to slightly emarginate, the caudal concavity (difference between
lengths of longest [upper] ray and shortest [middle] rays) 13 or more times in head length;
interorbital area with small embedded scales; head, body (including ventral parts), and
median fins covered with close-set, round to slightly oval, dark-edged blue spots;
distance between spots subequal to spot diameters; 2 to 7 developed gill rakers on lower
limb of first gill arch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectropomus areolatus
3b. Caudal fin emarginate, the caudal concavity 5 to 12 times in head length; no scales on
interorbital area; blue spots round to oblong; 4 to 10 developed gill rakers on lower limb
of first gill arch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Pectoral-fin rays 16 to 18; caudal-fin length 1.5 to 1.8 times in head length; pectoral and
pelvic fins 2.1 to 2.4 times in head length; head and body pale, with 4 or 5 saddle-like
dark brown or black blotches and a few small blue spots, the fins yellow; or head and
body brownish with numerous small blue spots and with or without faint dark saddle
blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectropomus laevis
4b. Pectoral-fin rays 15 to 17; caudal fin length 1.3 to 1.5 times in head length; pectoral and
pelvic fins 1.7 to 2.3 times in head length; no dark saddle-like blotches on body . . . . . . . . . ® 5
5a. Head and body covered (except ventrally) with small blue spots, about size of nostrils,
the distance between spots more than twice their diameter; median fins also covered
with blue spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectropomus leopardus
5b. Most blue spots on head and body more than twice size of nostrils; some spots on head
and body elongate (except juveniles) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Pelvic fins without blue spots; some spots on body of adults horizontally elongate; gill
raker at angle of first gill arch longer than longest gill filaments; pelvic fins 1.7 to 2.1 times
in head length; nostrils subequal . . . . . . . . . . . . . . . . . . . . . . . Plectropomus maculatus
6b. Pelvic fins with blue spots; some spots on body of adults vertically elongate; gill raker at
angle of first arch shorter than longest gill filaments; pelvic fins 1.9 to 2.3 times in head
length; rear nostrils of adults over 50 cm distinctly larger than anterior nostrils . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectropomus pessuliferus
Key to the species of Pseudanthias occurring in the area

1a. Dorsal and anal fins nearly covered by small scales; auxiliary scales (small scales
basally on large scales) present; third dorsal-fin spine longest, greatly elongated in
males (longer than head length) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Dorsal and anal fins naked or with only a few small scales basally; auxiliary scales
present or absent; third dorsal-fin spine, if longest, not longer than head . . . . . . . . . . . . . ® 3
2a. Band from eye to middle of pectoral fins (band yellow in females, orange in males) as
broad or broader than pupil and not distinctly edged in violet; outer edges of caudal-fin
lobes broadly edged in blue or violet; first 2 soft rays and adjacent membranes of pelvic
fins of males dark brownish red, in sharp contrast to pale rest of fins; spot on distal upper
part of pectoral fins of males brownish red and smaller than eye . . . . . . . . Pseudanthias huchtii
2b. Orange band from eye to middle of pectoral fins usually narrower than pupil and broadly
edged in violet; outer edge of caudal-fin lobes narrowly blue or violet; pelvic fins of males
not divided into outer brownish red and inner pale zones; spot on distal upper part of
pectoral fins of males pink to magenta, much larger than eye . . . . . . Pseudanthias squamipinnis
3a. Upper lip of males thickened at symphysis, forming a pointed fleshy protuberance (only
slightly developed in P. parvirostris); patch of villiform teeth on vomer roundish or
quadrangular; caudal fin of adults deeply forked . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3b. Upper lip of males not thickened at symphysis; vomerine tooth patch chevron-shaped or
subtriangular; caudal fin of adults slightly emarginate, slightly forked, or lunate . . . . . . . . . ® 13
4a. Lateral-line scales 44 to 52; pectoral-fin rays 15 to 19; a series of prominent papillae
along rear half of orbit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
4b. Lateral-line scales 53 to 64; pectoral-fin rays 18 to 22; papillae along rear half of orbit
present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
5a. Fourth to tenth dorsal-fin spines longest; 2 or 3 opercular spines . . . . . . . . . . . . . . . . . ® 6

5b. Third dorsal-fin spine longest; 3 opercular spines . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
6a. Lateral-line scales 41 to 44; no auxiliary scales; 3 opercular spines; snout length 4.6 to
5.6 times in head length; caudal fin yellow, the upper and lower edges violet . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias parvirostris
6b. Lateral-line scales 45 to 52; auxiliary scales present; 2 opercular spines; snout length
3.2 to 3.8 times in head length; caudal fin entirely violet or with yellow lobes . . . . . . . . . . . ® 7
7a. Lateral-line scales 45 to 49; pectoral-fin rays 15 to 17; females with a bright yellow band
dorsally on body, and caudal-fin lobes broadly yellow . . . . . . . . . . . . . . . Pseudanthias tuka
7b. Lateral-line scales 48 to 52 (rarely 48); pectoral-fin rays 16 to 19; no yellow band on
body, and no yellow on caudal fin . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias pascalus
8a. Opercular spines 2; dorsal-fin rays 15; body lavender-pink with 2 red stripes dorsally, the
lower as a series of contiguous blotches, and irregular yellow stripes on head which
break up into rows of small yellow spots on side of body . . . . . . . . . . Pseudanthias aurulentus
8b. Opercular spines 3; dorsal-fin rays 15 to 17; colour not as above . . . . . . . . . . . . . . . . . ® 9
9a. Lateral-line scales 44 to 48; body depth 3.1 to 3.3 times in standard length; body pink
with small yellow spots dorsally, shading ventrally to pale lavender; yellow stripe near
base of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias smithvanizi
9b. Lateral-line scales 49 to 52; body depth 3.4 to 4 times in standard length; body salmon
pink, shading to pinkish white ventrally, with a row of red blotches dorsally (except
anteriorly) and a broad red stripe along upper half of caudal peduncle . . . . . . . Pseudanthias lori
10a. No greatly elongated dorsal-fin spines (second dorsal-fin spine of males may be slightly
elongate); 2 opercular spines; rear half of orbit with several papillae; dorsal fin primarily
red . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias dispar
10b. Second and/or third dorsal-fin spines of adults greatly elongate; 3 opercular spines; no
orbital papillae; dorsal fin not primarily red . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
2466 Bony Fishes
11a. Second anal-fin spine slightly longer than third; second and third dorsal-fin spines
elongated in adults (the 2 spines nearly equal and with yellow membranous tips in
males); maximum number of preopercular serrae 36 (serrae increasing with size, in
general); maximum total length about 15 cm . . . . . . . . . . . . . . . . . . Pseudanthias bicolor
11b. Third anal-fin spine longer than second; second dorsal-fin spine elongated in adults (very
long in males); maximum number of preopercular serrae 24; maximum total length about
9 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Lateral-line scales 54 to 58; circumpeduncular scales 29 to 32; 3 to 5 irregular rows of
small teeth on palatines at widest place; pelvic fins of large males not reaching past
anal-fin base; dorsal and caudal fins yellow; head and body yellow dorsally, purple below
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias bartlettorum
12b. Lateral-line scales 56 to 62; circumpeduncular scales 32 to 36; 2 irregular rows of teeth
on palatines at widest place; pelvic fins of large males elongate, reaching past anal-fin
base; females orange-yellow, becoming red posteriorly; males violet, with top of head
and nape yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias regalis
13a. Anal-fin rays 9; first gill arch with 7 or 8 gill rakers on upper limb, 21 to 24 on lower limb;
pelvic fins elongated, in males reaching past base of anal fin; females lavender-pink with
broad yellow band dorsally on head and body and narrow magenta band across rear of
interorbital area; fins yellow with magenta margins; males with dorsal yellow area
invaded by lavender-magenta stripes and spots, and red and blue areas in dorsal and
caudal fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias ventralis
(Central Pacific to Great Barrier Reef; subspecifically different in Hawaii)
13b. Anal-fin rays 7 or 8; first gill arch with 8 to 12 gill rakers on upper limb, 22 to 29 on lower
limb; pelvic fins of males not reaching past anal-fin base; colour not as above . . . . . . . . . ® 14
14a. Pectoral-fin rays 15; dorsal-fin rays 14; lateral-line scales 35 (39 other side); pale brown
in alcohol with diffuse midlateral silvery stripe . . . . . . . . . . . . . . . Pseudanthias albofasciatus
(1 specimen, 75 mm standard length, taken off Hong Kong in 114 m; not yet recorded from the area)
14b. Pectoral-fin rays 16 to 20; dorsal-fin rays 15 to 18 (except P. hutomoi with 13 to 15 and
P. xanthomaculatus with 14); lateral-line scales 38 to 54 . . . . . . . . . . . . . . . . . . . . . ® 15
15a. First dorsal-fin spine as long as second spine; females lavender-pink, shading to yellow
posteriorly; median fins yellow; caudal fin of males orange, with large yellow elliptical
spot in basal half . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias pictilis
15b. First dorsal-fin spine distinctly shorter than second spine; caudal fin emarginate to
lunate; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
16a. Dorsal-fin rays 13 to 15 (modally 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
16b. Dorsal-fin rays 14 to 18 (none modally 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
17a. Eighth to thirteenth dorsal-fin rays with a single branch projecting as a short filament free
of membrane; eye diameter 2.8 or more times in head length; body with 3 small white
blotches dorsally, the first below last dorsal-fin spine; males develop a large irregular
whitish area below rear dorsal-fin rays . . . . . . . . . . . . . . . . . . . . . Pseudanthias hutomoi
17b. Posterior dorsal-fin rays without free filamentous tips; eye diameter 2.6 times in head
length; body rose with yellow blotch dorsally below middle of dorsal fin and another
behind eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias xanthomaculatus
(1 specimen, 45 mm standard length, taken off New Caledonia in 200 m)
18a. Third dorsal-fin spine of adults elongate, with a membranous filament at tip, 1.2 to 1.5
times in head length (spine of males, including filament, as long or longer than head
length); dorsal profile of head of males straight; females orange, with dusky yellow spot
on scales of upper two-thirds of body, the snout and chin yellow, the caudal fin yellow
except orange at base and violet on outer edges of lobes; males fuchsia with 2 broad
orange stripes, the dorsal, anal, and pelvic fins with a broad deep orange submarginal
band (on anal fin only anteriorly) . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias randalli
18b. Third dorsal-fin spine more than 1.5 times in head length; dorsal profile of head of males
slightly to moderately convex; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . ® 19
19a. Lateral-line scales 38 to 43; dorsal-fin rays 15 or 16 . . . . . . . . . . . . . . . . . . . . . . . ® 20

19b. Lateral-line scales 41 to 54; dorsal-fin rays 16 to 18 . . . . . . . . . . . . . . . . . . . . . . . ® 21
20a. Body depth 3.2 times in standard length; pectoral-fi n rays 18 or 19; first gill arch with 12
gill rakers on upper limb, 26 on lower limb; body orange-red anteriorly, shading to yellow
posteriorly and on caudal fin; narrow rose band from snout to subopercle . . . Pseudanthias cichlops
(1 specimen, 72 mm standard length, from Sumatra)
20b. Body depth 2.6 to 2.9 times in standard length; pectoral-fin rays 17 or 18; first gill arch
with 9 to 11 gill rakers on upper limb, 22 to 25 on lower limb; pinkish orange with a pale
yellow stripe, edged in violet, from front of upper lip through lower part of eye to
pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias lunulatus
21a. Third dorsal-fin spine clearly longest (at least in males and large females) . . . . . . . . . . . ® 22
21b. Third dorsal-fin spine not elongated (may be equal to longest spine) . . . . . . . . . . . . . . ® 26
22a. Females with midlateral lavender-edged orange-red band from eye to caudal fin, body
below band with alternating yellow and lavender-pink lines along scale rows, fins yellow;
males yellowish, the front third of body reddish; soft dorsal, anal, and caudal fins with
blue margin; lavender-edged red band from eye to pectoral-fin base; lateral-line scales
41 to 45; maximum size about 21 cm . . . . . . . . . . . . . . . . . . . . . Pseudanthias fasciatus
22b. Colour not as above; lateral-line scales 42 to 50; maximum size 15 cm . . . . . . . . . . . . . ® 23
23a. Lateral-line scales 42 to 45; pectoral-fin rays 17; lower limb of first gill arch with 24 or 25
gill rakers; body lavender-pink, the scale centres yellow; fins pale lavender, suffused with
yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias engelhardi
23b. Lateral-line scales 44 to 50; pectoral-fin rays 17 to 19; lower limb of first gill arch with 24
to 29 gill rakers; body and/or fins with distinctive colour pattern, at least in males . . . . . . . . ® 24
24a. Pelvic fins not reaching anal fin; dorsal-fin rays 15 to 17; lateral-line scales 44 to 48; lower
limb of first gill arch with 24 to 27 gill rakers; female pale orange-red, the scale centres
yellow to dusky yellow, the fins pale yellow; male with yellow-orange lines or rows of
spots along dorsal scale rows of body, red spot in rear part of spinous dorsal fin, and
broad red border on caudal fin . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias luzonensis
24b. Pelvic fins reaching to or beyond anal-fin origin; dorsal-fin rays 16 to 18; lateral-line
scales 45 to 50; lower limb of first gill arch with 25 to 29 gill rakers; colour not as above . . . . ® 25
25a. Body of females yellow, the scale edges magenta, with 2 magenta lines from eye to
pectoral-fin base thence to lower caudal peduncle; males orange-magenta with a large,
nearly square, lavender blotch on side of body preceded by a broad orange band curving
dorsally to nape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias pleurotaenia
25b. Coloration of females similar as above, but lighter yellow, and ventral part of body pale
lavender; males also similarly coloured, but no lavender blotch on body . . . . . Pseudanthias sheni
26a. Lateral-line scales 50 to 54; female brownish yellow, shading to pale lavender ventrally,
the scale centres pink, the dorsal-fin margin broadly lavender-pink; male
lavender-orange, with narrow orange bar below eighth dorsal-fin spine between pectoral
fins and lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias mooreanus
26b. Lateral-line scales 43 to 50; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . ® 27
27a. Female with emarginate caudal fin, the caudal concavity 4 to 6 times in head length, the
rear margin red; caudal fin of male convex to truncate, pink, shading distally to red, the
corners lavender; male with red spot between seventh and tenth dorsal-fin spines;
maximum length 19 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias hypselosoma
27b. Caudal fin emarginate to lunate, the caudal concavity 1 to 5 times in head length; colour
not as above; maximum length about 16 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 28
2468 Bony Fishes
28a. Dorsal-fin rays 15 or 16; caudal fin emarginate, the caudal concavity 2.3 to 4.6 times in
head length; body depth 2.4 to 2.9 times in standard length; female pink, the scale
centres yellow, forming irregular oblique lines on upper rear part of body; median fins
yellow; male lavender-pink, the scales edged with orange (especially dorsally), the body
ventrally with faint yellow stripes; caudal fin with irregular bars of orange-yellow and
violet, the corners violet-blue with yellow tips; dorsal fin with a dark red spot on fourth
and fifth interspinous membranes . . . . . . . . . . . . . . . . . . . . . . Pseudanthias bimaculatus
28b. Dorsal-fin rays 16 to 18; caudal fin emarginate to lunate, the concavity 0.8 to 2.6 times
in head length; body depth 2.7 to 3.2 times in standard length; colour not as above . . . . . . . ® 29
29a. Last dorsal-fin spine usually longest (but fourth to last subequal); lateral-line scales 46
to 52; scales dusky yellow and lavender-pink; pale lavender line from below eye to lower
pectoral-fin base; head and chest below this line whitish to lavender; first 4 dorsal-fin
membranes fuchsia with broad oblique orange band (more evident in males); males with
orange-red bar below lateral line and under ninth dorsal-fin spine; caudal-fin tips of
females bright red; caudal fin of male largely red, the filamentous tips yellow and
lavender . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias cooperi
29b. Fourth dorsal-fin spine longest (but adjacent spines nearly as long); lateral-line scales
42 to 49; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 30
30a. No auxiliary scales; dorsal-fin rays 16; lateral-line scales 42 to 47; female pink with
yellow spot or line on each scale, shading ventrally to whitish; rear edge of caudal fin
narrowly red, this colour broadening onto pointed lobe tips; male with a lavender-edged
red bar from below last 4 dorsal-fin spines to beneath pectoral fins, the body anterior to
bar, lavender-pink, posterior to bar, yellow shading dorsally to lavender-pink; caudal fin
yellow, upper and lower edges lavender, the lobe tips red . . . . . . . . . Pseudanthias rubrizonatus
30b. Auxiliary scales present on body; dorsal-fin rays 17 or 18; lateral-line scales 44 to 49;
female olive-grey, with irregular horizontal rows of small orange-yellow spots laterally
and ventrally; front of snout and chin reddish orange; dorsal fin orange-red distally;
caudal fin yellow; male dark olive-grey with 3 orange squares in a row on front half of
body, followed by horizontal series of small orange spots; caudal fin yellow with orange
lobes edged with blue distally; dark red (almost black) spot distally in upper part of
pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudanthias olivaceus
Key to the species of Pseudogramma occurring in the area

1a. Dermal tentacle on upper surface of eye; rows of red dots extending posteriorly from eye
and forming a circle opercular ocellus . . . . . . . . . . . . . . . . . . . . Pseudogramma australis
1b. No tentacle on top of eye; no rows of red dots behind eye . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Pectoral-fin rays 13, the middle rays unbranched; caudal-peduncle length (from anal-fin
base to caudal-fin base) 6.2 to 6.9 times in head length; first gill arch with 10 gill rakers
on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Pectoral-fin rays 14 to 18, the rays branched except upper and lowermost rays of some
specimens; caudal-peduncle length 4.3 to 5.5 times in head length; first gill arch with 10
to 13 gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Lateral scale series 43; pectoral-fin length 3.3 times in standard length; anal-fin rays 17;
body red, with pale pink blotches . . . . . . . . . . . . . . . . . . . . . . Pseudogramma erythreum
3b. Lateral scale series 52 or 53; pectoral-fin length 2.6 to 2.9 times in standard length;
anal-fin rays 18 or 19; body yellow, with a faint brown reticulum . . . . . . Pseudogramma pectoralis
4a. Dark brown spot on opercle; lateral-line scales of adults 31 to 41; rear ends of dorsal and
anal fins not reaching past vertical at caudal-fin base; pectoral fins not reaching vertical
at anal-fin origin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogramma polyacantha
4b. No dark spot on opercle; lateral-line scales 21 to 26; rear ends of dorsal and anal fins
project well past vertical at caudal-fin base; pectoral fins reaching past anal-fin origin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudogramma astigmum
Key to the species of Sacura occurring in the area

1a. Dorsal fin with X spines and 14 soft rays; body depth 2.2 to 2.3 times and pectoral fins
2.7 to 2.9 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sacura parva
(2 specimens, 17 and 67 mm standard length, taken in Timor Sea in 130 m)
1b. Dorsal fin with X spines and 15 soft rays; body depth 2.0 to 2.1 times and pectoral fins
2.3 to 2.5 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . Sacura speciosa
(1 specimen, 95 mm, off Sulawesi in 150 m)
Key to the species of Selenanthias occurring in the area
1a. Bony interorbital space wide, 3.4 to 3.8 times in head length; lower limb of first gill arch
with 19 to 21 gill rakers; large black spot at rear of anal fin . . . . . . . . . . . . Selenanthias analis
1b. Interorbital width 4.3 to 5.0 times in head length; lower limb of first gill arch with 22 to 24
gill rakers; no black spot in anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Body depth about 2.7 times in standard length; interorbital width 4.7 to 5.0 times in head
length; oblique rows of large cheek scales 7; body rose, with dark orange spot at base
of last 4 dorsal-fin spines; head orange, with oblique rose band across cheek . . Selenanthias barroi
2b. Body depth 2.3 to 2.6 times in standard length; interorbital width 4.3 to 4.5 times in head
length; cheek scale rows 6; life colour unknown . . . . . . . . . . . . . . . . . . Selenanthias myersi
Key to the species of Variola occurring in the area

1a. Rear margin of caudal fin with a black submarginal line and narrow white edge; dorsal,
anal, and pectoral fins without a distinct yellow posterior border; 13 to 16 gill rakers on
lower limb of first gill arch; pelvic fins usually not reaching anus; juveniles without a dark
stripe on body dorsally and no dark spot at base of upper caudal-fin rays . . . Variola albimarginata
1b. Caudal, dorsal, anal, and pectoral fins with a broad yellow rear margin; 15 to 18 gill
rakers on lower limb of first gill arch; pelvic fins reach beyond anus; juveniles with
irregular dark brown stripe along dorsal part of body and a dark spot at base of upper
caudal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Variola louti

The symbol 0is given when species accounts are included. A question mark indicates that presence in
Subfamily ANTHIINAE
Acanthistius ocellatus (Günther, 1859)
Caprodon schlegelii (Günther, 1859)1/
Dactylanthias haplodactylus (Bleeker, 1858) 2/
Holanthias borbonius (Valenciennes, 1828)
Holanthias chrysostictus (Günther, 1871)
(= Holanthias katayamai Randall, Maugé, and Plessis, 1979)
Holanthias rhodopeplus (Günther, 1871)
Holanthias tapui Randall, Maugé, and Plessis, 1979
Holanthias unimaculatus (Tanaka, 1917)
Hypoplectrodes annulatus (Günther, 1859)
Hypoplectrodes jamesoni (Ogilby, 1908)
Hypoplectrodes maccullochi Whitley, 1929
Luzonichthys earlei Randall, 1981
Luzonichthys taeniatus Randall and McCosker, 1992
Luzonichthys waitei (Fowler, 1931)
Luzonichthys whitleyi (Smith, 1955)
Luzonichthys williamsi Randall and McCosker, 1992
1/ Not yet known from the area, but should be expected; occurs in New South Wales at 31°51’S, Western Australia at
24°25’S, Taiwan Province of China, and Hawaii.
2/ Known from a single specimen (17 cm standard length) from Ambon, Indonesia.
2470 Bony Fishes
Plectranthias anthioides (Günther, 1871)

(= Plectranthias kamii Randall, 1980)
Plectranthias cirrhitoides Randall, 1980
Plectranthias foresti Fourmanoir, 1977
Plectranthias fourmanoiri Randall, 1980
Plectranthias inermis Randall, 1980
Plectranthias japonicus (Steindachner, 1884)
Plectranthias jothyi Randall, 1996
Plectranthias kelloggi (Jordan and Evermann, 1903)
Plectranthias knappi Randall, 1996
Plectranthias lasti Randall and Hoese, 1995
Plectranthias longimanus (Weber, 1913)
Plectranthias megalepis (Günther, 1880)
Plectranthias megalophthalmus (Fourmanoir and Randall, 1979)
Plectranthias nanus Randall, 1980
Plectranthias pallidus Randall and Hoese, 1980
Plectranthias retrofasciatus Randall and Fourmanoir, 1979
Plectranthias robertsi Randall and Hoese, 1980
Plectranthias rubrifasciatus Randall and Fourmanoir, 1979
Plectranthias sagamiensis (Katayama, 1963)
Plectranthias taylori Randall, 1980
Plectranthias wheeleri Randall, 1980
Plectranthias whiteheadi Randall, 1980
(= Plectranthias chungchowensis Shen and Lin, 1984)
Plectranthias winniensis (Tyler, 1966)
? Pseudanthias albofasciatus (Fowler and Bean, 1930)
Pseudanthias aurulentus Randall and McCosker, 1981
Pseudanthias bartlettorum Randall and Lubbock, 1981
Pseudanthias bicolor Randall, 1979
Pseudanthias bimaculatus (Smith, 1955)
Pseudanthias cichlops (Bleeker, 1853)
Pseudanthias cooperi (Regan, 1902)
(= Anthias kashiwae Tanaka, 1918)
Pseudanthias dispar (Herre, 1955)
Pseudanthias fasciatus (Kamohara, 1954)
(= Anthias rubrolineatus Fourmanoir and Rivaton, 1979)
Pseudanthias huchtii (Bleeker, 1857)
Pseudanthias hutomoi (Allen and Burhanuddin, 1976)
Pseudanthias hypselosoma Bleeker, 1878
(= Anthias truncatus Katayama and Masuda, 1983)
Pseudanthias lori Randall and Lubbock, 1981
Pseudanthias luzonensis (Katayama and Masuda, 1983)
Pseudanthias mooreanus (Herre, 1935)
Pseudanthias olivaceus Randall and McCosker, 1981
Pseudanthias parvirostris Randall and Lubbock, 1981
Pseudanthias pascalus (Jordan and Tanaka, 1927)
Pseudanthias pictilis (Randall and Allen, 1978)
Pseudanthias pleurotaenia (Bleeker, 1857)
Pseudanthias randalli (Lubbock and Allen, 1978)
Pseudanthias regalis Randall and Lubbock, 1981
Pseudanthias rubrizonatus Randall, 1983
Pseudanthias sheni Randall and Allen, 1989
Pseudanthias smithvanizi Randall and Lubbock, 1981
Pseudanthias squamipinnis (Peters, 1855)
Pseudanthias tuka (Herre and Montalban, 1927)
Pseudanthias ventralis Randall, 1979
Pseudanthias xanthomaculatus (Fourmanoir and Rivaton, 1979)
Rabaulichthys altipinnis Allen, 1984
Sacura parva Heemstra and Randall, 1979

Sacura speciosa Heemstra and Randall, 1979
Selenanthias analis Tanaka, 1918
(= Plectranthias maculatus Fourmanoir, 1982)
Selenanthias barroi (Fourmanoir, 1982)
Selenanthias myersi Randall, 1995
Serranocirrhitus latus Watanabe, 1949
Tosana niwae Smith and Pope, 1906
Tosanoides flavofasciatus Katayama and Masuda, 1980
Subfamily EPINEPHELINAE
0
Tribe EPINEPHELINI
Aethaloperca rogaa (Forsskål, 1775)
0 Anyperodon leucogrammicus (Valenciennes, 1828)
0 Cephalopholis aitha Randall and Heemstra, 1991
0 Cephalopholis argus Bloch and Schneider, 1801
0 Cephalopholis aurantia (Valenciennes, 1828)
0 Cephalopholis boenak (Bloch, 1790)
0 Cephalopholis cyanostigma (Valenciennes, 1828)
0 Cephalopholis formosa (Shaw and Nodder, 1812)
0 Cephalopholis igarashiensis Katayama, 1957
0 Cephalopholis leopardus (Lacepède, 1801)
0 Cephalopholis microprion (Bleeker, 1852)
0 Cephalopholis miniata (Forsskål, 1775)
0 Cephalopholis polleni (Bleeker, 1868)
0 Cephalopholis sexmaculata (Rüppell, 1830)
0 Cephalopholis sonnerati (Valenciennes, 1828)
0 Cephalopholis spiloparaea (Valenciennes, 1828)
0 Cephalopholis urodeta (Bloch and Schneider, 1801)
0 Cromileptes altivelis (Valenciennes, 1828)
0 ? Epinephelus akaara (Temminck and Schlegel, 1842)
0 Epinephelus amblycephalus (Bleeker, 1857)
0 Epinephelus areolatus (Forsskål, 1775)
0 Epinephelus awoara (Temminck and Schlegel, 1842)
0 Epinephelus bilobatus Randall and Allen, 1987
0 Epinephelus bleekeri (Vaillant, 1877)
0 Epinephelus bontoides (Bleeker, 1855)
0 Epinephelus bruneus Bloch, 1793
0 Epinephelus caeruleopunctatus (Bloch, 1790)
0 Epinephelus chlorostigma (Valenciennes, 1828)
0 Epinephelus coioides (Hamilton, 1822)
0 Epinephelus corallicola (Valenciennes, 1828)
0 Epinephelus cyanopodus (Richardson, 1846)
0 Epinephelus daemelii (Günther, 1876)
0 Epinephelus darwinensis Randall and Heemstra, 1991
0 Epinephelus epistictus (Temminck and Schlegel, 1842)
0 Epinephelus ergastularius Whitley, 1930
0 Epinephelus erythrurus (Valenciennes, 1828)
0 Epinephelus fasciatomaculosus (Peters, 1866)
0 Epinephelus fasciatus (Forsskål, 1775)
0 Epinephelus faveatus (Valenciennes, 1828)
0 Epinephelus fuscoguttatus (Forsskål, 1775)
0 Epinephelus heniochus Fowler, 1904
0 Epinephelus hexagonatus (Forster, 1801)
0 Epinephelus howlandi (Günther, 1873)
2472 Bony Fishes
0 Epinephelus irroratus (Forster, 1901)

0 Epinephelus lanceolatus (Bloch, 1790)
0 Epinephelus latifasciatus (Temminck and Schlegel, 1842)
0 Epinephelus longispinis (Kner, 1864)
0 Epinephelus macrospilos (Bleeker, 1855)
0 Epinephelus maculatus (Bloch, 1790)
0 Epinephelus magniscuttis Postel, Fourmanoir, and Guézé, 1963
0 Epinephelus malabaricus (Bloch and Schneider, 1801)
0 Epinephelus melanostigma Schultz, 1953
0 Epinephelus merra Bloch, 1793
0 Epinephelus miliaris (Valenciennes, 1830)
0 Epinephelus morrhua (Valenciennes, 1833)
0 Epinephelus multinotatus (Peters, 1876)
0 Epinephelus octofasciatus Griffin, 1926
0 Epinephelus ongus (Bloch, 1790)
0 Epinephelus perplexus Randall, Last, and Hoese, 1991
0 Epinephelus poecilonotus (Temminck and Schlegel, 1842)
0 Epinephelus polypekadion (Bleeker, 1849)
0 Epinephelus polystigma (Bleeker, 1853)
0 Epinephelus quoyanus (Valenciennes, 1830)
0 Epinephelus radiatus (Day, 1867)
0 Epinephelus retouti Bleeker, 1868
0 Epinephelus rivulatus (Valenciennes, 1830)
0 Epinephelus sexfasciatus (Valenciennes, 1828)
0 Epinephelus socialis (Günther, 1873)
0 Epinephelus spilotoceps Schlultz, 1953
0 Epinephelus stictus Randall and Allen, 1967
0 Epinephelus tauvina (Forsskål, 1775)
0 Epinephelus timorensis Randall and Allen, 1987
0 Epinephelus trophis Randall and Allen, 1987
? Epinephelus trimaculatus (Valenciennes, 1828)
0 Epinephelus tuamotuensis Fourmanoir, 1971

0 Epinephelus tukula Morgans, 1959
0 Epinephelus undulatostriatus (Peters, 1867)
0 Epinephelus undulosus (Quoy and Gaimard, 1824)
0 Gracila albomarginata (Fowler and Bean, 1930)
0 Plectropomus areolatus (Rüppell, 1830)
0 Plectropomus laevis (Lacepède, 1801)
0 Plectropomus leopardus (Lacepède, 1802)
0 Plectropomus maculatus (Bloch, 1790)
0 Plectropomus oligacanthus Bleeker, 1854
0 Plectropomus pessuliferus Fowler, 1904
0 Saloptia powelli Smith, 1964
0 Triso dermopterus (Temminck and Schlegel, 1842)
0 Variola albimarginata Baissac, 1952
0 Variola louti (Forsskål, 1775)
0 Niphon spinosus Cuvier, 1828
Tribe NIPHONINI
Tribe LIOPROPOMATINI
Liopropoma collettei Randall and Taylor, 1988
Liopropoma erythraeum Randall and Taylor, 1988
Liopropoma flavidum Randall and Taylor, 1988
Liopropoma incomptum Randall and Taylor, 1988
Liopropoma lunulatum (Guichenot, 1862)
Liopropoma mitratum Lubbock and Randall, 1978
Liopropoma multilineatum Randall and Taylor, 1988
Liopropoma pallidum (Fowler, 1938)

Liopropoma susumi (Jordan and Seale, 1906)
Liopropoma swalesi Fowler and Bean, 1930
Liopropoma tonstrinum Randall and Taylor, 1988
Rainfordia opercularis McCulloch, 1923
Tribe DIPLOPRIONI
Aulacocephalus temmincki Bleeker, 1857
Belonoperca chabanaudi Fowler and Bean, 1930
Diploprion bifasciatum Kuhl and van Hasselt, 1828
Tribe GRAMMISTINI
Aporops bilinearis Schultz, 1943
Grammistes sexlineatus (Thunberg, 1792)
Grammistops ocellatus Schultz, 1953
Pogonoperca punctata Valenciennes, 1830)
Pseudogramma astigmum Randall and Baldwin, 1997
Pseudogramma australis australis Randall and Balwin, 1997
Pseudogramma erythreum Randall and Baldwin, 1997
Pseudogramma pectoralis Randall and Baldwin, 1997
Pseudogramma polyacantha (Bleeker, 1856)
Suttonia lineata Gosline, 1960
Subfamily SERRANINAE
Chelidoperca hirundinacea (Valenciennes, 1831)
Chelidoperca margaritifera Weber, 1913
Chelidoperca pleurospilos (Günther, 1860)
References
Baldwin, C.C. and G.D. Johnson. 1993. Phylogeny of the Epinephelinae (Teleostei: Serranidae). Bull. Mar. Sci.,
52(1):240-283.
Heemstra, P.C. and D. Golani. 1993. Clarification of the Indo-Pacific groupers (Pisces: Serranidae) in the Mediterranean
Sea. Isr. J. Zool. , 39:381-390.
Heemstra, P.C. and J.E. Randall. 1993. FAO species catalogue. Vol. 16. Groupers of the world (family Serranidae,
subfamily Epinephelidae). An annotated and illustrated catalogue of the grouper, rockcod, hind, coral grouper
and lyretail species known to date. FAO Fish. Synop., (125)16:382 p.
Randall, J.E. 1980. Revision of the fish genus Plectranthias (Serranidae: Anthiinae), with descriptions of 13 new species.
Micronesica, 16(1):101-187.
Randall, J.E. 1996. Two new anthiine fishes of the genus Plectranthias (Perciformes: Serranidae), with a key to the
species. Micronesica, 29(2):113-131.
Randall, J.E. and C.C. Baldwin. 1997. Revision of the serranid fishes of the subtribe Pseudogrammina, with descriptions
of five new species. Indo-Pac. Fishes, (26):56 p.
Randall, J.E. and P.C. Heemstra. 1978. Reclassification of the Japanese cirrhitid fishes Serranocirrhitus latus and
Isobuna japonica to the Anthiinae. Japan. J. Ichthyol., 25(3):165-172.
Randall, J.E. and P.C. Heemstra. 1991. Revision of Indo-Pacific groupers (Perciformes: Serranidae: Epinephelinae),
with descriptions of five new species. Indo-Pac. Fishes, (20):296 p.
Randall, J.E. and R. Lubbock. 1981. A revision of the serranid fishes of the subgenus Mirolabrichthys (Anthiinae:
Anthias), with descriptions of five new species. Natural History Museum of Los Angeles County, Contributions
in Science, (333):27 p.
Randall, J.E. and J.E. McCosker. 1992. Revision of the fish genus Luzonichthys (Perciformes: Serranidae: Anthiinae),
with descriptions of two new species. Indo-Pac. Fishes, 21:1-21.
Randall, J.E. and L. Taylor. 1988. Review of the Indo-Pacific fishes of the serranid genus Liopropoma, with descriptions
of seven new species. Indo-Pac. Fishes, (16):47 p.
Randall, J.E., L.A. Mauge, and Y.B. Plessis. 1979. Two new anthiine fishes of the genus Holanthias from the southern
and western Pacific. Japan J. Ichthyol., 26(1):15-25.
The illustrations on the following species accounts that are signed DPV, EG, EH, and MMS are copyright of the J.L.B.
Smith Institute of Ichthyology.
click for next page

2474 Bony Fishes
Subfamily EPINEPHELINAE
Tribe EPINEPHELINI
Aethaloperca rogaa (Forsskål, 1775) (Plate I, 1)

Frequent synonyms / misidentifications: Cephalopholis rogaa (Forsskål, 1775) / None (rarely misiden-
tified).
FAO names: En - Redmouth grouper; Fr - Vielle roga; Sp - Cherna roga.
45 cm total length
Diagnostic characters: Body deep and compressed, its depth greater than head length and 2.1 to
2.4 times in standard length; body width 2.3 to 2.8 times in body depth; head length 2.5 to 2.75 times in
standard length. Dorsal head profile steep, straight, or slightly concave along snout and distinctly
convex from eye to dorsal fin; preorbital depth 6.5 to 9.2 times in head length; preopercle finely serrate,
the lower edge fleshy; rear nostrils round or oval, not much larger than front ones. Maxilla reaches past
eye; small canines at front of jaws, none elsewhere; 2 to 4 rows of small slender teeth at side of lower jaw;
palatine teeth present; ventroposterior corner of maxilla with a distinct bony protuberance; supramaxilla
slender. First gill arch with 8 to 10 gill rakers on upper limb, 15 to 17 on lower limb; longest raker slightly
longer than longest gill filaments. Dorsal fin with IX spines and 17 or 18 soft rays, the fin origin over opercle,
and the third or fourth spines longest; dorsal-fin membranes slightly incised between the spines; anal fin
with III spines and 8 or 9 soft rays; middle dorsal and anal-fin rays elongated in adults, giving these
fins an angular profile, with the rear margin almost vertical; caudal fin truncate, with 8 branched
rays in upper lobe and 7 in lower lobe; pectoral fins asymmetric, with 17 to 19 rays; pelvic fins subequal
to pectoral fins, extending beyond anus. Scales on body rough, with auxiliary scales; lateral-line scales 48
to 54; lateral scale series 94 to 104. Colour: dark brown to black, occasionally with an orange cast,
usually with a pale vertical bar on side of abdomen; distal part of spinous dorsal fin dark orange to
brownish red; inside of mouth, gill cavity and upper jaw membranes reddish orange. Juveniles with
a broad white posterior margin on caudal fin and a narrow white margin on soft dorsal fin.
Habitat, biology, and fisheries: Well-developed coral reefs in depths of 3 to at least 60 m. Usually seen
in or near caves and holes in the reef. Small fishes (including Pempheris sp., a common cave dweller) are
the primary food of this species, and it also feeds on stomatopods (Pseudosquilla sp.). Preliminary data
indicate that it spawns at any time of the year and matures (females?) at a length of about 35 cm standard
length. Occasionally seen in markets, but
this species is not common. Caught with
hook-and-line, spear, and probably in traps.
Distribution: Red Sea to South Africa and
east to the Gilbert Islands in the Central
Pacific; including Thailand, Japan, Philip-
pines, Indonesia, Australia, Papua New
Guinea, Palau, as well as Solomon,
Caroline, and Gilbert islands.
Anyperodon leucogrammicus (Valenciennes, 1828) (Plate I, 2)

Frequent synonyms / misidentifications: Serranus micronotatus Rüppell, 1830; S. urophthalmus
FAO names: En - Slender grouper; Fr - Mérou élégant; Sp - Mero elegante.
29.4 cm standard length
Diagnostic characters: Body and head elongate

and markedly compressed, its depth distinctly
less than head length, 3.1 to 3.7 times in standard
length; body width 2.3 to 2.8 times in body depth.
Head pointed, its length 2.3 to 2.5 times in standard
length; dorsal head profile almost straight, the interor- juvenile
bital region slightly concave, flat or slightly convex; 8 cm total length
preorbital narrow, its depth 14 to 17 times in head (from Randall and Kuiter, 1989)
length; preopercle rounded, finely serrate, the lower serrae only slightly enlarged, the lower edge fleshy;
ventral edge of interopercle with a shallow indentation; upper edge of operculum distinctly convex; diameter
of rear nostrils about twice that of anteriors. Maxilla reaches well past eye, the exposed part covered with
tiny scales; no teeth on palatines; canines at front of jaws rudimentary or absent; teeth at sides of lower
jaw subequal, in 2 or 3 irregular rows; no bony process on rear end of maxilla; supramaxilla well developed.
First gill arch with 7 to 9 gill rakers on upper limb, 14 to 17 on lower limb; rakers subequal to gill filaments
in length. Dorsal fin with XI spines and 14 to 16 soft rays, the fin origin over the opercle, the membranes
distinctly incised between the spines; anal fin with III spines and 8 or 9 soft rays; caudal fin rounded, with
8 branched rays in upper lobe and 7 in lower lobe; pectoral fins short, thin, transparent, and symmetrical,
with 15 to 17 rays; pelvic fins not reaching anus. Midlateral body scales rough; lateral-line scales 61 to 72;
lateral scale series 106 to 125; auxiliary scales present on body scales. Colour: adults greenish to
brownish grey with numerous orange-red spots on head (becoming smaller anteriorly), body, dorsal
fin, and basally on caudal fin; 4 longitudinal whitish bands or series of streaks often visible on
postorbital head and body; membranes of soft dorsal, anal, and caudal fins clear. Juveniles with
longitudinal pale bluish grey stripes on a gold background; a blue-edged black spot (or double
spot) at base of caudal fin and another at base of soft dorsal fin.
Size: Maximum total length at least 60 cm.
Habitat, biology, and fisheries: A coral-reef species usually found on protected reefs in depths of 5 to
80 m. Adults primarily piscivorous. The distinctive blue and gold striped juveniles are mimics of the wrasse
H a l i c h o e r es purpurescens
(Bloch and Schneider, 1801).
Often seen in markets but not
plentiful enough to be of com-
mercial importance. Caught
with hook-and-line, spear,
and probably in traps.
Distribution: R e d S e a to
Mozambique and east to Sa-
moa and the Phoenix Islands;
including Japan, Philippines,
Indonesia, Australia, and
Papua New Guinea.
2476 Bony Fishes
Cephalopholis argus Bloch and Schneider, 1801 (Plate I, 3)

Frequent synonyms / misidentifications: None / Cephalopholis cyanostigma (non Valenciennes, 1828).
FAO names: En - Peacock hind; Fr - Vielle ciosomoer; Sp - Cherna pavo real.
22 cm standard length
Diagnostic characters: Body depth distinctly less than head length, 2.7 to 3.2 times in standard length
(for specimens 10 to 40 cm standard length); head length 2.4 to 2.7 times in standard length. Eye small,
its diameter distinctly less than snout length; interorbital area flat to slightly convex; preopercle rounded,
finely serrate in young, virtually smooth in large adults, the lower edge fleshy; subopercle and interopercle
smooth. Maxilla scaly, reaching well past eye. First gill arch with 9 to 11 gill rakers on upper limb, 17 to 19
on lower limb; rudimentary rakers on lower limb difficult to distinguish from intercalated bony plates. Dorsal
fin with IX spines and 15 to 17 soft rays, the membranes indented between the spines; anal fin with
III spines and 9 soft rays; pectoral-fin rays 16 to 18; pectoral fins clearly longer than pelvic fins, 1.4 to
2.0 times in head length; pelvic fins not reaching anus, 1.9 to 2.4 times in head length; caudal fin well
rounded. Lateral body scales rough, with a few auxiliary scales in adults; lateral-line scales 46 to 51; lateral
scale series 95 to 110. Colour: dark brown, covered with small black-edged blue ocelli; often with 5
or 6 pale bars on rear part of body and a large pale area over the chest; rear margin of median fins
usually with a narrow white edge; distal part of pectoral fins sometimes maroon brown; triangular
membranes at tips of dorsal-fin spines orange-gold.
Habitat, biology, and fisheries: A common tropical species found in a variety of coral reef habitats from
tide pools to depths of at least 40 m. In the Gulf of Aqaba, Cephalopholis argus is found in social units
comprising up to 12 adults, including 1 dominant male. Each group occupies a specific area (up to
2 000 m2) that is defended by the territorial male and subdivided into secondary territories, each inhabited
by a single female. Primarily (75 to 95%) piscivorous, C. argus has been blamed for numerous cases of
ciguatera in the Pacific region. This common and widespread species is important to artisanal fisheries
throughout the Indo-West Pacific region. Caught with hook-and-line, spear, and in traps.
Distribution: C. argus is the
most widely distributed of the
groupers, occurring from the
Red Sea to South Africa and
east to French Polynesia and
the Pitcairn Group, including
Japan, northern Australia,
Lord Howe Island, Marque-
sas, and throughout Microne-
sia. Recently introduced to
Hawaii.
Cephalopholis cyanostigma (Valenciennes, 1828) (Plate I, 4)

Frequent synonyms / misidentifications: Cephalopholis kendalli Evermann and Seale, 1907;
C. xanthopterus Allen and Stark, 1975 / Cephalopholis miniata (non Forsskål, 1775); C. argus non Bloch
and Schneider, 1801; C. microprion (non Bleeker, 1852).
FAO names: En - Bluespotted hind; Fr - Vielle étoiles bleues; Sp - Cherna de pintas azules.
9.3 cm standard length juveniles

Diagnostic characters: Body somewhat compressed, its width 2.0 to 2.3 times in body depth; body depth
less than head length, 2.6 to 3.0 times in standard length (for specimens 8 to 23 cm standard length); head
length 2.3 to 2.6 times in standard length. Interorbital area flat to slightly convex, its width subequal to eye
diameter and maxilla width; preopercle rounded, finely serrate, the lower edge fleshy; subopercle and
interopercle smooth or with a few serrae; upper edge of operculum very convex, the rear edge almost
vertical; maxilla scaly, reaching well past eye. First gill arch with 7 to 9 gill rakers on upper limb, 14 to 18
on lower limb. Dorsal fin with IX spines and 15 to 17 soft rays, the fin membrane indented between the
spines; anal fin with III spines and 8 soft rays; caudal fin well rounded; pectoral-fin rays 15 to 18, the
fin length distinctly longer than pelvic-fin length, 1.5 to 1.8 times in head length; pelvic fins usually not
reaching anus, 1.8 to 2.1 times in head length. Body scales rough, with a few auxiliary scales in adults;
lateral-line scales 46 to 50; lateral scale series 92 to 106. Colour: adults brown to brownish red, with
numerous small black-edged blue ocelli on head, body, and fins, those on head and chest larger
and more distinctly black-edged than those on upper part of body, those on fins smaller (except
basally on pectoral fins) and also distinctly black-edged (spots entirely dark in preservative, the
ones on body faint); body often with dark chain-like bars; proximal two-thirds of pectoral fins with blue
ocelli becoming smaller distally, the distal part of fin orange-yellow with a blackish edge or white edge and
black submarginal line; median fins darker than body, the rear margin bluish white with a submarginal
blackish band. Juveniles with head, body, and pectoral fins dark brown, the other fins bright yellow; blue
ocelli on head and front of body indistinct or absent.
Habitat, biology, and fisheries: Coral reefs
and seagrass or algal beds at depths of 1 to
50 m. Feeds on crustaceans and fishes.
Probably of importance to artisanal fisheries.
Caught with hook-and-line, gill nets, spear,
and in traps.
Distribution: Tropical western Pacific, includ-
ing Philippines, Thailand, Indonesia, Papua
New Guinea, Palau, New Britain, Solomon
Islands, and north coast of Australia from Dam-
pier Islands off Western Australia to Capricorn
Islands of the southern Great Barrier Reef.
2478 Bony Fishes
Cephalopholis formosa (Shaw and Nodder, 1812)

Frequent synonyms / misidentifications: None / “Cephalopholis boelang” (Valenciennes, 1828);
C. boenak (non Bloch, 1790).
FAO names: En - Bluelined hind; Fr - Vielle lignes bleues; Sp - Cherna rayada.
Diagnostic characters: Body depth 2.5 to 2.9 times in standard length (for specimens 10 to 26 cm
standard length); head length 2.4 to 2.6 times in standard length. Preorbital depth 3.6 to 4.3% of standard
length, more than 1/2 maxilla width. Interorbital area convex; preopercle rounded, finely serrate, the
lower edge fleshy; subopercle and interopercle smooth; maxilla naked, reaching to or just past vertical at
rear edge of eye. First gill arch with 8 to 10 gill rakers on upper limb, 14 to 18 on lower limb. Dorsal fin with
IX spines and 15 to 17 soft rays, the membranes indented between the spines; anal fin with III spines
and 8 (rarely 7) soft rays, the second spine 12 to 14% of standard length, not reaching past tip of
third spine; caudal fin well rounded; pectoral-fin rays 16 to 18, pectoral fins usually longer than pelvic fins,
1.5 to 1.8 times in head length; pelvic fins reaching about to anus, 1.6 to 2.0 times in head length. Body
scales rough, without auxiliary scales; lateral-line scales 47 to 51; lateral scale series 91 to 109. Colour:
dark brown to yellowish brown, with slightly irregular dark blue lines on head, body and fins (those
on body and fins sometimes absent); snout, lips, and ventral part of head and chest with small dark
blue spots (blue markings become black in preservative, and may be lost on some specimens,
especially older museum material).
Habitat, biology, and fisheries: Like the chocolate hind (Cephalopholis boenak), the preferred habitat of
the bluelined hind is shallow dead or silty reefs; and this may account for the primarily continental
distributions of these 2 species. Too small to be of much commercial importance as a food fish, but often
seen in markets. Caught with hook-and-line, gill net, traps, and in trawls.
Distribution: Primarily conti-
nental and seems to avoid at-
olls or oceanic islands. Oc-
curs from western India to
Thailand, Indonesia, Philip-
pines, China, Taiwan Prov-
ince of China, Japan
(Honshu), and northern Aus-
tralia.
Cephalopholis igarashiensis Katayama, 1957 (Plate I, 5)

FAO names: En - Garish hind; Fr - Vielle voyant; Sp - Cherna chillona.
Diagnostic characters: Body depth greater than head

length, 2.0 to 2.4 times in standard length (for speci-
mens 13 to 29 cm standard length); body width 2.3 to
2.6 times in body depth; caudal peduncle depth equal
to or greater than its length; head length 2.3 to 2.5 times
in standard length. Dorsal head profile straight or
slightly concave to above eye, the nape markedly con-
vex; lateral line noticeably arched over pectoral fin.
Interorbital area flat; preopercle rounded, finely serrate,
the ventral serrae slightly enlarged, the lower edge irregu-
lar but hidden by skin; subopercle and interopercle finely juvenile
serrate; upper edge of operculum distinctly convex; maxilla reaching vertical at rear edge of eye. First gill
arch with 8 or 9 gill rakers on upper limb, 16 or 17 on lower limb. Dorsal fin with IX spines and 14 soft rays,
the membranes distinctly indented between the spines; anal fin with III spines and 9 soft rays; caudal fin
rounded; pectoral-fin rays 18 or 19, the fin length 1.6 to 1.8 times in head length; pelvic fins reach to or
beyond anus, 1.7 to 2.0 times in head length. Body scales rough, without auxiliaries; lateral-line scales 60
to 65; lateral scale series 101 to 117. Colour: head, body, and fins reddish orange; 7 lemon yellow
bars on dorsal part of body, extending onto dorsal fin; 3 broad yellow bands radiating from eye;
fins red, except for extension of yellow bars from body into dorsal fin and membranes at tips of
dorsal-fin spines, which are orange. Juveniles more yellowish, with a large black spot in dorsal fin,
pelvic fins and tips of interspinous dorsal-fin membranes blackish; tips of pelvic fins blackish in large
juveniles or small adults.
Habitat, biology, and fisheries: A rare, deep-water grouper; a specimen from Tahiti was taken in a depth
of 250 m. Like most groupers,
it is said to feed on fishes and
crustaceans. This species
seems too rare to be of com-
mercial importance. Caught
with hook-and-line and in
traps; marketed fresh.
Distribution: Tropical western
Pacific: southern Japan, Tai-
wan Province of China, Guam,
Philippines, South China Sea,
Samoa, and Tahiti.
2480 Bony Fishes
Cephalopholis miniata (Forsskål, 1775) (Plate I, 6)

Frequent synonyms / misidentifications: None / Cephalopholis sexmaculata (non Rüppell, 1830).
FAO names: En - Coral hind; Fr - Vielle de corail; Sp - Cherna estrellada.
standard length); head length 2.4 to 2.6 times in standard length. Interorbital area flat to slightly convex;
preopercle rounded, finely serrate, the lower edge fleshy; upper edge of operculum very convex; maxilla
scaly, reaching to or beyond vertical at rear edge of eye. First gill arch with 7 to 9 gill rakers on upper limb,
14 to 16 on lower limb. Dorsal fin with IX spines and 14 to 16 soft rays, the membranes distinctly
indented between the spines; anal fin with III spines and 9 (rarely 8) soft rays, the fin margin rounded
in adults; caudal fin well rounded; pectoral-fin rays 17 or 18, the fin length 1.4 to 1.75 times in head length;
pelvic fins usually not reaching anus, 1.9 to 2.3 times in head length. Midlateral body scales ctenoid; no
auxiliary scales; lateral-line scales 47 to 56; lateral scale series 94 to 114. Colour: orange-red to reddish
brown, covered with small (usually dark-edged) pale blue-grey spots; pectoral fins orange-yellow
distally; soft dorsal, caudal, and anal fins often with a narrow blue margin and blackish submarginal line;
pelvic fins orange-red, the distal edge dark bluish grey. Juveniles more yellowish, with faint pale blue spots,
fewer than in adults.
Habitat, biology, and fisheries: Well-developed exposed coral reefs in clear water at depths of 2 to 150 m.
Feeds mainly during early morning (07.00 to 09.00 h) and midafternoon (14.00 to 16.00 h). Most prey are
fishes that school just above the reef (predominantly Pseudanthias squamipinnis), and most of these are
caught by a quick rush (“ambush attack”) from the bottom. Cephalopholis miniata forms haremic groups
comprising a dominant male and 2 to 12 females. These groups occupy territories of up to 475 m2,
subdivided into secondary territories, which are defended by a single female. Females are mature at 25 cm
standard length. Although moderately small, C. miniata is a common species of considerable economic
importance to local fisheries. Caught with hook-and-line, spear, and in traps.
Pacific and Red Sea (but not
the Persian Gulf or French
Polynesia), from the African
coast (south to Durban, South
Africa) to the Line Islands in
the Central Pacific, including
most islands in Indian Ocean
and western Central Pacific,
southern Japan, Taiwan Prov-
ince of China, Philippines, In-
donesia, northern Australia,
Papua New Guinea, New
Caledonia, and the islands of
Micronesia.
Cephalopholis sexmaculata (Rüppell, 1830) (Plate I, 7)

Frequent synonyms / misidentifications: Cephalopholis coatesi Whitley, 1937; C. gibbus Fourmanoir,
1954 / Cephalopholis miniata (non Forsskål, 1775).
FAO names: En - Sixblotch hind; Fr - Vielle six taches; Sp - Cherna de seis manches.
dorsal head profile of large specimens (more than 30 cm standard length) distinctly concave above the
eyes; preopercle rounded, finely serrate in young, virtually smooth in large adults, the lower edge fleshy;
subopercle and interopercle with a few small serrae mostly hidden by skin; maxilla scaly, reaching past
eye. First gill arch with 7 to 9 gill rakers on upper limb, 14 to 16 on lower limb. Dorsal fin with IX spines
(the fin membranes distinctly indented between them) and 14 to 16 soft rays; anal fin rounded, with III
spines and 9 soft rays; caudal fin well rounded; pectoral-fin rays 16 to 18, pectoral fins clearly longer than
pelvic fins, 1.4 to 1.6 times in head length; pelvic fins 1.9 to 2.2 times in head length. Lateral body scales
rough, without auxiliary scales (just a few in very large specimens); lateral-line scales 49 to 54; lateral scale
series 95 to 108. Colour: orange-red, with small blue spots sparsely scattered on body but more
densely on head and median fins; head also with elongated blue spots and lines; body with 4 dark
bars (sometimes very faint) merging dorsally with blackish blotches at base of dorsal fin and
extending onto the fin; 2 similar but smaller dark bar/blotches on caudal peduncle; the spaces
between the dark body bars sometimes very pale; pectoral fins orange-red.
Habitat, biology, and fisheries: Coral reefs in depths of 10 to 150 m; a secretive species, generally seen
hiding in caves and crevices on the outer reef slope. The sixblotch hind is more active nocturnally in shallow
water and diurnally in deeper water. Feeds mainly on fishes. Probably of commercial interest to certain
local fisheries. Caught with hook-and-line, traps, and spear.
Indo-West Pacific region from
South Africa to French Poly-
nesia, including Christmas Is-
land, Indonesia, South China
Sea, Philippines, southern Ja-
pan, Palau, Queensland, New
Guinea, Solomon Islands,
Caroline Islands, Mariana Is-
lands, Marshall Islands, Line
Islands, Fiji Islands, Society
Islands, and the Marquesas.
2482 Bony Fishes
Cephalopholis sonnerati (Valenciennes, 1828) (Plate I, 8)

Frequent synonyms / misidentifications: Cephalopholis purpureus Fourmanoir, 1966 / “Cephalopholis
aurantius” (non Valenciennes, 1828), C. formosanus (Tanaka, 1911).
FAO names: En - Tomato hind; Fr - Vielle ananas; Sp - Cherna piña.
40 cm total length
Diagnostic characters: Body depth greater than or subequal to head length, 2.3 to 2.7 times in
standard length (for specimens 9 to 41 cm standard length); head length 2.5 to 2.7 times in standard
length. Dorsal head profile of adults straight to concave, the nape distinctly convex; interorbital area
flat to slightly convex; preopercle rounded, finely serrate, with a shallow notch just above the “corner”, the
lower edge serrate or irregular with broad spinules, usually exposed; subopercle and interopercle finely
serrate; maxilla reaches past eye. First gill arch with 7 to 9 gill rakers on upper limb, 14 to 16 on lower limb.
Dorsal fin with IX spines and 14 to 16 soft rays, the membranes distinctly indented between the spines;
anal fin with III spines and 9 soft rays; caudal fin rounded; pectoral-fin rays 18 to 20; pectoral fins subequal
to pelvic fins, 1.5 to 1.7 times in head length for specimens of 15 to 40 cm length (in specimens
larger than 40 cm, the pelvic fins are longer than the pectoral fins); pelvic fins reaching to beyond
anus. Lateral body scales rough; lateral-line scales 66 to 80; lateral scale series 115 to 134. Colour: adults
from Pacific Ocean generally pale reddish to yellowish brown, covered with small brownish red or
dark brown spots. Juveniles dark reddish brown to nearly black, the rear margin of caudal fin and
sometimes pectoral fins whitish; or body brownish orange with scattered pale greenish spots, and black
spot between upper and middle opercular spines; adults from Indian Ocean with orange-red to reddish
brown body, often with scattered small whitish or purple spots; purple network on head, maxilla, and lips;
pectoral fins orange distally; membranes of soft dorsal, caudal, anal, and pelvic fins dusky; dorsal-fin rays
orange distally; pelvic-fin tips blackish.
Habitat, biology, and fisheries: Coral reefs in depths of 20 to 100 m. Feeds on fishes and crustaceans.
Females mature at about 28 cm standard length and males at about 34 cm. A common and widespread
species of commercial importance throughout most of its range. Caught with hook-and-line, spear, and in
traps.
Distribution: I n d o - P a c i f i c
from east coast of Africa to
the Line Islands in the Central
Pacific; in the western Pacific,
Cephalopholis sonnerati
ranges from southern Japan
to southern Queensland.
Known from both continental
and insular localities, includ-
ing most islands in the tropical
Indian and Pacific Oceans,
but not reported from the Red
Sea or Persian Gulf.
Cephalopholis urodeta (Bloch and Schneider, 1801) (Plate II, 9)

Frequent synonyms / misidentifications: Cephalopholis nigripinnis (Valenciennes, 1828) / None.
FAO names: En - Darkfin hind; Fr - Vielle alle noire; Sp - Cherna alinegra.
Diagnostic characters: Body depth less than head length, 2.7 to 3.3 times in standard length (for
specimens 9 to 21 cm standard length); head length 2.4 to 2.7 times in standard length. Interorbital area
convex; preopercle rounded, finely serrate, the lower edge fleshy; subopercle and interopercle usually
smooth; maxilla reaching well past eye; snout and maxilla with minute cycloid scales. First gill arch with 7
to 9 gill rakers on upper limb, 15 to 17 on lower limb. Dorsal fin with IX spines and 14 to 16 soft rays; anal
fin with III spines and 9 (rarely 8) soft rays; caudal fin rounded; pectoral-fin rays 17 to 19, pectoral fins
distinctly longer than pelvic fins, 1.3 to 1.7 times in head length; pelvic fins usually not reaching anus, 1.8
to 2.3 times in head length. Lateral body scales rough, without auxiliary scales; lateral-line scales 54 to
68; lateral scale series 88 to 108. Colour: reddish brown to brownish red, darker posteriorly; body
sometimes with small pale spots and/or 4 faint, irregular, dark bars that bifurcate ventrally and another 2
dark bars on caudal peduncle; head with numerous, small, close-set orange/red spots (more evident on
paler parts of head) and sometimes with irregular reddish brown blotches; dark spot between upper and
middle opercular spines (more evident on juveniles); a pair of small dark spots on edge of lower lip in line
with the pair of anterior canine teeth; soft dorsal and anal fins with small orange-red spots; outer
triangular part of interspinous dorsal-fin membranes orange; pelvic fins orange-red, usually with a
dark blue edge. Caudal fin with 2 white to bluish white bands that converge posteriorly, the area
between the bands dark, often with pale spots, the outer corners of the fin red, with a whitish margin.
Pectoral fins reddish brown basally, becoming yellow distally.
Habitat, biology, and fisheries: A common coral reef species, usually seen in outer reef areas, but also
found in lagoons, back-reef areas and on the reef-top. Depth range 1 to 60 m. Feeds mainly on fishes
(68%) and crustaceans. Because of its small size, the darkfin hind is not of much interest as a food fish
(except perhaps to subsistence fisheries), but it does well in an aquarium and may find a market as an
aquarium fish. Caught with
hook-and-line, gill nets, and in
traps.
cific region from South Africa
to French Polynesia and the
Pitcairn Islands, probably oc-
curs at all tropical islands and
shallow banks of this region;
also known from southern Ja-
pan to northern coast of Aus-
tralia.
2484 Bony Fishes
Cromileptes altivelis (Valenciennes, 1828) (Plate II, 10)

FAO names: En - Humpback grouper; Fr - Mérou bossu; Sp - Mero jorobado.
Diagnostic characters: Body compressed, its depth less than head length, and 2.6 to 3.0 times in standard
length; body width 2.4 to 2.6 times in body depth; head length 2.5 to 2.8 in standard length; head depressed
anteriorly, the dorsal profile distinctly concave, rising steeply at the nape; preorbital narrow, its depth
3.0 to 5.0 times in eye diameter and 15 to 32 times in head length. Anterior nostril tubular; posterior nostril
a large, crescentic, vertical slit. Jaws with bands of villiform teeth; no canines; palatines with teeth; no
step or knob on ventral edge of maxilla; supramaxilla well developed. Preopercle finely serrate, the serrae
at the angle slightly enlarged, the lower edge smooth; opercle with middle spine inconspicuous, the upper
and lower spines rudimentary. Gill rakers short; first gill arch with 8 to 11 rakers on upper limb, and 13 to
17 on lower limb. Dorsal fin with X spines and 17 to 19 soft rays, the fin origin over opercle, the fin
membranes not incised between the spines, the posterior spines longest and the soft rays even longer;
anal fin with III spines and 9 or 10 soft rays; caudal fin rounded, with 8 branched rays in upper lobe and 7
in lower lobe; pectoral-fin rays 17 or 18. Scales on body smooth (the ctenii greatly reduced); lateral-line
scales 54 to 62; lateral scale series 106 to 122. Supraneural bones slender, the second more than 1/2
length of first; no trisegmental pterygiophores in dorsal or anal fins; rear edge of first dorsal pterygiophore
slightly excavated; epipleural ribs on vertebrae 1 to 8. Cranium elongate, depressed anteriorly and elevated
posteriorly; least interorbital width about 10% of cranium length; postorbital part of cranium elongated, 60%
or more of cranium length; supraoccipital crest not extending onto frontals. Colour: pale greenish brown,
with widely spaced small black spots on head, body, and fins; some spots on body and base of median
fins overlain by a large dusky blotch.
Habitat, biology, and fisheries: Well-developed coral reefs as well as dead or silty reef areas, from tide
pools to depths of 40 m. Growth in captivity is very slow. Juveniles are prized as aquarium fishes, and
adults are one of the most expensive fishes in fish markets. Caught with hook-and-line, spear, and in traps.
from southern Japan to Palau,
New Caledonia, and southern
Queensland (Australia); in the
eastern Indian Ocean from
the Nicobars to Broome,
Western Australia. Reports
f r o m t h e w e s t e r n I nd i an
Ocean are unsubstantiated.
Records from Hawaii are
probably based on released
aquarium fishes.
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Epinephelus akaara (Temminck and Schlegel, 1842) (Plate II, 11)

FAO names: En - Redspotted grouper; Fr - Mérou rouge tacheté; Sp - Mero de pintas rojas.
specimens 11 to 38 cm standard length); head length 2.3 to 2.6 times in standard length. Preopercle with
enlarged serrae at angle; upper edge of operculum straight; nostrils subequal; maxilla reaching about
to vertical at rear edge of eye; midside of lower jaw with 2 rows of teeth. First gill arch with 23 to 25 gill
rakers, of which 8 or 9 on upper limb and 15 to 17 on lower limb. Dorsal fin with XI spines and 15 to 17
soft rays, the third to sixth spines longest, 2.4 to 3.8 times in head length, the interspinous membranes
incised; anal fin with III spines and 8 soft rays; caudal fin rounded; pectoral-fin rays 17 to 19, the fin length
1.5 to 2.1 times in head length; pelvic fins not reaching anus, 1.9 to 2.3 times in head length. Lateral
body scales rough, with auxiliary scales in adults; lateral-line scales 50 to 54; lateral scale series 92 to
106. Colour: head and body pale brownish grey, covered (except ventrally) with small red, orange,
or gold spots; 6 faint oblique dark bars usually visible on body (at least dorsally), the first bar on nape,
the third bar confluent with a dark brown or black blotch on body at base of last 3 dorsal-fin spines, and
the last bar on caudal peduncle; dark body bars extend only onto base of dorsal fin; dorsal-fin margin
yellow or orange; a row of dusky yellow or orange spots (1 per membrane) along middle of spinous
dorsal fin and another row along base of fin; soft dorsal, caudal, and anal fins with faint red or orange
spots basally, the distal parts of these fins dusky with small faint white spots.
Habitat, biology, and fisheries: In Japanese waters, this species is common in rocky areas. Spawning
of pairs in shallow culture ponds has been reported, but the high mortality of larvae has hampered its use
in aquaculture. The redspotted grouper is of considerable commercial importance in Hong Kong and Japan
where it brings a high price in markets, and live specimens are sold for an even better price.
Distribution: Western Pa-
cific: from southern China,
East China Sea, Korea, and ? ?
southern Japan (Kyushu to
about 38°N on both coasts of
Honshu). Reported from Viet
Nam, but the illustration ap-
pears to be the rock grouper
(Epinephelus fasciatomacu-
losus). Records of E. akaara
from the Philippines and India
are unsubstantiated. May oc-
cur in the area.
2486 Bony Fishes
Epinephelus amblycephalus (Bleeker, 1857) (Plate II, 12)

FAO names: En - Banded grouper; Fr - Mérou bande; Sp - Cabrilla venda.
Diagnostic characters: Body depth less than head length,

2.5 to 3.0 times in standard length (for specimens 12 to 35 cm
standard length); head length 2.1 to 2.4 times in standard
length. Preopercle with 3 to 6 enlarged serrae at the
“corner”; upper edge of operculum slightly convex; nostrils
subequal or posterior nostril larger than anterior; maxilla
scaly, reaching about to vertical at rear edge of eye; midside
of lower jaw with 2 rows of teeth. First gill arch with 22 to 24
gill rakers, of which 8 on upper limb and 14 to 16 on lower juvenile
limb. Dorsal fin with XI spines and 15 or 16 soft rays, the third or fourth spine longest, 2.7 to 3.2 times in
head length, the interspinous membranes only slightly incised; anal fin with III spines and 8 soft rays;
caudal fin rounded; pectoral fins with 18 or 19 rays, the fin length equal to pelvic fins, 1.7 to 2.3 times in
head length. Lateral body scales ctenoid, with a few auxiliary scales in adults; lateral-line scales 47 to 52;
lateral scale series 90 to 121. Colour: head, body, and median fins pale grey; body with 4 broad dark
brown bars, the first 2 extending into spinous part of dorsal fin, the third and fourth bars extending
into soft dorsal and anal fins; a fifth dark bar at base of caudal fin; small black spots on dorsal parts
of body bars (mainly along the edges of the bars); dark brown saddle blotch on nape, with small
black spots mainly along edge of this blotch; cheeks, snout, interorbital area, jaws, and chest mostly
dark brown, with 2 or 3 white bands radiating from eye; irregular black bar on middle of caudal fin; maxillary
groove yellow; ventral parts of head and body often with a pinkish tinge. Small juveniles (6 cm standard
length) white, with black bars on body as described for adults, the fins pale yellow with small black
spots.
Habitat, biology, and fisheries: Usually found on offshore coral and rocky reefs in depths of 80 to 130 m.
The banded grouper is a popular and commercially important species in Hong Kong, but in Singapore it is
not a popular grouper in the market. Caught with hand lines. Sold fresh.
Distribution: Eastern Indian
Ocean and central western
Pacific from the Andaman
Sea to southern Japan, Tai-
wan Province of China,
China, Philippines, Viet Nam,
Malaysia, Thailand, Indone-
sia, New Guinea, and north-
western Australia; but
unknown east of New Guinea
except for one record from
Fiji.
Epinephelus areolatus (Forsskål, 1775) (Plate II, 13)

Frequent synonyms / misidentifications: None / Epinephelus chlorostigma (non Valenciennes, 1828).
FAO names: En - Areolate grouper; Fr - Mérou aréolé; Sp - Mero areolado.
specimens 14 to 31 cm standard length); head length 2.4 to 2.8 times in standard length. Interorbital area
convex; preopercle angular, with 2 to 7 enlarged serrae at the angle; upper edge of operculum straight or
slightly convex; nostrils subequal; maxilla extending to below rear half of eye, the lower edge with a distinct
step posteriorly; maxilla, lower jaw, and gular area scaly; midlateral part of lower jaw with 2 rows of teeth.
First gill arch with 23 to 25 gill rakers, of which 8 to 10 on upper limb and 14 to 16 on lower limb.
Pyloric caeca 11 to 17. Dorsal fin with XI spines and 15 to 17 soft rays, the third or fourth spines
longest and about equal to longest soft ray, the interspinous membranes moderately incised; anal fin with
III spines and 8 soft rays, the margin rounded or angular; caudal fin slightly convex in juveniles, truncate
or slightly emarginate in adults; pectoral-fin rays 17 to 19; pectoral fins longer than pelvic fins, 1.5 to
1.8 times in head length; pelvic fins 1.7 to 2.1 times in head length, reaching to or nearly to anus. Lateral
body scales rough; adults with auxiliary scales. Lateral-line scales 49 to 53; lateral scale series 97 to
116. Colour: head, body, and fins pale, covered with numerous close-set brown, brownish yellow
or greenish yellow spots, the largest about size of pupil, those on front of head smaller than those
on operculum. Pectoral fins pale, with small dark spots on the rays. Posterior edge of caudal fin
with a distinct white margin.
Habitat, biology, and fisheries: Usually found in seagrass beds or on fine sediment bottoms near rocky
reefs, dead coral, or alcyonarians. It has been taken in depths of 6 to 200 m. At New Caledonia, the areolate
grouper was reported to feed on penaeid prawns. Common in the markets of Hong Kong, Singapore, and
probably throughout its range. Caught with hook-and-line, traps, and in trawls.
cific from South Africa to Fiji,
including Andaman Islands,
Thailand, Malaysia, Viet Nam,
Indonesia, Philippines, Hong
Kong, China, Taiwan Prov-
ince of China, Japan, northern
Australia (and nearby is-
lands), Papua New Guinea,
and New Caledonia. It ap-
pears to be absent from Mi-
cronesia, Polynesia, and
most islands of the western
Indian Ocean.
2488 Bony Fishes
Epinephelus awoara (Temminck and Schlegel, 1842) (Plate II, 14)

FAO names: En - Yellow grouper; Fr - Mérou jaune; Sp - Mero amarillo.
standard length); head length 2.3 to 2.6 times in standard length. Interorbital area convex; dorsal head
profile strongly convex; preopercle subangular, with 2 to 5 strong spines at the angle; upper edge of
operculum straight, the uppermost spine rudimentary; maxilla reaches about to vertical at rear edge
of eye; midlateral part of lower jaw with 2 rows of small, subequal teeth. First gill arch with 22 to 26 gill
rakers, of which 8 or 9 on upper limb and 16 to 18 on lower limb. Pyloric caeca 12, in 3 bundles. Dorsal
fin with XI spines and 15 or 16 soft rays, the third or fourth spine longest but shorter than the longest soft
ray, the interspinous membranes deeply incised; anal fin with III spines and 8 soft rays; caudal fin convex;
pectoral-fin rays 17 to 19, the fin longer than pelvic fins, 1.6 to 1.9 times in head length. Lateral body scales
distinctly rough, with auxiliary scales in specimens larger than 30 cm standard length; lateral-line scales
49 to 55; lateral scale series 92 to 109. Colour: head and body pale greyish brown dorsally, usually golden
yellow ventrally; 4 broad dark bars on dorsal part of body, 1 on caudal peduncle and another often
visible on nape; head and body with numerous small yellow spots; body and median fins also with
small greyish white spots; soft dorsal, caudal, and sometimes the anal fin with prominent yellow
margin; paired fins dusky yellow. Dark bars on body may be faint or absent in large adults.
Habitat, biology, and fisheries: Occurs in rocky areas as well as on sandy-mud bottoms; juveniles are
common in tidepools, and adults are caught in depths of 10 to 50 m. In captivity, the yellow grouper is an
aggressive fish, spending much of its time chasing and biting other fishes, especially members of its own
species. Valuable food fish,
caught with trawls or hook-
and-line. Eggs have been ar-
tificially fertilized, with the
longest survival time for the
larvae of 15 days.
Distribution: Western North
Pacific: Korea, Japan, Taiwan
Province of China, China, Viet
Nam, and islands in the South
China Sea.
Epinephelus bleekeri (Vaillant, 1877) (Plate II, 15)

FAO names: En - Duskytail grouper; Fr - Mérou demideuil; Sp - Mero medioluto.
Diagnostic characters: Body elongate, its depth 3.0 to 3.5 times in standard length (for specimens
11 to 52 cm standard length); head length 2.4 to 2.7 times in standard length. Interorbital area flat to slightly
convex; preopercle angle with 2 to 9 enlarged serrae; adults with a notch above preopercle angle;
upper edge of operculum straight; maxilla scaly, reaching to or beyond vertical at rear edge of eye;
midlateral part of lower jaw with 2 rows of subequal teeth. First gill arch with 25 to 28 gill rakers, of which
9 to 11 on upper limb and 16 to 18 on lower limb. Dorsal fin with XI spines and 16 to 18 soft rays, third to
fifth spines longest, the interspinous membranes incised; anal fin with III spines and 8 or 9 (rarely 9) soft
rays; caudal fin truncate or slightly convex; pectoral-fin rays 17 to 19, the fin length 1.6 to 2.1 times in
head length; pelvic fins 1.9 to 2.5 times in head length. Lateral body scales rough; adults with a few small
auxiliary scales; lateral-line scales 49 to 53; lateral scale series 99 to 104. Colour: head and body
brownish, reddish brown or purplish grey, covered (except ventrally) with numerous reddish
orange, gold, or yellow spots; dorsal fin and upper third of caudal fin with spots like those on body;
lower two-thirds of caudal fin dusky; spots on body of some fish with a faint dark margin; pectoral and
pelvic fins and distal part of anal fin dusky; dark streak along maxillary groove. Juveniles (less than 11 cm
standard length) with 7 faint dark bars dorsally on body, the first 2 on nape, the last on caudal peduncle;
all bars more or less demarcated by small dark spots; no dark spots on head or fins.
Habitat, biology, and fisheries: Occurs on shallow rocky banks, but not reported from well-developed
coral reefs. Depth range reported as 30 to 104 m. An excellent eating fish, but apparently not abundant.
Caught with hook-and-line,
longlines, and trawls.
Pacific species occurring from
the Persian Gulf to Thailand,
Cambodia, Indonesia, Bor-
neo, Philippines, China (in-
cluding Hong Kong and
Taiwan Province), and north
coast of Australia. It is not
known from Japan, New
Guinea, or any of the islands
of Micronesia or Polynesia.
2490 Bony Fishes
Epinephelus bruneus Bloch, 1793

Frequent synonyms / misidentifications: Serranus moara Temminck and Schlegel, 1842 / None.
FAO names: En - Longtooth grouper; Fr - Mérou longues dents; Sp - Mero diente largo.
Diagnostic characters: Body elongate, its depth 3.0

to 3.6 times in standard length; head length 2.3 to 2.5
times in standard length. Interorbital area convex; pre-
opercle angular, with the serrae at angle distinctly
enlarged; upper spine on opercle inconspicuous; upper
edge of operculum convex; maxilla reaches past vertical
at rear edge of eye; tiny embedded scales on maxilla; no
step on ventral edge of maxilla; midlateral part of lower
jaw with 2 rows of well-developed canines, those juvenile 40 cm standard length
along sides of upper jaw slightly smaller; nostrils
small, subequal. Gill rakers shorter than gill filaments; first gill arch with 24 to 27 rakers, of which 9 to 11
on upper limb and 16 to 18 on lower limb. Dorsal fin with XI spines and 13 to 15 soft rays, the third or
fourth spine longest, but shorter than the longest soft ray, the interspinous membranes deeply incised;
anal fin with III spines and 8 soft rays; caudal fin rounded; pectoral-fin rays 17 to 19, the fin length 1.9 to
2.5 times in head length; pelvic fins 2.2 to 2.6 times in head length and ending well short of anus. Midlateral
body scales rough; no auxiliary scales on body; lateral-line scales 64 to 72; some anterior lateral-line
scales on adults with branched tubes; lateral scale series 93 to 126. Colour: juveniles pale yellowish
brown, the body with 6 irregular, oblique dark bars containing irregular pale spots; first bar extends
from nape to eye, the last on caudal peduncle; 3 dark brown bands radiating from lower part of eye;
some specimens with distal part of interspinous membranes greenish yellow. Adults (larger than 40 cm
standard length) dark greyish brown, the bars and bands replaced by faint dorsal blotches or absent
altogether, the body covered with small pale grey spots forming short horizontal lines and a mottled pattern;
lower margin of anal fin and lower corner of caudal fin with white edge.
Habitat, biology, and fisheries: Rocky reefs and mud bottom; adults found in depths of 20 to 200 m;
juveniles occur in shallow water. An excellent eating fish. In 1968, Epinephelus bruneus was one of the
most abundant species in
Hong Kong waters. Caught
with handlines, longlines, and
trawls.
Distribution: K n o w n o n l y
from Korea, Japan (north to
Hegura-jima Island, 37°50’N),
China (south to Hong Kong
and Hainan Island), Taiwan
Province of China, and Luzon,
Philippines. May be more
widespread in the area.
Epinephelus caeruleopunctatus (Bloch, 1790) (Plate II, 16)

Frequent synonyms / misidentifications: None / Epinephelus ongus (non Bloch, 1790); E. summana
(non Forsskål, 1775); E. corallicola (non Valenciennes, 1828).
FAO names: En - Whitespotted grouper; Fr - Vielle taches blanches; Sp - Mero nevero.
17 cm standard length 3 cm standard length

juveniles
Diagnostic characters: Body depth distinctly less than head length, 2.9 to 3.4 times in standard
length (for specimens 11 to 47 cm standard length); head length 2.3 to 2.5 times in standard length. Head
pointed and the dorsal profile almost straight; preopercle rounded, finely serrate; opercular spines
inconspicuous; upper edge of operculum straight, sinuous or slightly convex; maxilla naked, mostly
covered by upper lip; canines at front of jaws small or absent; midlateral part of lower jaw with 3 to 5 rows
of small teeth. In specimens more than 45 cm, the rear nostrils become vertically elongate, its length
5 or 6 times greater than diameter of anterior nostrils. First gill arch with 8 to 10 gill rakers on upper
limb, 13 to 17 on lower limb (juveniles); specimens larger than 25 cm with only 4 to 8 developed rakers on
lower limb, the rudiments difficult to count because of intercalated bony tooth plates; gill rakers shorter
than gill filaments, the raker at angle of gill arch about twice as long as adjacent rakers. Dorsal fin with XI
spines and 15 to 17 soft rays, the third or fourth spines longest, 2.7 to 3.6 times in head length, the
interspinous membranes incised; anal fin with III spines and 8 soft rays; caudal fin rounded; pectoral fins
large and fleshy, with 17 to 19 rays, the fin length 1.5 to 2.1 times in head length; pelvic fins end well
short of anus, their length 2.0 to 2.7 times in head length. Lateral body scales rough (1 specimen, 50 cm,
with “mostly smooth” lateral body scales), with auxiliary scales; lateral-line scales 51 to 61; lateral scale
series 86 to 109. Colour: adults brownish grey, the body covered with small pale spots overlain with
large pale blotches; oblique black saddle on rear half of peduncle; 4 or 5 indistinct black blotches
at base of dorsal fin; prominent black streak on maxillary groove. Large adults brownish, covered
with small, indistinct, contiguous pale spots. Juveniles (less than 25 cm) dark grey to black, covered
with prominent pupil-size white spots and smaller white dots.
Size: Largest specimen examined 59 cm total length; reported to 76 cm.
Habitat, biology, and fisheries: Coral reefs, usually in or near caves; juveniles found in tidepools. Caught
with hook-and-line, spear, and in traps. Prob-
ably of some importance to fisheries in areas
where it is common.
Distribution: Indo-West Pacific, from South
Africa to Fiji, including Thailand, Indonesia,
Philippines, Taiwan Province of China, Ja-
pan, Papua New Guinea, Australia (Queens-
land to New South Wales), Palau, Solomon
Islands, New Caledonia, Vanuatu Caroline
Islands, Marshall Islands, and Gilbert Is-
lands.
2492 Bony Fishes
Epinephelus chlorostigma (Valenciennes, 1828) (Plate III, 17)

Frequent synonyms / misidentifications: None / Epinephelus areolatus (non Forsskål, 1775).
FAO names: En - Brownspotted grouper; Fr - Mérou pintade; Sp - Mero pintado.
standard length); body width 1.8 to 2.2 times in body depth; head length 2.4 to 2.7 times in standard length;
caudal peduncle depth 3.0 to 3.6 times in head length. Interorbital slightly convex; preopercle angular,
with 4 to 7 enlarged serrae at angle; upper edge of operculum straight; maxilla reaches about to
vertical at rear edge of eye; maxilla scaly, with a low step on posterior part of ventral edge; midlateral part
of lower jaw with 2 to 4 rows of teeth, the inner ones about twice size of outer teeth; rear nostrils not
noticeably larger than anterior nostrils. First gill arch with 23 to 29 gill rakers, of which 8 to 11 on upper
limb and 15 to 18 on lower limb; gill rakers longer than gill filaments. Pyloric caeca 26 to 52. Dorsal fin
with XI spines and 16 to 18 soft rays, the third or fourth spines longest, 2.4 to 3.2 times in head length,
the interspinous membranes slightly to moderately incised; anal fin rounded or angular, with III spines
and 8 soft rays, the third spine longer than second; caudal fin truncate or slightly emarginate; pectoral-fin
rays 17 to 19; pectoral fins usually slightly longer than pelvic fins, 1.6 to 2.0 times in head length; pelvic
fins 1.8 to 2.3 times in head length. Lateral body scales rough, with auxiliary scales; lateral-line scales
48 to 53; lateral scale series 96 to 122. Colour: head, body, and fins with small, irregular, close-set
dark brown spots, the ground colour forming a pale network; caudal fin usually with white line along
rear margin; spots on pectoral fins mainly confined to rays.
Size: Maximum total length at least 75 cm; maximum weight 7 kg.
Habitat, biology, and fisheries: Coral reefs and also (in the South China Sea) on mud bottoms; depth
range 4 to 280 m. Feeds on fishes and crustaceans (mainly stomatopods and crabs). Females are mature
at 23 to 29 cm total length, and sexual transition occurs between 35 and 45 cm, but all females do not
change sex. Common in markets of Singapore and Hong Kong. Caught with hook-and-line, spear, traps,
and trawls.
Distribution: Red Sea to South Africa and eastwards to southern Japan, Papua New Guinea, New
Caledonia, New Ireland, Caroline Islands, American Samoa, and Fiji. The wide distribution of Epinephelus
chlorostigma has some puz-
zling gaps. There are no veri-
fiable records from the Co-
moros, the continental shelf
between Oman and Cambo-
dia, Indonesia, Philippines,
and Australia. Records from
the Persian Gulf are appar-
ently misidentifications of E.
polylepis.
Epinephelus coioides (Hamilton, 1822) (Plate III, 18)

Frequent synonyms / misidentifications: Epinephelus suillus (Valenciennes, 1828) / Epinephelus
malabaricus (non Bloch and Schneider, 1801); E. tauvina (non Forsskål, 1775).
FAO names: En - Orangespotted grouper; Fr - Mérou taches oranges; Sp - Mero de pintas naranjas.
Diagnostic characters: Body elongate, its depth 2.9 to 3.7 times in standard length (for specimens
10 to 78 cm standard length); body width 1.4 to 2.0 times in body depth; head length 2.3 to 2.6 times in
standard length. Interorbital width 5.0 to 6.2 times in head length; preopercle with enlarged serrae at
angle and a broad shallow notch just above angle; upper edge of operculum straight or somewhat convex;
maxilla reaches to or slightly past a vertical at rear edge of eye; upper jaw length 17 to 20% of standard
length; midlateral part of lower jaw with 2 or 3 rows of subequal teeth; nostrils subequal. First gill arch with
23 to 26 gill rakers, of which 8 to 10 on upper limb and 14 to 17 on lower limb; adults with small bony
platelets on lateral side of first gill arch. Pyloric caeca 50 to 60. Dorsal fin with XI spines and 14 to 16 soft
rays, the third or fourth spines longest, 2.9 to 4.0 times in head length; anal fin with III spines and 8 soft
rays, the third spine usually longer than the second; caudal fin rounded; pectoral-fin rays 18 to 20. Lateral
body scales rough, with minute auxiliary scales; lateral-line scales 58 to 65; lateral-line tubes of
anterior scales branched in adults; lateral scale series 100 to 118. Colour: head and body tan dorsally,
shading to whitish ventrally; numerous small brownish orange or reddish brown spots on head, body,
and median fins; body with 5 faint, irregular, oblique, dark bars which bifurcate ventrally; first dark
bar below anterior dorsal-fin spines, last bar on caudal peduncle; 2 dark spots on interopercle and another
1 or 2 at junction of sub- and interopercles. Orange spots turn brown on exposure to air and become fainter
(more diffuse) in preservative.
Size: Maximum length at least 95 cm.
Habitat, biology, and fisheries: Estuaries and offshore to depths of 100 m. Feeds on fishes, shrimps,
crabs, and cephalopods. Females mature at 25 to 30 cm (2 to 3 years old), and sexual transition occurs
at 55 to 75 cm. The subject of much recent research in aquaculture, the orangespotted grouper is often
misidentified as “Epinephelus tauvina” or “E. malabaricus” in the literature. Common and expensive in
markets of the region; sold fresh and kept alive at restaurants in Hong Kong and Taiwan Province of China.
traps, trawls, and lift nets.
Distribution: C o n t i n e n t a l
shores and large islands from
Red Sea to South Africa and
east to the western Pacific,
where it ranges from the
Ryukyu Islands of Japan to
Australia and out to the is-
lands of Palau and Fiji.
2494 Bony Fishes
Epinephelus cyanopodus (Richardson, 1846) (Plate III, 19)

Frequent synonyms / misidentifications: Epinephelus hoedtii (Bleeker, 1855); E. kohleri Schultz, 1953 /
None.
FAO names: En - Speckled grouper; Fr - Mérou bleu; Sp - Mero azul.
Diagnostic characters: Body deep and compressed, its depth subequal to head length, 2.4 to 2.7
times in standard length (for specimens 11 to 43 cm standard length); body width 1.9 to 2.8 times
in body depth. Dorsal head profile steep, the interorbital area distinctly convex; preopercle subangular,
finely serrate, serrae at rounded corner slightly enlarged; opercular spines inconspicuous; upper edge of
operculum straight; maxilla reaches about to vertical at rear edge of eye; midlateral part of lower jaw with
2 to 4 rows of teeth; rear nostrils of adults 2 or 3 times larger than anterior nostrils. First gill arch with 24
to 27 gill rakers, of which 9 or 10 on upper limb and 15 to 17 on lower limb. Pyloric caeca very numerous.
Dorsal fin with XI spines and 16 or 17 soft rays, the third or fourth spines longest, 2.2 to 2.8 times
in head length, the interspinous membranes not or only slightly incised; anal fin with III spines and
8 soft rays; caudal fin truncate or slightly emarginate; pectoral fins thin (not fleshy), with 18 to 20 rays,
subequal to pelvic fins, their length 1.7 to 2.0 times in head length. Lateral body scales rough, with auxiliary
scales; lateral-line scales 63 to 75; lateral scale series 128 to 147. Colour: adults usually pale bluish
grey, covered with black dots and a few scattered, irregular black spots; broad black submarginal
band on caudal fin of some large juveniles; pelvic fins usually black tipped. Juveniles with yellow
fins; body of small juveniles (to 12 cm standard length) mainly yellow, the head and front part of
body with a wash of bluish grey and faint dark dots.
Habitat, biology, and fisheries: Usually found on isolated coral heads in lagoons or bays, but also caught
at depths to 150 m on outer reef area. Swims out in the open, several metres above the bottom, and is
readily caught by anglers at night. Feeds on fishes and calappid crabs. In the 1960’s, the speckled grouper
was abundant at Macclesfield Bank and Pratas Reef in the South China Sea. It is caught with trawls,
handlines, and longlines. Marketed fresh.
Distribution: Western Pacific, from southern Japan to southern Queensland and east to Fiji and islands
of Micronesia; also known from Taiwan Province of China, Hong Kong, Viet Nam, Gulf of Thailand,
Indonesia, Philippines, Papua
New Guinea, New Caledonia,
and Lord Howe Island. The
only record from the Indian
Ocean (a specimen in the Vi-
enna Museum, NMW 40520,
29.3 cm standard length, from
Western Australia) is dubious;
the species is not known from
Western Australia, and there
are no specimens from there
in the Western Australian Mu-
seum.
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Epinephelus daemelii (Günther, 1876)

FAO names: En - Saddletail grouper; Fr - Mérou troussequin; Sp - Mero montado.
dorsal head profile almost straight; preopercle rounded, finely serrate; opercular spines inconspicu-
ous; upper edge of operculum distinctly convex. Maxilla reaches well past eye; canines at front of
jaws well developed, particularly the inner depressible teeth at symphysis of upper jaw; midlateral
part of lower jaw with 2 rows of large teeth. Rear nostrils of adults 2 to 4 times larger than anterior
nostrils. First gill arch with 25 to 28 gill rakers, of which 9 to 12 on upper limb and 15 to 19 lower limb;
longest gill raker shorter than longest gill filaments. Dorsal fin with XI spines and 14 soft rays, third to
last spines subequal and shorter than longest dorsal-fin rays, the interspinous membranes incised; anal
fin with III spines and 8 soft rays; caudal fin rounded; pectoral fins fleshy, with 17 to 19 rays, the fin length
1.6 to 2.1 times in head length; pelvic fins not reaching anus, their length 2.1 to 2.5 times in head length.
Lateral body scales smooth (except for area covered by pectoral fins), with auxiliary scales; lateral-line
scales 63 to 71; lateral scale series 111 to 126. Colour: variable, can change quickly from uniformly
dark brown or black to a pale greyish or brown with 4 irregular, oblique, dark bands; first band from
anterior dorsal-fin spines to upper edge of operculum; bands darkest dorsally, where they contain
small, irregular pale spots; ventrally, the bands break up into irregular dark markings; a fifth dark
band from nape to eye and a black saddle blotch on peduncle; median and pelvic fins with narrow
white margins on some fish. Large adults often uniformly dark brown or black (especially in estuaries),
the underside of head pale and a few pale spots on cheeks. Small juveniles with dark spots on edges of
dark body bars and extend onto median fins.
Size: Maximum length at least 122 cm; maximum weight 64 kg.
Habitat, biology, and fisheries: Rocky reefs from near shore to depths of at least 50 m. An aggressive
territorial species that may occupy a particular cave for life. Sex change (from female to male) occurs at a
length of 100 to 110 cm. Juveniles feed on crabs and fishes. An esteemed food fish, avidly sought by
anglers and spearfishermen.
Distribution: Temperate and
subtropical waters of south-
western Pacific: Australia
(from southern Queensland to
South Australia), Lord Howe
Island, Norfolk Island, Ker-
madec Islands, and New Zea-
land (North Island and Poor
Knights Islands).
2496 Bony Fishes
Epinephelus epistictus (Temminck and Schlegel, 1842) (Plate III, 20)

Frequent synonyms / misidentifications: Epinephelus praeopercularis Boulenger, 1887 / Epinephelus
magniscuttis non Postel, Fourmanoir, and Guézé, 1963; E. heniochus non Fowler, 1904.
FAO names: En - Dotted grouper; Fr - Mérou pâle; Sp - Mero palido.
standard length); head length 2.2 to 2.5 times in standard length. Interorbital area and dorsal head profile
slightly convex; preopercle angle produced, with 3 to 5 distinctly enlarged serrae; upper edge of
operculum straight or slightly convex; maxilla reaches to, almost to, or slightly past vertical at rear edge
of eye; adults with a step or abrupt bend on ventral edge of maxilla; midlateral part of lower jaw with 2 rows
of teeth, the inner ones slightly larger; nostrils of adults subequal, or rear nostrils about twice as large as
anteriors. First gill arch with 7 to 10 gill rakers on upper limb, 15 to 19 on lower limb; length of longest gill
raker subequal to longest gill filaments. Pyloric caeca 7 to 10. Dorsal fin with XI spines and 14 or 15
soft rays, the third or fourth spines longest, usually shorter than longest soft ray, the interspinous
membranes deeply incised; anal fin with III spines and 8 soft rays; caudal fin slightly to moderately rounded;
pectoral-fin rays 17 to 19, the fin length 1.6 to 2.1 times in head length; pelvic fins end well short of anus,
their length 2.0 to 2.6 times in head length. Lateral body scales rough, with a few auxiliary scales in adults;
lateral-line scales 57 to 70; lateral scale series 105 to 127. Colour: head and body pale brownish or
greenish grey, with conspicuous, small, brownish black spots on dorsolateral part of body, rear
part of head, and on median fins; some specimens with faint dark band from eye to end of operculum,
another from eye to notch of preopercle, and third a continuation of maxillary streak; pectoral-fin rays
brownish, the membranes clear. Juveniles with dark spots on head and body arranged in 3 longitudinal
rows.
Size: Maximum total length about 80 cm; maximum weight 7 kg.
Habitat, biology, and fisheries: Rocky and trawlable bottoms in depths of 71 to 290 m. Nothing has been
published on the biology of Epinephelus epistictus. Probably of some commercial importance, but separate
catch statistics are not available for this species. Caught with trawls and handlines.
Distribution: Mainly conti-
nental localities in the tropical
Indo-West Pacific region:
from Red Sea to South Africa
and across northern Indian
Ocean to Viet Nam, China (in-
cluding Hong Kong and Fujian
and Taiwan provinces), Ko-
rea, Japan, Ogasawara Is-
lands, Indonesia, Papua New
Guinea, and northern Austra-
lia.
Epinephelus ergastularius Whitley, 1930

Frequent synonyms / misidentifications: None / Epinephelus septemfasciatus (non Thunberg, 1793);
E. octofasciatus non Griffin, 1926.
FAO names: En - Sevenbar grouper; Fr - Mérou sept raies; Sp - Mero de siete bandas.
Diagnostic characters: Body depth 2.6 to 2.9 times in

standard length (for specimens 19 to 89 cm standard
length); head length 2.4 to 2.6 times in standard length.
Interorbital area and dorsal head profile slightly convex;
preopercle finely serrate, the serrae on the rounded
corner not enlarged and the ventral edge with 1 to 4
small serrae; upper edge of operculum almost straight;
maxilla reaches to or almost to vertical at rear edge of eye; juvenile
midlateral part of lower jaw with 2 rows of teeth; rear
nostrils of adults 2 to 4 times larger than anterior nostrils. First gill arch with 7 to 9 gill rakers on upper
limb, 14 or 15 on lower limb; outer side of first gill arch with small bony tooth plates. Dorsal fin with XI
spines and 14 or 15 soft rays, the third spine usually longest, 2.6 to 3.2 times in head length and subequal
to length of longest soft ray, the interspinous membranes deeply incised; anal fin with III spines and 9
or 10 soft rays; caudal fin slightly rounded in juveniles, truncate to slightly emarginate in adults; pectoral-fin
rays 18 or 19; the fin length 1.7 to 2.4 times in head length; pelvic fins distinctly shorter than pectoral fins.
Lateral body scales rough; no auxiliary scales; lateral-line scales 63 to 70; lateral scale series 103 to
116. Colour: body of juveniles brown, with 7 broad dark bars, the first from dorsal-fin origin to upper
edge of operculum, the last bar covering most of caudal peduncle, its upper half black; the pale
spaces separating second and third dark bars and that between fourth and fifth dark bars are
narrower than those between third and fourth or fourth and fifth and sixth dark bars; prominent black
maxillary streak; median and pelvic fins dark brown; pectoral fins hyaline brown, pelvic fins blackish brown.
Adults pinkish grey, with no trace of dark bars; fins darker than body and with white margins (except along
spinous part of dorsal fin).
Size: Maximum total length at least 157 cm;
maximum weight about 66 kg.
Habitat, biology, and fisheries: Adults are
caught in depths of 108 to 370 m; juveniles
are found in depths of 15 to 128 m. Com-
monly caught with hook-and-line off the
coast of New South Wales.
Distribution: Known only from east coast of
Australia between 18° and 36°S.
2498 Bony Fishes
Epinephelus fasciatomaculosus (Peters, 1866) (Plate III, 21)

Frequent synonyms / misidentifications: Epinephelus fasciatomaculatus (Peters, 1866) (misspelling) /
Epinephelus akaara (non Temminck and Schlegel, 1842); E. diacanthus (non Valenciennes, 1828);
E. sexfasciatus (non Valenciennes, 1828).
FAO names: En - Rock grouper; Fr - Mérou rocaille; Sp - Mero de las piedras.
slightly convex; preopercle angular, the serrae at the angle slightly enlarged; upper edge of opercu-
lum straight; maxilla reaches about to below rear third of eye or a little past eye; midlateral part of lower
jaw with 2 or 3 rows of subequal teeth; nostrils subequal. First gill arch with 7 or 8 gill rakers on upper limb,
14 to 16 on lower limb. Dorsal fin with XI spines and 15 to 17 soft rays, the third or fourth spines longest,
2.5 to 3.1 times in head length, and a little shorter than longest dorsal-fin rays, the interspinous membranes
1.5 to 1.8 times in head length; pelvic fins not reaching past anus, their length 1.9 to 2.3 times in head
length. Lateral body scales rough, with auxiliary scales; lateral-line scales 48 to 52; lateral scale series
92 to 106. Colour: head and body pale greyish brown, with numerous small dark brown, brownish
yellow or russet spots; 5 slightly oblique broad dark bars usually visible on body, the last covering
peduncle, the first 4 extending into dorsal fin; the first 2 bars are darker dorsally and extend to the
margin of spinous dorsal fin; ventral parts of head and body whitish or pinkish; soft dorsal, caudal, and
anal fins with faint pale spots and streaks; minute gold tag often visible at tips of some dorsal-fin spines.
Dark body bars faint in adults and may be lost in preservative, except for dorsal part of first 2 bars.
Habitat, biology, and fisheries: Shallow rocky areas. Feeds on fishes, prawns, crabs, worms, and
gastropods. Females mature at 12.5 cm standard length, and most sexual transition occurs in fish 2 or 3
years old (14 to 16 cm). Maximum age is 6 years. Of some commercial importance in Hong Kong and
Taiwan Province of China. Although a small species, it is excellent as a food fish and brings a good price.
Caught with trawls, gill nets, and hook-and-line.
from southern Japan to Ma-
laysia, including Taiwan Prov-
ince of China, Philippines,
Viet Nam, China, Hong Kong,
Hainan, and Sarawak.
Epinephelus fasciatus (Forsskål, 1775) (Plate III, 22)

Frequent synonyms / misidentifications: Epinephelus emoryi Schultz, 1853 / Epinephelus retouti non
Bleeker, 1868.
FAO names: En - Blacktip grouper; Fr - Mérou oriflamme; Sp - Mero banderilla.
standard length); head length 2.3 to 2.6 times in standard length. Preopercle rounded, the rear edge
serrate; upper edge of operculum straight; midlateral part of lower jaw with 2 to 4 rows of teeth; nostril
subequal. First gill arch with 6 to 8 gill rakers on upper limb, 15 to 17 on lower limb. Pyloric caeca 10 to
16. Dorsal fin with XI spines and 15 to 17 soft rays, the third to eleventh spines subequal and slightly
shorter than longest dorsal-fin ray, the interspinous membranes distinctly incised; anal fin with III spines
and 8 soft rays; caudal fin slightly to moderately rounded; Central-Pacific specimens often with
truncate caudal fins; pectoral-fin rays 18 to 20, the fin length 1.5 to 2.0 in head length; pelvic fins not
reaching past anus. Lateral body scales rough, with numerous auxiliary scales; nape and dorsopos-
terior part of head densely covered with minute auxiliary scales; lateral-line scales 49 to 75; lateral
scale series 92 to 135. Colour: ground colour pale greenish grey, to pale reddish yellow to scarlet;
body often with 5 or 6 faint dark bars, the last on peduncle; body scales (except ventrally) with pale
centre and dark rear margin, producing a faint checked pattern; usually with irregular pale blotches
and spots on body (a midlateral series diminishing in size posteriorly from behind pectoral-fin base to
caudal peduncle) and dark brown line along crease of dorsal-fin base; dorsal part of head and nape,
including upper jaw, dark red or reddish brown or with bands and blotches of similar colour; most specimens
with dark band from below eye to interopercle; rim of orbit black and often bordered by a pale bluish line.
Fins reddish orange, pale yellowish green or greenish brown, the outer triangular part of interspinous
membranes of dorsal fin black (dark red in fish from Western Australia and in some specimens from
deep water), with pale yellow or white spot behind tip of each spine.
Habitat, biology, and fisheries: Coral reefs and rocky bottom from shore to depths of 160 m. Feeds during
day and night on brachyuran crabs, fishes, shrimps, galatheid crabs, stomatopods, ophiuroids, and
octopus. Abundant in shallow water and widely used for food. Readily caught with hook-and-line, spear,
traps, and gill nets.
cific region from Red Sea to
Pitcairn Islands; in the west-
ern Pacific from Japan (south
of 33°) and Korea to southern
Queensland and Lord Howe
Island, and including virtually
all tropical islands of the re-
gion.
2500 Bony Fishes
Epinephelus faveatus (Valenciennes, 1828) (Plate III, 23)

Frequent synonyms / misidentifications: None / Epinephelus macrospilos (non Bleeker, 1855);
E. quoyanus (non Valenciennes, 1830); E. merra non Bloch, 1793; E. spilotoceps non Schultz, 1953;
E. hexagonatus (non Forster, 1801).
FAO names: En - Barredchest grouper; Fr - Mérou écharpe; Sp - Mero bandeado.
standard length); head length 2.3 to 2.6 times in standard length; caudal peduncle depth 3.8 to 4.3 times
in head length. Interorbital area flat or slightly concave, the dorsal head profile convex; preopercle rounded,
with shallow indentation just above corner and slightly enlarged serrae at the corner; upper edge of
operculum slightly convex; maxilla reaches past vertical at rear edge of eye; midlateral part of lower jaw
with 2 rows of teeth; nostrils subequal. First gill arch with 7 to 10 gill rakers on upper limb, 14 to 16 on lower
limb. Pyloric caeca about 25. Dorsal fin with XI spines and 16 to 18 soft rays, the third or fourth spine
longest, 2.6 to 3.9 times in head length and shorter than longest dorsal-fin rays, the interspinous
membranes incised; anal fin with III spines and 8 soft rays, the second and third spines subequal, 3.6
to 4.3 times in head length; caudal fin rounded, length of middle rays 1.65 to 1.9 times in head
length; pectoral-fin rays 17 to 19, the fin length 1.7 to 2.2 times in head length; pelvic fins reaching
anus, 2.2 to 2.6 times in head length. Lateral body scales smooth (except for area covered by
pectoral fin), with numerous auxiliary scales; lateral-line scales 48 to 52; lateral scale series 83 to
98. Colour: head, body, and fins pale, covered with close-set roundish brown spots of unequal size,
most of those on body larger than pupil; 4 groups of 2 or 3 spots at base of dorsal fin darker than
other spots on body; 2 oblique dark bands on side of chest; pectoral fins dusky, with dark blotch
at base and obscure dark spots, more distinct on inner surface of fin; spots on median fins darker
than those on body.
Habitat, biology, and fisheries: Shallow-water coral reefs and rocky areas. Apparently a small species;
females are mature at 17 cm standard length. Separate statistics are not available for this species, but
undoubtedly of importance in artisanal fisheries. Caught with hook-and-line, gill nets, lift nets, traps, and
spear.
Distribution: Northeastern
Indian Ocean: Southern India,
Sri Lanka, and southern Indo-
nesia (Bali and Lombok).
Epinephelus faveatus h a s
only recently been recognized
and distinguished from similar
species, so it is likely that its
distribution is continuous from
Pakistan to Indonesia.
Epinephelus fuscoguttatus (Forsskål, 1775)

Frequent synonyms / misidentifications: None / Epinephelus polyphekadion (non Bleeker, 1849)
(= E. microdon (Bleeker, 1856)).
FAO names: En - Brownmarbled grouper; Fr - Mérou marron; Sp - Mero manchado.
standard length); head length 2.3 to 2.5 times in standard length. Interorbital area flat or slightly
concave; dorsal head profile of adults indented at eyes and distinctly convex from there to
dorsal-fin origin; upper edge of operculum distinctly convex, descending almost vertically to rear
end of operculum; ventral edge of preorbital bone indented below nostrils. Maxilla extends well
posterior to eye; midlateral part of lower jaw with 3 or 4 rows of teeth, the inner teeth about twice length of
outer teeth; canines inconspicuous; nostrils close together; posterior nostrils triangular, 4 to 7 times
larger than anteriors in adults. First gill arch with 10 to 12 gill rakers on upper limb, 17 to 21 on
lower limb (but rudiments often difficult to count); gill rakers short and stout, raker at angle
subequal to longest gill filaments, other rakers distinctly shorter. Dorsal fin with XI spines and 14 or
15 soft rays, the third or fourth spines longest, 2.9 to 3.5 times in head length and obviously shorter than
longest dorsal-fin rays, the interspinous membranes distinctly incised; anal fin with III spines and 8 soft
rays; caudal fin rounded; pectoral-fin rays 18 to 20, the fin length 1.7 to 2.1 times in head length; pelvic
fins not reaching anus, 2.0 to 2.5 times in head length. Lateral body scales smooth, with auxiliary scales;
lateral-line scales 52 to 58; lateral scale series 102 to 115. Colour: pale yellowish brown, with 5 vertical
series of dark brown blotches that are very irregular in outline; head, body, and fins covered with
close-set small brown spots, those on the dark blotches much darker than spots in between
blotches; small black saddle spot on rear half of peduncle; 2 or 3 faint, dark bars at side of jaws.
Size: Maximum total length at least 95 cm (120 cm and 11 kg in the Philippines).
Habitat, biology, and fisheries: Shallow coral reefs and rocky bottoms to depths of 60 m; juveniles in
seagrass areas. Feeds on fishes, crabs, and cephalopods. Implicated in ciguatera fish poisonings at some
Pacific localities. Occasionally seen in local markets. Used in cage culture operations in Singapore. Caught
with hook-and-line, traps, and
spear.
cific region from Red Sea to
Phoenix Islands, including In-
donesia, Japan, Philippines,
New Guinea, tropical coast of
Australia, New Caledonia,
and most tropical islands of
the region; not known from
Persian Gulf, Asian mainland,
Hawaii or French Polynesia.
2502 Bony Fishes
Epinephelus heniochus Fowler, 1904

Frequent synonyms / misidentifications: Epinephelus hata Katayama, 1953 / Epinephelus epistictus
(non Temminck and Schlegel, 1842).
FAO names: En - Bridled grouper; Fr - Mérou bride; Sp - Mero embridado.
standard length); head length 2.2 to 2.4 times in standard length. Interorbital area slightly convex; dorsal
head profile convex; preopercle angular, with 2 to 4 distinctly enlarged serrae at the angle; upper
edge of operculum approximately straight; maxilla usually reaches 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, especially in upper jaw; midlateral part of lower jaw with 2 rows of teeth; rear nostrils about
twice as large as anterior nostrils. First gill arch with 7 to 9 gill rakers on upper limb, 14 to 16 on lower
limb. Pyloric caeca 7 or 8. Dorsal fin with XI spines and 14 or 15 soft rays, the third or fourth spines longest,
3.0 to 3.8 times in head length and shorter than longest dorsal-fin rays; anal fin with III spines and 8 soft
rays; caudal fin rounded; pectoral-fin rays 16 to 18, the fin length 1.6 to 1.9 times in head length; and pelvic
fins 1.9 to 2.4 times in head length. Lateral body scales rough, without auxiliary scales; lateral-line
scales 54 to 60; lateral scale series 89 to 100. Colour: head and body pale brown dorsally, shading
to whitish or pale pink ventrally; some specimens with minute brownish black dots on body and
rear part of head; faint dark brown stripe from eye to end of operculum, another darker stripe from
lower edge of eye to subopercle and a third from edge of preorbital to interopercle; pectoral fins
hyaline greyish yellow; lower part of caudal fin sometimes darker than rest of fin; margin of
interspinous dorsal fin membranes yellow.
Habitat, biology, and fisheries: Most specimens have been taken with trawls on mud or silty sand bottom
in depths of 40 to 235 m. Nothing has been published on the biology of this species. Reported as “common
but not abundant” in the South China Sea, and “not a very popular food fish” in the markets of Singapore.
Caught with trawls and vertical longlines.
Distribution: Tropical west-
ern Pacific: Singapore, Indo-
nesia, the Philippines, Gulf of
Thailand, Viet Nam, northern
Australia, and New Britain. Al-
though K atay ama’s type-
s p e c i m e n s o f Epinephelus
hata were purchased from the
Nagasaki fish market, the ab-
sence of other records of E.
heniochus from Japan indi-
cates that the types of E. hata
may have been caught south
of Japan.
Epinephelus hexagonatus (Forster, 1801) (Plate III, 24)

Frequent synonyms / misidentifications: None / Epinephelus merra non Bloch, 1793; E. macrospilos
(non Bleeker, 1855); E. spilotoceps non Schultz, 1953; E. quoyanus (non Valenciennes, 1830).
FAO names: En - Starspotted grouper; Fr - Mérou mélifère; Sp - Mero mielero.
standard length); head length 2.5 to 2.6 times in standard length. Preopercle rounded, the ventral serrae
slightly enlarged; upper edge of operculum convex; midlateral part of lower jaw with 3 to 5 rows of teeth;
nostrils subequal. First gill arch with 7 to 9 gill rakers on upper limb, 17 to 19 on lower limb. Dorsal fin with
XI spines and 15 to 17 soft rays, the fifth to ninth spines subequal (2.5 to 2.8 times in head length)
and slightly shorter than longest soft rays; anal fin with III spines and 8 soft rays, the second spine 2.1
to 2.5 times in head length, distinctly longer than third spine or depth of peduncle; caudal fin rounded;
pectoral-fin rays 17 to 19, the fin length 1.6 to 1.9 times in head length; pelvic fins not reaching past
anus, 1.8 to 2.1 times in head length. Lateral body scales rough, with auxiliary scales; lateral-line scales
61 to 70; lateral scale series 93 to 114. Colour: head and body covered with polygonal (mostly
hexagonal) brown spots that tend to merge, leaving only conspicuous triangular white dots at
corners of the polygons; dark spots on belly and ventral part of head more rounded and separated,
and often reddish brown; 4 or 5 brownish black saddle blotches (formed by groups of darker spots)
on dorsal part of body and caudal peduncle, the first 4 extending onto base of dorsal fin; irregular
dark bar, formed by darker polygonal spots, on lower part of body below each saddle blotch; large
brown or olive spot just behind eye, often joined to similar spot on opercle; fins with close-set dark
brown or reddish brown spots and white dots, except distal half of pectoral fins with faint dark
spots and no white dots; pelvic and anal fins with pale edge and dark brown submarginal band;
interspinous dorsal-fin membranes with dark brown triangle and short white or pale yellow filament
behind tip of each spine.
Habitat, biology, and fisheries: A common coral-reef species, usually found in shallow outer-reef areas
exposed to surge. Feeds mainly on fishes and crustaceans. Too small to be of commercial importance
except as a food fish in artisanal fisheries. Caught with hook-and-line, traps, spear, and gill nets.
West Pacific region from ?
Kenya to the Pitcairn Group. ?
Known from most tropical ?
Indo-Pacific islands (both on
and off the Pacific Plate), in-
cluding islands of the Great
Barrier Reef, but no verifiable
records from the mainland
coast of Australia, Indonesia,
the Philippines, Taiwan Prov-
ince of China, or Hawaii.
2504 Bony Fishes
Epinephelus irroratus (Forster, 1801) (Plate IV, 25)

Frequent synonyms / misidentifications: Epinephelus spiniger Günther, 1859; E. albopunctulatus
Boulenger, 1895 / None.
FAO names: En - Marquesan grouper; Fr - Mérou Marquises; Sp - Mero marquesano.
slightly convex; preopercle corner rounded and slightly indented, the lower serrae scarcely enlarged; upper
edge of operculum almost straight; maxilla reaching to or just past vertical at rear edge of eye; midlateral
part of lower jaw with 2 to 4 rows of teeth; rear nostril about twice size of anterior nostril. First gill arch with
6 to 8 gill rakers on upper limb, 13 to 16 on lower limb. Dorsal fin with XI spines and 16 soft rays, the
second spine elongated in adults, more than twice length of third spine, the interspinous mem-
branes not or only slightly incised; anal fin with III spines and 8 soft rays; caudal fin truncate to slightly
rounded; pectoral-fin rays 18 to 20, the fin length 1.6 to 1.9 times in head length; pelvic fins not reaching
past anus, 1.8 to 2 times in head length. Lateral body scales rough, with auxiliary scales; lateral-line scales
70 to 75; lateral scale series 117 to 136. Colour: reddish brown with white dot on each scale (dots
may not persist in preservative); maxillary streak dark reddish brown; spinous dorsal fin with
prominent dark red margin; white line along rear edges of median and pectoral fins.
Habitat, biology, and fisheries: Abundant in shallow water around fringing coral reefs. Nothing has been
published on the biology of this species, but it is of considerable interest to the artisanal fishery of the
Marquesas Islands. Caught
with hook-and-line, spear, and
traps.
Distribution: K no wn only
from the Marquesas Islands
and from a single specimen
taken at Minami Tori Shima
(Marcus Island), but the latter
record may be erroneous.
Epinephelus lanceolatus (Bloch, 1790)

Frequent synonyms / misidentifications: Promicrops lanceolatus (Bloch, 1790) / Epinephelus tauvina
(non Forsskål, 1775).
FAO names: En - Giant grouper; Fr - Mérou lancéolé; Sp - Mero lanceolado.
23.4 cm standard length juvenile

Diagnostic characters: Body robust, its width 1.5 to 1.75 times in body depth; body depth 2.4 to 3.4
times in standard length (for specimens 12 to 179 cm standard length); head length 2.2 to 2.7 times in
standard length; eye diameter 5.8 to 14 times in head length; interorbital width 3.3 (at 177 cm
standard length) to 6.2 (at 12 cm standard length) times in head length. Interorbital area flat to slightly
convex, the dorsal head profile convex; preopercle finely serrate, the corner rounded; upper edge of
operculum convex; maxilla reaching past eye; midlateral part of lower jaw with 2 or 3 rows of teeth
(at 20 to 25 cm standard length) increasing to 15 or 16 rows in a specimen of 177 cm standard
length; canine teeth at front of jaws small or absent; nostrils subequal. First gill arch of juveniles with
8 to 10 gill rakers on upper limb and 14 to 17 on lower limb; rudiments in adults are difficult to distinguish
from bony platelets covering gill arch. Dorsal fin with XI spines and 14 to 16 soft rays, the third to eleventh
spines subequal, shorter than longest soft rays and 3.1 to 5.7 times in head length; anal fin with III
spines and 8 soft rays; caudal fin rounded; pectoral-fin rays 18 to 20, the fin length 1.8 to 2.2 times in head
length; pelvic fins not reaching anus, 2.1 to 2.6 times in head length. Lateral body scales smooth, with
auxiliary scales; lateral-line scales 54 to 62, the anterior scales with branched tubules (except small
juveniles); lateral scale series 95 to 105. Colour: small juveniles (less than 15 cm) yellow, with 3
irregular black areas, the first from spinous dorsal fin to belly and chest and extending onto head,
the second from base of soft dorsal fin to anal fin and the last at base of caudal fin; subadults (25
to 60 cm) with irregular white or yellow spots on the black areas and fins with black spots; adults
(90 to 165 cm) dark brown with faint mottling, the fins with numerous small black spots; large adults
(180 to 250 cm) dark brown, fins darker.
Size: One of the largest groupers, attains 260 cm total length; maximum weight 288 kg.
Habitat, biology, and fisheries: Occurs in caves on coral reefs and around wrecks; juveniles and adults
also found in estuaries and harbours. Caught at depths of 100 m, but more often in shallow water. Feeds
on crustaceans (mainly spiny lobsters and large crabs), a variety of fishes including sharks and batoids,
and juvenile sea turtles. Not common enough to be of
commercial importance; rare in some places due to
spearfishing. Caught with hook-and-line and spear.
Distribution: Indo-Pacific region from Red Sea to South
Africa and eastwards to Hawaiian and Pitcairn islands,
including Japan, Taiwan Province of China, Philippines,
Indonesia, Thailand, Australia, and islands of the Central
Pacific.
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2506 Bony Fishes
Epinephelus latifasciatus (Temminck and Schlegel, 1842) (Plate IV, 26)

Frequent synonyms / misidentifications: Epinephelus grammicus (Day, 1867) / Epinephelus epistictus
(non Temminck and Schlegel, 1842); E. poecilonotus (non Temminck and Schlegel, 1842).
FAO names: En - Striped grouper; Fr - Mérou à bandes; Sp - Mero abanderado.

standard length (for specimens 13 to 62 cm standard
length); head length 2.3 to 2.6 times in standard length.
Interorbital area convex, the dorsal head profile convex;
preopercle angular, with 3 to 7 distinctly enlarged
serrae at angle; upper edge of operculum distinctly
convex; maxilla reaches past vertical at rear edge of eye; 7.6 cm standard length
midlateral part of lower jaw with 2 or 3 rows of teeth;
nostrils subequal. First gill arch with 8 to 11 gill rakers on juvenile
upper limb, 15 to 18 on lower limb. Dorsal fin with XI spines and 12 to 14 soft rays, the third or fourth
spine longest, 2.9 to 3.9 times in head and not much shorter than longest soft rays, the interspinous
membranes incised; anal fin with III spines and 8 soft rays; caudal fin truncate in large adults, the rear
margin convex in juveniles; pectoral-fin rays 17 to 19, the fin length 1.8 to 2.2 times in head length; pelvic
fins not reaching anus, 2.2 to 2.6 times in head length. Lateral body scales smooth; lateral-line scales 56
to 65; lateral scale series 91 to 106. Colour: juveniles lavender-grey or pale brownish, shading to
whitish ventrally; 2 black-edged white longitudinal bands, the upper band from above eye to anterior
dorsal-fin rays, and the lower band from below eye to lower caudal-fin rays; dorsal and caudal fins
with black spots and streaks; white bands disappearing on adults, the dark edges breaking into
dashes and spots; head and body of large adults uniformly grey.
Size: Maximum total length 157 cm; maximum weight 58.6 kg.
Habitat, biology, and fisheries: The preferred habitat seems to be bottoms of low relief; adults are taken
on coarse sand or rocky areas, while juveniles are found on silty-sand and mud bottom. Depths range from
20 to at least 230 m. Common in markets of the Persian Gulf, Hong Kong, Singapore, and Japan. Caught
with hook-and-line, longlines, trawls, and traps.
cific region, including the Red
Sea, Persian Gulf, Gulf of
Oman, Pakistan, coast of In-
dia, Viet Nam, Hong Kong,
China (Shanghai), Korea,
southern Japan, Taiwan Prov-
ince of China, and northwest
Australia. The banded grouper
seems to prefer continental lo-
calities, but it is not known from
the east coast of Africa, is-
lands of the Indian Ocean,
Philippines, or New Guinea.
Epinephelus longispinis (Kner, 1864) (Plate IV, 27)

Frequent synonyms / misidentifications: None / Epinephelus gaimardi (Valenciennes, 1830); E. fario
(nomen dubium); E. maculatus (non Bloch, 1790).
FAO names: En - Longspine grouper; Fr - Mérou longues épines; Sp - Mero espigón.
standard length); head length 2.4 to 2.6 times in standard length. Interorbital area flat to slightly convex,
the dorsal head profile convex; preopercle corner with enlarged serrae and a shallow indentation just
above the corner; upper edge of operculum straight or slightly convex; maxilla reaches to or past
vertical at rear edge of eye, the ventral edge with a blunt hook-like process distally in fishes larger
than 35 cm standard length; midlateral part of lower jaw with 2 rows of teeth; posterior nostrils not much
larger than anteriors. First gill arch with 8 to 11 gill rakers on upper limb, 15 to 17 on lower limb. Dorsal fin
with XI spines and 16 or 17 soft rays, the third or fourth spine longest, 2.1 to 2.6 times in head length
and distinctly longer than longest dorsal-fin rays, the interspinous membranes slightly incised
between anterior spines and not incised between posterior spines; anal fin with III spines and 8 soft
rays; caudal fin convex; pectoral-fin rays 17 to 19, the fin length 1.5 to 1.9 times in head length; pelvic fins
not reaching anus, 1.8 to 2.2 times in head length. Lateral body scales rough, with numerous auxiliary
scales; lateral-line scales 49 to 53; lateral scale series 98 to 121. Colour: head and body pale greyish
brown, covered with small, dark reddish brown spots that are round and widely spaced on head
and front half of body, but obliquely elongated, closer together and darker posteriorly; fins with
similar dark brown spots; a row of dark spots along distal margin of soft dorsal and caudal fins.
Juveniles with fewer dark spots on head and body, and 2 to 4 dark blotches on body at base of
dorsal fin and extending onto fin; ventral part of head and body with irregular white spots; white
blotch on upper part of operculum.
Habitat, biology, and fisheries: Usually found on coral reefs or rocky areas and occasionally on sandy
bottom; depths of capture range from 1 to 70 m. Feeds mainly on crustaceans, especially crabs and
stomatopods, and rarely on fish and squid. Caught with hook-and-line, spear, traps, and in trawls.
Distribution: Continental and
insular localities in Indian
Ocean from Kenya to South
Africa and east to the Watu-
bela Group of the eastern
Banda Sea. Not known from
the Red Sea or Persian Gulf.
2508 Bony Fishes
Epinephelus macrospilos (Bleeker, 1855) (Plate IV, 28)

