FAO Species Identification Guide For Fishery Purposes Western Central Pacific Vol 5

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iv
Table of Contents
Page
BONY FISHES
Order PERCIFORMES: PERCOIDEI
Menidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2791
Leiognathidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2792
Bramidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2824
Caristiidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2837
Emmelichthyidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2838
Lutjanidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2840
Caesionidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2919
Lobotidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2942
Gerreidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2946
Haemulidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2961
Sparidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2990
Lethrinidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3004
Nemipteridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3051
Polynemidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3090
Sciaenidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3117
Mullidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3175
Pempheridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3201
Glaucostomatidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3205
Leptobramidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3210
Bathyclupeidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3211
Toxotidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3212
Monodactylidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3216
Drepanidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3221
Chaetodontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3224
Pomacanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3266
Enoplosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3287
Pentacerotidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3288
Kyphosidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3290
Girellidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3297
Scorpididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3299
Microcanthidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3301
Arripididae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3304
Terapontidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3305
Kuhliidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3317
Cirrhitidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3321
Cheilodactylidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3329
Cepolidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3331
Order PERCIFORMES: LABROIDEI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3333
Cichlidae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3333
Pomacentridae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3337
INDEX . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3357
COLOUR PLATES
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Perciformes: Percoidei: Menidae 2791
Perciformes: Percoidei: Menidae MENIDAE

Moonfish
by D.J. Woodland
A single species in this family.
Mene maculata (Bloch and Schneider, 1801) MOD
Frequent synonyms / misidentifications: None / None.
FAO names: En - Moonfish; Fr - Luneur;
Sp - Lunero.
Diagnostic characters: Body very deep, al-
most triangular, extremely compressed;
breast sharp-edged; greatest body depth
below lateral line at least 4 times greater
than greatest body depth above lateral line.
Mouth small, almost vertical, protrusible; teeth
limited to villiform bands in jaws. Dorsal fin
single, long-based, and low, except for a short
peak anteriorly; spines thin and flexible and
difficult to distinguish from the 34th to 36th soft
rays; X spines in juveniles, reducing in number
with increasing age. Anal fin very long-
based, commencing at point of insertion of
pelvic fins, uniformly very low; II spines in
juveniles, but these and some leading soft
rays become overgrown with skin with in-
creasing age; free tips of visible rays splayed
out. Caudal fin deeply forked. Pectoral fins shorter than head length. Pelvic fins inserted slightly in front of
pectoral fins, with I short spine followed by 5 soft rays, the first 2 soft rays fused and greatly elongated. Lateral
line tracking upwards towards base of last ray of dorsal fin, often terminating there well above midline of caudal
peduncle but sometimes continuing onto base of upper lobe of caudal fin. Body covered with minute scales
invisible to the naked eye, easily brushed off. Colour: upper sides deep metallic blue, remainder silvery, with
a row of round to ovoid, dark slaty-blue spots above and below lateral line, sometimes a few additional spots
above and below these rows; pelvic fins with a trace of blue, other fins hyaline or slightly dusky.
Similar species occurring in the area
All other species with which Mene maculata might be confused have distinct spines in the dorsal and anal
fins, or distinct scales, or keels on the caudal peduncle, and the area of the sides above the midline (eye
to caudal-fin fork) is either greater than or about equal to the area below the midline.
Size: Maximum total length about 30 cm, commonly to 20 cm
Habitat, biology, and fisheries: Inhabits coastal waters near the bottom on both continental shelves and
around major island groups to depths of 200 m; seems to prefer deep water but occasionally enters river
estuaries. Schooling; feeds on small invertebrates. Caught in trawls and by beach seine, but mainly by
trawling, often incidentally, but schools may be sought out by echo survey; flesh excellent in quality but
small in quantity; dried in India and some other Asian countries but also marketed fresh. From 1990 to
1995, the FAO Yearbook of Fishery Statistics reports a range of yearly catch of 4 207 to 9 077 t of M.
maculata from the area (almost exclusively from the Philippines).
Distribution: From East Africa, as far south
as Durban, to the western Pacific; in Aus-
tralia, stragglers as far south as Shark Bay
(Western Australia) and Noosa R. on the
eastern Australian coast; northern limits of
southern Japan.
Reference
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.
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2792 Bony Fishes
LEIOGNATHIDAE
Slipmouths (ponyfishes)
by D.J. Woodland, S. Premcharoen, and A.S. Cabanban
D iagnostic characters: Small to medium-sized fishes (rarely exceeding 16 cm); body oblong or
rounded, moderately to markedly compressed laterally. Eyes moderate to large, preceded by a
short, snubnosed snout. Maxilla concealed under the preorbital, except for the widened posterior end
which curves downward and is tucked into a groove beneath the eye (giving these fishes the
appearance of being “down in the mouth”). Mouth highly protrusible, when extended forming a tube
directed either upwards (Secutor spp.), forward (Gazza spp.), or forward or downward (Leiognathus spp.).
Teeth small, punctate, arranged in 1 or several rows (Gazza spp. additionally with a pair of curved canines
at front of upper jaw and several in lower jaw). A pair of lateral elevated bony ridges on top of head between
eyes, each preceded by a single spine or a pair of small spines, and, medially, another ridge terminating
posteriorly in a bony crest, often referred to as the “nuchal spine”. All fin spines and soft rays fragile; a
single dorsal fin with VIII (rarely VII or IX) spines, the first one very small, and 16 or 17 soft rays; anal
fin with III spines and 14 soft rays; caudal fin deeply emarginate to forked; pectoral fins short; pelvic fins
very short. Trunk covered with small cycloid scales, except for breast and, less frequently, an area
just behind head in some species; head without scales in all but a few species which have a patch
of small scales below eye. Colour: silvery, with characteristic markings on the upper half of sides which
are useful for identification.
nuchal crest
top of head
scaleless
maxilla
maxilla
mouth protruded
Habitat, biology, and fisheries: Bottom-living fishes in shallow coastal waters, with several species
entering brackish waters, especially river estuaries, a few ranging up into fresh water. They occur at depths
as shallow as 0.5 m and down to around 160 m; deeper dwelling species have been reported to move away
from the bottom to midwater at night. All species so far examined possess a light producing organ containing
symbiotic luminescent bacteria; the transparent patch of skin associated with the organ is usually more
elaborately developed in males, indicating perhaps a reproductive as well as schooling role for the signal -
for these fishes live, by and large, in immense schools, often in poorly lit waters. They feed on copepods
and phytoplankton, with large fish feeding predominantly on benthic invertebrates. Members of the genus
Gazza, with their canine teeth, feed on small fishes and shrimps, while Secutor spp., with their upwardly
projecting mouths, feed only on organisms living in the water column. They appear to be short lived, 1 to 2
years, with a protracted breeding season. Leiognathids constituted an important part of the commercial trawl
catches of several Asian countries in the past, but in several areas catches have declined - declining from
20 to 30% of the total demersal catch to as little as 4 or 5% in the last 20 years. Fishing restrictions have
been imposed in some countries. For 1995, the FAO Yearbook of Fishery Statistics reports a total catch of
120 268 t of Leiognathidae from the Western Central Pacific. Commonly marketed fresh or dried-salted, but
surplus catches, especially of small fish, are often converted to fishmeal pellets, used fresh to feed ducks
or farmed fish, or used as manure. Often they are dumped as “nuisance” catch, e.g. by prawn trawlers.
Remarks: The genus Leiognathus is currently being revised by D.J. Woodland. Until the family is completely
revised some of the scientific names of the species used here must be regarded as provisional. The problem
is not so much one of species identification but of establishing to which species some of the very early names
should be applied.
Perciformes: Percoidei: Leiognathidae 2793
Similar families occurring in the area

Gerreidae: body shape similar and mouth also strongly
protrusible, but nuchal crest absent; scales much larger,
and clearly visible, scales also covering top of head and
breast; dorsal-fin spines IX or X (VIII, rarely VII or IX,
in Leiognathidae).
Carangidae: mouth not strongly protrusible; 2 dorsal
fins, or dorsal fin notched; first 2 anal-fin spines some-
what detached from third spine, sometimes deeply em-
Trachinotus sp.
bedded in skin; many scutes along lateral line (but
reduced in some species and absent in some genera).
IX spines
scutes
scales
scales
Pseudocaranx sp.
Gerreidae examples of Carangidae
Key to the species of Leiognathidae occurring in the area

Remarks on key characters: the species of this family are char-
acterized by their life colours (see colour plates at the end of this
volume). While the colours, and even the colour pattern in the
case of the fins, are lost on preservation, the pattern of darker
patches of colour once present on the upper sides is retained. It
is sometimes necessary to hold the fish at just the right angle to
the light to see this pattern. Most species show sexual dimor-
phism; this does not appear to influence the type of colour pattern
on the upper sides, but darker patches on the skin covering the
abdominal cavity area are often much more extensive in males
than females. There is, of course, intraspecies variation in col-
oration; for this reason, a number of specimens from a population cheek and breast cheek and
should be examined if possible. The presence or absence of fully scaly breast naked
scales on cheeks and breast (Fig. 1) is a valuable diagnostic Fig. 1
character, but these scales are lost easily and magnification and
moistening of the relevant area is almost mandatory if residual
scale pockets are to be detected.
1a. Mouthparts protract upwards (Fig. 2a); lower jaw ascends at angle of 85 to 90° when
mouth closed (Fig. 3a); ventral profile of body more convex than dorsal profile; upper half
of body with blue markings when fresh . . . . . . . . . . . . . . . . . . . . . . . . . (Secutor) ® 4
1b. Mouthparts protract forwards to downwards (Fig. 2b, c); lower jaw ascends at an angle
of 40 to 70° when mouth closed (Fig. 3b); dorsal profile of body either equal to or more
convex than ventral profile; upper half of body with dusky markings when fresh . . . . . . . . . . ® 2
a) Secutor b) Gazza and Leiognathus c) Leiognathus a) Secutor b) Gazza and Leiognathus
Fig. 2 Fig. 3
2794 Bony Fishes
2a. Canine teeth present anteriorly in jaws (Fig. 4) . . . . . . . . . (Gazza) ® 3

2b. No canines present, teeth small and weak . . . . . . . . (Leiognathus) ® 8
3a. Body scales absent anterior to a line running from base of

pectoral fins to first soft ray of dorsal fin . . . . . . . . . . Gazza achlamys
3b. Body scales present on upper sides, fully scaly above lateral
line through to margin of nuchal crest . . . . . . . . . . . . . Gazza minuta
4a. Cheeks scaly; breast fully scaly, including isthmus; lateral-line Fig. 4 Gazza
scale rows counted to end of caudal peduncle 43 to 60 . . . . . . . . . ® 5
4b. Cheeks without scales; ventral part of breast not scaly from origin of pelvic fins through
to, and including isthmus; lateral-line scale rows counted to end of caudal peduncle 60
to 115 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5a. Lateral-line scale rows counted to end of caudal peduncle 43 to 49; counting diagonally
from first dorsal-fin spine, 6 or 7 scale rows above lateral line, 13 to 15 below it . Secutor megalolepis
5b. Lateral-line scale rows counted to end of caudal peduncle 54 to 60; counting diagonally
from first dorsal-fin spine, 9 to 14 scale rows above lateral line, 18 to 26 below it . . Secutor ruconius
6a. Lateral-line scale rows counted to end of caudal peduncle 60 to 70 . . . . . . . . . Secutor hanedai
6b. Lateral-line scale rows counted to end of caudal peduncle in excess of 80. . . . . . . . . . . . . ® 7
7a. Body depth 2 to 2.5 times in standard length; on upper half of sides 17 to 22 vertical
markings, interrupted with a dot where they cross the lateral line . . . . . . . . . . . Secutor indicius
7b. Body becoming relatively deeper with growth, body depth 2.5 times in standard length
at 4 cm standard length, only 1.6 at 10 cm standard length; on upper half of sides 11 to
15 vertical markings, interrupted with a dot where they cross the lateral line . . . . Secutor insidiator
8a. Slender bodied, depth usually 2.9 to 4.4 times in standard length, though the body may
be slightly deeper (2.8) in some specimens longer than 10 cm standard length . . . . . . . . . . ® 9
8b. Deep bodied, depth usually much less than 3 times in standard length though values
around 2.9 common in specimens under 3 cm standard length . . . . . . . . . . . . . . . . . . ® 10
9a. Body very slender, its depth 3.5 to 4.4 times in standard length; 55 to 59 tubed lateral-line
scales; upper half of trunk marked with irregularly shaped, solid, dark patches, the
largest of them greater in area than pupil of eye . . . . . . . . . . . . . . . . Leiognathus elongatus
9b. Body less slender, its depth 2.8 to 3.6 times in standard length; around 42 or 43 tubed
lateral-line scales; upper half of trunk marked with fine scribbly lines . . . . Leiognathus stercorarius
10a. Cheeks fully scaly (but may be dislodged, in which case scale pockets visible under
magnification) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
10b. Cheeks not fully scaly (a few thin scales sometimes present in Leiognathus blochii) . . . . . . ® 12
11a. Lateral line continuing to base of caudal fin, with 51 to 55 tubed scales; body depth 1.9
to 2.1 times in standard length; outer half of spinous part of dorsal fin black . . Leiognathus rapsoni
11b. Lateral line terminating below last soft rays of dorsal fin, with around 38 tubed scales;
body depth 2 to 3 times in standard length; no black tip to spinous part of dorsal fin . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus moretoniensis
12a. No scales visible on breast, its appearance distinctly different from rest of sides, which
are scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
12b. Breast distinctly and usually totally covered in scales, its appearance same as rest of
sides, which are scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
13a. Second dorsal-fin spine greatly elongate, its length at least 0.7 times in body depth and
as long as twice body depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13b. Second dorsal-fin spine not greatly elongate, its length, at most, approximately 1/2 body
depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
14a. Second anal-fin spine elongate, its length at least 0.6 times in body depth and often equal
to body depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus longispinis
14b. Second anal-fin spine not markedly elongate, its length 0.3 to 0.5 times in body depth . . . . . ® 15
15a. Body deep, its depth equal to greater than 1/2 standard length, relatively deeper in larger
fish; pelvic fins long, clearly reaching more than half-way to origin of anal fin; dark vertical
bars on upper sides; in fresh specimens, a row of yellow blotches along midline of sides
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus fasciatus
15b. Body slender, its depth 0.3 to 0.45 times in standard length; pelvic fins short, not reaching
half-way to origin of anal fin; tightly packed, dark vermiculating lines on upper sides,
down almost to midline; no row of yellow spots along midline of sides . . . . . . . Leiognathus sp. 2
16a. Dark blotch astride nape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17

16b. No dark blotch astride nape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
17a. Second dorsal-fin spine elongate slightly, its length 1/2 depth of body or even longer, and
about 1/2 as long again as length of pectoral fins; upper half of body with mainly vertical,
dark brown, wavy lines, some broken into shorter segments and dots; a yellow brown
line across spinous parts of both dorsal and anal fins at a height corresponding to
margins of soft parts of these fins; yellow blotch about size of eye half-way between base
of pectoral fins and first anal-fin spine (said to be absent in some Australian populations)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus decorus
17b. Second dorsal-fin spine not elongate, its length much less than 1/2 depth of body and
less than or only fractionally longer than length of pectoral fins; upper half of body without
lines or other markings; tip of spinous part of dorsal fin black surrounded by yellow . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus nuchalis
(Japan and Taiwan Province of China; not yet recorded from the area)
18a. Body depth 2 to 2.2 times in standard length; dorsal and ventral profiles equally convex;
mandible nearly straight; above lateral line, sides with very faint vertical stripes, but these
disappearing with age; distal part or peak of spinous part of dorsal fin black . . . Leiognathus daura
18b. Body depth 1.6 to 1.8 times in standard length; dorsal profile much more convex than
ventral profile; mandible strongly concave; many fine vertical stripes extending from
dorsal margin to below lateral line; spinous part of dorsal fin hyaline, no black tip . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus equulus
19a. Mouthparts project forward rather than downward when protracted; teeth arranged in a
single row in each jaw (Fig. 5a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
19b. Mouthparts project downward when protracted; teeth arranged in villiform bands, though
these may contract to a single row laterally in young specimens (Fig. 5b) . . . . . . . . . . . . ® 21
a) b)
Fig. 5 lateral view of jaws
2796 Bony Fishes
20a. Body circular to ovate, its depth 1.7 to 2.1 times in standard length; many irregularly
shaped, thin, scribbly lines on upper sides; distal half (i.e. peak) of spinous part of dorsal
fin bright orange with a black line along its lower edge in life, the whole becoming dusky
in preserved specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus bindus
20b. Body elongate, its depth 2.3 to 2.9 times in standard length; upper half of sides with a
number of irregularly shaped blotches anteriorly, some enclosing pale centres, and with
broad bars and smaller patches posteriorly; spinous part of dorsal fin, clear yellow in life,
plain hyaline in preserved specimens . . . . . . . . . . . . . . . . . . . . . . . Leiognathus aureus
21a. Body very deep, its depth 1.7 to 2 times in standard length; body thick, not strongly
compressed laterally; lower margin of eye situated above upper edge of retracted upper
jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus splendens
21b. Body not very deep, its depth 2 to 3 times in standard length; body not thick, strongly
compressed laterally; lower margin of eye lower than upper edge of retracted upper jaw . . . . ® 22
22a. No dark blotch on nape; distal part of spinous dorsal fin without any dark markings . . . . . . . ® 23
22b. Dark blotch astride nape; distal part of spinous dorsal fin tipped with black . . . . . . . . . . . ® 26
23a. Second dorsal-fin spine greatly elongate, its length almost equal to greatest body depth,
often longer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus leuciscus
23b. Second dorsal-fin spine about 1/2 greatest body depth . . . . . . . . . . . . . . . . . . . . . . ® 24
24a. Depth of body 2 to 2.3 times in standard length; upper sides with vertical wavy lines
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus dussumieri
24b. Depth of body 2.3 to 3 times in standard length; upper sides with short grey contorted
lines, some more or less hook- or U-shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 25
25a. Upper half of sides with many densely packed, short grey contorted lines, serially
arranged and angling forward below lateral line . . . . . . . . . . . . . . . . . . Leiognathus berbis
25b. Sides above lateral line with several, sparsely distributed, short grey contorted lines;
below lateral line, a row of blotches . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus sp. 1
26a. Upper half of sides with 4 longitudinal rows of dark broken lines, the lowermost below
lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leiognathus pan
26b. Upper sides with a row of dark, wavy, vertical lines extending downward from base of
dorsal fin past lateral line; spinous dorsal and anal fin with yellow streak at midheight;
lateral line not covered with yellow scales . . . . . . . . . . . . . . . . . . . . . Leiognathus blochii
List of species occurring in the area

The symbol 0 is given when species accounts are included. A question mark indicates that presence in
the area is uncertain.
0
0
Gazza achlamys Jordan and Starks, 1917
Gazza minuta (Bloch, 1797)
0
0
Leiognathus aureus Abe and Haneda, 1972
Leiognathus berbis (Valenciennes, 1835)
0 Leiognathus bindus (Valenciennes, 1835)
0
0
Leiognathus blochii (Valenciennes, 1835)
Leiognathus daura (Cuvier, 1829)
0
0
Leiognathus decorus (de Vis, 1884)
Leiognathus dussumieri (Valenciennes, 1835)
0
Leiognathus edwardsi Evermann and Seale 1907
0
Leiognathus elongatus (Günther, 1860)
Leiognathus equulus (Forsskål, 1775)
0 Leiognathus fasciatus (Lacepède, 1803)
0
Leiognathus hataii Abe and Haneda, 1972
Leiognathus leuciscus (Günther, 1874)
0
0
Leiognathus longispinis (Valenciennes, 1835)
Leiognathus moretoniensis Ogilby, 1912
Leiognathus nuchalis (Temminck and Schlegel, 1845)
0
?
0
Leiognathus pan Wongratana, 1988
0
Leiognathus rapsoni Munro, 1964
Leiognathus splendens (Cuvier, 1829)
0
0
Leiognathus stercorarius Evermann and Seale, 1907
0
Leiognathus sp. 1
Leiognathus sp. 2
0
0
Secutor hanedai Mochizuki and Hayashi, 1989
0
Secutor indicius Monkolprasit, 1973
0
Secutor insidiator (Bloch, 1787)
0
Secutor megalolepis Mochizuki and Hayashi, 1989
Secutor ruconius (Hamilton-Buchanan, 1822)
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.
Jones, G. 1985. Revision of the Australian species of the fish family Leiognathidae. Aust. J. Mar. Freshwater Res.,
36:559-613.
2798 Bony Fishes
Gazza achlamys Jordan and Starks, 1917 (Plate I, 1)

FAO names: En - Naked toothpony; Fr - Sapsap nu; Sp - Motambo liso.
(after Jones, 1985)
Diagnostic characters: Body oval, moderately deep and somewhat compressed, its depth 1.9 to 2.2 times
in standard length. Mouth pointing forward when protracted, with distinct caninform teeth in both jaws.
Head scaleless; scales on body absent anterior to a line from origin of soft dorsal fin to behind
pectoral-fin base and thence on towards tip of depressed pelvic fins. Colour: silvery, back greyish to
bluish, with dark yellow, irregular marks extending to a little below lateral line; black dots all over ventral half
of body and head; snout tip grey; membrane of spinous dorsal fin black at its distal portion; edge of soft
part of dorsal fin grey; anal fin yellow anteriorly from second spine to third soft ray, tips of anal-fin rays dark
dusky; caudal fin dusky at posterior margin; pectoral and pelvic fins colourless.
Size: Maximum total length 17 cm, commonly to 12 cm.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over silty bottoms to depths of at least
20 m. Feeds mainly on small fishes, crustaceans, and polychaetes. Schooling. Caught mainly with bottom
trawls. Larger specimens may be marketed fresh or dried-salted but catches mostly converted to fishmeal
or dumped at sea.
Distribution: I n t h e I n d o -
Malayan region reported only
from peripheral localities such
as Sumatra, Jawa, Flores,
Sabah, Philippines, and
eastern half of New Guinea to
the Solomon Islands. In the
North Pacific reported from the
s o u t h e r n M a r i an as an d
Pohnpei. Records from the
east coast of Australia to as far
south as Moreton Bay need
confirmation. Outside the area,
reported from Taiwan Province
of China, Sri Lanka, and
southern India.
Gazza minuta (Bloch, 1797) (Plate I, 2)

FAO names: En - Toothpony; Fr - Sapsap dentu; Sp - Motambo dentudo.
Diagnostic characters: Body oval and somewhat compressed, its depth 1.9 to 2.7 times in standard
length; dorsal and ventral profiles equally convex, moderately deep. Mouth pointing forward when
protracted, with distinct caniniform teeth in both jaws. Gill rakers approximately equal in length to
corresponding gill lamellae, 17 to 20 on first gill arch. Head scaleless, but scales covering all of body
except for breast ahead of a line from base of pectoral fins to origin of anal fin. Tubed scales on lateral
line 45 to 51. Colour: silvery; back greyish, with dark yellow irregular marks extending to below lateral line.
Spinous dorsal-fin membrane black-edged; soft part of dorsal fin and anal fin edged with grey; anterior
part of anal fin yellow; caudal fin yellowish, its hind margin dusky; pectoral and pelvic fins colourless;
underside of pectoral-fin base with black dots.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over silty bottoms to depths of 75 m,
juveniles in shallower water around mangroves to 10 m; enters estuaries. Feeds on small fishes, prawns,
polychaetes, and crustacean larvae; juveniles feed on plankton and insects as well. Schooling. Caught
mainly with bottom trawls, but also in shore seines. Larger fish may be marketed fresh or dried-salted; but
mostly sold for consumption by domestic animals (e.g. ducks) or dumped at sea.
Distribution: Reported from
the east coast of Africa and the
Red Sea throughout the tropi-
cal coasts of the Indian Ocean
to the Indo-Malayan Archipel-
ago and beyond; in the Pacific
found in parts of Micronesia,
northward to Taiwan Province
of China and Okinawa, south-
ward to northern Australia,
eastward, at least as far as
Vanuatu, with unconfirmed re-
cords for Cook Islands and Ta-
hiti. (This distribution may
include a closely related, un-
described species.)
2800 Bony Fishes
Leiognathus aureus Abe and Haneda, 1972 (Plate I, 3)

FAO names: En - Golden ponyfish.
(after Jones, 1985)
Diagnostic characters: Body only moderately deep and compressed, its depth 2.3 to 2.9 times in
standard length; ventral profile much more convex than dorsal profile, especially due to deep head
relative to body depth (resembling Gazza spp.). Mouth pointing forward when protracted; single row
of small uniform teeth in each jaw, no canines. Gill rakers slightly more than 1/2 length of corresponding
gill lamellae, 19 to 22 in total on first gill arch. Spines in dorsal fin very slender, second and subsequent ones
all comparatively short, length of second dorsal-fin spine about 1.5 times diameter of eye, 0.75 times
in length of pectoral fins; anal-fin spines slender and short, length of second equal to eye diameter. Head
scaleless; breast fully scaly. Colour: silvery upper half with irregular grey-brown blotches and
marbling; in fresh specimens large gold luminous organ around esophagus; outer half or peak of spinous
part of dorsal fin pale yellow, margin of spinous part edged in black; other fins colourless.
Size: Maximum total length at least 10 cm, commonly to 6 cm.
Habitat, biology, and fisheries: Lives in deeper offshore coastal waters where taken by trawling at depths
greater than 70 m and down to 140 m. Probably treated as bycatch by trawlers.
Distribution: Indonesian and
Australian waters in Arafura
Sea area.
Leiognathus berbis (Valenciennes, 1835) (Plate I, 4)

Frequent synonyms / misidentifications: None / Leiognathus oblongus (Valenciennes, 1835).
FAO names: En - Berber ponyfish; Fr - Sapsap berbère; Sp - Motambo bérbero.
Diagnostic characters: Body moderately elongate and compressed, its depth 2.3 to 2.8 times in
standard length; dorsal and ventral profiles about equally convex, but a more or less distinct notch present
at nape. Mouth pointing downward when protracted. Dorsal-fin spines slender, the second equal to or, in
adults, slightly longer than 1/2 height of body; second anal-fin spine slightly less than 1/2 length of second
dorsal-fin spine. Head scaleless, but scales present on breast. Colour: belly silvery; back greenish to
brownish with light grey, crowded, irregular vermiculations extending on sides to slightly below
lateral line, where lines become serially arranged and angle forward; snout and underside of pectoral-fin
base dotted black; dorsal, anal, pectoral, and pelvic fins colourless, except for margins of dorsal and anal
fins, which are yellow; caudal fin dusky yellow.
Habitat, biology, and fisheries: Inhabits coastal inshore waters, where taken by bottom trawlers. Appar-
ently not a dominant species in catches in any area.
Distribution: In the Indian
Ocean from Madagascar to
the Red Sea and the Gulf of
Aden, along the Indian coasts
and off Sri Lanka, eastward to
Malaysia, Indonesia, and the
Philippines; north to Taiwan
and F ukien provinces of
China.
2802 Bony Fishes
Leiognathus bindus (Valenciennes, 1835) (Plate I, 5)

Frequent synonyms / misidentifications: Leiognathus virgatus Fowler, 1904 / None.
FAO names: En - Orangefin ponyfish; Fr - Sapsap voile orange; Sp - Motambo naranjón.
Diagnostic characters: Body deep and strongly compressed, its depth 1.8 to 1.9 times in standard
length; ventral profile at least as equally convex as dorsal profile, in larger specimens even more
strongly convex. Mouth pointing forward to slightly downward when protracted; single row of
uniform conical teeth in each jaw. Gill rakers long and slender, approximately equal in length to
corresponding gill lamellae, 19 to 23 on first gill arch. Head scaleless, but conspicuous scales present on
breast. Tubed scales on lateral line 45 to 49. Colour: body silvery, with short dark vermiculations on back;
tip of snout, head, and ventral half of body with grey dots which tend to be arranged in rows midlaterally;
spinous part of dorsal fin black at 1/2 height, above which the membrane between second and fifth
spines bears a bright orange blotch which turns yellow on preservation in formalin; soft dorsal-fin
margin grey; faint yellow on distal part of membrane of spinous anal fin; pectoral, pelvic, and caudal fins
colourless; underside of pectoral-fin base black.
Habitat, biology, and fisheries: Inhabits coastal inshore waters on bottoms of muddy sand at depths of 10
to 160 m. Feeds on calanoid copepods, ostracods, chaetognaths, and the larvae of annelids and fishes.
Schooling; abundant during spawning and recruitment seasons which are, respectively, September to
December and March to April in northern Australia. Caught mainly in bottom trawls along with Leiognathus
splendens and L. decorus. May be marketed fresh or dried-salted but in Southeast Asia more often used as
food for ducks or converted to
fishmeal; dumped as bycatch
in Australia.
Distribution: Reported to oc-
cur in the Red Sea (Port Su-
dan), Gulf of Arabia, coasts of
India and Sri Lanka, and else-
where in the eastern Indian
Ocean; eastward to the west-
ern Central Pacific.
Leiognathus blochii (Valenciennes, 1835) (Plate I, 6)

FAO names: En - Twoblotch ponyfish; Fr - Sapsap à deux taches; Sp - Motambo de dos manchas.
Diagnostic characters: Body oval, moderately elongate and compressed, its depth 2.3 to 3.2 times in
standard length; dorsal and ventral profiles almost equally convex. Snout pointed; mouth pointing down-
ward when protracted. Gill rakers slender, more than 1/2 length of corresponding gill lamellae, total gill rakers
on first gill arch 18 to 21. Second dorsal-fin spine less than 1/2 greatest body depth, fairly stout, long as head
without snout; second anal-fin spine also fairly stout and long, 0.8 to 0.9 times in second dorsal-fin spine. A
few weak scales on cheek below eye but these easily dislodged; conspicuous scales present on breast.
Lateral line runs through to base of caudal fin, tubed scales on lateral line 54 to 58. Colour: belly silvery;
back light brown with dark, irregular vertical lines extending down to midline; a brown blotch on
nape which becomes diffuse on preservation in formalin; tip of snout, head, and ventral half of body
with fine black dots; dorsal-fin membrane from about 1/2 its height to tips of second to fifth spines black; soft
part of dorsal and anal fins, as well as caudal fin, yellow with grey edges; pelvic and pectoral fins colourless,
underside of pectoral-fin base dotted black. In Australian populations the spinous part of the dorsal fin is
reputed to be traversed by a yellow streak at midheight, with the apex of the fin hyaline instead of black.
Habitat, biology, and fisheries: Inhabits coastal inshore waters on bottoms of muddy sand at depths of 10
to 90 m; enters estuaries. Feeds on plant detritus, algal tissue, polychaetes, and insects. Schooling. May
be marketed dried-salted, but in Southeast Asia more often used as food for ducks or converted to fishmeal.
Distribution: Pakistan and
down the west coast of India
(to about 12°S), and also in the
eastern Indian Ocean; in the
area recorded from Indonesia,
Philippines, Thailand, and er-
ratically distributed across
n o r t h e r n A u s t r a l i a f r om
Broome to Cape York. (Austra-
lian populations may represent
a distinct species.)
2804 Bony Fishes
Leiognathus daura (Cuvier, 1829) (Plate I, 7)

Frequent synonyms / misidentifications: Leiognathus gerroides (Bleeker, 1851) / None.
FAO names: En - Goldstripe ponyfish; Fr - Sapsap doré; Sp - Motambo dorado.
Diagnostic characters: Body rhomboid and compressed, its depth 2 to 2.5 times in standard length;
dorsal and ventral profiles about equally convex. Snout somewhat blunt; mouth pointing downward when
protracted. Gill rakers a little more than 1/2 length of corresponding gill lamellae, total gill rakers 18 to 22 on
first gill arch. Head and breast scaleless. Lateral line much less convex than dorsal profile, and terminating
below end of dorsal fin; tubed scales on lateral 59 to 65. Colour: back grey greenish, belly silvery; very faint
vertical lines above lateral line which disappear completely with increasing age; black dots all over
ventral half of body; a golden hue on dorsal half of body and on head; tip of snout black; distal half of fin
membrane between second and sixth dorsal-fin spines jet-black, lined above by creamy white; a
broad yellow band over the lateral line, which disappears gradually on preservation; outer half of anal
fin golden yellow; outer margin of lower caudal-fin lobe also yellow.
Habitat, biology, and fisheries: Inhabits very shallow coastal inshore waters on bottoms of muddy sand,
mostly around depths of 10 to 15 m. Schooling. Caught mainly in bottom trawls. May be marketed
dried-salted, but mostly treated as trash fish and either dumped or used as food for ducks or converted to
fishmeal.
Distribution: From the Gulf of
Aden along the coasts of India
and Sri Lanka to the western
Central Pacific.
Leiognathus decorus (de Vis, 1884) (Plate I, 8)

Frequent synonyms / misidentifications: None / Leiognathus brevirostris (Valenciennes, 1835); Equula
nuchalis Temminck and Schlegel, 1845.
FAO names: En - Yellowfinned ponyfish.
(after Randall, 1996)
Diagnostic characters: Body moderately deep and compressed, its depth 1.8 to 2.3 times in standard
length; dorsal profile more convex than ventral profile. Mouth pointing downward when protracted. Gill
rakers slightly less than 1/2 length of corresponding gill lamellae, total gill rakers on first gill arch 21 to 24.
Dorsal- and anal-fin spines strong, only slightly elongate, second dorsal-fin spine between 1/3 and
1/2 body depth. Head and breast scaleless. Colour: silvery, upper half with irregular dark brown wavy
to zig-zag vertical lines; brown blotch astride nape (fading in preserved specimens); axil of pectoral
fins with a yellow spot and another yellow spot about size of eye on abdomen half-way between bases of
pectoral fins and spinous anal fin (both spots absent in Australian specimens); outer half of spinous dorsal
and anal fins yellow, yellow margins to soft parts of both fins, caudal fin with dusky yellow margin, other fins
colourless.
Habitat, biology, and fisheries: Inhabits very shallow (to about 30 m), coastal inshore waters on silty-sand
bottoms near mangroves; enters large estuaries; juveniles in mangrove-lined creeks. Feeds on polychaetes,
small crustaceans (mysids, copepods, amphipods, ostracods, and cladocerans), large crustaceans
(macruran larvae), euphausids, molluscs, and detritus. Schooling; very abundant during spawning season
(September to December in northern Australia). Caught mainly in bottom trawls, but also by beach seines,
push nets, bamboo stake
traps, and dip nets. In South-
east Asia this species may be
m ar k ete d fresh or dried-
salted, but it is more often
used as food for ducks or con-
verted to fishmeal, while dis-
carded in Australia.
Distribution: So far recorded
only from Oman, Thailand, In-
donesia, New Guinea, and
northern Australia down to
25°S.
2806 Bony Fishes
Leiognathus dussumieri (Valenciennes, 1835) (Plate II, 9)

Frequent synonyms / misidentifications: None / Leiognathus equulus (Forsskål, 1775).
FAO names: En - Dussumier’s ponyfish; Fr - Sapsap vaguelette; Sp - Motambo de onda.
Diagnostic characters: Body moderately slender and compressed, its depth 2 to 2.3 times in standard
length; anterior part of dorsal profile more strongly arched than anterior part of ventral profile. Snout
pointed, slightly longer than eye diameter; mouth pointing downward when protracted. Pelvic fins
comparatively long, reaching first anal-fin spine. Head scaleless, but conspicuous scales present on
breast. About 60 tubed scales on lateral line. Colour: back greenish to brownish, belly silvery, fins yellowish
(often blue-green due to settlement of algae, and this colour becomes more prominent on preservation in
formalin); sides of body with grey, wavy, vertical lines descending from back to a little below lateral
line; an elongate yellow spot on abdomen half-way between bases of pectoral fins and spinous anal
fin.
Habitat, biology, and fisheries: Inhabits coastal waters demersally between depths of around 10 to 40 m.
Reported to favour bottoms of coral sand, but also to enter estuaries. Feeds on small crustaceans,
polychaetes, foraminiferans, bivalves, gastropods, and nematodes. Schooling. It is the dominant species in
catches from certain parts of southern India, but separate statistics not available for it. Caught mainly with
bottom trawls. As one of the larger species, large individuals probably marketed fresh, with surplus fish being
converted to fishmeal, used to feed ducks, or discarded.
Distribution: Recorded from
Madagascar, Réunion, the
coasts of India and Sri Lanka,
through Indonesia and the
Philippines. Possibly more
widely distributed but has been
confused with Leiognathus
equulus by some authors.
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Leiognathus elongatus (Günther, 1874) (Plate II, 10)

FAO names: En - Elongate ponyfish.
Diagnostic characters: Body moderately compressed, elongate, very slender, not deeper than head
length, its depth more than 3.3 times in standard length; ventral profile more convex than dorsal
profile. Mouth pointing downward when protracted. Supraorbital spine in front of eye narrow, terminating in
a single point. Cheek and breast covered with small scales, but cheek scales very easily lost, leaving no
trace of scale pockets. Tubed scales in lateral line 42 or 43. Colour: body silvery, back with irregular green
and dark marbling; horizontal yellow band at midheight of spinous part of dorsal fin, most of margin of
soft part orange; underside of pectoral-fin base with minute, dark dots; anal fin between second and
third spines yellow, as also margin of anterior part of fin; males with bluish longitudinal stripe along side of
belly.
Habitat, biology, and fisheries: Inhabits coastal inshore waters at depths of 30 to 130 m. Schools near the
bottom; sparsely distributed. Feeds on small crustaceans, polychaetes, and algae. Caught mainly with
bottom trawls. Marketed mostly dried-salted, but usually treated as a trash fish, being converted to fishmeal,
used to feed ducks, or discarded.
Distribution: From the Red
Sea along the east coast of
Africa to about 10°S and off
southwest India. Outside this
area, it occurs in the Andaman
Sea region, Indonesia, Thai-
land, the Philippines, and
South China, extending north-
ward to southern Japan and
south to northern Australia
and possible the Santa Cruz
Islands.
2808 Bony Fishes
Leiognathus equulus (Forsskål, 1775) (Plate II, 11)

FAO names: En - Common ponyfish; Fr - Sapsap commun; Sp - Motambo común.
Diagnostic characters: Body very deep, compressed, with a strongly humped back; body depth 1.7
to 1.9 times in standard length. Mouth pointing downward when protracted. Gill rakers short and fleshy,
less than 1/2 length of corresponding gill lamellae, total gill rakers on first gill arch 18 to 22. Head and breast
scaleless. Tubed scales on lateral line 61 to 66. Colour: in adults, back greyish, belly silvery and many
parallel close-set faint bars on back; usually a dark brown saddle on caudal peduncle; axil of pectoral fins
grey to black; margin of soft dorsal fin black; both caudal-fin lobes with broad dusky margins; pectoral, pelvic,
and anal fins colourless to yellowish. In juveniles (5 to 7 cm total length), thin, closely arranged, grey
vertical lines descending from back to about midheight; membrane between anal-fin spines conspicu-
ously yellow; posterior margin of caudal-fin lobes pale yellow and dusky; other fins hyaline; snout dotted
black.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over muddy bottoms to depths of 70 m;
juveniles around mangrove areas at depths of 1 to 10 m; enters estuaries. Feeds chiefly on polychaetes,
small crustaceans, and small fishes. Schooling. Caught near the bottom with bottom trawls, set nets, push
nets, dip nets, and beach seines. In Indian waters, schools near the surface during April-May when caught
with drift gill nets. Larger specimens sorted for human consumption and marketed fresh or dried-salted, but
surplus fish used to feed ducks, converted to fishmeal, or discarded.
Distribution: Widely distrib-
uted in the Indian Ocean from
the east coast of Africa to the
Red Sea, including oceanic is-
lands such as Réunion and
Comoros; then along the
coasts of India and the Indo-
Malayan Archipelago to the
West Pacific, as far east as
Samoa, as far north as the Ryu
Kyu Islands, and south to
southern Queensland.
Leiognathus fasciatus (Lacepède, 1803) (Plate II, 12)

FAO names: En - Striped ponyfish; Fr - Sapsap rayé; Sp - Motambo rayado.
Diagnostic characters: Body deep and compressed, the back more strongly arched than anterior
part of belly; body depth 1.7 to 2 times in standard length. Mouth pointing downward when protracted.
Gill rakers short and fleshy, less than 1/2 length of corresponding gill lamellae, total gill rakers on first gill
arch 18 to 22. Second dorsal-fin spine distinctly elongate. Head and breast scaleless. Tubed scales on
lateral line 60 to 66. Colour: body silvery, back less so than belly; about 11 widely spaced, dark vertical
lines on back superimposed by a few yellow blotches, lines fading gradually toward lower sides;
margin of soft dorsal fin grey; anal fin hyaline with a yellow margin; posterior margin of caudal-fin lobes black;
pectoral and pelvic fins hyaline; pectoral-fin axil yellow, base dotted black.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over muddy bottoms to depths of 120 m.
Feeds chiefly on polychaetes, small crustaceans, and small fishes. Schooling. Caught mainly with bottom
trawls, but also with various kinds of set nets and beach seines. Larger fish sorted for human con sumption
and marketed fresh or dried-salted; surplus fish used to feed ducks, converted to fishmeal, or discarded.
uted in the Indian Ocean from
the east coast of Africa and the
Red Sea, including oceanic is-
lands such as Mauritius and
the Seychelles, to the Indo-
Malayan Archipelago and the
West Pacific, as far east as
Samoa, north to Okinawa, and
south to northern Australia to
around 20°S.
2810 Bony Fishes
Leiognathus leuciscus (Günther, 1860) (Plate II, 13)

Frequent synonyms / misidentifications: Equulites novaehollandiae Munro, 1967 / None.
FAO names: En - Whipfin ponyfish; Fr - Sapsap fil; Sp - Motambo de hebra.
(after Jones, 1985)
Diagnostic characters: Body only moderately deep, compressed, its depth 2 to 3 times in standard
length; dorsal and ventral profiles almost equally convex. Mouth pointing downward when protracted. Gill
rakers approximately 1/2 length of corresponding gill lamellae, total gill rakers on first gill arch 16 to 19.
Second dorsal-fin spine elongate, and second anal-fin spine slightly elongate. Head scaleless; scales
present on breast. Tubed scales on lateral line 48 to 51. Colour: belly silvery; back with irregular, partly
semicircular, greyish green vermiculations; a row of oval yellow blotches below lateral line in larger
fish; membrane between dorsal-fin spines yellow at midheight, edge of soft part of dorsal fin also yellow,
superimposed with light grey; anal fin of similar colour to dorsal fin; posterior margin of caudal fin yellowish;
males usually with a triangular bluish patch on side of belly.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over bottoms of muddy sand at depths of
10 to 50 m. Feeds mainly on polychaetes, small shrimps, and other crustaceans. Schooling. Caught mainly
with bottom trawls, but also in set nets and traps, push nets, and beach seines. Marketed fresh or dried-salted
for human consumption; but surplus fish used to feed ducks, converted to fishmeal, or discarded.
Distribution: In the Indian
Ocean reported from Mada-
gascar and the Seychelles
eastward along the coasts of
India and Sri Lanka, to Indo-
nesia, Thailand, Philippines,
New Guinea, and northern
Australia south to Shark Bay
on the Western Australia
coast.
Leiognathus longispinis (Valenciennes, 1835) (Plate II, 14)

Frequent synonyms / misidentifications: Leiognathus smithursti (Ramsay and Ogilby, 1886) / None.
FAO names: En - Smithhurst’s ponyfish; Fr - Sapsap panache; Sp - Motambo de penacho.
Diagnostic characters: Body oval, compressed and deep, its depth 1.7 to 2 times in standard length;
anterior part of dorsal profile more strongly arched than anterior part of ventral profile. Mouth pointing
downward when protracted. Gill rakers short and fleshy, less than 1/2 length of corresponding gill lamellae,
total gill rakers on first gill arch 18 to 21. Second dorsal- and anal-fin spines greatly elongate (sometimes
reaching to caudal fin). Head and breast scaleless. Tubed scales on lateral line 64 to 70. Colour: belly
more silvery than back which shows a few faint, unevenly spaced, and horizontally elongate blotches;
a few yellow blotches below lateral line; dorsal and anal fins with yellow colouring, particularly on margins;
caudal fin faint yellow with a more intense yellow blotch on lower lobe; underside of pectoral-fin base with
minute black dots.
Habitat, biology, and fisheries: Inhabits coastal inshore waters at depths of 15 to 80 m. Feeds chiefly on
small crustaceans and nematodes. Schooling. Caught mainly with bottom trawls. Larger fish sorted for
human consumption and marketed fresh or dried-salted; surplus fish used to feed ducks, converted to
fishmeal, or discarded.
Distribution: From southern
India to the Indo-Malayan re-
gion and northern Australia to
about 20°S, northward to Phil-
ippines; reputed to occur in
Fiji.
2812 Bony Fishes
Leiognathus moretoniensis Ogilby, 1912 (Plate III, 15)

Frequent synonyms / misidentifications: Equulites hastatus (Ogilby, 1912) / None.
FAO names: En - Zig-zag-ponyfish.
(after Jones, 1985)
Diagnostic characters: Body moderately elongate and compressed, its depth variable from elongate to
moderately deep, 2 to 3 times in standard length; dorsal and ventral profiles about equally convex,
but a more or less distinct notch where head meets nape. Mouth projecting downward when protracted.
Gill rakers approximately 1/2 length of corresponding gill lamellae, total gill rakers on first gill arch 15 to 17.
Fin spines weak; second dorsal-fin spine varying from not elongate in populations south of the area
to substantially elongate in the tropics, 1.8 to 3.7 times in body depth, those with the long spine
occurring in the northern part of its range (southern form may prove to be a different species). Cheeks
scaly below eye; breast scaly. Colour: silvery, upper half with grey brown markings, broken into short
irregular lines immediately below base of dorsal fin, changing to broken chevrons or zig-zag lines
across lateral line; row of small opalescent spots extending back from pectoral fins, and sometimes an
additional row of yellow spots above this; dorsal fin sometimes with a yellow streak anteriorly at midheight
and continuing along margin of soft rays; soft-rayed portion of anal fin with narrow yellow margin; other fins
colourless.
Habitat, biology, and fisheries: Widely distributed across continental shelf waters of northern Australia,
living near the bottom in schools. Caught by bottom trawls, but discarded as bycatch.
Distribution: Across northern
Australia, from Shark Bay on
the Western Australia coast
out across the northwestern
shelf into the Arafura Sea,
down the east coast as far
south as Sydney.
Leiognathus pan Wongratana, 1988 (Plate III, 16)

FAO names: En - Shining ponyfish.
(after Wongratana, 1988)
Diagnostic characters: Body moderately elongate and compressed, its depth 2.3 to 2.9 times in
standard length; dorsal and ventral profiles about equally convex, but slightly concave where nape joins top
of head. Mouth pointing downward when protracted. Longest gill rakers 1/2 or little more than 1/2 length
of corresponding gill lamellae, total gill rakers on first gill arch 20 to 22. Dorsal-fin spines slender, second
not markedly elongate, noticeably shorter than 1/2 body depth; second anal-fin spine about 0.9 times in
length of second dorsal-fin spine. Head scaleless; breast scaly. Tubed scales on lateral line 48 to 54.
Colour: belly silvery; back iridescent blue-green, laterally washed with very pale orange; 4 longitudinal
rows of greatly elongate, greenish blue spots along upper side; dark blotch astride nape; spinous
part of dorsal fin with large black blotch on upper half from second to sixth spines; caudal fin washed with
pale orange on trailing edge of fin; otherwise fins hyaline.
Size: Maximum recorded total length 8 cm, commonly to 7 cm.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over bottoms of muddy sand at depths of
10 to 50 m. Caught mainly by bottom trawls. May be marketed dried-salted, but mostly treated as trash fish
and used to feed ducks, converted to fishmeal, or discarded.
Distribution: Thailand, both
Andaman Sea and Gulf of
Thailand coasts (presumably
then also on coasts of the Ma-
lay Peninsula, and perhaps
further afield).
2814 Bony Fishes
Leiognathus rapsoni Munro, 1964 (Plate III, 17)

FAO names: En - Papuan ponyfish.
(after Munro, 1964)
Diagnostic characters: Body oval and compressed, slightly more convex above than below, its depth 1.9
to 2.1 times in standard length; head profile slightly concave between eye and nape. Chin slightly concave.
Mouth pointing downward when protracted. A single row of fine teeth in both jaws. Total gill rakers on
first gill arch 24 to 28. Second spines of dorsal and anal fins longer than the others, 2.1 to 2.4 and 2.7 to 2.9
times, respectively, in body depth. Cheeks scaly below eye, breast scaly. Lateral line continued to base of
the caudal fin; tubed scales on lateral line 51 to 55. Colour: generally silvery, but upper half darker with
about 20 narrow, wavy, dark vertical bars; dark areas present on snout, operculum, and along middle of
sides; a dark line along base of dorsal fin; outer half of spinous dorsal fin black, other fins colourless.
Size: Maximum recorded total length 11 cm, commonly to 10 cm.
Habitat, biology, and fisheries: Has been trawled in a depth of about 10 to 15 m around river mouths in
the Gulf of Papua. No commercial fishery.
Distribution: K n o w n o n l y
from Papua New Guinea.
Leiognathus splendens (Cuvier, 1829) (Plate III, 18)

FAO names: En - Splendid ponyfish; Fr - Sapsap soleil; Sp - Motambo esplendor.
Diagnostic characters: Body very deep and compressed, its depth 1.8 to 2.4 times in standard length;
anterior part of dorsal profile more strongly arched than anterior part of ventral profile, with prominent
“brow” jutting out over eyes. Snout short (shorter than eye diameter) and blunt; mouth pointing slightly
downward when protracted. Gill rakers approximately equal in length to corresponding gill lamellae, total gill
rakers on first gill arch 23 to 28. Second dorsal and anal fins especially stout but not elongate. Head scaleless,
but prominent scales present on breast. Tubed scales on lateral line 49 to 56. Colour: belly silvery; back
greyish silvery, with faint, grey, wavy, vertical lines above lateral line in adults (in juveniles, a few
blotches instead of wavy lines). Scales of lateral line, bases of rays of pectoral fins, margins of dorsal
and anal fins bright yellow, as is whole membrane of anal fin from second spine to first soft ray and the
tip of the lower lobe of the caudal fin; usually a dark dusky area on outer third of anterior part of spinous
portion of dorsal fin.
Size: Maximum total length about 15 cm, commonly to 10 cm.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over bottoms of muddy sand to depths of
65 m; juveniles found in mangrove-lined creeks to depths of about 10 m. Feeds on plant detritus, copepods,
cyprids, and small molluscs. Schooling. In tropical Australia known to spawn in October but recruits present
in most months of the year. Caught mainly by bottom trawls (commonly, along with Leiognathus bindus and
L. decorus), but also in set nets, bamboo-stake traps, beach seines, and push and dip nets. Larger fish
sorted for human consumption and marketed fresh or dried-salted; surplus fish used to feed ducks, converted
to fishmeal, or dumped at sea.
Mauritius, Madagascar, Red
Sea, and the coasts of India to
the islands of the western
Central Pacific as far east as
Fiji, north to Okinawa, south to
northern parts of Australia to
2 0 ° S . P r o ba bl y o c c ur s
throughout Micronesia.
2816 Bony Fishes
Leiognathus stercorarius Evermann and Seale, 1907 (Plate III, 19)

Frequent synonyms / misidentifications: None / Leiognathus elongatus Smith and Pope, 1906.
FAO names: En - Slender ponyfish.
Diagnostic characters: Body elongate and moderately compressed, its depth 2.8 to 3.6 times in
standard length; dorsal and ventral profiles smoothly convex, slender but deeper than head length.
Mouth pointing downwards when protracted. Supraorbital spine in front of eye broad, terminating in 2 points.
Cheeks scaly below eye (scales deciduous but their pockets visible); breast scaly. Tubed scales in lateral
line 55 to 57. Colour: body silvery; back with irregular green and dark marbling; front of dorsal fin with
a horizontal yellow band, most of margin of hind part orange; axil of pectoral fin with minute dark dots or
dusky (covered by fin); anal fin between second and third spines yellow, as also margin of anterior part of
fin; males with a bluish longitudinal stripe along sides just below midline.
Size: Maximum total length 12 cm, commonly between 6 and 10 cm.
Habitat, biology, and fisheries: Inhabits coastal inshore waters over muddy bottoms especially around
mangrove areas to depths of 50 m; juveniles at depths of 1 to 10 m. Feeds on polychaetes and a variety of
small benthic animals. Schooling. May be caught in bottom trawls or with a variety of set nets and traps,
beach seines, lift and push nets. A small species that may be marketed dried-salted for human consumption,
or treated as trash fish and used to feed ducks, or converted to fishmeal.
Distribution: Southeast Asia
and Marianas: Philippines, In-
donesia, Malaysia, Thailand
(including Andaman Sea
coast), Yap, Guam, and Sai-
pan.
Leiognathus sp. 1 (Plate III, 20)