Frequent synonyms / misidentifications: None / Epinephelus quoyanus (non Valenciennes, 1830);
E. faveatus (non Valenciennes, 1828); E. corallicola (non Valenciennes, 1828); E. howlandi (non Günther,
1873).
FAO names: En - Snubnose grouper; Fr - Mérou tapis; Sp - Mero alfombrado.

standard length; head length 2.3 to 2.6 times in standard
length; caudal peduncle depth 3.2 to 3.7 times in head
length. Interorbital area flat or slightly concave, the dorsal
head profile of adults with a ventral bend at orbits; preoper-
cle rounded, with minute serrae mostly covered by skin
and a shallow indentation just above the “corner”; upper
edge of operculum straight or slightly convex; maxilla juvenile 5.5 cm standard length
reaches to or past vertical at rear edge of orbit, the ventral
edge smoothly curved at distal expansion; lower jaw strongly projecting, the midlateral part with 2 to 4 rows
of teeth; rear nostrils usually distinctly larger than anterior nostrils. First gill arch with 21 to 26 gill rakers,
of which 7 to 9 on upper limb and 14 to 17 on lower limb. Pyloric caeca 25 to 27. Dorsal fin with XI spines
and 15 to 17 soft rays, the third or fourth spines longest, 2.3 to 3.4 times in head length and shorter than
longest dorsal-fin rays; anal fin with III spines and 8 soft rays, the second and third spines subequal,
2.8 to 4.2 times in head length; caudal fin rounded, length of middle rays 1.5 to 2.0 times in head
length; pectoral-fin rays 17 to 20, the fin length 1.5 to 2 times in head length; pelvic-fin length 1.8 to 2.5
times in head length. Lateral body scales smooth, except for area covered by pectoral fins;
lateral-line scales 48 to 52; lateral scale series 86 to 103. Colour: head and body pale greyish brown,
with dark brown spots (centre of spots darker than the diffuse edges) that are large and well-sepa-
rated in young, becoming relatively smaller, more numerous and closer together in adults; median
and pelvic fins with similar dark spots; pectoral fins usually dusky with a few dark spots and white
line along the edge; some specimens with faint oblique dark line across lower part of chest; median
fins with narrow pale margin posteriorly; no dark spots on underside of lower jaw. Juveniles with
prominent white blotches as shown in figure.
Habitat, biology, and fisheries: Coral reefs to depths of at least 44 m. Feeds on crustaceans (mainly
crabs), fishes, octopus, and squid. Common in some areas and important in artisanal fisheries. Caught
with hook-and-line, spear, and traps.
Distribution: Indo-Pacific region from
Africa (Kenya to South Africa) to Central
Pacific, including Cocos-Keeling,
Indonesia, Okinawa, Western Australia
(Scott Reef), Great Barrier Reef,
Marquesas, and most western Pacific
islands (both on and off the Pacific Plate).
Not known from Red Sea, Persian Gulf,
Asian continent, or Hawaii.
Epinephelus maculatus (Bloch, 1790) (Plate IV, 29)

Frequent synonyms / misidentifications: Epinephelus medurensis (Günther, 1873) / Epinephelus
macrospilos (non Bleeker, 1855).
FAO names: En - Highfin grouper; Fr - Mérou haute voile; Sp - Mero aleta alta.

length. Interorbital area flat to slightly convex, the dorsal head
profile convex; preopercle with a shallow indentation just
above the enlarged serrae at the corner; upper edge of
operculum straight or slightly convex; maxilla reaches to or
past vertical at rear edge of eye, the ventral edge with a blunt
hook-like process distally in fishes larger than 35 cm juvenile 8 cm standard length
standard length; midlateral part of lower jaw with 2 rows of
teeth; nostrils subequal. First gill arch with 8 to 10 gill rakers on upper limb, 15 to 17 on lower limb. Pyloric
caeca 30 to 45. Dorsal fin with XI spines and 15 to 17 soft rays, the third or fourth spine longest, 2.1 to
2.6 times in head length and distinctly longer than dorsal-fin rays, the interspinous membranes
slightly incised between anterior spines and not incised posteriorly; anal fin with III spines and 8 soft
rays, the third spine longest, 3.3 to 4.1 times in head length and not longer than caudal peduncle
depth; caudal fin convex or rounded; pectoral-fin rays 17 to 19, the fin length 1.5 to 1.9 times in head
length; pelvic fins reaching to or near anus, 1.7 to 2.0 times in head length. Lateral body scales
distinctly rough, with numerous auxiliary scales; lateral-line scales 49 to 52; lateral scale series
102 to 120. Colour: head, body, and fins of adults pale brown, covered with small, round to
hexagonal, close-set, dark brown spots, extending onto ventral parts of body and head; 2 large
diffuse dusky areas on dorsal part of body and dorsal fin, the largest, extending over dorsal fin
from second to fifth spines, the second dark blotch from tenth spine to second soft ray, the dorsal
fin pale between these dark blotches, but with small dark spots. One resting colour phase is broadly
mottled with large dark and pale areas; in the pale areas, the usual dark brown spots are mostly
whitish, with small dark brown centres. Small juveniles (5 to 10 cm) yellowish brown, with well-
separated small black spots (mainly on head and fins) and prominent, white blotches and spots,
the largest on middle of dorsal fin.
Habitat, biology, and fisheries: Coral reefs at depths of 2 to 100 m. Prey comprises mainly small fishes,
crabs, and octopuses. Not common, but of
some interest in artisanal fisheries. Caught
with hook-and-line, spear, and traps.
Distribution: Eastern Indian Ocean and
Western Pacific from Cocos-Keeling Is-
lands, Indonesia, Hong Kong, Ryukyu Is-
lands, Ogasawara Islands, Philippines,
New Guinea, Great Barrier Reef, Lord
Howe Island, New Caledonia, New Ireland,
Fiji, Samoa, Palau, Caroline Islands, Mar-
shall Islands, and Gilbert Islands.
2510 Bony Fishes
Epinephelus magniscuttis Postel, Fourmanoir, and Guézé, 1963 (Plate IV, 30)
Frequent synonyms / misidentifications: Epinephelus pseudomorrhua Postel, Fourmanoir, and Guézé,
1963 / Epinephelus epistictus (non Temminck and Schlegel, 1842); E. poecilonotus (non Temminck and
Schlegel, 1842).
FAO names: En - Spotted grouper; Fr - Mérou grandes écailles; Sp - Mero bacalao.
Diagnostic characters: Body depth 2.7 to 3.2 times in standard length; head length 2.2 to 2.4 times
in standard length. Interorbital area flat to convex, the dorsal head profile almost straight; preopercle
angular, with 2 to 4 distinctly enlarged serrae at angle; upper edge of operculum slightly convex;
maxilla reaches to below rear half of eye, the ventral edge with a low step at the distal expansion; midlateral
part of lower jaw with 2 rows of teeth, the inner teeth larger than outer ones; posterior nostrils not much
larger than anteriors. First gill arch with 8 or 9 gill rakers on upper limb, 16 or 17 on lower limb. Pyloric
caeca 7. Dorsal fin with XI spines and 14 or 15 soft rays, the third or fourth spines longest, 2.5 to
3.0 times in head length and longer than longest dorsal-fin rays, the interspinous membranes deeply
1.7 to 2.0 times in head length; pelvic fins not reaching anus, 2.1 to 2.3 times in head length. Lateral body
scales distinctly rough and without auxiliary scales; lateral-line scales 55 to 62; lateral scale series 103 to
122. Colour: generally pale brown, with small dark brown (or greyish green) spots unevenly
scattered on dorsolateral parts of head (posterior to eyes) and body, dorsal fin, and caudal fin; no
spots on body below level of pectoral fins or on anal and paired fins. Juveniles are said to have 6 or
7 dark lines running horizontally along the body and these disappear with age.
Size: Maximum total length 150 cm; maximum weight about 50 kg.
Habitat, biology, and fisheries: Deep water (128 to 300 m) in the vicinity of coral reefs. Biology unknown.
An excellent food fish but apparently rare in the western Pacific region. Probably of some importance in
artisanal fisheries.
cific from South Africa,
Mozambique, Réunion, Mau-
ritius, New Caledonia, Philip-
pines, New Guinea, New
Ireland, and Fiji.
Epinephelus malabaricus (Bloch and Schneider, 1801) (Plate IV, 31)

Frequent synonyms / misidentifications: None / Epinephelus tauvina (non Forsskål, 1775).
FAO names: En - Malabar grouper; Fr - Mérou malabare; Sp - Mero malabárico.
juvenile 2.3 cm standard length

Diagnostic characters: Body elongate, its depth 3.0 to 3.7 times in standard length; body width 1.4
to 1.9 times in body depth; head length 2.3 to 2.6 times in standard length; interorbital width 4.5 to 6.5
times in head length, and 2.1 to 3.2 times in upper jaw length; preopercle with enlarged serrae at the
angle; upper edge of operculum almost straight; maxilla extends past vertical at rear edge of orbit; upper
jaw length 17 to 22% of standard length; snout length 1.7 to 2.0 times in upper jaw length; midlateral part
of lower jaw with 2 to 5 rows of teeth; large adults with rear nostrils slightly larger than anterior nostrils.
First gill arch with 23 to 27 gill rakers, of which 8 to 11 on upper limb and 14 to 18 on lower limb; rudiments
difficult to distinguish from small bony platelets on gill arch. Pyloric caeca numerous (more than 80
branches). Dorsal fin with XI spines and 14 to 16 soft rays, the third to fifth spines usually slightly longer
than posterior spines, 3.1 to 4.0 times in head length and distinctly shorter than longest soft rays; anal fin
with III spines and 8 soft rays, the third spine usually longest; caudal fin rounded; pectoral-fin rays 18 to
20, the fin length 1.7 to 2.2 times in head length; and pelvic fins 2.1 to 2.6 times in head length. Lateral
body scales rough, with auxiliary scales; lateral-line scales 54 to 64; anterior lateral-line tubes of large
adults with 2 to 4 branches; lateral scale series 101 to 117. Colour: head and body brownish, covered
with small, well-separated, blackish brown spots which extend onto chest, lower jaw and gular area,
and roof of mouth; head and body also with scattered white spots and blotches; 5 irregular, oblique,
dark brown bars (more or less interrupted by pale spots) often visible on body; fins with scattered small
black spots.
Size: Maximum size uncertain because of confusion with other species of large groupers; attains at least
115 cm total length and a weight of 25 kg.
Habitat, biology, and fisheries: Coral and rocky reefs, tidepools, estuaries, mangrove swamps, and
sandy/mud bottom from shore to a depth of 150 m. Feeds on fishes, crustaceans, and octopuses. One of
the most common groupers in markets of the Indo-West Pacific region, and widely used in the aquaculture
industry. Caught with trawls, longlines, traps, spear, and hook-and-line.
Distribution: Indo-Pacific area from Red
Sea and South Africa to Japan, Australia,
Palau, Yap, and Fiji. Occurs in continental
and insular localities: Indonesia, Singa-
pore, Philippines, Taiwan Province of
China, China, Papua New Guinea, New
Ireland, Caroline Islands, New Caledonia,
and Tonga. In Australia, it ranges from the
Northern Territory to New South Wales. Re-
cently found in eastern Mediterranean.
2512 Bony Fishes
Epinephelus merra Bloch, 1793 (Plate IV, 32)

Frequent synonyms / misidentifications: None / Epinephelus macrospilos (non Bleeker, 1855);
E. hexagonatus (non Forster, 1801); E. melanostigma non Schultz, 1953; E. spilotoceps non Schultz, 1953;
E. faveatus (non Valenciennes, 1828); E. quoyanus (non Valenciennes, 1830).
FAO names: En - Honeycomb grouper; Fr - Mérou gâteau de cire; Sp - Mero panal.

length; caudal peduncle depth 3.2 to 4.1 times in head
length. Preopercle rounded or subangular, the serrae at angle
enlarged; upper edge of operculum almost straight; maxilla
reaches past eye; midlateral part of lower jaw with 2 to 4 rows
of teeth, the inner teeth about twice length of outer ones; nostrils
subequal or rear nostrils larger. First gill arch with 6 to 9 gill juvenile
rakers on upper limb, 14 to 17 on lower limb. Pyloric caeca 8.
Dorsal fin with XI spines and 15 to 17 soft rays, the third to last spines subequal, the longest 2.4 to 3.2
times in head length; anal fin with III spines and 8 soft rays, second and third spines subequal, 2.1 to
3.0 times in head length and longer than depth of peduncle; caudal fin rounded; pectoral-fin rays 16
to 18, the fin length 1.5 to 1.8 times in head length; pelvic fins 1.8 to 2.2 times in head length. Lateral body
scales rough, with auxiliary scales; lateral-line scales 48 to 54; lateral scale series 98 to 114. Colour: head,
body, and fins pale, covered with close-set, dark brown or reddish brown spots, the interspaces
forming an irregular pale reticulum; spots on ventral part of body paler, more widely spaced and diffuse;
some midlateral spots often joined to form horizontal bands; 5 irregular dark bars may be displayed by
differential darkening of some body spots, but black dorsal blotches never present; dark spots on median
fins become smaller distally; pectoral fins covered with distinct small black spots, largely confined to
the rays (the best diagnostic colour character for Epinephelus merra); tips of interspinous dorsal-fin
membranes white or pale yellow, with a small submarginal black spot.
Habitat, biology, and fisheries: A coral-reef species typically found around patch reefs in depths less
than 20 m, but occasionally seen at 50 m. Feeds on fishes, crustaceans, and cephalopods. Females
mature at 14 cm standard length and sexual transition occurs at 18 to 21 cm. Important to artisanal fisheries
because of its abundance in shallow water. Caught with handlines, fish traps, and spear.
Distribution: Indo-Pacific region from
South Africa to French Polynesia. Primarily
an insular species; not known from the Red
Sea or Persian Gulf. Occurs at most tropi-
cal islands in the Indian Ocean and west-
ern Central Pacific, both on and off the
Pacific Plate, and ranges from Japan to
northwestern Australia, Queensland, Lord
Howe Island, and eastwards to the
Tuamotu Archipelago.
Epinephelus morrhua (Valenciennes, 1833) (Plate V, 33)

Frequent synonyms / misidentifications: Epinephelus cometae Tanaka, 1927 / Epinephelus
poecilonotus (non Temminck and Schlegel, 1842); E. radiatus (non Day, 1867); E. tuamotuensis non
Fourmanoir, 1971.
FAO names: En - Comet grouper; Fr - Mérou comète; Sp - Mero cometa.
in standard length. Preopercle with a shallow indentation just above enlarged serrae at the corner;
upper edge of operculum almost straight; maxilla reaches to or past vertical at rear edge of eye; midlateral
part of lower jaw with 2 rows of teeth. Adults with rear nostrils 2 or 3 times larger than anterior nostrils.
First gill arch with 8 to 10 gill rakers on upper limb, 15 to 18 on lower limb; longest gill rakers shorter than
longest gill filaments. Dorsal fin with XI spines and 14 or 15 soft rays, third or fourth spines longest,
2.6 to 3.3 times in head and about equal to longest dorsal-fin rays; anal fin with III spines and 7 or 8
soft rays, second and third spines subequal; caudal fin convex; pectoral-fin rays 17 or 18, the fin length
1.8 to 2.2 times in head length; pelvic fins not reaching anus, 2.0 to 2.7 times in head length. Lateral body
scales rough, some specimens with a few auxiliary scales; lateral-line scales 55 to 64; lateral scale series
108 to 125. Colour: head and body buff, with dark brown bands as follows: a bifurcate band begins
behind eye, the upper branch joining a dark brown saddle blotch on nape, the lower branch running
to lower opercular spine and continuing on body as a midlateral band that bifurcates above pectoral
fin, the upper branch running to a dark blotch at base of third to seventh dorsal-fin rays, the lower
branch curving up to base of last 4 dorsal-fin rays; another dark band from upper edge of operculum
to base of fifth to ninth dorsal-fin spines; dark band from lower edge of eye to pectoral-fin base,
thence as a broken band along lower part of body, curving up to dorsal part of peduncle; dark band
from maxillary groove to rear end of interopercle. Small dark brown spots often present in pale areas
between bands and usually arranged in series paralleling the bands; fins generally unmarked; pectoral fins
hyaline yellow.
Size: Maximum total length 73 cm; maximum weight 5 kg.
Habitat, biology, and fisheries: A common deep-reef species usually found in depths of 80 to 370 m.
Sexual transition probably occurs at about 40 to 45 cm standard length. Caught with hand-line, long-line,
gill net, and trawls.
western Indian Ocean to Cen-
tral Pacific, including An-
daman Sea, Indonesia, Viet
Nam, Philippines, Hong
Kong, southern Japan,
Ogasawara Islands, Mariana
Islands, Palau, Guam, Papua
New Guinea, Australia
(Northern Territory to south-
ern Queensland), New Ire-
land, New Caledonia,
Rotuma, Fiji, and Cook Islands.
2514 Bony Fishes
Epinephelus ongus (Bloch, 1790) (Plate V, 34)

Frequent synonyms / misidentifications: Epinephelus slacksmithi Whitley, 1959 / Epinephelus
summana (non Forsskål, 1775); E. caeruleopunctatus (non Bloch, 1790).
FAO names: En - Whitestreaked grouper; Fr - Mérou à flocons; Sp - Mero nubifero.

length. Dorsal head profile slightly convex; interorbital
area flat; preopercle rounded, the serrae small and
mostly covered by skin; upper edge of operculum
strongly convex, the rear edge almost vertical, the
upper spine extending posterior to lower spine; maxilla
reaches to or slightly past vertical at rear edge of eye; juvenile 11.2 cm standard length
small, embedded scales on maxilla; teeth small, 2 to 4
rows at midlateral part of lower jaw; canines at front of jaws small or absent. Rear nostril diameter about
twice that of anterior nostril. First gill arch with 8 to 10 gill rakers on upper limb, 15 to 18 on lower limb
(including 6 to 8 rudiments on each limb). Dorsal fin with XI spines and 14 to 16 soft rays, the third or fourth
spine longest, 2.6 to 3.4 times in head length, the interspinous membranes incised; anal fin with III spines
and 8 soft rays; caudal fin rounded; pectoral fins large and fleshy, with 15 to 17 rays, the fin length
1.4 to 1.7 in head length; pelvic fins end well short of anus, their length 2.0 to 2.3 times in head length.
Lateral body scales rough, with auxiliary scales; lateral-line scales 48 to 53; lateral scale series 95 to
109. Colour: body brown, with numerous small white spots which, in specimens larger than 10 cm
standard length, are horizontally elongate and, in adults, tend to form wavy white lines; several
round or irregular pale blotches (eye-sized or larger) usually superimposed over small white spots;
head brown, with numerous small white spots dorsally behind eyes; black maxillary streak usually
hidden by maxilla; median fins with small white spots and streaks, the posterior margin blackish
with a white edge; paired fins greyish brown. Juveniles of about 6 cm standard length are brown,
covered with small, dark-edged white spots which are round, except on front part of dorsal fin where
they are elongated; white spots on paired fins become fewer and fainter with growth, and are absent
in adults.
Habitat, biology, and fisheries: Occurs in shallow water on coral reefs and rocky substrata, primarily on
inner coastal and lagoon reefs, even in
brackish water, where it frequents ledges
and caves at depths of 5 to 25 m. Although
this cryptic species is not very common, it
is of some interest to fisheries in Japan
(and probably elsewhere). Caught with
hook-and-line, spear, and traps.
Distribution: Indo-Pacific region, from Af-
rica to the Ryukyu and Marshall islands and
south to Fiji, New Caledonia, and northern
Australia.
Epinephelus poecilonotus (Temminck and Schlegel, 1842)

Frequent synonyms / misidentifications: None / Epinephelus morrhua (non Valenciennes, 1833);
E. epistictus (non Temminck and Schlegel, 1842); E. magniscuttis non Postel, Fourmanoir, and Guézé,
1963; E. radiatus (non Day, 1867); E. tuamotuensis non Fourmanoir, 1971.
FAO names: En - Dotdash grouper; Fr - Mérou morse; Sp - Mero punto y linea.
Diagnostic characters: Body depth less than head length,

2.6 to 3.1 times in standard length. Preopercle angle with
2 to 5 enlarged serrae; upper edge of operculum slightly
convex; adults with a step on ventral edge of maxilla; midside
of lower jaw with 2 rows of teeth. First gill arch with 8 to 10 gill
rakers on upper limb, 15 to 18 on lower limb; longest raker
about equal to longest gill filaments. Pyloric caeca 8 or 9.
Dorsal fin with XI spines and 14 or 15 soft rays, the third or
juvenile
fourth spines longest, 2.6 to 3.1 times in head length and 11.2 cm standard length
subequal to longest dorsal-fin ray, the interspinous mem-
branes deeply incised; anal fin with III spines and 8 soft rays; caudal fin convex; pectoral fins not fleshy,
with 17 to 19 rays, the fin length 1.7 to 2.1 and pelvic fins 2.0 to 2.6 times in head length. Lateral body
scales rough, with auxiliary scales on adults; lateral-line scales 54 to 63; lateral scale series 110 to 121.
Colour: small juveniles (5 to 12 cm standard length) pale yellowish grey, with oval black blotch on
body between bases of third to ninth dorsal-fin spines and extending to edge of fin between first
and seventh spines; a semicircular pale band passes ventrally round the oval black blotch and
isolates it from a dark brown band that begins broadly on nape and bifurcates just behind
operculum, the upper branch curving dorsally and expanding over basal half of dorsal fin between
ninth spine and fourth soft ray, the lower branch also curving dorsally and expanding at base of
last 4 dorsal-fin rays. Second curved brown band, parallel to the one above, runs from interorbital
area and eye to a black saddle-spot on caudal peduncle. Third (and narrowest) dark brown band
runs from lower edge of eye to subopercle, thence as a series of dark dots along ventral part of
body to caudal fin. Fins pale yellow, except where dark markings occur on dorsal fin. On specimens
of 20 to 30 cm, the oval black blotch at base of dorsal-fin spines and the dark brown bands on body
break into series of black spots; faint dark band from dark maxillary streak to corner of preopercle.
On specimens 40 to 50 cm, most dark spots on body are faint or have disappeared and the dark
bands on head are fading too; the fins are yellowish brown, the triangular interspinous dorsal fin
margins abruptly orange-yellow or brownish yellow; soft dorsal, anal, and caudal fins blackish
distally with a bluish white edge.
Size: Maximum total length at least 63 cm; maximum
weight at least 4 kg.
Habitat, biology, and fisheries: Occurs in depths of 45
to 375 m. Females are mature at 35 cm standard length.
Caught with hook-and-line, vertical longline, and trawls.
Distribution: Indo-West Pacific: South Africa to Viet
Nam, Japan, Korea, and Fiji.
2516 Bony Fishes
Epinephelus polyphekadion (Bleeker, 1849)

Frequent synonyms / misidentifications: Epinephelus microdon Bleeker, 1856; E. dispar var. B
(Playfair, 1867) / Epinephelus fuscoguttatus (non Forsskål, 1775).
FAO names: En - Camouflage grouper; Fr - Merou camouflage; Sp - Mero disfrazado.
Diagnostic characters: Body depth 2.7 to 3.1 times in standard length. Interorbital area flat, the dorsal
head profile evenly convex; preopercle rounded, finely serrate; upper edge of operculum distinctly
convex; maxilla reaches to or beyond vertical at rear edge of eye; midlateral part of lower jaw with 2 or 3
rows of teeth; fixed canines at front of jaws inconspicuous. Rear nostril diameter about twice diameter
of front nostrils. First gill arch with 24 to 27 gill rakers, of which 8 to 10 on upper limb and 15 to 17
on lower limb. Dorsal fin with XI spines and 14 or 15 soft rays, third or fourth spines longest, 2.7
to 3.3 times in head and distinctly shorter than longest dorsal-fin ray; anal fin with III spines and 8
soft rays; caudal fin rounded; pectoral-fin rays 16 to 18, the fin length 1.7 to 2.1 times, pelvic fins 1.9 to 2.4
times in head length. Lateral body scales rough, with auxiliary scales. Lateral-line scales 47 to 52; lateral
scale series 95 to 113. Colour: head, body, and fins pale brown, covered with small dark brown spots;
head and body with irregular dark blotches (more distinct on live specimens) superimposed over
the dark spots; promiment black saddle blotch on caudal peduncle; dark spots extend all over head,
including lower jaw, lips, branchiostegal membranes, gular area, and inside of mouth; numerous
small white spots on fins (more distinct on live fish) and a few on head and body. Juveniles with a pair of
blackish spots on each side of snout and a black spot at margin of second and third interspinous dorsal
membranes.
Size: Maximum total length at least 75 cm; maximum weight at least 4 kg.
Habitat, biology, and fisheries: Coral reefs; most abundant at islands, particularly atolls. Feeds mainly
on crustaceans (primarily portunid crabs, but also some scyllarid and panularid lobsters) and fishes;
molluscs are less important food items. Females mature at 38 cm standard length, and sex change occurs
at about 42 cm standard length. Important in artisanal fisheries, but occasionally implicated in cases of
ciguatera. Now uncommon at localities with heavy spearfishing. Currently popular in the aquaculture
industry of Singapore, where it is known as “marble grouper”. Caught with hook-and-line, spear, and traps.
West Pacific region from Red
S e a a n d M o z a m b i q u e to
French Polynesia. In the west-
ern Pacific it ranges from
southern Japan to Taiwan
Province of China, Indonesia,
southern Queensland, and
Lord Howe Island and east-
wards to the Tuamotus. Re-
ported from most islands of
the western Central Pacific.
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Epinephelus polystigma (Bleeker, 1853) (Plate V, 35)

Frequent synonyms / misidentifications: Epinephelus rahanus Popta, 1918 / None.
FAO names: En - Whitedotted grouper; Fr - Mérou points blancs; Sp - Mero punteado blanco.

standard length. Interorbital area flat; snout short, its
length subequal to eye diameter; preorbital depth less
than greatest width of upper lip; preopercle rounded,
finely serrate; upper edge of operculum mostly
straight. Anterior nostrils funnel shaped, the rear mar-
gin expanded as a flap reaching rear nostril; rear 22 mm standard length
nostril diameter 2 to 4 times larger than front nostril juvenile
diameter; maxilla reaches past vertical at rear edge of eye, the lower edge with a rounded step at front of
widest part; midlateral part of lower jaw with 2 or 3 rows of small subequal teeth; 2 short stout canines at
front of both jaws. First gill arch with 21 to 24 gill rakers, of which 8 or 9 on upper limb and 13 to 16 on
lower limb; gill rakers shorter than gill filaments. Pyloric caeca 8. Dorsal fin with XI spines and 15 or 16
soft rays, the third or fourth spines longest, 2.7 to 3.3 times in head length and distinctly shorter
than longest dorsal-fin rays, the interspinous membranes distinctly incised; anal fin with III spines and
8 soft rays; caudal fin well rounded; pectoral-fin rays 16 to 18, the fin length 1.6 to 1.8 times in head length;
pelvic fins 2.0 to 2.2 times in head length. Lateral body scales rough, with auxiliary scales. Lateral-line
scales 49 to 55; lateral scale series 81 to 91. Colour: head, body, and fins dark brown, covered (except
on pelvic fins) with white or pale yellow dots (dots may be absent in preserved specimens); soft
dorsal, anal, caudal, and pectoral fins with a narrow orange-red margin. Juveniles (8 to 9 cm
standard length) with some larger, dark-edged, round or horizontally elongate, white spots on body
(in addition to white dots); a smaller juvenile (2.2 cm standard length) taken in the same collection
has dark-edged white spots, as large or larger than pupil, in about 3 irregular rows on body.
Size: Maximum total length at least 48 cm total length.
Habitat, biology, and fisheries: Known only from brackish or fresh-water areas; the usual habitat seems
to be mangrove-lined estuaries. A female of 19 cm standard length contained ripe ovaries. Appears to be
a rare species; consequently,
of little interest to fisheries.
spears, and traps.
from the western Pacific: Phil-
ippines, Indonesia, Papua
New Guinea, New Ireland,
Solomon Islands, and Cape
York, Queensland.
2518 Bony Fishes
Epinephelus quoyanus (Valenciennes, 1830) (Plate V, 36)

Frequent synonyms / misidentifications: Epinephelus megachir (Richardson, 1846) / Epinephelus
macrospilos (non Bleeker, 1855); E. hexagonatus (non Forster, 1801); E. faveatus (non Valenciennes,
1828); E. spilotoceps non Schultz, 1853.
FAO names: En - Longfin grouper; Fr - Mérou longues ailes; Sp - Mero aleta larga.
Diagnostic characters: Body depth 2.8 to 3.2 times in stand-

ard length; caudal-peduncle depth 3.1 to 3.6 times in head
length. Preopercle rounded or subangular; upper edge of
operculum almost straight; midlateral part of lower jaw with
2 or 3 rows of teeth; lower jaw barely projecting in front of
upper jaw. Rear nostrils diameter about twice that of anterior juvenile 5.9 cm standard length
nostrils. First gill arch with 6 to 8 gill rakers on upper limb, 14
to 16 on lower limb. Pyloric caeca 21. Dorsal fin with XI spines and 16 to 18 soft rays, the fourth spine
usually longest, 2.3 to 3.0 times in head length and shorter than longest dorsal-fin rays; anal fin with III
spines and 8 soft rays, the second and third spines subequal, 2.8 to 3.8 times in head length, subequal
to depth of peduncle; caudal fin rounded, middle rays 1.35 to 2.0 times in head length; pectoral-fin rays
17 to 19, middle rays 26 to 35% of standard length, 1.2 to 1.7 times in head length; pelvic fins 1.6 to
2.1 times in head length. Lateral body scales rough; auxiliary scales present; lateral-line scales 48 to 52;
lateral scale series 80 to 96. Colour: head and body pale, with large, close-set, hexagonal to roundish,
dark brown spots (some spots reddish brown or black), dorsally the spots are so close together
that the pale interspaces form a reticulum, but ventrally the spots are more separated and their
margins more diffuse; similar dark spots on median fins; chest with 2 dark brown bands joining
below pectoral-fin bases and nearly joined anteriorly to enclose a large pale area ventrally and
another pale area between upper dark band and dark blotch on pectoral-fin base; dark spots on head
smaller anteriorly, but still 3 or 4 times larger than nostrils; ventral edge of anal and caudal fins and
leading edge of pelvic fins with white line and blackish submarginal band; pectoral fins mostly
dusky with faint dark spots (more distinct on Australian fish).
Habitat, biology, and fisheries: Usually found on inshore silty reefs; no records below 50 m. The enlarged
fleshy pectoral fins appear to be related to its habit of sitting on the substrate, with its pectoral fins in contact
with the bottom. Feeds on crustaceans,
fishes, and worms. Of some economic im-
portance at Hong Kong and Taiwan Prov-
ince of China. Usually caught with gill nets
and handlines.
Distribution: Western Pacific; from south-
ern Japan to tropical coast of Australia,
including China (also Taiwan Province of
China and Hong Kong), Philippines, Viet
Nam, Thailand, Indonesia, New Guinea.
Not known from islands of Micronesia,
Melanesia, or the Central Pacific.
Epinephelus radiatus (Day, 1867) (Plate V, 37)

Frequent synonyms / misidentifications: Epinephelus döderleinii Franz, 1910 / Epinephelus morrhua
(non Valenciennes, 1833); E. poecilonotus (non Temminck and Schlegel, 1842).
FAO names: En - Obliquebanded grouper; Fr - Mérou zébré; Sp - Mero acebrado.
Diagnostic characters: Body depth 2.6 to 3 times in standard length; head length 2.1 to 2.3 times in
standard length. Preopercle angle with 2 to 5 enlarged serrae; upper edge of operculum almost straight;
midlateral part of lower jaw with 2 rows of teeth. Adults with rear nostril diameter 2 or 3 times that of front
nostrils. First gill arch with 8 or 9 gill rakers on upper limb, 16 to 18 on lower limb; longest gill raker shorter
than longest gill filaments. Dorsal fin with XI spines and 13 to 15 soft rays, the third spine longest, 2.6
to 3.0 times in head length and longer than soft rays, the interspinous membranes deeply incised; anal
fin with III spines and 8 soft rays, second and third spines subequal; caudal fin convex to moderately
rounded; pectoral-fin rays 17 or 18, the fin almost transparent, its length 1.7 to 2.2 times in head length;
pelvic fins 2.1 to 2.8 times in head length. Lateral body scales rough, without auxiliary scales; lateral-line
scales 52 to 66; lateral scale series 102 to 120. Colour: small adults (25 to 50 cm) buff, with 5 oblique
dark-edged pale bands as follows: first band from upper half of eye, curving and broadening on
nape; second band branching from first band just behind eye, bifurcating on operculum, the upper
branch continuing dorsally, broadening on body and extending more broadly over middle of
spinous dorsal fin; third band continued as lower branch of second band, curving dorsally from
end of operculum and expanding at base of last 2 spines and first 3 or 4 soft rays of dorsal fin (this
band with a ventral extension from an included pale circle covered by tip of pectoral fin); fourth
band runs from rear end of dorsal fin, branching at midside, with 1 branch going towards anal-fin
origin, the other to rear end of anal-fin base; fifth band on caudal peduncle (sometimes divided into
2 short branches at lower edge of peduncle); dark bands include small black spots and pale
blotches, especially dorsally; pale interspaces (between dark bands) with small dark brown spots,
mainly in a row along middle of interspaces; dark brown line from lower edge of eye to edge of
subopercle; faint dark band from maxillary groove and continuing to edge of interopercle. Large
adults (50 to 65 cm) with dark-edged bands replaced by series of dark spots (except for dark line
running posteriorly from lower edge of eye); no spots on ventral third of body; dorsal fin and most
of caudal fin covered with small dark spots. Juveniles (12 to 25 cm) dark brown with black-edged
pale brown bands (= white markings on smaller fish) enclosing numerous small black spots. Small
juveniles (5 to 10 cm) dark greenish brown, with dark-edged immaculate white bands represent in
the pale interspaces on larger specimens; fins translucent white, except spinous dorsal fin coloured
like body.
Habitat, biology, and fisheries: Adults occur in depths
of 80 to 383 m; juveniles found in depths of 18 to 20 m.
Apparently rare but of some commercial importance in
Japan. Caught with handlines and vertical longlines.
Distribution: From Red Sea to Japan, including Gulf of
Aden, Gulf of Oman, India, Sri Lanka, Réunion, Mauritius,
Chagos, northwestern Australia, Papua New Guinea,
Philippines, Taiwan Province of China, and Japan.
2520 Bony Fishes
Epinephelus retouti Bleeker, 1868 (Plate V, 38)

Frequent synonyms / misidentifications: Epinephelus truncatus Katayama, 1957 / Epinephelus
fasciatus (non Forsskål, 1775); E. irroratus (non Forster, 1801).
FAO names: En - Redtipped grouper; Fr - Mérou à bout rouge; Sp - Mero punteado.
Diagnostic characters: Body depth 2.5 to 3.1 times in stand-

ard length; head length 2.4 to 2.6 times in standard length.
Interorbital flat to slightly convex, the dorsal head profile slightly
convex; preopercle rounded, finely serrate; upper edge of
operculum sinuous; snout length 3.5 to 3.9 times in head
length; maxilla reaches below rear half of eye; midlateral part juvenile
of lower jaw with 3 or 4 rows of teeth; a pair of short fixed canines
at front of both jaws. Nostrils subequal. First gill arch with 21 to 24 gill rakers, of which 6 to 8 on upper limb
and 15 to 17 on lower limb. Pyloric caeca numerous (more than 40). Dorsal fin with XI spines and 16 or
17 soft rays, the third to fifth spines longest, 2.4 to 3.1 times in head length; anal fin with III spines
and 8 soft rays; caudal fin truncate to slightly convex; pectoral fins fleshy, with 19 to 20 rays, the fin
length 1.5 to 1.8 times in head length; pelvic fins 1.8 to 2.2 times in head length. Lateral body scales rough,
with numerous auxiliary scales; lateral-line scales 64 to 76; lateral scale series 120 to 141. Colour:
adults dull yellowish orange to brownish red, each scale on dorsolateral part of body with a dark
greenish grey spot; usually 5 faint dark bars on body, the second and third bars extending onto
spinous dorsal where they are dark brown; dorsal fin greenish brown, the soft-rayed part darker
than rest of fin, the outer triangular part of interspinous membranes dark red, set off from rest of
fin by an orange-yellow stripe; dorsal margin of caudal fin dark greenish brown; dark red or brown
line along base of dorsal fin; orbit narrowly edged, except anteriorly, with dark red (pale in
preservative); pale blue line adjacent to red rim of orbit and completely surrounding eye. Juveniles
with dorsal part of first 3 dark bars on body black, the second and third bars extending into dorsal
fin; dorsal part of head also black, with 4 irregular transverse whitish bands across dorsal surface.
Habitat, biology, and fisheries: Adults are found on coral reefs and outer reef slopes in depths of 70 to
220 m; juveniles occur in 20 to 40 m. Apparently rare, but may be of importance to artisanal fisheries.
Caught with handlines and vertical longlines.
Distribution: Islands of tropi-
cal Indo-Pacific region: from
w es t e r n In d i an O c ea n to
Tuamotus, including Christ-
mas Island (eastern Indian
Ocean), Indonesia (Bali Sea
off Lombok), South China sea,
Taiwan Province of China, Ja-
pan (Okinawa, Izu Islands),
Palau, New Caledonia, Line
Islands, American Samoa,
and Society Islands.
Epinephelus rivulatus (Valenciennes, 1830) (Plate V, 39)

Frequent synonyms / misidentifications: Epinephelus rhyncholepis (Bleeker, 1852); E. grammatophorus
Boulenger, 1903; E. homosinensis Whitley, 1944; E. spiramen Whitley, 1945 / None.
FAO names: En - Halfmoon grouper; Fr - Mérou demi-lune; Sp - Mero medialuna.
in standard length. Preopercle angular, with enlarged serrae at angle; upper edge of operculum straight,
almost horizontal; ventral edge of maxilla with a slight step at distal expansion; midlateral part of lower jaw
with 2 or 3 rows of teeth. Nostrils subequal. First gill arch with 20 to 24 gill rakers, of which 6 to 8 on upper
limb and 14 to 16 on lower limb. Pyloric caeca 26. Dorsal fin with XI spines and 16 to 18 soft rays, the third
or fourth spines longest, 2.4 to 3.2 times in head length, the interspinous membranes distinctly incised;
anal fin with III spines and 8 soft rays; caudal fin rounded; pectoral fins fleshy, with 17 to 19 rays, the fin
length 1.5 to 1.9 times in head length; pelvic fins 1.9 to 2.4 times in head length. Lateral body scales rough,
with auxiliary scales; nape and front of body above lateral line with minute auxiliary scales and
numerous pores; lateral-line scales 48 to 53; lateral scale series 86 to 98. Colour: body generally
reddish to greenish brown, with a small white or pale blue spot (about size of nostrils) on each
scale; 4 irregular dark bars usually visible on body and extending onto base of dorsal fin, and a
fifth dark bar on peduncle, the third and fourth bars run from soft dorsal fin to anal fin and are
usually joined midlaterally; pectoral fins dusky, with a dark red or reddish brown semicircular blotch
on base of rays; 2 dark red or reddish brown bands on chest and a dark brown spot on front of
isthmus. Head mostly dark brown or pale reddish, with irregular pale blotches and pale blue or
violet vermiculations; 4 white or pale spots along lower part of head (2 on side of lower jaw and
upper lip, 1 just behind end of maxilla and last on interopercle); fins greenish yellow or greyish
brown; dorsal fin usually with dark brown to black streak along base (may be only along base of
spinous part); margin of interspinous dorsal-fin membranes yellow to reddish next to spine,
becoming transparent posteriorly, with a pale stripe below triangular part of fin. Some fish with white
dots on operculum like those on body.
Size: Maximum total length at least 45 cm; maximum weight at least 2 kg.
Habitat, biology, and fisheries: Rocky bottoms or coral reefs in depths of 10 to 150 m. Females mature
at 22 cm, males at 25 cm standard length. Feeds on small fishes and crabs. Caught with handlines,
longlines, gill nets, traps, and
spear.
Distribution: South Africa to
tropical western Pacific and
temperate waters of Australia
and New Zealand, including
Indonesia, Philippines, Tai-
wan Province of China, and
Japan.
2522 Bony Fishes
Epinephelus sexfasciatus (Valenciennes, 1828) (Plate V, 40)

Frequent synonyms / misidentifications: None / Epinephelus diacanthus (non Valenciennes, 1828).
FAO names: En - Sixbar grouper; Fr - Mérou six raies; Sp - Mero de seis bandas.
in standard length; caudal peduncle depth 2.6 to 3.4 times in head length. Interorbital area flat or slightly
convex, the dorsal head profile convex; preopercle with 2 to 4 greatly enlarged serrae at the angle;
upper edge of operculum straight; maxilla reaches to or slightly past vertical at rear edge of eye;
midlateral part of lower jaw with 2 rows of subequal teeth. Nostrils subequal. First gill arch with 20 to 23
gill rakers, of which 7 or 8 on upper limb and 13 to 15 on lower limb; longest gill raker shorter than
longest gill filaments. Pyloric caeca 7 or 8. Dorsal fin with XI spines and 14 to 16 soft rays, the third or
fourth spines longest, 2.3 to 2.7 times in head length and about equal to longest soft ray, the interspinous
membranes distinctly incised; anal fin with III spines and 8 soft rays; caudal fin rounded; pectoral fins not
fleshy, with 17 to 19 rays, the fin length 1.5 to 1.7 times in head length; pelvic fins 1.8 to 2.2 times in head
length. Lateral body scales rough, with a few auxiliary scales; lateral-line scales 46 to 51; lateral scale
series 82 to 96. Colour: head and body pale greyish brown; 5 dark brown bars on body and 1 on nape
(dark bars usually more or less divided vertically by a narrow pale bar); scattered pale spots may
be present on body, and some faint small brown spots are often visible on the edges of the dark
bars; soft dorsal, caudal, and pelvic fins dusky grey, the pectoral fins greyish or dusky orange-red;
jaws and ventral parts of head sometimes pale reddish brown.
Habitat, biology, and fisheries: Usually found on silty sand or muddy bottoms at depths of 10 to 80 m.
This preference for soft-bottom habitats may account for its restricted distribution and absence at oceanic
islands. Females are mature at 13 cm standard length. Feeds on small fishes and crustaceans. Although
the sixbar grouper is a small species, it is common and readily caught in trawls; consequently, it is often
seen in local markets.
Distribution: Tropical west-
ern Pacific Ocean: Indonesia
(Sumatra, Java, Bali, Lombok,
Celebes, Moluccas), Singa-
pore, Malaysia, Thailand, Viet
Nam, Philippines, Papua New
Guinea, Louisiade Archipel-
ago, and Australia (north
coast from Western Australia
to Queensland).
Epinephelus socialis (Günther, 1873)

Frequent synonyms / misidentifications: None / Epinephelus ongus (non Bloch, 1790).
FAO names: En - Surge grouper; Fr - Mérou houleux; Sp - Mero oleado.
15.5 cm standard length juvenile 3.4 cm standard length

in standard length. Interorbital area flat or slightly convex, the dorsal head profile convex; preopercle
rounded, finely serrate; upper edge of operculum sinuous; maxilla reaches past vertical at rear edge
of eye; midlateral part of lower jaw with 3 or 4 rows of teeth. Nostrils subequal. First gill arch with 25 to 28
gill rakers, of which 8 to 10 on upper limb and 16 to 20 on lower limb. Pyloric caeca about 20. Dorsal fin
with XI spines and 14 to 16 soft rays, the third or fourth spines longest, 2.6 to 3.6 times in head length, the
interspinous membranes moderately incised; anal fin with III spines and 8 soft rays; caudal fin rounded;
pectoral fins fleshy, with 18 or 19 rays, the fin length 1.6 to 1.9 times in head length; pelvic fins 1.8 to 2.2
times in head length. Lateral body scales of juveniles rough; lateral body scales of subadults not so rough
and with some auxiliary scales; body scales of adults smooth (mostly embedded) with numerous auxiliary
scales; lateral-line scales 64 to 70; lateral scale series 97 to 111. Colour: head and body whitish,
covered (except ventral part of head, chest, and abdomen) with small close-set blackish brown
spots, those on rear part of body often coalesced to form irregular longitudinal bands; the dark
spots become relatively smaller with growth, and on a fish of 21 cm they are black dots about the
size of the rear nostrils; 4 large blackish blotches usually visible on body at base of dorsal fin and
a fifth black saddle blotch on peduncle. Median fins colored like body basally, becoming dark
greyish brown distally, the soft dorsal, caudal and anal fins with a white margin, small white spots,
and usually a black submarginal band; pectoral fins dark greyish brown with a white margin
posteriorly and small whitish spots; pelvic fins dark greyish brown, with a white leading edge.
Habitat, biology, and fisheries: A shallow-water species of coral reefs; more common on atolls than high
islands, and usually in outer reef areas exposed to heavy surge. Juveniles and occasionally adults occur
in tidepools. Feeds on crustaceans, (mainly grapsid crabs), octopuses, and fishes. Caught with spears,
traps, and handlines.
Distribution: Central-West Pacific:
Marcus Island, Marshall Islands, Phoenix
Islands, American Samoa, Cook, Line and
Society islands, Rapa, Tuamotus, and the
Pitcairn Group. Widely distributed over
Pacific Plate, but not known at Hawaii,
Marquesas, Caroline Islands, or at any
Melanesian islands. Reported from 2
localities off the Pacific Plate: Ogasawara
(Bonin) Islands, and Mariana Islands.
2524 Bony Fishes
Epinephelus spilotoceps Schultz, 1953 (Plate VI, 41)

Frequent synonyms / misidentifications: Epinephelus salonotus Smith and Smith, 1963 / Epinephelus
hexagonatus (non Forster, 1801); E. macrospilos (non Bleeker, 1855); E. melanostigma non Schultz, 1953;
E. merra non Bloch, 1793; E. quoyanus (non Valenciennes, 1830).
FAO names: En - Foursaddle grouper; Fr - Mérou quatre selles; Sp - Mero cuatro monturas.
Diagnostic characters: Body depth 3.1 to 3.6 times in standard length; caudal-peduncle depth 3.7
to 4.3 times in head length. Preopercle rounded, with a shallow notch, below which the serrae are
enlarged but covered with skin; upper edge of operculum almost straight; midlateral part of lower jaw with
2 to 4 rows of teeth. Nostrils subequal. First gill arch with 6 to 9 gill rakers on upper limb, 16 to 18 on lower
limb. Dorsal fin with XI spines and 14 to 16 soft rays, the third or fourth spine longest, 2.8 to 3.6 times
in head length and distinctly shorter than longest dorsal-fin rays; anal fin with III spines and 8 soft
rays, the second spine usually longest, 2.4 to 3.7 times in head length and usually more than
caudal-peduncle depth; caudal fin rounded, middle caudal-fin rays 1.7 to 2.0 times in head length;
pectoral-fin rays 17 to 19, middle rays 21 to 24% of standard length, 1.7 to 2.1 times in head length;
pelvic fins 1.9 to 2.4 times in head length. Lateral body scales rough, with auxiliary scales; lateral-line
scales 59 to 69; lateral scale series 86 to 100. Colour: head, body, and median fins pale, mostly
covered with close-set dark brown, olive, or reddish brown polygonal spots, the pale interspaces
forming a white reticulum; spots on ventral part of head and body more rounded, more separated and
often more reddish; large blackish brown saddle-blotch at base of last 4 dorsal-fin spines; 2 similar
but smaller dark blotches at base of soft dorsal-fin rays and a third on rear end of peduncle; these
dark saddle-blotches uniformly pigmented (on juveniles) or comprising a group of extra dark spots
separated by the white reticulum (on adults); spots on head progressively smaller and darker
anteriorly, those on snout blackish brown and about size of nostrils, with 3 or 4 irregular rows along
front of upper lip; pectoral fins with close-set reddish brown spots, larger and more distinct towards base
of fin, distal part of fin dull yellowish green; tips of dorsal-fin spines blackish, with a short white filament.
Habitat, biology, and fisheries: A shallow-water coral-reef species. No information on biology or fishery
statistics available for this species.
Distribution: Most (probably
all) islands of tropical Indo-
West Pacific region, from
Mozambique to the Central
Pacific (Line Islands). Primar-
ily an insular species; except
for a single specimen from
Hong Kong, there are no re-
cords from the Asian main-
land, nor from the Red Sea,
Persian Gulf, Sri Lanka, Phil-
ippines, Taiwan Province of
China, Japan, or Australia
(though it is found at Rowley
Shoals off Western Australia).
Epinephelus stictus Randall and Allen, 1987 (Plate VI, 42)

Frequent synonyms / misidentifications: None / Epinephelus diacanthus (non Valenciennes, 1828).
FAO names: En - Blackdotted grouper; Fr - Mérou pointe noirs; Sp - Mero punteado negro.
in standard length; caudal-peduncle depth 3.8 to 4.2 times in head length. Preopercle with 1 to 4
distinctly enlarged serrae at the angle; upper edge of operculum straight; maxilla reaches to or slightly
past vertical at rear edge of eye; midlateral part of lower jaw with 2 rows of teeth. Nostrils subequal. First
gill arch with 7 or 8 gill rakers on upper limb, 14 to 16 on lower limb. Pyloric caeca 12. Dorsal fin with XI
spines and 15 or 16 soft rays, the third or fourth spines longest, 2.7 to 3.4 times in head length and shorter
than longest soft rays, the interspinous membranes incised; anal fin with III spines and 8 soft rays; caudal
fin rounded; pectoral fins not fleshy, with 18 to 20 rays, the fin length 1.7 to 1.9 times in head length; pelvic
fins 2.1 to 2.6 times in head length. Lateral body scales rough, without auxiliary scales; lateral-line scales
48 to 51; lateral scale series 84 to 96. Colour: body and head yellowish brown dorsally; belly and rear
part of body whitish ventrally; chest and ventral parts of head pale reddish orange; body with 5 faint
oblique dark bars (may be lost in preservative); midlateral part of each bar may be darker,
representing a series of squarish dark blotches continued from the dark band running from eye to
end of operculum; numerous dark brown to black dots on dorsolateral parts of head and body,
sometimes concentrated within dark bars and bands. Fins pale, the rays darker than membranes; soft
dorsal and caudal fins with narrow dark brown margin; pelvic and anal fins often darker than other fins;
anal fin may also have a narrow dark margin; dark line along base of dorsal fin (indistinct anteriorly).
Habitat, biology, and fisheries: Mud or sand bottom in depths of 60 to 142 m. Although one of the most
common species of grouper caught by trawlers in the vicinity of Hong Kong in the 1960’s, the black-dotted
grouper was not of much commercial importance. The low price in the markets is a result of the small size
and poor edible quality of this species.
Distribution: Southern Ja-
pan, Hong Kong, Hainan Is- ?
land, Viet Nam, “South China
Sea”, and northwest Austra-
lia. The Japanese records as
“Epinephelus diacanthus“
may be based on the single
specimen reported from Ko-
chi City in 1954. The 1960 re-
cord from Taiwan Province of
China is dubious, as this spe-
cies has not been found there
in recent surveys of Taiwan-
ese serranids.
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2526 Bony Fishes
Epinephelus tauvina (Forsskål, 1775) (Plate VI, 43)

Frequent synonyms / misidentifications: Epinephelus elongatus Schultz, 1953; E. chewa Morgans,
1966 / Epinephelus coioides (non Hamilton, 1822); E. malabaricus (non Bloch and Schneider, 1801);
E. lanceolatus (non Bloch, 1790).
FAO names: En - Greasy grouper; Fr - Mérou loutre; Sp - Mero lutra.
Diagnostic characters: Body elongate, its depth 3.0 to 3.6 times in standard length; head large, its
length 2.1 to 2.4 times in standard length. Interorbital area narrow, flat to slightly concave, its width
6.8 to 8.1 times in head length; serrae at corner of preopercle slightly enlarged; upper edge of operculum
almost straight; maxilla reaching well past eye, the greatest width about twice suborbital depth (least
distance from eye to maxilla) and 5.4 to 6.5% of standard length; midlateral part of lower jaw with 2 to
5 rows of teeth. Rear nostrils distinctly larger than anterior nostrils. First gill arch with 8 to 10 gill rakers on
upper limb, 17 to 20 on lower limb; no bony platelets on gill arch. Pyloric caeca 16 to 18. Dorsal fin with
XI spines and 13 to 16 soft rays, the third to fifth spines longest, 3.1 to 4.7 times in head length and shorter
than longest dorsal-fin rays; anal fin with III spines and 8 soft rays; caudal fin rounded; pectoral-fin rays
18 or 19. Midlateral body scales rough in juveniles, smooth in adults, except for small patch covered by
pectoral fin; lateral-line scales 63 to 74; lateral scale series 95 to 112. Colour: head and body pale
greenish grey or brown, covered with dull orange-red to dark brown spots; spots on head
progressively smaller anteriorly; large black blotch (or group of black spots) often visible at base
of last 4 dorsal-fin spines and extending onto lower part of fin; 5 faint subvertical dark bars may be
present on body, 4 below dorsal fin and fifth on caudal peduncle (these bars may be represented by dusky
blotches at base of dorsal fin and a dark saddle-blotch on caudal peduncle); fins also covered with dark
spots, those on pectoral fins becoming smaller and less distinct distally; rear margin of caudal, anal, and
pectoral fins often with white edge; dark spots on median fins of juveniles are so close set that the pale
interspaces form a white reticulum.
Size: Maximum total length 75 cm; reports of larger sizes are probably based on misidentifications of
Epinephelus coioides, E. malabaricus, or E. lanceolatus.
Habitat, biology, and fisheries: Clear water areas on coral reefs; juveniles on reef flats and in tidepools,
but adults occur down to 50 m. Adults primarily piscivorous. Important to artisanal fisheries. Caught with
hook-and-line, spear, and traps. Sold fresh in local markets. Also important in aquaculture. From 1990 to
1995, the reported yearly aquaculture production of E. tauvina in Malaysia ranged from 144 to 1 006 t (FAO
Aquaculture Production Statistics).
Distribution: From Red Sea
to South Africa and eastwards
to Pitcairn Group; in western
Pacific, from Japan to New
South Wales and Lord Howe
Island. More common at is-
lands than along continental
shores. No verifiable records
from Persian Gulf, Asia main-
land, the Philippines, Indone-
s i a, n orthern or western
Australia.
Epinephelus tukula Morgans, 1959 (Plate VI, 45)

Frequent synonyms / misidentifications: Serranus dispar var. A Playfair, 1867 / None.
FAO names: En - Potato grouper; Fr - Mérou patate; Sp - Mero patata.
Diagnostic characters: Body depth 2.9 to 3.5 times in standard length; head length 2.3 to 2.6 times in
standard length. Interorbital area slightly convex; the dorsal head profile straight; preopercle rounded or
subangular, the serrae at corner slightly enlarged; upper edge of operculum almost straight; maxilla
reaches to or beyond vertical at rear edge of eye; midlateral part of lower jaw with 2 to 6 rows of teeth.
Nostrils subequal. First gill arch with 8 to 10 gill rakers on upper limb, 15 to 18 on lower limb. Dorsal fin
with XI spines and 14 or 15 soft rays, the third or fourth spines longest, 3.1 to 3.8 times in head length and
distinctly shorter than longest dorsal-fin rays; interspinous dorsal-fin membranes distinctly incised; anal fin
with III spines and 8 soft rays; caudal fin rounded; pectoral-fin rays 18 to 20, the fin length 1.6 to 2.1 times
in head length; pelvic fins usually not reaching anus, 1.9 to 2.4 times in head length. Lateral body scales
rough, with auxiliary scales in adults; lateral-line scales 62 to 70; lateral scale series 113 to 130. Colour:
body pale brownish grey with several dark brown to black widely-spaced blotches, mostly larger
than eye and varying in shape from round to oval or dumbbell-shaped; head with smaller dark brown
spots and streaks (many radiating from eye, especially posteriorly); dark spots on fins, smaller distally.
Large adults may be nearly black.
Size: One of the largest groupers; maximum total length 200 cm, weight to 90 kg.
Habitat, biology, and fisheries: Coral-reefs; juveniles in tidepools, and adults occur in depths of 10 to
150 m. Maturity occurs at 90 cm standard length. Feeds on a wide variety of reef fishes, skates, crabs,
and spiny lobsters. Epinephelus tukula is “exceedingly territorial, and is very aggressive towards unwel-
come intruders.” The large size and territorial behaviour of E. tukula make it especially vulnerable to
spearfishermen, hence spearfishing for this species is illegal in South African waters. Although it is not
common, it is an important species in the sportfishery of South Africa and Australia. Caught with
hook-and-line and spear.
Distribution: From the west-
ern Indian Ocean and Red
Sea to the western Pacific, in-
cluding Japan (Okinawa and
Honshu), Taiwan Province of
C h i n a , S o u t h C h in a S e a
(Paracel Islands), Western
Australia, and Queensland. It
is surprising that there are no
records of this distinctive spe-
cies from Indonesia, the Phil-
ippines, or New Guinea.
2528 Bony Fishes
Epinephelus undulatostriatus (Peters, 1867) (Plate VI, 46)

Frequent synonyms / misidentifications: Serranus guttulatus Macleay, 1879 / None.
FAO names: En - Maori grouper; Fr - Mérou Maori; Sp - Mero Maori.
in standard length. Interorbital area convex, the dorsal head profile almost straight; preopercle subangular,
the serrae at corner slightly enlarged; upper edge of operculum almost straight; maxilla reaches to or
slightly past vertical at rear edge of eye; midlateral part of lower jaw with 2 rows of teeth. Nostrils subequal.
First gill arch with 9 or 10 gill rakers on upper limb, 15 or 16 on lower limb. Pyloric caeca 11. Dorsal fin
with XI spines and 15 to 17 soft rays, the third or fourth spine longest, 2.2 to 2.8 times in head length and
longer than longest dorsal-fin rays; interspinous dorsal-fin membranes distinctly incised; anal fin with III
spines 8 soft rays; caudal fin rounded; pectoral fins fleshy, with 17 to 19 rays, the fin length 1.6 to
2.0 times in head length; pelvic fins not reaching anus (except in small juveniles), 1.8 to 2.0 times in head
length. Lateral body scales rough, with auxiliary scales; lateral-line scales 48 to 55; lateral scale series 98
to 108. Colour: body buff, covered with close-set wavy longitudinal brownish orange, golden brown,
or reddish brown stripes; some stripes broken into series of small spots; head pale greyish brown,
covered with small spots like those on body; maxillary streak orange brown; fin rays pale, with
small brown spots, the interradial membranes dark brown; median fins with narrow yellow margin;
pectoral fins diffusely yellow distally.
Habitat, biology, and fisheries: Coral reefs and rocky areas in depths of 27 to 80 m. An active swimmer
that will leave the bottom to take a floating bait. Feeds on fishes. An esteemed food fish and important
species in the hook-and-line
fishery of Australia’s east
coast. Caught with hook-and-
line and spear.
Distribution: E n d e m i c t o
Australia, from southern
Queensland (most northern
record, One Tree Island,
Great Barrier Reef at 23°30’S)
to Bateman’s Bay (35°44’S)
New South Wales. One speci-
men was reported from Kan-
garoo Island, South Australia.
Epinephelus undulosus (Quoy and Gaimard, 1824)

Frequent synonyms / misidentifications: Serranus lineatus Valenciennes, 1828, S. amboinensis
FAO names: En - Wavylined grouper; Fr - Mérou ondulé; Sp - Mero ondulado.
standard length. Interorbital area convex; preopercle angular, with a notch above the corner and serrae at
the corner enlarged; upper edge of operculum straight or slightly concave; maxilla reaches to vertical at
rear edge of eye; adults with a prominent knob or hook-like process on ventral edge of maxilla
(covered by upper lip); midlateral part of lower jaw with 2 rows of teeth. Nostrils subequal or the posteriors
slightly larger. Gill rakers longer and more numerous than other species of Epinephelus; first gill
arch with 32 to 38 rakers, of which 12 to 16 on upper limb and 20 to 23 on lower limb; 1 or 2
rudimentary rakers in juveniles, but none in adults. Dorsal fin with XI spines and 17 to 19 soft rays,
the third or fourth spines longest, 2.5 to 3.4 times in head length and longer than longest dorsal-fin
ray; interspinous membranes of dorsal fin not incised; anal fin with III spines and 8 soft rays; caudal
fin truncate to emarginate; pectoral-fin rays 18 or 19, the fins shorter than pelvic fins, 1.7 to 2.3 times in
head length; pelvic fins not reaching anus (except for juveniles), 1.6 to 2.2 times in head length. Lateral
body scales distinctly rough, with numerous auxiliary scales; lateral-line scales 63 to 76; lateral scale series
124 to 150. Colour: head, body, and fins purplish grey to brownish grey, with brown to golden-brown
dots on head and wavy longitudinal lines of the same colour on dorsal part of body (lines faint or
absent on large specimens); margin of spinous dorsal fin narrowly blackish.
Size: Maximum total length at least 73 cm (reported to attain 122 cm); maximum weight at least 7 kg.
Habitat, biology, and fisheries: Usually found on offshore banks (rather than coral reefs) at depths of 24
to 90 m. Feeds on a variety of small fishes, small crustaceans (especially stomatopods), shrimps, and
pelagic tunicates (Thalia and
Pyrosoma sp.). Females were
estimated to be mature at
55 cm total length. Caught with
hook-and-line, vertical long
lines, and trawls.
Distribution: Northern Indian
Ocean and western Central
Pacific: Indonesia, Sarawak,
New Guinea, Solomon Is-
lands, and the Philippines. Not
known from the Red Sea, Per-
sian Gulf, or Australia.
2530 Bony Fishes
Gracila albomarginata (Fowler and Bean, 1930) (Plate VI, 47)

FAO names: En - Masked grouper; Fr - Mérou bord rouge; Sp - Mero paranjero.
Diagnostic characters: Body oblong, somewhat compressed, its depth 2.6 to 3.3 times in standard length,
body width 1.8 to 2.3 times in body depth; head smaller than most other groupers, 2.9 to 3.2 times in
standard length; dorsal head profile evenly convex; interorbital area slightly convex and wide, its width
greater than eye diameter; preorbital narrow, its depth distinctly less than eye diameter, 10 to 15 times in
head length; preopercle rounded, finely serrate, the lower edge smooth and fleshy; ventral edge of
subopercle and interopercle smooth; upper edge of operculum distinctly convex; nostrils subequal. A pair
of small canines at front of both jaws; palatine teeth present; distal part of maxilla in adults with a
hook-shaped or step-like expansion on ventral edge; supramaxilla well developed. First gill arch with
8 to 10 gill rakers on upper limb, 14 to 16 on lower limb. Dorsal fin with IX slender spines and 14 to 16
soft rays, the fin origin over rear end of operculum; anal fin with III spines and 9 or 10 soft rays;
caudal fin truncate to slightly emarginate, with 8 branched rays in upper lobe and 7 in lower lobe;
soft dorsal and anal fins rounded; dorsal-fin membranes not (adults) or slightly (juveniles) incised
between the spines; pectoral fins rounded, with 18 or 19 rays, the middle rays longest. Midlateral body
scales rough. Colour: adults greenish or reddish brown to grey, with several narrow curved dark
bars on flanks and dark brown spot on side of caudal peduncle; 3 to 5 blue lines (often broken into
spots) across the head (the upper 2 lines may serve as margins for a mask-like dark band from
snout to end of opercle, enclosing eye); region under maxilla and along edge of gill opening often orange;
soft dorsal and anal fins with narrow blue margin. A transient colour phase has the caudal peduncle
white, with a prominent black spot, and a large white area flanked by black areas on dorsal part of
body. Juveniles brown or violet, with a bright reddish orange stripe in dorsal and anal fins and along
upper and lower edges of caudal peduncle and fin.
Habitat, biology, and fisheries: Usually found on outer reef slope at depths of 15 to 120 m. An active
swimmer that roams over the reef during the day. An excellent food fish, but not common enough to be of
commercial importance. Caught with hook-and-line, gill nets, and spear.
West Paciific from northern
Mozambique to French Poly-
nesia, including Taiwan Prov-
ince of China, Philippines,
Indonesia, New Guinea, New
Britain, New Caledonia, Soci-
ety Islands, Phoenix Islands,
Marshall Islands, Minami Tori
Shima, Guam, Caroline Is-
lands, Palau Islands, Loyalty
Islands, Samoa Islands,
Takaroa, Tuamotus, and Fan-
ning Island.
Plectropomus areolatus Rüppell, 1830 (Plate VI, 48)

Frequent synonyms / misidentifications: Plectropoma maculatum ? (non Bloch, 1790); Plectropomus
truncatus Fowler and Bean, 1930 / None.
FAO names: En - Squaretail coralgrouper; Fr - Mérou queue carrée; Sp - Mero troncon.
Diagnostic characters: Body elongate, robust, its depth 2.9 to 3.9 times in standard length; head length
2.7 to 3.1 times in standard length; snout 2.8 to 3.6 times in head length; suborbital depth 5.6 to 10 times
in head length. Interorbital area flat (rounded at edges of orbits), with small, embedded scales; preopercle
broadly rounded, with 3 large, ventrally-directed spines along lower half; interopercle and subopercle
smooth; opercle with 3 flat spines, the upper and lower spines covered by skin. Midlateral part of lower jaw
with 1 to 4 enlarged fixed canines. Nostrils subequal, set in a shallow groove running forward from eye.
First gill arch with 2 to 7 developed gill rakers on lower limb; gill raker at angle shorter than gill filaments
at angle. Dorsal fin with VII or VIII slender spines and 10 to 12 soft rays, the third or fourth spines longest,
3.2 to 4.2 times in head length, the longest soft ray 2.3 to 2.6 times in head length; base of spinous part
of dorsal fin subequal to that of soft-rayed part; anal fin with III slender spines and 8 soft rays, the first 1
or 2 spines embedded and difficult to see in large fish; caudal fin truncate to slightly emarginate, the
caudal concavity more than 13 times in head length; pectoral-fin rays 15 or 16; pectoral fins subequal
to pelvic fins, 2.0 to 2.4 times in head length. Lateral-line scales 83 to 97. Colour: head, body, and median
fins greenish grey to brown or brownish red, with numerous round to oval dark-edged blue spots
(the largest about equal to pupil); most spots within a spot diameter of adjacent spots; pelvic fins
with dark brown to blackish membranes; rear margin of caudal fin with a white edge and often with a
blackish submarginal band.
Habitat, biology, and fisheries: Coral reefs in lagoons and on outer reef at depths of 2 to 20 m. Feeds
on fishes. A wary grouper, difficult for a diver to approach. It is reported that for a few days before new
moon in May, large numbers gather in the seaward end of Ulong Channel, Palau to spawn and males may
display pale bodies with about 5 irregular dark saddles and dark dorsal and anal fins. Undoubtedly of
interest to artisanal fisheries. Caught with hook-and-line.
Distribution: Indo-Pacific;
except for the Red Sea and
Australia, records are limited
to insular localities: Chagos,
Maldives, Cocos-Keeling,
Rowley Shoals (Western Aus-
tralia), Indonesia, Philippines,
Ryukyus, Paracel Islands
(South China Sea), Palau,
Great Barrier Reef, Caroline,
Marshall, Samoa, and Phoe-
nix islands.
2532 Bony Fishes
Plectropomus laevis (Lacepède, 1801) (Plate VII, 49)

Frequent synonyms / misidentifications: Plectropomus melanoleucus (Lacepède, 1802) / Plectropomus
maculatus (non Bloch, 1790).
FAO names: En - Blacksaddled coralgrouper; Fr - Mérou sellé; Sp - Mero ensillado.
59.7 cm standard length 47.5 cm standard length

Diagnostic characters: Body elongate, robust, its depth
2.9 to 3.9 times in standard length; head length 2.7 to 3.1
times in standard length; suborbital depth 5.6 to 10 times
in head length. Interorbital area without scales; preopercle
broadly rounded, with 3 large, ventrally-directed spines
along lower half; interopercle and subopercle smooth;
opercle with 3 flat spines, the upper and lower spines juvenile 11 cm standard length
covered by skin. Midlateral part of lower jaw with 1 to 4
enlarged fixed canines. Nostrils subequal, set in a shallow groove running forward from eye. First gill arch
with 4 to 10 developed gill rakers on lower limb; gill raker at angle shorter than gill filaments at angle. Dorsal
fin with VII or VIII slender spines and 10 to 12 soft rays, the third or fourth spines longest, 3.2 to 4.2 times
in head length, the longest soft ray 2.3 to 3.3 times in head length; anal fin with III slender spines and 8
soft rays, the first 1 or 2 spines embedded and difficult see in large fish; caudal fin emarginate, the caudal
concavity 5 to 10 times in head length; pectoral-fin rays 16 to 18, the fin length about equal to pelvic
fins, 2.1 to 2.4 times in head length. Lateral-line scales 83 to 97. Colour: 2 colour forms: the pale
black-saddled form is whitish or pale yellowish with 5 dark brown to black, saddle-like or wedge-
shaped bars or blotches on dorsal part of head and body, the first just behind eyes, the second on
nape, the third to fifth under dorsal fin; small dark-edged blue spots may be visible on body (mainly
posteriorly); fins, caudal peduncle, snout, and jaws yellow; black blotch centrally at base of paired
fins; small juveniles (10 to 20 cm) with membranes between first and fifth dorsal-fin spines mostly
blackish. The dark form is brown, olivaceous, red, or nearly black, the dark bars (as on pale form)
are usually faint or absent; numerous, round, dark-edged blue spots (about 1/2 pupil diameter) on
head, body (except ventrally), soft dorsal, caudal, and anal fins, and base of pectoral fins; pectoral
fins dark brown distally, the rays darker than membranes, the rear edge white.
Size: Maximum total length about 125 cm; maximum weight 18 kg.
Habitat, biology, and fisheries: Coral reefs of Indo-West Pacific region in depths of 4 to 90 m. The pale
black-saddled juveniles may be mimics of the pufferfish Canthigaster valentini. Juveniles of both forms
[pale and dark] less than about 20 cm normally employ the pectoral fins for propulsion, holding the caudal
fin distinctly folded and the front section of the spinous dorsal erect. When swimming in this manner,
juveniles of Plectropomus laevis look remarkably similar to Canthigaster valentini. The pale black-saddled
form of P. laevis is usually smaller (8 to 57 cm standard length) than the dark reddish brown phase (15 to
100 cm standard length), and the smaller pale phase may be juveniles and females, with the larger dark
phase being males. This hypothesis would agree with a protogynous type of hermaphroditism which seems
to be the usual system for groupers and is known for P. leopardus. Maturity (at least for females) “seems
to occur” at 50 to 52 cm standard length (2.8 to 3.2 kg). The blacksaddled coralgrouper feeds on a variety
of large reef fishes, including groupers. This diet of large fishes is responsible for the high concentrations
of ciguatera toxins found in some specimens of this species. Despite the danger of ciguatera poisoning,
P. laevis is commonly used for food. Caught with hook-and-line, spear, and in fish traps.
Distribution: Indo-Pacific, from southern Mozambique
to the Tuamotus, Austral Islands, and Rapa, but not in
the Red Sea or Persian Gulf. Except for the east coasts
of Africa and Australia, records of P. laevis are from
insular localities, from Japan (Ryukyu Islands) to south-
ern Great Barrier Reef and most islands in the central
and western Pacific. Not recorded from the Asian coast
or Indonesia.
Plectropomus leopardus (Lacepède, 1802) (Plate VII, 50)

Frequent synonyms / misidentifications: Plectropoma leopardinus Cuvier, 1828 / Plectropoma
maculatus (non Bloch, 1790).
FAO names: En - Leopard coralgrouper; Fr - Saumonée léopard; Sp - Mero celestrial.
standard length; suborbital depth 5.6 to 10 times in head length. Interorbital area without scales; preopercle
broadly rounded, with 3 large, ventrally-directed spines along lower half; interopercle and subopercle
smooth; opercle with 3 flat spines, the upper and lower spines covered by skin. Midlateral part of lower jaw
with 1 to 4 enlarged fixed canines. Nostrils subequal for specimens less than 50 cm (at larger sizes, the
rear nostrils may be enlarged). First gill arch with 6 to 10 developed gill rakers on lower limb; gill raker at
angle longer than gill filaments at angle. Dorsal fin with VII or VIII slender spines and 10 to 12 soft rays,
the third or fourth spines longest, 3.2 to 4.2 times in head length, the longest soft ray 2.2 to 2.7 times in
head length; anal fin with III slender spines and 8 soft rays, the first 1 or 2 spines embedded in large fish;
caudal fin emarginate, the caudal concavity 5 to 12 times in head length and the fin length 1.3 to
1.5 times in head length; pectoral-fin rays 15 to 17, the fin length about equal to pelvic-fin length,
1.9 to 2.2 times in head length. Lateral-line scales 89 to 99. Colour: olivaceous to reddish brown,
orange-red, or red, with numerous, small blue spots (nostril sized and usually dark-edged) on head
and body (except ventrally) and median fins; more than 10 spots on cheek (below and behind eye
to rear edge of preopercle); blue ring (dark brown in alcohol) on edge of orbit (sometimes broken
into segments); pectoral fins reddish or hyaline with darker rays; an indistinct dark band at rear margin
of caudal fin and a white line usually visible along middle of rear edge.
Habitat, biology, and fisheries: Coral reefs at depths of 3 to 100 m. The leopard coralgrouper is a
protogynous hermaphrodite. On the Great Barrier Reef, the smallest mature female (21 cm standard
length) was estimated to be 2 years old, and the largest female (47 cm standard length) was age 4; the
smallest mature male (30 cm standard length) was age 3, and the largest male (54 cm standard length)
was estimated to be 5 years old. Spawning occurred in late November and early December. Feeds
throughout the day and is inactive at night; 96% of the prey comprised fishes. Small juveniles included a
few benthic crustaceans in their diet, but adults fed only on fishes, primarily atherinid and scarid fishes. A
popular food fish that rarely causes ciguatera. Caught with hook-and-line, spear, traps, and trawls.
Distribution: Western Pa-
cific, from southern Japan to
Australia (Queensland and
Western Australia) and east-
wards to the Caroline Islands
and Fiji, including Hong Kong,
Viet Nam, Philippines, Indo-
nesia, Palau, Papua New
Guinea, and New Caledonia.
2534 Bony Fishes
Plectropomus maculatus (Bloch, 1790) (Plate VII, 51)

FAO names: En - Spotted coralgrouper; Fr - Vielle Saint-Silac; Sp - Mero con pintas.
adult
2.7 to 3.1 times in standard length; suborbital depth 5.6 to 10 times in head length. Interorbital area without
scales; preopercle broadly rounded, with 3 large, ventrally-directed spines along lower half; interopercle
and subopercle smooth; opercle with 3 flat spines, the upper and lower spines covered by skin; midlateral
part of lower jaw with 1 to 4 enlarged fixed canines. Nostrils subequal, set in a shallow groove running
forward from eye. First gill arch with 6 to 9 developed gill rakers on lower limb; gill raker at angle distinctly
longer than gill filaments at angle. Dorsal fin with VII or VIII slender spines and 10 to 12 soft rays, the third
or fourth spines longest, 3.2 to 4.2 times in head length, the longest soft ray 2.2 to 2.5 times in head length;
base of spinous part of dorsal fin subequal to that of soft-rayed part; anal fin with III slender spines and 8
soft rays, the first 1 or 2 spines embedded in large fish; caudal fin emarginate, the caudal concavity 5
to 10 times and the fin length 1.3 to 1.5 times in head length; pectoral-fin rays 15 to 17; the fin length
about equal to pelvic-fin length, 1.7 to 2.2 times in head length. Lateral-line scales 83 to 97. Colour:
head, body, and median fins greenish grey, brown, red, or orange-red, with dark-edged blue spots;
spots on head and anterior part of body about 1/2 pupil diameter or larger (and many are oval or
horizontally elongated), becoming much smaller, more uniformly round, and more numerous on
rear part of body and on fins; few spots on head, 3 to 7 on cheeks (below and behind eye to rear
edge of preopercle); no blue spots on ventral parts of head and body; usually a single blue spot on
pectoral-fin base and another in axil; no blue spots on pelvic fins; rear margin of caudal fin with a white
edge.
Size: Maximum total length about 125 cm; maximum weight at least 25 kg.
Habitat, biology, and fisheries: Coral reefs at depths of 5 to 50 m. Common in markets. Caught with
hook-and-line, spear, and in trawls.
Distribution: K no wn only
from the western tropical Pa-
cific: Thailand, Singapore,
Philippines, Indonesia, Papua
New Guinea, Solomon Is-
lands, and Australia (Western
Australia to southern Queens-
land).
Plectropomus oligacanthus Bleeker, 1854 (Plate VII, 52)

Frequent synonyms / misidentifications: Plectropoma variegatum Castelnau, 1875 / None.
FAO names: En - Highfin coralgrouper; Fr - Mérou-loche cacatois; Sp - Mero vela.
2.7 to 3.1 times in standard length; suborbital depth 3.3 to 7.2 times in head length. Interorbital area without
scales; preopercle broadly rounded, with 3 large ventrally-directed spines along lower half; interopercle
and subopercle smooth; opercle with 3 flat spines, the upper and lower spines covered by skin. Midlateral
part of lower jaw with 1 to 4 enlarged fixed canines. First gill arch with 7 to 9 developed gill rakers on lower
limb, gill raker at angle subequal to gill filaments at angle. Dorsal fin with VII or VIII slender spines and
10 to 12 soft rays, the third or fourth spines longest, 3.2 to 4.2 times in head length; soft dorsal and anal
fins pointed anteriorly, the second to fourth rays elongated, 1.5 to 2.1 times in head length; anal fin
with III slender spines and 8 soft rays, the first 1 or 2 spines embedded and difficult see in large fish; caudal
fin emarginate, the caudal concavity 3 to 7 times in head length; pectoral-fin rays 14 to 16; the fin
shorter than pelvic fins, 2.1 to 2.4 times in head length. Lateral-line scales 86 to 96. Colour: head,
body, and fins reddish brown to red, with horizontal to oblique blue lines on rear of head and
anterodorsally on body; vertical blue lines (or dashes) anteriorly on side of body (at least in adults),
and numerous blue spots elsewhere on body and on caudal fin; dorsal and anal fins with blue lines
and spots; pectoral fins pale yellowish, with basal two-thirds of rays dark brown and a few blue
lines at base of fin; pelvic fins with brownish red rays and blue membranes.
Habitat, biology, and fisheries: Coral reefs in depths of 4 to 40 m. Very little has been published on the
biology of this rare species. Sometimes groups of 2 or 3 often forage together for rock- and sand-living
crustaceans and fishes. Well known in the Philippines, but it seems to be rare elsewhere. Of minor
importance in artisanal fisheries, and it has been implicated in cases of ciguatera fish poisonings. Caught
with hook-and-line, spear,
and in traps.
from western Pacific Ocean:
Philippines, Indonesia, New
Guinea, northeastern Austra-
lia (Cape York to northern
Great Barrier Reef), Palau,
Truk, Caroline Islands, Mar-
shall Islands, and the Solo-
mon Islands.
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2536 Bony Fishes
Plectropomus pessuliferus Fowler, 1904 (Plate VII, 53)

Frequent synonyms / misidentifications: Perca miniata var. C Forsskål, 1775; Plectropoma maculatum
var. D Playfair and Günther, 1867, P. maculatum var. A (in part) Boulenger, 1895 / Plectropoma maculatus
(non Bloch, 1790).
FAO names: En - Roving coralgrouper; Fr - Mérou-loche vagabonde; Sp - Mero errante.
2.7 to 3.1 times in standard length; suborbital depth 5.6 to 10 times in head length. Interorbital area without
scales; preopercle broadly rounded, with 3 large, ventrally-directed spines along lower half; interopercle
and subopercle smooth; opercle with 3 flat spines, the upper and lower spines covered by skin. Midlateral
part of lower jaw with 1 to 4 enlarged fixed canines. Nostrils set in a shallow groove running forward from
eye, nostrils subequal at less than about 30 cm standard length, the rear nostrils often enlarged in larger
fish. First gill arch with 7 to 10 developed gill rakers on lower limb; gill raker at angle shorter than gill
filaments at angle. Pyloric caeca 3, large and finger-like. Dorsal fin with VII or VIII slender spines and 10
to 12 soft rays, the third or fourth spines longest, 3.2 to 4.2 times in head length, the longest soft ray 2.3
to 3.0 times in head length; anal fin with III slender spines and 8 soft rays, the first 1 or 2 spines embedded
and difficult see in large fish; caudal fin emarginate (truncate in juveniles), the caudal concavity 4.6
to 6.2 times in head length; pectoral-fin rays 15 or 16, the fin about equal to pelvic fins, 1.9 to 2.3
times in head length. Lateral-line scales 85 to 104. Colour: head, body, and fins brown to orange-red,
with numerous small dark-edged blue spots; some spots on head and sides of body of adults are
elongated (those on body usually vertically elongate); spots few or absent on ventral part of body;
edge of orbit often blue (may be broken into segments).
Size: Maximum total length uncertain, at least 63 cm, perhaps 120 cm.
Habitat, biology, and fisheries: Coral reefs at depths of 25 to 147 m. Caught with hook-and-line, spear,
and in trawls.
Distribution: Red Sea,
Zanzibar, Maldives, St. Bran-
don’s Shoals, Sri Lanka,
Chagos, Andaman Islands,
Nazareth Bank, Sumatra, and
Fiji.
Saloptia powelli Smith, 1964 (Plate VII, 54)

FAO names: En - Golden grouper; Fr - Mérou d’or; Sp - Mero dorado.
Diagnostic characters: Body oblong, robust, its depth less than head length and 2.6 to 3.0 times in
standard length; body width 2.2 times in body depth; head length 2.5 to 2.6 times in standard length; dorsal
head profile slightly convex; interorbital area flat; suborbital depth 0.4 to 0.5 times eye diameter and 13
times in head length. Preopercle subangular, with 3 large curved spines (mostly hidden by skin) on
lower edge, the rear edge with minute serrae near the angle. Opercle with 3 flat, equidistant spines; upper
edge of operculum distinctly convex; subopercle and interopercle serrate. A pair of canines at front of both
jaws; lower jaw with 2 rows of teeth, but no enlarged canines at midside of jaw; vomer and palatines with
teeth; posterior part of maxilla evenly expanded (no step, hooks, or knob on ventral edge); supramaxilla
well developed. First gill arch with 8 or 9 on upper limb, 16 or 17 on lower limb. Dorsal fin with VIII spines
and 11 soft rays, the fin origin behind vertical at rear end of operculum, the fin membranes slightly incised
between the spines, the third spine longest; length of dorsal-fin base less than 50% of standard length;
anal fin with III spines and 8 soft rays; dorsal- and anal-fin spines strong, the anal-fin spines quite distinct;
caudal fin emarginate, with 7 branched rays in upper lobe and 6 in lower lobe; pectoral fins short and
rounded, the middle rays longest, subequal to pelvic fins, 2.2 to 2.4 times in head length. Body scales
small, distinctly rough, even on belly; no auxiliary scales; lateral-line scales 70 to 78; lateral scale series
115 to 133. A single curved supraneural bone anterior to tip of first neural spine; dorsal fin with 2
trisegmental pterygiophores, anal fin with 3; rear edge of first dorsal pterygiophore incised at tip of third
neural spine; epipleural ribs on vertebrae 1 to 9. Colour: head, body, and fins yellow to orange-yellow,
shading ventrally to white or pink, the snout, lips, and dorsal part of head suffused with red;
dorsal-fin spines of some specimens streaked with red.
Habitat, biology, and fisheries: Rocky reefs at depths of 140 to 367 m. Biology unknown. Although it is
not abundant, the golden grouper is an important food fish in the Ryukyu Islands of southern Japan and
also in the Mariana Islands.
to French Polynesia, includ-
ing Okinawa, Taiwan Prov-
ince of China, South China
Sea, Mariana Islands, Society
Islands, Cook Islands, Ameri-
can Samoa, Fiji, and Tuamo-
tus.
2538 Bony Fishes
Variola albimarginata Baissac, 1952 (Plate VII, 55)

Frequent synonyms / misidentifications: None / Variola louti (non Forsskål, 1775).
FAO names: En - White-edged lyretail; Fr - Croissant queue blanche; Sp - Mero rabiblanco.
Diagnostic characters: Body oblong, its depth less than head length and 2.8 to 3.5 times in standard
length; head length 2.6 to 2.8 times in standard length; interorbital area of adults convex; dorsal head profile
slightly convex; preorbital depth 0.6 to 1.4 times eye diameter and 6 to 10 times in head length; preopercle
rounded, finely serrate, the lower edge fleshy; opercle with 3 flat spines; upper edge of operculum almost
straight; subopercle and interopercle smooth; rear nostrils not much bigger than anterior nostrils. Maxilla
of adults with a distinct step on ventral edge; supramaxilla well developed; both jaws with a pair of large
canines at the front; 1 to 3 large canines at midside of lower jaw; palatines and vomer with teeth. First
gill arch with 7 to 9 gill rakers on upper limb, 13 to 16 lower limb (including 4 to 7 rudiments on each limb).
Dorsal fin with IX spines and 14 soft rays; anal fin with III spines and 8 soft rays; caudal fin lunate,
the upper and lower lobes produced, about twice length of middle rays; pectoral-fin rays 17 to 19,
the middle rays longest, 1.6 to 1.8 times in head length; pelvic fins usually not reaching anus, 1.4 to 1.8
times in head length. Midlateral body scales rough, without auxiliary scales; lateral-line scales 66 to 76;
lateral scale series 109 to 127. Colour: body brownish orange or reddish purple, with irregular red bands
alternating with yellow lines, the red bands containing small irregular pale blue to pink spots; head
orange-yellow, densely spotted with red; median fins with small pink and red spots; rear margin of caudal
fin usually dusky, with a narrow white edge; rear margin of dorsal and anal fins hyaline, without trace of
yellow; pectoral fins yellow, the basal half of rays often reddish or dark brown. Juveniles similar in colour
to adults, but with relatively fewer and larger pale blue or pink spots.
Size: Appears to be smaller than V. louti, the largest known specimen was 47 cm total length; probably
does not grow larger than 55 cm total length, at a maximum weight of 1 kg.
Habitat, biology, and fisheries: Coral reefs at depths of 4 to 200 m. Feeds on fishes. Females mature at
32 cm standard length. Because of its small size and rarity, of little interest to fisheries. Flesh excellent.
Caught with handline, traps, and spear.
Distribution: From east coast
of Africa (Zanzibar and Mafia
Island, Tanzania) to Samoa,
including South China Sea, In-
donesia, Philippines, Taiwan
Province of China, Ryukyu Is-
lands, tropical coast of Austra-
lia, Papua New Guinea, New
Ireland, New Caledonia,
Guam, Palau Islands, Mariana
Islands, Fiji, and Samoa.
Variola louti (Forsskål, 1775) (Plate VII, 56)

FAO names: En - Yellowedged lyretail; Fr - Croissant queue jaune; Sp - Mero luna creciente.
80 cm fork length
Diagnostic characters: Body oblong, its depth less than head

length and 2.8 to 3.3 times in standard length; head length 2.5
to 2.8 times in standard length; interorbital area of adults
convex; dorsal head profile slightly convex; preorbital depth 0.6
to 1.4 times eye diameter and 6 to 10 times in head length; < 7 cm standard length
preopercle rounded, finely serrate, the lower edge fleshy; op-
ercle with 3 flat spines; upper edge of operculum almost
straight; subopercle and interopercle smooth. Maxilla of adults
with a distinct step on ventral edge; supramaxilla well devel-
oped; both jaws with a pair of large canines at the front; 1 to 3
large canines at midside of lower jaw; palatines and vomer
with teeth. First gill arch with 7 to 10 gill rakers on upper limb, 7 cm standard length
15 to 18 lower limb (including 6 to 8 rudiments on each limb).
Dorsal fin with IX spines and 13 or 14 soft rays; anal fin with juveniles
III spines and 8 soft rays; caudal fin lunate, the upper and
lower lobes produced, about twice length of middle rays; pectoral-fin rays 16 to 19, the fin 1.6 to 1.8
times in head length; pelvic fins of adults reach past anus, 1 to 1.7 times in head length. Midlateral body
scales rough, without auxiliary scales; lateral-line scales 66 to 77; lateral scale series 113 to 135. Colour:
head, body, and median fins yellowish brown to orange-red (specimens from deep water more reddish)
with numerous small round or elongate spots of blue, lavender or pink; rear margin of median fins broadly
yellow; pectoral-fin rays red to brown, the distal third abruptly yellow. Large juveniles (10 to 25 cm)
with irregular black band along dorsal part of body ending below rear of dorsal fin and continuing
on head (up to eye) as 3 irregular black spots; black blotch at base of upper caudal-fin rays; head
and body (including black band) with small pale blue to pink spots; a pale yellow or white stripe
middorsally on head from tip of lower jaw to dorsal-fin origin. Small juveniles (less than 7 cm
standard length) lack the black band and black spots on dorsal part of head and body.
Size: Maximum total length 100 cm; maximum weight 12 kg.
Habitat, biology, and fisheries: Coral reefs at depths of 3 to 240 m; usually seen at depths below 15 m.
Prefers islands and offshore reefs, rather than continental shores. Feeds primarily on fishes, including a
variety of coral-reef species. Females mature at 33 cm standard length. Common and important food fish,
but often causes ciguatera fish poisoning. Caught with
handlines, spear, and traps.
Distribution: Tropical Indo-Pacific region from Red Sea
to Pitcairn Islands; known from most (probably all) tropi-
cal islands of the Indian Ocean and western Central
Pacific. In the western Pacific, it ranges from Japan
(southern Honshu) to the northern half of Australia (from
Shark Bay round to New South Wales).
2540 Bony Fishes
Cephalopholis aitha Randall and Heemstra, 1991

En - Rusty hind; Fr - Vielle rouillé; Sp - Cherna herrumbrosa.
Maximum total length about 18 cm. Inshore, in protected or silty reef areas at depths of 5 to 33 m.
Possibly taken in subsistence fisheries, but of no commercial importance. Caught in trawls and gill
nets. Papua New Guinea, Indonesia (Sulawesi, Flores), and the Philippines.
Cephalopholis aurantia (Valenciennes, 1828)

En - Golden hind; Fr - Vielle dorée; Sp - Cherna dorada.
Maximum total length at least 30 cm. Usually caught in depths over 100 m. Specimens are rare in
museums. Because of its small size and occurrence in moderately deep water, the golden hind is
of little commercial importance as a food fish. South Africa and islands of the western Indian Ocean
to the Central Pacific and Japan. Except for a single specimen caught off the coast of South Africa,
there are no confirmed records from continental localities.
Cephalopholis boenak (Bloch, 1790)

En - Chocolate hind; Fr - Vielle chocolat; Sp - Cherna chocolate.
Maximum total length 26 cm. Usually found in depths of 4 to 30 m on silty dead reefs in protected
waters; also taken in trawls to depths of 64 m. Feeds primarily on crustaceans. Spawns in pairs.
Too small to be of commercial importance as a food fish . Caught with hook-and-line, gill nets, traps,
and in trawls. Primarily continental in its distribution, occurring from Kenya to southern Mozambique
and eastwards to the western Pacific (Ryukyu Islands, Taiwan Province of China, China, Philip-
pines, Indonesia, Papua New Guinea, New Caledonia, and eastern Australia). Not known from the
Red Sea, Persian Gulf nor the islands of Micronesia or Polynesia, except for Palau.

Cephalopholis leopardus (Lacepède, 1801)

En - Leopard hind; Fr - Vielle léopard; Sp - Cherna leopardo.
Maximum total length 20 cm (larger sizes reported in the literature are unsubstantiated). Well-
developed coral reefs in depths of 3 to 38 m. Feeds on crustaceans. Like most small groupers, this
is a secretive species, which is usually seen hiding in caves and crevices in the reef; too small to
be of interest to fisheries. African coast to Central Pacific, including most islands of the Indian Ocean,
Viet Nam, Indonesia, Philippines, Taiwan Province of China, Okinawa, Papua New Guinea, northern
Australia, and most islands of the western Central Pacific.
Cephalopholis microprion (Bleeker, 1852)

En - Freckled hind; Fr - Vielle tache de rousseur; Sp - Cherna pecosa.
Maximum total length about 24 cm. Shallow-water silty reefs. Because of its small size, this species
is of minor importance to fisheries. Caught with hook-and-line and probably in traps. Tropical eastern
Indian Ocean and western Pacific: Andaman Sea, Gulf of Thailand, Indonesia, Philippine, Papua
New Guinea, Great Barrier Reef, Solomon Islands, and New Caledonia.
Cephalopholis polleni (Bleeker, 1868)

En - Harlequin hind; Fr - Vielle arlequin; Sp - Cherna arlequin.
Maximum total length 43 cm. Coral reefs in clear water on the edge of steep dropoffs; rarely seen
in depths less than 30 m, it has been taken at 120 m and undoubtedly occurs deeper. Probably
valuable as an attractive aquarium fish, but the species is apparently too rare and too small to be
of commercial importance as a food fish. Caught with hook-and-line and traps. Islands of the
Indo-Pacific region, from the western Indian Ocean to the Central Pacific (Palau and Line Islands);
also reported from the Cocos-Keeling Islands, Christmas Island, Indonesia (small islands), Philip-
pines, Okinawa, Guam, and New Britain. Not known from the Red Sea, Sri Lanka, Australia, larger
islands of Indonesia, or New Guinea.
2542 Bony Fishes
Cephalopholis spiloparaea (Valenciennes, 1828)

En - Strawberry hind; Fr - Vielle fraise; Sp - Cherna frutillera.
Maximum total length 22 cm. Common on coral reefs in depths of 15 to 108 m. Nothing published
on the biology of this species. Because of its small size and relatively deep habitat, the strawberry
hind is of little commercial importance. Indo-Pacific from Pinda, Mozambique (15°S) to French
Polynesia and the Pitcairn Group; ranges from the Ryukyu Islands to Heron Island at the southern
end of the Great Barrier Reef. Except for the Pinda specimens, this species is known only from
insular localities.
Epinephelus bilobatus Randall and Allen, 1987

En - Twinspot grouper; Fr - Mérou gemine; Sp - Mero de pintas gemelas.
Maximum total length at least 33 cm. Coral reefs and rocky bottom in depths of 4 to 50 m. There is
no published information on the biology of this species, or of its interest to fisheries. Epinephelus
bilobatus is known only from northwestern Australia.
Epinephelus bontoides (Bleeker, 1855)

En - Palemargin grouper; Fr - Mérou bord clair; Sp - Mero de margenes palidos.
Maximum total length at least 30 cm. Mud or rocky/cobble bottoms in depths of 2 to 30 m. Appears
to be rare. Nothing has been published on the biology of this species. Probably caught with
hook-and-line, trawls, and in traps. Not of interest to commercial fisheries. Western Pacific:
Indonesia, Philippines, Taiwan Province of China, and New Britain.
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Epinephelus corallicola (Valenciennes, 1828)

En - Coral grouper; Fr - Mérou corail; Sp - Mero de coral.
Maximum total length at least 49 cm total length.
Shallow silty reefs and estuarine areas. Ovary
well developed in a specimen of 30 cm standard
length. Apparently too rare to be of commercial
importance. Western Pacific from Thailand, Hong
Kong, and Taiwan Province of China to Australia
and eastward to the Solomon and Mariana is-
lands, including Indonesia, Philippines, Papua
New Guinea, and Palau.
13.3 cm standard length juvenile 16.2 cm standard length
Epinephelus darwinensis Randall and Heemstra, 1991

En - Darwin grouper; Fr - Mérou Darwin; Sp - Mero de Darwin.
Maximum total length at least 62 cm. Depth of capture 107 m. Biology and interest to fisheries
unknown. Apparently rare; a single specimen caught off Darwin, Australia.
Epinephelus erythrurus (Valenciennes, 1828)

En - Cloudy grouper; Fr - Mérou nébuleux; Sp - Mero nublado.
Maximum total length about 43 cm. In harbours and estuaries with muddy or silty-sand bottoms.
Nothing has been published on the biology of this species; females mature at 15 cm. This species
is of minor commercial importance. Caught with trawls, hook-and-line, and traps. Known from
Pakistan, India, Laccadive (Lakshadweep) Islands, Sri Lanka, Gulf of Thailand, Indonesia, and
Singapore.

2544 Bony Fishes
Epinephelus howlandi (Günther, 1873)

En - Blacksaddle grouper; Fr - Mérou selle noir; Sp - Mero montura negra.
Maximum total length at least 44 cm. Usually found in rocky areas or on coral reefs at depths of 1
to 37 m. Biology unknown. Appears to be rare and is thus of little commercial importance. Tropical
western Central Pacific from Ryukyu Islands to New Guinea, the Great Barrier Reef, Lord Howe
Island, New Caledonia, Vanuatu, Caroline Islands, Palau, Marshall and Mariana islands, Howland
Island, and Samoa.
Epinephelus melanostigma Schultz, 1953

En - Oneblotch grouper; Fr - Mérou dossard; Sp - Mero espaldaron.
Maximum total length at least 33 cm. Shallow coral reefs, also taken over soft bottom. Because of
its small size and apparent rarity, Epinephelus melanostigma is of little or no commercial importance;
caught with handlines, traps, and trawls. Indo-West Pacific from South Africa to Central Pacific,
including Indonesia, Philippines, Hong Kong, Taiwan Province of China, Okinawa, Papua New
Guinea (Bougainville), Solomon Islands, Palau, Caroline Islands, Gilbert Islands, Samoa, Phoenix
Islands, Baker, Howland, Cook Islands, and Line Islands. Reports from Australia have not been
verified.
Epinephelus miliaris (Valenciennes, 1830)

En - Netfin grouper; Fr - Mérou abeille; Sp - Mero colmenar.
Maximum total length 53 cm. Juveniles (10 to 25 cm standard length) over mud bottom, in seagrass
beds, mangrove swamps and on coral reefs in depths of 1 to 16 m; adults on coral reefs in depths
of 18 to 180 m. Females mature at 25 cm stand-
ard length. Apparently rare, hence of minor impor-
tance in local fisheries. Western Indian Ocean to
the Gilbert Islands and Samoa. An insular species
reported from East Burma Sea, Andaman Islands,
Indonesia (Sulawesi [Celebes] only), South China
Sea, Ryukyu Islands, New Guinea, New Ireland,
New Georgia, Solomon and Santa Cruz islands,
Palau, Caroline Islands, and Fiji.
adult 30 cm standard length juvenile 28 mm standard length

Epinephelus multinotatus (Peters, 1876)

En - Whiteblotched grouper; Fr - Vielle plate grise; Sp - Mero de lunares.
Maximum total length 100 cm. Juveniles on inshore coral reefs, and adults to depths of 90 m.
Females mature at 46 cm standard length. Feeds on fishes and crabs. Probably of some commercial
importance in local fisheries. Caught with hook-and-line and traps. Known only from Indian Ocean
(but not Red Sea) including Western Australia (from Monte Bello Islands to Shark Bay).
Epinephelus octofasciatus Griffin, 1926

En - Eightbar grouper; Fr - Mérou huit raies; Sp - Mero de ocho bandas.
Maximum total length at least 130 cm; maximum weight about 80 kg. Occurs in depths of 150 to
350 m. The apparent rarity of this species may be due to its deep-water habitat. Caught with vertical
longlines. Indo-West Pacific from South Africa to Japan, Australia, and New Zealand.
Epinephelus perplexus Randall, Last, and Hoese, 1991

En - Puzzling grouper; Fr - Mérou curieux; Sp - Mero acertijo.
Maximum total length at least 55 cm. Known from a single specimen caught in a depth between
129 and 137 m off southern Queensland, Australia.

2546 Bony Fishes
Epinephelus timorensis Randall and Allen, 1987

En - Yellowspotted grouper; Fr - Mérou taches jaunes; Sp - Mero de pintas amarillas.
Maximum total length at least 32 cm. Known from depths of 73 to 210 m off coral reefs. Nothing
published on biology or fishery of this rare species. Western Australia, American Samoa (Tutuila),
and Phoenix Islands (Canton Island).
Epinephelus trophis Randall and Allen, 1987

En - Plump grouper; Fr - Mérou rondelet; Sp - Mero rollizo.
Maximum total length unknown. Only 2 specimens known, from the base of drilling rig at a depth of
130 m on Dillon Shoals in the Timor Sea off Western Australia.
Epinephelus tuamotuensis Fourmanoir, 1971

En - Reticulate grouper; Fr - Mérou réseau; Sp - Mero reticular.
Maximum total length at least 76 cm. Outer slope of coral reefs in depths of 120 to 250 m. Feeds
on fishes. Probably of some importance in artisanal fisheries, but not commonly caught because of
its preference for deep water. Tuamotu Islands, Society Islands, Pitcairn Group, and Rapa.

Triso dermopterus (Temminck and Schlegel, 1842)

En - Oval grouper; Fr - Mérou ovale; Sp - Mero ovalado.
Maximum total length at least 68 cm. Found on rocky or soft (silty sand or mud) bottoms at depths
of 22 to 103 m. Juveniles feed in the water column on zooplankton. Feeding by adults has not been
observed. Of minor importance in the commercial fishery of Japan and Hong Kong. No data are
available on the landings of this species. Caught with hook-and-line and trawls. Anti-tropical in the
eastern Indian Ocean and western Pacific; known from Korea, Japan, Hong Kong, and Fujian and
Taiwan provinces of China, off the coast of Western Australia (north of 19°30’S) and off eastern
Australia between 25° and 32°S.
adult
2548 Bony Fishes
Tribe NIPHONINI
Niphon spinosus Cuvier, 1828

FAO names: En - Rough grouper; Fr - Mérou rugueux; Sp - Mero aspero.
juvenile 30 cm standard length

(after Takara, 1930)
Diagnostic characters: Body elongate, compressed, its depth 2.9 to 4.2 times in standard length; head
large, 2.5 to 2.8 times in standard length; dorsal head profile straight; interorbital area slightly convex, its
width 4.9 to 6.8 times in head length; preorbital serrate; opercle with 3 strong spines; upper edge of
operculum convex; preopercle angular, the rear edge serrate, with a large posteriorly-directed spine
at angle and a few smaller spines on lower edge; suborbital depth less than eye diameter, 11 times
in head length. Rear nostril an elongate slit, close to anterior nostril. Mouth large, lower jaw projecting;
band of villiform teeth on jaws, vomer, and palatines; no teeth on tongue. No supramaxilla. Gill rakers long
and slender, 7 to 9 on upper limb, 13 to 16 on lower limb. Dorsal fin with XIII spines and 10 or 11 soft
rays, the fin divided to base before last spine, which is much longer than twelfth spine; fourth spine
longest, longer than longest dorsal-fin ray and 2 to 2.6 times in head length; anal fin with III spines and 6
to 8 soft rays; caudal fin emarginate, with rounded corners, 8 branched rays in upper lobe and 7 in
lower lobe; pectoral-fin rays 15 to 17, the fin length about equal to pelvic fin, 1.8 to 2.4 times in head length.
Scales small and rough; lateral-line scales 84 to 93; lateral scale series 140 to 163. Colour: Head and
body brownish grey, the fins darker; juveniles silvery with dark lateral band from snout through eye to upper
part of caudal peduncle and another, median, dark band from snout along top of head, dorsal part of body,
and continued over front half of soft-dorsal fin; upper and lower thirds of caudal fin dusky, the corners and
middle part of fin white.
Habitat, biology, and fisheries: Rocky bottom in depths of 50 to 100 m. Nothing published on the biology
or fisheries of this species. Of some importance as a food fish in Japan. Caught with trawls and longlines.
Distribution: Japan, Korea,
and the Philippines (Sulu
Sea).
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Perciformes: Percoidei: Centrogeniidae 2549
CENTROGENIIDAE
(= Centrogenysidae)
False scorpionfish
by A.C. Gill
A single species in this family.
Centrogenys vaigiensis (Quoy and Gaimard, 1824)

Frequent synonyms / misidentifications: Centropristes scorpaenoides C u v i e r in Cuvier and
Valenciennes, 1829; Sebastes stoliczkae Day, 1875 (including combinations of these in Myriodon,
Gennadius, and Rhabdosebastes) / None.
FAO names: En - False scorpionfish.
Diagnostic characters: A perch-like species; moderately deep bodied (greatest body depth 34 to 39%
standard length). Head large, 42 to 44% standard length. Posterior rim of anterior nostril with large
fringed flap. Opercle with 2 spines, the lower spine strong and conspicuous, the upper spine (near dorsal
edge of operculum) weak and concealed by skin and scales; preopercle finely serrate posteriorly, with
3 or 4 large, antrorse (forward pointing) spines on ventral margin. Mouth moderately large and
protrusible, reaching to anterior half of eye; supramaxilla present. A band of villiform teeth in each jaw; a
chevron-shaped band of villiform teeth on vomer; an elongate patch of villiform teeth on each palatine; no
teeth on tongue. Seven branchiostegal rays; branchiostegal membranes not united. Gill rakers short or
rudimentary, the first gill arch with 15 to 19 gill rakers, of which 3 to 6 on upper limb, and 12 to 14 on lower
limb. Dorsal fin with XIII or XIV spines and 9 to 11 branched, segmented rays, the origin of fin
situated over middle of head above posterior edge of preopercle; dorsal-fin membranes project slightly
beyond spine tips. Anal fin short based, with III spines and 5 branched, segmented rays; second
anal-fin spine strong and very long (28 to 32% standard length), reaching to or beyond posterior
margin of anal fin when depressed. Caudal fin rounded to truncate, with 6 (rarely 5) and 6 branched
rays in upper and lower part, respectively, and 4 to 6 unbranched rays in upper and lower part.
Pelvic fins positioned behind pectoral fins, consisting of I spine and 5 branched, segmented rays, the
inner ray broadly bound to body by membrane. Pectoral-fin rays 12 to 14, with only the upper ray
unbranched. Scales on body ctenoid, extending onto snout; lateral line relatively low on body, extending
from gill opening to caudal-fin base, consisting of 36 to 44 tubed scales, with additional 4 to 6 tubed scales
on caudal fin; tubed scales on body each with a dorsal branch of tube extending to edge of scale; soft
portions of dorsal and anal fins with basal covering of scales. Vertebrae 11+14; lower hypurals (hypurals
1 and 2) fused into a single plate. Tooth plates of fifth ceratobranchials broadly overlapping and
2550 Bony Fishes
asymmetrical. Colour: head and body pale grey to pale brown, paler ventrally, with dark grey to brown
mottling; fins pale grey or pale brown to white or clear, with large dark grey to dark brown spots, these
forming irregular bands (particularly on pectoral and pelvic fins, and on anterior part of anal fin).

Cirrhitidae: similar in general appearance, but easily
distinguished in having X dorsal-fin spines, cycloid
scales, branchiostegal membranes joined and form-
ing a free fold across isthmus, 6 branchiostegal rays,
10+16 vertebrae, the lower 5 to 7 pectoral-fin rays
unbranched and usually thickened with deeply in-
cised membranes, and in lacking large antrorse
spines on the ventral edge of the preopercle.
Scorpaenidae: distinguished by the more extensive
head spination/ornamentation, particularly in having Cirrhitidae
a suborbital stay (or ridge), a spine-like extension of
third infraorbital bone toward the preopercle (usually
bound to the preopercle in most species).
Serranidae: sometimes resemble Centrogenys (particularly certain benthic anthiines), but easily distin-
guished in having 3 opercular spines, 6 to 22 segmented anal-fin rays, usually fewer dorsal-fin spines
(usually XI or fewer; only Niphon and Acanthistius have as many as XIII), and at least some canine teeth
in the jaws.
Scorpaenidae Serranidae

Habitat, biology, and fisheries: A sedentary species that occurs on rocky reefs and rubble bottoms in
shallow coastal waters. Feeds during the day on small fishes, shrimps, and crabs. Of no commercial
importance, although occa-
sionally enters the aquarium
fish trade.
Distribution: West Pacific
and eastern Indian oceans,
from Nicobar Islands east to
New Guinea and northern
Australia, north to the Ryukyu
Islands.
Reference
Weber, M. and L.F. de Beaufort. 1931. The fishes of the Indo-Australian Archipelago. Vol. 6. Leiden, E.J. Brill, 448 p.
Perciformes: Percoidei: Ostracoberycidae 2551
OSTRACOBERYCIDAE
Ostracoberycids
by K.E. Carpenter
D iagnostic characters: Body ovoid and moderately compressed (size to around 19 cm standard
length), its height 37 to 44% of standard length. Eye large, longer than snout length; a very strong
spine extending back from lower limb of preopercle beyond base of pectoral fins; juveniles up to
10 cm standard length with a median supraoccipital spine. Gill rakers on first gill arch 4 to 9 on upper limb
and angle, and 9 to 17 on lower limb. Two separate (but close) dorsal fins, the first with IX spines and
the second with 9 or 10 soft rays. Anal fin with III spines and 7 soft rays. Caudal fin truncate. Lateral
line with 48 to 58 scales. Colour: brownish on upper sides and whitish on lower sides; fins mostly hyaline,
without prominent markings, the first dorsal fin more dusky.
2 separate dorsal fins, the 1st with IX spines
and the second with 9-10 soft rays
prominent preopercular spine

extends posteriorly beyond
base of pectoral fins
anal fin with III spines and 7 soft rays
Habitat, biology, and fisheries: Demersal or near bottom on the continental slope at depths between 250
and 710 m. Very little is known about the biology of these fishes. Caught incidentally in deep-bottom trawls.

The prominent spine extending posteriorly from the preopercle, general body shape, 2 distinct dorsal fins,
and pelvic-fin formula distinguishes ostracoberycids from all other fishes in the area. Other fishes with a
prominent preopercular spine either have a single dorsal fin and very different body shape (e.g. some
Engraulidae and Pomacanthidae) or an armoured head and very different body shape (e.g. Dactylopteridae
and Triglidae of the subfamily Peristediinae), or different pelvic- and anal-fin spine and ray counts.
Ostracoberycids superficially resemble fishes of the order
Beryciformes but have the typical perciform pelvic-fin con-
figuration of I spine and 5 soft rays not found in beryciforms.
They also differ from all beryciforms with the combination
of an elongate preopercular spine, and 2 separate dorsal
fins or anal fin with III spines.
Ostracoberycids most closely resemble holocentrid fishes
of the genus Sargocentron and some Neoniphon that also
posses an elongate preopercular spine. However, the pre-
opercular spine in these beryciform fishes does not extend
beyond the base of the pectoral fins and, they posses IV
anal-fin spines (III in ostracoberycids), a pelvic-fin ray
configuration of I spine and 7 soft rays (I spine and 5 soft
rays in ostracoberycids), and forked caudal fins (truncate Sargocentron (Holocentridae)
in ostracoberycids). In addition, holocentrids with promi-
nent preopercular spines are nearshore fishes while ostra-
coberycids are found on the continental slope.
2552 Bony Fishes
Key to the species of Ostracoberycidae occurring in the area

1a. Lower limb of first gill arch with 9 to 13 gill rakers; juveniles with median supraoccipital
spine that curves forward . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ostracoberyx paxtoni
(southern Queensland)
1b. Lower limb of first gill arch with 15 to 17 gill rakers; juveniles with median supraoccipital
spine that curves slightly posteriorly . . . . . . . . . . . . . . . . . . . . . .Ostracoberyx dorygenys
(southern Japan, Philippines, western Indian Ocean)

Ostracoberyx dorygenys Fowler, 1934
Ostracoberyx paxtoni Quero and Ozouf-Costaz, 1991
Reference
Quero, J.-C. and C. Ozouf-Costaz. 1991. Ostracoberyx paxtoni, nouvellle espèce des côtes est de l’Australie.
Remarques sur les modifications morphologiques des Ostracoberyx au cours de leur croissance (Perciformes,
Ostracoberycidae). Cybium, 15(1):43-54.
Perciformes: Percoidei: Callanthiidae 2553
CALLANTHIIDAE
Groppos (also goldies and splendid perches)
by W.D. Anderson, Jr.
D iagnostic characters: Body oblong, somewhat compressed (size to about 25 cm). Eyes moderate to
large, their diameter longer than snout length. Nasal organ without lamellae. Opercular spines 1
or 2. Mouth terminal and oblique; jaws almost equal. Jaws with caniniform, conical, and villiform teeth;
vomer usually with teeth; palatine with or without teeth; pterygoids and tongue without teeth. Bran-
chiostegal rays 6. First gill arch with 7 to 12 gill rakers on upper limb and 17 to 28 on lower limb (total 24
to 38). Dorsal fin not incised at junction of spinous and soft rays. Soft rays of dorsal and anal fins somewhat
elongated, posteriormost rays sometimes noticeably produced. Caudal fin lunate to almost truncate, upper
and lower lobes frequently produced in larger individuals. Dorsal fin with XI spines and 9 to 11 soft rays.
Anal fin with III spines and 9 to 11 soft rays. Principal caudal-fin rays 15 (8 in upper lobe, 7 in lower lobe),
or 17 (9 in upper lobe, 8 in lower lobe). Pectoral-fin rays 17 to 23 (usually 19 to 22). Pelvic fins with I spine
and 5 soft rays. Membranes of dorsal and anal fins without scales. Axillary scales of pelvic fins and scaly
interpelvic process typically well developed. Most of head, including maxillae and usually dentaries,
covered with scales. Scales moderate to large, ctenoid, without ctenial bases in posterior field. Series of
modified scales with unique ornamentation along body midlaterally. Lateral line running along base
of dorsal fin and terminating near base of ultimate soft dorsal-fin ray or continuing posteriorly on
dorsolateral surface of caudal peduncle. Tubed lateral-line scales 14 to 42. Vertebrae 10+14=24.
Colour: magnificently coloured, with various shades of red, orange, yellow, blue, and purple being
prominent.
lateral line running along dorsal-fin base
caudal-fin
lobes
often
produced
1-2 opercular
spines (after Rivaton, 1989)
Habitat, biology, and fisheries: Usually found near the bottom over rocky areas and on reefs (frequently
near a reef-sand boundary) in depths of 4 to 660 m. Carnivorous. Probably protogynous hermaphrodites.
Remarks: In the past considered as members of the family Serranidae, but species of Callanthiidae share
derived characters that are not found in the Serranidae.
Serranidae: 3 opercular spines (versus 1 or 2 in Callan- 3 opercular
thiidae); nasal organ with lamellae (versus nasal organ spines
devoid of lamellae); 6 or 7 branchiostegal rays (versus 6);
midlateral body scales without ornamentation (versus
series of midlateral body scales with unique ornamenta-
tion); lateral line running at least a few scale rows below
dorsal fin, usually uninterrupted and usually extending to
at least base of caudal fin (versus lateral line running
along base of dorsal fin and terminating near base of
ultimate soft dorsal-fin ray or continuing posteriorly on Serranidae (subfamily Anthiinae)
dorsolateral surface of caudal peduncle).
2554 Bony Fishes
Key to the species of Callanthiidae occurring in the area

Remarks on key characters: counts of lateral-line scales are of tubed scales along the body.
1a. Lateral-line scales 34 to 41 (usually 35 to 40); dorsal fin with XI spines and 10 or 11 soft
rays (almost always 11); anal fin with III spines and 10 or 11 soft rays (almost always
11); first gill arch with 7 to 10 gill rakers on upper limb and 23 to 28 on lower limb (total
30 to 38, usually 32 to 36); principal caudal-fin rays 17 (9 in upper lobe, 8 in lower lobe);
2 opercular spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Callanthias australis
1b. Lateral-line scales 14 to 17; dorsal fin with XI spines and 9 soft rays; anal fin with III
spines and 9 soft rays; first gill arch with 8 or 9 gill rakers on upper limb and 18 to 20 on
lower limb (total 26 to 29); principal caudal-fin rays 15 (8 in upper lobe, 7 in lower lobe);
1 opercular spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Grammatonotus crosnieri

0 Callanthias australis Ogilby, 1899
0 Grammatonotus crosnieri (Fourmanoir, 1981)
Reference
Anderson, W.D., Jr. and G.D. Johnson. 1984. A new species of Callanthias (Pisces: Perciformes: Percoidei:
Callanthiidae) from the southeastern Pacific Ocean. Proc. Biol, Soc. Wash., 97(4):942-950.
Perciformes: Percoidei: Callanthiidae 2555
Callanthias australis Ogilby, 1899

Frequent synonyms / misidentifications: Callanthias platei australis Ogilby, 1899 / Callanthias allporti
Günther, 1876.
FAO names: En - Glorious groppo.
(after Rivaton, 1989)
Diagnostic characters: Body slender to moderately deep. Head moderate. Eyes moderate. Opercular
spines 2. First gill arch with 7 to 10 gill rakers on upper limb and 23 to 28 on lower limb (total 30 to
38, usually 32 to 36). Dorsal fin with XI spines and 10 or 11 soft rays (almost always 11). Anal fin
with III spines and 10 or 11 soft rays (almost always 11). Principal caudal-fin rays 17 (9 in upper
lobe, 8 in lower lobe); branched caudal-fin rays 15 (8 in upper lobe, 7 in lower lobe); both lobes of
caudal fin well produced in some specimens (these possibly males). Pectoral-fin rays 18 to 23 (usually 21
or 22). Tubed lateral-line scales 34 to 41 (usually 35 to 40). Colour: body and head rose, mostly orange
with admixture of yellow, or mostly yellow; dorsal, anal, and pelvic fins yellow or mostly yellow; caudal fin
yellow proximally and rosy or purplish distally or almost entirely purplish red; dorsalmost and ventralmost
caudal-fin rays, when produced, yellow distally or for their entire lengths; pectoral fins yellow, or with
considerable yellow basally and most of fins rosy or purplish red.
Size: Maximum standard length about 25 cm; commonly to 18 cm.
Habitat, biology, and fisheries: Occurs in depths of 15 to 365 m. A schooling species that seeks the
cover of caves and crevices at night and when disturbed. Presumably feeds on plankton. No information
available on fisheries.
Distribution: Occurs in wa-
ters off New Zealand and in
temperate waters off Austra-
lia; also known from a collec-
tion made northeast of
Rockhampton (Queensland)
and from another collection in
the Chesterfield Island region
of the Coral Sea; may reach
Indonesian waters.
Remarks: Closely related to
Callanthias japonicus from
J ap an es e w ater s and C.
platei from the eastern South
Pacific; all 3 taxa may be sub-
species of a single wide-rang-
ing species.
2556 Bony Fishes
Grammatonotus crosnieri (Fourmanoir, 1981)

Frequent synonyms / misidentifications: Callanthias crosnieri Fourmanoir, 1981 / None.
FAO names: En - Uncle Phoo’s groppo.
Diagnostic characters: Body depth moderate to fairly deep. Head moderate. Eyes fairly large. One
opercular spine. First gill arch with 8 or 9 gill rakers on upper limb and 18 to 20 on lower limb (total
26 to 29). Both lobes of caudal fin extremely well produced in some specimens (these possibly males).
Dorsal fin with XI spines and 9 soft rays. Anal fin with III spines and 9 soft rays. Principal caudal-fin
rays 15 (8 in upper lobe, 7 in lower lobe); branched caudal-fin rays 13 (7 in upper lobe, 6 in lower
lobe). Pectoral-fin rays 18 to 20. Tubed lateral-line scales 14 to 17. Colour: upper body dull orange or
yellow, lower body pale to rosy; head orange or yellow above, pale below; fins mostly dull orange to yellow.
Habitat, biology, and fisheries: Known from depths of 170 to 210 m. No other information available.
from the Philippines, off the
southwestern coast of Luzon.
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Perciformes: Percoidei: Pseudochromidae 2557
PSEUDOCHROMIDAE
Dottybacks (eelblennies)
by A.C. Gill
D iagnostic characters: Perch-like to eel-like fishes (size to 45 cm), moderately deep bodied to
elongate, usually moderately compressed. Mouth moderate, weakly protrusible; maxilla usually not
extending posteriorly beyond middle of eye. Branchiostegal rays 6, with a single ray articulating with
the posterior ceratohyal (epihyal). Dorsal fin long based, composed of III or fewer spines (very
rarely IV spines in some species) and 21 to 74 segmented rays. Lateral line modified (disjunct,
posteriorly truncated or multiple). Adductor mandibulae (jaw) musculature with a superficial, strap-like
A1 section, which inserts on the medial face of the maxilla and originates on the vertical part of the
preopercle. Hypurals 1 and 2 fused to each other; hypurals 3 and 4 fused to each other and to urostylar
complex; no interarcual cartilage between epibranchial 1 and infrapharyngobranchial 2; anterior part of
suspensorium weakly attached to posterior part.
Habitat, biology, and fisheries: Cryptic diurnal inhabitants of coral reefs and rock bottoms; a few species
live in large sponges; occur from shallow intertidal areas to depths of about 100 m. Most are generalist
micropredators feeding on small fishes, crustaceans, small molluscs, and worms. Usually occur singly or
in pairs. Many if not most species are sequential hermaphrodites. Eggs are ornamented with filaments
and, sometimes, hook-like structures; eggs are arranged in a ball or mass, owing to the filaments of
adjacent eggs entangling with each other; the egg mass is guarded by the male in a burrow or cave until
the larvae hatch and become free swimming. The brighter coloured species (particularly Pseudochromis
diadema, P. paccagnellae, and P. porphyreus) are of commercial importance in the aquarium fish trade.
Remarks: The generic assignment of species should be regarded as provisional; some species presently
included herein in Pseudochromis and Pseudoplesiops will eventually be assigned to new genera.

Callanthiidae: resemble some pseudochromids, but have more dorsal-fin spines (X or XI versus III or
fewer).
Notograptidae: resemble some pseudochromids, but have mostly spinous rays in the dorsal and anal fins,
a larger mouth (extending well beyond the posterior margin of the eye versus to anterior or middle of eye),
and a conspicuous barbel-like lobe on the middle of the lower lip.
Callanthiidae Notograptidae
2558 Bony Fishes
Opistognathidae: resemble some pseudochromids, but have more dorsal-fin spines (IX to XII versus III
or fewer), a much larger mouth (extending well beyond posterior edge of eye versus to anterior or middle
of eye), and a different configuration of the pelvic fins (composed of I spine and 5 segmented rays, the
inner 3 weak and branched, the outer 2 stout and unbranched).
Pholidichthyidae: resemble some pseudochromids, but have a slightly inferior mouth (lower jaw projecting
in pseudochromids), and a single nostril (versus 2 in pseudochromids) on each side of the head.
Pholidichthyidae
Opistognathidae
Plesiopidae: resemble some pseudochromids, but have more dorsal-fin spines (IX to XXVI versus III or
fewer), and usually larger mouth (extending to posterior part of eye or beyond versus to anterior or middle
of eye).
Serranidae: resemble certain pseudochromids, but usually have more dorsal-fin spines (VII to XIII in
species within the area versus III or fewer in pseudochromids; among serranids, only Rypticus from the
east Pacific and Atlantic has fewer than VII [II to IV] dorsal-fin spines), have 6 or 7 branchiostegal rays,
and usually 3 (sometimes only 2 apparent) spines on opercle.
Plesiopidae Serranidae
Key to the genera of Pseudochromidae occurring in the area

Remarks on key characters: generic characters given in the key are based only on species occurring in
the area.
1a. Body elongate, body depth at anal-fin origin less than 16% standard length; segmented
dorsal-fin rays 49 to 74; segmented anal-fin rays 38 to 66; pectoral-fin rays 9 to 11;
branchiostegal membranes united posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Body not elongate, body depth at anal-fin origin greater than 20% standard length;
segmented dorsal-fin rays 21 to 38; segmented anal-fin rays 10 to 21; pectoral-fin rays 15
to 20; branchiostegal membranes not united posteriorly . . . . . . . . . . . . . . . . . . . . . . . ® 4
2a. Pelvic fins present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blennodesmus

2b. Pelvic fins absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Dorsal fin with none or I spine, and with 57 to 74 segmented rays; ocellated black spot
either absent or on opercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Congrogadus
3b. Dorsal fin with II spines (second minute and buried in flesh behind first spine) and 49
segmented rays; ocellated spot on shoulder above gill opening . . . . . . . . . . . . . . . Haliophis
4a. Anterior lateral line consisting of a single tubed scale, and an intermittent series of
centrally pitted scales; posterior lateral line consisting of an intermittent series of
centrally pitted scales; pelvic fins with I spine and 3 or 4, usually unbranched, segmented
rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops
4b. Anterior lateral line consisting of a series of tubed scales; posterior lateral line consisting
of a series of tubed scales; pelvic fins with I spine and 5 branched, segmented rays . . . . . . . . ® 5
5a. Vomerine teeth small and arranged in broad triangular patch; segmented dorsal-fin rays
31 to 38; dorsal and anal fins with well-developed scaly sheaths . . . . . . . . . . . . . . Ogilbyina
5b. Vomerine teeth relatively large and arranged in a chevron-shaped patch; combination of
other characters not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Tubed scales in anterior lateral line 43 to 62 . . . . . . . . . . . . . . . . . . . . . . . . . Labracinus

6b. Tubed scales in anterior lateral line 17 to 40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Opercle with serrations ventral to subopercle junction; at least some scales on cheek
and/or operculum ctenoid; vertebrae 11+17 (rarely 11+18); supraneural bones 2, or 3 with
the third much smaller than other 2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Assiculus
7b. Combination of first 2 characters not as above; vertebrae 10+16; supraneural bones 3, of
approximately equal size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Dark oblique lines present on at least some anterior body scales; segmented dorsal-fin
rays 23 (rarely 22 or 24); segmented anal-fin rays 14 (rarely 13 or 15); circumpeduncular
scales 16 (rarely 14, 15, or 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cypho
8b. No dark oblique lines on anterior body scales; combination of meristic characters not as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis
Key to the species of Pseudochromidae occurring in the area

Remarks on key characters: “scales in lateral series” refer to the number of scales in the anterior lateral
line plus the number of posteroventrally sloping scale rows on the midside of the caudal peduncle, the
latter beginning with the scale row immediately following the row including the last anterior lateral-line
scale, and ending with the row including the scale at the hypural edge.
1a. Body elongate, body depth at anal-fin origin less than 16% standard length; segmented
dorsal-fin rays 49 to 74; segmented anal-fin rays 38 to 66; pectoral-fin rays 9 to 11;
branchiostegal membranes united posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Body usually not elongate, body depth at anal-fin origin usually greater than 20% standard
length (15% in Pseudoplesiops sp. 3); segmented dorsal-fin rays 21 to 38; segmented
anal-fin rays 10 to 21; pectoral-fin rays 15 to 20; branchiostegal membranes not united
posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
2a. Pelvic fins present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Blennodesmus scapularis

2b. Pelvic fins absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Dorsal fin with none or I spine, and with 57 to 74 segmented rays; ocellated black spot
either absent or on opercle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3b. Dorsal fin with II spines (second minute and buried in flesh behind first spine) and 49
segmented rays; ocellated spot on shoulder above gill opening . . . . . . . . . Haliophis aethiopus
4a. Pale round spots present on body; gill membranes fused to isthmus . . . . . . . . . . . . . . . . ® 5
4b. No pale round spots on body; gill membranes fused together but free from isthmus . . . . . . . . ® 7
2560 Bony Fishes
5a. White horizontal stripe from eye to lateral-line origin . . . . . . . . . . . . . . Congrogadus spinifer
5b. No white horizontal stripe from eye to lateral-line origin, although a few horizontally aligned
white spots sometimes present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Anterior part of body with 5 or 6 rows of pupil-diameter sized pale spots, the ventralmost
row of spots not encroaching onto anal-fin ray bases . . . . . . . . . . . . Congrogadus malayanus
6b. Anterior part of body with 3 or 4 rows of eye-diameter sized pale spots, the ventralmost
row of spots encroaching onto anal-fin ray bases . . . . . . . . . . . . Congrogadus amplimaculatus
7a. Dorsal fin without spines, with 68 to 76 segmented rays; sensory pores on head not
surrounded by black spots . . . . . . . . . . . . . . . . . . . . . . . . . . . Congrogadus subducens
7b. Dorsal fin with I spine and 58 to 62 segmented rays; each sensory pore on head
surrounded by small black spot . . . . . . . . . . . . . . . . . . . . . . . Congrogadus hierichthys
8a. Anterior lateral line consisting of a single tubed scale, and an intermittent series of
centrally pitted scales; posterior lateral line consisting of an intermittent series of centrally
pitted scales; pelvic fins with I spine and 3 or 4, usually unbranched, segmented rays . . . . . . . ® 9
8b. Anterior lateral line consisting of a series of tubed scales; posterior lateral line consisting
of a series of tubed scales (sometimes absent in juveniles of some species); pelvic fins
with I spine and 5 branched, segmented rays . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 21
9a. Scales with distinct centres and radii in all fields . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
9b. Scales without distinct centres, with radii confined to anterior field . . . . . . . . . . . . . . . . ® 11
10a. Dorsal fin with I weak spine and 22 or 23 segmented rays; anal fin with I weak spine and
13 or 14 segmented rays; transverse scale rows from gill opening to base of caudal fin 26
to 29; posterior rim of eye broadly edged with dark grey to black . . . . . . . . . Pseudoplesiops rosae
10b. Dorsal fin with II weak spines and 24 to 26 segmented rays; anal fin with II or III weak
spines and 14 to 16 segmented rays; transverse scale rows from gill opening to base of
caudal fin 33 to 39; narrow red to grey or black ring around edge of eye . . . . Pseudoplesiops typus
11a. Posterior body scales ctenoid; transverse scale rows from upper edge of gill opening to
base of caudal fin 30 to 42 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
11b. All body scales cycloid; transverse scale rows from upper edge of gill opening to base of
caudal fin 53 to 66 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
12a. Segmented dorsal-fin rays 23 or 24; segmented anal-fin rays 13 to 15 (usually 14) . . . . . . . ® 13
12b. Segmented dorsal-fin rays 25 to 29; segmented anal-fin rays 15 to 18 . . . . . . . . . . . . . . ® 14
13a. Predorsal scales extending anteriorly to or almost to posterior part of eyes; eye diameter
10 to 13% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops knighti
13b. Predorsal scales extending anteriorly to above posterior edge of preopercle; eye diameter
8 to 10% standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops howensis
14a. A prominent fleshy keel or flap on chin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
14b. No prominent fleshy keel or flap on chin, at most a weak ridge . . . . . . . . . . . . . . . . . . ® 16
15a. Large dark spot on opercle . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops revellei

15b. No large dark spot on opercle . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops sp. 1
16a. Circumpeduncular scales 20; 2 dark, oblique bars on head and anterior part of body
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops collare
16b. Circumpeduncular scales 16; head without dark oblique bars . . . . . . . . . . . . . . . . . . . ® 17
17a. Segmented dorsal-fin rays 25 or 26 (usually 25); segmented anal-fin rays 14 to 16 (usually
15); dark spots, if present, on bases of each segmented ray of dorsal and anal fins; pelvic
fins usually with small, conspicuous dark spots . . . . . . . . . . . . . . . . . Pseudoplesiops annae
17b. Segmented dorsal-fin rays 27 to 29 (usually 28); segmented anal-fin rays 16 to 18; dark
spots present on bases of alternate segmented rays of dorsal and anal fins; pelvic fins
without dark spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops sp. 2
18a. Body depth at dorsal-fin origin 15% standard length; vertebrae 14+18 . . . . . . Pseudoplesiops sp. 3
18b. Body depth at dorsal-fin origin 19 to 29% standard length; vertebrae 12-13+17-18 (rarely
19) = 30 (rarely 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
19a. Four anal-fin pterygiophores precede second haemal spine; vertebrae 13+17; body depth
at dorsal-fin origin 19 to 21% standard length . . . . . . . . . . . . . . . . . . . Pseudoplesiops sp. 4
19b. Three (rarely 2) anal-fin pterygiophores precede second haemal spine; vertebrae usually
12+18 (rarely 13+17 or 12+19); body depth at dorsal-fin origin 21 to 29% standard length . . . . ® 20
20a. Two (rarely 1) anal-fin pterygiophores precede first haemal spine; body depth at dorsal-fin
origin 21 to 25% standard length; pectoral-fin rays 16 to 18 (usually 17) . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudoplesiops multisquamatus
20b. One (rarely 2) anal-fin pterygiophores precede first haemal spine; body depth at dorsal-fin
origin 25 to 29% standard length; pectoral-fin rays 17 to 19 . . . . . . . . . . . . Pseudoplesiops sp. 5
21a. Vomerine teeth small and arranged in broad triangular patch; segmented dorsal-fin rays
31 to 38; dorsal and anal fins with well-developed scaly sheaths . . . . . . . . . . . . . . . . . ® 22
21b. Vomerine teeth relatively large and arranged in a chevron-shaped patch; combination of
other characters not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 24
22a. Scales above anal-fin origin 15 to 20 (formula: 14 to 16 + 1 + 2 or 3 = 15 to 20); cir-

cumpeduncular scales 19 or 20; dark brown (bright blue to dark purple or black in life)
spots present on cheeks, operculum, pectoral-fin base, and breast . . . . . . . . . . Ogilbyina salvati
22b. Scales above anal-fin origin 21 to 28 (formula: 17 to 23 + 1 + 2 to 5 = 21 to 28); cir-
cumpeduncular scales 22 to 26; coloration not as above . . . . . . . . . . . . . . . . . . . . . . ® 23
23a. Body relatively shallow, dorsal-fin origin to pelvic-fin origin 25 to 31% standard length; gill
rakers on lower limb of first gill arch 10 to 12 (usually 11); first dorsal-fin pterygiophore
inserted either between first and second neural spines or anterior to first neural spine . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ogilbyina novaehollandiae
23b. Body relatively deep, dorsal-fin origin to pelvic-fin origin 32 to 36% standard length; gill
rakers on lower limb of first gill arch 11 to 13 (usually 12); first dorsal-fin pterygiophore
inserted between second and third neural spines . . . . . . . . . . . . . . Ogilbyina queenslandiae
24a. Tubed scales in anterior lateral line 43 to 62 . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 25

24b. Tubed scales in anterior lateral line 17 to 40 . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 26
25a. Horizontal scale rows above anal-fin origin to dorsal-fin base 30 to 31; pseudobranch
filaments 23 or 24; circumpeduncular scales 32; in preservative a series of narrow (less
than 1 scale wide) dark brown oblique bars on sides of body beneath posterior two-thirds
of dorsal fin; a large (subequal to pupil) dark brown spot positioned on basal third of dorsal
fin between segmented rays 21 and 24 . . . . . . . . . . . . . . . . . . . . Labracinus atrofasciatus
25b. Horizontal scale rows above anal-fin origin to dorsal-fin base 23 to 30 (usually 24 to 28);
pseudobranch filaments 12 to 20; circumpeduncular scales 24 to 30 (usually 26 to 28);
bars on sides of body, if present, broad (several scales wide) and confined to beneath
anterior half of dorsal fin; spots if present at posterior of dorsal fin small (less than 1/2 pupil
diameter) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Labracinus cyclophthalmus
26a. Opercle with serrations ventral to subopercle junction; at least some scales on cheek
and/or operculum ctenoid; vertebrae 11+17 (rarely 11+18); supraneural bones 2, or 3 with
the third much smaller than other 2 . . . . . . . . . . . . . . . . . . . . . . . . . Assiculus punctatus
26b. Combination of first 2 characters not as above; vertebrae 10+16; supraneural bones 3, of
approximately equal size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 27
2562 Bony Fishes
27a. Dark oblique lines present on at least some anterior body scales; segmented dorsal-fin
rays 23 (rarely 22 or 24); segmented anal-fin rays 14 (rarely 13 or 15); circumpeduncular
scales 16 (rarely 14, 15, or 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 28
27b. No dark oblique lines on anterior body scales; combination of meristic characters not as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 29
28a. Central dark lines on scales on anterior part of body aligning to form oblique bars . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cypho purpurascens
28b. Central dark lines on scales not aligning to form oblique bars . . . . . . . . . . . . . . . . Cypho sp.
29a. Segmented dorsal-fin rays 20 to 22 (usually 22); segmented anal-fin rays 10 to 13 (usually
11 or 12); total gill rakers on first gill arch 18 to 24, of which 5 to 8 on upper limb, and 13
to 17 on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 30
29b. Combination of meristic characters not as above . . . . . . . . . . . . . . . . . . . . . . . . . ® 35
30a. Posterior part of body uniformly blue grey or reddish purple to magenta in life . . . . . . . . . . ® 31
30b. Posterior part of body entirely or partly bright yellow in life (yellow area usually noticeably
paler than anterior or dorsoanterior part of body in preservative) . . . . . . . . . . . . . . . . . ® 32
31a. Head and body uniformly reddish purple to magenta in life (pale brown to purplish grey or
brown in preservative) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis porphyreus
31b. Snout, dorsal contour of head, and sometimes dorsal contour of body to beneath posterior
part of dorsal fin bright yellow in life (pale brown in preservative), the remainder of head
and body dark blue-grey to magenta (brown in preservative) . . . . . . . . Pseudochromis aurifrons
32a. Dorsal contour of head and body magenta in life (greyish brown in preservative), the
remainder of head and body bright yellow (pale brown in preservative) . . . Pseudochromis diadema
32b. Head entirely reddish purple to magenta in life . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 33
33a. Posterior part of body bright purple (pinkish grey in preservative) with bright yellow (pale
yellow in preservative) saddle-like marking extending from anterior part of dorsal fin to
upper caudal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis ephippiatus
33b. Head and body anterior to a more or less vertical line through vicinity of anal-fin origin
magenta in life (greyish brown to brown or grey in preservative), the remainder of body
bright yellow (pale brown to brown or pale greyish brown in preservative) . . . . . . . . . . . . .® 34
34a. Scales in lateral series 33 to 39 (usually 34 to 38) . . . . . . . . . . . . Pseudochromis paccagnellae

34b. Scales in lateral series 36 to 42 (usually 38 to 42) . . . . . . . . . . . . . . . . Pseudochromis sp. 1
35a. Segmented dorsal-fin rays 22; segmented anal-fin rays 13; scales in lateral series 28 to
32; dorsal and anal fins with well-developed scaly sheaths . . . . . . . . Pseudochromis marginatus
35b. Character combination not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 36
36a. Dorsal fin with II spines and 25 or 26 (usually 26) segmented rays; segmented anal-fin
rays 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis veliferus
36b. Character combination not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 37
37a. At least some scales on cheek and/or operculum ctenoid . . . . . . . . . . . . . . . . . . . . ® 38

37b. All scales on cheek and operculum cycloid . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 40
38a. Curved dark brown to black bar extending from nape through eye to interopercle . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis splendens
38b. Coloration not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 39
39a. Predorsal scales 17 to 22, extending anteriorly to point ranging from above midanterior
part of eye to just behind posterior nostrils; caudal fin rounded to truncate . . . . Pseudochromis sp. 2
39b. Predorsal scales 16 to 30, and 25 to 30 in specimens larger than 3 cm standard length,
extending anteriorly to point ranging from above midanterior part of eye to slightly anterior
of anterior nostrils (anterior to posterior nostrils in specimens larger than 3 cm standard
length); caudal fin emarginate in small (less than 3 cm) specimens, becoming strongly
emarginate to lunate in larger specimens . . . . . . . . . . . . . . . . . . Pseudochromis polynemus
40a. Pectoral fins black; well-developed denticles on outer ceratobranchial-1 gill rakers mainly
confined to raker tips; 3 or 4 (usually 4) dorsal-fin pterygiophores inserting anterior to
neural spine 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 41
40b. Combination of first 2 characters not as above; 3 dorsal-fin pterygiophores inserting
anterior to neural spine 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 42
41a. Pectoral-fin rays 17 to 19 (usually 18 or 19); horizontal scale rows above anal-fin origin to
anterior lateral line 14 to 17; circumpeduncular scales 18 to 21 . . . . . . . . Pseudochromis paranox
41b. Pectoral-fin rays 16 or 17; horizontal scale rows above anal-fin origin to anterior lateral
line 12 or 13; circumpeduncular scales 16 . . . . . . . . . . . . . . . . . . . . Pseudochromis sp. 3
42a. Palatine tooth patches directed strongly inward behind posterolateral arms of vomerine
tooth patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 43
42b. Palatine tooth patches more or less contiguous with posterolateral arms of vomerine tooth
patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 48
43a. Upper part of body with a dark longitudinal stripe . . . . . . . . . . . . . . . . . . . . . . . . . ® 44

43b. No dark stripe on upper part of body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 45
44a. Dark stripe on upper part of body extends below anterior lateral line to upper caudal-fin
rays; scales in lateral series 33 or 34; scales between lateral lines 3; circumpeduncular
scales 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis colei
44b. Dark stripe on upper part of body crosses middle of anterior lateral line to join basal stripe
on midposterior part of dorsal fin; scales in lateral series 38 to 42; scales between lateral
lines 4 to 6; circumpeduncular scales 20 to 24 . . . . . . . . . . . . . Pseudochromis perspicillatus
45a. No prominent dark spot in front of each anterior nostril; posterodorsal corner of operculum
without dark spot; caudal fin rounded . . . . . . . . . . . . . . . . Pseudochromis quinquedentatus
45b. A prominent dark spot in front of each anterior nostril; indistinct to distinct grey to black
spot present on posterodorsal corner of operculum; caudal fin rounded, becoming strongly
emarginate in large specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 46
46a. Anal fin with broad dark distal stripe; pelvic fins pale, with anterior margin broadly dark;
pale bar extending from behind eye to posterior edge of upper jaw . . . . . . Pseudochromis steenei
46b. Coloration not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 47
47a. Dark spot on operculum indistinct and small, smaller than pupil; preopercle edge not dark
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis howsoni
47b. Dark spot on operculum distinct and large, approximately equal to eye size; preopercle
edge dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis moorei
48a. Circumpeduncular scales 16 to 23 (rarely fewer than 18) . . . . . . . . . . . . . . . . . . . . . ® 49

48b. Circumpeduncular scales 15 to 18 (usually 16) . . . . . . . . . . . . . . . . . . . . . . . . . . ® 56
49a. Segmented dorsal-fin rays 23 to 25 (usually 24) . . . . . . . . . . . . . . . . . . . . . . . . . . ® 50

49b. Segmented dorsal-fin rays 25 to 31 (usually 26 to 30) . . . . . . . . . . . . . . . . . . . . . . . ® 52
2564 Bony Fishes
50a. Anal-fin spines slender and wealy pungent to flexible, the second spine about as stout as
the third . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis flammicauda
50b. Anal-fin spines relatively stout and pungent, the second spine much stouter than the third . . . . ® 51
51a. Segmented anal-fin rays 12 or 13 (usually 13); scales in lateral series 31 to 36; predorsal
scales 18 to 24; scales of body behind pectoral-fin base each with a large pale yellow to
pale brown central spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudochromis sp. 4
51b. Segmented anal-fin rays 13 or 14 (usually 14); scales in lateral series 36 to 39; predorsal
scales 14 to 18; no large pale spots on body scales . . . . . . . . . . . . Pseudochromis ransonneti
52a. Body with 8 to 15 brown (black in life) straight-edged stripes . . . . . . . . . Pseudochromis cometes
52b. If dark stripes present on body, these consisting of closely spaced spots so that stripes
wider at base of each scale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 53
53a. Well-developed denticles on ceratobranchial-1 outer rakers arranged in 2 rows running

most of raker lengths; dorsal and anal fins usually with weakly to well-developed scaly
sheaths present on at least posterior part of fins . . . . . . . . . . . . . . . . Pseudochromis fuscus
53b. Well-developed denticles on ceratobranchial-1 outer rakers mainly confined to raker tips;
dorsal and anal fins without scaly sheaths . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 54
54a. Segmented anal-fin rays 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis pictus

54b. Segmented anal-fin rays 13 to 15 (usually 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 55
55a. Second anal-fin spine much stouter than the third . . . . . . . . . . . . . Pseudochromis bitaeniatus
55b. Second anal-fin spine about as stout as the third . . . . . . . . . . . . . . . Pseudochromis wilsoni
56a. Segmented dorsal-fin rays 21 to 23 (usually 22); segmented anal-fin rays 12 to 14 (rarely
12 or 14); 2 epurals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 57
56b. Combination of fin-ray counts not as above; 3 epurals . . . . . . . . . . . . . . . . . . . . . . . ® 60
57a. Males with dark grey to black horseshoe-shaped mark on caudal fin; females with
conspicuous dark grey to black spot covering posttemporal pores (immediately above gill
opening); scales in lateral series 27 to 34 (usually 29 to 32); predorsal scales 10 to 15
(usually 11 to 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis tapeinosoma
57b. Males without dark horseshoe-shaped mark on caudal fin; females with inconspicuous
grey to dusky grey spot covering posttemporal pores; scales in lateral series 29 to 40
(usually 30 to 38); predorsal scales 11 to 19 (usually 13 to 16) . . . . . . . . . . . . . . . . . . ® 58
58a. Scales in lateral series 33 to 40 (usually 35 to 38); anterior lateral-line scales 28 to 34

(usually 29 to 33); scales below anterior lateral line 11 to 14 (usually 12 or 13) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis coccinicauda
58b. Scales in lateral series 29 to 37 (usually 31 to 35); anterior lateral-line scales 23 to 31
(usually 24 to 29); scales below anterior lateral line 9 to 13 (usually 10 to 12) . . . . . . . . . . ® 59
59a. Males with lower part of head and breast and scales of ascending portion of anterior lateral
line and of scale row immediately below horizontal portion of anterior lateral line abruptly
yellowish brown to bright yellow (pale brown to brown in preservative), the remainder of
head and body dark grey to black (dark brown to black in preservative); scales in lateral
series 30 to 37 (usually 31 to 36) . . . . . . . . . . . . . . . . . . . . . Pseudochromis cyanotaenia
59b. Males dark grey to black (dark brown to black in preservative), with lower part of head and
body in front of anal fin yellowish (pale brown to brown in preservative); scales in lateral
series 29 to 35 (usually 30 to 33) . . . . . . . . . . . . . . . . . . . . . . . . . . Peudochromis sp. 5
60a. Segmented dorsal-fin rays 25 to 27 (usually 26), with only the last 2 to 9 rays branched . . . . . ® 61
60b. Character combination not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 62
61a. Body more or less uniformly pale to dark; horizontal scales above anal-fin origin to
dorsal-fin base 14 to 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis elongatus
61b. Body pale with 5 to 8 dark stripes; horizontal scales above anal-fin origin 11 or 12 . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis striatus
62a. Fin spines weakly pungent to flexible; second anal-fin spine about as stout as the third;
lower lip varying from incomplete with weak symphysial interruption to complete . . . . . . . . ® 63
62b. Fin spines stout; second anal-fin spine slightly to much stouter than the third; lower lip
incomplete . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 65
63a. Anterior lateral-line scales 25 or 26; anterior lateral line terminating beneath segmented
dorsal-fin ray 15 or 16; 4 consecutive dorsal-fin pterygiophores inserting in a 1:1 associa-
tion with interneural spaces immediately behind neural spine 4 . . . . . . . Pseudochromis kolythrus
63b. Anterior lateral-line scales 26 to 34 (usually 28 to 31); anterior lateral line terminating
beneath segmented dorsal-fin ray 17 to 25 (usually 19 to 24); 1 to 4 (usually 1 to 3)
consecutive dorsal-fin pterygiophores inserting in a 1:1 association with interneural
spaces immediately behind neural spine 4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 64
64a. Dorsoanterior part of caudal peduncle usually with indistinct to distinct grey to black spot;
1 or 2 (modally 1) consecutive dorsal-fin pterygiophores inserting in a 1:1 association with
interneural spaces immediately behind neural spine 4; pectoral-fin rays 18 to 20 . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis jamesi
64b. Coloration not as above; 2 or 3 (modally 3) consecutive dorsal-fin pterygiophores inserting
in a 1:1 association with interneural spaces immediately behind neural spine 4; pectoral-
fin rays 16 to 19 (usually 17 or 18) . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis luteus
65a. Third anal-fin spine length 10 to 14% standard length; distance from dorsal-fin origin to
pelvic-fin origin 30 to 35% standard length; scales in lateral series 30 to 34 . . . . Pseudochromis fowleri
65b. Third anal-fin spine length 6 to 9% standard length; distance from dorsal-fin origin to
pelvic-fin origin 23 to 32% standard length; scales in lateral series 33 to 42 . . . . . . . . . . . . ® 66
66a. Segmented anal-fin rays 16; scales below anterior lateral line 10 to 12 (usually 10 or 11)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis flavopunctatus
66b. Segmented anal-fin rays 11 to 15; scales below anterior lateral line 11 to 15 (usually 12
to 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 67
67a. Caudal fin rounded in small (less than 3.5 cm) specimens, becoming pointed (rounded
with middle rays produced) in larger specimens; in preservative, predorsal contour and
nape pale pinkish brown; dorsal contour of body and caudal peduncle with dark, grey-
brown reticulation surrounding pale spots . . . . . . . . . . . . . . . . . Pseudochromis reticulatus
67b. Caudal fin rounded, truncate, or emarginate; coloration not as above . . . . . . . . . . . . . . ® 68
68a. Segmented anal-fin rays 11 to 14 (usually 13) . . . . . . . . . . . . . Pseudochromis marshallensis

68b. Segmented anal-fin rays 13 to 15 (usually 14 or 15) . . . . . . . . . . . . . . . . . . . . . . . . ® 69
69a. Segmented dorsal-fin rays 22 to 25 (usually 24) . . . . . . . . . . . . Pseudochromis andamanensis

69b. Segmented dorsal-fin rays 25 to 27 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 70
70a. No dark spot on axil of pectoral fins; segmented dorsal-fin rays 25 . . . . . . . . Pseudochromis litus
70b. Dark spot on axil of pectoral fins; segmented dorsal-fin rays usually 26 or 27 (rarely 25) . . . . . ® 71
2566 Bony Fishes
71a. Head and anterior part of body bright golden-orange (yellowish brown in preservative),
becoming bluish grey posteriorly, with caudal fin bluish grey basally (brown in preserv-
ative); segmented dorsal-fin rays 27; distance from anal-fin origin to middle dorsal-fin ray
(spines included in enumerating middle ray) 27 to 29% standard length; caudal peduncle
depth (measured between bases of last dorsal- and anal-fin rays) 16 to 17% standard
length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudochromis aurulentus
71b. Head and body bluish grey (brown in preservative), paler ventrally, with caudal fin, caudal
peduncle, and posterior part of body behind middle of anal fin abruptly bright yellow (pale
brown in preservative); segmented dorsal-fin rays 25 to 27 (usually 26); distance from
anal-fin origin to middle dorsal-fin ray (spines included in enumerating middle ray) 22 to
27% standard length; caudal peduncle depth (measured between bases of last dorsal-
and anal-fin rays) 14 to 16% standard length . . . . . . . . . . . . . . . . . . . Pseudochromis pylei

0 Assiculus punctatus Richardson, 1846
Blennodesmus scapularis Günther, 1871
Congrogadus amplimaculatus (Winterbottom, 1980)
Congrogadus heirichthys Jordan and Richardson, 1908
Congrogadus malayanus (Weber, 1909)
0
Congrogadus spinifer (Borodin, 1933)
Congrogadus subducens (Richardson, 1843)
0 Cypho purpurascens (De Vis, 1884)
Cypho sp.
Haliophis aethiopus Winterbottom, 1985
0
Labracinus atrofasciatus (Herre, 1933)
Labracinus cyclophthalmus (Müller and Troschel, 1849)
0
Ogilbyina novaehollandiae (Steindachner, 1880)
Ogilbyina queenslandiae (Saville-Kent, 1893)
Ogilbyina salvati (Plessis and Fourmanoir, 1966)
Pseudochromis andamanensis Lubbock, 1980
Pseudochromis aurifrons Lubbock, 1980
Pseudochromis aurulentus Gill and Randall, 1998
Pseudochromis bitaeniatus (Fowler, 1931)
Pseudochromis coccinicauda (Tickell, 1888)
Pseudochromis colei Herre, 1933
0
Pseudochromis cometes Gill and Randall, 1998
Pseudochromis cyanotaenia Bleeker, 1857
Pseudochromis diadema Lubbock and Randall, 1978
Pseudochromis elongatus Lubbock, 1980
Pseudochromis ephippiatus Gill, Pyle, and Earle, 1996
Pseudochromis flammicauda Lubbock and Goldman, 1976
Pseudochromis flavopunctatus Gill and Randall, 1998
0
Pseudochromis fowleri Herre, 1934
Pseudochromis fuscus Müller and Troschel, 1849
Pseudochromis howsoni Allen, 1995
Pseudochromis kolythrus Gill and Winterbottom, 1993
Pseudochromis litus Gill and Randall, 1988
Pseudochromis luteus Aoyagi, 1943
Pseudochromis marginatus Lubbock, 1980
Pseudochromis marshallensis Schultz, 1953
Pseudochromis moorei Fowler, 1931
Pseudochromis paccagnellae Axelrod, 1973
Pseudochromis paranox Lubbock and Goldman, 1976
Pseudochromis perspicillatus Günther, 1862
0
Pseudochromis pictus Gill and Randall, 1998
0
Pseudochromis polynemus Fowler, 1931
Pseudochromis porphyreus Lubbock and Goldman, 1974
Pseudochromis pylei Randall and McCosker, 1989
Pseudochromis quinquedentatus McCulloch
Pseudochromis ransonneti Steindachner, 1870
Pseudochromis reticulatus Gill and Woodland, 1992
Pseudochromis splendens Fowler, 1931
Pseudochromis steenei Gill and Randall, 1992
Pseudochromis striatus Gill, Shao, and Chen, 1995
Pseudochromis tapeinosoma Bleeker, 1853
Pseudochromis veliferus Lubbock, 1980
Pseudochromis wilsoni (Whitley, 1929)
Pseudochromis sp. 1
Pseudochromis sp. 2
Pseudochromis sp. 3
Pseudochromis sp. 4
Pseudochromis sp. 5
Pseudoplesiops annae (Weber, 1913)
Pseudoplesiops collare Gill, Randall, and Edwards, 1991
Pseudoplesiops howensis Allen, 1987
Pseudoplesiops knighti Allen, 1987
Pseudoplesiops multisquamatus Allen, 1987
Pseudoplesiops revellei Schultz, 1953
0
Pseudoplesiops rosae Schultz, 1943
Pseudoplesiops typus Bleeker, 1858
Pseudoplesiops sp. 1
References
Fowler, H.W. 1931. Contributions to the biology of the Philippine Archipelago and adjacent regions. The fishes of the
families Pseudochromidae, Lobotidae, Pempheridae, Priacanthidae, Lutjanidae, Pomadasyidae, and
Theraponidae, collected by the United States Bureau of Fisheries Steamer “Albatross”, chiefly in Philippine seas
and adjacent waters. Bull. U.S. Natl. Mus., 100(11):1-388.
Wintersbottom, R. 1986. Revision and vicariance biogeography of the subfamily Congrogadinae (Pisces: Perciformes:
Pseudochromidae). Indo-Pac. Fishes, (9):34 p.
2568 Bony Fishes
Assiculus punctatus Richardson, 1846

Frequent synonyms / misidentifications: Pseudochromis punctatus (Richardson, 1846) / None.
FAO names: En - Bluespotted dottyback.
Diagnostic characters: Body moderately deep; distance from dorsal-fin origin to pelvic-fin origin 29 to
34% standard length. Lower lip uninterrupted at symphysis. Vomerine teeth relatively large, arranged
in a chevron. Total gill rakers on first gill arch 13 to 17, of which 4 to 6 on upper limb, and 9 to 11 on lower
limb. Dorsal fin with III spines and 23 (rarely 22 or 24) mostly unbranched, segmented rays. Anal
fin with III spines and 13 (rarely 12 or 14) mostly unbranched, segmented rays. Caudal fin rounded,
the upper part with 5 (rarely 4) procurrent rays and 9 principal rays, the lower part with 4 (rarely 5)
procurrent rays and 8 principal rays. Pectoral fins with 17 or 18 (rarely 15 or 16) segmented rays. Pelvic
fins with I spine and 5 branched, segmented rays. Lateral line disjunct, consisting of an anterodorsal
series of 33 to 43 (usually 36 to 40) tubed scales extending from gill opening, and a peduncular series
of 6 to 9 (rarely 5 or 10) tubed scales. Circumpeduncular scales 17 to 21 (usually 19 or 20). Dorsal and
anal fins without distinct scaly sheaths. Vertebrae 11+17 (rarely 11+18); supraneural (predorsal) bones
either 2, or 3 with the third a rudiment. Colour: head, body, and fins either uniformly grey to dark grey or
brown to olive, sometimes with ventral part of head and breast reddish brown to bright yellow; small, bright
blue spots on head and body; dorsal, anal, caudal, and pelvic fins with bright blue spots or bright blue
irregular stripes and short streaks; pectoral fins clear.
Habitat, biology, and fisheries: Inhabits coastal reefs and weedbed areas, often in relatively silty waters,
at depths ranging to 30 m. A cryptic reef species, usually found in association with highly eroded limestone
reef and rocks. Of considerable potential as an aquarium fish.
Australia, from Shark Bay to
the Gulf of Carpentaria.
Congrogadus subducens (Richardson, 1843)

FAO names: En - Carpet eelblenny.
Diagnostic characters: Body elongate and laterally compressed; body depth at anal-fin origin 7 to 16%
standard length. Dorsal fin without spines, with 68 to 76 branched, segmented rays. Anal fin without
spines, with 57 to 66 branched, segmented rays. Caudal fin confluent with dorsal and anal fins, with 5
branched rays in upper part, and 5 branched rays in lower part of fin; no unbranched rays. Pectoral fins
with 9 to 11 segmented rays. Pelvic fins absent. Lateral line composed of 42 to 68 pored scales extending
from gill opening. Cheek, opercle, and body covered with small, elliptical cycloid scales. Colour: head and
body variegated green or reddish brown to brown, with irregular pale spots (up to eye diameter in size);
pale spots sometimes forming reticulations or bands; large ocellated dark spot usually present on
opercle; dorsal, anal, and caudal fins similar to body coloration; pectoral fins clear.
Habitat, biology, and fisheries: Cryptic inhabitant of shallow coral and rock reefs, often in silty areas
adjacent to seagrass beds. Sometimes enters the aquarium fish trade.
Distribution: Nicobar Islands
to southern coast of
Queensland, Australia, north
to the Ryukyu Islands and
south to the central coast of
Western Australia.
2570 Bony Fishes
Cypho purpurascens (De Vis, 1884)

Frequent synonyms / misidentifications: Pseudochromis mccullochi Myers, 1932; P. mccullochi
perpulcher Whitley, 1959 / None.
FAO names: En - Obliquelined dottyback.
Diagnostic characters: Body moderately slender; distance from dorsal-fin origin to pelvic-fin origin 27 to
30% standard length. Lower lip either uninterrupted at symphysis or with weak interruption only.
Vomerine teeth relatively large, arranged in a chevron. Total gill rakers on first gill arch 14 to 19 (usually
15 or 16), of which 4 to 7 (usually 4 or 5) on upper limb, and 10 to 13 (usually 11 or 12) on lower limb.
Dorsal fin with III slender spines and 23 (rarely 22 or 24) segmented rays, with at least some anterior
rays unbranched. Anal fin with III slender spines and 14 (rarely 13 or 15) segmented rays. Caudal
fin rounded, the upper part with 6 (rarely 5 or 7) procurrent rays and 9 principal rays, the lower part
with 5 or 6 (rarely 4) procurrent rays and 8 principal rays. Pectoral fins with 17 to 19 segmented rays.
Pelvic fins with I spine and 5 branched, segmented rays. Lateral line disjunct, consisting of an
anterodorsal series of 23 to 32 (usually 25 to 29) tubed scales extending from gill opening, and a
peduncular series of 3 to 9 (usually 5 to 7) tubed scales. Circumpeduncular scales 16 (rarely 17).
Dorsal and anal fins without distinct scaly sheaths. Vertebrae 10+16; supraneural (predorsal) bones
3, of approximately equal size. Colour: head and body either bright orange to bright red (males) or brown
to olive with large pink to red area above anterior part of anal fin (females); scattered pink to bright green
spots on top of head and snout; mauve to bright blue line extending around posteroventral edge of eye;
cheek and operculum with scattered mauve to dark blue bars and irregular markings; females usually with
large bright yellow spot on cheek behind eye; blue to magenta or black bars following oblique scale
rows on anterior part of body, these becoming broken and indistinct posteriorly; dorsal, anal, and caudal
fins similar to adjacent body coloration, often adorned with blue, red, or brown stipes and/or spots; males
sometimes with 1 to 3 large, gold-edged black spots on midanterior part of dorsal fin.
Habitat, biology, and fisheries: Cryptic inhabitant of coral and rock reefs, occurring in tidal pools, lagoonal
reefs, and reef slopes to depths of at least 40 m. Sometimes enters the aquarium fish trade.
Distribution: Southwest
Pacific, from the Great Barrier
Reef and Papua New Guinea
east to Tonga.
Labracinus cyclophthalmus (Müller and Troschel, 1849)

Frequent synonyms / misidentifications: Cichlops melanotaenia Bleeker, 1852; C. spilopterus Bleeker,
1853; C. hellmuthii Bleeker, 1854; C. japonicus Gill, 1859; C. trispilos Bleeker, 1855; Dampieria
melanostigma Fowler, 1931; D. ocellifera Fowler, 1946 (including various combinations of these in
Cichlops, Dampieria, and Labracinus) / Labracinus lineatus (Castenau, 1875); Pseudochromis fuscus
Müller and Troschel, 1849.
FAO names: En - Red dottyback.
Diagnostic characters: Body moderately deep to deep; distance from dorsal-fin origin to pelvic-fin origin
31 to 38% standard length. Lower lip uninterrupted at symphysis. Vomerine teeth relatively large,
arranged in a chevron. Total gill rakers on first gill arch 18 to 22 (rarely 17 or 23), of which 6 to 9 (rarely
10) on upper limb, and 11 to 14 (rarely 15) on lower limb. Dorsal fin with II slender spines and 25 (rarely
24 or 26) mostly branched, segmented rays. Anal fin with III spines and 14 (rarely 15) branched,
segmented rays. Caudal fin rounded, the upper part with 5 or 6 (usually 6) procurrent rays and 9 principal
rays, the lower part with 5 (rarely 4 or 6) procurrent rays and 8 principal rays. Pectoral fins with 18 or 19
(rarely 17 or 20) segmented rays. Pelvic fins with I spine and 5 branched, segmented rays. Lateral
line disjunct, consisting of an anterodorsal series of 43 to 62 (usually 48 to 62) tubed scales
extending from gill opening, and a peduncular series of 12 to 24 (usually 18 to 22) tubed scales.
Circumpeduncular scales 24 to 30 (usually 26 to 28). Dorsal and anal fins with distinct scaly sheaths.
Vertebrae 11+17; supraneural (predorsal) bones 3, of approximately equal size. Colour: head and body
varying from pale grey to dark grey, olive, or bright red, usually darker dorsally; cheeks and preorbital area
usually with blue to dark grey oblique bars; scales of dorsoanterior part of body each with a bluish to black
spot; dark spots sometimes coalescing to form stripes along dorsal-fin base or along alternate scale rows
on posterior part of body; upper part of body sometimes with a series of white to bright yellow, short, oblique
bars; dorsal fin greyish to dark grey or bright red, usually with outer margin blue to dark grey, and with a
series of reddish to black rows of spots anteriorly, which align posteriorly to form stripes; females often
with spots forming comma-shaped markings; males sometimes with 1 to several large, dark grey to black,
horizontally elongate blotches on anterior part of dorsal fin; anal fin brownish or bluish to dark grey or red,
usually with 3 to 12 pale stripes and blue to dark grey outer margin; caudal fin bluish to bright red or black
with blue to black margin.
Habitat, biology, and fisheries: Inhabits
rock and coral reefs in various habitats, from
tidal pools and lagoonal reefs to reef slopes,
often in relatively silty areas, to depths of at
least 20 m. Frequently enters the aquarium
fish trade, and has been bred in captivity.
Distribution: Widely distributed through-
out the Indo-Malaysian Archipelago, from
southern Japan south to the Northwest Shelf
of Australia, and east to New Ireland, Papua
New Guinea.
2572 Bony Fishes
Ogilbyina queenslandiae (Saville-Kent, 1893)

Frequent synonyms / misidentifications: None / Dampieria longipinna Ogilby, 1908; Pseudochromis
novaehollandiae Steindachner, 1879; P. veliferus Lubbock, 1980.
FAO names: En - Queensland dottyback.
Diagnostic characters: Body moderately deep to deep; distance from dorsal-fin origin to pelvic-fin
origin 32 to 36% standard length. Lower lip uninterrupted at symphysis. Vomerine teeth small,
arranged in a broad, triangular patch. Total gill rakers on first gill arch 16 to 20 (usually 18 or 19), of
which 5 to 8 (usually 6 or 7) on upper limb, and 11 to 13 (usually 12) on lower limb. Dorsal fin with III
(rarely IV) slender spines and 32 to 37 (usually 34 to 36) mostly unbranched, segmented rays. Anal
fin with III slender spines and 18 to 21 (usually 19 or 20) segmented rays. Caudal fin usually rounded
to spade-shaped, often with posterior margin truncate or irregular, the upper part with 6 (rarely 7) procurrent
rays and 9 principal rays, the lower part with 5 (rarely 6) procurrent rays and 8 principal rays. Pectoral fins
with 18 to 20 (usually 19) segmented rays. Pelvic fins with I spine and 5 branched, segmented rays.
Lateral line disjunct, consisting of an anterodorsal series of 38 to 50 (usually 40 to 45) tubed scales
extending from gill opening, and a peduncular series of 4 to 18 (usually 10 to 14) tubed scales.
Circumpeduncular scales 24 to 26. Dorsal and anal fins with distinct scaly sheaths. Vertebrae 10+16;
supraneural (predorsal) bones 2. Colour: males with head and anterior part of body reddish, often with
short dark bars extending from beneath anterior part of dorsal fin, the posterior part of body olive to dark
grey; females brown to dark grey anteriorly, often with short dark bars extending from beneath anterior
part of dorsal fin, and reddish posteriorly, with area above anal fin bright yellow to orange; dorsal, anal,
and caudal fins similar to adjacent body coloration, often with blue or red stripes and/or spots, and irregular
yellow markings on caudal fin.
Habitat, biology, and fisheries: Inhabits rocky and coral reefs in various habitats, including tidal pools,
lagoon areas, and reef slopes, at depths ranging to 15 m. Frequently enters the aquarium fish trade, and
has been bred in captivity.
Distribution: Great Barrier
Reef and adjacent coast of
Queensland, Australia.
Pseudochromis cyanotaenia Bleeker, 1857

Frequent synonyms / misidentifications: Pseudochromis kikaii Aoyagi, 1941 / Pseudochromis
tapeinosoma Bleeker, 1853.
FAO names: En - Bluebarred dottyback.
Diagnostic characters: Body relatively slender; distance from dorsal-fin origin to pelvic-fin origin 25 to
28% standard length. Lower lip either uninterrupted at symphysis, or with weak interruption only.
Vomerine teeth relatively large, arranged in a chevron. Total gill rakers on first gill arch 13 to 16, of which
3 to 5 on upper limb, and 10 or 11 (rarely 9 or 12) on lower limb. Dorsal fin with III slender spines
(anterior spine easily overlooked) and 22 (rarely 21 or 23) mostly branched, segmented rays. Anal
fin with III slender spines (anterior spine easily overlooked) and 13 (rarely 12) mostly branched,
segmented rays. Caudal fin rounded, the upper part with 7 or 8 (rarely 6) procurrent rays and 9 principal
rays, the lower part with 6 or 7 (rarely 8) procurrent rays and 8 principal rays. Pectoral fins with 17 to 19
(rarely 16 or 20) segmented rays. Pelvic fins with I spine and 5 branched, segmented rays. Lateral
line disjunct, consisting of an anterodorsal series of 24 to 31 (usually 25 to 29) tubed scales
extending from gill opening, and a peduncular series of 6 to 11 tubed scales. Circumpeduncular scales
16. Dorsal and anal fins without distinct scaly sheaths. Vertebrae 10+16; supraneural (predorsal) bones
3, of approximately equal size. Colour: females and juveniles with head and body brownish to greyish
brown, paler ventrally on head and breast, sometimes becoming reddish on caudal peduncle; dorsal and
anal fins greyish basally and clear on outer parts; caudal fin yellowish grey to bright orange-red basally,
becoming bright yellow to clear posteriorly; males with lower part of head and lower part of body
anterior to about middle of pectoral fins yellowish grey to bright yellow; scales of ascending portion
of anterior lateral line and of scale row beneath horizontal portion of anterior lateral line yellowish
grey to bright yellow; remainder of head and body dark grey to black, often with bluish or greenish
sheen, and sometimes with several short, bluish grey to bright blue bars above anal fin and on caudal
peduncle; dorsal, anal, and caudal fins dark grey to black basally, with remainder of fin pale blue to clear.
Habitat, biology, and fisheries: Inhabits various reef habitats, including tidal pools, lagoonal patch reefs,
and reef slopes, at depths
ranging to 30 m; usually most
abundant in high current or
surgeareas. Frequentlyenters
the aquarium fish trade, and
has been bred in captivity.
Distribution: P e n i n s u l a r
Malaysia north to Hong Kong
and the Ryukyu Islands, south
to Western Australia, and east
to Vanuatu and the Gilbert
Islands.
2574 Bony Fishes
Pseudochromis fuscus (Müller and Troschel, 1849)

Frequent synonyms / misidentifications: Pseudochromis aureus Seale, 1910; P. xanthochir Bleeker,
1855 / Labracinus cyclophthalmus (Müller and Troschel, 1849); L. lineatus (Castenau, 1875).
FAO names: En - Brown dottyback.
30 to 36% standard length. Lower lip interrupted at symphysis. Total gill rakers on first gill arch 17 to
21 (rarely 16), of which 5 to 7 (rarely 8 or 9) on upper limb, and 12 to 14 (rarely 11 or 15) on lower limb.
Dorsal fin with III relatively stout spines and 25 to 29 (usually 26 or 27) segmented rays, usually
with at least some anterior rays unbranched. Anal fin with III stout spines and 14 (rarely 13 or 15)
segmented rays. Caudal fin rounded to truncate or emarginate, the upper part with 6 or 7 (rarely 5)
procurrent rays and 9 principal rays, the lower part with 5 or 6 (rarely 4 or 7) procurrent rays and 8 principal
rays. Pectoral fins with 18 or 19 (rarely 17 or 20) segmented rays. Pelvic fins with I spine and 5 branched,
segmented rays. Lateral line disjunct, consisting of an anterodorsal series of 17 to 36 (usually 25
to 33) tubed scales extending from gill opening, and a peduncular series of 4 to 14 (usually 6 to 11)
tubed scales. Circumpeduncular scales 20 (rarely as few as 16 or as many as 23). Dorsal and anal
fins with weakly to strongly developed scaly sheaths. Vertebrae 10+16; supraneural (predorsal)
bones 3. Colour: bright yellow to dark grey, 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.
Habitat, biology, and fisheries: Known from a variety of reef habitats, from tidal pools and shallow lagoon
reefs, to outer reef slopes, at depths ranging to 30 m. Usually found in association with branching corals
(e.g. Acropora and Pocillopora). Sometimes enters the aquarium fish trade.
Distribution: Sri Lanka east
to Vanuatu, north to at least
Hong Kong and the Ryukyu
Islands, and south to the
central coast of Western
Australia and the southern
Great Barrier Reef.
Pseudochromis polynemus Fowler, 1931

FAO names: En - Threadfinned dottyback.
32 to 36% standard length. Lower lip interrupted at symphysis. Total gill rakers on first gill arch 17 or
18, of which 4 or 5 on upper limb, and 12 or 13 on lower limb. Dorsal fin with III relatively stout spines
and 27 to 29 mostly unbranched, segmented rays. Anal fin with III stout spines and 14 or 15
segmented rays. Caudal fin emarginate to lunate, the upper part with 6 procurrent rays and 9 principal
rays, the lower part with 6 procurrent rays and 8 principal rays. Pectoral fins with 16 to 18 segmented rays.
Pelvic fins with I spine and 5 branched, segmented rays. Lateral line disjunct, consisting of an
anterodorsal series of 27 to 32 tubed scales extending from gill opening, and a peduncular series
of 8 to 17 tubed scales. Circumpeduncular scales 16. Dorsal and anal fins usually with indistinct scaly
sheaths posteriorly. Vertebrae 10+16; supraneural (predorsal) bones 3, of approximately equal size.
Colour: head and body brownish to bluish grey, paler ventrally, with scales of cheek, operculum
and body excluding dorsal contour of nape each with a large, central dull yellow to bright orange
spot; spots becoming paler ventrally on body and tending to coalesce to form stripes; pale yellow to bright
yellow ring around orbit, ending ventrally in short broad bar, which extends behind and below
posterior edge of upper jaw; dorsal and anal fins grey with series of white to yellow spots, and posterior
part of fins abruptly white or clear; caudal fin bluish grey basally, becoming clear or whitish posteriorly;
pectoral fins clear; pelvic fins white to grey with large bright yellow to bright red spot basally.
Habitat, biology, and fisheries: Occurs around corals and rocks on reef slopes and dropoffs in depths
of 2 to 50 m, often in association with large sponges. Of considerable potential as an aquarium fish.
Distribution: Currently known
only from northern Sulawesi
and Belau.
2576 Bony Fishes
Pseudochromis porphyreus Lubbock and Goldman, 1974

FAO names: En - Strawberry dottyback.
32% standard length. Lower lip interrupted at symphysis. Total gill rakers on first gill arch 19 to 22
(rarely 18 or 23), of which 5 to 8 on upper limb, and 13 to 16 on lower limb. Dorsal fin with III spines and
22 (rarely 21) mostly branched, segmented rays. Anal fin with III spines and 12 (rarely 10 or 11)
mostly branched, segmented rays. Caudal fin truncate to emarginate, the upper part with 6 or 7 (usually
6) procurrent rays and 9 principal rays, the lower part with 6 (rarely 5 or 7) procurrent rays and 8 principal
rays. Pectoral fins with 16 to 18 (rarely 19) segmented rays. Pelvic fins with I spine and 5 branched,
segmented rays. Lateral line disjunct, consisting of an anterodorsal series of 19 to 28 (usually 21 to
25) tubed scales extending from gill opening, and a peduncular series of 0 to 10 tubed scales.
Circumpeduncular scales 16 (rarely 18). Dorsal and anal fins without distinct scaly sheaths.
Vertebrae 10+16; supraneural (predorsal) bones 3, of approximately equal size. Colour: head and body
reddish purple to magenta; basal parts of dorsal and anal fins pink to magenta, the outer parts clear;
basal and central part of caudal fin magenta, the remainder of fin clear, giving rounded appearance to fin.
Size: Maximum total length of captured specimens about 6.5 cm; aquarium specimens known to grow to
at least 8 cm total length.
Habitat, biology, and fisheries: Lives in small caves and among rubble and coral on reef slopes and
dropoffs at depths ranging from 3 to 65 m. This species and 2 of its close relatives (Pseudochromis diadema
and P. paccagnellae) are popular aquarium fishes.
Distribution: Central and
western Pacific, from the
R y uk y u Is la nds south to
eastern Indonesia, and east
to American Samoa.
Pseudoplesiops typus Bleeker, 1858

Frequent synonyms / misidentifications: Pseudoplesiops sargenti Schultz, 1953 / None.
FAO names: En - Ringeyed dottyback.
32% standard length. Lower lip uninterrupted at symphysis. Total gill rakers on first gill arch 11 to 16
(usually 12 to 14), of which 2 to 4 (usually 3 or 4) on upper limb, and 9 to 12 (usually 9 or 10) on lower
limb. Dorsal fin with II slender spines (easily overlooked) and 24 (rarely 25 or 26) mostly unbranched,
segmented rays. Anal fin with II or III spines (easily overlooked) and 14 to 16 (usually 15) mostly
unbranched, segmented rays. Caudal fin rounded, the upper part with 4 (rarely 5) procurrent rays and 9
principal rays, the lower part with 3 or 4 procurrent rays and 8 principal rays. Pectoral fins with 16 to 18
segmented rays. Pelvic fins with I spine and 4 unbranched, segmented rays. Lateral line represented
by a single tubed scale at gill opening, followed by an intermittent series of centrally pitted scales
extending beneath base of dorsal fin, and a second intermittent series of centrally pitted scales on caudal
peduncle; transverse scale rows from upper edge of gill opening to base of caudal fin 33 to 39.
Circumpeduncular scales 20 (rarely 19 or 22). Predorsal scales 10 to 16, extending anteriorly to
about midway between dorsal-fin origin and posterior edge of eye. Dorsal and anal fins without distinct
scaly sheaths. Posterior body scales ctenoid in small specimens, becoming cycloid in specimens
larger than about 3.5 cm standard length, with distinct centres and radii in all fields. Vertebrae 11+17
to 18; supraneural (predorsal) bones 3, of approximately equal size. Colour: head and body pale bluish
grey to pale yellow or bright pink; narrow red to dark grey or black ring around eye; dorsal, anal, and
caudal fins similar to body coloration, often with blue or red edging on fins; pectoral fins clear; pelvic fins
pale blue to white.
Habitat, biology, and fisheries: Cryptic inhabitant of coral reefs, in depths ranging from 1 to at least 30 m.
Of some potential as an aquarium fish.
Distribution: Indonesia and
the Philippines south to the
Northwest Shelf of Australia,
and east to the Marshall and
Solomon islands.
click for next page

2578 Bony Fishes
PLESIOPIDAE
Roundheads (prettyfins, longfins)
by R.D. Mooi
D iagnostic characters: Small to medium-sized fishes (3 to 30 cm total length); body elongate to oblong.
Snout short. Preopercular sensory canal open (except in Beliops and Acanthoplesiops), giving the
preopercle a “double-bordered” appearance (not always obvious externally). No opercular spines.
Notch in the posterolateral margin of branchiostegal membranes just dorsal to third branchiostegal
ray (not obvious in Calloplesiops). A single dorsal fin with IX to XXVI spines and 2 to 11 segmented rays,
some species with deeply incised fin membranes between the spines (not in Assessor or Callople-
siops); anal fin with III to XVI spines and 2 to 11 segmented rays; caudal fin usually rounded, sometimes
elongate to lanceolate or forked; pelvic fins with I spine and 2 or 4 segmented rays, the first ray
bifurcate, often elongate and sometimes considerably thickened; pectoral fins with 14 to 30 rays;
Branchiostegal rays 6. Scales often cycloid anteriorly and ctenoid posteriorly; scales in lateral series 23 to
more than 100; lateral line disjunct and in 2 or more parts, with a more anterior dorsal lateral line
running near dorsal-fin base, a posterior lateral line running midlaterally onto caudal peduncle, and
occasionally a ventral lateral line as well (except Steeneichthys with a single pored lateral-line scale and
the remainder almost wholly replaced by scales with sensory papillae, and Acanthoplesiops with a single
dorsal lateral line). Parasphenoid keel present; dorsal-fin pterygiophores articulate with spines via
closed bony ring (except in some species of Paraplesiops); each pelvic bone with a well-developed subpelvic
shelf as well as a subpelvic concavity posterior to subpelvic processes visible in ventral view; extensor proprius
pelvicus muscle inserts onto second to fourth pelvic-fin rays (except in the subfamily Acanthoclininae where
the muscle is absent); hypurals 1 and 2 (and sometimes parhypural) fused into autogenous plate, hypurals 3
and 4 fused to each other and to urostylar complex. Colour: variable, even changeable for individuals from
solid or spotted body colour to barred, but usually dark background colour on body (although Fraudella is
orange); dorsal-fin spines often with yellow or red tips.
incised membranes between dorsal-fin spines
disjunct or interrupted
lateral line, in 1, 2, or
sometimes 3 parts
“double-bordered” preopercle
1st pelvic-fin ray bifurcate and usually

notch in branchiostegal membrane elongate, sometimes thickened
Habitat, biology, and fisheries: On coral reefs and tide pools; occur near the surface in the surf zone to
a depth of 30 m, sometimes deeper. Some species mostly nocturnal, feeding on crustaceans and
molluscs. Can be schooling in caves and overhangs, in loose aggregations or schools around coral heads,
or mostly solitary. Eggs with hair-like filaments that either entangle with one another to form a mass or
adhere eggs to a hard surface; eggs are guarded by male in a crevice or cave, or are mouthbrooded. No
commercial catch, although some of the larger Australian species take baited line. Some species,
particularly the comet or marine betta (Calloplesiops), are popular in the aquarium trade and have been
successfully bred in captivity.
Remarks: Two of the listed species (Acanthoplesiops psilogaster and Beliops batanensis) have been
reported just north of the area at Batan Island, Philippines, but are included because they might occur.
Based on colour variation, some authors have suggested that there are 2 species of Calloplesiops:
C. altivelis (figured here) with larger spots which do not merge into longitudinal stripes on the dorsal and
anal fins, and C. argus with smaller spots which merge to produce longitudinal stripes on the dorsal and
anal fins. These colour morphs co-occur, and an indepth study has yet to be undertaken to determine
specific status; such information might be important for conservation of this popular aquarium fish.
Perciformes: Percoidei: Plesiopidae 2579

Pseudochromidae: superficially similar in having a disjunct lateral line or lateral line reduced to a single
anterior pored scale, some members with a reduced number of pelvic-fin rays and with pelvic fins frequently
elongate. But this family is markedly different in having 3 or fewer dorsal-fin spines, and these often weak;
21 or more segmented dorsal-fin rays and 13 or more segmented anal-fin rays.
Serranidae: 3 opercular spines; lateral line complete, except some members of the tribe Grammistini
(Pseudogramma, Aporops, Suttonia ). The latter have been misidentified as Plesiopidae, but differ in
having VIII or fewer dorsal-fin spines, short pelvic fins with I spine and 5 rays, 3 opercular spines, and 1
preopercular spine.
21 or more segmented dorsal-fin rays VIII or fewer dorsal-fin spines

III or fewer
dorsal-fin 3 opercular spines
spines
short
pelvic fin
13 or more segmented anal-fin rays
Pseudochromidae Serranidae (tribe Grammistini)
Callanthiidae: somewhat similar in having a dorsally placed lateral line which runs near base of the dorsal
fin and can continue along dorsal margin of caudal peduncle, with a series of posterior midlateral scales
with sensory papillae giving the appearance of a disjunct lateral line in 2 parts. Readily differentiated by
pelvic fins with I spine and 5 segmented rays which are not elongate; caudal fin lunulate; 1 or 2 opercular
spines; tubed lateral line short (Grammatonotus, 14 to 18 scales) or extending beyond dorsal-fin base onto
dorsal portion of caudal peduncle (Callanthias).
Opistognathidae: lateral line high on body, close to dorsal-fin base and ending below middle of fin, pelvic
fins with I spine and 5 segmented rays but outer 2 unbranched, hence giving appearance of 4 branched
rays only. Otherwise easily distinguished from Plesiopidae by the following: head and mouth large, with
maxilla extending well beyond eye, eyes relatively large and high on head, dorsal fin with weak spines.
dorsal lateral line onto caudal peduncle

1-2 opercle weak dorsal-fin spines
spines lunate truncate dorsal lateral line
caudal fin large eye
large mouth
pelvic fins with

I spine and 5 rays
Callanthiidae Opistognathidae
Key to the species of Plesiopidae occurring in the area

1a. Dorsal fin with more than XV spines; anal fin with more than VI spines . . . . . . . . . . . . . . ® 2
1b. Dorsal fin with fewer than XV spines; anal fin with III spines . . . . . . . . . . . . . . . . . . . ® 6
2580 Bony Fishes
2a. Three lateral lines with tubed scales: 1 dorsal line near base of dorsal fin, 1 midlateral,
and 1 ventral line near base of anal fin (Fig. 1); body pale to dark brown with a number
of narrow dark bars, opercle with a dark brown ocellus, head dark brown dorsally and
sharply delimited through the eye and bordered by a white stripe or merging with the pink
to red cheeks and throat, dorsal and anal fins red or brown with pale tips (Fig. 1) . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Belonepterygion fasciolatum
2b. One or 2 lateral lines with tubed scales: 1 dorsal line near base of dorsal fin and 1
midlateral line (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
dorsal midlateral
3 lateral lines
Fig. 1 Belonepterygion fasciolatum Fig. 2 possible positions of lateral line
3a. Two lateral lines with tubed scales, the dorsalmost extending from upper angle of opercle
to base of last dorsal-fin ray, the midlateral line extending forward from middle of
caudal-fin base to about level of anus; body scales with membranous central lobe
without cteni (Fig. 3a); no pale spot on pectoral-fin base; live coloration unknown (Fig. 4)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Beliops batanensis
3b. Single lateral line with tubed scales extending form upper angle of opercle to no more
than half-way along dorsal-fin base; body scales without central lobe or, if present,
supported by elongate cteni (Fig. 3b); pectoral-fin base with pale spot (Acanthoplesiops) . . . . . ® 4
central elongate
lobe cteni
a) b)
Fig. 3 body scales Fig. 4 Beliops batanensis
4a. Dark band of about eye diameter

originating on upper jaw, continuing
through eye to middle of spinous
dorsal fin, giving appearance of a
mask; most scales on posterior half
of body with a flap-like lobe sup-
ported by elongate cteni (Fig. 3b),
and at least some midlateral scales
bilobed; belly completely scaled; live
coloration unknown (Fig. 5) . . . . .
. . . . . . . . . . . . Acanthoplesiops echinatus
4b. Body uniformly dark laterally, with- Fig. 5 Acanthoplesiops echinatus
out dark stripe through eye; scales
on posterior half of body typically
ctenoid, without flap-like lobes; belly
completely scaled or anterior two-
thirds of belly unscaled . . . . . . . . . . . . ® 5
5a. Belly completely scaled; symphysis of lower jaw with terminal pair of pores (each dentary
with 4 pores) (Fig. 6a); live coloration unknown (Fig. 7) . . . . . . . . . . . . Acanthoplesiops hiatti
5b. Anterior two-thirds of belly unscaled; symphysis of lower jaw without terminal pair of
pores (each dentary with 3 pores) (Fig. 6b); body olive-grey, dorsal, anal, and pelvic fins
olive-grey with yellow-orange tips, caudal-fin grey with yellow-orange margin; colour in
ethanol similar to Acanthoplesiops hiatti . . . . . . . . . . . . . . . . . Acanthoplesiops psilogaster
4 pores 3 pores
Fig. 7 Acanthoplesiops hiatti

a) Acanthoplesiops hiatti b) Acanthoplesiops psilogaster
6a. Dorsal fin with IX spines; a single tu-
bed lateral-line scale at upper end of
operculum (Fig. 8) . . . . Steeneichthys plesiopsus
6b. Dorsal fin with XI to XIII spines; 2
disjunct lateral lines with tubed scales,
1 from upper end of operculum to last
dorsal-fin ray, the other midlateral on
caudal peduncle . . . . . . . . . . . . . . . . . ® 7
Fig. 8 Steeneichthys plesiopsus
7a. Head scaled forward to, but not onto, snout; maxilla scaled; caudal fin forked; vomer
toothless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Assessor) ® 8
7b. Head scaled only to occiput; maxilla unscaled; caudal fin rounded or elongate; vomer
toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
8a. Head, body, and all fins except pectoral fins dark blue; total gill rakers on first gill arch
33 to 36, of which 23 to 24 on lower limb (Fig. 9) . . . . . . . . . . . . . . . . . . Assessor macneilli
8b. Head, body, and all fins except pectoral fins bright yellow, with an orange stripe from eye to
opercular angle, dorsal and anal fins with submarginal orange stripe and black margin; total
gill rakers on first gill arch 23 to 27, of which 16 to 19 on lower limb (Fig. 10) . . . . Assessor flavissimus
Fig. 9 Assessor macneilli Fig. 10 Assessor flavissimus
9a. Preopercular margin serrated with numer-

ous teeth along its length; dorsal fin with
XIII spines (sometimes XII); teeth on
tongue; body orange (Fig. 11) . . . Fraudella carassiops
9b. Preopercular margin smooth, or at most
denticulate at angle (overlying skin some-
times fimbriate giving appearance of serra-
tion, but bone is smooth); dorsal fin with XII
or fewer spines; tongue without teeth in all
Fig. 11 Fraudella carassiops
remaining genera except Paraplesiops . . . . . . ® 10
2582 Bony Fishes
10a. Head, body, and fins (except pectoral fins) black with white spots; posterior rays of dorsal
fin with black ocellus bordered by yellow; dorsal-fin membranes weakly incised; no teeth
on tongue (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Calloplesiops altivelis
10b. Head, body, and fins often dark, but never black; posterior rays of dorsal fin never with
ocellus; membranes between dorsal-fin spines deeply incised; tongue with or without
teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
11a. Tongue with teeth; upper lateral-line scales 28 to 40; scales with normal circulae and
radiating lines in anterior field only (Fig. 13a); segmented dorsal-fin rays 9 or 10 (rarely 9)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .(Paraplesiops) ® 12
(Great Barrier Reef only)
11b. Tongue without teeth; fewer than 24 upper lateral-line scales; scales with distinctive
unmarked centres and radiating lines in all fields (Fig. 13b); segmented dorsal-fin rays 7
or 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Plesiops) ® 13
no centre unmarked centre
a) b)
Fig. 12 Calloplesiops altivelis Fig. 13
12a. Body dark brown with 8 to 10 narrow black vertical bands, wider than interspaces; head
with several blue spots smaller than pupil diameter on cheek and opercle, and a larger
blue ocellus of pupil diameter ventrally on operculum; upper lateral line with 28 to 32
tubed scales; 7 to 12 predorsal scales (Fig. 14) . . . . . . . . . . . . . . . . . . Paraplesiops poweri
12b. Body with 4 broad black bands, interspace bands white; head with numerous small
iridescent blue spots smaller than pupil diameter; small iridescent blue spots along bases
of dorsal and anal fins; pectoral and caudal fins yellow; upper lateral line with 34 to 40
tubed scales; 20 to 29 predorsal scales (Fig. 15) . . . . . . . . . . . . . . . . Paraplesiops bleekeri
Fig. 15 Paralesiops bleekeri

Fig. 14 Paralesiops poweri
13a. Dark blue ocellus of slightly less

than eye diameter on ventral por-
tion of opercle; head, body, and
caudal fin with small blue spots
(Fig. 16) . . . . . . . . . . Plesiops corallicola
(throughout area except far east, New Caledonia, and
Great Barrier Reef)
13b. No ocellus on opercle; occasion-
ally pale spots scattered on head,
but no spots on body or caudal fin . . . . ® 14
Fig. 16 Plesiops corallicola
14a. Dorsal fin with XI spines (very rarely X or XII, but always only 1 supernumerary spine
on first pterygiophore); total number of branches on 6 ventralmost pectoral-fin rays fewer
than 15, usually 12 (very rarely more than 15) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
14b. Dorsal fin with XII spines (very rarely XI or XIII, but always 2 supernumerary spines on
first pterygiophore); total number of branches on 6 ventralmost pectoral-fin rays usually
more than 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
15a. Pectoral fins with 19 to 24 rays; total gill rakers on first gill arch usually 11 or 12 (sometimes
15); 6 to 8 predorsal scale rows (very rarely 5) (Fig. 17) . . . . . . . . . . . . Plesiops coeruleolineatus
15b. Pectoral fins with 26 to 28 rays; total gill rakers on first gill arch 13 to 18; 5 or 6 predorsal
scale rows (usually 5) (Fig. 18) . . . . . . . . . . . . . . . . . . . . . . . . . . Plesiops polydactylus
Fig. 17 Plesiops coeruleolineatus Fig. 18 Plesiops polydactylus
16a. Pectoral fins with 25 to 27 rays; 4 to

6 predorsal scale rows (usually 5);
dark spot at dorsal angle of opercle
(Fig. 19) . . . . . . . . . . . . . Plesiops auritus
16b. Pectoral fins with 24 or fewer rays
(very rarely 25); 6 to 11 predorsal
scale rows (infrequently 6); no dark
spot at dorsal angle of opercle . . . . . . . ® 17 Fig. 19 Plesiops auritus
17a. Three or more posterior upper lateral-line scales with anterior pore not covered by
preceding scale (Fig. 20); pectoral-fin rays 17 to 19 (sometimes 20); number of dentary
pores on 1 side usually fewer than 20, always fewer than 25 . . . . . . . . . . . . . . . . . . . ® 18
17b. Three or fewer upper lateral-line scales with anterior pore not covered by preceding
scale; pectoral-fin rays usually 20 or more (sometimes 19); usually more than 20 dentary
pores on 1 side when greater than 3.5 cm standard length . . . . . . . . . . . . . . . . . . . . ® 20
18a. Anterior part of head densely speckled with small brown spots; sensory pores on head
sparse and large, especially those of the interorbital (about 1 mm); only 5 large pores on
each dentary (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plesiops facicavus
18b. Anterior part of head not speckled; sensory pores relatively numerous and small,
especially in the interorbital area; 6 or more dentary pores (very rarely 5) . . . . . . . . . . . . ® 19
pore covered pore uncovered
Fig. 20 lateral-line scales Fig. 21 Plesiops facicavus

2584 Bony Fishes
19a. Predorsal scale rows 9 to 11 (usually 10

or 11); cheek scale rows 4 or 5 (usually enlarged
5); 5 to 9 sensory pores on each dentary small
pore pore
(rarely more than 7); anteroventral pre-
opercular sensory pore enlarged
(Figs 22a, 23) . . . . . . . . . . . . Plesiops gracilis
19b. Predorsal scale rows 6 to 8; cheek scale
rows 3 or 4; 5 to 23 sensory pores on each
dentary (usually more than 7); anteroven-
tral preopercular sensory pore not en-
larged (Figs 22b, 24) . . . . . . Plesiops cephalotaenia a) Plesiops b) Plesiops
gracilis cephalotaenia
Fig. 23 Plesiops gracilis Fig. 24 Plesiops cephalotaenia
20a. Nape with a mottled or dendritic pattern; body usually pale with dark spots, often
congregated to form 4 to 6 bands, 4 on body and 2 on caudal peduncle (body sometimes
dark); tips of dorsal-fin spines yellow; pectoral fins with 19 to 21 rays (usually 20);
predorsal scale rows 8 to 11 (usually 9 or 10) (Fig. 25) . . . . . . . . . . . . . Plesiops oxycephalus
20b. Nape without a mottled or dendritic pattern; body usually dark without bars; tips of
dorsal-fin spines red or pale, not yellow; pectoral fins with 20 to 24 rays (rarely 19 or 25),
usually 22 or more; predorsal scale rows 6 to 8 (usually 7) . . . . . . . . . . . . . . . . . . . . ® 21
21a. Tips of dorsal-fin spines red (except in specimens 3 cm standard length); total number
of branches on ventral 6 pectoral-fin rays rarely more than 20 (usually 12); branches on
ventralmost pectoral-fin ray 1 or 2 (very rarely 3) (Fig. 26) . . . . . . . . . . . . Plesiops verecundus
21b. Tips of dorsal-fin spines pale, but not red; total number of branches on ventral 6
pectoral-fin rays 20 to 35, very rarely fewer than 20; branches on ventralmost pectoral-fin
ray 2 to 5 (usually 3 or 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
Fig. 25 Plesiops oxycephalus Fig. 26 Plesiops verecundus

few exposed
22a. Skin bearing sensory pores posterior to eye and many exposed cheek scales
cheek scales
on skin of preopercle not expanded (Fig. 27a);
cheek scale rows 5 (4 to 6), visible as far anteriorly
as posterior tip of maxilla; few sensory pores im-
mediately posterior to tip of maxilla, suborbital
sensory pore series not continuous with those of
preopercle or dentary (Fig. 28) . . . . . . . Plesiops insularis many
22b. Skin bearing sensory pores posterior to eye and few or no a) Plesiops pores
b) Plesiops
on skin of preopercle expanded, resulting in a pores insularis genaricus
Fig. 27
smaller area of visible cheek scales (Fig. 27b);
cheek scale row 4 (3 to 5), not visible as far anteriorly as posterior tip of maxilla; many
sensory pores present immediately posterior to maxilla, resulting in a continuous preop-
ercular, dentary, and suborbital pore series (Fig. 29) . . . . . . . . . . . . . . . . Plesiops genaricus
Fig. 28 Plesiops insularis Fig. 29 Plesiops genaricus

Acanthoplesiops echinatus Smith-Vaniz and Johnson, 1990
Acanthoplesiops hiatti Schultz, 1953
Acanthoplesiops psilogaster Hardy, 1985
Assessor flavissimus Allen and Kuiter, 1976
Assessor macneilli Whitley, 1935
Beliops batanensis Smith-Vaniz and Johnson, 1990
Belonepterygion fasciolatum (Ogilby, 1889)
Calloplesiops altivelis (Steindachner, 1903)
Fraudella carassiops Whitley, 1935
Paraplesiops bleekeri Günther, 1861
Paraplesiops poweri Ogilby, 1908
Plesiops auritus Mooi, 1996
Plesiops cephalotaenia Inger, 1955
Plesiops coeruleolineatus Rüppell, 1835
Plesiops corallicola Bleeker, 1853
Plesiops facicavus Mooi, 1996
Plesiops genaricus Mooi and Randall, 1991
Plesiops gracilis Mooi and Randall, 1991
Plesiops insularis Mooi and Randall, 1991
Plesiops oxycephalus Bleeker, 1855
Plesiops polydactylus Mooi, 1996
Plesiops verecundus Mooi, 1996
Steeneichthys plesiopsus Allen and Randall, 1985
References
Mooi, R.D. 1993. Phylogeny of the Plesiopidae (Pisces: Perciformes) with evidence for the inclusion of the
Acanthoclinidae. Bull. Mar. Sci., 52(1):284-326.
Mooi, R.D. 1996. Revision, phylogeny, and discussion of biology and biogeography of the fish genus Plesiops
(Perciformes: Plesiopidae). Roy. Ontario Mus. Life Sci. Contrib., (159):108 p.
Smith-Vaniz, W. and G.D. Johnson. 1990. Two new species of Acanthoclininae (Pisces: Plesiopidae) with a synopsis
and phylogeny of the subfamily. Proc. Acad. Natl. Sci. Philadelphia, 142:211-260.
2586 Bony Fishes
NOTOGRAPTIDAE
Bearded eelblennies
by R.D. Mooi
D iagnostic characters: Body elongate, eel-like (to about 20 cm total length). Head somewhat
flattened and long, with eye appearing to be far forward. Mouth large, with maxilla extending well
beyond posterior margin of eye. Teeth small and conical, forming broad toothed surfaces on palatines
and jaws. Upper and lower lips complete (i.e. not interrupted at symphysis), each with a median
lobe, the lower of which is elongate and barbel-like. Branchiostegal rays 6. Gill rakers absent.
Dorsal, anal, and caudal fins confluent; dorsal fin with LXII to LXIX spines and 1 or 2 segmented
rays; anal fin with XXXVII to XLIII spines and 1 or 2 segmented rays; spines in dorsal and anal fins
increase in stoutness posteriorly; caudal-fin rays reduced to a single unsegmented ray dorsally and
ventrally and 6 dorsal and 5 ventral segmented rays; pelvic fins with I spine and 2 segmented rays,
usually unbranched (often incorrectly reported as 1 branched ray). Lateral line of enlarged dermal
ossicles high on body along length of dorsal fin; remaining scales very small and cycloid, deeply embedded
in skin. Skeleton highly modified; median ethmoid laterally compressed and plate-like, extending anterior
of vomer; most suspensorial bones horizontally elongate, with palatine extending far posteriorly beneath
ectopterygoid; hyomandibula weakly associated with remainder of suspensorium; only 4 infraorbital bones;
gill arches reduced, remaining bones elongate; no supraneural bones; dorsal-fin pterygiophores insert in
1:1 ratio with interneural spaces; all spine-bearing dorsal- and anal-fin pterygiophores articulate with spines
via a complete, interlocking bony ring; 71 to 75 total vertebrae; epineural ribs confined to first 3 vertebrae,
pleural ribs usually confined to vertebrae 3 to 6; caudal elements reduced, parhypural and hypurals 1 and
2 fused into a single element, hypurals 3 and 4 fused to each other and to urostylar complex, there are no
free uroneurals, hypural 5 absent, epurals absent, free caudal cartilages absent. Gut a simple tube.
Colour: variable, from pale with numerous spots to dark-bodied.
short median lobe on upper lip dorsal fin with more than LX spines
large mouth
barbel-like
median lobe reduced pelvic fins
on lower lip anal fin with more than XXX spines
Habitat, biology, and fisheries: Inshore reefs, rocky crevices, and sand-weed areas. Seldom seen. At
least some species appear to specialize on alpheid shrimps. Eggs with long filaments from hook-like
protuberances, a similar morphology to other reef groups that exhibit egg-guarding, suggesting that
notograptids might share this behaviour. No commercial catch, and not encountered frequently enough
to be important for the aquarium trade.
Remarks: There are 5 nominal species, Notograptus gregoryi, N. guttatus, N. kauffmani, N. livingstonei,
and Blanchardia maculata. The latter was considered a synonym of N. guttatus by Ogilby, and this status
will be maintained here; the family contains only a single genus, Notograptus. The validity of the remaining
species has not been recently investigated. N. kauffmani is based on a small specimen that differs only
in colour, and is very likely a juvenile of N. guttatus. Recognition of N. livingstonei is based on what appear
to be minor colour differences from N. guttatus. N. gregoryi is not recorded from the area (presently known
only from Western Australia), but is included in the key because accurate distribution data for these
seldomly encountered fishes are not available. The family is found on the coasts of southern Papua New
Guinea and northern Australia.
Pseudochromidae: members of the pseudochromid sub-
family Congrogadinae resemble notograptids in having
an elongate, eel-like body form, long dorsal and anal fins
that are confluent with the caudal fin in some species,
and reduced (often absent) pelvic fins. Congrogadines
are easily distinguished by the 2 or fewer spines in the
dorsal fin, lack of spines in the anal fin, a smaller mouth
which does not reach the posterior margin of eye or Congrogadinae
extends only slightly beyond, a short, toothless palatine,
and no barbel-like median lobes on the lips.
Perciformes: Percoidei: Notograptidae 2587
Pholidichthyidae: superficially similar to notograptids with an elongate, eel-shaped body, reduced pelvic
fins, long dorsal and anal fins confluent with caudal fin. They are readily differentiated by their lack of
dorsal- or anal-fin spines (segmented rays only), smaller mouth, single nostril on each side (rather than
2), presence of gill rakers, lack of palatine teeth, and no barbel-like median lobes on the lips.
Ophidiidae and Bythitidae: elongate with long dorsal and anal fins that can be confluent with the caudal
fin; pelvic fins reduced and often far forward, even under chin; some species with true barbels in pairs, not
associated with lips as in notograptids; dorsal and anal fins without spines; body shape less eel-like than
in notograptids.
Pholidichthyidae Ophidiidae
Blenniidae: conceivably, some elongate members of this family with long dorsal and anal fins (e.g.
Xiphasia) could be confused with notograptids, but all differ in having II or fewer spines in the anal fin,
dorsal-fin spines are usually flexible, and barbel-like median lobes on the lips absent.
Key to the species of Notograptidae occurring in the area

1a. Body a uniform chocolate brown without
spots; median fins also dark except for
pale margins; paired fins yellowish; head
paler with cheeks and opercle with a few
oblong spots (Fig. 1) . . . . . . Notograptus gregoryi
(Western Australia; not yet recorded from the area)
Fig. 1 Notograptus gregoryi
1b. Body spotted or with wide longitudinal
stripes . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Body with 2 dark wide stripes or bands on a paler background: a dorsal band extending
from top of head onto dorsal fin to caudal fin, and a lateral band extending from eye
gradually angling downward onto anal fin to caudal fin; head without spots, a small bar
from ventral eye onto maxilla and lower lip (Fig. 2) . . . . . . . . . . . . . . Notograptus kauffmani
2b. Body spotted; numerous oblong spots on cheek and dorsal portion of head . . . . . . . . . . . ® 3
3a. Dorsal fin without spots or with spots only on anterior membranes (Fig. 3) . . . Notograptus guttatus
3b. Dorsal fin with spots over entire length . . . . . . . . . . . . . . . . . . . . Notograptus livingstonei
Fig. 2 Notograptus kauffmani Fig. 3 Notograptus guttatus

Notograptus guttatus Günther, 1867
Notograptus kauffmani Tyler and Smith, 1970
Notograptus livingstonei Whitley, 1931
Reference
Gill, A.C. and R.D. Mooi. 1993. Monophyly of the Grammatidae and of the Notograptidae, with evidence for their
phylogenetic positions among perciforms. Bull. Mar. Sci., 52(1):327-350.
2588 Bony Fishes
OPISTOGNATHIDAE
Jawfishes
by W.F. Smith-Vaniz
D iagnostic characters: Generally small, moderately elongate fishes (most under 12 cm total length, a
single species to about 50 cm), with a tapering narrow body. Head bulbous. Eyes relatively large
and high on head. Mouth large; in some species the maxilla extends to or well beyond posterior margin
of gill flap. Moderate canine-like teeth in a single row along sides of jaws; several rows of smaller teeth
may also be present anteriorly. Dorsal fin shallowly notched (if at all) between spinous and soft
portions, with IX to XII usually flexible spines and 10 to 22 segmented rays; anal fin with II or III slender
spines and 10 to 20 segmented rays; caudal fin rounded, the middle 12 to 14 rays branched in most species;
pelvic fins positioned anterior to pectoral fins, with I spine and 5 segmented rays, the outer 2
segmented rays unbranched and stout, the inner rays branched and weaker. Lateral line high on
body, incomplete, and usually ending below about middle of dorsal fin; lateral-line tubes or canals usually
embedded in skin, rather than occurring on scales. Scales cycloid (smooth), usually absent from cheeks.
Colour: most species mottled with various shades of brown; a black spot on spinous dorsal fin often
present.
II-III slender spines
maxilla very long in pelvic fin

some species
Habitat, biology, and fisheries: Most jawfishes occur in relatively shallow depths (2 to 30 m) on sandy
substrates adjacent to coral reefs, but some species have been trawled in depths exceeding 200 m.
Jawfishes live in burrows, which they construct themselves, and usually are found with only their heads
protruding from the burrow. They normally venture from their burrows only a short distance to feed on
crustaceans, zooplankton, and/or other invertebrate prey, or to defend their territories from conspecific
intruders. When treatened by potential predators they nearly always back tail first into their burrows. Some
species are solitary but most live in small colonies. All species are mouth brooders and their eggs, which
clump together in a sticky egg ball, can be set down in the burrow by the parent during brief feeding forays;
once the eggs hatch no additional care is provided. Jawfishes are not of commercial importance, although
they are occasionally caught by hook-and-line fishermen, taken in trawls, or by spear fishermen. They are
reported to be good to eat.
Remarks: No key to species of Opistognathus is given here because there are 16 undescribed species
known from the area that are currently under study. These new species will be treated in a separate
publication by the present author. The 12 species of Stalix are all small (22 to 60 mm standard length) and
rare, with fewer than 35 total collections known.
Perciformes: Percoidei: Opistognathidae 2589

The arrangement of the pelvic-fin rays, consisting of
I spine and 5 segmented rays (the outer 2 un-
branched and stout, inner 3 branched and weak), will
distinguish the jawfishes from all other families. Addi-
tional distinguishing characters of the superficially
similar Cepolidae are the following:
Cepolidae (including Owstoniidae): dorsal-fin spines
0 to IV (IX to XII in Opistognathidae); caudal fin
moderately to strongly lanceolate (rounded in Opis- Cepolidae
tognathidae).
Key to the genera of Opisthognathidae occurring in the area
1a. First 5 to 9 dorsal-fin spines transversely forked distally (posteriorly spines successively
less deeply and more narrowly forked); skin covering spinous dorsal fin forming a broad
dorsal hood to accommodate transversely forked spines (Fig. 1) . . . . . . . . . . . . . . . . . Stalix
1b. Anterior dorsal-fin spines not transversely forked; skin covering spinous dorsal fin not
forming a broad dorsal hood (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . Opistognathus
broad dorsal hood
Fig. 1 Stalix moenensis (3.9 cm standard length) Fig. 2 Opistognathus

Opistognathus castelnaui Bleeker, 1859
Opistognathus darwinensis Macleay, 1878
Opistognathus decorus Smith-Vaniz and Yoshino, 1972
Opistognathus dendriticus (Jordan and Richardson, 1908)
Opistognathus evermanni (Jordan and Snyder, 1902)
Opistognathus eximus (Ogilby, 1908)
Opistognathus jacksoniensis Macleay, 1881
Opistognathus latitabundus (Whitley, 1937)
Opistognathus macrolepis Peters, 1866
Opistognathus nigromarginatus Rüppell, 1830
Opistognathus papuensis Bleeker, 1868
Opistognathus reticulatus (McKay, 1969)
Opistognathus solorensis Bleeker, 1853
Stalix dicra Smith-Vaniz, 1989
Stalix eremia Smith-Vaniz, 1989
Stalix histrio Jordan and Snyder, 1902
Stalix immaculata Xu and Zhan, 1980
Stalix moenensis (Popta, 1922)
Stalix versluysi (Weber, 1913)
References
Smith-Vaniz, W.F. 1989. Revision of the jawfish genus Stalix (Pisces: Opistognathidae), with descriptions of four new
species. Proc. Acad. Nat. Sci. Phila., 141:375-407.
Smith-Vaniz, W.F. and T. Yoshino. 1985. Review of Japanese jawfishes of the genus Opistognathus (Opistognathidae)
with description of two new species. Japan. J. Ichthyol., 32(1):18-27.
click for next page

2590 Bony Fishes
PRIACANTHIDAE
Bigeyes
by W.C. Starnes
D iagnostic characters: Deep-bodied, laterally compressed percoid fishes (size to 36 cm) with ex-
tremely large eyes (about 1/2 head length) and upturned mouth. Weak spine on posterior opercle
and prominent to remnant spine at angle of preopercle. Branchiostegal rays 6. Total gill rakers on first
gill arch 17 to 32. Dorsal fin with X spines and 11 to 15 soft rays; spinous and soft-rayed portions of fin
continuous, relatively short to long, soft-rayed portion broadly rounded to broadly pointed. Anal-fin rays
relatively short to long and broadly rounded to broadly pointed; anal fin with III spines and 10 to 16 soft
rays. Caudal fin rounded, emarginate, or lunate, with 16 principal rays. Pectoral fins relatively short, with
17 to 21 rays. Pelvic fins with I spine and 5 soft rays, the fins short to very long, broadly attached to belly
by membrane and positioned in advance of pectoral fins. Head and body mostly covered with extremely
adherent, rough, spiny scales (bearing true spines which are integral part of scale rather than cteni on
individual detachable bases). Scales much modified, varying among genera and species. Scales on
branchiostegal rays. Spinules on fin spines. Scales in lateral series (all lateral-line scales including
pored scales on caudal-fin base) 38 to 115. Vertebrae 23. Some species with modifications of skull and
swimbladder, including connections between these components. Colour: head, iris of eye, and body
generally reddish, sometimes with silvery blotches or, in some species, occasionally a pattern of red and
silver-white bars; fins reddish to dusky or black, occasionally yellowish in some species; some species
with dark spots or speckling on fin membranes.
single dorsal fin with X spines and 11-15 soft rays
eyes
very
large
pelvic fins in advance

of pectoral fins
anal fin with III spines

pelvic fins attached to and 10-16 soft rays
belly by membrane
Habitat, biology, and fisheries: Generally epibenthic fishes occurring near coral reefs or rock formations
but occasionally in more open areas at depths of 5 to 400 m, or deeper. Probably most active nocturnally
but known to feed diurnally as well. Feed primarily on crustaceans, small cephalopods, polychaetes, and
small fishes. Eggs, larvae, and early juvenile stages pelagic, transforming on settling to suitable habitats.
Occur solitarily or in small aggregations, but some species may form large aggregations at times based
on trawl catches. Not important in most fishery areas but some species occasionally common in trawl
catches of southeast Asian waters. Generally incidental in trawls or hook-and-line fisheries elsewhere.
Flesh is said to be of good quality. From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports a
range of yearly catch of around 23 100 to 52 000 t of Priacanthus spp. from the Western Central Pacific
(Indonesia, Thailand).
Perciformes: Percoidei: Priacanthidae 2591
dorsal fin deeply incised

Holocentridae: also with large eyes (particularly in Myripristis) and
reddish colour, but readily distinguishable from bigeyes by spines
on opercular margin, spinous and soft-rayed portions of dorsal fin
nearly separate, deeply forked caudal fin with 18 or 19 rays, pelvic
fins with origin behind pectoral-fin origin and having I spine and
usually 7 soft rays and not attached to belly by membrane, and IV
anal-fin spines.
Berycidae: also with large eyes and reddish coloration but readily 7 pelvic-fin IV anal-fin spines
distinguishable from bigeyes by a short-based dorsal fin with only rays
IV to VII spines, anal fin with IV spines, caudal fin deeply forked, Holocentridae
and pelvic fins having origin behind pectoral fins and with 7 to 13
soft rays.
Pempheridae: also with large eyes and reddish to coppery colour but with dorsal-fin base short, IV or V
spines and 8 or 9 soft rays, and anal fin with very long base, III spines and 22 or more soft rays.
IV-VII dorsal-fin spines IV-V dorsal-fin spines
IV anal-fin spines III anal-fin spines

7-13 pelvic-fin rays
Berycidae Pempheridae
Identification note
Scales in lateral series are counted in straight line at midbody from behind opercle onto caudal fin, joining
lateral line in anterior peduncle area and including all pored scales onto caudal-fin base (Fig. 1)
Key to the species of Priacanthidae occurring in the area
Note: some specimens which would to key to Priacanthus macracanthus herein may represent problematic
forms which resemble that species but may comprise 1 or more additional taxa (see remarks in species
account of P. macracanthus).
1a. Body profile very deep and broadly ovate, its depth 1.7 to 1.9 times in standard length;
anal-fin rays 10 or 11; dorsal-fin rays 11 or 12; scales in lateral series 36 to 51 . . . (Pristigenys) ® 2
1b. Body profile less deep, its depth 2 to 3.1 times in standard length; anal-fin rays 13 to 16;
dorsal-fin rays 12 to 15; scales in lateral series 56 to 115 . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Dorsal-fin rays 12; anal-fin rays 11; midlateral scales with 8 to 20 spinules on posterior
margin; sides light coloured with numerous narrow red bars (Fig. 1) . . . . . . . . Pristigenys meyeri
2b. Dorsal-fin rays 11; anal-fin rays 10; midlateral scales with 24 to 42 spinules on posterior
margin; sides red with widely spaced light bars (Fig. 2) . . . . . . . . . . . . . Pristigenys niphonia
location of scales in
lateral series counts
Fig. 1 Pristigenys meyeri Fig. 2 Pristigenys niphonia

2592 Bony Fishes
3a. Scale rows between dorsal fin and lateral line at highest point 16 to 20; pelvic fins very
long except in large adults (30 cm standard length or larger) exceeding head length
(Fig. 3); soft dorsal and anal fins long and broadly pointed except in very large specimens
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cookeolus japonicus
3b. Scales rows between dorsal fin and lateral line at highest point fewer than 16; pelvic fins
short, less than or equal to head length; soft dorsal and anal fins moderately long,
broadly rounded to very broadly pointed (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
pelvic fins
long
Fig. 3 Cookeolus Fig. 4
4a. Posterior portion of preopercle lacking
scales (Fig. 5a) and notably striate; ante -
rior profile nearly symmetrical, extremity
of lower jaw when mouth tightly closed
about level with midline of body (Fig. 6);
soft dorsal, anal, and caudal fins usually
with small dark specks in membranes . .
. . . . . . . . . . . . . Heteropriacanthus cruentatus
4b. Posterior portion of preopercle with scales a) Heteropriacanthus b) Priacanthus
(Fig. 5b); anterior profile more asymmetri-
cal, extremity of lower jaw usually above Fig. 5
level of midline of body (Fig. 7); fins plain
or with larger dusky spots . . . . . . . . . . . . . ® 5
Fig. 6 Heteropriacanthus cruentatus Fig. 7 Priacanthus hamrur
5a. Pelvic-fin membranes with several

large purplish black spots (Fig. 8);
dorsal-fin rays usually 12 . . Priacanthus tayenus
5b. Pelvic-fin membranes lacking spots
or with single dark basal blotch; dor-
sal-fin rays usually 13 or more . . . . . . . . ® 6
Fig. 8 Priacanthus tayenus

6a. Pelvic fins usually with a dark

blotch basally (Fig. 7); preopercu-
lar spine very short, virtually lack-
ing in specimens over 12 cm total
length (Fig. 9a) . . . . . . . . . . . . . . . ® 7
6b. Pelvic fins lacking dark blotch
ba s al l y ; pr e op ercular spine a) Priacanthus hamrur b) Priacanthus macracanthus
reaching to or beyond opercular Fig. 9
margin (Fig. 9b) . . . . . . . . . . . . . . ® 12
7a. Caudal-fin margin concave, outer rays slightly (Fig. 7) to much longer than remainder of
rays; anal-fin rays usually 15 or 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7b. Caudal-fin margin truncate, convex (Fig. 12), or very slightly emarginate; anal-fin rays
usually 14 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
8a. Total gill rakers on first gill arch 24 to 26; body depth at sixth dorsal-fin spine about 2.6
to 2.8 times in standard length (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . Priacanthus hamrur
8b. Total gill rakers on first gill arch 29 to 31; body depth at sixth dorsal-fin spine 3 or more
times in standard length (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . Priacanthus prolixus
(Indian Ocean, perhaps also Hong Kong; not yet recorded from the area)
9a. Total gill rakers on first gill arch 25 to 28; pectoral fins bright yellow in life . . . . Priacanthus zaiserae
9b. Total gill rakers on first gill arch 23 or fewer; pectoral fins not bright yellow . . . . . . . . . . . . ® 10
10a. First 2 spinous dorsal-fin membranes with black blotch (Fig. 11); length of second dorsal-fin
spine about 2 times in length of tenth spine; scales in lateral series 67 to 74 . . . Priacanthus sagittarius
10b. First 2 spinous dorsal-fin membranes not darker than succeeding; length of second
dorsal-fin spine 1.5 to 1.7 times in length of tenth spine; scales in lateral series 72 to 93 . . . . ® 11
Fig. 10 Priacanthus prolixus Fig. 11 Priacanthus sagittarius

11a. Length of pectoral fins about 1.9 to 2.2 times in head length (measured from tip of upper
jaw to tip of opercular spine, Fig. 12); body depth at sixth dorsal-fin spine 2.6 to 2.9
(usually about 2.7) times in standard length . . . . . . . . . . . . . . . . . . . . Priacanthus blochii
11b. Length of pectoral fins about 1.3 to 1.6 times in head length (Fig. 13); body depth at sixth
dorsal-fin spine 2.3 to 2.7 (usually about 2.5) times in standard length . . . . . . Priacanthus alalaua
Fig. 12 Priacanthus blochii Fig. 13 Priacanthus alalaua

2594 Bony Fishes
12a. Dorsal-, anal-, and pelvic-fin membranes with yellow-brown spots (grey in preservative);
soft portions of dorsal and anal fins not distinctly whiter posteriorly; body not tapering
1/
from just behind head (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . Priacanthus macracanthus
12b. Dorsal-, anal-, and pelvic-fin membranes lacking spots; soft portions of dorsal and anal
fins creamy white on posterior half; body generally tapering from just behind head
(Fig. 15) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Priacanthus fitchi
Fig. 14 Priacanthus macracanthus Fig. 15 Priacanthus fitchi

0 Cookeolus japonicus (Cuvier, 1829)
0 Heteropriacanthus cruentatus (Lacepède, 1801)
0
0
Priacanthus alalaua Jordan and Evermann, 1904
0
Priacanthus blochii Bleeker, 1853
0
Priacanthus fitchi Starnes, 1988
0
Priacanthus hamrur (Forsskål, 1775)
Priacanthus macracanthus Cuvier, 1829 1/
Priacanthus prolixus Starnes, 1988 2/
0
?
0
Priacanthus sagittarius Starnes, 1988
0
Priacanthus tayenus Richardson, 1846
Priacanthus zaiserae Starnes and Moyer, 1988
0
0
Pristigenys meyeri (Günther, 1871)
Pristigenys niphonia (Cuvier, 1829)
References
Senta, T. 1977. Species and size composition of priacanthid fishes in the South China Sea and adjacent waters. Bull.
Pac. Fish., 42:25-31.
Starnes, W.C. 1984. Priacanthidae. In FAO species identification sheets for fishery purposes. Western Indian Ocean
Starnes, W.C. 1988. Revision, phylogeny, and biogeographic comments on the circumglobal marine percoid fish family
Priacanthidae. Bull. Mar. Sci., (43(2):117-203.
1/ See remarks on species account.

2/ Occurrence in the area questionable; based on a specimen ostensibly from Hong Kong; all other known records are
from the northwestern Indian Ocean.
Heteropriacanthus cruentatus (Lacepède, 1801)

Frequent synonyms / misidentifications: Priacanthus cruentatus (Lacepède, 1801) / Various
Priacanthus spp.
FAO names: En - Glasseye; Fr - Beauclaire de roche; Sp - Catalufa de roca.
Diagnostic characters: Body deep, ovate, and laterally compressed. Anterior profile symmetrical, tip
of protruding lower jaw about on level with midline of body when mouth tightly closed. Well-developed
spine at angle of preopercle. Small teeth on dentaries, vomer, palatines, and premaxillaries. Total gill rakers
on first gill arch 21 to 25. Dorsal fin with X spines and 11 to 13 soft rays. Anal fin with III spines and 13 or
14 soft rays. Caudal fin truncate. Pectoral fins with 18 or 19 rays. Scales covering most of head and body
but scales lacking on posterior portion of preopercle. Scales modified, those of midlateral area with
posterior field elevated above a separate flange, broadly pointed, with spinules confined to
posterior margin. Scales in lateral series 78 to 96; pored lateral-line scales 63 to 81; vertical scale rows
(dorsal-fin origin to anus) 56 to 68. Swimbladder with pair of posterior extensions only. Colour: entire
body and head pinkish red or blotched with red and silver; iris of eye red; fins reddish, membranes of
spinous dorsal fin and margin of caudal fin sometimes dusky; caudal, soft dorsal, and anal fins with
elliptical dark specks.
Size: Maximum total length to about 35 cm.
Habitat, biology, and fisheries: Inhabits shallow reef areas, particularly in insular areas, where it may be
common in both lagoons and seaward areas, commonly at depths of 20 m or less. Secretive by day and
foraging at night. Feeds on octupuses, shrimps, stomatopods, crabs, small fishes, and polychaetes. Caught
primarily on hook-and-line and in traps. Marketed mostly fresh.
Distribution: Circumtropical
and into subtropical waters.
Young occasionally in tem-
perate waters due to postlar-
val transport.
2596 Bony Fishes
Priacanthus hamrur (Forsskål, 1775)

Frequent synonyms / misidentifications: None / Possibly Priacanthus prolixus Starnes, 1988.
FAO names: En - Moontail bullseye; Fr - Beauclaire miroir; Sp - Catalufa espejuelo.
Diagnostic characters: Body deep, ovate, and laterally compressed; body depth 2.6 to 2.8 times in
standard length. Anterior profile slightly asymmetrical, tip of protruding lower jaw usually above midline
of body. Small teeth on dentaries, vomer, palatines, and premaxillaries. Spine at angle of preopercle
reduced or non-existent in specimens over 12.5 cm total length. Total gill rakers on first gill arch
24 to 26. Dorsal fin with X spines and 13 to 15 soft rays. Anal fin with III spines and 13 to 16 soft rays.
Caudal fin with concave margin, slightly emarginate to lunate. Pectoral-fin rays 17 to 20. Scales
covering most of head and body onto base of caudal fin. Scales modified, the posterior field elevated
as a separate flange with spinules both on surface and posterior margin. Scales in lateral series 79
to 96; pored lateral-line scales 70 to 90. Vertical scale rows (dorsal-fin origin to anus) 48 to 57.
Swimbladder with pair of anterior and posterior protrusions, the former associated with specialized
recesses in posterior of skull. Colour: red on body, head, and iris of eye; sometimes silvery white with
pattern of broad reddish bars on head and body; row of small dark spots sometimes evident along lateral
line; fins red to light pink, with light dusky to nearly black in dorsal-, anal-, and caudal-fin membranes; fins
occasionally with yellow.
Habitat, biology, and fisheries: Occurs near reefs and rocky areas at depths ranging from less than 20
to 250 m or more, but probably most common at 30 to 50 m. Prefers outer reef slopes to more sheltered
environments. Occasionally taken in moderate numbers in trawls and by hook-and-line. Marketed mostly
fresh.
the Indo-West Pacific from
Red Sea and South Africa to
northern Australia and south-
ern Japan and eastward to at
least Tuamotu in French Poly-
nesia.
Priacanthus macracanthus Cuvier, 1829 BIR

Frequent synonyms / misidentifications: None / May represent 2 or more forms (see Remarks).
FAO names: En - Brownspot bigeye (formerly reported as “red bigeye”); Fr - Beauclaire Pacifique;
Sp - Catalufa Pacífico.
Diagnostic characters: Body relatively deep, ovate, and laterally compressed. Anterior profile slightly
asymmetrical, tip of protruding lower jaw generally slightly above midline of body. Small teeth on dentaries,
vomer, palatines, and premaxillaries. Spine at angle of preopercle well developed in specimens of all
sizes. First gill arch with 23 to 29 gill rakers (highest gill-raker count in eastern Australian waters). Dorsal
fin with X spines and 12 to 14 soft rays. Anal fin with III spines and 13 or 14 soft rays. Caudal-fin margin
truncate to very slightly concave. Pectoral fins generally short but averaging longer in some Indonesian
specimens; pectoral-fin rays 18 or 19. Scales covering most of body, head, and caudal-fin base. Scales
modified with posterior field elevated as a separate flange with spinules both on surface and
posterior edge. Scales in lateral series 79 to 87; pored lateral-line scales 72 to 82. Vertical scale rows
(dorsal-fin origin to anus) 45 to 52. Swimbladder with both anterior and posterior projections, the
former associated with specialized recessed areas on rear of skull. Colour: body, head, and iris silvery
pink to reddish; fins light pinkish; dorsal, anal, and pelvic fins with rusty brown spots which are about
1/2 diameter of pupil of eye; specimens tentatively referable to this species from off eastern Australia have
smaller spots.
Size: Maximum total length possibly 35 cm.
Habitat, biology, and fisheries: As presently construed, Priacanthus macracanthus occurs over a range of
depths from 15 to 400 m or more. Occurs, at times, both in reef or rocky areas and in more open bottom areas.
Occasionally important in trawl fisheries of South China and Andaman seas and to lesser extent in southern
Australia; separate statistics are available from Taiwan Province of China, but not from the Western Central
Pacific. Most commonly taken at dusk and dawn; also taken by hook-and-line. Juvenile aggregations in midwater
are reported from the Philippines. Marketed fresh, dried, salted, and as fish balls.
Distribution: As presently construed, in Indo-West Pacific and generally confined to region from both east
and west coasts of Australia northward to Andaman Sea and southern Japan.
Remarks: The variation in pectoral-fin size, gill rakers, and fin coloration described above plus other
morphological and ecological factors may be indicative of polytypy among populations collectively treated
here as P. macracanthus. Some specimens may represent problematic forms which resemble P. macra-
canthus but may comprise 1 or more addi-
tional taxa. Problematic specimens are
generally from deeper trawl (e.g. 400 m)
collections (thus far from northern and east-
ern Australian waters) as opposed to those
from shallower stations which are more
“typical” macracanthus. Resolution of this
problem has been hampered by insufficient
material.
2598 Bony Fishes
Priacanthus tayenus Richardson, 1846

FAO names: En - Purplespot bigeye; Fr - Beauclaire tache pourpre; Sp - Catalufa mota purpúreo.
Diagnostic characters: Body moderately deep, moderately elongate, and laterally compressed. Anterior
profile slightly assymetrical, extremity of protruding lower jaw usually slightly above level of midline of body.
Small teeth on dentaries, vomer, palatines, and premaxillaries. Well-developed spine at angle of
preopercle. Total gill rakers on first gill arch 21 to 24. Dorsal fin with X spines and 11 to 13 soft rays. Anal
fin with III spines and 12 to 14 soft rays. Caudal fin more or less truncate in smaller specimens but
becoming very lunate in some (possibly males) but not all larger specimens. Pectoral-fin rays 17 to
19. Scales covering most of body, head, and caudal-fin base. Scales modified; scales of midlateral region
with elevated posterior field reduced and lacking spinules in larger specimens. Scales in lateral
series 56 to 73; pored lateral-line scales 51 to 67. Vertical scale rows (dorsal-fin origin to anus) 40 to 50.
Swimbladder with both anterior and posterior projections, the former associated with modified recesses in
skull. Colour: body, head, and iris of eye pink to reddish or silvery white with pink tinges; fins pinkish;
pelvic fins with characteristic small deep purple to inky black spots in membranes with 1 or 2 larger
spots in connective membrane to belly.
Size: A smaller species, maximum total length perhaps 29 cm.
Habitat, biology, and fisheries: Inhabits coastal waters in both rocky reef and, occasionally, more open
areas at depths of 20 to 200 m or more. Adults apparently aggregate at times and are periodically relatively
common in the trawl fisheries of the Andaman and South China seas. Recruit, at least periodically, into
aggregations at about 12 cm total length and attain about 24 cm by following year. Marketed fresh, dried,
and as fish balls.
Distribution: In coastal ar-
eas of northern Indian Ocean
from Persian Gulf eastward
and into West Pacific from
northern Australia and Solo-
mon Islands northward to Tai-
wan Province of China.
?
Cookeolus japonicus (Cuvier, 1829)

En - Longfin bulleye; Fr - Beauclaire longe aile; Sp - Catalufa aleta larga.
Maximum total length about 60 cm (largest member of family). In deeper waters off rocky coasts or
insular areas in association with holes and ledges at depths of 60 to 400 m. Feeds on crustaceans
and small fishes; lifespan of 9 or more years. Principally caught incidentally on deep handlines or
other rigs; probably rare in markets. Circumtropical into subtropical regions; young occasionally
occurring in temperate waters as result of postlarval transport.
Priacanthus alalaua Jordan and Evermann, 1904

En - Alalaua; Fr - Beauclaire alalaua; Sp - Catalufa alalaua.
Maximum total length about 32 cm (eastern Pacific, probably less in western Pacific). Inhabits
deeper reefs or rocky areas usually at depths of 100 to 275 m; occasionally taken at depths of 10
to 50 m at night. Also trawled from relatively open bottom areas in Hawaiian waters. Apparently rare
in western Pacific region, occasionally taken on hook-and-line. Tropical eastern Pacific and Hawaii.
In the western Pacific, thus far known only from Guam.
Priacanthus blochii Bleeker, 1853

En - Shortfin bigeye; Fr - Beauclaire aile court; Sp - Catalufa aleta corto.
Maximum total length about 30 cm. Coastal in shallow reef and rocky areas at depths of 15 to 30 m.
Probably rare in trawl catches; occasionally taken by hook-and-line. Frequently misidentified as
Priacanthus hamrur, P. sagittarius, or Heteropriacanthus cruentatus. Widespread in Indo-West
Pacific from Gulf of Aden (and perhaps Red Sea) and Seychelles eastward to Samoa.
?
2600 Bony Fishes
Priacanthus fitchi Starnes, 1988

En - Whitefin bigeye; Fr - Beauclaire aile blanc; Sp - Catalufa aleta blanca.
So far as known, a small species, not exceeding about 24 cm total length. A deeper water species
known from depths of 150 to 400 m or more. Occasionally taken in deeper trawls in moderate
numbers, particularly along the northwestern shelf of Australia. Presently known in Indo-West
Pacific from northwestern Australia and western Sumatra to southern Japan; also known from near
Mascarene Islands in western Indian Ocean.
? ?
Priacanthus sagittarius Starnes, 1988

En - Arrowfin bigeye; Fr - Beauclaire aile sagitté; Sp - Catalufa aleta saeta.
Maximum total length about 34 cm. Occurs in rocky and reef areas and occasionally in more open
areas at depths of 60 to 100 m or more. Moderate numbers occasionally taken in trawls, particularly
juveniles in 10 to 15 cm total length range. Frequently misidentified as Priacanthus blochii (Bleeker,
1853). Wide ranging in Indo-West Pacific from Red Sea and Reunion eastward to Japan, eastern
Australia, and Samoa.
Priacanthus zaiserae Starnes and Moyer, 1988

En - Yellowfin bigeye; Fr - Beauclaire aile juane; Sp - Catalufa aleta amarillo.
Maximum total length perhaps 30 cm. Reported in rocky areas from depths of 30 to over 320 m.
Occasionally taken by hook-and-line and sometimes appears in markets in Philippines. So far as
known, restricted to western Pacific in area of southern Japan southward into Philippines.
Pristigenys meyeri (Günther, 1871)

En - Redstripe bigeye; Fr - Beauclaire raie rouges; Sp - Catalufa raya roja.
Maximum total length perhaps 30 to 34 cm. The few specimens known are from rocky areas at
depths of 100 to 200 m. May be taken uncommonly by deep hook-and-line fishing. Pristigenys
multifasciata Yoshino and Iwai, 1973 is a junior synonym. Thus far known from only the western
Pacific from southern Japan southward to New Guinea and eastward to Samoa.
Pristigenys niphonia (Cuvier, 1829)

En - Whiteband bigeye; Fr - Beauclaire bande blanc; Sp - Catalufa faja blanca.
Maximum total length about 34 cm. Occurs at depths of 80 to 100 m or more in rocky habitats. Taken
occasionally by hook-and-line and possibly trawls. Widely distributed in Indo-West Pacific from Red
Sea and South Africa eastward to southern Japan, eastern Australia, and Samoa.
click for next page

2602 Bony Fishes
APOGONIDAE
Cardinalfishes
by G.R. Allen
D iagnostic characters: Small (to 20 cm, usually under 12 cm) percoid fishes; body ovate to elongate,
moderately compressed. Eyes large, their diameter exceeding snout length. Rear margin of cheek
(preopercle) with characteristic double edge, which is variously serrated or smooth. Mouth large,
lower jaw often protruding. Jaws with bands of small villiform teeth; teeth present on vomer, but palatine
teeth may be absent; enlarged canines on premaxillae, dentaries, or vomer in some species. Two separate
dorsal fins, the first consisting of VI to VIII rigid spines, the second with I rigid spine and 8 to 14
soft rays. Anal fin with II rigid spines and 8 to 18 soft rays. Caudal fin rounded to forked, frequently
emarginate or truncate. Pelvic fins with I spine and 5 soft rays. Pectoral fins with 10 to 20 rays.
Branchiostegal rays 7. Scales relatively large in marine species, except absent in Gymnapogon; scales
usually ctenoid, but cycloid in a few species, about 9 to 37 lateral-line scales in most marine species, but
absent in at least 1 species of Siphamia. Colour: highly variable, frequently shades of black, brown, red,
or yellow; many species exhibit a pattern of dark bars or stripes on a lighter ground colour.
2 dorsal fins
Apogon
double-edged
preopercle II anal-fin Pterapogon
spines
Rhabdamia
Habitat, biology, and fisheries: Inhabit coral and rocky reefs and adjacent habitats including sand-rubble
patches and seagrass beds; several species frequently shelter among the spines of sea urchins (usually
Diadema) or Crown-of-thorns starfish (Acanthaster). The genus Glossamia of Australia and New Guinea
is restricted to fresh waters. Cardinalfishes are among the few marine fishes which exhibit oral egg brooding
(by males). Generally not important economically, but a few species are seen in the aquarium trade and
species of Rhabdamia are occasionally used as tuna bait.
single dorsal fin

Ambassidae (= Chandidae): differ from Apogonidae in
having a single (although deeply notched) dorsal fin and
III anal-fin spines.
III anal-fin spines
Ambassidae
Perciformes: Percoidei: Apogonidae 2603
Key to the genera of marine Apogonidae occurring in the area

1a. Jaws without enlarged canine teeth . . . . . . . . . . . . . . . ® 2
1b. Jaws with enlarged canine teeth (Fig. 1) . . . . . . . . . . . . ® 13
2a. First dorsal fin with VIII spines (Fig. 2) . . . . . . . . . . . . Neamia

2b. First dorsal fin with VI or VII spines . . . . . . . . . . . . . . . ® 3
3a. Lower side of body with silvery, bioluminescent band
(Fig. 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Siphamia Fig. 1
3b. Lower side of body without silvery, bioluminescent band . . . . . ® 4
VIII spines in
1st dorsal fin
silvery,
bioluminescent
band
Fig. 2 Neamia Fig. 3 Siphamia
4a. Body scales entirely cycloid and deciduous; fragile, elongate fishes; semitransparent in
life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
4b. Body scales ctenoid; ovate to elongate fishes; not usually semitransparent, except for a
few species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
5a. Soft dorsal-fin rays 12 or 13 (Fig. 4) . . Lachneratus
5b. Soft dorsal-fin rays 9 . . . . . . . . . . . . . . ® 6
6a. Rear edge of preopercle with 1 to 3
small spines near angle (Fig. 5) . . . . . Cercamia
6b. Rear edge of preopercle smooth
(Fig. 6) . . . . . . . . . . . . . . . . . . Rhabdamia
Fig. 4 Lachneratus
1-3 spines smooth

Fig. 5 Cercamia Fig. 6 Rhabdamia
7a. Anal-fin rays 13 to 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8

7b. Anal-fin rays 8 to 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
2604 Bony Fishes
8a. Soft dorsal-fin rays 14 (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterapogon

8b. Soft dorsal-fin rays 9 (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Archamia
Fig. 7 Pterapogon Fig. 8 Archamia

9a. Secondary rays on uppermost and lower-
most edge of caudal fin spiny (Fig. 9) . . . Sphaeramia
9b. Secondary rays on uppermost and lower-
most edge of caudal fin soft . . . . . . . . . . . ® 10
10a. Rear edge of preopercle or ridge immedi-
ately in front of it, or both edge and ridge
serrated (sometimes weakly) (Fig. 10) . . . . Apogon
10b. Rear edge of preopercle and ridge imme-
diately in front of it smooth . . . . . . . . . . . . ® 11
11a. Lateral line complete, extending from up- spiny

per edge of operculum to base of caudal secondary
fin (Fig. 11) . . . . . . . . . . . . . . . Apogonichthys rays
11b. Lateral line incomplete, ending well be-
fore base of caudal fin (Figs 12 and 13) . . . . . ® 12 Fig. 9 Sphaeramia
serrated
Fig. 10 Apogon Fig. 11 Apogonichthys

12a. Palatine teeth present (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Foa
12b. Palatine teeth absent (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Fowleria
lateral line lateral line

incomplete incomplete
Fig. 12 Foa Fig. 13 Fowleria

13a. Scales absent; a distinct spine at angle of preopercle (Fig. 14) . . . . . . . . . . . . . Gymnapogon
13b. Scales present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
14a. Body covered with strong ctenoid scales; colour pattern usually consisting of conspicu-
ous dark stripes on a light ground (Fig. 15) . . . . . . . . . . . . . . . . . . . . . . . Cheilodipterus
14b. Body covered with deciduous cycloid scales; colour pattern usually lacking conspicuous
dark stripes, although series of thin dark lines may be present . . . . . . . . . . . . . . . . . . ® 15
distinct
preopercular
spine Fig. 14 Gymnapogon Fig. 15 Cheilodipterus
15a. Rear corner of upper jaw with tiny, downward projecting spine; front nostril lacking skin
flap on posterior margin (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudamiops
15b. Rear corner of upper jaw without downward projecting spine; front nostril with skin flap
on posterior margin (Fig. 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pseudamia
skin flap
tiny
downward
projecting Fig. 16 Pseudamiops Fig. 17 Pseudamia
spine
List of marine species occurring in the area

The symbol 0
Apogon abogramma Fraser and Lachner, 1985
Apogon albimaculosus Kailola, 1976
Apogon albomarginatus Smith and Radcliffe, 1912
Apogon amboinensis Bleeker, 1853
Apogon angustatus Smith and Radcliffe, 1912
Apogon apogonides (Bleeker, 1856)
Apogon atrogaster (Smith and Radcliffe, 1911)
Apogon aureus (Lacepède, 1803)
Apogon bandanensis Bleeker, 1854
Apogon brevicaudata Weber, 1909
Apogon capricornis Allen and Randall, 1993
Apogon caudicinctus Randall and Smith, 1988
Apogon carinatus Cuvier, 1828
Apogon cavitiensis (Jordan and Seale, 1907)
Apogon ceramensis Bleeker, 1852
Apogon cheni Hayashi, 1990
Apogon chrysopomus Bleeker, 1854
Apogon chrysotaenia Bleeker, 1851
Apogon coccineus Rüppell, 1835
Apogon compressus (Smith and Radcliffe, 1911)
Apogon cookii Macleay, 1881
0
Apogon crassiceps Garman, 1903
Apogon cyanosoma Bleeker, 1853
2606 Bony Fishes
Apogon dispar Fraser and Randall, 1976

Apogon diversa (Smith and Radcliffe, 1912)
Apogon doederleini Jordan and Snyder, 1901
Apogon doryssa (Jordan and Seale, 1906)
Apogon ellioti Day, 1875
Apogon endekataenia Bleeker, 1852
Apogon evermanni Jordan and Snyder, 1904)
Apogon exostigma (Jordan and Starks, 1906)
Apogon flavus Allen and Randall, 1993
Apogon fleurieu (Lacepède, 1802)
Apogon fraenatus Valenciennes, 1832
Apogon fragilis Smith, 1961
Apogon franssedai Allen, Kuiter, and Randall, 1995
Apogon fuscomaculatus Allen and Morrison, 1996
Apogon fuscus Quoy and Gaimard, 1825
Apogon gilberti Jordan and Seale, 1905
Apogon griffini (Seale, 1910)
Apogon guamensis Valenciennes, 1832
Apogon hartzfeldii Bleeker, 1852
Apogon hyalosoma Bleeker, 1852
Apogon kallopterus Bleeker, 1856
Apogon kiensis Jordan and Snyder, 1901
Apogon komodoensis Allen, 1998
Apogon lateralis Valenciennes, 1832
Apogon leptacanthus Bleeker, 1856
Apogon luteus Randall and Kulbicki, 1998
Apogon margaritophorus Bleeker, 1854
Apogon melanoproctus Fraser and Randall, 1976
Apogon melanopus Weber, 1911
Apogon melas Bleeker, 1848
Apogon moluccensis Valenciennes, 1832
Apogon multilineatus (Bleeker, 1865)
Apogon multitaeniatus Cuvier, 1828
Apogon nanus Allen, Kuiter, and Randall, 1994
Apogon neotes Allen, Kuiter, and Randall, 1994
Apogon niger Doderlein, 1884
Apogon nigripinis Cuvier, 1828
Apogon nigrocinctus (Smith and Radcliffe, 1911)
Apogon nigrofasciatus Lachner, 1953
Apogon notatus (Houttuyn, 1782)
Apogon novemfasciatus Cuvier, 1828
Apogon ocellicaudus Allen, Kuiter, and Randall, 1994
Apogon pallidofasciatus Allen, 1987
Apogon parvulus Smith and Radcliffe, 1912
Apogon perlitus Fraser and Lachner, 1985
Apogon poecilopterus Kuhl and van Hasselt, 1828
Apogon quadrifasciatus Cuvier, 1828
Apogon rhodopterus Bleeker, 1852
Apogon rubrimacula Randall and Kulbicki, 1998
Apogon ruepelli Günther, 1859
Apogon sangiensis Bleeker, 1857
Apogon sealei (Fowler, 1918)
Apogon selas Randall and Hayashi, 1990
Apogon semilineatus Schlegel, 1846
Apogon semiornatus Peters, 1876
Apogon septemstriatus Günther, 1880
Apogon striatus (Smith and Radcliffe, 1912)
Apogon taeniophorus Regan, 1908
Apogon taeniopterus Bennett, 1835
Apogon timorensis Bleeker, 1854

Apogon trimaculatus Cuvier, 1828
Apogon unicolor Doderlein, 1884
Apogon unitaeniatus Allen, 1995
Apogon ventrifasciatus Allen, Kuiter, and Randall, 1994
Apogonichthys ocellatus (Weber, 1913)
Apogonichthys perdix Bleeker, 1854
Archamia biguttata Lachner, 1951
Archamia buruensis (Bleeker, 1856)
Archamia dispilus Lachner, 1951
Archamia fucata (Cantor, 1850)
Archamia leai Waite, 1916
Archamia macroptera (Cuvier, 1828)
Archamia melasma Lachner and Taylor, 1960
Archamia zosterophora (Bleeker, 1856)
Cercamia cladara Randall and Smith, 1988
Cercamia eremia Allen, 1987
Cheilodipterus alleni Gon, 1993
Cheilodipterus artus Smith, 1961
Cheilodipterus intermedius Gon, 1993
Cheilodipterus isostigmus (Schultz, 1940)
Cheilodipterus macrodon (Lacepède, 1801)
Cheilodipterus nigrotaeniatus Smith and Radcliffe, 1912
Cheilodipterus parazonatus Gon, 1993
Cheilodipterus quinquelineatus Cuvier, 1828
Cheilodipterus singapurensis Bleeker, 1859
Cheilodipterus zonatus Smith and Radcliffe, 1912
Foa brachygramma (Jenkins, 1902)
Foa fo (Jordan and Seale, 1906)
Fowleria abocellata Goren and Karplus, 1980
Fowleria aurita (Valenciennes, 1831)
Fowleria flammea Allen, 1993
Fowleria marmorata (Alleyne and Macleay, 1877)
Fowleria punctulata (Rüppell, 1838)
Fowleria variegata (Valenciennes, 1832)
Gymnapogon annona (Whitley, 1936)
Gymnapogon philippinus (Herre, 1939)
Gymnapogon urospilotus Lachner, 1953
Gymnapogon vanderbilti (Fowler, 1938)
Lachneratus phasmaticus Fraser and Struhsaker, 1991
Neamia octospina Smith and Radcliffe, 1912
Pseudamia amblyuropterus (Bleeker, 1856)
Pseudamia gelatinosa Smith, 1955
Pseudamia hayashii Randall, Lachner, and Fraser, 1985
Pseudamia niger Allen, 1992
Pseudamia rubra Randall and Ida, 1993
Pseudamia zonata Randall, Lachner, and Fraser, 1985
Pseudamiops gracilicauda (Lachner, 1953)
0
0
Pterapogon kauderni (Koumans, 1933)
Pterapogon mirifica (Mees, 1966)
Rhabdamia cypselura Weber, 1909
Rhabdamia gracilis (Bleeker, 1856)
Rhabdamia spilota Allen and Kuiter, 1994
2608 Bony Fishes
Siphamia argentea Lachner, 1953

Siphamia argyrogaster (Weber, 1909)
Siphamia corallicola Allen, 1993
Siphamia cuniceps Whitley, 1941
Siphamia cuprea Lachner, 1909
Siphamia elongata Lachner, 1953
Siphamia fistulosa (Weber, 1909)
Siphamia fuscolineata Lachner, 1953
Siphamia guttulatus (Alleyne and Macleay, 1877)
Siphamia jebbi Allen, 1993
Siphamia majimae Matsubara and Iwai, 1958
Siphamia ovalis Lachner, 1953
Siphamia roseigaster (Ramsay and Ogilby, 1886)
Siphamia tubifer (Weber, 1909)
Siphamia tubulata (Weber, 1909)
Siphamia versicolor (Smith and Radcliffe, 1911)
Siphamia woodi (McCulloch, 1921)
0
0
Sphaeramia nematoptera (Bleeker, 1856)
Sphaeramia orbicularis (Kuhl and van Hasselt, 1828)
References
Allen, G.R. 1993. Cardinalfishes (Apogonidae) of Madang Province, Papua New Guinea, with descriptions of three new
species. Revue fr. Aquariol., 20(1):9-20.
Fowler, H.W. and B.A. Bean. 1930. Contributions to the biology of the Philippine Archipelago and adjacent regions. The
fishes of the families Amiidae, Chandidae, Duleidae, and Serranidae, obtained by the United States Bureau of
Fisheries steamer “Albatross” in 1907 to 1910, chiefly in the Philippine Islands and adjacent seas. US Nat. Mus.
Bull., 100(10):1-388.
Apogon cyanosoma Bleeker, 1853

En - Yellowstriped cardinalfish.
Maximum total length about 6 cm. Coral reefs of lagoons and seaward slopes, usually in clear water
at depths to about 20 m. Usually in small to large aggregations that shelter among or near live coral
during the day; feeds on small shrimps and crabs. Brooding males incubate up to 1 500 to 2 100
eggs at one time. Sometimes seen in the aquarium trade; captured with hand nets or surround nets.
Widespread in the Indo-Pacific from the Red Sea to Australia and the Marshall Islands, northward
to Japan.
Pterapogon kauderni (Koumans, 1933)

En - Banggai cardinalfish.
Maximum total length about 8 cm. Sheltered inshore sandy areas with seagrass at depths to 16 m.
Usually in small to large aggregations. The young shelter among the spines of Diadema sea urchins
or in sea anemones; feeds on small shrimps and crabs, also zooplankton. Brooding males incubate
up to 10 to 15 eggs at one time; lacks a pelagic larval stage and is the only marine fish that orally
broods young. A very popular, but high priced, aquarium fish due to its beauty and unusual biology.
Has been bred in captivity. Known only from the Banggai Islands off central-eastern Sulawesi.
Pterapogon mirifica (Mees, 1966)

En - Sailfin cardinalfish.
Maximum total length about 13 cm. Coral reefs of lagoons and sheltered inshore areas at depths
to about 15 m. Occurs solitarily or in pairs; shelters among live coral or reef crevices during the day;
feeds on small shrimps and crabs. Sometimes seen in the aquarium trade; captured with hand nets
or surround nets. Known only from northwestern Australia.
2610 Bony Fishes
Sphaeramia nematoptera (Bleeker, 1856)

En - Pajama cardinalfish.
Maximum total length about 8 cm. Coral reefs of lagoons and sheltered inshore areas at depths to
about 12 m. Usually in small aggregations that shelter among live coral during the day; feeds on
small benthic shrimps and crabs, also zooplankton. A popular aquarium fish; captured with hand
nets or surround nets. Western Indonesia to the northern Great Barrier Reef and northward to the
Ryukyu Islands.
Sphaeramia orbicularis (Kuhl and van Hasselt, 1828)

En - Coral cardinalfish.
Maximum total length about 10 cm. Sheltered inshore areas including mangroves, rock formations,
piers, and debris to depths of about 2 or 3 m. Usually in small to large aggregations that shelter in
the shade during the day; feeds mainly on crabs, floating insects, and copepods. Brooding males
incubate up to 11 000 eggs at one time. Sometimes seen in the aquarium trade; captured with hand
nets or surround nets. Widespread in the Indo-Pacific from East Africa to New Caledonia and
Kiribati, northward to the Ryukyu Islands.
Perciformes: Percoidei: Epigonidae 2611
EPIGONIDAE
Deepwater cardinalfishes
by O. Gon
D iagnostic characters: Small to medium-sized fishes (to about 50 cm). Body varies from elongate and
subcylindrical or compressed, to short and stocky. Eyes large, round to oval; margin of infraorbital
bones smooth (or infraorbital bones 1 to 4 serrate in Sphyraenops). Opercle with 1 or 3 (Sphyraenops)
spines, weak (rarely absent) to stout; posterior edge of opercular bones smooth, rarely poorly ossified, or
serrate (Florenciella, Rosenblattia, and Sphyraenops). Mouth large, oblique; maxilla narrow, not
reaching beyond level of middle of eye. Teeth in jaws, vomer, and palatines usually small, conical, in 1
to several series (palatines of Epigonus parini toothless); in some species enlarged caniniform teeth
protruding forward at tip of lower jaw (E. glossodontus) or both jaws (Florenciella and Rosenblattia). Two
separate dorsal fins, the first with VI to VIII spines, the second with a single spine and 8 to 11 soft
rays; anal fin with I to III spines and 7 to 10 soft rays; caudal fin emarginate to forked; pectoral-fin rays
14 to 23. Branchiostegal rays 7 (or 6 in Sphyraenops). Scales weakly to strongly ctenoid, and deciduous
to firmly attached; lateral line complete and extending onto caudal fin, with 33 to 56 tubular scales
(counted to end of hypural plates). Vertebrae 10-11+14-15=25. Colour: reddish brown to blackish.
eyes large lateral line extends

onto caudal fin
(after Mayer, 1974)
Habitat, biology, and fisheries: Contains 5 or 6 genera with about 30 species. Epigonus, with 25 species,
is the largest genus. Engybenthic, found around the world on continental and insular slopes, seamounts,
and oceanic rises, from northern cold-temperate to subantarctic waters, at depths of 75 to 3 700 m.
Carnivorous, feeding on planktonic organisms, including copepods, euphausiids, shrimps, and small
myctophids. Bycatch of trawl fisheries.
Similar families and genera in the area

Acropomatidae: II or III (never I) anal-fin spines; maxilla wide; lateral line not extending onto caudal fin;
canine teeth usually present; opercle usually with 2 spines.
Scombropidae: always III anal-fin spines; second dorsal fin and anal fin with 11 to 14 soft rays; maxilla
scaly, wide, and with large supramaxilla; jaws with large canines; scales cycloid, deciduous; lateral line
not extending onto caudal fin.
Acropomatidae Scombropidae
2612 Bony Fishes
Bathysphyraenops simplex (incertae sedis; provisionally placed in the Acropomatidae): always III anal-fin
spines; long pectoral fins, reaching beyond anal-fin origin; 6 branchiostegal rays; 5 pyloric caeca; maxilla
wide; opercle with 2 spines; other opercular bones each with a small spine; angle of preopercle serrate;
lateral line not extending onto caudal fin.
Howella brodiei (incertae sedis; provisionally placed in the Acropomatidae): always III anal-fin spines;
long pectoral fins, reaching beyond anal-fin origin; maxilla wide; lateral line interrupted, not extending onto
caudal fin; opercular bones armed with spines and/or serrae; scales large, ctenoid and adherent; no
caniniform teeth.
lateral line interrupted
(after Parr, 1933) (after Norman, 1930)

Bathysphyraenops Howella
Key to the species of Epigonidae occurring in the area serrate 3 opercular spines
1a. Three strong spines on opercle (Fig. 1); anal infraorbitals
fin with III spines and 7 soft rays; orbital edge
of infraorbitals 1 to 4 serrate (Fig. 1); bran-
chiostegal rays 6 . . . . . . . . . Sphyraenops bairdianus
1b. A single opercular spine, weak to strong; anal
fin with II spines and 9 (rarely 8 or 10) soft
rays; orbital edge of infraorbitals 1 to 4
smooth; branchiostegal rays 7 . . . . . . . (Epigonus) ® 2
2a. Opercular spine strong; first dorsal fin with VII Fig. 1 Sphyraenops bairdianus
spines; total gill rakers on first gill arch 21 to
(adapted from Suda and Tominaga, 1983)
23; pyloric caeca 9 to 11; greatest body depth
15 to 18.5% standard length; no intestinal light
organ . . . . . . . . . . . . . . . . Epigonus atherinoides
2b. Opercular spine weak; first dorsal fin with VIII spines; total gill rakers on first gill arch 17
to 21; pyloric caeca 7 or 8; greatest body depth 21 to 25.5% standard length; intestinal
light organ present . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Epigonus macrops

0 Epigonus atherinoides (Gilbert, 1905)
0 Epigonus macrops (Brauer, 1906)
0 Sphyraenops bairdianus Poey, 1860
References
Abramov, A.A. 1992. Species composition and distribution of Epigonus (Epigonidae) in the world ocean. J. Ichthyol.,
32(5):94-108.
Mayer, G.F. 1974. A revision of the cardinal fish genus Epigonus (Perciformes, Apogonidae), with descriptions of two
new species. Bull. Mus. Comp. Zool., 146(3):147-203.
Perciformes: Percoidei: Epigonidae 2613
Epigonus atherinoides (Gilbert, 1905)

En - Slender deepwater cardinalfish.
Maximum standard length at least 16 cm. Depth range 410 to 630 m. Occurs near the Philippines
(east of Luzon); also known from Hawaii.
(after Gilbert, 1905)
Epigonus macrops (Brauer, 1906)

En - Luminous deepwater cardinalfish.
Maximum standard length at least 21 cm. Depth range 550 to 1 300 m, usually from 640 to 920 m;
juveniles pelagic and found at 120 to 550 m. In the area, found off southern Viet Nam.
(after Uyeno et al., 1983)
Sphyraenops bairdianus Poey, 1860

En - Triplespine deepwater cardinalfish.
Maximum standard length at least 9.2 cm standard length. Caught at the surface (juveniles) and
between 380 and 1 600 m. Juveniles probably pelagic. Records from the northernmost part of the
area, the continental slope of northwestern Australia, and off Vanuatu, Tahiti, and French Polynesia.
click for next page

2614 Bony Fishes
Bony Fishes
Sillaginidae SILLAGINIDAE
Sillagos
by R.J. McKay
D iagnostic characters: Moderate-sized (to 51 cm), elongate perciform fishes. Opercle with small sharp
spine; lower part of preopercle horizontal. Snout long and 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 to 12 slender
spines; the second with I spine and 16 to 27 soft rays; anal fin long, with II weak spines and 14 to 27
soft rays. Scales small, ctenoid; lateral line slightly arched. Swimbladder frequently complex. Colour:
silvery to sandy grey or green, sometimes with black spots on body and pectoral-fin base.
2 dorsal fins, the 1st with IX-XII spines,

the 2nd with I spine and 16-27 soft rays
snout long and

conical
opercle with a small

sharp spine
anal fin with II spines and
14-27 soft rays
Habitat, biology, and fisheries: The family is widespread throughout the Indian Ocean and the western
Pacific Ocean. All species are inshore, bottom-dwelling, schooling fishes with a few species found in deeper
water to about 180 m. They are commonly taken by beach seine net and line in shallow sandy bays and
frequently enter estuaries, penetrating into fresh water for brief periods. Their flesh is very white, tender,
and of exceedingly delicate flavour. It contains very little oil and is easily digested. Steamed sillago fillets
are highly esteemed as food for invalids and infants. From 1990 to 1995, the FAO Yearbook of Fishery
Statistics reports a range of yearly catch of around 12 100 to 16 600 t from the Western Central Pacific.