Frequent synonyms / misidentifications: None / ? Leiognathus lineolatus (Valenciennes, 1835).
FAO names: En - Ornate ponyfish; Fr - Sapsap écriture; Sp - Motambo ornado.
Diagnostic characters: Body moderately elongate and compressed, its depth 2.4 to 3 times in standard
length; dorsal and ventral profiles about equally convex; nape without a distinct notch. Mouth pointing
downward when protracted. Gill rakers approximately 1/2 length of corresponding gill lamellae, total gill
rakers on first gill arch 17 or 18. Dorsal-fin spines slender, the second about 1/2 body depth; second anal-fin
spine slender, about same length as pectoral fins. Head scaleless; anterior part of breast below head
scaleless but conspicuous scales present elsewhere on breast. Tubed scales on lateral line 51 to 54.
Colour: belly silvery, back greenish to brownish with relatively sparse, irregular, grey vermiculations
extending down to a little below lateral line, the lowest marks in the form of blotches in a row; snout
and underside of pectoral-fin base dotted black; dorsal, pectoral, and pelvic fins colourless except for some
yellow on spinous parts and along margins of dorsal and anal fins; margin of caudal fin yellow.
Habitat, biology, and fisheries: Inhabits inshore coastal waters over muddy bottoms, especially around
mangroves; juveniles in shallows, 1 to 10 m, adults in deeper water to at least 40 m. Feeds on small
crustaceans, chaetognaths, nematodes, bivalves, and gastropods. Schooling. Caught mainly with bottom
trawls but also in beach seines. Larger fish sometimes sorted for human consumption and marketed fresh
or dried-salted; surplus fish
used to feed ducks, converted
to fishmeal, or discarded.
Distribution: Madagascar,
Red Sea, the Gulf of Arabia,
and the coasts of India and Sri
Lanka, eastward to Thailand,
Indonesia, and the Philip-
pines; no positive records for
New Guinea or Australia.
2818 Bony Fishes
Secutor hanedai Mochizuki and Hayashi, 1989 (Plate IV, 22)

Frequent synonyms / misidentifications: None / Secutor insidiator (Bloch, 1787).
FAO names: En - Haneda’s ponyfish.
Diagnostic characters: Body oval, deep and compressed, its depth 1.9 to 2.2 times in standard length.
Head strongly concave above eye. Mouth pointing upward when protracted. Body scales small, 33 to 42
rows between bases of pectoral and pelvic fins. Total gill rakers on first gill arch 22 to 26. Cheeks scaleless;
breast without scales ventrally, including isthmus. Lateral line terminating below end of dorsal fin, but if
scale-row count were continued to end of caudal peduncle total count would be 60 to 70 scale rows.
Colour: belly silvery, back greenish to brownish, with a series of about 9 columns of dark blotches,
spots, and horizontal bars extending to midline of sides; a black, curved band from lower margin of eye
to posterior angle of lower jaw; dorsal-fin membrane black at tip of leading spines; other fins reported as
hyaline but caudal fin, likely to be partly yellow.
Habitat, biology, and fisheries: Inhabits coastal waters above bottoms of muddy sand to depths of 40 m;
common near mangroves, enters estuaries, but also occurs up to at least 6 km offshore. Diet unknown, but
probably copepods and mysids. Schooling. Caught mainly in bottom trawls. May be marketed dried-salted,
but mostly treated as trash fish, and either discarded or used as food for domestic animals (e.g. ducks) or
as food in fish farms.
Distribution: Southern parts
of South China Sea and Straits
of Malacca; probably more
widely distributed in Indo-Ma-
layan Archipelago.
Secutor indicius Monkolprasit, 1973 (Plate IV, 23)

Frequent synonyms / misidentifications: None / Secutor insidiator (Bloch, 1787).
FAO names: En - Dots-and-dashes ponyfish.
(after Monkolprasit, 1973)
Diagnostic characters: Body oval, deep and very compressed, its depth 2.2 to 2.6 times in standard
length. Head strongly concave above eye. Mouth pointing upward when protracted. Gill rakers long and
slender, approximately equal in length to corresponding gill lamellae, total gill rakers on first gill arch 24 to
30. Body scales very small; cheeks scaleless; breast without scales ventrally, including isthmus. Lateral
line reported as terminating below middle to last third of dorsal fin; tubed scales on lateral line up
to this point 80 to 92, but if scale-row count were continued to end of caudal peduncle total count
would be 87 to 111 scale rows. Colour: belly silvery, back light blue, with 17 to 22 dark, vertical
markings on upper half, these terminating above lateral line and resuming below it, with row of dots
along lateral line corresponding to points where vertical markings cross it, vertical markings as
dashes or string of small dots; narrow black line from lower margin of eye to posterior angle of lower jaw;
dorsal-fin membrane between second and fifth spines black at upper third reducing in width posteriorly with
a narrow yellow band immediately below it; soft parts of dorsal and anal fins colourless; caudal fin pale yellow
with a dusky trailing edge; underside of pectoral-fin base black, membrane pale yellow.
Size: Maximum total length 10 cm, commonly to almost 10 cm.
Habitat, biology, and fisheries: Inhabits coastal waters over bottoms of muddy sand at depths of 20 to
70 m; also enters estuaries. Feeds mostly on copepods, mysids, and plant detritus. Schooling. Caught mainly
in bottom trawls. Marketed dried-salted or fresh; in Southeast Asia mostly treated as trash fish and used as
food for domestic animals, e.g. ducks, or as food in fish farms.
Distribution: Coasts border-
ing on the South China Sea:
Taiwan Province of China,
Gulf of Thailand, Philippines,
Indonesia, and northern coast
of Papua New Guinea. Re-
c o r d s b a s e d o n tr a w l er
catches landed at major fish-
ing ports and trawl surveys,
probably distributed through-
out whole of Indo-Malayan re-
gion.
2820 Bony Fishes
Secutor insidiator (Bloch, 1787) (Plate IV, 24)

Frequent synonyms / misidentifications: Leiognathus insidiator (Bloch, 1787) / None.
FAO names: En - Pugnose ponyfish; Fr - Sapsap boxeur; Sp - Motambo boxeador.
Diagnostic characters: Body oval, deep and compressed, its depth 1.6 to 2.5 times in standard length.
Head strongly concave above eye. Mouth pointing upward when protracted. Gill rakers long and slender,
approximately equal in length to corresponding gill lamellae, total gill rakers on first gill arch 24 to 29. Body
scales small, 22 to 28 rows between bases of pectoral and pelvic fins; cheeks scaleless; breast without
scales ventrally, including isthmus. Lateral line terminating below end of dorsal fin, but if scale-row
count were continued to end of caudal peduncle total count would be 84 to 107 scale rows. Colour:
belly silvery, back greenish to brownish, with a series of 11 to 15 columns of bars and spots extending
to a little below lateral line; a black curved band from lower margin of eye to posterior angle of lower jaw;
dorsal-fin membrane between second and sixth spines black at upper third; soft part of dorsal and anal fin
colourless; caudal fin partly yellow; underside of pectoral-fin base dotted black.
Size: Maximum total length 10.5 cm, commonly to 8 cm.
Habitat, biology, and fisheries: Inhabits inshore coastal waters over bottoms of muddy sand to depths of
about 50 m; enters estuaries. Feeds on copepods, mysids, and plant detritus. Schooling, Caught mainly in
bottom trawls. Marketed dried-salted or fresh; in Southeast Asia mostly treated as trash fish and used as
food for domestic animals, e.g. ducks, or as food in fish farms. In Australia discarded as bycatch.
Distribution: From the coasts
of East Africa and the Red Sea
eastward along the coasts of
India to the Indo-Malayan Ar-
c h i p e l a g o , s o u t h w a r d to
around 25°S in northern Aus-
tralia.
Secutor megalolepis Mochizuki and Hayashi, 1989 (Plate IV, 25)

Frequent synonyms / misidentifications: None / Secutor ruconius (Hamilton-Buchanan, 1822).
FAO names: En - Bigscaled ponyfish.
Diagnostic characters: Body oval, very deep and compressed, its depth 1.6 to 1.8 times in standard
length. Head strongly concave above eye. Mouth pointing upward when protracted. Total gill rakers on
first gill arch 21 to 25. Body scales comparatively much larger than in other species in the genus, 9 to
11 rows between bases of pectoral fins and pelvic fins; scales on cheek; breast fully scaly, including
isthmus. Lateral line terminating below last few rays of soft dorsal fin, but if scale-row count were
continued to end of caudal peduncle total count would be 43 to 49 scale rows. Colour: silvery, with
dark markings forming about 10 irregular “bars” on upper half sides, anteriorly the “bars” made up
of columns of short bars and spots; a prominent curved black band running from lower margin of eye to
posterior angle of lower jaw; dorsal-fin membrane black at tip of leading spines; other fins reported as hyaline
but caudal fin, likely to be partly yellow.
Habitat, biology, and fisheries: Inhabits inshore coastal waters over bottoms of muddy sand at depths of
less than 50 m; enters estuaries, inhabiting brackish to fresh waters. Feeds on copepods, mysids, and plant
detritus. Caught mainly in bottoms trawls, but also in stake traps, dip nets, and beach seines. In Southeast
Asia, marketed dried-salted or fresh; but mostly treated as trash fish and used as food for ducks or for farmed
fish, or converted to fishmeal. In Australia discarded as bycatch.
the coasts of Thailand, Malay-
sia, and Indonesia which bor-
der the South China Sea, but
also from northern Australia;
presumably present in inter-
mediate areas such as New
Guinea, eastern Indonesia,
and the Philippines.
2822 Bony Fishes
Secutor ruconius (Hamilton-Buchanan, 1822) (Plate IV, 26)

Frequent synonyms / misidentifications: Secutor interruptus (Valenciennes, 1835) / None.
FAO names: En - Deep pugnose ponyfish; Fr - Sapsap bouledogue; Sp - Motambo buldog.
Diagnostic characters: Body oval, compressed and very deep, its depth 1.4 to 1.7 times in standard
length. Head strongly concave above eye. Mouth pointing upward when protracted. Gill rakers long and
slender, approximately equal in length to corresponding gill lamellae, total gill rakers on first gill arch 18 to
28. Body scales comparatively large, 10 to 16 rows between bases of pectoral and pelvic fins; scales
on cheek; breast fully scaly, including isthmus. Lateral line terminating below about middle of soft
portion of dorsal fin; tubed scales on lateral line up to this point 28 to 32, but if scale-row count were
continued to end of caudal peduncle total count would be 54 to 60 scale rows. Colour: silvery, with
about 10 bluish vertical bars on back extending to a little below lateral line; a prominent curved black
band running from lower margin of eye to beyond posterior angle of lower jaw; membrane between second
and fifth dorsal-fin spines black on upper third portion; soft part of dorsal and anal fins colourless; caudal fin
partly yellow; underside of pectoral-fin base black.
Size: Maximum total length almost 10 cm, commonly to 6 cm.
Habitat, biology, and fisheries: Inhabits inshore coastal waters over bottoms of muddy sand to depths of
60 m; both young and adults enter estuaries. Feeds on copepods, mysids, and plant detritus. Found to spawn
in March in northern Australia, but spawning may be serial. Caught mainly in bottom trawls, but also in beach
seines. In Southeast Asia, marketed fresh or dried-salted, but mostly treated as trash fish and converted to
fishmeal or used as food for ducks or for farmed fish. In Australia, discarded as bycatch.
Distribution: From the coast
of East Africa and the oceanic
islands of Seychelles, Réun-
ion, and Mauritius, to the Red
Sea, coasts of India to Malay-
sia, Thailand, Indonesia, Phil-
ippines, northward to Taiwan
Province of China, and south-
ward to around 20°S in north-
ern Australia.
Leiognathus sp. 2 (Plate IV, 21)

En - Vermiculated ponyfish.
Maximum total length 15 cm. Demersal at depths of 15 to 170 m on the continental shelf of northern
Australia from Exmouth Gulf to Cairns; range must extend at least into adjacent continental shelf
area of New Guinea. Of minor importance to fisheries; discarded as trashfish by trawlers operating
in Australian waters.
(after Jones, 1985)
click for next page

2824 Bony Fishes
BRAMIDAE
Pomfrets
by P.R. Last and M. Moteki
D iagnostic characters: Small to large (to 70 cm), compressed, ovate to elongate fusiform perciform
fishes. Anterior profile of head arched (except in Taractes); body form usually changing greatly with
growth and varying considerably between genera. Eye moderately large, variable in position from near
dorsal surface (in Taractes and Pterycombus) to more centrally on head. Spines absent from opercular
and preopercular margins, usually present in young. Mouth of moderate size, terminal or superior,
extending to below eye; angle of jaw very oblique, between 45 and 70o to horizontal axis. Dentition
variously reduced; teeth small or minute, villiform, conical, or caniniform, usually present on palatines and
absent on vomer. Total gill rakers on first gill arch 7 to 24. Branchiostegal rays 6 to 8. Dorsal fin single,
its origin from head to just behind pectoral-fin base; dorsal and anal fins of adults moderately stiff,
scaly, and not fully depressible (except for Pteraclis and Pterycombus where fins scaleless, greatly
enlarged, and entirely recessible into deep grooves formed by rows of enlarged, sheath-like basal
scales), anterior rays of dorsal and anal fins often forming distinct lobes; dorsal-fin rays 31 to 57; anal-fin
rays 21 to 50. Caudal fin of adults strongly forked or with biconcave central margin, upper lobe
sometimes extended; principal caudal-fin rays 17. Pectoral fins falcate in adults, with 17 to 23 rays.
Pelvic fins thoracic or jugular, with I spine and 5 soft rays. Scales strongly adhesive, variable in size
over body, frequently larger below lateral line than above, sometimes enlarged on caudal peduncle; maxilla
scaly; snout, lower jaw, and opercular and preopercular margins naked; scales of young and some
adults bearing spiny keels; horizontal scale rows on body 34 to 95. Vertebrae 36 to 54. Colour: body mostly
uniformly black, bluish, brownish, or silvery; fins usually similar but sometimes darker or with yellowish
spots or bars.
single dorsal fin
extended
upper
lobe
oblique jaw
oblong scales
scaly maxilla
(after Mead, 1972)
Habitat, biology, and fisheries: Mostly epipelagic in the open ocean. Occurring in schools mostly
shallower than 200 m but Eumegistus is frequently found deeper (to at least 600 m). Seasonal migrations
between high and low latitudes are reported in temperate species (i.e. Brama japonica and B. brama).
Midlevel predators feeding on small midwater fishes, crustaceans, squids, and salps. Important prey for
some tunas and billfishes. The basis of longline fishery off Spain and Portugal yielding in excess of 10 000 t
annually. Large unexploited resources appear to exist in other areas, particularly in temperate parts of the
Pacific Ocean. Excellent food fishes which deserve the title of “chicken of the sea”.
Perciformes: Percoidei: Bramidae 2825

Caristiidae: body soft, covered with thin, deciduous scales sail-like
(rather than thick, adhesive scales); head bones thin and fin
fragile (rather than firm); pelvic fins folding into shallow
groove (no equivalent groove in bramids). Like Pteraclis
and Pterycombus, the dorsal fin is sail-like, but can be
recessed between skin flaps rather than into a groove of
enlarged scales.
Diretmidae: head with pronounced bony ridges (rather
than absent); eye extremely large, diameter exceeding
postorbital head length (rather than much less); anterior
rays of dorsal and anal fins never forming a lobe; pelvic
fins with I spine and 6 soft rays (rather than I spine and 5
soft rays).
belly groove
Lamprididae: pectoral-fin base almost horizontal (rather
than oblique or vertical); pectoral and pelvic fins similar in Caristiidae
size (pectoral fins much larger than pelvic fins in bramids);
fins reddish.
eye large
pectoral-fin base
bony ridges horizontal
Diretmidae Lamprididae
Stromateidae (Pampus only): mouth small (not reaching below eye); gill membranes attached to isthmus
(rather than separate); scales small, deciduous; no pelvic fins.
Carangidae (Parastromateus niger only): mouth small (not reaching below eye); small scute-like scales
on caudal peduncle; no pelvic fins.
scute-like scales
mouth
very
small mouth
small
gill membrane
attached to isthmus
no pelvic fins
no pelvic fins
Stromateidae (Pampus sp.) Carangidae (Parastromateus niger)

2826 Bony Fishes
Key to the species of Bramidae occurring in the area

Remarks on key characters: until recently, our knowledge of the taxonomy of Indo-Pacific members of the
family have been poor. Members of each of the genera are quite distinct from each other but some species
are harder to identify. Bramids undergo dramatic changes in morphology as they grow and it is often difficult
to find identifying features that will distinguish species at all stages of growth. External meristics can be
used but even these overlap. In some cases, vertebrae need to be counted to distinguish species - all
vertebrae including the urostyle and small postcranial centrum are counted.
1a. Dorsal and anal fins very long, scaleless, and can
be recessed into grooves formed from rows of sheath of
enlarged, sheath-like scales (Fig. 1) . . . . . . . scales
. . . . . . . . . . . . . . . . . . (Pteraclis, Pterycombus) ® 2 long fins
1b. Dorsal and anal fins scaly and not recessible in
adults . . (Brama, Eumegistus, Taractes, Taractichthys) ® 4
2a. Dorsal-fin origin over or behind hind margin of eye

(Fig. 2a); none of the first 5 dorsal-fin spines obvi-
ously thicker than the rest (Figs 2a and 3) . . . .
. . . . . . . . . . . . . . . . . . . . . . . Pterycombus petersii
2b. Dorsal-fin origin forward of eye (Fig. 2b); one of
the first 5 dorsal-fin spines obviously thicker than
the rest (Fig. 2b) . . . . . . . . . . . . . . . . (Pteraclis) ® 3
Fig. 1 Pterycombus
fin origin thickened

spine
fin
origin
a) Pterycombus b) Pteraclis
Fig. 2 Fig. 3 Pterycombus petersii
3a. Dorsal fin with 54 to 57 rays; anal fin with 47 to 50 rays; vertebrae 51 to 54 (Fig. 4) . . . Pteraclis velifera
3b. Dorsal fin with 49 to 52 rays; anal fin with 40 to 44 rays; vertebrae 45 to 48 (Fig. 5) . . . Pteraclis aesticola
Fig. 4 Pteraclis velifera Fig. 5 Pteraclis aesticola

4a. Snout pointed, lower jaw greatly pro- area almost flat,
broad
jecting (Fig. 7); interorbit broad, al-
most flat between eyes in frontal view distinctly
(Fig. 6a) . . . . . . . . . . . . . . . (Taractes) ® 5 rounded,
arched
4b. Snout not especially pointed, mouth
terminal or almost so (Fig. 12); interor-
bit relatively narrow, convex between
eyes in frontal view (Fig. 6b) . . . . .
. . . . . (Brama, Eumegistus, Taractichthys) ® 6
5a. Scales on caudal peduncle greatly en-

larged, forming a sharp keel in adults; a) b)
pectoral fins more than 36% standard Fig. 6 frontal view
length in specimens exceeding 5 cm;
anal-fin rays mostly fewer than 23 (21
to 23) (Fig. 7) . . . . . . . . . . Taractes rubescens
5b. Scales on caudal peduncle not greatly enlarged, not forming a keel; pectoral fins less than
36% standard length in specimens exceeding 5 cm; anal-fin rays mostly more than 23 (23
to 26) (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taractes asper
snout
keel
pointed
lower jaw
projecting
Fig. 7 Taractes rubescens Fig. 8 Taractes asper
upper lip not connected to upper lip joined to

snout at tip snout at tip
6a. Upper lip free, not connected to snout
near anterior nostril (Fig. 9a) . . . . .
. . . . . . . . . (Eumegistus, Taractichthys) ® 7
6b. Upper lip not free anteriorly, con-
nected to snout near anterior nostril
(Fig. 9b) . . . . . . . . . . . . . . . (Brama) ® 9
a) b)
Fig. 9
7a. Ridge on belly originating just behind pelvic-fin bases, interpelvic space almost flat
(Fig. 10a); no deep transverse grooves at origins of caudal-fin lobes; anal fin with 23 to 25
rays (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Eumegistus illustris
7b. Ridge on belly originating forward of pelvic-fin origins, interpelvic space distinctly convex
(Fig. 10b); deep transverse grooves at origins of caudal-fin lobes (Fig. 11); anal fin with
26 or more rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Taractichthys) ® 8
interpelvic
interpelvic area
anus anus
area flat rounded
a) Eumegistus ventral ridge b) Taractichthys ventral

ridge
Fig. 10 ventral view
2828 Bony Fishes
groove
Fig. 11 caudal fin Fig. 12 Eumegistus illustris
8a. Scales in horizontal series 40 or fewer (34 to 40); predorsal scales fewer than 31 (21 to
29) (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taractichthys steindachneri
8b. Scales in horizontal series more than 40 (usually 42 or more); predorsal scales more than
31 (33 to 37) (Fig. 14) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Taractichthys longipinnis
Fig. 13 Taractichthys steindachneri Fig. 14 Taractichthys longipinnis
9a. First few rays of anal fin not forming a distinct lobe (Fig. 15a); scales in longitudinal series
49 to 68; scales below lateral line 12 to 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
9b. First few rays of anal fin forming a well-developed lobe (except in small juveniles)
(Fig. 15b); scales in longitudinal series 71 to 90; scales below lateral line 18 to 24 . . . . . . . . ® 13
10a. Predorsal scales 23 to 28; anal fin with 23 to 25 rays (rarely 25); sharply demarcated, pale
saddle on undersurface of caudal peduncle (Fig. 16) . . . . . . . . . . . . . . . Brama pauciradiata
10b. Predorsal scales 32 to 43; anal fin with more than 25 rays (rarely 25 in B. dussumieri); no
distinct saddle on undersurface of caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . ® 11
no pronounced anal-fin lobe pale saddle

lobe pronounced
a) b)
Fig. 15 anal fin Fig. 16 Brama pauciradiata
11a. Pectoral fins situated relatively close to ventral surface, distance from pectoral-fin base to
pelvic-fin base less than 12.5% standard length (Fig. 17); scales in longitudinal series
usually more than 59 (60 to 68); anal-fin rays usually 28 or fewer (25 to 29) (Fig. 17)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brama dussumieri
11b. Pectoral fins situated well away from ventral surface, distance from pectoral-fin base to
pelvic-fin base exceeding 12.5% standard length (Fig. 18); scales in longitudinal series
fewer than 59 (48 to 57); anal-fin rays mainly 29 or more (28 to 31) . . . . . . . . . . . . . . . . ® 12
Fig. 17 Brama dussumieri Fig. 18 Brama orcini
12a. Distance from snout to pectoral-fin

origin usually exceeding 30%
standard length in specimens
smaller than 20 cm standard
length; length of upper caudal-fin
lobe less than 40% standard
length in specimens less than
8 cm standard length; total verte-
brae 37 to 40 (mostly 38 or 39)
(Fig. 18) . . . . . . . . . . . . . . Brama orcini
12b. Specimens exceeding 10 cm un-
known; distance from snout to pec-
toral-fin origin usually less than
30% standard length; length of up- Fig. 19 Brama myersi
per caudal-fin lobe more than 50%
standard length; total vertebrae 40
or more (40 to 42) (Fig. 19) . . . Brama myersi
13a. Head very prominently arched; anal-fin rays mostly 30 or more (rarely 29); total vertebrae
41 to 45 (rarely 41) (Fig. 20) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brama brama
13b. Head moderately arched; anal-fin rays mostly 29 or fewer (rarely 30); total vertebrae 39
to 42 (rarely 42) (Fig. 21) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Brama australis
very arched moderately arched
Fig. 20 Brama brama Fig. 21 Brama australis

2830 Bony Fishes

The symbol 0 is given when species accounts are included.
0
0
Brama australis Valenciennes, 1840
0
Brama brama (Bonnaterre, 1788)
0
Brama dussumieri Cuvier, 1831
Brama myersi Mead, 1972
0
0
Brama orcini Cuvier, 1831
Brama pauciradiata Moteki, Fujita, and Last, 1995
0 Eumegistus illustris Jordan and Jordan, 1922
0
0
Pteraclis aesticola (Jordan and Snyder, 1901)
Pteraclis velifera (Pallas, 1769)
0 Pterycombus petersii (Hilgendorf, 1878)
0
0
Taractes asper Lowe, 1843
Taractes rubescens (Jordan and Jordan, 1887)
0 Taractichthys longipinnis (Lowe, 1843)
0 Taractichthys steindachneri (Döderlein, 1883)
Reference
Mead, G.W. 1972. Bramidae. Dana Rept, 81:1-166.
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Brama dussumieri Cuvier, 1831

Frequent synonyms / misidentifications: Brama leucotaenia Fowler, 1938 / Brama orcini Cuvier, 1831;
B. myersi Mead, 1972.
FAO names: En - Lesser bream; Fr - Castagnole mignonne; Sp - Japuta menuda.
Diagnostic characters: Body very deep, very compressed; forehead slightly arched. Upper lip fused to
head just forward of anterior nostril. Eye subequal or slightly larger than snout, located near middle of head
in adults. Palatine and vomerine teeth small and sometimes absent. Total gill rakers in first gill arch 13 to
17. Dorsal fin originating about above pectoral-fin base; dorsal and anal fins mostly covered by scales, no
sheath of scales at their bases into which fins can be depressed; no anal-fin lobe; dorsal-fin rays 31 to 36
(mainly 32 to 34); anal-fin rays 25 to 29 (rarely 25 or 29). Upper caudal-fin lobe greatly extended in
specimens less than 12 cm standard length. Pectoral fins positioned relatively low on side (distance
to pelvic fins 9.9 to 11.7% standard length in specimens exceeding 5 cm standard length); pectoral-fin rays
18 to 22 (mainly 19 or 20). Pelvic fins relatively long (more than 11% standard length in specimens
greater than 10 cm, rarely less than 15% standard length in specimens less than 10 cm). Interpelvic area
almost flat, narrow, greatest width slightly more than 1/2 eye diameter in adults. Scales in horizontal
series 60 to 68; rows above lateral line 12 to 18, rows below lateral line 13 to 16; predorsal scales 32 to
42; scales below lateral line oblique and very tall; scales of adults without spines. Vertebrae 40 to 43
(mainly 41 or 42). Colour: uniform silvery blue, slightly darker dorsally; underside of pectoral-fin base
pale or dusky; vertical fins dark; pectoral and pelvic fins translucent.
Size: Maximum standard length about 19 cm, commonly to 15 cm.
Habitat, biology, and fisheries: Pelagic in upper oceanic waters and common throughout range. Probably
the dominant member of the genus in tropical seas where it appears to replace temperate species such
a s Brama brama a n d B.
japonica. Spawns throughout
the year. Available only in
small quantities as bycatch of
small-scale, traditional line
fisheries.
uted in equatorial seas be-
tween latitudes of 35° in
depths to about 200 m.
Range appears to be limited
by the location of the 24°C
isotherm.
2832 Bony Fishes
Brama orcini Cuvier, 1831

Frequent synonyms / misidentifications: Collybus drachme Snyder, 1904 / Brama dussumieri Cuvier,
1831; B. myersi Mead, 1972.
FAO names: En - Bigbelly pomfret.
Diagnostic characters: Body very deep, very compressed; forehead slightly arched. Upper lip fused to
head just forward of anterior nostril. Eye subequal or slightly larger than snout, located near middle of head
in adults. Palatine and vomerine teeth small and sometimes absent. Dorsal fin originating about above
pectoral-fin base; dorsal and anal fins mostly covered by scales, no sheath of scales at their bases into
which fins can be depressed; no anal-fin lobe; dorsal-fin rays 33 to 36 (mainly 34 or 35); anal-fin rays 28
to 31 rays (rarely 31). Upper caudal-fin lobe greatly extended. Pectoral fins positioned relatively high
on side (distance to pelvic fins 13.3 to 16.6% standard length in specimens exceeding 50 cm standard
length); pectoral-fin rays 18 to 21 (mainly 19 or 20). Pelvic fins relatively short (mostly less than 11%
standard length in specimens greater than 100 cm, rarely greater than 15% standard length in specimens
less than 100 cm). Interpelvic area almost flat, narrow, greatest width slightly more than 1/2 eye diameter
in adults. Scales in horizontal series 48 to 57; rows above lateral line 10 to 15, rows below lateral line
13 to 16; predorsal scales 33 to 43; scales below lateral line oblique and very tall; scales of adults without
spines. Total gill rakers in first gill arch 12 to 16. Vertebrae 37 to 40 (mainly 38 or 39). Colour: uniform
brownish or silvery, slightly darker dorsally; underside of pectoral-fin base black; vertical fins dark;
pectoral and pelvic fins translucent.
Size: Maximum standard length about 30 cm, commonly exceeding 20 cm.
Habitat, biology, and fisheries: Shallow, epipelagic pomfret occurring primarily in the vicinity of land
masses. Largest tropical member of the genus. Major prey species of yellowfin tuna, Thunnus albacares.
Taken incidentally as bycatch of small-scale, traditional line fisheries. It is likely to be the most frequently
caught bramid in a region in which catch data for the family is poor. Like Brama dussumieri, it is both good
to eat and acceptable to
domestic markets of the
Indo-Pacific.
Distribution: Widespread in
Indian and Pacific oceans in
tropical seas between lati-
tudes of 30°. Mostly captured
within 100 m of surface and in
water temperatures above
18.3° C.
Brama australis Valenciennes, 1840

En - Southern rays bream.
Maximum standard length about 53 cm. Potentially valuable food fish in temperate parts of the
Pacific Ocean where it is caught on tuna longline. Found in southern sector of the area but rarely
caught. Possibly circumglobal in the southern hemisphere below 25°S.
Brama brama (Bonnaterre, 1788)

En - Rays bream.
Maximum standard length about 50 cm. Possibly in great abundance in the South Pacific but
primarily a temperate species that is caught only occasionally in the area. Forms a major winter
fishery off Spain and Portugal where it is targeted by longliners or jigged near the surface.
Considered to be widespread in the North Atlantic, Indian and South Pacific oceans, mostly in
latitudes higher than 30°.
Brama myersi Mead, 1972

En - Myers pomfret.
Maximum standard length 8 cm. A dwarf species of no commercial significance. Not well known but
possibly widespread in the tropical Indian and Pacific oceans between latitude 30°.
? ?
?
?
2834 Bony Fishes
Brama pauciradiata Moteki, Fujita, and Last, 1995

En - Shortfin pomfret.
Maximum standard length 8 cm. Another dwarf tropical species of no commercial significance.
Known from eastern Indian, and western and Central Pacific oceans.
? ?
Eumegistus illustris Jordan and Jordan, 1922

En - Brilliant pomfret.
Maximum standard length about 55 cm. Seldom caught on lines but trawled occasionally. Appears
to live deeper than other bramids. Several records from depths exceeding 400 m and a single record
from 620 m. Known only from the Pacific Ocean, mainly from warm-temperate latitudes.
(after Masuda et al., 1984)
Pteraclis aesticola (Jordan and Snyder, 1901)

En - Pacific fanfish.
Maximum standard length about 43 cm. Rarely caught. Occurs primarily in warm-temperate and
tropical parts of the Pacific Ocean.
(after Mead, 1972)

Pteraclis velifera (Pallas, 1769)

En - Southern fanfish.
Maximum standard length about 53 cm. Rarely caught. Occurs mainly in temperate parts of the
Indian and Pacific oceans, and possibly the South Atlantic.
Pterycombus petersii (Hilgendorf, 1878)

En - Prickly fanfish.
Maximum standard length about 32 cm. Rarely caught. Widely distributed in both tropical and
temperate seas throughout the Pacific and Indian oceans.
Taractes asper Lowe, 1843

En - Flathead pomfret.
Maximum standard length about 46 cm. Rarely caught in warm seas. Widely distributed in temperate
latitudes of the North Atlantic, North Pacific, and in the southern hemisphere, off southern Australia,
New Zealand, Chile, and South Africa.
2836 Bony Fishes
Taractes rubescens (Jordan and Jordan, 1887)

En - Knifetail pomfret.
Maximum standard length at least 70 cm. Caught occasionally as bycatch of the tuna longline
fishery. Probably circumtropical but yet to be recorded from the Indian Ocean.
?
?
Taractichthys longipinnis (Lowe, 1843)

En - Bigscale pomfret.
Maximum standard length about 1 m. Rarely caught in the tropical Indo-Pacific. Widespread in the
Atlantic Ocean but occurring mainly in temperate southern parts of the Indian and Pacific oceans.
Taractichthys steindachneri (Döderlein, 1883)

En - Sickle pomfret.
Maximum standard length at least 60 cm. Caught occasionally as bycatch of the tuna longline
fishery. Widespread mainly in warm-temperate and tropical parts of the Indian and Pacific oceans.
click for next page

Perciformes: Percoidei: Caristiidae 2837
CARISTIIDAE
Manefishes
by J.R. Paxton
D iagnostic characters: L a r g e ( t o
60 cm) perciform fishes, with deep,
strongly compressed soft body. Head
short. Eye large. Snout usually short,
very steep in large specimens. Mouth
moderate, jaws reaching level of posterior
margin of eye in large specimens. Teeth
as small canines in jaws and vomer, pre-
sent or absent on palatines. Fin spines
poorly developed; a single dorsal fin
with 28 to 36 elements, all elongate,
origin over head; anal fin with 17 to 22
elements; caudal fin with 17 principal
rays and procurrent spines; pelvic-fin
origin thoracic, the fin with I spine and 5
soft rays, elongate, longer than pecto-
ral-fin rays, folding into shallow
groove; pectoral fins with 16 to 21 rays.
Scales cycloid, deciduous. Total verte-
brae 33 to 40. Colour: body pinkish
brown to brown, fins black.
Habitat, biology, and fisheries: Juve-
niles epi- and mesopelagic, adults meso- and bathypelagic. Feed in association with siphonophores on
zooplankton and small fishes. Rare oceanic fishes of no commercial importance.
Remarks: One or 2 genera with perhaps 4 or more species in tropical and temperate latitudes throughout
the world ocean, except the northeastern Pacific. The family is in need of revision, with undescribed taxa
questions about the 2 nominal genera and 5 nominal species.

Bramidae: body firm, covered with thick, adhesive scales (rather than thin, deciduous scales); pelvic fins
shorter than pectoral fins; colour brownish black, bluish black, or silvery.
Diretmidae: dorsal- and anal-fin rays not exceptionally elongate, but covered with small, spinose scales;
pelvic fins with laminar (rather than normal) spine and 6 soft rays.
Bramidae Diretmidae

Caristius macropus (Bellotti, 1903)
Platyberyx cf. opalescens Zugmayer, 1911
Reference
Heemstra, P.C. 1986. Family Caristiidae. In Smiths’ sea fishes, edited by M.M. Smith and P.C. Heemstra.
Johannesburg, Macmillan South Africa, pp. 636-637.
2838 Bony Fishes
EMMELICHTHYIDAE
Rovers (rubyfishes)
by K.E. Carpenter
D iagnostic characters (for species found in the area): Moderate-sized (to 58 cm) fusiform perciform
fishes. Body moderately slender, body depth 3.2 to 6 times in standard length. Upper and lower profiles
of head nearly equally sloping and gently curved, head length 2.9 to 3.9 times in standard length. Rear
margin of opercle with 2 flat spines and sometimes an additional smaller spine dorsally. Eye large, its bony
orbit diameter 2.5 to 3.7 times in head length, nearly bisected by a horizontal from tip of snout to middle
of caudal-fin base. Mouth small, upper jaw highly protrusible, premaxilla with a long median
ascending process and prominent midlateral process; maxilla expanded distally, scaly, and not
covered by preorbital bone; a distinct supramaxillary bone dorsal to maxilla; jaws toothless or with a
few small teeth in front, vomer and palatines toothless or with a few small teeth. Dorsal fin deeply notched,
or separated by a gap with 0 to 3 short isolated spines, with XI to XIII spines and 10 to 12 soft rays.
Anal fin with III spines and 9 to 10 soft rays. Caudal fin deeply forked. Pelvic fins with I spine and 5 soft
rays. Body and most of head covered with small finely ctenoid scales. Colour: typically reddish, reddish
brown, pinkish, silvery pink, or orange-red, lighter ventrally; Erythrocles taeniatus with a distinct red stripe
laterally.
Habitat, biology, and fisheries: Found near the bottom in

depths between 100 and 400 m, although sometimes found
in shallower waters, especially at night. They typically feed
on small mesopelagic fishes and zooplankton such as ser-
gistid shrimps. Because of their generally deep distribution
and rarity, they are not considered important to fisheries.
Emmelichthyids are tropical and temperate species but in
the Indo-Pacific typically found between the latitudes of
20°N and 45°S. The distribution of emmelichthyids in the head
Western Central Pacific is patchy with each species re-
ported from a single general location. Emmelichthys karnellai is reported from Guam, E. strusakeri is
reported from the Philippines, Erythrocles scintillans is reported from Tahiti, E. schlegeli is reported from
the central South China Sea, and E. taeniatus is known only from New Caledonia.

Caesionidae: maxilla scaleless and partially covered
by preorbital bone; no supramaxilla; most caesionids
have 14 to 22 soft dorsal-fin rays (10 to 12 soft
dorsal-fin rays in emmelichthyids) except
Dipterygonotus balteatus w h i c h m o s t c l o s e l y
resembles an emmelichthyid (and previously was
included in the Emmelichthyidae), however, D.
balteatus most often has XIV dorsal-fin spines (rarely
XII or XIII; emmelichthyids with XI to XIII dorsal-fin
spines). Caesionidae
Percifomres: Percoidei: Emmelichthyidae 2839
Lutjanidae: maxilla scaleless and partially covered by pre-

orbital bone; no supramaxilla; teeth typically well developed
in jaws, vomer, and palatines (toothless or with a few small
teeth in emmelichthyids).
Lutjanidae
Key to species of Emmelichthyidae occurring in the area
1a. Dorsal fin continuous but deeply notched before last spine; dorsal-fin spines XI (Fig. 1a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Erythrocles) ® 2
1b. First and second dorsal fins separated, the gap between them with 0 to III isolated
spines; typically XII or XIII (rarely XI) dorsal-fin spines (Fig. 1b) . . . . . . . . (Emmelichthys) ® 4
a) Erythrocles b) Emmelichthys
Fig. 1 dorsal fins
2a. Fleshy protuberance on rear margin of gill cavity . . . . . . . . . . . . . . . . Erythrocles schlegelii

(in the area so far only reported from South China Sea)
2b. No fleshy protuberance on rear margin of gill cavity . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Most frequently XII (less frequently XI) dorsal-fin spines; a distinct red stripe midlaterally
on body when fresh . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythrocles taeniatus
(New Caledonia)
3b. Most frequently XI (less frequently X) dorsal-fin spines; no red strip on sides when fresh
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Erythrocles scintillans
(in the area so far only reported from Tahiti)
4a. Gap between first and second dorsal fin with I to III short isolated spines . . Emmelichthys strusakeri
(in the area so far only reported from the Philippines)
4b. Gap between first and second dorsal fin without isolated spines, scales continuous
across gap between dorsal fins . . . . . . . . . . . . . . . . . . . . . . . . Emmelichthys karnellai
(in the area so far only reported from Guam)

Emmelichthys karnellai Heemstra and Randall, 1977
Emmelichthys strusakeri Heemstra and Randall, 1977
Erythrocles scintillans Jordan and Thompson, 1912
Erythrocles schlegelii Richardson, 1846
Erythrocles taeniatus Randall and Rivaton, 1992
References
Heemstra, P.C. and J.E. Randall. 1977. A revision of the Emmelichthyidae (Pisces: Perciformes). Aust. J. Mar.
Freshwater Res., 28:361-396.
Nor, L.A., N.N. Kykharev, A.K. Zaytiev. 1985. The biology of Erythrocles schlegelii (Richardson) (Emmelichthyidae)
of the South China Sea. J. Ichthyol., 25(2):146-149.
Randall, J.E. and J. Rivaton. 1992. Erythrocles taeniatus, a new emmelichthyid fish from New Caledonia. Copeia,
1992(4):1028-1032.
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2840 Bony Fishes
LUTJANIDAE
Snappers (jobfishes)
by W.D. Anderson, Jr. and G.R. Allen
D iagnostic characters: Typical perch-like fishes, oblong in shape, moderately compressed (size to
160 cm). Eye usually moderate. Premaxillae usually moderately protrusible (fixed in Aphareus and
Randallichthys). Two nostrils on each side of snout. Mouth terminal and fairly large. Maxilla slipping for
most or all of its length under lacrimal when mouth closed. Supramaxilla absent. Jaws usually with
more or less distinct canines (canines absent in Aphareus, Parapristipomoides, and Pinjalo). Vomer and
palatines usually with teeth. Pterygoids usually toothless. Cheek and operculum scaly; maxilla with or
without scales; snout, lacrimal, and lower jaw naked. Opercular spines 2. Branchiostegal rays 7. Dorsal
fin single, spinous portion sometimes deeply incised posteriorly where it joins soft portion. Caudal fin
truncate to deeply forked. Dorsal fin with X to XII spines and 10 to 19 soft rays. Anal fin with III spines and
7 to 11 soft rays. Principal caudal-fin rays 17 (9 in upper lobe, 8 in lower lobe of fin). Pectoral-fin rays 14
to 19. Pelvic fins with I spine and 5 soft rays. Scales moderate to rather small, ctenoid. Lateral line complete.
Pelvic axillary process usually well developed. Vertebrae 24 (10+14). Colour: highly variable; mainly from
red through yellow to blue; often with blotches, lines, or other patterns.
dorsal fin with X-XII spines

scales on cheek and and 10-19 soft rays
opercle but no scales
between eye and mouth
preopercle usually
serrated
anal fin with III spines
and 7-11 soft rays
Habitat, biology, and fisheries: Found worldwide in warm seas; juveniles of several species of Lutjanus
enter estuaries and the lower reaches of fresh-water streams; a few Indo-Pacific species of Lutjanus are
inhabitants of fresh waters. Mostly bottom-associated fishes, occurring from shallow inshore areas to
depths of about 500 m, mainly over reefs or rocky outcrops. Active predators, mostly nocturnal, feeding
on fishes, crustaceans (especially crabs, shrimps, stomatopods, lobsters), molluscs (gastropods, cepha-
lopods), and pelagic urochordates; plankton is particularly important in the diets of those species with
reduced dentition and numerous well-developed gill rakers. Gonochoristic (sexes separate), reaching
sexual maturity at about 40 to 50% of maximum length, with big females producing large numbers of eggs.
Populations in continental waters have extended spawning throughout the summer, whereas those
occurring around islands spawn throughout the year with peaks in spring and fall; lutjanids are batch
spawners, with individual females usually spawning several times in a reproductive season. Spawning is
apparently at night, on some occasions coinciding with spring tides. Courtship terminates in a spiral swim
upward, with gametes released just below the surface. Eggs and larvae identified as lutjanid are pelagic;
the larvae avoid surface waters during the day, but display a more even vertical distribution at night.
Long-lived, slow-growing fishes with relatively low rates of natural mortality and with considerable
vulnerability to overfishing. Snappers are important to artisanal fisheries, but seldom the prime interest of
major commercial fishing activities; many are fine food fishes, frequently found in markets, although some
species are ciguatoxic in certain areas. They are caught with handlines, traps, a variety of nets, and trawls.
The species that reach large sizes are important recreational fishes in some areas, including the waters
off the eastern coast of Australia. For 1995, the FAO Yearbook of Fishery Statistics reports a total catch
of around 93 200 t of Lutjanidae from the Western Central Pacific.
Perciformes: Percoidei: Lutjanidae 2841

Caesionidae: mouth small (fairly large in Lutjanidae); premaxillae extremely protrusible (premaxillae
moderately protrusible or fixed in Lutjanidae); teeth small or minute (jaws usually with distinct canines in
Lutjanidae); teeth absent on premaxilla, vomer, and palatines in some species (teeth typically well
developed on those bones in Lutjanidae); caudal fin deeply forked, with pointed lobes (caudal-fin shape
variable in Lutjanidae -frequently truncate, emarginate, or lunate).
Haemulidae: scales present on snout and lacrimal, those on lacrimal often embedded (snout and lacrimal
naked in Lutjanidae); preoperculomandibular canal of lateralis system opening under chin through median
longitudinal groove or enlarged pores or both (no chin pores in Lutjanidae); usually no teeth on vomer, no
teeth on palatines (teeth usually present on those bones in Lutjanidae); vertebrae 26 or 27 (24 in
Lutjanidae).
Caesionidae Haemulidae
Lethrinidae: gill membranes broadly united to each other (not united in Lutjanidae); preopercular margin
typically smooth (usually serrate in Lutjanidae); branchiostegal rays 6 (7 in Lutjanidae); soft rays in dorsal
fin 9 or 10 (usually 11 or more in Lutjanidae); no teeth on vomer or palatines (teeth usually present on
those bones in Lutjanidae); most species lack scales on cheek (cheek scales present in Lutjanidae); lips
frequently fleshy (usually not fleshy in Lutjanidae).
Nemipteridae: preopercular margin typically smooth (usually serrate in Lutjanidae); soft rays in dorsal fin
9 (almost always 10 or more in Lutjanidae); soft rays in anal fin usually 7, rarely 8 (almost always 8 or more
in Lutjanidae); branchiostegal rays 6 (7 in Lutjanidae); no teeth on vomer or palatines (teeth usually present
on those bones in Lutjanidae).
Lethrinidae Nemipteridae
Sparidae: preopercular margin typically smooth

(usually serrate in Lutjanidae); branchiostegal rays
6 (7 in Lutjanidae); teeth in jaws variable-conical,
incisiform, or molariform (no incisiform teeth in Lut-
janidae, molariform teeth found in jaws of only a
single eastern Pacific species of Lutjanidae, submo-
lariform teeth in jaws of the western Pacific lutjanid
Symphorichthys spilurus).
Sparidae
2842 Bony Fishes
Key to the genera of Lutjanidae occurring in the area

Notes: species names are given when a genus includes a single species. Counts of gill rakers include
rudiments, if present.
1a. Dorsal and anal fins without scales; dorsal fin with X spines and 10 or 11 soft rays . . . . . . . . ® 2
1b. Soft dorsal and anal fins with scales or sheathed with scales basally; dorsal fin with X to
XII spines and 11 to 19 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
2a. Maxilla with scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Maxilla without scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
3a. Spinous portion of dorsal fin deeply incised at its junction with soft portion; dorsal fin with
X spines and 11 (very infrequently 10) soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . Etelis
3b. Spinous portion of dorsal fin not deeply incised at its junction with soft portion; dorsal fin
with X spines and 10 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Last soft ray of both dorsal and anal fins shorter than next to last soft ray . . . . . . . . . Paracaesio
4b. Last soft ray of both dorsal and anal fins about equal to or slightly longer than next to last
soft ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapristipomoides squamimaxillaris
5a. Premaxillae essentially not protrusible, attached to snout at symphysis by a frenum . . . . . . . ® 6
5b. Premaxillae protrusible, not attached to snout by frenum . . . . . . . . . . . . . . . . . . . . . . ® 7
6a. Vomer without teeth (small juveniles may have minute teeth on vomer); teeth in jaws very
small, no caniniform teeth; pectoral fins somewhat shorter than head; lateral surface of
maxilla smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aphareus
6b. Vomer with teeth; jaws with some caniniform teeth; pectoral fins about 1/2 to 2/3 length
of head; lateral surface of maxilla with a series of well-developed longitudinal ridges . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Randallichthys filamentosus
7a. Dorsal fin with X spines and 11 (rarely 10) soft rays; last soft ray of both dorsal and anal
fins longer than next to last soft ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7b. Dorsal fin with X spines and 10 soft rays; last soft ray of both dorsal and anal fins shorter
than next to last soft ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . → 9
8a. Groove present on snout below nostrils (Fig. 1); pectoral fins less than 1/2 length of head
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aprion virescens
8b. No groove on snout; pectoral fins a little shorter than head to somewhat longer than head
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pristipomoides
9a. Upper lip with a median fleshy protrusion, well developed in adults (Fig. 2); spines of
dorsal and anal fins strong, very robust in large adults . . . . . . . . . . . Lipocheilus carnolabrum
9b. Upper lip without a median fleshy protrusion . . . . . . . . . . . . . . . . . . . . . . . . Paracaesio
groove
frontal view lateral view
Fig. 1 Aprion Fig. 2 head of Lipocheilus carnolabrum
10a. Vomer without teeth; dorsal fin with X spines and 14 to 19 soft rays; 1 or more anterior
soft dorsal-fin rays produced as filaments (at least in juveniles) . . . . . . . . . . . . . . . . . . ® 11
10b. Vomer with teeth; dorsal fin with X to XII spines and 11 to 16 soft rays; none of anterior
soft dorsal-fin rays produced as filaments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
11a. Anterior profile quite steep; dorsal fin with X spines and 17 to 19 soft rays; upper and
lower pharyngeals enlarged and bearing large molariform teeth . . . . . . . Symphorichthys spilurus
11b. Anterior profile sloping more gently; dorsal fin with X spines and 14 to 17 soft rays; upper
and lower pharyngeals not particularly enlarged, not bearing molariform teeth . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Symphorus nematophorus
12a. First gill arch with 60 or more gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . Macolor
12b. First gill arch with 20 or fewer gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Upper and lower profiles of head equally rounded; eye set toward middle of head; mouth
rather small, somewhat upturned; no fang-like canines at anterior ends of jaws . . . . . . . . Pinjalo
13b. Upper and lower profiles of head not equally rounded, upper profile evenly rounded to
steeply sloped, and lower profile flattened; eye closer to upper profile of head than to
lower; mouth larger, usually not upturned; some fang-like canines usually present at
anterior ends of jaws . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus
Key to the species of Aphareus occurring in the area

Remark on key character: counts of gill rakers include rudiments, if present.
1a. First gill arch with 6 to 12 gill rakers on upper limb and 15 to 18 on lower limb (total 22 to
28); colour of body varying from steel blue to purplish brown . . . . . . . . . . . . . Aphareus furca
1b. First gill arch with 16 to 19 gill rakers on upper limb and 32 to 35 on lower limb (total 49
to 52); colour of body varying from blue-grey or mauve to reddish . . . . . . . . . Aphareus rutilans
Key to the species of Etelis occurring in the area

Remark on key character: counts of gill rakers include rudiments, unless otherwise stated.
1a. Total gill rakers on first gill arch 17 to 22 (total number excluding rudiments, 10 to 13);
upper lobe of caudal fin short, 26 to 30% standard length in specimens larger than
120 mm standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Etelis carbunculus
1b. Total gill rakers on first gill arch 23 to 36 (total number excluding rudiments, 20 to 33);
upper lobe of caudal fin longer, 30 to 75% standard length in individuals larger than
130 mm standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Total gill rakers on first gill arch 32 to 36; upper lobe of caudal fin relatively short, 30 to
34% standard length; length of pelvic fin 17 to 18% standard length . . . . . . . . . Etelis radiosus
2b. Total gill rakers on first gill arch 23 to 28; upper lobe of caudal fin short to very long, 33 to
75% standard length in individuals larger than 130 mm standard length; length of pelvic fin
19 to 23% standard length in individuals larger than 80 mm standard length . . . . . . Etelis coruscans
Key to the species of Lutjanus occurring in the area

1a. Preorbital (“suborbital”) space broad
narrow preorbital
(distance between upper jaw and preorbital space
eye) very narrow, 9.2 to 16.3 space
times in head length (Fig. 3a);
body slender, usually 3 (some-
times 2.9) or more times in stand-
ard length; dorsal-fin spines
usually XI (occasionally X o r
rarely XI); soft dorsal-fin rays 12 . . . . . . ® 2
1b. Preorbital (“suborbital”) space a) b)
wider, 3.3 to 8.9 times in head
length (Fig. 3b); body deeper, 2.1 Fig. 3 lateral view of head
to 3.1 times, but usually less than
3 times, in standard length; dor-
sal-fin spines variable, X to XII;
soft dorsal-fin rays occasionally
12 (usually 13 or more) . . . . . . . . . . ® 3
2844 Bony Fishes
2a. Body depth 3.5 to 3.8 times in standard length; tongue usually smooth without teeth; a
dark band from snout to caudal-fin base and 2 pearly spots above lateral line, 1 below
spinous portion and the other below soft portion of dorsal fin . . . . . . . . . . . Lutjanus biguttatus
(central Indian Ocean to Melanesia)
2b. Body depth 2.9 to 3.3 times in standard length; tongue with a patch of fine granular teeth;
colour generally silvery white with a broad yellow stripe along middle of side to caudal-fin
base, and narrow yellowish lines, corresponding with longitudinal scale rows . . . Lutjanus lutjanus
(eastern Africa to western Pacific)
3a. Ground colour pale (mainly yellow in life) with a series of 4 or 5 longitudinal stripes (blue
in life, often brownish in preservative) on side . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3b. Colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
4a. Dorsal-fin spines XI or XII . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus bengalensis