Branchiostegidae: a single continuous dorsal fin; mouth large with fleshy lips.
Pinguipedidae (= Parapercidae, Mugiloididae): dorsal-fin spines short; spinous dorsal fin sometimes joined
to soft dorsal fin; pelvic-fin base in advance of pectoral-fin base.
a single dorsal fin
Branchiostegidae Pinguipedidae
Perciformes: Percoidei: Sillaginidae 2615
Key to the species of Sillaginidae occurring in the area

1a. Pelvic-fin spine very small and
situated at base of a thickened
club-shaped outer pelvic-fin ray pelvic-fin
(Fig. 1a); swimbladder re- spine
duced, no duct-like process
club-shaped outer
(Fig. 1b); no modified caudal pelvic-fin ray
vertebrae present (subgenus
Sillaginopodys) . . . . . Sillago chondropus a) pelvic fin b) swimbladder
1b. No club-shaped outer pelvic-fin Fig. 1 Sillago chondropus

ray . . . . . . . . . . . . . . . . . . . . → 2
2 posterior extensions
2a. Swimbladder divided posteriorly into 2
tapering extensions projecting below
vertebral column into tail musculature
(Fig. 2) . . . . . . . . . . . . (subgenus Sillago) → 3
2b. Swimbladder with a single posterior ex-
tension below vertebral column and en-
tering the tail section (Figs 3 and 4) . . . Fig. 2 swimbladder (subgenus Sillago)
. . . . . . . . . . . . . . (subgenus Parasillago) → 4
3a. Body with a longitudinal row of dark spots below lateral line and a series of dark
saddle-like blotches on back . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago intermedius
3b. Body uniform in coloration . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago sihama
4a. A dark brown, dusky, or blue-black spot or blotch on or just preceding pectoral-fin base . . . . . → 5
4b. No dark mark at pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 8
5a. Body with dark blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 6

5b. Body uniform colour in adults (juveniles to 9 cm have darker blotches along sides and
back) with snout bluish in some large specimens; first dorsal fin with XI spines, second
dorsal fin with I spine and 16 to 18 soft rays; anal fin with II spines and 15 to 17 soft rays;
lateral-line scales 60 to 69; vertebrae 14-15 + 5-8 + 11-14 (total 32 to 34); swimbladder
with rudimentary tubules projecting anteriorly and a series of sawtooth-like pockets
laterally (Fig. 3a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago ciliata
(eastern Australia)
6a. Upper and lower dark blotches on sides joined at least posteriorly; first dorsal fin with XI
spines, second dorsal fin with I spine and 19 to 21 soft rays; anal fin with II spines and
19 or 20 soft rays; lateral-line scales 71 to 75; vertebrae 34 to 36; swimbladder with a
short median extension anteriorly and a complex anterolateral extension that continues
posteriorly to the duct-like process (Fig. 3b) . . . . . . . . . . . . . . . . . . . . . Sillago maculata
(eastern Australia)
6b. Upper and lower dark blotches separate; swimbladder without complex anterolateral
extensions extending well posteriorly to end of abdomen . . . . . . . . . . . . . . . . . . . . . → 7
anterolateral extensions continues

rudimentary tubules projecting anteriorly short median posteriorly to duct-like process
extension anteriorly
a) Sillago ciliata b) Sillago maculata
Fig. 3 swimbladder
2616 Bony Fishes
7a. Swimbladder with 4 rudimentary anterolateral extensions that are normally convoluted
(Fig. 4a); usually 19 or 20 anal-fin rays; usually 35 vertebrae . . . . . . . . . . . . . . Sillago burrus
(northern coast of Australia, southern New Guinea, and Indonesia)
7b. Swimbladder with 3 rudimentary anterolateral extensions that may be simple or convo-
luted (Fig. 4b); usually 18 anal-fin rays; vertebrae 34 . . . . . . . . . . . . . . . . . . Sillago aeolus
(western Pacific, except Australia and southern New Guinea)
8a. Swimbladder with anterolateral extensions recurved posteriorly (Fig. 4c) . . . . . . . Sillago asiatica
8b. Swimbladder without posteriorly recurved anterolateral extensions . . . . . . . . . . . . . . . . ® 9
1 anterior anterolateral extensions
4 rudimentary anterolateral extensions 3 rudimentary anterolateral extensions recurved posteriorly
extension
a) Sillago burrus b) Sillago aeolus c) Sillago asiatica
Fig. 4 swimbladder
9a. Lateral-line scales 61 or less . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
9b. Lateral-line scales 64 or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
10a. Anal-fin rays 14 to 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago analis
(southern New Guinea and northern Australia)
10b. Anal-fin rays 19 to 21 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago macrolepis
(Indonesia to Philippines, excluding Australia)
11a. Anal-fin rays 21 to 24; 5 scales between lateral line and base of first dorsal-fin spine . . Sillago lutea
(India and Sri Lanka to northern Australia)
11b. Anal-fin rays 17 or 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Dorsal-fin rays 17; cheek scales ctenoid; vertebrae 13 + 9-11 + 9-11 (total 33); eye
diameter 19 to 23% of head length; no wide brilliant silvery longitudinal band on side . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago ingenuua
(Thailand, India, and northern Australia)
12b. Dorsal-fin rays 17 or 18; cheek scales with upper row cycloid and lower 2 rows ctenoid;
eye diameter 28 to 29% of head length; a wide brilliant silvery longitudinal band on side
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Sillago argentifasciata
(Lumbucan Island, Philippines)
The symbol 0 is given when species acounts are included.
0
0
Sillago aeolus Jordan and Evermann, 1902
0
Sillago analis Whitley, 1943
0
Sillago argentifasciata Martin and Montalban, 1935
0
Sillago asiatica McKay, 1983
0
Sillago burrus Richardson, 1842
0
Sillago chondropus Bleeker, 1849
0
Sillago ciliata Cuvier, 1829
0
Sillago ingenuua McKay, 1985
0
Sillago intermedius Wongratana, 1977
0
Sillago lutea McKay, 1985
0
Sillago macrolepis Bleeker, 1859
0
Sillago maculata Quoy and Gaimard, 1824
Sillago sihama (Forsskål, 1775)
Reference
McKay, R.J. 1992. FAO species catalogue. Vol. 14. Sillaginid fishes of the world. (Family Sillaginidae). An annotated
and illustrated catalogue of the sillago, smelt or Indo-Pacific whiting species known to date. FAO Fish. Synop.,
(125)Vol.14:87 p.
Sillago aeolus Jordan and Evermann, 1902

Frequent synonyms / misidentifications: None / Sillago maculata Quoy and Gaimard, 1824.
FAO names: En - Oriental sillago; Fr - Pêche-madame oriental; Sp - Silago oriental.
(from Martin and Montalban, 1935)
Diagnostic characters: First dorsal fin with XI spines;

second dorsal fin with I spine and 18 to 20 soft rays; anal
fin with II slender spines and 17 to 19 (usually 18) soft
rays. Lateral line with 67 to 72 scales. Vertebrae 13-14 +
4-7 + 14-16 (total 34). Swimbladder with 3 rudimentary
anterolateral extensions instead of 4; differs from swimbladder
Sillago maculata in lacking well-developed anterolateral
extensions reaching to level of anus. Colour: very similar
to S. burrus in coloration but has the most posterior
midlateral dark brown blotch elongate and reaching caudal
flexure.
Habitat, biology, and fisheries: Coastal inshore waters, most common in embayments on silty bottom.
Juveniles burrow in the sand. Small local fisheries exist throughout the range of this species, particularly
where bottom trawls are employed. The flesh is prone to spoil rapidly and the Oriental sillago is not
considered as highly as the inshore sillago’s.
Distribution: S i n g a p o r e ,
Thailand, China (including
Hong Kong and Taiwan Prov-
ince), Philippines, and south-
ern Japan. Possibly dis-
tributed throughout the Indo-
West Pacific from Delagoa
Bay (South Africa) to Oki-
nawajima (Japan), but not re-
corded from Australia or
southern New Guinea.
2618 Bony Fishes
Sillago analis Whitley, 1943

Frequent synonyms / misidentifications: Sillago nierstraszi Hardenberg, 1941 (possible senior
synonym) / None.
FAO names: En - Goldenlined sillago; Fr - Pêche-madame doré; Sp - Silago aureolineado.

second dorsal fin with I spine and 16 to 18 soft rays; anal fin
with II slender spines and 14 to 17 soft rays. Lateral line
with 54 to 61 scales. Vertebrae 13-14 + 4-8 + 11-15 (total
33 to 34). Colour: no dark spot at pectoral-fin base; body
light silvery, slightly darker to dusky above; a dull
golden-silver to golden-yellow band longitudinally on sides swimbladder
below lateral line; pelvic and anal fins pale yellow to bright
yellow; pectoral fins with a darker dusting of fine black-brown
spots.
Size: Maximum standard length 45 cm; commonly to 35 cm.
Habitat, biology, and fisheries: Shallow mangrove creek
shorelines, protected inlets and tidal areas. An important
fishery has developed in northern Australia.
Distribution: S h a r k B a y ,
Western Australia, Northern
Territory, Queensland south
to Moreton Bay, and southern
coast of New Guinea.
Sillago argentifasciata Martin and Montalban, 1935

FAO names: En - Silverbanded sillago; Fr - Pêche-madame ceinture d’or; Sp - Silago de bandas
plateadas.
(from Martin and Montalban, 1935)
Diagnostic characters: First dorsal fin with XI spines; second dorsal fin with I spine and 17 or 18 soft
rays; anal fin with II slender spines and 17 soft rays. Lateral line with 66 scales. Cheek with 3 rows of
scales, those on upper row cycloid, and on lower 2 rows ctenoid. Colour: no irregular dark blotches
on sides; a wide, brilliant, silvery, longitudinal band on each side of body.
Size: Not known.
Habitat, biology, and fisheries: Unknown.
Distribution: Lumbucan Island, Philippines.
2620 Bony Fishes
Sillago asiatica McKay, 1983

FAO names: En - Asian sillago; Fr - Pêche-madam asiate; Sp - Silago asiatico.

second dorsal fin with I spine and 20 or 21 soft rays; anal
fin with II slender spines and 21 to 23 soft rays. Lateral line
with 67 to 70 scales. Vertebrae 13-14 + 6-7 + 13-16 (total
34). Swimbladder with 3 anterior extensions, the middle
one projecting forwards and the anterolateral ones swimbladder
recurved backwards up to 1/2 length of swimbladder;
a single posterior extension. Colour: head and body
pale sandy brown to light fawn, an indistinct pale midlateral band is present on some specimens; belly
paler, almost white; opercle and preopercle transparent with a crescentic patch of fine black-brown spots
in a pigmented area, the shape of the gill arches on the inside of the gill cover showing through; fins hyaline,
the margins of unpaired fins finely spotted with brown; upper and lower margins of caudal fin dark brown
to almost black.
ies: Inshore waters. Biology
unknown. Taken as an inci-
dental catch with other sillago
species.
Distribution: Gulf of Thailand
and Taiwan Province of
China. This species is possi-
bly widespread.
Sillago burrus Richardson, 1842

Frequent synonyms / misidentifications: Sillago maculata burra Richardson, 1842 / None.
FAO names: En - Western trumpeter sillago; Fr - Pêche-madam clairon; Sp - Silago tromperero
occidental.
(after Richardson, 1843)

fin with II slender spines and 18 to 20 (rarely 18) soft rays.
Lateral line with 69 to 76 scales. Vertebrae 13-14 + 20-22
(total 34 to 36, rarely 34). Swimbladder with 4 anterolat- swimbladder
eral extensions that are more complex than S. aeolus, but
much shorter than Sillago maculata. Colour: very similar
in coloration to S. aeolus and S. maculata but in the latter species the upper and lower blotches are
frequently joined, at least posteriorly; the upper blotches are generally larger; the black spot at the base
of the pectoral fins is more distinct; the belly is not silver, and the operculum is dull or with the inner dark
blotch showing through (inside of operculum of S. burrus is white). S. aeolus is best distinguished from S.
burrus by means of vertebrae and anal-fin counts.
Habitat, biology, and fisheries: Sillago burrus is most abundant on silty-sand or muddy substrates.
Commonly taken by trawl net in shallow coastal waters to a depth of 32 m. Marketed fresh.
Distribution: Northern coast
of Australia, southern New
Guinea, and Indonesia.
2622 Bony Fishes
Sillago chondropus Bleeker, 1849

FAO names: En - Clubfoot sillago; Fr - Pêche-madame diablotin; Sp - Silago pateta.
Diagnostic characters: First dorsal fin with XI or XII

spines; second dorsal fin with I spine and 20 or 21 soft
rays; anal fin with II slender spines and 22 or 23 soft rays;
first pelvic-fin ray modified into a laterally compressed
thickened club-like structure. Lateral line with 66 to 73
scales. Vertebrae 12-13 + 22-23 (total 35). Swimbladder
reduced in size, no duct-like process from the ventral swimbladder
surface to the urogenital aperture.
Habitat, biology, and fisheries: Shallow coastal waters.
Captured by seine and marketed fresh.
Distribution: South Africa,
Mozambique, West Pakistan,
India, Myanmar, Indonesia,
New Guinea, Thailand, Philip-
pines, and Taiwan Province of
China. Not recorded from
southern New Guinea or Aus-
tralia.
Sillago ciliata Cuvier, 1829

FAO names: En - Sand sillago; Fr - Pêche-madame sable; Sp - Silago arena.
(after Grant, 1972)

fin with II slender spines and 15 to 17 soft rays. Lateral
line with 60 to 69 scales. Vertebrae 14-15 + 5-8 + 11-14
(total 32 to 34). Anterior part of swimbladder with rudimen-
tary tubules projecting anteriorly and a series laterally that
diminish in size and become sawtooth-like posteriorly;
shape of swimbladder not distinguishable from Sillago swimbladder
analis. Colour: a dark spot at base of pectoral fins;
coloration of adult specimens uniform without darker bars
or blotches.
Habitat, biology, and fisheries: An onshore species occurring on coastal beaches, sandbars, and surf
zones as well as open bays, estuaries, and coastal lakes. Feeds mostly on polychaetes and crustaceans.
An important commercial species and an esteemed angling fish in eastern Australia. Taken by beach seine
and tunnel-net. Marketed fresh.
Australia from Cape York,
Queensland (rare), south-
wards along the coast and the
Great Barrier Reef to eastern
Victoria, and the east coast of
Tasmania; also known from
Lord Howe Island, New Cale-
donia, and Woodlark Island
(Papua).
2624 Bony Fishes
Sillago ingenuua McKay, 1985

Frequent synonyms / misidentifications: Sillago argentifasciata (non Martin and Montalban, 1935) /
None.
FAO names: En - Bay sillago; Fr - Pêche-madame halanda; Sp - Silago de bahia.

second dorsal fin with I spine and 17 soft rays; anal fin
with II slender spines and 17 soft rays. Cheek scales
ctenoid. Lateral line with 66 to 70 scales. Vertebrae 13
+ 9-11 + 9-11 (total 33). Swimbladder with a short
median anterior extension and about 5 small pointed
anterolateral projections. Colour: no black spot on swimbladder
pectoral-fin base; no wide distinct silvery lateral
band; peritoneum black-brown.
Habitat, biology, and fisheries: Inshore coastal wa-
ters. Taken by prawn trawl. Marketed fresh.
Distribution: Known from In-
dia, Gulf of Thailand, Taiwan
Province of China, and north-
ern Australia from Shark Bay
around the northern coast to
Adolphus Passage in north-
eastern Queensland.
Sillago intermedius Wongratana, 1977

Frequent synonyms / misidentifications: None / Sillago maculata maculata Quoy and Gaimard, 1834.
FAO names: En - Intermediate sillago; Fr - Pêche-madame murda; Sp - Silago intermedio.
(after Wongratana, 1977)

second dorsal fin with I spine and 21 or 22 soft rays;
anal fin with II slender spines and 21 or 22 soft rays.
Lateral line with 67 to 70 scales. Vertebrae 14 + 5 + 15
(total 34). Swimbladder with 2 posterior extensions;
anterior margin with 2 divergent blind tubes that extend
to basioccipital above auditory capsule; an anterolat-
eral extension on each side, each sending a blind tubule
anteriorly and then curving posteriorly along abdominal swimbladder
wall as a simple tube to terminate just posterior to the
duct-like process. Colour: sides of body just below
lateral line with longitudinal row of dusky black
spots, and a series of saddle-like dusky black
blotches.
Habitat, biology, and fisheries: An inshore species on open silty bottom. Taken locally and marketed
fresh.
Distribution: Thailand and
India.
2626 Bony Fishes
Sillago lutea McKay, 1985

Frequent synonyms / misidentifications: None / Sillago macrolepis Bleeker, 1859.
FAO names: En - Mud sillago; Fr - Pêche-madame de vase; Sp - Silago de fango.
Diagnostic characters: First dorsal fin with X I

spines; second dorsal fin with I spine and 20 to 22 soft
rays; anal fin with II slender spines and 21 to 24
soft rays. Lateral line with 67 to 72 scales. Verte-
brae 13-14 (normally 13) + 4-11 + 10-17 (total 33 to
35). Swimbladder with median anterior extension and
with or without rudimentary anteriorly directed antero-
lateral projections; posterior extension single. Col-
our: body light sandy brown above, pale brown to
swimbladder
whitish below, with an indistinct silvery midlateral
band; margins of scales may be slightly darker, giving
a vague meshwork pattern to the body above lateral
line; fins hyaline, first dorsal-fin membrane tipped with
a fine dusting of black; no dark spot at pectoral-fin
base.
Habitat, biology, and fisheries: Inshore waters on muddy or very silty substrates. Large catches are
taken by prawn trawlers but as the species attains sexual maturity at 10 cm standard length (ripe females
10.4 to 12 cm) and grows to only 16 cm; the catch is of no commercial importance at present.
Distribution: Exmouth Gulf,
Western Australia, north-
wards and eastwards to Prin-
cess Charlotte Bay, Cape
York; also known from India
and Sri Lanka.
Sillago macrolepis Bleeker, 1859

FAO names: En - Largescale sillago; Fr - Pêche-madame grandes ecailles; Sp - Silago escamoso.
(after Bleeker, 1877)

fin with II slender spines and 19 to 21 soft rays. Lateral
line with 51 to 56 scales. Vertebrae 14 + 20 (total 34); no
haemal bridge overlying swimbladder. Swimbladder with-
out extensions. Colour: yellowish, darker above, with dif-
fuse silvery longitudinal midlateral band; dorsal fins dusky
with narrow blackish margin; juveniles with a series of
small brown spots, 1 on each side along back at base of swimbladder
dorsal fins; first dot at commencement of spinous dorsal
fin, second about middle of spinous dorsal fin, third below
fourth dorsal-fin ray, fourth below eleventh dorsal-fin ray,
and last spot below end of soft-rayed dorsal fin.
Habitat, biology, and fisheries: Enters estuaries and may penetrate fresh waters. No fishery exists for
this species.
Distribution: Recorded from
the Indonesian Archipelago,
New Britain, Solomon Is-
lands, and the Philippine Is-
lands.
2628 Bony Fishes
Sillago maculata Quoy and Gaimard, 1834

FAO names: En - Trumpeter sillago; Fr - Pêche-madame trompette; Fr - Silago trompetero.
(after Grant, 1972)
Diagnostic characters: First dorsal fin with XI

spines; second dorsal fin with I spine and 19 to 21
soft rays; anal fin with II slender spines and 19 or 20
soft rays. Lateral line with 71 to 75 scales. Vertebrae
13-15 + 8-11 + 10 to 14 (total 34 to 36). Anterolat-
eral extensions of swimbladder recurved poste-
riorly to reach level of anus. Colour: a black spot
at pectoral-fin base, dark blotches on back and swimbladder
side of body; the upper and lower blotches fre-
quently joined, at least posteriorly, the upper
blotches generally larger; operculum dull or with an
inner dark blotch showing through.
Habitat, biology, and fisheries: Found on silty and muddy substrates in the deeper water of bays, but
also frequenting the mouths of rivers, estuaries, and mangrove creeks. The juveniles are most abundant
in estuaries and shallow water during the summer months, moving into deeper water as they mature. Taken
by trawlers mainly as a bycatch of the prawn fishery. Marketed fresh.
Distribution: East Coast of
Australia.
Sillago sihama (Forsskål, 1775)

FAO names: En - Silver sillago; Fr - Pêche-madame argenté; Sp - Silago plateado.

second dorsal fin with I spine and 20 to 23 soft rays;
anal fin with II slender spines and 21 to 23 soft rays.
Lateral line with 66 to 72 scales. Vertebrae 14 + 2-8 +
12-18 (total 34). Swimbladder with 2 anterior exten-
sions extending forward and diverging to termi-
nate on each side of the basioccipital above the swimbladder
auditory capsule; 2 lateral extensions commence
anteriorly, each sending a blind tubule anterolat-
erally and then extending along the abdominal wall
below the investing peritoneum to just posterior of the duct-like process; 2 posterior tapering
extensions of the swimbladder project into caudal region, one usually longer than the other. The lateral
extensions are normally convoluted and have blind tubules arising along their length. Colour: body light
tan, silvery yellow-brown, sandy-brown, or honey coloured; paler brown to silvery white below; a midlateral,
silvery, longitudinal band normally present; dorsal fins dusky terminally with or without rows of dark brown
spots on the second dorsal-fin membrane; caudal fin dusky terminally; no dark blotch at base of pectoral
fins; other fins hyaline, the anal fin frequently with a whitish margin; after long preservation coloration may
become a uniform light brown.
Habitat, biology, and fisheries: A nearshore species inhabiting shallow water along beaches, sandbars,
mangrove creeks, and estuaries; very rarely captured by prawn trawling vessels. Captured by line, seine
net, cast-net, and traps operated from beaches. Marketed fresh and salted.
Distribution: A wide-ranging
species throughout the Indo-
West Pacific region from
Knysna (South Africa) to Ja-
pan.
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2630 Bony Fishes
BRANCHIOSTEGIDAE
(= Malacanthidae)
Tilefishes (also, quakerfish, blanquillos, burrowfishes, amadais, horseheads, and sand tilefishes)
by J.K. Dooley
D iagnostic characters (subfamily Branchiosteginae, genus Branchiostegus): Body quadriform and

square-headed; body depth 22 to 30% (usually 27%) standard length. Predorsal ridge (a raised
seam in front of dorsal fin) reduced, but always present. Predorsal length 27 to 37% (usually 32%)
standard length; head length 24 to 33% (usually 28%) standard length; head depth 82 to 108%
(usually 95%) head length; suborbital depth 14 to 35% (usually 19 to 25%) head length; orbit
diameter 20 to 37% (usually 24 to 30%) head length. Preopercle finely serrated on upper limb to
angle or just below, lower limb with few or no serrae; no enlarged spine at preopercular angle;
preopercle angle 85° to 115°; opercle with a single soft, blunt spine. Jaws slightly oblique, extending
posteriorly from in front of orbit under rear nostril to below rear rim of pupil; mouth terminal to slightly
inferior, jaws each side with 4 or 5 mandibular pores (usually 5). Total gill rakers on first gill arch 18 to
24. Dorsal and anal fins long and continuous; length of dorsal-fin base plus anal-fin base 80 to 97%
(usually less than 90%) standard length. Dorsal fin with VI to VIII (usually VII) spines and 14 to 16
(usually 15) soft rays. Anal fin with II spines and 11 to 13 (usually 12) soft rays. Caudal fin rounded,
truncate, or double emarginate (never lunate or forked), with 17 principal rays, sometimes with elongate
tips. Scales ctenoid (in pockets) over most of body, cycloid in head region. Pored lateral-line scales
usually 47 to 51 (67 to 72 only in B. serratus); scales above lateral line 6 to 11, below lateral line 16 to
31. Vertebrae 10+14; supraoccipital skull crest well elevated and elongate; well-formed foramen in
the ceratohyal; first haemal spine over second anal-fin ray with parapophyses fused medially
forming an arch for rear of swimbladder; ratio of anal-fin supports to haemal spines 0.86:1;
predorsal-fin supports always 0-0-2-; procurrent caudal-fin rays 10 or 11 (usually 10) in upper lobe, and
9 or 10 (usually 9) in lower lobe; 1 uroneural and parhypural, 3 epurals, and 2+3 autogenous hypurals.
Highly complex adductor mandibulae (jaw) musculature, with 5 major subdivisions (see figure of
jaw muscles below). Larvae pelagic, with numerous head spines and serrated ridges, no rostral head
spines, and without elongate fin spines.
A3b“ supraoccipital
skull antero-posteriorly crest well
A 3b compressed elevated
A1b A3 a
A2
Branchiostegus japonicus
deep jaw muscles, with overlaying subdivisional lateral view of skull
predorsal ridge (A1 a) removed (Branchiostegus serratus)
(modified from Marino and Dooley, 1982)
dorsal view
subfamily Branchiosteginae (Branchiostegus)
1st haemal spine with

parapophyses fused medially
lateral view
vertebrae postlarval stage (10 mm)
(Branchiostegus sawakinensis) (Branchiostegus japonicus)
Perciformes: Percoidei: Branchiostegidae 2631
Diagnostic characters (subfamily Malacanthinae, genera Hoplolatilus, Malacanthus): Body elongate,

fusiform or nearly cylindrical, with blunt or rounded snout; body depth 15 to 29% (usually 18 to
25%) standard length (Hoplolatilus ), or 12 to 20% (usually 14 to 17%) standard length (Malacanthus ).
No predorsal ridge. Predorsal length 19 to 34% (usually 22 to 30%) standard length; head length 19
to 32% (usually 21 to 29%) standard length; head depth 57 to 86% head length (Hoplolatilus ), or 49
to 64% head length (Malacanthus ); suborbital depth 3 to 7% head length (Hoplolatilus ), or 5 to 20%
head length (Malacanthus ); orbit diameter 22 to 31% head length (Hoplolatilus ), or 13 to 29% head
length (Malacanthus ). Preopercle serrate (Hoplolatilus ), or smooth (Malacanthus ); an enlarged
spine at angle of preopercle (Hoplolatilus ), or spine absent (Malacanthus ); opercle with single sharp
pointed spine (not found in Branchiosteginae: only a single soft blunt spine). Mouth terminal to
slightly inferior, jaws slightly oblique, extending posteriorly to eye or just under eye in Malacanthus , and
to posterior rim of orbit and beyond in Hoplolatilus ; jaws each side with 4 to 6 mandibular pores (4 in
Malacanthus , usually 5 or 6 in Hoplolatilus ). Total gill rakers on first gill arch 16 to 28 (usually 16 to
19, moderate to elongate) in Hoplolatilus , and 6 to 20 (usually 9 to 15, reduced and blunt) in
Malacanthus . Dorsal and anal fins long and continuous; length of dorsal-fin base plus anal-fin base
80 to 135% (usually more than 90%) standard length. Dorsal-fin spines III to X in Hoplolatilus , and
I to IV (usually II or IV) in Malacanthus ; dorsal-fin rays 13 to 34 (Hoplolatilus ), or 37 to 55
(Malacanthus ). Anal-fin spines I or II (I in Malacanthus ; usually II in Hoplolatilus , only H. cuniculus
with I); anal-fin rays 12 to 20 in Hoplolatilus , and 37 to 55 (usually 39 to 51) in Malacanthus . Caudal
fin falcate with rounded or pointed lobes, or truncate (may have elongate rays dorsally); caudal fin
with 17 principal rays. Scales ctenoid in pockets over most of body, mostly cycloid in head region; pored
lateral-line scales 89 to 141 in Hoplolatilus , 116 to 181 in Malacanthus ; scales above lateral line 10
to 20 in Hoplolatilus , 7 to 17 in Malacanthus ; scales below lateral line 34 to 49 in Hoplolatilus , 31 to
53 in Malacanthus . Vertebrae 10-11+14; supraoccipital skull crest much reduced to a small pointed
process; first haemal spine over first or second anal-fin ray in Hoplolatilus , and over anal-fin rays
12 to 18 in Malacanthus ; first haemal spine formed from parahypophyses fused only at their tips,
forming a broad elliptical arch (unlike Branchiosteginae where they are fused medially forming a
curved arch for rear of swimbladder); ratio of anal-fin supports to haemal spines 0.96 to 1.46
(Hoplolatilus ) or 2.8 to 3.9 (Malacanthus) ; predorsal-fin supports 0-0-2- ( Hoplolatilus ) or 0-0-1-, 0-1-,
or 2- (Malacanthus ); procurrent caudal-fin rays in upper fin lobe 10 to 12 (Malacanthus ) or 11 to 13
(Hoplolatilus ), and in lower lobe 9 to 13 (Hoplolatilus ) or 10 to 12 (Malacanthus ); a uroneural and
parhypural, 3 epurals, and 2+3 autogenous hypurals. Highly complex adductor mandibulae (jaw)
musculature but less complex than Branchiosteginae, with only 4 major subdivisions (lacking
subdivision A3ß) (cf. figure of jaw muscles on previous page). Larvae pelagic, with numerous enlarged
head spines and serrated ridges.
no predorsal ridge
sharp spines
subfamily Malacanthinae (Hoplolatilus)
no predorsal ridge
sharp spine
subfamily Malacanthinae (Malacanthus)
supraoccipital crest
skull elongate low, almost absent elongate
rostral spine
elongate
lateral view of skull
head of 15 mm larval opercular
(Malacanthus plumieri) spine
Hoplolatilus fronticinctus
2632 Bony Fishes
Habitat, biology, and fisheries: Tilefishes are a moderately deep, soft-bottom (mud or sand, or mixed
sand, mud, and shell) dwelling group. They are usually caught at depths of 60 to 200 m in the area. Usually
found living in burrows along the margins of continents or oceanic islands. The genus Branchiostegus is
most abundant in the eastern Pacific and Indian oceans and Red Sea (15 nominal species). One species
(B. semifasciatus) is also found in the eastern Atlantic off West Africa. The group all have pelagic larvae with
elaborate patterns of head and scale spination until metamorphosis takes place when they become benthic
prejuveniles (spination occurs only between 3 and 13 mm total length in B. japonicus). Tilefishes feed mainly
on invertebrates (molluscs, polychaete worms, shrimps, squids, small crabs), although they may feed on
small fish. They are very high quality food fishes and several species are commercially important. They are
caught by hook-and-line or bottom longline and in trawls. For example, the fishery of B. japonicus is one of
the more important fisheries of the East China Sea (Japanese landings have ranged from 6 000 to 12 000 t
per year). The South China Sea is the location of another important tilefish fishery. Total annual tilefish
landings for the Western Central Pacific probably exceed 25 000 t, but statistical data are not available for
the area. Tilefishes are marketed fresh, dried, salted, or canned.
Remarks: The tilefishes are comprised of 2 morphologically different and evolutionarily distinct groups
which are distinct enough to be considered as separate families (Branchiostegidae and Malacanthidae).
However, for reasons of conformity in this field guide, these groups are tentatively treated herein as
subfamilies following the classification in Nelson’s (1994) “Fishes of the world”, and Eschmeyer’s (1998)
“Catalog of fishes”. However, the family name Malacanthidae used in these works must then be considered
a junior synonym of Branchiostegidae. The subfamily Malacanthinae is also sometimes known as Latilinae.
Representatives of several families look superficially similar to tilefishes, but can be separated from them
as follows:
Coryphaenidae: dorsal fin extends forward to nape.
Pinguipedidae: always I anal-fin spine (versus I or II in Branchiostegidae).
Coryphaenidae Pinguipedidae
Nemipteridae: caudal fin always forked (usually not forked in Branchiostegidae, except some Hoplolatilus);
dorsal-fin spines X (versus VI to X); anal-fin spines III (versus I or II); anal-fin rays 7 or 8 (versus 11 to 55).
Serranidae: anal-fin spines III (versus I or II in Branchiostegidae); 1 to 3 opercular spines (versus a single
sharp pointed spine or a single soft blunt spine).
Nemipteridae Serranidae
Labridae: with prominant nipping canines.

Scaridae: mouth smaller; scales larger; lateral line discontinuous.
Labridae Scaridae
Lutjanidae: anal-fin spines III (versus I or II in Branchiostegidae); pointed head profile.

Caesionidae: caudal fin forked; mouth smaller; dorsal-fin spines X to XV (versus VI to X in Branchiostegi-
dae); anal-fin spines III (versus I or II).
Lutjanidae Caesionidae
Key to the subfamilies of Branchiostegidae occurring in the area

1a. Predorsal ridge present (may be as an enlarged flap); body depth 21 to 36% (usually 27%)
standard length; dorsal-fin elements (i.e. spines plus soft rays) 22 to 36; anal-fin elements
14 to 28; length of dorsal-fin base plus anal-fin base 80 to 109% (usually about 90%)
standard length; body shape robust or quadriform . . . . . . . . . . . . . . . . . . Branchiosteginae
(a single genus, Branchiostegus, occurring in the area)
1b. No predorsal ridge; body depth 12 to 26% (usually 27%) standard length; dorsal-fin
elements 22 to 64; anal-fin elements 14 to 56; length of dorsal-fin base plus anal-fin base
80 to 135% (usually over 90%) standard length; body elongate or fusiform . . . . . . . Malacanthinae
Key to the species of Branchiostegus occurring in the area (as modified from Dooley and Kailola, 1988)
Remarks on key characters: the 16 species of Branchiostegus presently known (14 in the area) are often
difficult to distinguish by single external characters except their living colour pattern. As their coloration does
not preserve well, identification becomes more difficult with the specimen’s age or preservation. Neverthe-
less, meristic data such as pored lateral-line scales and morphological characters such as head, body, and
fin proportions and internal characters are useful for species identification.
1a. Pored lateral-line scales 67 to 72 (modally 70); dorsal fin with VII (rarely VI) spines and
15 soft rays; body with 18 or 19 dark tapered vertical bars; operculum, predorsal ridge,
and area above pectoral-fin axil without dark pigment (Fig. 1) . . . . . . . . Branchiostegus serratus
(eastern Australia)
1b. Pored lateral-line scales 47 to 51; dorsal fin with VI spines and 16 (rarely 15) soft rays, or
with VII spines and 15 (rarely 14) soft rays; body with 16 to 20 dark tapered vertical bars,
light vertical bars, or no bars; operculum, predorsal ridge, with or without dark pigment . . . . . . ® 2
2a. Body with 6 or 7 rows of dark spots, 1 per scale, between lateral line and pectoral-fin base
(spots may be faint); a dark spot on each dorsal-fin membrane along base of dorsal fin;
caudal-fin margin double emarginate; lower half of caudal fin grey with yellow spots, and
usually with 2 thin yellow stripes in central part of caudal fin; anal fin pale with clear areas
between soft rays (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . Branchiostegus sawakinensis
(Red Sea, South Africa, Philippines, northern and northwestern Australia)
2b. Body without rows of dark spots; no dark spots along base of dorsal fin between each ray . . . . . ® 3
Fig. 1 Branchiostegus serratus Fig. 2 Branchiostegus sawakinensis

3a. Jaws do not reach past a vertical line through anterior rim of orbit . . . . . . . . . . . . . . . . . ® 4
3b. Jaws reach beyond a vertical line through anterior rim of orbit . . . . . . . . . . . . . . . . . . . ® 6
2634 Bony Fishes
4a. Preopercle indented above angle; predor-

sal ridge light coloured; caudal fin with nu-
merous yellow spots, yellow rays, and light
upper and lower margins; body whitish or
silvery with some underlying pink; dorsal
and anal fins clear with dark margin; broad
white pelvic fins; no markings on cheek or
snout (Fig. 3) . . . . . . . . . . Branchiostegus albus Fig. 3 Branchiostegus albus
(after Abe, 1965)
(South Korea, Japan, East China Sea, Taiwan Province of
China, South China Sea to Viet Nam)
4b. Preopercle not indented above angle; predorsal ridge dark coloured; caudal fin with
distinct yellow stripes and/or dark lower lobe and dark upper and lower margins; body light
pink or silvery; dorsal fin clear (may have a yellow or black margin); anal fin clear; pelvic
fins colourless and not broad; markings on cheek or snout may be present . . . . . . . . . . . . ® 5
5a. Caudal-fin margin double emarginate, the fin with dark lower lobe and 2 central yellow
stripes; 2 silvery or golden areas extending from suborbital rim to near upper jaw; snout
top of head, and upper body red; sides and belly silvery white; a black upper margin along
pectoral and soft dorsal fins; upper spinous dorsal-fin membrane with a thin yellow margin
(Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Branchiostegus australiensis
(known from 1 specimen collected in Shark Bay, Western Australia; not yet recorded from the area)
5b. Caudal-fin margin somewhat truncate, the fin with a dark lower lobe and a single central
dark stripe, remainder of fin yellowish; suborbital and snout bright yellow, the snout with
a triangular orange spot on lower medial portion; top of head and upper body dusky brown
(in alcohol); sides and belly silvery; no black margin along upper pectoral or soft dorsal
fins; spinous dorsal fin without yellow margin (Fig. 5) . . . . . . . . . . . . Branchiostegus ilocanus
[known only from Herre’s (1928) description of a single specimen from the Philippines]
Fig. 4 Branchiostegus australiensis Fig. 5 Branchiostegus ilocanus

(from Herre, 1928)
6a. Distinct suborbital bars extending to near or over upper jaw . . . . . . . . . . . . . . . . . . . . ® 7
6b. No distinct suborbital markings extending to near or over upper jaw . . . . . . . . . . . . . . . . ® 9
7a. A single narrow, pearly or silvery anterior subor-

bital bar extending to maxilla; head yellowish;
dorsal fin may have small dark spot between first
and second spines; upper dorsal-fin membrane
yellow, narrowing posteriorly; base of dorsal-fin
membrane also yellow; yellow dorsal-fin bands
separated by narrow white streaks and central
translucent areas; anal-fin membrane dusky with
small white blotches along base between each
ray; caudal fin double emarginate, its lower lobe
dusky with several small yellow spots and 2
central yellow stripes; upper caudal-fin margin
white near tip, with 3 or 4 radiating yellow Fig. 6 Branchiostegus auratus
stripes; upper margin of pectoral fins not dark; (after Abe, 1965)
pelvic fins yellowish with white anterior margin;
body reddish silver with no longitudinal stripes
(Fig. 6) . . . . . . . . . . Branchiostegus auratus
(East China Sea, Japan, South Korea, Taiwan Province of China, South China Sea; not yet recorded from the area)
7b. Two or 3 silvery or white-coloured suborbital bars; remaining colour not as above . . . . . . . . . ® 8
8a. Three suborbital bars: a single pearl-coloured band extending from anterior suborbital to
snout, a wider pearl-coloured bar extending from anterior suborbital to maxilla, a third
silvery bar extending from posterior suborbital down scaled portion of cheek to bran-
chiostegal membranes; snout bright pink; upper margin of spinous dorsal-fin membrane
black; upper margin of soft dorsal-fin membrane yellow, the basal portion pearly; caudal
fin subtruncate or somewhat double emarginate; 6 diagonal yellow stripes on upper
two-thirds of caudal fin; body silvery pink with large dark spot above lateral line near upper
opercular margin; predorsal ridge dark (Fig. 7) . . . . . . . . . . . . . . . . Branchiostegus vittatus
[known only from Herre’s (1926) description of a single specimen from the Philippines]
8b. Two suborbital bars: parallel silvery bars extending from below the orbit and narrowing to
over maxilla; head silvery; dorsal-fin membranes with a series of dark medial spots,
becoming larger on soft-ray membranes, remaining membranes pink or clear; distal half
of anal-fin membrane dusky, the proximal portion translucent (may have small elliptical
white spots); caudal fin double emarginate, lower two-thirds dusky with 4 or 5 yellow
longitudinal stripes; upper third of caudal fin pink with dark upper margin; upper margin of
pectoral fins dark; pelvic fins translucent with white anterior margin; body pink with 2 dark
stripes along entire length; predorsal ridge light coloured (Fig. 8) . . . . . Branchiostegus argentatus
(East China Sea, Japan, South Korea, Taiwan Province of China, South China Sea, Nhatrang, Viet Nam)
Fig. 7 Branchiostegus vittatus Fig. 8 Branchiostegus argentatus

(after Herre, 1926) (after Yoshino, 1984)
9a. Large triangular silvery marking from posterior orbital to midpreopercle; a silver triangular
marking at upper opercular opening; upper body red with several irregular yellow mark-
ings near midbody; silvery iris usually with dark blotch along upper rim; caudal-fin
membrane dusky, darker ventrally; 2 broad yellow parallel stripes in centre of caudal fin,
with 5 or 6 yellow stripes on upper lobe; upper margin of caudal fin dark (reddish) (Fig. 9)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Branchiostegus japonicus
(East China Sea, South Korea, Japan, South China Sea, Taiwan Province of China, South Viet Nam)
9b. No large silvery triangular marking on posterior orbital, nor at upper opercular opening;
body without irregular yellow markings middorsally; no dark blotch along upper rim of iris;
caudal-fin membrane light, dusky or dark ventrally; yellow stripes present or absent on
caudal fin; upper margin of caudal fin light or dark . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. A dusky yellow area on dorsal-fin membrane between first and third spines; caudal fin
yellow above, greenish grey below with 2 parallel yellow stripes in centre (Fig. 10) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Branchiostegus sp. A
(Okinawa, Japan; not yet recorded from the area)
10b. No dusky yellow area on dorsal-fin membrane between first and third spines; caudal fin
light above and light dusky or dark ventrally, yellow stripes present or absent . . . . . . . . . . ® 11
Fig. 9 Branchiostegus japonicus Fig. 10 Branchiostegus sp. A

(after Abe, 1965)
2636 Bony Fishes
11a. Predorsal ridge and surrounding area light (pale yellowish); dorsal and anal fins high (13%
and 12% standard length, respectively); faint vertical body bars; caudal fin more truncate,
the lower margin slightly emarginate, the fin without medial yellow stripes, but with single
grey stripe (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Branchiostegus gloerfelti
(eastern Indian Ocean: western central Sumatra; not yet recorded from the area)
11b. Predorsal ridge entirely dark or dark with a pale medial seam; dorsal-fin height 9 to 11%
standard length; anal-fin height 7 to 9% standard length; may have faint reddish body
markings corresponding to underlying myomeres; caudal-fin margin double emarginate,
the fin with 2 or more medial yellow stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Caudal fin with black triangular patch on lower lobe, the lower margin light; upper caudal-fin
lobe yellow with some grey, the upper margin dusky, central part of fin with 2 broad yellow
stripes; body depth 23 to 26% (usually 25%) standard length; jaws extend to a vertical line
drawn from posterior pupil rim; predorsal ridge dark (Fig. 12) . . . . . . . . . . . Branchiostegus wardi
(eastern Australia, Gulf of Papua to southern Queensland, Sydney, and New Caledonia)
12b. Caudal fin dusky, lacking black triangular patch on lower lobe, the lower margin dusky; upper
caudal-fin lobe light with dusky upper margin; caudal fin with several light yellow radiating
stripes corresponding to some of the rays; body depth 25 to 27% standard length; jaws extend
to under anterior or middle of pupil; predorsal ridge pale with dark edges . . . . . . . . . . . . . . . ® 13
Fig. 11 Branchiostegus gloerfelti Fig. 12 Branchiostegus wardi

13a. Interorbital space 22 to 24% head length; jaws extending posteriorly to midorbit; suborbital
depth 21% head length; predorsal ridge low, pale centrally with dark edges; preopercular
angle about 90° (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Branchiostegus paxtoni
(northwest of Port Hedland, Western Australia; not yet recorded from the area)
13b. Interorbital space 28 to 29% head length; jaws extending posteriorly to under anterior
edge of pupil; suborbital depth 19 to 21% head length; predorsal ridge very prominent and
dark; preopercular angle about 110° (Fig. 14) . . . . . . . . . . . . . . Branchiostegus hedlandensis
(north of Port Hedland, Western Australia; not yet recorded from the area)
Fig. 13 Branchiostegus paxtoni Fig. 14 Branchiostegus hedlandensis
Key to the genera of Malacanthinae occurring in the area

1a. Body depth 12 to 20% standard length (usually 14 to 17%); head depth 49 to 64% head
length; suborbital depth 5 to 20% head length; preopercle smooth (no serrae), no spine
at angle; opercle with a single sharp spine; no predorsal ridge; dorsal fin with I to IV spines
and 43 to 60 soft rays; a single anal-fin spine; length of dorsal-fin base plus anal-fin base
112 to 135% (usually 125%) standard length; total gill rakers on first gill arch 6 to 20
(usually 9 to 15); pored lateral-line scales 116 to 181; predorsal-fin supports in the
formulae 0-0-1-, 0-1-, or 2-; first haemal spine fused only at tip and positioned over anal-fin
rays 12 to 18; vertebrae 10+14; all inhabit burrows . . . . . . . . . . . . . . . . . . . . Malacanthus
1b. Body depth 15 to 29% standard length (usually 18 to 25%); head depth 57 to 86% head
length; suborbital depth 3 to 7% head length; preopercle serrate, spine at angle (may be
very reduced or prominent); opercle with sharp spine at angle; no predorsal ridge; dorsal
fin with III to V (or VIII to X) spines and 13 to 34 soft rays; anal-fin spines I or II; length
of dorsal-fin base and anal-fin base 80 to 100% (usually 90%) standard length; total gill
rakers on first gill arch 16 to 28; pored lateral-line scales 89 to 140; predorsal-fin supports
in the formula 0-0-2-; first haemal spine fused only at tip and positioned over second
anal-fin ray; vertebrae 10 or 11+14; inhabit burrows except 2 known species (H. fron-
ticinctus and H. starcki live in self-constructed mound of coral rubble) . . . . . . . . . . Hoplolatilus
Key to the species of Hoplolatilus occurring in the area

(modified from Randall and Dooley, 1974, Klausewitz, et al., 1978, and Randall, 1981)
1a. Caudal fin emarginate or forked; vertebrae 10+14; total gill rakers on first gill arch 20 to
29; opercular spine smaller than pupil; width of maxilla less that diameter of pupil . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (subgenus Hoplolatilus) ® 2
1b. Caudal fin truncate, not emarginate or forked, its upper lobe with elongate rays; vertebrae
11+14; total gill rakers on first gill arch 16 to 19; opercular spine larger than pupil; width of
maxilla about equal to or greater than diameter of pupil . . . . . . . . (subgenus Asymmetrurus) ® 7
(a single representative of this subgenus, H. fourmanoiri, occurring in the area)
2a. Dorsal fin with X spines and 13 soft rays; anal fin with II spines and 12 soft rays; lateral-line
scales 89 to 97; an elongate projection of white skin along inner edge of clavicle under
operculum near pectoral-fin base; posterior margin of preopercle with 18 or less serrae . . . . . . ® 3
2b. Dorsal fin with III to IX spines and 18 to 34 soft rays; anal fin with I or II spines and 12 to
20 soft rays; lateral-line scales 85 to 140; without an elongate projection of white skin
along inner edge of clavicle under operculum near pectoral-fin base; posterior margin of
preopercle with 35 or more serrae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Lateral-line scales 85 to 92; 16 to 20 coarse preopercular serrae; a spine at preopercular

angle, about 1/2 as large to nearly as large as opercular spine; total gill rakers on first gill
arch 26 to 29; pectoral-fin rays 16 to 17; caudal fin deeply emarginate (forked); body
olive-brown with blue (brown upon preservation) saddle-shaped spot on caudal peduncle;
a broad blue area on side of cheek, chest, and abdomen; a narrow deep blue band across
snout extending nearly to eyes; dorsal and anal fins yellowish with pink margins; spinous
dorsal fin yellow; caudal fin mostly yellow, with faint thin medial white streak (Fig. 15) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplolatilus fronticinctus
(Mauritius; Madras, India; Luzon, Philippines; Palau Islands and Solomon Islands)
3b. Lateral-line scales 90 to 97; 14 to 16 coarse preopercular serrae; no enlarged spine at
preopercular angle; total gill rakers on first gill arch 20 or 21; pectoral-fin rays 16; caudal
fin forked; body salmon dorsally, white or bluish white ventrally; a yellow longitudinal stripe
may appear on back along dorsal-fin base, extending to caudal peduncle; snout and upper
head olive (may have some yellow); a yellow-orange bar from eye to mouth; cheek with 2
irregular (wavy) longitudinal blue bands separated by light yellow bands; dorsal and anal
fins light yellow; pectoral fins pale, pelvic fins pale or light yellow; upper and lower
caudal-fin lobes various colours (yellow, green, blue), the central area transparent; can
undergo a very rapid (24 colours in 15 seconds) colour change when frightened (Fig. 16)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplolatilus chlupatyi
(Philippines: Batangas, Luzon)
Fig. 15 Hoplolatilus fronticinctus Fig. 16 Hoplolatilus chlupatyi

2638 Bony Fishes
4a. Dorsal fin with III to V spines and 29 to 34 soft rays; anal fin with I spine and 19 or 20 soft
rays; lateral-line scales 116 to 140; body depth 5.2 to 6.3 times in standard length; caudal
fin forked; colour: body light olive-brown on back, fading to light yellow ventrally; a blue
area on dorsal and postorbital area of head (fades upon death); dorsal-fin base coloured
like upper body, remainder of fin bluish; caudal peduncle and upper and lower caudal-fin
lobes bright yellow (Fig. 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplolatilus cuniculus
(widespread Indo-Pacific: Tahiti, western Pacific to western Indian Ocean)
4b. Dorsal fin with VIII or IX spines and 18 to 23 soft rays; anal fin with I or II spines and 14
to 16 soft rays; lateral-line scales 95 to 121; body depth 3.8 to 5.5 times in standard length;
caudal fin deeply emarginate to forked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Dorsal fin with IX spines and 18 or 19 soft rays; anal fin with I spine and 15 or 16 soft rays;
total gill rakers on first gill arch 20 to 24; lateral-line scales 105 to 121; body depth 5.3 to
5.5 times in standard length; caudal fin deeply emarginate; colour: head and nape blue;
body mostly magenta fading to white on belly; dorsal portion of body and base of dorsal
fin brilliant blue; remainder of dorsal fin reddish orange; anal fin light orange with a pale
blue margin; upper and lower lobes of caudal fin deep magenta, central area blue fading
to magenta posteriorly (Fig. 18) . . . . . . . . . . . . . . . . . . . . . . . . Hoplolatilus purpureus
(Guadalcanal, Solomon Islands, Philippines, and Indonesia)
5b. Dorsal fin with VIII or IX spines and 16 to 23 soft rays; anal fin with I or II spines and 14
to 16 soft rays; total gill rakers on first gill arch 21 to 27; lateral-line scales 95 to 118; body
depth 3.7 to 5.1 times in standard length; caudal fin emarginate to forked . . . . . . . . . . . . . ® 6
Fig. 17 Hoplolatilus cuniculus Fig. 18 Hoplolatilus purpureus

6a. Dorsal fin with IX spines and 16 soft rays; anal fin with I spine and 14 soft rays; total gill
rakers on first gill arch 21 to 24; lateral-line scales 95 to 104; body depth 5 to 5.1 times in
standard length; caudal fin emarginate; colour: an orange-red band from tip of snout to
eye, arching dorsally to back then curving ventrally to middle of caudal peduncle, then
broadening into a triangular area and becoming much darker (reddish black); head and
body above band tan to red-brown, except on dorsal area of caudal peduncle where it is
white; head and body below band white to pink-white; dorsal and anal fins white; dorsal
with an red-orange margin, anal with a light blue margin; caudal fin white above and below
with a dark triangle (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hoplolatilus marcosi
(Luzon, Philippines, Guadalcanal and New Georgia, Solomon Islands)
6b. Dorsal fin with VIII spines and 21 to 24 soft rays; anal fin with II spines and 15 or 16 soft rays;
total gill rakers on first gill arch 22 to 27; lateral-line scales 100 to 118; body depth 3.7 to 4.6
times in standard length; caudal fin deeply forked; colour: body tan with a large bright blue area
over head area from below upper eye to thorax past base of pectoral fins; dorsal and anal fins
light yellow, the dorsal fin with some blue; caudal fin bright yellow except upper and lower and
central posterior margins which are whitish; young completely blue (Fig. 20) . . . . . Hoplolatilus starcki
(Mariana Islands, Caroline Islands, Palau Islands, Marshall Islands, and Tuamotu Archipelago)
Fig. 19 Hoplolatilus marcosi Fig. 20 Hoplolatilus starcki

7a. A prominent spine at preopercular angle, 1/2 or greater than length of opercular spine; dorsal
fin with X spines and 21 to 23 soft rays; anal fin with II spines and 18 or 19 soft rays; total gill
rakers on first gill arch 16 to 19; lateral-line scales 96 to 101; maxilla reaching a vertical at
posterior rim of orbit; colour: body grey, darker dorsally; top of head violet-black; large irregular
light (yellow) areas separated by dark areas; a large, nearly triangular black spot from caudal
peduncle extending to near central caudal-fin margin (Fig. 21) . . . . . . . . . Hoplolatilus fourmanoiri
(South Viet Nam, Guadalcanal, Solomon Islands, Philippines)
7b. A short broad-based spine at preopercular angle; dorsal fin with X spines and 22 soft rays;
anal fin with II spines and 20 soft rays; total gill rakers on first gill arch 17; lateral-line scales
92; maxilla reaching beyond a vertical at posterior rim of orbit; an irregular dark stripe
slightly above middle of side of body, ending at middle of caudal fin; an irregular row of
dark spots on body above stripe (Fig. 22) . . . . . . . . . . . . . . . . . . . . . . Hoplolatilus oreni
(known only from the type specimen collected in the Red Sea; not yet recorded from the area)
Fig. 21 Hoplolatilus fourmanoiri Fig. 22 Hoplolatilus oreni
Key to the species of Malacanthus occurring in the area

1a. Dorsal-fin elements (i.e. spines plus soft rays) 53 to 65; total gill rakers on first gill arch 9
to 20 (modally 15); broad midbody longitudinal band from operculum to tip of caudal fin,
widening on tail to include most of middle and lower portion, lower portion with a small
white rectangular area (Fig. 23) . . . . . . . . . . . . . . . . . . . . . . . Malacanthus brevirostris
1b. Dorsal-fin elements 46 to 51 (modally 49); total gill rakers on first gill arch 6 to 14; caudal
fin with characteristic 2 dark parallel bands (Fig. 24) . . . . . . . . . . . . Malacanthus latovittatus
Fig. 23 Malacanthus brevirostris Fig. 24 Malacanthus latovittatus

The symbol 0 is given when species accounts are included. A question mark indicates that presence in
0
Subfamily BRANCHIOSTEGIDAE
0
Branchiostegus albus Dooley, 1978
Branchiostegus argentatus (Cuvier, 1830)
? Branchiostegus auratus (Kishinouye, 1907) 1/
? Branchiostegus australiensis Dooley and Kailola, 1988 2/
? Branchiostegus gloerfelti Dooley and Kailola, 1988 3/
? Branchiostegus hedlandensis Dooley and Kailola, 1988 4/
1/ Marginal to the area: known from southern Japan and south coast of Korea to the East China Sea to Taiwan Province
of China and possibly northern portions of the South China Sea. Maximum standard length 29 cm. Found living along
with B. japonicus, B. argentatus , and B. albus within part of their ranges (East China Sea), although depth preferences
may differ somewhat (B. auratus usually caught at greater depths, 225 to 290 m, but at 30 to 290 m off Taiwan Province
of China).
2/ Not yet recorded from the area. Known only from the holotype, 26.6 cm standard length, collected southeast of Shark
Bay (26°21’S, 112°35’E), Western Australia at a depth of 164 to 170 m. Potentially commercial food fish.
3/ Not yet recorded from the area. Known only from the holotype, 23.5 cm standard length, trawled off central Sumatra
(about 00°22’S, 98°44’E). Might be expected elsewhere in Indonesia. A potentially commercial species.
4/ Not yet recorded from the area. Known only from Port Hedland (18°32’S, 118°17’E), Western Australia. Size range
22.5 to 26 cm standard length. Collected by bottom trawls at a depth of 200 to 204 m. A potentially commercial species.
2640 Bony Fishes
Branchiostegus ilocanus Herre, 1928 5/

0 Branchiostegus japonicus (Houttuyn, 1782) 6/
0 Branchiostegus sawakinensis Amirthalingam, 1969

? Branchiostegus paxtoni Dooley and Kailola, 1988
0 Branchiostegus serratus Dooley and Paxton, 1975

7/
0 Branchiostegus wardi Whitley, 1932

Branchiostegus vittatus Herre, 1926
? Branchiostegus sp. A from Okinawa, sensu Yoshino and Hayashi

Subfamily MALACANTHINAE
Hoplolatilus (Hoplolatilus) chlupatyi Klausewitz, McCosker, Randall, and Zetzsche, 1978
Hoplolatilus (Hoplolatilus) cuniculus Randall and Dooley, 1974
Hoplolatilus (Asymmetrurus) fourmanoiri Smith, 1963
Hoplolatilus (Hoplolatilus) fronticinctus Günther, 1887
Hoplolatilus (Hoplolatilus) luteus Allen and Kuiter, 1989
Hoplolatilus (Hoplolatilus) marcosi Burgess, 1978
? Hoplolatilus (Asymmetrurus) oreni (Clark and Ben Tuvia, 1973)
Hoplolatilus (Hoplolatilus) purpureus Burgess, 1978
Hoplolatilus (Hoplolatilus) starcki Randall and Dooley, 1974
0
0
Malacanthus brevirostris Guichenot, 1848
Malacanthus latovittatus (Lacepède, 1802)
References
Burgess, W.E. 1978. Two new species of tilefishes (family Branchiostegidae) from the western Pacific. Trop. Fish Hobb.,
26(263,5):43-47.
Dooley, J.K. 1978. Systematics and biology of the tilefishes (Perciformes: Branchiostegidae and Malacanthidae) with
descriptions of two new species. NOAA. Tech. Rept. NMFS Circ., (411):78 p.
Dooley, J.K. and P.J. Kailola. 1988. Four new tilefishes from the northeastern Indian Ocean, with a review of the genus
Branchiostegus. Japan. J. Ichthyol., 35(3):247-260.
Dooley, J.K. and J.R. Paxton. 1975. A new species of tilefish (family Branchiostegidae) from eastern Australia. Proc.
Linn. Soc. N. S. W., 99(3):151-156.
Dooley, J.K. and N. Rau. 1982. A remarkable tilefish record and comments on the Phillipine tilefishes. Japan. J.
Ichthyol., 28(4):450-452.
Hayashi, Y. 1976. Studies on the growth of the red tilefish in the East China Sea. II. Estimation of age and growth from
otolith readings. Bull. Japan. Soc. Sci. Fish., 42(11):1243-1250.
Kuiter, R.H. 1992. Tropical reef-fishes of the western Pacific. Indonesian and adjacent waters. Jakarta, Indonesia,
Penerbit Pt Gramedia Pustaka Utama, 314 p.
Marino, R.P. and J.K. Dooley. 1982. Phylogenetic relationships of the tilefish family Branchiostegidae (Perciformes)
based on comparative myology. J. Zool. Soc. London , 196:151-163.
Okiyama, M. (ed.). 1993. An atlas of the early stage fishes in Japan. Germany, Koeltz Sci. Books, pp. 452-454.
Randall, J.E. 1981. A review of the Indo-Pacific sand tilefish genus Hoplolatilus (Perciformes: Malacanthidae).
Aquarium, 4(12):39-46.
Randall, J.E. and J.K. Dooley. 1974. Revision of the Indo-Pacific branchiostegid fish genus Hoplolatilus , with
descriptions of two species. Copeia, 1974(2):457-471.
Yoshino, T., W. Hiramatsuu, O. Tabata, and Y. Hayashi. 1984. First record of the tilefish Branchiostegus argentatus
(Cuvier) from Japanese waters, with a discussion on the validity of B. auratus (Kishinouye). Galaxea, 3:145-151.
5/ This species has not been verified after the original description by Herre (1928), based on a 27 cm specimen (original
specimen lost) purchased from a market at Narvacan, Ilocos Sur Province (Luzon, Philippines). A recent (1995)
examination of fishes from numerous Philippine markets failed to reveal any specimens of Branchiostegus.
6/ Not yet recorded from the area. So far, known only from northwest of Port Hedland, Western Australia (19°01’S;
117°12’E). Size range 20.5 to 25.5 cm standard length. Caught in bottom trawls in depths of 200 to 202 m. Potential
commercial food fish.
7/ This species has not been verified after the original description by Herre (1926), based on a single specimen, 24 cm
standard length, collected from a Manila fish market (original specimen lost). A recent (1995) examination of fishes
from numerous Philippine markets failed to reveal any specimens of Branchiostegus. Caudal-fin margin of this species
may not be as rounded as drawn by Herre (see Fig. 7 in above key; figure redrawn from original).
Branchiostegus albus Dooley, 1978

Frequent synonyms / misidentifications: None / Branchiostegus argentatus (Cuvier, 1830); B. japonicus
(Houttuyn, 1782).
FAO names: En - White tilefish; Sp - Blanquillo de blanco.
(after Abe, 1965)
Diagnostic characters: Preopercle finely serrated to its angle with a slight indention above the angle
is about 105°. Eye high on head and proportionately small, 20 to 24% head length. Posterior margin
of jaw not quite or just reaching a vertical at anterior eye margin. Caudal fin double emarginate. Pelvic
fins comparatively broad, with rounded posterior margin. First haemal spine characteristically very
recurved. Colour: body without dark vertical bars; upper body silvery white with a pink hue (fresh
specimens); belly white; snout yellow-pink, cheek with a broad silvery band from suborbit to upper jaw;
dorsal-fin membrane translucent with some areas of white or pink; a dark band along margin of spinous
dorsal fin; pectoral fins translucent; pelvic fins milky white with some overlying dusky tint; predorsal ridge
prominent and pale yellowish; caudal fin dusky, lower lobe not dark, the rays yellow at base, the
membranes with small yellow spots, the dorsal and ventral edges mainly white.
Size: Maximum standard length perhaps 60 cm; commonly between 20 and 40 cm standard length.
Habitat, biology, and fisheries: Found on muddy or sand bottom, usually associated with burrows; depths
generally under 100 m. Caught by hook-and-line, bottom longlines, or trawls.
( H o n s h u ) , N o r th K o r ea
(Pusan), East China Sea (in-
cluding Shanghai and Taiwan
Province of China), South
China Sea (including Hong
Kong and Macao) to Viet Nam.
2642 Bony Fishes
Branchiostegus argentatus (Cuvier, 1830)

Frequent synonyms / misidentifications: None / Branchiostegus japonicus (Houttuyn, 1782); B. albus
Dooley, 1978.
FAO names: En - Silver tilefish; Sp - Blanquillo de plata.
(after Yoshino, 1984)
Diagnostic characters: Body depth shallow, 22 to 25% (usually 24%) standard length; body width
narrow, 10 to 12% standard length; predorsal length short, 28 to 31% (usually 30%) standard length.
Preopercle finely serrate only to angle, no indentation at angle as in Branchiostegus albus, lower limb
smooth; preopercular angle 85° to 95°. Suborbital shallow, 14 to 24% (usually 19%) head length (a
character shared in the area only with B. wardi, suborbital 16 to 22% head length, usually 19%). Eye
large, 25 to 36% (usually 30%) head length (proportionally largest among species of Branchiostegus).
Jaws extend posteriorly to nearly under a vertical from posterior pupil. Dorsal fin comparatively low,
8% standard length. Caudal fin usually double emarginate. Colour: body dusky above lateral line, silvery
below with a hint of yellow-orange stripes; belly white with a rose hue; numerous (2 or more
prominent) longitudinal orange stripes on dorsal part of body, only faint ventrally; head with 2
prominant pearly silvery bands from suborbital to upper jaws; predorsal ridge dark; dorsal-fin
membrane pink with a series of dark elliptical blotches dorsally (most prominant in rayed portion),
disappearing before posterior edge; a series of faint yellow blotches basally along dorsal-fin
membrane; anal fin whitish with a series of faint white spots basally, the distal half of fin dusky;
caudal fin with dark leading upper edge, pink underneath, the lower portion dusky with 6 bright
yellow stripes and several small spots below lowest stripe; upper edge of pectoral fins dark, and
dusky below; pelvic fins white anteriorly, transparent posteriorly; much of the coloration lost upon preser-
vation.
Habitat, biology, and fisheries: Found benthically in relatively shallow depths (between 50 and 65 m).
Caught by hook-and-line, bottom longlines, and trawls.
Distribution: Southern Japan
to Viet Nam (Nhatrang), includ-
ing Taiwan Province of China,
East China Sea, coast of
China, Hong Kong, and South
China Sea.
Branchiostegus japonicus (Houttuyn, 1782)

Frequent synonyms / misidentifications: None / Branchiostegus auratus (Kishinouye, 1907);
B. argentatus (Cuvier, 1830); B. albus Dooley, 1978.
FAO names: En - Red tilefish; Sp - Blanquillo de roja.
(after Abe, 1965)
Diagnostic characters: (Most meristic and morphometric data overlap other species of Branchiostegus in
the area.) Head large, 26 to 30% (usually 28%) standard length. Preopercle angle 90° to 95°, without
notch above angle. Eye large, 28% head length. Jaws slightly oblique, extending posteriorly to under
anterior third of pupil. Colour: body pink-red with overlying golden tint and underlying faint yellowish
blotches, several irregular light reddish blotches dorsally at midbody (appears as though mucus
coating has been lost in that area revealing the underlying golden reddish coloration); predorsal
ridge dark; head without suborbital silvery bars, but with characteristic large triangular postsubor-
bital silvery white area (may have another smaller triangular area at upper margin of operculum);
caudal fin with 5 or 6 bright yellow stripes (2 prominent midcaudal stripes plus 3 or 4 smaller, fainter
stripes dorsally on fin).
Size: Maximum recorded total length 37.8 cm; commonly between 20 and 33 cm standard length; maximum
weight estimated to be 485 g (females) and 714 g (males), for an estimated age of 8 years.
Habitat, biology, and fisheries: Found associated with burrows on sandy-muddy, or shell-sand-mud
bottoms in depths of 30 to 265 m; commonly caught at depths of 80 to 200 m in East China and Yellow seas.
Bottom temperatures 8° to 18°C (winter) 14° to 22°C (autumn). Spawning peaks appear to be twice a year
(July and October) in the East China Sea. Larvae appear in October off Sado Straits in Japan at depths of
0 to 50 m (19° to 21°C). Caught by bottom longlines and trawls. Catches have fluctuated from 500 t before
1956 to a peak of 12 460 t in 1970. Catches of the Red tilefish, although comprising 90% of all tilefish
catches in the East China Sea, have declined since 1970, averaging about 6 000 t in recent years. The
catch/effort in 1967 to 1980 declined from 9 to 4.4 fish per longline (1978 to 1980 catch per day averaged
about 270 kg per 120 longlines, with 130 hooks/longline), indicating the need for fishery management at
least for the male stock as males are generally larger, more voracious, and more easily caught. Marketed
fresh, canned, and salted.
Distribution: Sea of Japan
(central Honshu, Japan), por-
tions of the East China Sea,
South China Sea, and coast of
China to South Viet Nam and
waters around the Philippines.
Previous records from South
Africa are probably attribut-
able to either B. sawakinensis
or B. doliatus.
2644 Bony Fishes
Branchiostegus sawakinensis Amirthalingam, 1969

Frequent synonyms / misidentifications: None / Branchiostegus doliatus (Cuvier, 1830); B. japonicus
(Houttuyn, 1782).
FAO names: En - Freckled tilefish; Fr - Tile taches de son; Sp - Barko barko.
Diagnostic characters: Preopercle angle 95° to 100°. Jaws extending posteriorly to under middle of
pupil. Dorsal-fin height low, about 7% standard length. Possesses (along with only Branchiostegus
wardi) a unique arrangement of jaw muscles (adductor mandibulae complex) in having subdivisions
of the A1b(cf. figure of jaw muscles on family sheet). Colour: body with varied reddish and yellow
pastel shades and 5 to 7 rows of dark spots from below lateral line to beneath pectoral fins;
rose-violet around eye and silvery yellow below; predorsal ridge darkly pigmented; a golden area above
opercle; dorsal fin with a small dark rectangular markings along its base; anal-fin membrane with small,
silver-white triangles along its base; upper portion of caudal fin light with yellow rays, lower portion dark grey
with 10 to 15 small yellow spots (noted on Philippine specimens).
Size: Maximum standard length 39 cm, commonly to 25 cm standard length; preserved weight 1 kg.
Habitat, biology, and fisheries: Found over muddy bottoms at 80 m in the Red Sea (called locally “shawra”
or “theena” and regarded to be occasionally poisonous there), 45 m over sandy bottom from South Africa,
and captured by trawls at 120 to 180 m over sandy bottom off the Philippines. Not common in the markets
of Cebu City (Philippines). Probably inhabits burrows. Captures by hook-and-line, and by trawls. Marketed
fresh or salted.
Distribution: Recent records
from the Philippines and
northwestern Australia. Also
disjunctly known from the
western Indian Ocean to
South Africa, where it has
been erroneously identified as
B. japonicus or B. doliatus.
May be widespread through-
out the Indian Ocean and
western Central Pacific.
Branchiostegus serratus Dooley and Paxton, 1975

Frequent synonyms / misidentifications: None / Branchiostegus wardi Whitley, 1932.
FAO names: En - Australian barred tilefish.
Diagnostic characters: Snout profile steep, nearly vertical. Jaws oblique reaching posteriorly to below
centre of pupil. Total gill rakers on first gill arch 18 to 20. Caudal fin double emarginate. Pored lateral-line
scales 67 to 72 (highest among all species of Branchiostegus). Colour: body silvery white with
overlying yellow-orange and 18 or 19 dark violet tapering bars extending from back to below lateral
line; no bars under orbit; predorsal ridge light yellow; snout yellow-orange; iris golden; cheeks
silver-white with an oblique darker band on preopercle; spinous dorsal fin dusky with a narrow
yellow upper margin; soft dorsal-fin membrane with a series of elliptical spots, fading posteriorly;
anal-fin membrane translucent; pectoral fins opaque with an upper black edge (first 2 rays); pelvic
fins translucent; caudal fin with wide central yellow area; dorsal, ventral, and posterior caudal-fin
margins black.
Size: Maximum standard length 41 cm; commonly between 25 and 30 cm standard length.
Habitat, biology, and fisheries: Bottom-dwelling, well documented from depths of 110 to 150 m; maximum
depth recorded 162 m. Shallower eastern Australian trawling surveys (60 to 110 m) conducted in the
Australian area earlier, failed to catch any specimens of B. serratus. Often caught in the same trawl samples
with B. wardi (another eastern Australian species not known until collected in 1928). Both species are now
common in the Sydney fish markets. Stomach analysis revealed a wide variety of food items: fish remains,
bivalves, gastropods, crabs, amphipods, stomatopods, and polychaete worms. Deviations of the expected
50/50 sex ratio suggest protogynous sex reversal (other explanations possible); specimens under 38 m
standard length exhibit a 50/50 sex ratio, while larger specimens are generally males. Ovaries collected in
June were filled with 0.2 to 0.9 mm ova. About 25 kg per day can be found in the Sydney market. Marketed
fresh.
Cape Moreton (Queensland)
and the coast of New South
Wales (Australia) from Coff’s
Harbour 30°20’S, to Lake Il-
lawarra 34°30’S.
2646 Bony Fishes
Branchiostegus wardi Whitley, 1932

Frequent synonyms / misidentifications: None / Branchiostegus serratus Dooley and Paxton, 1975.
FAO names: En - Ward’s tilefish.
Diagnostic characters: Suborbital shallow, 16 to 22% (usually 19%) head length, a character in the
area shared only with Branchiostegus argentatus (suborbital 14 to 24% head length, usually 19%). Snout
blunt. Pored lateral-line scales 48 to 51 (unlike 67 to 72 in B. serratus); caudal fin double emarginate.
Colour: body without vertical bars, dark brown dorsally, silver-white ventrally; snout purple; top of
head dark; predorsal ridge dark; cheeks silver without suborbital bars; spinous membrane of dorsal fin
light to dusky, with some yellow; soft dorsal-fin membrane dusky, without distinct markings; pectoral
fins with dark dorsal margin, upper portion somewhat dusky, the lower clear; pelvic fins dusky; caudal fin
with dark triangle on lower lobe, the dorsal and ventral edges yellow; 2 parallel medial yellow bands
separated by medial dusky area, remainder of fin yellowish with some dusky areas.
Size: Maximum standard length 41 cm; commonly between 22 and 33 cm standard length.
Habitat, biology, and fisheries: Caught mainly by bottom trawls in depths of 90 to 110 m off southern
Queensland and New South Wales; also caught at a depth of 250 m near the edge of a barrier reef off
Nouméa, New Caledonia. Often caught in the same trawl with B. serratus. A benthic carnivore feeding on
similar organisms as B. serratus. As with B. serratus, there is a disparity in the sex ratio in relation to size;
specimens under 30 cm standard length were predominantly female, while those above 30 cm standard
length were males. Either only males grow to larger sizes, or there may be protogynous sex reversal; no
seasonal changes in ovaries were observed (perhaps indicating multiple spawning during the year).
Marketed fresh in Sydney.
Distribution: Southern Queens-
land (26°25’S) to New South
Wales (33°30’S); also known
off New Caledonia.
Malacanthus brevirostris Guichenot, 1848

Frequent synonyms / misidentifications: Malacanthus hoedtii Bleeker, 1859; M. parvipinnis Vaillant and
Sauvage, 1875; Dikellorhynchus incredibilis, Smith, 1956 (postlarvae) / Malacanthus latovittatus
(Lacepède, 1802).
FAO names: En - Quakerfish; Sp - Blanquillo.
Diagnostic characters: Body elongate, its depth 12 to 16% (usually 14%) standard length. Preopercle edge
smooth, angle 105°. Opercle with a single sharp spine, size about equal to pupil diameter. Snout short,
29 to 37% (usually 32%) head length. Jaws extending posteriorly to anterior rim of orbit. Total gill rakers
on first gill arch 9 to 20 (usually 15). Dorsal fin with I to V (usually II) spines and 52 to 60 (usually 57)
soft rays. Anal fin with I spine and 46 to 55 (usually 51) soft rays. Caudal fin truncate to slightly rounded
(upper lobe without elongate rays). Pored lateral-line scales 146 to 181. Colour: dorsal portions of body
light olive green, belly silver-white; dorsal fin faint pink with a narrow yellow upper margin; upper and
lower lobes of caudal fin yellow, separated by a central white band and with characteristic 2 dark
parallel bands.
Size: Maximum standard length 38.8 cm; commonly between 15 and 30 cm standard length.
Habitat, biology, and fisheries: Lives in self-constructed burrows in pairs at depths between 5 and 33 m.
Commonly found in coral rubble or sandy bottom. When frightened, it dives into burrow head first. Caught
mostly by spear, hook-and-line, and traps. Occasionally found in markets throughout the range. Marketed
fresh or salted.
Distribution: Found through-
out the Indo-Pacific, and
westward to the eastern Pa-
cific (Costa Rica, Panama,
Columbia).
2648 Bony Fishes
Malacanthus latovittatus (Lacepède, 1802)

Frequent synonyms / misidentifications: None / juveniles of Labroides dimidiatus Valenciennes, 1839
(family Labridae).
FAO names: En - Blue tilefish; Fr - Matajuel bleu; Sp - Matajuelo azul.
Diagnostic characters: Body elongate (somewhat deeper than in Malacanthus brevirostris), its depth 15
to 20% (usually 17%) standard length. Snout long, its length 37 to 47% (usually 44%) head length. Jaws
extending posteriorly only to below anterior nostril, well in front of eye. Preopercle edge smooth, the
angle about 110° to 120°, opercle with a single sharp spine about equal in size to pupil diameter. Gill rakers
on first gill arch reduced, their total number 6 to 14 (usually 9). Dorsal fin with III or IV (usually IV) spines
and 43 to 47 (usually 45) soft rays. Anal fin with I spine and 37 to 40 (usually 39) soft rays; caudal fin
truncate, with elongate elements in upper lobe. Pored lateral-line scales 116 to 132 (usually 124). Colour:
head blue, iris yellow; body olive-grey to violet-blue above, bluish white on belly; a broad midbody
longitudinal band running from operculum to tip of caudal fin, widening on tail to include most of
middle and lower portion; lower portion of fin with a small white rectangular area; dorsal fin
grey-brown with a pale band along upper margin; anal and pelvic fins white; pectoral fins bluish; juveniles
with colour distinctly different from adult, closely resembling that of Labroides dimidiatus, a cleaner wrasse
(family Labridae).
Size: Maximum standard length 38 cm; commonly to 35 cm standard length.
Habitat, biology, and fisheries: Lives in shallow waters (6 to 10 m), near edges of reefs. Inhabits
self-constructed burrows of sand and rubble. Caught by hook-and-line, spears, and traps. Marketed fresh
or salted.
Distribution: Throughout
Indo-Pacific, but not known
from Hawaii; extends as far
north as Honshu (Japan), but
not yet recorded from Australia
in the south Pacific. Found in
hyposaline waters of the Goldie
River, New Guinea.
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Perciformes: Percoidei: Lactariidae 2649
LACTARIIDAE
False trevally (milk trevally)
by J.M. Leis
Lactarius lactarius (Bloch and Schneider, 1801) TRF

Frequent synonyms / misidentifications: Lactarius delicatulus Valenciennes, 1833 / None.
FAO names: En - False trevally; Fr - Péliau chanos; Sp - Pagapa.
Diagnostic characters: Body of moderate depth, strongly compressed. Head large, contained 2.8 to 3.2
times in standard length. Mouth large and oblique (maxilla reaching past middle of eye), with
protruding lower jaw; a pair of small, sharp canine teeth at front of each jaw. Total gill rakers on first
gill arch 16 to 21. Two dorsal fins of about equal height, the first with VII or VIII spines, the second
with I spine and 19 to 23 soft rays. Anal fin with III spines and 25 to 28 soft rays (more soft rays
than in dorsal fin); no detached spines in front of anal fin. Caudal fin forked. Pectoral fins long and
pointed. Pelvic fins immediately below base of pectoral fins, and less than 1/2 as long. Caudal peduncle
not exceptionally narrowed. Cycloid scales moderate in size and easily shed; no scutes along lateral
line; 74 to 80 lateral-line scales. Colour: silver with blue iridescence above; silver-white below; a dusky
blotch on upper edge of gill cover; fins pale yellow to colourless; dorsal, anal, and caudal fins sometimes
with dusky margin; blue and yellow fade quickly following death.
Carangidae: first 1 or 2 anal-fin spines separate from remainder
of fin; caudal peduncle exceptionally narrowed; most species
have many spiny scutes (modified scales) along hind-part of
lateral line; other scales small and adherent; anal fin does not scutes
have more soft rays than dorsal fin.
Size: Maximum standard length about 35 cm; commonly to
25 cm. Carangidae
Habitat, biology, and fisheries: A coastal, schooling species over soft bottoms in depths of less than
100 m. Feeds on bottom-living invertebrates and fishes. Taken in seines, gill nets, trawls, and by
hook-and-line. Moderately important in coastal fisheries; marketed mostly fresh, but also dried-salted. From
1990 to 1995, the FAO Yearbook of Fishery Statistics reports a range of yearly catch of around 6 420 to
9 140 t of Lactarius lactarius from the Western Central Pacific.
Distribution: Over continental shelves from the Persian
Gulf to Taiwan Province of China, Philippines, Solomon
Islands, Fiji, and Australia.
Reference
Leis, J.M. 1994. Larvae, adults and relationships of the
monotypic perciform fish family Lactariidae. Rec. Austr.
Mus., 46(2):131-144.
2650 Bony Fishes
Pomatomidae POMATOMIDAE
Bluefishes
by B.B. Collette
Pomatomus saltatrix (Linnaeus, 1766)
Frequent synonyms / misidentifications: Pomatomus saltator (Linnaeus, 1766); Temnodon saltator
(Valenciennes, 1833) / None.
FAO names: En - Bluefish; Fr - Tassergal; Sp - Anchova de banco.
Diagnostic characters: A large species with a sturdy, compressed body and large head. Mouth large and
terminal, lower jaw sometimes slightly projecting; jaw teeth prominent, sharp, compressed, in a single
series. Two dorsal fins, the first short and low, with VII or VIII feeble spines connected by a
membrane, the second long with I spine and 23 to 28 soft rays; anal fin a little shorter than soft dorsal fin,
with II or III spines and 23 to 27 soft rays; caudal fin forked, but not deeply so; pectoral fins short, not
reaching to origin of soft dorsal fin. Scales small, covering head and body and bases of fins; lateral line
complete, almost straight. Colour: back greenish blue, sides and belly silvery; dorsal and anal fins pale
green tinged with yellow; pectoral fins bluish at base; caudal fin dull greenish tinged with yellow.
Similar families occurring in the area scutes
Carangidae: 2 detached spines in front of anal fin; also, scutes
on caudal peduncle in many species, and detached finlets
behind dorsal and anal fins in Elagatis and Decapterus.
Rachycentridae: spines of dorsal fin shorter, isolated, not con-
nected by a membrane; body not elongate; 2 silvery stripes on
sides; teeth smaller. Carangidae
Epigonidae: anal fin with II spines and 9 soft rays; jaws lacking
caniniform teeth.
Size: Maximum total length 1.1 m; commonly to 60 cm.
Habitat, biology, and fisheries: A powerful, swift fish, the
young hunting in schools, the adults in loose groups, often
attacking shoals of mullet or other fishes and destroying num- Rachycentridae
bers apparently far in excess of feeding requirements. Caught
mainly with gill nets, lines, and purse seines; commonly taken
on hook-and-line by sportsfishermen; marketed mostly fresh.
Distribution: Atlantic and Indo-West Pacific, but absent from
the eastern Pacific and the northwest Pacific. Barely enters the
area. Found along the southern coast of Australia from Onslow,
Western Australia (21°38’S) to Maryborough, Queensland Epigonidae
(25°32’S). Records from the Northern Territory and from
Indonesia appear to be erroneous.
Reference
Goodbred, C.O. and J.E. Graves. 1996. Genetic relationships
among geographically isolated populations of bluefish
Pomatomus saltatrix. Mar. Freshwater Res.,
47:347-355.
Perciformes: Percoidei: Scombropidae 2651
SCOMBROPIDAE
Gnomefishes
by P.C. Heemstra
D iagnostic characters (largely based on Scombrops boops): Body elongate (to 1.5 m total length),
slightly compressed, body length less than head length, contained 3.2 to 3.8 times in standard
length. Head (including maxilla) and body covered with cycloid scales. Eyes large, their diameter more
than snout length, 27 to 30% head length. Preopercle finely serrate, with ventral lobe. No spines on
opercle. Mouth large, maxilla (with large supramaxilla) reaching to below rear half of eye. Lower jaw with
2 or 3 canines at front, followed by a row of 8 or 9 large, knife-like spaced teeth; upper jaw with 12 or 13
large canines and 2 or 3 fang-like teeth just in front of vomer tooth patch; 1 or 2 rows of compressed canines
on palatines and a patch of smaller teeth on vomer; 2 elongate patches of small slender teeth on tongue.
Branchiostegal membranes separate, free of isthmus, with 7 rays. First gill arch with 2 or 3 gill rakers on
upper limb and 12 to 15 on lower limb. Two dorsal fins, the first with VIII or IX spines, the second
with I spine and 13 or 14 soft rays. Anal fin with III spines (first minute) and 12 or 13 soft rays.
Caudal fin forked. Pectoral fins about 2/3 head length, with 15 rays, not reaching past vertical at origin of
second dorsal fin. Origin of pelvic fins below base of pectoral fins; pelvic fins with I spine and 5 soft rays.
Soft dorsal, anal, and caudal fins scaly. Lateral line complete, with 51 to 55 pored scales to base of caudal
fin. Swimbladder present. Vertebrae 10+16. Colour: adults golden purplish brown; body of juveniles
reddish brown or yellowish dorsally, paler ventrally.
Scombrops boops
Habitat, biology, and fisheries: Adults occur in rocky areas near the bottom at depths of 200 to 700 m;
juveniles found in tidepools and shallow coastal water. Biology little known. Caught with hook-and-line and
in trawls.
Remarks: This family contains a single genus with about 4 species.
It has not yet been recorded from the Western Central Pacific;
Scombrops boops, is known from Japan to the East China Sea and
probably occurs in the area.
Acropomatidae
Acropomatidae: anal fin with II or III spines and 6 to 9 soft rays;
scales large, less than 50 in lateral line; pectoral fins about equal to
head length; vertebrae 10+15.
Apogonidae: anal fin with II spines; second dorsal fin with 8 or 9 soft
rays.
Sphyraenidae: snout much longer than eye diameter; dorsal fins
widely separated; body more elongate, its depth 6 to 10 times in
standard length.
Apogonidae
? Scombrops boops (Houttuyn, 1782)
Reference
Mochizuki, K. 1988. Family Scombropidae. In The fishes of the Japanese
Archipelago, edited by H. Masuda, K. Amaoka, C. Araga, T. Uyeno, Sphyraenidae
and T. Yoshino. Tokyo, Tokai University Press, p. 152.
2652 Bony Fishes
Echeneidae ECHENEIDAE
Remoras (sharksuckers, discfishes)
by B.B. Collette
D iagnostic characters: Perciform fishes with fusiform, elongate body (size to about 90 cm), charac-
terized by a transversely laminated, oval-shaped cephalic disc, this structure homologous with
spinous dorsal fin; skull wide, depressed to support disc. Opercle without spines, premaxillae not
protractile, gill membranes free from isthmus. Jaws broad, the lower projecting beyond the upper; villiform
teeth present in jaws and vomer (i.e. centrally on roof of mouth), usually on tongue and in certain species
on palatines (i.e. laterally on roof of mouth). Dorsal and anal fins long, lacking spines; dorsal-fin rays
range from 18 to 45, anal-fin rays from 18 to 41; pectoral fins set high on body, pointed or rounded, with
18 to 32 rays; pelvic fins far forward, close together, narrowly or broadly attached to underside of body,
with I spine and 5 soft rays; caudal fin slightly forked, emarginate, or slightly rounded (in large specimens
of some species), juveniles of some species with an elongate median caudal-fin filament. Scales small,
cycloid (smooth), usually embedded in the skin. No swimbladder. Colour: in life colours subdued, pale
brown, greyish to black, sometimes light to whitish or with light and dark horizontal stripes on trunk.
sucking disc dorsal and anal fins long,

lacking spines
Habitat, biology, and fisheries: Remoras attach themselves to many different marine vertebrates
including sharks, rays, tarpons, barracudas, sailfishes, marlins, swordfishes, jacks, basses, groupers,
ocean sunfish, sea turtles, whales, and dolphins; they may also attach to ships and various floating objects.
Some remoras have a great preference or specificity toward certain hosts. Remora australis, the
whalesucker, is only known from marine mammals. R. osteochir, the marlinsucker, is almost always found
attached to spearfishes, particularly the sailfish and white marlin. The preferred host of R. albescens, the
white sucker, is the manta ray. Species of the genus Echeneis are often free-swimming and occur in
shallow, inshore waters. Remora and Remorina are almost always captured on their host where they may
be found attached to the body, in the mouth, or in the gill cavity. Remoras feed on parasitic copepods
attached to their host and food scraps dropped by the host. Although remoras are not considered to be of
any commercial importance, at least Echeneis naucrates is taken in coastal fisheries along with other
species and sold in local markets.

No other family of fishes has a cephalic sucking disc.
The cobia (Rachycentron canadum, family Rachy-
centridae) bears some resemblance to the remoras.
It has been postulated that a cobia-like ancestor
may have given rise to the echeneid fishes.
Rachycentridae
Perciformes: Percoidei: Echeneidae 2653
Key to the species of Echeneidae occuring in the area

Note: the Echeneidae is divisible into 2 subfamilies, 4 genera, and 8 species, 7 of which occur in the
Western Central Pacific.
1a. Body very elongate, its depth contained 8 to 14 times in standard length; pectoral fins
pointed; usually a dark longitudinal band on sides, bordered with white; anal-fin base
long, anal-fin rays 29 to 41; caudal fin lanceolate in young, the middle rays filamentous,
almost truncate in adults, the lobes pronounced (Fig. 1) . . . . . . . . . (subfamily Echeneinae) ® 2
1b. Body not elongate, its depth contained 5 to 8 times in standard length; pectoral fins
rounded; colour nearly uniform, without bands; anal-fin base short, anal-fin rays 18 to 28;
caudal fin forked in young, becoming emarginate or truncate in adults (Fig. 2) . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (subfamily Remorinae) ® 3
Fig. 1 Echeneis Fig. 2 Remora
2a. Sucking disc large, with 18 to 28 laminae; vertebrae 30 . . . . . . . . . . . . . Echeneis naucrates

2b. Sucking disc small, with 9 to 11 laminae; vertebrae 39 to 41 . . . . . . . . . Phtheirichthys lineatus
3a. Pelvic fins broadly attached to abdomen; disc laminae 15 to 28; vertebrae 27; colour light
to dark brown; hosts include sharks, billfishes, or cetaceans, depending on the species
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Remora) ® 4
3b. Pelvic fins narrowly attached to abdomen; disc laminae 13 or 14; vertebrae 26; colour
whitish; usual host, manta rays . . . . . . . . . . . . . . . . . . . . . . . . . . . Remorina albescens
4a. Disc laminae 25 to 28; total gill rakers on first gill arch 17 to 20 . . . . . . . . . . . Remora australis
4b. Disc laminae 15 to 20; total gill rakers on first gill arch 11 to 17 . . . . . . . . . . . . . . . . . . ® 5
5a. Posterior border of disc located far posterior to tips of adpressed pectoral fins . . . Remora osteochir
5b. Posterior border of disc located anterior to tips of adpressed pectoral fins . . . . . . . . . . . . ® 6
6a. Disc laminae 15 to 18; dorsal-fin rays 27 to 34; total gill rakers on first gill arch less than
21 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Remora brachyptera
6b. Disc laminae 16 to 20; dorsal-fin rays 21 to 27; total gill rakers on first gill arch 28 to 37
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Remora remora
0 Echeneis naucrates Linnaeus, 1758
Phtheirichthys lineatus (Menzies, 1791)
Remora australis (Bennett, 1840)
Remora brachyptera (Lowe, 1839)
Remora osteochir (Cuvier, 1829)
Remora remora (Linnaeus, 1758)
Remorina albescens (Temminck and Schlegel, 1845)
References
Cressey, R.F. and E.A. Lachner. 1970. The parasitic copepod diet and life history of diskfishes (Echeneidae). Copeia,
1970:310-318.
Lachner, E.A. 1986. Echeneididae. In Fishes of the North-eastern Atlantic and the Mediterranean 3, edited by P.J.P.
Whitehead et al. UNESCO, pp.1329-1334.
2654 Bony Fishes
Echeneis naucrates Linnaeus, 1758

FAO names: En - Sharksucker; Fr - Rémora commun; Sp - Pegatimón.
Diagnostic characters: An elongate fish, depth of body contained 8 to 14 times in standard length.
Jaws broad, the lower projecting beyond the upper. First dorsal fin replaced by a transversal, laminated,
oval cephalic disc with 21 to 28 laminae; second dorsal fin and anal fin long, lacking spines, the anal fin
with 31 to 41 soft rays; pectoral fins short, high on body, pointed; caudal fin lanceolate in young, the
middle rays elongate and filamentous; caudal fin almost truncate in adults, with the upper and lower lobes
longer than the middle rays. Colour: dark longitudinal band on sides bordered with white; juveniles with
upper and lower margins of fins white.
Size: Maximum standard length about 1 m.
Habitat, biology, and fisheries: Unlike most other remoras, Echeneis naucrates is often found free-swim-
ming and occurs in shallow inshore waters. It will attach temporarily to a wide variety of hosts such as
sharks and sea turtles and also to ships. Sometimes used by natives as an aid to fishing. A line is tied to
the caudal peduncle of the remora and then it is released; upon attaching to another fish, the remora and
its host are hauled in by the fishermen. Taken with drift nets and trawls. Occasionally marketed fresh.
tropical and temperate seas,
except for Pacific American
coast.
Perciformes: Percoidei: Rachycentridae 2655
Perciformes: Percoidei: Rachycentridae RACHYCENTRIDAE

Cobia
by B.B. Collette
Rachycentron canadum (Linnaeus, 1766) CBA
FAO names: En - Cobia; Fr - Mafou; Sp - Cobie.
Diagnostic characters:
Body elongate, subcylin-
drical; head broad and
depressed. Mouth large,
terminal, with projecting
lower jaw; villiform teeth in
jaws and on roof of mouth
and tongue. First dorsal
fin consisting of VII to IX
(usually VIII) short but
strong isolated spines,
not connected by a membrane; second dorsal fin long, with 26 to 33 soft rays, the anterior rays somewhat
elevated in adults; anal fin similar to dorsal fin, but shorter, with II or III spines and 22 to 28 soft rays;
caudal fin lunate in adults, upper lobe longer than lower (caudal fin rounded in young, the central rays
much prolonged); pectoral fins pointed, becoming more falcate with age. Scales small, embedded in thick
skin; lateral line slightly wavy anteriorly. Colour: back and sides dark brown, with 2 sharply defined
narrow silvery bands; belly yellowish.
Pomatomidae: spines of dorsal fin connected by a membrane;
also, body and head deeper and no stripes on sides; teeth large
and very sharp.
Carangidae (jacks): usually, II detached spines visible in front
of anal fin; also, distinctly elongate carangid species have either Pomatomus (Pomatomidae)
scutes on lateral line (Decapterus, Trachurus) or detached scutes finlet
finlets behind dorsal and anal fins (Decapterus, Elagatis).
Size: Maximum total length 2 m, commonly to 1.1 m; maximum
weight 50 kg.
Habitat, biology, and fisheries: Pelagic, but also found over
shallow coral reefs and off rock shores, occasionally in estuar-
ies. Feeds on crabs, squids, and fishes. Caught mainly with Decapterus (Carangidae)
handlines, and by trolling pelagically offshore over the continen-
2-rayed finlet
tal shelf. Excellent food and game fish but does not occur in large
enough numbers to support a commercial fishery. Marketed
mostly fresh. From 1990 to 1995, the FAO Yearbook of Fishery
Statistics reports a range of yearly catch of around 1 290 to
1 750 t of Rachycentron canadum from the Western Central
Pacific.
Elagatis (Carangidae)
Distribution: Widespread in the Indo-West Pacific, but absent
from the eastern Pacific and from the
Pacific Plate, except marginally. Within the
western Pacific, found around the northern
coast of Australia, north to New Guinea,
Indonesia, Borneo, the Philippines, and
Japan.
Reference
Smith, J.W. 1975. Life history of cobia,
Rachycentron canadum (Osteichthyes:
Rachycentridae), in North Carolina
waters. Brimleyana, 23:1-23.
2656 Bony Fishes
Coryphaenidae CORYPHAENIDAE
Dolphinfishes (“dolphins”)
by B.B. Collette
D iagnostic characters: Elongate, compressed fishes (size to 2.1 m). Mouth large, with many fine teeth
in bands; 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 fins fitting into a groove on body. Scales small and
cycloid. Colour: in life very variable, sides with golden hues and back brilliant metallic greens and blues;
many small, black spots on head and body; specimens less than 15 cm have dark vertical bars.
dorsal fin
begins on nape dorsal and anal fins
almost to caudal fin
bony crest
in male
Habitat, biology, and fisheries: Both species are open ocean pelagic species. They feed on flyingfishes
and small fishes and crustaceans associated with floating Sargassum weed. Taken by both commercial
and recreational fisheries wherever they occur. The flesh is highly esteemed.

None. No other fishes have a combination of characters such as dorsal fin from nape almost to caudal fin;
anal fin from about midpoint of body almost to caudal fin; no sharp spines in dorsal and anal fins; caudal
fin deeply forked; and pelvic fins well developed.
Key to the species of Coryphaenidae

1a. Greatest body depth in adults less than 25% of stand-
ard length; tooth patch on tongue small and oval
(Fig. 1a); dorsal-fin rays 58 to 66; lateral-line scales
200 or more; vertebrae 31. . . . . . . . . . Coryphaena hippurus
1b. Greatest body depth in adults more than 25% of
standard length; tooth patch on tongue broad and
trapezoidal (Fig. 1b); dorsal-fin rays 52 to 59; lateral- a) Coryphaena b) Coryphaena
line scales 200 or fewer; vertebrae 33 . . . Coryphaena equiselis hippurus equiselis
Fig. 1 tooth patch on tongue

0
The symbol is given when species accounts are included.
0 Coryphaena equiselis Linnaeus, 1758
0 Coryphaena hippurus Linnaeus, 1758
References
Gibbs, R.H., Jr. and B.B. Collette. 1959. On the identification, distribution, and biology of the dolphins, Coryphaena
hippurus and C. equiselis. Bull. Mar. Sci. Gulf, Carib., 9(2):117-152.
Palko, B.J., G.L. Beardsley, and W.J. Richards. 1982. Synopsis of the biological data on dolphin fishes, Coryphaena
hippurus Linnaeus and Coryphaena equiselis Linnaeus. NOAA Tech. Rep. NMFS Circ., (443):28 p.
Perciformes: Percoidei: Coryphaenidae 2657
Coryphaena equiselis Linnaeus, 1758

Frequent synonyms / misidentifications: Coryphaena equisetis Osbeck, 1765 / Coryphaena hippurus
Linnaeus, 1758.
FAO names: En - Pompano dolphinfish; Fr - Coryphène dauphin; Sp - Dorado.
Diagnostic characters: Body elongate and compressed, greatest body depth in adults more than 25%
of standard length; young fish (up to 30 cm) have head profile slightly convex. Tooth patch on tongue
broad and trapezoidal; bands of teeth on jaws and roof of mouth (vomer and palatines). A single
dorsal fin extending from just behind eyes almost to caudal fin, with 52 to 59 soft rays; a convex
anal fin extending from anus almost to caudal fin; pectoral fins about 1/2 of head length; caudal fin
deeply forked. Lateral-line scales 200 or fewer. Vertebrae 33. Colour: back brilliant metallic blue-green
in life; fading rapidly after death to green tinge; sides silvery with a golden sheen and numerous black
spots; dorsal fin dark; in juveniles, entire margin of caudal fin white, pelvic fins not pigmented.
Habitat, biology, and fisheries: Pelagic, inhabiting open waters, but also approaching the coast. Probably
resembles Coryphaena hippurus in following ships and concentrating below floating objects. Feeds on
small fishes and squids. Caught mainly by trolling and with floating lines. Marketed fresh.
most tropical and subtropical
seas but frequently misidenti-
fied as juvenile or female
C. hippurus.
2658 Bony Fishes
Coryphaena hippurus Linnaeus, 1758

FAO names: En - Common dolphinfish; Fr - Coryphène commune; Sp - Dorado común.
Diagnostic characters: Body elongate and compressed, greatest body

depth in adults less than 25% of standard length; young fish (up to 30 cm)
have a slender, elongate body with head profile slightly convex; in larger males
(30 to 200 cm) the head profile becomes vertical with development of a bony
crest; tooth patch on tongue small and oval; bands of teeth on jaws and
roof of mouth (vomer and palatines). A single dorsal fin extending from
above eyes almost to caudal fin, with 58 to 66 soft rays; a concave anal
fin extending from anus almost to caudal fin; pectoral fins more than 1/2
of head length; caudal fin deeply forked. Lateral-line scales 200 or more.
Vertebrae 31. Colour: back brilliant metallic blue-green in life, after death
fading to grey with a green tinge; sides silvery with a golden sheen, and 1 row
of dark spots or golden blotches running below dorsal fin and 1, 2, or more
rows on and below lateral line, some scattered irregularly; dorsal and anal fins
black, the latter with a white edge; pectoral fins pale; caudal fin silvery with a diagram showing
golden sheen; in juveniles, only tips of caudal-fin lobes white, pelvic fins black. development of bony
Size: Maximum total length 2.1 m; commonly to 1 m. crest in males
Habitat, biology, and fisheries: Pelagic, inhabiting open waters, but also approaching the coast; follows
ships and forms small concentrations below floating objects. Feeds mainly on fishes, but also on
crustaceans and squids. Breeds in the open sea, probably approaching the coast as water temperatures
rise. Caught by trolling and on tuna longlines; also occasionally with drift nets. Marketed fresh; a very highly
appreciated food fish. For 1995, the FAO Yearbook of Fishery Statistics reports a total catch of 4 847 t of
Coryphaena hippurus from the area.
tropical and subtropical seas.
click for next page

Perciformes: Percoidei: Carangidae 2659
CARANGIDAE
Jacks and scads (also trevallies, queenfishes, runners, amberjacks, pilotfishes, pampanos, etc.)
by W.F. Smith-Vaniz
D iagnostic characters: Body highly variable in shape, ranging from elongate and fusiform to deep and
strongly compressed; caudal peduncle of medium width to notably slender, in some species with a
moderate lateral keel, bilateral paired keels or dorsal and ventral grooves. Head varying from moderately
long and rounded to short, deep and very compressed. Eye small to large, with adipose eyelid negligible
to strongly developed. Snout pointed to blunt; lower jaw protruding to subtended (included). Teeth in jaws
in rows or bands, either small to minute or an enlarged row of recurved canines present; teeth on roof of
mouth (vomer, palatines) or tongue present or absent depending on species or developmental stage. Gill
openings large, gill membranes not united, free from isthmus; branchiostegal rays 6 to 10 (usually 7); gill
rakers moderate in length and number to long and numerous, their number decreasing with growth in some
species; opercular bones smooth (but with spines in larvae and small juveniles). Two dorsal fins that are
separate in small juveniles, the first of moderate height or very low, with IV to VIII spines (spines
obsolete or embedded in adults of some species), the second dorsal fin with I spine and 18 to 44
soft rays and the anterior lobe scarcely produced to extremely long; anal fin with II anterior spines (only
I spine in Elagatis and Seriolina) that are separate from rest of fin by a gap (becoming embedded
in adults of some species) followed by I spine and 15 to 39 soft rays, with the anterior lobe low to
elongate; pectoral fins with about 14 to 24 soft rays, either long and falcate or short and pointed or rounded;
pelvic fins with I spine and 5 soft rays, moderately long in some species to becoming rudimentary in others
(absent in Parastromateus); caudal fin forked, with the lobes equal in most species. Scales small,
sometimes difficult to see, and cycloid (smooth to touch), but strongly lanceolate to needle-like in a few
species, usually absent from some areas of head and covering body (but absent on certain body areas in
some species) and sometimes extending onto fins. Lateral line arched (curved) or elevated above
pectoral fins and straight posteriorly, extending onto caudal fin; scutes (enlarged, thickened, and
often pointed scales in lateral line) present and prominent, or reduced in some species and absent
in some genera. Vertebrae 10-11+14-16 (usually 10+14; total 24 to 26). Colour: darker above (green or
blue to blackish) and paler below (silvery to white or yellow-golden), some species almost entirely silvery
when alive, others with dark or coloured bars or stripes on head, body or fins, and some able to change
patterns; the young of many species with bars or spots.
Atropus Trachinotus
Decapterus
Caranx Scomberoides
examples of body shapes

2660 Bony Fishes
Habitat, biology, and fisheries: Mostly schooling species (but Alectis generally solitary); some species
have largely continental distributions and occur primarily in brackish environments (especially young),
others such as Elagatis and Naucrates are pelagic, usually found at or near the surface, mostly in oceanic
waters, often far offshore. This is one of the most important families of commercial fishes, and all species are
used for food. For 1995, FAO’s Yearbook of Fishery Statistics reports a total catch of around 959 300 t of
Carangidae from the Western Central Pacific. Caught commercially with trawls, also with purse seines, traps,
and on line gear. The larger species of Trachinotus, Seriola, and Caranx are highly regarded as sportfish.
The Carangidae is distinguished from all similar families
in having the first 2 anal-fin spines detached from rest of
fin (caution: these spines sometimes partially or com-
pletely embedded in large adults of several genera, espe- anal fin
cially Seriola, Alectis, and Caranx; however, they can be
found, without much difficulty by dissection). The pres- II detached spines, Carangidae
ence of enlarged, thickened scutes in the straight part of often embedded in skin
lateral line in some genera easily distinguishes them from
other families. Additional distinguishing characters of
similar families (especially to those carangid genera lack-
more dorsal-fin series of
ing scutes on the lateral line), are the following: spines finlets
Scombridae: dorsal-fin spines IX to XXVII (IV to VIII in
Carangidae); no scutes developed along posterior part of
lateral line; series of finlets present behind dorsal and anal
fins (Scomberoides, the only carangid that lacks scutes
and at the same time has a series of finlets, is further
distinguished by having II detached, depressible spines
in front of anal fin). no scutes here
Scombridae
Stromateidae: Pampus, the only genus in the area, can
be distinguished from carangids as follows: IV blade-like anal-fin spines, not detached from fin (barely
visible in adults, may be embedded in skin); pelvic fins minute or absent in adults (readily apparent in
carangids, except in adults of Parastromateus). Pampus is very similar in body shape to Parastromateus,
but has the lateral line strongly arched anteriorly (weakly arched in Parastromateus) and the straight part
does not form a slight keel on the caudal peduncle or has 8 to 19 weak scutes as in Parastromateus.
no keel here
keel of
scutes
IV blade-like anal-fin spines
(barely visible in adults)
Pampus (Stromateidae) Parastromateus niger (Carangidae)
Gempylidae (especially Lepidocybium and Ruvettus species): base of first dorsal fin longer than that of
second excluding finlets (equal or shorter than second in Carangidae); a series of dorsal and anal finlets
present in Lepidocybium and Ruvettus.
Lepidocybium Ruvettus
Gempylidae
Pomatomidae: both jaws with a series of strong com-

pressed teeth; no grooves on caudal peduncle (present
in Seriola which is superficially similar). dorsal view
no groove caudal-peduncle groove
Pomatomidae lateral view
Rachycentridae: head broad and depressed, lower jaw Seriola (Carangidae)

projecting; body more slender; first dorsal fin with VIII
or IX short, free spines, each depressible in a groove,
II or III anal-fin spines, none detached from fin.
Centrolophidae, particularly the genus Hyperoglyphe:
III anal-fin spines, none detached from fin; adults with
margin of preopercle usually moderately denticulate
(smooth in Carangidae); jaw teeth all conical; simple
caudal fin, not deeply forked.
Lactariidae: III anal-fin spines, none detached from fin; Rachycentridae
anal fin has more soft rays than dorsal fin; caudal
peduncle not exceptionally narrowed.
anal-fin rays outnumbering

denticulated III anal-fin spines, not dorsal-fin rays
detached from fin
Hyperoglyphe (Centrolophidae) Lactariidae
Identification note
Dentition: Dentition has traditionally been used by past workers to finely subdivide a number of presumably
monophyletic species-groups centered around Caranx under different generic or subgeneric designations.
One such group is the “catch-basket” category “Carangoides.” The Indo-Pacific species of Carangoides
(sensu lato) exhibit a wide range of dentition types, including some that appear to be morphologically
intermediate. In other cases, genera long recognized as valid appear to be most closely related to a
subgroup within Carangoides. It is tempting to recognize a more inclusive genus Caranx, with Caran-
goides recognized as one of several subgenera. However, Caranx would then probably become undefin-
able as a monophyletic group, thus the status quo is maintained for the present with some reluctance.
Two nominal genera, Pseudocaranx and Ulua, are especially likely candidates for synonymization.
Fin spines: The 2 detached anterior anal-fin spines and some of the anterior spines of the first dorsal fin
(especially the first spine) frequently become completely embedded in the skin in large individuals of many
carangids (all spines of the first dorsal fin become embedded in Alectis and Parastromateus at a relatively
small size). Even in those genera with a relatively high spinous dorsal fin, the first spine is usually small
and closely appressed to the second spine and thus can easily be overlooked.
Breast squamation: Many species of Carangoides, Caranx, and Uraspis have the breast only partially
scaly, and the pattern of breast squamation is an important character used to distinguish species. The
pattern of breast squamation is sometimes difficult to observe in fresh specimens; observation is facilitated
by gently scraping the breast with a knife to remove mucous and allowing the breast to partially dry,
hastened by blowing air on the area.
Gill-raker counts: In species with relatively numerous gill rakers (e.g. Decapterus and Trachurus) great
care must be taken not to overlook rakers at either end of the gill arch. It is suggested that a small knife
be used to free the upper limb of the gill arch where it joins the skull. With a little practice this can be done
2662 Bony Fishes
without leaving any stub with rakers attached. Once this has been accomplished, the gill rakers are much
easier to see. In some genera (e.g. Caranx and Seriola ) the number of developed rakers decreases with
growth with a concomitant increase in the number of rudiments (tubercles or short rakers with the diameter
of their bases greater than their height). When rudimentary rakers are included in the gill-raker counts, and
large specimens are being examined, it is very important that all of the tubercles are counted. In all cases
the raker in the angle of the gill arch is included in the count of lower-limb rakers.
Lateral-line scutes: In many carangids, size and configuration of the scales and scutes on the lateral line
is variable and there may be a gradual transition from one type to another. Scutes are here defined as
modified scales that either have their posterior margin with a small to moderate projecting spine or the
scale has a raised horizontal ridge and ends in an apex not exceeding a 90° angle. All scutes should be
counted, including those extending onto the caudal-fin base. In order to observe and accurately count the
lateral-line scales and scutes, good lighting and some magnification is recommended. In some species it
may also be necessary to remove small body scales that tend to overgrow or otherwise obscure the lateral
line.
Measurements: The curved part of the lateral line is measured as a chord of the arch extending from the
upper edge of the opercle to its junction with the straight part. The straight part of the lateral line is measured
from its junction with the curved part to its termination on the caudal-fin base (end of the last scute). In
cases where the junction of the curved and straight parts is very gradual, the curved part is considered to
begin with the scale or scute that has 3/4 of its height above the central axis of the straight part.
Key to the genera and subgenera of Carangidae occurring in the area

1a. Posterior straight part of lateral
line with hardened scutes; in
scutes
adults, pectoral fins long and fal-
cate, in most genera longer than
head (Fig. 1) (about equal to head
length in Selar a n d Trachurus,
and shorter than head length in
some Decapterus spp.) . . . . . . . . . . . ® 2
1b. No scutes in lateral line (only
pored scales, not enlarged); pec-
toral fins relatively short, shorter
than head (about 50 to 90% of
head length) . . . . . . . . . . . . . . . . ® 19 Fig. 1 Caranx
2a. Pored scales in curved lateral line scute-like, expanded dorsoventrally (Fig. 2) (caution:
in large fish may be obscured by overgrowth of smaller scales); dorsal accessory lateral
line normally extends posteriorly at least to below origin of second dorsal fin, usually
farther posteriorly (Figs 2 and 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachurus
2b. No enlarged scute-like scales in curved lateral line; dorsal accessory lateral line
terminating before origin of spinous dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
accessory 1st dorsal-fin spine
lateral line
dorsal view accessory lateral line
Fig. 2 Trachurus Fig. 3 Trachurus

3a. Pelvic fins, if present (absent in specimens larger than about 10 cm fork length),
positioned distinctly anterior to a vertical line through pectoral-fin base; soft anal-fin rays
35 to 39 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parastromateus
3b. Pelvic fins (always present) not positioned distinctly anterior to a vertical through
pectoral-fin base; soft anal-fin rays 15 to 31 . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Scales on body minute, inconspicu-

ous and embedded giving the im-
pression of naked skin; in smaller filamentous
fish, anterior soft rays of dorsal and rays
anal fins filamentous (Fig. 4) . . . . . . . . Alectis
4b. Scales on body small but conspicu-
ous, not embedded; in smaller fish,
anterior soft rays of dorsal and anal
fins not filamentous . . . . . . . . . . . . . . ® 5
5a. Second dorsal and anal fins with 1

or more distinctly separate finlets . . . . . . ® 6
5b. Second dorsal and anal fins without
finlets . . . . . . . . . . . . . . . . . . . . . ® 7 Fig. 4 Alectis
6a. Single detached terminal 2-rayed single finlet

finlet in dorsal and anal fins (Fig. 5);
shoulder girdle (cleithrum) margin
with 2 papillae, the lower papilla
larger (Fig. 6); maximum scute
height smaller than eye diameter . . . Decapterus
6b. Posterior soft dorsal- and anal-fin
rays consisting of 6 to 10 detached
finlets; shoulder girdle margin Fig. 5 Decapterus
smooth; maximum scute height
larger than eye diameter (Fig. 7) . . . Megalaspis
large scutes
small finlets
papilla
cleithral
margin
large
papilla
Fig. 7 Megalaspis
gill chamber after lifting operculum

small
Fig. 6 Decapterus papilla
7a. Shoulder girdle (cleithrum) margin with a furrow

ventrally, a large papilla immediately above it and cleithral
a smaller papilla near upper edge (Fig. 8) . . . . . . . . . Selar margin
7b. Shoulder girdle margin smooth . . . . . . . . . . . . . . . ® 8
8a. Upper jaw without teeth . . . . . . . . . . . . . . . . . . . ® 9 large

8b. Upper jaw with 1 or 2 rows or a band of minute papilla
teeth (caution: teeth difficult to detect in some
Carangoides) . . . . . . . . . . . . . . . . . . . . . . . . ® 10 furrow
gill chamber after lifting operculum
Fig. 8 Selar
2664 Bony Fishes
9a. Lower jaw with a series of minute teeth; a prominent black

opercular spot encroaching on shoulder; adipose eyelid
well developed posteriorly . . . . . . . . . . . . . . . . . . . Selaroides
9b. Lower jaw with a few feeble teeth in young (smaller than
10 cm fork length), absent in adults; no black opercular
spot; adipose eyelid poorly developed . . . . . . . . . . . Gnathanodon
10a. Tongue, roof and floor of mouth white, the rest dark (Fig. 9);
anal-fin spines reabsorbed or reduced and immovable; no
teeth on vomer or palatines . . . . . . . . . . . . . . . . . . . . Uraspis
10b. Lining of mouth not distinctly black and white; anal-fin
spines normal and movable; teeth present on vomer and
palatines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
Fig. 9 Uraspis
11a. Fleshy adipose eyelid com-
pletely covering eye except for
a vertical slit centred on pupil
(Fig. 10a); terminal ray of dor-
sal and anal fins finlet-like, a
little more separated from
other rays but not detached,
and about twice length of pe-
nultimate ray . . . . . . . . . . . . . . Atule
11b. Fleshy adipose eyelid, if pre-
adipose
sent, not well developed ante- eyelid
riorly, most of anterior half of a) Atule b) Alepes
pupil exposed; terminal ray of
Fig. 10
dorsal and anal fins not finlet-
like (except terminal ray length
1.5 times the length of penulti-
m a t e r a y i n l a r g e Alepes
djedeba) . . . . . . . . . . . . . . . . ® 12
12a. Both jaws with a single row of numerous, comb-like teeth; adipose eyelid well developed
on posterior half of eye only (Fig. 10b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alepes
12b. Dentition not as above; adipose eyelid, if present, variously developed . . . . . . . . . . . . . ® 13
13a. Upper jaw anteriorly with 2 irregular rows of short conical teeth, posteriorly inner surface
of jaw paved with blunt teeth (Fig. 11a); snout shorter than eye diameter . . . . . . . . . . “Alepes”
13b. Dentition not as above; snout usually equal or larger than eye diameter . . . . . . . . . . . . ® 14
14a. Upper jaw with an outer series of moderate to strong canines and an inner band of fine
teeth (Fig. 11b); lower jaw with a single row of teeth . . . . . . . . . . . . . . . . . . . . . . . ® 15
14b. Dentition not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
2 rows of short
conical teeth
canines
conical
teeth
inner fine
surface teeth
of jaw
a) “Alepes” b) Caranx c) Pseudocaranx

Fig. 11 ventral view of teeth in upper jaw (teeth on roof of mouth not shown)
15a. Body generally deep, with dorsal profile more convex than ventral; total gill rakers 20
to 31 on first gill arch; segmented dorsal- and anal-fin rays never produced as filaments;
2 to 4 canines anteriorly in each jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx
15b. Body shallow with dorsal and ventral profiles equally convex; total gill rakers 34 to 39
on first gill arch; segmented dorsal- and anal-fin rays produced as filaments in adult
males; no canines anteriorly in either jaw . . . . . . . . . . . . . . . . . . . . . . . . . . Pantolabus
16a. Both jaws with single series of short, conical teeth (upper jaw sometimes with an inner
row of conical teeth anteriorly) (Fig. 11c); breast completely scaly . . . . . . . . . . . Pseudocaranx
16b. Both jaws with a band of teeth, at least anteriorly; breast naked ventrally (most species)
to completely scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
17a. Belly with a deep median groove, accommodating pelvic fins, anus, and anal-fin spines
(Fig. 12a); pelvic fins conspicuously long and black, tip of appressed fins extending
almost to origin of anal fin; curved lateral line short, chord of curved part of lateral line
contained 1.5 to 2 times in straight part (Fig. 12b) . . . . . . . . . . . . . . . . . . . . . . . Atropus
17b. Belly without median groove; pelvic fins not conspicuously long and black; curved lateral
line moderate in most species, with chord of curved part of lateral line contained less
than 1.5 times in straight part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
median groove curved part of

lateral line
pelvic fins long,

black
a) underside of belly
b) lateral view
Fig. 12 Atropus
18a. Gill rakers long, feather-like, and project into mouth along side of tongue (Fig. 13); total
gill rakers 54 to 86 on first gill arch; lower jaw becoming prominent in large adults, with
the angle of “chin” projecting beyond upper jaw (Fig. 14) . . . . . . . . . . . . . . . . . . . . . Ulua
18b. Gill rakers of normal length and shape; total gill rakers 21 to 37 on first gill arch; shape
of lower jaw not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides
gill
rakers
tongue
Fig. 13 Ulua Fig. 14 Ulua

2666 Bony Fishes
19a. Bases of soft dorsal and anal fins unequal in length, anal-fin base shorter and only about
45 to 70% of dorsal-fin base length (Fig. 15); caudal-peduncle grooves present, dorsally
and ventrally (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
19b. Base of soft anal fin as long as, or only slightly shorter than, base of dorsal fin; no
caudal-peduncle grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
spines
caudal-peduncle groove
dorsal view
lateral view
Fig. 15 Naucrates Fig. 16 Naucrates
20a. Terminal 2-rayed finlet present in 2-rayed finlet

dorsal and anal fins (Fig. 17); up-
per jaw ending distinctly before
eye (to below anterior margin of
eye in young) . . . . . . . . . . . . . Elagatis
20b. No finlets in dorsal and anal fins;
upper jaw ending below anterior
margin of eye to posterior margin
of eye . . . . . . . . . . . . . . . . . . . ® 21 Fig. 17 Elagatis
21a. Upper jaw broadly rounded posteriorly and usually terminating below posterior margin
of eye (Fig. 18a); gill rakers on first gill arch mostly consisting of rudiments, 4 to 10 total
elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriolina
21b. Upper jaw truncate or slightly rounded posteriorly and terminating below about anterior
margin of eye to middle of eye (Fig. 18b, c); gill rakers on first gill arch mostly well
developed, 11 to 29 total elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
a) Seriolina b) Naucrates c) Seriola
Fig. 18
22a. First dorsal fin with IV or V spines; fleshy keel

soft anal-fin rays 15 to 17; fleshy
keel laterally on caudal peduncle
well developed (Fig. 19) . . . . . . . Naucrates
22b. First dorsal fin with VII or VIII
spines (caution: anterior spines
may become completely embed-
ded in large individuals); soft
anal-fin rays 18 to 22 (except 15 groove
to 17 in S. hippos); fleshy keel on
caudal peduncle absent to mod- Fig. 19 Naucrates
erately developed (S. lalandi) . . . . . Seriola
23a. Posterior soft dorsal- and anal-fin rays consisting of semi-detached finlets (Fig. 20);
distal quarter to half of rays not connected by interradial membrane (unattached portion
of rays increasing with growth); lower jaw of adults with 2 rows of conical teeth separated
by a shallow groove; upper lip joined to snout at midline by a bridge of skin (frenum),
except crossed by a shallow groove in very young . . . . . . . . . . . . . . . . . . . Scomberoides
23b. Posterior soft dorsal- and anal-fin rays not consisting of semi-detached finlets (Fig. 21);
lower jaw without teeth or with band of small villiform teeth; upper lip separated from
snout at midline by a continuous deep groove . . . . . . . . . . . . . . . . . . . . . . . Trachinotus
semi-detached finlets
Fig. 20 Scomberoides Fig. 21 Trachinotus
Key to the species of Alectis occurring in the area

1a. Profile of nape and head
broadly rounded; suborbital
de pt h r el atively narrow,
contained 1.7 to 3 times in
upper jaw length (Fig. 22);
gill rakers (excluding
rudiments) on lower limb of
first arch 12 to 17 . . . . . Alectis ciliaris
1b. Profile of nape and head
somewhat angular; suborbi-
tal depth relatively broad,
contained 0.8 to 1 times in
upper jaw length (Fig. 23);
gill rakers (excluding rudi-
ments) on lower limb of first Fig. 22 Alectis ciliaris Fig. 23 Alectis indica
arch 21 to 26 . . . . . . . Alectis indica
2668 Bony Fishes
Key to the species of Alepes occurring in the area

1a. Interradial membranes of spinous dorsal fin black; total gill rakers 24 to 30 on first arch
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alepes melanoptera
1b. Interradial membranes of spinous dorsal fin transparent to dusky; total gill rakers 32 to
47 on first arch (except 27 to 30 in A. apercna) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Total gill rakers 27 to 30 on first gill arch; upper jaw with supramaxilla relatively small
and without an anterior spine-like projection (Fig. 24a) . . . . . . . . . . . . . . . . Alepes apercna
2b. Total gill rakers 32 to 47 on first gill arch; upper jaw with supramaxilla relatively large
and with an anterior spine-like projection (Fig. 24b, c) . . . . . . . . . . . . . . . . . . . . . . . ® 3
a) Alepes apercna b) Alepes djedaba c) Alepes vari
Fig. 24 upper jaw (supramaxilla shaded)
3a. Total gill rakers on first gill arch 38 to 47, of which 10 to 14 on upper limb, and 27 to 33
on lower limb; lateral line with 31 to 36 scales and 39 to 51 scutes (total 77 to 85); scutes
larger (Fig. 25a); ultimate ray of dorsal and anal fins about 1.3 to 1.5 times the length
of penultimate ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alepes djedaba
3b. Total gill rakers on first gill arch 32 to 38, of which 9 to 12 on upper limb, and 23 to 26
on lower limb; lateral line with 42 to 50 scales, 48 to 69 scutes (total 86 to 119); scutes
smaller (Fig. 25b); ultimate and penultimate rays of dorsal and anal fins of equal length . Alepes vari
a) Alepes djedaba b) Alepes vari
c) Alepes apercna d) Alepes melanoptera

Fig. 25 lateral line
Key to the species of Carangoides occurring in the area

Note: species of Carangoides that have variable patterns of breast squamation may key out under both
sections of couplets when this character is utilized.
1a. Breast completely scaly or with a small, median naked area ventrally, scarcely if at all
visible in lateral view (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Breast partially to completely naked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
small naked area
small
naked area
lateral view ventral view (showing examples of variation)
Fig. 26
2a. Second dorsal fin with a conspicuous black blotch or submarginal band; vomerine tooth
patch anchor-shaped, with a long posteromedian extension (Fig. 27a, b) . . . . . . . . . . . . ® 3
2b. Second dorsal fin without a conspicuous black blotch or submarginal band; vomerine
tooth patch without a distinct posteromedian extension (Fig. 27c) . . . . . . . . . . . . . . . . ® 4
vomerine
tooth patch
vomerine vomerine
tooth patch tooth patch
a) Carangoides praeustus b) Carangoides equula c) Carangoides plagiotaenia
Fig. 27 tooth patches on roof of mouth and upper jaw
3a. Second dorsal fin with a conspicuous black blotch anteriorly (Fig. 28); soft anal-fin rays
18 to 20; total gill rakers on first gill arch 32 to 37 . . . . . . . . . . . . . . . Carangoides praeustus
3b. Second dorsal fin with a submarginal black band (Fig. 29); soft anal-fin rays 21 to 24;
total gill rakers on first gill arch 27 to 32 . . . . . . . . . . . . . . . . . . . . . Carangoides equula
Fig. 28 Carangoides praeustus Fig. 29 Carangoides equula
4a. Soft anal-fin rays 18 to 20; poste-

rior margin of preopercle outlined
in black, at least in adults
(Fig. 30); scutes 11 to 18 . . . .
. . . . . . . . . . . . Carangoides plagiotaenia
4b. Soft anal-fin rays 21 to 24; poste-
rior margin of preopercle not
black; scutes 20 to 30 . . . Carangoides bajad Fig. 30 Carangoides plagiotaenia
5a. Naked area of breast separated from naked base of pectoral fins by a broad band of
scales (Fig. 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5b. Naked area of breast uninterrupted to naked base of pectoral fins (Fig. 37) . . . . . . . . . . . ® 13
6a. Second dorsal fin with a conspicuous black blotch anteriorly (Fig. 28); vomerine tooth
patch anchor-shaped, with a long posteromedian extension (Fig. 27a) . . . Carangoides praeustus
6b. Second dorsal fin without a conspicuous black blotch; vomerine tooth patch without a
distinct posterior extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Soft dorsal-fin rays 25 to 34; soft anal-fin rays 21 to 26 . . . . . . . . . . . . . . . . . . . . . . ® 8

7b. Soft dorsal-fin rays 17 to 23; soft anal-fin rays 15 to 19 . . . . . . . . . . . . . . . . . . . . . . ® 10
2670 Bony Fishes
8a. Naked area of breast extends posteroventrally well beyond origin of pelvic fins (Fig. 31a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides fulvoguttatus
8b. Naked area of breast does not extend posteroventrally beyond origin of pelvic fins
(Fig. 31b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
naked area naked

long area short
a) Carangoides fulvoguttatus b) Carangoides ferdau, C. orthogrammus

Fig. 31
9a. In life, typically 5 or 6 distinct dusky bands on sides of adults and yellow or orange spots
on sides, if present, small, numerous and mostly above lateral line (Fig. 32); lips not
papillose in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides ferdau
9b. In life, dark bands usually absent on sides of adults and several relatively large, oblong
yellow spots with dark centres present mostly below lateral line (Fig. 33); lips finely
papillose in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides orthogrammus
bands typically bands usually

present absent
oblong
yellow spots
Fig. 32 Carangoides ferdau Fig. 33 Carangoides orthogrammus
10a. Straight part of lateral line slightly longer than curved part (Fig. 34); scutes 37 to 45
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides oblongus
10b. Straight part of lateral line slightly shorter than curved part (Fig. 35); scutes 16 to 38 . . . . . ® 11
straight part longer straight part shorter
Fig. 34 Carangoides oblongus Fig. 35 Carangoides dinema
11a. No small dark blotches on dorsum between bases of dorsal-fin rays . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides caeruleopinnatus
11b. Small dark blotches (becoming larger posteriorly) on dorsum between bases of dorsal-
fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Soft dorsal-fin rays 20 to 22 (rarely 19); large black spot usually present on shoulder;
naked area of breast extends posteroventrally well beyond origin of pelvic fins (Fig. 36a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides humerosus
12b. Soft dorsal-fin rays 17 to 19; no large black spot present on shoulder; naked area of
breast typically does not extend posteroventrally beyond origin of pelvic fins (Fig. 36b)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides dinema
naked area naked area

long shorter
a) Carangoides humerosus b) Carangoides dinema

Fig. 36
13a. Soft dorsaI-fin rays 25 to 32 (rarely 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13b. Soft dorsal-fin rays 17 to 23 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
14a. Profile of snout angular and, in specimens larger than about 30 cm fork length, horizontal
line from tip of snout distinctly below level of eye (Fig. 37a); soft anal-fin rays 21 to 26
(rarely 25 or 26); total gill rakers (including rudiments) on first gill arch 22 to 27, of which
6 to 8 on upper limb, and 17 to 21 on lower limb; vertebrae 10+14 . . . . Carangoides fulvoguttatus
14b. Profile of snout moderately rounded and, in specimens larger than about 30 cm fork
length, horizontal line from tip of snout at or through level of eye (Fig. 37b), soft anal-fin
rays 24 to 26 (usually 25); total gill rakers (including rudiments) on first gill arch 27 to
31, of which 7 to 9 on upper limb, and 19 to 22 on lower limb; vertebrae 10+15 . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides gymnostethus
a) Carangoides fulvoguttatus b) Carangoides gymnostethus

Fig. 37
15a. Small area naked of scales anteriorly just above pectoral-fin base (Fig. 38a) . . . . . . . . . . ® 16
15b. Area anteriorly just above pectoral-fin base completely scaly (Fig. 38b) . . . . . . . . . . . . . ® 17
no naked area
naked area here
a) Carangoides malabaricus b) Carangoides armatus

Fig. 38
2672 Bony Fishes
16a. Total gill rakers (including rudiments) on first gill arch 32 to 38, of which 8 to 12 on upper
limb, and 21 to 27 on lower limb; in life, tongue greyish brown to brown . . Carangoides malabaricus
16b. Total gill rakers (including rudiments) on first gill arch 27 to 31, of which 6 to 9 on upper
limb, and 19 to 22 on lower limb; in life, tongue white to pale grey . . . . Carangoides talamparoides
17a. Small dark blotches (becoming larger posteriorly) on back between bases of dorsal-fin
rays; naked area of breast typically does not extend posteroventrally beyond origin of
pelvic fins (Fig. 36b); soft dorsal-fin rays 17 to 19 . . . . . . . . . . . . . . . . . Carangoides dinema
17b. Colour pattern not as above; naked area of breast extends posteroventrally well beyond
origin of pelvic fins (Fig. 38b); soft dorsal-fin rays 18 to 23 . . . . . . . . . . . . . . . . . . . . ® 18
18a. Total gill rakers (including rudiments) on first gill arch 31 to 37 . . . . . . . . . Carangoides armatus
18b. Total gill rakers (including rudiments) on first gill arch 20 to 27 . . . . . . . . . . . . . . . . . . ® 19
19a. Adults with a steep dorsal head profile, and with a distinct break (“bump”) in profile in the
interorbital region (Fig. 39a); 3 to 8 (usually 5 to 7) of central soft rays of dorsal and anal
fins elongated in mature males (about 17 cm fork length); eye diameter about equal to,
or larger than snout length . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides hedlandensis
19b. Dorsal head profile not as steep in adults, and no distinct break (“bump”) in profile in the
interorbital region (Fig. 39b, c); central soft rays of dorsal and anal fins not elongated in
mature males; eye diameter slightly to much smaller than snout length . . . . . . . . . . . . . ® 20
20a. Dorsal profile of snout gently sloped, then abruptly vertical just above mouth cleft
(Fig. 39b); soft dorsal-fin rays 18 to 20; soft anal-fin rays 15 to 17 . . . . . Carangoides chrysophrys
20b. Dorsal profile of snout not as above (Fig. 39c), soft dorsal-fin rays 20 to 23 (usually 22
or 23); soft anal-fin rays 16 to 20 (usually 18 or 19) . . . . . . . . . . Carangoides caeruleopinnatus
bump
a) Carangoides hedlandensis b) Carangoides chrysophrys c) Carangoides caeruleopinnatus
Fig. 39
Key to the species of Caranx occurring in the area

Note: species of Caranx that have variable patterns of breast squamation will key out under both sections
of a couplet when this character is utilized.
1a. Breast completely scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Breast naked ventrally, frequently with small patch of prepelvic scales (Fig. 40) . . . . . . . . . ® 6
2a. In life, body coloration essentially uniform grey to brown; lobe of dorsal fin relatively long,
contained 2.3 to 5.3 times in fork length; profile of head relatively steep and angular
(Fig. 41) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx lugubris
2b. Body coloration not as above; lobe of dorsal fin contained 4.2 to 8.8 times in fork length;
profile of head not noticeably steep and angular . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
prepelvic
scale patch naked
area
pelvic fins
Fig. 40 ventral view Fig. 41 Caranx lugubris
3a. Small black spots scattered on head and body (forming at about 16 to 22 cm fork length);
snout length contained 9.2 to 12.7 times in fork length; total gill rakers (including
rudiments) on first gill arch 25 to 29 (usually 26 or 27), and total soft dorsal- and anal-fin
rays 39 to 44 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx melampygus
3b. No small black spots scattered on head and body; snout length contained 13.1 to 18.4
times in fork length; total gill rakers (including rudiments) on first gill arch 22 to 25, except
24 to 27 in C. heberi which has 34 to 38 total soft dorsal- and anal-fin rays . . . . . . . . . . . . ® 4
4a. No small black spot present on upper margin of opercle; upper lobe of caudal fin
frequently with distal half noticeably dark or black, especially in juveniles; in adults,
adipose eyelid only slightly developed (Fig. 42a) . . . . . . . . . . . . . . . . . . . . Caranx heberi
4b. A small, black spot present on upper margin of opercle; upper lobe of caudal fin usually
uniformly pigmented; in adults, adipose eyelid well developed, especially posteriorly
(Fig. 42b, c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. In adults, dorsal-fin lobe without white tip; dorsal profile of head strongly convex and a
black spot on upper margin of opercle, in adults, at least 1/2 the diameter of pupil
(Fig. 42b); in specimens larger than 15 cm fork length, postorbital head length longer,
contained 5.7 to 7.3 times in fork length, and dorsal-fin lobe shorter, contained 5.7 to 8.8
times in fork length; vertebrae 10+14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx tille
5b. In adults, dorsal-fin lobe with white tip; dorsal profile of head moderately convex and
black spot on upper margin of opercle, its size in adults no larger than twice the diameter
of pupil (Fig. 42c); in specimens larger than 15 cm fork length, postorbital head length
shorter, contained 6.4 to 8.2 times in fork length, and dorsal-fin lobe longer, contained 5
to 6.6 times in fork length; vertebrae 10+15 . . . . . . . . . . . . . . . . . . . . Caranx sexfasciatus
opercular opercular
adipose eyelid spot spot
a) Caranx heberi b) Caranx tille c) Caranx sexfasciatus
Fig. 42
2674 Bony Fishes
6a. Naked area of breast uninterrupted to naked base of pectoral fins (Fig. 43); curved part
of lateral line short, chord of curved part contained 2.5 to 3.3 times in straight part; scutes
larger (Fig. 44a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx bucculentus
6b. Naked area of breast separated from naked base of pectoral fins by a broad band of
scales; curved part of lateral line moderate, with chord of curved part contained less than
1.5 times in straight part; scutes smaller (Fig. 44b) . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
curved
part very
short
a) Caranx bucculentus
naked area b) Caranx ignobilis
Fig. 43 Caranx bucculentus Fig. 44 lateral line
7a. Total gill rakers (including rudiments) 20 to 24 on first gill arch; general body colour
silvery to black; in specimens larger than 15 cm fork length, body depth contained 2.5 to
3.2 times in fork length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx ignobilis
7b. Total gill rakers (including rudiments) 23 to 30 (rarely 23) on first gill arch; general body
colour bronze to yellow-green; in specimens larger than 15 cm fork length, body depth
contained 2.7 to 3.8 times in fork length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Total soft dorsal- and anal-fin rays 34 to 38 (rarely

38); upper lobe of caudal fin frequently with distal brilliant white spot
half noticeably dark or black (especially in juve-
niles), and posterior margin of lower lobe without
a narrow white border; no pale spot on shoulder
just behind posterodorsal margin of opercle;
adults without small black spots . . . . . . . . Caranx heberi
8b. Total soft dorsal- and anal-fin rays 37 to 41 (rarely
37); upper lobe of caudal fin usually uniformly
pigmented, and posterior margin of lower lobe
with a narrow white border; in life a conspicuous
pale spot, approximately the diameter of pupil, on
shoulder just behind posterodorsal margin of op-
ercle (Fig. 45); adults with small black spots on
body above lateral line (forming at about 25 cm
fork length) . . . . . . . . . . . . . . . . . . Caranx papuensis
Fig. 45 Caranx papuensis
Key to the species of Decapterus occurring in the area

1a. Posterior end of upper jaw concave above, rounded and produced below (Fig. 46a);
straight part of lateral line with 14 to 29 scales (Fig. 47a) . . . . . . . . . . . Decapterus macrosoma
1b. Posterior end of upper jaw straight above, straight to slightly concave below (Fig. 46b-e);
straight part of lateral line with 0 to 15 scales (except 18 to 39 scales in D. macarellus) . . . . . ® 2
a) Decapterus macrosoma b) Decapterus macarellus
c) Decapterus tabl d) Decapterus russelli e) Decapterus kurroides
Fig. 46
2a. Posterior end of upper jaw noticeably slanted anteroventrally (Fig. 46b); straight part of
lateral line with 18 to 39 scales and 24 to 40 scutes = 52 to 67 total (Fig. 47b); pectoral
fins usually shorter in adults, 58 to 72% head length; oral valve (membrane) at symphysis
of upper jaw conspicuously white (Fig. 48) . . . . . . . . . . . . . . . . . . . Decapterus macarellus
2b. Posterior end of upper jaw not as noticeably slanted anteroventrally (Fig. 46c-e); straight
part of lateral line with 0 to 10 scales and 30 to 40 scutes = 30 to 49 total (except 5 to 15
scales and 32 to 42 scutes = 41 to 50 total in D. muroadsi; Fig. 47c); pectoral fins usually
longer in adults, 71 to 105% head length (except 62 to 83% in D. tabl); oral valve
(membrane) at symphysis of upper jaw dusky or transparent (except white in
D. muroadsi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
curved part of lateral line straight part

14-29 scutes
scales 18-39
scales
a) Decapterus macrosoma b) Decapterus macarellus
5-15 scales 4-10 scales
c) Decapterus muroadsi d) Decapterus tabl
0-4 scales no scales
e) Decapterus russelli f) Decapterus kurroides

2676 Bony Fishes
oral valve
3a. In life, caudal fin with upper lobe greenish yellow (or membrane)
and lower lobe grey; oral valve (membrane) at white
symphysis of upper jaw conspicuously white in
adults (Fig. 48); gill rakers on lower limb of first gill
arch 36 to 42 . . . . . . . . . . . . . . . Decapterus muroadsi
3b. In life, caudal fin with upper and lower lobes both
hyaline, brownish or red; oral valve (membrane)
at symphysis of upper jaw transparent or dusky;
gill rakers on lower limb of first gill arch 26 to 33 Fig. 48 ventral view of upper
(except 30 to 39 in D. russelli) . . . . . . . . . . . . . . . ® 4 jaw and roof of mouth
4a. Scutes relatively small (Fig. 47d); curved part of

lateral line with 61 to 73 scales; straight part of
lateral line with 4 to 12 anterior scales; total lat-
eral-line scales and scutes (excluding scales on
caudal fin) 103 to 118; in life, caudal fin red;
posterodorsal margin of opercular membrane mi-
nutely serrated in large adults (Fig. 49) . . . . Decapterus tabl
4b. Scutes relatively large (Fig. 47e, f); curved part of
lateral line with 42 to 62 scales; straight part of
lateral line with 0 to 4 anterior scales; total lateral-
line scales and scutes (excluding scales on cau-
dal fin) 77 to 102; in life, caudal fin dusky or red; Fig. 49 Decapterus tabl
posterodorsal margin of opercular membrane
smooth in adults . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. In life, caudal fin hyaline to dusky; gill rakers on lower limb of first gill arch usually 32 to
39; soft anal-fin rays (including finlet) 25 to 29 (rarely 25); eye usually smaller . . Decapterus russelli
5b. In life, caudal fin red; gill rakers on lower limb of first gill arch usually 26 to 32; soft anal-fin
rays (including finlet) 22 to 26 (rarely 26); eye usually larger . . . . . . . . . . Decapterus kurroides
Key to the species of Pseudocaranx occurring in the area

1a. Scales in curved part of lateral line
57 to 78; no scales on preorbital
bone below and in front of eye and
on expanded part of maxilla; pos-
terior end of upper jaw essentially
vertical (Fig. 50a); total gill rakers
(including rudiments) 29 to 35 on
first gill arch . . . . . . . Pseudocaranx dentex
1b. Scales in curved part of lateral line
37 to 48; scales present on preorbi-
tal bone below and in front of eye
and on expanded part of maxilla;
posterior end of upper jaw slanted a) Pseudocaranx dentex b) Pseudocaranx wrighti
posteroventrally (Fig. 50b); total gill
rakers (including rudiments) 36 to Fig. 50
44 on first gill arch . . . . Pseudocaranx wrighti
(western and southern Australia; not yet recorded
from the area, but easily confused with P. dentex and
therefore included here)
Key to the species of Scomberoides occurring in the area

1a. Total gill rakers 8 to 15 on first gill
arch; scales on midbody below
lateral line broadly oblong or
blunty lanceolate (Fig. 51a, b);
lobe of dorsal fin uniformly pig-
mented (Figs 52 and 53) . . . . . . . . . ® 2
1b. Total gill rakers 21 to 27 on first
gill arch; scales on midbody be-
low lateral line sharply lanceo-
a) Scomberoides commersonnianus b) Scomberoides tala
late or needle-like (Fig. 51c, d);
distal half of dorsal-fin lobe
abruptly and heavily pigmented
(Figs 55 and 56) . . . . . . . . . . . . . . ® 3
2a. Upper jaw extends well beyond

posterior margin of eye, espe-
cially in adults (Fig. 52); in adults,
teeth of inner and outer rows in c) Scomberoides lysan d) Scomberoides tol
lower jaw subequal in size
(Fig. 54a); in life, large oval Fig. 51 scales on midbody below lateral line
blotches above or touching lat-
eral line . . Scomberoides commersonnianus
2b. Upper jaw extends slightly beyond posterior margin of eye (Fig. 53); in adults, inner row
of teeth in lower jaw distinctly larger than those in outer row (Fig. 54b); in life, vertically
elongate blotches intersecting lateral line . . . . . . . . . . . . . . . . . . . . . . Scomberoides tala
Fig. 52 Scomberoides commersonnianus Fig. 53 Scomberoides tala
a) Scomberoides commersonnianus
b) Scomberoides tala
Fig. 54 teeth in lower jaw (teeth in inner row black)
3a. Scales on midbody below lateral line sharply lanceolate (Fig. 51c); in adults, upper jaw
extends to or slightly beyond posterior margin of eye (Fig. 55); in life, a double series of
6 to 8 dusky, roundish blotches above and below lateral line, occasionally connected by
narrow isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scomberoides lysan
3b. Scales on midbody below lateral line slender, needle-like (Fig. 51d); in adults, upper jaw
does not extend to posterior margin of eye (Fig. 56); in life, oval or vertically oblong dark
blotches, the first 4 or 5 intersecting lateral line . . . . . . . . . . . . . . . . . . . . Scomberoides tol
2678 Bony Fishes
Fig. 55 Scomberoides lysan Fig. 56 Scomberoides tol
Key to the species of Selar occurring in the area

1a. Curved part of lateral line with 48 to 56 scales; curved part of lateral line moderate, with
chord of curved part contained 0.7 to 1.2 times in straight part; scutes smaller (Fig. 57a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Selar crumenophthalmus
1b. Curved part of lateral line with 21 to 24 scales; curved part of lateral line short, chord of
curved part contained 2.1 to 3 times in straight part; scutes larger (Fig. 57b) . . . . . . . Selar boops
48-56 scales
21-24
scales
a) Selar crumenophthalmus b) Selar boops
Key to the species of Seriola occurring in the area

1a. Soft dorsal- and anal-fin rays 22 to 25 and 15 to 17 (rarely 17), respectively; in specimens
larger than about 20 cm fork length, total gill rakers (excluding rudiments) on first gill arch
10 to 12; in life, papillae surrounding broad band of teeth in both jaws engorged with
blood giving teeth a red appearance . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola hippos
1b. Soft dorsal- and anal-fin rays 27 to 35 and 18 to 22, respectively; in specimens larger
than about 20 cm fork length, total gill rakers (excluding rudiments) on first gill arch 11
to 29; in life, papillae surrounding broad band of teeth in both jaws not engorged with
blood, appearance of teeth white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Upper jaw moderately slender posteriorly, with slender supramaxilla (Fig. 58a); caudal
fin yellowish; in adults, a moderate cutaneous keel laterally on caudal peduncle; verte-
brae 11+14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola lalandi
2b. Upper jaw relatively broad posteriorly, with broad supramaxilla (Fig. 58b-d); caudal fin
dark to dusky sometimes with a lighter posterior margin; in adults, cutaneous keel on
caudal peduncle absent to slightly developed; vertebrae 10+14 . . . . . . . . . . . . . . . . . . ® 3
supramaxilla supramaxilla supramaxilla supramaxilla

narrow broad broad broad
a) Seriola lalandi b) Seriola rivoliana c) Seriola dumerili d) Seriola hippos
Fig. 58 lateral view of head (supramaxilla shaded)
3a. In adults, length of dorsal-fin lobe about 1.3 to 1.6 times longer than pectoral fins
(Fig. 59a) and 18 to 22% of fork length; in specimens larger than about 20 cm fork length,
total gill rakers (excluding rudiments) 22 to 26 on first gill arch; first pterygiophore of anal
fin straight in specimens larger than about 10 cm fork length (Fig. 60a) . . . . . . . Seriola rivoliana
3b. In adults, length of dorsal-fin lobe about equal to, or slightly longer than pectoral fins
(Fig. 59b) and 13 to 18% of fork length; in specimens larger than about 20 cm fork length,
total gill rakers (excluding rudiments) 11 to 19 on first gill arch; first pterygiophore of anal
fin moderately curved (Fig. 60b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola dumerili
vertebrae
a) Seriola rivoliana
pterygiophore
pterygiophore curved
straight ventral
margin
of belly
anterior
margin of
first 2 spines detached from anal fin anal fin
(embedded in skin in large individuals)
b) Seriola dumerili
a) Seriola rivoliana b) Seriola dumerili
Fig. 59 Fig. 60
Key to the species of Trachinotus occurring in the area

1a. One to 7 spots in a longitudinal row on or
near lateral line (spots absent on speci-
mens smaller than about 10 to 13 cm
fork length); soft dorsal-fin rays 21 to 25 . . . . . ® 2
1b. No spots in a longitudinal row on or near
lateral line; soft dorsal-fin rays 18 to 20
(except 21 to 23 in T. africanus) . . . . . . . . . ® 4
2a. In adults, all spots equal to, or smaller
than eye diameter, and with about half of
spot below lateral line (Fig. 61); gill rak-
ers (including rudiments) on lower limb Fig. 61 Trachinotus baillonii
of first gill arch 15 to 19; vomerine and
palatine tooth patches typically type “A”
(Fig. 62a) . . . . . . . . . . . . Trachinotus baillonii
2b. In adults, anterior 2 spots larger than eye diameter and at least two-thirds of spot above
lateral line (Figs 63 and 64); gill rakers (including rudiments) on lower limb of first gill arch
11 to 15; vomerine and palatine tooth patches usually type “B” or “C” (Fig. 62b, c) . . . . . . . . ® 3
premaxilla
vomer
palatines
vomer
palatines
a) type “A” b) type “B” c) type “C”

(Trachinotus baillonii) (Trachinotus botla) (Trachinotus copperingi )
Fig. 62 tooth patches on roof of mouth and upper jaw
2680 Bony Fishes
3a. Soft anal-fin rays 19 to 21; in adults only 1 dark spot above pectoral fins, and in large
specimens spots oval-shaped (Fig. 63); dorsal-fin lobe usually longer than anal-fin lobe
in specimens larger than about 25 cm fork length; pelvic fins long, their length contained
1.5 to 1.7 times in pectoral-fin length in specimens larger than about 25 cm fork length;
vomerine tooth patch usually type “B” (Fig. 62b) . . . . . . . . . . . . . . . . . . . Trachinotus botla
3b. Soft anal-fin rays 22 to 24; in adults 2 dark spots above pectoral fins, and in large
specimens spots more vertically elongate (Fig. 64); dorsal-fin lobe consistently shorter
than anal-fin lobe in specimens larger than about 25 cm fork length; pelvic fins relatively
short, their length contained 1.7 to 2.2 times in pectoral-fin length in specimens larger
than about 25 cm fork length; vomerine tooth patch usually type “C” (Fig. 62c) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachinotus copperingi
Fig. 63 Trachinotus bolta Fig. 64 Trachinotus copperingi
4a. Soft dorsal- and anal-fin rays 21 to 23 and 19 to 21, respectively; body often with a black
oval-shaped blotch in axillary base under pectoral fins . . . . . . . . . . . . . Trachinotus africanus
4b. Soft dorsal- and anal-fin rays 18 to 20 and 16 to 18, respectively; body uniformly
pigmented in axillary base under pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. First predorsal bone inverted teardrop- or oval-shaped (Fig. 65a, this character is easily
observed by simple dissection along midline of nape); height of dorsal-fin lobe longer,
35 to 60% of fork length in specimens 10 to 40 cm fork length . . . . . . . . . . Trachinotus blochii
5b. First predorsal bone shaped like an inverted “L” with the arm projecting anteriorly
(Fig. 65b, c); anal-fin lobe yellow without a brownish anterior margin; height of dorsal-fin
lobe moderate, 24 to 37% of fork length in specimens 10 to 40 cm fork length . . . . . . . . . . ® 6
teardrop- or oval-shaped inverted L-shape inverted L-shape
a) Trachinotus blochii b) Trachinotus mookalee c) Trachinotus anak
Fig. 65 anterior part of skeleton in lateral view

(hyperossified bones shown in black)
6a. T e e t h in a na r r o w ba n d on
tongue, persisting to about 50 cm
fork length; supraoccipital bone of
skull becoming broad and sau-
sage-shaped (Fig. 66a) in speci-
mens larger than about 30 cm fork dorsal
length; profile of snout evenly supraoccipital bone view
rounded in specimens larger than supraoccipital bone
about 20 cm fork length; preorbi-
tal and nasal bones never exhibit-
ing hyperostosis . . . . Trachinotus mookalee
6b. No teeth on tongue; supraoccipi-
tal bone of skull thin and blade-
like in adults (Fig. 66b); profile of
lateral
snout essentially straight immedi- view
ately above upper lip in speci-
mens larger than about 20 cm fork a) Trachinotus mookalee b) Trachinotus anak
length; preorbital and nasal bones Fig. 66 skull
hyperossified in specimens larger Note: the skull of Trachinotus africanus is used here for
than about 30 cm fork length T. anak; the supraoccipital bone shape is nearly identical in
(Fig. 65c) . . . . . . . . . . Trachinotus anak both species
Key to the species of Trachurus occurring in the area

1a. Dorsal accessory lateral line immediately below base of dorsal fin terminates below fifth
to eleventh (usually seventh to ninth) soft dorsal-fin ray; total scales and scutes in lateral
line 71 to 89 (usually more than 75); curved part of lateral line essentially parallel to axis
of body for most of its length (Fig. 67) . . . . . . . . . . . . . . . . . . . . . . . . Trachurus declivis
1b. Dorsal accessory lateral line immediately below base of dorsal fin terminates below first
to fifth (usually first or second) soft dorsal-fin ray; total scales and scutes in lateral line
67 to 81 (rarely more than 76); curved part of lateral line slanted for most of its length
(Fig. 68) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachurus novaezelandia
dorsal accessory
lateral line
Fig. 67 Trachurus declivis Fig. 68 Trachurus novaezelandiae
Key to the species of Ulua occurring in the area

1a. Total gill rakers on first gill arch 54 to 61, of which 16 to 21 on upper limb, and 37 to 41
on lower limb; tongue with central band of villiform teeth; in specimens smaller than about
15 cm fork length, first anal-fin ray produced but not filamentous, not extending beyond
caudal peducle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulua aurochs
1b. Total gill rakers on first gill arch 74 to 86, of which 23 to 27 on upper limb, and 51 to 61
on lower limb; tongue without central band of villiform teeth; in specimens smaller than
about 15 cm fork length, first anal-fin ray filamentous, extending beyond caudal peduncle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulua mentalis
2682 Bony Fishes
Key to the species of Uraspis occurring in the area

1a. Naked area of breast extends
uninterrupted to naked base
of pectoral fins (Fig. 69a);
scales in curved part of lateral
line 61 to 82 . . . . . . . . Uraspis uraspis
1b. Naked area of breast sepa- continuous
rated from naked base of pec- continuous broad band
naked area naked area
toral fins by a broad band of of scales
scales (Fig. 69b); scales in
curved part of lateral line 48 to a) Uraspis uraspis b) Uraspis helvola and U. secunda
66 . . . . . Uraspis helvola and U. secunda
Fig. 69
(no verified records from the area; adults of these
2 species are virtually impossible to distinguish)

The generic limits and intrafamilial relationships of carangids are not well established; a few of the generic
units recognized here are subject to change and are used for convenience. At least 5 additional species,
“Caranx” koheru Hector, Pseudocaranx wrighti (Whitley), Seriola quinqueradiata Temminck and
Schlegel, Trachurus japonicus (Temminck and Schlegel), and T. murphyi Nichols, occur in the western
Pacific Ocean outside of the area of coverage.
The symbol 0 is given when species accounts are included. Species with a question mark have not yet
been recorded from the area but should be watched for.
0 Alectis ciliaris (Bloch, 1788)
0 Alectis indica (Rüppell, 1830)
0 Alepes apercna Grant, 1987
0 Alepes djedaba (Forsskål, 1775)
0 “Alepes” kleinii (Bloch, 1793)
0 Alepes melanoptera Swainson, 1839
1/
0 Alepes vari (Cuvier, 1833)

0 Atropus atropos (Schneider, 1801)
0 Atule mate (Cuvier, 1833)
0 Carangoides armatus (Rüppell, 1830)
0 Carangoides bajad (Forsskål, 1775)
0 Carangoides caeruleopinnatus (Rüppell, 1830)
0 Carangoides chrysophrys (Cuvier, 1833)
0 Carangoides dinema Bleeker, 1851
0 Carangoides equula (Temminck and Schlegel, 1844)
0 Carangoides ferdau (Forsskål, 1775)
0 Carangoides fulvoguttatus (Forsskål, 1775)
0 Carangoides gymnostethus (Cuvier, 1833)
0 Carangoides hedlandensis (Whitley, 1933)
0 Carangoides humerosus (McCulloch, 1915)
0 Carangoides malabaricus (Bloch and Schneider, 1801)
0 Carangoides oblongus (Cuvier, 1833)
0 Carangoides orthogrammus (Jordan and Gilbert, 1882)
0 Carangoides plagiotaenia Bleeker, 1857
0 Carangoides praeustus (Bennett, 1830)
0 Carangoides talamparoides Bleeker, 1852
0 Caranx bucculentus Alleyne and Macleay, 1877
0 Caranx ignobilis (Forsskål, 1775)
0 Caranx heberi (Bennett, 1830)
0 Caranx lugubris Poey, 1860
1/ Generic allocation of this species uncertain; might eventually be assigned to a separate, monotypic genus.
0 Caranx melampygus Cuvier, 1833

0 Caranx papuensis Alleyne and Macleay, 1877
0 Caranx sexfasciatus Quoy and Gaimard, 1825
0 Caranx tille Cuvier, 1833
0 Decapterus kurroides Bleeker, 1855
0 Decapterus macarellus (Cuvier, 1833)
0 Decapterus macrosoma Bleeker, 1851
0 Decapterus muroadsi (Temminck and Schlegel, 1844)
0 Decapterus russelli (Rüppell, 1830)
0 Decapterus tabl Berry, 1967
0 Elagatis bipinnulata (Quoy and Gaimard, 1825)
0 Gnathanodon speciosus (Forsskål, 1775)
0 Megalaspis cordyla (Linnaeus, 1758)
0 Naucrates ductor (Linnaeus, 1758)
0 Pantolabus radiatus (Macleay, 1881)
0 Parastromateus niger (Bloch, 1795) 2/
0 Pseudocaranx dentex (Bloch and Schneider, 1801)

0 Scomberoides commersonnianus Lacepède, 1801
0 Scomberoides lysan (Forsskål, 1775)
0 Scomberoides tala (Cuvier, 1832)
0 Scomberoides tol (Cuvier, 1832)
0 Selar boops (Cuvier, 1833)
0 Selar crumenophthalmus (Bloch, 1793)
0 Selaroides leptolepis (Cuvier, 1833)
0 Seriola dumerili (Risso, 1810)
0 Seriola hippos Günther, 1876
0 Seriola lalandi Valenciennes, 1833
0 Seriola rivoliana Valenciennes, 1833
0 Seriolina nigrofasciata (Rüppell, 1829)
0 Trachinotus africanus Smith, 1967
0 Trachinotus anak Ogilby, 1909
0 Trachinotus baillonii (Lacepède, 1801)
0 Trachinotus blochii (Lacepède, 1801)
0 Trachinotus botla (Shaw, 1803)
0 Trachinotus copperingi (Günther, 1884)
0 Trachinotus mookalee Cuvier, 1832
0 Trachurus declivis (Jenyns, 1841)
0 Trachurus novaezelandiae Richardson, 1843
0 Ulua aurochs (Ogilby, 1915)
0 Ulua mentalis (Cuvier, 1833)
0? Uraspis helvola (Forster, 1801)
0? Uraspis secunda (Poey, 1860)
0 Uraspis uraspis (Günther, 1860)
References
Gunn, J.S. 1990. A revision of selected genera of the family Carangidae (Pisces) from Australia. Rec. Aust. Mus., Supp.
(12):77 p.
Gushiken, S. 1983. Revision of the carangid fishes of Japan. Galaxea, 2:135-264.
Lin, P.-L. and K.-T. Shao. 1999. A review of the carangid fishes (family Carangidae) from Taiwan with descriptions of
five new records. Zool. Stud. , 38(1):33-68.
2/ Many previous authors have assigned this species to a separate family, Apolectidae (or Formionidae)
click for next page

2684 Bony Fishes
Alectis ciliaris (Bloch, 1788)

Frequent synonyms / misidentifications: Atlantic Ocean: Alectis crinitus (Mitchill, 1826); Hawaii:
Carangoides ajax Snyder, 1904 / None.
FAO names: En - African pompano; Fr - Cordonnier fil; Sp - Pámpano de hebra.
61 cm fork length
Diagnostic characters: Body deep, becoming more

elongate with growth, and very compressed. Profile
of nape and head broadly rounded; suborbital
depth relatively narrow, contained 1.7 to 3 times in
upper jaw length; upper jaw extends to below middle
to posterior third of eye. Both on jaws with bands of
villiform teeth, becoming obsolete with age. Gill rak-
ers (excluding rudiments) 4 to 6 on upper limb and
12 to 17 on lower limb of first gill arch (total 18 to
22). Dorsal fin with VII short spines (embedded and juvenile
not apparent at about 17 cm fork length) followed by 13 cm standard length
I spine and 18 to 20 soft rays; anal fin with II spines
(embedded and not apparent with growth) followed by I spine and 15 to 17 soft rays; anterior soft rays of
dorsal and anal fins extremely long and filamentous in young; pectoral fins falcate, longer than head; pelvic
fins elongate in young. Lateral line anteriorly with a strong and moderately long arch, its posterior (straight)
part with 12 to 30 scutes; body superficially naked, scales minute and embedded where present.
Vertebrae 10+14; no hyperostosis. Colour: mostly silvery with a light metallic bluish tinge on upper third
of body and head; juveniles with 5 chevron-shaped dark bars on body, and a black blotch at base of third
to sixth soft dorsal-fin rays, filaments black distally.
Size: Maximum total length possibly 130 to 150 cm, but reliable data not available; commonly to 100 cm
fork length, largest specimen examined 109 cm fork length and 16.5 kg. World angling record 18.8 kg.
Habitat, biology, and fisheries: Adults are usually solitary and frequent shallow coastal waters to a depth
of about 100 m; young usually pelagic and drifting. Small pelagic juveniles with very filamentous dorsal
and anal fins resemble medusae, and this “mimicry” may gain them some protection from predators. The
diet consists of slow-swim-
ming or sedentary crusta-
ceans; small crabs and occa-
sionally small fishes
sometimes eaten. Adults are
mainly taken on hook-and-
line, especially on light tackle.
Juveniles are often taken in
beach seines.
Distribution: Circumtropical
in marine waters; widespread
in the tropical Indo-West Pa-
cific.
Alectis indica (Rüppell, 1830)

FAO names: En - Indian threadfish; Fr - Cordonnier plume; Sp - Pámpano indico.
69 cm fork length
Diagnostic characters: Body deep, becoming more elongate with growth, and very compressed. Profile
of nape and head somewhat angular; suborbital depth relatively broad, contained 0.8 to 1 times in upper
jaw length; upper jaw extends to slightly before eye (to below anterior margin of eye in young). Both jaws
with bands of villiform teeth, becoming obsolete with age. Gill rakers (excluding rudiments) 8 to 11 on
upper limb and 21 to 26 on lower limb of first gill arch (total 29 to 37). Dorsal fin with VI short spines
(embedded and not apparent at about 17 cm fork length) followed by I spine and 18 to 20 soft rays; anal
fin with II spines (embedded and not apparent with growth) followed by I spine and 15 to 17 soft rays;
anterior soft rays of dorsal and anal fins extremely long and filamentous in young; pectoral fins falcate,
longer than head; pelvic fins elongate in young. Lateral line anteriorly with a strong and moderately long
arch, its posterior (straight) part with 6 to 11 scutes; body superficially naked, scales minute and
embedded where present. Vertebrae 10+14; supraoccipital and distal ends of supraneural (predor-
sal) bones hyperossified and enlarged distally in specimens larger than 25 cm fork length. Colour:
mostly silvery with dusky green tinge dorsally; juveniles with dark bars on body; second dorsal fin pale with
leading edges and tips of rays dusky; anal fin pale yellow and caudal fin dusky green.
Size: Maximum reported total length 150 cm; South African spear fishing record 21.4 kg.
Habitat, biology, and fisheries: Adults are reported to concentrate in large shoals and are wide ranging
in tropical coastal waters including reef areas; juveniles are solitary and often enter estuaries. The diet
consists of a wide range of fishes, small squids, and crustaceans. Adults are mainly taken on hook-and-line,
especially on light tackle. Juveniles are often taken in beach seines.
coastal waters of tropical
Indo-West Pacific, from East
Africa, including the Mala-
gasy Republic, Red Sea, and
Persian Gulf to Papua New
Guinea; north to the Ryukyu
Islands and south to Austra-
lia.
2686 Bony Fishes
Alepes apercna Grant, 1987

FAO names: En - Smallmouth scad.
22.5 cm fork length
(drawn by T.D. Pedersen)
Diagnostic characters: Body oblong, compressed, with dorsal and ventral profiles almost evenly convex;
snout bluntly rounded. Eye diameter about equal to snout length, with adipose eyelid well developed on
posterior half of eye only. Upper jaw narrowly rounded posteriorly, and supramaxilla relatively small
without an anterior spine-like extension; both jaws with a single row of numerous, comb-like teeth. Gill
rakers (including rudiments) 8 to 10 on upper limb and 18 to 21 on lower limb of first gill arch (total 27
to 30). Shoulder girdle (cleithrum) margin smooth, without papillae. Two separate dorsal fins, the first with
VII spines, the second with I spine and 24 to 26 soft rays; anal fin with II detached spines followed by I
spine and 20 to 22 soft rays; spinous dorsal fin moderately high, longest spine height about equal to length
of soft dorsal-fin lobe; ultimate and penultimate rays of dorsal and anal fins of equal length. Lateral line
strongly arched anteriorly, with junction of curved and straight parts below vertical from between second
dorsal-fin origin and third soft ray; chord of curved part of lateral line contained 1.7 to 2.3 times in straight
part (to caudal-fin base); scales in curved part of lateral line 35 to 48; 0 to 3 scutes in curved part; straight
part with 0 to 2 scales and 50 to 64 scutes; total scales and scutes in lateral line (excluding scales on
caudal fin) 90 to 113. Vertebrae 10+14. Colour: in life, green to bluish green above, silvery to white below;
a diffuse dusky blotch on margin of opercle, not bordered above by a white spot; interradial membranes
of spinous dorsal fin hyaline to dusky; spinous and second dorsal, anal, and caudal fins dusky yellow,
with anterior rays of second dorsal and anal fins often with white tips; caudal-fin lobes often with dark tips;
pelvic fins white and pectoral fins hyaline; juveniles with 7 dark bands on body.
Size: A small species, largest specimen examined 25 cm fork length and 29.5 cm total length.
Habitat, biology, and fisheries: An inshore species. Feeds primarily on small invertebrates, primarily
microscopic crustaceans and molluscs. Caught mainly on hook-and-line.
Remarks: Grant (1987, Fishes of Australia) used the above scientific name (taken from manuscript
paratypes deposited in the Queensland Museum) together with a very brief description and photograph,
incorrectly crediting the species to Smith-Vaniz; according to the rules of the International Code of
Zoological Nomenclature,
this action qualifies as an
original description and be-
cause the “description” was
solely Grant’s he becomes
the author of the species.
from Australia: Exmouth Gulf,
western Australia (22°05’S) to
Wide Bay, Queensland
(25°52’S); may also occur in
the Gulf of Papua from where
not yet reported.
Alepes djedaba (Forsskål, 1775)

Frequent synonyms / misidentifications: Atule djedaba (Forsskål, 1775) [this name has frequently been
misspelled “A. djeddaba”]; Caranx djedaba (Forsskål, 1775) / Alepes vari (Cuvier, 1833).
FAO names: En - Shrimp scad; Fr - Sélar subari; Sp - Jurel subari.
23.5 cm fork length
snout pointed. Eye diameter about equal to snout length, with adipose eyelid well developed on posterior
half of eye only. Upper jaw slightly concave posteriorly, and supramaxilla relatively large with an
anterior spine-like extension; both jaws with a single row of numerous, comb-like teeth. Gill rakers
(including rudiments) 10 to 14 on upper limb and 27 to 33 on lower limb of first gill arch (total 38 to
47). Shoulder girdle (cleithrum) margin smooth, without papillae. Two separate dorsal fins, the first with
VIII spines, the second with I spine and 23 to 25 soft rays; anal fin with II detached spines followed by I
of soft dorsal-fin lobe; ultimate ray of dorsal and anal fins about 1.3 to 1.5 times the length of penultimate
ray. Lateral line strongly arched anteriorly, with junction of curved and straight parts below vertical from
between origin of second dorsal fin and third soft ray; chord of curved part of lateral line contained 2 to 2.6
times in straight part (to caudal-fin base); scales in curved part of lateral line 31 to 36; 0 to 3 scutes in
curved part; straight part with 0 to 2 scales and 39 to 51 scutes; total scales and scutes in lateral line
(excluding scales on caudal fin) 77 to 85. Vertebrae 10+14. Colour: in life, greyish green above, silvery
to white below; a distinct black blotch on margin of opercle, bordered above by a smaller white spot;
interradial membranes of spinous dorsal fin pale to dark dusky; dorsal-fin lobe dusky, pale distally;
caudal fin yellowish, except upper lobe often dusky to black distally; other fins pale.
Size: Largest specimen examined 29 cm fork length; seldom exceeds 25 cm fork length.
Habitat, biology, and fisheries: Rather common near inshore reefs, even in moderately turbid water,
often forming large schools. Sexual maturity is attained by about 17 cm fork length. The diet consists
primarily of invertebrates, shrimps, copepods, and larvae of decapod crustaceans; at larger sizes also
feeds on small fishes. Caught mainly on hook-and-line.
Distribution: Occurs through-
out the western Indian Ocean,
but appears to have a re-
stricted distribution in the
western Pacific Ocean where
it is known only from Thailand,
Sumatra, and the Philippines.
Also occurs in the eastern part
of the Mediterranean Sea
(coasts of Israel, Lebanon,
and Egypt); a recent immi-
grant from the Red Sea via the
Suez Canal.
2688 Bony Fishes
“Alepes” kleinii (Bloch, 1793)

Frequent synonyms / misidentifications: Caranx para Cuvier, 1833; C. microchir Cuvier, 1833;
C. miyakamii Wakiya, 1924 / Caranx kalla Cuvier, 1833 (this name has generally been misapplied to
“Alepes” kleinii); Alepes kalla (Cuvier, 1833).
FAO names: En - Banded scad; Fr - Comère rubari; Sp - Macarela bandoneada.
12 cm fork length
Diagnostic characters: Body oval, strongly compressed, with ventral profile distinctly more convex than
dorsal profile; snout pointed. Eye diameter slightly larger than snout length, with adipose eyelid well
developed on posterior half of eye only. Upper jaw broad and slightly concave posteriorly, extending to
anterior margin of pupil; upper jaw anteriorly with 2 irregular rows of short conical teeth, posteriorly
inner surface of jaw paved with blunt teeth; lower jaw with a single row of short, conical teeth except
2 rows anteriorly. Gill rakers (including rudiments) 10 to 12 on upper limb and 27 to 32 on lower limb of
first gill arch (total 38 to 44). Shoulder girdle (cleithrum) margin smooth, without papillae. Two separate
dorsal fins, the first with VIII spines, the second with I spine and 23 to 26 soft rays; anal fin with II detached
spines followed by I spine and 19 to 22 soft rays; spinous dorsal fin moderately high, longest spine height
about equal to length of soft dorsal-fin lobe; upper lobe of caudal fin longer than lower lobe in large adults.
Lateral line strongly arched anteriorly, with junction of curved and straight parts below vertical from fourth
to sixth soft ray of second dorsal fin; chord of curved part of lateral line contained 1.5 to 2.2 times in straight
caudal fin) 72 to 86. Vertebrae 10+14. Colour: in life, bluish grey to green above, silvery below; large
black spot on upper margin of opercle and adjacent area of shoulder; dark bands sometimes evident
on sides above lateral line; caudal fin dusky to bright yellow, upper lobe with dark fringe; other fins mostly
pale to hyaline.
Size: A relatively small species; largest specimen examined 16 cm fork length, commonly to about 14 cm
fork length.
Habitat, biology, and fisheries: An inshore, coastal species. Feeds primarily on planktonic crustaceans
and larval fishes. Caught on hook-and-line; also with gill nets and other artisanal gear.
Distribution: In the western
Indian Ocean, this locally
abundant species is appar-
ently restricted to coastal wa-
ters from Pakistan to Sri
Lanka. Elsewhere in the Indo-
West Pacific, known from the
east coast of India, Taiwan
Province of China, Okinawa,
Japan, the Philippines, Papua
New Guinea, and Australia.
Alepes melanoptera Swainson, 1839

Frequent synonyms / misidentifications: Caranx malam (BIeeker, 1851); Alepes malam (Bleeker,
1851); Caranx nigripinnis Day, 1876; C. pectoralis Chan, 1968 / None.
FAO names: En - Blackfin scad; Fr - Sélar aile noire; Sp - Jurel alinegro.
16.7 cm fork length
snout bluntly rounded. Eye diameter about equal to snout length, with adipose eyelid well developed on
posterior half of eye only. Upper jaw narrowly rounded posteriorly, and supramaxilla relatively small
without an anterior spine-like extension; both jaws with a single row of numerous, comb-like teeth. Gill
rakers (including rudiments) 7 to 9 on upper limb and 17 to 24 on lower limb of first gill arch (total 24 to
30). Shoulder girdle (cleithrum) margin smooth, without papillae. Two separate dorsal fins, the first with
VII spines, the second with I spine and 23 to 26 soft rays; anal fin with II detached spines followed by I
of soft dorsal-fin lobe; ultimate and penultimate rays of dorsal and anal fins of equal length. Lateral line
strongly arched anteriorly, with junction of curved and straight parts below vertical from between second
dorsal-fin origin and third soft ray; chord of curved part of lateral line contained 2.3 to 2.7 times in straight
caudal fin) 95 to 114. Vertebrae 10+14. Colour: in life, grey-blue above, silvery to white below; a diffuse
dusky blotch on margin of opercle, not bordered above by a white spot; interradial membranes of spinous
dorsal fin black; second dorsal fin greyish, lobe pale distally; caudal fin dusky yellow, with darker trailing
edges, other fins pale.
Size: A small species, largest specimen examined 21 cm fork length.
Habitat, biology, and fisheries: An inshore species. Feeds primarily on small invertebrates, shrimps,
and copepods. Caught mainly on hook-and-line.
Indian Ocean, known only
from the Persian Gulf east-
ward to Sri Lanka; elsewhere
in the In do -We s t Pacific
land southern China, Indone-
sia, and the Philippines.
2690 Bony Fishes
Alepes vari (Cuvier, 1833)

Frequent synonyms / misidentifications: Caranx macrurus (Bleeker, 1852); Alepes macrurus (Bleeker,
1852); A. glabra (Fowler, 1904) / None.
FAO names: En - Herring scad; Fr - Sélar harengule; Sp - Jurel arenque.
19 cm fork length
snout pointed. Eye diameter about equal to snout length, with adipose eyelid well developed on posterior
half of eye only. Upper jaw nearly straight posteriorly, and supramaxilla relatively large with an anterior
spine-like extension; both jaws with a single row of numerous, comb-like teeth. Gill rakers (including
rudiments) 9 to 12 on upper limb and 23 to 26 on lower limb of first gill arch (total 32 to 38). Shoulder girdle
(cleithrum) margin smooth, without papillae. Two separate dorsal fins, the first with VIII spines, the second
with I spine and 24 to 27 soft rays; anal fin with II detached spines followed by I spine and 20 to 23 soft
rays; spinous dorsal fin moderately high, longest spine height about equal to length of soft dorsal-fin lobe;
ultimate and penultimate rays of dorsal and anal fins of about equal length. Lateral line strongly arched
anteriorly, with junction of curved and straight parts below vertical from between origin of second dorsal
fin and third soft ray; chord of curved part of lateral line contained 1.9 to 2.5 times in straight part (to
caudal-fin base); scales in curved part of lateral line 42 to 50; 0 to 2 scutes in curved part; straight
part with 0 to 7 scales and 48 to 69 scutes; total scales and scutes in lateral line (excluding scales
on caudal fin) 86 to 119. Vertebrae 10+14. Colour: in life, ash blue above, silvery to white below; a diffuse
dusky blotch on margin of opercle, not bordered above by a white spot; interradial membranes of spinous
dorsal fin pale to dark dusky; second dorsal fin, anal, and caudal fins dusky; mature males develop black
pigmentation in spinous dorsal fin, lobes of soft dorsal and anal fins, and in pelvlc fins that is lacking in
females.
Size: A relatively large species, commonly attaining 30 cm fork length; largest specimen examined 46 cm
fork length and 56 cm total length.
Habitat, biology, and fisheries: Common in shallow coastal waters where it oftens swims near the
surface. Feeds primarily on shrimps, copepods, decapod crustaceans, and small fishes. Caught mainly
on hook-and-line.
Indian Ocean apparently con-
fined to coastal waters from
the Red Sea to Sri Lanka;
elsewhere in the Indo-West
Pacific known from the east
coast of India, Bay of Bengal,
Gulf of Thailand, Taiwan
Province of China, Okinawa,
Indonesia, northern Australia,
and New Guinea.
Atropus atropos (Schneider, 1801)

Frequent synonyms / misidentifications: Atropus atropus (misspelling) / Carangoides armatus
(Rüppell, 1830); C. hedlandensis (Whitley, 1933).
FAO names: En - Cleftbelly trevally; Fr - Pompano à quille noire; Sp - Pámpano de quilla.
, 21 cm fork length
Diagnostic characters: Body strongly compressed, almost ovate; nape strongly convex; belly with a
deep median groove, accommodating pelvic fins, anus, and anal-fin spines. Eye diameter slightly
larger than snout length. Upper jaw with a narrow band of small teeth, the band widest anteriorly; lower
jaw with a single series of small teeth, except 2 or 3 irregular rows anteriorly. Gill rakers (including
rudiments) 8 to 11 on upper limb and 19 to 22 on lower limb of first gill arch (total 29 to 34). Two separate
dorsal fins, the first with VIII spines, the second with I spine and 19 to 22 soft rays; anal fin with II detached
spines followed by I spine and 17 or 18 soft rays; lobe of second dorsal fin shorter than head length; dorsal
fin sexually dimorphic, in mature males (only) 6 to 12 of the central rays produced into filaments
of greatly varying lengths; pelvic fins conspicuously long, tip of appressed fin extending almost to
anal-fin origin. Lateral line anteriorly with a moderate regular arch, with junction of curved and straight
parts below vertical from sixth or seventh soft rays of second dorsal fin; chord of curved part of lateral line
contained 1.5 to 2 times in straight part; straight part of lateral line with 31 to 37 scutes. Breast naked
ventrally to behind pelvic fins; laterally, naked area of breast extends diagonally to naked base of pectoral
fins. Vertebrae 10+14. Colour: in life, head and body bluish green above, silvery below; membranes of
pelvic fins black, with the rays white basally; other fins pale; young with indistinct dark bands and
opercular spot, the latter prominent in adults.
Size: Largest specimen examined 23 cm fork length and 26.5 cm total length; probably does not exceed
30 cm total length.
Habitat, biology, and fisheries: Common, in shallow coastal waters where it oftens swims near the
surface. Feeds primarily on
shrimps, copepods, decapod
crustaceans, and small
fishes. Caught mainly with
bottom trawls, traps, and on
hook-and-line.
West Pacific in coastal waters
of the Arabian Gulf to Sri
Lanka, east coast of India,
Malaysia, Gulf of Thailand,
Okinawa, and the Philippines.
2692 Bony Fishes
Atule mate (Cuvier, 1833)

Frequent synonyms / misidentifications: Caranx mate Cuvier, 1833; C. affinis Rüppell, 1836;
Decapterus politus (Jenkins, 1904); D. lundini Jordan and Seale, 1906; D. normani Bertin and Dollfus,
1948 / None.
FAO names: En - Yellowtail scad; Fr - Sélar queue jaune; Sp - Jurel rabo amarillo.
28.5 cm fork length
Diagnostic characters: Body elongate oval, moderately compressed, with dorsal and ventral profiles
almost evenly convex; snout pointed. Eye moderate, diameter shorter than snout in adults, with adipose
eyelid well developed and completely covering eye except for a vertical slit centred on pupil. Upper
jaw extending to below anterior third of eye; both jaws with a single series of small conspicuous teeth,
except in large specimens upper jaw with 2 or 3 rows of small canines anteriorly. Gill rakers (including
rudiments) 10 to 13 on upper limb and 26 to 31 on lower limb of first gill arch (total 37 to 44). Shoulder
girdle (cleithrum) margin smooth, without papillae. Two separate dorsal fins, the first with VIII spines,
the second with I spine and 22 to 25 soft rays; anal fin with II detached spines followed by I spine and 18
to 21 soft rays; spinous dorsal fin moderately high, longest spine height about equal to length of soft
dorsal-fin lobe; terminal dorsal and anal rays finlet-like in adults, about twice length of adjacent rays
and a little more separated but joined by interradial membrane; pectoral fins long and falcate, tip of
appressed fin extending to junction of curved and straight parts of lateral line. Lateral line gently arched
anteriorly, with junction of curved and straight parts below vertical from sixth to eighth soft rays
of second dorsal fin; chord of curved part of lateral line contained 1 to 1.4 times in straight part (to
caudal-fin base), scales in curved part of lateral line 39 to 57; straight part with 0 to 10 scales and
36 to 49 scutes; total scales and scutes in lateral line (excluding scales on caudal fin) 92 to 103. Vertebrae
10+14. Colour: in life, bright olive-green dorsally, yellowish green laterally and whitish ventrally; dorso-
laterally 9 or 10 faint, grey bars, wider than pale interspaces; a black spot, slightly smaller than eye,
on upper margin of opercle and adjacent area of shoulder; dorsal and caudal fins dusky greenish
yellow; anal fin pale yellow; pectoral and pelvic fins hyaline to whitish.
Size: Largest specimen examined 20 cm fork length and 30 cm total length; commonly to about 26 cm
fork length.
Habitat, biology, and fisheries: Occurs in schools at depths to about 50 m in inshore waters throughout
its range; feeds primarily on
crustaceans and planktonic
invertebrates. Caught mainly
on hook-and-line, and with
various types of artisanal
gear.
Distribution: Broadly distrib-
uted throughout the Indian
Ocean; elsewhere in the Indo-
West Pacific from southern
Japan and Australia eastward
to Hawaii.
Carangoides armatus (Rüppell, 1830)

Frequent synonyms / misidentifications: Caranx armatus (Rüppell, 1830); Carangoides ciliarius of
authors; Caranx schlegeli Wakiya, 1924 / Carangoides hedlandensis (Whitley, 1933).
FAO names: En - Longfin trevally; Fr - Carangue sap-sap; Sp - Jurel sap-sap.
, 22 cm fork length
Diagnostic characters: Body strongly compressed and deep. Head profile very steep in adults but
relatively straight from snout to nape. Eye diameter about equal to, or smaller than, snout length. Both
jaws with bands of villiform teeth, the bands widest anteriorly; vomerine tooth patch wedge-shaped without
a posteromedian, extension. Gill rakers (including rudiments) 10 to 15 on upper limb and 20 to 24 on
lower limb of first gill arch (total 31 to 37). Two separate dorsal fins, the first with VIII spines, the second
with I spine and 19 to 22 soft rays; anal fin, with II detached spines followed by I spine and 16 to 18 soft
rays; lobe of second dorsal fin elongate and filamentous, longer than head length; dorsal fin sexually
dimorphic, in mature males (only) larger than 21 cm fork length, 3 to 12 of the central rays produced
into filaments of greatly varying lengths. Lateral line anteriorly with a moderate regular arch, with
junction of curved and straight parts below vertical from tenth to twelfth soft rays of second dorsal fin; chord
of curved part of lateral line longer than straight part of lateral line, contained 0.7 to 0.9 times in straight
part; straight part of lateral line with 11 to 24 weak scutes and 25 to 43 total elements (including anterior
scales). Breast naked ventrally to behind origin of pelvic fins; laterally, naked area of breast extends
diagonally to naked base of pectoral fins. Vertebrae 10+14. Colour: bluish grey above, silvery below;
blackish blotch on upper margin of opercle; spinous dorsal fin blackish; second dorsal and anal fins pale
to dusky, leading edge of lobes dusky to blackish; caudal fin grey with leading and trailing edges black;
pelvic fins generally black in specimens smaller than 10 cm fork length, but becoming paler with increasing
size until at 20 cm pale or only slightly dusky.
Size: Largest specimen examined 50 cm fork length.
Habitat, biology, and fisheries: Inhabits rocky and coral coastlines as well as shallow bays; small groups
of adults often swim along the edges of reefs. Juveniles may occur in estuarine areas. Sexual maturity is
attained at 21 or 22 cm fork length. Caught mainly on hook-and-line; also by spears.
Distribution: Western Indian
Ocean from South Africa
northward to southern India
and Sri Lanka, including the
Red Sea, Gulf of Oman, and
west coast of Malagasy Re-
public; in the western Pacific
land, Hong Kong, Thailand,
Okinawa, and Japan.
2694 Bony Fishes
Carangoides bajad (Forsskål, 1775)

Frequent synonyms / misidentifications: Caranx auroguttatus Cuvier, 1833; Carangoides auroguttatus
(Cuvier, 1833) / Carangoides gymnostethus (Cuvier, 1833).
FAO names: En - Orangespotted trevally; Fr - Carangue lentigine; Sp - Jurel lentejuela.
38 cm fork length
Diagnostic characters: Body oblong and compressed; dorsal profile more convex than ventral profile;
dorsal profile of head slightly elevated to nape, almost straight. Eye diameter smaller than snout length.
Both jaws with narrow bands of villiform teeth, the bands widest anteriorly; vomerine tooth patch triangular,
without a posteromedian extension. Gill rakers (including rudiments) 7 to 9 on upper limb and 18 to 21 on
rays, lobe of second dorsal fin shorter than head length. Lateral line anteriorly with a relatively low
arch, with junction of curved and straight parts below vertical from eleventh to fifteenth soft rays of second
dorsal fin; chord of curved part of lateral line longer than straight part of lateral line, contained 0.7 to 0.9
times in straight part; straight part of lateral line with 14 to 26 scales followed by 20 to 30 scutes. Breast
completely scaly or with a narrow naked area anteroventrally, scarcely if at all visible in lateral view.
Vertebrae 10+14. Colour: brassy dorsally, shading to silvery white on sides, with numerous conspicu-
ous orange-yellow spots; no dark opercular spot; capable of rapidly changing colour to almost entirely
orange.
Size: Largest specimen examined 49 cm fork length and 55 cm total length.
Habitat, biology, and fisheries: Caught on hook-and-line, with gill nets, and other artisanal gear.
Distribution: In the Indian
Ocean known from the Red
Sea, Gulf of Aden, Persian
Gulf, and the Gulf of Oman;
Pacific known from the Gulf of
Thailand, Okinawa, Indone-
sia, Philippines, and New Brit-
ain.
Carangoides caeruleopinnatus (Rüppell, 1830)

Frequent synonyms / misidentifications: Caranx caeruleopinnatus Rüppell, 1830; Carangoides
altissimus Jordan and Seale, 1907; C. formosanus Jordan and Snyder, 1910; C. uii Wakiya, 1924; Citula
diversa Whitley, 1940 / None.
FAO names: En - Coastal trevally; Fr - Carangue côtière; Sp - Jurel costero.
26.7 cm fork length
Diagnostic characters: Body strongly compressed, almost ovate; dorsal profile of body more strongly
convex than ventral profile, nape moderately curved. Eye diameter slightly smaller than snout length. Both
jaws with bands of villiform teeth, the bands widest anteriorly; vomerine tooth patch ovate, without a
posteromedian extension. Gill rakers (including rudiments) 5 to 8 on upper limb and 15 to 19 on lower
limb of first gill arch (total 21 to 27). Two separate dorsal fins, the first with VII spines, the second
with I spine and 20 to 23 (usually 22 or 23) soft rays; anal fin with II detached spines followed by I spine
and 16 to 20 soft rays; lobe of second dorsal fin filamentous in young, becoming shorter with age,
in mature adults distinctly shorter than head length (in specimens larger than 25 cm fork length, height
of second dorsal-fin lobe usually shorter than head length, and slightly to distinctly shorter than anal-fin
lobe). Lateral line anteriorly with a moderately regular arch, with junction of curved and straight parts below
vertical from between twelfth to fourteenth soft rays of second dorsal fin; chord of curved part of lateral line
longer than straight part of lateral line, contained 0.55 to 0.8 times in straight part; straight part of lateral
line with 16 to 20 scutes, and 31 to 50 total elements (including anterior scales). Breast naked ventrally
to distinctly behind origin of pelvic fins; laterally, naked area of breast typically extends diagonally
to naked base of pectoral fins (rarely, naked areas of breast and pectoral-fin base interrupted
laterally by a narrow band of scales). Colour: in life, bluish green above, silvery grey below; sides with
numerous, small yellow spots; small black blotch on upper margin of opercle; second dorsal, anal, and
caudal fins dusky, the latter sometimes yellowish; pectoral fins pale yellow; pelvic fins hyaline to pale grey.
Size: Largest specimen examined 36 cm fork length; commonly to 25 cm fork length.
Habitat, biology, and fisheries: Adults are commonly found over deeper coastal reefs, but are rarely
close to the shore. A rather sluggish fish, as compared to other Carangoides species. Taken incidental to
artisanal fisheries throughout
its range. Caught on hook-
and-line, gill nets, and traps.
Distribution: C o m m o n i n
coastal waters throughout the
western Indian Ocean; else-
where in the Indo-West Pa-
cific broadly distributed
throughout the eastern Indian
and western Pacific oceans
from Japan to Australia.
2696 Bony Fishes
Carangoides chrysophrys (Cuvier, 1833)

Frequent synonyms / misidentifications: C a ranx chrysophrys C uv ier, 1833; Carangoides
chrysophryoides Bleeker, 1851; Caranx typus Gilchrist and Thompson, 1917 / None.
FAO names: En - Longnose trevally; Fr - Carangue tapir; Sp - Jurel tapir.
27 cm fork length
Diagnostic characters: Body subovate in juveniles, oblong in adults and compressed; nape only slightly
elevated, dorsal profile of snout gently sloped, then abruptly vertical just above mouth cleft. Eye
diameter smaller than snout length. Both jaws with bands of small villiform teeth, the bands widest
anteriorly and in large specimens some outer teeth conical; vomerine tooth patch ovate, without a
limb of first gill arch (total 21 to 26). Two separate dorsal fins, the first with VIII spines, the second with
I spine and 18 to 20 soft rays; anal fin with II detached spines followed by I spine and 15 to 17 soft rays;
lobe of second dorsal fin falcate but usually shorter than head length, height of fin lobe becoming shorter
with age. Lateral line anteriorly with a moderate regular arch, with junction of curved and straight parts
below vertical from between twelfth to fourteenth soft rays of second dorsal fin; chord of curved part of
lateral line longer than straight part of lateral line, contained 0.5 to 0.6 times in straight part; straight part
of lateral line with 20 to 37 weak scutes and 32 to 42 total elements (including anterior scales). Breast
naked posteroventrally to distinctly behind pelvic fins; laterally, naked area of breast extends
diagonally to naked base of pectoral fins. Vertebrae 10+14. Colour: in life, generally silvery with head
and body greenish above and silvery with yellow-green reflections below; opercle with a small black spot
on upper margin; dorsal and anal fins ranging in colour from whitish to pale yellow to dusky; interradial
membranes of soft anal-fin rays often with a white spot basally; caudal and pectoral fins pale to dusky
yellow; large adults sometimes very dark, head and fins blackish; these fish perhaps exhibiting nuptial or
spawning coloration.
Size: Largest specimen examined 38.5 cm fork length and 44 cm total length; reported to attain 60 cm
total length.
Habitat, biology, and fisheries: Inhabitats coastal waters in depths to 90 m, but most abundant at 30 to
60 m. Feeds on small demersal fishes and epibenthic crustaceans. Caught on hook-and-line, bottom
trawls, gill nets, and traps.
coastal waters of the Indian
Ocean, including the Red Sea
and Persian Gulf; elsewhere
in the Indo-West Pacific from
Okinawa to Australia and
eastward to New Britain.
Carangoides dinema Bleeker, 1851

Frequent synonyms / misidentifications: Caranx dinema (Bleeker, 1851); Carangichthys dinema
(Bleeker, 1851); Caranx deani Jordan and Snyder, 1907 / None.
FAO names: En - Shadow trevally; Fr - Carangue crépuscule; Sp - Jurel crepúsculo.
43 cm fork length
Diagnostic characters: Body strongly compressed, almost ovate; nape slightly elevated, almost straight
in profile. Eye diameter slightly smaller than snout length. Both jaws with bands of small teeth, the bands
widest anteriorly; upper jaw also with an irregular outer series of moderately large teeth (in large specimens
outer row teeth may be enlarged in both jaws); vomerine tooth patch ovate, without a posteromedian
extension. Gill rakers (including rudiments) 7 to 9 on upper limb and 16 to 19 on lower limb of first gill arch
(total 24 to 28). Two separate dorsal fins, the first with VIII spines, the second with I spine and 17 to
19 soft rays; anal fin with II detached spines followed by I spine and 15 to 17 soft rays; lobe of second
dorsal fin elongate, longer than head length, in adults about equal or exceeding the length of second
dorsal-fin base. Lateral line anteriorly with a moderate regular arch, with junction of curved and straight
parts below vertical from between tenth or twelfth soft rays of second dorsal fin; chord of curved part of
lateral line slightly longer than straight part of lateral line, contained 0.7 to 0.95 times in straight
part; straight part of lateral line with 0 to 6 scales followed by 23 to 30 scutes. Breast naked ventrally
only to origin of pelvic fins; laterally, naked area of breast usually separated from naked base of
pectoral fins by a moderate to narrow band of scales (rarely, naked areas of breast and pectoral-fin
base united laterally by a narrow naked strip). Vertebrae 10+14. Colour: in life, bluish green above, silvery
white below; small brown blotches (becoming larger posteriorly) on back between bases of second
dorsal-fin rays; opercular blotch brownish and diffuse; spinous dorsal fin pale to dusky; second dorsal-fin
lobe dusky and ray tips yellowish; anal fin with distal margin whitish blue; caudal fin with upper lobe
yellowish, and with trailing edges and tip of lower lobe pale; pectoral fins hyaline and pelvic fins whitish to
dusky.
Size: Largest specimen examined 53 cm fork length, 58.5 cm total length, and 2.6 kg.
Habitat, biology, and fisheries: Inhabitats coastal waters. Caught on hook-and-line, with bottom trawls
and various kinds of artisanal
gear.
Ocean known only from
South Africa to Tanzania;
Pacific known from southern
Japan, Okinawa, Indonesia, ?
P h i l i p p i n e s , P a p u a N ew
Guinea, Tonga Islands, and
Samoa.
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2698 Bony Fishes
Carangoides equula (Temminck and Schlegel, 1844)

Frequent synonyms / misidentifications: Caranx equula Temminck and Schlegel, 1844; Kaiwarinus
equula (Temminck and Schlegel, 1844); Carangoides acutus Kotthaus, 1974; ? Caranx dasson Jordan
and Snyder, 1907 (see Remarks) / None.
FAO names: En - Whitefin trevally; Fr - Carangue aile blanche; Sp - Jurel aliblanco.
22 cm fork length
Diagnostic characters: Body compressed, almost rhomboidal; dorsal and ventral profiles about equally
convex, dorsal profile of snout and nape almost straight. Eye diameter slightly smaller than snout length.
Both jaws with narrow bands of small teeth, the outer teeth slightly larger; vomerine tooth patch
anchor-shaped, with a distinct posteromedian extension. Gill rakers (including rudiments) 7 to 10
on upper limb and 18 to 23 on lower limb of first gill arch (total 27 to 32). Two separate dorsal fins, the
first with VII spines, the second with I spine and 23 to 25 soft rays; anal fin with II detached spines followed
by I spine and 21 to 24 soft rays; spinous dorsal fin moderately high, longest spine height about equal
to length of soft dorsal-fin lobe. Lateral line anteriorly with a moderate regular arch, with junction of
curved and straight parts below vertical from between twelfth and fifteenth soft rays of second dorsal fin;
chord of curved part of lateral line longer than straight part of lateral line, contained 0.5 to 0.8 times in
straight part; straight part of lateral line with 0 to 6 scales followed by 22 to 32 scutes. Breast completely
scaly or with a very small naked area anteroventrally. Vertebrae 10+14. Colour: in life, head and body
bluish grey to green dorsally, silvery white below; juveniles with 5 to 7 dark bands on sides; second dorsal
and anal fins with a submarginal brown to blackish band, white distally; caudal fin dusky yellow;
pectoral and pelvic fins pale yellowish to white.
Size: Largest specimen examined 34.5 cm fork length, 37.5 cm total length.
Habitat, biology, and fisheries: In northern Australian waters largely restricted to shelf slope habitats in
depths of 100 to 200 m. Feeds on a variety of fishes, crustaceans, and cephalopods. Taken primarily with
bottom trawls.
Ocean known only from South
Africa, off Somalia, and the ?
Gulf of Oman; elsewhere in
the Indo-West Pacific defi-
nitely known from Okinawa,
southern Japan, and Austra-
lia. If Carangoides dasson is
conspecific (see Remarks),
the species also occurs at Ha-
waii and Easter Island.
Remarks: Specimens from
Hawaii and Easter Island (for
which the name C. dasson is
available) have more slender bodies and larger eyes than do specimens from other localities but share all
the diagnostic features of C. equula. Additional study is required to determine if C. equula and C. dasson
represent 2 closely related allopatric species.
Carangoides ferdau (Forsskål, 1775)

Frequent synonyms / misidentifications: C a r a n x ferdau ( F o r s s k å l , 1 7 7 5 ) ; Carangoides
hemigymnostethus Bleeker, 1851; Caranx gilberti Jordan and Seale, 1906 / None.
FAO names: En - Blue trevally; Fr - Carangue tachetée; Sp - Jurel manchado.
30 cm fork length
snout bluntly rounded. Upper jaw highly protractile; lips of large adults not noticeably fleshy, both
jaws with narrow bands of villiform teeth, becoming obsolescent with age; vomerine tooth patch ovate,
without a posteromedian extension. Gill rakers (including rudiments) 7 to 10 on upper limb and 17 to 20
on lower limb of first gill arch (total 24 to 29). Two separate dorsal fins, the first with VIII spines, the
second with I spine and 26 to 34 soft rays; anal fin with II detached spines followed by I spine and 21
to 26 soft rays; lobe of second dorsal fin falcate, especially in small adults, but usually shorter than head
length. Lateral line anteriorly with a very slight arch, with junction of curved and straight parts below vertical
from between fifteenth to twentieth soft rays of second dorsal fin; chord of curved part of lateral line longer
than straight part of lateral line, contained 0.6 to 0.85 times in straight part; straight part of lateral line with
10 to 30 scales followed by 21 to 37 small scutes. Breast naked ventrally to origin of pelvic fins; laterally,
naked area of breast separated from naked base of pectoral fins by a moderate band of scales. Vertebrae
10+14. Colour: adults typically with 5 or 6 dusky bands on sides that usually persist in market
specimens; in life, head and body generally silvery, blue-green above paler below; numerous, incon-
spicuous golden spots often present on sides mainly above level of pectoral fins; leading edge and
tips of dorsal- and anal-fin lobes dark blue to dusky, rest of fin pale yellow-green except distal margin of
anal fin whitish. Caudal fin yellow-green with trailing edge and lobe tips dark; pelvic fins hyaline to whitish;
caudal fin yellow-green with trailing edge and lobe tips dark; pelvic fins hyaline to whitish.
Size: Largest specimen examined 47 cm fork length and 53 cm total length; reported to attain 70 cm total
length. South African spearfishing record 8 kg.
Habitat, biology, and fisheries: Reported to prefer coastal waters adjacent to sandy beaches but also
found to depths of 60 m, often near reefs. Feeds primarily on prawns, crabs, and small fishes. Caught
mainly on hook-and-line; also
with gill nets and traps.
Distribution: Broadly dis-
tributed throughout the Indian
Ocean, and western and Cen-
tral Pacific ocean from south-
ern Japan to Australia and
eastward to Hawaii.
2700 Bony Fishes
Carangoides fulvoguttatus (Forsskål, 1775)

Frequent synonyms / misidentifications: Caranx fulvoguttatus (Forsskål, 1775); Turrum emburyi
Whitley, 1932 / Carangoides gymnostethus (Cuvier, 1833).
FAO names: En - Yellowspotted trevally; Fr - Carangue pailleteé; Sp - Jurel centellante.
74 cm fork length
Diagnostic characters: Body subovate and compressed, becoming elongate-ovate and slightly subcylin-
drical with age; profile of head and nape slightly angular becoming more steep with age. In adults
mouth cleft distinctly below level of eye. Eye diameter smaller than snout length. Both jaws with bands
of villiform teeth, the bands widest anteriorly; vomerine tooth patch ovate, without a posteromedian
extension. Gill rakers (including rudiments) 6 to 8 on upper limb and 17 to 21 on lower limb of first gill
arch (total 22 to 27). Two separate dorsal fins, the first with VIII spines, the second with I spine and 25 to
30 soft rays; anal fin with II detached spines followed by I spine and 21 to 26 (rarely 25 or 26 soft rays);
lobe of second dorsal fin shorter than head length. Lateral line anteriorly with a low regular arch, with
junction of curved and straight parts below vertical from between thirteenth to sixteenth soft rays of second
times in straight part; straight part of lateral line with 18 to 27 scales followed by 15 to 21 small scutes.
Breast naked ventrally to distinctly behind origin of pelvic fins; breast squamation variable laterally,
either separated from naked base of pectoral fins by a moderate to very narrow band of scales or
naked area of breast uninterrupted to naked base of pectoral fins (only 23 of 47 specimens examined
for this character had the uninterrupted pattern of breast squamation). Vertebrae 10+14. Colour: adults
iridescent blue-green dorsally, shading to silvery on sides, with many small golden or brassy spots
mainly above midline; large individuals often with 3 irregular black blotches (sometimes very
indistinct or absent) on flanks, the first below dorsal-fin lobe, the second at inflection point of lateral
line, and the third slightly anterior to midpoint of straight part of lateral line; opercular spot dusky
and inconspicuous; dorsal and anal fins dusky yellow, the latter with leading edge and distal margin whitish
blue; caudal fin olive-yellow with upper leading edge and trailing edges dusky; pelvic fins whitish blue.
Size: Largest specimen examined 80 cm fork length, 87.5 cm total length, and 6.4 kg. South African
spearfishing record 10 kg. Reliably reported to attain 100 cm fork length.
Habitat, biology, and fisheries: Prefers rocky and coral reefs, but is occasionally found over offshore
banks in depths to 100 m. Feeds mainly on small invertebrates and fishes. Caught mainly on hook-and-line
and by spearing. Also with
general types of artisanal
gear, including gill nets and
traps. A good sportsfish.
uted through the Indian and
western Pacific oceans, in-
cluding Okinawa, Palau, and
Australia.
Carangoides gymnostethus (Cuvier, 1833)

Frequent synonyms / misidentifications: Caranx gymnostethus (Cuvier, 1833); Carangoides
gymnostethoides Bleeker, 1851 / Carangoides fulvoguttatus (Forsskål, 1775).
FAO names: En - Bludger; Fr - Carangue balo; Sp - Jurel balo.
52.4 cm fork length
Diagnostic characters: Body ovate and compressed, becoming elongate-ovate and slightly subcylindrical
with age; profile of head and nape gently convex becoming less steep with age. In adults mouth
cleft at level with lower margin of eye. Diameter of eye smaller than snout length. Both jaws with bands
of villiform teeth, the bands widest anteriorly; vomerine tooth patch ovate without a posteromedian
extension. Gill rakers (including rudiments) 7 to 9 on upper limb and 19 to 22 on lower limb of first gill
arch (total 27 to 31). Two separate dorsal fins, the first with VIII spines, the second with I spine and 28 to
32 soft rays; anal fin with II detached spines followed by I spine and 24 to 26 (usually 25) soft rays; lobe
of second dorsal fin shorter than head length. Lateral line anteriorly with a low regular arch, with junction
of curved and straight parts below vertical from between sixteenth to twentieth soft rays of second dorsal
fin; chord of curved part of lateral line longer than straight part of lateral line, contained 0.65 to 0.9 times
in straight part; straight part of lateral line with 14 to 25 scales followed by 20 to 31 small scutes. Breast
naked ventrally to distinctly behind origin of pelvic fins; laterally, naked area of breast extends diagonally
to naked base of pectoral fins, the naked area somewhat constricted below pectoral-fin base.
Vertebrae 10+15. Colour: in life, adults olive-green above, silvery white below with a few brown or golden
spots sometimes present midlaterally; opercular spot dusky and inconspicuous; dorsal, anal, and caudal
fins pale olive-green to greenish grey, leading edge and distal margin of anal fin white; pelvic and pectoral
fins pale green to hyaline.
Size: Largest specimen examined 76.5 cm fork length, 83 cm total length, and 7.2 kg. South African
spearfishing record 14.5 kg. Reported to attain at least 90 cm total length.
Habitat, biology, and fisheries: Most common over slightly deeper offshore reefs. Larger individuals tend
to be solitary, while juveniles form small schools. Feeds mainly on small prey, including shrimps. Caught
mainly on hook-and-line and with several types of artisanal gear, including gill nets and traps. A good
sportsfish.
and western Pacific oceans,
including the Ryukyu Islands,
Australia, and Kapingama-
rangi Atoll.
2702 Bony Fishes
Carangoides hedlandensis (Whitley, 1933)

Frequent synonyms / misidentifications: Caranx hedlandensis (Whitley, 1933); C. plumbeus of authors /
Carangoides armatus (Rüppell, 1830).
FAO names: En - Bumpnose trevally; Fr - Carangue nez bossu; Sp - Jurel hocicón.
, 20 cm fork length
Diagnostic characters: Body strongly compressed and deep; head profile extremely steep in adults,
and with a distinct break in contour “bump” in the interorbital region which becomes more
pronounced with increasing size. Eye diameter about equal to or larger than snout length. Both jaws
with bands of villiform teeth, the bands widest anteriorly; vomerine tooth patch wedge-shaped without a
posteromedian extension. Gill rakers (including rudiments) 6 to 11 on upper limb, and 14 to 17 on lower
limb of first gill arch (total 20 to 27). Two separate dorsal fins, the first with VIII spines, the second with I
spine and 20 to 22 soft rays; anal fin with II detached spines followed by I spine and 16 to 18 soft rays;
lobe of second dorsal fin elongate and filamentous, longer than head length; dorsal fin sexually
dimorphic, in mature males (only) larger than 16.5 cm fork length, 3 to 8 of the central rays produced
into filaments of greatly varying lengths. Lateral line anteriorly with a moderate regular arch, with
junction of curved and straight parts below vertical from between tenth and twelfth soft rays of second
times in straight part; straight part of lateral line with 17 to 29 weak scutes and 29 to 41 total elements
(including anterior scales). Breast naked ventrally to behind origin of pelvic fins; laterally, naked area
of breast extends diagonally to naked base of pectoral fins. Vertebrae 10+14. Colour: greenish blue
above with dusky tinge, shading to silvery grey below; blackish blotch on upper margin of opercle; spinous
dorsal fin, elongated dorsal-fin rays and edges of caudal fin blackish; filamentous lobe of anal fin either
blackish or pale brownish, elongated rays always pale brownish; pelvic fins generally black in specimens
smaller than 10 cm fork length, at larger than 15 cm fork length pelvic fins variable in colour from pale to
dusky blackish.
Habitat, biology, and fisheries: A coastal demersal species. Caught on hook-and-line, with bottom trawls
and several types of artisanal
gear.
Indian Ocean, known from
Durban (South Africa), the
Seychelles, southern India
and Sri Lanka; elsewhere reli-
ably reported from the Bay of
Bengal, Gulf of Thailand, Tai-
wan Province of China, Oki-
nawa, Japan, the Philippines,
Indonesia, Australia, New
Guinea, and Samoa.
Carangoides humerosus (McCulloch, 1915)

Frequent synonyms / misidentifications: Caranx humerosus McCulloch, 1915 / None.
FAO names: En - Epaulet trevally.
12.7 cm fork length

Diagnostic characters: Body oblong to elongate oval, compressed; dorsal profile more convex than
ventral profile; head profile gently sloping to convex, in large males profile with a distinct bulge that is
absent in females and juveniles of both sexes. Eye diameter larger than snout length. Both jaws with bands
of small teeth, the bands widest anteriorly; upper jaw also with an irregular outer series of moderately large
teeth (in large specimens outer row teeth may be enlarged in both jaws); vomerine tooth patch triangular,
second with I spine and 20 to 22 (rarely 19) soft rays; anal fin with II detached spines followed by I spine
and 17 to 19 soft rays; lobe of second dorsal fin falcate, slightly longer than to slightly shorter than head
length, in adults shorter than the length of second dorsal-fin base. Lateral line anteriorly with a moderate
regular arch, with junction of curved and straight parts below vertical from between eleventh to thirteenth
soft rays of second dorsal fin; chord of curved part of lateral line slightly longer than straight part of lateral
line, contained 0.7 to 0.9 times in straight part; straight part of lateral line with 0 to 10 scales followed by
22 to 33 scutes. Breast naked posteroventrally to well behind origin of pelvic fins; laterally, naked
area of breast extends uninterrupted to naked base of pectoral fins. Vertebrae 10+14. Colour: in
life, head and body greenish above, silvery below, small blue to black blotches on dorsum between
bases of second dorsal-fin rays; adults and juveniles often with 5 or 6 broad, dark bands separated by
narrow interspaces, and a large black spot on shoulder; axil of pectoral fins black; opercular spot
diffuse; distal parts of branchiostegal rays black in adult males, white in females and juveniles; spinous
dorsal fin black, second dorsal fin, and caudal fins dusky; pectoral and pelvic fins white to hyaline.
Size: Relatively small species, largest specimen examined 25 cm fork length.
Habitat, biology, and fisheries: In Australia in depths shallower than 50 m. Feeds on a variety of
demersal or bottom associated fishes, crustaceans, and cephalopods. Caught primarily with bottom trawls.
Distribution: K n o w n f r o m ?
A u s t r a l i a , E x m o u th G u lf
(22°05’S) to Bustard Head,
Queensland (24°08’S), south-
ern Papua New Guinea, and
reported (based on question-
able collection records) from
Taiwan Province of China.
2704 Bony Fishes
Carangoides malabaricus (Bloch and Schneider, 1801)

Frequent synonyms / misidentifications: Caranx malabaricus (Bloch and Schneider, 1801);
Carangoides rectipinnis Williams, 1958; C. rhomboides Kotthaus, 1974 / Carangoides talamparoides
Bleeker, 1852.
FAO names: En - Malabar trevally; Fr - Carangue monique; Sp - Jurel malabárico.
24.7 cm fork length
Diagnostic characters: Body strongly compressed, almost ovate; dorsal profile of head strongly elevated
to nape, almost straight. Eye diameter distinctly smaller than snout length. Both jaws with bands of small
villiform teeth, anteriorly some outer teeth conical; vomerine tooth patch roughly triangular, without a
limb of first gill arch (total 32 to 38). Two separate dorsal fins, the first with VIII spines, the second with I
spine and 20 to 23 soft rays; anal fin with II detached spines followed by I spine and 17 to 19 soft rays;
lobe of second dorsal fin shorter than head length. Lateral line anteriorly with a moderate regular arch, with
junction of curved and straight parts below vertical from between twelfth and fourteenth soft rays of second
dorsal fin; chord of curved part of lateral line longer than straight part of lateral line, contained 0.5 to 1 times
in straight part; straight part of lateral line with 19 to 36 weak scutes and 31 to 55 total elements (including
anterior scales). Breast naked ventrally to distinctly behind pelvic fins often to origin of second anal
fin; laterally, naked area of breast extends diagonally to naked base of pectoral fins, including small
area anteriorly just above pectoral-fin base. Vertebrae 10+14. Colour: in life, generally silvery with
bluish grey above, silvery white below; opercle with a small black spot on upper margin; tongue greyish
brown to brown; caudal fin, soft dorsal and anal fins pale greenish yellow to dusky; interradial membranes
of soft anal-fin rays often with a white spot basally.
Habitat, biology, and fisheries: Frequents coral and rocky reefs but may also be found in shallow sandy
bays (mainly the juveniles). Moderately tolerant of turbid waters. Feeds on crustaceans, small squids, and
fishes. Caught on hook-and-line, bottom trawls, gill nets, and traps.
Indian ocean broadly distrib-
uted in coastal waters, from
South Africa to Sri Lanka, in-
cluding the west coast of the
Malagasy Republic; else-
cific known from Straits of
Malacca, Gulf of Thailand, ?
Okinawa, Japan (rare), Indo-
nesia, and Australia.
Carangoides oblongus (Cuvier, 1833)

Frequent synonyms / misidentifications: Caranx oblongus Cuvier, 1833; Carangichthys oblongus
(Cuvier, 1833); Caranx gracilis (Ogilby, 1915); C. tanakai Wakiya, 1924 / None.
FAO names: En - Coachwhip trevally; Fr - Carangue postillon; Sp - Jurel postillón.
28 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile more convex than ventral profile; head
profile gently convex. Eye diameter smaller than snout length. Both jaws with bands of small teeth, the
bands widest anteriorly; upper jaw also with an irregular outer series of moderately large teeth (in
large specimens outer row teeth may be enlarged in both jaws); vomerine tooth patch triangular,
without a posteromedian extension. Gill rakers (including rudiments) 7 to 9 on upper limb and 17 to 21 on
with I spine and 20 to 22 soft rays; anal fin with II detached spines followed by I spine and 18 or 19 soft
rays; lobe of second dorsal fin elongate, longer than head length, in adults about equal or exceeding the
length of second dorsal-fin base. Lateral line anteriorly with a moderate regular arch, with junction of curved
and straight parts below vertical from between eighth or ninth soft rays of second dorsal fin; chord of
curved part of lateral line slightly shorter than straight part of lateral line, contained 1 to 1.3 times in
straight part; straight part of lateral line with 0 to 2 scales followed by 37 to 45 scutes. Breast naked
ventrally to origin of pelvic fins; laterally, naked area of breast separated from naked base of
pectoral fins by a moderate to narrow band of scales. Vertebrae 10+14. Colour: in life, head and body
bluish green above, silvery below, small blue to black blotches on dorsum between bases of second
dorsal-fin rays; opercular spot diffuse or absent; spinous dorsal fin pale to dusky; second dorsal-fin lobe
and upper lobe of caudal fin dusky blue; anal fin and remainder of dorsal and caudal fins yellowish; pectoral
and pelvic fins pale yellow.
Size: Largest specimen examined 41 cm fork length; 46 cm total length.
Habitat, biology, and fisheries: Caught on hook-and-line, with bottom trawls and various kinds of artisanal
gear.
uted throughout the western
Indian Ocean, but no records
from the Persian Gulf to the
west coast of India; else-
cific known from Thailand,
Japan, Indonesia, the Philip-
pines, Australia, New Guinea,
Solomon Islands, and Fiji.
2706 Bony Fishes
Carangoides orthogrammus Jordan and Gilbert, 1881

Frequent synonyms / misidentifications: Carangoides gymnostethoides evermanni Nichols, 1921;
C. jordani Nichols, 1922; C. nitidus J.L.B. Smith, 1972; Caranx ferdau jordani of authors / Carangoides
ferdau (Forsskål, 1775).
FAO names: En - Island trevally; Fr - Carangue des îles; Sp - Jurel isleño.
27 cm fork length
snout usually slightly angular. Upper jaw highly protractile, lips noticeably fleshy in large adults. Both
jaws with narrow bands of villiform teeth, becoming obsolesent with age; vomerine tooth patch ovate,
second with I spine and 28 to 31 soft rays; anal fin with II detached spines followed by I spine and 24
to 26 soft rays; lobe of second dorsal fin falcate, especially in small adults, but usually shorter than head
length. Lateral line anteriorly with a very slight arch with junction, of curved and straight parts below second
dorsal fin between fifteenth to nineteenth soft rays; chord of curved part of lateral line slightly longer to
about equal straight part of lateral line, contained 0.8 to 1 times in straight part; straight part of lateral line
with 21 to 34 scales followed by 19 to 31 small scutes. Breast naked ventrally to origin of pelvic fins,
occasionally with a small patch of prepelvic scales; laterally, naked area of breast separated from
naked base of pectoral fins by a moderate band of scales. Vertebrae 10+14. Colour: in life, head and
body generally silvery, brassy to greenish blue above, paler below; adults with several relatively large,
elliptical yellow spots, often with dusky centres, present on sides; dusky bands usually not present
on sides, especially in postmortem specimens; leading edge and tips of dorsal- and anal-fin lobes dark
to dusky, rest of fin pale brownish except distal margin of anal fin whitish; caudal fin dusky brown with
trailing edge and lobe tips darker; pelvic fins whitish.
Size: Largest specimen examined 63 cm fork length and 70 cm total length; commonly exceeds 40 cm
fork length.
Habitat, biology, and fisheries: This species has a nearly ubiquitous occurrence at oceanic islands and is
virtually absent from inshore, neritic areas. Inhabits lagoon and seaward reefs to depths exceeding 50 m.
Occurs in pairs or in schools of up to a few dozen individuals and frequents sandy channels where it feeds on
sand-dwelling crustaceans. Caught on hook-and-line, with trawls, and various kinds of artisanal gear.
Ocean known from Kenya,
Aldabra, Seychelles, Chagos
Archipelago, and Cocos-
Keeling Islands; elsewhere in
the Indo-West Pacific known
from Okinawa, Indonesia,
Australia, and eastward to
Hawaii and the Marquesas
Islands, and the Revillagigedo
Islands in the eastern Pacific
Ocean.
Carangoides plagiotaenia Bleeker, 1857

Frequent synonyms / misidentifications: Caranx plagiotaenia (Bleeker, 1857); C. vomerinus Playfair,
1867; C. compressus Day, 1870; C. brevicarinatus Klunzinger, 1871 / None.
FAO names: En - Barcheek trevally; Fr - Carangue grimée; Sp - Jurel maquillado.
30 cm fork length
Diagnostic characters: Body oblong and compressed; dorsal and ventral profiles equally convex, but in
large adults dorsal profile of head almost straight; lower jaw somewhat enlarged and projecting beyond
upper jaw. Upper jaw with a narrow band of small teeth; lower jaw with an irregular row of small teeth,
except a narrow band anteriorly; vomerine tooth patch triangular, without a posteromedian extension. Gill
rakers (including rudiments) 8 to 14 on upper limb and 19 to 27 on lower limb of first gill arch (total 27 to
40). Two separate dorsal fins, the first with VIII spines, the second with I spine and 22 to 24 soft rays; anal
fin with II detached spines followed by I spine and 18 to 20 soft rays; lobe of second dorsal fin shorter than
head length. Lateral line anteriorly with a relatively low arch, with junction of curved and straight parts below
vertical from between thirteenth and fifteenth soft rays of second dorsal fin; chord of curved part of lateral
line longer than straight part of lateral line, contained 0.5 to 0.6 times in straight part; straight part of lateral
line with 20 to 26 scales followed by 11 to 18 relatively small scutes. Breast completely scaly. Vertebrae
10+14. Colour: in life, generally silvery, greyish above, paler below; no opercular spot but adults with
posterior margin of preopercle distinctly dark to black; sides sometimes with 6 or 7 dusky oblique
bands, mostly above midline; large adults occasionally with scattered dark blotches or a few small yellow
spots on sides; caudal, second dorsal, and anal fins dusky to dark; leading edge of pelvic fins and distal
margin of anal fin with a narrow white border; pectoral fins pale.
Size: Largest specimen examined 39 cm fork length and 41.5 cm total length.
Habitat, biology, and fisheries: Appears to be largely confined to outer reef areas and is often taken
while bottom fishing. No information available on dietary preferences. Caught on hook-and-line, with
trawls, and several types of artisanal gear.
the Indian Ocean, except ab-
sent from Persian Gulf; else-
where in Indo-West Pacific
known from southern Japan
to eastern Australia and east-
ward to the Marshall Islands,
Fiji, and Samoa.
2708 Bony Fishes
Carangoides praeustus (Bennett, 1830)

Frequent synonyms / misidentifications: Caranx praeustus Bennett, 1830; C. ire Cuvier, 1833;
C. melanostethos Day, 1865 / None.
FAO names: En - Brownback trevally; Fr - Carangue à dos brun; Sp - Jurel pardo.
14 cm fork length
Diagnostic characters: Body elongate and compressed; dorsal and ventral profiles similar and gently
convex, dorsal profile of head nearly straight. Eye diameter slightly smaller than snout length. Both jaws
with an irregular row of small conical teeth; upper jaw with a narrow band of teeth anteriorly, the outer
series slightly enlarged; vomerine tooth patch anchor-shaped with a distinct posteromedian exten-
sion. Gill rakers (including rudiments) 9 to 11 on upper limb and 22 to 26 on lower limb of first gill arch
(total 32 to 37). Two separate dorsal fins, the first with VIII spines, the second with I spine and 21 to 24
soft rays; anal fin with II detached spines followed by I spine and 18 to 20 soft rays; spinous dorsal fin
moderately high, longest spine nearly equal height of soft dorsal-fin lobe. Lateral line anteriorly with
a moderate regular arch, with junction of curved and straight parts below vertical from between seventh
and eleventh soft rays of second dorsal fin; chord of curved part of lateral line slightly longer than, to slightly
shorter than straight part of lateral line, contained 0.9 to 1.2 times in straight part; straight part of lateral
line with 4 to 12 scales followed by 23 to 34 scutes. Breast usually completely scaly or nearly so (see
“Remarks”); laterally, naked area of breast separated from naked base of pectoral fins by a broad
band of scales (naked area of breast scarcely visible in lateral view). Vertebrae 10+14. Colour: in
life, bluish grey above, silvery white below; no dark opercular spot; distal half of second dorsal-fin lobe
abruptly black, sometimes with a white margin, widest anteriorly; remainder of fin pale dusky; caudal
fin pale yellowish, other fins hyaline to whitish.
Size: A relatively small species, largest specimen examined 19.5 cm fork length and 22 cm total length;
commonly to about 16 cm fork length.
Habitat, biology, and fisheries: Inhabits coastal waters throughout its range. Caught on hook-and-line,
with bottom trawls, and several types of artisanal gear.
Distribution: In the Indian Ocean from the
Persian Gulf to the Bay of Bengal; else-
where in the Indo-West Pacific known from
the Gulf of Thailand, Indonesia, Borneo,
and the Philippines.
Remarks: This species, as currently recog-
nized, consists of 2 populations that occur
in different geographic regions (Persian
Gulf to Bay of Bengal versus WCP area -
see distribution map) that differ in gill-raker
counts and also tend to have different pat-
terns of breast squamation as follows (con-
dition in non-WCP population in parentheses): total gill rakers on first gill arch 32 to 37 (40 to 47), of which
9 to 11 (11 to 15) on upper limb and 22 to 26 (28 to 32) on lower limb; squamation pattern: breast usually
completely scaly or nearly so, if partially naked midventrally naked area rarely extending entire length of
breast (breast typically with narrow naked midventral area extending entire length of breast). If future
studies show that these 2 “geographic races” are actually represented by different sister species, the oldest
available name for the Indian Ocean species is Carangoides ire (Cuvier).
Carangoides talamparoides Bleeker, 1852

Frequent synonyms / misidentifications: None / Carangoides malabaricus (Bloch and Schneider,
1801).
FAO names: En - Imposter trevally; Fr - Carangue cavallys; Sp - Jurel impostor.
26 cm fork length
Diagnostic characters: Body strongly compressed, almost ovate; dorsal profile of head strongly elevated
to nape, almost straight in profile. Eye diameter distinctly smaller than snout length. Both jaws with bands
of small villiform teeth, anteriorly some outer teeth conical; vomerine tooth patch roughly triangular, without
a posteromedian extension. Gill rakers (including rudiments) 6 to 9 on upper limb and 19 to 22 on lower
limb of first gill arch (total 27 to 31). Two separate dorsal fins, the first with VIII spines, the second with
I spine and 20 to 23 soft rays; anal fin with II detached spines followed by I spine and 17 to 19 soft rays;
lobe of second dorsal fin shorter than head length. Lateral line anteriorly with a moderate regular arch, with
junction of curved and straight parts below vertical from between twelfth and fourteenth soft rays of second
times in straight part; straight part of lateral line with 20 to 32 weak scutes, and 32 to 52 total elements
(including anterior scales). Breast naked ventrally to distinctly behind pelvic fins, often to origin of
second anal fin; laterally, naked area of breast extends diagonally to naked base of pectoral fins,
including small area anteriorly just above pectoral-fin base. Vertebrae 10+14. Colour: in life, generally
silvery, bluish grey above, silvery white below; opercle with a small black spot on upper margin; tongue
white to pale grey; soft dorsal and anal fins dusky; caudal fin with central rays dusky yellow with black
distal margin.
Habitat, biology, and fisheries: Inhabits coastal waters throughout its range. Caught on hook-and-line,
with bottom trawls and several types of artisanal gear.
Indian Ocean known from the
Gulf of Oman eastward to Sri
Lanka; elsewhere in the Indo-
West Pacific known from the
Gulf of Thailand, Sumatra,
Borneo, Philippines, Guam,
and Australia.
2710 Bony Fishes
Caranx bucculentus Alleyne and Macleay, 1877

FAO names: En - Bluespotted trevally.
19.4 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile strongly convex anteriorly, ventral profile
only slightly convex. Adipose eyelid moderately developed, small anteriorly, posterior adipose eyelid
extends onto eye to near rear border of pupil. End of upper jaw extends to posterior edge of pupil or a little
beyond. Upper jaw with outer row of strong canines widely spaced in adults, and an inner band of small
villiform teeth, widest at symphysis; lower jaw with a single row of strong conical teeth widely spaced in
adults. Gill rakers (including rudiments) 7 to 10 on upper limb and 17 to 21 on lower limb of first gill arch
(total 26 to 31). Two separate dorsal fins, the first with VIII spines, the second with I spine and 18 or 19
soft rays; anal fin with II detached spines followed by I spine and 15 to 17 soft rays; total soft dorsal- and
anal-fin rays 34 to 36; dorsal-fin lobe contained 3.4 to 5.1 times in fork length in specimens larger than
15 cm fork length. Curved part of lateral line short and strongly arched, chord of curved part
contained 2.5 to 3.3 times in straight part (to caudal-fin base); straight part of lateral line with no anterior
scales and 33 to 39 strong scutes. Breast naked posteroventrally to distinctly behind origin of pelvic
fins and diagonally to naked base of pectoral fins. Vertebrae 10+14. Colour: in life pale olive green
above, silvery white below; adults with small blue spots on upper half of body; large dark spot at upper
end of opercle and conspicuous black spot at upper base of pectoral fins; fins pale yellow-green.
Size: Largest specimen examined 56 cm fork length and 65 cm total length.
Habitat, biology, and fisheries: Feeds primarily on fishes. Caught on hook-and-line and commonly taken
in trawls.
Taiwan Province of China, In-
donesia (Ambon), Australia,
and Papua New Guinea.
Caranx heberi (Bennett, 1830)

Frequent synonyms / misidentifications: Caranx sem Cuvier, 1833; C. williamsi J.L.B. Smith, 1968;
C. sansun of authors / None.
FAO names: En - Blacktip trevally; Fr - Carangue cocole; Sp - Jurel cocolí.
55 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile strongly convex to second dorsal fin,
ventral profile only slightly convex. Adipose eyelid moderately developed, small anteriorly, posterior
adipose eyelid extends onto eye to rear border of pupil. End of upper jaw extends to posterior edge of pupil
or a little beyond. Upper jaw with outer row of strong canines widely spaced in adults, and an inner band
of small villiform teeth, widest at symphysis; lower jaw with a single row of strong conical teeth widely
spaced in adults. Gill rakers (including rudiments) 6 to 8 on upper limb and 17 to 19 on lower limb of
first gill arch (total 24 to 27, but rarely 24). Two separate dorsal fins, the first with VIII spines, the second
rays; total soft dorsal- and anal-fin rays 34 to 38; dorsal-fin lobe contained 5.3 to 7.4 times in fork length
in specimens larger than 15 cm fork length. Straight part of lateral line with 0 to 4 anterior scales followed
by 30 to 40 strong scutes. Breast squamation variable, completely scaly (uncommon) to naked
ventrally and typically with a small to large patch of prepelvic scales. Vertebrae 10+14. Colour: in
life, adults with head and body dark bronze to yellow-green above, silvery bronze to yellow-green
below; no dark spot on upper margin of opercle; caudal fin bright yellow or olive-yellow to dusky,
typically with distal half of fin darker or black (especially in juveniles); other fins bright yellow to
yellow-olive, sometimes dusky without any yellow hue; in juveniles and young adults black pigment
in upper caudal-fin lobe typically present.
Size: Maximum verified size 78 cm fork length; weight to 7.7 kg.
Habitat, biology, and fisheries: Prefers clean, open coastal waters; adults are particularly common in
small aggregations over rocky reefs and are seldom found in turbid or estuarine waters. Feeds primarily
on fishes and crustaceans. Caught mainly on hook-and-line; also with gill nets and other artisanal gear.
uted in coastal waters
throughout most of the Indian
Ocean, including the Persian
Gulf, but no verified records
from the Red Sea; elsewhere
in the Indo-West Pacific
known from Indonesia, the
Arafura Sea, Ambon, New
Guinea, and New Britain.
2712 Bony Fishes
Caranx ignobilis (Forsskål, 1775)

FAO names: En - Giant trevally; Fr - Carangue têtue; Sp - Jurel gigante.
59.5 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile strongly convex anteriorly, ventral
profile only slightly convex. Adipose eyelid moderately developed, small anteriorly, posterior adipose eyelid
extends onto eye to rear border of pupil. End of upper jaw extends to posterior edge of pupil or a little
beyond. Upper jaw with outer row of strong canines widely spaced in adults, and an inner band of small
villiform teeth, widest at symphysis; lower jaw with a single row of strong conical teeth widely spaced in
adults. Gill rakers (including rudiments) 5 to 7 on upper limb and 15 to 17 on lower limb of first gill arch
(total 20 to 24). Two separate dorsal fins, the first with VIII spines, the second with I spine and 18 to 21
soft rays; anal fin with II detached spines followed by I spine and 15 to 17 soft rays; total soft dorsal- and
anal-fin rays 34 to 38; dorsal-fin lobe contained 4.4 to 6.2 times in fork length in specimens larger than
15 cm fork length. Straight part of lateral line with 0 to 4 anterior scales followed by 26 to 38 strong scutes.
Breast naked ventrally, typically with a small to large patch of prepelvic scales. Vertebrae 10+14.
Colour: in life, adults with head and body silvery grey to black above, usually paler below; fins
usually uniformly grey to black, except leading edge and tips of dorsal- and anal-fin lobes paler; no dark
spot on upper margin of opercle; mature males usually darker than females, sometimes head and fins
completely black; juveniles and young adults generally with more silvery bodies and paler fins, fish from
turbid coastal waters often with yellow fins, the anal fin usually brightest.
Size: Maximum verified size 146 cm total length and 52.6 kg; commonly to about 80 cm fork length. World
angling record (IGFA) 66 kg (145 lb 8 oz).
Habitat, biology, and fisheries: Abundant in all marine habitats; juveniles often encountered in estuaries.
Feeds primarily on fishes. Caught mainly on hook-and-line and by spearing; also with gill nets and other
artisanal gear. An excellent sports fish.
West Pacific from southern
Japan to Australia, and east-
ward to Hawaii and the Mar-
quesas Islands.
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Caranx lugubris Poey, 1860