4b. Dorsal-fin spines X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Four stripes on side, belly more or less abruptly whitish, frequently with thin grey lines;
scale rows on cheek 5 or 6; upper pectoral-fin rays darkish . . . . . . . . . . . . . Lutjanus kasmira
(eastern Africa to central Pacific)
5b. Five stripes on side, belly not abruptly whitish and without thin lines; scale rows on cheek
10 or 11; upper pectoral-fin rays pale . . . . . . . . . . . . . . . . . . . . Lutjanus quinquelineatus
(central Indian Ocean to western Pacific)
6a. Longitudinal scale rows above lateral line obliquely positioned (Fig. 4a) . . . . . . . . . . . . . ® 7
6b. Longitudinal scale rows above lateral line entirely horizontal (Fig. 4b) or some rows rising
obliquely from below middle part of dorsal fin (Fig. 4c) . . . . . . . . . . . . . . . . . . . . . . . ® 31
scale rows scale rows
oblique horizontal
a) b)
scale rows
oblique
c)
Fig. 4
7a. Vomerine tooth patch triangular or dia-

mond-shaped with a medial posterior ex-
tension (Fig. 5c) . . . . . . . . . . . . . . . . . . . ® 8
7b. Vomerine tooth patch crescentic to trian-
gular without a posterior extension . . . . . . . . ® 13
a) b) c)
Fig. 5 shapes of the vomerine tooth patch

8a. Axil of pectoral fins with distinct black spot on upper portion; a series of 8 or 9 relatively
broad orange or yellow stripes on side; soft dorsal-fin rays usually 15 (occasionally 14,
rarely 16); soft anal-fin rays 9 . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus carponotatus
(India to Melanesia and Australia)
8b. Axil of pectoral fins without black spot; colour not as above; soft dorsal-fin rays usually
13 or 14 (rarely 12); soft anal-fin rays usually 8 (rarely 9) . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. A large black spot usually present on upper side, juveniles sometimes with an ocellated
spot and/or a series of 4 to 7 broad dark stripes on side (adults of L. fulviflamma with
yellow stripes) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
9b. Black spot absent; a series of narrow, yellowish longitudinal lines on side, those on upper
back slanting upward toward dorsal-fin base, sometimes an enlarged darker stripe from
eye to middle of caudal-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
10a. Soft dorsal-fin rays usually 14; a relatively wide gap between temporal scale bands of
each side (Fig. 6a); spot on upper side situated mainly above lateral line; young
specimens with series of 4 to 7 broad stripes (blackish to orange or yellow-brown in life)
on side, these persisting as thin stripes in adults from the western Indian Ocean . . Lutjanus russelli
10b. Soft dorsal-fin rays usually 13; little or no gap between temporal scale bands of each side
(Fig. 6b); spot on upper side situated mostly below lateral line or bisected by it, spot
sometimes very elongated; young specimens without series of 4 to 7 broad dark stripes
on side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus fulviflamma
(eastern Africa to central South Pacific)
11a. Midlateral stripe usually broader and darker than other stripes on side; transverse scale
rows on cheek 7 to 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus vitta
(western Indian Ocean to western Pacific)
11b. Midlateral stripe not broader or darker than other stripes on side, yellow in life and faint
or absent in preserved specimens; transverse scale rows on cheek usually 6 or 7
(occasionally 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Predorsal scales extending to mid-interorbital level; a blunt, flattened spine on upper margin
of opercle, above the main centrally located spine (Fig. 7a); interorbital width 4.4 to 6.5 in
head length; total gill rakers on first gill arch (including rudiments) 18 to 21 . . . . . . . Lutjanus madras
(eastern Africa to Indonesia and the Philippines)
12b. Predorsal scales extending to level of rear part of orbit; blunt spine above central opercle
spine absent (Fig. 7b); interorbital width 6.5 to 6.9 in head length; total gill rakers on first
gill arch (including rudiments) 15 or 16 . . . . . . . . . . . . . . . . . . . . . . . Lutjanus mizenkoi
(Indonesia to Samoa)
flattened
spine
a) Lutjanus russelli b) Lutjanus fulviflamma a) Lutjanus madras b) Lutjanus mizenkoi
Fig. 6 dorsal view of head Fig. 7 lateral view of head
13a. Total gill rakers (including rudiments) on first gill arch (including rudiments) 25 to 30 . . . . . . ® 14
13b. Total gill rakers (including rudiments) on first gill arch (including rudiments) 14 to 23 . . . . . . ® 15
2846 Bony Fishes
14a. Dorsal fin with X spines and 14 soft rays; longitudinal scale rows below lateral line
parallel to axis of body (Fig. 8a); caudal fin emarginate; colour generally pale with a
golden-brown midlateral stripe, slightly narrower than eye, and a series of oblique
golden-brown lines ascending from lateral line to base of dorsal fin . . . . . . . . . . Lutjanus adetii
(eastern Australia and New Caledonia)
14b. Dorsal fin with X spines and 13 or 14 soft rays; scale rows below lateral line ascending
obliquely (Fig. 8b); caudal fin distinctly forked with rounded lobes; colour deep red to
grey, fins red or dark brown to blackish . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus gibbus
(eastern Africa to central West Pacific)
scale rows oblique
scale rows
a) Lutjanus adetii horizontal b) Lutjanus gibbus
Fig. 8
15a. Soft anal-fin rays 10; dorsal fin with XI spines and 16 rays (rarely 15); colour pattern
consisting of 3 dark brown to red transverse bars (may be indistinct in large adults) . . . Lutjanus sebae
15b. Soft anal-fin rays 8 or 9; dorsal-fin elements variable, the fin with X or XI spines and 12
to 16 soft rays; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
16a. Preopercular notch distinctive (moder-

ately to well developed, Fig. 9a) . . . . . . . . . ® 17
16b. Preopercular notch indistinct (shallow or
absent, Fig. 9b) . . . . . . . . . . . . . . . . . . ® 21
17a. Soft dorsal-fin rays 15 or 16; body rela-

tively deep, 2.1 to 2.4 times in standard notch strong notch weak
length; head usually with numerous wavy a) b)
lines (bluish in life); a chalky spot often
present below junction of spinous and Fig. 9 lateral view of head
soft parts of dorsal fin, bordered with
black in juveniles, but lost with age; lips
thick in large adults . . . . . . . . . Lutjanus rivulatus
17b. Soft dorsal-fin rays 13 or 14; body usually more slender, 2.3 to 2.8 times in standard
length; colour not as above; lips not thick in adults . . . . . . . . . . . . . . . . . . . . . . . . ® 18
18a. Caudal fin and distal third of dorsal fin blackish or dusky brown with a narrow white
border . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus fulvus
(eastern Africa to central West Pacific)
18b. Caudal fin yellow or grey basally and yellow distally (tan to medium brown in preserv-
ative) without narrow white border; distal third of dorsal fin not noticeably darker than
remainder of fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
19a. A small (much less than pupil size) chalk-white spot on back just above lateral line at
level of anterior part of soft dorsal fin; colour generally brown to purplish; body depth 2.4
to 2.5 times in standard length; snout 2.6 times and preorbital 4.1 to 5.2 times in head
length; total gill rakers on first gill arch 16 to 19; dorsal-fin spines X . . . . . . . . Lutjanus stellatus
19b. No white spot on back, often a large brownish spot in this position, but usually faint or
absent in fresh dead or preserved specimens; colour generally pink, sometimes with faint
yellow stripes (absent in preservative) on side, margin of dorsal fin sometimes blackish;
snout 2.8 to 3.2, preorbital 5.3 to 6.6 times, both in head length; total gill rakers on first
gill arch 20 to 23; dorsal-fin spines X or XI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
20a. Dorsal-fin spines X; no yellow stripes on side of body . . . . . . . . . . . . . . . . Lutjanus boutton

(Indo-Malayan Archipelago to New Guinea)
20b. Dorsal-fin spines XI; 10 to 12 faint yellow stripes on side of body . . . . . . . Lutjanus rufolineatus
(West Pacific to Samoa and Tonga)
21a. Eight broad transverse bands or saddles on upper half of body and a large black blotch
in centre of caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus semicinctus
(Indonesia to central South Pacific)
21b. Colour pattern not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
22a. Colour pattern consisting of a series of 5 dark stripes on whitish ground colour; 2 or 3
uppermost stripes crossed by dark vertical bars forming a network of light and dark
squares; a large dark spot at base of caudal fin . . . . . . . . . . . . . . . . . . Lutjanus decussatus
(central Indian Ocean to western Pacific)
23a. Nostrils set in a prominent groove running forward nostrils in

from eye in specimens exceeding about 20 cm stand- groove
ard length (Fig. 10); specimens under this size fre-
quently with 2 whitish spots on upper back, anterior
spot below last 4 dorsal-fin spines and posterior one
under last 6 dorsal-fin rays and meeting that of other
side across top of caudal peduncle; colour generally
dark brown on upper back grading to tan or light
brownish (white or pink in life) ventrally; dorsal and
caudal fins dusky; outer portion of anal and pelvic
fins distinctly blackish; upper third of pectoral fins Fig. 10 Lutjanus bohar
dusky brown; tongue with a patch of fine granular teeth
. . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus bohar
23b. Nostrils not set in a groove at all sizes; colour pattern not as above; tongue smooth or
with a patch of granular teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 24
24a. Caudal fin with a distinctive crescentic black marking, remainder of body and fins
uniformly yellowish tan (yellow in life) with a silvery sheen on lower sides . . . . . Lutjanus lunulatus
(central Indian Ocean to Melanesia)
24b. Caudal fin without a distinctive black marking (although a dark smudge or blotch present
on middle caudal-fin rays of L. bitaeniatus under about 20 cm standard length); colour
of body and fins variable . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 25
25a. A black spot on upper side at level of lateral line below soft dorsal fin (faint or occasionally
absent in large adults); remainder of body and fins mainly pale (fins yellow, body pink or
yellow in life); tongue with a patch of find granular teeth, although sometimes absent in
juveniles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus monostigma
25b. Black spot on upper side of body absent, although a saddle or spot sometimes present
on upper portion of caudal peduncle; tongue smooth . . . . . . . . . . . . . . . . . . . . . . . ® 26
2848 Bony Fishes
26a. Dorsal-fin spines XII; a series of 5 or 6 dusky stripes (yellow in life, may be faint in
preservative); longitudinal rows of scales below lateral line rising obliquely . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus dodecacanthoides
(Indonesia and the Philippines)
26b. Dorsal-fin spines X or XI; colour not as above; longitudinal rows of scales below lateral
line parallel to axis of body or rising obliquely . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 27
27a. Axil of pectoral fins black (Fig. 11); colour overall deep red
in life; posterior dorsal- and anal-fin rays elongated to form pectoral-
pointed fins; soft anal-fin rays 8 . . . . . . . . . . . Lutjanus timorensis fin axil
(eastern Indian Ocean to western Pacific) black
27b. Axil of pectoral fins without black marking; colour variable,
although often red in life; posterior dorsal- and anal-fin rays
low and rounded or tall and pointed, but specimens having
latter condition usually with 9 soft anal-fin rays . . . . . . . . . . . ® 28
Fig. 11 Lutjanus timorensis
28a. Dorsal-fin spines X; soft anal-fin rays usually 8 (rarely 9); tongue with a patch of fine,
granular teeth; colour variable, pink to grey-brown (tan to brown in preservative);
juveniles without black saddle on upper caudal peduncle . . . . . . . . . . . . . . . . . . . . . ® 29
28b. Dorsal-fin spines usually XI (rarely X); soft anal-fin rays usually 9 (occasionally 8);
tongue smooth; colour largely reddish (brown in preservative); juveniles usually with a
black saddle on upper caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 30
29a. Interorbital width 4.9 to 5.2 times in head length; body relatively deep, its depth 2.3 to 2.5
times in standard length; snout-forehead profile straight or convex; colour generally red
or pink (brown to yellowish in preservative), fins pale except in juveniles which may have
a crescentic blotch in middle of caudal fin . . . . . . . . . . . . . . . . . . . . . Lutjanus bitaeniatus
(eastern Indian Ocean and Indonesia)
29b. Interorbital width 5.5 to 7.6 times in head length; body generally more slender, its depth
2.5 to 2.8 times in standard length; snout-forehead profile concave in specimens over
about 15 to 20 cm standard length; colour generally grey-brown, reddish or pink
ventrally, dorsal and caudal fins dusky brown or black, frequently with a narrow white
border posteriorly; juveniles with a broad, black, midlateral stripe . . . . . . . . Lutjanus lemniscatus
30a. Mouth relatively small, maxilla length much less than distance between bases of last
dorsal- and anal-fin rays (Fig. 12a); interorbital width 3.5 to 4.8 times in head length
(specimens in excess of 15 cm standard length); some longitudinal scale rows below
lateral line slanting obliquely in posterior direction toward dorsal profile; head profile
convex (in specimens over about 15 cm standard length) . . . . . . . . . . . Lutjanus erythropterus
(northwestern Indian Ocean to western Pacific)
30b. Mouth larger, maxilla length about equal to distance between bases of last dorsal- and
anal-fin rays (Fig. 12b); interorbital width 5.1 to 6.6 times in head length (specimens in
excess of 12 cm standard length); longitudinal scale rows below lateral line horizontal,
although some rows may slant obliquely in juveniles under about 10 cm standard length;
head profile straight or slightly concave . . . . . . . . . . . . . . . . . . . . . Lutjanus malabaricus
a) Lutjanus erythropterus b) Lutjanus malabaricus

Fig. 12
31a. Colour pattern consisting of 4, relatively wide dusky brown or blackish stripes on pale
ground; only juveniles to 15 cm standard length known . . . . . . . . . . . . . Lutjanus maxweberi
(fresh-water streams and mangroves of the Philippines, Indonesia, and New Guinea)
32a. Vomerine tooth patch triangular with medial posterior extension (Fig. 5a, b); preorbital
space narrow, 8.6 to 10.3 times in head length; a prominent black spot, larger than eye,
bisected by the lateral line below posterior part of spinous dorsal fin . . . . . . Lutjanus ehrenbergii
32b. Vomerine tooth patch crescentic to triangular without a medial posterior extension
(Fig. 5c); preorbital space wider, 4.7 to 4.9 times in head length; black spot on back
present or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 33
33a. A large black spot on upper back usually present, if absent ground colour pale . . . . . . . . . ® 34
33b. Black spot on upper back absent, ground colour dark . . . . . . . . . . . . . . . . . . . . . . . ® 35
34a. Ground colour pale, each scale on side often with a brownish spot forming longitudinal
rows on side; large spot on back, if present, situated mainly above lateral line; interorbital
width 5.6 to 7.3 times in head length; preorbital space 4.9 to 6 times in head length;
tongue with a patch of fine granular teeth . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus johni
34b. Ground colour dusky brown; large spot on back bisected equally by the lateral line;
interorbital width 5 to 5.6 times in head length; preorbital space about 6.8 times in head
length; tongue smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus fuscescens
(fresh waters of China, the Philippines, Indonesia, and New Guinea)
35a. Body depth 2.5 to 2.9 (average about 2.7) times in standard length; least depth of caudal
peduncle 3 to 3.5 times in head length; longitudinal scale rows on upper back parallel to
lateral line anteriorly and some rows usually ascend obliquely below posterior dorsal-fin
spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus argentimaculatus
(a marine species also dwelling in brackish estuaries and lower reaches of fresh-water streams; eastern Africa to
central Pacific)
35b. Body depth 2.2 to 2.6 times in standard length; least depth of caudal peduncle 2.5 to 3
times in head length; longitudinal scale rows on upper back entirely parallel to lateral line
(Fig. 4c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lutjanus goldiei
(fresh-water species; southern New Guinea north to Japan)
Key to the species of Macolor occurring in the area

Remark on key character: counts of gill rakers include rudiments, if present.
1a. First gill arch with 37 to 42 gill rakers on upper limb and 71 to 81 on lower limb (total 110
to 122); anal fin with III spines and usually 10 (occasionally 11) soft rays; pelvic fins very
long and pointed in young, but short and rounded in adults . . . . . . . . . . . . Macolor macularis
1b. First gill arch with 26 to 38 gill rakers on upper limb and 60 to 71 on lower limb (total 89
to 107); anal fin with III spines and usually 11 (occasionally 10) soft rays; pelvic fins short
and rounded at all sizes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Macolor niger
Key to the species of Paracaesio occurring in the area

Remark on key character: counts of lateral-line scales are of tubed scales; teeth are measured from points
where they emerge from jaws to their distal tips.
1a. Lateral-line scales 68 to 72; caudal fin forked . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Lateral-line scales 48 to 50; caudal fin lunate or emarginate . . . . . . . . . . . . . . . . . . . ® 4
2850 Bony Fishes
2a. Several prominent exserted canine teeth at anterior ends of upper and lower jaws
(longest anterior tooth in upper jaw 6.3 to 8.7% postorbital head length, longest anterior
tooth in lower jaw 6.5 to 6.9% postorbital head length); maxilla without scales . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracaesio paragrapsimodon
2b. Teeth at anterior ends of jaws smaller, rarely prominent (longest anterior tooth in upper
jaw 2.4 to 5.9% postorbital head length; longest anterior tooth in lower jaw 1.3 to 6%
postorbital head length); maxilla with or without scales . . . . . . . . . . . . . . . . . . . . . . . ® 3
preopercle
3a. General body colour dark grey, almost
dark purplish brown, dark violet, or preopercle always
bluish; preopercle with 1 to 3 rows with 1-3 without
rows of scales
of scales peripheral to and usually scales
distinctly separated from main
group of cheek scales; maxilla
without scales (Fig. 13a) . . Paracaesio sordida
3b. Caudal fin, upper part of caudal
peduncle, and upper side of body
to anterior end of dorsal fin yellow;
remainder of body mostly blue;
a) Paracaesio sordida b) Paracaesio xanthura
preopercle almost always without
scales peripheral to main group of Fig. 13
cheek scales; maxilla with or with-
out scales (Fig. 13b) . . . Paracaesio xanthura
4a. Maxilla without scales; caudal fin emarginate; general body colour light brown dorsally
and silver ventrally with 4 or 5 broad brown to dark grey vertical bars on body extending
from dorsal surface to midline or below midline laterally . . . . . . . . . . . . . . Paracaesio stonei
4b. Maxilla with scales; caudal fin lunate to emarginate (middle of distal margin of caudal fin
of P. kusakarii becoming rounded with age) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Yellow band running from anterior end of lateral line obliquely to about middle of dorsal
fin; in life sometimes with 8 broad dark vertical bars; nape without pronounced hump;
middle of distal margin of caudal fin not rounded with age . . . . . . . . . . . Paracaesio gonzalesi
5b. No oblique yellow band on side of body; 4 broad dark vertical bars on upper side of body;
large individuals with pronounced hump on nape as a result of higher supraoccipital
crest; middle of distal margin of caudal fin becoming rounded with age . . . . . Paracaesio kusakarii
Key to the species of Pinjalo occurring in the area

Remark on key character: caudal concavity is the horizontal distance between the tips of the longest and
shortest caudal-fin rays.
1a. Dorsal fin with XI spines and 14 or 15 soft rays; anal fin with III spines and 9 or (usually)
10 soft rays; longest anal-fin spine 2.1 to 2.65 times in head length; pectoral-fin length
3.05 to 3.9 times in standard length; pelvic-fin length 1.4 to 1.65 times in head length;
caudal fin deeply emarginate, caudal concavity 2.1 to 3.6 times in head length; diagonal
dark lines following scale rows on nape and upper half to two-thirds of body . . . . . Pinjalo pinjalo
1b. Dorsal fin with XII spines and 13 soft rays; anal fin with III spines and 8 or 9 soft rays;
longest anal-fin spine 2.7 to 3.2 times in head length; pectoral-fin length 2.5 to 3.05 times
in standard length; pelvic-fin length 1.65 to 1.8 times in head length; caudal fin slightly
emarginate, caudal concavity 3.55 to 5.45 times in head length; no diagonal dark lines
on nape and body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pinjalo lewisi
Key to the species of Pristipomoides occurring in the area

Remarks on key characters: counts of gill rakers include rudiments, if present; counts of lateral-line scales
are of tubed scales.
1a. Lateral-line scales 48 to 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Lateral-line scales 57 to 74 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Two golden stripes bordered with blue on snout and cheek; transverse vermiculations
on top of head (Fig. 14a); suborbital wide, its width 7 times in head length at 15 cm
standard length, 5.5 times at 25 cm, 4 times at 40 cm . . . . . . . . . . . Pristipomoides multidens
2b. No golden stripes on snout and cheek; longitudinal vermiculations on top of head
(Fig. 14b); suborbital narrow, its width 8.4 times in head length at 15 cm standard length,
7.3 times at 25 cm, 5.8 times at 40 cm . . . . . . . . . . . . . . . . . . . . . . Pristipomoides typus
3a. Total gill rakers on first gill arch 27 to 33; lateral-line scales 67 to 74 . . . . . . . . . . . . . . .®4
3b. Total gill rakers on first gill arch 17 to 27; lateral-line scales 57 to 67 . . . . . . . . . . . . . . .®5
4a. Vomerine tooth patch with backward prolongation in midline (Fig. 15a; vomerine tooth
patch sometimes diamond-shaped); patch of teeth on tongue . . . . . . . . Pristipomoides sieboldii
4b. Vomerine tooth patch triangular in shape, without backward prolongation in midline
(Fig. 15b); no teeth on tongue . . . . . . . . . . . . . . . . . . . . . . . . . Pristipomoides auricilla
a) Pristipomoides b) Pristipomoides b) Pristipomoides

multidens typus a) Pristipomoides sieboldii auricilla
Fig. 14 dorsal view of head Fig. 15 shapes of vomerine tooth patch
5a. Lateral-line scales 63 to 67; side of body with alternating oblique red and yellow bars
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pristipomoides zonatus
5b. Lateral-line scales 57 to 63; side of body without red and yellow bars . . . . . . . . . . . . . . . ® 6
6a. Total gill rakers on first gill arch 17 to 21; depth of body 2.8 to 3.1 times in standard length;
side of body with numerous irregular dark markings, including zigzag line that runs at
about level of lateral line anteriorly and then onto dorsolateral part of caudal peduncle
posteriorly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pristipomoides argyrogrammicus
6b. Total gill rakers on first gill arch 22 to 27; depth of body 3 to 3.8 times in standard length;
side of body without irregular dark markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Length of upper jaw 2.4 to 2.7 times in head length; caniniform teeth at anterior ends of
jaws not appreciably enlarged; pyloric caeca 7 to 9 (usually 8) . . . . . Pristipomoides filamentosus
7b. Length of upper jaw 2.1 to 2.4 times in head length; canine teeth at anterior ends of both
jaws usually enlarged; pyloric caeca 4 to 6 (usually 5) . . . . . . . . . . . Pristipomoides flavipinnis
2852 Bony Fishes

0 Aphareus furca (Lacepède, 1801)
0 Aphareus rutilans Cuvier, 1830
0 Aprion virescens Valenciennes, 1830
0 Etelis carbunculus Cuvier, 1828
0 Etelis coruscans Valenciennes, 1862
0 Etelis radiosus Anderson, 1981
0 Lipocheilus carnolabrum (Chan, 1970)
0 Lutjanus adetii (Castelnau, 1873)
0 Lutjanus argentimaculatus (Forsskål, 1775)
0 Lutjanus bengalensis (Bloch, 1790)
0 Lutjanus biguttatus (Valenciennes, 1830)
0 Lutjanus bitaeniatus (Valenciennes, 1830)
0 Lutjanus bohar (Forsskål, 1775)
0 Lutjanus boutton (Lacepède, 1803)
0 Lutjanus carponotatus (Richardson, 1842)
0 Lutjanus decussatus (Cuvier, 1828)
0 Lutjanus dodecacanthoides (Bleeker, 1854)
0 Lutjanus ehrenbergii (Peters, 1869)
0 Lutjanus erythropterus (Bloch, 1790)
0 Lutjanus fulviflamma (Forsskål, 1775)
0 Lutjanus fulvus (Schneider, 1801)
0 Lutjanus fuscescens (Valenciennes, 1830)
0 Lutjanus gibbus (Forsskål, 1775)
0 Lutjanus goldiei (Macleay, 1882)
0 Lutjanus johnii (Bloch, 1792)
0 Lutjanus kasmira (Forsskål, 1775)
0 Lutjanus lemniscatus (Valenciennes, 1828)
0 Lutjanus lunulatus (Park, 1797)
0 Lutjanus lutjanus (Bloch, 1790)
0 Lutjanus madras (Valenciennes, 1831)
0 Lutjanus malabaricus Schneider, 1801
0 Lutjanus maxweberi Popta, 1921
0 Lutjanus mizenkoi Allen and Talbot, 1985
0 Lutjanus monostigma (Cuvier, 1828)
0 Lutjanus quinquelineatus (Bloch, 1790)
0 Lutjanus rivulatus (Cuvier, 1828)
0 Lutjanus rufolineatus (Valenciennes, 1830)
0 Lutjanus russelli (Bleeker, 1849)
0 Lutjanus sebae (Cuvier, 1828)
0 Lutjanus semicinctus (Quoy and Gaimard, 1824)
0 Lutjanus stellatus Akazaki, 1983
0 Lutjanus timorensis (Quoy and Gaimard, 1824)
0 Lutjanus vitta (Quoy and Gaimard, 1824)
0 Macolor macularis Fowler, 1931
0 Macolor niger (Forsskål, 1775)
0 Paracaesio gonzalesi Fourmanoir and Rivaton, 1979
0 Paracaesio kusakarii Abe, 1960
0 Paracaesio paragrapsimodon Anderson and Kailola, 1992
0 Paracaesio sordida Abe and Shinohara, 1962
0 Paracaesio stonei Raj and Seeto, 1983
0 Paracaesio xanthura (Bleeker, 1869)
0 Parapristipomoides squamimaxillaris (Kami, 1973)
0 Pinjalo lewisi Randall, Allen, and Anderson, 1987

0 Pinjalo pinjalo (Bleeker, 1850)
0 Pristipomoides argyrogrammicus (Valenciennes, 1832)
0 Pristipomoides auricilla (Jordan, Evermann, and Tanaka, 1927)
0 Pristipomoides filamentosus (Valenciennes, 1830)
0 Pristipomoides flavipinnis Shinohara, 1963
0 Pristipomoides multidens (Day, 1871)
0 Pristipomoides sieboldii (Bleeker, 1857)
0 Pristipomoides typus Bleeker, 1852
0 Pristipomoides zonatus (Valenciennes, 1830)
0 Randallichthys filamentosus (Fourmanoir, 1970)
0 Symphorichthys spilurus (Günther, 1874)
0 Symphorus nematophorus (Bleeker, 1860)
References
Allen, G.R. 1985. FAO species catalogue. Vol. 6. Snappers of the world. An annotated and illustrated catalogue of
lutjanid species known to date. FAO Fish. Synop., (125)6:208 p.
Allen, G.R. and F.H. Talbot. 1985. Review of the snappers of the genus Lutjanus (Pisces: Lutjanidae) from the
Indo-Pacific, with the description of a new species. Indo-Pac. Fishes, (11):87 p.
Anderson, W.D. Jr. 1987. Systematics of the fishes of the family Lutjanidae (Perciformes: Percoidei), the snappers. In
Tropical snappers and groupers: biology and fisheries management, edited by J.J. Polovina and S. Ralston.
Boulder, Colorado, Westview Press, pp. 1-31.
Anderson, W.D. Jr., P.J. Kailola, and B.B. Collette. 1992. Two new snappers (Teleostei: Lutjanidae: Apsilinae):
Paracaesio paragrapsimodon Anderson and Kailola from the western Pacific and P. waltervadi Anderson and
Collette from the western Indian Ocean. Proc. Biol. Soc. Wash., (105)3:443-461.
Randall, J.E., G.R. Allen, and W.D. Anderson Jr. 1987. Revision of the lutjanid genus Pinjalo, with description of a new
species. Indo-Pac. Fishes, (14):17 p.
click for next page

2854 Bony Fishes
Aphareus furca (Lacepède, 1801) (Plate V, 27)

FAO names: En - Smalltoothed jobfish; Fr - Vivaneau tidents; Sp - Pargo boquidulce.
Diagnostic characters: Body elongate, laterally compressed. Nostrils on each side of snout close
together. Lower jaw protruding. Premaxillae not protrusible, fixed by a frenum. Maxilla extending to
below middle of eye or slightly beyond. Teeth in jaws extremely small; roof of mouth usually toothless
(small juveniles may have minute teeth in semicircular pattern on vomer). Maxilla without scales
or longitudinal ridges. Interorbital region flattened. Gill openings extending far anterior to orbit.
First gill arch with 6 to 12 gill rakers on upper limb, 15 to 18 on lower limb (total 22 to 28). Dorsal fin
continuous, not deeply incised near junction of spinous and soft portions. Last soft ray of both dorsal
and anal fins well produced, longer than next to last ray. Caudal fin forked. Pectoral fins somewhat
shorter than head, reaching posteriorly to about vertical through anterior part of soft dorsal fin. Dorsal fin
with X spines and 11 (infrequently 10 or 12) soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin
rays 15 or 16. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales 71 to 74.
Colour: variable: head and body may be steel blue overall with dorsal and anal fins mainly yellow and
other fins white to yellow to dark grey; or back and upper sides purplish brown, blue-grey on sides, silvery
sheen on head and lower sides, edges of preopercle and opercle outlined with black, fins whitish to
yellow-brown; on occasion reported to have brilliant yellow on head.
Habitat, biology, and fisheries: Inhabits reefs at depths between about 1 and 120 m. Occurs solitarily or
in small groups. Feeds mainly on fishes, but also eats crustaceans. Individuals with brilliant yellow on head
may be nuptial males. Caught mainly with handlines or vertical longlines; also speared by divers. Often
seen in markets where it is sold fresh.
the Indo-Pacific from Hawaii
to East Africa and from south-
ern Japan southward to Aus-
tralia; also known from Cocos
Island in the eastern Pacific.
Remarks: The spelling usu-
ally given for the specific
name is furcatus, but in the
original description it is furca.
The word furca is a noun in
apposition and properly re-
tains the original spelling with
whatever generic name it is
combined.
Aphareus rutilans Cuvier, 1830 (Plate V, 28)

FAO names: En - Rusty jobfish; Fr - Vivaneau rouillé; Sp - Pargo bermellón.
together. Lower jaw protruding. Premaxillae not protrusible, fixed by a frenum. Maxilla extending to
below middle of eye or slightly beyond. Teeth in jaws extremely small, minute in large adults; roof of
mouth toothless. Maxilla without scales or longitudinal ridges. Interorbital region flattened. Gill
openings extending far anterior to orbit. First gill arch with 16 to 19 gill rakers on upper limb, 32 to
35 on lower limb (total 49 to 52). Dorsal fin continuous, not deeply incised near junction of spinous and
soft portions. Last soft ray of both dorsal and anal fins well produced, longer than next to last ray.
Caudal fin forked. Pectoral fins somewhat shorter than head. Dorsal fin with X spines and 11
(infrequently 10) soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 15 or 16. Membranes
of dorsal and anal fins without scales. Tubed lateral-line scales 69 to 75. Colour: blue-grey or mauve
to overall reddish; margin of maxilla black; in some specimens, at least, inside of mouth, gill chamber,
and gills shining silver; fins yellowish to reddish except pelvic and anal fins sometimes whitish.
Habitat, biology, and fisheries: Inhabits reefs and rocky bottom areas to depths of at least 250 m. At
Vanuatu (New Hebrides) spawning occurs mainly during spring and summer, with peak activity in
November and December. Caught with handlines and bottom longlines. Locally abundant and an important
market fish in some areas. It is one of the principal species in the Hawaiian offshore handline fishery.
Marketed fresh.
to East Africa and from the
Ryukyu and Ogasawara is-
lands southward to Australia.
2856 Bony Fishes
Aprion virescens Valenciennes, 1830 (Plate V, 29)

FAO names: En - Green jobfish; Fr - Vivaneau job; Sp - Pargo verde.
Diagnostic characters: Body elongate, robust. Nostrils on each side of snout close together. Very distinct
horizontal groove on snout ventral to nostrils. Mouth terminal; jaws about equal. Maxilla extending to
vertical through about anterior margin of orbit or somewhat short of orbit. Teeth in both jaws in bands, with
strong canines anteriorly; teeth on roof of mouth in chevron- or crescent-shaped patch on vomer and in
elongate band on each palatine. Maxilla without scales or longitudinal ridges. Interorbital region
flattened. Preopercular edge smooth, sometimes denticulated in juveniles. First gill arch with 7 or 8 gill
rakers on upper limb, 13 to 16 on lower limb (total 20 to 24). Dorsal fin continuous, not deeply incised
near junction of spinous and soft portions. Last soft ray of both dorsal and anal fins well produced,
longer than next to last ray. Caudal fin forked. Pectoral fins short, less than 1/2 length of head,
about equal to length of snout. Dorsal fin with X spines and 11 soft rays. Anal fin with III spines and
8 soft rays. Pectoral-fin rays 16 to 18. Membranes of dorsal and anal fins without scales. Tubed
lateral-line scales 48 to 50. Colour: dark green, bluish, or grey-blue.
Habitat, biology, and fisheries: Inhabits coastal waters, from the surface down to depths of 120 m. Mainly
a solitary species. Feeds mostly on fishes, but also on shrimps, crabs, and cephalopods. Spawning
recorded in January and November off East Africa. Caught mainly with trolled lures, handlines, bottom
longlines, and bottom trawls. One of the main species in the Hawaiian offshore handline fishery. Significant
in some subsistence fisheries. A fine food fish. Frequently seen in markets where it is sold mostly fresh,
but also dried and salted.
uted in the Indo-Pacific from
Hawaii to East Africa and from
southern Japan southward to
Australia.
Etelis carbunculus Cuvier, 1828 (Plate V, 30)

Frequent synonyms / misidentifications: Etelis marshi (Jenkins, 1903) / Other species of Etelis.
FAO names: En - Ruby snapper; Fr - Vivaneau rubis; Sp - Pargo rubí.
Diagnostic characters: Body relatively elongate, laterally compressed. Nostrils on each side of snout
close together. Lower jaw protruding slightly. Premaxillae protrusible. Maxilla extending to below middle
of eye. Both upper and lower jaws with conical teeth; 1 to several enlarged canines on each side of both
jaws; vomer and palatines with teeth, those on vomer in a chevron-shaped patch. Maxilla with scales,
but without longitudinal ridges. Interorbital region flattened. First gill arch with 5 to 8 gill rakers on
upper limb, 11 to 14 on lower limb (total number, including rudiments, 17 to 22). Dorsal fin
continuous, but spinous portion of fin deeply incised at its junction with soft portion. Last soft ray
of both dorsal and anal fins produced, longer than next to last ray. Caudal fin forked, the lobes
relatively short (about 25 to 30% standard length) compared with other species of Etelis. Pectoral
fins fairly long; in specimens more than about 19 cm standard length, length of pectoral fins about 80 to
90% of head length. Dorsal fin with X spines and 11 soft rays. Anal fin with III spines and 8 soft rays.
Pectoral-fin rays 15 to 17. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales
48 to 50. Colour: mainly pink to red, becoming white on lower sides and belly.
Size: Maximum standard length 80 cm or more, commonly to 50 cm.
Habitat, biology, and fisheries: Inhabits rocky bottoms at depths between about 90 and 400 m. Feeds
on fishes and larger invertebrates such as squids, shrimps, and crabs; also takes planktonic organisms,
including pelagic urochordates. At Vanuatu (New Hebrides) spawning occurs throughout most of the year,
with peak activity during November. Caught mainly with bottom longlines and deep handlines. An important
food fish in some areas, and one of the principal species in the Hawaiian offshore handline fishery.
Marketed fresh or frozen.
Australia; also reported from
off northern New Zealand.
2858 Bony Fishes
Etelis coruscans Valenciennes, 1862 (Plate V, 31)

Frequent synonyms / misidentifications: None / Other species of Etelis.
FAO names: En - Flame snapper; Fr - Vivaneau la flamme; Sp - Pargo de llama.
close together. Lower jaw protruding slightly. Premaxillae protrusible. Maxilla extending to below middle
of eye. Usually one to a few canine or canine-like teeth on each side of both jaws anteriorly, followed by
a series of conical teeth; vomer and palatines with teeth, those on vomer in a chevron-shaped patch.
Maxilla with scales, but without longitudinal ridges. Interorbital region flattened. First gill arch with
8 to 10 gill rakers on upper limb, 15 to 18 on lower limb (total number, including rudiments, 23 to
28). Dorsal fin continuous, but spinous portion of fin deeply incised at its junction with soft portion.
Last soft ray of both dorsal and anal fins produced, longer than next to last ray. Caudal fin deeply
forked, the lobes short to long; upper lobe of caudal fin extremely long in some specimens (in
specimens more than about 13 cm standard length, upper lobe of caudal fin 33 to 75% standard
length). Pectoral fins fairly long; in specimens more than about 18 cm standard length, length of pectoral
fins about 80 to 90% of head length. Dorsal fin with X spines and 11 soft rays. Anal fins with III spines
and 8 soft rays. Pectoral-fin rays 15 or 16. Membranes of dorsal and anal fins without scales. Tubed
lateral-line scales 47 to 50. Colour: back and upper sides deep pink to red; lower sides and belly pink
to white; fins pink to red.
on small fishes, squids, and crustaceans. Caught mainly with deep handlines. An important food fish in
some areas (for example southern Japan). Marketed fresh or frozen.
Australia (Queensland and
New South Wales) and Lord
Howe Island; also known from
off the Kermadec Islands,
northeast of New Zealand.
Remarks: Two morphs are
present in this species: long-
and short-tailed forms. Cau-
dal-fin length may be sexually
dimorphic, but there are no data available that would support or refute this. Both forms have been caught
in the vicinity of Fiji, where the short-tailed form is generally restricted to seamounts and the long-tailed
form, although found on seamounts, occurs more commonly on island slopes.
Etelis radiosus Anderson, 1981

Frequent synonyms / misidentifications: None / Other species of Etelis.
FAO names: En - Scarlet snapper; Fr - Vivaneau écarlate; Sp - Pargo escarlata.
close together. Lower jaw protruding. Premaxillae protrusible. Maxilla extending to vertical near posterior
margin of orbit. Small conical teeth in jaws with a few anterior ones in each jaw enlarged, sometimes into
canine-like teeth; vomer and palatines with teeth, those on vomer in a rounded arch-shaped or chevron-
shaped patch. Maxilla with scales, but without longitudinal ridges. Interorbital region flattened. First
gill arch with 11 to 15 gill rakers on upper limb, 20 to 22 on lower limb (total, including rudiments,
32 to 36). Dorsal fin continuous, but spinous portion of fin deeply incised at its junction with soft
portion. Last soft ray of both dorsal and anal fins produced, longer than next to last ray. Caudal fin
lunate, with a notch at the middle of the distal margin of the fin. Pectoral fins fairly long, a little shorter
than head. Dorsal fin with X spines and 11 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin
rays 16. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales 50 or 51. Colour:
mainly red, lighter on lower sides and belly.
Size: Maximum standard length about 80 cm, commonly to about 50 cm.
mainly on fishes. Caught mainly with deep handlines. Marketed fresh.
Samoa to Sri Lanka and from
the Ryukyu Islands south-
ward to Australia.
2860 Bony Fishes
Lipocheilus carnolabrum (Chan, 1970) (Plate V, 32)

FAO names: En - Tang’s snapper; Fr - Vivaneau verrue; Sp - Pargo trompudo.
Diagnostic characters: Body moderately deep, its depth 38 to 42% standard length, laterally compressed.
Nostrils on each side of snout close to each other and to eye. Upper jaw protruding when mouth closed.
Adults with a thick fleshy protrusion at anterior end of upper lip. Premaxillae protrusible. Maxilla
reaching vertical through middle of eye. Upper and lower jaws both with an outer row of conical teeth (a
few at anterior end of each jaw enlarged into canines, canines of lower jaw frequently well exserted) and
an inner band of villiform teeth; vomer and palatines with teeth, those on vomer in chevron-shaped patch;
no teeth on tongue. Maxilla without scales or longitudinal ridges. Interorbital region somewhat
flattened to slightly convex. First gill arch with 5 to 7 gill rakers on upper limb, 11 to 14 on lower
limb (total 17 to 21). Dorsal fin continuous, not deeply incised near junction of spinous and soft portions.
Spines of dorsal and anal fins strong, very robust in large adults. Last soft ray of dorsal and anal
fins not produced, shorter than next to last ray. Caudal fin moderately forked, lobes not produced
into filaments. Pectoral fins long, reaching vertical through base of second anal-fin spine or beyond.
Dorsal fin with X spines and 10 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 15
or 16. Membranes of dorsal and anal fins without scales except for a few scales basally at posterior
end of each fin. Tubed lateral-line scales 49 to 54. Colour: yellow with a tinge of brown on upper part
of head and anterior part of lips and a silvery sheen on lower side of body; juveniles frequently with about
5 vertical bars.
Size: Maximum standard length more than 50 cm, commonly to 30 cm.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between 90 and 340 m. Probably
feeds on fishes and larger invertebrates. Caught mainly with longlines and handlines; also taken with
bottom trawls. A moderately common constituent of the longline fishery on the continental shelf off South
China.
the Indo-West Pacific from
Vanuatu to the Arabian Sea
and from the Ryukyu Islands
southward to northern
Australia.
Lutjanus adetii (Castelnau, 1873) (Plate V, 33)

FAO names: En - Yellowbanded snapper; Fr - Vivaneau de nuit; Sp - Pargo nachero.
Diagnostic characters: Body relatively deep, its depth 2.5 to 2.7 times in standard length. Snout somewhat
pointed; preorbital bone relatively broad, wider than eye diameter; preopercular notch and knob moderately
well developed; vomerine tooth patch triangular, without a medial posterior extension; tongue with a
patch of granular teeth. First gill arch with 27 to 29 gill rakers, of which 19 or 20 (including rudiments)
on lower limb. Dorsal fin with X spines and 14 soft rays; anal fin with III spines and 8 soft rays; posterior
profile of dorsal and anal fins rounded. Caudal fin emarginate or slightly forked. Pectoral fins with 17 rays.
Scale rows on back rising obliquely above lateral line. Colour: back and upper sides olive-brownish; lower
sides and belly whitish to pink; a prominent yellow to golden brown stripe along middle of sides; eyes
surrounded by yellow or orange.
Size: Maximum total length about 50 cm, commonly to about 30 cm. Matures at about 20 to 30 cm.
Habitat, biology, and fisheries: Mainly a coral reef species, sometimes forming large aggregations
around rocky outcrops during daylight hours. At New Caledonia spawning occurs from August to February
with peak activity during November to January. Mainly of interest to recreational fishermen, but sometimes
appearing in fish markets, particularly at New Caledonia. Good eating and usually marketed fresh. Caught
with hook-and-line; also with gill nets.
Distribution: Restricted to
the east coast of Australia and
New Caledonia. The main
Australian distribution is off
southern Queensland be-
tween Cape Moreton (about
27°S) and the Capricorn
Group (23°S).
2862 Bony Fishes
Lutjanus argentimaculatus (Forsskål, 1775) (Plate V, 34) RES

FAO names: En - Mangrove red snapper; Fr - Vivaneau des mangroves; Sp - Pargo de manglar.
Diagnostic characters: Body moderately deep, its depth 2.5 to 3.1 times in standard length. Snout
somewhat pointed; preorbital bone relatively broad, wider than eye diameter; preopercular notch and knob
poorly developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue with a
patch of granular teeth. First gill arch with 16 to 20 gill rakers, of which 9 to 12 (including rudiments) on
lower limb. Dorsal fin with X spines and 13 or 14 soft rays; anal fin with III spines and 8 soft rays;
posterior profile of dorsal and anal fins rounded. Caudal fin emarginate to nearly truncate. Pectoral fins
with 16 or 17 rays. Scale rows on back more or less parallel to lateral line, or parallel below spinous
part of dorsal fin and sometimes rising obliquely posteriorly, or rarely with entirely oblique rows.
Colour: back and sides greenish brown to reddish; belly silvery or whitish; specimens from deep water
frequently overall reddish; juveniles with a series of about 8 whitish bars crossing sides, and 1 or 2 blue
lines across cheek.
Habitat, biology, and fisheries: Juveniles and young adults found in mangrove estuaries and in the lower
reaches of fresh-water streams. Eventually they migrate offshore to deeper reef areas, sometimes
penetrating to depths in excess of 100 m. Habitat frequently consists of areas of abundant shelter in the
form of caves or overhanging ledges. Feeds mainly on fishes and crustaceans. Spawning occurs
throughout the year, at least in lower latitudes. An important market species throughout the Indo-Pacific
region, but never found in large quantities. Caught mainly with handlines, bottom longlines, and trawls.
Marketed mostly fresh; also dried-salted. From 1990 to 1995, the FAO Yearbook of Fishery Statistics
reports a range of yearly catch of around 4 300 to 12 700 t of Lutjanus argentimaculatus from the Western
Central Pacific.
Samoa and the Line Islands
to East Africa, and from Aus-
tralia northward to the Ryukyu
Islands. Has dispersed into
the eastern Mediterranean
(off Lebanon) via the Suez
Canal, but is not well estab-
lished there.
Lutjanus bengalensis (Bloch, 1790) (Plate VI, 35)

Frequent synonyms/ misidentifications: None / None.
FAO names: En - Bengal snapper; Fr - Vivaneau du Bengale; Sp - Pargo de Bengala.
Diagnostic characters: Body fusiform, slender to moderately deep, its depth 2.5 to 2.9 times in standard
length. Snout somewhat pointed; preorbital bone relatively narrow; its width usually less than eye diameter;
preopercular notch and knob well developed; vomerine tooth patch crescentic, without a medial
posterior extension; tongue smooth, without teeth. First gill arch with 26 to 28 gill rakers, of which 17 to 19
(including rudiments) on lower limb. Dorsal fin with XI or XII spines and 12 to 14 soft rays; anal fin with
III spines and 8 soft rays; posterior profile and anal fins moderately pointed. Caudal fin emarginate. Pectoral
fins with 16 or 17 rays. Scale rows on back rising obliquely above lateral line. Colour: back and sides
yellowish; belly whitish; sides with a series of 4 bright blue stripes; no black spot on back below middle
of dorsal fin; fins yellow to whitish.
Habitat, biology, and fisheries: Mainly a coral reef species found at depths between about 10 and 25 m;
sometimes forming aggregations around rocky outcrops and coral heads during daylight hours. Feeds
mainly on fishes and crustaceans. Found in small quantities in most markets and not considered important,
although the flesh is of good quality. Caught mainly with handlines, traps, and gill nets. Marketed mostly
fresh.
Distribution: East Africa and
the Red Sea to the Molucca
Islands in the Indonesian Ar-
chipelago.
2864 Bony Fishes
Lutjanus biguttatus (Valenciennes, 1830) (Plate VI, 36)

FAO names: En - Twospot banded snapper; Fr - Vivaneau à bande blanche; Sp - Pargo de banda bianca.
Diagnostic characters: Body fusiform, very slender, its depth 3.5 to 3.8 times in standard length.
Snout profile low, sloping very gently; preorbital bone narrow, its width less than 1/2 of eye diameter;
preopercular notch and know poorly developed; vomerine tooth patch triangular, with a medial posterior
extension, or diamond-shaped; tongue smooth without teeth. First gill arch with 23 to 25 gill rakers, of which
16 to 19 (including rudiments) on lower limb. Dorsal fin with XI spines and 12 soft rays; anal fin with III
spines and 8 soft rays; posterior profile of dorsal and anal fins somewhat pointed or angular. Caudal fin
truncate. Pectoral fins with 15 or 16 rays. Scale rows on back rising obliquely above lateral line. Colour:
back and upper sides dark brown; lower sides and belly golden-brown to yellowish; a broad pearly white
stripe from below eye to base of caudal fin; 2 small white spots on upper back, 1 below base of
seventh dorsal-fin spine and 1 below middle of soft part of dorsal fin.
Habitat, biology, and fisheries: Inhabits coral reefs at depths between about 5 and 25 m. Sometimes
occurs in large schools of more than 100 individuals. Feeds mainly on small fishes and crustaceans. An
important market fish in some areas such as Sri Lanka, but generally marketed in small quantities. Caught
mainly with handlines, traps, and gill nets. Marketed mostly fresh.
Distribution: M ai nly the
Indo-Australian Archipelago
from the Solomon Islands to
Sumatra, and from off Cape
York Peninsula, Australia,
northward to the Philippines;
also occurring in the central
and eastern Indian Ocean as
far west as the Maldives.
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Lutjanus bitaeniatus (Valenciennes, 1830)

FAO names: En - Indonesian snapper; Fr - Vivaneau de l’Indonesia; Sp - Pargo de Indonesia.
adult
juvenile
Diagnostic characters: Body relatively deep, its greatest depth 2.3 to 2.5 times in standard length.
Snout somewhat pointed, dorsal profile of head sloping steeply; preorbital bone relatively narrow, its width
usually slightly less than eye diameter; preopercular notch and knob poorly developed; vomerine tooth
patch crescentic, without a medial posterior extension; tongue with a patch of granular teeth. First gill
arch with 18 or 19 gill rakers, of which 11 or 12 (including rudiments) on lower limb. Dorsal fin with X
spines and 13 or 14 soft rays; anal fin with III spines and 8 or 9 soft rays; posterior profile of dorsal and
anal fins rounded. Caudal fin slightly emarginate or truncate. Pectoral fins with 16 or 17 rays. Scale rows
on back rising obliquely above lateral line. Colour: back and upper sides red; lower sides and belly pink
or silvery white; fins reddish except caudal fin sometimes dusky brown; juveniles with a black stripe along
middle of sides and a blackish crescentic marking at base of caudal fin; belly, anal, and pelvic fins of young
frequently yellowish.
Size: Maximum total length of known specimens about 30 cm, but probably grows larger.
Habitat, biology, and fisheries: Inhabits deeper reef areas in depths between about 40 and 65 m. Occurs
solitarily or in small groups. Of potential interest, but presently caught in low numbers, mainly by
experimental trawlers. The flesh is of good quality.
Distribution: Known only
from a few specimens col-
lected at Indonesia (off Suma-
tra and Sulawesi) and west of
Kuri Bay, Western Australia.
2866 Bony Fishes
Lutjanus bohar (Forsskål, 1775) (Plate VI, 37)

FAO names: En - Twospot red snapper; Fr - Vivaneau chin rouge; Sp - Pargo de dos manchas.
Diagnostic characters: Body moderately deep, its greatest depth 2.4 to 2.9 times in standard length.
Snout somewhat pointed, dorsal profile of head rounded; preorbital bone relatively broad; its width usually
greater than eye diameter; a deep groove or pit from nostrils to front of eye; preopercular notch and
knob moderately developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue
with a patch of granular teeth. First gill arch with 22 or 23 gill rakers, of which 16 (including rudiments) on
posterior profile of dorsal and anal fins rounded. Caudal fin slightly emarginate. Pectoral fins with 16 or 17
rays. Scale rows on back rising obliquely above lateral line. Colour: back and upper sides dark brown or
blackish; lower sides and belly reddish, with darker horizontal stripes (between each scale row); fins mainly
dusky except pectoral fins pink with dorsal edge prominently black; young and some adults with 2
silvery white spots on back, 1 below middle of dorsal fin and 1 below base of last soft dorsal-fin rays.
Size: Maximum total length about 75 cm, commonly to 50 cm. Matures at about 50 to 55 cm.
Habitat, biology, and fisheries: Inhabits coral reefs, including sheltered lagoons and outer reefs, usually
at depths between about 10 and 70 m. Usually found solitarily, often adjacent to steep outer reef slopes.
Feeds mainly on fishes, but also takes shrimps, crabs, stomatopods, amphipods, gastropods, and
urochordates. Spawning occurs throughout the year, at least in lower latitudes. Estimated maximum age:
13 years. An important market fish in many localities and also important to subsistence fisheries. However,
large specimens from oceanic areas in the western Pacific are often poisonous (ciguatera). Caught mainly
with handlines and bottom longlines. Marketed mostly fresh; also dried-salted.
the Marquesas and Line is-
lands to East Africa, and from
Australia northward to the
Ryukyu Islands. More com-
mon around oceanic islands
than in continental areas.
Lutjanus boutton (Lacepède, 1803) (Plate VI, 38)

FAO names: En - Moluccan snapper; Fr - Vivaneau des Moluques; Sp - Pargo de Molucas.
somewhat pointed; eye large (3.5 to 4.2 times in head length); preorbital bone narrow, its width much less
than eye diameter; preopercular notch and knob well developed; vomerine tooth patch in a crescen-
tic band without a medial posterior extension; tongue smooth, without teeth. First gill arch with 13 to
15 gill rakers, of which 20 to 23 (including rudiments) on lower limb. Dorsal fin with X spines and 13 or
14 soft rays; anal fin with III spines and 8 soft rays; posterior profile of dorsal and anal fins rounded.
Caudal fin emarginate. Pectoral fins with 16 or 17 rays. Scale rows on back rising obliquely above lateral
line. Colour: head and breast dusky reddish brown; back salmon pink and lower half of side metallic
yellow; some specimens with black spot, slightly smaller than eye, on back at level of junction between
spinous and segmented dorsal-fin rays, its lower edge in contact with lateral line; dorsal and caudal fin
light pink, caudal sometimes with yellow suffusion; anal and pelvic fins yellow; pectoral fins translucent
yellow with small brown spot at base of uppermost rays.
Habitat, biology, and fisheries: Inhabits coral reefs at depths between about 5 and 20 m. It is usually
seen alone or in groups of up to 20 to 30 individuals, which shelter close to the bottom during the day. The
diet includes fishes, shrimps, crabs, other crustaceans, cephalopods, and some planktonic items. Caught
mainly with handlines and bottom longlines. Marketed fresh.
Distribution: The limits of
d i s t r i b u t i o n o f Lutjanus
boutton are uncertain due to
i t s c o n f u s i o n w i t h L.
rufolineatus. It is currently
known from the Philippines,
Sulawesi, Flores, Molucca
Islands, New Guinea, and
New Britain.
2868 Bony Fishes
Lutjanus carponotatus (Richardson, 1842) (Plate VI, 39)

FAO names: En - Spanish flag snapper; Fr - Vivaneau drapeau; Sp - Pargo abanderado.
somewhat pointed, dorsal profile of head steeply sloped; preorbital bone about equal to eye diameter, or
slightly wider; preopercular notch and knob poorly developed; vomerine tooth patch triangular, with
a medial posterior extension, or diamond-shaped; tongue with a patch of granular teeth. First gill
arch with 15 to 18 gill rakers, of which 8 to 11 (including rudiments) on lower limb. Dorsal fin with X spines
and 14 to 16 soft rays; anal fin with III spines and 9 soft rays; posterior profile of dorsal and anal fins
rounded. Caudal fin emarginate. Pectoral fins with 15 to 17 rays. Scale rows on back rising obliquely above
lateral line. Colour: back and upper sides blue-grey or brownish; lower sides and belly white or yellow-
white; a series of 8 or 9 orange, yellow, or golden-brown stripes on sides; fins yellowish; pectoral fin
with a distinct black spot at base of uppermost rays and in axil. Trawl-captured specimens from deeper
water may be pink with yellow stripes.
Habitat, biology, and fisheries: Inhabits coral reefs in both sheltered lagoons and on outer reef slopes
in depths between about 2 and 35 m. Also trawled to depths of about 80 m. Often seen in schools of up to
20 to 30 individuals. Occasionally seen in markets. More important as a subsistence or recreational fish.
Caught mainly with handlines or rod and reel (off Australia); also with traps and gill nets. Marketed fresh.
Distribution: Western Pacific
and northeastern Indian
Ocean, from northern Australia
to India.
Lutjanus decussatus (Cuvier, 1828) (Plate VI, 40)

FAO names: En - Checkered snapper; Fr - Vivaneau damier; Sp - Pargo ajedrezado.
Diagnostic characters: Body moderately deep, its depth 2.6 to 3.1 times in standard length. Dorsal profile
of head moderately sloped; preorbital bone usually about equal to eye diameter or slightly narrower;
preopercular notch and knob poorly developed; vomerine tooth patch crescentic, without a medial
posterior extension; tongue with a patch of granular teeth. First gill arch with 14 to 16 gill rakers, of
which 8 to 10 (including rudiments) on lower limb. Dorsal fin with X spines and 13 or 14 soft rays; anal
fin with III spines and 8 or 9 rays; posterior profile of dorsal and anal fins rounded, with 16 or 17 rays.
Caudal fin emarginate. Scale rows on back rising obliquely above lateral line. Colour: generally whitish
with a “checker-board” pattern on upper half of sides, consisting of dark brown bars and stripes
surrounding rectangular, whitish “windows”; lower half of sides with 2 dark brown stripes; a large black
spot covering most of caudal-fin base.
Habitat, biology, and fisheries: Inhabits coral reefs usually at depths between about 5 and 30 m. Occurs
both solitarily and in schools. A small snapper of little importance to commercial fisheries, regularly seen
in markets and caught by subsistence fishermen, mainly with handlines, traps, and gill nets. Marketed
mostly fresh.
Distribution: Western Pa-
c i f i c a n d e a s t e r n In di a n
Ocean from New Guinea to
Sri Lanka and southern Indian
and northward to the Ryukyu
Islands.
2870 Bony Fishes
Lutjanus dodecacanthoides (Bleeker, 1854) (Plate VI, 41)

FAO names: En - Sunbeam snapper; Fr - Vivaneau soleil; Sp - Pargo solrayo.
of head steeply sloped; eye relatively large; preorbital bone slightly narrower than eye diameter; preoper-
cular notch and knob poorly developed; vomerine tooth patch crescentic, without a medial posterior
extension; tongue smooth without teeth. First gill arch with 22 gill rakers, of which 15 on lower limb. Dorsal
fin with XII spines and 12 or 13 soft rays; anal fin with III spines and 8 soft rays; posterior profile of
dorsal and anal fins rounded. Caudal fin slightly emarginate or truncate. Pectoral fins with 17 rays. Scale
rows on back rising obliquely above lateral line. Colour: back brownish; sides slightly pink with silver sheen,
grading to white on belly and underside of head; series of 6 orange horizontal stripes on sides, the 4
uppermost slanted posteriorly toward dorsal-fin base; brown to golden spot at base of uppermost
pectoral-fin rays, sometimes invading axil.
Size: Maximum total length about 30 cm.
Habitat, biology, and fisheries: Inhabits coral reefs, presumably in shallow water (less than about 30 m).
A rare species, appearing infrequently in markets. Caught mainly with handlines, traps, and gill nets.
from Indonesia and the Philip-
pines.
Lutjanus ehrenbergii (Peters, 1869) (Plate VI, 42)

FAO names: En - Ehrenberg’s snapper; Fr - Vivaneau encrier; Sp - Pargo manchado.
Diagnostic characters: Body moderately deep to somewhat slender, its depth 2.5 to 3 times in standard
length. Dorsal profile of head moderately sloped; preorbital bone very narrow, its width about 1/2 of eye
diameter; preopercular notch and knob poorly developed; vomerine tooth patch triangular, with a medial
posterior extension; tongue with a patch of granular teeth. First gill arch with 16 to 21 gill rakers, of which
15 (including rudiments) on lower limb. Dorsal fin with X spines and 13 or 14 soft rays; anal fin with III
spines and 8 or 9 soft rays; posterior profile of dorsal and anal fins angular or somewhat rounded. Caudal
fin truncate or slightly emarginate. Pectoral fins with 15 or 16 rays. Scale rows on back parallel to lateral
line. Colour: back and upper sides dark brown; lower sides and belly whitish with a silver sheen; often a
series of 4 or 5 narrow yellow stripes on sides below lateral line; a prominent round, black spot on back
below posterior part of spinous portion of dorsal fin.
Size: Maximum total length of about 35 cm, commonly to 20 cm. Matures at about 12 cm.
Habitat, biology, and fisheries: Inhabits coral reefs at depths between about 5 and 20 m. Juveniles
frequent inshore areas over sand, silt, or coral rubble bottoms, occasionally in mangrove-lined streams
and estuaries. A small species, commonly utilized in subsistence fisheries and also seen in markets.
Caught mainly with handlines, traps, and gill nets. Marketed mostly fresh.
Solomon and Mariana islands
to East Africa.
2872 Bony Fishes
Lutjanus erythropterus (Bloch, 1790)

FAO names: En - Crimson snapper; Fr - Vivaneau cramoisi; Sp - Pargo carmesi.
Diagnostic characters: Body moderately deep, its depth 2.5 to 3 times in standard length. Dorsal profile
of head sloped; mouth relatively small, length of upper jaw smaller than distance between base of
last dorsal- and anal-fin rays; preorbital bone usually narrower than eye diameter; preopercular notch
and knob poorly developed; vomerine tooth patch crescentic or triangular, without a medial posterior
extension; tongue smooth, without teeth. First gill arch with 18 or 19 gill rakers, of which 13 or 14 (including
rudiments) on lower limb. Dorsal fin with X spines and 12 to 14 soft rays; anal fin with III spines and 8
or 9 soft rays; posterior profile of dorsal and anal fins rounded. Caudal fin emarginate. Pectoral fins with
16 or 17 rays. Scale rows on back rising obliquely above lateral line. Colour: overall pink or red, including
fins; juveniles with a broad, oblique, black band extending from mouth to beginning of dorsal fin and a
large black spot at base of caudal fin; frequently with narrow red stripes.
Habitat, biology, and fisheries: Inhabits trawling grounds and reefs to depths of at least 100 m. An
excellent food fish, appearing regularly in markets, but usually in small quantities. Caught mainly with
handlines and bottom trawls. Marketed fresh or dried-salted.
Australia and New Guinea to
the Gulf of Oman, and north-
ward to southern Japan.
Lutjanus fulviflamma (Forsskål, 1775) (Plate VII, 43)

FAO names: En - Blackspot snapper; Fr - Vivaneau gibelot; Sp - Pargo tintero.
Diagnostic characters: Body moderately deep to somewhat slender, its depth 2.6 to 2.9 times in standard
length. Dorsal profile of head moderately sloped; preorbital bone about equal in width to eye diameter or
slightly less; preopercular notch and knob poorly developed; vomerine tooth patch triangular, with a medial
posterior extension or diamond-shaped; tongue with a patch of granular teeth. First gill arch with 16 to 19
gill rakers, of which 9 to 12 (including rudiments) on lower limb. Dorsal fin with X spines and 12 to 14
soft rays; anal fin with III spines and 8 soft rays; posterior profile of dorsal and anal fins rounded to
somewhat angular. Caudal fin truncate or slightly emarginate. Pectoral fins with 15 to 17 rays. Scale rows
on back rising obliquely above lateral line. Colour: back and upper sides brown; lower sides whitish or
light brown; whitish to yellow on belly; a series of 6 or 7 yellow stripes on sides; a prominent black
spot at level of lateral line below base of anterior part of soft portion of dorsal fin; fins yellowish.
Habitat, biology, and fisheries: Inhabits coral reefs at depths between 3 and 35 m. Juveniles sometimes
found in brackish water or mangrove estuaries or in the lower reaches of fresh-water streams. Feeds mainly
on fishes, shrimps, crabs, and other crustaceans. At New Caledonia and East Africa spawning occurs
mainly from August to March (spring and summer). A small species, commonly utilized in subsistence
fisheries and also seen frequently in markets, usually fresh. Caught mainly with handlines, traps, and gill
nets.
the Indo-Pacific from Samoa
to East Africa, and from Aus-
tralia northward to the Ryukyu
Islands.
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2874 Bony Fishes
Lutjanus fulvus (Schneider, 1801) (Plate VII, 44)

FAO names: En - Blacktail snapper; Fr - Vivaneau queue noire; Sp - Pargo rabo negro.
Diagnostic characters: Dorsal profile of head steeply sloped; preorbital bone about equal in width to eye
diameter; preopercular notch and knob well developed; vomerine tooth patch crescentic, without a
medial posterior extension; tongue smooth, without teeth. First gill arch with 16 to 20 gill rakers, of which
10 to 13 (including rudiments) on lower limb. Dorsal fin with X spines and 14 soft rays; anal fin with III
spines and 8 soft rays; posterior profile of dorsal and anal fins rounded. Caudal fin slightly emarginate.
Pectoral fins with 16 rays. Scale rows on back rising obliquely above lateral line. Colour: back and sides
grey to brown; often with a series of narrow yellow of golden-brown stripes, 1 per scale rows, on sides;
belly and underside of head whitish; dorsal fin brown to reddish, with a narrow blackish band near margin,
broader on soft part of dorsal fin; caudal fin blackish, dorsal and caudal fins with a narrow white
border; pelvic and anal fins yellowish.
Habitat, biology, and fisheries: Inhabits coral reefs in lagoons and on outer reef slopes in depths of about
2 to 40 m. Juveniles sometimes found in shallow mangrove swamps and the lower parts of fresh-water
streams. Feeds at night on fishes, shrimps, crabs, holoturians, and cephalopods. Spawning occurs
throughout the year, at least in lower latitudes. Commonly seen in markets, usually fresh. Caught with
handlines, traps, and gill nets. Sometimes causes ciguatera poisoning, particularly in the Pacific portion
of its range.
the Indo-Pacific from the Mar-
quesas and Line islands to
East Africa, and from Austra-
lia to southern Japan. Intro-
duced to Hawaii.
Lutjanus fuscescens (Valenciennes, 1830)

FAO names: En - Freshwater snapper; Fr - Vivaneau d’eau douce; Sp - Pargo de agua dulce.
adult
juvenile
Diagnostic characters: Dorsal profile of head moderately sloped; preorbital width about equal to, or
slightly narrower than eye diameter; preopercular notch and knob poorly developed; vomerine tooth
patch crescentic, without a medial posterior extension; tongue smooth, without teeth. First gill arch
with 18 gill rakers, of which 12 (including rudiments) on lower limb. Dorsal fin with X spines and 13 to
Caudal fin truncate or slightly emarginate. Pectoral fins with 16 or 17 rays. Scale rows on back rising parallel
to lateral line. Colour: back and sides greyish brown or olive, whitish on belly; a prominent black spot
on back, bisected by lateral line, below anterior soft dorsal-fin rays; juveniles with a series of diffuse
blackish bars on sides.
Habitat, biology, and fisheries: Inhabits fresh-water streams and brackish, mangrove-lined estuaries.
Not reported from purely marine habitats. Apparently rare and of little interest to fisheries.
from the Solomon Islands,
New Guinea, Indonesia, the
Philippines, and China.
2876 Bony Fishes
Lutjanus gibbus (Forsskål, 1775) (Plate VII, 45)

FAO names: En - Humpback red snapper; Fr - Vivaneau pagaie; Sp - Pargo jorobado.
Diagnostic characters: Body relatively deep, its depth 2.2 to 2.5 times in standard length. Dorsal profile
of head steeply sloped; preorbital bone broad, much wider than eye diameter; preopercular notch and
knob well developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue
smooth, without teeth. First gill arch with 25 to 30 gill rakers, of which 15 to 20 (including rudiments) on
posterior profile of dorsal and anal fins pointed. Caudal fin distinctly forked with rounded lobes. Pectoral
fins with 16 or 17 rays. Scale rows on back rising obliquely both above and below the lateral line. Colour:
red or grey, darker on back and upper portion of head; an orange hue on lower part of opercle and in
pectoral fin axil; fins red or frequently dark brown to blackish; soft part of dorsal fin, anal and caudal fins
with a narrow white margin; juveniles with a large round black spot at base of caudal fin.
Size: Maximum total length about 50 cm, commonly to 35 cm. Matures at about 30 cm.
Habitat, biology, and fisheries: Mainly inhabits coral reefs, sometimes forming large aggregations, which
are mostly stationary during daylight hours. Normal depth distribution ranges from about 6 to at least 30 m.
Feeds on fishes and a variety of invertebrates including shrimps, crabs, lobsters, stomatopods, cephalo-
pods, echinoderms, and ophiuroids. Off East Africa spawning occurs mainly during spring and summer.
Caught mainly with handlines, traps, and gill nets. Commonly seen in markets, usually offered fresh.
Sometimes causes ciguatera poisoning, particularly around the Pacific islands.
the Society and Line islands
to East Africa, and Australia to
southern Japan.
Lutjanus goldiei (Macleay, 1882)

FAO names: En - Papuan black snapper; Fr - Vivaneau de Papua; Sp - Pargo de Papua.
adult
juveniles
Diagnostic characters: Body relatively deep, its depth 2.2 to 2.8 times in standard length. Dorsal
profile of head steeply sloped; preorbital bone relatively broad, usually much greater than eye diameter;
preopercular notch and knob poorly developed; vomerine tooth patch crescentic, without a medial posterior
extension; tongue with a patch of granular teeth. First gill arch with 14 or 15 gill rakers, of which 9 (including
rudiments) on lower limb. Dorsal fin with X spines and 13 or 14 soft rays; anal fin with III spines and 8
soft rays; posterior profile of dorsal and anal fins rounded. Caudal fin truncate. Pectoral fins with 16 or 17
rays. Scale rows on back parallel to lateral line anteriorly. Colour: back and upper sides dark brown
to charcoal grey, lower sides and belly golden brown to whitish; there are also distinct colour phases with
the entire body blackish or with a series of 6 or 7 broad greyish bars on sides; fins dusky brown to blackish,
sometimes with a yellow hue.
Habitat, biology, and fisheries: Inhabits large fresh-water streams. Not reliably reported from marine
habitats but may occur in brackish estuaries. Occasionally seen in the fish market at Port Moresby. A
popular sport fish forming the basis of a safari-angling industry. Caught mainly with handlines, rod and
reel, traps, and gill nets; also with spears.
from southern Papua New
Guinea between the Port
Moresby district and the Fly
River.
2878 Bony Fishes
Lutjanus johnii (Bloch, 1792) (Plate VII, 46)

FAO names: En - John’s snapper; Fr - Vivaneau ziebelo; Sp - Pargo jaspeado.
of head steeply sloped; preorbital width equal to eye diameter or larger (4.9 to 6 times in head length);
posterior extension; tongue with a patch of granular teeth. First gill arch with 17 or 18 gill rakers, of which
11 (including rudiments) on lower limb. Dorsal fin with X spines and 13 or 14 soft rays; anal fin with III
spines and 8 soft rays; posterior part of dorsal and anal fins rounded. Caudal fin truncate or slightly
emarginate. Pectoral fins with 16 or 17 rays. Scale rows on back parallel to lateral line. Colour: generally
yellow with a bronze to silvery sheen, grading to silvery white on belly and underside of head; centre of
each scale often with a reddish brown spot, giving an overall appearance of series of horizontal lines
on side of body; a round black spot, larger than eye, on back, mainly above lateral line, below anterior soft
dorsal-fin rays (sometimes absent in large adults).
Habitat, biology, and fisheries: Little information on habitat of adults, although they probably frequent
coral reef areas. Juveniles in brackish mangrove estuaries. Large adults trawled to depths of 80 m. Feeds
on fishes and benthic invertebrates including shrimps, crabs, and cephalopods. Spawning has been
reported during September in the Andaman Sea. Frequently found in markets. Caught mainly with
handlines, bottom longlines, traps, and bottom trawls. Excellent quality flesh, usually marketed fresh or
sometimes dried-salted.
the Fiji Islands to East Africa
and from Australia to the
Ryukyu Islands.
Lutjanus kasmira (Forsskål, 1775) (Plate VII, 47)

FAO names: En - Common bluestripe snapper; Fr - Vivaneau à raies bleues; Sp - Pargo de rayas azules.
of head steeply sloped; preorbital width usually greater than eye diameter, but sometimes less in small
specimens; preopercular notch and knob well developed; vomerine tooth patch crescentic, without a
medial posterior extension; tongue smooth, without teeth. First gill arch with 20 to 22 gill rakers, of which
13 or 14 (including rudiments) on lower limb. Dorsal fin with X spines and 14 or 15 soft rays; anal fin
with III spines and 7 or 8 soft rays; posterior profile of dorsal and anal fins somewhat angular. Caudal fin
slightly emarginate. Pectoral fins with 15 or 16 rays. Scale rows on back rising obliquely above lateral line.
Colour: back and sides bright yellow; lower sides and underside of head white; 4 bright blue stripes on
side; several faint greyish stripes on lowermost part of sides; fins yellow.
Habitat, biology, and fisheries: Inhabits coral reefs, occurring in both shallow lagoons and on outer reef
slopes to depths of at least 60 m, but occurring in 180 and 265 m at the Marquesas Islands and Red Sea
respectively. Frequently found in large aggregations around coral formations, caves, or wrecks during
daylight hours. Feeds on fishes, shrimps, crabs, stomatopods, cephalopods, and planktonic crustaceans.
Spawning occurs throughout most of the year in lower latitudes with peak activity reported for November
and December in the Andaman Sea. Eggs measure from 0.78 to 0.85 mm in diameter and hatch in about
18 hours at 22° to 25°C. Frequently found in markets. It is one of the principal species in the Hawaiian
offshore handline fishery, but commands a relatively low price. Caught mainly with handlines, traps, and
gill nets. Usually offered fresh.
lia to southern Japan.
2880 Bony Fishes
Lutjanus lemniscatus (Valenciennes, 1828) (Plate VII, 48)

FAO names: En - Yellowstreaked snapper; Fr - Vivaneau à raies jaunes; Sp - Pargo de rayas amarillas.
of head steeply sloped; snout profile slightly concave; preorbital bone usually much greater than eye
diameter, at least in larger specimens; preopercular notch and knob poorly developed; vomerine tooth
patch crescentic, without a medial posterior extension; tongue with a patch of granular teeth. First gill
arch with 18 to 21 gill rakers, of which 10 to 14 (including rudiments) on lower limb. Dorsal fin with X
spines and 13 or 14 soft rays; anal fin with III spines and 8 soft rays; posterior profile of dorsal and anal
fins rounded. Caudal fin truncate or slightly emarginate. Pectoral fins with 16 rays. Scale rows on back
rising obliquely above lateral line. Colour: back and upper sides grey-brown or olive; sides grey, brown
or reddish; belly and underside of head whitish; dorsal and caudal fins brownish, remaining fins pink or
reddish; juveniles with a broad black horizontal band from snout tip to caudal-fin base.
Habitat, biology, and fisheries: Inhabits offshore reefs to depths of 70 to 80 m. Juveniles sometimes
encountered in the vicinity of coral reefs, often located close to the shore where silting is moderate and
visibility reduced. Feeds on fishes and a variety of benthic invertebrates. An important market species in
many localities. Caught mainly with handlines, traps, and occasionally with bottom trawls. Usually offered
fresh, sometimes dried-salted.
Distribution: Indo-West Pa-
cific from Australia and New
Guinea to Sri Lanka and
southern India, and north-
ward to the Philippines.
Lutjanus lunulatus (Park, 1797) (Plate VII, 49)

FAO names: En - Lunartail snapper; Fr - Vivaneau queue lune; Sp - Pargo raboluna.
of head steeply sloped; preorbital width about equal to eye diameter; preopercular notch and knob poorly
developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue with a
patch of granular teeth. First gill arch with 15 to 18 gill rakers, of which 8 to 10 (including rudiments) on
lower limb. Dorsal fin with X spines and 13 or 14 soft rays; anal fin with III spines and 8 or 9 soft rays;
posterior profile of dorsal and anal fins somewhat angular. Caudal fin slightly emarginate. Pectoral fins
with 16 or 17 rays. Scale rows on back rising obliquely above lateral line. Colour: back and upper side
brown to reddish pink; reddish pink to mauve on middle of side; rear half of maxilla, lower part of preopercle,
breast and abdomen creamy yellow; dorsal fin reddish pink to clear; caudal fin with a broad, crescentic
black band with the posterior portion of fin pink; pectoral, anal, and pelvic fins yellow.
Habitat, biology, and fisheries: Inhabits coral reefs at depths between about 10 and 30 m. Usually seen
solitarily or in small groups. Infrequently seen in markets. Caught mainly with handlines, traps, and gill
nets. Usually offered fresh.
and northern Indian Ocean from
Vanuatu and the Philippines to
the northeastern Arabian Sea.
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2882 Bony Fishes
Lutjanus lutjanus (Bloch, 1790) (Plate VII, 50)

Frequent synonyms / misidentifications: Lutjanus lineolatus (Rüppell, 1829) / None.
FAO names: En - Bigeye snapper; Fr - Vivaneau gros yeux; Sp - Pargo de Madras.
Diagnostic characters: Body fusiform, slender, its depth 2.9 to 3.3 times in standard length. Dorsal
profile of head gently sloped; preorbital bone very narrow, much less than eye diameter; preopercular
notch and knob poorly developed; vomerine tooth patch triangular, with a medial posterior extension;
tongue with a patch of granular teeth. First gill arch with 24 to 26 gill rakers, of which 17 to 19 (including
rudiments) on lower limb. Dorsal fin with X to XII spines and 12 soft rays; anal fin with III spines and
8 soft rays; posterior profile of dorsal and anal fins angular. Caudal fin truncate or slightly emarginate.
Pectoral fins with 16 or 17 rays. Scale rows on back rising obliquely above lateral line. Colour: upper back
golden brown; sides silvery white; a broad yellow to brownish stripe from eye to caudal-fin base; a series
of yellow horizontal lines (1 per scale row) on lower half of body, and similar lines running obliquely above
lateral line; fins pale yellow to whitish.
Habitat, biology, and fisheries: Inhabits offshore coral reefs and trawling grounds to depths of at least
90 m. Frequently seen in large schools of more than 100 individuals. Feeds on fishes and crustaceans.
Spawning has been reported during March and November in the Gulf of Aden and off East Africa
respectively, and from January to June in the Gulf of Suez. Estimated maximum age at least 11 years.
One of the more common snappers found in the markets. Forms about 10 to 20% of the trawl catch in the
Gulf of Suez. Caught mainly with handlines and with bottom trawls in some areas. Usually offered fresh.
the Indo-West Pacific from the
Solomon Islands to East Africa
and from Australia to southern
Japan.
Lutjanus madras (Valenciennes, 1831) (Plate VIII, 51)

Frequent synonyms / misidentifications: None / Lutjanus lutjanus (Bloch, 1790).
FAO names: En - Indian snapper; Fr - Vivaneau madras; Sp - Pargo madraseño.
Diagnostic characters: Body fusiform, somewhat slender, its depth 2.6 to 3.1 times in standard
length. Dorsal profile of head moderately to gently sloped; preorbital width about equal to 2/3 of eye
diameter; preopercular notch and knob poorly developed; vomerine tooth patch triangular with a medial
posterior extension or diamond-shaped; tongue with a patch of granular teeth. First gill arch with 18 to 21
gill rakers, of which 12 to 15 (including rudiments) on lower limb. Dorsal fin with X spines and 13 (rarely
11 or 14) soft rays; anal fin with III spines and 8 (rarely 9) soft rays; posterior profile of dorsal and anal
fins angular. Caudal fin truncate or slightly emarginate. Pectoral fins with 16 or 17 rays. Scale rows on
back rising obliquely above lateral line. Colour: upper back brownish; sides whitish with a series of
fine yellow horizontal lines, 1 per scale row, on lower half of body, and similar brownish lines
running obliquely above lateral line; fins yellow except pelvic fins frequently white or faintly yellow.
Habitat, biology, and fisheries: Inhabits coral and rocky reefs at depths between about 5 and 90 m.
Common in some markets, for example at Sri Lanka. Caught mainly with handlines, traps, and gill nets.
Usually offered fresh.
and Indian Ocean from New
Guinea and the Philippines to
the Laccadive Islands; also
found in the Seychelles.
2884 Bony Fishes
Lutjanus malabaricus Schneider, 1801 (Plate VIII, 52)

FAO names: En - Malabar blood snapper; Fr - Vivaneau malabar; Sp - Pargo malabárico.
profile of head steeply sloped; mouth large, maxilla length about equal to distance between bases of
last dorsal- and anal-fin rays; snout profile straight or slightly concave; preorbital bone much broader
than eye diameter; preopercular notch and knob poorly developed; vomerine tooth patch crescentic or
triangular, without a medial posterior extension; tongue smooth, without teeth. First gill arch with 18 to 20
gill rakers, of which 12 to 14 (including rudiments) on lower limb. Dorsal fin with XI spines and 12 to 14
soft rays; anal fin with III spines and 8 or 9 soft rays; posterior profile of dorsal and anal fins slightly
rounded to angular. Caudal fin truncate. Pectoral fins with 16 or 17 rays. Scale rows on back rising obliquely
above lateral line. Colour: back and sides red or red-orange, lighter on lower parts; fins reddish;
juveniles with a broad, oblique band of brown or black from upper jaw to beginning of dorsal fin, and a
prominent black band across caudal peduncle with a pearly white anterior border; young also with a series
of narrow reddish horizontal lines (sometimes absent) on sides.
Habitat, biology, and fisheries: Inhabits both coastal and offshore reefs. In Australia it frequently forms
mixed shoals with Lutjanus erythropterus. Depth range from about 12 to 100 m. Feeds mainly on fishes
and benthic crustaceans. Spawning occurs throughout most of the year in lower latitudes with peak activity
during spring and summer at New Caledonia. The von Bertalanffy growth coefficient (K) and exponential
rate of mortality (M) were estimated as 0.31 and 0.42 respectively for the Vanuatu (New Hebrides)
population (Brouard and Grandperrin, 1984). Estimated maximum age 10 to 12 years. An excellent food
fish, found regularly in markets but usually in small quantities. One of the principal market fishes of Kuwait
and other Persian Gulf ports. Also constitutes about 5% of trawl catch on the Northwest Shelf of Australia
in combination with L. erythropterus and L. sebae (1971 to 1976 data). Caught mainly with handlines,
bottom longlines, and bottom
trawls. Marketed fresh or
dried-salted.
the Indo-West Pacific from the
Fiji Islands to the Arabian Sea
and Persian Gulf, and from
Australia to southern Japan.
Lutjanus maxweberi Popta, 1921

FAO names: En - Pygmy snapper; Fr - Vivaneau nain; Sp - Pargo enano.
Diagnostic characters: Body relatively deep, its depth 2.2 to 2.7 times in standard length. Dorsal profile
of head steeply sloped; preorbital bone narrow, its width slightly less than eye diameter; preopercular
notch and knob poorly developed; vomerine tooth patch crescentic or triangular, without a medial
posterior extension; tongue smooth, without teeth. First gill arch with 16 to 19 gill rakers, of which 10 to 12
(including rudiments) on lower limb. Dorsal fin with X spines and 13 soft rays; anal fin with III spines
and 8 soft rays; posterior profile of dorsal and anal fins rounded to slightly angular. Caudal fin emarginate.
Pectoral fins with 16 rays. Scale rows on back parallel to lateral line. Colour: whitish with 4 dark brown
to blackish stipes on sides; older specimens becoming overall brownish except whitish on ventral
surface.
Size: Maximum total length of known specimens 15 cm, but adults unknown.
Habitat, biology, and fisheries: Apparently restricted to fresh-water streams and brackish estuaries,
although adult specimens are unknown. A poorly known species, of little interest to fisheries.
Distribution: Philippines, In-
donesia (Sulawesi), and New
Guinea.
2886 Bony Fishes
Lutjanus mizenkoi Allen and Talbot, 1985

FAO names: En - Samoan snapper; Fr - Vivaneau de Samoa; Sp - Pargo de Samoa.
Diagnostic characters: Body relatively slender, its depth 2.9 to 3 times in standard length). Dorsal profile
of head gently sloped; preorbital bone narrower than eye diameter; preopercular notch and knob
poorly developed; vomerine tooth patch diamond-shaped; tongue with granular teeth. First gill arch with
15 or 16 gill rakers, of which 9 or 10 (including rudiments) on lower limb. Dorsal fin with X spines and
Caudal fin truncate or slightly emarginate. Pectoral fins with 16 rays. Scale rows on back rising obliquely
above lateral line. Colour: back and sides reddish grading to pink; ventral portion of head and body
white; a series of faint, yellow horizontal lines, 1 per scale row on sides; fins whitish or yellow.
Habitat, biology, and fisheries: Inhabits offshore reefs at depths between about 100 and 150 m. Of
potential interest to fisheries if sufficient stocks can be located. A small, but excellent food fish occasionally
found in markets in Western Samoa; caught mainly with deep handlines. Usually offered fresh.
fro m Samoa, Papua New
Guinea, and Indonesia (Su-
lawesi).
Lutjanus monostigma (Cuvier, 1828) (Plate VIII, 53)

FAO names: En - Onespot snapper; Fr - Vivaneau eglefin; Sp - Pargo eglefino.
Diagnostic characters: Body moderately deep to somewhat slender, its depth 2.6 to 3 times in standard
length. Dorsal profile of head gently to moderately sloped; preorbital width about equal to eye diameter;
posterior extension; tongue smooth, without teeth. First gill arch with 18 or 19 gill rakers, of which 10 or
11 (including rudiments) on lower limb. Dorsal fin with X spines and 13 (rarely 14) soft rays; anal fin
with III spines and 8 or 9 soft rays; posterior profile of dorsal and anal fins rounded to somewhat angular.
Caudal fin truncate or slightly emarginate. Pectoral fins with 15 to 17 rays. Scale rows on back rising
obliquely above lateral line. Colour: generally yellowish to pinkish with dusky scale margins; grey or
brown on upper back and dorsal portion of head; a black spot, sometimes faint or absent, on back
below anterior soft dorsal-fin rays; fins yellowish.
Habitat, biology, and fisheries: Inhabits coral reef areas, usually close to shelter in the form of caves,
large coral formations, and wreckage. Solitary in habit or occurring in small groups. Depths range from
about 5 to 30 m. Feeds mainly on fishes and benthic crustaceans. Spawning has been reported during
February and November off East Africa. An important market fish in many areas, particularly in Oceania,
although the flesh is sometimes poisonous (ciguatera). Caught mainly with handlines, traps, and gill nets.
Marketed mostly fresh.
lia to the Ryukyu Islands.
2888 Bony Fishes
Lutjanus quinquelineatus (Bloch, 1790) (Plate VIII, 54)

FAO names: En - Fivelined snapper; Fr - Vivaneau à cinq bandes; Sp - Pargo de cinco lineas.
of head steeply sloped; preorbital width usually less than eye diameter; preopercular notch and knob
well developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue smooth,
without teeth. First gill arch with 20 to 23 gill rakers, of which 13 to 15 (including rudiments) on lower limb.
Dorsal fin with X spines and 13 to 15 soft rays; anal fin with III spines and 8 soft rays; posterior profile
of dorsal and anal fins rounded to somewhat angular. Caudal fin truncate or slightly emarginate. Pectoral
fins with 16 or 17 rays. Scale rows on back rising obliquely above lateral line. Colour: upper part of head
brownish; sides and belly bright yellow; a series of 5 bright blue stripes on sides; a round black spot,
eye-sized or larger, below anteriormost soft dorsal-fin rays at level of lateral line; fins yellow.
Habitat, biology, and fisheries: Inhabits sheltered lagoons and exposed, outer slope coral reefs at depths
ranging from about 2 and 40 m. Frequently encountered in large aggregations including 100 or more
individuals. At New Caledonia spawning occurs over most of the year with peak activity during summer
(November to January). A good eating fish commonly found in markets throughout its range, also important
in artisanal fisheries. Caught mainly with handlines, traps, and gill nets; also sought by sport fishermen off
the Queensland coast of Australia.
the Indo-Pacific from the Fiji
Islands to the Gulf of Oman
and the Persian Gulf, and
from Australia to southern
Japan.
Lutjanus rivulatus (Cuvier, 1828) (Plate VIII, 55)

FAO names: En - Blubberlip snapper; Fr - Vivaneau maori; Sp - Pargo maori.
Diagnostic characters: Body very deep, its depth 2 to 2.4 times in standard length. Dorsal profile of head
steeply sloped; preorbital bone broad, about twice the eye diameter; preopercular notch and knob
moderately developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue
smooth, without teeth. First gill arch with 17 or 18 gill rakers, of which 11 or 12 (including rudiments) on
lower limb. Lips greatly thickened in adults. Dorsal fin with X spines and 15 or 16 soft rays; anal fin
with III spines and 8 soft rays; posterior profile of dorsal fin rounded; posterior profile of anal fin distinctly
pointed. Caudal fin truncate or slightly emarginate. Pectoral fins with 17 rays. Scale rows on back rising
obliquely above lateral line. Colour: generally brown with a reddish tinge; each scale on side with a pale
brown border and 2 to 3 small bluish white spots in central portion; head with numerous undulating blue
lines; lips tan; fins largely yellowish to dusky grey-brown. Juveniles with a series of 3 to 8 brown bars on
sides and a chalky white spot, with a broad blackish margin, present below anterior soft dorsal-fin rays at
level of lateral line.
Habitat, biology, and fisheries: Occasionally encountered on coral reefs or shallow inshore flats. Also
occurs in deeper water to at least 100 m. Found solitarily or in small groups of up to 15 to 20 fish. Feeds
on fishes, cephalopods and benthic crustaceans. An excellent food fish frequently found in markets and
important to artisanal fisheries. Caught with handlines, traps, and gill nets, occasionally trawled. Marketed
mostly fresh.
the Indo-Pacific from Tahiti to
lia to southern Japan.
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2890 Bony Fishes
Lutjanus rufolineatus (Valenciennes, 1830)

Frequent synonyms / misidentifications: None / Lutjanus boutton (Lacepède, 1803).
FAO names: En - Yellowlined snapper.
(from Bleeker, 1873-76)
somewhat pointed; eye large (3.4 to 4.2 times in head length); preorbital bone narrow, its width much less
than eye diameter; preopercular notch and knob well developed; vomerine tooth patch in a crescentic band
without a medial posterior extension; tongue smooth, without teeth. First gill arch with 20 to 23 gill rakers,
of which 13 to 15 (including rudiments) on lower limb. Dorsal fin with XI spines and 13 or 14 soft rays;
anal fin with III spines and 8 soft rays; posterior profile of dorsal and anal fins rounded. Caudal fin
emarginate. Pectoral fins with 16 or 17 rays. Scale rows on back rising obliquely above lateral line. Colour:
generally pink or reddish, white or silvery white ventrally; a series of about 10 to 12 faint yellow stripes
on side; some specimens with a black spot, eye size or smaller, below anterior part of soft dorsal
fin at level of lateral line; spinous dorsal fin whitish with a yellow margin; remaining fins yellowish, although
pelvic fins sometimes white; axil of pectoral fins brown on dorsal portion.
Habitat, biology, and fisheries: Inhabits coral reefs at depths between about 15 and 50 m. Usually seen
in schools of up to 100 or more individuals, frequently well above the bottom. The diet includes fishes,
shrimps, crabs, other crustaceans, cephalopods, and some planktonic items. At Samoa, spawning occurs
throughout the year with peak activity from March to June. Occasionally seen in markets. An important
subsistence fish in many areas. Caught mainly with handlines and bottom longlines. Marketed fresh.
Ocean from Samoa to Sumatra
and northward to southern Ja-
pan; also in the eastern Indian
Ocean off northwestern Austra-
lia.
Lutjanus russelli (Bleeker, 1849) (Plate VIII, 56)

FAO names: En - Russell’s snapper; Fr - Vivaneau hublot; Sp - Pargo ojo de buey.
Diagnostic characters: Body moderately deep to somewhat slender, its depth 2.6 to 2.8 times in standard
length. Dorsal profile of head steeply to moderately sloped; preorbital width about equal to, or slightly less
than eye diameter; preopercular notch and knob poorly developed; vomerine tooth patch triangular,
with a medial posterior extension or diamond-shaped; tongue with a patch of granular teeth. First gill
arch with 13 to 18 gill rakers, of which 7 to 11 (including rudiments) on lower limb. Dorsal fin with X spines
and 14 soft rays; anal fin with III spines and 8 soft rays; posterior profile of dorsal and anal fins rounded
(anal fin sometimes angular). Caudal fin truncate or slightly emarginate. Pectoral fins with 16 or 17 rays.
Scale rows on back rising obliquely above lateral line. Colour: back and upper side brownish; lower
sides and belly pink to whitish with a silvery sheen; a black spot, mainly above lateral line, below anterior
rays of soft dorsal fin; adults from the Indian Ocean usually with 7 or 8 narrow golden brown stripes on
sides; juveniles from the western Pacific whitish, with 4 black stripes on sides and with a round black spot
on upper back.
Habitat, biology, and fisheries: Inhabits offshore coral reefs and also inshore rock and coral reefs. Large
adults are sometimes trawled well offshore in depths to about 70 to 80 m. Juveniles frequent brackish
mangrove estuaries and lower reaches of fresh-water streams. A common market species throughout its
range; also a component of artisanal fisheries. Caught with handlines, traps, and bottom trawls. Marketed
mostly fresh.
the Fiji Islands to East Africa,
and from Australia to south-
ern Japan.
2892 Bony Fishes
Lutjanus sebae (Cuvier, 1828)

FAO names: En - Emperor red snapper; Fr - Vivaneau bourgeois; Sp - Pargo imperial.
Diagnostic characters: Body very deep, its depth 2.1 to 2.4 times in standard length. Dorsal profile
of head steeply sloped; snout profile straight or slightly convex; preorbital bone broad, much wider than
eye diameter; preopercular notch and knob moderately developed; vomerine tooth patch crescentic or
triangular, without a medial posterior extension; tongue smooth, without teeth. First gill arch with 16 to 19
gill rakers, of which 10 to 12 (including rudiments) on lower limb. Dorsal fin with XI spines and 15 or 16
soft rays; anal fin with III spines and 10 soft rays; posterior profile of dorsal and anal fins distinctly
pointed. Pectoral fins with 17 rays. Caudal fin slightly forked. Scale rows on back rising obliquely above
lateral line. Colour: generally red or pink in adults; juveniles and smaller adults pink with 3 dark red
bands, the first from first dorsal-fin spine through eye to tip of snout; the second from middle of spinous
part of dorsal fin to pelvic fins; and the third from base of last dorsal-fin spine running obliquely downward
across caudal peduncle and along lower edge of caudal fin.
Size: Maximum total length to at least 100 cm, commonly to 60 cm.
Habitat, biology, and fisheries: Occurs in the vicinity of coral reefs, often over adjacent sand flats. Also
trawled in deeper water on relatively flat bottoms. Small juveniles are frequently commensal with sea
urchins, or sometimes found in mangrove areas. Adults range from about 10 to depths of at least 100 m.
Feeds on fishes, crabs, stomatopods, other benthic crustaceans, and cephalopods. At New Caledonia
spawning occurs mainly during spring and summer. Estimated maximum age to at least 12 years. Often
found in markets and a valuable component of artisanal fisheries. It is a popular sport angling fish off the
Queensland coast of Australia. Caught with handlines, traps, and bottom trawls. Marketed mostly fresh,
but also dried-salted.
cific and Indian Ocean from
Australia to southern Japan,
and westward to East Africa
and the southern Red Sea.
Lutjanus semicinctus (Quoy and Gaimard, 1824) (Plate VIII, 57)

FAO names: En - Blackbanded snapper; Fr - Vivaneau à bandes noires; Sp - Pargo cinteado.
Diagnostic characters: Body moderately deep to relatively slender, its depth 2.6 to 3.1 times in
standard length. Dorsal profile of head moderately sloped; preorbital width about equal to eye diameter
or slightly greater; preopercular notch and knob poorly developed; vomerine tooth patch crescentic, without
a medial posterior extension; tongue with a patch of granular teeth. First gill arch with 14 to 19 gill rakers,
of which 8 to 12 (including rudiments) on lower limb. Dorsal fin with X spines and 13 soft rays; anal fin
with III spines and 8 or 9 soft rays; posterior profile of dorsal and anal fins rounded to angular. Caudal fin
truncate or slightly emarginate. Pectoral fins with 16 rays. Scale rows on back rising obliquely above lateral
line. Colour: generally pink grading to white on lower half of body; snout and forehead grey; a series of
7 narrow brown bars on upper back extending about half-way down sides; posterior half of caudal
peduncle and adjacent caudal-fin base black; fins mainly brownish, sometimes with a reddish hue, except
pelvic and pectoral fins pale.
Habitat, biology, and fisheries: Inhabits coral reefs at depths ranging from about 10 to 30 m. Usually
found solitarily or in small groups. Mainly important to artisanal fisheries, and occasionally seen in markets.
Caught mostly with handlines and traps. Offered fresh.
Distribution: Mainly south-
ern Oceania from Tahiti to
New Guinea, and extending
northward to the Philippines.
2894 Bony Fishes
Lutjanus stellatus Akazaki, 1983 (Plate VIII, 58)

FAO names: En - Star snapper; Fr - Vivaneau étoié; Sp - Pargo estrellado.
profile of head moderately sloped; preorbital bone broad, about wide the eye diameter; preopercular notch
and knob moderately developed; vomerine tooth patch crescentic, without a medial posterior extension;
tongue smooth, without teeth. First gill arch with 16 to 19 gill rakers, of which 11 to 13 (including rudiments)
on lower limb. Dorsal fin with X spines and 13 to 15 soft rays; anal fin with III spines and 8 or 9 soft
rays; posterior profile of dorsal and anal fins rounded. Caudal fin truncate or slightly emarginate. Pectoral
fins with 16 to 18 rays. Scale rows on back rising obliquely above lateral line. Colour: back and upper
sides dark brown to purplish; lower sides purple to yellow-brown grading to pale orange on belly; a small
white spot above lateral line at level of first soft dorsal-fin rays; a blue stripe from snout to opercle;
upper edge of opercle yellowish; fins greyish basally but mostly yellowish; juveniles yellowish with blue
longitudinal lines on sides; a white spot bordered with black at level of lateral line below last dorsal-fin
spines; a pair of horizontal blue lines below eye; fins yellow.
Habitat, biology, and fisheries: Occurs in the vicinity of coral and rocky reefs. Usually found solitarily or
in small groups. A good eating fish found in markets of southern Japan, Taiwan Province of China, and
Hong Kong. Caught by handlines, set nets, and gill nets. Usually offered fresh.
from the northwestern tropical
and subtropical Pacific, from
the vicinity of Hong Kong. May
enter the northern part of the
area.
Lutjanus timorensis (Quoy and Gaimard, 1824)

Frequent synonyms / misidentifications: None / Lutjanus malabaricus Schneider, 1801.
FAO names: En - Timor snapper; Fr - Vivaneau de Timor; Sp - Pargo de Timor.
adult
juvenile
profile of head steeply sloped; preorbital width greater than eye diameter; preopercular notch and knob
poorly developed; vomerine tooth patch crescentic, without a medial posterior extension; tongue smooth,
without teeth. First gill arch with 19 or 20 gill rakers, of which 12 or 13 (including rudiments) on lower limb.
Dorsal fin with X spines and 14 or 15 soft rays; anal fin with III spines and 8 soft rays; posterior profile
of dorsal and anal fins pointed. Caudal fin truncate or slightly emarginate. Pectoral fins with 17 rays. Scale
rows on back rising obliquely above lateral line. Colour: back and upper sides reddish; lower sides and
belly red or pinkish to silvery white; fins reddish; axil of pectoral fins black. Juveniles with a broad blackish
or brownish band from upper jaw to beginning of dorsal fin and a black saddle preceded by a pearly white
border on upper edge of caudal peduncle; sides with a series of fine reddish brown lines.
Habitat, biology, and fisheries: Inhabits coral and rubble reefs at depths of about 20 m to at least 130
m. Usually occurs solitarily. A poorly known red snapper of potential interest to fisheries, although it appears
to be relatively uncommon in market catches. Caught mainly with deep handlines and occasionally trawled.
Marketed mainly fresh.
Distribution: Mainly west-
ern Pacific Ocean from Fiji Is-
lands to Malay Peninsula;
also recorded from the An-
daman Sea off Thailand.
2896 Bony Fishes
Lutjanus vitta (Quoy and Gaimard, 1824) (Plate IX, 59)

FAO names: En - Brownstripe snapper or Brownstripe red snapper; Fr - Vivaneau à bande brune;
Sp - Pargo bitilla.
Diagnostic characters: Body moderately deep to relatively slender, its depth 2.6 to 3 times in standard
length. Dorsal profile of head moderately sloped; preorbital width about equal to eye diameter;
preopercular notch and knob poorly developed; vomerine tooth patch triangular with a medial posterior
extension or diamond-shaped; tongue with a patch of granular teeth. First gill arch with 15 to 19 gill rakers,
of which 9 to 12 (including rudiments) on lower limb. Dorsal fin with X spines and 12 or 13 soft rays;
anal fin with III spines and 8 or 9 soft rays; posterior profile of dorsal and anal fins angular. Caudal fin
slightly emarginate or truncate. Pectoral fins with 15 or 16 rays. Scale rows on back rising obliquely above
lateral line. Colour: back and upper sides brown, lower sides and belly whitish or pink; narrow longitudinal
brown lines, 1 per scale row, on sides, those above lateral line slanted posteriorly toward dorsal-fin base;
a dark brown to blackish stripe along middle of side from eye to upper half of caudal peduncle; fins
yellow except pelvics whitish. Juveniles and subadults with an intensely black midlateral stripe and an oval
black spot, eye-sized or greater, lying in the middle of the stripe below last dorsal-fin spine.
Habitat, biology, and fisheries: Inhabits the vicinity of coral reefs, also in areas of flat bottom with
occasional low coral outcrops, sponges, and sea whips at depths between about 10 and 40 m. Occurs
solitarily or in groups of up to about 30 individuals. Feeds on fishes, shrimps, crabs and other benthic
invertebrates. At New Caledonia spawning occurs over most of the year with peak activity during spring
and summer. Eggs have a diameter of 0.78 to 0.84 mm and hatch in about 26 hours at 26.5°C. The larvae
grow at a rate of 1.2 to 1.7 mm per day during the first 1 or 2 months.
Distribution: Mainly western
Pacific and eastern Indian
Ocean from New Caledonia
and Gilbert Islands to south-
ern India, extending north-
ward to southern Japan; also
found in the Seychelles.
Macolor macularis Fowler, 1931

Frequent synonyms / misidentifications: None / Macolor niger (Forsskål, 1775).
FAO names: En - Midnight snapper; Fr - Vivaneau minuit; Sp - Pargo medianoche.
D i a g n o s t i c c h a r a c ters :
Body relatively deep, laterally
compressed. Dorsal profile
of head convex in adults, in
oblique straight line in
young. Lacrimal (suborbital
region) relatively broad. Nos-
trils on each side of snout not
very close together. Mouth
large and terminal, lower jaw
p rotruding slightly wh en
mouth closed. Premaxillae
protrusible. Maxilla reaching
vertical through middle of adult
eye. Row of conical teeth in
each jaw, enlarged anteriorly,
and inner bands of villiform
teeth on sides of upper jaw
and front of lower jaw; vomer-
ine tooth patch broadly V-
s h a p e d . Maxilla without
scales or longitudinal
ridges. Preopercle with
deep notch (or gash) re-
ceiving elongate interoper- juvenile
cular spine which is
distinct in specimens more
than about 20 cm standard
length. First gill arch with 37 to 42 gill rakers on upper limb, 71 to 81 on lower limb (total 110 to 122).
Dorsal fin continuous, but spinous portion distinctly notched in young; largest individuals without notch
or with barely perceptible notch. Dorsal and anal fins distinctly pointed posteriorly. Last soft ray of
both dorsal and anal fins not produced, shorter than next to last ray. Caudal fin rounded in
specimens less than about 20 mm standard length, emarginate in individuals more than about
10 cm standard length. Pectoral fins fairly long; in specimens more than about 10 cm standard length,
pectoral fins 95 to 110% head length. Pelvic fins very long and pointed in young, but short and rounded
in adults. Dorsal fin with X spines and 13 or 14 (usually 13) soft rays. Anal fin with III spines and
10 or 11 (usually 10) soft rays. Pectoral-fin rays 16 to 18 (usually 17). Soft dorsal and anal fins with
scales basally. Tubed lateral-line scales 50 to 55. Colour: adults mostly brownish black to black;
individuals less than about 20 cm total length black on upper sides (usually with 6 to 10 white spots)
and white below, black vertical bar through eye, broad black horizontal band from eye to rear edge
of opercle, and black stripe from pectoral region to distal end of lower caudal-fin lobe.
Size: Maximum standard length at least 45 cm.
Habitat, biology, and fisheries: Inhabits coastal waters in depths between about 5 and 90 m; apparently
does not form large schools.
Feeds largely on fishes and
crustaceans. Caught with
handlines, gill nets, and traps;
also speared by divers. Fre-
quently seen in markets,
mainly fresh.
from Ryukyu Islands south-
ward to Australia and Melane-
sia. May be more widespread;
confused in the literature with
Macolor niger.
click for next page