Frequent synonyms / misidentifications: Caranx ascensionis Cuvier, 1833 [Smith-Vaniz and Randall,
1994, Bull. Zool. Nomen., 15 (4), have petitioned the International Commission of Zoological Nomenclature
to conserve the name C. lugubris]; C. ishikawai Wakiya, 1924 / None.
FAO names: En - Black jack; Fr - Carangue noire; Sp - Jurel negro.
54 cm fork length
(after Masada et al., 1984)
profile only slightly convex; profile of head relatively steep and angular. Adipose eyelid moderately
developed, weak anteriorly, posterior eyelid extends onto eye to rear half of pupil. End of upper jaw extends
to below middle of eye. Upper jaw with outer row of strong canines widely spaced in adults, and an inner
band of small villiform teeth, widest at symphysis; on lower limb of first gill arch jaw with a single row of
strong conical teeth widely spaced in adults. Gill rakers (including rudiments) 6 to 8 on upper limb and 17
to 22 on lower limb of first gill arch (total 23 to 30). Two separate dorsal fins, the first with VIII spines, the
second with I spine and 20 to 22 soft rays; anal fin with II detached spines followed by I spine and 16 to
19 soft rays; total soft dorsal- and anal-fin rays 37 to 41; dorsal-fin lobe contained 2.3 to 5.3 times in
fork length in specimens larger than 15 cm fork length. Straight part of lateral line with 26 to 32 strong
scutes. Breast completely scaly. Vertebrae 10+14. Colour: in life, head, body, and fins essentially
uniform grey to brown, median fins and lateral-line scutes usually dark brown to black; upper end
of opercle frequently with a dark spot, much smaller than pupil diameter.
Size: Largest specimen examined 73.5 cm fork length (maximum weight not recorded).
Habitat, biology, and fisheries: Uncommon in shoal areas, confined to clear, offshore waters usually at
depths of 25 to 65 m. Its early life history is unknown. Feeds primarily on fishes. Caught primarily on
hook-and-line; also with traps and gill nets.
Distribution: A circumtropi-
cal species, very widespread
but restricted to oceanic insu-
lar habitats.
2714 Bony Fishes
Caranx melampygus Cuvier, 1833

Frequent synonyms / misidentifications: Caranx stellatus Eydoux and Souleyet, 1841; C. medusicola
Jordan and Starks, 1895 / None.
FAO names: En - Bluefin trevally; Fr - Carangue aile bleue; Sp - Jurel de aleta azul.
50 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile moderately convex to second dorsal
fin, ventral profile only slightly convex. Adipose eyelid weakly developed, small anteriorly, posterior adipose
eyelid extends onto eye to rear border of pupil in large adults. End of upper jaw extends to below anterior
margin of eye. Upper jaw with outer row of strong canines widely spaced in adults, and an inner band of
small villiform teeth, widest at symphysis; on lower limb of first gill arch jaw with a single row of strong
conical teeth widely spaced in adults. Gill rakers (including rudiments) 5 to 9 on upper limb and 17 to
21 on lower limb of first gill arch (total 25 to 29, usually 26 or 27). Two separate dorsal fins, the first with
VIII spines, the second with I spine and 21 to 24 soft rays; anal fin with II detached spines followed by I
spine and 17 to 20 soft rays, total soft dorsal- and anal-fin rays 39 to 44; dorsal-fin lobe contained 4.2 to
6.7 times in fork length in specimens larger than 15 cm fork length. Straight part of lateral line with 0 to 10
anterior scales followed by 27 to 42 strong scutes. Breast completely scaly. Vertebrae 10+14. Colour:
adults with head and dorsal half of body brassy, suffused with blue, and covered with small blue
black spots (forming at about 16 to 22 cm fork length and increasing in number with size); ventrally
body generally silvery white; second dorsal, anal, and caudal fins electric blue; pelvic fins whitish and
pectoral fins mostly pale yellow; in juveniles and young adults, head and body silvery grey and fins
pale to dark dusky, except pectoral fins yellow.
Size: Maximum verified size 80 cm total length and 6.8 kg; reported to attain 100 cm total length.
Habitat, biology, and fisheries: Invariably associated with reef habitats throughout its broad range; most
common in offshore areas. Feeds primarily on other fishes; reported to move up onto reef flats at night to
feed. Caught primarily on hook-and-line and by spearing; also with traps and gill nets. An excellent sports
fish.
Distribution: Common
throughout most of the
western Indian Ocean, ex-
cept apparently unrecorded
from the Persian Gulf; else-
where generally distributed
throughout the Indo-West
Pacific and tropical Eastern
Pacific.
Caranx papuensis Alleyne and Macleay, 1877

Frequent synonyms / misidentifications: Caranx regularis Garman, 1903; C. celetus Smith, 1968 /
C. melampygus Cuvier, 1833 and “C. sansun.”
FAO names: En - Brassy trevally; Fr - Carangue bronze; Sp - Jurel bronceado.
39 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile moderately convex to second dorsal
fin, ventral profile only slightly convex. Adipose eyelid weakly developed. End of upper jaw extends to
below middle of eye. Upper jaw with outer row of strong canines widely spaced in adults, and an inner
band of small villiform teeth, widest at symphysis; on lower limb of first gill arch jaw with a single row of
strong conical teeth, widely spaced in adults. Gill rakers (including rudiments) 7 to 9 on upper limb and
18 to 21 on lower limb of first gill arch (total 26 to 30). Two separate dorsal fins, the first with VIII spines,
the second with I spine and 21 to 23 soft rays; anal fin with II detached spines followed by I spine and 16
to 19 soft rays; total soft dorsal- and anal-fin rays 37 to 41; dorsal-fin lobe contained 4.1 to 5.6 times in fork
length in specimens larger than 15 cm fork length. Straight part of lateral line with 0 to 3 anterior scales
followed by 31 to 39 strong scutes. Breast naked ventrally, usually with a small patch of prepelvic
scales. Vertebrae 10+14. Colour: in life, adults with head and body brassy to yellowish green above,
silvery below; a conspicuous pale spot, approximately diameter of pupil, on shoulder just behind
posterodorsal margin of opercle; small black spots (forming at about 25 cm fork length) scattered
on body above lateral line, becoming more numerous with age; caudal fin with upper lobe uniformly
dusky, on lower limb of first gill arch lobe dusky to bright yellow with a distinct, narrow white border;
other fins pale to dusky yellow; anal and pelvic fins also with narrow white distal margins.
Size: Maximum verified size 68 cm fork length and 6.4 kg; commonly to at least 55 cm fork length.
Habitat, biology, and fisheries: Predominantly a seaward reef species although juveniles have been
taken in estuaries; common on mainland reefs but rarely captured in offshore areas. Feeds primarily on
other fishes. Caught primarily on hook-and-line and by spearing; also with gill nets and other artisanal gear.
Ocean from South Africa to
Zanzibar, the Malagasy Re- ?
public, Mauritius, Seychelles,
and Sri Lanka; elsewhere in ?
the Indo-West Pacific from
Ryukyu Islands to Australia
and eastward to the Mariana
and Marquesas islands.
2716 Bony Fishes
Caranx sexfasciatus Quoy and Gaimard, 1825

Frequent synonyms / misientifications: Caranx marginatus Gill, 1863; C. elacate Jordan and Evermann,
1904 / None.
FAO names: En - Bigeye trevally; Fr - Carangue vorace; Sp - Jurel voraz.
62 cm fork length
Diagnostic characters: Body oblong, compressed; dorsal profile moderately convex anteriorly,
ventral profile only slightly convex; postorbital head length contained 6.4 to 8.2 times in fork length in
specimens larger than 15 cm fork length. Adipose eyelid well developed, moderate anteriorly, posterior
eyelid extends onto eye to rear border of pupil. End of upper jaw extends to beyond posterior margin of
eye. Upper jaw with outer row of strong canines widely spaced in adults, and an inner band of small villiform
teeth, widest at symphysis; on lower limb of first gill arch jaw with a single row of strong conical teeth widely
spaced in adults. Gill rakers (including rudiments) 6 to 8 on upper limb and 15 to 19 on lower limb of first
gill arch (total 21 to 25). Two separate dorsal fins, the first with VIII spines, the second with I spine and 19
to 22 soft rays; anal fin with II detached spines followed by I spine and 14 to 17 soft rays; total soft dorsal-
and anal-fin rays 33 to 39; dorsal-fin lobe contained 5 to 6.6 times in fork length in specimens larger
than 15 cm fork length. Straight part of lateral line with 0 to 3 anterior scales followed by 27 to 36 strong
scutes. Breast completely scaly. Vertebrae 10+15. Colour: in life, adults with head and body silvery
olive to iridescent blue-green above, silvery olive to whitish below; small blackish spot, much smaller
than pupil diameter, at upper angle of opercle (this spot evident on specimens of about 14 cm fork
length); second dorsal fin olive to blackish, the lobe with a white tip (white tip becomes more
obvious with increasing size); anal and caudal fins yellowish to black; lateral-line scutes dark to black.
In juveniles and young adults, head, body, and scutes more silvery and fins paler (grey to yellow) except
upper half of dorsal-fin lobe and upper lobe of caudal fin usually dark.
Size: Maximum verified size 78 cm fork length and 7.7 kg; commonly to about 60 cm fork length.
Habitat, biology, and fisheries: A reef-associated species commonly caught in large numbers hand-lin-
ing around lights at night in waters inside the reef; juveniles may occur in estuaries. Feeds primarily on
fishes and crustaceans. Caught mainly on hook-and-line; also with gill nets, purse seines, and other
artisanal gear.
uted throughout the tropical
Indo-West Pacific, including
Hawaii; also occurs in the
eastern Pacific Ocean and
from Ecuador to Mexico, in-
cluding the Galapagos Is-
lands.
Caranx tille Cuvier, 1833

Frequent synonyms / misidentifications: Caranx cynodon Bleeker, 1851 / Caranx sexfasciatus (Quoy
and Gaimard, 1825).
FAO names: En - Tille trevally; Fr - Carangue tille; Sp - Jurel tille.
43 cm fork length
profile only slightly convex; postorbital head length contained 5.7 to 7.3 times in fork length in specimens
larger than 15 cm fork length. Adipose eyelid well developed, moderate anteriorly, posterior eyelid extends
onto pupil. End of upper jaw extends beyond posterior margin of eye. Upper jaw with outer row of strong
canines widely spaced in adults, and an inner band of small villiform teeth, widest at symphysis; on lower
limb of first gill arch jaw with a single row of strong conical teeth widely spaced in adults. Gill rakers
(including rudiments) 6 to 8 on upper limb and 15 to 17 on lower limb of first gill arch (total 22 to 25). Two
separate dorsal fins, the first with VIII spines, the second with I spine and 20 to 22 soft rays; anal fin with
II detached spines followed by I spine and 16 to 18 soft rays; total soft dorsal- and anal-fin rays 36 to 40;
dorsal-fin lobe contained 5.7 to 8.8 times in fork length in specimens larger than 15 cm fork length.
Straight part of lateral line with 0 to 2 anterior scales followed by 33 to 42 strong scutes. Breast completely
scaly. Vertebrae 10+14. Colour: in life, adults with head and body dark olive-green to bluish grey above,
shading to silvery white below; blackish spot, at least 1/2 diameter of pupil, at upper margin of opercle
(this spot evident on specimens of about 20 cm fork length); second dorsal fin olive-grey to
blackish, the lobe without a white tip; anal and caudal fins yellow-olive to blackish; lateral-line scutes
grey, except dark in caudal peduncle region; in juveniles and young adults, head and body pale olive
to silvery grey and fins uniformly pale to dusky.
Size: Maximum verified size 69 cm fork length and 4.3 kg.
Habitat, biology, and fisheries: A coastal species with a preference for inshore neritic waters, mainly
around rocky and coral reefs. Feeds primarily on other fishes and crustaceans. Caught mainly on
hook-and-line; also with gill nets, purse seines, and other artisanal gear.
Distribution: South Africa
northward at least to
Zanzibar, northeast coast of ?
Malagasy Republic, and Sri ?
West Pacific known from Oki-
nawa, Guam, the Philippines,
Indonesia, northern Australia,
New Guinea, and Fiji.
2718 Bony Fishes
Decapterus kurroides Bleeker, 1855

Frequent synonyms / misidentifications: Decapterus kurroides akaadsi Abe, 1958 / None.
FAO names: En - Redtail scad; Fr - Comète maouane; Sp - Macarela maouana.
24.2 cm fork length
(drawn by M. Wheaton)
Diagnostic characters: Body elongate, moderately slender and slightly compressed. Adipose eyelid well
developed, completely covering eye except for a vertical slit centred on pupil; Scales on top of head extend
forward to anterior margin of pupil. Oral valve (membrane) at symphysis of upper jaw dusky or transparent.
Posterior end of upper jaw straight above, slightly concave and not noticeably slanted anterovertrally.
Upper jaw with a narrow band of minute teeth, widest anteriorly; lower jaw with a single irregular series of
minute teeth. Gill rakers (including rudiments) 9 to 12 on upper limb and 26 to 32 on lower limb of first gill
arch (total 36 to 41). Shoulder girdle (cleithrum) margin with 2 small papillae, the lower papilla larger.
Two widely separate dorsal fins, the first with VIII spines, the second with I spine and 28 to 30 soft rays
(including finet); anal fin with II detached spines followed by I spine and 22 to 26 soft rays (including finlet);
terminal dorsal- and anal-fin rays each consisting of a widely detached finlet; pectoral fins moderately
long (88 to 105% of head length), tip of appressed fins typically extending to or slightly beyond a vertical
line from second dorsal-fin origin. Lateral line anteriorly with a low regular arch, with junction of curved
and straight parts below vertical from eleventh to thirteenth soft rays of second dorsal fin; scales in curved
part of lateral line 47 to 55; scutes in curved part 0 to 2; straight part without anterior scales, and
with 31 to 36 scutes; total scales and scutes in lateral line (excluding scales on caudal fin) 80 to 86.
Vertebrae 10+14. Colour: in life, bluish green above, silvery white below; moderate black blotch on margin
of opercle near upper edge; caudal fin red; spinous dorsal and second dorsal-fin lobe sometimes dark,
anal and pelvic fins pale dusky to white.
fork length.
Habitat, biology, and fisheries: A deep-water, schooling species taken in depths of 100 to 300 m, but
usually below 150 m. Feeds primarily on smaller planktonic invertebrates. Caught with midwater trawls.
Distribution: D i s t r i b u t i o n
poorly known, due to the
deep-water habitat prefer-
ence of this species. In the
Indian Ocean known from off
Kenya, Tanzania, and Reun-
ion Island; elsewhere in the
Indo-West Pacific known from
Japan, Taiwan Province of
China, Indonesia, the Philip-
pines, and Australia.
Decapterus macarellus Cuvier, 1833

Frequent synonymys / misidentifications: Decapterus jacobeus (Cuvier, 1833); D. pinnulatus (Eydoux
and Souleyet, 1841); D. canonoides Jenkins, 1903 / Decapterus macrosoma Bleeker, 1851 (a valid species,
but this name has often been misapplied to D. macarellus).
FAO names: En - Mackerel scad; Fr - Comète maquereau; Sp - Macarela caballa.
19.5 cm fork length
Diagnostic characters: Body very elongate, slender and nearly round in cross-section. Adipose eyelid
well developed, completely covering eye except for a vertical slit centred on pupil. Scales on top of head
extend forward to anterior margin of pupil. Oral valve (membrane) at symphysis of upper jaw conspicu-
ously white in adults. Posterior end of upper jaw straight above, moderately rounded and noticeably
slanted anteroventrally. Both jaws without teeth. Gill rakers (including rudiments) 10 to 13 on upper limb
and 34 to 41 on lower limb of first gill arch (total 45 to 54). Shoulder girdle (cleithrum) margin with 2
small papillae, the lower papilla larger. Two widely separate dorsal fins, the first with VIII spines, the
second with I spine and 31 to 37 soft rays (including finlet); anal fin with II detached spines followed by I
spine and 27 to 31 soft rays (including finlet); terminal dorsal- and anal-fin rays each consisting of a
widely detached finlet; pectoral fins short (58 to 72% of head length), tip of appressed fins falling
considerably short of a vertical line from second dorsal-fin origin. Lateral line anteriorly with a low
regular arch, with junction of curved and straight parts below vertical from eleventh to thirteenth soft rays
of second dorsal fin; scales in curved part of lateral line 58 to 75, no scutes in curved part; straight
part with 18 to 39 scales, followed by 24 to 40 scutes; total scales and scutes in lateral line (excluding
scales on caudal fin) 110 to 138. Vertebrae 10+14. Colour: in life, bluish green above, silvery below; small
black blotch on margin of opercle near upper edge; caudal fin yellow-green; dorsal-fin lobe sometimes dark
distally, anal and pelvic fins pale to whitish.
fork length.
Habitat, biology, and fisheries: A schooling species, occurring mostly in open water and common in
insular habitats. Sometimes near the surface but usually taken in depths between 40 and 200 m. Feeds
primarily on smaller plank-
tonic invertebrates. Caught
with purse seines and trawls.
Distribution: A circumtropi-
cal species. Widespread
throughout the tropical Indian
Ocean, including the Red Sea
but not the Persian Gulf; in the
Indo-West Pacific to the Mar-
quesas Islands, north to
southern Japan and south to
Australia.
2720 Bony Fishes
Decapterus macrosoma Bleeker, 1851

Frequent synonyms / misidentifications: Decapterus afuerae Hildebrand, 1946; D. lajang (a synonym
of D. russelli, but this name has often been misapplied to D. macrosoma) / Decapterus macarellus (Cuvier,
1833).
FAO names: En - Shortfin scad; Fr - Comère fine; Sp - Macarela alicorta.
19.8 cm fork length
Diagnostic characters: Body very elongate, slender and nearly rounded. Eye moderate, with adipose
eyelid well developed, completely covering eye except for a vertical slit centred on pupil. Scales on top of
head do not extend forward to beyond posterior margin of pupil. Oral valve (membrane) at symphysis of
upper jaw dusky or transparent. Posterior end of upper jaw concave above, rounded and produced
below. Upper jaw without teeth and lower jaw with a single series of minute teeth. Gill rakers (including
rudiments) 10 to 12 on upper limb and 34 to 38 on lower limb of first gill arch (total 43 to 50). Shoulder
girdle (cleithrum) margin with 2 small papillae, the lower papillae larger. Two widely separate dorsal
fins, the first with VIII spines, the second with I spine and 33 to 39 soft rays (including finlet); anal fin with
II detached spines followed by I spine and 27 to 31 soft rays (including finlet); terminal dorsal- and anal-fin
rays each consisting of a widely detached finlet; pectoral fins short (61 to 75% of head length), tip of
appressed fins falling considerably short of a vertical line from second dorsal-fin origin. Lateral line
anteriorly with a low regular arch, with junction of curved and straight parts below vertical from eleventh to
thirteenth soft rays of second dorsal fin; scales in curved part of lateral line 58 to 72; no scutes in
curved part; straight part with 14 to 29 scales followed by 24 to 40 scutes; total scales and scutes in
lateral line (excluding scales on caudal fin) 110 to 126. Vertebrae 10+14. Colour: in life, metallic blue
above, silvery below; small black blotch on margin of opercle near upper edge; caudal fin hyaline to dusky
and dorsal-fin lobe sometimes dark distally; other fins mostly pale.
Size: The smallest Indo-West Pacific species of Decapterus. Maximum fork length about 30 cm, but rarely
exceeding 25 cm fork length.
Habitat, biology, and fisheries: A schooling species, in depths between 30 and at least 170 m, reported
to occur much deeper. Feeds primarily on smaller planktonic invertebrates. Caught with purse seines and
trawls.
the tropical Indo-West Pacific,
from East Africa, including the
Red Sea but not the Persian
Gulf, north to southern Japan
and south to Australia and
eastward to Hawaii; also pre-
sent in the eastern Pacific
Ocean, from the Gulf of Cali-
fornia and Peru.
Decapterus muroadsi (Temminck and Schlegel, 1844)

Frequent synonyms / misidentifications: Caranx scombrinus Valenciennes, 1844; Decapterus hypodus
Gill, 1862 / Decapterus macarellus (Cuvier, 1833).
FAO names: En - Amberstripe scad.
33.6 cm fork length
head do not extend forward to anterior margin of pupil. Oral valve (membrane) at symphysis of upper
jaw white. Posterior end of upper jaw straight above, not noticeably slanted anteroventrally. Upper
jaw with irregular series of minute teeth anteriorly, becoming obsolete with age; lower jaw with a single
series of minute teeth. Gill rakers (including rudiments) 13 to 15 on upper limb and 36 to 42 on lower limb
of first gill arch (total 47 to 56). Shoulder girdle (cleithrum) margin with 2 small papillae, the lower
papillae larger. Two widely separate dorsal fins, the first with VIII spines, the second with I spine and 29
to 33 soft rays (including finlet); anal fin with II detached spines followed by I spine and 25 to 28 soft rays
(including finlet); terminal dorsal- and anal-fin rays each consisting of a widely detached finlet;
pectoral fins short (71 to 93% of head length), tip of appressed fins falling usually short of a vertical line
from second dorsal-fin origin. Lateral line anteriorly with a low regular arch, with junction of curved and
straight parts below vertical from eleventh to thirteenth soft rays of second dorsal fin; scales in curved
part of lateral line 54 to 76; no scutes in curved part; straight part with 5 to 15 scales followed by 32
to 42 scutes; total scales and scutes in lateral line (excluding scales on caudal fin) 102 to 121. Vertebrae
10+14. Colour: in life, bluish green above, silvery below; small black blotch on margin of opercle near
upper edge; amber stripe often present on sides; caudal fin with lower lobe dusky and upper lobe
greenish yellow; dorsal-fin lobe usually dark distally; other fins mostly pale.
Size: Maximum fork length at least 45 cm; commonly to about 30 cm fork length.
Habitat, biology, and fisheries: A pelagic schooling species. Feeds primarily on smaller planktonic
invertebrates. Caught with purse seines and trawls.
Distribution: Broadly dis-
tributed, warm-temperate
Indo-West Pacific species
known from Australia, south-
ern Japan, northern Hawaiian
Islands, Rapa, Easter Island,
Nazca Ridge, the eastern Pa-
cific Ocean (Gulf of California
to Peru), and an apparently
relict population at St. Helena
Is. in the mid-South Atlantic
Ocean.
2722 Bony Fishes
Decapterus russelli (Rüppell, 1830)

Frequent synonyms / misidentifications: Decapterus kiliche (Cuvier, 1833); D. dayi Wakiya, 1928; D. lajang
Bleeker, 1885 (this name has often been misapplied to D. macrosoma) / Decapterus maruadsi (Temminck
and Schlegel, 1844) (a questionably valid species, apparently restricted to Japan and the South China
Sea, but this name has often been misapplied to D. russelli); D. tabl Berry, 1968.
FAO names: En - Indian scad; Fr - Comère indienne; Sp - Macarela indica.
23.5 cm fork length

Diagnostic characters: Body elongate, moderately slender and slightly compressed. Eye moderate, with
adipose eyelid well developed, completely covering eye except for a vertical slit centred on pupil. Scales
on top of head do not extend forward to beyond centre of pupil. Oral valve (membrane) at symphysis of
upper jaw dusky or transparent. Posterior end of upper jaw straight above, slightly concave and not
noticeably slanted anteroventrally. Both jaws with an irregular row of minute teeth anteriorly and a single
series posteriorly. Gill rakers (including rudiments) 10 to 14 on upper limb and 30 to 39 on lower limb of
first gill arch (total 41 to 53). Shoulder girdle (cleithrum) margin with 2 small papillae, the lower papilla
larger. Two widely separate dorsal fins, the first with VIII spines the second with I spine and 28 to 33 soft
rays (including finlet); anal fin with II detached spines followed by I spine and 25 to 29 soft rays (including
finlet); terminal dorsal- and anal-fin rays each consisting of a widely detached finlet; pectoral fins
76.5 to 97% of head length, tip of appressed fins falling short of to slightly beyond a vertical line from second
dorsal-fin origin. Lateral line anteriorly with a low regular arch, with junction of curved and straight parts
below vertical from eleventh to thirteenth soft rays of second dorsal fin; scales in curved part of lateral
line 42 to 62, scutes in curved part 0 to 4, straight part with 0 to 4 scales, followed by 30 to 40 scutes;
total scales and scutes in lateral line (excluding scales on caudal fin) 77 to 102. Vertebrae 10+14.
Colour: in life, bluish green above, silvery below; small black blotch on margin of opercle near upper edge;
caudal fin hyaline to dusky brown, second dorsal fin hyaline basally, light dusky distally; other fins mostly
hyaline, except pelvic fins of adult males slightly dark.
Size: Largest specimen examined 35 cm fork length; commonly to about 20 cm fork length.
Habitat, biology, and fisheries: The most common species of Decapterus in inshore waters in depths not
exceeding 100 m. Reaches sexual maturity during the first year of life at about 12 cm total length. Fish of 15 cm
total length (average weight 50 g) and 2 to 3 years of age are the most frequent in catches. Feeds primarily on
smaller planktonic invertebrates. Caught with purse seines and trawls.
Distribution: Broadly distributed through-
out the Indian Ocean and from Japan to
Australia in the western Pacific Ocean.
Remarks: Two somewhat different mor-
phological “forms” of Decapterus are here
recognized as D. russelli. The more abun-
dant form is characterized by a more ro-
bust body, larger scutes, fewer scales in
the curved part of the lateral line and tends
to have more gill rakers. The more elon-
gate “lajang” form attains a larger size and
usually has shorter pectoral fins. Some
morphologically intermediate specimens exist and because the “lajang” form tends to be more typical of
fish from insular habitats, the observed differences are believed to be environmentally determined.
Decapterus tabl Berry, 1968

Frequent synonyms / misidentifications: None / In Japan, the name Decapterus russelli has often been
misapplied to this relatively common species. The misapplication of the name originates with Bertin and
Dollfus (1948, Mem.Mus.natn.Hist.Nat.) who made an invalid neotype selection for D. russelli (Rüppell).
FAO names: En - Roughear scad; Fr - Comère queue rouge; Sp - Macarela rabo colorado.
35.5 cm fork length

head extend forward to anterior margin of pupil. Posterodorsal margin of opercular membrane minutely
serrated in adults. Oral valve (membrane) at symphysis of upper jaw dusky or transparent. Posterior
end of upper jaw straight above, slightly concave and not strongly slanted anteroventrally. Upper
jaw with an irregular series of minute teeth anteriorly that become obsolete with age. Lower jaw with a
single series of minute teeth. Gill rakers (including rudiments) 10 to 12 on upper limb and 30 to 33 on lower
limb of first gill arch (total 41 to 45). Shoulder girdle (cleithrum) margin with 2 small papillae, the lower
papilla larger. Two widely separate dorsal fins, the first with VIII spines, the second with I spine and 30
to 34 soft rays (including finlet); anal fin with II detached spines followed by I spine and 24 to 26 soft rays
(including finlet); terminal dorsal- and anal-fin rays each consisting of a widely detached finlet;
pectoral fins 62 to 83% of head length, tip of appressed fins falling slightly to distinctly short of a vertical
line from second dorsal-fin origin. Lateral line anteriorly with a low regular arch, with junction of curved and
straight parts below vertical from eleventh to fourteenth soft rays of second dorsal fin; scales in curved
part of lateral line 61 to 73; no scutes in curved and straight part with 4 to 12 scales, followed by
30 to 40 scutes; total scales and scutes in lateral line (excluding scales on caudal fin) 103 to 118.
Vertebrae 10+14. Colour: in life, metallic blue to greenish above, silvery below; small black blotch on
margin of opercle near upper edge; caudal fin bright red and tips of soft dorsal-fin rays tinged with
red.
Size: The largest Indo-West Pacific species of Decapterus. Maximum fork length about 48 cm, commonly
to 35 cm fork length.
Habitat, biology, and fisheries: Most captures of this deep-water, schooling species are from depths of
200 to 360 m. Feeds primarily on smaller planktonic invertebrates. Caught with purse seines and trawls.
Distribution: Distribution poorly known, due to the deep-water habitat preference of this species. In the
Indian Ocean known from a
single collection off Kenya.
Elsewhere in the Indo-West
Pacific known from Japan and
Okinawa, Indonesia, Austra-
lia, and the Hawaiian Archi-
pelago (Necker Island). Also
occurs in the western Atlantic
O c e a n : B e r m u d a, S ou th
America off Colombia and
Verlezuela, and at St. Helena
in the mid-South Atlantic.
2724 Bony Fishes
Elagatis bipinnulata (Quoy and Gaimard, 1825)

FAO names: En - Rainbow runner; Fr - Comère saumon; Sp - Macarela salmón.
30 cm fork length
Diagnostic characters: Body greatly elongate, almost fusiform. Head and snout pointed; mouth small,
upper jaw ending distinctly before eye (to below anterior margin of eye in young). Teeth in jaws in
villiform bands, minute teeth also on roof of mouth and on tongue. Dorsal fin with VI spines, followed by
I spine and 25 to 30 soft rays including a detached terminal 2-rayed finlet; anal fin with only I spine,
slightly detached from rest of fin and covered by skin in fish of larger sizes, followed by a second
spine continuous with the following 18 to 22 soft rays, including a detached 2-rayed finlet; anal-fin
base relatively short, contained about 1.5 times in second dorsal-fin base; pectoral fins short, about 2 times
in head length and about as long as pelvic fins; caudal fin deeply forked. Lateral line with a slight anterior
arch. No scutes; caudal peduncle grooves present. Vertebrae 10+14. Colour: dark olive-blue or green
dorsally and white ventrally; 2 narrow light blue or bluish white stripes along sides, with a broader
olive or yellowish stripe between them; fins dark with an olive or yellow tint.
Size: Maximum fork length 107 cm (possibly to 120 cm); commonly to 80 cm fork length. Maximum weight
10.5 kg.
Habitat, biology, and fisheries: A pelagic species, usually found at or near the surface, over reefs or
sometimes far offshore; may form sizeable schools. Feeds on invertebrates and small fishes. Caught
mainly on hook-and-line; also with gill nets. An excellent sports fish.
circumtropical species.
Gnathanodon speciosus (Forsskål, 1775)

Frequent synonyms / misidentifications: Caranx speciosus (Forsskål, 1775) / None.
FAO names: En - Golden trevally; Fr - Carangue royale jaune; Sp - Jurel dorado.
49.3 cm fork length
Diagnostic characters: Body compressed, oblong, lips noticeably papillose and upper jaw strongly
protactile. Eye diameter smaller than snout length. Upper jaw without teeth; lower jaw with a few
feeble teeth in young (smaller than 10 cm fork length) absent in adults. Gill rakers (including
dorsal fins, the first with VII spines, the second with I spine and 18 to 20 soft rays; anal fin with II detached
spines followed by I spine and 15 to 17 soft rays; lobe of second dorsal fin shorter than head length. Lateral
line anteriorly with a moderate regular arch, with junction of curved and straight parts below vertical from
between ninth and fourteenth soft rays of second dorsal fin; chord of curved part of lateral line longer to
slightly shorter than straight part of lateral line, contained 0.75 to 1.3 times in straight part; straight part of
lateral line with 17 to 24 scales followed by 17 to 26 scutes. Breast completely scaly. Vertebrae 10+14.
Colour: juveniles and young adults silvery to yellow with 7 to 11 black bands, usually alternating
broad and narrow; the first band oblique through eye, the second crossing back and opercle; upper margin
of opercle black; all fins yellow, tips of caudal-fin lobes black; dark opercular spot also fades with
age and body becomes yellow-green dorsally, pale yellow below.
Size: Maximum size not known but reported to attain 110 cm fork length. South African spearfishing record
14.5 kg.
Habitat, biology, and fisheries: Found inshore, including rocky reefs, deep lagoons, and seaward reefs.
A bottom feeder that uses its highly protractile mouth to root the sand and small crevices for crustaceans,
molluscs, and small fishes. Young display “piloting” habits with sharks and other large fish such as
groupers; their maneuverability protects them from their host, and the association probably affords them
some protection from other
predators. Caught mainly with
gill nets and by spearing.
uted throughout the Indo-
West Pacific, including
Hawaii; also occurs in the
eastern Pacific Ocean, from
Peru to Baja California.
2726 Bony Fishes
Megalaspis cordyla (Linnaeus, 1758)

FAO names: En - Torpedo scad; Fr - Comère torpille; Sp - Macarela torpedo.
28 cm fork length
Diagnostic characters: Body elongate, subcylindrical, a little compressed posteriorly, and caudal
peduncle strongly compressed with a marked medial keel; snout and lower jaw pointed. Eye moderate,
with well developed adipose eyelid completely covering eye except for a vertical slit centred on pupil. Upper
jaw extending posteriorly to centre of eye. Upper jaw with small villiform teeth, outer teeth moderately
enlarged; lower jaw with a single row of small teeth, except a narrow band anteriorly. Gill rakers (including
rudiments) 8 to 11 on upper limb and 18 to 22 on lower limb of first gill arch (total 26 to 32). Shoulder girdle
(cleithrum) margin smooth, without papillae. Two separate dorsal fins, the first with VIII spines, the second
with I spine and 18 to 20 soft rays, posterior 7 to 9 rays consisting of detached finlets; anal fin with
II detached spines followed by I spine and 16 or 17 soft rays, posterior 8 to 10 rays consisting of
detached finlets; spinous dorsal fin moderately high, longest spine height about equal to length of soft
dorsal-fin lobe. Lateral line strongly arched anteriorly, with junction of curved and straight parts
below vertical from fourth or fifth spine of dorsal fin; chord of curved part of lateral line contained 3.6
to 4.7 times in straight part (to caudal-fin base); scales in curved part of lateral line 21 to 28; straight part
with 51 to 59 very large scutes. Breast naked ventrally and laterally in a triangular area for about one
third distance to pectoral-fin base. Vertebrae 10+14. Colour: head and body bluish grey to green dorsally,
sides and belly silvery; large black opercular spot; dorsal and anal fins pale to yellow, distally dusky;
pectoral and pelvic fins pale, with upper half dusky; caudal fin dark, especially leading and trailing edges
of fin.
Size: Maximum reported total length 80 cm; commonly attains 30 to 40 cm total length and 3 to 4 kg.
Habitat, biology, and fisheries: A pelagic, schooling species. Feeds primarily on other fishes. Caught
on hook-and-line, with beach seines, trawls, purse seines, and traps.
West Pacific from Japan to
Australia and eastward to Fiji.
Records of the species from
Hawaii are erroneous.
Naucrates ductor (Linnaeus, 1758)

FAO names: En - Pilotfish; Fr - Poisson pilote; Sp - Pez piloto.
32.5 cm fork length
Diagnostic characters: Body elongate, shallow and barely compressed, with nearly equal upper and
lower profiles, but head profile tapering sharply above anterior half of upper jaw to produce a nearly blunt
snout. Upper jaw very narrow, posteriorly and extending to about below anterior margin of eye. Teeth
minute, in a band in upper and lower jaws. Gill rakers (including rudiments) 6 or 7 on upper limb and 15 to
20 on lower limb of first gill arch (total 21 to 27). Dorsal fin with IV or V spines (first spine minute and/or
last spine may be reduced and embedded in fish larger than 20 cm fork length), followed by I spine and
25 to 29 soft rays; anal fin with II spines slightly separated from rest of fin (first spine may be reduced
and embedded) followed by I spine and 15 to 17 soft rays; second dorsal-fin lobe short, contained 7.1 to
8.2 times in in fork length; anal-fin base short, contained 1.6 to 1.9 times in second dorsal-fin base. No
scutes; caudal peduncle with a well-developed lateral, fleshy keel on each side and dorsal and
ventral peduncle grooves. Vertebrae 10+15. Colour: in live fish, 6 or 7 black bands against a light
silvery background, but there also is a transient coloration (possibly aggressive display) with bands
disappearing and most of fish silvery white with 3 broad blue patched in tandem across back; in fresh or
preserved specimens, head dark, 5 or 6 dark, broad body bands and a similar band at end of caudal
peduncle, bands 3 to 6 extending through soft dorsal- and anal-fin membranes, and the bands persistent
at all sizes; rest of body bluish (fresh) or light to dusky; white tips prominent on upper and lower lobes
of caudal fin, and smaller white tips on anterior lobes of second dorsal and anal fins; fins otherwise
dusky to dark.
Size: Maximum size 63 cm fork length and 70 cm total length; commonly to 35 cm fork length; weight 0.5 kg
at 33 cm fork length.
Habitat, biology, and fisheries: Primarily pelagic in oceanic waters. This species has a semi-obligate
relationship with large sharks, rays and other fishes, turtles, and flotsam. Juveniles are frequently found
in association with jellyfish; the larvae are widespread in epipelagic oceanic waters. Maturity is reached
by at least 23 cm fork length. Caught with pelagic trawls.
circumtropical species.
click for next page

2728 Bony Fishes
Pantolabus radiatus (Macleay, 1881)

Frequent synonyms / misidentifications: Caranx radiatus Macleay, 1881; Absalom radiatus (Macleay,
1881) / None.
FAO names: En - Fringefin trevally.
, 19.2 cm fork length

Diagnostic characters: Body elongate oval, compressed; dorsal and ventral profiles equally convex;
snout pointed. Eye diameter about equal or larger than snout length, with adipose eyelid well developed
on posterior half of eye only. Upper jaw extending to below middle of centre of pupil, and with posterior
margin distinctly concave. Upper jaw with an outer row of moderately enlarged conical teeth bordered
by an inner band of small, sharply pointed teeth; lower jaw with a single row of moderately enlarged
conical teeth, sometimes with a few inner teeth anteriorly. Gill rakers (including rudiments) 11 to
13 on upper limb and 25 to 28 on lower limb of first gill arch (total 36 to 41). Two separate dorsal fins, the
first with VIII spines, followed by I spine and 23 to 26 soft rays; anal fin with II detached spines followed
by I spine and 18 to 20 soft rays; spinous dorsal fin relatively high, equal or subequal lobe of second dorsal
fin in females; second dorsal fin and anal fin with well-developed scaly basal sheaths; adult males
with all soft dorsal- and anal-fin rays produced as filaments (females without filaments). Lateral line
moderately arched anteriorly, with junction of curved and straight parts below a vertical from between fourth
and sixth soft rays of second dorsal fin; chord of curved part of lateral line contained 1.6 to 2.2 times in
straight part (to caudal-fin base); scales in curved part 33 to 41; straight part with 0 to 9 scales and 38 to
49 scutes; total scales and scutes in lateral line (excluding scales on caudal fin) 77 to 85. Breast completely
scaly, except sometimes a small naked patch on ventral midline anteriorly. Vertebrae 10+14. Colour:
olive-green to bluish green above, silvery to white below; black spot on operculum; spinous and soft dorsal
fins, and anal fin orange-yellow; caudal fin pale to bright yellow with tip of upper lobe black; pelvic fins white
and pectoral fins pale orange to hyaline.
Size: Largest specimen examined 21.5 cm fork length, 24.5 cm total length.
Habitat, biology, and fisheries: In northern Australia in coastal waters, sometimes entering river mouths,
rarely found in depths ex-
ceeding 30 m. Diet consists
primarily of epibenthic crusta-
ceans. Caught mainly with
bottom trawls.
from Papua New Guinea and
Australia: Port Headland,
Western Australia (20°18’S)
to Port Curtis, Queensland
(23°55’S).
Parastromateus niger (Bloch, 1795)

Frequent synonyms / misidentifications: Apolectus niger (Bloch, 1795); Formio niger (Bloch, 1795);
Citula halli Evermann and Seale, 1907/ None.
FAO names: En - Black pomfret; Fr - Castagnoline noire; Sp - Palometa negra.
Diagnostic characters: Body deep and compressed; dorsal

and ventral profiles of body strongly and equally convex.
Mouth terminal with upper jaw unrestricted dorsally and ending
below and slightly before anterior margin of eye. Both jaws with
a single row of small conical teeth. Gill openings unrestricted
laterally and ventrally. Dorsal fin with VI or V short spines
(embedded and not apparent in adults) followed by I spine
and 41 and 44 soft rays; anal fin with II spines (embedded
and not apparent in adults) followed by I spine and 35 to 39 juvenile
soft rays; profile of second dorsal and anal fins nearly 5.9 cm fork length
identical, with elevated, broadly rounded anterior lobes; pelvic fin
pelvic fins absent in specimens larger than about 10 cm present
fork length, and in juveniles positioned distinctly anterior to a vertical line through pectoral-fin
base; pectoral fins long and falcate. Lateral line very weakly arched anteriorly, with junction of straight and
curved parts below posterior third of dorsal fin; straight part of lateral line with 8 to 19 weak scutes, forming
a slight keel on caudal peduncle; scales small and deciduous, and almost completely covering dorsal
and anal fins; breast completely scaly. Vertebrae 10+14. Colour: in life, adults uniformly silvery grey
to bluish brown (yellowish brown when deciduous scales missing); fins with dark edges; young with
dark vertical bars and long black jugular pelvic fins.
Size: Maximum total length about 55 cm; commonly to 30 cm total length.
Habitat, biology, and fisheries: Generally near the bottom during the day, rising to near the surface at
night. Often occurs in large schools, and not uncommonly observed swimming on their sides. Usually found
in depths of 15 to 40 m, generally over
muddy bottoms. Feeds on zooplankton.
The principal gear used is gill nets, but boat
seines and trawls also take black pomfrets.
Distribution: Pelagic on the continental
shelf from South Africa, Mozambique,
Kenya, the Arabian Sea, Bay of Bengal,
Indonesia, the Philippines, China, south-
ern Japan, and Australia. Most abundant
on the west coast of India and in Indonesia.
2730 Bony Fishes
Pseudocaranx dentex (Bloch and Schneider, 1801)

Frequent synonyms / misidentifications: Caranx dentex (Bloch and Schneider, 1801); C. georgianus
Cuvier, 1833; Usacaranx georgianus (Cuvier, 1833); Caranx nobilis Macleay, 1881; C. cheilio Snyder,
1904; C. natalensis Gilchrist and Thompson, 1911 / None.
FAO names: En - White trevally; Fr - Carangue dentue; Sp - Jurel dentón.
20 cm fork length
Diagnostic characters: Body elongate, moderately deep and compressed, with dorsal and ventral profiles
similar. Eye relatively small with adipose eyelid poorly developed. Lips noticeably papillose and upper jaw
projecting beyond lower in large adults; end of upper jaw essentially vertical. Both jaws with a single
row of blunt conical teeth, upper jaw sometimes with an inner series of conical teeth anteriorly. Gill
rakers (including rudiments) 9 to 11 on upper limb and 20 to 24 on lower limb of first gill arch (total
29 to 35). Two separate dorsal fins, the first with VIII spines, the second with I spine and 24 to 28 soft
rays; anal fin with II detached spines followed by I spine and 20 to 24 soft rays; terminal dorsal and anal-fin
rays enlarged and a little separated from adjacent rays but not detached; spinous dorsal fin high, longest
spine height slightly longer than length of soft dorsal-fin lobe. Lateral line with a weak and extended anterior
arch, with junction of curved and straight parts of lateral line below vertical from twelfth to fourteenth soft
rays of second dorsal fin; chord of curved part of lateral line contained 0.6 to 0.85 times in straight part (to
caudal-fin base); curved part of lateral line with 57 to 78 scales; straight part of lateral line with 2 to 27
anterior scales and 16 to 30 scutes; total scales and scutes in lateral line (excluding scales on caudal fin)
91 to 115; no scales on preorbital bone below and in front of eye and on expanded part of maxilla;
breast completely scaly. Vertebrae 10+14-15. Colour: pale greenish to steely blue above, silvery below;
a yellow stripe usually present along sides (wider posteriorly) and at bases of soft dorsal and anal fins;
caudal fin and soft dorsal fins dusky yellow; a diffuse black spot on upper margin of opercle.
Size: Maximum fork length about 82 cm; commonly to 40 cm fork length; weight to 10.7 kg.
Habitat, biology, and fisheries: Found mainly on banks and shelf slope habitats at depths of 80 to 200 m
throughout much of its range, but at some localities juveniles and occasionally adults may also frequent
the shoreline. Fishes, molluscs, and crustaceans constitute the bulk of the diet of this opportunist bottom
feeder. Caught mainly with bottom trawls.
Distribution: A broadly distributed, anti-tropical species. In the Indo-Pacific known from off South Africa,
Japan, Australia, New Zealand, Rapa, Pitcairn, Hawaiian Archipelago, and Easter Island. Also occurs on
both sides of the Atlantic Ocean (Bermuda, off North Carolina, southern Brazil, Azores, Madeira, Canary,
Cape Verde, Ascension, and St. Helena
islands) and in the Mediterranean Sea.
Remarks: Pseudocaranx wrighti, which is
known only from western and southern
Australia and thus outside the WCP area,
is included in the key to species of Pseudo-
caranx. Australian fishery biologists need
to be able to distinguish both of these spe-
cies, which in the past have often been
confused.
Scomberoides commersonnianus Lacepède, 1802

Frequent synonyms / misidentifications: Chorinemus commersonnianus (Lacepède, 1802) / The name
Scomberoides lysan (Cuvier) had been generally misapplied to this species prior to Smith-Vaniz and
Staiger’s (1973, Proc. Cal. Acad. Sci.) revision of Scomberoides.
FAO names: En - Talang queenfish; Fr - Sauteur talang; Sp - Jurel saltador.
47 cm fork length
Diagnostic characters: Body oblong to elliptical, strongly compressed; ventral profiles similar, snout blunt
with dorsal profile of head and scales on midbody nape slightly convex. Upper lip joined to snout at midline
by a bridge of skin (frenum), except crossed by a shallow groove in very young. Upper jaw extends well
beyond posterior margin of eye in adults. Lower jaw with 2 rows of teeth separated by a shallow
groove, adults with teeth in inner row about equal in size to those in outer row. Gill rakers (excluding
dorsal fins, the first with VI or VII short spines, depressible into a shallow groove on dorsal midline, followed
by I spine and 19 to 21 soft rays; anal fin with II detatched spines followed by 16 to 19 soft rays; posterior
soft dorsal-and anal-fin rays consisting of semidetached finlets, bases of anal and second dorsal fins
about equal in length; height of second dorsal-fin lobe 14.4 to 19.8% of fork length in specimens larger
than 15 cm fork length; pelvic fins about equal in length or shorter than pectoral fins. Lateral line only slightly
irregular, weakly to moderately convex above pectoral fins, becoming straight posteriorly. No scutes;
scales on midbody below lateral line, partially embedded and broadly oblong. No caudal peduncle
grooves. Vertebrae 10+16. Colour: in life, head and body dusky green to bluish dorsally, grey to silvery
below, large individuals often golden, especially ventrally; sides of adults with 5 to 8 large, plumbeous
blotches above or touching lateral line, first 2 may intersect lateral line; dorsal and anal fins dusky
to dark, uniformly pigmented; pectoral fins of adults frequently with a dusky blotch ventrally.
Size: Largest specimen examined 94 cm fork length; reported to attain 120 cm total length. African
spearfishing record 14.4 kg.
Habitat, biology, and fisheries: Although periodically entering estuaries, generally intolerant of low
salinities, and turbid water; swims in small groups and usually frequents reefs and offshore islands. As is
true of all members of the genus, the spines of the first dorsal and anal fins are venomous (especially the
latter) and capable of inflicting painful stings. Primarily a daytime feeder on fishes, cephalopods, and other
pelagic prey; young use spe-
cialized juvenile dentition to
aggressively remove scales
and epidermal tissue from
other fishes. Caught with drift
set nets, gill nets, seines, and
on hook-and-line.
Gulf of Thailand, Okinawa, In-
donesia, the Philippines, Aus-
tralia, and Papua New Guinea.
2732 Bony Fishes
Scomberoides lysan (Forsskål, 1775)

Frequent synonyms / misidentifications: Chorinemus sanctipetri Cuvier, 1832; C. orientalis Temminck and
Schlegel, 1844 / The name Scomberoides lysan (Cuvier) had been generally misapplied to S. commersonnianus
prior to Smith-Vaniz and Staiger’s (1973, Proc. Cal. Acad. Sci.) revision of Scomberoides.
FAO names: En - Doublespotted queenfish; Fr - Sauteur sabre; Sp - Jurel sable.
52 cm fork length
Diagnostic characters: Body oblong to elliptical, strongly compressed; dorsal and ventral profiles nearly
equal, snout pointed with dorsal profile of head and nape slightly concave. Upper lip joined to snout at
midline by a bridge of skin (frenum), except crossed by a shallow groove in very young. Upper jaw extends
to or slightly beyond posterior margin of eye in adults. Lower jaw with 2 rows of teeth separated by a
shallow groove, adults with teeth in inner row equal in size to those in outer row. Gill rakers (excluding
dorsal fins, the first with VI or VII short spines, depressible into a shallow groove on dorsal midline, followed
by I spine and 19 to 21 soft rays; anal fin with II detached spines followed by 17 to 19 soft rays; posterior
soft dorsal- and anal-fin rays consisting of semidetached finets, bases of anal and second dorsal fins
about equal in length; height of second dorsal-fin lobe 9.1 to 14.2% of fork length in specimens larger than
15 cm fork length; pelvic fins about equal in length or shorter than pectoral fins. Lateral line only slightly
irregular, weakly to moderately convex above pectoral fins becoming straight posteriorly. No scutes; scales
on midbody below lateral line partially embedded and sharply lanceolate. No caudal peduncle
grooves. Vertebrae 10+6. Colour: in life, body grey-green dorsally, silvery grey to midline and silvery white
ventrally; adults with a series of 6 to 8 dusky roundish blotches above and below lateral line,
occasionally connected by narrow isthmus (blotches faint or absent in young); distal half of
dorsal-fin lobe abruptly and heavily pigmented; anal-fin lobe white or pale yellow, often interradial
membranes of several anterior rays black, especially in large individuals.
Size: Largest specimen examined 58.5 cm fork length.
Habitat, biology, and fisheries: Often travels in small schools and inhabits inshore waters from shallow
lagoons to offshore areas from the surface to depths of 100 m. Reported to be attracted by hitting the
surface of the water repeatedly in rapid succession. As a true of all members of the genus, the spines of
the first dorsal and anal fins are venomous (especially the latter) and capable of inflicting painful stings.
Feeds primarily on fishes and small crustaceans; young use specialized juvenile dentition to aggressively
remove scales and epidermal tissue from other fishes. Caught with drift set nets, gill nets, seines, and on
hook-and-line.
ocean, except no confirmed
records from the Persian Gulf;
widespread elsewhere in the
Indo-West Pacific, from Oki-
nawa to Australia and east-
ward to Hawaii.
Scomberoides tala (Cuvier, 1832)

Frequent synonyms / misidentifications: Chorinemus tala Cuvier, 1832; C. hainanensis Chu and
Cheng, 1958 / None.
FAO names: En - Barred queenfish; Fr - Sauteur carsia; Sp - Jurel carsia.
46 cm fork length
Diagnostic characters: Body oblong to elliptical, strongly compressed; snout and nuchal profile nearly
straight. Upper lip joined to snout at midline by a bridge of skin (frenum), except crossed by a shallow
groove in very young. Upper jaw extends slightly beyond margin of eye in adults. Lower jaw with 2
rows of teeth separated by a shallow groove, adults with teeth in inner row distinctly larger than
those in outer row. Gill rakers (excluding rudiments) 1 to 3 on upper limb and 7 to 11 on lower limb of
first gill arch (total 11 to 15). Two separate dorsal fins, the first with VI or VII short spines, depressible
into a shallow groove on dorsal midline, followed by I spine and 19 to 21 soft rays; anal fin with II detached
spines followed by 16 to 19 soft rays; posterior soft dorsal-and anal-fin rays consisting of semide-
tached finlets; bases of anal and second dorsal fins about equal in length; height of second dorsal-fin lobe
14.6 to 18% of fork length in specimens larger than 15 cm fork length; pelvic fins about equal in length or
shorter than pectoral fins. Lateral line only slightly irregular, weakly to moderately convex above pectoral
fins becoming straight posteriorly. No scutes; scales on midbody below lateral line partially embedded
and bluntly lanceolate. No caudal peduncle grooves. Vertebrae 10+16. Colour: in life, head and body
greenish grey dorsally, grey to silvery below; sides of adults with 4 to 8 vertically elongate plumbeous
blotches, most of which intersect lateral line; dorsal and anal fins dusky to dark and uniformly
pigmented; pectoral fins yellow and pelvic fins white in adults.
Habitat, biology, and fisheries: An inshore species whose biology is poorly known. As is true of all
members of the genus, the spines of the first dorsal and anal fins are venomous (especially the latter) and
capable of inflicting painful stings. Adults feed primarily on other fishes. Caught with drift set nets, gill nets,
seines, and on hook-and-line.
Ocean known only from Sri
Lanka, but probably also oc-
curs on the west coast of In-
dia. Elsewhere known from
the east coast of India, An-
daman Islands, Gulf of Thai-
land, Malaysia, Indonesia, the
Philippines, Australia, and the
Solomon Islands.
2734 Bony Fishes
Scomberoides tol (Cuvier, 1832)

Frequent synonyms / misidentifications: Chorinemus tol Cuvier, 1832 / None.
FAO names: En - Needlescaled queenfish; Fr - Sauteur leurre; Sp - Jurel saltarín.
45 cm fork length
Diagnostic characters: Body oblong to elliptical, strongly compressed; dorsal and ventral profiles nearly
equal, snout pointed with dorsal profile of head and nape slightly concave. Upper lip joined to snout at
midline by a bridge of skin (frenum), except crossed by a shallow groove in very young. Upper jaw extends
to posterior margin of pupil in adults. Lower jaw with 2 rows of teeth separated by a shallow groove,
adults with teeth in inner row equal in size to those in outer row. Gill rakers (excluding rudiments) 4 to
7 on upper limb and 17 to 20 on lower limb of first gill arch (total 21 to 26). Two separate dorsal fins, the
first with VI or VII short spines, depressible into a shallow groove on dorsal midline, followed by I spine
and 19 to 21 soft rays; anal fin with II detached spines followed by 18 to 20 soft rays; posterior soft
dorsal-and anal-fin rays consisting of semidetached finlets, bases of anal and second dorsal fins about
equal in length; height of second dorsal-fin lobe 7.7 to 11.2% of fork length in specimens larger than 15
cm fork length; pelvic fins about equal in length or shorter than pectoral fins. Lateral line only slightly
irregular, weakly to moderately convex above pectoral fins becoming straight posteriorly. No scutes; scales
on midbody below lateral line partially embedded, slender and needle-like. No caudal peduncle
grooves. Vertebrae 10+6. Colour: in life, body bluish dorsally, white ventrally; adults with 5 to 8 oval or
vertically oblong black blotches, the first 4 or 5 of which intersect lateral line (blotches faint or
absent in young); distal half of dorsal-fin lobe abruptly and heavily pigmented; anal-fin lobe usually
immaculate, white in life.
Habitat, biology, and fisheries: Usually found in small schools near the surface in coastal waters. As is
true of all members of the genus, the spines of the first dorsal and anal fins are venomous (especially the
latter) and capable of inflicting painful stings. Feeds primarily on fishes; young use specialized juvenile
dentition to aggressively remove scales and epidermal tissue from other fishes. Caught with drift set nets,
gill nets, and seines.
uted throughout most of the
Indian and western Pacific
oceans, from Japan to Aus-
tralia, and eastward to Fiji.
Selar boops (Cuvier, 1833)

FAO names: En - Oxeye scad; Fr - Sélar oeil de boeuf; Sp - Chicharro ojo buey.
22 cm fork length
(after Evermann and Seale, 1907)
Diagnostic characters: Body elongate and moderately compressed, with lower profile slightly more
convex than upper. Eye very large, shorter than snout length and with a well developed adipose
eyelid completely covering eye except for a vertical slit centred on pupil. Upper jaw moderately broad
posteriorly and extending to below anterior margin of pupil. Teeth small and recurved, upper jaw with a
narrow band, tapering posteriorly; lower jaw with an irregular single row. Gill rakers (including rudiments)
8 to 12 on upper limb and 25 to 29 on lower limb of first gill arch. Shoulder girdle (cleithrum) margin
with a deep furrow, a large papilla immediately above it and a smaller papilla near upper edge. Dorsal
fin with VIII spines, followed by I spine and 23 to 25 soft rays; anal fin with II spines separated from rest
of fin, followed by I spine and 19 to 21 soft rays; dorsal and anal fins without a detached terminal finlet;
pectoral fins shorter than head. Scales moderately small and cycloid (smooth to touch), covering body
except for a small area behind pectoral fins, scutes relatively large, maximum height contained 1.3 to
1.6 times in eye diameter; chord of the curved part of lateral line contained 2.1 to 3 times in straight
part (to caudal-fin base); scales in curved part of lateral line 21 to 24; 0 to 4 scutes in curved part;
straight part with no anterior scales and 37 to 46 scutes; total scales and scutes in lateral line (excluding
scales on caudal fin) 62 to 69. Vertebrae 10+14. Colour: in fresh fish, upper third of body and top of head
bluish green to green; lower two-thirds of body and head silvery or whitish; a narrow, yellowish stripe
may be present from edge of opercle to upper part of caudal peduncle; blackish areas above and
below pupil with a reddish area sometimes present; a small elongated, blackish opercular spot on edge
near upper margin; first dorsal fin dusky on margins with rest of fin clear; second dorsal, anal, and caudal
fins pale green with ducky margins; pectoral fins pale green to hyaline; pelvic fins white.
Size: Maximum length reported 26 cm total length; attains at least 24 cm fork length.
Habitat, biology, and fisheries: Co-occurs with Selar crumenophthalmus on continental shelf waters of
northern Australia where common between 20 and 100 m; also one confirmed record from off Portugal
where trawled in 366 to 439 m. Feeds primarily on planktonic or benthic invertebrates, including shrimps,
crabs, and foraminifers; also
on fishes. Caught on hook-
and-line, with beach seines,
trawls, purse seines, and
traps.
Distribution: Restricted to
the Indo-West Pacific except
for one confirmed record from
Sezimbra Bay, Portugal; else-
where recorded from the Phil-
ippines, Palau, Indonesia,
and northern Australia.
2736 Bony Fishes
Selar crumenophthalmus (Bloch, 1793)

Frequent synonyms / misidentifications: Trachurops crumenophthalmus (Bloch, 1793) / None.
FAO names: En - Bigeye scad; Fr - Sélar coulisou; Sp - Chicharro ojón.
27.9 cm fork length
Diagnostic characters: Body elongate and moderately compressed, with lower profile slightly more
convex than upper. Eye very large, shorter than snout length and with a well-developed adipose
eyelid completely covering eye except for a vertical slit centred on pupil. Upper jaw moderately broad
posteriorly and extending to below anterior margin of pupil. Teeth small and recurved, upper jaw with a
narrow band, tapering posteriorly; lower jaw with an irregular single row. Gill rakers (including rudiments)
9 to 12 on upper limb and 27 to 31 on lower limb of first gill arch. Shoulder girdle (cleithrum) margin
with a deep furrow, a large papilla immediately above it and a smaller papilla near upper edge. Dorsal
fin with VIII spines, followed by I spine and 24 to 27 soft rays; anal fin with II spines separated from rest
of fin, followed by I spine and 21 to 23 soft rays; dorsal and anal fins without a detached terminal finlet;
pectoral fins shorter than head. Scales moderately small and cycloid (smooth to touch), covering body
except for a small area behind pectoral fins, scutes relatively small, maximum height contained 2.1 to
2.9 times in eye diameter; chord of the curved part of lateral line contained 0.7 to 1.2 times in straight
part (to caudal-fin base); scales in curved part of lateral line 48 to 56; 0 to 4 scutes in curved part;
straight part with 0 to 11 anterior scales and 29 to 42 scutes; total scales and scutes in lateral line (excluding
scales on caudal fin) 84 to 94. Vertebrae 10+14. Colour: in fresh fish, upper third of body and top of head
metallic blue or bluish green; tip of snout dusky or blackish; lower two-thirds of body and head silvery or
whitish; a narrow, yellowish stripe may be present from edge of opercle to upper part of caudal
peduncle; blackish areas above and below pupil with a reddish area sometimes present; a small elongated,
blackish opercular spot on edge near upper margin; first dorsal fin dusky on margins with rest of fin clear;
second dorsal fin dusky over most of fin with dorsal lobe blackish; anal fin clear or slightly dusky along
base; caudal fin dusky with tip of upper lobe dark; pectoral fins clear or slightly dusky near base and with
a yellowish tint sometimes present; pelvic fins clear.
Size: Maximum length not well established, but unverified report of 60 cm standard length; documented
record of 27 cm standard length; commonly to about 24 cm fork length and 0.23 kg.
Habitat, biology, and fisheries: Found in small to large schools, mainly inshore or in shallow water; at
times over shallow reefs and in turbid
water, but ranging in depth to about 170 m.
Feeds primarily on planktonic or benthic
invertebrates, including shrimps, crabs,
and foraminifers; also on fishes. Caught on
hook-and-line, with beach seines, trawls,
purse seines, and traps.
Distribution: Worldwide in tropical and
subtropical marine waters; broadly distrib-
uted throughout the Indo-West Pacific from
Japan to Australia and eastward to Hawaii
where an important commercial species.
Selaroides leptolepis (Cuvier, 1833)

FAO names: En - Yellowstripe scad; Fr - Sélar à bande dorée; Sp - Chicharro banda dorada.
12 cm fork length
Diagnostic characters: Body elongate, oblong and compressed; dorsal and ventral profiles equally
convex. Eye diameter about equal to slightly smaller than snout length, with adipose eyelid moderately
developed on posterior half of eye. Upper jaw strongly protractile with posterior end of jaw concave
above, concave and produced below. Upper jaw without teeth; lower jaw with a series of minute teeth.
Gill rakers (including rudiments) 10 to 14 on upper limb and 27 to 32 on lower limb of first gill arch (total
40 to 46). Shoulder girdle (cleithrum) margin smooth, without papillae. Two separate dorsal fins, the
first with VIII spines, the second with I spine and 24 to 26 soft rays; anal fin with II detached spines followed
by I spine and 21 to 23 soft rays; spinous dorsal fin moderately high, longest spine height about equal
length of soft dorsal-fin lobe. Lateral line anteriorly with a moderate regular arch, with junction of curved
and straight parts below vertical from tenth to twelfth soft rays of second dorsal fin; chord of curved part
of lateral line longer than straight part of lateral line, contained 0.6 to 0.8 times in straight part; straight part
of lateral line with 13 to 25 scales followed by 24 to 29 relatively small scutes. Breast completely scaly.
Vertebrae 10+14. Colour: in life, metallic blue above, silvery white below, with a broad yellow stripe
from upper margin of eye to caudal peduncle; prominent black opercular spot encroaching onto
shoulder; dorsal, anal, and caudal fins pale to dusky yellow; pelvic fins white.
Size: A small species, largest specimens examined 18.5 cm fork length.
Habitat, biology, and fisheries: Occurs in large demersal schools over soft bottom habitats at depths
shallower than 50 m. Females attain sexual maturity by about 11 cm fork length. Ostracods, gastropods,
and euphausids are common prey but it also feeds on small fishes. Reported to feed at night. Caught with
seines and bottom trawls; also with traps and gill nets.
Ocean confined to coastal
waters from the Persian Gulf
to the Bay of Bengal; else-
cific known from the Gulf of
Thailand, Okinawa, Japan,
the Philippines, Indonesia,
and Australia.
2738 Bony Fishes
Seriola dumerili (Risso, 1810)

Frequent synonyms / misidentifications: Australia: Seriola simplex Ramsey and Ogilby, 1886; Japan:
S. purpurescens Temminck and Schlegel, 1844; Indonesia: S. tapeinometapon Bleeker, 1853 / None.
FAO names: En - Greater amberjack; Fr - Sériole couronée; Sp - Medregal coronado.
29 cm fork length
Diagnostic characters: Body elongate, moderately shallow and slightly compressed, with upper profile
slightly more convex than lower. Upper jaw very broad posteriorly (with very broad supramaxilla), and
extending to below posterior margin of pupil. Teeth minute, in a broad band in both jaws. Gill rakers
(excluding rudiments) decreasing in number with growth, 5 or 6 on upper limb, 15 or 16 on lower
limb, and 20 to 24 total on first gill arch at sizes 2 to 7 cm fork length; in specimens larger than
about 20 cm fork length, total gill rakers 11 to 19. Dorsal fin with VII spines (first spine minute or
missing in large fish), followed by I spine and 29 to 35 soft rays; anal fin with II detached spines (these
spines reduced or completely embedded in large fish), followed by I spine and 18 to 22 soft rays; in adults,
length of dorsal-fin lobe about equal to, or slightly longer than, pectoral fins and 13 to 18% of fork length;
anal-fin base contained 1.4 to 1.7 times in second dorsal-fin base; pelvic fins longer than pectoral fins. No
scutes; caudal peduncle with dorsal and ventral grooves present. First pterygiophore of anal fin
moderately curved in specimens larger than about 10 cm fork length. Vertebrae 10+14. Colour: bluish
grey or olivaceous above, sides and belly silvery white, sometimes brownish or with a pinkish tinge; usually
a darker nuchal bar through eye to dorsal-fin origin; often an amber stripe from eye along middle of body;
caudal fin dark or dusky with a lighter narrow posterior margin, extreme tip of lower caudal-fin lobe
sometimes light or white; small juveniles (2 to 17 cm fork length) with a prominent dark nuchal bar extending
from eye to dorsal-fin origin; 5 dark body bars, becoming irregularly split vertically, that do not extend into
the membranes of the second dorsal and anal fins, and a sixth bar at end of caudal peduncle; papillae
surrounding broad bands of teeth in both jaws not engorged with blood, appearance of teeth white.
Size: Maximum total length 188 cm; weight to 80.6 kg. Verified angling record of 150 cm fork length and
67.6 kg; commonly to 100 cm fork length and 15 kg.
Habitat, biology, and fisheries: Occurs both epibenthically and pelagically. Smaller fish (less than 3 kg)
may be taken in shallow water (less than 10 m). Larger fish usually occur in 18 to 72 m and have been,
taken as deep as 360 m; they are often found on reefs or at deep offshore holes or drop-offs, usually in
small or moderate-sized schools, but may be solitary. In some areas large individuals have reported to
cause cases of ciguatera when eaten. Small juveniles associate with floating plants or debris in oceanic
and offshore neritic waters. Known to feed primarily on fishes and also invertebrates, and to take live, dead,
and artificial bait. Caught with
pelagic and bottom trawls, as
well as on hook-and-line.
Distribution: I n t h e I n d o -
West Pacific known from Ja-
p an , A u s tralia, Johnston
Island, and Hawaii. Also
known from the Indian Ocean,
both sides of the Atlantic
Ocean, and the Mediterra-
nean Sea; absent from the
eastern Pacific Ocean.
Seriola hippos Günther, 1876

FAO names: En - Samson fish.
53.5 cm fork length

Diagnostic characters: Body elongate, moderately shallow and slightly compressed, with upper profile
slightly more convex than lower; juveniles with very blunt head, becoming less pronounced with age. Upper
jaw very broad posteriorly (with very broad supramaxilla), extending to below posterior margin of
pupil. Teeth minute, in a broad band in both jaws. Gill rakers (excluding rudiments) decreasing in
number with growth, 2 to 5 on upper limb, 9 to 12 on lower limb, and 11 to 17 total on first gill arch at
sizes 2 to 7 cm fork length; in specimens larger than about 20 cm fork length, total gill rakers
(excluding rudiments) 10 to 12. Dorsal fin with VIII spines (first spine minute or missing in large fish),
followed by I spine and 22 to 25 soft rays; anal fin with II detached spines (these spines reduced or
completely embedded in large fish), followed by I spine and 15 to 17 soft rays; in adults, length of dorsal-fin
lobe about equal to, or slightly longer than, pectoral fins and 14 to 18% of fork length; anal-fin base
contained 1.6 to 1.7 times in second dorsal-fin base; pelvic fins longer than pectoral fins. No scutes; caudal
peduncle with dorsal and ventral grooves present. First pterygiophore of anal fin moderately curved in
specimens larger than about 10 cm fork length. Vertebrae 10+14. Colour: in fresh adults, generally silvery
to bluish green above, sides and belly silvery white, on death changing to brownish, paler below; an amber
stripe from eye along middle of body may be present or absent; spinous and second dorsal fins dusky to
dark, and anal and caudal fins generally dark; juveniles with 5 broad, dark bands on body and a dark blotch
on head above and behind eye; papillae surrounding broad bands of teeth in both jaws often engorged
with blood giving teeth a red appearance.
Size: Maximum total length 173 cm; weight to 53 kg.
Habitat, biology, and fisheries: Occurs in small schools or pairs around rocky reefs or wrecks in depths
to 70 m. Feeds primarily on other fishes. It is an especially inquisitive fish and will frequently follow another
hooked fish to the side of the boat. An excellent sport fish mainly caught by anglers using handlines or
strong rods.
Distribution: A t em perate
s p e c i e s k n o w n fr o m th e
southern half of Australia,
Shark Bay, Western Australia
(25°21’S) to Moreton Bay,
Queensland (27°39’S), Nor-
folk Island, and northern New
Zealand.
2740 Bony Fishes
Seriola lalandi Valenciennes, 1833

Frequent synonyms / misidentifications: South Africa: Seriola pappei (Castelnau, 1861) and
S. banisteri Smith, 1959; Japan: S. auerovittata Temminck and Schlegel, 1844; Australia: S. grandis
Castelnau, 1872; eastern Pacific: S. dorsalis (Gill, 1864) / None.
FAO names: En - Yellowtail amberjack; Fr - Sériole chicard; Sp - Medregal rabo amarillo.
Diagnostic characters: Body elongate, moderately slender and slightly compressed, with upper and lower
profiles similar. Upper jaw moderately slender posteriorly (with moderately slender supramaxilla),
extending to below about anterior margin of pupil. Teeth minute, in a broad band in both jaws. Gill
rakers (excluding rudiments) 7 to 10 on upper limb and 15 to 20 on lower limb of first gill arch (total 22 to
29). Dorsal fin with VII spines (first spine minute or missing in large fish), followed by I spine and 30 to 35
soft rays; anal fin with II detached spines (these spines reduced or completely embedded in large fish),
followed by I spine and 19 to 22 soft rays; in adults, length of dorsal-fin lobe subequal or slightly
shorter than pectoral fins and 11 to 13.4% of fork length, anal-fin base contained 1.6 to 1.8 times in
second dorsal-fin base; pelvic fins longer than pectoral fins. No scutes; caudal peduncle with a slight
lateral fleshy keel on each side, and dorsal and ventral grooves present. First pterygiophore of anal fin
moderately curved in specimens larger than about 10 cm fork length. Vertebrae 11+14. Colour: usually
blue to olivaceous above, sides and belly silver to white, sometimes with a rosy tinge; a narrow bronze
stripe from snout extending through eye and along midside of body, darker on head, becoming yellow
posteriorly; spinous dorsal fin dusky; second dorsal fin and anal fin dusky olive basally, yellow distally;
caudal fin olivaceous yellow, pectoral and pelvic fins yellowish; juveniles (to about 20 cm fork length)
with many irregular dusky body bars, slightly wider than paler interspaces that do not extend into the
membranes of the second dorsal and anal fins.
Size: Maximum size not well established but known to attain at least 150 cm fork length and 50 kg. If
Seriola banisteri is conspecific, as believed, then the maximum verified size is 193 cm total length and
58.4 kg.
Habitat, biology, and fisheries: Congregates in large offshore shoals in depths of 50 m, but occasional
fish will venture into surf zones in pursuit of prey. Feeds primarily on small fishes and squids. An excellent
sport fish. Caught with seines, bottom trawls, and on hook-and-line.
Distribution: A circumglobal
species restricted to subtropi-
cal waters, and consisting of
a series of apparently disjunct
populations, many of which
until recently were considered
to represent distinct species.
In the Indo-Pacific known
from South Africa, Walters
Shoals, Amsterdam Island,
Japan, Australia, New Zea-
land, New Caledonia, Rapa,
Pitcairn Island, Easter Island,
and Hawaii.
Seriola rivoliana Valenciennes, 1833

Frequent synonyms / misidentifications: South Africa: Seriola songoro Smith, 1959; Mozambique:
S. bovinoculata Smith, 1959; eastern Pacific: S. colburni Evermann and Clark, 1928; western Atlantic:
S. falcata Cuvier, 1833 / None.
FAO names: En - Almaco jack; Fr - Sériole limon; Sp - Medregal limón.
39 cm fork length
Diagnostic characters: Body elongate, moderately deep and slightly compressed, with upper profile more
convex than lower. Upper jaw very broad posteriorly (with very broad supramaxiila), extending to
below posterior midpoint of pupil. Teeth minute in a broad band in both jaws. Gill rakers (excluding
rudiments) decreasing slightly in number with growth, 6 to 9 on upper limb, 18 to 20 on lower limb,
and 24 to 29 total on first gill arch at sizes of 2 to 7 cm fork length; in specimens larger than about
20 cm fork length, total gill rakers 22 to 26. Dorsal fin with VII spines (first spine minute or missing in
large fish), followed by I spine and 27 to 33 soft rays; anal fin with II detached spines (these spines reduced
or completely embedded in large fish), followed by I spine and 18 to 22 soft rays; in adults, length of
dorsal-fin lobe about 1.3 to 1.6 times longer than pectoral fins and 18 to 22% of fork length; anal-fin
base contained 1.5 to 1.6 times in second dorsal-fin base; pelvic fins longer than pectoral fins. No scutes;
caudal peduncle with dorsal and ventral grooves present. First pterygiophore of anal fin straight
in specimens larger than about 10 cm fork length. Vertebrae 10+14. Colour: brown or olivaceous to
bluish green above, sides and belly lighter, sometimes with brassy or lavender reflections; the nuchal bar
often persistent in adults, and a faint amber lateral stripe extending backward from eye frequently present;
caudal fin dark with a lighter narrow posterior margin; juveniles (from 2 to 18 cm fork length) with a dark
nuchal bar extending from the eye to the dorsal-fin origin; 6 dark body bars, each with a light narrow irregular
area through their centre vertically, that do not extend into the membranes of the second dorsal and anal
fins, and a seventh bar at the end of caudal peduncle; tip of anal fin white; pectoral, pelvic, and caudal fins
dusky.
Size: Maximum weight 24 kg, data on maximum length not available. Verified angling record of 103 cm
total length and 15.7 kg; commonly to 65 cm fork length and 4.6 kg.
Habitat, biology, and fisheries: Adults are pelagic and epibenthic, possibly more oceanic than other
Seriola species, and rarely caught in inshore waters. Juveniles are pelagic and occur offshore, under
floating plants and debris when small. Known to feed on fishes, to strike trolled artificial bait and
bottom-fished dead bait. Caught mainly with seines, bottom trawls, and on hook-and-line.
Distribution: Circumtropical in
marine waters, entering tem-
perate waters in some areas.
Broadly distributed in the In-
dian Ocean, except unre-
corded from the Persian Gulf;
Pacific from southern Japan,
Australia, and eastward to Ha-
waii.
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2742 Bony Fishes
Seriolina nigrofasciata (Rüppell, 1829)

Frequent synonyms / misidentifications: Zonichthys nigrofasciata (Rüppell, 1829) / None.
FAO names: En - Blackbanded trevally; Fr - Sériole amourez; Sp - Medregal listado.
40 cm fork length
Diagnostic characters: Body elongate,

moderately shallow and slightly com-
pressed, with head profile rising steeply
to interorbital, then becoming smoothly
convex to origin of spinous dorsal fin.
Upper jaw broadly rounded posteri-
orly (with moderately slender su-
pramaxilla), usually extending to
below posterior margin of eye. Teeth
minute, in a broad band in both jaws. Gill
rakers on first gill arch mostly con-
juvenile 12.5 cm fork length
sisting of rudiments, 4 to 10 total ele-
ments. Dorsal fin with VII or VIII short
spines (spines weak, especially several posterior ones which may be embedded), followed by I spine and
30 or 37 soft rays; anal fin with I detached spine (usually embedded), followed by I spine and 15 to 18 soft
rays; length of dorsal-fin lobe slightly longer than pectoral fins and 15 to 20% of fork length; anal-fin base
contained 2.1 to 2.3 times in second dorsal-fin base; pelvic fins longer than pectoral fins. No scutes; caudal
peduncle with a slight lateral fleshy keel on each side, and dorsal and ventral grooves present. First
pterygiophore of anal fin moderately curved in specimens larger than about 10 cm fork length; vertebrae
11+13. Colour: in life, head and body bluish grey to black dorsally, white to dusky below; young with 5 to
7 dark oblique bands and blotches on upper body that fade with age; spinous dorsal fin black; second
dorsal and anal fins dusky brown, tips of anterior lobes white, except in large adults; caudal and pelvic fins
yellowish brown to black.
Size: Maximum size not well established, reported to attain 70 cm total length. South African angling record
5.2 kg.
Habitat, biology, and fisheries: A non-schooling, generally solitary species seldom seen close to shore.
Favoured habitat appears to be offshore reefs near continental shelfs at depths of 20 to 150 m. Feeds on
demersal fishes, cephalo-
pods, and prawns. Caught
with seines, traps, gill nets,
and on hook-and-line; also by
spearing.
West Pacific known from
southern Japan, Indonesia,
Philippines, and Australia.
Trachinotus africanus Smith, 1967

FAO names: En - African pompano; Fr - Pompaneau africain; Sp - Pámpano africano.
61 cm fork length
Diagnostic characters: Body ovate in young to subovate in large adults and compressed; profile of snout
broadly rounded. Both jaws with small villiform bands of teeth (absent in adults); tongue with a narrow
patch of teeth, persisting to about 50 cm fork length. Gill rakers (including rudiments) 7 to 10 on upper limb
and 11 to 14 on lower limb of first gill arch. Two separate dorsal fins, the first with VI short spines (the
anterior spines often becoming completely embedded in large adults) followed by I spine and 21 to 23 soft
rays; anal fin with II detached spines (becoming embedded in large adults) followed by I spine and 19 to
21 soft rays; height of second dorsal-fin lobe 19 to 25% of fork length in specimens of 10 to 40 cm fork
length; pelvic fins shorter than pectoral fins. Lateral line only slightly irregular, weakly convex above
pectoral fins, becoming straight posteriorly. No scutes or caudal-peduncle grooves. First predorsal
(supraneural) bone shaped like an inverted “L” with the arm projecting anteriorly, this character is
easily observed by simple dissection along midline of nape; supraoccipital bone of skull thin and
blade-like in adults. Vertebrae 10+14. Colour: in life, adults bluish dorsally, ventral half of body silvery,
the 2 areas separated by an indistinct orange-pink stripe; second dorsal-fin base and fin lobe dusky, distal
half of fin yellow; anal fin bright yellow with fin lobe orange; caudal fin, pectoral, and pelvic fins yellowish;
juveniles silvery with yellow hues, dorsal- and anal-fin lobes dusky.
Size: Maximum verified size 83 cm fork length, 92 cm total length and about 9.2 kg; Durban Aquarium fish
reported to have exceeded 25 kg. South African spearfishing record 14.1 kg.
Habitat, biology, and fisheries: Prefers shallow coastal areas marked with reefs or rocky outcrops, but
is tolerant of low salinities. Adults usually encountered singly or in pairs, although shoals of up to 100
individuals may congregate during the spring breeding season (South Africa). Young make extensive use
of sheltered bays as nursery areas. Molluscs (rock mussels) are a preferred food source, but crabs are
also taken. Caught with traps, gill nets, and handlines; also by spearing.
Indian Ocean known from
South Africa northward to
Durban, Gulf of Aden, Oman,
and eastward to Karachi,
Pakistan. Elsewhere in the
Indo-West Pacific known from
a single confirmed record
from Bali, Indonesia.
2744 Bony Fishes
Trachinotus anak Ogilby, 1909

FAO names: En - Oyster pompano.
53.5 cm fork length

Diagnostic characters: Body ovate in young to subovate in large adults and compressed; in profile of
snout broadly rounded, in adults becoming nearly straight to interorbital region. Both jaws with bands of
small villiform teeth; tongue toothless (except 2 or 3 slender teeth rarely on small specimens). Gill
rakers (including rudiments) 3 to 6 on upper limb and 8 to 10 on lower limb of first gill arch. Two separate
dorsal fins, the first with VI short spines (the anterior spines often becoming completely embedded in large
adults), followed by I spine and 18 to 20 soft rays; anal fin with II detached spines (becoming embedded
in large adults), followed by I spine and 16 to 18 soft rays; height of second dorsal-fin lobe 30 to 37% of
fork length in specimens 10 to 40 cm fork length; pelvic fins shorter than pectoral fins. Lateral line only
slightly irregular, weakly convex above pectoral fins, becoming straight posteriorly. No scutes or caudal-
peduncle grooves. First predorsal bone (supraneural) shaped like an inverted “L” with the arm
projecting anteriorly, this character is easily observed by a simple dissection along midline of nape;
supraoccipital bone of skull thin and blade-like in adults; preorbital and nasal bones hyperossified
in specimens larger than about 30 cm fork length. Vertebrae 10+14. Colour: in life, head and body
generally silvery, greenish to bluish grey dorsally, paler below, large adults sometimes with body mostly
bronze or greenish golden; second dorsal and caudal fins dusky orange to nearly black, leading edges and
fin tips darkest; anal fin bright to dirty yellow, lobe without a brownish anterior margin; pelvic fins
pale to bright yellow; pectoral fins dark.
Size: Largest specimen examined 88 cm fork length and about 100 cm total length.
Habitat, biology, and fisheries: Inhabits shallow coastal waters. Diet consists primarily of benthic
molluscs. Caught primarily
with gill nets and by hook-
and-line.
from Australia (Darwin to
Moreton Bay, Queensland),
New Caledonia, and Taiwan
Province of China (Takao).
Trachinotus baillonii (Lacepède, 1801)

FAO names: En - Smallspotted dart; Fr - Pompaneau muscadin; Sp - Pámpano abotonado.
27.5 cm fork length
Diagnostic characters: Body elongate to ovate and strongly compressed; dorsal and ventral profiles
almost equally convex, snout blunt. Both jaws with bands of small villiform teeth; vomerine tooth patch
usually chevron-shaped and palatine tooth patch elongate; tongue without teeth. Gill rakers (including
rudiments) 7 to 13 on upper limb and 15 to 19 on lower limb of first gill arch. Two separate dorsal fins, the
first with VI short spines, followed by I spine and 21 to 25 soft rays; anal fin with II detached spines, followed
by I spine and 20 to 24 soft rays; dorsal-fin lobe consistently shorter than anal-fin lobe in specimens
larger than about 25 cm fork length; pelvic fins relatively short, their length contained 1.9 to 2.3
times in pectoral-fin length in specimens larger than about 25 cm fork length. Lateral line only slightly
irregular, weakly convex above pectoral fins, becoming straight posteriorly. No scutes or caudal-peduncle
grooves. Vertebrae 10+14. Colour: in life, adults silvery blue to grey above, silvery white below, sides
with 1 to 6 relatively small black spots (spots absent on fish smaller than about 10 to 13 cm fork
length), the number of spots generally increasing with age (spots individually and bilaterally
variable in number, location and intensity) in a longitudinal row on or near lateral line; in adults, all
spots typically equal to or smaller than eye diameter, and with about half of spot below lateral line;
caudal, second dorsal, and anal fins grey to black, the lobes usually darkest; pectoral fins pale to dusky
yellow; pelvic fins pale yellow to orange-yellow.
Size: Largest specimen examined 41 cm fork length, 53.5 cm total length, and about 0.9 kg.
Habitat, biology, and fisheries: Usually found in surge zone along sandy beaches. Feeds primarily on
crustaceans and worms. Caught with traps, gill nets, and handlines.
Ocean, except unrecorded
from the Persian Gulf; else-
cific known from southern
Japan southward to northern
and eastern Australia, and
eastward to the Marshall,
Line, and Society islands.
2746 Bony Fishes
Trachinotus blochii (Lacepède, 1801)

FAO names: En - Snubnose pompano; Fr - Pompaneau lune; Sp - Pámpano lunero.
55.5 cm fork length
broadly rounded, in adults becoming nearly straight in interorbital region. Both jaws with bands of small
villiform teeth; tongue toothless (except 2 or 3 slender teeth rarely on small specimens). Gill rakers
(including rudiments) 5 to 8 on upper limb and 8 to 10 on lower limb of first gill arch. Two separate dorsal
fins, the first with VI short spines (the anterior spines often becoming completely embedded in large adults),
followed by I spine and 18 to 20 soft rays; anal fin with II detached spines (becoming embedded in large
adults), followed by I spine and 16 to 18 soft rays; height of second dorsal-fin lobe 35 to 60% of fork length
in specimens of 10 to 40 cm fork length; pelvic fins shorter than pectoral fins. Lateral line only slightly
irregular, weakly convex above pectoral fins, becoming straight posteriorly. No scutes or caudal-peduncle
grooves. First predorsal (supraneural) bone shaped like an inverted teardrop- or oval-shaped, this
character is easily observed by simple dissection along midline of nape; supraoccipital bone of skull
thin and blade-like in adults. Vertebrae 10+14. Colour: in life, head and body generally silvery, blue-grey
above, paler below; large adults sometimes with most of body golden orange, especially snout and lower
half of body; second dorsal fin dark, lobe of fin dusky orange; anal fin dusky to dirty orange, lobe
with a brownish anterior margin; caudal fin dark to dirty orange, leading edges of fin darkest; pelvic
fins white to dirty orange; pectoral fins dark; juveniles silvery with pale fins, except lobes of median fins
and anterior half of pelvic fins brownish to dirty orange.
Size: Largest specimen examined 55.5 cm fork length, 65 cm total length, and about 5 kg.
Habitat, biology, and fisheries: Inhabits coral and rocky reef areas in shallow coastal waters. Feeds on
benthic molluscs. Caught mainly with traps, gill nets, and handlines.
Oc ean; elsewhere in the
Indo-West-Pacific from
southern Japan to northern
Australia, and eastward to
Samoa, Tonga, Marina, and
Marshall islands.
Trachinotus botla (Shaw, 1803)

Frequent synonyms / misidentifications: Trachinotus russelli (Cuvier, 1832) (see Remarks) / None.
FAO names: En - Largespotted dart; Fr - Pompaneau pierrot; Sp - Pámpano pastilla.
34.3 cm fork length
usually a small oval or a narrow triangle and palatine tooth patch short; tongue without teeth. Gill rakers
(including rudiments) 6 to 9 on upper limb and 11 to 15 on lower limb of first gill arch. Two separate
dorsal fins, the first with VI short spines, followed by I spine and 22 to 24 soft rays; anal fin with II detached
spines, followed by I spine and 19 to 21 soft rays; dorsal-fin lobe usually longer than anal-fin lobe in
specimens larger than about 25 cm fork length; pelvic fins relatively long, their length contained
1.5 to 1.7 times in pectoral-fin length in specimens larger than about 25 cm fork length. Lateral line
only sightly irregular, weakly convex above pectoral fins, becoming straight posteriorly. No scutes or
caudal-peduncle grooves. Vertebrae 10+14. Colour: in life, adults bluish black above, silvery below, sides
usually with 5 or 6 relatively large plumbeous spots (spots absent on fish smaller than about 10 to
13 cm fork length), the number of spots generally increasing with age (spots individually and bilaterally
variable in number, location, and intensity), in a longitudinal row on or near lateral line; in adults, only 1
of anterior 2 spots is above the pectoral fins and both are typically larger than eye diameter, all
spots broadly oval-shaped, and with at least two-thirds of spot above lateral line; second dorsal and
anal fins dusky to bluish black, the lobes usually darkest; caudal fin dusky with leading edges and most of
lobes blue-black; pectoral fins pale, upper two-thirds sometimes dark; pelvic fins white.
Size: Largest specimen examined 48 cm fork length, 61 cm total length, and 0.7 kg. South African angling
record 2.3 kg.
Habitat, biology, and fisheries: Inhabits shallow coastal waters, often in rough surf zone along sandy
beaches. Feeds mainly on crabs, mussels, and worms. Caught with seines, gill nets, on hook-and-line,
and by spearing.
Distribution: Apparently restricted to the Indian Ocean where found along the African coast from South
Africa to Kenya, Malagasy
Republic (west coast), India,
Sri Lanka, Myanmar, Java,
and Western Australia.
Remarks: Scomber b o tl a
Shaw, as originally proposed,
was a composite species;
validation and priority of name
is based on the action of Day
(1876:233, The fishes of In-
dia, Pt. 2) who as “first re-
viser” listed it as a synonym of
Trachinotus russelli.
2748 Bony Fishes
Trachinotus copperingi Günther, 1884

Frequent synonyms / misidentifications: Trachinotus velox Ogilby, 1908 / Trachinotus botla (Shaw,
1803) or T. russelli (Cuvier, 1832).
FAO names: En - Swallowtail dart.
25.4 cm fork length
(drawn by K. Ackoff)
varying from chevron-shaped to a large triangle and palatine tooth patch usually short; tongue without
teeth. Gill rakers (including rudiments) 5 to 9 on upper limb and 12 to 15 on lower limb of first gill
arch. Two separate dorsal fins, the first with VI short spines, followed by I spine and 23 to 25 soft rays;
anal fin with II detached spines, followed by I spine and 22 to 24 soft rays; dorsal-fin lobe consistently
shorter than anal-fin lobe in specimens larger than about 25 cm fork length; pelvic fins relatively
short, their length contained 1.7 to 2.2 times in pectoral-fin length in specimens larger than about
25 cm fork length. Lateral line only sightly irregular, weakly convex above pectoral fins, becoming straight
posteriorly. No scutes or caudal-peduncle grooves. Vertebrae 10+14. Colour: in life, adults bluish black
above, silvery below, sides usually with 6 or 7 relatively elongate plumbeous spots (spots absent on
fish smaller than about 10 to 13 cm fork length), the number of spots generally increasing with age
(spots individually and bilaterally variable in number, location, and intensity), in a longitudinal row on or
near lateral line; in adults, anterior 2 spots are above the pectoral fins and at least some spots larger
than eye diameter, all spots vertically elongate ovals, and with at least two-thirds of spot above
lateral line; second dorsal and anal fins dusky to bluish black, the lobes usually darkest; caudal fin dusky
with leading edges and most of lobes blue-black; pectoral fins pale, upper two-thirds sometimes dark;
pelvic fins white.
Size: Largest specimen examined 34.5 cm fork length.
Habitat, biology, and fisheries: Inhabits shallow coastal waters, often in rough surf zone along sandy
beaches. Feeds mainly on crabs, mussels, and worms. Caught with seines, gill nets, on hook-and-line,
and by spearing.
Distribution: Essentially an
Australian endemic species,
known only from the north-
east coast (19°S) to New
South Wales (34°S), and Lord
Howe Island.
Trachinotus mookalee Cuvier, 1832

FAO names: En - Indian pompano; Fr - Pompaneau indien; Sp - Pámpano indico.
58 cm fork length
broadly rounded, in adults becoming nearly straight to interorbital region. Both jaws with bands of small
villiform teeth, tongue with a narrow band of teeth, persisting to about 50 cm fork length. Gill rakers
(including rudiments) 5 to 8 on upper limb and 8 to 10 on lower limb of first gill arch. Two separate dorsal
fins, the first with VI short spines (the anterior spines often becoming completely embedded in large adults),
followed by I spine and 18 to 20 soft rays; anal fin with II detached spines (becoming embedded in large
adults), followed by I spine and 16 to 18 soft rays; height of second dorsal-fin lobe 24 to 34% of fork length
in specimens 10 to 40 cm fork length; pelvic fins shorter than pectoral fins. Lateral line only slightly irregular,
weakly convex above pectoral fin, becoming straight posteriorly. No scutes or caudal-peduncle grooves.
First predorsal bone (supraneural) shaped like an inverted “L” with the arm projecting anteriorly,
this character is easily observed by a simple dissection along midline of nape; supraoccipital bone
becoming broad and sausage-shaped in specimens larger than about 30 cm fork length. Vertebrae
10+14. Colour: in life, head and body generally silvery, greenish to bluish grey dorsally, paler below, large
adults sometimes with body mostly bronze or greenish golden; second dorsal and caudal fins dusky yellow,
leading edges and fin tips darkest; anal fin bright to dirty yellow, lobe without a brownish anterior
margin; pelvic fins pale yellow to white; pectoral fins dark; juveniles silvery with pale yellow fins, except
distal half of dorsal-fin lobe black.
Size: Largest specimen examined 77 cm fork length, 90 cm total length, and about 8.1 kg.
Habitat, biology, and fisheries: Inhabits shallow coastal waters. Caught with traps, gill nets, handlines,
and other artisanal gear.
Indian Ocean known from the
Gulf of Oman and the Persian
Gulf eastward to Sri Lanka;
Pacific known from Singa-
pore, Gulf of Thailand, and
Hong Kong.
2750 Bony Fishes
Trachurus declivis (Jenyns, 1842)

Frequent synonyms / misidentifications: None / Trachurus novaezelandiae Richardson, 1848.
FAO names: En - Greenback scad.
32 cm fork length
Diagnostic characters: Body elongate and nearly cylindrical, with upper and lower profiles about equal.
Eye moderate, with adipose eyelid well developed, usually covering most of eye except for a vertical oval
centred on pupil. Upper jaw moderately broad and extending to below anterior margin of eye. Teeth small,
in a single row in both jaws. Gill rakers (including rudiments) 11 to 16 on upper limb and 32 to 42 on lower
limb of first gill arch (total 53 to 55). Shoulder girdle (cleithrum) margin with a small furrow at upper
end, but no papillae present. Two separate dorsal fins, the first with VIII spines, followed by I spine and
29 to 35 soft rays; anal fin with II detached spines followed by I spine and 24 to 29 soft rays; pectoral fins
about equal to or slightly longer than head length. Scales in curved as well as straight part of lateral
line enlarged and scute-like (caution: in large Trachurus these scales may be obscured by an
overgrowth of smaller scales); total scales and scutes in lateral line 71 to 89, usually more than 75;
curved part of lateral line essentially parallel to axis of body for most of its length. Dorsal accessory
lateral line immediately below base of dorsal fin terminates below fifth to eleventh (usually seventh
to ninth) soft dorsal-fin ray. Vertebrae 10+14. Colour: greenish above, silvery below; caudal fin greyish;
a prominent black spot on upper margin of opercle.
Size: Attains 19 to 23 cm fork length by age of 2 years and reaches maturity by 3 or 4 years and about
27 cm fork length and 250 g; maximum recorded size 47 cm fork length at an age of 16 years.
Habitat, biology, and fisheries: Occurs in large schools and generally not present where surface water
temperature exceeds 17°C. Feeds mostly during the day on krill and other planktonic crustaceans or lantern
fishes (Myctophidae) at edge of continental shelf from surface to 500 m. Caught with demersal trawls and
purse seines.
Distribution: T e m p e r a t e
species known from southern
half of Australia, Shark Bay,
to Wide Bay, Queensland
(25°52’S), and from New Zea-
land.
Trachurus novaezelandiae Richardson, 1848

Frequent synonyms / misidentifications: Trachurus mccullochi Nichols, 1920 / Trachurus declivis
(Jenyns, 1842).
FAO names: En - Yellowtail horse mackerel.
23 cm fork length
Diagnostic characters: Body elongate and slightly compressed, with upper and lower profiles about equal.
Eye moderate, with adipose eyelid well developed, usually covering most of eye except for a vertical oval
centred on pupil. Upper jaw moderately broad and extending to below anterior margin of eye. Teeth small,
in a single row in both jaws. Gill rakers (including rudiments) 11 to 17 on upper limb and 35 to 45 on lower
limb of first gill arch (total 53 to 61). Shoulder girdle (cleithrum) margin with a small furrow at upper
end, but no papillae present. Two separate dorsal fins, the first with VIII spines, followed by I spine and
27 to 33 soft rays; anal fin with II detached spines followed by I spine and 22 to 29 soft rays; pectoral fins
about equal to or slightly longer than head length. Scales in curved as well as straight part of lateral
line enlarged and scute-like (caution: in large Trachurus these scales may be obscured by an
overgrowth of smaller scales); total scales and scutes in lateral line 67 to 81, rarely more than 76; curved
part of lateral line slanted for most of its length. Dorsal accessory lateral line immediately below
base of dorsal fin terminates below first to fifth (usually first or second) soft dorsal-fin ray. Vertebrae
10+14. Colour: brassy green above, silvery below (sometimes with iridescent brown vertical bands in fresh
specimens), with yellow tinges on scutes in straight lateral line; caudal and second dorsal fins yellowish;
a prominent black spot on upper margin of opercle.
Habitat, biology, and fisheries: Occurs in large schools on coastal reefs. Feeds mainly on planktonic
crustaceans. Caught with demersal trawls and purse seines.
Distribution: T e m p e r a t e
species known from southern
half of Australia, Exmouth
Gulf, Western Australia
(22°05’S) to Wide Bay,
Queensland (25°52’S), and
from New Zealand.
2752 Bony Fishes
Ulua aurochs (Ogilby, 1915)

Frequent synonyms / misidentifications: Citula aurochs Ogilby, 1915; Carangoides aurochs (Ogilby,
1915) / None.
FAO names: En - Silvermouth trevally.
, 16.5 cm fork length
Diagnostic characters: Body oval, strongly compressed; lower jaw protudes slightly beyond upper jaw.
Eye diameter larger than snout length. Both jaws with a very narrow band of small villiform teeth; tongue
with central band of villiform teeth. Gill rakers (including rudiments) 16 to 21 on upper limb and 37
to 41 on lower limb of first gill arch (total 54 to 61); gill rakers long, feather-like, and project into
mouth along side of tongue. Two separate dorsal fins, the first with VIII spines, the second with I spine
and 20 to 22 soft rays; anal fin with II detached spines followed by I spine and 17 or 18 soft rays; lobe of
second dorsal fin in young elongate, longer than head length; in large males (only), first 5 to 8 soft
dorsal-fin rays produced as filaments; anterior rays of anal fin slightly produced in both sexes but not
as filaments. Lateral line anteriorly with a moderate arch, with junction of curved and straight parts below
vertical between tenth and twelfth soft rays of second dorsal fin; chord of curved part of lateral line slightly
longer to slightly shorter than straight part of lateral line, contained 0.8 to 1.2 times in straight part; straight
part of lateral line with 0 to 5 scales followed by 26 to 38 scutes. Breast naked ventrally to behind pelvic
fins; laterally, naked area of breast extends diagonally to naked base of pectoral fins. Vertebrae 10+14.
Colour: in life, body blue to silvery blue dorsally, silvery below; dorsal surface of head dark blue to black;
diffuse dark blotch on upper margin of opercle; dorsal-fin filaments and inner rays of anal fin black; other
fins dusky to hyaline; cheeks, lower jaw, inside of mouth and tongue bright silver; juveniles with 5 or 6 dark
bands on body.
Size: Maximum reported fork length 26 cm.
Habitat, biology, and fisheries: Inhabits shallow coastal waters; in northern Autralian waters largely
restricted to depths shallower than 70 m. Feeds on small crustaceans and cephalopods. Caught primarily
with trawls.
Distribution: K n o w n f r o m
Australia, off Cape Leveque,
to Townsville, Queensland
(19°40’S), Papua New
Guinea, and Taiwan Province
of China.
Ulua mentalis (Cuvier, 1833)

Frequent synonyms / misidentifications: Caranx mentalis Cuvier, 1833; C. mandibularis Macleay,
1833; Ulua mandibularis (Macleay, 1833) / None.
FAO names: En - Longrakered trevally; Fr - Carangue démêloir; Sp - Jurel peinero.
- 26.5 cm fork length
Diagnostic characters: Body strongly compressed; dorsal profile more strongly convex than ventral
profile, lower jaw becoming prominent in large adults, with the angle of “chin” projecting beyond
upper jaw. Both jaws with a very narrow band of small villiform teeth; tongue without central band of
villiform teeth. Gill rakers (including rudiments) 23 to 27 on upper limb and 51 to 61 on lower limb
of first gill arch (total 74 to 86); gill rakers extremely long, feather-like, and project into mouth along
side of tongue. Two separate dorsal fins, the first with VIII spines, the second with I spine and 21 or 22
soft rays; anal fin with II detached spines followed by I spine and 17 or 18 soft rays; lobe of second dorsal
fin elongate in young, longer than head length becoming shorter than head in large adults. Lateral line
anteriorly with a moderate arch, with junction of curved and straight parts below vertical between tenth and
twelfth soft rays of second dorsal fin; chord of curved part of lateral line slightly longer to slightly shorter
than straight part of lateral line, contained 0.8 to 1.2 times in straight part; straight part of lateral line with
0 to 5 scales followed by 26 to 38 scutes. Breast naked ventrally to behind pelvic fins; laterally, naked area
of breast extends diagonally to naked base of pectoral fins. Vertebrae 10+14. Colour: in life, head and
body blue-green dorsally, silvery below; diffuse dark blotch on upper margin of opercle in large specimens,
faint or absent in young; adults with a dusky mark on cheek at angle of jaws; spinous dorsal and caudal
fins dusky to black; second dorsal and anal fins pale green except leading edges and distal margins of fins
dusky; cheeks, lower jaw, inside of mouth and tongue silver in small specimens; juveniles with 7 or 8 dark
bands on body.
Size: Largest specimen examined 75.5 cm fork length, 85 cm total length, and 6.4 kg; commonly to about
60 cm total length, reported to attain 100 cm total length.
Habitat, biology, and fisheries: Inhabits shallow coastal waters. Feeds on crustaceans and fishes.
Caught with gill nets, traps, handlines, and other types of artisanal gear.
Distribution: African coast
southward to Tanzania, north-
west coast of Malagasy Re-
public, Red Sea, Gulf of Aden,
Oman, Persian Gulf and Sri
Gulf of Thailand, Taiwan
Province of China, Indonesia,
the Philippines, Australia, and
Papua New Guinea.
2754 Bony Fishes
Uraspis helvola (Forster, 1801)

Frequent synonyms / misidentifications: Caranx helvolus (Forster, 1801); C. micropterus Rüppell,
1836; Leucoglossa candens Jordan, Evermann, and Wakiya, 1927 / See Remarks.
FAO names: En - Whitetongue jack; Fr - Carangue langue blanche; Sp - Jurel lengua blanca.
29 cm fork length
Diagnostic characters: Body oblong and compressed; dorsal profile strongly convex, ventral profile slightly
convex to isthmus, then nearly straight to origin of second dorsal fin; snout broadly rounded. Upper jaw extending
posteriorly to below anterior margin of eye to middle of eye. Teeth in both jaws small, pointed and usually
recurved, in a narrow band, an irregular row or a single row, becoming uniserial with increasing age. Gill rakers
(including rudiments) 5 to 8 on upper limb and 13 to 17 on lower limb of first gill arch (total 19 to 24). Two dorsal
fins, the first with VIII short, slender spines (the posterior 2 or 3 embedded and not apparent with growth)
followed by I spine and 25 to 30 soft rays; anal fin with II detached spines (embedded and not apparent in
all but very young) followed by I spine and 19 to 22 soft rays; lobe of second dorsal fin shorter than head
length; pelvic fins very long in young but becoming relatively shorter with age; chord of curved part of lateral
line contained 0.8 to 1.3 times in straight part of lateral line (ratio generally increasing with increasing size
of fish); curved part of lateral line with 48 to 66 scales and straight part of lateral line with 23 to 40 scutes;
in fish smaller than about 20 cm fork length some of the scutes with spines directed anteriorly (antrorse), the
number of antrorse spines decreasing with growth. Breast naked ventrally to origin of pelvic fins; laterally
naked area of breast separated from naked base of pectoral fins by a broad band of scales. Vertebrae
10+14. Colour: tongue, roof and floor of mouth white or cream-coloured, the rest blue-black; head dusky
to black, with a large, diffuse black opercular blotch; body dusky to black dorsally, lighter below and with
6 or 7 wide, dark bands and narrow pale interspaces (faint bands evident in specimens as large as 25 cm;
in specimens smaller than 10 cm bands may extend onto second dorsal and anal fins); second dorsal and
anal fins hyaline to black; the anal fin commonly, and the second dorsal fin occasionally with tips of the
falcate lobe and fin margin pale; pelvic fins generally black in specimens smaller than 10 cm fork length,
rapidly becoming pale whitish at larger sizes; caudal fin pale yellowish dusky with trailing edges and tips
of lobes blackish.
Size: Maximum fork length 46 cm.
Habitat, biology, and fisheries: A benthic-pelagic species inhabiting shelf waters along continental coasts
and around islands. Typically occurs in small demersal schools. Caught mainly with bottom trawls, seines,
traps, and on hook-and-line.
Distribution: Widely distributed in the Indo-West Pacific but rarely collected; verified records from the
WCP area are pending. Confirmed records are from the Red Sea, Arabian Sea, off Oman and Sri Lanka,
and Hawaii; also known from Ascension and St. Helena in the South Atlantic and from offshore islands in
the northeastern tropical Pacific Ocean.
Remarks: Adults of Uraspis helvola and
U. secunda are virtually impossible to dis-
tinguish although juvenile characters in-
volving allometric growth patterns suggest
that they are distinct species. If sub-
sequent studies indicated that these 2
nominal species are conspecific, the old-
est available name is Uraspis helvola.
Uraspis secunda (Poey, 1860)

Frequent synonyms / misidentifications: Caranx hullianus McCulloch, 1909; Uraspis reversa Jordan,
Evermann, and Wakiya, 1927; U. wakiyai Williams, 1961; U. cadenati Blache and Rossignol, 1962 / See
Remarks.
FAO names: En - Cottonmouth jack; Fr - Carangue cotor; Sp - Jurel volantín.
32.6 cm fork length
Diagnostic characters: Meristic and colour pattern characters are identical and broadly overlap those of
Uraspis helvola (see Remarks), and only the following major diagnostic characters (which apply to both
species) are repeated here. In fish smaller than about 20 cm fork length some of the scutes with
spines directed anteriorly (antrorse), the number of antrorse spines decreasing with growth. Breast
naked ventrally to origin of pelvic fins; laterally naked area of breast separated from naked base of
pectoral fins by a broad band of scales. Colour: tongue, roof and floor of mouth white or
cream-coloured, the rest blue-black.
Size: Maximum fork length 43.5 cm.
Habitat, biology, and fisheries: Apparently an oceanic species; at surface, pelagic, and benthic; solitary
and in small schools. Can produce a grunting sound when caught. Caught with trawls, purse seines,
longlines, and on hook-and-line.
Distribution: In the Indo-
West Pacific known from ?
South Africa and Tanzania,
Japan to Taiwan Province of
China, Northwest shelf and
Tasman Sea off Australia, and
Hawaii; also known from both
sides of the Atlantic Ocean
and from offshore islands in ? ?
the northeastern tropical Pa-
cific Ocean. Records of ? ?
Uraspis not identified to spe- ? ?
cies from Samoa, the Tonga
Islands, and Rapa may apply
to this species.
Remarks: Adults of U. secunda and U. helvola are virtually impossible to distinguish although juvenile
characters involving allometric growth patterns suggest that they are different species. If subsequent
studies indicated that these 2 nominal species are conspecific, the oldest available name is Uraspis
helvola.
2756 Bony Fishes
Uraspis uraspis (Günther, 1860)

Frequent synonyms / misidentifications: Caranx uraspis (Günther, 1860); C. quptae Chaudhuri, 1909;
Leucoglossa herklotsi Herre, 1932; Uraspis pectoralis Fowler, 1938 / See Remarks.
FAO names: En - Whitemouth jack; Fr - Carangue paia; Sp - Jurel paia.
23 cm fork length
Diagnostic characters: Body oblong and compressed; dorsal profile strongly convex, ventral profile
slightly convex to isthmus, then nearly straight to origin of second dorsal fin; snout broadly rounded. Upper
jaw extending posteriorly to below anterior margin of eye to middle of eye. Teeth in both jaws small, pointed
and usually recurved, in a narrow band, an irregular row, or a single row, becoming uniserial with increasing
age. Gill rakers (including rudiments) 5 to 7 on upper limb and 13 to 16 on lower limb of first gill arch (total
18 to 22). Two dorsal fins, the first with VIII short slender spines (the posterior 2 or 3 embedded and not
apparent with growth) followed by I spine and 25 to 30 soft rays; anal fin with II detached spines (embedded
and not apparent in all but very young) followed by I spine and 17 to 22 soft rays; lobe of second dorsal
fin shorter than head length; pelvic fins very long in young but becoming shorter with age; chord of curved
part of lateral line contained 0.7 to 0.98 times in straight part of lateral line (ratio generally increasing with
increasing size of fish but always less than 1); curved part of lateral line with 61 to 82 scales and straight
part of lateral line with 24 to 39 scutes; in fish smaller than about 20 cm fork length some of the scutes with
spines directed anteriorly (antrorse), the number of antrorse spines decreasing with growth. Breast naked
ventrally to origin of pelvic fins; laterally naked area of breast extends diagonally to naked base of
pectoral fins. Vertebrae 10+14. Colour: tongue, roof and floor of mouth white or cream-coloured,
the rest blue-black; body and head dusky to black dorsally, shading to dusky or pale grey ventrally;
juveniles and occasionally adults with 6 dusky or blue-black bars, which are about twice the width of pale
interspaces and much more distinct on ventral half of body below the level of lateral line (bars evident on
specimens as large as 22.5 cm fork length and absent on specimens as small as 12 cm); second dorsal
fin pale hyaline proximally with distal areas dusky posteriorly; anal fin yellow-white hyaline with distal areas
dusky, especially posteriorly; in small specimens of 8 to 13 cm fork length, pelvic fins whitish with distal
half of one third black, at sizes larger than 13 cm entire fin becoming pale white; caudal fin pale to dusky
with trailing edges dusky.
Size: Apparently a small species; maximum recorded size 28 cm fork length.
Habitat, biology, and fisheries: Forms
small demersal schools in depths from 50
to at least 130 m. Caught with bottom
trawls, longlines, and traps.
Distribution: In the western Indian Ocean
known from the Red Sea, Persian Gulf,
and in coastal waters eastward to Sri
Lanka; elsewhere in the Indo-West Pacific
known from the east coast of India, Straits
of Malacca, Gulf of Thailand, Hong Kong,
the Philippines, Indonesia, Australia,
Papua New Guinea, and Hawaii (Oahu).
click for next page

INDEX OF SCIENTIFIC AND VERNACULAR NAMES
Explanation of the System
Italics : Valid scientific names (genera and species).