2898 Bony Fishes
Macolor niger (Forsskål, 1775)

Frequent synonyms / misidentifications: Macolor macolor (Lesson, 1827); M. typus Bleeker, 1860 /
Macolor macularis Fowler, 1931.
FAO names: En - Black-and-white snapper; Fr - Vivaneau plate; Sp - Pargo blanco y negro.
Diagnostic characters: Body
relatively deep, laterally com-
pressed. Dorsal profile of
head convex in adults, in
oblique straight line in young.
Lacrimal (suborbital region)
relatively broad. Nostrils on
each side of snout not very
close together. Mouth large and
terminal, lower jaw protruding
slightly when mouth closed.
Premaxillae protrusible. Maxilla
reaching vertical slightly ante-
rior to middle of eye. Row of
conical teeth in each jaw, en-
larged anteriorly, and inner
adult
bands of villiform teeth on sides
of upper jaw and front of lower
ja w; v o m erin e too th patch
b r o a d l y V - s h a p e d . Maxilla
without scales or longitudinal
ridges. Preopercle with deep
notch (or gash) receiving
elongate interopercular spine
which is absent in specimens
less than about 16 cm stand- juvenile
ard length. First gill arch with
26 to 38 gill rakers on upper
limb, 60 to 71 on lower limb (total 89 to 107). Dorsal fin continuous, but spinous portion distinctly
notched in young, notch shallower in large specimens. Dorsal and anal fins distinctly pointed
posteriorly. Last soft ray of both dorsal and anal fins not produced, shorter than next to last ray.
Caudal fin rounded and with central rays elongated in small juveniles, slightly notched at interme-
diate sizes, and emarginate in adults. Pectoral fins fairly long; in specimens more than about 95 mm
standard length, pectoral fins 90 to 110% head length. Pelvic fins short and rounded at all sizes. Dorsal
fin with X spines and 13 to 15 (usually 14) soft rays. Anal fin with III spines and 10 or (usually) 11
soft rays. Pectoral-fin rays 16 to 18 (usually 17). Soft dorsal and anal fins with scales basally. Tubed
lateral-line scales 49 to 58. Colour: adults mostly brownish black to black; individuals less than
about 24 cm standard length have black-and-white bands on body, about 5 (4 to 7) white spots on
upper side, and white at tips of caudal-fin lobes and in middle of fin.
Habitat, biology, and fisheries: Inhabits coastal waters in depths between about 2 and 90 m; in the
Ryukyu Islands, adults reported to occur in large schools in depths usually less than 30 m. Feeds largely
on fishes and crustaceans.
Caught with handlines, gill
nets, and traps; also speared
by divers. Frequently seen in
markets, mainly fresh.
Samoa to East Africa and
from central Japan southward
to Australia.
Paracaesio gonzalesi Fourmanoir and Rivaton, 1979

FAO names: En - Vanuatu snapper; Fr - Vivanette vanuatu; Sp - Panchote vanuatú.
Diagnostic characters: Body moderately deep, laterally compressed. Nostrils on each side of snout close
together. Lower jaw protruding slightly when mouth closed. Anterior end of upper lip without a thick
fleshy protrusion. Premaxillae protrusible. Maxilla reaching vertical through anterior part of eye. Both
upper and lower jaws with an outer row of small conical teeth, larger anteriorly, and with an inner band of
small villiform teeth; vomer and palatines with very small, essentially villiform teeth, those on vomer in
chevron-shaped patch; no teeth on tongue. Maxilla with scales, but without longitudinal ridges.
Interorbital region rather strongly convex. First gill arch with 11 gill rakers on upper limb and 20 on
lower limb (total 31). Dorsal fin continuous, not deeply incised near junction of spinous and soft portions.
Last soft ray of dorsal and anal fins not produced, shorter than next to last ray. Caudal fin lunate
to emarginate. Pectoral fins long, longer than head. Dorsal fin with X spines and 10 soft rays. Anal fin
with III spines and 8 soft rays. Pectoral-fin rays 16. Membranes of dorsal and anal fins without scales.
Tubed lateral-line scales 48 or 49. Colour: back and upper sides light brown; silvery white on lower sides
and belly; yellow band running from origin of lateral line obliquely to about middle of dorsal fin; in
life sometimes with 8 broad dark vertical bars on side; upper lobe of caudal fin violet grey, lower lobe bright
yellow.
Size: Maximum standard length about 42 cm.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between about 140 and 250 m.
Caught mainly with handlines. A good food fish. Marketed fresh.
Distribution: Tropical west-
ern Pacific. Known only from
the Fiji Islands, Vanuatu, and
the Philippines, but probably
more widespread.
Remarks: According to some
of the literature, this species
lacks maxillary squamation,
but the holotype, at least, has
scaly maxillae. Maxillary
squamation may be a plastic
character in Paracaesio gon-
zalesi, as it is in P. xanthura.
2900 Bony Fishes
Paracaesio kusakarii Abe, 1960 (Plate IX, 60)

FAO names: En - Saddleback snapper; Fr - Vivanette sellée; Sp - Panchote sillevo.
fleshy protrusion. Premaxillae protrusible. Maxilla reaching vertical through anterior part of eye. Both
upper and lower jaws with an outer row of conical teeth (some of these enlarged into small canines
anteriorly) and with an inner band of small villiform teeth; vomer and palatines with villiform teeth, those
on vomer in chevron-shaped patch; no teeth on tongue. Maxilla with scales, but without longitudinal
ridges. Interorbital region rather strongly convex. Large individuals with pronounced hump on nape
as a result of higher supraoccipital crest. First gill arch with 7 to 12 gill rakers on upper limb, 19 or
20 on lower limb (total 26 to 32). Dorsal fin continuous, not deeply incised near junction of spinous and
soft portions. Last soft ray of dorsal and anal fins not produced, shorter than next to last ray. Caudal
fin lunate to emarginate; middle of distal margin of caudal fin becoming rounded with age. Pectoral
fins long, longer than head. Dorsal fin with X spines and 10 soft rays. Anal fin with III spines and 8 soft
rays. Pectoral-fin rays 16. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales
48 to 50. Colour: back and upper side of body light brown; remainder of body silvery white; 4 broad dark
vertical bars on upper side of body; fins greyish, whitish, or yellowish.
Caught mainly with handlines. Marketed fresh.
cific from Samoa to Taiwan
Province of China and Singa-
pore and from the Ryukyu and
Ogasawara islands south-
ward to New Caledonia and
northeastern Australia.
Paracaesio paragrapsimodon Anderson and Kailola, 1992

FAO names: En - Toothy.
(after Anderson, Kailola, and Collette, 1992)
Diagnostic characters: Body relatively elongate (depth of body 33 to 35% standard length), laterally
compressed. Nostrils on each side of snout close together. Mouth terminal, jaws about equal. Anterior
end of upper lip without a thick fleshy protrusion. Premaxillae protrusible. Maxilla reaching posteriorly
to a vertical well past anterior border of eye. Upper jaw with an outer row of caniniform teeth, some
very well-developed exserted canines anteriorly, and an inner band of very small teeth; lower jaw with
outer row of conical teeth, some very well-developed strongly exserted canines anteriorly, and an
inner band of smaller conical teeth; vomer and palatines with small teeth, those on vomer in roughly
triangular patch; no teeth on tongue. Maxilla without scales or longitudinal ridges. Interorbital region
rather strongly convex. First gill arch with 9 gill rakers on upper limb, 20 or 21 on lower limb (total
29 or 30). Dorsal fin continuous, not incised near junction of spinous and soft portions. Last soft ray of
dorsal and anal fins not produced, shorter than next to last ray. Caudal fin deeply forked. Pectoral
fins long, longer than head. Dorsal fin with X spines and 10 soft rays. Anal fin with III spines and 8 soft
rays. Pectoral-fin rays 17. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales
69 or 70. Colour: body deep metallic blue; proximal halves of soft dorsal and soft anal fins pale yellow,
with yellow extending onto dorsalmost part of back and dorsal part of caudal peduncle; yellow along middle
of each caudal-fin lobe and at posterior margin of caudal fin.
Size: Maximum standard length unknown, but reaches at least 28 cm.
Habitat, biology, and fisheries: Depth of capture 200 m, available for only 1 collection. No other
information available.
cific; known from only 2 speci-
mens, one collected off the
Caroline Islands, the other off
southern Papua New Guinea.
2902 Bony Fishes
Paracaesio sordida Abe and Shinohara, 1962 (Plate IX, 61)

FAO names: En - Dirty ordure snapper.
together. Mouth terminal, jaws about equal. Anterior end of upper lip without a thick fleshy
protrusion. Premaxillae protrusible. Maxilla reaching vertical through anterior part of eye. Both upper and
lower jaws with an outer row of conical teeth (some of these enlarged into caniniform teeth anteriorly) and
with an inner band of very small teeth; vomer and palatines with very small teeth, those on vomer in a
triangular or chevron-shaped patch; no teeth on tongue. Maxilla without scales or longitudinal ridges.
Interorbital region convex. First gill arch with 9 to 11 gill rakers on upper limb, 19 to 24 on lower
limb (total 28 to 34). Dorsal fin continuous, not deeply incised near junction of spinous and soft portions.
Last soft ray of dorsal and anal fins not produced, shorter than next to last ray. Caudal fin deeply
forked. Pectoral fins long, usually longer than head. Dorsal fin with X spines and 10 soft rays. Anal fin
with III spines and 8 or 9 (usually 8) soft rays. Pectoral-fin rays 16 or (usually) 17. Membranes of dorsal
and anal fins without scales. Tubed lateral-line scales 69 to 72. Preopercle with 1 to 3 rows of scales
peripheral to and usually distinctly separated from main group of cheek scales. Colour: overall dark
grey, dark purplish brown, dark violet, or bluish; lower sides and belly about as dark as overall coloration,
silvery, or whitish; fins variable: dark grey, reddish brown, or dorsal and caudal fins brownish to slightly
yellow with other fins mostly whitish to translucent.
Caught mainly with handlines, but in relatively low numbers. An important food fish in some areas. Usually
marketed fresh.
the Marquesas Islands and
Pitcairn Island in the east to
the Gulf of Aqaba and East
Africa in the west and ranging
northward to the Ryukyu Is-
lands.
Remarks: Paracaesio i s a
feminine noun; therefore, ad-
jectival specific names for
species of Paracaesio (such
as sordida) correctly termi-
nate in “a”.
Paracaesio stonei Raj and Seeto, 1983

FAO names: En - Cocoa snapper; Fr - Vivanettte cacao; Sp - Panchote cacao.
fleshy protrusion. Premaxillae protrusible. Maxilla reaching vertical through about middle of eye. Both
upper and lower jaws with an outer row of conical teeth (teeth larger anteriorly) and with an inner band of
much smaller teeth; vomer and palatines with villiform teeth, those on vomer in triangular patch; no teeth
on tongue. Maxilla without scales or longitudinal ridges. Interorbital region convex. First gill arch
with 10 or 11 gill rakers on upper limb, 17 or 18 on lower limb (total 27 to 29). Dorsal fin continuous,
not deeply incised near junction of spinous and soft portions. Spines of dorsal and anal fins robust, at
least on large individuals. Last soft ray of dorsal and anal fins not produced, shorter than next to
last ray. Caudal fin emarginate. Pectoral fins long, longer than head. Dorsal fin with X spines and 10
soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 16. Membranes of dorsal and anal
fins without scales. Tubed lateral-line scales 48 to 50. Colour: general body coloration light brown
dorsally and silver ventrally with 4 or 5 broad brown to dark grey vertical bars on body extending from
dorsal surface to midline or below midline laterally; fins grey to whitish except dorsal and caudal fins
with variable amounts of yellow.
Habitat, biology, and fisheries: Known from depths ranging from 200 to 320 m. Caught with deep
handlines.
Distribution: Known from off
the Ryukyu Islands, north-
eastern Australia, Vanuatu,
Fiji, and Samoa; also reported
from off Papua New Guinea;
probably more widely distrib-
uted.
2904 Bony Fishes
Paracaesio xanthura (Bleeker, 1869) (Plate IX, 62)

Frequent synonyms / misidentifications: ? Paracaesio pedleyi Mcculloch and Waite, 1916; P. tumida
(Tanaka, 1917); P. cantharoides (Barnard, 1937) / Other species of Paracaesio.
FAO names: En - Yellowtail blue snapper; Fr - Vivanette queue jaune; Sp - Panchote rabo amarillo.
together. Mouth terminal, jaws about equal. Anterior end of upper lip without a thick fleshy
protrusion. Premaxillae protrusible. Maxilla reaching vertical through anterior part of eye. Both upper and
lower jaws with an outer row of conical teeth (some of these enlarged into caniniform teeth anteriorly) and
with an inner band of very small teeth; vomer and palatines with very small teeth, those on vomer in a
chevron-shaped or triangular patch; no teeth on tongue. Maxilla with or without scales and without
longitudinal ridges. Interorbital region convex. First gill arch with 7 to 12 gill rakers on upper limb,
20 to 23 on lower limb (total 28 to 35). Dorsal fin continuous, not deeply incised near junction of spinous
and soft portions. Last soft ray of dorsal and anal fins not produced, shorter than next to last ray.
Caudal fin deeply forked. Pectoral fins long, usually longer than head in specimens greater than 12 cm
standard length. Dorsal fin with X spines and 10 soft rays. Anal fin with III spines and 8 or (rarely) 9
soft rays. Pectoral-fin rays 16 to 18 (usually 17). Membranes of dorsal and anal fins without scales.
Tubed lateral-line scales 68 to 72. Preopercle almost always without scales peripheral to main group
of cheek scales. Colour: mainly bluish; broad yellow band on upper side beginning near anterior end
of dorsal fin, continuing over most of caudal peduncle and out onto both lobes of caudal fin; dorsal
fin grey to yellowish, other fins greyish or whitish.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between about 5 and 150 m;
sometimes forming large schools. Feeds largely on zooplankton. Caught mainly with handlines and bottom
longlines. A food fish in some areas. Marketed fresh or frozen.
the Austral Islands to East Af-
rica and from southern Japan
southward to southeastern
Australia.
Remarks: Paracaesio xan-
thura, a complex of variable
populations, may compose a
single species or 2 (or more)
species. Paracaesio pedleyi,
from Lord Howe Island, Mid-
dleton Reef, and eastern Aus-
tralian waters, is included here with some reservation. Paracaesio is a feminine noun; therefore, adjectival
specific names for species of Paracaesio (such as xanthura) correctly terminate in “a”.
Parapristipomoides squamimaxillaris (Kami, 1973) (Plate IX, 63)

FAO names: En - Scalemouth jobfish; Fr - Colas écailleuse; Sp - Panchito escamudo.
Diagnostic characters: Body elongate, its depth 28 to 33% standard length, laterally compressed. Nostrils
on each side of snout close together. Lower jaw protruding slightly when mouth closed. Anterior end
of upper lip without a thick fleshy protrusion. Premaxillae protrusible. Maxilla reaching vertical through
anterior part of eye. Jaw teeth poorly developed, no canines present; both upper and lower jaws with
an outer row of small conical teeth and an inner band of villiform teeth that is best developed anteriorly;
vomer and palatines with teeth, those on vomer in roughly an oval-shaped patch; no teeth on tongue.
Maxilla with scales, but without longitudinal ridges. Interorbital region very slightly convex. First
gill arch with 10 to 12 gill rakers on upper limb, 22 to 24 on lower limb (total 32 to 36). Dorsal fin
continuous, not deeply incised near junction of spinous and soft portions. Last soft ray of dorsal and anal
fins about equal to or slightly longer than next to last soft ray. Caudal fin well forked. Pectoral fins
fairly long, about as long as head. Dorsal fin with X spines and 10 soft rays. Anal fin with III spines and
8 soft rays. Pectoral-fin rays 16. Membranes of dorsal and anal fins without scales except for a few
scales that may occur at bases of posteriormost soft ray of each fin. Tubed lateral-line scales 53
to 55. Colour: silvery pink, darker pink on back and upper sides; spinous part of dorsal fin with faint mottling
of light yellow; upper lobe of caudal fin bright yellow, lower lobe pink; pectoral-fin rays edged with pink,
with a little yellow in upper rays; other fins mainly whitish.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths from about 130 to at least 460 m.
Caught by deep handlining. Marketed fresh.
Distribution: Known from off
Easter Island, Rapa, Tonga,
New Caledonia, and from the
Chesterfield Islands region of
the Coral Sea.
2906 Bony Fishes
Pinjalo lewisi Randall, Allen, and Anderson, 1987 (Plate IX, 64)
Frequent synonyms / misidentifications: ? Pinjalo microphthalmus Lee, 1987 / Pinjalo pinjalo
(Bleeker, 1850).
FAO names: En - Red pinjalo.
(after Randall, Allen, and Anderson, 1987)
Diagnostic characters: Body moderately deep, laterally compressed. Dorsal profile of head convex.
Nostrils on each side of snout not very close together. Mouth small and oblique, forming an angle of about
45° with the horizontal; lower jaw protruding slightly when mouth closed. Premaxillae protrusible. Maxilla
falling short of, reaching, or extending slightly beyond vertical through anterior edge of orbit. No caniniform
teeth; an outer row of small fixed incurved conical teeth in each jaw; an inner band of depressible villiform
teeth in upper jaw and front of lower jaw; minute teeth in a triangular or chevron-shaped patch on vomer
and in a narrow band on palatines; no teeth on tongue. Maxilla without scales or longitudinal ridges.
Interorbital region strongly convex. First gill arch with 7 or 8 gill rakers on upper limb, 16 or 17 on
lower limb. Dorsal fin continuous, without a notch between spinous and soft portions. Dorsal- and anal-fin
spines rather slender. Last soft ray of both dorsal and anal fins not produced, shorter than next to
last ray. Caudal fin slightly emarginate, the caudal concavity 3.55 to 5.45 times in head length.
Pectoral fins relatively long, 2.5 to 3.05 times in standard length. Pelvic fins relatively short, 1.65
to 1.8 times in head length. Dorsal fin with XII spines and 13 soft rays. Anal fin with III spines and
8 or 9 soft rays. Pectoral-fin rays 17 or 18. Dorsal and anal fins with scales basally. Tubed lateral-line
scales 48 to 50. Colour: dorsal half of body red, shading ventrally to pinkish white, no diagonal dark
lines on nape and body; head red except areas covered with scales are pinkish white; a curved pale
pink spot dorsally on caudal peduncle, broadest on top of peduncle (spot about as wide as diameter
of pupil), this spot extending diagonally downward and forward to within a scale row of lateral line
(spot may not be evident in all individuals); fins red, anal and paired fins lighter than dorsal and
caudal fins; margin of dorsal fin and to a lesser extent soft portion of anal fin blackish; posterior
margin of caudal fin blackish, and upper and lower margins narrowly dark.
Habitat, biology, and fisheries: Inhabits reefs and rocky bottoms in depths between about 40 and 150 m.
Feeds on benthic and plank-
tonic invertebrates and possi-
bly small fishes. Mainly taken
with handlines, traps, and bot-
tom trawls. Marketed fresh or
dried-salted.
uted in the Indo-West Pacific
from Fiji to the Laccadive Is-
lands and northward from
southern Indonesia to the
Ryukyu Islands.
Pinjalo pinjalo (Bleeker, 1850) (Plate IX, 65)

Frequent synonyms / misidentifications: ? Pinjalo microphthalmus Lee, 1987 / None.
FAO names: En - Pinjalo; Fr - Pinjalo; Sp - Pinjalo.
(after Randall, Allen, and Anderson, 1987)
Diagnostic characters: Body moderately deep, laterally compressed. Dorsal profile of head convex.
Nostrils on each side of snout not very close together. Mouth small and oblique, forming an angle of about
45o with the horizontal; lower jaw protruding slightly when mouth closed. Premaxillae protrusible. Maxilla
falling short of, reaching, or extending slightly beyond vertical through anterior edge of orbit. No caniniform
teeth; an outer row of small fixed incurved conical teeth in each jaw; an inner band of depressible villiform
teeth in upper jaw and front of lower jaw; minute teeth in a triangular or chevron-shaped patch on vomer
and in a narrow band on palatines; no teeth on tongue. Maxilla without scales or longitudinal ridges.
Interorbital region strongly convex. First gill arch with 6 to 8 gill rakers on upper limb, 16 to 18 on
lower limb. Dorsal fin continuous, without a notch between spinous and soft portions. Dorsal- and anal-fin
spines rather slender. Last soft ray of both dorsal and anal fins not produced, shorter than next to
last ray. Caudal fin deeply emarginate, the caudal concavity 2.1 to 3.6 times in head length. Pectoral
fins relatively short, 3.05 to 3.9 times in standard length. Pelvic fins relatively long, 1.4 to 1.65 times
in head length. Dorsal fin with XI spines and 14 or 15 soft rays. Anal fin with III spines and 9 or
(usually) 10 soft rays. Pectoral-fin rays 17 to 19. Dorsal and anal fins with scales basally. Tubed
lateral-line scales 48 to 51. Colour: body pink to silvery lavender dorsally, shading to pale pink or
silvery white ventrally, with diagonal yellowish brown to brownish red lines following scale rows
on nape and dorsal one-half to two-thirds of body; head purplish pink to brownish red dorsally,
pink to silvery white ventrally; pale spot sometimes present dorsally on caudal peduncle of small
individuals; dorsal and caudal fins reddish, usually suffused with yellow, the outer margin blackish;
anal and pelvic fins light red to yellow; pectoral fins light red.
Habitat, biology, and fisheries: Inhabits reefs and rocky bottoms to depths of about 60 m. Feeds on
benthic and planktonic invertebrates and possibly small fishes. Caught mainly with handlines, traps, and
bottom trawls. Marketed fresh
or dried-salted.
uted in the Indo-West Pacific
from the Gulf of Papua
(Papua New Guinea) to the
Persian Gulf and northward
from southern Indonesia to
Taiwan Province of China.
2908 Bony Fishes
Pristipomoides argyrogrammicus (Valenciennes, 1832) (Plate IX, 66)

Frequent synonyms / misidentifications: Pristipomoides amoenus (Snyder, 1911) / None.
FAO names: En - Ornate jobfish; Fr - Colas orné; Sp - Panchito adornado.
together. Jaws about equal or upper jaw protruding slightly. Premaxillae protrusible. Maxilla extending to
vertical through anterior one-third of eye or slightly beyond. Jaws with bands of fine teeth, the outer ones
slightly enlarged, but not as prominent canines; vomer and palatines with teeth, those on vomer more or
less in chevron-shaped patch; no teeth on tongue. Maxilla without scales or longitudinal ridges.
Interorbital region flattened to slightly convex. First gill arch with 5 or 6 gill rakers on upper limb,
and soft portions. Last soft ray of both dorsal and anal fins well produced, longer than next to last
ray. Caudal fin forked. Pectoral fins fairly long, about 90% of head length. Dorsal fin with X spines and
11 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 15 or 16. Membranes of dorsal and
anal fins without scales. Tubed lateral-line scales 58 to 62. Colour: upper part of head reddish; back
mainly yellow; sides and belly silvery, sometimes pinkish; body with many bright blue spots and vermicu-
lations; zigzag line running at about level of lateral line anteriorly and then onto dorsolateral part of
caudal peduncle posteriorly; dorsal and caudal fins yellowish.
Size: Maximum standard length approaching 40 cm, commonly to 25 cm.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between about 70 and 350 m. Feeds
on small fishes, crustaceans, and squids. Caught mainly with handlines. A good food fish sometimes seen
in markets where it is sold fresh.
the Indo-Pacific from the
Society Islands to East Africa
and from New Caledonia
northward to southern Japan.
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Pristipomoides auricilla (Jordan, Evermann, and Tanaka, 1927) (Plate X, 67)

Frequent synonyms / misidentifications: None / Other species of Pristipomoides.
FAO names: En - Goldflag jobfish; Fr - Colas drapeau; Sp - Panchito abanderado.
together. Jaws about equal or lower jaw protruding slightly. Premaxillae protrusible. Maxilla extending to
vertical through anterior part of eye or slightly beyond. Upper and lower jaws both with an outer row of
conical and canine teeth and an inner band of villiform teeth; vomer and palatines with teeth, those on
vomer in triangular patch; no teeth on tongue. Maxilla without scales or longitudinal ridges. Interorbital
region flattened. First gill arch with 8 to 11 gill rakers on upper limb, 17 to 21 on lower limb (total
27 to 32). Dorsal fin continuous, not deeply incised near junction of spinous and soft portions. Last soft
ray of both dorsal and anal fins well produced, longer than next to last ray. Caudal fin forked. Pectoral
fins long, equal to or somewhat shorter than head length. Dorsal fin with X spines and 11 soft rays. Anal
fin with III spines and 8 soft rays. Pectoral-fin rays 15 or 16. Membranes of dorsal and anal fins without
scales. Tubed lateral-line scales 67 to 74. Colour: body purplish or brownish violet; sides with
numerous yellow spots or faint yellow chevron-shaped bands; dorsal fin yellowish to yellowish brown;
upper lobe of caudal fin yellow. Sexual dichromatism: males over 27 cm (fork length) with much yellow
on lower lobe of caudal fin, usually forming a distinct blotch; females with or without yellowish colour on
lower lobe of caudal fin, but if yellow present, not forming a distinct blotch.
Size: Maximum standard length 40 cm or more, commonly to about 25 cm.
Habitat, biology, and fisheries: Occurs over rocky bottoms between about 80 and 360 m; off Guam,
caught most abundantly from about 180 to 270 m. Feeds on fishes and salps. Caught mainly with deep
handlines. A common food fish in some areas such as Samoa and Guam. Marketed fresh.
and Rapa to Mauritius and
northward from Australia and
New Caledonia to Japan.
2910 Bony Fishes
Pristipomoides filamentosus (Valenciennes, 1830) (Plate X, 68)

Frequent synonyms / misidentifications: Pristipomoides microlepis (Bleeker, 1869) / Other species of
Pristipomoides.
FAO names: En - Crimson jobfish; Fr - Colas fil; Sp - Panchito hebra.
vertical through about anterior margin of pupil. Upper and lower jaws both with an outer row of conical to
small canine teeth and an inner band of villiform teeth, caniniform teeth at anterior ends of jaws not
appreciably enlarged; vomer and palatines with teeth, those on vomer in triangular patch; no teeth on
tongue. Maxilla without scales or longitudinal ridges. Interorbital region flattened. First gill arch with
7 or 8 gill rakers on upper limb, 15 to 18 on lower limb (total 22 to 26). Dorsal fin continuous, not deeply
incised near junction of spinous and soft portions. Last soft ray of both dorsal and anal fins well
produced, longer than next to last ray. Caudal fin forked. Pectoral fins long, about equal to or somewhat
shorter than head length. Dorsal fin with X spines and 11 soft rays. Anal fin with III spines and 8 (rarely
7 or 9) soft rays. Pectoral-fin rays 15 or 16. Membranes of dorsal and anal fins without scales. Tubed
lateral-line scales 57 to 62. Colour: back and sides variable, ranging from brownish to lavender or reddish
purple; snout and interorbital space with narrow yellow lines and blue spots often persisting as dark spots
in preservative; dorsal and caudal fins light blue or lavender with reddish orange margins.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between 40 and 360 m; off Guam,
caught most abundantly between 180 and 270 m. Feeds on small fishes, shrimps, crabs, amphipods,
ascidians, and salps. Around Hawaii, spawning occurs from March to December with peak activity from
May to September. Estimated maximum age is 18 years. Caught mainly with bottom longlines and deep
handlines. It is one of the principal species in the Hawaiian offshore handline fishery, commanding a high
price. A good quality food fish. Marketed mainly fresh.
and Tahiti to East Africa and
northward from eastern
Australia (Queensland and
New South Wales) and Lord
Howe Island to southern
Japan.
Pristipomoides flavipinnis Shinohara, 1963 (Plate X, 69)

FAO names: En - Goldeneye jobfish; Fr - Colas oeil doré; Sp - Panchito de ojo dorado.
together. Lower jaw protruding slightly. Premaxillae protrusible. Maxilla extending to vertical through about
anterior margin of pupil. Upper and lower jaws both with an outer row of conical and canine teeth and an
inner band of villiform teeth, canines at anterior ends of both jaws usually enlarged; vomer and
palatines with teeth, those on vomer in triangular patch; no teeth on tongue. Maxilla without scales or
longitudinal ridges. Interorbital region flattened. First gill arch with 6 to 9 gill rakers on upper limb,
ray. Caudal fin forked. Pectoral fins long, about equal to or somewhat shorter than head length. Dorsal
fin with X spines and 11 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 16.
Membranes of dorsal and anal fins without scales. Tubed lateral-line scales 59 to 63. Colour: back
and upper sides pale lavender or pinkish; lower sides and belly silvery; network of dark spots on top of
head; margin of dorsal fin yellow when fresh.
Size: Maximum standard length about 50 cm or more, commonly to about 35 cm.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between about 90 and 360 m; off
Guam, caught most abundantly between 180 and 270 m. Feeds primarily on benthic fishes and to a lesser
extent on crustaceans, squids, and pelagic tunicates. At Vanuatu (New Hebrides), spawning occurs
throughout the year with peak activity from December to February. Caught mainly with bottom longlines
and handlines. An important market fish in some areas. Marketed mainly fresh.
Distribution: Pacific Ocean
from Tahiti to southeastern
Asia and northward from Aus-
tralia to the Ryukyu Islands.
2912 Bony Fishes
Pristipomoides multidens (Day, 1871) (Plate X, 70)

Frequent synonyms / misidentifications: None / Pristipomoides typus Bleeker, 1852.
FAO names: En - Goldbanded jobfish; Fr - Colas à bandes dorées; Sp - Panchito de bandas doradas.
together. Jaws about equal or lower jaw protruding very slightly. Premaxillae protrusible. Maxilla extending
to vertical through about middle of eye. Upper and lower jaws both with an outer row of conical teeth: some
of the anteriormost teeth enlarged into canines, and an inner band of villiform teeth; vomer and palatines
with teeth, those on vomer in chevron-shaped patch; no teeth on tongue. Maxilla without scales or
longitudinal ridges. Interorbital region flattened. First gill arch with 6 to 8 gill rakers on upper limb,
ray. Caudal fin forked. Pectoral fins long, a little shorter than length of head. Dorsal fin with X spines
and 11 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 15 or 16. Membranes of dorsal
and anal fins without scales. Tubed lateral-line scales 48 to 50. Colour: yellowish to rosy, with about
6 broken golden stripes on sides; side of snout and cheek with 2 golden stripes bordered with blue
(persisting as brownish stripes in preservative); top of head with a series of transverse brownish
yellow vermiculations; dorsal fin with yellowish stripes or rows of spots.
Size: Maximum standard length more than 70 cm, commonly to 40 cm.
on fishes, shrimps, crabs, lobsters, stomatopods, squids, gastropods, and urochordates. Spawning has
been reported during May and August in the South China Sea and throughout most of the year at Samoa
and Vanuatu (New Hebrides) with peak activity during December and January at the latter locality.
Estimated maximum age is 14 years. Caught mainly with handlines. An excellent food fish, often seen in
markets, but usually in small quantities. Marketed fresh.
Samoa to the Red and Ara-
bian seas and to East Africa
and southward from southern
Japan to Australia.
Pristipomoides sieboldii (Bleeker, 1857) (Plate X, 71)

Frequent synonyms / misidentifications: ? Pristipomoides microdon (Steindachner, 1877) / Other
species of Pristipomoides.
FAO names: En - Lavender jobfish; Fr - Colas lavande; Sp - Panchito lavanda.
together. Lower jaw protruding slightly. Premaxillae protrusible. Maxilla extending to vertical through
anterior part of eye. Upper and lower jaws both with an outer row of conical teeth (a few anterior teeth in
upper jaw enlarged into small canines) and an inner band of villiform teeth; vomer and palatines with teeth,
vomerine tooth patch with backward prolongation in midline (patch sometimes diamond-shaped);
teeth present on tongue. Maxilla without scales or longitudinal ridges. Interorbital region flattened.
First gill arch with 9 to 12 gill rakers on upper limb, 19 to 22 on lower limb (total 28 to 33). Dorsal fin
continuous, not deeply incised near junction of spinous and soft portions. Last soft ray of both dorsal
and anal fins well produced, longer than next to last ray. Caudal fin forked. Pectoral fins long:
somewhat shorter than, about equal to, or somewhat longer than length of head. Dorsal fin with X spines
and 11 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin rays 16 or 17. Membranes of dorsal
and anal fins without scales. Tubed lateral-line scales 69 to 74. Colour: silvery with lavender hue;
dark spots on top of head, more conspicuous in young; margin of dorsal fin orange; caudal fin purplish with
inner margin of fork pale.
Size: Maximum standard length 50 cm or more, commonly to about 30 cm.
Habitat, biology, and fisheries: Occurs over rocky bottoms between about 140 and 360 m; off Guam,
caught most abundantly from about 180 to 270 m. Feeds mainly on fishes, shrimps, crabs, polychaetes,
cephalopods, and urochordates. Caught mainly with bottom longlines and deep handlines. A common food
fish in some areas such as Japan. It is one of the principal species in the Hawaiian offshore handline
fishery. Marketed mostly fresh.
to East Africa and northward
to southern Japan. A single
specimen, apparently of this
species, obtained from Vema
Seamount in the eastern
South Atlantic.
2914 Bony Fishes
Pristipomoides typus Bleeker, 1852

FAO names: En - Sharptooth jobfish; Fr - Colas dentu; Sp - Panchito dentón.
vertical through about middle of eye or slightly short of middle of eye. Upper and lower jaws both with an
outer row of conical teeth, a few of the anteriormost teeth enlarged into canines, and an inner band of
villiform teeth; vomer and palatines with teeth, those on vomer in chevron-shaped patch; no teeth on
tongue. Maxilla without scales or longitudinal ridges. Interorbital region flattened. First gill arch with
8 or 9 gill rakers on upper limb, 15 to 17 on lower limb (total 23 to 26). Dorsal fin continuous, not deeply
incised near junction of spinous and soft portions. Last soft ray of both dorsal and anal fins well
produced, longer than next to last ray. Caudal fin forked. Pectoral fins long, a little shorter than length
of head. Dorsal fin with X spines and 11 soft rays. Anal fin with III spines and 8 soft rays. Pectoral-fin
rays 15 or 16. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales 48 to 50.
Colour: body and fins rosy; top of head with longitudinal brownish yellow vermiculations and spots
of brownish yellow; dorsal fin with wavy yellow lines.
on benthic invertebrates and fishes. Estimated maximum age is 11 years. Caught mainly with handlines.
A good food fish, sometimes seen in markets. Usually marketed fresh.
Distribution: W e s te rn P a-
cific from New Guinea to Su-
matra and northward to the
Ryukyu Islands; also known
from the Andaman Sea arm of
the eastern Indian Ocean; re-
ports from the western Indian
Ocean and Western Samoa
need verification.
Pristipomoides zonatus (Valenciennes, 1830) (Plate X, 72)

Frequent synonyms / misidentifications: Rooseveltia brighami (Seale, 1901) / None.
FAO names: En - Obliquebanded snapper; Fr - Colas bagnard; Sp - Panchito rayado.
vertical through about middle of eye or slightly short of middle of eye. Upper and lower jaws both with an
outer row of sharp conical teeth (a few of anteriormost teeth in upper jaw enlarged and caniniform) and an
inner band of villiform teeth; vomer and palatines with teeth, those on vomer in chevron-shaped patch; no
teeth on tongue. Maxilla without scales or longitudinal ridges. Interorbital region flattened to slightly
convex. First gill arch with 5 to 8 gill rakers on upper limb, 12 to 14 on lower limb (total 17 to 21).
Dorsal fin continuous, not deeply incised near junction of spinous and soft portions. Last soft ray of both
dorsal and anal fins well produced, longer than next to last ray. Caudal fin forked. Pectoral fins long,
a little shorter than length of head. Dorsal fin with X spines and 11 (occasionally 10) soft rays. Anal fin
with III spines and 8 soft rays. Pectoral-fin rays 15 or 16. Membranes of dorsal and anal fins without
scales. Tubed lateral-line scales 63 to 67. Colour: overall pink or reddish with alternating oblique red
and yellow bars on side of body; dorsal and caudal fins yellow, other fins pink.
Habitat, biology, and fisheries: Occurs over rocky bottoms at depths between about 70 and 350 m; most
abundant between about 100 and 200 m. Feeds on fishes, shrimps, crabs, cephalopods, miscellaneous
benthic invertebrates, and pelagic organisms, including urochordates. Caught mainly with handlines. It is
one of the principal species in the Hawaiian offshore handline fishery. A good food fish occasionally seen
in markets where it is sold
fresh.
and Tahiti to East Africa and
from southern Japan south-
ward to Australia and Lord
Howe Island.
2916 Bony Fishes
Randallichthys filamentosus (Fourmanoir, 1970) (Plate X, 73)

Frequent synonyms / misidentifications: Etelis filamentosus Fourmanoir, 1970; E. nudimaxillaris
Yoshino and Araga, 1975 / Species of Etelis.
FAO names: En - Randall’s snapper; Fr - Vivaneau de Randall; Sp - Pargo de Randall.
close together and to eye. Lower jaw protruding. Premaxillae not protrusible, fixed by a frenum. Maxilla
ending posteriorly somewhat short of vertical through middle of eye. Jaws each with a series of small
conical teeth, larger anteriorly where on each side one to a few teeth may be canine or canine-like; larger
teeth on jaws needle sharp; vomer and palatines with teeth, those on vomer in roughly chevron-shaped
patch with apex blunt and directed anteriorly. Maxilla without scales, but with a series of about 4 to 9
well-developed longitudinal bony ridges. Interorbital region flattened. Gill openings extending well
anterior to orbit. First gill arch with 5 to 9 gill rakers on upper limb, 14 to 16 on lower limb (total 19
to 23). Dorsal fin continuous, indented just anterior to junction of spinous and soft portions, but
not as deeply incised as in species of Etelis. Last soft ray of dorsal fin shorter than next to last in
young, becoming slightly longer than next to last ray in older individuals; last soft ray of anal fin
equal to or slightly longer than next to last. Caudal fin lunate; lower lobe of caudal fin with some
rays excessively produced and filamentous in young. Pectoral fins short, 55 to 66% of head length.
Dorsal fin with X spines and 11 soft rays. Anal fin with III spines and 9 soft rays. Pectoral-fin rays
16 or 17. Membranes of dorsal and anal fins without scales. Tubed lateral-line scales 48 or 49. Larger
individuals quite oily. Colour: young with several dark bars; adults with head and iris rosy; body rosy with
suffusion of yellowish pigment dorsolaterally; dorsal fin mostly greyish; other fins, except pelvic fins, mostly
rosy with varying amounts of yellow; pelvic fins mostly yellowish.
Habitat, biology, and fisheries: Inhabits rocky bottoms in relatively deep water (150 to 380 m). Caught
mainly with handlines, but in relatively small quantities. Flesh is of good quality. Marketed fresh.
from off Hawaii, Cook Is-
lands, New Caledonia,
Guam, Ogasawara Islands,
and Okinawa, but probably
occurring in many other parts
of the western and central Pa-
cific.
Symphorichthys spilurus (Günther, 1874) (Plate X, 74)

FAO names: En - Sailfin snapper; Fr - Vivaneau voilier; Sp - Pargo velero.
D i a g n o s t i c c h a r a ct e rs :
Body deep, laterally com-
pressed. Head extremely
blunt (i.e. anterior profile
quite steep) in adults. Lacri-
mal (suborbital region) very
broad in adults. Nostrils on
each side of snout not very
close together. Mouth termi-
nal, jaws about equal. Pre-
maxillae protrusible. Maxilla
reaching vertical through
about middle of eye. Upper
jaw with a row of submolar
teeth; lower jaw with about
3 rows of smaller submo-
lars; anterior ends of both
jaws with an outer row of
s h o r t c a n i n e s, sl ig h t ly
larger than the other teeth;
vomer toothless; upper and
lower pharyngeal bones en-
l ar g ed an d bearin g en-
larged molariform teeth.
Maxilla without scales or adult
longitudinal ridges. First
gill arch with 4 to 7 gill rak- juvenile
ers on upper limb, 14 or 15
on lower limb (total 18 to
22). Dorsal fin continuous, notched slightly near junction of spinous and soft portions. Spinous part of
dorsal fin distinctly lower than soft part. One or more anterior soft rays of dorsal and anal fins
produced into filaments. Last soft ray of dorsal and anal fins not produced. Caudal fin emarginate
to lunate in adults, truncate in juveniles. Pectoral fins somewhat longer than head. Dorsal fin with X
spines and 17 to 19 soft rays. Anal fin with III spines and 11 soft rays. Pectoral-fin rays 16 or 17. Soft
dorsal and anal fins sheathed in scales basally. Tubed lateral-line scales 52 to 59. Colour: overall
yellowish, with a series of bright blue stripes on sides and head; a prominent black spot with a pale
margin on dorsal surface of caudal peduncle; sometimes with an orange bar across interorbital region
and an orange bar above pectoral-fin base; juveniles light brown above and whitish below with broad black
stripe along midbody.
Size: Maximum standard length about 50 cm, commonly to about 25 cm.
Habitat, biology, and fisheries: Found over sand bottoms in the vicinity of coral reefs at depths between
about 5 and 60 m. Usually seen solitarily. Feeds on fishes and sand-dwelling crustaceans and molluscs.
Caught mainly with handlines and bottom trawls. Occasionally seen in markets.
Distribution: Western mar-
gin of the Pacific including
New Caledonia, the Great
Barrier Reef of Australia, New
Guinea, Sulawesi (Celebes),
Admiralty Islands, Palau Is-
lands, the Philippines, and the
Ryukyu Islands; also known
from Rowley Shoals in the
eastern Indian Ocean off
Broome, Western Australia.
2918 Bony Fishes
Symphorus nematophorus (Bleeker, 1860)

Frequent synonyms / misidentifications: Glabrilutjanus nematophorus (Bleeker, 1860); Paradicichthys
venenatus Whitley, 1930 / None.
FAO names: En - Chinamanfish; Fr - Vivaneau diable; Sp - Pargo diablo.
adult
juvenile
Diagnostic characters: Body moderately deep, laterally compressed. Anterior profile relatively steep, but
not nearly as steep as in Symphorichthys. Eye located well above mouth. Lacrimal (suborbital region)
broad. Deep groove running obliquely downward from eye to anterior nostril or a little beyond. Nostrils
on each side of snout not very close together. Mouth terminal, jaws almost equal. Premaxillae protrusible. Maxilla
reaching vertical through about middle of eye. Teeth in jaws in narrow bands, with the outer teeth enlarged;
canine teeth present at anterior end of upper jaw; vomer toothless; tongue with a patch of granular teeth.
Maxilla without scales or longitudinal ridges. First gill arch with 4 to 6 gill rakers on upper limb, 10 to
15 on lower limb (total 15 to 21). Dorsal fin continuous, notched slightly near junction of spinous and soft
portions. Spinous part of dorsal fin distinctly lower than soft part. One or more anterior soft rays of dorsal
fin produced into long filaments in juveniles. Last soft ray of dorsal and anal fins not produced. Caudal
fin emarginate. Pectoral fins 70 to 90% head length. Dorsal fin with X spines and 14 to 17 soft rays. Anal
fin with III spines and 9 or 10 soft rays. Pectoral-fin rays 16 or 17. Soft dorsal and anal fins sheathed in
scales basally. Tubed lateral-line scales 50 to 56. Colour: adults mainly reddish, frequently with lighter
blotching or transverse bars; young brown or yellowish on back and upper sides, yellowish below, and with
bright blue stripes on side.
Size: Maximum standard length about 1 m, commonly to 35 cm.
Habitat, biology, and fisheries: Inhabits coral reefs from shallow water to depths of at least 50 m. Occurs
solitarily. Feeds mainly on fishes. Caught mainly with handlines; also speared by divers. An excellent food
fish, but in Queensland it is considered dangerous due to the chance of ciguatera poisoning. Frequently
seen in markets.
Distribution: W e st e r n P a-
cific from New Caledonia
throughout the Indo-Austra-
lian Archipelago (from New
Guinea and northern Austra-
lia to the Malay Peninsula)
and northward to the Ryukyu
Islands; also known from the
eastern Indian Ocean off
northwestern Australia.
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Perciformes: Percoidei: Caesonidae 2919
CAESIONIDAE
Fusiliers
by K.E. Carpenter
D iagnostic characters: Oblong to fusiform, moderately compressed, medium-sized to small (to about
50 cm) lutjanoid fishes; longitudinal axis from tip of snout to middle of caudal fin passing through
centre of eye. Eye moderately large, its diameter longer than snout length. Mouth small and highly
protrusible; 1 or 2 finger-like postmaxillary processes on dorsoposterior surface of premaxilla (Figs 1 and
2); angle of jaw oblique, about 40° to horizontal. Dentition variously reduced; small or minute conical
teeth; premaxillae, vomer, and palatines with or without teeth. Caudal fin deeply forked. Margin of dorsal
and anal fins more or less evenly sloping; third or fourth dorsal-fin spines longest; second or third anal-fin
spines longest, remaining spines and rays gradually decreasing in length (except in Dipterygonotus with
dorsal fin profile not evenly sloping, last IV-V dorsal-fin spines small and nearly separate, connected only
at their bases by membrane, and dorsal-fin rays much longer than these spines). Dorsal fin with X to XV
slender weak spines and 8 to 22 soft rays; anal fin with III spines and 9 to 13 soft rays; pelvic fins
with I spine and 5 soft rays; pectoral fins with 16 to 24 rays; caudal fin distinctly forked, with pointed
lobes. Branchiostegal rays 7. Scales moderate to small, weakly ctenoid; lateral-line scales 45 to 88; scale
rows on body running horizontally; dorsal and anal fins with scales except for Gymnocaesio
gymnoptera and Dipterygonotus balteatus. Ascending premaxillary process a separate ossification from
premaxilla; ethmo-maxillary ligament absent; a separate A1’ section of the adductor mandibulae which
originates on the subocular shelf. Supraneural configuration 0/0/0+2/1+1/, /0+0/0+2/1+1/, or /0+0/2/1+1/.
Epineurals 10 to 15. Procurrent caudal-fin rays typically 7 to 10. Hypurals 1-2 and 3-4 typically fused in all
species (except some juveniles). Openings in external wall of pars jugularis 2 to 5. Colour: sides with or
without longitudinal stripes; caudal fin either without markings, with a blackish blotch on tips of lobes, or
with a longitudinal blackish streak in middle of each lobe; axil of pectoral fins black.
X-XV spines
8-22 soft rays
mouth small,
protractile
9-13 soft rays
Habitat, biology, and fisheries: Mostly on coral reefs; occur near the surface to depths of 60 m. Feed by
picking zooplankton. Schooling, often in groups with more than one species. Major importance in coral-reef
fisheries where drive-in nets are employed. From 1990 to 1995, the FAO Yearbook of Fishery Statistics
reports a yearly catch of around 24 600 and 55 400 t of Caesionidae from the Western Central Pacific.
However, much larger catches are expected in small-scale fisheries for which statistics are not available.
Marketed mostly fresh but sometimes dried and as fermented fish paste.
2920 Bony Fishes

Lutjanidae: difficult to distinguish from Caesionidae
on the basis of any single external character, but
most members of the Lutjanidae are deeper bodied,
have the eye well above the horizontal axis of the
body, and lack a strongly forked caudal fin; those
lutjanid genera that have the horizontal passing near
the centre of the eye either have scale rows running
obliquely upward, fewer than 9 anal-fin rays, no
scales on dorsal and anal fins, or have the caudal
fin much less deeply forked. Paracaesio (Lutjanidae)
Pinjalo (Lutjanidae) Lutjanus (Lutjanidae)
Nemipteridae: eye above horizontal axis of body in most species; 9 dorsal-fin rays (13 to 22 in all caesionids
except for Dipterygonotus balteatus, which usually has 10 soft rays but rarely 8 or 9) and only 7 or 8 anal-fin
rays (9 to 13 in Caesionidae).
less than 10 rays
7-8 rays
Nemipterus (Nemipteridae) Pentapodus (Nemipteridae)
Lethrinidae: eye always above, and usually well above horizontal axis of body; base of soft part of dorsal
fin generally shorter than base of spinous part (soft part generally longer than spinous part in Caesionidae);
8 to 10 soft rays in anal fin (usually 11 or more in caesionids except in Dipterygonotus balteatus which
has 9 to11); usually enlarged canines in front of jaws, sometimes lateral molars.
Emmelichthyidae: superficially similar but caught in deep water; maxilla fully scaly (not scaly in Caesioni-
dae); dorsal fin with XI spines and usually 12 soft rays.
canines
Lethrinidae Emmelichthyidae
Key to the species of Caesionidae occurring in the area

Remarks on key characters: there has been much confusion with species names of caesionids because
preserved specimens are difficult to identify. The species of this family are easily distinguished on the basis
of colours of living or fresh specimens, but these colour patterns usually do not preserve well. The
morphological features of caesionids are often difficult to use in identification due to the feebleness of the
structures. Scales readily detach, and rays are slender and easily broken; teeth are small and weak and
often require staining for proper characterization. When these elements are not broken or detached, they
are useful taxonomically. The overlaps in counts, however, are such that, based on meristics alone, there
is often a possibility of misidentification. For this reason, when life colours are not known, it is better to
base identifications on a number of specimens.
1a. A single postmaxillary process; posterior end of maxilla blunt, its greatest depth posterior
to end of premaxilla (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Caesio) ® 2
1b. Two postmaxillary processes; posterior end of maxilla tapered, its greatest depth ante-
rior to end of premaxilla (Fig. 2) . . . . . . . . . . (Dipterygonotus, Gymnocaesio, Pterocaesio) ® 6
2 postmaxillary
only 1 postmaxillary processes
process
end of premaxilla end of premaxilla maxilla

maxilla maxilla maxilla mouth closed mouth protracted
mouth closed mouth protracted
Fig. 1 Caesio Fig. 2 Dipterygonotus
2a. Anal fin usually with 11 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3

2b. Anal fin usually with 12 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Dorsal fin usually with 15 soft rays; supratemporal band of scales confluent at dorsal
midline (Fig. 3); caudal fin without any prominent blackish markings (Fig. 3) . . . . . . Caesio cuning
3b. Dorsal fin usually with 14 soft rays; supratemporal band of scales interrupted at dorsal
midline by a thin scaleless zone (Fig. 4); tips of caudal-fin lobes with a blackish blotch
(Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caesio lunaris
supratemporal supratemporal
band of scales band of scales
continuous interrupted
Fig. 3 Caesio cuning Fig. 4 Caesio lunaris

2922 Bony Fishes
4a. Lateral-line scales 57 to 65; scale rows on spinous portion of dorsal fin horizontal
(Fig. 5a); caudal fin not yellow, each lobe with a median blackish streak (Fig. 6) . Caesio caerulaurea
4b. Lateral-line scales 51 to 61; upper scale rows on spinous portion of dorsal fin usually
oblique (Fig. 5b); caudal fin yellow in life, without blackish markings . . . . . . . . . . . . . . . . ® 5
scale rows oblique

scale rows horizontal
a) b)
Fig. 5 dorsal fin Fig. 6 Caesio caerulaurea
5a. Body yellow dorsally, blue on side, the demarcation horizontal from interorbital space
across upper third of body; predorsal and supratemporal region not considerably darker
than dorsal part of caudal peduncle in alcohol-preserved specimens (Fig. 7) . . . Caesio xanthonota
5b. Body yellow dorsally, blue on side, the demarcation oblique from slightly anterior to origin
of dorsal fin to lower posterior part of caudal peduncle (in western and Central Pacific
the yellow region in adults is restricted to caudal peduncle); predorsal region, especially
supratemporal and interorbital region, considerably darker than dorsal part of caudal
peduncle in alcohol-preserved specimens (Fig. 8) . . . . . . . . . . . . . . . . . . . . . Caesio teres
Fig. 7 Caesio xanthonota Fig. 8 Caesio teres
6a. Dorsal and anal fins scaly; premaxilla with small conical teeth, sometimes restricted to
front of jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Pterocaesio) ® 7
6b. Dorsal and anal fins without scales; premaxilla without teeth . (Dipterygonotus, Gymnocaesio) ® 15
7a. Dorsal fin with XI or XII (rarely X)

spines and 19 to 22 soft rays; a
blackish streak in each caudal-fin
lobe (Fig. 9) . . . . . . . . . . Pterocaesio tile
7b. Dorsal fin with X or XI (usually X)
spines and 14 to 16 soft rays; tips
of caudal-fin lobes with a black
blotch . . . . . . . . . . . . . . . . . . . . ® 8
Fig. 9 Pterocaesio tile

8a. Dorsal peduncular scales usually 11 (rarely 12) (Fig. 10); lateral-line scales 62 to 75;
pectoral-fin rays 17 to 21 (most frequently 19 or 20); scales above lateral line to dorsal-fin
origin usually 8 or 9 (Fig. 11); scales below lateral line to anal-fin origin usually 13 to 17
(Fig. 11); side with or without stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
8b. Dorsal peduncular scales usually 12 or 13 (rarely 11 or 14) (Fig. 10); lateral-line scales
66 to 88; pectoral-fin rays 20 to 24 (always most frequently 21 or above); scales above
lateral line to dorsal-fin origin 9 to 11 (Fig. 11); scales below lateral line to anal-fin origin
usually 16 to 18 (Fig. 11); side with 1 or more longitudinal stripes . . . . . . . . . . . . . . . . ® 11
upper peduncular scales upper scale rows

(circumference scale row count from just above lateral
line on 1 side to just above lateral line on other side) lateral line
lower peduncular scales

(circumference scale row count including lateral lower scale rows
line row on 1 side and below to and including
lateral line row on other side)
Fig. 11 counts of scale rows above and below lateral line
Fig. 10 scale counts on caudal peduncle
(do not include lateral line or midline scale rows)
9a. Pectoral-fin rays usually 19 to 21 (most frequently 20); 3 light and 3 dark stripes on upper
side in life (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterocaesio trilineata
9b. Pectoral-fin rays 17 to 20 (most frequently 19); side without stripes or with at most, 2
stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Body without stripes on side, its colour reddish or greenish blue . . . . . . . . . Pterocaesio pisang
10b. Two yellow stripes on side in life, one on dorsal midline on nape and along base of dorsal
fin, and the other, which is broader anteriorly, directly below lateral line for most its
length, except above lateral line on caudal peduncle (Fig. 13) . . . . . . . . Pterocaesio chrysozona
Fig. 12 Pterocaesio trilineata Fig. 13 Pterocaesio chrysozona
11a. Lateral-line scales 74 to 88; pec-

toral-fin rays 21 to 23 (most fre-
quently 22); a broad yellow stripe
on side in life, wider anteriorly,
covering lateral line for most its
length (Fig. 14) . . . . . Pterocaesio lativittata
11b. Lateral-line scales 66 to 80; pec-
toral-fin rays either 20 to 22 (most
frequently 21) or 22 to 24 (most
frequently 23); either a thin yellow
stripe on side or a large yellow Fig. 14 Pterocaesio lativittata
blotch above pectoral-fin base . . . . . . ® 12
2924 Bony Fishes
12a. Pectoral-fin rays 22 to 24 (most frequently 23); 2 thin yellow stripes on side, the lower
stripe covering lateral line for most its length, above lateral line on caudal peduncle, the
upper stripe mostly 1 or 2 scales below the dorsal profile of the body (Fig. 15) . . . Pterocaesio marri
12b. Pectoral-fin rays 20 to 22 (most frequently 21); 1 or 2 yellow lines or a large yellow blotch
on side (if there are 2 lines, the lower one is distinctly below lateral line for most its length,
except on caudal peduncle where it is above lateral line) . . . . . . . . . . . . . . . . . . . . . ® 13
13a. A large yellow blotch above pectoral fins in life; no stripes on side (Fig. 16) . . . Pterocaesio randalli
13b. No large yellow blotch above pectoral fins; 1 or 2 yellow stripes on side . . . . . . . . . . . . . ® 14
Fig. 15 Pterocaesio marri Fig. 16 Pterocaesio randalli
14a. A single thin yellow stripe on side covering lateral line for most its length, on caudal
peduncle above lateral line (Fig. 17) . . . . . . . . . . . . . . . . . . . . . . . Pterocaesio tessellata
14b. Two thin longitudinal stripes on side, the lower distinctly below lateral line for most its length,
on caudal peduncle above lateral line, the upper following dorsal profile (Fig. 18) . .Pterocaesio digramma
Fig. 17 Pterocaesio tessellata Fig. 18 Pterocaesio digramma
15a. Dorsal fin with X or XI (usually X) spines and 14 to 16 (usually 15) soft rays, the fin not
deeply notched; anal fin with III spines and 11 to 13 (usually 12) soft rays; pectoral-fin
rays 20 to 22 (Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocaesio gymnoptera
15b. Dorsal fin with XII to XV (usually XIV) spines and 8 to 11 (usually 10) soft rays, the fin deeply
notched, the last few spines joined only at base by fin membrane; anal fin with III spines and
9 to 11 (usually 10) soft rays; pectoral-fin rays 16 to 19 (Fig. 20) . . . . . . . . Dipterygonotus balteatus
Fig. 19 Gymnocaesio gymnoptera Fig. 20 Dipterygonotus balteatus


The symbol 0is given when species accounts are included.
0
0
Caesio caerulaurea Lacepède, 1801
0
Caesio cuning (Bloch, 1791)
0
Caesio lunaris Cuvier, 1830
Caesio teres Seale, 1906
0 Caesio xanthonota Bleeker, 1853
0 Dipterygonotus balteatus (Valciennes, 1830)
0 Gymnocaesio gymnoptera (Bleeker, 1856)
0 Pterocaesio chrysozona (Cuvier, 1830)
0
0
Pterocaesio digramma (Bleeker, 1865)
0
Pterocaesio lativittata Carpenter, 1987
0
Pterocaesio marri Schultz, 1953
0
Pterocaesio pisang (Bleeker, 1853)
0
Pterocaesio randalli Carpenter, 1987
Pterocaesio tessellata Carpenter, 1987
0
0
Pterocaesio tile (Cuvier, 1830)
Pterocaesio trilineata Carpenter, 1987
References
Carpenter, K.C. 1987. Revision of the Indo-Pacific fish family Caesionidae (Perciformes: Lutjanoidea), with descriptions
of five new species. Indo-Pac. Fish., (15):56 p.
Carpenter, K.C. 1988. Fusilier fishes of the world. An annotated and illustrated catalogue of caesionid species known
to date. FAO Fish. Synop., (125)Vol. 8:75 p.
2926 Bony Fishes
Caesio caerulaurea Lacepède, 1801 (Plate XI, 75)

Frequent synonyms / misidentifications: None / Caesio varilineata non Carpenter, 1987.
FAO names: En - Blue-and-gold fusilier; Fr - Caesio azuror; Sp - Fusilero azur.
Diagnostic characters: Body moderately deep, fusiform, elongate, and moderately compressed. Ratio of
eye diameter to head length usually around 3.4 to 4.2; a single postmaxillary process; small conical
teeth in jaws, vomer, and palatines. Dorsal fin with X spines and 15 (rarely 14 or 16) soft rays; anal fin
with III spines and 12 (rarely 13) soft rays; pectoral fins with 20 to 22 (rarely 19) rays. Scales in lateral
line 57 to 65 (most frequently around 61); upper peduncular scale rows usually 11, lower peduncular
scale rows usually 15; scale rows above lateral line to origin of dorsal fin 8 to 10 (most frequently 9); scale
rows below lateral line to origin of anal fin usually 15 to 17; scale rows on cheek usually 4; predorsal scales
usually 22 to 25; dorsal and anal fins scaly, the spinous part of dorsal fin with about 3/4 of its greatest
height covered with scales, the scale rows on spinous part of dorsal fin horizontal; supratemporal
bands of scales often interrupted at dorsal midline by a scaleless zone, always a V-shaped scaleless
zone anteriorly at midline intruding between the supratemporal band of scales. Colour: upper body
bluish, lower body white to pale bluish; a single yellow stripe directly above lateral line except on
caudal peduncle where it is about 1 scale above lateral line, the yellow stripe 2 or 3 scales wide,
bordered directly above and below by a white or light blue stripe which is about 1 scale wide
(juveniles sometimes have another black stripe between the yellow stripe and each of the whitish outer
stripes); caudal-fin lobes with a black median streak, the tips not markedly darker than the streak; outer
margin of each caudal-fin lobe often bordered in white, and inner margins pale; pectoral, pelvic, and anal
fins white; axil of pectoral fins black, and a black triangular patch on outside upper base; dorsal fin light
blue to pale with a black distal border.
Habitat, biology, and fisheries: Inhabits coastal areas, primarily around coral reefs to depths of around
40 m. Feeds by picking zooplankton. Schooling, often feeding in large midwater aggregations; commonly
schools together with Caesio varilineata, C. striata, and Pterocaesio tile. Moderately important in coastal
fisheries. Common in markets in Indonesia and the Philippines. Caught by drive-in nets, gill nets, traps,
trawls, and handlines. Marketed fresh and sometimes dried-salted. Juveniles are important as tuna baitfish
in some areas.
the tropical Indo-West Pacific,
from East Africa, including the
Red Sea but not the Persian
Gulf, to Samoa; north to south-
ern Japan and south to Mauri-
tius and New Caledonia.
Caesio cuning (Bloch, 1791) (Plate XI, 76)

Frequent synonyms / misidentifications: Caesio erythrogaster Cuvier, 1830 / None.
FAO names: En - Redbelly yellowtail fusilier; Fr - Caesio ventre rouge; Sp - Fusilero vientre colorado.
Diagnostic characters: Body fairly deep and compressed. A single postmaxillary process; small,
conical teeth in jaws, vomer, and palatines. Dorsal fin with X spines and 15 (rarely 14 or 16) soft rays;
anal fin with III spines and 11 (rarely 10 or 12) soft rays; pectoral fins with 18 or 19 (rarely 17 or 20)
rays. Scales in lateral line 45 to 51 (most frequently 49); upper peduncular scales usually 9 or 10, lower
peduncular scales usually 13 or 14; scales above lateral line to origin of dorsal fin usually 8 or 9; scales
below lateral line to origin of anal fin usually 15 to 17; 4 or 5 scale rows on cheek; predorsal scales usually
21 to 25; dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest spinous height covered with
scales; supratemporal band of scales confluent at dorsal midline. Colour: caudal fin, upper caudal
peduncle and posterior portion of back yellow; upper body, where not yellow, greyish blue; lower
sides and belly white or pinkish; pectoral, pelvic, and anal fins white to pink; axil and upper base of pectoral
fins black; dorsal fin yellow posteriorly and greyish blue anteriorly.
Habitat, biology, and fisheries: Inhabits coastal areas, usually over rocky and coral reefs to depths of
around 60 m. Of all the caesionids, Caesio cuning appears most tolerant of murky water; it is usually the
most abundant caesionid in reef areas characterized by low underwater visibility. This species ranges
widely between reefs as it is often captured by trawl net over soft bottom. A schooling fish, found in small
to large aggregations. Feeds primarily on zooplankton in midwater. A moderately important food fish in
many areas. Common in markets in the Gulf of Thailand where it is taken in trawls; also caught in Indonesia,
the Philippines, and Papua New Guinea by a variety of methods including drive-in nets, fish traps, and gill
nets. Marketed mostly fresh.
Distribution: Tropical east-
ern Indian Ocean to the west-
ern Pacific, from Sri Lanka to
Vanuatu, and from southern
Japan to northwestern and
northeastern Australia.
2928 Bony Fishes
Caesio lunaris Cuvier, 1830 (Plate XI, 77)

FAO names: En - Lunar fusilier; Fr - Caesio croissant; Sp - Fusilero luna.
Diagnostic characters: Body fairly deep, fusiform, and moderately compressed. A single postmaxillary
process; small conical teeth in jaws, vomer, and palatines. Dorsal fin with X spines and 14 (rarely 13
or 15) soft rays; anal fin with III spines and 11 (rarely 10) soft rays; pectoral fins with 19 or 20 (rarely
18 or 21) rays. Scales in lateral line 45 to 53 (most frequently 49); upper peduncular scales usually 10 or
11, lower peduncular scales usually 14 or 15; scales above lateral line to origin of dorsal fin usually 8 or
9; scales below lateral line to origin of anal fin usually 15 to 19; usually 4 scale rows on cheek; predorsal
scales usually 20 to 23; dorsal and anal fins scaly, the dorsal fin with about 3/4 of its greatest spinous
height covered with scales; supratemporal band of scales generally interrupted at dorsal midline by
a narrow scaleless zone. Colour: body bluish, belly paler than upper sides; tips of caudal-fin lobes,
axil of pectoral fins, and upper base of pectoral fins black; caudal fin blue (except in juveniles where
caudal fin and portions of caudal peduncle often yellow); pectoral, pelvic, and anal fins white to pale blue;
dorsal fin bluish.
Size: Maximum size about 40 cm.
Habitat, biology, and fisheries: Found in coastal areas, mainly on or near coral reefs. They feed on
zooplankton in large aggregations. Unlike most other caesionids, this species changes colour during
development. As juveniles, they stay close to the reef and school with other juvenile caesionids; commonly
with Caesio cuning. During this stage, they typically have a yellow caudal fin and peduncle. As adults, they
mostly feed a little further off the reef than other fusiliers, in deep, clear water; returning to the reef only at
night to shelter. Adults loose the yellow coloration, perhaps because it is conspicuous in clear, deep water.
Of minor importance to fisheries in most areas. Caught primarily by drive-in net, seines, and fish traps.
the Indo-West Pacific, from
East Africa, including the Red
Sea and the Persian Gulf, to
southern Japan and the Solo-
mon Islands.
Caesio teres Seale, 1906 (Plate XI, 78)

Frequent synonyms / misidentifications: None / Caesio xanthonota non Bleeker, 1853.
FAO names: En - Yellow-and-blueback fusilier; Fr - Fusilier à dos jaune et bleu; Sp - Fusilero amarillo
azulado.
Diagnostic characters:
Body moderately deep,
f u s i f o r m , a n d c o m-
pressed. A single post-
m a x i l l a r y p r o c e s s;
small teeth in jaws,
vomer, and palatines.
Dorsal fin with X spines
and 15 (rarely 14 or 16)
soft rays; anal fin with
III s p i n e s a n d 1 2
(rarely 13) soft rays;
pectoral fins usually with
20 to 22 (most frequently
21) rays. Scales in lat-
eral line 51 to 61 (most
frequently 55); upper
peduncular scales 11 to
13, lower peduncular
scales usually 15 or 16;
scales above lateral line
to origin of dorsal fin usu-
ally 8 to 10 (most fre-
quently 8); scales below
lateral line to origin of
anal fin usually 17 to 20;
4 or 5 scale rows on
cheek; predorsal scales
usually 21 to 25; dorsal
and anal fins scaly, the
dorsal fin with about 2/3
of its greatest spinous
height covered with scales; supratemporal band of scales interrupted at dorsal midline by a narrow
scaleless zone. Colour: bright yellow on caudal fin, caudal peduncle, and body above a diagonal
from just anterior to dorsal-fin origin to ventral origin of caudal peduncle, except in large individuals
particularly in the western Pacific where the yellow does not extend as far anteriorly; remainder of
upper two-thirds of body bright blue, the lower third silvery white; axil and upper base of pectoral fins black;
pectoral, pelvic, and anal fins white; dorsal fin bluish proximally and yellow distally.
Habitat, biology, and fisheries: Found primarily around coral reefs, with a preference for coraline lagoons.
A rapidly moving schooling fish which ranges widely around reefs. Feeds on zooplankton in large midwater
groups. Schools together with other caesionids; most often with Caesio xanthonota. It spawns in large
aggregations around the full moon. Eggs pelagic, spherical, unpigmented, usually with a single transparent
oil globule, and a diameter of
0.77 to 0.78 mm. Of minor to
moderate importance in
coastal fisheries. Caught by
drive-in nets, gill nets, handli-
nes, and traps. Marketed
mostly fresh.
Distribution: W i d e s p r e a d
tropical Indo-West Pacific,
from East Africa, not including
the Red Sea or the Persian
Gulf, to the Line Islands.
2930 Bony Fishes
Caesio xanthonota Bleeker, 1853 (Plate XI, 79)

Frequent synonyms / misidentifications: None / Caesio teres non Seale, 1906.
FAO names: En - Yellowback fusilier; Fr - Fusilier à dos jaune; Sp - Fusilero de lomo amarillo.
Diagnostic characters: Body moderately deep, fusiform, and compressed. A single postmaxillary
process; small conical teeth in jaws, vomer, and palatines. Dorsal fin with X spines and 15 (rarely 14) soft
rays; anal fin with III spines and 12 (rarely 11) soft rays; pectoral fins with 20 to 22 (most frequently
21) rays. Scales in lateral line 52 to 59 (most frequently 56); upper peduncular scales 11 or 12, lower
peduncular scales usually 15; scales above lateral line to origin of dorsal fin usually 9 or 10; scales below
lateral line to origin of anal fin usually 18 or 19; 4 or 5 scale rows on cheek; predorsal scales usually 21 to
23; dorsal and anal fins scaly, the dorsal fin with about 2/3 of its greatest spinous height covered with
scales; supratemporal band of scales interrupted at dorsal midline by a narrow scaleless zone. Colour:
upper third of body and head, including interorbital region, and caudal fin bright yellow, middle
third blue, lower third white; axil and upper base of pectoral fins black; pectoral, pelvic, and anal fins white;
dorsal fin yellow.
Habitat, biology, and fisheries: Inhabits coastal areas, primarily around coral reefs. A schooling fish;
feeds on zooplankton in large midwater aggregations. This species ranges widely among reefs during the
day but shelters on the reef at night. It sometimes schools together with Caesio teres, which has a similar
colour pattern. C. xanthonota appears to prefer the coraline lagoon habitat more than most other species
of caesionids. Of minor to moderate importance in coastal fisheries. Caught by gill net, traps, and handline.
Distribution: Primarily Indian
Ocean, from East Africa, not
including the Red Sea or the
Persian Gulf, to Indonesia.
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Dipterygonotus balteatus (Valenciennes, 1830) (Plate XI, 80)

Frequent synonyms / misidentifications: Dipterygonotus leucogrammicus Bleeker, 1849 / None.
FAO names: En - Mottled fusilier; Fr - Fusilier marbré; Sp - Fusilero jaspeado.
Diagnostic characters: Body slender, fusiform, elongate, and moderately compressed. Two postmaxil-
lary processes; small conical teeth on dentary and vomer; premaxilla and palatines without teeth.
Dorsal fin deeply notched, with XIV spines and 8 to 11 soft rays; anal fin with III spines and usually
9 or 10 soft rays; pectoral fins with 16 to 19 rays; procurrent caudal-fin rays typically 7 or 8. Scales in
lateral line 68 to 80; upper peduncular scales 11 to 14, lower peduncular scales 15 to 18; scales above
lateral line to origin of dorsal fin 9 to 11; scales below lateral line to origin of anal fin 15 to 18; usually 6 to
9 scale rows on cheek; predorsal scales usually 29 to 34; dorsal and anal fins without scales,
supratemporal band of scales indistinct, scales in interorbital region near ascending premaxillary process.
Colour: upper body brownish bronze; a thin, straight, tan stripe about 1 scale wide from orbit to caudal
fin, directly above lateral line for anterior half of its length, about 2 scales above lateral line on caudal
peduncle; above and parallel to this stripe 2 thin, irregular, and usually interrupted stripes of same colour;
lower body silvery white; dorsal, anal, pelvic, and pectoral fins clear to pinkish; axil of pectoral fins black;
caudal fin tan to pinkish; when captured at night, the entire body and fins are often reddish.
Size: The smallest of all fusiliers, attaining a maximum size of about 14 cm.
Habitat, biology, and fisheries: This is the only caesionid that is primarily nearshore pelagic rather than
most typically found near coral reefs. In the markets of the Philippines, Dipterygonotus balteatus is most
frequently found mixed with sardines and anchovies. As juveniles, they school together with other juvenile
caesionids on coral reefs. Feeds on zooplankton. Occasionally marketed fresh along with sardines and
anchovies in the Philippines. Caught at night using lights and dip nets.
Distribution: Widespread
Indo-Pacific, from East Africa,
not including the Red Sea or
Persian Gulf, to the Solomon
Islands.
2932 Bony Fishes
Gymnocaesio gymnoptera (Bleeker), 1856 (Plate XI, 81)

FAO names: En - Slender fusilier; Fr - Fusilier élégant; Sp - Fusilero elegante.
Diagnostic characters: Body slender, fusiform, elongate, and moderately compressed. Two postmaxil-
lary processes; posterior end of maxilla tapered, its greatest depth anterior to posterior end of the
premaxilla; small conical teeth on dentary and vomer; premaxilla and palatines without teeth. Dorsal
fin with X (rarely XI) spines and 15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely
11 or 13) soft rays; pectoral fins with 20 to 22 rays. Scales in lateral line 64 to 74 (most frequently 70);
upper peduncular scales usually 11, lower peduncular scales usually 14 or 15; scales above lateral line to
origin of dorsal fin 7 to 9; scales below lateral line to origin of anal fin usually 13 or 14; usually 4 scale rows
on cheek; predorsal scales usually 23 to 26; dorsal and anal fins without scales. Colour: upper body
bluish green, the centres of scales lighter, giving a striped appearance; a single yellow or brown stripe
about 1 scale wide covering lateral line for most of its length, running dorsal to lateral line on caudal
peduncle; often a bright blue longitudinal band directly below lateral line stripe covering up to 1/3 of side;
body silvery white ventrally; axil of pectoral fins black; pectoral, pelvic, dorsal, and anal fins white; caudal
fin dusky, the tips of lobes black.
Habitat, biology, and fisheries: Inhabits coastal areas ranging widely around coral reefs. Feeds on
zooplankton in large midwater aggregations. A schooling fish, often in groups with members of the genus
Pterocaesio. Of minor importance to fisheries. Occasionally caught with traps and drive-in nets and
marketed fresh in the Philippines. Used as a baitfish for tuna fisheries.
Indo-West Pacific, from East
Africa, including the Red Sea,
to Fiji.
Pterocaesio chrysozona (Cuvier, 1830) (Plate XI, 82)

FAO names: En - Goldband fusilier; Fr - Caesio ceinture d’or; Sp - Fusilero cinta dorada.
Diagnostic characters: Body fusiform, elongate, and moderately compressed. Two postmaxillary
processes; small conical teeth in jaws, vomer, and palatines. Dorsal fin with X (rarely XI) spines and
15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely 11 or 13) soft rays; pectoral fins with
17 to 20 (most frequently 19) rays. Scales in lateral line usually 64 to 69; upper peduncular scales
usually 11, lower peduncular scales usually 15; scales above lateral line to origin of dorsal fin 7 to 9; scales
below lateral line to origin of anal fin usually 14 to 16; usually 4 scale rows on cheek; predorsal scales
usually 23 to 26; dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest spinous height
covered with scales. Colour: upper body light blue to brownish, lower body white to pinkish; a bright
yellow band directly below lateral line for most of its length, from behind eye to base of caudal fin,
2 to 3 scales wide anteriorly, tapering to 1 scale in width on caudal peduncle where it is above lateral
line; a less conspicuous yellow stripe along dorsal midline; fins white to pinkish; axil of pectoral fins black;
dorsal fin slightly dusky distally; tips of caudal-fin lobes black.
Habitat, biology, and fisheries: Ranges widely around coral reefs in schools. Often found in groups with
other caesionids. Feeds on zooplankton in large midwater aggregations. Moderately important as a food
fish in some areas. Common in the markets of Philippines. Caught by drive-in net, gill net, traps, handline,
and, occasionally by trawl. A valuable tuna baitfish in some areas.
Africa, including the Red Sea
to eastern Australia.
2934 Bony Fishes
Pterocaesio digramma (Bleeker, 1865) (Plate XII, 83)

FAO names: En - Doublelined fusilier; Fr - Fusilier deux bandes jaunes; Sp - Fusilero de dos bandas.
processes; small conical teeth in jaws, vomer, and palatines. Dorsal fin with X spines and 15 (rarely
14 or 16) soft rays; anal fin with III spines and 12 (rarely 11) soft rays; pectoral fins usually with 20 to
22 rays. Scales in lateral line 66 to 75 (most frequently 72); upper peduncular scales usually 12 or
13, lower peduncular scales usually 16 or 17; scales above lateral line to origin of dorsal fin 9 to 11; scales
below lateral line to origin of anal fin usually 17 or 18; 4 or 5 scale rows on cheek; predorsal scales usually
26 to 28; dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest spinous height covered
with scales. Colour: body blue to greenish dorsally, white ventrally; 2 longitudinal stripes laterally, the
lower running from behind upper orbit about 1 scale below the lateral line to a vertical at about
middle of soft portion of dorsal fin where it crosses and usually passes above lateral line for the
length of caudal peduncle; upper stripe 1 to 2 scales below dorsal profile for most of length of body,
ending dorsally on caudal peduncle; fins white to pinkish; axil of pectoral fins black; dorsal fin slightly
dusky distally; tips of caudal-fin lobes black.
Habitat, biology, and fisheries: Found in coastal areas, primarily around coral reefs. A schooling fish,
often in mixed species groups containing other species of Pterocaesio. Feeds on zooplankton in midwater
aggregations.
Interest to Fisheries: Moderately important as a food fish. This species is one of the most common
caesionids in Philippine markets. Caught mostly by drive-in net, gill net, and traps. Marketed mostly fresh;
sometimes dried salted.
Distribution: Primarily West
Pacific, from Indonesia and
western Australia to New
Caledonia, north to southern
Japan.
Pterocaesio lativittata Carpenter, 1987 (Plate XII, 84)

Frequent synonyms / misidentifications: None / Pterocaesio chrysozona non Cuvier, 1830.
FAO names: En - Wideband fusilier; Fr - Fusilier à bande large; Sp - Fusilero de banda ancha.
processes; small conical teeth in jaws and vomer; palatines without teeth. Dorsal fin with X spines and
15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely 13) soft rays; pectoral fins with 21 to
23 rays. Scales in lateral line 74 to 88; upper peduncular scales 12 to 14, lower peduncular scales
usually 16 or 17; scales above lateral line to origin of dorsal fin usually 9 or 10; scales below lateral line to
origin of anal fin usually 16 to 18; usually 4 scale rows on cheek; predorsal scales 23 to 30; dorsal and
anal fins scaly, the dorsal fin with about 1/2 of its spinous height covered with scales; supratemporal band
of scales confluent at dorsal midline. Colour: upper body bluish or reddish, lighter ventrally; a bright yellow
band straddling lateral line for most of its length, from tip of snout to base of caudal fin, 3 to 5 scales
wide anteriorly, tapering to 1 scale in width on caudal peduncle where it is above lateral line; fins
white to pinkish; dorsal fin slightly dusky distally and yellowish near base; axil of pectoral fins with a black
blotch; tips of caudal-fin lobes black.
Size: Maximum size estimated to be about 20 cm.
Habitat, biology, and fisheries: Inhabits coastal areas around coral reefs. A schooling fish, sometimes
in groups with other species of Pterocaesio. Feeds on zooplankton in midwater aggregations. Of minor
importance in fisheries; rarely seen in markets.
Distribution: Eastern Indian
Ocean and western Pacific,
from the Chagos Archipelago
to Papua New Guinea.
2936 Bony Fishes
Pterocaesio marri Schultz, 1953 (Plate XII, 85)

Frequent synonyms / misidentifications: Pterocaesio kohleri Schultz, 1953 / Pterocaesio digramma
(non Bleeker, 1865).
FAO names: En - Marr’s fusilier; Fr - Fusilier de Marr; Sp - Fusilero de Marr.
processes; small conical teeth in jaws; vomer and palatines without teeth. Dorsal fin with X (rarely
XI) spines and 15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely 11 or 13 soft rays;
pectoral fins with 22 to 24 soft rays. Scales in lateral line usually 70 to 75; upper peduncular scales
usually 12 or 13, lower peduncular scales usually 16 or 17; scales above lateral line to origin of dorsal fin
9 to 11; scales below lateral line to origin of anal fin usually 16 to 18; usually 5 scale rows on cheek;
predorsal scales usually 26 to 30; dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest
spinous height covered with scales; supratemporal band of scales confluent at dorsal midline. Colour:
body blue to greenish dorsally, white ventrally; 2 longitudinal stripes laterally, about 1 scale wide each,
the lower running from behind supratemporal membrane, covering lateral line to a vertical at about
middle of soft portion of dorsal fin where it passes above lateral line for the length of caudal
peduncle; upper stripe 1 to 2 scales below dorsal profile for most of length of body, ending dorsally on
caudal peduncle; fins white to pinkish, dorsal fin slightly dusky distally; axil of pectoral fins and tips of
caudal-fin lobes black.
Habitat, biology, and fisheries: Ranges widely around coral reefs in schools. This species appears to
prefer clear waters of oceanic islands or reefs far from large land masses. Feeds on zooplankton in
midwater aggregations. Of minor importance to fisheries. Caught fairly often by drive-in net in western
Palawan (Philippines) and in other areas occasionally by gill net and traps. Sometimes used as baitfish in
tuna fisheries.
Africa, not including the Red
Sea or the Persian Gulf, to the
Marquesas Islands.
Pterocaesio pisang (Bleeker, 1853) (Plate XII, 86)

FAO names: En - Banana fusilier; Fr - Fusilier banane; Sp - Fusilero banana.
processes; small conical teeth in jaws, vomer, and palatines. Dorsal fin with X (rarely XI) spines and
15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely 11 or 13) soft rays; pectoral fins with
18 to 20 rays. Scales in lateral line usually 63 to 71; upper peduncular scales usually 11, lower
peduncular scales usually 15; scales above lateral line to origin of dorsal fin 8 to 10; scales below lateral
line to origin of anal fin usually 14 or 15; usually 4 scale rows on cheek; predorsal scales usually 23 to 26;
dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest spinous height covered with scales.
Colour: body coloration variable, dark red to silvery, paler ventrally; lateral line darker than background
coloration; no stripes or bands on side; snout often yellowish; axil of pectoral fins black; tips of caudal-fin
lobes dark red to black.
Habitat, biology, and fisheries: Ranges widely around coral reefs. A schooling fish, sometimes in groups
with other species of Pterocaesio. Feeds on zooplankton in midwater aggregations. A common food fish
in the Philippines; of minor importance in other areas. Caught mostly by drive-in net, gill net, and traps.
Marketed fresh and sometimes as dried salted. An important tuna baitfish.
Sea or the Persian Gulf, and
eastward to Fiji.
2938 Bony Fishes
Pterocaesio randalli Carpenter, 1987 (Plate XII, 87)

FAO names: En - Randall’s fusilier; Fr - Fusilier de Randall; Sp - Fusilero de Randall.
processes; small conical teeth in jaws; vomer and palatines without teeth. Dorsal fin with X spines
and 15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 soft rays; pectoral fins with 20 to 22
rays. Scales in lateral line 69 to 80; upper peduncular scales usually 11 to 13, lower peduncular
scales usually 15 or 16; scales above lateral line to origin of dorsal fin usually 9 or 10; scales below lateral
line to origin of anal fin usually 16 or 17; 4 scale rows on cheek; predorsal scales 22 to 27; dorsal and
anal fins scaly, the dorsal fin with about 1/2 of its spinous height covered with scales; supratemporal band
of scales confluent at dorsal midline. Colour: body blue-green to reddish dorsally, lighter ventrally; a large
yellow blotch of variable size above pectoral fins, roughly rhomboid in shape, with the oblique side
slanting anteriorly, typically starting posterior to upper rim of orbit, covering the operculum above pectoral
base and 2 or 3 scales above lateral line, stretching posteriorly to about a vertical at anal-fin origin, 6 or 7
scales in height and about 14 scales in width; pectoral, pelvic, and anal fins white to pinkish; dorsal fin
blue-green to pinkish; axil of pectoral fins and tips of caudal-fin lobes black.
Habitat, biology, and fisheries: Ranges widely around coral reefs in schools. Often in groups with other
species of Pterocaesio. Feeds on zooplankton in midwater aggregations. Of minor importance to fisheries.
Occasionally caught by drive-in net, gill net, traps, and handline.
Ocean and West Pacific, from
the Andaman Sea to the Phil-
ippines.
Pterocaesio tessellata Carpenter, 1987 (Plate XII, 88)

FAO names: En - Onestripe fusilier; Fr - Fusilier à une bande; Sp - Fusilero de una bande.
processes; small conical teeth in jaws and vomer; palatines without teeth. Dorsal fin with X (rarely XI)
spines and 15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely 11 or 13) soft rays;
pectoral fins with 20 to 22 rays. Scales in lateral line usually 69 to 75; upper peduncular scales
usually 11 to 13, lower peduncular scales 15 to 17; scales above lateral line to origin of dorsal fin usually
9 or 10; scales below lateral line to origin of anal fin usually 16 to 18; usually 5 scale rows on cheek;
predorsal scales 22 to 29; dorsal and anal fins scaly, the dorsal fin with about 1/2 of its greatest spinous
height covered with scales; supratemporal band of scales confluent at dorsal midline. Colour: upper body
light bluish green; body white ventrally; centres of scales on upper two-thirds of body lighter than distal
portion, giving a checkered appearance (in many individuals the scale rows are uneven, giving a mosaic
appearance); a single yellow longitudinal stripe laterally, about 1 scale wide, covering lateral line
for most of its length, usually running dorsal to lateral line on caudal peduncle; pectoral, pelvic, and
anal fins white to pinkish; axil of pectoral fins black; dorsal fin light bluish green; caudal fin dusky, the
tips of lobes black.
Habitat, biology, and fisheries: Inhabits coastal areas around coral reefs. A schooling fish, often found
in groups with other species of Pterocaesio. Feeds on zooplankton in midwater aggregations. Moderately
important as a food fish. Common in markets in the Philippines. Caught primarily by drive-in net, gill net,
and traps. Marketed mostly fresh.
Distribution: C e n t r a l a n d
eastern Indian Ocean and
western Pacific, from the
Maldives to Vanuatu.
2940 Bony Fishes
Pterocaesio tile (Cuvier), 1830 (Plate XII, 89)

FAO names: En - Darkbanded fusilier; Fr - Fusilier ligne olive; Sp - Fusilero aceitunero.
Diagnostic characters: Body fusiform, elongate, and compressed. Two postmaxillary processes; small
conical teeth in jaws, vomer, and palatines. Dorsal fin with XI or XII (rarely X) spines and 19 to 22
soft rays; anal fin with III spines and 13 soft rays; pectoral fins with 22 to 24 rays. Scales in lateral line 69
to 76 (most frequently 71); upper peduncular scales usually 11 or 12, upper peduncular scales usually 15
or 16; scales above lateral line to origin of dorsal fin 7 or 8; scales below lateral line to origin of anal fin
usually 16 or 17; usually 4 scales on cheek; predorsal scales usually 27 to 30; dorsal and anal fins scaly,
the dorsal fin with about 1/2 of its greatest spinous height covered with scales. Colour: scales above lateral
line bluish green in their centre and black on their margins, giving the appearance of a checkered pattern;
lateral line covered for most of its length by a black stripe about 1 scale wide (on caudal peduncle
this stripe is above lateral line); a brilliant light blue zone below black stripe, usually covering middle
third of body, but sometimes restricted to anterior part of body, and sometimes absent; lower third of body
white to pinkish; pectoral, pelvic, and anal fins white to pinkish; axil and upper base of pectoral fins black;
dorsal fin light bluish green to pinkish; caudal fin with a black streak within each lobe, the upper streak
continuous with lateral stripe.
Habitat, biology, and fisheries: Ranges widely around coral reefs. A schooling fish, sometimes in groups
with other caesionids. Feeds on zooplankton in midwater aggregations. Moderately important in small-
scale fisheries. Caught by drive-in net, gill net, traps, and handline. Marketed fresh and dried-salted.
Important as tuna baitfish.
Sea or the Persian Gulf, to the
Tuamotu Archipelago, as far
north as southern Japan and
south to Mauritius and the
Austral Islands.
Pterocaesio trilineata Carpenter, 1987 (Plate XII, 90)

FAO names: En - Threestripe fusilier; Fr - Fusilier à trois bandes; Sp - Fusilero de tres bandas.
processes; small conical teeth in jaws; vomer and palatines without teeth. Dorsal fin with X (rarely
XI) spines and 15 (rarely 14 or 16) soft rays; anal fin with III spines and 12 (rarely 11) soft rays; pectoral
fins usually with 19 to 21 rays. Scales in lateral line usually 63 to 69; upper peduncular scales usually
11, lower peduncular scales usually 15; scales above lateral line to origin of dorsal fin usually 8 or 9; scales
below lateral line to origin of anal fin 14 to 17 (most frequently 15); usually 4 scale rows on cheek; predorsal
scales 20 to 30; dorsal and anal fins scaly, the dorsal fin with about 1/3 of its spinous height covered
with scales; supratemporal band of scales confluent at dorsal midline. Colour: 3 brown to yellow and 3
bluish white longitudinal stripes on upper side; lowest dark stripe about 2 scales wide, originating on
snout and upper orbit, and covering lateral line for most of its length, running dorsal to lateral line on caudal
peduncle; middle dark stripe about 2 scales wide, originating on snout, running about midway between
dorsal profile and lower dark stripe; uppermost stripe originating on snout and straddling dorsal profile for
about 1 scale to either side; bluish white stripes about 1 to 1 ½ scales wide, covering spaces between dark
stripes and just below lowest dark stripe; body whitish ventrally; pectoral, pelvic, and anal fins white; axil
of pectoral fins black; dorsal fin light bluish; caudal fin dusky with lobe tips black.
Habitat, biology, and fisheries: Ranges widely around coral reefs in schools. Feeds on zooplankton in
midwater aggregations. Not common as a food fish. A fairly important tuna baitfish.
Distribution: W e st e r n P a-
cific, from the Philippines to
Fiji.
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2942 Bony Fishes
LOBOTIDAE
Tripletails
by K.E. Carpenter
D iagnostic characters: Moderately deep bodied and compressed fishes (size to over 1 m). Head length
shorter than body depth; profile of head from eye to just posterior to nape generally concave.
Preopercle finely to coarsely serrate; opercle with 1 or 2 flat spines. Mouth terminal to slightly superior,
extending to anterior or middle portion of eye; in jaws, an outer row of small conical teeth and an inner
band of smaller teeth; roof of mouth without teeth. Single dorsal fin with XI to XIII spines and 13 to
16 soft rays; soft-rayed portion broadly rounded and extending beyond base of caudal fin. Anal fin
with III spines, second anal-fin spine enlarged (in Coius) or not enlarged (Lobotes), and 8 to 11 soft
rays, the soft-rayed portion broadly rounded and extending beyond base of caudal fin. Caudal fin
rounded. Scales ctenoid, 42 to 70 in lateral line. Colour: brownish, olive, or yellowish and sometimes
mottled, or whitish with 4 to 10 dark brownish bars.
Habitat, biology, and fisheries: Coastal and estuarine, some entering fresh water; some associated with
floating objects well out to sea. Feed mostly on small fish and invertebrates. Caught by traps, hook-and-line,
trawls, gill nets, and seines.
Remarks: The species of Coius are sometimes separated from Lobotes and variously placed in either the
Coiidae or Datnioididae. Datnioides is a junior synonym of Coius. Two other species of Coius occur in the
area but are restricted to fresh water. These are C. microlepis (Bleeker, 1853) and C. undecimradiatus
Roberts and Kottelat, 1994.

Serranidae: certain serranid species look similar to tripletails but all serranids can be distinguished by the
usually 3 flat spines on upper part of opercle (1 or 2 flat spines in Lobotes); in addition, those species that
tend to look like tripletails typically have fewer than XI dorsal-fin spines.
Haemulidae: certain species of haemulids look similar to tripletails but these can be distinguished by the
presence of chin pores.
Serranidae Haemulidae
Perciformes: Percoidei: Lobotidae 2943
Lutjanidae: certain species of lutjanids can look similar

to species of Coius but these can be readily distin-
guished by the truncate, emarginate, or forked caudal
fins, the usual absence of an enlarged second dorsal-
fin spine (second dorsal-fin spine enlarged compared
to third spine in Coius), and teeth usually present on
vomer and palatines (absent in Lobotidae).
Lutjanidae
Key to the species of Lobotidae occurring in the area

1a. Third anal-fin spine longer than second anal-fin spine; 42 to 45 scales in lateral line
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lobotes surinamensis
1b. Second anal-fin spine longer than third anal-fin spine; 55 to 70 scales in lateral line . . . . . . . ® 2
2a. Total gill rakers on first gill arch 20 to 23; anal fin usually with 9 soft rays . . . Coius quadrifasciatus
(India to Viet Nam, Indonesia, and Borneo)
2b. Total gill rakers on first gill arch 15 to 17; anal fin usually with 10 soft rays . . . . . . Coius campbelli
(Irian Jaya and Papua New Guinea)

0
0
Coius campbelli (Whitley, 1938)
Coius quadrifasciatus (Sevastianof, 1809)
0 Lobotes surinamensis (Bloch, 1790)
References
Heemstra, P.C. 1984. Lobotidae. In FAO species identification sheets for fishery Purposes. Western Indian Ocean
(Fishing Area 51), edited by W. Fischer and G. Bianchi. Vol. 2. Rome, Italy (unpaginated).
Roberts, T.R. and M. Kottelat. 1994. The Indo-Pacific tigerperches, with a new species from the Mekong basin (Pisces:
Coiidae). Ichthyol. Explor. Freshwaters, 5(3):257-266.
2944 Bony Fishes
Lobotes surinamensis (Bloch, 1790)

FAO names: En - Tripletail; Fr - Croupia roche; Sp - Dormilona.
Diagnostic characters: Body moderately compressed and deep, its depth 2 to 2.5 times in standard
length. Head length shorter than body depth. Opercle with 1 or 2 flat spines. Mouth slightly superior,
reaching posteriorly to about middle of eye; upper jaw slightly protrusible; jaws with an outer row of short,
close-set canines and an inner band of smaller teeth; no teeth on vomer, palatines, or tongue;
preopercle coarsely serrate. Branchiostegal rays 6. First gill arch with 6 or 7 gill rakers on upper limb,
13 to 15 on lower limb. A single dorsal fin with XI to XIII strong spines and 13 to 16 soft rays; anal
fin with III spines and 11 or 12 soft rays; soft dorsal and anal fins large and broadly rounded, reaching
past base of caudal fin (appearing as 3 tails together with rounded caudal fin). Caudal peduncle deep,
its depth greater than its length. Scales ctenoid and fairly large, covering body and head except for
preorbital region and jaws; lateral-line scales 42 to 45. Colour: juveniles brownish and often mottled darker;
adults uniform dark brown or olive to mottled yellowish.
Size: Maximum total length 100 cm; commonly to 50 cm.
Habitat, biology, and fisheries: In coastal areas and estuaries; also found offshore around floating
objects. Juvenile coloration is similar to dead mangrove leaves and they mimic fallen leaves by floating
sideways. Feeds mostly on benthic invertebrates and small fish. Caught by beach seines, gill nets, trawls,
traps, and hook-and line. Valued as both a food and sport fish.
Distribution: Worldwide in
tropical and subtropical wa-
ters. In the Indo-Pacific, from
East Africa through all coun-
tries of Southeast Asia north
to Taiwan Province of China
and southern Japan, northern
Australia to southern
Queensland, New Guinea to
New Britain, and south to Fiji.
Reported as rare visitors in
Ponape, Hawaii, and Tahiti.
Perciformes: Percoidei: Lobotidae 2945
Coius campbelli (Whitley, 1938)

En - New Guinean tigerperch.
Maximum standard length 32 cm. Occurs in brackish coastal lagoons and rivers mouths, and in
fresh-water rivers far inland from the coast. Restricted to coastal waters and drainages of New
Guinea and the Gulf of Papua. Feeds on small fish and invertebrates. The Southeast Asian species
of Coius occurring in brackish water look similar and have a variable number of bars on the sides.
These were previously considered a single species, C. quadrifasciatus, but the New Guinean form
is now considered distinct from the form occurring elsewhere in Asia.
(after Roberts and Kottelat, 1994)
Coius quadrifasciatus (Sevastianof, 1809)

En - Barred tigerfish.
Maximum total length 30 cm. Occurs in brackish coastal lagoons and rivers mouths, and in
fresh-water rivers far inland from the coast. From India around the mainland coast of Southeast
Asia to southern Viet Nam and Sumatra and Borneo. Feeds on small fish and invertebrates.
(after Day, 1875)
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2946 Bony Fishes
GERREIDAE
Mojarras (silverbiddies)
by D.J. Woodland
D iagnostic characters: Body laterally compressed, oblong, oval, or with markedly elevated back (size
to about 35 cm). Mouth terminal, strongly protractile, pointing downward when extended. Bands
of minute, acute teeth in both jaws; no incisors, canines, or molars. Dorsal fin long, single, with IX
(occasionally X) spines, the first very short (except in Parequula, not known from the area), and a
similar number (9 to 11 in Gerres) or larger number (12 to 15 in Pentaprion and 17 in Parequula) of
soft rays; base of dorsal fin sheathed in a row of deciduous scales. Anal fin usually with III (less
commonly II) spines, but V or VI in Pentaprion, the first spine very short in all species except Parequula;
6 to 8 anal-fin rays, but 12 to 14 in Pentaprion and 16 to 18 in Parequula. Pectoral fins long and pointed.
Caudal fin markedly to very deeply forked. Scales large, obvious but deciduous, cycloid or finely ctenoid,
extending over sides of head. Colour: predominantly brilliant silver, faint olive to brown dorsally, with dusky
markings on sides in some species; dusky bars on sides occur only in juveniles in some species; margin
and/or tip of spinous part of dorsal fin often black.
large scales
scaly sheath
mouth
protractile
Habitat, biology, and fisheries: Near the bottom in coastal waters of all warm seas, down to 70 m; a few
temperate species; especially abundant in very shallow bays, estuaries, and coastal lagoons. Tolerant of
hyper- and hyposaline conditions, a few entering fresh water; some species characteristically found in clear
water over bottoms of clean sand, others in turbid waters on muddy bottoms; a couple of species inhabit
open areas of sand on coral reefs where they forage singly or in small schools, but most live in large
schools. Feed on small invertebrates gleaned from the bottom by taking up mouthfuls of substrate and
expelling sand and debris. Temperate species have a single spawning season in the warmer months; some
species spawn biannually; larvae from a mixture of species abundant throughout the whole year in tropical
coastal lagoons in some localities. Caught with set, lift, cast, and seine nets, and as bycatch with bottom
trawls; flesh of good quality, but softens very quickly unless chilled; marketed fresh or as processed frozen
products.
nuchal
Similar families occurring in the area crest incisors
Leiognathidae: bony ridges with median
“nuchal” crest on top of head; scales minute,
barely visible. canines
Sparidae: mouth not strongly protractile;
large incisor, canine, and/or molar-shaped molars molars
teeth in jaws.
Argyrops Diplodus
examples of teeth
Leiognathidae Sparidae
Perciformes: Percoidei: Gerreidae 2947
Identification note
1. The number of lateral-line scales can be a useful diagnostic character. These scales may be divided
into those that occur anterior to the line of flexure at the end of the hypural bone (i.e. up to the point to
which “standard length” is measured in fishes in general) and those that occur posterior to this point -
the latter residing in the scales covering the base of the caudal fin. If these scales are lost, the number
up to the line of flexure may still be determined from the scale pockets, but posterior to this point an
accurate estimate of the number is very difficult. For this reason the numbers of lateral-line scales
recorded in this work are quoted in 2 parts, those anterior to the line of flexure and those posterior to
it; the former figure carries the greater weight as a diagnostic character.
2. The transverse scale-row counts have been made from the base of the fifth dorsal-fin spine,
diagonally backward to immediately in front of the first anal-fin spine; the row of scales sheathing the
base of the dorsal fin is not included in the count. Excluding these sheath scales, the most dorsal row
of scales on the sides is often composed of scales which are shorter in height than the scales below
them; these have been included in the count expressed in fractions, e.g. 4.5 + 1 + 9 means that,
counting from the base of the fifth dorsal-fin spine, there are 5 scale rows above the row of lateral-line
pored scales, with the most dorsal row composed of scales only 1/2 the height of those immediately
ventral to them (with a further 9 scale rows below the lateral line).
3. The premaxillae (upper jaws) have not been included in measurements of standard length or length
of head.
Key to the species of Gerreidae occurring in the area

1a. Anal fin with V or VI spines and 12 to 14 soft rays, the fin base longer than length of base
of soft portion of dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . Pentaprion longimanus
1b. Anal fin with III (occasionally II) spines, the first very short, and 6 to 8 soft rays, the fin
base shorter than length of base of soft portion of dorsal fin . . . . . . . . . . . . . . . . . . . . ® 2
2a. Dorsal fin with X spines and 9 soft rays1/. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3

2b. Dorsal fin with IX spines and 10 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Lateral-line scales 38 to 40, and 3 or 4 more smaller pored scales on base of caudal fin;
lower margin of preopercle not serrated . . . . . . . . . . . . . . . . . . . . . . . . Gerres japonicus
3b. Lateral-line scales 34 to 37, and 2 more smaller pored scales on base of caudal fin; lower
margin of preopercle finely serrated . . . . . . . . . . . . . . . . . . . . . . . . Gerres decacanthus
4a. Scales in interorbital area extending on a broad front

down to level of posterior margin of posterior nostrils,
extensively covering the skin over the median groove
into which the extensions of premaxillary bones (up-
per jaw) retract when mouth closed (Fig. 1a) . . . . . . . . . ® 5
4b. Scales in interorbital area not extending on a broad
front down to nostrils, but extending as a pair of a) premaxillary groove scaly
horns, 1 on each side of midline, leaving the skin over
premaxillary eye
premaxillary groove unscaly (Fig. 1b) . . . . . . . . . . . . . ® 6 groove
upper jaw
5a. Scales absent from a small, isolated, median patch
of skin (a little larger than posterior nostril) in interor-
bital area . . . . . . . . . . . . . . . . . . . . Gerres baconensis
5b. No small, isolated patch of skin lacking scales in
interorbital area . . . . . . . . . . . . . . . . Gerres subfasciatus
b) premaxillary groove naked

and upper jaw protruded
Fig. 1 dorsal view of heads of
2 species of Gerres
1/ Gerres decacanthus sometimes with IX spines.