Italics
: Synonyms (genera and species), misidentifications.
ROMAN : Family names.
ROMAN : Names of divisions, classes, subclasses, orders, suborders, and subfamilies.
Roman : FAO and local names.
2758 The Living Marine Resoursed of the Western Central Pacific
Agujón de quilla . .. . . . . . . . . . . . . . 2155

A Agujon needlefish .. . . . . . . . . . . . . . 2159
Abe’s flyingfish . . . . . . . . . . . . . . . . 2169 Agujón ocelado . .. . . . . . . . . . . . . . 2158
abei, Cheilopogon . . . . . . . . . . . . . . 2169 Agujón sable . . . .. . . . . . . . . . . . . . 2154
Ablabys taenianotus . . . . . . . . . . . . . 2309 Aiguille crocodile . .. . . . . . . . . . . . . . 2160
Ablennes hians . . . . . . . . . . . . . . . . 2154 Aiguille voyeuse . .. . . . . . . . . . . . . . 2159
Absalom radiatus . . . . . . . . . . . . . . . 2728 Aiguillette ocellée .. . . . . . . . . . . . . . 2158
Absent-minded scorpionfish . . . . . . . . . . 2347 aitha, Cephalopholis . . . . . . . . . . . . . 2540
Abudefduf biocellatus . . . . . . . . . . . . 2349 ajax, Carangoides . . . . . . . . . . . . . . . 2684
Abudefduf glaucus . . . . . . . . . . . . . . 2349 akaadsi, Decapterus kurroides . . . . . . . . 2718
Abudefduf leucozona . . . . . . . . . . . . . 2349 akaara, Epinephelus . . . . . . . . . 2485, 2498
Acanthaster . . . . . . . . . . . . . . . . . . 2602 Alalaua . . . . . . . . . . . . . . . . . . . . . 2599
Acanthistius . . . . . . . . . . . . . . . . . . 2550 alalaua, Priacanthus . . . . . . . . . . . . . 2599
acanthomoplate, Bovitrigla . . . . . . . . . . 2371 alata, Liza . . . . . . . . . . . . 2086, 2088-2089
acanthomoplate, Pterygotrigla . . . . 2360, 2371 albaiensis, Scorpaenodes . . . . . . . . . . . 2336
Acanthoplesiops . . . . . . . . . . . . . . . 2578 albescens, Remora . . . . . . . . . . . . . . 2652
Acanthoplesiops psilogaster . . . . . . . . . 2578 albimaculatus, Hirundichthys . . . . . . . . 2177
acanthopodus, Macrurocyttus . . . . . . . . 2258 albimarginata, Variola . . . . . . . . . . . . 2538
ACANTHURIDAE . . . . . . . . . . . . . . . 2260 albomarginata, Gracila . . . . . . . . . . . 2530
See Vol. 6 albopunctulatus, Epinephelus . . . . . . . . . 2504
Acanthurids . . . . .
. . . . . . . . . . . . . 2260 albus, Branchiostegus . . . . . . . . . 2641-2643
Acropoma . . . . . .
. . . . . . . . . . . . . 2436 alcocki, Mugil . . . . . . . . . . . . . . . . . 2093
ACROPOMATIDAE .
2436, 2444, 2611-2612, 2651 Aldrichetta . . . . . . . . . . . . . . . . . . 2069
Acropomatids . . . .
. . . . . . . . . . . . . 2436 Aldrichetta forsteri . . . . . . . . . . 2079, 2087
Acropora . . . . . .
. . . . . . . . . 2353, 2574 Alectis . . . . . . . . . . . . . . 2660-2661, 2667
acus melanotus, Tylosurus . . . . . . . . . . 2159 Alectis ciliaris . . . . . . . . . . . . . . . . . 2684
acus, Tylosurus . . . . . . . . . . . . . . . . 2160 Alectis crinitus . . . . . . . . . . . . . . . . . 2684
Acute-jawed mullet . . . . . . . . . . . . . . 2099 Alectis indica . . . . . . . . . . . . . . . . . 2685
acutus acutus, Hyporhamphus (Reporhamphus) Alepes . . . . . . . . . . . . . . . . . . . . . 2668
. . . . . . . . . . . . . . . . . . . . . . . 2194 Alepes apercna . . . . . . . . . . . . . . . . 2686
acutus pacificus, Hyporhamphus . . . . . . 2194 Alepes djedaba . . . . . . . . . . . . . . . . 2687
acutus, Carangoides . . . . . . . . . . . . . 2698 Alepes glabra . . . . . . . . . . . . . . . . . 2690
acutus, Hyporhamphus (Reporhamphus) acutus Alepes kalla . . . . . . . . . . . . . . . . . . 2688
. . . . . . . . . . . . . . . . . . . . . . . 2194 Alepes kleinii . . . . . . . . . . . . . . . . . 2688
Adioryx caudimaculatus . . . . . . . . . . . 2249 Alepes macrurus . . . . . . . . . . . . . . . . 2690
Adioryx ruber . . . . . . . . . . . . . . . . . 2251 Alepes malam . . . . . . . . . . . . . . . . . 2689
ADRIANICHTHYIDAE . .
2147, 2149, 2197, 2200 Alepes melanoptera . . . . . . . . . . . . . . 2689
Adrianichthyids . . . . . .
. . . . . . . . . . 2197 Alepes vari . . . . . . . . . . . . . . . 2687, 2690
adusta, Myripristis . . . .
. . . . . . . . . . 2236 alepis, Prionotus . . . . . . . . . . . . . . . 2373
Aeoliscus strigatus . . . .
. . . . . . . . . . 2282 Alfonsino . . . . . .. . . . . . . . . . . . . . 2221
aeolus, Sillago . . . . . .
. . . . . . 2617, 2621 Alfonsino besugo . . . . . . . . . . . . . . . 2222
Aeschrichthys goldiei . . .
. . . . . . . . . . 2080 Alfonsino palometón . . . . . . . . . . . . . . 2221
Aethaloperca rogaa . . . .
. . . . . . . . . . 2474 Alfonsinos . . . . . . . . . . . . . . . . . . . 2218
affinis, Caranx . . . . . .
. . . . . . . . . . 2692 Allanetta afra . . . . . . . . . . . . . . . . . 2136
affinis, Centroberyx . . .
. . . . . . . . . . 2223 Allanetta crenolepis . . . . . . . . . . . . . . 2134
affinis, Hyporhamphus (Reporhamphus) . . 2194 Allanetta duodecimalis . . . . . . . . . . . . 2122
affinis, Liza . . . . . . . . . . . . . . . . . . 2085 Allanetta klunzingeri . . . . . . . . . . . . . 2133
afra, Allanetta . . . . . . . . . . . . . . . . . 2136 Allanetta mugiloides . . . . . . . . . . . . . . 2131
African pompano . . . . . . . . . . . 2684, 2743 Allanetta ovalaua . . . . . . . . . . . . . . . 2135
African sailfin flyingfish . . . . . . . . . . . . 2178 Allanetta regina . . . . . . . . . . . . . . . . 2128
africanus, Trachinotus . . . . . . . . . . . . 2743 Allanetta valenciennei . . . . . . . . . . . . . 2138
afuerae, Decapterus . . . . . . . . . . . . . . 2720 Alligator pipefish . . . . . . . . . . . . . . . . 2276
Agonostoma diemensis . . . . . . . . . . . . 2079 allporti, Callanthias . . . . . . . . . . . . . . 2555
Agonostomus forsteri . . . . . . . . . . . . . 2079 Almaco jack . . . . . . . . . . . . . . . . . . 2741
Aguja imperial . . . . . . . . . . . . . . . . . 2159 Alón buho . . . . . . . . . . . . . . . . . . . 2286
Agujeta congaturí . . . . . . . . . . . . . . . 2191 Alón estrellado . . . . . . . . . . . . . . . . . 2289
Agujeta de Dussumier . . . . . . . . . . . . . 2189 Alón oriental . . . . . . . . . . . . . . . . . . 2287
Agujeta de Lutke . . . . . . . . . . . . . . . 2188 altissimus, Carangoides . . . . . . . . . . . . 2695
Agujeta de Quoy . . . . . . . . . . . . . . . 2190 altivelis, Calloplesiops . . . . . . . . . . . . 2578
Agujeta saltona . . . . . . . . . . . . . . . . 2186 altivelis, Cromileptes . . . . . . . . . . . . . 2484
Agujete manchada . . . . . . . . . . . . . . 2187 Amadais . . . . . . . . . . . . . . . . . . . . 2630
Index of Scientific and Vernacular names 2759
amaena, Myripristis . . . . . . . . . . . . . 2237 areolatus, Plectropomus . . . . . . . . . . . 2531

amarulus, Mugil . . . . . . . . . . . . . . . . 2106 argalus platyura, Platybelone . . . . . . . . 2155
AMBASSIDAE . . . . . . . . . . . . 2433, 2602 argentatus, Branchiostegus . 2641-2642-2643, 2646
Ambassids . . . . . . . . . . . . . . . . . . . 2434 argentea, Liza . . . . . . . . . . . . . 2087, 2092
Amberjacks . . . . . . . . . . . . . . . . . . 2659 argentifasciata, Sillago . . . . . . . . 2619, 2624
Amberstripe scad . . . . . . . . . . . . . . . 2721 argus, Calloplesiops . . . . . . . . . . . . . 2578
Ambiserrula jugosa . . . . . . . . . . . . . . 2391 argus, Cephalopholis . . . . . . . . . . 2476-2477
amblycephalus, Epinephelus . . . . . . . . . 2486 argyromus, Myripristis . . . . . . . . . . . . 2237
amboinensis, Serranus . . . . . . . . . . . . 2529 argyrotaenia, Haplocheilus . . . . . . . . . . 2138
Amphipoda . . . . . . . . . . . . . . . . . . 2276 Aristocratic bunquelovely . . . .
. . . . . . . 2441
amplexus, Corythoichthys . . . . . . . . . . 2273 armatus, Carangoides . . . . .
2691, 2693, 2702
amplisquamiceps, Neomerinthe . . . . 2322-2323 armatus, Caranx . . . . . . . . .
. . . . . . . 2693
anak, Trachinotus . . . . . . . . . . . . . . 2744 Armoured gurnards . . . . . . .
. . . . . . . 2359
analis, Sillago . . . . . . . . . . . . . 2618, 2623 Armoured sea robins . . . . . .
. . . . . . . 2359
Anchova de banco . . . . . . . . . . . . . . 2650 Arrhamphus . . . . . . . . . . .
. . . 2162, 2180
andamanense, Holocentrum . . . . . . . . . 2249 Arrhamphus sclerolepis krefftii . . . . . . . 2193
andertoni, Pterygotrigla . . . . . . . 2360, 2372 Arrhamphus sclerolepis sclerolepis . . . . . 2193
angusticeps, Cypselurus . . . . . . . . . . . 2173 Arrowfin bigeye . . . . . . . . . . . . . . . . 2600
ANOMALOPIDAE . . . . . . . . 2210-2212, 2215 ascensionis, Caranx . . . . . . . . . . . . . . 2713
Anomalops . . . . . . . . . . . . . . . . . . 2212 Asian sillago . . . . . . . . . . . . . . . . . . 2620
ANOPLOGASTRIDAE . . . . . . . . . 2210-2211 asiatica, Sillago . . . . . . . . . . . . . . . . 2620
antennata, Pterois . . . . . . . . . . . . . . 2329 asper, Atherion elymus . . . . . . . . . . . . 2118
ANTHIINAE . . . . . . . . . . . . . . . . . . 2442 asper, Rogadius . . . . . . . . . . . . . . . . 2412
Anthiines . . . . . . . . . . . . . . . . . . . . 2442 Assessor . . . . . . . . . . . . . . . . . . . . 2578
anticanus, Craterocephalus . . . . . . . . . . 2129 Assiculus punctatus . . . . . . . . . . . . . . 2568
Antigonia . . . . . . . . . . . . . . . . . . . 2261 asteroblepa, Leptosynanceia . . . . . . . . . 2318
Antipodocottus . . . . . . . . . . . . . . . . 2427 Atherin têtu . . . . . . . . . . . . . . . . . . 2125
Antipodocottus elegans . . . . . . . . 2425, 2428 Atherina bleekeri . . . . . . . . . . . . . . . 2138
Antipodocottus mesembrinus . . . . 2425, 2428 Atherina breviceps . . . . . . . . . . . . . . 2113
Antrorse spined gurnard . . . . . . . . . . . 2378 Atherina forskalii . . . . . . . . . . . . . . . 2125
Anyperodon leucogrammicus . . . . . . . . 2475 Atherina honoriae . . . . . . . . . . . . . . . 2130
aper, Capros . . . . . . . . . . . . . . . . . 2261 Atherina insularum . . . . . . . . . . . . . . 2124
apercna, Alepes . . . . . . . . . . . . . . . . 2686 Atherina lineata . . . . . . . . . . . . . . . . 2126
aphanes, Rhinopias . . . . . . . . . . . . . . 2332 Atherina morrisi . . . . . . . . . . . . . . . . 2125
aphrozoicus, Atherion elymus . . . . . . . . 2118 Atherina panatela . . . . . . . . . . . . . . . 2139
APHYONIDAE . . . . . . . . . . . . . . . . 2427 Atherina regina . . . . . . . . . . . . . . . . 2128
See Vol. 3 Atherina valenciennei . . . . . . . . . . . . . 2138
apia, Ocosia . . . . . . . .
. . . . . . . . . . 2324 Atherine de Barnes . . . . . . . . . . . . . . 2133
APISTINAE . . . . . . . . . . . . . . . . . . 2292 Athérine samoan . . . . . . . . . . . . . . . . 2136
Apistops caloundra . . . . . . . . . . . . . . 2309 Athérine tropicale . . . . . . . . . . . . . . . 2122
Apistus bottae . . . . . . .
. . . . . . . . . . 2352 Atherinichthys punctatus . . . . . . . . . . . 2131
Apistus carinatus . . . . .. . . . . . . . . . 2310 ATHERINIDAE . . . . . . . . . . . . 2070, 2113
APLOACTINIDAE . . . . . . . . 2292-2293, 2354 ATHERINIFORMES . . . . . . . . . . . 2109-2148
Aploactinids . . . . . . . .
. . . . . . . . . . 2293 atherinoides, Epigonus . . . . . . . . . . . . 2613
APLOCHEILICHTHYINAE . . . . . . . . . . 2200 Atherinomorus . . . . . . . . . . . . 2115, 2120
APLOCHEILIDAE . . . . . 2147, 2150, 2197, 2200 Atherinomorus balabacensis . . . . . 2119, 2128
Aplocheilus panchax . . . . . . . . . . . . . 2197 Atherinomorus capricornensis . 2120, 2125, 2137
Apogon cyanosoma . . . . . . . . . . . . . . 2608 Atherinomorus cylindricus . . . . . . . 2121-2122
APOGONIDAE . . . . . . 2433, 2436, 2602, 2651 Atherinomorus duodecimalis . . 2119, 2121-2122-
Apolectus niger . . . . . .. . . . . . . . . . 2729 2124, 2128
Aporops . . . . . . . . . .. . . . . . . . . . 2579 Atherinomorus eendrachtensis . . . . . . . . 2123
appendiculatus, Belone . . . . . . . . . . . . 2159 Atherinomorus endrachtensis . 2119-2123, 2126,
arabica, Pterygotrigla . . . . . . . . . . . . 2373 2129, 2131
arabica, Trigla . . . . . .. . . . . . . . . . 2373 Atherinomorus insularum . . . . . . . . . . 2124
archiepiscopus, Ostichthys . . . . . . . . . . 2246 Atherinomorus lacunosus . . . 2113, 2120, 2123-
archipelagicus, Hemiramphus . . . . . . . . 2186 2125, 2127, 2137
arcticeps, Cheilopogon . . . . . . . . . . . . 2169 Atherinomorus lineatus . . . . . . . 2123, 2126
arenarius, Platycephalus . . . . . . . . . . . 2407 Atherinomorus ogilbyi . . . . . 2120, 2123, 2125,
arenicola, Thysanophrys . . . . . . . . . . . 2417 2127, 2131
Areolate grouper . . . . . . . . . . . . . . . 2487 Atherinomorus reginae . . . . . . . . . . . . 2128
areolatus, Epinephelus . . . . . . . . 2487, 2492 Atherinosoma microstoma . . . . . . . . . . 2130
Atherinosoma tropicalis . . . . . . . . . . . 2137 barnesi, Hypoatherina . . . . . . . . . . . . 2133

Atherion elymus . . . . . . . . . . . . . . . 2118 Barracudas . . . . . . . .
. . . . . . . . . . 2151
Atherion elymus aphrozoicus . . . . . . . . . 2118 Barramundi . . . . . . . .
. . . . . . . . . . 2431
Atherion elymus asper . . . . . . . . . . . . 2118 Barred queenfish . . . . .
. . . . . . . . . . 2733
Atherion elymus freyi . . . . . . . . . . . . . 2118 Barredchest grouper . . . .
. . . . . . . . . . 2500
Atherion maccullochi . . . . . . . . . . . . 2118 Bartail flathead . . .
. . . .
. . . . . . . . . . 2409
atlanticus, Hoplostethus . . . . . . . . . . . 2215 batanensis, Beliops . . . .
. . . . . . . . . . 2578
atrisignis, Cheilopogon . . . . . . . . . . . . 2169 Bathysphyraenops simplex . . . . . . . . . . 2612
atropos, Atropus . . . . . . . . . . . . . . . 2691 BATRACHOIDIDAE . . . . . . . . . . . . . . 2427
Atropus atropos . . . . . . . . . . . . . . . . 2691 See also Vol. 3
Atropus atropus . . . . . . . . . . . . . . . . 2691 Bay sillago . . . . . . . . . . . . . . . . . . . 2624
atropus, Atropus . . . . . . . . . . . . . . . 2691 Bearded eelblennies . . . . . . . . . . . . . . 2586
Bearded ghoulfish . . . . . . . . . . . . . . . 2316
Atule djedaba . . . . . . . . . . . . . . . . . 2687
Bearded rougefish . . . . . . . . . . . . . . . 2350
Atule djeddaba . . . . . . . . . . . . . . . . 2687 Bearded silverside . . . . . . . . . . . . . . . 2118
Atule mate . . . . . . . . . . . . . . . . . . 2692 Bearhead flyingfish . . . . . . . . . . . . . . 2169
auerovittata, Seriola . . . . . . . . . . . . . 2740 Beauclaire aile blanc . . . . . . . . . . . . . 2600
AULOSTOMIDAE . . . . . 2263, 2265, 2277-2278 Beauclaire aile court . . . . . . . . . . . . . . 2599
Aulostomus chinensis . . . . . . . . . . . . 2277 Beauclaire aile juane . . . . . . . . . . . . . 2600
Aulostomus valentini . . . . . . . . . . . . . 2277 Beauclaire aile sagitté . . . . . . . . . . . . . 2600
aurantia, Cephalopholis . . . . . . . . . . . 2540 Beauclaire alalaua . . . . . . . . . . . . . . . 2599
aurantius, Cephalopholis . . . . . . . . . . 2482 Beauclaire bande blanc . . . . . . . . . . . . 2601
auratus, Branchiostegus . . . . . . . . . . . 2643 Beauclaire de roche . . . . . . . . . . . . . . 2595
aureus, Pseudochromis . . . . . . . . . . . . 2574 Beauclaire longe aile . . . . . . . . . . . . . 2599
auroch, Citula . . . . . . . . . . . . . . . . . 2752 Beauclaire miroir . . . . . . . . . . . . . . . . 2596
aurochs, Carangoides . . . . . . . . . . . . . 2752 Beauclaire Pacifique . . . . . . . . . . . . . . 2597
aurochs, Ulua . . . . . . . . . . . . . . . . . 2752 Beauclaire raie rouges . . . . . . . . . . . . . 2601
auroguttatus, Carangoides . . . . . . . . . . 2694 Beauclaire tache pourpre . . . . . . . . . . . 2598
auroguttatus, Caranx . . . . . . . . . . . . . 2694 beauforti, Cymbacephalus . . . . . . . . . . 2394
Australian barred tilefish . . . . . . . . . . . 2645 Beautimous wampeejaw . . . . . . . . . . . . 2440
Australian neglected halfbeak . . . . . . . . . 2195 Bécasse de mer . . . . . . . . . . . . . . . . 2280
australis, Callanthias . . . . . . . . . . . . . 2555 belanak, Mugil . . . . . . . . . . . . . . . . . 2091
australis, Callanthias platei . . . . . . . . . 2555 Beliops . . . . . . . . . . . . . . . . . . . . . 2578
australis, Centropogon . . . . . . . . . . . . 2311 Beliops batanensis . . . . . . . . . . . . . . 2578
australis, Mugil . . . . . . . . . . . . . . . . 2087 bellus, Dendrochirus . . . . . . . . . . . . . 2312
australis, Remora . . . . . . . . . . . . . . . 2652 Belone appendiculatus . . . . . . . . . . . . 2159
awoara, Epinephelus . . . . . . . . . . . . . 2488 Belone ciconia . . . . . . . . . . . . . . . . . 2157
axillaris, Mugil . . . . . . . . . . . . . . . . 2107 Belone leiuroides . . . . . . . . . . . . . . . 2156
axillaris, Myripristis . . . . . . . . . . . . . 2242 Belone leiurus . . . . . . . . . . . . . . . . . 2157
Belone platyura . . . . . . . . . . . . . . . . 2155
B BELONIDAE . . . . . . . . . . . 2151, 2180, 2278
Backspot flyingfish . . . . . . . . . . . . . . 2170 BELONIFORMES . . . . . . . . . . . . 2149-2196
baillonii, Trachinotus . . . . . . . . . . . . 2745 Bembradium . . . . . . . . . . . . . . . . . 2383
bairdianus, Sphyraenops . . . . . . . . . . . 2613 Bembras . . . . . . . . . . . . . . . . . . . . 2383
bajad, Carangoides . . . . . . . . . . . . . . 2694 BEMBRIDAE . . . . . . . 2360, 2383, 2386, 2425
Balabac Island silverside . . . . . . . . . . . 2119 berndti, Myripristis . . . . . . . . . . . . . . 2238
balabacensis, Atherinomorus . . . . 2119, 2128 BERYCIDAE . . . . . . . . 2214, 2216, 2218, 2591
Bali halfbeak . . . . . . . . . . . . . . . . . . 2194 BERYCIFORMES . . . . . . . . 2210-2256, 2551
balinensis, Hyporhamphus (Reporhamphus) . 2194 Beryx . . . . . . . . . . . . . . . . . 2218, 2222
Beryx commun . . . . . . . . . . . . . . . . . 2221
ballieu, Sebastapistes . . . . . . . . . . . . . 2345
Banded grouper . . . . . . . . . . . . . . . . 2486 Beryx decadactylus . . . . . . . . . . . 2221-2222
Banded pipefish . . . . . . . . . . . . . . . . 2274 Beryx long . . . . . . . . . . . . . . . . . . . 2222
Banded scad . . . . . . . . . . . . . . . . . 2688 Beryx mollis . . . . . . . . . . . . . . . . . . 2222
Banggai cardinalfish . . . . . . . . . . . . . . 2609 Beryx splendens . . . . . . . . . . . . . 2221-2222
banisteri, Seriola . . . . . . . . . . . . . . . 2740 biaculeatus, Syngnathoides . . . . . . . . . . 2276
barbata, Tetraroge . . . . . . . . . . . . . . 2350 bicoarctatus, Trachyrhamphus . . . . . . . . 2276
Barbel flyingfish . . . . . . . . . . . . . . . . 2176 bicolor, Rondeletia . . . . . . . . . . . . . . 2204
BARBOURISIIDAE . . . . . . . . 2204-2205-2206 Bigeye gurnard . . . . . . . . . . . . . . . . 2377
Barcheek trevally . . . . . . . . . . . . . . . 2707 Bigeye scad . . . . . . . . . . . . . . . . . . 2736
Barchin scorpionfish . . . . . . . . . . . . . . 2346 Bigeye trevally . . . . . . . . . . . . . . . . . 2716
Barko barko . . . . . . . . . . . . . . . . . . 2644 Bigeyes . . . . . . . . . . . . . . . . . . . . 2590
Barnes’ silverside . . . . . . . . . . . . . . . 2133 Bigmouth scorpionfish . . . . . . . . . . . . . 2340
Bignose fishes . . . . . . . . . . . . . . . . . 2209 bontoides, Epinephelus . . . . . . . . . . . . 2542

Bigscale scorpionfish . . . . . . . . . . . . . 2322 Bony flyingfish . . . . . . . . . . . . . . . . . 2177
Bigscales . . . . . . . . . . . . . . . . . . . 2201 boops, Scombrops . . . . . . . . . . . . . . . 2651
bilobatus, Epinephelus . . . . . . . . . . . . 2542 boops, Selar . . . . . . . . . . . . . . . . . . 2735
biocellatus, Abudefduf . . . . . . . . . . . . 2349 borbonicus, Mugil . . . . . . . . . . . . . . . 2107
biocellatus, Dendrochirus . . . . . . . . . . 2313 borbonicus, Myripristis . . . . . . . . . . . . 2241
bipinnulata, Elagatis . . . . . . . . . . . . . 2724 borneensis, Mugil . . . . . . . . . . . . . . . 2088
Black jack . . . . . . . . . . . . . . . . . . . 2713 bosschei, Cymbacephalus . . . . . . . . . . 2395
Black pomfret . . . . . . . . . . . . . . . . . 2729 botche, Myripristis . . . . . . . . . . . . . . 2239
Black scorpionfish . . . . . . . . . . . . . . . 2314 botla, Scomber . . . . . . . . . . . . . . . . 2747
Black-barred halfbeak . . . . . . . . . . . . . 2187 botla, Trachinotus . . . . . . . . . . . 2747-2748
Blackbanded flathead . . . . . . . . . . . . . 2411 bottae, Apistus . . . . . . . . . . . . . . . . . 2352
Blackbanded trevally . . . . . . . . . . . . . 2742 bovinoculata, Seriola . . . . . . . . . . . . . 2741
Blackbreasted pipefish . . . . . . . . . . . . 2274 Bovitrigla . . . . . . . . . . . . . . . . . . . 2360
Blackdotted grouper . . . . . . . . . . . . . . 2525
Blackfin scad . . . . . . . . . . . . . . . . . 2689
Bovitrigla acanthomoplate . . . . . . . . . . 2371
Blackfin squirrelfish . . . . . . . . . . . . . . 2244 Bovitrigla leptacanthus . . . . . . . . . . . . 2375
Blackfinned halfbeak . . . . . . . . . . . . . 2195 brachiusculus, Grammicolepis . . . . . . . . 2260
Blackfooted firefish . . . . . . . . . . . . . . 2326 Brachybembras . . . . . . . . . . . . . . . . 2383
Blacksaddle grouper . . . . . . . . . . . . . 2544 Brachypterois serrulatus . . . . . . . . . . . 2310
Blacksaddled coralgrouper . . . . . . . . . . 2532 brachypterus, Dendrochirus . . . . . . . . . 2313
Blacksea wasp . . . . . . . . . . . . . . . . 2350 brachypterus, Parexocoetus . . . . . . . . . 2178
Blackspot waspfish . . . . . . . . . . . . . . 2318 BRANCHIOSTEGIDAE . . . . . . . . 2614, 2630
Blackspotted gurnard . . . . . . . . . . . . . 2373 BRANCHIOSTEGINAE . . . . . . . . . . . . 2630
Blacktip grouper . . . . . . . . . . . . . . . . 2499 Branchiostegus . . . . . . 2630, 2632-2633, 2642
Blacktip soldierfish . . . . . . . . . . . . . . 2239 -2643, 2645
Blacktip trevally . . . . . . . . . . . . . . . . 2711 Branchiostegus albus . . . . . . . . . . 2641-2643
Blanchardia maculata . . . . . . . . . . . . 2586 Branchiostegus argentatus . 2641-2642-2643, 2646
Blanquillo . . . . . . . . . . . . . . . . . . . 2647 Branchiostegus auratus . . . . . . . . . . . 2643
Blanquillo de blanco . . . . . . . . . . . . . . 2641 Branchiostegus doliatus . . . . . . . . 2643-2644
Blanquillo de plata . . . . . . . . . . . . . . . 2642 Branchiostegus japonicus 2632, 2641-2643-2644
Blanquillo de roja . . . . . . . . . . . . . . . 2643 Branchiostegus sawakinensis . . . . . 2643-2644
Blanquillos . . . . . . . . . . . . . . . . . . . 2630 Branchiostegus semifasciatus . . . . . . . . 2632
bleekeri, Atherina . . . . . . . . . . . . . . . 2138 Branchiostegus serratus . . . . 2630, 2645-2646
bleekeri, Epinephelus . . . . . . . . . . . . 2489 Branchiostegus wardi . . . . . . 2642, 2644-2646
BLENNIIDAE . . . . . . . . . . . . . . . . . 2587 Brassy trevally . . . . . . . . . . . . . . . . . 2715
See also Vol. 6 breve, Melapedalion . . . . . . . . . . . . . 2195
blochii, Priacanthus . . . . . . . . . . . . . 2599 brevicarinatus, Caranx . . . . . . . . . . . . 2707
blochii, Trachinotus . . . . . . . . . . . . . 2746 breviceps, Atherina . . . . . . . . . . . . . . 2113
Blotched flathead . . . . . . . . . . . . . . . 2397 breviceps, Mugil . . . . . . . . . . . . . . . . 2098
Blotcheye soldierfish . . . . . . . . . . . . . 2238 brevifrons, Scorpaenopsis . . . . . . . . . . 2340
Blotchfin scorpionfish . . . . . . . . . . . . . 2340 brevipinnis, Prognichthys . . . . . . . . . . 2178
Bludger . . . . . . . . . . . . . . . . . . . . 2701 brevirostris, Malacanthus . . . . . . . 2647-2648
Blue eyes . . . . . . . . . . . . . . . . . . . 2109 Brick soldierfish . . . . . . . . . . . . . . . . 2237
Blue tilefish . . . . . . . . . . . . . . . . . . 2648 Bridled grouper . . . . . . . . . . . . . . . . 2502
Blue trevally . . . . . . . . . . . . . . . . . . 2699 Broadband flathead . . . . . . . . . . . . . . 2405
Blueback blue-eye . . . . . . . . . . . . . . . 2110 Broadbared lionfish . . . . . . . . . . . . . . 2329
Bluebarred dottyback . . . . . . . . . . . . . 2573 Broadhead flathead . . . . . . . . . . . . . . 2417
Bluefin gurnard . . . . . . . . . . . . . . . . 2364 Broadmouthed mullet . . . . . . . . . . . . . 2090
Bluefin trevally . . . . . . . . . . . . . . . . . 2714 Brochete de mer . . . . . . . . . . . . . . . . 2432
Bluefish . . . . . . . . . . . . . . . . . . . . 2650 brodiei, Howella . . . . . . . . . . . . . . . 2612
Bluefishes . . . . . . . . . . . . . . . . . . . 2650 Broussonnet’s mullet . . . . . . . . . . . . . 2095
Bluelined hind . . . . . . . . . . . . . . . . . 2478 broussonneti, Mugil . . . . . . . . . . . . . 2095
Bluespot mullet . . . . . . . . . . . . . . . . 2107 Brown dottyback . . . . . . . . . . . . . . . . 2574
Bluespotted cornetfish . . . . . . . . . . . . . 2279 Brownback trevally . . . . . . . . . . . . . . 2708
Bluespotted dottyback . . . . . . . . . . . . . 2568 Brownbanded pipefish . . . . . . . . . . . . . 2273
Bluespotted hind . . . . . . . . . . . . . . . 2477 Brownmarbled grouper . . . . . . . . . . . . 2501
Bluespotted trevally . . . . . . . . . . . . . . 2710 Brownmargined flathead . . . . . . . . . . . . 2392
Bluestripe pipefish . . . . . . . . . . . . . . . 2275 Brownspot bigeye . . . . . . . . . . . . . . . 2597
Bluetail mullet . . . . . . . . . . . . . . . . . 2102 Brownspot squirrelfish . . . . . . . . . . . . . 2250
Boarfishes . . . . . . . . . . . . . . . . . . . 2261 Brownspotted grouper . . . . . . . . . . . . . 2492
boelang, Cephalopholis . . . . . . . . . . . 2478 bruneus, Epinephelus . . . . . . . . . . . . . 2490
boenak, Cephalopholis . . . . . . . . 2478, 2540
bruuni, Oxyporhamphus convexus . . . . . 2192 Canthigaster valentini . . . . . . . . . . . . 2532

bucculentus, Caranx . . . . . . . . . . . . . 2710 Cape blue-eye . . . . . . . . . . . . . . . . . 2112
buchanani, Valamugil . . . . . . . . 2102, 2107 CAPRELLIDAE . . . . . . . . . . . . . . . . 2276
Bulbheaded scorpionfish . . . . . . . . . . . 2341 capreoli, Craterocephalus . . . . . . 2129, 2131
Bullheaded gurnard . . . . . . . . . . . . . . 2371 Capricorn silverside . . . . . . . . . . . . . . 2120
Bullrout . . . . . . . . . . . . . . . . . . . . 2324 capricornensis, Atherinomorus 2120, 2125, 2137
Bumphead scorpionfish . . . . . . . . . . . . 2335 capricornensis, Pranesus . . . . . . . . . . . 2120
Bumpnose trevally . . . . . . . . . . . . . . . 2702 CAPROIDAE . . . . . . . . . . . . . . 2260-2261
Bunquelovelies . . . . . . . . . . . . . . . . 2438 Capros aper . . . . . . . . . . . . . . . . . . 2261
burra, Sillago maculata . . . . . . . . . . . . 2621 CARACANTHIDAE . . . . . . . 2292-2293, 2353
Burrowfishes . . . . . . . . . . . . . . . . . 2630 Caracanthids . . . . . . . . . . . . . . . . . . 2353
burrus, Sillago . . . . . . . . . . . . 2617, 2621 Carangichthys dinema . . . . . . . . . . . . 2697
Butterfly scorpionfish . . . . . . . . . . . . . 2312 Carangichthys oblongus . . . . . . . . . . . 2705
bynoensis, Scorpaena . . . . . . . . . . . . 2346 CARANGIDAE . . . . . . . . . 2650, 2655, 2659
BYTHITIDAE . . . . . . . . . . . . . 2427, 2587 Carangids . . . . . . . . . . . . . . . . . . . 2661
See also Vol. 3 Carangoides . . . . . . . . . . . 2661, 2668, 2695
Carangoides acutus . . . . . . . . . . . . . . 2698
C Carangoides ajax . . . . . . . . . . . . . . . 2684
Cabete aleta manchada . . . . . . . . . . . . 2368 Carangoides altissimus . . . . . . . . . . . . 2695
Cabete tintero . . . . . . . . . . . . . . . . . 2373 Carangoides armatus . . . . . . 2691, 2693, 2702
Cabrilla venda . . . . . . . . . . . . . . . . . 2486 Carangoides aurochs . . . . . . . . . . . . . 2752
cadenati, Uraspis . . . . . . . . . . . . . . . 2755 Carangoides auroguttatus . . . . . . . . . . . 2694
caeruleomaculatus, Mugil . . . . . . . . . . 2107 Carangoides bajad . . . . . . . . . . . . . . 2694
caeruleopinnatus, Carangoides . . . . . . . 2695 Carangoides caeruleopinnatus . . . . . . . . 2695
caeruleopinnatus, Caranx . . . . . . . . . . 2695 Carangoides chrysophryoides . . . . . . . . . 2696
caeruleopunctatus, Epinephelus . . . 2491, 2514 Carangoides chrysophrys . . . . . . . . . . . 2696
CAESIONIDAE . . . . . . . . . . . . . . . . 2633 Carangoides ciliarius . . . . . . . . . . . . . 2693
See also Vol. 6 Carangoides dasson . . . . . . . . . . . . . . 2698
calcarifer, Lates . . . . .
. . . . . . 2429, 2431 Carangoides dinema . . . . . . . . . . . . . 2697
Caledonian stinger . . . .
. . . . . . . . . . 2315 Carangoides equula . . . . . . . . . . . . . . 2698
caledonicus, Inimicus . .
. . . . . . . . . . 2315 Carangoides ferdau . . . . . . . . . . 2699, 2706
Callanthias . . . . . . . .
. . . . . . . . . . 2579 Carangoides formosanus . . . . . . . . . . . 2695
Callanthias allporti . . . .
. . . . . . . . . . 2555 Carangoides fulvoguttatus . . . . . . . 2700-2701
Callanthias australis . . .
. . . . . . . . . . 2555 Carangoides gymnostethoides . . . . . . . . 2701
Callanthias crosnieri . . .
. . . . . . . . . . 2556 Carangoides gymnostethoides evermanni . . 2706
Callanthias japonicus . .
. . . . . . . . . . 2555 Carangoides gymnostethus . . . 2694, 2700-2701
Callanthias platei . . . . .
. . . . . . . . . . 2555 Carangoides hedlandensis . . . 2691, 2693, 2702
Callanthias platei australis . . . . . . . . . . 2555 Carangoides hemigymnostethus . . . . . . . 2699
CALLANTHIIDAE . . . . . 2443, 2553, 2557, 2579 Carangoides humerosus . . . . . . . . . . . 2703
CALLIONYMIDAE . . . . . . . . . . 2423, 2426 Carangoides ire . . . . . . . . . . . . . . . . 2708
See also Vol. 6 Carangoides jordani . . . . . . . . . . . . . 2706
Calloplesiops . . . . . . . . . . . . . . . . . 2578 Carangoides malabaricus . . . . . . 2704, 2709
Calloplesiops altivelis . . . . . . . . . . . . . 2578 Carangoides nitidus . . . . . . . . . . . . . . 2706
Calloplesiops argus . . . . . . . . . . . . . . 2578 Carangoides oblongus . . . . . . . . . . . . 2705
caloundra, Apistops . . . . . . . . . . . . . 2309 Carangoides orthogrammus . . . . . . . . . 2706
Camouflage grouper . . . . . . . . . . . . . 2516 Carangoides plagiotaenia . . . . . . . . . . 2707
canadum, Rachycentron . . . . . . . 2652, 2655 Carangoides praeustus . . . . . . . . . . . . 2708
candens, Leucoglossa . . . . . . . . . . . . . 2754 Carangoides rectipinnis . . . . . . . . . . . . 2704
Candil acorazado . . . . . . . . . . . . . . . 2246
Candil chocolate . . . . . . . . . . . . . . . . 2250
Carangoides rhomboides . . . . . . . . . . . 2704
Candil de aleta negra . . . . . . . . . . . . . 2244
Carangoides talamparoides . . . . . 2704, 2709
Candil de lomo manchado . . . . . . . . . . 2241 Carangoides uii . . . . . . . . . . . . . . . . 2695
Candil guru-guru . . . . . . . . . . . . . . . 2243 Carangue à dos brun . . . . . . . . . . . . . 2708
Candil ojo manchado . . . . . . . . . . . . . 2238 Carangue aile blanche . . . . . . . . . . . . . 2698
Candil piñón . . . . . . . . . . . . . . . . . . 2242 Carangue aile bleue . . . . . . . . . . . . . . 2714
Candil platero . . . . . . . . . . . . . . . . . 2249 Carangue balo . . . . . . . . . . . . . . . . . 2701
Candil rechoncho . . . . . . . . . . . . . . . 2239 Carangue bronze . . . . . . . . . . . . . . . 2715
Candil rubí . . . . . . . . . . . . . . . . . . . 2251 Carangue cavallys . . . . . . . . . . . . . . . 2709
Candil sable . . . . . . . . . . . . . . . . . . 2252 Carangue cocole . . . . . . . . . . . . . . . . 2711
Candil samara . . . . . . . . . . . . . . . . . 2245 Carangue côtière . . . . . . . . . . . . . . . 2695
Candil sombreado . . . . . . . . . . . . . . . 2236 Carangue cotor . . . . . . . . . . . . . . . . 2755
canonoides, Decapterus . . . . . . . . . . . . 2719 Carangue crépuscule . . . . . . . . . . . . . 2697
Carangue démêloir . . . . . . . . . . . . . . 2753 Caranx mate . . . . . . . . . . . . . . . . . . 2692

Carangue dentue . . . . . . . . . . . . . . . 2730 Caranx medusicola . . . . . . . . . . . . . . 2714
Carangue des îles . . . . . . . . . . . . . . . 2706 Caranx melampygus . . . . . . . . . . 2714-2715
Carangue grimée . . . . . . . . . . . . . . . 2707 Caranx melanostethos . . . . . . . . . . . . . 2708
Carangue langue blanche . . . . . . . . . . . 2754 Caranx mentalis . . . . . . . . . . . . . . . . 2753
Carangue lentigine . . . . . . . . . . . . . . 2694 Caranx microchir . . . . . . . . . . . . . . . 2688
Carangue monique . . . . . . . . . . . . . . 2704 Caranx micropterus . . . . . . . . . . . . . . 2754
Carangue nez bossu . . . . . . . . . . . . . 2702 Caranx miyakamii . . . . . . . . . . . . . . . 2688
Carangue noire . . . . . . . . . . . . . . . . 2713 Caranx natalensis . . . . . . . . . . . . . . . 2730
Carangue paia . . . . . . . . . . . . . . . . . 2756
Carangue pailleteé . . . . . . . . . . . . . . 2700
Caranx nigripinnis . . . . . . . . . . . . . . 2689
Carangue postillon . . . . . . . . . . . . . . 2705 Caranx nobilis . . . . . . . . . . . . . . . . . 2730
Carangue royale jaune . . . . . . . . . . . . 2725 Caranx oblongus . . . . . . . . . . . . . . . 2705
Carangue sap-sap . . . . . . . . . . . . . . . 2693 Caranx papuensis . . . . . . . . . . . . . . . 2715
Carangue tachetée . . . . . . . . . . . . . . 2699 Caranx para . . . . . . . . . . . . . . . . . . 2688
Carangue tapir . . . . . . . . . . . . . . . . 2696 Caranx pectoralis . . . . . . . . . . . . . . . 2689
Carangue têtue . . . . . . . . . . . . . . . . 2712 Caranx plagiotaenia . . . . . . . . . . . . . 2707
Carangue tille . . . . . . . . . . . . . . . . . 2717 Caranx plumbeus . . . . . . . . . . . . . . . 2702
Carangue vorace . . . . . . . . . . . . . . . 2716 Caranx praeustus . . . . . . . . . . . . . . . 2708
Caranx . . . . . . . . . . . . . 2660-2662, 2672 Caranx quptae . . . . . . . . . . . . . . . . . 2756
Caranx affinis . . . . . . . . . . . . . . . . . 2692 Caranx radiatus . . . . . . . . . . . . . . . . 2728
Caranx armatus . . . . . . . . . . . . . . . . 2693 Caranx regularis . . . . . . . . . . . . . . . 2715
Caranx ascensionis . . . . . . . . . . . . . . 2713 Caranx sansun . . . . . . . . . . . . . 2711, 2715
Caranx auroguttatus . . . . . . . . . . . . . 2694 Caranx schlegeli . . . . . . . . . . . . . . . . 2693
Caranx brevicarinatus . . . . . . . . . . . . 2707 Caranx scombrinus . . . . . . . . . . . . . . 2721
Caranx bucculentus . . . . . . . . . . . . . 2710 Caranx sem . . . . . . . . . . . . . . . . . . 2711
Caranx caeruleopinnatus . . . . . . . . . . . 2695 Caranx sexfasciatus . . . . . . . . . . 2716-2717
Caranx celetus . . . . . . . . . . . . . . . . 2715 Caranx speciosus . . . . . . . . . . . . . . . 2725
Caranx cheilio . . . . . . . . . . . . . . . . 2730 Caranx stellatus . . . . . . . . . . . . . . . . 2714
Caranx chrysophrys . . . . . . . . . . . . . . 2696 Caranx tanakai . . . . . . . . . . . . . . . . 2705
Caranx compressus . . . . . . . . . . . . . . 2707 Caranx tille . . . . . . . . . . . . . . . . . . 2717
Caranx cynodon . . . . . . . . . . . . . . . . 2717 Caranx typus . . . . . . . . . . . . . . . . . 2696
Caranx dasson . . . . . . . . . . . . . . . . 2698 Caranx uraspis . . . . . . . . . . . . . . . . 2756
Caranx deani . . . . . . . . . . . . . . . . . 2697 Caranx vomerinus . . . . . . . . . . . . . . . 2707
Caranx dentex . . . . . . . . . . . . . . . . . 2730 Caranx williamsi . . . . . . . . . . . . . . . 2711
Caranx dinema . . . . . . . . . . . . . . . . 2697 carbunculus, Thysanophrys . . . . . . . . . 2418
Caranx djedaba . . . . . . . . . . . . . . . . 2687 Cardinalfishes . . . . . . . . . . . . . . . . . 2602
Caranx elacate . . . . . . . . . . . . . . . . 2716 cardinalis, Scorpaena . . . . . . . . . . . . . 2334
Caranx equula . . . . . . . . . . . . . . . . 2698 carinata, Liza . . . . . . . . . . . . . . . . . 2085
Caranx ferdau . . . . . . . . . . . . . . . . . 2699 carinatus, Apistus . . . . . . . . . . . . . . . 2310
Caranx ferdau jordani . . . . . . . . . . . . 2706 Carpet eelblenny . . . . . . . . . . . . . . . . 2569
Caranx fulvoguttatus . . . . . . . . . . . . . 2700 Castagnoline noire . . . . . . . . . . . . . . . 2729
Caranx georgianus . . . . . . . . . . . . . . 2730 Catalufa alalaua . . . . . . . . . . . . . . . . 2599
Caranx gilberti . . . . . . . . . . . . . . . . 2699 Catalufa aleta amarillo . . . . . . . . . . . . . 2600
Caranx gracilis . . . . . . . . . . . . . . . . 2705 Catalufa aleta blanca . . . . . . . . . . . . . 2600
Caranx gymnostethus . . . . . . . . . . . . . 2701 Catalufa aleta corto . . . . . . . . . . . . . . 2599
Catalufa aleta larga . . . . . . . . . . . . . . 2599
Caranx heberi . . . . . . . . . . . . . . . . 2711
Catalufa aleta saeta . . . . . . . . . . . . . . 2600
Caranx hedlandensis . . . . . . . . . . . . . 2702 Catalufa de roca . . . . . . . . . . . . . . . . 2595
Caranx helvolus . . . . . . . . . . . . . . . . 2754 Catalufa espejuelo . . . . . . . . . . . . . . . 2596
Caranx hullianus . . . . . . . . . . . . . . . 2755 Catalufa faja blanca . . . . . . . . . . . . . . 2601
Caranx humerosus . . . . . . . . . . . . . . 2703 Catalufa mota purpúreo . . . . . . . . . . . . 2598
Caranx ignobilis . . . . . . . . . . . . . . . 2712 Catalufa Pacífico . . . . . . . . . . . . . . . . 2597
Caranx ire . . . . . . . . . . . . . . . . . . . 2708 Catalufa raya roja . . . . . . . . . . . . . . . 2601
Caranx ishikawai . . . . . . . . . . . . . . . 2713 caudimaculatum, Sargocentron . . . . . . . 2249
Caranx kalla . . . . . . . . . . . . . . . . . 2688 caudimaculatus, Adioryx . . . . . . . . . . . 2249
Caranx lugubris . . . . . . . . . . . . . . . 2713 caudimaculatus, Holocentrus . . . . . . . . . 2249
Caranx macrurus . . . . . . . . . . . . . . . 2690 Celebes flathead . . . . . . . . . . . . . . . . 2419
Caranx malabaricus . . . . . . . . . . . . . 2704 celebicus, Thysanophrys . . . . . . . . . . . 2419
Caranx malam . . . . . . . . . . . . . . . . 2689 celetus, Caranx . . . . . . . . . . . . . . . . 2715
Caranx mandibularis . . . . . . . . . . . . . 2753 CENTRISCIDAE . . . . . . . . . . . . . 2280-2281
Caranx marginatus . . . . . . . . . . . . . . 2716 Centriscus cristatus . . . . . . . . . . . . . . 2282
Centriscus scutatus . . . . . . . . . . . . . . 2282 Chato japones . . . . ... . . . . . . . . . . 2402

Centroberyx . . . . . . . . . . . . . . . . . . 2218 Chato puñalero . . . . .. . . . . . . . . . . 2415
Centroberyx affinis . . . . . . . . . . . . . . 2223 Chato rodrigo . . . . ... . . . . . . . . . . 2403
Centroberyx druzhinini . . . . . . . . . . . 2224 Cheekspot scorpionfish .. . . . . . . . . . . 2338
Centroberyx gerrardi . . . . . . . . . . . . . 2223 cheilio, Caranx . . . ... . . . . . . . . . . 2730
Centroberyx lineatus . . . . . . . . . . . . . 2224 Cheilopogon abei . . . .. . . . . . . . . . . 2169
CENTROGENIIDAE . . . . . . . . . . 2293, 2549 Cheilopogon arcticeps . . . . . . . . . . . . 2169
Centrogenys . . . . . . . . . . . . . . . . . . 2550 Cheilopogon atrisignis .. . . . . . . . . . . 2169
Centrogenys vaigiensis . . . . . . . . 2293, 2549 Cheilopogon cyanopterus . . . . . . . . . . . 2170
CENTROGENYSIDAE . . . . . . . . . . . . 2549 Cheilopogon dorsomacula . . . . . . . . . . 2170
CENTROLOPHIDAE . . . . . . . . . . . . . 2661 Cheilopogon furcatus . . . . . . . . . . . . . 2170
See also Vol. 6 Cheilopogon intermedius . . . . . . . . . . . 2171
Centropogon australis . . . . . . . . . . . . 2311 Cheilopogon katoptron . . . . . . . . . . . . 2171
CENTROPOMIDAE . . . . . . . . . . 2429, 2443 Cheilopogon pitcairnensis . . . . . . . . . . 2171
CENTROPOMINAE . . . . . . . . . . . . . . 2429 Cheilopogon rapanouiensis . . . . . . . . . 2172
Centropristes scorpaenoides . . . . . . . . . 2549 Cheilopogon spilonotopterus . . . . . . . . . 2172
Cephalopholis . . . . . . . . . . . . . . . . 2449 Cheilopogon spilopterus . . . . . . . . . . . 2172
Cephalopholis aitha . . . . . . . . . . . . . 2540 Cheilopogon suttoni . . . . . . . . . . . . . 2173
Cephalopholis argus . . . . . . . . . . 2476-2477 Cheilopogon unicolor . . . . . . . . . . . . . 2173
Cephalopholis aurantia . . . . . . . . . . . 2540 Chelidonichthys kumu . . . . . 2364-2365, 2368
Cephalopholis aurantius . . . . . . . . . . . 2482 Chelidonichthys spinosus . . . . 2364-2365, 2368
Cephalopholis boelang . . . . . . . . . . . . 2478 Chelidoperca . . . . . . . . . . . . . . . . . 2451
Cephalopholis boenak . . . . . . . . 2478, 2540 Cherna alinegra . . . . . . . . . . . . . . . . 2483
Cephalopholis coatesi . . . . . . . . . . . . . 2481 Cherna arlequin . . . . . . . . . . . . . . . . 2541
Cephalopholis cyanostigma . . . . . . 2476-2477 Cherna chillona . . . . . . . . . . . . . . . . 2479
Cephalopholis formosa . . . . . . . . . . . . 2478 Cherna chocolate . . . . . . . . . . . . . . . 2540
Cephalopholis formosanus . . . . . . . . . . 2482 Cherna de pintas azules . . . . . . . . . . . . 2477
Cephalopholis gibbus . . . . . . . . . . . . . 2481 Cherna de seis manches . . . . . . . . . . . 2481
Cephalopholis igarashiensis . . . . . . . . . 2479 Cherna dorada . . . . . . . . . . . . . . . . . 2540
Cephalopholis kendalli . . . . . . . . . . . . 2477 Cherna estrellada . . . . . . . . . . . . . . . 2480
Cephalopholis leopardus . . . . . . . . . . . 2541 Cherna frutillera . . . . .
. . . . . . . . . . . 2542
Cephalopholis microprion . . . . . . 2477, 2541 Cherna herrumbrosa . . . . . . . . . . . . . 2540
Cephalopholis miniata . . . . . 2477, 2480-2481 Cherna leopardo . . . . . . . . . . . . . . . . 2541
Cherna pavo real . . . . . . . . . . . . . . . 2476
Cephalopholis nigripinnis . . . . . . . . . . 2483 Cherna pecosa . . . . . . . . . . . . . . . . 2541
Cephalopholis polleni . . . . . . . . . . . . 2541 Cherna piña . . . . . . . . . . . . . . . . . . 2482
Cephalopholis purpureus . . . . . . . . . . . 2482 Cherna rayada . . . . . . . . . . . . . . . . . 2478
Cephalopholis rogaa . . . . . . . . . . . . . 2474 Cherna roga . . . . . . . . . . . . . . . . . . 2474
Cephalopholis sexmaculata . . . . . . 2480-2481 chewa, Epinephelus . . . . . . . . . . . . . . 2526
Cephalopholis sonnerati . . . . . . . . . . . 2482 Chicharro banda dorada . . . . . . . . . . . . 2737
Cephalopholis spiloparaea . . . . . . . . . . 2542 Chicharro ojo buey . . . . . . . . . . . . . . 2735
Cephalopholis urodeta . . . . . . . . . . . . 2483 Chicharro ojón . . . . . .. . . . . . . . . . . 2736
Cephalopholis xanthopterus . . . . . . . . . 2477 chiltonae, Thysanophrys . . . . . . . . . . . 2420
cephalotus, Mugil . . . . . . . . . . . . . . . 2096 chinensis, Aulostomus . . . . . . . . . . . . 2277
cephalus, Mugil . . . . . . . . . . . . . 2095-2096 Chinese trumpetfish . . . . . . . . . . . . . . 2277
CEPOLIDAE . . . . . . . . . . . . . . . . . 2589 chlorostigma, Epinephelus . . . . . . 2487, 2492
See also Vol. 5 Chocolate hind . . . . . . . . . . . . . 2478, 2540
ceramensis, Mugil . . . . . . . . . . . . . . . 2089 chondropus, Sillago . . . . . . . . . . . . . . 2622
Cestraeus . . . . . . . . . . . . . . . . . . . 2069 Choridactylus multibarbus . . . . . . . . . . 2311
Cestraeus goldiei . . . . . . . . . . . . . . . 2080 Chorinemus commersonnianus . . . . . . . . 2731
Cestraeus oxyrhynchus . . . . . . . . . . . . 2081 Chorinemus hainanensis . . . . . . . . . . . 2733
Cestraeus plicatilis . . . . . . . . . . . . . . 2082 Chorinemus orientalis . . . . . . . . . . . . . 2732
CETOMIMIDAE . . . . . . 2204-2206, 2208-2209 Chorinemus sanctipetri . . . . . . . . . . . . 2732
Cetomimus . . . . . . . . . . . . . . . . . . 2206 Chorinemus tala . . . . . . . . . . . . . . . . 2733
ceylonensis, Mugil . . . . . . . . . . . . . . 2102 Chorinemus tol . . . . . . . . . . . . . . . . 2734
Chaenomugil leuciscus . . . . . . . . . . . . 2099 chryseres, Myripristis . . . . . . . . . . . . . 2240
Chaenomugil nauticus . . . . . . . . . . . . 2099 chrysophryoides, Carangoides . . . . . . . . 2696
CHANDIDAE . . . . . . . . . . . . . 2433, 2602 chrysophrys, Carangoides . . . . . . . . . . 2696
chaptalii, Mugil . . . . . . . . . . . . . . . . 2096 chrysophrys, Caranx . . . . . . . . . . . . . 2696
chaptalii, Neomyxus . . . . . . . . . . . . . 2099 Cichlops . . . . . . . . . . . . . . . . . . . . 2571
Chato guiñón . . . . . . . . . . . . . . . . . 2420 Cichlops hellmuthii . . . . . . . . . . . . . . 2571
Chato índico . . . . . . . . . . . . . . . . . . 2409
Cichlops japonicus . .. . . . . . . . . . . . 2571 Coral cardinalfish . . . . .. . . . . . . . . . 2610

Cichlops melanotaenia . . . . . . . . . . . . 2571 Coral crouchers . . . . . .. . . . . . . . . . 2353
Cichlops spilopterus .. . . . . . . . . . . . 2571 Coral grouper . . . . . . .. . . . . . . . . . 2543
Cichlops trispilos . . .
. . . . . . . . . . . . 2571 Coral hind . . . . . . . . .. . . . . . . . . . 2480
ciconia, Belone . . . .
. . . . . . . . . . . . 2157 Coral reef halfbeak . . . .. . . . . . . . . . 2194
ciliaris, Alectis . . . .
. . . . . . . . . . . . 2684 Coral scorpionfish . . . . .. . . . . . . . . . 2339
ciliarius, Carangoides. . . . . . . . . . . . 2693 corallicola, Epinephelus .. . . 2491, 2508, 2543
ciliata, Sillago . . . . .
. . . . . . . . . . . . 2623 Cordonnier fil . . . . . . .. . . . . . . . . . 2684
CIRRHITIDAE . . . . . . . . . . . . . . . . . 2550 Cordonnier plume . . . . .. . . . . . . . . . 2685
See also Vol. 5 cordyla, Megalaspis . . . .. . . . . . . . . . 2726
cirrhosa, Scorpaenopsis . . .. . . . 2341, 2343 Corneta colorada . . . . .. . . . . . . . . . 2279
cirrhostomus, Mugil . . . . . .. . . . . . . . 2083 Corneta pintada . . . . . .. . . . . . . . . . 2279
Citula auroch . . . . . . . . .. . . . . . . . 2752 Cornetfishes . . . . . . . .. . . . . . . . . . 2278
Cornette à taches bleues .. . . . . . . . . . 2279
Citula diversa . . . . . . . . .. . . . . . . . 2695
Cornette rouge . . . . . . .. . . . . . . . . . 2279
Citula halli . . . . . . . . . .. . . . . . . . 2729
coruscans, Parmops . . .. . . . . . . . . . 2212
Cleaner wrasse . . . . . . . .. . . . . . . . 2648
Cleftbelly trevally . . . . . . .. . . . . . . . 2691 Coryphaena equiselis . . .. . . . . . . . . . 2657
Cleidopus . . . . . . . . . . .. . . . . . . . 2214 Coryphaena equisetis . . .. . . . . . . . . . 2657
Cloudy grouper . . . . . . . .. . . . . . . . 2543 Coryphaena hippurus . . .. . . . . . . 2657-2658
Clubfoot sillago . . . . . . . .. . . . . . . . 2622 CORYPHAENIDAE . . . .. . . . . . 2632, 2656
Clupea . . . . . . . . . . . .. . . . . . . . 2125 Coryphène commune . . .. . . . . . . . . . 2658
Coachwhip trevally . . . . . .. . . . . . . . 2705 Coryphène dauphin . . . .. . . . . . . . . . 2657
Coastal trevally . . . . . . . .. . . . . . . . 2695 Corythoichthys amplexus .. . . . . . . . . . 2273
coatesi, Cephalopholis . . . .. . . . . . . . 2481 Corythoichthys flavofasciatus . . . . . . . . 2273
Cobia . . . . . . . . . . . . .. . . . 2652, 2655 Corythoichthys haematopterus . . . . . . . . 2273
Cobie . . . . . . . . . . . . .. . . . . . . . 2655 Corythoichthys nigripectus . . . . . . . . . . 2274
coccineus, Minous . . . . . .. . . . . . . . 2320 Corythoichthys schultzi . . . . . . . . . . . . 2274
Coccotropsis . . . . . . . . .. . . . . . . . 2354 Corythoichthys spp. . . . . . . . . . . . . . 2265
Cociella crocodila . . . . . .. . . . . . . . 2393 Cottapistus cottoides . . . . . . . . . . . . . 2312
Cociella hutchinsi . . . . . .. . . . . . . . 2392 cotticeps, Scorpaenopsis . . . . . . . . . . . 2341
Cociella punctata . . . . . . .. . . . . . . . 2393 COTTIDAE . . . . . . . . . . . 2425, 2427-2428
Cockatoo waspfish . . . . . .. . . . . . . . 2309 cottoides, Cottapistus . . . . . . . . . . . . . 2312
coioides, Epinephelus . . . .. . . . 2493, 2526 Cottonmouth jack . . . . . . . . . . . . . . . 2755
colburni, Seriola . . . . . . .. . . . . . . . 2741 Cottunculus gyrinoides . . . . . . . . . . . . 2427
Comère fine . . . . . . . . . .. . . . . . . . 2720 Craterocephalus . . . . . . . . . . . 2116, 2131
Comère indienne . . . . . . .. . . . . . . . 2722 Craterocephalus anticanus . . . . . . . . . . 2129
Comère queue rouge . . . . .. . . . . . . . 2723 Craterocephalus capreoli . . . . . . . 2129, 2131
Comère rubari . . . . . . . . .. . . . . . . . 2688 Craterocephalus honoriae . . . . . . . 2130-2131
Comère saumon . . . . . . . .. . . . . . . . 2724 Craterocephalus marjoriae . . . . . . . . . . 2132
Comère torpille . . . . . . . .. . . . . . . . 2726 Craterocephalus mugiloides . . 2123, 2129, 2131
Comet betta . . . . . . . . . .. . . . . . . . 2578 Craterocephalus munroi . . . . . . . . . . . 2132
Comet grouper . . . . . . . .. . . . . . . . 2513 Craterocephalus pauciradiatus . . . . 2129, 2131
cometae, Epinephelus . . . . .. . . . . . . . 2513 crenilabis, Crenimugil . . . . . . . . . 2083-2084
Comète maouane . . . . . . .. . . . . . . . 2718 Crenimugil . . . . . . . . . . . . . . . . . . 2100
Comète maquereau . . . . . .. . . . . . . . 2719 Crenimugil crenilabis . . . . . . . . . . 2083-2084
commersonii, Fistularia . . .. . . . . 2278-2279 Crenimugil heterocheilus . . . . . . . . . . . 2084
commersonii, Hemiramphus . . . . . . . . . 2187 Crenimugil labiosus . . . . . . . . . . . . . . 2100
commersonnianus, Chorinemus . . . . . . . 2731 crenolepis, Allanetta . . . . . . . . . . . . . 2134
commersonnianus, Scomberoides . . . 2731-2732 crenolepis, Hypoatherina . . . . . . . . . . . 2134
Common dolphinfish . . . . . . . . . . . . . 2658 Crenulated silverside . . . . . . . . . . . . . 2134
compressus, Caranx . . . . . . . . . . . . . . 2707 crinitus, Alectis . . . . . . . . . . . . . . . . 2684
Congaturi halfbeak . . . . . . . . . . . . . . 2191 cristatus, Centriscus . . . . . . . . . . . . . 2282
CONGROGADINAE . . . . . . . . . . . . . . 2586 crocodila, Cociella . . . . . . . . . . . . . . 2393
Congrogadines . . . . . . . . . . . . . . . . 2586 Crocodile fish . . . . . . . . . . . . . . . . . 2394
Congrogadus subducens . . . . . . . . . . . 2569 crocodilus crocodilus, Tylosurus . . . . . . . 2160
convexus bruuni, Oxyporhamphus . . . . . 2192 crocodilus fodiator, Tylosurus . . . . . . . . 2160
convexus convexus, Oxyporhamphus . . . . 2192 crocodilus, Tylosurus . . . . . . . . . . . . . 2159
convexus, Oxyporhamphus convexus . . . . 2192 crocodilus, Tylosurus crocodilus . . . . . . . 2160
Cookeolus japonicus . . . . . . . . . . . . . 2599 Croissant queue blanche . . . . . . . . . . . 2538
cookii, Scorpaena . . . . . . . . . . . . . . . 2334 Croissant queue jaune . . . . . . . . . . . . . 2539
copperingi, Trachinotus . . . . . . . . . . . 2748 Cromileptes altivelis . . . . . . . . . . . . . 2484
crosnieri, Callanthias . . . . . . . . . . . . . 2556 Darkfin hind . . . . . . . .

. . . . . . . . . . 2483
crosnieri, Grammatonotus . . . . . . . . . . 2556 Darkspotted scorpionfish .
. . . . . . . . . . 2346
Crown-of-thorns starfish . . . . . .. . . . . . 2602 Daruma stinger . . . . . .
. . . . . . . . . . 2315
cruentatus, Heteropriacanthus . . . 2595, 2599 Darwin grouper . . . . . .
. . . . . . . . . . 2543
cruentatus, Priacanthus . . . . . . . . . . . . 2595 darwinensis, Epinephelus . . . . . . . . . . 2543
crumenophthalmus, Selar . . . . . . . 2735-2736 dasson, Carangoides . . . . . . . . . . . . . 2698
crumenophthalmus, Trachurops . . . . . . . 2736 dasson, Caranx . . . . . . . . . . . . . . . . 2698
Culion silverside . . . . . . . . . .
. . . . . . 2128 dayi, Decapterus . . . . . . . . . . . . . . . . 2722
cunnesius, Valamugil . . . . . . 2102-2103-2108 deani, Caranx . . . . . . . . . . . . . . . . . 2697
curtus, Parabembras . . . . . . . . . . . . . 2383 decadactylus, Beryx . . . . . . . . . . . 2221-2222
Curvedspine scorpionfish . . . . . . . . . . . 2323 Decapterus . . . . . . . 2650, 2655, 2661, 2675,
cuvieri, Inimicus . . . . . . . . . . . . . . . 2316 2720, 2722-2723
cyanodorsalis, Pseudomugil . . . . . . 2110-2111 Decapterus afuerae . . . . . . . . . . . . . . 2720
cyanopodus, Epinephelus . . . . . . . . . . 2494 Decapterus canonoides . . . . . . . . . . . . 2719
cyanopterus, Cheilopogon . . . . . . . . . . 2170 Decapterus dayi . . . . . . . . . . . . . . . . 2722
cyanosoma, Apogon . . . . . . . . . . . . . 2608 Decapterus hypodus . . . . . . . . . . . . . . 2721
cyanostigma, Cephalopholis . . . . . . 2476-2477 Decapterus jacobeus . . . . . . . . . . . . . 2719
cyanostigma, Sebastapistes . . . . . . . . . . 2344 Decapterus kiliche . . . . . . . . . . . . . . . 2722
cyanotaenia, Pseudochromis . . . . . . . . . 2573 Decapterus kurroides . . . . . . . . . . . . . 2718
cyclophthalmus, Labracinus . . . . . 2571, 2574 Decapterus kurroides akaadsi . . . . . . . . 2718
cylindricus, Atherinomorus . . . . . . 2121-2122 Decapterus lajang . . . . . . . . . . . 2720, 2722
cylindricus, Mugil . . . . . . . . .. . . . . . 2107 Decapterus lundini . . . . . . . . . . . . . . 2692
Cymbacephalus beauforti . . . . . . . . . . 2394 Decapterus macarellus . . . . . . . . . 2719-2721
Cymbacephalus bosschei . . . . . . . . . . . 2395 Decapterus macrosoma . . . . . 2719-2720, 2722
Cymbacephalus nematophthalamus . . . . . 2396 Decapterus maruadsi . . . . . . . . . . . . . 2722
Cymbacephalus parilis . . . . . . . . . . . . 2397 Decapterus muroadsi . . . . . . . . . . . . . 2721
Cymbacephalus staigeri . . . . . . . . . . . 2397 Decapterus normani . . . . . . . . . . . . . . 2692
cynodon, Caranx . . . . . . . . . . . . . . . 2717 Decapterus pinnulatus . . . . . . . . . . . . 2719
Cypho purpurascens . . . . . . . . . . . . . 2570 Decapterus politus . . . . . . . . . . . . . . 2692
cypho, Vespicula . . . . . . . . . . . . . . . 2351 Decapterus russelli . . . . . . . 2720, 2722-2723
CYPRINODONTIFORMES . . . . . . . 2197-2200 Decapterus tabl . . . . . . . . . . . . . 2722-2723
Cypselurus angusticeps . . . . . . . . . . . 2173 declivis, Trachurus . . . . . . . . . . . 2750-2751
Cypselurus hexazona . . . . . . . . . . . . . 2174 Decoy scorpionfish . . . . . . . . . . . . . . 2317
Cypselurus naresii . . . . . . . . . . . . . . 2174 Deepwater cardinalfishes . . . . . . . . . . . 2611
Cypselurus oligolepis . . . . . . . . . . . . . 2174 Deepwater firefish . . . . . . . . . . . . . . . 2330
Cypselurus opisthopus . . . . . . . . . . . . 2175 Deepwater flatheads . . . . . . . . . . . . . . 2383
Cypselurus poecilopterus . . . . . . . . . . . 2175 Deepwater scorpionfish . . . . . . . . . . . . 2347
Cypselurus simus . . . . . . . . . . . . . . . 2175 delicatulus, Lactarius . . . . . . . . . . . . . 2649
Cyttomimus . . . . . . . . . . . . . . . . . . 2259 Demi-bec bagnard . . . . . . . . . . . . . . . 2187
Cyttopsis . . . . . . . . . . . . . . . . . . . 2259 Demi-bec congaturi . . . . . . . . . . . . . . 2191
Demi-bec de Dussumier . . . . . . . . . . . . 2189
D Demi-bec de Lutke . . . . . . . . . . . . . . 2188
Demi-bec de Quoy . . . . . . . . . . . . . . . 2190
dactyliophorus, Doryrhamphus (Doryrhamphus) Demi-bec saltou . . . . . . . . . . . . . . . . 2186
. . . . . . . . . . . . . . . . . . . . . . . 2274 Dendrochirus bellus . . . . . . . . . . . . . . 2312
Dactyloptena gilberti . . . . . . . . . . . . . 2285 Dendrochirus biocellatus . . . . . . . . . . . 2313
Dactyloptena jordani . . . . . . . . . . . . . 2285 Dendrochirus brachypterus . . . . . . . . . 2313
Dactyloptena macracantha . . . . . . . . . . 2286 Dendrochirus zebra . . . . . . . . . . . . . . 2314
Dactyloptena orientalis . . . . . . . . 2287, 2289 Dentatherina merceri . . . . . . . . . . . . . 2145
Dactyloptena papilio . . . . . . . . . . . . . 2288 DENTATHERINIDAE . . . . . . . . . . . . . 2145
Dactyloptena peterseni . . . . . . . . . . . . 2289 dentex, Caranx . . . . . . . . . . . . . . . . 2730
Dactyloptena tiltoni . . . . . . . . . . . . . . 2290 dentex, Pseudocaranx . . . . . . . . . . . . 2730
DACTYLOPTERIDAE . . 2262, 2283, 2360, 2551 depressifrons, Vespicula . . . . . . . . . . . 2352
daemelii, Epinephelus . . . . . . . . . . . . 2495 Dermogenys . . . . . . . . . . . . . . . . . . 2181
Daicocus peterseni . . . . . . . . . . . . . . 2289 dermopterus, Triso . . . . . . . . . . . . . . 2547
Dampieria . . . . . . . . . . . . . . . . . . . 2571 detrusus, Kumococius . . . . . . . . . . . . . 2403
Dampieria longipinna . . . . . . . . . . . . 2572 Devil firefish . . . . . . . . . . . . . . . . . . 2330
Dampieria melanostigma . . . . . . . . . . . 2571 diabolus, Scorpaenopsis . . . . . . . . 2342-2343
Dampieria ocellifera . . . . . . . . . . . . . 2571 diacanthus, Epinephelus . . . . 2498, 2522, 2525
danae, Melamphaes . . . . . . . . . . . . . 2201 Diadema . . . . . . . . . . . . . . . . 2602, 2609
Darkbar flyingfish . . . . . . . . . . . . . . . 2174 Diadema setosum . . . . . . . . . . . . . . . 2282
diadema, Mugil . . . . . . . . . . . . . . . . 2086 duodecimalis, Atherinomorus . . 2119, 2121-

diadema, Pseudochromis . . . . . . . 2557, 2576 2122-2124, 2128
diadema, Pteromugil . . . . . . . . . . . . . 2086 duodecimalis, Pranesus . . . . . . . . . . . . 2122
Diamond mullet . . . . . . . . . . . . . . . . 2086 Duskytail grouper . . . . . . . . . . . . . . . 2489
Diamond-scale mullet . . . . . . . . . . . . . 2094 Dussumier’s halfbeak . . . . . . . . . . . . . 2189
didactylus, Inimicus . . . . . . . . . . . . . 2316 dussumieri, Hyporhamphus . . . . . . . . . 2190
diemensis, Agonostoma . . . . . . . . . . . . 2079 dussumieri, Hyporhamphus (Reporhamphus)
Dikellorhynchus incredibilis . . . . . . . . . 2647 . . . . . . . . . . . . . . . . . . . 2189, 2194
dimidiatus, Labroides . . . . . . . . . . . . 2648 dussumieri, Mugil . . . . . . . . . . . . . . . 2093
dinema, Carangichthys . . . . . . . . . . . . 2697 Dwarf dories . . . . . . . . . . . . . . . . . . 2258
dinema, Carangoides . . . . . . . . . . . . . 2697 Dwarf flathead . . . . . . . . . . . . . . . . . 2398
dinema, Caranx . . . . . . . . . . . . . . . . 2697 Dwarf lionfish . . . . . . . . . . . . . . . . . 2313
DIRETMIDAE . . . . . . . . . . 2210-2211, 2216 Dwarf scorpionfish . . . . . . . . . . . . . . . 2337
Discfishes . . . . . . . . . . . . . . . . . . . 2652 Dwarf stingfish . . . . . . . . . . . . . . . . . 2320
dispar var. A, Serranus . . . . . . . . . . . . 2527 Dwarf thronyhead . . . . . . . . . . . . . . . 2328
dispar var. B, Epinephelus . . . . . . . . . . 2516
E
diversa, Citula . . . . . . . . . . . . . . . . . 2695
diversidans, Ratabulus . . . . . . . . . . . . 2410 Eastern keelback mullet . . . .
. . . . . . . . 2085
Dixiphichthys ferculum . . . . . . . . . . . . 2376 ECHENEIDAE . . . . . . . . .
. . . . . . . . 2652
Dixiphichthys hoplites . . . . . . . . . . . . 2374 Echeneis . . . . . . . . . . . .
. . . . . . . . 2652
Dixiphistes macrorhynchus . . . . . . . . . . 2376 Echeneis naucrates . . . . . .
. . . . 2652, 2654
Dixiphistops megalops . . . . . . . . . . . . 2377 Ectreposebastes . . . . . . . .
. . . . . . . . 2291
djedaba, Alepes . . . . . . . . . . . . . . . . 2687 Ectreposebastes imus . . . . .
. . . . . . . . 2314
djedaba, Atule . . . . . . . . . . . . . . . . . 2687 Eelblennies . . . . . . . . . .
. . . . . . . . 2557
djedaba, Caranx . . . . . . . . . . . . . . . 2687 eendrachtensis, Atherinomorus . . . . . . . . 2123
djeddaba, Atule . . . . . . . . . . . . . . . . 2687 Egg-laying toothcarps . . . . . . . . . . . . . 2197
Eightbar grouper . . . . . . . . . . . . . . . . 2545
dobula, Mugil . . . . . . . . . . . . . . . . . 2096
döderleinii, Epinephelus . . . . . . . . . . . 2519
elacate, Caranx . . . . . . . . . . . . . . . . 2716
doliatus, Branchiostegus . . . . . . . . 2643-2644
Elagatis . . . . . . . . . . 2650, 2655, 2659-2660
Dolphinfishes . . . . . . . . . . . . . . . . . 2656 Elagatis bipinnulata . . . . . . . . . . . . . 2724
Dolphins . . . . . . . . . . . . . . . . . . . . 2656 Elates . . . . . . . . . . . . . . . . . . . . . 2385
Dorado . . . . . . . . . . . . . . . . . . . . . 2657 Elates ransonnetii . . . . . . . . . . . . . . . 2398
Dorado común . . . . . . . . . . . . . . . . . 2658 elegans, Antipodocottus . . . . . . . 2425, 2428
Dories . . . . . . . . . . . . . . . . . . . . . 2259 Ellochelon vaigiensis . . . . . . . . . . . . . 2094
dorsalis, Seriola . . . . . . . . . . . . . . . . 2740 elongatus, Epinephelus . . . . . . . . . . . . 2526
dorsomacula, Cheilopogon . . . . . . . . . . 2170 elongatus, Myxus . . . . . . . . . . . . . . . 2097
Doryrhamphus (Doryrhamphus) dactyliophorus elymus aphrozoicus, Atherion . . . . . . . . . 2118
. . . . . . . . . . . . . . . . . . . . . . . 2274 elymus asper, Atherion . . . . . . . . . . . . 2118
Doryrhamphus (Doryrhamphus) excisus excisus elymus freyi, Atherion . . . . . . . . . . . . . 2118
. . . . . . . . . . . . . . . . . . . . . . . 2275 elymus, Atherion . . . . . . . . . . . . . . . 2118
Doryrhamphus (Doryrhamphus) multiannulatus emburyi, Turrum . . . . . . . . . . . . . . . 2700
. . . . . . . . . . . . . . . . . . . . . . . 2275 EMMELICHTHYIDAE . . . . . . . . . . . . . 2437
Doryrhamphus (Doryrhamphus) pessuliferus See also Vol. 5
. . . . . . . . . . . . . . . . . . . . . . . 2275 emoryi, Epinephelus . . . . . . . . . . . . . . 2499
Doryrhamphus spp. . . . . . . . . . . . . . 2265 Endracht land silverside . . . . . . . . . . . . 2123
Dotdash grouper . . . . . . . . . . . . . . . 2515 endrachtensis, Atherinomorus . . . . 2119-2123,
Dotted grouper . . . . . . . . . . . . . . . . 2496 2126, 2129, 2131
Dottybacks . . . . . . . . . . . . . . . . . . 2557 endrachtensis, Platycephalus . . . . . . . . . 2408
Double-ended pipefish . . . . . . . . . . . . 2276 endrachtensis, Pranesus . . . . . . . 2123, 2127
Doublespotted queenfish . . . . . . . . . . . 2732 engeli, Valamugil . . . . . . . . 2102, 2104-2106
Doubletooth solderfish . . . . . . . . . . . . 2254 ENGRAULIDAE . . . . . . . . . . . . 2113, 2551
Dragon’s beard fish . . . . . . . . . . . . . . 2329 See also Vol. 3
DREPANEIDAE . . . . . . . . . . . . . . . . 2259 Epaulet trevally . . . . . . . . . . . . . . . . 2703
See also Vol. 5 EPIGONIDAE . . . . . . . . . . 2436, 2611, 2650
Druzhinin’s alfonsino . . . . . . . . . . . . . 2224 Epigonus . . . . . . . . . . . . . . . . . . . 2611
druzhinini, Centroberyx . . . . . . . . . . . 2224 Epigonus atherinoides . . . . . . . . . . . . 2613
ductor, Naucrates . . . . . . . . . . . . . . . 2727 Epigonus glossodontus . . . . . . . . . . . . 2611
dumerili, Seriola . . . . . . . . . . . . . . . 2738 Epigonus macrops . . . . . . . . . . . . . . 2613
Duncker’s pipefish . . . . . . . . . . . . . . . 2275 Epigonus parini . . . . . . . . . . . . . . . . 2611
dunckeri, Halicampus . . . . . . . . . . . . 2275 EPINEPHELINAE . . . . . . . . . . . 2442, 2474
duodecimalis, Allanetta . . . . . . . . . . . . 2122 EPINEPHELINI . . . . . . . . . . . . . . . . 2474
Epinephelus . . . . . . . . . . . . . . 2451, 2529 Epinephelus morrhua . . . . .

2513, 2515, 2519
Epinephelus akaara . . . . . . . . . 2485, 2498 Epinephelus multinotatus . . .
. . . . . . . 2545
Epinephelus albopunctulatus . . . . . . . . . 2504 Epinephelus octofasciatus . . .
. . . 2497, 2545
Epinephelus amblycephalus . . . . . . . . . 2486 Epinephelus ongus . . . . . . .
2491, 2514, 2523
Epinephelus areolatus . . . . . . . . 2487, 2492 Epinephelus perplexus . . . . .
. . . . . . . 2545
Epinephelus awoara . . . . . . . . . . . . . 2488 Epinephelus poecilonotus . . .
. . . 2506, 2510,
Epinephelus bilobatus . . . . . . . . . . . . 2542 2513, 2515, 2519
Epinephelus bleekeri . . . . . . . . . . . . . 2489 Epinephelus polylepis . . . . . . . . . . . . . 2492
Epinephelus bontoides . . . . . . . . . . . . 2542 Epinephelus polyphekadion . . . . . 2501, 2516
Epinephelus bruneus . . . . . . . . . . . . . 2490 Epinephelus polystigma . . . . . . . . . . . 2517
Epinephelus caeruleopunctatus . . . 2491, 2514 Epinephelus praeopercularis . . . . . . . . . 2496
Epinephelus chewa . . . . . . . . . . . . . . 2526 Epinephelus pseudomorrhua . . . . . . . . . 2510
Epinephelus chlorostigma . . . . . . 2487, 2492 Epinephelus quoyanus . . . . . 2500, 2503, 2508,
Epinephelus coioides . . . . . . . . . 2493, 2526 2512, 2518, 2524
Epinephelus cometae . . . . . . . . . . . . . 2513 Epinephelus radiatus . . . . . . 2513, 2515, 2519
Epinephelus corallicola . . . . 2491, 2508, 2543 Epinephelus rahanus . . . . . . . . . . . . . 2517
Epinephelus cyanopodus . . . . . . . . . . . 2494 Epinephelus retouti . . . . . . . . . . 2499, 2520
Epinephelus daemelii . . . . . . . . . . . . . 2495 Epinephelus rhyncholepis . . . . . . . . . . . 2521
Epinephelus darwinensis . . . . . . . . . . . 2543 Epinephelus rivulatus . . . . . . . . . . . . 2521
Epinephelus diacanthus . . . . 2498, 2522, 2525 Epinephelus salonotus . . . . . . . . . . . . . 2524
Epinephelus dispar var. B . . . . . . . . . . 2516 Epinephelus septemfasciatus . . . . . . . . . 2497
Epinephelus döderleinii . . . . . . . . . . . . 2519 Epinephelus sexfasciatus . . . . . . . 2498, 2522
Epinephelus elongatus . . . . . . . . . . . . 2526 Epinephelus slacksmithi . . . . . . . . . . . . 2514
Epinephelus emoryi . . . . . . . . . . . . . . 2499 Epinephelus socialis . . . . . . . . . . . . . 2523
Epinephelus epistictus . . . . . . . . 2496, 2502, Epinephelus spilotoceps . . . . . . . 2500, 2503,
2506, 2510, 2515 2512, 2518, 2524
Epinephelus ergastularius . . . . . . . . . . 2497 Epinephelus spiniger . . . . . . . . . . . . . 2504
Epinephelus erythrurus . . . . . . . . . . . 2543 Epinephelus spiramen . . . . . . . . . . . . . 2521
Epinephelus fario . . . . . . . . . . . . . . . 2507 Epinephelus stictus . . . . . . . . . . . . . . 2525
Epinephelus fasciatomaculatus . . . . . . . . 2498 Epinephelus suillus . . . . . . . . . . . . . . 2493
Epinephelus fasciatomaculosus . . . 2485, 2498 Epinephelus summana . . . . . . . . 2491, 2514
Epinephelus fasciatus . . . . . . . . 2499, 2520 Epinephelus tauvina . . . 2493, 2505, 2511, 2526
Epinephelus faveatus . . . 2500, 2508, 2512, 2518 Epinephelus timorensis . . . . . . . . . . . . 2546
Epinephelus fuscoguttatus . . . . . . 2501, 2516 Epinephelus trophis . . . . . . . . . . . . . . 2546
Epinephelus gaimardi . . . . . . . . . . . . 2507 Epinephelus truncatus . . . . . . . . . . . . . 2520
Epinephelus grammatophorus . . . . . . . . 2521 Epinephelus tuamotuensis . . . 2513, 2515, 2546
Epinephelus grammicus . . . . . . . . . . . . 2506 Epinephelus tukula . . . . . . . . . . . . . . 2527
Epinephelus hata . . . . . . . . . . . . . . . 2502 Epinephelus undulatostriatus . . . . . . . . 2528
Epinephelus heniochus . . . . . . . 2496, 2502 Epinephelus undulosus . . . . . . . . . . . . 2529
Epinephelus hexagonatus . . . . . . 2500, 2503, epistictus, Epinephelus . . . . . . . . 2496, 2502,
2512, 2518, 2524 2506, 2510, 2515
Epinephelus hoedtii . . . . . . . . . . . . . . 2494 equiselis, Coryphaena . . . . . . . . . . . . 2657
Epinephelus homosinensis . . . . . . . . . . 2521 equisetis, Coryphaena . . . . . . . . . . . . . 2657
Epinephelus howlandi . . . . . . . . 2508, 2544 equula, Carangoides . . . . . . . . . . . . . 2698
Epinephelus irroratus . . . . . . . . 2504, 2520 equula, Caranx . . . . . . . . . . . . . . . . 2698
Epinephelus kohleri . . . . . . . . . . . . . . 2494 equula, Kaiwarinus . . . . . . . . . . . . . . 2698
Epinephelus lanceolatus . . . . . . . 2505, 2526 ergastularius, Epinephelus . . . . . . . . . . 2497
Epinephelus latifasciatus . . . . . . . . . . 2506 Erosa erosa . . . . . . . . . . . . . . . . . . 2315
Epinephelus longispinis . . . . . . . . . . . 2507 erosa, Erosa . . . . . . . . . . . . . . . . . . 2315
Epinephelus macrospilos . . . . . . . 2500, 2503, erythrurus, Epinephelus . . . . . . . . . . . 2543
2508-2512, 2518, 2524 Estuarine hardyhead . . . . . . . . . . . . . 2130
Epinephelus maculatus . . . . . . . 2507, 2509 Estuarine stonefish . . . . . . . . . . . . . . 2348
Epinephelus magniscuttis . . . 2496, 2510, 2515 Euleptorhamphus viridis . . . . . . . 2180, 2193
Epinephelus malabaricus . . . 2493, 2511, 2526 evermanni, Carangoides gymnostethoides . . 2706
Epinephelus medurensis . . . . . . . . . . . 2509 excisus excisus, Doryrhamphus (Doryrhamphus)
Epinephelus megachir . . . . . . . . . . . . 2518 . . . . . . . . . . . . . . . . . . . . . . . 2275
Epinephelus melanostigma . . 2512, 2524, 2544 excisus, Doryrhamphus (Doryrhamphus) excisus
Epinephelus merra . . . . 2500, 2503, 2512, 2524 . . . . . . . . . . . . . . . . . . . . . . . 2275
Epinephelus microdon . . . . . . . . 2501, 2516 Exocet à dos tacheté . . . . . . . . . . . . . 2170
Exocet à grandes écailles . . . . . . . . . . . 2174
Epinephelus miliaris . . . . . . . . . . . . . 2544 Exocet à tache blanche . . . . . . . . . . . . 2177
Exocet aile jaune . . .. . . . . . . . . . . . 2175 Flyingfishes . . . . . . . . . . . . . . 2162, 2180

Exocet alle courte . . .. . . . . . . . . . . . 2178 fodiator, Tylosurus crocodilus . . . . . . . . 2160
Exocet barbu . . . . .. . . . . . . . . . . . 2176 Formio niger . . . . . . . . . . . . . . . . . 2729
Exocet bécune . . . .. . . . . . . . . . . . 2173 formosa, Cephalopholis . . . . . . . . . . . 2478
Exocet bouledogue . .. . . . . . . . . . . . 2176 formosanus, Carangoides . . . . . . . . . . . 2695
Exocet casque . . . . .. . . . . . . . . . . . 2177 formosanus, Cephalopholis . . . . . . . . . 2482
Exocet codène . . . .. . . . . . . . . . . . 2170 forskalii, Atherina . . . . . . . . . . . . . . . 2125
Exocet de Sutton . . .. . . . . . . . . . . . 2173 forsteri, Agonostomus . . . . . . . . . . . . . 2079
Exocet marbré . . . . .. . . . . . . . . . . . 2172 forsteri, Aldrichetta . . . . . . . . . . 2079, 2087
Exocet matelot . . . .. . . . . . . . . . . . 2179 Fortesque . . . . . . . . . . . . . . . . . . . 2311
Exocet miroir . . . . .. . . . . . . . . . . . 2177 Foursaddle grouper . . . . . . . . . . . . . . 2524
Exocet pharaon . . . .. . . . . . . . . . . . 2174 Fowler’s scorpionfish . . . . . . . . . . . . . 2342
Exocet planeur . . . .. . . . . . . . . . . . 2169 fowleri, Scorpaenopsis . . . . . . . . . . . . 2342
Exocet tacheté . . . .. . . . . . . . . . . . 2170
Exocet voiler africain .. . . . . . . . . . . . 2178
Fraudella . . . . . . . . . . . . . . . . . . . 2578
Freckled hind . . . . . . . . . . . . . . . . . 2541
Exocet voilier . . . . .. . . . . . . . . . . . 2178
Freckled tilefish . . . . . . . . . . . . . . . . 2644
Exocet volant . . . . .. . . . . . . . . . . . 2176
Freespine flathead . . . . . . . . . . . . . . . 2410
EXOCOETIDAE . . . .. . . . . . . . 2162, 2180
Freshwater mullet . . . . . . . . . . . . . . . 2098
Exocoetus monocirrhus . . . . . . . . . . . 2176
freyi, Atherion elymus . . . . . . . . . . . . . 2118
Exocoetus obtusirostris . . . . . . . . . . . . 2176 Fringe-eyed flathead . . . . . . . . . . . . . . 2396
Exocoetus volitans . . . . . . . . . . . . . . 2176 Fringefin trevally . . . . . . . . . . . . . . . . 2728
Fringelip flathead . . . . . . . . . . . . . . . 2421
F Fringelip mullet . . . . . . . . . . . . . . . . 2083
falcata, Seriola . . . . . . . . .
. . . . . . . 2741 frondosa, Rhinopias . . . . . . . . . . . . . 2333
False scorpionfish . . . . . . . .
. . . 2293, 2549 fulvoguttatus, Carangoides . . . . . . . 2700-2701
False stonefish . . . . . . . . .
. . . . . . . 2342 fulvoguttatus, Caranx . . . . . . . . . . . . . 2700
False trevally . . . . . . . . . .
. . . . . . . 2649 furcatus, Cheilopogon . . . . . . . . . . . . 2170
Fangtooths . . . . . . . . . . .
. . . . . . . 2210 fuscoguttatus, Epinephelus . . . . . . 2501, 2516
Fantail mullet . . . . . . . . . .
. . . . . . . 2105 fuscus, Pseudochromis . . . . . . . . 2571, 2574
far, Hemiramphus . . . . . . .
. . . 2187, 2193
fario, Epinephelus . . . . . . .
. . . . . . . 2507 G
fasciatomaculatus, Epinephelus . . . . . . . 2498
fasciatomaculosus, Epinephelus . . . 2485, 2498 gaimardi, Epinephelus . . . . . . . . . . . . 2507
fasciatus, Epinephelus . . . . . . . . 2499, 2520 gaimardi, Hemiramphus . . . . . . . . . . . 2191
Galactacma scorpionfish . . . . . . . . . . . 2345
Fathead sculpins . . . . . . . . . . . . . . . 2427
faveatus, Epinephelus . . 2500, 2508, 2512, 2518 galactacma, Sebastapistes . . . . . . . . . . 2345
Gammarida . . . . . . . . . . . . . . . . . . 2276
ferculum, Dixiphichthys . . . . . . . . . . . . 2376 Gargariscus prionocephalus . . . . . . . . . 2366
ferdau jordani, Caranx . . . . . . . . . . . . 2706 Gargariscus semidentatus . . . . . . . . . . . 2366
ferdau, Carangoides . . . . . . . . . 2699, 2706 Garish hind . . . . . . . . . . . . . . . . . . 2479
ferdau, Caranx . . . . . . . . . . . . . . . . 2699 GASTEROSTEIFORMES . . . . . . . . 2262-2282
ferrandi, Mugil . . . . . . . . . . . . . . . . 2087 gavialoides, Tylosurus . . . . . . . . . . . . 2161
Ficklespotted flyingfish . . . . . . . . . . . . 2175 GEMPYLIDAE . . . . . . . . . . . . . . . . . 2660
Fijian silverside . . . . . . . . . . . . . . . . 2135 See also Vol. 6
Fistularia commersonii . . . . . . . . 2278-2279 Gennadius . . . . . . . . . . . . . . . . . . . 2549
Fistularia petimba . . . . . . . . . . . 2278-2279 George’s halfbeak . . . . . . . . . . . . . . . 2196
FISTULARIIDAE . . . . . 2263, 2265, 2277-2278 georgianus, Caranx . . . . . . . . . . . . . . 2730
fitchi, Priacanthus . . . . . . . . . . . . . . 2600 georgianus, Usacaranx . . . . . . . . . . . . 2730
Flammeo opercularis . . . . . . . . . . . . . 2244 georgii, Rhynchorhamphus . . . . . . . . . 2196
Flammeo sammara . . . . . . . . . . . . . . 2245 georgii, Valamugil . . . . . . . . . . . . . . 2105
Flashlight fishes . . . . . . . . . . . . . . . . 2212 gerrardi, Centroberyx . . . . . . . . . . . . . 2223
Flat needlefish . . . . . . . . . . . . . . . . . 2154 Ghost flatheads . . . . . . . . . . . . . . . . 2422
Flat-tail mullet . . . . . . . . . . . . . . . . . 2087 Ghost pipefishes . . . . . . . . . . . . . . . . 2263
Flateared helmet gurnard . . . . . . . . . . . 2285 Giant grouper . . . . . . . . . . . . . . . . . 2505
Flathead mullet . . . . . . . . . . . . . . . . 2096 Giant seaperch . . . . . . . . . . . . . . . . 2431
Flatheads . . . . . . . . . . . . . . . . . . . 2385 Giant trevally . . . . . . . . . . . . . . . . . . 2712
flavofasciatus, Corythoichthys . . . . . . . . 2273 Gibberfishes . . . . . . . . . . . . . . . . . . 2203
Florenciella . . . . . . . . . . . . . . . . . . 2611 GIBBERICHTHYIDAE . . . . . . . . . 2201, 2203
flosmaris, Iso . . . . . . . . . . . . . . . . . 2141 gibbifrons, Scorpaena . . . . . . . . . . . . 2335
Flute mouths . . . . . . . . . . . . . . . . . 2278 gibbosa, Scorpaenopsis . . . . . . . . . 2342-2343
FLUVIPHYLACINAE . . . . . . . . . . . . . 2200 gibbus, Cephalopholis . . . . . . . . . . . . . 2481
Flying gurnards . . . . . . . . . . . . . . . . 2283
Flying halfbeak . . . . . . . . . . . . . . . . 2192
gilberti, Caranx . . . . . . . . . . . . . . . . 2699
gilberti, Dactyloptena . . . . . . . . . . . . . 2285 gymnostethoides evermanni, Carangoides . . 2706

glabra, Alepes . . . . . . . . . . . . . . . . . 2690 gymnostethoides, Carangoides . . . . . . . . 2701
Glasseye . . . . . . . . . . . . . . . . . . . 2595 gymnostethus, Carangoides . . 2694, 2700-2701
Glassfishes . . . . . . . . . . . . . . . . . . 2433 gymnostethus, Caranx . . . . . . . . . . . . . 2701
GLAUCOSOMATIDAE . . . . . . . . . . . . 2443 gyrinoides, Cottunculus . . . . . . . . . . . 2427
See also Vol. 5
glaucus, Abudefduf . . . . . . . . . . . . . . 2349 H
Glider flyingfish . . . . . . . . . . . . . . . . 2169 haematopterus, Corythoichthys . . . . . . . 2273
Glorious groppo . . . . . . . . . . . . . . . . 2555 HAEMULIDAE . . . . . . . . . . . . . . . . . 2443
Glossamia . . . . . . . . . . . . . . . . . . . 2602 See also Vol. 5
glossodontus, Epigonus . . . . . . . . . . . 2611 hainanensis, Chorinemus . . . . . . . . . . . 2733
Gnathanodon speciosus . . . . . . . . . . . 2725 Hairy scorpionfish . . . . . . . . . . . . . . . 2337
Gnomefishes . . . . . . . . . . . . . . . . . 2651 Hairyfish . . . . . . . . . . . . . . . . . . . . 2208
Gobid fishes . . . . . . . . . . . . . . . . . . 2353 Half fringelip mullet . . . . . . . . . . . . . . 2084
gobio, Hypoatherina . . . . . . . . . . . . . 2136 Halfbeaks . . . . . . . . . . . . . . . 2151, 2180
Gobiodon . . . . . . . . . . . . . . . . . . . 2353 Halfmoon grouper . . . . . . . . . . . . . . . 2521
Goblinfish . . . . . . . . . . . . . . . . . . . 2352 Halfscaled scorpionfish . . . . . . . . . . . . 2335
Golden grouper . . . . . . . . . . . . . . . . 2537 Halicampus dunckeri . . . . . . . . . . . . . 2275
Golden hind . . . . . . . . . . . . . . . . . . 2540 Halicampus spp. . . . . . . . . . . . . . . . 2265
Golden trevally . . . . . . . . . . . . . . . . 2725 Halichoeres purpurescens . . . . . . . . . . 2475
Goldenlined sillago . . . . . . . . . . . . . . 2618 halli, Citula . . . . . . . . . . . . . . . . . . 2729
Goldie river mullet . . . . . . . . . . . . . . . 2080 hamrur, Priacanthus . . . . . . . . . 2596, 2599
goldiei, Aeschrichthys . . . . . . . . . . . . . 2080 Haplocheilus argyrotaenia . . . . . . . . . . 2138
goldiei, Cestraeus . . . . . . . . . . . . . . . 2080 Hardyhead silverside . . . . . . . . . . . . . 2125
Goldies . . . . . . . . . . . . . . . . . . . . 2553 Hardyheads . . . . . . . . . . . . . . . . . . 2113
Gracila albomarginata . . . . . . . . . . . . 2530 Harlequin hind . . . . . . . . . . . . . . . . . 2541
gracilis, Caranx . . . . . . . . . . . . . . . . 2705 Harris’ flathead . . . . . . . . . . . . . . . . . 2401
gracilis, Macrorhamphosus . . . . . . . . . . 2280 harrisii, Inegocia . . . . . . . . . . . . . . . 2401
Gracimugil ramsayi . . . . . . . . . . . . . . 2092 haswelli, Hoplichthys . . . . . . . . . . . . . 2422
Grammatonotus . . . . . . . . . . . . . . . 2579 hata, Epinephelus . . . . . . . . . . . . . . . 2502
Grammatonotus crosnieri . . . . . . . . . . 2556 Hawaiian silverside . . . . . . . . . . . . . . 2124
grammatophorus, Epinephelus . . . . . . . . 2521 Hawaiian surf sardine . . . . . . . . . . . . . 2142
GRAMMICOLEPIDIDAE . . . . . . . . 2260-2261 hawaiiensis, Iso . . . . . . . . . 2141-2142-2143
Grammicolepis brachiusculus . . . . . . . . 2260 heberi, Caranx . . . . . . . . . . . . . . . . 2711
grammicus, Epinephelus . . . . . . . . . . . 2506 hedlandensis, Carangoides . . . 2691, 2693, 2702
GRAMMISTINI . . . . . . . . . . . . . . . . 2579 hedlandensis, Caranx . . . . . . . . . . . . . 2702
Grammoplites knappi . . . . . . . . . . . . 2399 hellmuthii, Cichlops . . . . . . . . . . . . . . 2571
Grammoplites scaber . . . . . . . . . . 2399-2400 Helmet gurnards . . . . . . . . . . . . . . . . 2283
grandis, Seriola . . . . . . . . . . . . . . . . 2740 helodes, Kalyptatherina . . . . . . . . . . . . 2144
Greasy grouper . . . . . . . . . . . . . . . . 2526 helodes, Pseudomugil . . . . . . . . . . . . . 2144
Greater amberjack . . . . . . . . . . . . . . 2738 helvola, Uraspis . . . . . . . . . . . . . 2754-2755
Greenback mullet . . . . . . . . . . . . . . . 2093 helvolus, Caranx . . . . . . . . . . . . . . . 2754
Greenback scad . . . . . . . . . . . . . . . . 2750
hemigymnostethus, Carangoides . . . . . . . 2699
gregoryi, Notograptus . . . . . . . . . . . . 2586 hemilepidota, Scorpaena . . . . . . . . . . . 2335
Grey goblinfish . . . . . . . . . . . . . . . . 2319
Grondin aile bleue . . . . . . . . . . . . . . . 2364
Heminodus philippinus . . . . . . . . . . . . 2367
HEMIRAMPHIDAE . . . . . . . 2151, 2162, 2180
Grondin aile tachetée . . . . . . . . . . . . . 2368
Grondin encré . . . . . . . . . . . . . . . . . 2373 Hemiramphus . . . . . . . . . . . . . . . . . 2188
Grondin volant cocarde . . . . . . . . . . . . 2286 Hemiramphus archipelagicus . . . . . . . . 2186
Grondin volant étoilé . . . . . . . . . . . . . 2289 Hemiramphus commersonii . . . . . . . . . . 2187
Grondin volant oriental . . . . . . . . . . . . 2287 Hemiramphus far . . . . . . . . . . . 2187, 2193
Groppos . . . . . . . . . . . . . . . . . . . . 2553 Hemiramphus gaimardi . . . . . . . . . . . 2191
Groupers . . . . . . . . . . . . . . . . . . . 2442 Hemiramphus lutkei . . . . . . . . . . . . . 2188
gruzovi, Inimicus . . . . . . . . . . . . . . . 2316 Hemiramphus marginatus . . . . . . 2186, 2188
Guam scorpionfish . . . . . . . . . . . . . . 2336 Hemiramphus robustus . . . . . . . . . . . . 2193
guamensis, Scorpaenodes . . . . . . 2336, 2339 Hemirhamphodon . . . . . . . . . . . . . . 2181
guentheri, Setarches . . . . . . . . . . . . . 2347 hemisticta, Otohime . . . . . . . . . . . . . . 2373
guezei, Myripristis (Holotrachys) . . . . . . 2248 hemisticta, Pterygotrigla . . . . 2360, 2373, 2380
Gurnards . . . . . . . . . . . . . . . . . . . 2359 heniochus, Epinephelus . . . . . . . 2496, 2502
guttatus, Notograptus . . . . . . . . . . . . 2586 herklotsi, Leucoglossa . . . . . . . . . . . . . 2756
guttulatus, Serranus . . . . . . . . . . . . . . 2528 Herring scad . . . . . . . . . . . . . . . . . . 2690
Gymnapogon . . . . . . . . . . . . . . . . . 2602 heterocheilus, Crenimugil . . . . . . . . . . 2084
Heteropriacanthus cruentatus .
. . . 2595, 2599 Hypoatherina . . . 2116, 2120, 2133-2134, 2138
heterurus, Parapterois . . . . .
. . . . . . . 2326 Hypoatherina barnesi . . . . . . . . . . . . . 2133
hexagona, Myripristis . . . . .
. . . 2239, 2254 Hypoatherina crenolepis . . . . . . . . . . . 2134
hexagonatus, Epinephelus . . .
. . . 2500, 2503, Hypoatherina gobio . . . . . . . . . . . . . . 2136
2512, 2518, 2524 Hypoatherina ovalaua . . . . . . . . . . . . 2135
hexazona, Cypselurus . . . . . . . . . . . . 2174 Hypoatherina panatela . . . . . . . . . . . . 2139
hians, Ablennes . . . . . . . . . . . . . . . . 2154 Hypoatherina temminckii . . . 2123, 2133, 2136
Highfin coralgrouper . . . . . . . . . . . . . . 2535 Hypoatherina tropicalis . . . . . . . . . . . 2137
Highfin grouper . . . . . . . . . . . . . . . . 2509 Hypoatherina uisila . . . . . . . . . . 2133, 2136
HIPPOCAMPINAE . . . . . . . . . . . 2264-2265 Hypoatherina valenciennei . . . . . . 2134, 2138
Hippocampus histrix . . . . . . . . . . . . . 2272 Hypoatherina woodwardi . . . . . . . 2134, 2138
Hippocampus kuda . . . . . . . . . . . . . . 2272 hypodus, Decapterus . . . . . . . . . . . . . 2721
Hippocampus spp. . . . . . . . . . . . . . . 2265 Hypoplectrodes . . . . . . . . . . . . . . . . 2459
Hippocampus whitei . . . . . . . . . . . . . 2272 Hyporhamphus (Hyporhamphus) limbatus
hippos, Seriola . . . . . . . . . . . . . . . . 2739 . . . . . . . . . . . . . . . . . . . 2191, 2195
hippurus, Coryphaena . . . . . . . . . 2657-2658 Hyporhamphus (Hyporhamphus) melanopterus
hirsutus, Scorpaenodes . . . . . . . . . . . . 2337 . . . . . . . . . . . . . . . . . . . . . . . 2195
Hirundichthys albimaculatus . . . . . . . . 2177 Hyporhamphus (Hyporhamphus) neglectissimus
Hirundichthys oxycephalus . . . . . . . . . 2177 . . . . . . . . . . . . . . . . . . . . . . . 2195
Hirundichthys speculiger . . . . . . . . . . 2177 Hyporhamphus (Hyporhamphus) neglectus . 2195
histrix, Hippocampus . . . . . . . . . . . . . 2272 Hyporhamphus (Reporhamphus) acutus acutus
hoedtii, Epinephelus . . . . . . . . . . . . . 2494 . . . . . . . . . . . . . . . . . . . . . . . 2194
hoedtii, Malacanthus . . . . . . . . . . . . . 2647 Hyporhamphus (Reporhamphus) affinis . . . 2194
Holanthias . . . . . . . . . . . . . . . . . . 2459 Hyporhamphus (Reporhamphus) balinensis . 2194
HOLOCENTRIDAE . . . . . . . 2214, 2225, 2591 Hyporhamphus (Reporhamphus) dussumieri
Holocentrids . . . . . . . . . . . . . . 2225, 2551 . . . . . . . . . . . . . . . . . . . 2189, 2194
HOLOCENTRINAE . . . . . . . . . . . . . . 2225 Hyporhamphus (Reporhamphus) quoyi . . . . 2190
Holocentrum andamanense . . . . . . . . . . 2249 Hyporhamphus acutus pacificus . . . . . . . 2194
Holocentrus caudimaculatus . . . . . . . . . 2249 Hyporhamphus dussumieri . . . . . . . . . . 2190
Holocentrus opercularis . . . . . . . . . . . 2244 Hyporhamphus neglectissimus . . . . . . . . 2191
Holocentrus ruber . . . . . . . . . . . . . . . 2251 Hyporhamphus neglectus . . . . . . . . . . 2191
Holocentrus sammara . . . . . . . . . . . . . 2245 Hyporhamphus unifasciatus . . . . . . . . . 2191
Holocentrus spiniferus . . . . . . . . . . . . 2252
hololepis, Zenion . . . . . . . . . . . . . . . 2258 I
Holotrachys major . . . . . . . . . . . . . . 2247 igarashiensis, Cephalopholis . . . . . . . . . 2479
homosinensis, Epinephelus . . . . . . . . . . 2521 ignobilis, Caranx . . . . . . . . . . . . . . . 2712
Honeycomb grouper . . . . . . . . . . . . . 2512 Imposter trevally . . . . . . . . . . . . . . . . 2709
honoriae, Atherina . . . . . . . . . . . . . . 2130 imus, Ectreposebastes . . . . . . . . . . . . . 2314
honoriae, Craterocephalus . . . . . . . 2130-2131 incisa, Strongylura . . . . . . . . . . . . . . 2156
honoriae, Stenatherina . . . . . . . . . . . . 2130 Inconspicuous blue-eye . . . . . . . . . . . . 2111
HOPLICHTHYIDAE . . . . 2360, 2383, 2386, 2422 inconspicuus, Pseudomugil . . . . . . 2110-2111
Hoplichthys haswelli . . . . . . . . . . . . . 2422 incredibilis, Dikellorhynchus . . . . . . . . . 2647
hoplites, Dixiphichthys . . . . . . . . . . . . 2374 Indian pompano . . . . . . . . . . . . . . . . 2749
hoplites, Pterygotrigla . . . . . . . . 2360, 2374 Indian scad . . . . . . . . . . . . . . . . . . 2722
Hoplolatilus . . . . . . . . . . . 2631-2632, 2637 Indian threadfish . . . . . . . . . . . . . . . . 2685
Hoplostethus atlanticus . . . . . . . . . . . 2215 indica, Alectis . . . . . . . . . . . . . . . . . 2685
Hornlip mullet . . . . . . . . . . . . . . . . . 2100 indicus, Platycephalus . . . . . . . . . 2408-2409
horrida, Synanceia . . . . . . . . . . . . . . 2348 Indonesian flyingfish . . . . . . . . . . . . . . 2171
Horseheads . . . . . . . . . . . . . . . . . . 2630 Inegocia harrisii . . . . . . . . . . . . . . . 2401
Hound needlefish . . . . . . . . . . . . . . . 2160 Inegocia isacanthus . . . . . . . . . . . . . . 2402
Howella . . . . . . . . . . . . . . . . . . . . 2436 Inegocia japonica . . . . . . . . . . . . . . . 2402
Howella brodiei . . . . . . . . . . . . . . . . 2612 ingenuua, Sillago . . . . . . . . . . . . . . . 2624
HOWELLIDAE . . . . . . . . . . . . . . . . 2436 Inimicus . . . . . . . . . . . . . . . . . . . . 2316
howlandi, Epinephelus . . . . . . . . 2508, 2544 Inimicus caledonicus . . . . . . . . . . . . . 2315
hullianus, Caranx . . . . . . . . . . . . . . . 2755 Inimicus cuvieri . . . . . . . . . . . . . . . . 2316
humerosus, Carangoides . . . . . . . . . . . 2703 Inimicus didactylus . . . . . . . . . . . . . . 2316
humerosus, Caranx . . . . . . . . . . . . . . 2703 Inimicus gruzovi . . . . . . . . . . . . . . . . 2316
Humpback grouper . . . . . . . . . . . . . . 2484 Inimicus japonicus . . . . . . . . . . . . . . 2316
Hunchbacked goblinfish . . . . . . . . . . . . 2351 Inimicus sinensis . . . . . . . . . . . . . . . 2317
hutchinsi, Cociella . . . . . . . . . . . . . . 2392 Inimicus smirnovi . . . . . . . . . . . . . . . 2316
Hyperoglyphe . . . . . . . . . . . . . . . . . 2661 insularum, Atherina . . . . . . . . . . . . . . 2124
insularum, Atherinomorus . . . . . . . . . . 2124 Jurel lentejuela . . . . . . . . . . . . . . . . . 2694

Intermediate flyingfish . . . . . . . . . . . . . 2171 Jurel malabárico . . . . . . . . . . . . . . . . 2704
Intermediate sillago . . . . . . . . . . . . . . 2625 Jurel manchado . . . . . . . . . . . . . . . . 2699
intermedius, Cheilopogon . . . . . . . . . . 2171 Jurel maquillado . . . . . . . . . . . . . . . . 2707
intermedius, Sillago . . . . . . . . . . . . . 2625 Jurel negro . . . . . . . . . . . . . . . . . . . 2713
iop, Scorpaenopsis . . . . . . . . . . . . . . 2341 Jurel paia . . . . . . . . . . . . . . . . . . . 2756
Iracundus signifer . . . . . . . . . . . . . . 2317 Jurel pardo . . . . . . . . . . . . . . . . . . . 2708
ire, Carangoides . . . . . . . . . . . . . . . 2708 Jurel peinero . . . . . . . . . . . . . . . . . . 2753
ire, Caranx . . . . . . . . . . . . . . . . . . 2708 Jurel postillón . . . . . . . . . . . . . . . . . 2705
irroratus, Epinephelus . . . . . . . . 2504, 2520 Jurel rabo amarillo . . . . . . . . . . . . . . . 2692
isacanthus, Inegocia . . . . . . . . . . . . . 2402 Jurel sable . . . . . . . . . . . . . . . . . . . 2732
Jurel saltador . . . . . . . . . . . . . . . . . 2731
ishikawai, Caranx . . . . . . . . . . . . . . . 2713
Jurel saltarín . . . . . . . . . . . . . . . . . . 2734
Island trevally . . . . . . . . . . . . . . . . . 2706
Jurel sap-sap . . . . . . . . . . . . . . . . . 2693
Iso flosmaris . . . . . . . . . . . . . . . . . 2141 Jurel subari . . . . . . . . . . . . . . . . . . 2687
Iso hawaiiensis . . . . . . . . . . 2141-2142-2143 Jurel tapir . . . . . . . . . . . . . . . . . . . 2696
Iso nesiotes . . . . . . . . . . . . . . . . . . 2143 Jurel tille . . . . . . . . . . . . . . . . . . . . 2717
Iso rhothophilus . . . . . . . . . . . . . . . 2141 Jurel volantín . . . . . . . . . . . . . . . . . 2755
Iso spp. . . . . . . . . . . . . . . . . . . . . 2140 Jurel voraz . . . . . . . . . . . . . . . . . . . 2716
ISONIDAE . . . . . . . . . . . . . . . . . . . 2140
K
J Kai soldierfish . . . . . . . . . .
. . . . . . . 2248
Jacks . . . . . . . . . . . . . . . . . 2655, 2659 kaianus, Ostichthys . . . . . . .
. . . . . . . 2248
jacobeus, Decapterus . . . . . . . . . . . . . 2719 Kaiwarinus equula . . . . . . .
. . . . . . . 2698
Jaggedhead armoured gurnard . . . . . . . . 2366 kalla, Alepes . . . . . . . . . . .
. . . . . . . 2688
Japanese flathead . . . . . . . . . . . . . . . 2402 kalla, Caranx . . . . . . . . . .
. . . . . . . 2688
Japanese soldierfish . . . . . . . . . . . . . 2247 Kalyptatherina helodes . . . . .
. . . . . . . 2144
japonica, Inegocia . . . . . . . . . . . . . . 2402 Kanda . . . . . . . . . . . . . .
. . . . . . . 2104
japonicus, Branchiostegus . 2632, 2641-2643-2644 kandavensis, Mugil . . . . . . .
. . . . . . . 2104
japonicus, Callanthias . . . . . . . . . . . . 2555 katoptron, Cheilopogon . . . . .
. . . . . . . 2171
japonicus, Cichlops . . . . . . . . . . . . . . 2571 kauderni, Pterapogon . . . . . .
. . . . . . . 2609
japonicus, Cookeolus . . . . . . . . . . . . . 2599 kauffmani, Notograptus . . . .
. . . . . . . 2586
japonicus, Inimicus . . . . . . . . . . . . . . 2316 Keeltail needlefish . . . . . . . .
. . . . . . . 2155
japonicus, Lateolabrax . . . . . 2429, 2431, 2444 kelaartii, Mugil . . . . . . . . .
. . . . . . . 2106
japonicus, Mugil . . . . . . . . . . . . . . . 2096 kelloggi, Scorpaenodes . . . . .
. . . . . . . 2337
japonicus, Ostichthys . . . . . . . . . . . . . 2247 kendalli, Cephalopholis . . . . .
. . . . . . . 2477
javanicus, Mugil . . . . . . . . . . . . . . . 2093 kikaii, Pseudochromis . . . . . .
. . . . . . . 2573
Jawfishes . . . . . . . . . . . . . . . . . . . 2588 kiliche, Decapterus . . . . . . .
. . . . . . . 2722
jerdoni, Mugil . . . . . . . . . . . . . . . . . 2093 kleinii, Alepes . . . . . . . . . .
. . . . . . . 2688
jordani, Carangoides . . . . . . . . . . . . . 2706 klunzingeri, Allanetta . . . . . .
. . . . . . . 2133
jordani, Caranx ferdau . . . . . . . . . . . . 2706 klunzingeri, Liza . . . . . . . .
. . . . . . . 2085
jordani, Dactyloptena . . . . . . . . . . . . 2285 knappi, Grammoplites . . . . .
. . . . . . . 2399
jugosa, Ambiserrula . . . . . . . . . . . . . 2391 kohleri, Epinephelus . . . . . . .
. . . . . . . 2494
Jumping halfbeak . . . . . . . . . . . . . . . 2186 krefftii, Arrhamphus sclerolepis . . . . . . . 2193
Jurel aliblanco . . . . . . . . . . . . . . . . . 2698
Jurel alinegro . . . . . . . . . . . . . . . . . 2689
kuda, Hippocampus . . . . . . . . . . . . . . 2272
KUHLIIDAE . . . . . . . . . . . . . . . . . . 2444
Jurel arenque . . . . . . . . . . . . . . . . . 2690
See also Vol. 5
Jurel balo . . . . . . . . . . . . . . . . . . . 2701
Jurel bronceado . . . . . . . . . . . . . . . . 2715 Kumococius detrusus . . . . . . . . . . . . . 2403
Jurel carsia . . . . . . . . . . . . . . . . . . 2733 Kumococius rodericensis . . . . . . . . . . . 2403
Jurel centellante . . . . . . . . . . . . . . . . 2700 kumu, Chelidonichthys . . . . . 2364-2365, 2368
Jurel cocolí . . . . . . . . . . . . . . . . . . 2711 kumu, Trigla . . . . . . . . . . . . . . . . . . 2364
Jurel costero . . . . . . . . . . . . . . . . . . 2695 kuntee, Myripristis . . . . . . . . . . . . . . 2241
Jurel crepúsculo . . . . . . . . . . . . . . . . 2697 kurroides akaadsi, Decapterus . . . . . . . . 2718
Jurel de aleta azul . . . . . . . . . . . . . . . 2714 kurroides, Decapterus . . . . . . . . . . . . 2718
Jurel dentón . . . . . . . . . . . . . . . . . . 2730 Kutaflammeo opercularis . . . . . . . . . . . 2244
Jurel dorado . . . . . . . . . . . . . . . . . . 2725 Kutaflammeo sammara . . . . . . . . . . . . 2245
Jurel gigante . . . . . . . . . . . . . . . . . . 2712
Jurel hocicón . . . . . . . . . . . . . . . . . 2702 L
Jurel impostor . . . . . . . . . . . . . . . . . 2709 labiosus, Crenimugil . . . . . . . . . . . . . 2100
Jurel isleño . . . . . . . . . . . . . . . . . . 2706 labiosus, Oedalechilus . . . . . . . . . . . . 2100
Jurel lengua blanca . . . . . . . . . . . . . . 2754 labiosus, Plicomugil . . . . . . . . . . . . . . 2100
Labracinus . . . . . . . . . . . . . . . . . . 2571 lineatus, Centroberyx . . . . . . . . . . . . . 2224

Labracinus cyclophthalmus . . . . . 2571, 2574 lineatus, Labracinus . . . . . . . . . 2571, 2574
Labracinus lineatus . . . . . . . . . 2571, 2574 lineatus, Serranus . . . . . . . . . . . . . . . 2529
LABRIDAE . . . . . . . . . . . . . . 2632, 2648 Lined silverside . . . . . . . . . . . . . . . . 2126
See also Vol.6 Liocranium praepositum . . . . . . . . . . . 2318
Labroides dimidiatus . . . . . .
. . . . . . . 2648 Lionfish . . . . . . . . . . . . . . . . . . . . . 2332
LACTARIIDAE . . . . . . . . .
2437, 2649, 2661 Lionfishes . . . . . . . . . . . . . . . 2291, 2331
Lactarius delicatulus . . . . . .
. . . . . . . 2649 Liopropoma . . . . . . . . . . . . . . . . . . 2460
Lactarius lactarius . . . . . . .
. . . . . . . 2649 LIOPROPOMATINI . . . . . . . . . . . . . . 2442
lactarius, Lactarius . . . . . . .
. . . . . . . 2649 liorhynchus, Peristedion . . . . . . . . . . . 2370
lacunosus, Atherinomorus . . .
2113, 2120, 2123- Lioscorpius . . . . . . . . . . . . . . . . . . 2291
2125, 2127, 2137 Lioscorpius longiceps . . . . . . . . . . . . . 2319
laevis, Plectropomus . . . . . . . . . . . . . 2532 LIPARIDAE . . . . . . . . . . . . . . . . . . 2428
lajang, Decapterus . . . . . . . . . . 2720, 2722 Lisa chiraya . . . . . . . . . . . . . . . . . . 2108
lalandi, Seriola . . . . . . . . . . . . . . . . 2740 Lisa de aleta larga . . . . . . . . . . . . . . . 2103
lanceolatus, Epinephelus . . . . . . . 2505, 2526 Lisa de mancha azul . . . . . . . . . . . . . . 2107
lanceolatus, Promicrops . . . . . . . . . . . 2505 Lisa diamante . . . . . . . . . . . . . . . . . 2086
Lanternbellies . . . . . . . . . . . . . . . . . 2436 Lisa godeya . . . . . . . . . . . . . . . . . . 2088
Lappies . . . . . . . . . . . . . . . . . . . . 2344 Lisa labride . . . . . . . . . . . . . . . . . . 2083
Largescale flyingfish . . . . . . . . . . . . . . 2174 Lisa lomo verde . . . . . . . . . . . . . . . . 2093
Largescale mullet . . . . . . . . . . . . . . . 2088 Lisa mopiro . . . . . . . . . . . . . . . . . . 2094
Largescale sillago . . . . . . . . . . . . . . . 2627 Lisa morruda . . . . . . . . . . . . . . . . . . 2100
Largescaled gurnard . . . . . . . . . . . . . 2382 Lisa ojo de perdiz . . . . . . . . . . . . . . . 2104
Largescaled spiny flathead . . . . . . . . . . 2406 Lisa otomebora . . . . . . . . . . . . . . . . 2089
Largespined flathead . . . . . . . . . . . . . 2416 Lisa pardete . . . . . . . . . . . . . . . . . . 2096
Largespotted dart . . . . . . . . . . . . . . . 2747 Lisa rabo azul . . . . . . . . . . . . . . . . . 2102
Lateolabrax japonicus . . . . . 2429, 2431, 2444 Lisa tade . . . . . . . . . . . . . . . . . . . . 2091
Lates calcarifer . . . . . . . . . . . . 2429, 2431 littoralis, Pomacentrus . . . . . . . . . . . . 2349
latifasciatus, Epinephelus . . . . . . . . . . 2506 littoralis, Scorpaenodes . . . . . . . . . 2338-2339
LATILINAE . . . . . . . . . . . . . . . . . . 2632 Livebearing toothcarps . . . . . . . . . . . . 2199
LATINAE . . . . . . . . . . . . . . . . . . . 2429 livingstonei, Notograptus . . . . . . . . . . . 2586
latovittatus, Malacanthus . . . . . . . 2647-2648 Liza . . . . . . . . . . . . . . . . . . . . . . 2070
Leaf goblinfish . . . . . . . . . . . . . . . . . 2352 Liza affinis . . . . . . . . . . . . . . . . . . 2085
Leaf scorpionfish . . . . . . . . . . . . . . . 2349 Liza alata . . . . . . . . . . . . 2086, 2088-2089
leiura, Strongylura . . . . . . . . . . . . . . 2157 Liza argentea . . . . . . . . . . . . . 2087, 2092
leiuroides, Belone . . . . . . . . . . . . . . . 2156 Liza carinata . . . . . . . . . . . . . . . . . 2085
leiurus, Belone . . . . . . . . . . . . . . . . 2157 Liza klunzingeri . . . . . . . . . . . . . . . . 2085
Leopard coralgrouper . . . . . . . . . . . . . 2533 Liza macrolepis . . . . . . . . . 2088-2090, 2093
Leopard hind . . . . . . . . . . . . . . . . . 2541 Liza melinoptera . . 2086, 2088-2089-2090, 2094
leopardinus, Plectropoma . . . . . . . . . . 2533 Liza oligolepis . . . . . . . . . . . . . . . . . 2090
leopardus, Cephalopholis . . . . . . . . . . 2541 Liza parmata . . . . . . . 2089-2090, 2093-2094
leopardus, Plectropomus . . . . . . . . 2532-2533 Liza parsia . . . . . . . . . . . . 2088, 2091, 2093
Lepidocybium . . . . . . . . . . . . . . . . . 2660 Liza planiceps . . . . . . . . . . . . . 2091, 2093
Lepidotrigla spiloptera . . . . . . . . . . . . 2368 Liza ramsayi . . . . . . . . . . . . . . 2087, 2092
leptacanthus, Bovitrigla . . . . . . . . . . . . 2375 Liza subviridis . . . . . . . . . . 2088-2091, 2093
leptacanthus, Pterygotrigla . . . . . . . . . 2375 Liza tade . . . . . . . . . . . . . . . . . . . . 2091
leptacanthus, Trigla . . . . . . . . . . . . . . 2375 Liza vaigiensis . . . . . . . 2089, 2091, 2093-2094
leptolepis, Selaroides . . . . . . . . . . . . . 2737 Liza waigiensis . . . . . . . . . . . . . . . . 2094
Leptosynanceia asteroblepa . . . . . . . . . 2318 Lobed river mullet . . . . . . . . . . . . . . . 2082
leuciscus, Chaenomugil . . . . . . . . . . . . 2099 LOBOTIDAE . . . . . . . . . . . . . . . . . . 2444
leuciscus, Neomyxus . . . . . . . . . . . . . 2099 See also Vol. 5
leucogaster, Richardsonichthys . . . . . . . 2333 Longarm mullet . . . . . . . . . . . . . . . . 2103
Leucoglossa candens . . . . . . . . . . . . . 2754 Longfin bulleye . . . . . . . . . . . . . . . . . 2599
Leucoglossa herklotsi . . . . . . . . . . . . . 2756 Longfin grouper . . . . . . . . . . . . . . . . 2518
leucogrammicus, Anyperodon . . . . . . . . 2475 Longfin mullet . . . . . . . . . . . . . . . . . 2103
leucozona, Abudefduf . . . . . . . . . . . . 2349 Longfin trevally . . . . . . . . . . . . . . . . . 2693
limbatus, Hyporhamphus (Hyporhamphus) Longfingered scorpionfish . . . . . . . . . . . 2336
. . . . . . . . . . . . . . . . . . . 2191, 2195 Longfinned halfbeak . . . . . . . . . . . . . . 2193
Limpidwing flyingfish . . . . . . . . . . . . . 2173 Longfinned mullet . . . . . . . . . . . . . . . 2106
lineata, Atherina . . . . . . . . . . . . . . . 2126 Longfins . . . . . . . . . . . . . . . . . . . . 2578
lineatus, Atherinomorus . . . . . . . 2123, 2126 longiceps, Lioscorpius . . . . . . . . . . . . 2319
longimanus, Mugil . . . . . . . . . . . . . . . 2106
longimanus, Setarches . . . . . . . . . . . . 2348 Macrurocyttus . . . . . . . . . . . . . . . . . 2258

longipinna, Dampieria . . . . . . . . . . . . 2572 Macrurocyttus acanthopodus . . . . . . . . 2258
longispinis, Epinephelus . . . . . . . . . . . 2507 macrurus, Alepes . . . . . . . . . . . . . . . 2690
longispinis, Paracentropogon . . . . . . . . 2325 macrurus, Caranx . . . . . . . . . . . . . . . 2690
Longnose gurnard . . . . . . . . . . . . . . . 2376 maculata burra, Sillago . . . . . . . . . . . . 2621
Longnose trevally . . . . . . . . . . . . . . . 2696 maculata maculata, Sillago . . . . . . . . . . . 2625
Longrakered trevally . . . . . . . . . . . . . . 2753 maculata, Blanchardia . . . . . . . . . . . . 2586
Longsnout flathead . . . . . . . . . . . . . . 2420 maculata, Sillago . . . . . . . . 2617, 2621, 2628
Longsnout stinger . . . . . . . . . . . . . . . 2316 maculata, Sillago maculata . . . . . . . . . 2625
Longspine grouper . . . . . . . . . . . . . . 2507 maculatum var. A, Plectropoma . . . . . . . 2536
Longspine snipefish . . . . . . . . . . . . . . 2280 maculatum var. D, Plectropoma . . . . . . . 2536
Longtooth grouper . . . . . . . . . . . . . . . 2490 maculatum, Plectropoma . . . . . . . 2531, 2536
louti, Variola . . . . . . . . . . . . . . 2538-2539 maculatus, Epinephelus . . . . . . . 2507, 2509
lugubris, Caranx . . . . . . . . . . . . . . . 2713 maculatus, Plectropoma . . . . . . . 2533, 2536
Luminous deepwater cardinalfish . . . . . . . 2613 maculatus, Plectropomus . . . . . . . 2532, 2534
lundini, Decapterus . . . . . . . . . . . . . . 2692 maculatus, Pranesus . . . . . . . . . . . . . 2125
lunulata, Pterois . . . . . . . . . . . 2329, 2331 Mafou . . . . . . . . . . . . . . . . . . . . . 2655
lutea, Sillago . . . . . . . . . . . . . . . . . 2626 magniscuttis, Epinephelus . . . 2496, 2510, 2515
LUTJANIDAE . . . . . . . 2438-2439, 2443, 2633 major, Holotrachys . . . . . . . . . . . . . . 2247
See also Vol. 5 majusculus, Pseudomugil . . . . . . 2112, 2144
Lutjanids . . . . . . . . . . . . . . . . . . . . 2438 Malabar grouper . . . . . . . . . . . . . . . . 2511
Lutke’s halfbeak . . . . . . . . . . . . . . . . 2188 Malabar trevally . . . . . . . . . . . . . . . . 2704
lutkei, Hemiramphus . . . . . . . . . . . . . 2188 malabaricus, Carangoides . . . . . . 2704, 2709
Luzonichthys . . . . . . . . . . . . . . . . . 2461 malabaricus, Caranx . . . . . . . . . . . . . 2704
lysan, Scomberoides . . . . . . . . . . 2731-2732 malabaricus, Epinephelus . . . 2493, 2511, 2526
MALACANTHIDAE . . . . . . . . . . 2630, 2632
M MALACANTHINAE . . . . . . . 2631-2632, 2636
Macarela alicorta . . . .
. . . . .
. . . . . . 2720 Malacanthus . . . . . . . . . . . . . 2631, 2639
Macarela bandoneada .
. . . . .
. . . . . . 2688 Malacanthus brevirostris . . . . . . . . 2647-2648
Macarela caballa . . . .
. . . . .
. . . . . . 2719 Malacanthus hoedtii . . . . . . . . . . . . . . 2647
Macarela indica . . . . .
. . . . .
. . . . . . 2722 Malacanthus latovittatus . . . . . . . . 2647-2648
Macarela maouana . . .
. . . . .
. . . . . . 2718 Malacanthus parvipinnis . . . . . . . . . . . 2647
Macarela rabo colorado .
. . . . .
. . . . . . 2723
Macarela salmón . . . .
. . . . .
. . . . . . 2724
malam, Alepes . . . . . . . . . . . . . . . . . 2689
Macarela torpedo . . . .
. . . . .
. . . . . . 2726 malam, Caranx . . . . . . . . . . . . . . . . 2689
macarellus, Decapterus . . . . .
. . . 2719-2721 malayanus, Myxus . . . . . . . . . . . . . . . 2095
maccullochi, Atherion . . . . . .
. . . . . . 2118 mandibularis, Caranx . . . . . . . . . . . . . 2753
Mackerel scad . . . . . . . . . . .
. . . . . . 2719 mandibularis, Ulua . . . . . . . . . . . . . . 2753
Manyspotted flyingfish . . . . . . . . . . . . . 2172
macracantha, Dactyloptena . . .
. . . . . . 2286
Maori grouper . . . . . . . . . . . . . . . . . 2528
macracanthus, Priacanthus . . .
. . 2591, 2597
Marao lisero . . . . . . . . . . . . . . . . . . 2160
macracanthus, Suggrundus . . .
. . . . . . 2416 Marao ojón . . . . . . . . . . . . . . . . . . . 2159
macrocheilos, Mugil . . . . . . .
. . . . . . 2083 Marble grouper . . . . . . . . . . . . . . . . 2516
macrochir, Scorpaenopsis . . . .
. . . . . . 2343 marginatus, Caranx . . . . . . . . . . . . . . 2716
macrolepidota, Pterygotrigla . . .
. . . . . . 2382 marginatus, Hemiramphus . . . . . . 2186, 2188
macrolepidota, Uradia . . . . . .
. . . . . . 2382 Margined flyingfish . . . . . . . . . . . . . . . 2170
macrolepidotus, Mugil . . . . . .
. . . . . . 2095 Marignan à oeillères . . . . . . . . . . . . . . 2238
macrolepis, Liza . . . . . . . .
2088-2090, 2093 Marignan aile noire . . . . . . . . . . . . . . 2244
macrolepis, Onigocia . . . . . .
. . . . . . . 2404 Marignan ardoisé . . . . . . . . . . . . . . . 2241
macrolepis, Sillago . . . . . . .
. . . . 2626-2627 Marignan chocolat . . . . . . . . . . . . . . . 2250
macrops, Epigonus . . . . . . .
. . . . . . . 2613 Marignan cuirassé . . . . . . . . . . . . . . . 2246
MACRORHAMPHOSIDAE . . .
. . . . 2280-2281 Marignan ombré . . . . . . . . . . . . . . . . 2236
Macrorhamphosus gracilis . . .
. . . . . . . 2280 Marignan pomme de pin . . . . . . . . . . . . 2242
Macrorhamphosus scolopax . .
. . . . . . . 2280 Marignan potelé . . . . . . . . . . . . . . . . 2239
Macrorhamphosus velitaris . . .
. . . . . . . 2280 Marignan rouge et argent . . . . . . . . . . . 2249
macrorhynchus, Dixiphistes . .
. . . . . . . 2376 Marignan rouget . . . . . . . . . . . . . . . . 2251
macrorhynchus, Parapterygotyrigla . . . . . 2376 Marignan sabre . . . . . . . . . . . . . . . . 2252
macrorhynchus, Pterygotrigla . . . . 2360, 2376 Marignan tacheté . . . . . . . . . . . . . . . 2245
macrosoma, Decapterus . . . . 2719-2720, 2722 Marignan violacé . . . . . . . . . . . . . . . . 2243
macrospilos, Epinephelus . . . 2500, 2503, 2508- Marine betta . . . . . . . . . . . . . . . . . . 2578
2509, 2512, 2518, 2524 Marine sailfin silverside . . . . . . . . . . . . 2144
MACRUROCYTTIDAE . . . . . . . . . 2257-2258 marjoriae, Craterocephalus . . . . . . . . . 2132
Marlinsucker . . . . . . . .. . .
. . . 2652 . . . . Mero areolado . . . . . . . . . . . . . . . . . 2487
marmoratus, Sebastiscus . . . .
. . . 2347 . . . . Mero aspero . . . . . . . . . . . . . . . . . . 2548
Marquesan grouper . . . . . . .
. . . 2504 . . . . Mero azul . . . . . . . . . . . . . . . . . . . 2494
maruadsi, Decapterus . . . . .
. . . 2722 . . . . Mero bacalao . . . . . . . . . . . . . . . . . 2510
Masked grouper . . . . . . . . .
. . . 2530 . . . . Mero bandeado . . . . . . . . . . . . . . . . 2500
Matajuel bleu . . . . . . . . . .
. . . 2648 . . . . Mero banderilla . . . . . . . . . . . . . . . . 2499
Matajuelo azul . . . . . . .. . .
. . . 2648 . . . . Mero celestrial . . . . . . . . . . . . . . . . . 2533
mate, Atule . . . . . . . . . . .
. . . 2692 . . . . Mero colmenar . . . . . . . . . . . . . . . . . 2544
mate, Caranx . . . . . . . . . .
. . . 2692 . . . . Mero cometa . . . . . . . . . . . . . . . . . . 2513
maunaloae, Symphysanodon . .
. . . 2440 . . . . Mero con pintas . . . . . . . . . . . . . . . . 2534
mauritiana, Sebastapistes . . .
. . . 2345 . . . . Mero cuatro monturas . . . . . . . . . . . . . 2524
mcadamsi, Parascorpaena . . .
. . . 2326 . . . . Mero de coral . . . . . . . . . . . . . . . . . 2543
mccullochi perpulcher, Pseudochromis . . . 2570 Mero de Darwin . . . . . . . . . . . . . . . . 2543
mccullochi, Pseudochromis . . . . . . . . . . 2570 Mero de las piedras . . . . . . . . . . . . . . 2498
Mero de lunares . . . . . . . . . . . . . . . . 2545
mccullochi, Trachurus . . . . . . . . . . . . 2751 Mero de margenes palidos . . . . . . . . . . 2542
Medregal coronado . . . . . . . . . . . . . . 2738
Mero de ocho bandas . . . . . . . . . . . . . 2545
Medregal limón . . . . . . . . . . . . . . . . 2741
Mero de pintas amarillas . . . . . . . . . . . 2546
Medregal listado . . . . . . . . . . . . . . . . 2742
Mero de pintas gemelas . . . . . . . . . . . . 2542
Medregal rabo amarillo . . . . . . . . . . . . 2740
Mero de pintas naranjas . . . . . . . . . . . . 2493
medurensis, Epinephelus . . . . . . . . . . . 2509 Mero de pintas rojas . . . . . . . . . . . . . . 2485
medusicola, Caranx . . . . . . . . . . . . . . 2714 Mero de seis bandas . . . . . . . . . . . . . 2522
megacephalus, Ratabulus . . . . . . . . . . . 2410 Mero de siete bandas . . . . . . . . . . . . . 2497
megachir, Epinephelus . . . . . . . . . . . . 2518 Mero diente largo . . . . . . . . . . . . . . . 2490
Megalaspis cordyla . . . . . . . . . . . . . . 2726 Mero disfrazado . . . . . . . . . . . . . . . . 2516
megalepis, Neomerinthe . . . . . . . . . . . 2322 Mero dorado . . . . . . . . . . . . . . . . . . 2537
MEGALOMYCTERIDAE . . . . . . . . 2207-2209 Mero elegante . . . . . . . . . . . . . . . . . 2475
megalops, Dixiphistops . . . . . . . . . . . . 2377 Mero embridado . . . . . . . . . . . . . . . . 2502
megalops, Parapterygotyrigla . . . . . . . . 2377 Mero ensillado . . . . . . . . . . . . . . . . . 2532
megalops, Phenacoscorpius . . . . . . . . . 2328 Mero errante . . . . . . . . . . . . . . . . . . 2536
megalops, Pterygotrigla . . . . . . . . . . . 2377 Mero espaldaron . . . . . . . . . . . . . . . . 2544
Melamphaes danae . . . . . . . . . . . . . . 2201 Mero espigón . . . . . . . . . . . . . . . . . 2507
Melamphaes simus . . . . . . . . . . . . . . 2201 Mero jorobado . . . . . . . . . . . . . . . . . 2484
MELAMPHAIDAE . . . . . . . . . . . 2201, 2203 Mero lanceolado . . . . . . . . . . . . . . . . 2505
melampygus, Caranx . . . . . . . . . . 2714-2715 Mero luna creciente . . . . . . . . . . . . . . 2539
melanoleucus, Plectropomus . . . . . . . . . 2532 Mero lutra . . . . . . . . . . . . . . . . . . . 2526
melanoptera, Alepes . . . . . . . . . . . . . 2689 Mero malabárico . . . . . . . . . . . . . . . . 2511
melanopterus, Hyporhamphus (Hyporhamphus) Mero manchado . . . . . . . . . . . . . . . . 2501
. . . . . . . . . . . . . . . . . . . . . . . 2195 Mero Maori . . . . . . . . . . . . . . . . . . . 2528
melanostethos, Caranx . . . . . . . . . . . . 2708 Mero marquesano . . . . . . . . . . . . . . . 2504
melanostictus, Myripristis . . . . . . . . . . 2239 Mero medialuna . . . . . . . . . . . . . . . . 2521
melanostigma, Dampieria . . . . . . . . . . 2571 Mero medioluto . . . . . . . . . . . . . . . . 2489
melanostigma, Epinephelus . . 2512, 2524, 2544 Mero mielero . . . . . . . . . . . . . . . . . . 2503
Mero montado . . . . . . . . . . . . . . . . . 2495
melanotaenia, Cichlops . . . . . . . . . . . . 2571 Mero montura negra . . . . . . . . . . . . . . 2544
melanotus, Tylosurus acus . . . . . . . . . . 2159 Mero nevero . . . . . . . . . . . . . . . . . . 2491
Melapedalion . . . . . . . . . . . . . 2162, 2180 Mero nubifero . . . . . . . . . . . . . . . . . 2514
Melapedalion breve . . . . . . . . . . . . . . 2195 Mero nublado . . . . . . . . . . . . . . . . . 2543
meleagris, Sebastes . . . . . . . . . . . . . . 2336 Mero oleado . . . . . . . . . . . . . . . . . . 2523
melinoptera, Liza . . 2086, 2088-2089-2090, 2094 Mero ondulado . . . . . . . . . . . . . . . . . 2529
mentalis, Caranx . . . . . . . . . . . . . . . 2753 Mero ovalado . . . . . . . . . . . . . . . . . 2547
mentalis, Ulua . . . . . . . . . . . . . . . . 2753 Mero palido . . . . . . . . . . . . . . . . . . 2496
mento, Parexocoetus . . . . . . . . . . . . . 2178 Mero panal . . . . . . . . . . . . . . . . . . . 2512
Mercer’s tusked silverside . . . . . . . . . . . 2145 Mero paranjero . . . . . . . . . . . . . . . . 2530
merceri, Dentatherina . . . . . . . . . . . . 2145 Mero patata . . . . . . . . . . . . . . . . . . 2527
meristocystis, Oxyporhamphus . . . . . . . . 2192 Mero pintado . . . . . . . . . . . . . . . . . . 2492
Mero abanderado . . . . . . . . . . . . . . . 2506 Mero punteado . . . . . . . . . . . . . . . . . 2520
Mero acebrado . . . . . . . . . . . . . . . . 2519 Mero punteado blanco . . . . . . . . . . . . . 2517
Mero acertijo . . . . . . . . . . . . . . . . . 2545 Mero punteado negro . . . . . . . . . . . . . 2525
Mero aleta alta . . . . . . . . . . . . . . . . 2509 Mero punto y linea . . . . . . . . . . . . . . . 2515
Mero aleta larga . . . . . . . . . . . . . . . . 2518 Mero rabiblanco . . . . . . . . . . . . . . . . 2538
Mero alfombrado . . . . . . . . . . . . . . . 2508 Mero reticular . . . . . . . . . . . . . . . . . 2546
Mero amarillo . . . . . . . . . . . . . . . . . 2488 Mero rollizo . . . . . . . . . . . . . . . . . . 2546
Mero troncon . . . . . . . . . . . . . . . . . 2531 Mérou taches oranges . . .