2948 Bony Fishes
6a. Second dorsal-fin spine laterally compressed and greatly elongate, tapering to a slender,
flexible filament whose tip when depressed extends past level of origin of first anal-fin
spine; tip often broken . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6b. Second dorsal-fin spine not greatly elongate, when depressed its tip not reaching level
of origin of first anal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7a. Usually 5 or 5 ½ (rarely 4 ½) scale rows between lateral line and base of fifth dorsal-fin spine;
typically 44 or 45 (rarely 43 or 46) lateral-line scales to base of caudal fin . . . . . .Gerres filamentosus
7b. Usually 4 or 4 ½ (rarely 5) scale rows between lateral line and base of fifth dorsal-fin spine;
typically 42 or 43 (rarely 41 or 44) lateral-line scales to base of caudal fin . . . . .Gerres macracanthus
7c. Usually 4 ½ scale rows between lateral line and base of caudal fin; typically 39 or 40
lateral-line scales to base of caudal fin . . . . . . . . . . . . . . . . . . . . . . . Gerres infasciatus
8a. Body very slender, the depth 3.1 to 3.4 times in standard length . . . . . . . . . . .Gerres oblongus
8b. Body comparatively deep, the depth 1.9 to 3 times in standard length . . . . . . . . . . . . . . . ® 9
9a. Body very deep, the depth 1.9 to 2.3 times in standard length; second anal-fin spine
robust and long, its length subequal with length of base of anal fin . . . . . . . . . . . . . . . . ® 10
9b. Body comparatively slender, the depth 2.3 to 3 times in standard length; second anal-fin
spine relatively slender and short, its length clearly shorter than length of base of anal
fin, often less than 1/2 length of base of anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
10a. Tip of depressed pectoral fins reaching well past vertical line through origin of first
anal-fin spine; body depth 1.9 to 2.3 times in standard length; second dorsal-fin spine
longer than length of head minus snout . . . . . . . . . . . . . . . . . . . . . . . Gerres abbreviatus
10b. Tip of depressed pectoral fins just reaching to vertical line through origin of first anal-fin
spine; body depth 2.1 to 2.3 times in standard length; second dorsal-fin spine shorter
than length of head minus snout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gerres poeti
11a. Tip of depressed pectoral fins clearly not reaching past level of anus . . . . . . . . . . . . . . . ® 12
11b. Tip of depressed pectoral fins reaching past level of anus (almost reaching to or reaching
or past level of first anal-fin spine) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
12a. Lateral-line scales 35 to 39 to base of caudal fin, and 3 or 4 more pored scales in scaly
sheath covering base of caudal fin; 3 ½ scale rows between lateral line and base of fifth
dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gerres oyena
12b. Lateral-line scales 40 to 43 to base of caudal fin, and 2 or 3 more pored scales in scaly
sheath covering base of caudal fin; 4 ½ to 5 scale rows between lateral line and base of
fifth dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gerres macrosoma
13a. Caudal fin long, much longer than length of head; lateral -line scales 42 to 44 to base of
caudal fin, and 3 to 5 more in scaly sheath covering base of caudal fin; 4 ½ to 5 ½ scale
rows between lateral line and base of fifth dorsal-fin spine . . . . . . . . . . . . . . Gerres argyreus
13b. Caudal fin short, shorter than length of head; lateral-line scales 33 to 41 to base of caudal
fin, and 2 or 3 more in scaly sheath covering base of caudal fin; 3 to 4 ½ scale rows
between lateral line and base of fifth dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . ® 14
14a. Lateral-line scales 33 to 36 to base of caudal fin, and 2 or 3 more on scaly sheath
covering base of caudal fin; 3 or 3 ½ scale rows between lateral line and base of fifth
dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gerres decacanthus
14b. Lateral-line scales 37 to 41 to base of caudal fin, and 2 or 3 more on scaly sheath
covering base of caudal fin; 4 or 4 ½ scale rows between lateral line and base of fifth
dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gerres kapas

0
0
Gerres abbreviatus Bleeker 1850
0
Gerres argyreus (Schneider, 1801)
Gerres baconensis (Evermann and Seale, 1907)
Gerres chrysops Iwatsuki, Kimura, and Yoshino, 19992/
0
0
Gerres decacanthus Bleeker, 1865
Gerres filamentosus Cuvier, 1829
Gerres infasciatus Iwatsuki and Kimura, 19983/
0
0
Gerres japonicus Bleeker, 1854
0
Gerres kapas Bleeker, 1854
Gerres macrosoma Bleeker, 1854
0
0
Gerres macracanthus Bleeker, 1854
0
Gerres oblongus Cuvier, 1830
Gerres oyena (Forsskål, 1775)
0 Gerres poeti Cuvier in Cuvier and Valenciennes, 18304/
0 Gerres subfasciatus Cuvier, 1830
0 Pentaprion longimanus (Cantor, 1850)
Reference
Woodland, D.J. 1984. Gerreidae. 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).
2/ Recently described from 3 specimens taken in the northern Gulf of Thailand. See species account of Gerres
filamentosus to follow.
3/ Recently described from the northeastern Gulf of Thailand. Difficult to distinguish from G. decacanthus and not included
in the identification key; reported as being slightly less deep-bodied and having slightly longer dorsal- and anal-fin
spines.
4/ Cuvier’s G. poieti was based on a pre-Linnaean published picture of a fish; there were no type specimens, and the
picture is too crude to identify the fish to species. On the other hand, Cuvier’s Gerres poeti has a type specimen, and
is easily identifiable. Cuvier himself expressed doubt about the 2 fish being the same. He used the French conditional
tense to remark that IF the indigenous names of these 2 fish were the same, then “they might well be” the same fish.
My research shows that the indigenous names he refers to [“poia-poieti” (S. India) and “poeti” (Malay)] are not variations
of the same name. Were it not for this ‘indecision’ by Cuvier, one could argue that the first spelling used (poieti) has
priority, and that the subsequent “poeti” is a mis-spelling of “poieti”. Following this line of argument, Gerres poeti
currently has no valid scientific name, and it would need to be redescribed with a new name. While it could be done,
most taxonomists would agree that this FAO publication is not the right place to be describing/naming a new species.
Alternatively, the entry for this species could read “Gerres sp. (undescribed)”, with “Gerres poeti” given as a “Frequent
Synonym...”. HOWEVER, I do not think it is necessary to give Gerres poeti a new name. Under the ICZN Rules, a
difference in spelling of just a single letter between two scientific names is sufficient to validate both of them. Further,
under the same Rules, one cannot conclude that a published name is a misspelling by reference to a source other
than the original publication; so we cannot correct the spelling “poieti” in Cuvier 1829 to “poeti” (even though it would
be logical to do so because “poeti” is the spelling used for the fish in the pre-Linnaean publication on which Gerres
poieti is based). [D.J. Woodland]
2950 Bony Fishes
Gerres abbreviatus Bleeker, 1850

FAO names: En - Deepbody silverbiddy; Fr - Blanche elévée; Sp - Mojarra chata.
Diagnostic characters: Body compressed, deep, with an obtusely angled, elevated back; body
depth 1.9 to 2.3 times in standard length. Anterodorsal profile nearly a straight line from tip of snout to
base of first dorsal-fin spine, ascending at an angle of about 45° to the horizontal. Dorsal- and anal-fin
spines strong, particularly second anal-fin spine; second dorsal-fin spine longer than head minus snout;
second anal-fin spine either subequal to (0.8 times) or much longer than length of base of anal fin; pectoral
fins long, tip of depressed fins reaching to or past level of first anal-fin spine; caudal fin strongly forked
but short, longest rays subequal to head length, lobes broadly rounded without pointed tips. Typically
fewer than 38 lateral-line scales to base of caudal fin, and 2 or 3 more pored scales in scaly sheath on
base of fin; 3 ½ to 4 scale rows between base of fifth dorsal-fin spine and lateral line. Colour: silvery,
tinged with brown dorsally; indistinct, fine, dark stripes following scale rows in older fish; many specimens,
particularly juveniles, with 7 to 11 slender, dark bars along sides; dorsal fin edged with black; trailing edge
of caudal fin dusky; anal and pelvic fins yellowish.
Habitat, biology, and fisheries: Lives in small schools on sandy bottoms, juveniles in the littoral zone,
larger fish down to 40 m. Small juveniles feed on zooplankton, larger fish on small polychaetes, bivalves,
crustaceans, and fishes. One of the more common gerreids in markets in Southeast Asia. Caught in set
traps, seines, and by trawls.
Usually marketed fresh, but
the flesh deteriorates rapidly;
large catches processed into
fish balls and marketed fro-
zen.
Distribution: From the west
coast of India to the Indo-Ma-
layan Archipelago and Palau,
north to South China Sea, and
south to northern Australia.
Gerres argyreus (Forster in Bloch and Schneider, 1801)

Frequent synonyms / misidentifications: Gerres acinaces Bleeker, 1854 / Gerres oyena (Forsskål, 1775).
FAO names: En - Longtail silverbiddy; Fr - Blanche gouvernail; Sp - Mojarra timonera.
Diagnostic characters: Body compressed, slender, its depth 2.5 to 2.9 times in standard length;
anterodorsal profile convex, except for slightly concave interorbital area; snout pointed, long, meas-
ured from tip to leading edge of eye almost 1.2 times length of eye; prominent, scaleless patch of
skin encircling eye. Dorsal fin deeply notched, last dorsal-fin spine about 0.6 to 0.7 times in first soft ray;
second and third anal-fin spines slender, length moderate, 0.5 to 0.6 times in second dorsal-fin spine;
pectoral fins moderately long, tip of depressed fins just reaching to level of first anal-fin spine; caudal fin
extremely deeply forked and very long, length of longest ray greater than distance from base of
pelvic-fin spine to first anal-fin spine. Lateral-line scales 41 to 44 to base of caudal fin, and 1 to 3 more
pored scales in scaly sheath on base of fin; 4 ½ to 5 scale rows between base of fifth dorsal-fin spine and
lateral line. Colour: silvery, 5 or more faint slender bars on midsides in some specimens, narrow black
margin to anterior point of spinous dorsal fin, trailing edge of caudal fin with a narrow, darker margin.
Habitat, biology, and fisheries: Inhabits inshore areas favouring clear, shallow water over bottoms of
clean sand; in the area, the most common gerreid on coral reef flats and lagoons. Juveniles school; adults
live singly or in small groups. Larger fish found picking over bottom for small invertebrates in areas swept
by swift currents. Caught with bottom trawls down to about 40 m, also by beach seine and handline.
Marketed fresh; large catches may be used as duck food or fertiliser in some areas.
Distribution: From the east
coast of Africa and islands of
the Indian Ocean to those of
the Central Pacific; at least as
far east as Samoa and Tonga,
throughout Micronesia, north
to Ryukyu Islands, south to
northern Australia and New
Caledonia.
2952 Bony Fishes
Gerres filamentosus Cuvier, 1829

Frequent synonyms / misidentifications: Gerres punctatus Cuvier, 1830; ? G. macracanthus Bleeker,
1854 / None.
FAO names: En - Whipfin silverbiddy; Fr - Blanche fil; Sp - Mojarra de hebra.
Diagnostic characters: Body compressed, elevated back, deep, its depth 2 to 2.5 times in standard
length in larger specimens, up to 3 times in smaller ones. Anterodorsal profile ascending steeply in a
gentle curve at an angle of about 40° to horizontal axis. Second dorsal-fin spine laterally compressed,
produced into a filament whose tip extends to at least level of first anal-fin spine, except in very
smallest of specimens, only equal to head length in specimens less than 6 cm standard length (filament
often lost, but compressed condition of spine indicates original condition); third dorsal-fin spine also
laterally compressed, as long as distance from tip of snout to preopercular margin; second anal-fin spine
much shorter than length of base of anal fin; pectoral fins long, tip of depressed fins reaching to level of
first anal-fin spine; caudal fin deeply forked, its longest rays 3 times length of median rays and slightly
longer than length of head. Scales on lateral line 43 to 46 to base of caudal fin, and 2 to 5 more pored
scales in scaly sheath on base of fin. Colour: silvery, with a touch of lightest brown dorsally; 7 to 10
columns of pale brown ovoid spots on upper half of sides, coalesced as bars in smaller specimens;
iris silvery with a dark ovoid patch in anterodorsal quarter; dorsal fin hyaline except for end of filamentous
spine, which is black; anal, caudal, pectoral, and pelvic fins dusky hyaline, tips of anal fin, lower lobe of
caudal and pelvic fins white.
Habitat, biology, and fisheries: Lives in shallow coastal waters to depths of at least 50 m, on sandy
bottoms, including around coralline areas, also entering lower fresh-water reaches of rivers. Schooling;
feeds on small organisms such as crustaceans, polychaetes, and foraminiferans living on sand or
muddy-sand bottoms. The commonest gerreid species in markets in many tropical areas. Caught mainly
with beach seines and bottom trawls. Flesh excellent, but spoils rapidly.
Distribution: Widespread in all warm seas of the Indo-Pacific from the east coast of Africa through the
Indo-Malayan Archipelago to the West Pacific islands; south to northern Australia, north to Ryukyu Islands.
Remarks: Gerres infasciatus Iwatsuki and Kimura, 1998
was recently described from 3 specimens collected in the
northern Gulf of Thailand. It is very similar in body shape
to G. filamentosus adults of similar size (12.5 to 14 cm
standard length), but lacks the dark markings on sides,
has the tips of first and second dorsal-fin rays yellow, and
shows fewer lateral-line scales (39 or 40, plus 3 or 4
pored scales on caudal-fin base). Also similar therefore
to G. macracanthus.
Gerres kapas Bleeker, 1851

Frequent synonyms / misidentifications: None / Gerres oyena (Forsskål, 1775).
FAO names: En - Cotton silverbiddy.
Diagnostic characters: Body compressed, oblong, moderately deep, its depth 2.3 to 2.4 times in
standard length; anterodorsal profile a smooth, convex curve from tip of snout to base of first dorsal-fin
spine. Dorsal-fin spines slender, fin notched, last spine 3/4 length of first soft ray, second dorsal-fin spine
neither elongate nor exceptionally short, contained about 0.6 to 0.7 times in length of head (excluding
jaws); second and third anal-fin spines stouter than dorsal-fin spines, the second more so than the third,
both spines long, the third more so than the second, the third longer than 1/2 length third dorsal-fin spine;
pectoral fins moderately long, tip of depressed fins almost reaching to or just past level of first
anal-fin spine; caudal fin deeply forked, but lobes relatively short, length of longest ray almost
identical to length of head. Scales on lateral line 37 to 41 to base of caudal fin, and 2 or 3 more pored
scales in scaly sheath on base of caudal fin; 4 to 4.5 scales between lateral line and base of fifth dorsal-fin
spine. Colour: silvery, bluish, or greenish shading dorsally, juveniles with variable dark blotches, spinous
dorsal fin with a narrow black margin, anal and pelvic fins yellow, others hyaline.
Habitat, biology, and fisheries: Lives around river mouths and along beaches; seems to prefer murkier
waters. Schools; feeds on small invertebrates gleaned from bottom sediments. Caught with trawl, seine,
and cast nets. Common in markets, but not highly esteemed as flesh deteriorates rapidly.
Distribution: Indo-Malayan
region: Malaysia, Singapore,
Thailand, Indonesia, New
Gu in ea , P hilippines, and
northwestern shelf of Western
Australia.
2954 Bony Fishes
Gerres macrosoma Bleeker, 1854

FAO names: En - Largebodied silverbiddy.
Diagnostic characters: Body compressed, oblong, deep, its depth 2.5 to 3 times in standard length;
anterodorsal profile sloping at an angle of about 30° to horizontal, slightly convex with a small concavity
above anterior third of eye. Dorsal-fin spines slender, fin notched, second dorsal-fin spine neither elongate
nor exceptionally short but short relative to length of base of spinous part of fin, its length very slightly less
than 1/2 body depth but much shorter than twice the length of second anal-fin spine; second and third
anal-fin spines slender and long, their lengths about 0.7 times in second dorsal-fin spine; tip of pectoral
fins not reaching anywhere near to level of first anal-fin spine, reaching anus; caudal fin very deeply
forked, with long pointed lobes, the longest rays clearly greater than length of head and subequal
to body depth. Scales on lateral line 40 to 43 to base of caudal fin, and 2 or 3 more pored scales in
scaly sheath on base of caudal fin; 4.5 to 5 scales between lateral line and base of fifth dorsal-fin
spine. Colour: silvery, pale brownish olive above, spinous part of dorsal fin with a narrow blackish margin,
other fins hyaline
Habitat, biology, and fisheries: Coastal, marine species, not recorded as entering estuaries. Schools;
no doubt feeds on small invertebrates gleaned from substrate with its protrusible mouth. Common in
Indonesia and Philippines. Marketed fresh.
Distribution: Central Indo-
Malayan region (an unverified
record for Samoa); Malaysia,
Indonesia, Philippines, New
Guinea, and Queensland.
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Gerres oblongus Cuvier, 1830

Frequent synonyms / misidentifications: Gerres gigas Günther, 1862 / None.
FAO names: En - Slender silverbiddy; Fr - Blanche élégante; Sp - Mojarra elegante.
Diagnostic characters: Body compressed, elongate, slender, its depth 3.1 to 3.4 times in standard
length; anterodorsal profile very slightly convex, ascending at an angle of less than 30° to horizontal axis;
eye comparatively large, its diameter 3 times in length of head (excluding upper jaw); snout long,
subequal to eye diameter. Fin spines slender; in larger fish, leading dorsal- and anal-fin spines relatively
longer than other fin spines and rays, in fish longer than about 8 cm standard length profile of spinous part
of dorsal fin sweeping rapidly upwards from about midway along its length, second dorsal-fin spine, for
example, at least twice length of sixth dorsal-fin spine; second anal-fin spine short, about 2/3 length of
base of anal fin, even shorter in smaller fish; pectoral fins short, tip of depressed fins reaching almost to
or just past level of anus, nowhere near level of base of first anal-fin spine; caudal fin very deeply forked,
but lobes not exceptionally long, longest caudal-fin ray about 1/3 standard length. Scales on lateral
line 44 to 46 to base of caudal fin, and 2 to 5 more pored scales in scaly sheath on base of caudal fin;
5.5 to 6 scales between lateral line and base of fifth dorsal-fin spine (transverse series of scales
inclined at a low angle to horizontal). Colour: silvery, blue dorsally in life, but lilac brown in some market
specimens; up to 5 columns of horizontally elongated spots on sides of trunk appearing as faint bars in
some cases; dusky tip to dorsal fin, other fins pale.
Habitat, biology, and fisheries: Judging by the localities at which this species has been caught, its habitat
is restricted to coral reefs. Schooling, at least while small. In Palau, adults congregate to spawn over
shallow sandy patches near the outer reef edge around certain full moons. Feeds on small organisms living
in sand. Caught with set nets and beach seines. Flesh reported to be excellent.
uted throughout the tropical
Indo-West Pacific region:
Mozambique, Kenya, Sey-
chelles, Laccadive Islands,
Maldives, Sri Lanka, South In-
dia, Andaman Islands, Indo-
nesia, Philippines, Palau,
Marianas, Rotorua, Fiji,
Tonga, Samoa, north to the
Ryukyu Islands and New
Guinea, south to coasts of
Queensland and New Caledo-
nia.
2956 Bony Fishes
Gerres oyena (Forsskål, 1775)

Frequent synonyms / misidentifications: None / Gerres argyreus (Schneider, 1801).
FAO names: En - Common silverbiddy; Fr - Blanche commune; Sp - Mojarra común.
Diagnostic characters: Body compressed, oblong, slender, its depth 3 to 3.3 times in standard
length; anterodorsal profile gently convex. Dorsal fin distinctly notched, last spine 2/3 length of first soft
ray; second dorsal-fin spine neither elongate nor exceptionally short, contained about 0.6 to 0.7 times in
length of head (excluding jaws); second and third anal-fin spines slender and short, e.g. second
slightly less than 1/2 length of second or third dorsal-fin spines; caudal fin very deeply forked, but
lobes relatively short, length of longest ray subequal with head length; pectoral fins short, tip of depressed
fins only reaching level between bases of first and second dorsal-fin ray. Scales on lateral line 35 to 39
to base of caudal fin, and 2 to 4 more pored scales in scaly sheath on base of caudal fin; 3.25 to 3.5
scales between lateral line and base of fifth dorsal-fin spine. Colour: silvery, with a trace of pale olive
above lateral line; 7 or 8 faint dusky bars on sides of body in young, absent in older fish; prominent black
tip to spinous part of dorsal fin, usually confined to distal part of membrane between second and fifth spines
but sometimes continuing as a black margin along whole of dorsal fin.
Habitat, biology, and fisheries: Inhabits inshore areas on silty-sand bottoms down to at least 30 m; found
along sandy beaches but commonly in estuaries and brackish lagoons. Schools; feeds by sifting small
invertebrates from mouthfuls of substrate. Fished mainly with seines and bottom trawls. Marketed fresh;
also used as fishmeal.
the tropical Indo-West Pacific
Region from the east coast of
Africa and the Red Sea west-
ward to Samoa in the south
Pacific and the Caroline Is-
lands in the north Pacific; ex-
tends northward to southern
Japan and southward to tropi-
cal Australia and to New Cale-
donia.
Gerres poeti Cuvier in Cuvier and Valenciennes, 1830

Frequent synonyms / misidentifications: None / non Gerres poieti Cuvier, 1829.
FAO names: En - Strongspine silverbiddy; Fr - Blanche armie; Sp - Mojarra espinuda.
Diagnostic characters: Body compressed, deep, with an elevated back; body depth 2.1 to 2.3 times
in standard length. Anterodorsal profile ascending at an angle of about 45° to the horizontal in 2 steps:
one corresponding to curve of head, the other to curve of nape up to base of first dorsal-fin spine. Dorsal-
and anal-fin spines strong, second anal-fin spine exceptionally robust; dorsal-fin rays long, the first
about 1.25 times eye diameter; second anal-fin spine as long as or slightly shorter than anal-fin base;
pectoral fins moderately long, tip of depressed fins just reaching to level of first anal-fin spine; caudal
fin strongly forked but short, longest rays slightly longer than head length, lobes broad with pointed
tips. Lateral-line scales 38 to 40 to base of caudal fin, and 2 or 3 more pored scales in scaly sheath on
base of fin; 4 scale rows between base of fifth dorsal-fin spine and lateral line. Colour: upper sides of head
and trunk tinged with golden brown, grading to silver on lower sides; about 10 diffuse dusky bars spread
across sides of trunk in some specimens; fins mostly hyaline: a fine black line outlining margin of membrane
of dorsal fin; tips of first few rays of anal fin and trailing edge of caudal fin dusky.
Habitat, biology, and fisheries: Lives in very shallow coastal waters; common about estuaries and coastal
lagoons over bottoms of muddy sand. Schools; feeds from the bottom on small invertebrates. Caught with
stake nets, lift nets, and beach seines. Marketed fresh.
Distribution: Occurs discon-
tinuously from southwestern
coast of India to central Indo-
Malayan region: Indonesia,
Philippines, and New Guinea.
2958 Bony Fishes
Pentaprion longimanus (Cantor, 1850)

FAO names: En - Longfin silverbiddy; Fr - Blanche à pagaies; Sp - Mojarra alona.
Diagnostic characters: Body compressed, ovoid, moderately deep, its depth 2.5 to 2.8 times in
standard length. Dorsal fin with IX or X spines and 12 to 15 soft rays; anal fin with V or VI spines and
12 to 14 soft rays, its base longer than base of soft portion of dorsal fin; caudal fin deeply forked with
pointed lobes, longer than head; pectoral fins long, tip of depressed fins reaching a level half-way along
base of anal fin. Scales on lateral line 44 to 48 to base of caudal fin; all scales shed easily. Colour: silvery,
with a mirror-like stripe from snout to caudal peduncle; fins dusky yellow; market specimens pale pink or
grey when scales lost.
Habitat, biology, and fisheries: Inhabits inshore areas, on muddy-sand bottoms, to depths of 70 m. Feeds
on small benthic invertebrates. Forms very large schools. Caught mainly with bottom trawls, in very large
quantities in some areas. Marketed fresh, but shabby look makes them unattractive as food fish; used in
large quantities for fishmeal and as a duck food in parts of Southeast Asia.
Distribution: West coast of
India, Sri Lanka, Indo-Ma-
layan area including Indone-
sia, Philippines, northern
Australia, north to Taiwan
Province of China and Ryukyu
Islands.
Gerres baconensis (Evermann and Seale, 1907)

En - Scalynosed silverbiddy.
Maximum total length about 25 cm. Coastal, in bays and along open beaches. Schooling; no doubt
feeds on small invertebrates living in surface layer of the substrate, as other gerreids do. Caught
with beach seines; marketed fresh. In the WCP area known only from the Philippines (Sulu
Archipelago north to Luzon), but also recorded from Ogaswara Islands.
Gerres decacanthus (Bleeker, 1865)

En - Tenspined silverbiddy.
Maximum total length about 15 cm. Recorded for continental localities only, not for offshore coral
islands. Schooling; no doubt similar feeding habits to other gerreids. Known only from South China
Sea area.
Gerres japonicus Bleeker, 1854

En - Japanese silverbiddy.
Maximum total length about 25 cm. Coastal, along the shore and in bays. Schooling; common in
the northern part of its range, which is outside the WCP area. Feeds from the bottom by taking up
mouthfuls of substrate and sieving out the small invertebrates. Caught with set and seine nets.
Marketed fresh; flesh delicious, but full of bones. From Mie and Nagasaki prefectures in South
Japan, and Korea southward to Taiwan Province of China, Hainan, and, reputedly, Luzon, Philip-
pines.
2960 Bony Fishes
Gerres macracanthus Bleeker, 1854

En - Longspined silverbiddy.
Maximum total length 20 cm. Shallow coastal waters to depths of at least 30 m on sandy bottoms,
entering estuaries. Schooling; feeds by sieving small vertebrates from sand. Caught with gill nets
and beach seines. Reported from East Africa to the central Indo-Malayan area, as far east as the
Philippines and north to Taiwan Province of China.
Gerres subfasciatus Cuvier, 1830

En - Southern silverbiddy.
Maximum total length about 26 cm. Coastal, down to 40 m, but very common in small and large
estuaries, preferring murky waters over muddy bottoms. In schools of up to a few hundred
individuals; feeds on any small invertebrates living in bottom sediments. Taken as bycatch by
estuarine prawn trawlers, or by cast net for bait. Marketed fresh in small quantities. Within Australia
confined to tropical and warm-temperate waters; not present along southern coastline; probably
occurs in southern New Guinea.
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Perciformes: Percoidei: Haemulidae 2961
HAEMULIDAE
(= Pomadasyidae)
Grunts (also sweetlips, rubberlips, hotlips, and velvetchins)
by R.J. McKay
D iagnostic characters: Oblong, compressed, perch-like fishes (size to 120 cm). Head profile strongly
convex; scales present on entire head except tip of snout, lips, and chin. Mouth small or moderate,
lips thick in Plectorhinchus, tip of upper jaw hidden when mouth closed; chin with 2 pores anteriorly
and a median pit (Pomadasys), or 6 pores and no pit (Plectorhinchus). Teeth conical, in a narrow band
in each jaw, the outer series enlarged, but no canines; palate toothless. Preopercle with a slightly concave
and serrated posterior margin; opercle with 1 distinct spine. Dorsal fin single, with IX to XV strong spines
and 12 to 26 soft rays; anal fin with III spines (the second often very strong) and 7 to 9 soft rays; caudal
fin truncate or emarginate (rounded in juveniles); pelvic fins below base of pectoral fins, with I spine and
5 soft rays; pectoral fins long. Scales ctenoid (rough to touch), small or moderate. Colour: highly variable,
ranging from uniformly coloured to banded, blotched, and spotted; juveniles may differ markedly from adults
in shape, development of lips, and coloration.
dorsal-fin spines
often strong
scales
between eye
and mouth
margin of preopercle 2nd anal-fin spine

concave and serrate often strong
mouth small in
Plectorhinchus
Habitat, biology, and fisheries: Generally frequenting coastal reefs (Plectorhinchus), inshore bays, and
estuaries (Pomadasys). Feed mostly at dawn or dusk, moving out from the reef to feed on open sandy
bottoms, but may be taken throughout the day by line, spear, or net. All are good table fishes with white
flesh, the larger grunts are considered fine sport fishes. For 1995, the FAO Yearbook of Fishery Statistics
reports a total catch of around 12 000 t of Haemulidae from the area (Indonesia, Malaysia, Singapore).
Remarks: The genus Hapalogenys has been removed from the family Haemulidae. It is very close to the
2 species of the family Dinopercidae, but lacks intrinsic muscles on the posterior part of the swimbladder.
Two species of Hapalogenys occur in the area and are placed here for convenience until better accom-
modated.

Lutjanidae: no pores on chin, canine teeth usu-
ally present, hind part of the upper jaw exposed
when mouth is closed, teeth usually present on
roof of mouth, second anal-fin spine weak.
Lethrinidae: no pores on chin, no scales on pre-
opercle in Lethrinus, and its hind edge not ser-
rated.
no scales no scales
Lutjanidae Lethrinidae
2962 Bony Fishes
Nemipteridae: no pores on chin, hind margin of

suborbital exposed.
Sparidae: no pores on chin, edge of preopercle
smooth, suborbital space scaleless; molar teeth
sometimes present.
hind margin of
suborbital exposed
Nemipteridae Sparidae
Key to the species of Haemulidae occurring in the area 6 pores

2 pores
1a. Chin with 2 pores followed by a pit contain- pit
ing a pore on each side (Fig. 1a) . . . . (Pomadasys) ® 4
1b. Chin with 6 pores and no median pit contain-
ing a pore on each side (Fig. 1b) . . . . . . . . . . . ® 2
2a. Chin with a covering of short barbels or fur-

like papillae; an antrorse spine before first
dorsal-fin spine; caudal fin rounded, becom-
ing almost truncate in adults . . . . (Hapalogenys) ® 11
2b. Chin naked, frequently swollen with age; no a) Pomadasys b) other genera
antrorse spine; caudal rounded or lanceo-
late in juveniles, becoming truncate or emar- Fig. 1 ventral view of head
ginate in adults . . . . . . . . . . . . . . . . . . . . ® 3
3a. Dorsal fin with IX or X spines and 21 to 26 soft rays; 16 to 20 scales between lateral line
and dorsal-fin origin; juveniles with longitudinal black bands, lower part of head and body
yellowish; swimbladder with 3 short anterior horn-like extensions directed forwards . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Diagramma) ® 12
3b. Dorsal fin with XI or more spines and 14 to 22 soft rays; 10 to 17 scales between lateral
line and dorsal-fin origin; swimbladder without horn-like extensions anteriorly . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Plectorhinchus) ® 13
4a. A vertical black band about 8 scales wide over nape, terminating about 3 scales below
lateral line; about 6 black blotches posterior to the vertical band; spinous part of dorsal
fin with a large black blotch . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomadasys maculatus
5a. Gill cover produced posteriorly as a rounded lobe reaching to middle of pectoral fins;
preopercle produced into a posterior lobe; circumpeduncular scales 16 . . . . . . Pomadasys auritus
5b. Gill cover and preopercle normal; circumpeduncular scales more than 16 . . . . . . . . . . . . . ® 6
6a. Body with distinct black or brown horizontal

stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6b. Body without distinct stripes in adults, juveniles
may have 3 indistinct stripes or numerous hori-
zontal lines of contiguous spots . . . . . . . . . . . . . ® 8
7a. Body with 4 distinct black stripes, the lower one

behind eye to middle of tail base (Fig. 2) . . . .
. . . . . . . . . . . . . . . . . . Pomadasys andamanensis
(known from Phuket Island, Andaman Sea; not yet recorded from
the area)
7b. Body with 6 brown stripes that bifurcate anteri-
orly; the bands becoming more numerous with Fig. 2 Pomadasys andamanensis
growth . . . . . . . . . . . . . . . . . Pomadasys furcatum
8a. Body with dark spots or blotches arranged as horizontal rows or vertical interrupted bars;
dorsal fins with distinct spots or dark blotches; dorsal-fin rays usually 14 . . . . . . . . . . . . . ® 9
8b. Body without dark spots or blotches, juveniles of P. trifasciatus have 3 darker stripes of
which the middle one is broadest; opercle with distinct dark spot; dorsal fins unspotted
or with a few vague darker blotches on base only; dorsal-fin rays usually 13 . . . . . . . . . . . ® 10
9a. Body with pairs of spots or dull blotches arranged into vertically interrupted bars; old
specimens may be uniformly coloured with dark blotches or spots on dorsal fins;
circumpeduncular scales 20 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomadasys kaakan
9b. Body with conspicuous rounded spots, scattered on body or forming lines or rows of
spots on back; circumpeduncular scales 21 or 22 . . . . . . . . . . . . . . . . Pomadasys argenteus
10a. Second anal-fin spine usually longer than length of snout plus diameter of eye; eye
diameter usually less than depth of caudal peduncle; first gill arch with middle gill rakers
about 2/3 length of gill lamellae; circumpeduncular scales usually 18 or 19 (rarely 20);
lining of abdominal cavity pale . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomadasys argyreus
10b. Second anal-fin spine about equal to or shorter than length of snout plus diameter of eye;
eye diameter about equal to depth of caudal peduncle; first gill arch with middle gill rakers
about equal to length of gill lamellae; circumpeduncular scales usually 21 to 23 (rarely
19 or 20); lining of abdominal cavity black . . . . . . . . . . . . . . . . . . . Pomadasys trifasciatus
11a. Body with 5 broad black to dark brown vertical bands; vertical fins with black margin . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hapalogenys analis
11b. Body with 4 horizontal dark brown bands on upper part; vertical fins without black margin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hapalogenys kishinouyei
12a. Adults have body and vertical fins blue-grey with yellow or orange spots becoming larger
and coalescing on head to form irregular stripes; inside mouth bright orange to red . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diagramma pictum
12b. Adults have body bronze grey with mauve tints and numerous small dark brown spots
on caudal and soft dorsal fins; some scattered darker blotches may be found on body
and head; anal fins dark grey; no yellow spots at any stage; inside mouth flesh-pi nk . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Diagramma labiosum
13a. Body uniform grey, brownish or olive, with or without a paler caudal fin, red opercular
membrane or dark tips to fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13b. Body not uniformly coloured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
14a. Anterior half of soft dorsal fin black; caudal-fin lobes, particularly lower, tipped with black;
pectoral, pelvic, and anal fins dusky to black; dorsal-fin spines usually XIII (rarely XII);
dorsal-fin rays 16 to 20 . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus albovittatus (adult)
14b. Soft dorsal fin uniformly coloured . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15
15a. Dorsal fin with XIV (very rarely XIII) spines and 15 to 17 (mostly 16) soft rays; pored
lateral-line scales 46 to 55; base of soft dorsal fin shorter than head; juveniles with caudal
fin pink . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus gibbosus
15b. Dorsal fin with XII (rarely XIII) spines and 18 to 21 (mostly 19 or 20) soft rays; pored
lateral-line scales 54 to 58; base of soft dorsal fin equal to or (usually) longer than head;
juveniles with faint blue lines below eye and along body . . . . . . . . . . . . Plectorhinchus schotaf
16a. Body with distinct spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17

16b. Body with large blotches, longitudinal stripes and/or small spots . . . . . . . . . . . . . . . . . ® 20
2964 Bony Fishes
17a. Soft dorsal-fin rays 15 to 17 (mostly 16); colour greyish brown, with large round black
spots on posterior two-thirds of body . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus pictus
17b. Soft dorsal-fin rays 18 to 22 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
18a. Gill rakers on lower limb of first gill arch 16 to 19; body with orange, gold, or rust-coloured
spots fading with age; head with narrow, undulating yellow to rust coloured lines . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus flavomaculatus (adult)
18b. Gill rakers on lower limb of first gill arch 22 or more . . . . . . . . . . . . . . . . . . . . . . . . ® 19
19a. Body with numerous dark brown to black spots about as large as pupil; pelvic fins dusky
with a basal red brown blotch or streaks, not spotted; total gill rakers on first gill arch 33
to 37; longest dorsal-fin ray 12 to 14% of standard length, about 1/2 length of soft
dorsal-fin base or less . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus picus (adult)
19b. Body with numerous dark brown spots, generally larger than pupil; pelvic fins spotted,
darkening with age; total gill rakers on first gill arch 36 to 43; longest dorsal-fin ray 16 to
25% of standard length, almost equal to length of soft dorsal-fin base in small specimens,
more than 1/2 length of soft dorsal-fin base in adults . . . . . Plectorhinchus chaetodonoides (adult)
20a. Body with large blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 21

20b. Body without large blotches . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
21a. Gill rakers on lower limb of first gill arch 26 to 32 . . . . . Plectorhinchus chaetodonoides (juvenile)
21b. Gill rakers on lower limb of first gill arch 25 or less . . . . . . . . . . . . . . . . . . . . . . . . ® 22
22a. Twelve dorsal-fin spines; light areas with or without dark spots; black areas break up into
dark spots; pectoral fins pale . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus picus (juvenile)
22b. Thirteen dorsal-fin spines; light areas with darker spots that elongate and change into
dark horizontal stripes; pectoral fins black or with black blotches . Plectorhinchus vittatus (juvenile)
23a. Regular dark or light horizontal stripes along body . . . . . . . . . . . . . . . . . . . . . . . . ® 24

23b. Yellow stripes along head and body or on head only . . . . . . . . . . . . . . . . . . . . . . . ® 28
24a. Upper stripes horizontal on body across nape, interorbital, and snout . Plectorhinchus vittatus (adult)
24b. Upper stripes horizontal on body but bending downward and converging on snout . . . . . . . ® 25
25a. Total gill rakers on first gill arch 19 to 22, of which 11 to 15 on lower limb; juveniles with
3 or 4 white stripes on a dark brown or black body, changing to about 4 dark brown
stripes above pectoral-fin base, none below except 2 below eye on head; pelvic fins with
anterior half except margin black or dark brown . . . . . . . . . . . . . . . . Plectorhinchus lessonii
25b. Total gill rakers on first gill arch 26 or more, of which 17 or more on lower limb . . . . . . . . . ® 26
26a. Three narrow light stripes on body fading with age; upper anterior part of soft dorsal fin
dark brown to black; tips of caudal-fin lobes blackish . . . . . Plectorhinchus albovittatus (juvenile)
26b. Usually more than 4 light stripes on body; pelvic fins without an area of black pigmenta-
tion; stripes splitting and increasing with growth . . . . . . . . . . . . . . . . . . . . . . . . . . ® 27
27a. Dorsal-fin spines XIII; stripes increase in number and become diagonal with growth;
upper base of pectoral fins with a splash of carmine or scarlet; caudal fin yellow with
black spots and posterior margin . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus lineatus
27b. Dorsal-fin spines XII (rarely XIII); stripes increase in number, but remain horizontal and
develop a dark edge; stripes flow onto yellow caudal fin in a diagonal manner and fade
with growth; fins yellow, unspotted . . . . . . . . . . . . . . . . . . . . . Plectorhinchus polytaenia
28a. Pale blue or white horizontal stripes edged with dark brown . . . . Plectorhinchus polytaenia (adult)
28b. Stripes not dark edged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 29
29a. Pale blue and bright yellow horizontal stripes on head and body; all fins uniform yellow
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus chrysotaenia
29b. Pale blue and bright yellow horizontal stripes on head only; dorsal fin not uniform yellow . . . . ® 30
30a. Body with yellow, golden, or rusty spots fading away with age; dorsal fin with XIII (rarely
XII) spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus flavomaculatus
30b. Body with narrow yellow lines becoming diagonal posteriorly; dorsal-fin spines XII (rarely
XI or XIII) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Plectorhinchus multivittatus

0 Diagramma labiosum Macleay, 1883
0 Diagramma pictum (Thunberg, 1792)
0 Hapalogenys analis Richardson, 1845
0 Hapalogenys kishinouyei Smith and Pope, 1906
0 Plectorhinchus albovittatus (Rüppell, 1838)
0 Plectorhinchus chaetodonoides Lacepède, 1801
0 Plectorhinchus chrysotaenia (Bleeker, 1855)
0 Plectorhinchus flavomaculatus (Cuvier in Cuvier and Valenciennes, 1830)
0 Plectorhinchus gibbosus Lacepède, 1802
0 Plectorhinchus lessonii (Cuvier in Cuvier and Valenciennes, 1830)
0 Plectorhinchus lineatus (Linnaeus, 1758)
0 Plectorhinchus multivittatus (Macleay, 1878)
0 Plectorhinchus pictus (Tortonese, 1936)
0 Plectorhinchus picus (Cuvier in Cuvier and Valenciennes, 1830)
0 Plectorhinchus polytaenia (Bleeker, 1852)
0 Plectorhinchus schotaf (Forsskål, 1775)
0 Plectorhinchus vittatus (Linnaeus, 1758)
1/
0 Pomadasys argenteus (Forsskål, 1775)

? Pomadasys andamanensis McKay and Satapoomin, 1994
0 Pomadasys argyreus (Valenciennes in Cuvier and Valenciennes, 1833)

0 Pomadasys auritus (Cuvier in Cuvier and Valenciennes, 1830)
0 Pomadasys furcatum (Bloch and Schneider, 1801)
0 Pomadasys kaakan (Cuvier in Cuvier and Valenciennes, 1830)
0 Pomadasys maculatus (Bloch, 1793)
0 Pomadasys trifasciatus Fowler, 1937
1/ Occurrence in the area uncertain. Recently described from Thailand (Phuket Island, Andaman Sea).
2966 Bony Fishes
Diagramma labiosum Macleay, 1883

Frequent synonyms / misidentifications: None / Diagramma pictum (Thunberg, 1792).
FAO names: En - Grey sweetlips.
Diagnostic characters: Lips fleshy, moderately swollen with age. Chin with 6 pores but no median pit.
Total gill rakers on first gill arch 18 to 21 (6 to 8 on upper limb, 1 at corner, and 12 to 14 on lower
limb). Dorsal fin with IX or X spines and 21 to 26 soft rays. Scales ctenoid (rough to touch). Lateral
line with about 50 to 74 tubed scales. Colour: grey bronze to blue-grey with numerous small dark brown
spots on caudal and soft dorsal fins; scattered darker blotches may be present on body and head; no yellow
spots on head at any size; juveniles dark brown to black above, divided into 3 or more broad longitudinal
stripes by narrower pearl blue interspaces; silvery yellow below; dorsal and caudal fins yellow with black
patches and broken stripes; stripes increase with age and break up into small rounded spots before fading
on body and sometimes on caudal fin.
Habitat, biology, and fisheries: Coastal inshore waters near reefs and inside coral lagoons. Taken by
handline and spear. A common species marketed fresh.
Distribution: Bali, Indonesia
to southern New Guinea and
northern Australia from Hout-
m an A b rolhos Islands to
northern New South Wales.
Diagramma pictum (Thunberg, 1792) (Plate XIII, 91 and 92)

Frequent synonyms / misidentifications: Diagramma punctatum Rüppell, 1830; Plectorhinchus pictus
(Thunberg, 1792) / None.
FAO names: En - Painted sweetlips; Fr -Diagramme voilier; Sp - Burro velero.
large adult
juvenile subadults
limb). Dorsal fin with IX or X spines and 20 to 24 soft rays. Scales ctenoid (rough to touch). Lateral
line with about 54 to 73 tubed scales. Colour: adults blue-grey with bright yellow to golden orange spots
sometimes forming lines on head; inside mouth bright orange to red; dorsal fins bordered with black; anal
and pelvic fins tipped with black; juveniles dark brown to black above, divided into 3 or more broad
longitudinal stripes by narrower pearl blue interspaces; silvery yellow below; dorsal and caudal fins yellow
with black patches and broken stripes; stripes increase with age and break up into small rounded spots
before fading on body and sometimes on caudal fin.
Habitat, biology, and fisheries: Coastal inshore waters near reefs and open sandy areas to depth of
80 m. Feeds on bottom inver-
tebrates and small fishes.
Taken by handline and spear.
A common species, marketed
fresh; a small quantity is
salted.
cific from south Natal to
southern Japan and Fiji, ex-
cluding northern Australia
and most of southern New
Guinea.
2968 Bony Fishes
Hapalogenys analis Richardson, 1845

Frequent synonyms / misidentifications: Hapalogenys mucronatus (Eydoux and Souleyet, 1850) /
None.
FAO names: En - Broadbanded velvetchin.
(after Matsubara, 1933)
Diagnostic characters: Chin with 6 pores but no median pit, covered with fleshy villi that frequently
obscure anterior pores. Dorsal fin with XI spines, preceded by an antrorse spine under skin, and 15
or 16 soft rays. Anal fin with III spines, second spine long and strong, and 9 soft rays. Scales ctenoid (rough
to touch). Lateral line with about 54 to 58 tubed scales. Colour: pale brown with 6 vertical broad dark brown
bands; pelvic fins black; spinous dorsal and anal fin interspinous membranes dark brown to black; soft
dorsal, caudal, and soft anal fins yellowish with black margin.
Habitat, biology, and fisheries: Coastal inshore waters near reefs. Feeds on bottom invertebrates and
small fishes. Taken by handline, spear, and occasionally bottom trawl. Uncommon.
from Singapore to southern
Japan.
Hapalogenys kishinouyei Smith and Pope, 1906

FAO names: En - Striped velvetchin.
(after Sainsbury, Kailola, and Leyland, 1985)
Diagnostic characters: Chin with 6 pores but no median pit, covered with fleshy villi that frequently
obscure anterior pores. Dorsal fin with XI spines, an antrorse spine under skin, and 14 soft rays. Anal
fin with III spines, second spine long and strong, and 9 soft rays. Scales ctenoid (rough to touch). Lateral
line with about 58 tubed scales. Colour: pale greyish brown with 4 brown stripes on upper half of body; a
brown band across interorbital behind eye, a broader band across nape; pelvic fins brown in juveniles,
fading with age; spinous dorsal and anal fin interspinous membranes brown; soft dorsal, caudal, and soft
anal fins pale.
Habitat, biology, and fisheries: Coastal inshore waters near reefs. Feeds on bottom invertebrates and
small fishes. Taken by handline, spear, and occasionally bottom trawl. Uncommon.
Distribution: Northern West-
ern Australia, Philippines, Tai-
wan Province of China, South
Korea, and southern Japan.
2970 Bony Fishes
Plectorhinchus albovittatus (Rüppell, 1838) (Plate XIII, 93 to 95)

Frequent synonyms / misidentifications: Plectorhinchus giganteus (Günther, 1879); P. obscurum
(Günther, 1872); P. harrawayi (Smith, 1952) / None.
FAO names: En - Giant sweetlips.
adult
Diagnostic characters: Lips fleshy, greatly swol-

len with age. Chin with 6 pores but no median pit.
Total gill rakers on first gill arch 26 to 31 (6 to 9
on upper limb, 1 at corner, and 19 to 21 on lower
limb). Dorsal fin with XII or XIII (mostly XIII)
spines and 16 to 20 soft rays. Scales ctenoid
(rough to touch). Lateral line with about 54 to 56
tubed scales. Colour: juveniles brown, paler below,
with 3 narrow white bands on body descending juvenile
down nape and snout; spinous dorsal pale brown or
cream, with a narrow dark brown edge, rayed dorsal fin dark brown with a whitish horizontal stripe and a
narrower one above, both reaching posterior margin of fin, last 2 or 3 short rays white, with the posterior
half ray usually dark brown; caudal fin dark brown below, white above, with tips dark brown to black-brown
and a dark brown bar diagonally across upper lobe; anal fin dark brown, posterior rays light brown; pectoral
fins hyaline to very pale brown; pelvic fins tipped with dusky brown. Adults grey to grey-brown with pectoral,
pelvic, and anal fins blackish, lower lobe and upper tip of caudal fin black, anterior half of soft dorsal fin
black, spinous dorsal fin dusky to black; lower lip and chin white; belly silvery grey to white.
Habitat, biology, and fisheries: Coastal inshore waters near reefs, sheltering in the deeper part of surge
channels during the day. Taken by handline and spear. Large fish are marketed fresh, a small quantity is
salted.
cific from East Africa to Red
Sea, India, Indonesia,
Queensland, and northern
New South Wales, Polynesia,
Philippines, and southern Ja-
pan.
Plectorhinchus chaetodonoides Lacepède, 1801 (Plate XIII, 96 to 98)

FAO names: En - Harlequin sweetlips; Fr - Diagramme arlequin; Sp - Burro arlequin.
adult preadult
intermediate phase juvenile
limb). Dorsal fin with XII (rarely XI) spines and 18 to 20 soft rays; height of soft part of dorsal fin about
equal to length of its base. Scales ctenoid (rough to touch). Lateral line with about 52 to 59 tubed scales.
Colour: juveniles brownish with large, well-defined creamy white blotches on body that include brown
spots with age; colour gradually changing into a greyish background with large, deep brown spots.
Size: Maximum total length about 65 cm; commonly to 40 cm.
Habitat, biology, and fisheries: Common in coastal inshore waters near reefs. Taken by handline and
spear. Marketed fresh, a small quantity is salted. Large fish have coarse and dry flesh.
c i fi c from E as t A frica to
Ryukyu Islands and wide
ranging throughout South Pa-
cific eastwards to Rapa.
2972 Bony Fishes
Plectorhinchus chrysotaenia (Bleeker, 1855)

Frequent synonyms / misidentifications: Plectorhinchus celebicus Bleeker, 1873 / None.
FAO names: En - Goldbanded sweetlips.
(after Bleeker, 1876-77)
limb). Dorsal fin with XIII spines and 18 to 21 soft rays. Scales ctenoid (rough to touch). Lateral line
with about 53 to 58 tubed scales. Colour: body and head bright blue-green to sky blue with 5 to 7 bright
yellow longitudinal bands often separated by thinner yellow bands or lines of spots; fins bright yellow to
orange; juveniles similar to adults, but with fewer bands.
Habitat, biology, and fisheries: Coastal inshore waters near reefs. Taken by handline and spear.
Marketed fresh, a small quantity is salted.
cific from Indonesia, New
Guinea, Queensland to Philip-
pines and southern Japan.
Plectorhinchus flavomaculatus (Cuvier in Cuvier and Valenciennes, 1830) (Plate XIV, 99 and 100)
Frequent synonyms / misidentifications: Plectorhinchus reticulatus Günther, 1859 / None.
FAO names: En - Lemon sweetlips; Fr - Diagramme citron; Sp - Burro limón.
adult
Diagnostic characters: Lips fleshy,

moderately swollen with age. Chin
with 6 pores but no median pit. To-
tal gill rakers on first gill arch 26 to
34 (11 to 14 on upper limb, 1 at
corner, and 16 to 19 on lower limb).
Dorsal fin with XII or XIII (mostly
XIII) spines and 19 to 22 soft rays.
Scales ctenoid (rough to touch). Lat-
eral line with about 53 to 60 tubed
scales. Colour: bluish grey, juveniles
with golden or rusty orange stripes
along the body that break into a reticu-
late pattern and later form small spots
that may be lost with age or remain as
intermediate phase
lines, or lines of spots on head.
Size: Maximum total length about
72 cm, commonly to 40 cm.
Habitat, biology, and fisheries: Coastal inshore waters near reefs, seagrass beds, and sandbanks. Taken
by handline and spear. A common species marketed fresh, a small quantity is salted.
Distribution: South Africa to
Red Sea including Madagas-
car, Mauritius, and Comoro Is-
lands to Indonesia, and
western, northern, and east-
ern coast of Australia to Phil-
ippines and Kii Peninsula,
Japan.
2974 Bony Fishes
Plectorhinchus gibbosus Lacepède, 1802 (Plate XIV, 101)

Frequent synonyms / misidentifications: Plectorhinchus nigrus (Cuvier in Cuvier and Valenciennes,
1830) / None.
FAO names: En - Harry hotlips; Fr - Diagramme noire; Sp - Burro negro.
Diagnostic characters: Lips fleshy, greatly swollen with age. Chin with 6 pores but no median pit. Total
gill rakers on first gill arch 24 to 30 (8 to 10 on upper limb, 1 at corner, and 17 to 19 on lower limb).
Dorsal fin with XIV spines (rarely XIII) and 15 to 17 soft rays. Scales ctenoid (rough to touch). Lateral
line with about 46 to 55 tubed scales. Colour: juveniles brownish, caudal fin pink or transparent; adults
uniform dark grey to brownish or blackish; large examples are sometimes mottled with light brown.
Habitat, biology, and fisheries: Coastal inshore waters, common near reefs and estuaries. Taken by
handline and spear. Marketed fresh, a small quantity is salted.
cific from Natal northwards to
Indonesia, Australia, Philip-
pines, China, and Polynesia.
Plectorhinchus lessonii (Cuvier in Cuvier and Valenciennes, 1830) (Plate XIV, 102)
Frequent synonyms / misidentifications: None / Plectorhinchus diagrammus (Linnaeus, 1758).
FAO names: En - Lesson’s sweetlips.
adult
Diagnostic characters: Lips fleshy, moder-

ately swollen with age. Chin with 6 pores
but no median pit. Total gill rakers on
first gill arch 19 to 22 (5 to 7 on upper
limb, 1 at corner, and 11 to 15 on lower
limb). Dorsal fin with XII (rarely XIII)
spines and 19 to 21 soft rays. Scales
ctenoid (rough to touch). Lateral line with
about 54 to 58 tubed scales. Colour: juve- juvenile
niles dark brown with 2 white stripes on body
and through caudal fin, increasing to 4; adult with about 4 brown stripes on body above pectoral fins and
2 brown stripes below eye on head, belly white; middle of pectoral fins black in juveniles, becoming
colourless with base red-brown; pelvic fins pale with the anterior basal half of membranes black, becoming
brown; dorsal, caudal, and anal fins yellow, spotted and margined with black.
Habitat, biology, and fisheries: Coastal and offshore waters near coral reefs. Taken by handline and
spear. Marketed fresh.
Distribution: Zanzibar, Red
Sea to Sri Lanka, Indonesia,
Queensland, Polynesia to
Philippines, Taiwan Province
of China, and southern Ja-
pan.
2976 Bony Fishes
Plectorhinchus lineatus (Linnaeus, 1758) (Plate XIV, 103)

Frequent synonyms / misidentifications: Plectorhinchus goldmanni (Bleeker, 1853) / None.
FAO names: En - Lined sweetlips.
limb). Dorsal fin with XIII spines and 18 to 20 soft rays. Scales ctenoid (rough to touch). Lateral line
with about 54 to 58 tubed scales. Colour: juveniles with few horizontal to slightly oblique dark brown bands
which subdivide with age until the body appears to be dark chocolate brown above, with a network of fine
white lines running up and back, the broad brown bars breaking up into smaller and smaller spots on lower
sides; belly white; lips and fins bright yellow, anal and caudal fins dotted with black; pectoral fins with a
splash of bright crimson or scarlet red at the base.
Habitat, biology, and fisheries: Coastal and offshore waters near coral reefs. Taken by handline and
spear. Marketed fresh, a small quantity is salted.
cific from Andaman Sea to In-
donesia, New Guinea,
Queensland, New Caledonia,
Philippines, China (including
Taiwan Province) to Ryukyu
Islands.
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Plectorhinchus multivittatus (Macleay, 1878)