. . . . . . . . . . 2493
Mero vela . . . . . . . . . . . . . . . . . . . 2535 Mérou tapis . . . . . . . .
. . . . . . . . . . 2508
Mérou à bandes . . . . . . . . . . . . . . . . 2506 Mérou troussequin . . . . .
. . . . . . . . . . 2495
Mérou á bout rouge . . . . . . . . . . . . . . 2520 Mérou zébré . . . . . . . .
. . . . . . . . . . 2519
Mérou à flocons . . . . . . . . . . . . . . . . 2514 Mérou-loche cacatois . . .
. . . . . . . . . . 2535
Mérou abeille . . . . . . . . . . . . . . . . . 2544 Mérou-loche vagabonde . .
. . . . . . . . . . 2536
Mérou aréolé . . . . . . . . . . . . . . . . . 2487 merra, Epinephelus . . . .
2500, 2503, 2512, 2524
Mérou bande . . . . . . . . . . . . . . . . . 2486 mesembrinus, Antipodocottus . . . . 2425, 2428
Mérou bleu . . . . . . . . . . . . . . . . . . 2494 meyeri, Pristigenys . . . . . . . . . . . . . . 2601
Mérou bord clair . . . . . . . . . . . . . . . . 2542 microchir, Caranx . . . . . . . . . . . . . . . . 2688
Mérou bord rouge . . . . . . . . . . . . . . . 2530 microdon, Epinephelus . . . . . . . . 2501, 2516
Mérou bossu . . . . . . . . . . . . . . . . . 2484 micronotatus, Serranus . . . . . . . . . . . . 2475
Mérou bride . . . . . . . . . . . . . . . . . . 2502 microphthalmus, Myripristis . . . . . . . . . 2243
Merou camouflage . . . . . . . . . . . . . . 2516 microprion, Cephalopholis . . . . . . 2477, 2541
Mérou comète . . . . . . . . . . . . . . . . . 2513 micropterus micropterus, Oxyporhamphus . 2196
Mérou corail . . . . . . . . . . . . . . . . . . 2543 micropterus similis, Oxyporhamphus . . . . 2196
Mérou curieux . . . . . . . . . . . . . . . . . 2545
Mérou d’or . . . . . . . . . . . . . . . . . . . 2537
micropterus, Caranx . . . . . . . . . . . . . 2754
Mérou Darwin . . . . . . . . . . . . . . . . . 2543 micropterus, Oxyporhamphus . . . . . . . . 2192
Mérou demi-lune . . . . . . . . . . . . . . . 2521 micropterus, Oxyporhamphus micropterus . 2196
Mérou demideuil . . . . . . . . . . . . . . . . 2489 microstoma, Atherinosoma . . . . . . . . . . 2130
Mérou dossard . . . . . . . . . . . . . . . . 2544 miles, Pterois . . . . . . . . . . . . . 2330, 2332
Mérou écharpe . . . . . . . . . . . . . . . . 2500 miliaris, Epinephelus . . . . . . . . . . . . . 2544
Mérou élégant . . . . . . . . . . . . . . . . . 2475 Milk trevally . . . . . . . . . . . . . . . . . . 2649
Mérou gâteau de cire . . . . . . . . . . . . . 2512 miniata var. C, Perca . . . . . . . . . . . . . 2536
Mérou gemine . . . . . . . . . . . . . . . . . 2542 miniata, Cephalopholis . . . . . 2477, 2480-2481
Mérou grandes écailles . . . . . . . . . . . . 2510 MINOINAE . . . . . . . . . . . . . . . . . . . 2293
Mérou haute voile . . . . . . . . . . . . . . . 2509 Minor scorpionfish . . . . . . . . . . . . . . . 2338
Mérou houleux . . . . . . . . . . . . . . . . 2523 minor, Scorpaenodes . . . . . . . . . . . . . 2338
Mérou huit raies . . . . . . . . . . . . . . . . 2545 Minous . . . . . . . . . . . . . . . . 2320, 2354
Mérou jaune . . . . . . . . . . . . . . . . . . 2488 Minous coccineus . . . . . . . . . . . . . . . 2320
Mérou lancéolé . . . . . . . . . . . . . . . . 2505 Minous monodactylus . . . . . . . . . . . . 2319
Mérou longues ailes . . . . . . . . . . . . . . 2518 Minous pusillus . . . . . . . . . . . . . . . . 2320
Mérou longues dents . . . . . . . . . . . . . 2490 Minous trachycephalus . . . . . . . . . . . . 2320
Mérou longues épines . . . . . . . . . . . . . 2507 Minous versicolor . . . . . . . . . . . . . . . 2321
Mérou loutre . . . . . . . . . . . . . . . . . . 2526 Mirapinna . . . . . . . . . . . . . . . . . . . 2208
Mérou malabare . . . . . . . . . . . . . . . . 2511 MIRAPINNIDAE . . . . . . . . . 2207-2208-2209
Mérou Maori . . . . . . . . . . . . . . . . . . 2528 mirifica, Pterapogon . . . . . . . . . . . . . 2609
Mérou Marquises . . . . . . . . . . . . . . . 2504 Mirrorwing flyingfish . . . . . . . . . . . . . . 2177
Mérou marron . . . . . . . . . . . . . . . . . 2501 miyakamii, Caranx . . . . . . . . . . . . . . 2688
Mérou mélifère . . . . . . . . . . . . . . . . 2503 moara, Serranus . . . . . . . . . . . . . . . . 2490
Mérou morse . . . . . . . . . . . . . . . . . 2515 mollis, Beryx . . . . . . . . . . . . . . . . . 2222
Mérou nébuleux . . . . . . . . . . . . . . . . 2543
Mérou ondulé . . . . . . . . . . . . . . . . . 2529
mombasae, Pterois . . . . . . . . . . . . . . 2330
MONOCENTRIDIDAE . . . . . . . . . . . . . 2214
Mérou oriflamme . . . . . . . . . . . . . . . 2499
Mérou ovale . . . . . . . . . . . . . . . . . . 2547 monocirrhus, Exocoetus . . . . . . . . . . . 2176
Mérou pâle . . . . . . . . . . . . . . . . . . 2496 monodactylus, Minous . . . . . . . . . . . . 2319
Mérou patate . . . . . . . . . . . . . . . . . 2527 mookalee, Trachinotus . . . . . . . . . . . . 2749
Mérou pintade . . . . . . . . . . . . . . . . . 2492 Moolgarda (Planiliza) ordensis . . . . . . . . 2086
Mérou pointe noirs . . . . . . . . . . . . . . 2525 Moontail bullseye . . . . . . . . . . . . . . . 2596
Mérou points blancs . . . . . . . . . . . . . . 2517 MORONIDAE . . . . . . . . . . 2429, 2431, 2444
Mérou quatre selles . . . . . . . . . . . . . . 2524 morrhua, Epinephelus . . . . . 2513, 2515, 2519
Mérou queue carrée . . . . . . . . . . . . . . 2531 morrisi, Atherina . . . . . . . . . . . . . . . 2125
Mérou réseau . . . . . . . . . . . . . . . . . 2546 mossambica, Parascorpaena . . . . . . . . . 2327
Mérou rocaille . . . . . . . . . . . . . . . . . 2498 moultoni, Scorpaena . . . . . . . . . . . . . 2326
Mérou rondelet . . . . . . . . . . . . . . . . 2546 Mozambique scorpionfish . . . . . . . . . . . 2327
Mérou rouge tacheté . . . . . . . . . . . . . 2485 Mud flathead . . . . . . . . . . . . . . . . . . 2391
Mérou rugueux . . . . . . . . . . . . . . . . 2548 Mud sillago . . . . . . . . . . . . . . . . . . . 2626
Mérou sellé . . . . . . . . . . . . . . . . . . 2532 Mugil alcocki . . . . . . . . . . . . . . . . . 2093
Mérou selle noir . . . . . . . . . . . . . . . . 2544 Mugil amarulus . . . . . . . . . . . . . . . . 2106
Mérou sept raies . . . . . . . . . . . . . . . 2497 Mugil australis . . . . . . . . . . . . . . . . 2087
Mérou six raies . . . . . . . . . . . . . . . . 2522 Mugil axillaris . . . . . . . . . . . . . . . . . 2107
Mérou taches jaunes . . . . . . . . . . . . . 2546 Mugil belanak . . . . . . . . . . . . . . . . . 2091
Mugil borbonicus . . . . . . . . . . . . . . . 2107 multiocellata, Pterygotrigla . . . . . . 2360, 2378

Mugil borneensis . . . . . . . . . . . . . . . 2088 multiradiatus, Myripristis . . . . . . . . . . . 2241
Mugil breviceps . . . . . . . . . . . . . . . . 2098 Munro’s hardyhead . . . . . . . . . . . . . . 2132
Mugil broussonneti . . . . . . . . . . . . . . 2095 munroi, Craterocephalus . . . . . . . . . . . 2132
Mugil caeruleomaculatus . . . . . . . . . . . 2107 murdjan, Myripristis . . . . . . . . . 2238, 2242
Mugil cephalotus . . . . . . . . . . . . . . . 2096 muroadsi, Decapterus . . . . . . . . . . . . . 2721
Mugil cephalus . . . . . . . . . . . . . 2095-2096 Murray’s armoured gurnard . . . . . . . . . . 2369
Mugil ceramensis . . . . . . . . . . . . . . . 2089 murrayi, Paraheminodus . . . . . . . . . . . 2369
Mugil ceylonensis . . . . . . . . . . . . . . . 2102 murrayi, Satyrichthys . . . . . . . . . . . . . 2369
Mugil chaptalii . . . . . . . . . . . . . . . . 2096 Myriodon . . . . . . . . . . . . . . . . . . . 2549
Mugil cirrhostomus . . . . . . . . . . . . . . 2083 MYRIPRISTINAE . . . . . . . . . . . . . . . 2225
Mugil cylindricus . . . . . . . . . . . . . . . 2107 Myripristis . . . . . . . . . 2225, 2227, 2239, 2591
Mugil diadema . . . . . . . . . . . . . . . . 2086 Myripristis (Holotrachys) guezei . . . . . . . 2248
Mugil dobula . . . . . . . . . . . . . . . . . 2096 Myripristis adusta . . . . . . . . . . . . . . . 2236
Mugil dussumieri . . . . . . . . . . . . . . . 2093 Myripristis amaena . . . . . . . . . . . . . . 2237
Mugil ferrandi . . . . . .. . . . . . . . . . . 2087 Myripristis argyromus . . . . . . . . . . . . . 2237
Mugil japonicus . . . . . . . . . . . . . . . . 2096 Myripristis axillaris . . . . . . . . . . . . . . 2242
Mugil javanicus . . . . . . . . . . . . . . . . 2093 Myripristis berndti . . . . . . . . . . . . . . 2238
Mugil jerdoni . . . . . . . . . . . . . . . . . 2093 Myripristis borbonicus . . . . . . . . . . . . 2241
Mugil kandavensis . . . . . . . . . . . . . . 2104 Myripristis botche . . . . . . . . . . . . . . . 2239
Mugil kelaartii . . . . . . . . . . . . . . . . 2106 Myripristis chryseres . . . . . . . . . . . . . 2240
Mugil longimanus . . . . . . . . . . . . . . . 2106 Myripristis hexagona . . . . . . . . . 2239, 2254
Mugil macrocheilos . . . . . . . . . . . . . . 2083 Myripristis kuntee . . . . . . . . . . . . . . . 2241
Mugil macrolepidotus . . . . . . . . . . . . . 2095 Myripristis melanostictus . . . . . . . . . . . 2239
Mugil nortoni . . . . . . . . . . . . . . . . . 2105 Myripristis microphthalmus . . . . . . . . . . 2243
Mugil oeur . . . . . . . . . . . . . . . . . . 2096 Myripristis multiradiatus . . . . . . . . . . . 2241
Mugil oligolepis . . . . .. . . . . . . . . . . 2089 Myripristis murdjan . . . . . . . . . . 2238, 2242
Mugil ophuysenii . . . . . . . . . . . . . . . 2103 Myripristis parvidens . . . . . . . . . . . . . 2242
Mugil öur . . . . . . . . . . . . . . . . . . . 2096 Myripristis pralinia . . . . . . . . . . 2241, 2254
Mugil peronii . . . . . . . . . . . . . . . . . 2087 Myripristis randalli . . . . . . . . . . . . . . 2254
Mugil strongylocephalus . . . . . . . . . . . 2106 Myripristis tiki . . . . . . . . . . . . . . . . . 2255
Mugil sudanensis . . . . . . . . . . . . . . . 2093 Myripristis trachyacron . . . . . . . . . . . . 2255
Mugil suppositus . . . . . . . . . . . . . . . 2108 Myripristis violacea . . . . . . . . . . . . . . 2243
Mugil tade . . . . . . . .. . . . . . . . . . . 2091 Myripristis vittata . . . . . . . . . . . . . . . 2255
Mugil troscheli . . . . . . . . . . . . . . . . 2088 Myripristis woodsi . . . . . . . . . . . . . . 2256
MUGILIDAE . . . . . . . . . . . 2069, 2109, 2113 MYSIDACEA . . . . . . . . . . . . . . . . . . 2276
MUGILIFORMES . . . . . . . . . . . . 2069-2108 Myxus . . . . . . . . . . . . . . . . . . . . . 2069
mugiloides, Allanetta . . . . . . . . . . . . . 2131 Myxus elongatus . . . . . . . . . . . . . . . 2097
mugiloides, Craterocephalus . . 2123, 2129, 2131 Myxus malayanus . . . . . . . . . . . . . . . 2095
MUGILOIDIDAE . . . . . . . . . . . . . . . . 2614 Myxus petardi . . . . . . . . . . . . . . . . . 2098
Mulet à grandes écailles . . . . . . . . . . . 2088
Mulet à grosse tête . . . . . . . . . . . . . . 2096 N
Mulet à queue bleue . . . . . . . . . . . . . . 2102 Naga halfbeak . . . . . . . . . . . . . . . . . 2196
Mulet à tache bleue . . . . . . . . . . . . . . 2107 naga, Rhynchorhamphus . . . . . . . . . . . 2196
Mulet boxeur . . . . . . . . . . . . . . . . . 2083 Nannygai . . . . . . . . . . . . . . . . . . . . 2223
Mulet chiraya . . . . . . . . . . . . . . . . . 2108 nanum, Plectrogenium . . . . . . . . . . . . 2328
Mulet diamant . . . . . . . . . . . . . . . . . 2086 naresii, Cypselurus . . . . . . . . . . . . . . 2174
Mulet dos vert . . . . . . . . . . . . . . . . . 2093 Narrowhead flyingfish . . . . . . . . . . . . . 2173
Mulet labéon . . . . . . . . . . . . . . . . . . 2100 nasutus, Rhinomugil . . . . . . . . . . . . . 2101
Mulet longue aile . . . . . . . . . . . . . . . 2103 nasutus, Squalomugil . . . . . . . . . . . . . 2101
Mulet mopiro . . . . . . . . . . . . . . . . . 2094
Mulet oeil de perdrix . . . . . . . . . . . . . . 2104
natalensis, Caranx . . . . . . . . . . . . . . . 2730
Mulet otomebora . . . . . . . . . . . . . . . 2089 Naucrates . . . . . . . . . . . . . . . . . . . 2660
Mulet tade . . . . . . . . . . . . . . . . . . . 2091 Naucrates ductor . . . . . . . . . . . . . . . 2727
Mullets . . . . . . . . . . . . . . . . . 2069, 2109 naucrates, Echeneis . . . . . . . . . 2652, 2654
multiannulatus, Doryrhamphus (Doryrhamphus) nauticus, Chaenomugil . . . . . . . . . . . . 2099
. . . . . . . . . . . . . . . . . . . . . . . 2275 Needlefishes . . . . . . . . . . . . . . 2151, 2180
multibarbus, Choridactylus . . . . . . . . . 2311 Needlescaled queenfish . . . . . . . . . . . . 2734
multifasciata, Pristigenys . . . . . . . . . . . 2601 neglecta, Scorpaenopsis . . . . . . . . . . . 2343
multinotatus, Epinephelus . . . . . . . . . . 2545 Neglected halfbeak . . . . . . . . . . . . . . 2195
multiocellata, Parapterygotyrigla . . . . . . 2378 neglectissimus , Hyporhamphus (Hyporhamphus)
. . . . . . . . . . . . . . . . . . . . . . . 2195
neglectissimus, Hyporhamphus . . . . . . . 2191 Oceanic flying halfbeak . . . . . . . . . . . . 2196

neglectus, Hyporhamphus . . . . . . . . . . 2191 Oceanic two-wing flyingfish . . . . . . . . . . 2176
neglectus, Hyporhamphus (Hyporhamphus) . 2195 Ocellated scorpionfish . . . . . . . . . . . . . 2326
nematophthalamus, Cymbacephalus . . . . 2396 Ocellated waspfish . . . . . . . . . . . . . . . 2310
nematoptera, Sphaeramia . . . . . . . . . . 2610 ocellifera, Dampieria . . . . . . . . . . . . . 2571
NEMIPTERIDAE . . . . . . . . . . . . . . . 2632 Ocosia apia . . . . . . . . . . . . . . . . . . 2324
See also Vol. 5 Ocosia zaspilota . . . . . . . . . . . . . . . . 2325
Neocentropogon trimaculatus . . . . . . . . 2321 octofasciatus, Epinephelus . . . . . . 2497, 2545
Neomerinthe amplisquamiceps . . . . 2322-2323 Oedalechilus labiosus . . . . . . . . . . . . . 2100
Neomerinthe megalepis . . . . . . . . . . . 2322 oeur, Mugil . . . . . . . . . . . . . . . . . . 2096
Neomerinthe procurva . . . . . . . . . 2322-2323 Ogilby’s silverside . . . . . . . . . . . . . . . 2127
Neomerinthe rotunda . . . . . . . . . . . . . 2323 ogilbyi, Atherinomorus . . . . 2120, 2123, 2125,
Neomyxus chaptalii . . . . . . . . . . . . . 2099 2127, 2131
Neomyxus leuciscus . . . . . . . . . . . . . 2099 ogilbyi, Pranesus . . . . . . . . . . . 2120, 2127
Neomyxus sclateri . . . . . . . . . . . . . . . 2099 Ogilbyina queenslandiae . . . . . . . . . . . 2572
Neoniphon . . . . . . . . . . . 2229, 2244, 2551 oligacanthus, Plectropomus . . . . . . . . . 2535
Neoniphon opercularis . . . . . . . . . . . . 2244 oligolepis, Cypselurus . . . . . . . . . . . . . 2174
Neoniphon sammara . . . . . . . . . . . . . 2245 oligolepis, Liza . . . . . . . . . . . . . . . . 2090
nesiotes, Iso . . . . . . . . . . . . . . . . . . 2143 oligolepis, Mugil . . . . . . . . . . . . . . . 2089
Netfin grouper . . . . . . . . . . . . . . . . . 2544 Oneblotch grouper . . . . . . . . . . . . . . . 2544
Network pipefish . . . . . . . . . . . . . . . . 2273 ongus, Epinephelus . . . . . . . 2491, 2514, 2523
nierstraszi, Sillago . . . . . . . . . . . . . . 2618 Onigocia . . . . . . . . . . . . . . . . . . . . 2385
niger, Apolectus . . . . . . . . . . . . . . . . 2729 Onigocia macrolepis . . . . . . . . . . . . . 2404
niger, Formio . . . . . . . . . . . . . . . . . 2729 Onigocia pedimacula . . . . . . . . . . . . . 2405
niger, Parastromateus . . . . . . . . . . . . 2729 Onigocia spinosa . . . . . . . . . . . . . . . 2406
niger, Tetraroge . . . . . . . . . . . . . . . 2350 opercularis, Flammeo . . . . . . . . . . . . . 2244
nigripectus, Corythoichthys . . . . . . . . . 2274 opercularis, Holocentrus . . . . . . . . . . . 2244
nigripinnis, Caranx . . . . . . . . . . . . . . 2689 opercularis, Kutaflammeo . . . . . . . . . . . 2244
nigripinnis, Cephalopholis . . . . . . . . . . 2483 opercularis, Neoniphon . . . . . . . . . . . . 2244
nigrofasciata, Seriolina . . . . . . . . . . . 2742 OPHIDIIDAE . . . . . . . . . . . . . . 2427, 2587
nigrofasciata, Zonichthys . . . . . . . . . . . 2742 See also Vol. 3
Niphon . . . . . . . . . . . . . . . . . . . . 2550 ophuysenii, Mugil . . . . . . . . . . . . . . . 2103
Niphon spinosus . . . . . . . . . . . 2429, 2548 opisthopterus, Scopeloberyx . . . . . . . . . 2201
niphonia, Pristigenys . . . . . . . . . . . . . 2601 opisthopus, Cypselurus . . . . . . . . . . . . 2175
NIPHONINI . . . . . . . . . . . . . . . . . . 2548 OPISTOGNATHIDAE . . . . . . 2558, 2579, 2588
nitidus, Carangoides . . . . . . . . . . . . . 2706 Opistognathus . . . . . . . . . . . . . . . . . 2588
nobilis, Caranx . . . . . . . . . . . . . . . . 2730 Orange roughy . . . . . . . . . . . . . . . . . 2215
Noline scorpionfish . . . . . . . . . . . . . . 2328 Orange scorpionfish . . . . . . . . . . . . . . 2322
Nomorhamphus . . . . . . . . . . . . . . . 2181 Orangespotted grouper . . . . . . . . . . . . 2493
normani, Decapterus . . . . . . . . . . . . . 2692 Orangespotted trevally . . . . . . . . . . . . 2694
Northern scorpionfish . . . . . . . . . . . . . 2334 Orbicular velvetfishes .. . . . . . . . . . . . 2353
Northern snubnose garfish . . . . . . . . . . 2193 orbicularis, Sphaeramia . . . . . . . . . . . 2610
nortoni, Mugil . . . . . . . . . . . . . . . . . 2105 ordensis, Moolgarda (Planiliza) . . . . . . . 2086
Notched flathead . . . . . . . . . . . . . . . 2404 Oriental helmet gurnard . . . . . . . . . . . . 2287
Notesthes robusta . . . . . . . . . . . . . . . 2324 Oriental sillago . . . . . . . . . . . . . . . . . 2617
NOTOGRAPTIDAE . . . . . . . . . . 2557, 2586 orientalis, Chorinemus . . . . . . . . . . . . 2732
Notograptus . . . . . . . . . . . . . . . . . . 2586 orientalis, Dactyloptena . . . . . . . 2287, 2289
Notograptus gregoryi . . . . . . . . . . . . . 2586 Orphie carène . . . . . . . . . . . . . . . . . 2155
Orphie plate . . . . . . . . . . . . . . . . . . 2154
Notograptus guttatus . . . . . . . . . . . . . 2586
Notograptus kauffmani . . . . . . . . . . . 2586 orthogrammus, Carangoides . . . . . . . . . 2706
Notograptus livingstonei . . . . . . . . . . . 2586 Oryzias spp. . . . . . . . . . . . . . . . . . . 2150
novaehollandiae, Pseudochromis . . . . . . 2572 osteochir, Remora . . . . . . . . . . . . . . . 2652
novaezelandiae, Trachurus . . . . . . 2750-2751 Ostichthys . . . . . . . . . . . . . . . 2225, 2230
Ostichthys archiepiscopus . . . . . . . . . . 2246
O Ostichthys japonicus . . . . . . . . . . . . . 2247
Ostichthys kaianus . . . . . . . . . . . . . . 2248
Obliquebanded grouper . . . . . . . . . . . . 2519 Ostichthys pillwaxii . . . . . . . . . . . . . . 2246
Obliquelined dottyback . . . . . . . . . . . . 2570 OSTRACOBERYCIDAE . . . . . . . . . . . 2551
oblongus, Carangichthys . . . . . . . . . . . 2705 Ostracoberycids . . . . . . . . . . . . . . . . 2551
oblongus, Carangoides . . . . . . . . . . . . 2705 otaitensis, Thysanophrys . . . . . . . . . . . 2421
oblongus, Caranx . . . . . . . . . . . . . . . 2705 Otohime . . . . . . . . . . . . . . . . . . . . 2360
obtusirostris, Exocoetus . . . . . . . . . . . 2176
Otohime hemisticta . . . . . . . . . . . . . . 2373 Parapterygotyrigla multiocellata . . . . . . . 2378

Otomebora mullet . . . .. .. . .
. . . 2089. . . Parascorpaena . . . . . . . . . . . . . . . . 2326
öur, Mugil . . . . . . . .. .. . .
. . . 2096. . . Parascorpaena mcadamsi . . . . . . . . . . 2326
Oval grouper . . . . . .. .. . .
. . . 2547. . . Parascorpaena mossambica . . . . . . . . . 2327
ovalaua, Allanetta . . . .. .. . .
. . . 2135. . . Parascorpaena picta . . . . . . . . . 2327, 2346
ovalaua, Hypoatherina .. .. . .
. . . 2135. . . Parastromateus . . . . . . . . . . . . . 2659-2661
ovalauus, Pranesus . . .. .. . .
. . . 2135. . . Parastromateus niger . . . . . . . . . . . . . 2729
OWSTONIIDAE . . . . .. .. . .
. . . 2589. . . Parazen . . . . . . . . . . . . . . . . . . . . 2257
Oxeye scad . . . . . . .. .. . .
. . . 2735. . . PARAZENIDAE . . . . . . . . . . . . . 2257-2258
oxycephala, Scorpaenopsis .. . .
2341, 2343 . . Parexocoetus brachypterus . . . . . . . . . . 2178
oxycephalus, Hirundichthys . . .
. . . . 2177. . Parexocoetus mento . . . . . . . . . . . . . 2178
Oxyporhamphus . . . . . . . . .
2162, 2180 . . parilis, Cymbacephalus . . . . . . . . . . . . 2397
Oxyporhamphus convexus bruuni . . . . . 2192 . parini, Epigonus . . . . . . . . . . . . . . . 2611
Oxyporhamphus convexus convexus . . . . 2192 parmata, Liza . . . . . . . 2089-2090, 2093-2094
Oxyporhamphus meristocystis . . . . . . . . 2192 Parmops coruscans . . . . . . . . . . . . . . 2212
Oxyporhamphus micropterus . . . . . . . . 2192 parsia, Liza . . . . . . . . . . . 2088, 2091, 2093
Oxyporhamphus micropterus micropterus . 2196 parvidens, Myripristis . . . . . . . . . . . . . 2242
Oxyporhamphus micropterus similis . . . . 2196 parvipinnis, Malacanthus . . . . . . . . . . . 2647
oxyrhynchus, Cestraeus . . . . . . . . . . . 2081 parvipinnis, Scorpaenodes . . . . . . . . . . 2339
Oyster pompano . . . . . . . . . . . . . . . . 2744 patriciae, Rogadius . . . . . . . . . . . . . . 2411
pauciradiatus, Craterocephalus . . . 2129, 2131
P pauli, Pterygotrigla . . . . . . . . . . . . . . 2360
paccagnellae, Pseudochromis . . . . 2557, 2576 Peacock hind . . . . . . . . . . . . . . . . . 2476
pacificus, Hyporhamphus acutus . . . . . . 2194 Pêche-madam asiate . . . . . . . . . . . . . 2620
Pagapa . . . . . . . . .. . . . . . . . . . . 2649 Pêche-madam clairon . . . . . . . . . . . . . 2621
Painted scorpionfish . . .. . . . . . . . . . . 2327 Pêche-madame argenté . . . . . . . . . . . . 2629
Pajama cardinalfish . . .. . . . . . . . . . . 2610 Pêche-madame ceinture d’or . . . . . . . . . 2619
Palemargin grouper . . .. . . . . . . . . . . 2542 Pêche-madame de vase . . . . . . . . . . . . 2626
Palometa negra . . . . .. . . . . . . . . . . 2729 Pêche-madame diablotin . . . . . . . . . . . 2622
Pámpano abotonado . .. . . . . . . . . . . 2745 Pêche-madame doré . . . . . . . . . . . . . 2618
Pámpano africano . . . .. . . . . . . . . . . 2743 Pêche-madame grandes ecailles . . . . . . . 2627
Pámpano de hebra . . .. . . . . . . . . . . 2684 Pêche-madame halanda . . . . . . . . . . . 2624
Pámpano de quilla . . .. . . . . . . . . . . 2691 Pêche-madame murda . . . . . . . . . . . . 2625
Pámpano indico . . . . .. . . . . . . 2685, 2749 Pêche-madame oriental . . . . . . . . . . . . 2617
Pámpano lunero . . . . .. . . . . . . . . . . 2746 Pêche-madame sable . . . . . . . . . . . . . 2623
Pámpano pastilla . . . .. . . . . . . . . . . 2747 Pêche-madame trompette . . . . . . . . . . . 2628
Pampanos . . . . . . . .. . . . . . . . . . . 2659 pectoralis, Caranx . . . . . . . . . . . . . . . 2689
Pampus . . . . . . . . .. . . . . . . . . . . 2660 pectoralis, Uraspis . . . . . . . . . . . . . . 2756
Panatela silverside . . .. . . . . . . . . . . 2139 pedimacula, Onigocia . . . . . . . . . . . . 2405
panatela, Atherina . . .. . . . . . . . . . . 2139 PEGASIDAE . . . . . . . . . . . . . . . . . . 2262
panatela, Hypoatherina .. . . . . . . . . . . 2139 Pegatimón . . . . . . . . . . . . . . . . . . . 2654
panatela, Stenatherina .. . . . . . . . . . . 2139 Pejerrey cabezón . . . . . . . . . . . . . . . 2125
panchax, Aplocheilus .. . . . . . . . . . . 2197 Pejerrey samoano . . . . . . . . . . . . . . . 2136
Pantolabus radiatus . .. . . . . . . . . . . 2728 Pejerrey tropical . . . . . . . . . . . . . . . . 2122
papilio, Dactyloptena . .. . . . . . . . . . . 2288 Pejerry de Barnes . . . . . . . . . . . . . . . 2133
Papillose flathead . . . .. . . . . . . . . . . 2418 Péliau chanos . . . . . . . . . . . . . . . . . 2649
pappei, Seriola . . . . .. . . . . . . . . . . 2740 PEMPHERIDAE . . . . . . . . . . . . 2219, 2591
papuensis, Caranx . . .. . . . . . . . . . . 2715 See also Vol. 5
para, Caranx . . . . . .. . . . . . . . . . . 2688 Pempheris sp. . . . . . . . . . . . . . . . . . 2474
Perca de mar . . . . . . . . . . . . . . . . . 2432
Parabembras curtus . .. . . . . . . . . . . 2383
PARABEMBRIDAE . . .. . . . . . . . . . . 2383 Perca miniata var. C . . . . . . . . . . . . . 2536
Paracentropogon longispinis . . . . . . . . 2325 Perchlets . . . . . . . . . . . . . . . . . . . . 2433
PERCICHTHYIDAE . . . . . . . . . . . . . . 2436
Paracentropogon vespa . . . . . . . . . . . 2325
PERCIFORMES . . . . . . . . . . . . 2261, 2429
Paraheminodus murrayi . . . . . . . . . . . 2369 See also Vols 5 and 6
PARAPERCIDAE . . . . . . . . . . . . . . . 2614 PERCOIDEI . . . . . . . . . . . . . . . . . . 2429
See also Vol. 6 PERCOPHIDAE . . . . . . . . . . . . . . . . 2386
Paraplesiops . . . . . . . . . .. . . . . . . 2578 See also Vol. 6
Parapterois heterurus . . . . . . . . . . . . 2326 PERISTEDIIDAE . . . . . . . . . . . . . . . 2359
Parapterygotrigla . . . . . . . .
. . . . . . . 2360 PERISTEDIINAE . 2359, 2367, 2370, 2381, 2551
Parapterygotyrigla macrorhynchus . . . . . 2376 Peristedion liorhynchus . . . . . . . . . . . 2370
Parapterygotyrigla megalops . . . . . . . . . 2377 Peristedion undulatus . . . . . . . . . . . . . 2366
peronii, Mugil . . . . . . . . . . . . . . . . . 2087 Plectropoma maculatum var. D . . . . . . . . 2536

perplexus, Epinephelus . . . . . . . . . . . . 2545 Plectropoma maculatus . . . . . . . . 2533, 2536
perpulcher, Pseudochromis mccullochi . . . 2570 Plectropoma variegatum . . . . . . . . . . . 2535
perusii, Valamugil . . . . . . . . . . . 2104-2106 Plectropomus . . . . . . . . . . . . . . . . . 2464
pessuliferus, Doryrhamphus (Doryrhamphus) 2275 Plectropomus areolatus . . . . . . . . . . . . 2531
pessuliferus, Plectropomus . . . . . . . . . . 2536 Plectropomus laevis . . . . . . . . . . . . . . 2532
petardi, Myxus . . . . . . . . . . . . . . . . 2098 Plectropomus leopardus . . . . . . . . 2532-2533
petardi, Trachystoma . . . . . . . . . . . . . 2098 Plectropomus maculatus . . . . . . . 2532, 2534
peterseni, Dactyloptena . . . . . . . . . . . . 2289 Plectropomus melanoleucus . . . . . . . . . 2532
peterseni, Daicocus . . . . . . . . . . . . . . 2289 Plectropomus oligacanthus . . . . . . . . . . 2535
petimba, Fistularia . . . . . . . . . . . 2278-2279 Plectropomus pessuliferus . . . . . . . . . . 2536
Pez piloto . . . . . . . . . . . . . . . . . . . 2727 Plectropomus truncatus . . . . . . . . . . . . 2531
PHALLOSTETHIDAE . . . . . . 2146-2147, 2150, PLESIOPIDAE . . . . . . . . . . . . . 2558, 2578
2198, 2200 plicatilis, Cestraeus . . . . . . . . . . . . . . 2082
Pharao flyingfish . . . . . . . . . . . . . . . . 2174 Plicomugil labiosus . . . . . . . . . . . . . . 2100
Phenacoscorpius megalops . . . . . . . . . 2328 plumbeus, Caranx . . . . . . . . . . . . . . . 2702
Philippine armoured gurnard . . . . . . . . . 2367 Plumbstriped stingfish . . . . . . . . . . . . . 2321
Philippine snubnose halfbeak . . . . . . . . . 2195 Plump grouper . . . . . . . . . . . . . . . . . 2546
philippinus, Heminodus . . . . . . . . . . . 2367 Pocillopora . . . . . . . . . . . . . . . . . . 2574
PHOLIDICHTHYIDAE . . . . . . . . 2558, 2587 Poecilia sphenops . . . . . . . . . . . . . . . 2199
See also Vol. 6 POECILIIDAE . . . . . . . 2147, 2150, 2198-2199
picta, Parascorpaena . . . . . . . . . 2327, 2346 Poeciliids . . . . . . . . . . . . . . . . . . . . 2199
picta, Pterygotrigla . . . . . . . . . . . 2372-2373 POECILIINAE . . . . . . . . . . . . . . . . . 2200
pillwaxii, Ostichthys . . . . . . . . . . . . . 2246 Poecillopora . . . . . . . . . . . . . . . . . . 2353
Pilotfish . . . . . . . . . . . . . . . . . . . . 2727 poecilonotus, Epinephelus . . . . . . 2506, 2510,
Pilotfishes . . . . . . . . . . . . . . . . . . . 2659 2513, 2515, 2519
Pineapple fishes . . . . . . . . . . . . . . . . 2214 poecilopterus, Cypselurus . . . . . . . . . . 2175
Pinecone soldierfish . . . . . . . . . . . . . . 2242 Poisson arme . . . . . . . . . . . . . . . . . 2341
PINGUIPEDIDAE . . . . . . . . . . . . . . . 2632 Poisson balance . . . . . . . . . . . . . . . . 2349
See also Vol. 6 Poisson pilote . . . . . . . . . . . . . . . . . 2727
pinguis, Pranesus . . . . . 2120, 2124-2125, 2127 Poisson scorpion feuillu . . . . . . . . . . . . 2333
pinnulatus, Decapterus . . . . . . . . . . . . 2719 Poisson volant . . . . . . . . . . . . . . . . . 2330
Pipefishes . . . . . . . . . . . . . . . . . . . 2264 Poisson-pierre . . . . . . . . . . . . . . . . . 2349
Pitcairn flyingfish . . . . . . . . . . . . . . . 2171 politus, Decapterus . . . . . . . . . . . . . . 2692
pitcairnensis, Cheilopogon . . . . . . . . . . 2171 Polka dot waspfish . . . . . . . . . . . . . . . 2325
plagiotaenia, Carangoides . . . . . . . . . . 2707 polleni, Cephalopholis . . . . . . . . . . . . 2541
plagiotaenia, Caranx . . . . . . . . . . . . . 2707 polylepis, Epinephelus . . . . . . . . . . . . 2492
Plain helmet gurnard . . . . . . . . . . . . . 2290 polynemus, Pseudochromis . . . . . . . . . . 2575
Plaintail firefish . . . . . . . . . . . . . . . . 2331 polyphekadion, Epinephelus . . . . . 2501, 2516
planiceps, Liza . . . . . . . . . . . . 2091, 2093 Polyprion . . . . . . . . . . . . . . . . . . . 2436
platei australis, Callanthias . . . . . . . . . 2555 POLYPRIONIDAE . . . . . . . . . . . . . . . 2436
platei, Callanthias . . . . . . . . . . . . . . 2555 polystigma, Epinephelus . . . . . . . . . . . 2517
Platybelone argalus platyura . . . . . . . . . 2155 POMACANTHIDAE . . . . . . . . . . . . . . 2551
Platycéphale clin d’oeil . . . . . . . . . . . . 2420 See also Vol. 5
Platycéphale cocarde . . . . . . . . . . . . . 2403 Pomacentrus littoralis . . . . . . . . . . . . 2349
Platycéphale indian . . . . . . . . . . . . . . 2409 POMATOMIDAE . . . . . . . . 2650, 2655, 2661
Platycéphale japonais . . . . . . . . . . . . . 2402 Pomatomus saltator . . . . . . . . . . . . . . 2650
Platycéphale poignard . . . . . . . . . . . . 2415 Pomatomus saltatrix . . . . . . . . . . . . . 2650
PLATYCEPHALIDAE . . . 2292-2293, 2360, 2383, Pompaneau africain . . . . . . . . . . . . . . 2743
2385, 2422 Pompaneau indien . . . . . . . . . . . . . . . 2749
Platycephalids . . . . . . . . . . . . . . . . . 2386 Pompaneau lune . . . . . . . . . . . . . . . . 2746
Platycephalus arenarius . . . . . . . . . . . 2407 Pompaneau muscadin . . . . . . . . . . . . . 2745
Platycephalus endrachtensis . . . . . . . . . 2408 Pompaneau pierrot . . . . . . . . . . . . . . 2747
Platycephalus indicus . . . . . . . . . 2408-2409 Pompano à quille noire . . . . . . . . . . . . 2691
platyura, Belone . . . . . . . . . . . . . . . . 2155 Pompano dolphinfish . . . . . . . . . . . . . 2657
platyura, Platybelone argalus . . . . . . . . 2155 Popeyed scorpionfish . . . . . . . . . . . . . 2333
Plectranthias . . . . . . . . . . . . . 2442, 2461 Poromitra . . . . . . . . . . . . . . . . . . . 2201
PLECTROGENIIDAE . . . . . . . . . . . . . 2383 porphyreus, Pseudochromis . . . . . 2557, 2576
Plectrogenium nanum . . . . . . . . . . . . 2328 Potato grouper . . . . . . . . . . . . . . . . . 2527
Plectropoma leopardinus . . . . . . . . . . . 2533 powelli, Saloptia . . . . . . . . . . . . . . . 2537
Plectropoma maculatum . . . . . . . . . . . 2531 praeopercularis, Epinephelus . . . . . . . . . 2496
Plectropoma maculatum var. A . . . . . . . . 2536
praepositum, Liocranium . . . . . . . . . . 2318 Pseudochromis porphyreus . . . . . . 2557, 2576

praeustus, Carangoides . . . . . . . . . . . 2708 Pseudochromis punctatus . . . . . . . . . . . 2568
praeustus, Caranx . . . . . . . . . . . . . . . 2708 Pseudochromis tapeinosoma . . . . . . . . . 2573
pralinia, Myripristis . . . . . . . . . 2241, 2254 Pseudochromis veliferus . . . . . . . . . . . 2572
Pranesus capricornensis . . . . . . . . . . . 2120 Pseudochromis xanthochir . . . . . . . . . . 2574
Pranesus duodecimalis . . . . . . . . . . . . 2122 Pseudogramma . . . . . . . . . . . . 2468, 2579
Pranesus endrachtensis . . . . . . . 2123, 2127 Pseudogrammini . . . . . . . . . . . . . . . 2442
Pranesus maculatus . . . . . . . . . . . . . . 2125 pseudomorrhua, Epinephelus . . . . . . . . . 2510
Pranesus ogilbyi . . . . . . . . . . . 2120, 2127 Pseudomugil cyanodorsalis . . . . . . 2110-2111
Pranesus ovalauus . . . . . . . . . . . . . . 2135 Pseudomugil helodes . . . . . . . . . . . . . 2144
Pranesus pinguis . . . . . 2120, 2124-2125, 2127 Pseudomugil inconspicuus . . . . . . . 2110-2111
Pranesus vaigiensis . . . . . . . . . . . . . . 2125 Pseudomugil majusculus . . . . . . . 2112, 2144
Pranesus valenciennei . . . . . . . . . . . . 2138 Pseudomugil signifer . . . . . . . . . . . . . 2112
praslin, Sargocentron . . . . . . . . . 2250-2251 PSEUDOMUGILIDAE . . . . . . . . . 2109, 2144
Prettyfins . . . . . . . . . . . . . . . . . . . 2578 Pseudomugilids . . . . . . . . . . . . . . . . 2109
PRIACANTHIDAE . . . . . . . . . . . . . . 2590 Pseudoplesiops . . . . . . . . . . . . . . . . 2557
Priacanthus alalaua . . . . . . . . . . . . . 2599 Pseudoplesiops sargenti . . . . . . . . . . . . 2577
Priacanthus blochii . . . . . . . . . . . . . . 2599 Pseudoplesiops typus . . . . . . . . . . . . . 2577
Priacanthus cruentatus . . . . . . . . . . . . 2595 Pseudosquilla sp. . . . . . . . . . . . . . . . 2474
Priacanthus fitchi . . . . . . . . . . . . . . 2600 psilogaster, Acanthoplesiops . . . . . . . . . 2578
Priacanthus hamrur . . . . . . . . . 2596, 2599 PSYCHROLUTIDAE . . . . . . . . . 2425, 2427
Priacanthus macracanthus . . . . . 2591, 2597 Pterapogon kauderni . . . . . . . . . . . . . 2609
Priacanthus prolixus . . . . . . . . . . . . . 2596 Pterapogon mirifica . . . . . . . . . . . . . . 2609
Priacanthus sagittarius . . . . . . . . . 2599-2600 Pterois . . . . . . . . . . . . . . . . . . . . . 2331
Priacanthus spp. . . . . . . . . . . . . . . . 2590 Pterois antennata . . . . . . . . . . . . . . . 2329
Priacanthus tayenus . . . . . . . . . . . . . 2598 Pterois lunulata . . . . . . . . . . . . 2329, 2331
Priacanthus zaiserae . . . . . . . . . . . . . 2600 Pterois miles . . . . . . . . . . . . . . 2330, 2332
Priapium fishes . . . . . . . . . . . . . . . . 2146 Pterois mombasae . . . . . . . . . . . . . . . 2330
prionocephalus, Gargariscus . . . . . . . . 2366 Pterois radiata . . . . . . . . . . . . . . . . . 2331
Prionotus alepis . . . . . . . . . . . . . . . . 2373 Pterois russellii . . . . . . . . . . . . 2329, 2331
Pristigenys meyeri . . . . . . . . . . . . . . 2601 Pterois volitans . . . . . . . . . . . . . 2330-2332
Pristigenys multifasciata . . . . . . . . . . . 2601 Pteromugil diadema . . . . . . . . . . . . . . 2086
Pristigenys niphonia . . . . . . . . . . . . . 2601 Pterygotrigla . . . . . . . . . . . . . . . . . 2360
pristiger, Rogadius . . . . . . . . . . . . . . 2412 Pterygotrigla acanthomoplate . . . . 2360, 2371
Procetichthys . . . . . . . . . . . . . . . . . 2206 Pterygotrigla andertoni . . . . . . . . 2360, 2372
procurva, Neomerinthe . . . . . . . . . 2322-2323 Pterygotrigla arabica . . . . . . . . . . . . . 2373
Prognichthys brevipinnis . . . . . . . . . . . 2178 Pterygotrigla hemisticta . . . . 2360, 2373, 2380
Prognichthys sealei . . . . . . . . . . . . . . 2179 Pterygotrigla hoplites . . . . . . . . . 2360, 2374
prolixus, Priacanthus . . . . . . . . . . . . 2596 Pterygotrigla leptacanthus . . . . . . . . . . 2375
Promicrops lanceolatus . . . . . . . . . . . . 2505 Pterygotrigla macrolepidota . . . . . . . . . 2382
Protoblepharon rosenblatti . . . . . . . . . . 2212 Pterygotrigla macrorhynchus . . . . 2360, 2376
Psammoperca vaigiensis . . . . . . . . . . . 2432 Pterygotrigla megalops . . . . . . . . 2360, 2377
Pseudanthias . . . . . . . . . . . . . . . . . 2464 Pterygotrigla multiocellata . . . . . . 2360, 2378
Pseudanthias squamipinnis . . . . . . . . . 2480 Pterygotrigla pauli . . . . . . . . . . . . . . 2360
Pseudocaranx . . . . . . . . . . 2661, 2676, 2730 Pterygotrigla picta . . . . . . . . . . . 2372-2373
Pseudocaranx dentex . . . . . . . . . . . . . 2730 Pterygotrigla ryukyuensis . . . . . . 2360, 2379
Pseudocaranx wrighti . . . . . . . . . . . . 2730 Pterygotrigla spinosa . . . . . . . . . . . . . 2380
PSEUDOCHROMIDAE . . . . . 2557, 2579, 2586 Pterygotrigla tagala . . . . . . . . . . 2360, 2380
Pseudochromids . . . . . . . . . . . . . . . 2558 Pufferfish . . . . . . . . . . . . . . . . . . . . 2532
Pseudochromis . . . . . . . . . . . . . . . . 2557 punctata, Cociella . . . . . . . . . . . . . . . 2393
Pseudochromis aureus . . . . . . . . . . . . 2574 punctatus, Assiculus . . . . . . . . . . . . . 2568
Pseudochromis cyanotaenia . . . . . . . . . 2573 punctatus, Atherinichthys . . . . . . . . . . . 2131
Pseudochromis diadema . . . . . . . 2557, 2576 punctatus, Pseudochromis . . . . . . . . . . 2568
Pseudochromis fuscus . . . . . . . . 2571, 2574 punctulatus, Tylosurus . . . . . . . . . . . . 2161
Pseudochromis kikaii . . . . . . . . . . . . . 2573 Purplespot bigeye . . . . . . . . . . . . . . . 2598
Pseudochromis mccullochi . . . . . . . . . . 2570 purpurascens, Cypho . . . . . . . . . . . . . 2570
Pseudochromis mccullochi perpulcher . . . . 2570 purpurescens, Halichoeres . . . . . . . . . . 2475
Pseudochromis novaehollandiae . . . . . . 2572 purpurescens, Seriola . . . . . . . . . . . . . 2738
Pseudochromis paccagnellae . . . . 2557, 2576 purpureus, Cephalopholis . . . . . . . . . . . 2482
Pseudochromis polynemus . . . . . . . . . . 2575 pusillus, Minous . . . . . . . . . . . . . . . 2320
Puzzling grouper . . . . . . . . . . . . . . . 2545 Reef needlefish . . . . . . . . . . . . . . . . 2156

Pygmy scorpionfish . . . . . . . . . . . . . . 2339 Reef stonefish . . . . . . . . . . . . . . . . . 2349
Pyrosoma sp. . . . . . . . . . . . . . . . . . 2529 Reeftop pipefish . . . . . . . . . . . . . . . . 2273
regina, Allanetta . . . . . . . . . . . . . . . . 2128
Q regina, Atherina . . . . . . . . . . . . . . . . 2128
Quakerfish . . . . . . . . . . . .
. . . 2630, 2647 reginae, Atherinomorus . . . . . . . . . . . 2128
Queenfishes . . . . . . . . . . .
. . . . . . . 2659 regularis, Caranx . . . . . . . . . . . . . . . 2715
Queensland dottyback . . . . .
. . . . . . . 2572 Remora . . . . . . . . . . . . . . . . . . . . 2652
queenslandiae, Ogilbyina . . .
. . . . . . . 2572 Remora albescens . . . . . . . . . . . . . . . 2652
Quoy’s halfbeak . . . . . . . . .
. . . . . . . 2190 Remora australis . . . . . . . . . . . . . . . 2652
quoyanus, Epinephelus . . . . .
2500, 2503, 2508, Rémora commun . . . . . . . . . . . . . . . 2654
2512, 2518, 2524 Remora osteochir . . . . . . . . . . . . . . . 2652
quoyi, Hyporhamphus (Reporhamphus) . . 2190 Remoras . . . . . . . . . . . . . . . . . . . . 2652
quptae, Caranx . . . . . . . . . . . . . . . . 2756 Remorina . . . . . . . . . . . . . . . . . . . 2652
Rendahl’s hardyhead . . . . . . . . . . . . . 2129
R Reticulate grouper . . . . . . . . . . . . . . . 2546
RACHYCENTRIDAE . . . 2650, 2652, 2655, 2661 retouti, Epinephelus . . . . . . . . . 2499, 2520
Rachycentron canadum . . . . . . . 2652, 2655 reversa, Uraspis . . . . . . . . . . . . . . . . 2755
Radial firefish . . . . . . . . . . . . . . . . . 2331 Rhabdamia . . . . . . . . . . . . . . . . . . 2602
radiata, Pterois . . . . . . . . . . . . . . . . 2331 Rhabdosebastes . . . . . . . . . . . . . . . . 2549
radiatus, Absalom . . . . . . . . . . . . . . . 2728 Rhaphiobelone robusta . . . . . . . . . . . . 2156
radiatus, Caranx . . . . . . . . . . . . . . . 2728 Rhinomugil nasutus . . . . . . . . . . . . . 2101
radiatus, Epinephelus . . . . . 2513, 2515, 2519 Rhinopias aphanes . . . . . . . . . . . . . . 2332
radiatus, Pantolabus . . . . . . . . . . . . . 2728 Rhinopias frondosa . . . . . . . . . . . . . . 2333
Raggy scorpionfish . . . . . . . . . . . . . . 2344 rhomboides, Carangoides . . . . . . . . . . . 2704
rahanus, Epinephelus . . . . . . . . . . . . . 2517 rhothophilus, Iso . . . . . . . . . . . . . . . 2141
Rainbow runner . . . . . . . . . . . . . . . . 2724 rhothophilus, Tropidostethops . . . . . . . . 2141
Ramsay’s mullet . . . . . . . . . . . . . . . . 2092 rhothophilus, Tropidostethus . . . . . . . . . 2141
ramsayi, Gracimugil . . . . . . . . . . . . . 2092 rhyncholepis, Epinephelus . . . . . . . . . . 2521
ramsayi, Liza . . . . . . . . . . . . . 2087, 2092 Rhynchorhamphus . . . . . . . . . . . . . . 2196
ramsayi, Trachystoma . . . . . . . . . . . . 2092 Rhynchorhamphus georgii . . . . . . . . . . 2196
Randall’s soldierfish . . . . . . . . . . . . . . 2254 Rhynchorhamphus naga . . . . . . . . . . . 2196
randalli, Myripristis . . . . . . . . . . . . . 2254 Ricefishes . . . . . . . . . . . . . . . . . . . 2149
ransonnetii, Elates . . . . . . . . . . . . . . 2398 Richardsonichthys leucogaster . . . . . . . . 2333
Rapanoui flyingfish . . . . . . . . . . . . . . 2172 Ridgeheads . . . . . . . . . . . . . . . . . . 2201
rapanouiensis, Cheilopogon . . . . . . . . . 2172 riefflei, Satyrichthys . . . . . . . . . . . . . 2381
raphidoma, Tylosurus . . . . . . . . . . . . . 2160 Ringeyed dottyback . . . . . . . . . . . . . . 2577
Rascacio de Suez . . . . . . . . . . . . . . . 2345 rivoliana, Seriola . . . . . . . . . . . . . . . 2741
Rascacio ocelado . . . . . . . . . . . . . . . 2310 rivulatus, Epinephelus . . . . . . . . . . . . 2521
Rascacio serrano . . . . . . . . . . . . . . . 2347 robusta, Notesthes . . . . . . . . . . . . . . 2324
Rascasse de Suez . . . . . . . . . . . . . . 2345 robusta, Rhaphiobelone . . . . . . . . . . . . 2156
Rascasse ocellèe . . . . . . . . . . . . . . . 2310 robustus, Hemiramphus . . . . . . . . . . . 2193
Rascasse serran . . . . . . . . . . . . . . . 2347 Rock grouper . . . . . . . . . . . . . . . . . 2498
Ratabulus diversidans . . . . . . . . . . . . 2410 Rockfishes . . . . . . . . . . . . . . . . . . . 2291
Ratabulus megacephalus . . . . . . . . . . . 2410 rodericensis, Kumococius . . . . . . . . . . 2403
Razorfishes . . . . . . . . . . . . . . . . . . 2281 rogaa, Aethaloperca . . . . . . . . . . . . . 2474
Rearfin flyingfish . . . . . . . . . . . . . . . . 2175 rogaa, Cephalopholis . . . . . . . . . . . . . 2474
rectipinnis, Carangoides . . . . . . . . . . . 2704 Rogadius asper . . . . . . . . . . . . . . . . 2412
Red bigeye . . . . . . . . . . . . . . . . . . 2597 Rogadius patriciae . . . . . . . . . . . . . . 2411
Red cornetfish . . . . . . . . . . . . . . . . . 2279 Rogadius pristiger . . . . . . . . . . . . . . 2412
Red dottyback . . . . . . . . . . . . . . . . . 2571 Rogadius serratus . . . . . . . . . . . . . . . 2413
Red gurnard . . . . . . . . . . . . . . . . . . 2365 Rogadius welanderi . . . . . . . . . . . . . . 2414
Red tilefish . . . . . . . . . . . . . . . . . . . 2643 Rompeolas . . . . . . . . . . . . . . . . . . . 2141
Redcoat . . . . . . . . . . . . . . . . . . . . 2251
Rondeletia bicolor . . . . . . . . . . . . . . 2204
Redmouth grouper . . . . . . . . . . . . . . 2474 RONDELETIIDAE . . . . . . . . . . . . 2204-2206
Redmouth whalefishes . . . . . . . . . . . . 2204
Redsmooth scorpionfish . . . . . . . . . . . 2348
rosenblatti, Protoblepharon . . . . . . . . . 2212
Redspotted grouper . . . . . . . . . . . . . . 2485 Rosenblattia . . . . . . . . . . . . . . . . . . 2611
Redstripe bigeye . . . . . . . . . . . . . . . 2601 rotunda, Neomerinthe . . . . . . . . . . . . 2323
Redtail scad . . . . . . . . . . . . . . . . . . 2718 Rough flathead . . . . . . . . . . . . . . . . 2400
Redtipped grouper . . . . . . . . . . . . . . 2520 Rough grouper . . . . . . . . . . . . . . . . . 2548
Redvelvet whalefish . . . . . . . . . . . . . . 2205 Rough humpbacked scorpionfish . . . . . . . 2343
Roughear scad . . . .
. . . . . . . . . . . . 2723 Saumonée léopard . . . . . . . . . . . . . . 2533
Roughies . . . . . . .
. . . . . . . . . . . . 2215 Sauteur carsia . . . . . . . . . . . . . . . . . 2733
Roughskull soldierfish .
. . . . . . . . . . . . 2255 Sauteur leurre . . . . . . . . . . . . . . . . . 2734
Round scorpionfish . .
. . . . . . . . . . . . 2323 Sauteur sabre . . . . . . . . . . . . . . . . . 2732
Roundheads . . . . . .
. . . . . . . . . . . . 2578 Sauteur talang . . . . . . . . . . . . . . . . . 2731
Roving coralgrouper . .
. . . . . . . . . . . . 2536 sawakinensis, Branchiostegus . . . . . 2643-2644
ruber, Adioryx . . . . .
. . . . . . . . . . . . 2251 Sawcheek scorpionfish . . . . . . . . . . . . 2310
ruber, Holocentrus . .
. . . . . . . . . . . . 2251 scaber, Grammoplites . . . . . . . . . . 2399-2400
rubrum, Sargocentron . . . . . . . . . 2250-2251 scaber, Scorpaenodes . . . . . . . . . . . . . 2339
Runners . . . . . . . . . . . . . . . . . . . . 2659 Scads . . . . . . . . . . . . . . . . . . . . . 2659
russelli, Decapterus . . . . . . . 2720, 2722-2723 SCARIDAE . . . . . . . . . . . . . . . . . . 2632
russelli, Trachinotus . . . . . . . . . . 2747-2748 See also Vol. 6
russellii, Pterois . . . . . . . . . . . . 2329, 2331 Scarlet soldierfish . . . .
. . . . .
. . . . . 2254 .
Rusty hind . . . . . . . . . . . . . . . . . . . 2540 schlegeli, Caranx . . . .. . . . .
. . . . . 2693 .
Ruvettus . . . . . . . . . . . . . . . . . . . . 2660 Schultz’ pipefish . . . . .
. . . . .
. . . . . 2274 .
Rypticus . . . . . . . . . . . . . . . . . . . . 2558 schultzi, Corythoichthys . . . . .
. . . . . 2274 .
Ryukyu gurnard . . . . . . . . . . . . . . . . 2379 sclateri, Neomyxus . . . . . . . .
. . . . . 2099 .
ryukyuensis, Pterygotrigla . . . . . . 2360, 2379 sclerolepis krefftii, Arrhamphus . . . . . . 2193 .
sclerolepis sclerolepis, Arrhamphus . . . . . 2193
sclerolepis, Arrhamphus sclerolepis . . . . . 2193
S scolopax, Macrorhamphosus . . . . . . . . . 2280
Sabre squirrelfish . . . . . . . . . . . . . . . 2252 Scomber botla . . . . . . . . . . . . . . . . . 2747
Sacura . . . . . . . . . . . . . . . . . . . . . 2469 Scomberoides . . . . . . . 2660, 2677, 2731-2732
Saddletail grouper . . . . . . . . . . . . . . . 2495 Scomberoides commersonnianus . . . 2731-2732
sagittarius, Priacanthus . . . . . . . . 2599-2600 Scomberoides lysan . . . . . . . . . . . 2731-2732
Sailfin cardinalfish . . . . . . . . . . . . . . . 2609 Scomberoides tala . . . . . . . . . . . . . . . 2733
Sailfin flyingfish . . . . . . . . . . . . . . . . 2178 Scomberoides tol . . . . . . . . . . . . . . . 2734
Sailfin silversides . . . . . . . . . . . . . . . 2144 SCOMBRIDAE . . . . . . . . . . . . . . . . 2660
Sailor flyingfish . . . . . . . . . . . . . . . . 2179 See also Vol. 6
salonotus, Epinephelus . . . . . . . . . . . . 2524 scombrinus, Caranx . . . . . . . . . . . . . . 2721
Saloptia powelli . . . . . . . . . . . . . . . . 2537 SCOMBROPIDAE . . . . . . . . 2437, 2611, 2651
saltator, Pomatomus . . . . . . . . . . . . . 2650 Scombrops boops . . . . . . . . . . . . . . . 2651
saltator, Temnodon . . . . . . . . . . . . . . 2650 Scopeloberyx . . . . . . . . . . . . . . . . . 2201
saltatrix, Pomatomus . . . . . . . . . . . . . 2650 Scopeloberyx opisthopterus . . . . . . . . . 2201
Sammara squirrelfish . . . . . . . . . . . . . 2245 Scorpaena . . . . . . . . . . . . . . . . . . . 2293
sammara, Flammeo . . . . . . . . . . . . . . 2245 Scorpaena bynoensis . . . . . . . . . . . . . 2346
sammara, Holocentrus . . . . . . . . . . . . 2245 Scorpaena cardinalis . . . . . . . . . . . . . 2334
sammara, Kutaflammeo . . . . . . . . . . . . 2245 Scorpaena cookii . . . . . . . . . . . . . . . 2334
sammara, Neoniphon . . . . . . . . . . . . 2245 Scorpaena gibbifrons . . . . . . . . . . . . . 2335
Samoan silverside . . . . . . . . . . . . . . . 2136 Scorpaena hemilepidota . . . . . . . . . . . 2335
Samoan surf sardine . . . . . . . . . . . . . 2143 Scorpaena moultoni . . . . . . . . . . . . . . 2326
Samson fish . . . . . . . . . . . . . . . . . . 2739 SCORPAENIDAE . . . . . 2291, 2354, 2426, 2550
sanctipetri, Chorinemus . . . . . . . . . . . . 2732 Scorpaenids . . . . . . . . . . . . . . . 2292-2293
Sand flathead . . . . . . . . . . . . . . . . . 2407 SCORPAENIFORMES . . . . . . . . . 2283-2428
Sand mullet . . . . . . . . . . . . . . . . . . 2097 Scorpaenodes . . . . . . . . . . . . . . . . . 2339
Sand sillago . . . . . . . . . . . . . . . . . . 2623 Scorpaenodes albaiensis . . . . . . . . . . . 2336
Sand tilefishes . . . . . . . . . . . . . . . . . 2630 Scorpaenodes guamensis . . . . . . . 2336, 2339
Sandy-Bay cod . . . . . . . . . . . . . . . . 2334 Scorpaenodes hirsutus . . . . . . . . . . . . 2337
sansun, Caranx . . . . . . . . . . . . 2711, 2715 Scorpaenodes kelloggi . . . . . . . . . . . . 2337
Sardinella . . . . . . . . . . . . . . . . . . . 2125 Scorpaenodes littoralis . . . . . . . . . 2338-2339
Sargassum . . . . . . . . . . . 2265, 2432, 2656 Scorpaenodes minor . . . . . . . . . . . . . 2338
sargenti, Pseudoplesiops . . . . . . . . . . . 2577 Scorpaenodes parvipinnis . . . . . . . . . . 2339
Sargocentron . . . . . . . . . . 2225, 2231, 2551 Scorpaenodes scaber . . . . . . . . . . . . . 2339
Sargocentron caudimaculatum . . . . . . . 2249 Scorpaenodes varipinnis . . . . . . . . . . . 2340
Sargocentron praslin . . . . . . . . . . 2250-2251 scorpaenoides, Centropristes . . . . . . . . . 2549
Sargocentron rubrum . . . . . . . . . 2250-2251 Scorpaenoids . . . . . . . . . . . . . . . . . 2293
Sargocentron spiniferum . . . . . . . . . . . 2252 Scorpaenopsis . . . . . . . . . . . . . 2341, 2343
Sargocentron tiere . . . . . . . . . . . . . . 2253 Scorpaenopsis brevifrons . . . . . . . . . . . 2340
Satyrichthys murrayi . . . . . . . . . . . . . 2369 Scorpaenopsis cirrhosa . . . . . . . . 2341, 2343
Satyrichthys riefflei . . . . . . . . . . . . . . 2381 Scorpaenopsis cotticeps . . . . . . . . . . . . 2341
Satyrichthys spp. . . . . . . . . . . . . . . . 2369 Scorpaenopsis diabolus . . . . . . . . . 2342-2343
Scorpaenopsis fowleri . . . . . . . . .
. . . 2342 Seriola rivoliana . . . . . . . . . . . . . . . 2741
Scorpaenopsis gibbosa . . . . . . . . .
2342-2343 Seriola simplex . . . . . . . . . . . . . . . . 2738
Scorpaenopsis iop . . . . . . . . . . .
. . . 2341 Seriola songoro . . . . . . . . . . . . . . . . 2741
Scorpaenopsis macrochir . . . . . . .
. . . 2343 Seriola tapeinometapon . . . . . . . . . . . . 2738
Scorpaenopsis neglecta . . . . . . . . .
. . . 2343 Sériole amourez . . . . . . . . . . . . . . . . 2742
Scorpaenopsis oxycephala . . . . . .
2341, 2343 Sériole chicard . . . . . . . . . . . . . . . . . 2740
Scorpaenopsis venosa . . . . . . . .
. . . . 2344 Sériole couronee . . . . . . . . . . . . . . . . 2738
Scorpionfishes . . . . . . . . . . . . .
. 2291-2293 Sériole limon . . . . . . . . . . . . . . . . . . 2741
Sculpins . . . . . . . . . . . . . . . .
. . . . 2425 Seriolina . . . . . . . . . . . . . . . . . . . . 2659
scutatus, Centriscus . . . . . . . . .
. . . . 2282 Seriolina nigrofasciata . . . . . . . . . . . . 2742
Sea basses . . . . . . . . . . . . . .
. . . . 2442 SERRANIDAE . . . 2293, 2429, 2436, 2438-2439,
Sea perches . . . . . . . . . . . . . .
. . . . 2429 2442, 2550, 2553, 2558, 2579, 2632
Sea robins . . . . . . . . . . . . . . .
2292, 2359 Serranids . . . . . . . . . . . . 2438, 2442, 2558
Sea urchins . . . . . . . . . . . . . .
2282, 2609 SERRANINAE . . . . . . . . . . . . . . . . . 2442
Seadragons . . . . . . . . . . . . . .
. . . . 2262 Serranus amboinensis . . . . . . . . . . . . . 2529
Seahorses . . . . . . . . . . . . . . .
. . . . 2264 Serranus dispar var. A . . . . . . . . . . . . 2527
sealei, Prognichthys . . . . . . . . .
. . . . 2179 Serranus guttulatus . . . . . . . . . . . . . . 2528
Seamoths . . . . . . . . . . . . . . .
. . . . 2262 Serranus lineatus . . . . . . . . . . . . . . . 2529
Seaweeds . . . . . . . . . . . . . . .
. . . . 2265 Serranus micronotatus . . . . . . . . . . . . 2475
Sebastapistes ballieu . . . . . . . . .
. . . . 2345 Serranus moara . . . . . . . . . . . . . . . . 2490
Sebastapistes cyanostigma . . . . . .
. . . . 2344 Serranus urophthalmus . . . . . . . . . . . . 2475
Sebastapistes galactacma . . . . . .
. . . . 2345 Serrate razorfish . . . . . . . . . . . . . . . . 2282
Sebastapistes mauritiana . . . . . . .
. . . . 2345 Serrated flathead . . . . . . . . . . . . . . . 2413
Sebastapistes strongia . . . . . . . .
. . . . 2346 serratus, Branchiostegus . . . . 2630, 2645-2646
Sebastapistes tinkhami . . . . . . . .
. . . . 2346 serratus, Rogadius . . . . . . . . . . . . . . 2413
Sebastes meleagris . . . . . . . . . .
. . . . 2336 serrulatus, Brachypterois . . . . . . . . . . . 2310
Sebastes stoliczkae . . . . . . . . . .
. . . . 2549 Setarches . . . . . . . . . . . . . . . . . . . 2291
Sebastiscus marmoratus . . . . . . .
. . . . 2347 Setarches guentheri . . . . . . . . . . . . . . 2347
Sebastiscus tertius . . . . . . . . . .
. . . . 2347 Setarches longimanus . . . . . . . . . . . . 2348
secunda, Uraspis . . . . . . . . . . .
. 2754-2755 setosum, Diadema . . . . . . . . . . . . . . . 2282
seheli, Valamugil . . . . . . . . . . .
2102, 2107 Sevenbar grouper . . . . . . . . . . . . . . . 2497
Selar . . . . . . . . . . . . . . . . . .
. . . . 2678 sexfasciatus, Caranx . . . . . . . . . . 2716-2717
Sélar à bande dorée . . . . . . . . . .
. . . . 2737 sexfasciatus, Epinephelus . . . . . . 2498, 2522
Sélar aile noire . . . . . . . . . . . .
. . . . 2689 sexmaculata, Cephalopholis . . . . . . 2480-2481
Selar boops . . . . . . . . . . . . . .
. . . . 2735 Shadow trevally . . . . . . . . . . . . . . . . 2697
Sélar coulisou . . . . . . . . . . . . .
. . . . 2736 Shadowfin soldierfish . . . . . . . . . . . . . 2236
Selar crumenophthalmus . . . . . .
. 2735-2736 Shark mullet . . . . . . . . . . . . . . . . . . 2101
Sélar harengule . . . . . . . . . . . .
. . . . 2690 Sharksucker . . . . . . . . . . . . . . . . . . 2654
Sélar oeil de boeuf . . . . . . . . . .
. . . . 2735 Sharksuckers . . . . . . . . . . . . . . . . . 2652
Sélar queue jaune . . . . . . . . . . .
. . . . 2692 Sharpeared helmet gurnard . . . . . . . . . . 2288
Sélar subari . . . . . . . . . . . . . .
. . . . 2687 Sharpnose halfbeak . . . . . . . . . . . . . . 2194
Selaroides leptolepis . . . . . . . . .
. . . . 2737 Sharpnosed river mullet . . . . . . . . . . . . 2081
Selenanthias . . . . . . . . . . . . .
. . . . 2469 Shawra . . . . . . . . . . . . . . . . . . . . . 2644
sem, Caranx . . . . . . . . . . . . . .
. . . . 2711 Shelf beauties . . . . . . . . . . . . . . . . . 2438
semidentatus, Gargariscus . . . . . .
. . . . 2366 Shortfin bigeye . . . . . . . . . . . . . . . . . 2599
semifasciatus, Branchiostegus . . . .
. . . . 2632 Shortfin flyingfish . . . . . . . . . . . . . . . 2178
septemfasciatus, Epinephelus . . . .
. . . . 2497 Shortfin scad . . . . . . . . . . . . . . . . . . 2720
Seriatopora . . . . . . . . . . . . . . . . . . 2342 Shortspined waspfish . . . . . . . . . . . . . 2309
Shoulderbar soldierfish . . . . . . . . . . . . 2241
Seriola . . . . . . . . . . . . . . 2660-2662, 2678
Shrimp scad . . . . . . . . . . . . . . . . . . 2687
Seriola auerovittata . . . . . . . . . . . . . . 2740 Shrimpfish . . . . . . . . . . . . . . . . . . . 2282
Seriola banisteri . . . . . . . . . . . . . . . . 2740 Shrimpfishes . . . . . . . . . . . . . . . . . . 2281
Seriola bovinoculata . . . . . . . . . . . . . 2741 signifer, Iracundus . . . . . . . . . . . . . . 2317
Seriola colburni . . . . . . . . . . . . . . . . 2741 signifer, Pseudomugil . . . . . . . . . . . . . 2112
Seriola dorsalis . . . . . . . . . . . . . . . . 2740 sihama, Sillago . . . . . . . . . . . . . . . . 2629
Seriola dumerili . . . . . . . . . . . . . . . . 2738 Silago arena . . . . . . . . . . . . . . . . . . 2623
Seriola falcata . . . . . . . . . . . . . . . . . 2741 Silago asiatico . . . . . . . . . . . . . . . . . 2620
Seriola grandis . . . . . . . . . . . . . . . . 2740 Silago aureolineado . . . . . . . . . . . . . . 2618
Seriola hippos . . . . . . . . . . . . . . . . . 2739 Silago de bahia . . . . . . . . . . . . . . . . 2624
Seriola lalandi . . . . . . . . . . . . . . . . 2740 Silago de bandas plateadas . . . . . . . . . . 2619
Seriola pappei . . . . . . . . . . . . . . . . . 2740 Silago de fango . . . . . . . . . . . . . . . . 2626
Seriola purpurescens . . . . . . . . . . . . . 2738 Silago escamoso . . . . . . . . . . . . . . . 2627
Silago intermedio . . . . . . . .
. . . . . . . 2625 songoro, Seriola . . . . . . . . . . . . . . . . 2741
Silago oriental . . . . . . . . . .
. . . . . . . 2617 sonnerati, Cephalopholis . . . . . . . . . . . 2482
Silago pateta . . . . . . . . . .
. . . . . . . 2622 Sorsogona tuberculata . . . . . . . . . . . . 2415
Silago plateado . . . . . . . . .
. . . . . . . 2629 SPARIDAE . . . . . . . . . . . . . . . . . . 2443
Silago tromperero occidental . .
. . . . . . . 2621 See also Vol. 5
Silago trompetero . . . . . . . .
. . . . . . . 2628 speciosus, Caranx . . . . . . . . . . . . . . . 2725
SILLAGINIDAE . . . . . . . . .
. . . . . . . 2614 speciosus, Gnathanodon . . . . . . . . . . . 2725
Sillaginopodys . . . . . . . . .
. . . . . . . 2615 Speckled grouper . . . . . . . . . . . . . . . 2494
Sillago aeolus . . . . . . . . . .
. . . 2617, 2621 speculiger, Hirundichthys . . . . . . . . . . 2177
Sillago analis . . . . . . . . . .
. . . 2618, 2623 Speigler’s mullet . . . . . . . . . . . . . . . . 2108
Sillago argentifasciata . . . . .
. . . 2619, 2624 speigleri, Valamugil . . . . . . 2095, 2107-2108
Sillago asiatica . . . . . . . . .
. . . . . . . 2620 Sphaeramia nematoptera . . . . . . . . . . . 2610
Sillago burrus . . . . . . . . . .
. . . 2617, 2621 Sphaeramia orbicularis . . . . . . . . . . . . 2610
Sillago chondropus . . . . . . .
. . . . . . . 2622 sphenops, Poecilia . . . . . . . . . . . . . . 2199
Sillago ciliata . . . . . . . . . .
. . . . . . . 2623 SPHYRAENIDAE . . . . . . . . . . . 2151, 2651
Sillago ingenuua . . . . . . . .
. . . . . . . 2624 See also Vol. 3
Sillago intermedius . . . . . . .
. . . . . . . 2625 Sphyraenops . . . . . . . . . . . . . . . . . . 2611
Sillago lutea . . . . . . . . . . .
. . . . . . . 2626 Sphyraenops bairdianus . . . . . . . . . . . 2613
Sillago macrolepis . . . . . . .
. . . . 2626-2627 spilonotopterus, Cheilopogon . . . . . . . . 2172
Sillago maculata . . . . . . . .
2617, 2621, 2628 spiloparaea, Cephalopholis . . . . . . . . . . 2542
Sillago maculata burra . . . . .
. . . . . . . 2621 spiloptera, Lepidotrigla . . . . . . . . . . . . 2368
Sillago maculata maculata . . .
. . . . . . . 2625 spilopterus, Cheilopogon . . . . . . . . . . . 2172
Sillago nierstraszi . . . . . . . .
. . . . . . . 2618 spilopterus, Cichlops . . . . . . . . . . . . . 2571
Sillago sihama . . . . . . . . .
. . . . . . . 2629 spilotoceps, Epinephelus . . . . 2500, 2503, 2512,
Sillagos . . . . . . . . . . . . .
. . . . . . . 2614 2518, 2524
Silver sillago . . . . . . . . . . .
. . . . . . . 2629 Spineblotch scorpionfish . . . . . . . . . . . . 2345
Silver tilefish . . . . . . . . . . .
. . . . . . . 2642 spiniferum, Sargocentron . . . . . . . . . . 2252
Silverbanded sillago . . . . . . .
. . . . . . . 2619 spiniferus, Holocentrus . . . . . . . . . . . . 2252
Silvermouth trevally . . . . . . .
. . . . . . . 2752 spiniger, Epinephelus . . . . . . . . . . . . . 2504
Silversides . . . . . . . . . . . .
. . . . . . . 2113 spinosa, Onigocia . . . . . . . . . . . . . . . 2406
Silverspot squirrelfish . . . . . .
. . . . . . . 2249 spinosa, Pterygotrigla . . . . . . . . . . . . . 2380
similis, Oxyporhamphus micropterus . . . . 2196 spinosa, Trigla . . . . . . . . . . . . . . . . . 2365
simplex, Bathysphyraenops . . . . . . . . . 2612 spinosus, Chelidonichthys . . . 2364-2365, 2368
simplex, Seriola . . . . . . . . . . . . . . . . 2738 spinosus, Niphon . . . . . . . . . . . 2429, 2548
simus, Cypselurus . . . . . . . . . . . . . . 2175 Spiny flathead . . . . . . . . . . . . . . . . . 2403
simus, Melamphaes . . . . . . . . . . . . . . 2201 Spiny flatheads . . . . . . . . . . . . . . . . 2422
sinensis, Inimicus . . . . . . . . . . . . . . . 2317 Spinyfins . . . . . . . . . . . . . . . . . . . . 2211
Siphamia . . . . . . . . . . . . . . . . . . . 2602 spiramen, Epinephelus . . . . . . . . . . . . 2521
Sixbar grouper . . . . . . . . . . . . . . . . . 2522 splendens, Beryx . . . . . . . . . . . . 2221-2222
Sixblotch hind . . . . . . . . . . . . . . . . . 2481 Splendid perches . . . . . . . . . . . . . . . 2553
slacksmithi, Epinephelus . . . . . . . . . . . 2514 Splitfins . . . . . . . . . . . . . . . . . . . . 2436
Slender alfonsino . . . . . . . . . . . . . . . 2222 Spongyjawed needlefish . . . . . . . . . . . . 2161
Slender deepwater cardinalfish . . . . . . . . 2613 Spotfin flyingfish . . . . . . . . . . . . . . . . 2170
Slender grouper . . . . . . . . . . . . . . . . 2475 Spottail needlefish . . . . . . . . . . . . . . . 2158
Slender scorpionfish . . . . . . . . . . . . . . 2319 Spotted armoured gurnard . . . . . . . . . . 2381
Slimeheads . . . . . . . . . . . . . . . . . . 2215 Spotted bullheaded gurnard . . . . . . . . . . 2375
Slopefishes . . . . . . . . . . . . . . . . . . 2438 Spotted coralgrouper . . . . . . . . . . . . . 2534
Smalleyed flathead . . . . . . . . . . . . . . 2395 Spotted flathead . . . . . . . . . . . . . . . . 2393
Smallmouth scad . . . . . . . . . . . . . . . 2686 Spotted grouper . . . . . . . . . . . . . . . . 2510
Smallscale scorpionfish . . . . . . . . . . . . 2343 Spotted gurnard . . . . . . . . . . . . . . . . 2372
Smallspined flathead . . . . . . . . . . . . . 2399 Spotted hardyhead . . . . . . . . . . . . . . 2131
Smallspotted dart . . . . . . . . . . . . . . . 2745 Spotted stonefish . . . . . . . . . . . . . . . 2317
smirnovi, Inimicus . . . . . . . . . . . . . . 2316 Spotwing gurnard . . . . . . . . . . . . . . . 2368
Smooth razorfish . . . . . . . . . . . . . . . 2282 Spotwing helmet gurnard . . . . . . . . . . . 2286
Smoothnose armoured gurnard . . . . . . . . 2370 Squalomugil nasutus . . . . . . . . . . . . . 2101
Snipefishes . . . . . . . . . . . . . . . . . . 2280 squamipinnis, Pseudanthias . . . . . . . . . 2480
Snubnose grouper . . . . . . . . . . . . . . . 2508 Squaretail coralgrouper . . . . . . . . . . . . 2531
Snubnose pompano . . . . . . . . . . . . . . 2746 Squaretail mullet . . . . . . . . . . . . . . . . 2094
Soapfishes . . . . . . . . . . . . . . . . . . 2442 Squirrelfishes . . . . . . . . . . . . . . . . . 2225
socialis, Epinephelus . . . . . . . . . . . . . 2523 staigeri, Cymbacephalus . . . . . . . . . . . 2397
Soldierfishes . . . . . . . . . . . . . . . . . . 2225 Stained flyingfish . . . . . . . . . . . . . . . . 2172
SOLENOSTOMIDAE 2263, 2265, 2277, 2280-2281 Stalix . . . . . . . . . . . . . . . . . . . . . 2588
Stareyed stonefish . . . . . . . . . . . . . . 2318 SYNGNATHIDAE . . . . . . . . 2263-2264, 2278

Stargazing stonefish .. . . . . . . . . . . . . 2351 Syngnathids . . . . . . . . . . . . . . . . . . 2265
Starry helmet gurnard . . . . . . . . . . . . . 2289 SYNGNATHINAE . . . . . . . . . . . . . . . 2264
Starspotted grouper . . . . . . . . . . . . . . 2503 Syngnathoides biaculeatus . . . . . . . . . . 2276
Steeneichthys . . . . . . . . . . . . . . . . . 2578
stellatus, Caranx . . . . . . . . . . . . . . . 2714 T
Stenatherina honoriae . . . . . . . . . . . . 2130 tabl, Decapterus . . . . . . . . . . . . . 2722-2723
Stenatherina panatela . . . . . . . . . . . . 2139 Tade mullet . . . .. . . . . .
. . . . . . . . 2091
Stenatherina temminckii . . . . . . . 2136, 2139 tade, Liza . . . . .. . . . . .
. . . . . . . . 2091
STEPHANOBERYCIFORMES . . . . . 2201-2209 tade, Mugil . . . .. . . . . .
. . . . . . . . 2091
stictus, Epinephelus . . . . . . . . . . . . . 2525 Taenianotus . . . .. . . . . .
. . . . . . . . 2354
Stingfishes . . . . . . . . . . . . . . . . . . . 2291 Taenianotus triacanthus . . .
. . . . . . . . 2349
stoliczkae, Sebastes . . . . . . . . . . . . . . 2549 taenianotus, Ablabys . . . . .
. . . . . . . . 2309
Stonefishes . . . . . . . . . . . . . . . . . . 2291 Taeniomembras tropicalis . . .
. . . . . . . . 2137
Stour longtom . . . . . . . . . . . . . . . . . 2161 Tagala gurnard . . . . . . . .
. . . . . . . . 2380
Stoutspine waspfish . . . . . . . . . . . . . . 2324 tagala, Pterygotrigla . . . . .
. . . . 2360, 2380
Straighthead soldierfish . . . . . . . . . . . . 2246 Tahitian squirrelfish . . . . . .
. . . . . . . . 2253
Strawberry dottyback . . . . . . . . . . . . . 2576 tala, Chorinemus . . . . . . .
. . . . . . . . 2733
Strawberry hind . . . . . . . . . . . . . . . . 2542
tala, Scomberoides . . . . . .
. . . . . . . . 2733
strigatus, Aeoliscus . . . . . . . . . . . . . . 2282 talamparoides, Carangoides . . . . 2704, 2709 .
Striped grouper . . . . . . . . . . . . . . . . 2506 Talang queenfish . . . . . . . . . . . . . . 2731
.
Striped stingfish . . . . . . . . . . . . . . . . 2320
STROMATEIDAE . . . . . . . . . . . . . . . 2660
tanakai, Caranx . . . . . . . . . . . . . . . 2705
.
See also Vol. 6 tapeinometapon, Seriola . . . . . . . . . . 2738
.
strongia, Sebastapistes . . . . . . . . . . . . 2346 tapeinosoma, Pseudochromis . . . . . . . . 2573
Tapetails . . . . . . . . . . . . . . . . . . . . 2208
strongylocephalus, Mugil . . . . . . . . . . . 2106
Tassergal . . . . . . . . . . . . . . . . . . . 2650
Strongylura incisa . . . . . . . . . . . . . . 2156
tauvina, Epinephelus 2493, 2505, 2511, 2515, 2526
Strongylura leiura . . . . . . . . . . . . . . 2157
tayenus, Priacanthus . . . . . . . . . . . . . 2598
Strongylura strongylura . . . . . . . . . . . 2158 TELMATHERINIDAE . . . . . . . . . . . . . 2144
Strongylura urvillii . . . . . . . . . . . . . . 2161 temminckii, Hypoatherina . . . 2123, 2133, 2136
strongylura, Strongylura . . . . . . . . . . . 2158 temminckii, Stenatherina . . . . . . . 2136, 2139
strongylura, Tylosurus . . . . . . . . . . . . 2158 Temnodon saltator . . . . . . . . . . . . . . 2650
Stylophora . . . . . . . . . . . . . . . . . . 2353 Temperate ocean-basses . . . . . . . . . . . 2436
subducens, Congrogadus . . . . . . . . . . . 2569 tertius, Sebastiscus . . . . . . . . . . . . . . 2347
subviridis, Liza . . . . . . . . . 2088-2091, 2093 Testolín de aleta azul . . . . . . . . . . . . . 2364
sudanensis, Mugil . . . . . . . . . . . . . . . 2093 Tetraroge barbata . . . . . . . . . . . . . . . 2350
Suggrundus macracanthus . . . . . . . . . 2416 Tetraroge niger . . . . . . . . . . . . . . . . 2350
Suggrundus sp. 1 . . . . . . . . . . . . . . . 2392 TETRAROGINAE . . . . . . . . . . . . . . . 2292
Suggrundus sp. 2 . . . . . . . . . . . . . . . 2411 Tetrarogine scorpaenids . . . . . . . . . . . . 2354
Suggrundus sundaicus . . . . . . . . . . . . 2416 Thalia sp. . . . . . . . . . . . . . . . . . . . 2529
suillus, Epinephelus . . . . . . . . . . . . . . 2493 Theena . . . . . . . . . . . . . . . . . . . . . 2644
Sumatran silverside . . . . . . . . . . . . . . 2138 Thorny flathead . . . . . . . . . . . . . . . . 2412
summana, Epinephelus . . . . . . . 2491, 2514 Thorny seahorse . . . . . . . . . . . . . . . . 2272
sundaicus, Suggrundus . . . . . . . . . . . . 2416 Threadfinned dottyback . . . . . . . . . . . . 2575
suppositus, Mugil . . . . . . . . . . . . . . . 2108 Three-by-two garfish . . . . . . . . . . . . . . 2193
Surf sardine . . . . . . . . . . . . . . . . . . 2141 Threefinger scorpionfish . . . . . . . . . . . . 2311
Surf sardines . . . . . . . . . . . . . . . . . 2140 Threespotted waspfish . . . . . . . . . . . . 2321
Surfette commune . . . . . . . . . . . . . . . 2141 Thysanophrys . . . . . . . . . . . . . . . . . 2385
Surge grouper . . . . . . . . . . . . . . . . . 2523 Thysanophrys arenicola . . . . . . . . . . . 2417
Surgeonfishes . . . . . . . . . . . . . . . . . 2260 Thysanophrys carbunculus . . . . . . . . . . 2418
Sutton’s flyingfish . . . . . . . . . . . . . . . 2173 Thysanophrys celebicus . . . . . . . . . . . 2419
suttoni, Cheilopogon . . . . . . . . . . . . . 2173 Thysanophrys chiltonae . . . . . . . . . . . 2420
Suttonia . . . . . . . . . . . . . . . . . . . . 2579 Thysanophrys otaitensis . . . . . . . . . . . 2421
Swallowtail dart . . . . . . . . . . . . . . . . 2748 tiere, Sargocentron . . . . . . . . . . . . . . 2253
Swordspine gunard . . . . . . . . . . . . . . 2374 Tiki soldierfish . . . . . . . . . . . . . . . . . 2255
Symphysanodon maunaloae . . . . . . . . . 2440 tiki, Myripristis . . . . . . . . . . . . . . . . 2255
Symphysanodon typus . . . . . . . . . 2440-2441 Tile taches de son . . . . . . . . . . . . . . . 2644
SYMPHYSANODONTIDAE . . . . . . . . . . 2438 Tilefishes . . . . . . . . . . . . . . . . . . . . 2630
Synanceia horrida . . . . . . . . . . . . . . 2348 Tille trevally . . . . . . . . . . . . . . . . . . 2717
Synanceia verrucosa . . . . . . . . . . . . . 2349 tille, Caranx . . . . . . . . . . . . . . . . . . 2717
SYNANCEIINAE . . . . . . . . . . . . 2292-2293 tiltoni, Dactyloptena . . . . . . . . . . . . . 2290
timorensis, Epinephelus . . . . . . . . . . . 2546 tuamotuensis, Epinephelus . . . 2513, 2515, 2546

tinkhami, Sebastapistes . . . . . . . . . . . . 2346 tuberculata, Sorsogona . . . . . . . . . . . . 2415
Tinselfishes . . . . . . . . . . . . . . . . . . 2260 Tuberculated flathead . . . . . . . . . . . . . 2415
tol, Chorinemus . . . . . . . . . . . . . . . . 2734 tukula, Epinephelus . . . . . . . . . . . . . 2527
tol, Scomberoides . . . . . . . . . . . . . . . 2734 Turrum emburyi . . . . . . . . . . . . . . . . 2700
Tomato hind . . . . . . . . . . . . . . . . . . 2482 Twinspot grouper . .. . . . . . . . . . . . . 2542
Torpedo scad . . . . . . . . . . . . . . . . . 2726 Twinspot lionfish . . .. . . . . . . . . . . . . 2313
Trachicephalus uranoscopus . . . . . . . . 2351 Tylosurus . . . . . .. . . . . . . . . . . . . 2160
TRACHICHTHYIDAE . . . 2210-2212, 2215, 2219 Tylosurus acus . . .. . . . . . . . . . . . . 2160
Trachichthyids . . . . . . . . . . . . . . . . . 2215 Tylosurus acus melanotus . . . . . . . . . . 2159
trachinoides, Vespicula . . . . . . . . . . . . 2352 Tylosurus crocodilus . . . . . . . . . . . . . 2159
Trachinotus . . . . . . . . . . . . . . 2660, 2679 Tylosurus crocodilus crocodilus . . . . . . . 2160
Trachinotus africanus . . . . . . . . . . . . 2743 Tylosurus crocodilus fodiator . . . . . . . . 2160
Trachinotus anak . . . . . . . . . . . . . . . 2744 Tylosurus gavialoides . . . . . . . . . . . . . 2161
Trachinotus baillonii . . . . . . . . . . . . . 2745 Tylosurus punctulatus . . . . . . . . . . . . 2161
Trachinotus blochii . . . . . . . . . . . . . . 2746 Tylosurus raphidoma . . . . . . . . . . . . . 2160
Trachinotus botla . . . . . . . . . . . . 2747-2748 Tylosurus strongylura . . . . . . . . . . . . . 2158
Trachinotus copperingi . . . . . . . . . . . 2748 typus, Caranx . . . . . . . . . . . . . . . . . 2696
Trachinotus mookalee . . . . . . . . . . . . 2749 typus, Pseudoplesiops . . . . . . . . . . . . . 2577
Trachinotus russelli . . . . . . . . . . 2747-2748 typus, Symphysanodon . . . . . . . . . 2440-2441
Trachinotus velox . . . . . . . . . . . . . . . 2748
Trachurops crumenophthalmus . . . . . . . . 2736 U
Trachurus . . . . . . 2655, 2661, 2681, 2750-2751 uii, Carangoides . . . . . . . . . . . . . . . . 2695
Trachurus declivis . . . . . . . . . . . 2750-2751 uisila, Hypoatherina . . . . . . . . . 2133, 2136
Trachurus mccullochi . . . . . . . . . . . . . 2751 Ulua . . . . . . . . . . . . . . . . . . 2661, 2681
Trachurus novaezelandiae . . . . . . . 2750-2751 Ulua aurochs . . . . . . . . . . . . . . . . . 2752
trachyacron, Myripristis . . . . . . . . . . . 2255 Ulua mandibularis . . . . . . . . . . . . . . 2753
trachycephalus, Minous . . . . . . . . . . . 2320 Ulua mentalis . . . . . . . . . . . . . . . . . 2753
Trachyrhamphus bicoarctatus . . . . . . . . 2276 Uncle Phoo’s groppo . . . . . . . . . . . . . 2556
Trachystoma petardi . . . . . . . . . . . . . 2098 undulatostriatus, Epinephelus . . . . . . . . 2528
Trachystoma ramsayi . . . . . . . . . . . . . 2092 undulatus, Peristedion . . . . . . . . . . . . . 2366
Trevallies . . . . . . . . . . . . . . . . . . . 2659 undulosus, Epinephelus . . . . . . . . . . . 2529
TRIACANTHIDAE . . . . . . . . . . . . . . . 2280 unicolor, Cheilopogon . . . . . . . . . . . . 2173
See also Vol. 6 unifasciatus, Hyporhamphus . . . . . . . . . 2191
triacanthus, Taenianotus . . . . . . . . . . . 2349 Uradia macrolepidota . . . . . . . . . . . . . 2382
Trigla arabica . . . . . . . . . . . . . . . . . 2373 uranoscopus, Trachicephalus . . . . . . . . 2351
Trigla kumu . . . . . . . . . . . . . . . . . . 2364 Uraspis . . . . . . . . . . . . . 2661, 2682, 2755
Trigla leptacanthus . . . . . . . . . . . . . . 2375 Uraspis cadenati . . . . . . . . . . . . . . . 2755
Trigla spinosa . . . . . . . . . . . . . . . . . 2365 Uraspis helvola . . . . . . . . . . . . . 2754-2755
TRIGLIDAE . . . . 2283, 2292, 2359, 2422, 2551 Uraspis pectoralis . . . . . . . . . . . . . . . 2756
TRIGLINAE . . . . . . . . . . . . . . . . . . 2359
Uraspis reversa . . . . . . . . . . . . . . . . 2755
trimaculatus, Neocentropogon . . . . . . . . 2321 Uraspis secunda . . . . . . . . . . . . . 2754-2755
Triplespine deepwater cardinalfish . . . . . . 2613
Uraspis uraspis . . . . . . . . . . . . . . . . 2756
Triso dermopterus . . . . . . . . . . . . . . 2547 Uraspis wakiyai . . . . . . . . . . . . . . . . 2755
trispilos, Cichlops . . . . . . . . . . . . . . . 2571
Trompetero . . . . . . . . . . . . . . . . . . 2280
uraspis, Caranx . . . . . . . . . . . . . . . . 2756
Trompetero chino . . . . . . . . . . . . . . . 2277 uraspis, Uraspis . . . . . . . . . . . . . . . . 2756
Trompette chinoise . . . . . . . . . . . . . . 2277 urodeta, Cephalopholis . . . . . . . . . . . . 2483
trophis, Epinephelus . . . . . . . . . . . . . 2546 urophthalmus, Serranus . . . . . . . . . . . . 2475
Tropical silverside . . . . . . . . . . . . . . . 2122 Urville’s needlefish . . . . . . . . . . . . . . . 2161
Tropical two-wing flyingfish . . . . . . . . . . 2176 urvillii, Strongylura . . . . . . . . . . . . . . 2161
tropicalis, Atherinosoma . . . . . . . . . . . 2137 Usacaranx georgianus . . . . . . . . . . . . 2730
tropicalis, Hypoatherina . . . . . . . . . . . 2137
tropicalis, Taeniomembras . . . . . . . . . . 2137 V
Tropidostethops rhothophilus . . . . . . . . . 2141 vaigiensis, Centrogenys . . . . . . . . 2293, 2549
Tropidostethus rhothophilus . . . . . . . . . 2141 vaigiensis, Ellochelon . . . . . . . . . . . . . 2094
troscheli, Mugil . . . . . . . . . . . . . . . . 2088 vaigiensis, Liza . . . . . . 2089, 2091, 2093-2094
Trumpeter sillago . . . . . . . . . . . . . . . 2628 vaigiensis, Pranesus . . . . . . . . . . . . . . 2125
Trumpetfishes . . . . . . . . . . . . . . . . . 2277 vaigiensis, Psammoperca . . . . . . . . . . . 2432
truncatus, Epinephelus . . . . . . . . . . . . 2520 Valamugil . . . . . . . . . . . . 2069-2070, 2103
truncatus, Plectropomus . . . . . . . . . . . 2531 Valamugil buchanani . . . . . . . . . 2102, 2107
Valamugil cunnesius . . . . . . . 2102-2103-2108 Volador de Sutton . . . . . . . . . . . . . . . 2173

Valamugil engeli . . . . . . . . 2102, 2104-2106 Volador escamudo . . . . . . . . . . . . . . . 2174
Valamugil georgii . . . . . . . . . . . . . . 2105 Volador espejo . . . . . . . . . . . . . . . . . 2177
Valamugil perusii . . . . . . . . . . . . 2104-2106 Volador farsón . . . . . . . . . . . . . . . . . 2174
Valamugil seheli . . . . . . . . . . . 2102, 2107 Volador jaspeado . . . . . . . . . . . . . . . 2172
Valamugil speigleri . . . . . . . 2095, 2107-2108 Volador manchado . . . . . . . . . . . . . . 2170
valenciennei, Allanetta . . . . . . . . . . . . 2138 Volador marinero . . . . . . . . . . . . . . . 2179
valenciennei, Atherina . . . . . . . . . . . . 2138 Volador ñato . . . . . . . . . . . . . . . . . . 2176
valenciennei, Hypoatherina . . . . . 2134, 2138 Volador picudo . . . . . . . . . . . . . . . . . 2173
Volador planeador . . . . . . . . . . . . . . . 2169
valenciennei, Pranesus . . . . . . . . . . . . 2138
valentini, Aulostomus . . . . . . . . . . . . . 2277
volitans, Exocoetus . . . . . . . . . . . . . . 2176
valentini, Canthigaster . . . . . . . . . . . . 2532
volitans, Pterois . . . . . . . . . . . . . 2330-2332
vari, Alepes . . . . . . . . . . . . . . 2687, 2690
vomerinus, Caranx . . . . . . . . . . . . . . 2707
variegatum, Plectropoma . . . . . . . . . . . 2535 W
Variola . . . . . . . . . . . . . . . . . . . . 2469
Variola albimarginata . . . . . . . . . . . . 2538 Waigeo silverside . . ... . . . . . . . . . . 2121
Variola louti . . . . . . . . . . . . . . 2538-2539 waigiensis, Liza . . . ... . . . . . . . . . . 2094
varipinnis, Scorpaenodes . . . . . . . . . . . 2340 Waigieu seaperch . . ... . . . . . . . . . . 2432
veliferus, Pseudochromis . . . . . . . . . . . 2572 wakiyai, Uraspis . . . ... . . . . . . . . . . 2755
Wampeejawed fishes ... . . . . . . . . . . 2438
velitaris, Macrorhamphosus . . . . . . . . . 2280
Ward’s tilefish . . . . ... . . . . . . . . . . 2646
velox, Trachinotus . . . . . . . . . . . . . . . 2748
Velvetfishes . . . . . . . . . . . . . . . . . . 2354
wardi, Branchiostegus . . . . . 2642, 2644-2646
Waspfishes . . . . . . . . . . . . . . 2291, 2354
venosa, Scorpaenopsis . . . . . . . . . . . . 2344 Wavylined grouper . . . . . . . . . . . . . . . 2529
verrucosa, Synanceia . . . . . . . . . . . . . 2349 Weedy scorpionfish . . . . . . . . . . . . . . 2332
versicolor, Minous . . . . . . . . . . . . . . 2321 Weedy stingfish . . . . . . . . . . . . . . . . 2341
vespa, Paracentropogon . . . . . . . . . . . 2325 Welander’s flathead . . . . . . . . . . . . . . 2414
Vespicula cypho . . . . . . . . . . . . . . . 2351 welanderi, Rogadius . . . . . . . . . . . . . 2414
Vespicula depressifrons . . . . . . . . . . . 2352 Western trumpeter sillago . . . . . . . . . . . 2621
Vespicula trachinoides . . . . . . . . . . . . 2352 Whalefishes . . . . . . . . . . . . . . . . . . 2206
Vielle alle noire . . . . . . . . . . . . . . . . 2483 Whalesucker, . . . . . . . . . . . . . . . . . 2652
Vielle ananas . . . . . . . . . . . . . . . . . 2482 White sucker . . . . . . .. . . . . . . . . . . 2652
Vielle arlequin . . . . . . . . . . . . . . . . . 2541 White tilefish . . . . . . .
. . . . . . . . . . . 2641
Vielle chocolat . . . . . . . . . . . . . . . . . 2540 White trevally . . . . . .. . . . . . . . . . . 2730
Vielle ciosomoer . . . . . . . . . . . . . . . . 2476 White-edged lyretail . . .. . . . . . . . . . . 2538
Vielle de corail . . . . . . . . . . . . . . . . . 2480 White’s seahorse . . . . . . . . . . . . . . . 2272
Vielle dorée . . . . . . . . . . . . . . . . . . 2540 Whiteband bigeye . . . . . . . . . . . . . . . 2601
Vielle étoiles bleues . . . . . . . . . . . . . . 2477 Whitebellied rougefish . .. . . . . . . . . . . 2333
Vielle fraise . . . . . . . . . . . . . . . . . . 2542 Whiteblotched grouper . . . . . . . . . . . . 2545
Vielle léopard . . . . . . . . . . . . . . . . . 2541 Whitedotted grouper . . . . . . . . . . . . . . 2517
Vielle lignes bleues . . . . . . . . . . . . . . 2478 Whiteface waspfish . . . . . . . . . . . . . . 2325
Vielle plate grise . . . . . . . . . . . . . . . . 2545 Whitefin bigeye . . . . . . . . . . . . . . . . 2600
Vielle roga . . . . . . . . . . . . . . . . . . . 2474 Whitefin trevally . . . . .
. . . . . . . . . . . 2698
Vielle rouillé . . . . . . . . . . . . . . . . . . 2540 whitei, Hippocampus . . . . . . . . . . . . . 2272
Vielle Saint-Silac . . . . . . . . . . . . . . . 2534 Whitemouth jack . . . . . . . . . . . . . . . . 2756
Vielle six taches . . . . . . . . . . . . . . . . 2481 Whitespot flyingfish . . .. . . . . . . . . . . 2177
Vielle tache de rousseur . . . . . . . . . . . 2541 Whitespotted grouper . . . . . . . . . . . . . 2491
Vielle taches blanches . . . . . . . . . . . . 2491 Whitestreaked grouper . . . . . . . . . . . . 2514
Vielle voyant . . . . . . . . . . . . . . . . . . 2479 Whitetip soldierfish . . . .
. . . . . . . . . . . 2255
violacea, Myripristis . . . . . . . . . . . . . 2243 Whitetongue jack . . . . . . . . . . . . . . . 2754
Violet soldierfish . . . . . . . . . . . . . . . . 2243 Whitley’s silverside . . .. . . . . . . . . . . 2137
viridis, Euleptorhamphus . . . . . . 2180, 2193 williamsi, Caranx . . . . . . . . . . . . . . . 2711
vittata, Myripristis . . . . . . . . . . . . . . 2255 Wood’s soldierfish . . . .. . . . . . . . . . . 2256
Volador . . . . . . . . . . . . . . . . . . . . 2176 woodsi, Myripristis . . . . . . . . . . . . . . 2256
Volador aletón . . . . . . . . . . . . . . . . . 2178 woodwardi, Hypoatherina . . . . . . 2134, 2138
Volador aletón africano . . . . . . . . . . . . 2178 wrighti, Pseudocaranx . . . . . . . . . . . . 2730
Volador barbudo . . . . . . . . . . . . . . . . 2176
Volador bordiblanco . . . . . . . . . . . . . . 2170 X
Volador casquete . . . . . . . . . . . . . . . 2177
Volador de ala amarilla . . . . . . . . . . . . 2175 xanthochir, Pseudochromis . . . . . . . . . . 2574
Volador de ala corta . . . . . . . . . . . . . . 2178 xanthopterus, Cephalopholis . . . . . . . . . 2477
Volador de dorso manchado . . . . . . . . . 2170 Xenolepidichthys . . . . . . . . . . . . . . . 2260
Volador de mancha blanca . . . . . . . . . . 2177 Xiphasia . . . . . . . . . . . . . . . . . . . . 2587
Yellowtail scad . . . . . . . . . . . . . . . . . 2692

Y Yellowtailed flathead . . . . . . . . . . . . . . 2408
Yellow grouper . . . . . . . . . . . . . . . . 2488
Yellow seahorse . . . . . . . . . . . . . . . . 2272 Z
Yellow waspfish . . . . . . . . . . . . . . . . 2312 zaiserae, Priacanthus . . . . . . . . . . . . . 2600
Yellowbar pipefish . . . . . . . . . . . . . . . 2275 zaspilota, Ocosia . . . . . . . . . . . . . . . 2325
Yellowedged lyretail . . . . . . . . . . . . . . 2539 Zebra lionfish . . . . . . . . . . . . . . . . . 2314
Yelloweye mullet . . . . . . . . . . . . . . . 2079 zebra, Dendrochirus . . . . . . . . . . . . . 2314
Yellowfin bigeye . . . . . . . . . . . . . . . . 2600
ZEIDAE . . . . . . . . . . . . . . . . 2257, 2259
Yellowfin needlefish . . . . . . . . . . . . . . 2157
Zeids . . . . . . .. . . . . . . . . . . . . . . 2259
Yellowfin soldierfish . . . . . . . . . . . . . . 2240
ZEIFORMES . . . . . . . . . . . . . . . 2257-2261
Yellowing flyingfish . . . . . . . . . . . . . . 2175
Yellowspotted grouper . . . . . . . . . . . . 2546
Zenarchopterus . . . . . . . . . . . . . . . . 2181
Yellowspotted scorpionfish . . . . . . . . . . 2344 Zenion . . . . . . . . . . . . . . . . . . . . . 2258
Yellowspotted trevally . . . . . . . . . . . . . 2700 Zenion hololepis . . . . . . . . . . . . . . . 2258
Yellowstripe scad . . . . . . . . . . . . . . . 2737 Zenopsis . . . . . . . . . . . . . . . . . . . . 2259
Yellowstriped cardinalfish . . . . . . . . . . . 2608 Zeus . . . . . . . . . . . . . . . . . . . . . . 2259
Yellowtail amberjack . . . . . . . . . . . . . 2740 Zonichthys nigrofasciata . . . . . . . . . . . 2742
Yellowtail horse mackerel . . . . . . . . . . . 2751
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2790 List of Colour Plates
LIST OF COLOUR PLATES
PLATE I 28. Epinephelus macrospilos

1. Aethaloperca rogaa 29. Epinephelus maculatus
2. Anyperodon leucogrammicus 30. Epinephelus magniscuttis
3. Cephalopholis argus 31. Epinephelus malabaricus
4. Cephalopholis cyanostigma 32. Epinephelus merra
5. Cephalopholis igarashiensis
6. Cephalopholis miniata
7. Cephalopholis sexmaculata
8. Cephalopholis sonnerati PLATE V
33. Epinephelus morrhua
34. Epinephelus ongus
35. Epinephelus polystigma
PLATE II 36. Epinephelus quoyanus
9. Cephalopholis urodeta 37. Epinephelus radiatus
10. Cromileptes altivelis 38. Epinephelus retouti
11. Epinephelus akaara 39. Epinephelus rivulatus
12. Epinephelus amblycephalus 40. Epinephelus sexfasciatus
13. Epinephelus areolatus
14. Epinephelus awoara
15. Epinephelus bleekeri
16. Epinephelus caeruleopunctatus PLATE VI
41. Epinephelus spilotoceps
42. Epinephelus stictus
43. Epinephelus tauvina
PLATE III 44. Epinephelus trimaculatus
17. Epinephelus chlorostigma 45. Epinephelus tukula
18. Epinephelus coioides 46. Epinephelus undulatostriatus
19. Epinephelus cyanopodus 47. Gracila albomarginata
20. Epinephelus epistictus 48. Plectropomus areolatus
21. Epinephelus fasciatomaculosus
22. Epinephelus fasciatus
23. Epinephelus faveatus PLATE VII
24. Epinephelus hexagonatus 49. Plectropomus laevis
50. Plectropomus leopardus
51. Plectropomus maculatus
52. Plectropomus oligacanthus
PLATE IV 53. Plectropomus pessuliferus marisrubri
25. Epinephelus irroratus 54. Saloptia powelli
26. Epinephelus latifasciatus 55. Variola albimarginata
27. Epinephelus longispinis 56. Variola louti
COLOUR PLATES
Family Serranidae (pages 2442 to 2548)
Photographs by J.E. Randall
Abbreviation used: SL = standard lenght

PLATE I
1. Aethaloperca rogaa 29.5 cm SL, Bahrain 2. Anyperodon leucogrammicus 24.2 cm SL, Palau
3. Cephalopholis argus 23.2 cm SL, Teavaraa Pass, 4. Cephalopholis cyanostigma 20.6 cm SL, Ambon Bay,
Tahiti Indonesia
5. Cephalopholis igarashiensis 14.9 cm SL, Naha, 6. Cephalopholis miniata 25.4 cm SL, Ishigaki,
Okinawa, Ryukyu Islands Ryukyu Islands
7. Cephalopholis sexmaculata 25.6 cm SL, Hiva Oa, 8.Cephalopholis sonnerati 16.2 cm SL, Tutuila,
Marquesas Islands American Samoa
PLATE II
9. Cephalopholis urodeta 15.3 cm SL, Tetiaroa Atoll, 10. Cromileptes altivelis 9.3 cm SL, One Tree Island,
Society Islands Great Barrier Reef, Australia
11. Epinephelus akaara 29.8 cm SL, 12. Epinephelus amblycephalus 32.2 cm SL,
Hong Kong Dumaguete, Negros, Philippines
13. Epinephelus areolatus 26.2 cm SL, Nuweiba, 14. Epinephelus awoara 20.8 cm SL, Keelung,
Sinai, Egypt Taiwan Province of China
15. Epinephelus bleekeri 14.8 cm SL, Batu Nampar, 16. Epinephelus caeruleopunctatus 23.7 cm SL,
Lombok, Indonesia Alite Reef, Solomon Islands
PLATE III
17. Epinephelus chlorostigma 32.7 cm SL, 18. Epinephelus coioides 17.5 cm SL,
Port Sudan, Sudan Jakarta, Indonesia
19. Epinephelus cyanopodus 14.5 cm SL, Manila, 20. Epinephelus epistictus 44 cm SL, Ambon,
Philippines Indonesia
21. Epinephelus fasciatomaculosus 8.5 cm SL, 22. Epinephelus fasciatus 21.3 cm SL,
Yeh-Liu, Taiwan Province of China Teavaraa Pass, Tahiti
23. Epinephelus faveatus 22.5 cm SL, Batu Nampar, 24. Epinephelus hexagonatus 16 cm SL, Enewetak,
Lombok, Indonesia Marshall Islands
PLATE IV
25. Epinephelus irroratus 27.9 cm SL, Ua Huka, 26. Epinephelus latifasciatus 27.4 cm SL,
Marquesas Islands Cochin, India
27. Epinephelus longispinis 30.2 cm SL, 28. Epinephelus macrospilos 18.6 cm SL,
Batu Nampar, Lombok, Indonesia Sodwana Bay, Natal, South Africa
29. Epinephelus maculatus 26.3 cm SL, 30. Epinephelus magniscuttis 33.5 cm SL,
Enewetak, Marshall Islands Sodwana Bay, Natal, South Africa
31. Epinephelus malabaricus 31.7 cm SL, 32. Epinephelus merra 9.3 cm SL,
Cochin, India Lord Howe Island, Australia
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PLATE V
33. Epinephelus morrhua 22.4 cm SL, Naha, 34. Epinephelus ongus 22.2 cm SL, Pohnpei,
Okinawa, Ryukyu Islands Caroline Islands
35. Epinephelus polystigma 8.8 cm SL, Ambon Bay, 36. Epinephelus quoyanus 10 cm SL, Poka, Ambon,
Indonesia Indonesia
37. Epinephelus radiatus 29.5 cm SL, Naha, 38. Epinephelus retouti 30.4 cm SL, Papeete,
Okinawa, Ryukyu Islands Tahiti
39. Epinephelus rivulatus 20.6 cm SL, Naha, 40. Epinephelus sexfasciatus 20.8 cm SL, Cebu,
Okinawa, Ryukyu Islands Philippines
PLATE VI
41. Epinephelus spilotoceps 18.8 cm SL, Kwajalein, 42. Epinephelus stictus Western Australia
Marshall Islands (painting by R. Swainston)
43. Epinephelus tauvina 19.2 cm SL, Enewetak, 44. Epinephelus trimaculatus 27 cm SL,
Marshall Islands Keelung, Taiwan Province of China
45. Epinephelus tukula 8.8 cm SL, Kovalam, 46. Epinephelus undulatostriatus 26.4 cm SL,
India One Tree Island, Great Barrier Reef, Australia
47. Gracila albomarginata 23 cm SL, Majuro, 48. Plectropomus areolatus 35 cm SL, Enewetak,
Marshall Islands Marshall Islands
PLATE VII
49. Plectropomus laevis 59.7 cm SL, Enewetak, 50. Plectropomus laevis 47.5 cm SL, Enewetak,
Marshall Islands Marshall Islands
51. Plectropomus leopardus 20.8 cm SL, 52. Plectropomus maculatus 40.2 cm SL

Gill Ayer, Lombok, Indonesia
53. Plectropomus oligacanthus 24A cm SL, 54. Saloptiapowelli 38.8 cm SL,

Alite Reef, Solomon Islands Guam
55. Variola albimarginata 22 cm SL, 56. Variola louti 27.3 cm SL, Enewetak,
Mauritius Marshall Islands

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FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES

THE LIVING MARINE RESOURCES OF THE

THE LIVING MARINE RESOURCES OF THE

WESTERN CENTRAL PACIFIC

with the support of the

South Pacific Forum Fisheries Agency (FFA)

FOOD AND AGRICULTURE ORGANIZATION OF THE UNITED NATIONS

The designations employed and the presentation of material in

All rights reserved. No part of this publication may be reproduced by

Carpenter, K.E.; Niem, V.H. (eds)

Production staff, Species Identification and Data Programme, Marine Resources

Project manager: P. Oliver.

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Mugiliformes: Mugilidae 2069

Mugiliformes: Mugilidae Order MUGILIFORMES

Similar families occurring in the area

ridge of medial lobes

frontal view of head

Fig. 7 Sicamugil hamiltoni Fig. 8 Liza sp.

Fig. 9 Rhinomugil nasutus Fig. 10 Liza sp.

Fig. 12 Oedalechilus labiosus Fig. 13 Neomyxus leuciscus Fig. 14 Liza sp.

adipose eyefold adipose eyefold adipose eyefold

12a. Anal fin usually with 8 (rarely 9) soft

sulcus fat body

papillate denticulate area

teeth b) spatulate teeth

Fig. 20 Myxus elongatus Fig. 21 scales

16a. Anal fin with 8 soft rays in adults; caudal fin

17a. Predorsal scales with keel (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liza affinis

14 transverse scales 10 transverse scales

Fig. 23 Liza argentea Fig. 24 Liza alata

19a. Adults usually with 10 anal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . Liza argentea

Fig. 25 Liza macrolepis Fig. 26 Liza planiceps

Fig. 27 inner surface of upper lip Fig. 28 Liza spp.

a) Liza subviridis b) Liza melinoptera

18-20 scales in transverse 19-20 scales in

16 scales in transverse 16 scales in transverse

Fig. 31 lateral view

List of species occurring in the area

0 Liza macrolepis (Smith, 1846)

0 Liza planiceps (Valenciennes, 1836)

Aldrichetta forsteri (Valenciennes, 1836)

Cestraeus goldiei (Macleay, 1883)

Cestraeus oxyrhynchus Valenciennes, 1836

Cestraeus plicatilis Valenciennes, 1836

Crenimugil crenilabis (Forsskål, 1775)

(from Senou, 1988)

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2084 Bony Fishes

Crenimugil heterocheilus (Bleeker, 1855)

(from Senou, 1988)

Liza affinis (Günther, 1861)

Liza alata (Steindachner, 1892)

Liza argentea (Quoy and Gaimard, 1825)

Liza macrolepis (Smith, 1846) LZM

(from Senou, 1988)

Liza melinoptera (Valenciennes, 1836)

(from Senou, 1988)

Liza parmata (Cantor, 1850)

(after Day, 1876)

Liza planiceps (Valenciennes, 1836)

Liza ramsayi (Macleay, 1883)