Frequent synonyms / misidentifications: None / Plectorhinchus ordinalis (Scott, 1962).
FAO names: En - Manybanded sweetlips.
limb). Dorsal fin with XII (rarely XI or XIII) spines and 18 to 21 soft rays. Scales ctenoid (rough to
touch). Lateral line with about 56 to 58 tubed scales. Colour: blue-grey above, white below; 8 or 9 bright
yellow or yellow-orange longitudinal bands on head, extending onto body and remaining distinct on lower
half, but becoming broken into a series of dots and short bands on upper half of body; 2 distinct rows of
similarly coloured spots along entire length of dorsal fin; all other fins transparent light yellow.
Marketed fresh. Uncommon.
Distribution: Northern Aus-
tralia from Shark Bay to Torres
Straits, and Taiwan Province
of China.
2978 Bony Fishes
Plectorhinchus pictus (Tortonese, 1936) (Plate XIV, 104)

Frequent synonyms / misidentifications: Plectorhinchus fangi (Whitley, 1951); Plectorhynchus sinensis
Chu, Wu, and Jin, 1977 / Plectorhinchus cinctus (Temminck and Schlegel, 1843).
FAO names: En - Trout sweetlips; Fr - Diagramme truité; Sp - Burro trucha.
limb). Dorsal fin with XII spines and 15 to 17 (mostly 16) soft rays. Scales ctenoid (rough to touch).
Lateral line with about 48 to 58 tubed scales. Colour: greyish brown, most of the body with rounded black
spots about equal in size to interspaces, larger than interspaces in young; pectoral fins olive without spots,
pelvic fins dusky anteriorly without spots or blackish, anal and caudal fins dark brown to blackish, the outer
three-fifths of the caudal fin lighter in juveniles; juveniles have 3 longitudinal brownish bands which may
be broken into blotches and with numerous dark spots about equal to eye diameter between the 2
uppermost bands.
Distribution: Gulf of Oman
eastwards along the Indian
coast, Gulf of Manaar, Thai-
land and China.
Plectorhinchus picus (Cuvier in Cuvier and Valenciennes, 1830) (Plate XIV, 105 and 106; Plate XV, 107)
Frequent synonyms / misidentifications: Plectorhinchus punctatissimus (Playfair, 1867) / None.
FAO names: En - Spotted sweetlips.
adult
Diagnostic characters: L i p s f l e s h y ,
greatly swollen with age. Chin with 6
pores but no median pit. Total gill rak-
ers on first gill arch 33 to 37 (8 to 13 on
upper limb, 1 at corner, and 22 to 26 on
lower limb). Dorsal fin with XII spines
and 18 to 22 soft rays. Scales ctenoid
(rough to touch). Lateral line with about 62
to 70 tubed scales. Colour: juveniles are juvenile
creamy white below, blackish above with
white snout and a pair of white saddles and
spots. Adults become covered with dark brown spots on head, back, sides, and fins; dorsal fin dark brown
to blackish with a median horizontal grey stripe on membranes only and with a series of large black spots
and sometimes with a narrow dark margin; pectoral and pelvic fins with the base and axil vermillion in some
specimens; inside of mouth, hind edge of maxilla, gill-cover membrane and base of pectoral fins vermillion,
scarlet, or red-brown; belly pale mauve grey.
Habitat, biology, and fisheries: Coastal waters near reefs. Taken by handline and spear. Sometimes
abundant. Marketed fresh, a small quantity is salted.
cific from East Africa, Great
Barrier Reef to southern Ja-
pan and eastwards to the So-
ciety Islands and Rapa.
2980 Bony Fishes
Plectorhinchus polytaenia (Bleeker, 1852) (Plate XV, 108)

FAO names: En - Ribboned sweetlips; Fr - Diagramme rubanne; Sp - Burro listado.
limb). Dorsal fin with XII (rarely XIII) spines and 19 to 22 soft rays. Scales ctenoid (rough to touch).
Lateral line with about 54 to 60 tubed scales. Colour: brown to yellowish grey with 5 to 9 fairly narrow grey
or white longitudinal stripes outlined with dark brown on body and continuing around snout; fins yellow,
soft dorsal, caudal, and pectoral fins with darker stripes disappearing with age; eye and lips yellowish;
mouth, tongue, and gill rakers scarlet; chin white; juveniles have fins striped and fewer stripes on body.
cific from the west coast of
I n d i a , W e s te r n A u s tr a li a
northwards to Torres Straits
and to the Philippines.
Plectorhinchus schotaf (Forsskål, 1775) (Plate XV, 109)

FAO names: En - Minstrel sweetlips; Fr - Diagramme ménestrel; Sp - Burro trovador.
limb). Dorsal fin with XII (rarely XIII) spines and 18 to 21 soft rays. Scales ctenoid (rough to touch).
Lateral line with about 54 to 58 tubed scales. Colour: grey, silver, or greyish brown, pale grey to whitish
below; head darker dorsally with opercular membrane crimson, scarlet red to orange-red; base of pectoral
fins dark red; inside of mouth and tongue bright reddish; juveniles sometimes with narrow faint blue lines
on body and below eye; fins usually dark. Freshly caught fish often have irregular broad pale bars and
blotches which disappear after death.
cific from Port St. Johns, Tran-
skei, East Africa to northern
Australia, the Philippines, and
southern Japan. Red Sea
population differs slightly in
coloration.
2982 Bony Fishes
Plectorhinchus vittatus (Linnaeus, 1758) (Plate XV, 110)

Frequent synonyms / misidentifications: Plectorhinchus orientalis (Bloch, 1793) / None.
FAO names: En - Oriental sweetlips; Fr - Diagramme oriental; Sp - Burro oriental.
adult
Diagnostic characters: Lips fleshy, greatly

swollen with age. Chin with 6 pores but no
median pit. Total gill rakers on first gill arch
29 to 34 (9 to 11 on upper limb, 1 at corner,
and 20 to 23 on lower limb). Dorsal fin with
XIII (rarely XII or XIV) spines and 17 to 20
soft rays. Scales ctenoid (rough to touch).
Lateral line with about 55 to 65 tubed scales.
Colour: juveniles have connected black
blotches and spots that gradually break up
into horizontal stripes; pectoral fins black in juvenile
juveniles becoming uniform yellow in adults;
tail spotted with age. Adults with 6 to 12 broad
dark brown, blue-brown or black stripes that persist on the belly and join horizontally across the nape and
snout with anterior part of pale interspaces yellow forming 2 bright yellow stripes across interorbital and 2
around snout; eye usually yellow; fins yellow with black margins to vertical fins and black spots, pectoral
fins uniform yellow with red-brown, chocolate, or blackish base, pelvic fins yellow with base red-brown,
scarlet, or dark brown.
cific from East Africa to Indo-
nesia, northern Australia,
Thailand, Philippines, and
Polynesia.
Pomadasys argenteus (Forsskål, 1775) (Plate XV, 111) GRL

Frequent synonyms / misidentifications: Pomadasys hasta (Bloch, 1790) / Pomadasys opercularis
Playfair in Playfair and Günther, 1867.
FAO names: En - Silver grunt; Fr - Grondeur argenté; Sp - Corocoro plateado.
adult
Diagnostic characters: Body depth 2.7 to 2.8

times in standard length. Head blunt, its upper
profile convex; mouth small; maxilla reaching to
eye; teeth in jaws arranged in villiform (brush-like)
bands; 2 pores and a median pit on chin. Dorsal
fin with XII spines and 14 soft rays; anal fin
with III spines and 7 soft rays. Scales ctenoid
(rough to touch); lateral line with 45 to 50 scales;
5 scales between lateral line and dorsal-fin
origin; 21 or 22 scales around caudal peduncle,
9 above lateral line, 10 or 11 below. Swimbladder
without anterior extensions. Colour: juveniles with juvenile
body pale brownish, lighter below, back with ir-
regular longitudinal streaks on alternate scale
rows; dorsal fin with dusky membranes; a dark spot on gill cover. Adults with silvery body, dark blotches
on dorsal fin and numerous scattered dark brown to blackish spots on body.
Habitat, biology, and fisheries: Coastal inshore waters in open bays and estuaries. Taken by handline,
set net, trap, and spear. Mar-
keted fresh, a small quantity is
salted. From 1990 to 1995, the
FAO Yearbook of Fishery Sta-
tistics reports a range of yearly
catch of 537 to 1 239 t of Po-
madasys argenteus from the
area (Malaysia only).
Distribution: R e d S e a t o
southern Japan, including
northern Australia from
Exmouth Gulf to Clarence
River, New South Wales.
2984 Bony Fishes
Pomadasys argyreus (Valenciennes in Cuvier and Valenciennes, 1833)

FAO names: En - Bluecheek silver grunt; Fr - Grondeur à joues bleus; Sp - Corocoro mejilla azul.
Diagnostic characters: Body depth 2.3 to 2.7 times in standard length. Head blunt, its upper profile
convex; mouth small; maxilla reaching to eye; teeth in jaws arranged in villiform (brush-like) bands; 2 pores
and a median pit on chin. Dorsal fin with XII spines and 13 soft rays; anal fin with III spines and 7
soft rays. Scales ctenoid (rough to touch); lateral line with 42 to 47 scales; 6 scales between lateral line
and dorsal-fin origin; 18 to 20 scales around caudal peduncle, 7 to 9 above lateral line, 9 below.
Swimbladder more or less truncate anteriorly, without anterior extensions. Colour: body silvery; gill cover
with a large blue-black blotch extending onto opercle and becoming indistinct in adults; peritoneum pale
sandy brown; alimentary canal pale (not black).
Habitat, biology, and fisheries: Coastal inshore waters in open bays and on trawling grounds. Taken by
handline, set net, bottom trawl, trap, and spear. Marketed fresh, a small quantity is salted.
Distribution: India, Sri Lanka
to Papua New Guinea, ex-
cluding northern Australia.
Pomadasys auritus (Cuvier in Cuvier and Valenciennes, 1830)

FAO names: En - Longhead grunt.
Diagnostic characters: Body depth 3.2 times in standard length. Head large, its upper profile convex, gill
cover and angle of preopercle lengthened as convex rounded lobes, gill cover extending to above
one-third to one-half of pectoral fins; mouth small; maxilla reaching to eye; teeth in jaws arranged in
villiform (brush-like) bands; 2 pores and a median pit on chin. Dorsal fin with XII spines and 14 soft
rays; anal fin with III spines and 7 soft rays. Scales ctenoid (rough to touch); lateral line with 43 scales;
5 scales between lateral line and dorsal-fin origin; 16 scales around caudal peduncle, 7 above lateral
line, 7 below. Swimbladder morphology unknown. Colour: body silvery to light burnished gold; dorsal fins
with well defined blackish spots on interspinous membranes; adults with a blackish spot on tip of each
scale on back and upper sides.
Size: Maximum total length about 52 cm; none captured less than 43 cm.
Habitat, biology, and fisheries: Unknown, perhaps from deeper waters of continental shelf. A rare
species. Specimens wanted by museums.
Distribution: China, Thai-
land, Singapore, Malaysia,
and New South Wales, Aus-
tralia.
2986 Bony Fishes
Pomadasys furcatum (Bloch and Schneider, 1801) (Plate XV, 112)

Frequent synonyms / misidentifications: Rhonciscus furcatus (Bloch and Schneider, 1801) / None.
FAO names: En - Banded grunt; Fr - Goret à six bandes; Sp - Ronco de seis bandas.
Diagnostic characters: Body depth 2.9 to 3 times in standard length. Head profile convex; mouth small;
maxilla reaching to hind nostril or eye; teeth in jaws arranged in villiform (brush-like) bands; 2 pores and
a median pit on chin. Dorsal fin with XII spines and 14 to 16 soft rays; anal fin with III spines and
8 or 9 soft rays; soft portions of dorsal and anal fins with a row of interradial scales. Scales ctenoid
(rough to touch); lateral line with 52 to 54 scales; 8 or 9 scales between lateral line and dorsal-fin origin;
25 or 26 scales around caudal peduncle, 11 above lateral line, 12 or 13 below. Swimbladder without
anteriolateral extensions. Colour: body light purplish brown above, paler below, with about 6 longitudinal
dark stripes that are usually bifurcate anteriorly; the stripes becoming progressively more bifurcate or
double with age.
Habitat, biology, and fisheries: Coastal waters near reefs. Caught with handlines and gill nets. Marketed
fresh.
Distribution: Malaysia, south-
ern Indonesia, Thailand to
China.
Pomadasys kaakan (Cuvier in Cuvier and Valenciennes, 1830) (Plate XV, 113)
Frequent synonyms / misidentifications: None / Pomadasys hasta (Bloch, 1790).
FAO names: En - Javelin grunt; Fr - Grondeur javelot; Sp - Corocoro jabalina.
adult
Diagnostic characters: Body depth 2.5 to 3

times in standard length. Head blunt, its upper
profile convex; mouth small; maxilla reaching to
eye; teeth in jaws arranged in villiform (brush-
like) bands; 2 pores and a median pit on chin.
Dorsal fin with XII spines and 13 to 15 (usu-
ally 14 ½) soft rays; anal fin with III spines
and 7 soft rays. Scales ctenoid (rough to
touch); lateral line with 43 to 50 scales; 7 scales
between lateral line and dorsal-fin origin; 20 subadult
scales around caudal peduncle, 9 above lat-
eral line, 9 below. Swimbladder without anteriolateral horn-like extensions. Colour: juveniles brilliant
silvery green above to golden silver on sides and silvery white on belly with 12 or more faint vertical bars
comprised of small dark brown spots or irregular blotches; dorsal fin spotted with black-brown spots on
lower half of spinous dorsal fin and in 3 rows on rayed dorsal fin; margin of soft dorsal fin darker and lower
tip of caudal fin milk white. Adults uniform golden green above, silvery below with traces of the vertical
bars present only when alive and spots on dorsal fins indistinct or absent.
Habitat, biology, and fisheries: Coastal inshore waters in open bays and estuaries, penetrating rivers to
the upper limit of brackish water. Taken by handline, set net, trap, and spear. Marketed fresh, a small
quantity is salted or smoked.
cific from Transkei to Red
Sea, Persian Gulf, India, Indo-
nesia to China (including Tai-
wan Province), including
Exmouth Gulf to Moreton Bay.
2988 Bony Fishes
Pomadasys maculatus (Bloch, 1793) (Plate XV, 114)

FAO names: En - Saddle grunt (= blotched grunt, Fishing Areas 57/71); Fr - Grondeur selle; Sp - Corocoro
montura.
convex; mouth small; maxilla reaching to eye; teeth in jaws arranged in villiform (brush-like) bands; 2 pores
and a median pit on chin. Dorsal fin with XII spines and 13 to 15 soft rays; anal fin with III spines
and 7 (rarely 6 or 8) soft rays. Scales ctenoid (rough to touch); lateral line with 50 to 55 scales; 7 to 9
scales between lateral line and dorsal-fin origin; 22 to 25 scales around caudal peduncle, 9 or 10
above lateral line, 11 or 12 below. Colour: silvery grey, head purplish or brownish, nape and back with
a series of incomplete cross bars (one on nape most conspicuous); spinous part of dorsal fin with a large
black blotch; dorsal and caudal fins edged with black, other fins yellowish.
Habitat, biology, and fisheries: Coastal inshore waters in open bays, trawling grounds, and estuaries.
Taken by bottom trawl, handline, set net, trap, and spear. Frequently abundant on prawn trawl grounds.
Marketed fresh, a small quantity is salted or smoked.
Distribution: East coast of
Afric a, Madagascar, Red
Sea, Gulf of Aden, Persian
Gulf, Pakistan, India, Sri
Lanka to northern half of Aus-
tralia from Shark Bay to More-
ton Bay, New Guinea,
Philippines to southern Ja-
pan.
Pomadasys trifasciatus Fowler, 1937

FAO names: En - Blackeared grunt.
juvenile
convex; mouth small; maxilla reaching to anterior margin of eye or pupil; teeth in jaws arranged in villiform
(brush-like) bands; 2 pores and a median pit on chin. Dorsal fin with XII (rarely XIII) spines and 13
(rarely 12 or 14) soft rays; anal fin with III spines and 7 soft rays. Scales ctenoid (rough to touch);
lateral line with 43 to 50 (usually 47) scales; 6 scales between lateral line and dorsal-fin origin; 19 to
23 scales around caudal peduncle, 8 to 10 above lateral line, 9 to 11 below (mostly 9 above, 2 close
to, and 11 below, lateral line). Colour: juveniles with 3 dark stripes, the middle one wide, becoming faded
or absent with growth; a distinct black-brown spot on gill cover; dorsal fins without dark spots or blotches;
adults plain or with faint stripes.
Habitat, biology, and fisheries: Coastal inshore waters in open bays and mouths of rivers. Taken by
bottom trawl and set net. No
fishery has yet developed.
Abundant on some prawn
trawling grounds.
Distribution: Bay of Bengal
to Sumatra, Bali, Lombok,
Joseph Bonaparte Gulf to
Bowen, Gulf of Thailand, and
Philippines. Possibly more
widespread.
click for next page

2990 Bony Fishes
SPARIDAE
Porgies (seabreams)
by K.E. Carpenter
D iagnostic characters: Body oblong, moderately deep and compressed (size to 75 cm). Head large,
often with a steep upper profile; snout scaleless, cheeks scaly, preopercle with or without scales,
without spines or serrations on margin; opercle scaly, without spines. Mouth subhorizontal and slightly
protrusible, upper jaw never reaching backward beyond a vertical line through middle of eye; hind
tip of premaxilla overlapping maxilla; maxilla in sheath formed by first 2 circumorbital bones when
mouth closed; jaw teeth well developed, differentiated into either conical (canine-like), or flattened
(incisor-like), and often rounded (molar-like); roof of mouth (vomer and palatines) toothless. Gill rakers
variable, 7 to 20 on lower limb of first gill arch. Dorsal fin single, with X to XIII spines and 9 to 17 soft
rays (last ray double, counted as 1), the spiny and soft portions not separated by a notch, anterior spines
sometimes elongate or filamentous; anal fin with III spines and 7 to 15 soft rays (last ray double, counted
as 1), the spines, especially the second, often stout; caudal fin moderately deeply emarginate or forked;
pectoral fins usually long and pointed; pelvic fins below or just behind pectoral-fin bases, with I spine and
5 soft rays, and with an axillary scale at their base (axillary pelvic process). Scales cycloid (smooth) or
weakly ctenoid (rough to touch); a single continuous lateral line extending backward to base of caudal fin.
Colour: overall colour highly variable, from pinkish or reddish to yellowish or greyish, often with silvery or
golden reflections, often with dark or coloured spots, stripes, or bars.
cheek scaly
axillary scale
maxilla
premaxilla
Habitat, biology, and fisheries: Seabreams inhabit tropical and temperate coastal waters. They are
demersal inhabitants of the continental shelf and the slope. The smaller species, as well as the young of
large species, usually form aggregations, while large adults are less gregarious or solitary and occur in
deeper waters. Occasionally they are found in estuaries which are used as nurseries. Hermaphroditism is
widespread in this family. Most seabreams are excellent food fish and are of notable commercial importance.
From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports a range of yearly catch of around 2 170
to 4 020 t of Sparidae from the Western Central Pacific.
Perciformes: Percoidei: Sparidae 2991

Haemulidae: edge of preopercle serrate; suborbital space scaly; at least 2 conspicuous pores beneath chin;
never with molar teeth.
Lutjanidae: edge of preopercle usually serrate, and often excavated to accommodate a bony knob; never
molar or incisor-like teeth; roof of mouth usually toothed (except in Aphareus).
Lobotidae: edge of preopercle strongly toothed; never molar teeth; dorsal, anal, and caudal fins rounded,
appearing as a single tri-lobed fin (‘triple-tails’).
serrate
scales
strongly
serrate toothed
Haemulidae Lutjanidae Lobotidae

Serranidae: edge of preopercle serrate; suborbital space scaly; similar species lack axillary scale of pelvic
fins; caudal fin generally rounded, never forked; also never molar teeth.
Lethrinidae: posterior tip of maxilla overlapping premaxilla (posterior tip of premaxilla overlapping maxilla in
Sparidae); cheek and preopercular flange scaleless (but scales present on cheek in Wattsia, Monotaxis,
Gymnocranius, and Gnathodentex); 8 to 11 soft dorsal-fin rays (9 to 17 in Sparidae); incisor-like teeth never
present.
cheek and
preopercular
flange
scaleless
Serranidae Lethrinidae (Lethrinus)

Kyphosidae: head entirely scaly, except for snout; teeth in jaw incisor-like, close-set, and of a peculiar
hockey-stick shape, with their base set horizontally, resembling a radially striated bone inside mouth.
Cirrhitidae and Cheilodactylidae: lower pectoral-fin rays simple and thickened.
teeth
horizontal
tooth bases
ventral view of individual tooth

upper jaw (lateral view)
Cirrhitidae
incisor-like hockey
stick-shaped teeth
Kyphosidae
Cheilodactylidae
2992 Bony Fishes
lateral teeth pointed
Key to the species of Sparidae occurring in the area
1a. Only conical teeth laterally in jaws (Fig. 1) . . . (Dentex) ® 2
1b. Lateral teeth include molars . . . . . . . . . . . . . . . ® 3
canines
2a. Dorsal fin with XII spines and 10 soft rays
(Fig. 2) . . . . . . . . . . . . . . . . . . . Dentex tumifrons
2b. Dorsal fin with XIII spines and 9 soft rays
(Fig. 3) . . . . . . . . . . . . . . . . . Dentex fourmanoiri
Fig. 1 Dentex
Fig. 2 Dentex tumifrons Fig. 3 Dentex fourmanoiri
3a. Interorbital area scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4

3b. Interorbital area not scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
4a. No elongate or extended dorsal-fin spines (Fig. 4a, b) . . . . . . . . . . . . . . . . . Pagrus auratus
4b. Dorsal fin with third and fourth spines elongate and filamentous . . . . . . . . . . . . . . . . . . ® 5
a) small adult b) large adult
Fig. 4 Pagrus auratus
5a. First 2 dorsal-fin spines minute (Fig. 5a);

8 soft rays in anal fin (Fig. 6) . . . . Argyrops spinifer
5b. First 2 dorsal-fin spines well developed
(Fig. 5b); 9 soft rays in anal fin . . . . (Evynnis) ® 6
a) Argyrops b) Evynnis
Fig. 5 dorsal-fin spines Fig. 6 Argyrops spinifer
6a. Transverse series of scales between lateral line and fourth dorsal-fin spine usually 5 ½
(rarely 4 ½); vomer with conical teeth (Fig. 7) . . . . . . . . . . . . . . . . . . . . . Evynnis japonica
6b. Transverse series of scales between lateral line and fourth dorsal-fin spine usually 6 ½
(rarely 5 ½); vomer toothless . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Evynnis cardinalis
(only unconfirmed and doubtful reports from the Philippines)
7a. Transverse series of scales between lateral line and fourth dorsal-fin spine 5 ½ or 6 ½;
dorsal-fin spines moderately slender, not strongly appearing alternately broad and
narrow on each side; 1 enlarged molar posteriorly on each jaw (Figs 8a and 9) . Rhabdosargus sarba
7b. Transverse series of scales between lateral line and fourth dorsal-fin spine 3 ½ or 4 ½;
dorsal-fin spines strong, appearing alternately broad and narrow on each side; no single
greatly enlarged molar posteriorly in jaws (Fig. 8b) . . . . . . . . . . . . . . . (Acanthopagrus) ® 8
front
teeth
enlarged
posterior
molar
a) Rhabdosargus b) Acanthopagrus
Fig. 7 Evynnis japonica Fig. 8 right half of upper jaw and roof of mouth
8a. Pelvic, anal, and caudal fins, particularly the spines and soft rays, blackish; margin of
soft-rayed part of anal fin rounded posteriorly (Fig. 10) . . . . . . . . . . . . . Acanthopagrus berda
8b. Pelvic and anal fins and ventral part of caudal fin, particularly the spines and soft rays,
yellowish and whitish, sometimes with blackish areas on inter-ray membranes; margin
of soft-rayed part of anal fin nearly straight . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
Fig. 9 Rhabdosargus sarba Fig. 10 Acanthopagrus berda

2994 Bony Fishes
9a. Transverse series of scales between lateral line and median dorsal-fin spines typically
4 ½ (rarely 3 ½) (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthopagrus australis
(southeastern Australia, from central Queensland south to eastern Victoria)
9b. Transverse series of scales between lateral line and median dorsal-fin spines typically
3 ½ (rarely 4 ½) (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthopagrus latus
(in the area, northern Australia, and northwestern South China Sea, and northernmost Luzon)
Fig. 11 Acanthopagrus australis Fig. 12 Acanthopagrus latus

The symbol 0 is given when species accounts are included. A question mark indicates that presence in
the area is uncertain.
0 Acanthopagrus australis (Günther, 1859)
0 Acanthopagrus berda (Forsskål, 1775)
0 Acanthopagrus latus (Houttuyn, 1782)
? Acanthopagrus palmaris (Whitley, 1935)
0 Argyrops spinifer (Forsskål, 1775)
0 Dentex fourmanoiri Akuzaki and Séret, 1999
0 Dentex tumifrons (Temminck and Schlegel, 1842)
0 Evynnis japonica Tanaka, 1931
? Evynnis cardinalis (Lacepède, 1802)
0 Pagrus auratus (Bloch and Schneider, 1801)

0 Rhabdosargus sarba (Forsskål, 1775)
References
Akazaki, M. and B. Séret. 1999. Dentex fourmanoiri , a new species of sea bream (Sparidae: Denticinae) from off New
Caledonia. In Proc. 5th Indo-Pac. Fish Conf. (Nouméa, 3-8 Nov.1997), edited by B. Séret and J.-Y. Sire. Paris,
Soc. Fr. Ichtyol., pp. 177-183.
Gloerfelt-Tarp, T. and P. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Australia . Jakarta,
Kuiter, R.H. 1993. Coastal fishes of South-eastern Australia. Honolulu, Univ. Hawaii Press, 437 p.
Acanthopagrus australis (Günther, 1859)

FAO names: En - Australian seabream.
(after Grant, 1975)
Diagnostic characters: Body fairly deep and compressed, its depth about 2.1 to 2.3 times in standard
length. Head 2.8 to 3 times in standard length, its upper profile straight (sometimes a bulge above eye);
snout pointed; eye moderate in size. In both jaws, 4 to 6 large, moderately compressed teeth in front,
followed by 4 or 5 rows of molar-like teeth; upper lateral teeth of outer row conical and blunt. First gill
arch with 9 gill rakers on lower limb. Dorsal fin with XI or XII spines and 10 or 11 soft rays, fourth to sixth
spines longest (spines appear alternately broad and narrow on either side). Anal fin with III spines and 8
soft rays, first spine shorter than eye diameter, second spine flattened laterally, much longer and
stronger than third; margin of soft-rayed part of anal fin nearly straight. Pelvic fins with I strong spine.
Caudal fin slightly forked, with rounded lobes. Scales large, 43 to 46 in lateral line (to base of caudal fin);
4 ½ (rarely 3 ½) scale rows between lateral line and fourth dorsal-fin spine; a scaly sheath at base of
dorsal and anal fins; a long pelvic axillary process. Colour: body silvery bronze; pelvic and anal fins and
ventral part of caudal fins, particularly the spines and soft rays yellowish and whitish, sometimes
with blackish areas on inter-ray membranes; a dark blotch at origin of lateral line and a dark spot at upper
base of pectoral fins.
Habitat, biology, and fisheries: Most common in estuaries but also found on inshore reefs, and sandy,
muddy, and rocky coastal areas. Feeds on a wide variety of benthic crustaceans and molluscs and small
fish. One of the most common fishes caught by anglers where it occurs throughout its range. A popular food
fish in Australia, caught mostly by hook-and-line by anglers and commercially caught with tunnel nets, gill
nets, and to a lesser extent as bycatch in trawling and trapping fisheries.
Distribution: Australia, from
Townsville south on the east
coast and from Barrow Island
south on the west coast.
2996 Bony Fishes
Acanthopagrus berda (Forsskål, 1775) (Plate XVI, 115)

Frequent synonyms / misidentifications: Mylio berda (Forsskål, 1775) / None.
FAO names: En - Picnic seabream; Fr - Pagre picnic; Sp - Sargo picnic.
Diagnostic characters: Body fairly deep and compressed, its depth about 2 times in standard length. Head
3 or 4 times in standard length, its upper profile straight (sometimes a bulge above eye); snout pointed;
eye moderate in size; ventral profile almost straight to anus. In both jaws, 4 to 6 large, moderately
compressed teeth in front, followed by 3 to 5 rows of molar-like teeth; upper lateral teeth of outer row
conical and blunt. Gill rakers 9 to 11 on lower limb of first gill arch. Dorsal fin with XI (rarely XII) spines and
10 to 13 (usually 11 or 12) soft rays, fourth to sixth spines longest (spines appear alternately broad and
narrow on either side). Anal fin with III spines and 8 or 9 soft rays, first spine shorter than eye diameter,
second spine flattened laterally, much longer and stronger than the third; margin of soft-rayed part
of fin convex, rounded. Pelvic fins with I strong spine. Caudal fin slightly forked, with rounded lobes. Scales
large, 43 to 45 in lateral line (to base of caudal fin); usually 3 ½ scale rows between lateral line and fourth
dorsal-fin spine; a scaly sheath at base of dorsal and anal fins; a long pelvic axillary process. Colour: grey,
dark silver grey or dull olive brown with silvery or brassy reflections upper part of body and base of scales
darkest, lower part of head and body paler, a dark edge along opercle; spinous dorsal fin with a dark edge
and spines often silvery; pectoral fins dusky with a yellow tinge, soft dorsal, anal, pelvic, and caudal fins
blackish; caudal fin grey with darker shading.
Habitat, biology, and fisheries: A bottom-living fish, very shy, cunning. Found mainly on rough and muddy
sand grounds in coastal waters, particularly in estuaries, from shallow water to depths of about 50 m.
Juveniles usually occur in more sheltered shallow bays and estuaries. Feeds on a wide variety of bottom
invertebrates (worms, molluscs, crustaceans, echinoderms, etc.). Not intensively fished. Caught with bottom
trawls and handlines. Mar-
keted fresh; the flesh is excel-
lent.
Indo-West Pacific from South
Africa to India, extending to
western Pacific, Japan, and
northern Australia.
Acanthopagrus latus (Houttuyn, 1782) (Plate XVI, 116)

Frequent synonyms / misidentifications: Sparus latus Houttuyn, 1782; Mylio latus (Houttuyn, 1782) /
None.
FAO names: En - Yellowfin seabream; Fr - Pagre à nageoires jaunes; Sp - Sargo aleta amarilla.
Diagnostic characters: Body fairly deep and compressed, its depth about 2 times in standard length. Head
3 times in standard length, its upper profile notably convex and angular due to a bulge at eye; eye moderate
in size. In both jaws, 4 to 6 large, moderately compressed teeth in front, followed by 3 to 5 rows of
molar-like teeth; upper lateral teeth of outer row conical and blunt. Dorsal fin with XI spines and 11 or 12
soft rays, fourth to sixth spines longest (spines appear alternately broad and narrow on either side). Anal
fin with III spines and 8 or 9 soft rays, the first spine shorter than eye diameter, second spine flattened
laterally, much longer and stronger than third; margin of soft-rayed part of anal fin nearly straight.
Pelvic fins with I strong spine. Caudal fin slightly forked, with tips sharp. Scales large, about 48 to 50 in
lateral line (to base of caudal fin); 3 ½ (rarely 4 ½) rows between lateral line and fourth dorsal-fin spine;
a scaly sheath at base of dorsal and anal fins; a long axillary pelvic process. Colour: pale grey to whitish,
darker above, belly usually yellowish; golden streaks along longitudinal rows of scales; a diffuse dark blotch
at origin of lateral line; a dark interorbital band and a dark edge along opercle; dorsal fin greyish to hyaline,
pelvic and anal fins yellowish and whitish, sometimes with blackish areas on inter-ray membranes;
caudal fin greyish with dark blackish margin, lower part of fin with yellowish tinge; caudal fin with a
black margin, lower portion of fin with yellowish tinge.
Habitat, biology, and fisheries: Inhabits shallow coastal waters to depths of about 50 m; enters estuaries.
Carnivorous, feeds on invertebrates, mainly echinoderms, worms, crustaceans, and molluscs. Caught with
bottom trawls and lines. Mainly exploited by artisanal fisheries. Marketed fresh, whole.
cific from the Persian Gulf and
along the coasts of India; east-
ward extending to the Philip-
pines, Japan, and northern
Australia.
2998 Bony Fishes
Argyrops spinifer (Forsskål, 1775) (Plate XVI, 117 and 118)

FAO names: En - King soldierbream; Fr - Spare royal; Sp - Sargo real.
(after Sainsbury et al., 1985)
Diagnostic characters: Body deep, strongly compressed. Upper profile of head steep and almost straight
from upper jaw to eye; eye moderately large. At front of jaws, 4 upper and 4 to 6 lower enlarged canines,
followed in both jaws by 2 rows of molars, those of outer row bluntly conical anteriorly; small molars behind
canines. Gill rakers 10 or 11 on lower limb of first gill arch. Dorsal fin with XI or XII spines and 10 or 11 soft
rays, the first 2 spines very short, third to fifth spines and sometimes to seventh spine flattened and
much elongated (in young reaching to level of caudal fin, shorter in old fish). Anal fin with III spines (second
and third subequal) and 8 soft rays. Caudal fin deeply emarginate to forked, with pointed lobes. Scales
large, 50 to 54 in lateral line, scaly area between eyes narrow, ending in a point; soft dorsal and anal fins
with low basal scaly sheaths. Colour: body mainly silvery pinkish, darker on head; usually dark red on
margin of upper part of opercle; all fins red; young fish with several vertical red bars on body.
Habitat, biology, and fisheries: This common species inhabits a wide range of bottoms (usually at depths
common between 5 and 100 m). The young individuals occur in very shallow waters of sheltered bays, the
larger ones at greater depths. Feeds on bottom-living invertebrates, mainly molluscs. Caught with bottom
trawls, longlines, handlines, stake traps, and fish traps. Mostly exploited by artisanal fisheries. Usually
marketed fresh.
Distribution: W i d e s p r ea d
Africa, including the Red Sea
and Persian Gulf, to the Indo-
Malayan Archipelago to south-
e r n J a pa n a nd n or t he r n
Australia.
Dentex fourmanoiri Akazaki and Séret, 1999

FAO names: En - Fourmanoir’s seabream.
(after Akazaki and Séret, 1999)
Diagnostic characters: Body moderately deep and compressed, body depth greater than head length;
profile of head with a bulge near eye; eye moderately large, 2.4 to 3.35 times in head length. Front of upper
jaw with 4 strong canines and lower jaw with 4 to 6 canines; a single series of conical teeth laterally
in jaws. Dorsal fin with XIII spines and 9 soft rays, none of the spines extended or filamentous; third to
fifth spines longest. Anal fin with III spines and 8 soft rays, second spine is slightly longer than third spine.
Caudal fin forked. Scales large, 45 to 50 in lateral line. Dorsal and anal fins with low scaly sheath. Colour:
body reddish to slightly purplish, with silvery reflections, sometimes pronounced along centres of scales to
form silvery indistinct stripes; dorsal and caudal fin pale yellowish with bright yellow distal margin; pectoral,
pelvic, and anal fins pale.
Size: Maximum standard length so far recorded is 22.6 cm, and it likely gets larger.
Habitat, biology, and fisheries: Found so far mostly over soft bottoms at depths between 210 and 290 m.
Caught by bottom trawl and bottom longline.
Distribution: So far known
only from New Caledonia and
Chesterfield Islands.
3000 Bony Fishes
Dentex tumifrons (Temminck and Schlegel, 1842) (Plate XVI, 119)

Frequent synonyms / misidentifications: Taius tumifrons (Temminck and Schlegel, 1842); Dentex
spariformis (Ogilby, 1910) / None.
FAO names: En - Yellowback seabream.
Diagnostic characters: Body deep and compressed; head large, slightly deeper than long, its profile with
bulge near eye, more prominent in larger fish; eye moderate in size, 2.8 to 4.4 times in head length, close
to front profile. Front of upper jaw with 4 strong canines and lower jaw with 4 to 6 canines; a single
series of conical teeth laterally in jaws. Dorsal fin with XII spines end 10 soft rays, none of the spines
extended or filamentous; third to fifth spines longest. Anal fin with III spines and 8 soft rays, second spine
equal to or slightly longer than third. Caudal fin forked. Scales large, 46 to 50 in lateral line; dorsal and anal
fins with low scaly sheath. Colour: back and upper sides orange-brown, lower sides and belly silvery; 3
golden yellow saddle-like blotches on back, the first at origin of dorsal fin; dorsal, anal, and caudal fins
orange-red.
Size: Maximum total length 40 cm.
Habitat, biology, and fisheries: Lives on mud and muddy-sand bottoms, from depths of 50 to 250 m.
Spawns in late spring. Feeds on a wide range of bottom-living invertebrates and on fish. Caught mainly with
bottom trawls and bottom longlines. Marketed mostly fresh, whole; also dried-salted.
Distribution: Northern part of
South China Sea, including
northern coasts of the Philip-
pines and northward to Japan,
and southern Indonesia to
northern Australia.
Evynnis japonica Tanaka, 1931

FAO names: En - Crimson seabream.
Diagnostic characters: Body robust and deeper than head, strongly compressed, depth of head much
greater than its length; upper profile of head oblique, often with a bulge near eye; eye moderate in size, close
to front profile. Front of upper jaw with 4 canines and lower jaw with 4 to 6 canines; 2 lateral rows of
weak molar teeth in jaws; vomer with conical teeth. Dorsal fin with XII spines and 10 soft rays, the first
2 spines short but well developed, third and fourth (and sometimes fifth) spines elongated as fine filaments,
spines becoming slightly shorter toward tail; soft part of dorsal fin slightly higher than posterior part of spinous
fin. Anal fin with III spines and 9 soft rays, first spine short, second and third about equal in length but
second a little shorter. Caudal fin slightly forked, with pointed lobes. Scales large, 54 to 61 in lateral line;
transverse series of scales between lateral line and median dorsal-fin spines usually 6 ½ (rarely 5 ½);
interorbital area scaly. Colour: pale, silvery pink, but more red on head, upper sides, fins and especially on
filaments of dorsal fin; rows of pale blue spots along scale rows.
Size: Maximum total length 45 cm, commonly 15 to 25 cm.
Habitat, biology, and fisheries: Occurs over a wide range of bottom types but is more common close to
reefs or on rough bottoms at depths greater than 100 m. Marketed mostly fresh, whole.
Distribution: T e m p er a te
West Pacific from Hokkaido to
East China Sea; a single re-
cord from off Bali.
3002 Bony Fishes
Pagrus auratus (Bloch and Schneider, 1801) (Plate XVI, 120)

Frequent synonyms / misidentifications: Sparus major (Temminck and Schlegel, 1843); Chrysophrys
major (Temminck and Schlegel, 1842); C. auratus (Bloch and Schneider, 1801) / None.
FAO names: En - Silver seabream.
(after Grant, 1975)
Diagnostic characters: Body robust, oblong, moder-

ately compressed; upper profile of head convex with a
bulge above eye in large specimens; eye moderate in
size. Lower jaw slightly shorter than upper; 4 enlarged
canines in front of upper jaw and 4 to 6 enlarged
canines in lower jaw; 2 series of moderately large
molar teeth on lateral side of jaws. Dorsal fin single,
with XII strong spines and 10 to 12 soft rays, the
spines not elongated into filaments first spine about
1/2 length of second, which is about 1/2 as long as
third, third to seventh spines longest, other spines small adult
gradually decreasing in length along fin. Anal fin with
(after Randall et al., 1990)
III stout spines and 7 to 9 soft rays, first spine short,
about 1/2 length of second; second and third spines about equal in length. Caudal fin forked, with pointed
lobes. Scales moderately large, lateral line with 53 to 64 scales; interorbital area scaly; absent from bases
of soft dorsal and anal fins. Colour: head and upper body red-brown, sides and belly silvery; numerous
small bright blue spots on upper sides; fins red or faint red; caudal fin with a white lower margin.
Size: Maximum total length 130 cm, commonly 20 to 40 cm.
Habitat, biology, and fisheries: Bottom-living at depths of 10 to 150 m, often on rough grounds, but also
on softer bottoms. Adult fish migrate into shallower parts of their depth range to spawn in late spring and
summer; juvenile fish occur mainly in the shallower areas. Feeds on a wide range of bottom-living
invertebrates, including echinoderms, worms, molluscs and crustaceans; also on fishes. Caught mainly with
bottom trawls and bottom long lines. A popular food fish throughout its range. It is particularly high priced in
Japan where it is much sought for ceremonial banquets and eaten raw. It is also prepared into a wide range
of food products.
Distribution: Northeastern
part of South China Sea (ex-
cluding the Philippines) and
northward to Japan; southern
part of Australia and New Zea-
land.
?
Remarks: In the northern part
of its range it is often referred ?
to as a distinct species Pagrus
major, and it is possible this
will be proven correct.
Rhabdosargus sarba (Forsskål, 1775) (Plate XVI, 121)

Frequent synonyms / misidentifications: Sparus sarba (Forsskål, 1775) / None.
FAO names: En - Goldlined seabream; Fr - Sargue doré; Sp - Sargo dorado.
Diagnostic characters: Body deep (depth about 2 times in standard length), compressed. Head large, its
length about 3 to 3.3 times in standard length; upper profile convex, most strongly arched from snout
to origin of dorsal fin; eye moderate to small in large specimens; mouth almost horizontal, low. At front of
jaws, 4 to 6 upper, and 6 to 8 lower, enlarged, compressed teeth, becoming subconical in large adults,
followed by 4 or 5 rows of molar teeth in posterior part of upper jaw and, 2 to 4 rows in lower jaw, the last
molar in each jaw largest. Gill rakers short, few, 7 to 9 on lower limb of first gill arch. Dorsal fin single, with
XI or XII slender spines and 13 (rarely 12) to 15 soft rays, third and fourth spines longest. Anal fin with III
spines and 11 soft rays, second and third spines subequal; pectoral fins long. Pelvic fins not reaching anus.
Caudal fin forked. Scales cycloid (smooth), more than 50 in lateral line; 5 ½ or 6 ½ scale rows between
lateral line and fourth dorsal-fin spine; preopercle flange naked; dorsal and anal fins with a low scaly
sheath; a long axillary pelvic process. Colour: overall bright silver grey; each scale with a golden centre so
as to form longitudinal lines on body; belly with a bright, yellow band starting at pelvic fin and flaring upward
and backward; dorsal fin hyaline at base, dusky at margin; pectoral and pelvic fins dusky yellowish green;
anal fin with hyaline base, yellow toward margin; lower caudal-fin lobe yellow with tip and lower edge whitish.
Habitat, biology, and fisheries: A bottom-living coastal fish, to a depth of 60 m, sometimes entering
estuaries. In the southern part of the area, spawning takes place near river mouths; after a short planktonic
period, the young fish move into the estuaries, which act as nurseries, and move out into deeper waters with
growth. Feeds on bottom invertebrates, mainly molluscs. Caught mainly with bottom trawls, gill nets,
longlines, handlines, and stake traps. Marketed mostly fresh.
Africa to India, including the
Red Sea and Persian Gulf, the
southern half of Australia, and
from the South China Sea to
Japan.
click for next page

3004 Bony Fishes
LETHRINIDAE
Emperors (emperor snappers)
by K.E. Carpenter
D iagnostic characters: Perch-like sparoid fishes (size to 100 cm). Dorsal fin continuous, with X
spines and 9 or 10 soft rays; anal fin with III spines and 8 to 10 soft rays; pectoral fins with 13
to 15 soft rays; pelvic fins thoracic, with I spine and 5 soft rays; caudal fin emarginate or forked, with 7 to
9 procurrent caudal-fin rays. Scales finely ctenoid and moderate in size; cheek naked in Lethrinus and
scaly in remaining genera. Mouth small to moderate, terminal; lips often soft and fleshy; upper jaw
protrusible, ascending process of premaxilla confluent with articular process, usually longer than or almost
equal to alveolar ramus; postmaxillary process absent; palato-premaxillary ligament very well developed;
maxilla mostly concealed below infraorbital bones, not articulating broadly with distal tip of
premaxilla, without a supplementary bone; ethmo-maxillary ligament absent; an outer row of canine teeth
in front of both jaws, teeth on sides conical or molar-like; an inner row of villiform teeth anteriorly; vomer
and palatines toothless. Gill membranes broadly united to one another but separated from isthmus; gills
4, slit behind the fourth present; pseudobranchs present; gill rakers short and knob-like; 4 branchiostegal
rays inserting on ceratohyal, the fifth at interspace between ceratohyal and epihyal, the sixth on epihyal;
second epibranchial tooth plate present, the third absent. Two openings in pars jugularis; subocular shelf
reduced or absent. Three predorsal bones present in the following configuration: first predorsal, first neural
spine, second and third predorsal, second neural spine, first pterygiophore supporting the first 2 dorsal-fin
spines and second pterygiophore supporting third dorsal-fin spine, third neural spine, third pterygiophore
supporting fourth dorsal-fin spine, fourth neural spine; 2 or 3 trisegmental pterygiophores in dorsal and
anal fins; 11 epipleural ribs; accessory subpelvic keel absent; postpelvic process well developed. Pyloric
caeca few, usually 3. Colour: body and head colour silvery, grey, olive, greenish, bluish, tan, brown, or
reddish, often with dark blotches or bars; head sometimes with bluish streaks and spots; sometimes bright
red markings on or near base of pectoral fins, on pelvic-, anal-, and dorsal-fin membranes near base of
fins, on opercular and preopercular margins, and on head, these markings often variable within species;
fins clear, pale, bluish, yellowish, or reddish, often blotched, the edge of fins often reddish.
9 soft rays
cheek scaleless
8 soft rays Lethrinus
10 spines
10 soft rays
rows of scales
on cheek 10 soft rays
III spines Gymnocranius
Perciformes: Percoidei: Lethrinidae 3005
Habitat, biology, and fisheries: Coastal fishes of the tropical Indo-West Pacific and West Africa, often
found over sandy bottoms and including seagrass beds, mangrove swamps, coral reefs, and rocky reefs,
to depths of 100 m. They usually occur solitary or in small groups and form large aggregations while
spawning. The diet consists of echinoderms, crustaceans, molluscs, fishes, and polychaetes. Moderate to
significantly important in fisheries; in certain countries, members of this genus are the most important catch
by weight. Caught primarily by handline, traps, and trawls. For 1995, the FAO Yearbook of Fishery Statistics
reports a total catch of 31 518 t of Lethrini dae from the Western Central Pacific.

Lutjanidae: teeth usually present on vomer and palatines; cheek always scaly (naked in Lethrinus but scaly
in other genera of Lethrinidae); preopercular edge usually serrate (typically smooth in Lethrinidae); usually
11 or more soft dorsal-fin rays (9 or 10 in Lethrinidae).
Nemipteridae: anal fin with 7 soft rays (except 8 in Nemipterus virgatus); cheek always scaly (naked in
Lethrinus but scaly in other genera of Lethrinidae); no molar-like teeth in jaws (present in some species of
Lethrinidae).
palatine jaw
tooth teeth
bands
vomerine
tooth
patch
roof of mouth 3 rows

preopercular edge of scales
usually serrate examples of dentition
on roof of mouth
Lutjanidae (Lutjanus) Nemipterus
Sparidae: posterior tip of premaxilla overlapping maxilla at hind end of mouth (maxilla overlapping premaxilla
in Lethrinidae); more than X dorsal-fin spines (always X in Lethrinidae); cheek always scaly (naked in
Lethrinus but scaly in the other genera of Lethrinidae).
Haemulidae: scales always present between eye and mouth (absent in that area in Lethrinidae); 2 or more
pores present on chin; second anal-fin spine often very strong.
pores on chin
maxilla
maxilla
premaxilla premaxilla
scales ventral view of head
Pomadasys Plectorhinchus
Sparidae Lethrinidae Haemulidae
3006 Bony Fishes
Key to the genera of Lethrinidae

1a. Cheek naked (Fig. 1); 9 soft rays in dorsal fin; 8 soft rays in anal fin (Fig. 2) . . . . . . . . Lethrinus
1b. Cheek with 3 to 6 vertical rows of scales (Fig. 3); 10 soft rays in dorsal fin; usually 9 or
10 soft rays in anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
9 soft rays
8 soft rays
no scales here
Fig. 1 Lethrinus Fig. 2 Lethrinus Fig. 3 Wattsia
2a. Inner surface of pectoral-fin base densely scaled (Fig. 4a); sides of jaws with round, flat
molars (Figs 5a and 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Monotaxis grandoculis
2b. Inner surface of pectoral-fin base scaleless (Fig. 4b); sides of jaws with canines and
villiform teeth (except 1 species, Gymnocranius euanus, which has molars) (Fig. 5b) . . . . . . . ® 3
pectoral fin pectoral fin

folded folded canine-like teeth
forward forward
conical
teeth
villiform
teeth villiform
teeth
scales on no scales on molar-like teeth

inner base inner base
a) b) a) Monotaxis b) Gnathodentex
Fig. 4 Fig. 5 roof of mouth
3a. Outer surface of maxilla smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocranius

3b. Outer surface of maxilla with denticulated ridge (Fig. 7) . . . . . . . . . . . . . . . . . . . . . . ® 4
denticulated
ridge
(lateral teeth not shown)

Fig. 6 Monotaxis grandoculis Fig. 7
4a. Caudal-fin lobes pointed; anal fin with 8 or 9 soft rays; pectoral fins with 15 rays (Fig. 8)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gnathodentex aurolineatus
4b. Caudal-fin lobes rounded; anal fin with 10 soft rays; pectoral fins with 14 rays (Fig. 9) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Wattsia mossambica
Fig. 8 Gnathodentex aurolineatus Fig. 9 Wattsia mossambica
Key to the species of Gymnocranius occurring in the area

1a. Caudal fin strongly forked (Fig. 10a), the median rays shorter than eye diameter; lower edge
of eye intersected by line from tip of snout to middle of caudal-fin fork (Figs 11 and 12) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Gymnocranius elongatus
1b. Caudal fin moderately forked (Fig. 10b), the median rays about equal to, or longer than,
eye diameter; lower edge of eye above line from tip of snout to middle of caudal-fin fork
(Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
a) b)
Fig. 10 Fig. 11
Fig. 12 Gymnocranius elongatus Fig. 13

3008 Bony Fishes
2a. Longitudinal scale rows between lateral line and base of middle dorsal-fin spines 4 ½;
molars present on sides of jaws (Figs 14a and 15) . . . . . . . . . . . . . . . Gymnocranius euanus
2b. Longitudinal scale rows between lateral line and base of middle dorsal-fin spines 5 ½
(Fig. 16); molars absent, usually villiform to conical teeth present on sides of jaws (Fig. 14b) . . . . . ® 3
a) b)
Fig. 14 Fig. 15 Gymnocranius euanus
3a. Head with a prominent diamond-shaped, blackish patch of scales surrounded by a white
border above and behind each eye (Fig. 17) . . . . . . . . . . . . . . . . . . Gymnocranius audleyi
3b. Head without the marking described in 3a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
5½ scale rows
patch
Fig. 16 Gymnocranius Fig. 17 Gymnocranius audleyi

4a. Snout with prominent blue-edged yellow
band from front of eye to above upper lip
(encompassing nostrils); 3 or 4 oblique,
blue (brown in preservative) lines across
cheek; body moderately deep, the maxi-
mum depth about 2.3 to 2.4 times in
standard length (Fig. 18) . . . Gymnocranius frenatus
4b. Snout without the marking described in
4a; cheek may have wavy, blue lines, but
they are more or less horizontal rather
than oblique; body depth variable, 1.9 to Fig. 18 Gymnocranius frenatus
3 times in standard length . . . . . . . . . . . . . ® 5
5a. Inner edge of caudal-fin fork slightly convex with fin tips blunt (Fig. 19a); live fish with
longitudinal rows of dark spots on upper side (Fig. 20) . . . . . . . . . . . . . . . Gymnocranius sp.
5b. Inner edge of caudal-fin fork straight and fin tips pointed (Fig. 19b); no longitudinal rows
of dark spots on upper side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
a) b)
Fig. 19 Fig. 20 Gymnocranius sp.
6a. Body comparatively deep, its depth about

1.9 to 2.2 (occasionally 2.3) times in stand-
ard length; no blue spots or wavy blue lines
on cheek; usually with several irregular dark
bars on sides and dark subocular bar
(Fig. 21) . . . . . . . . . . . . . . Gymnocranius griseus
6b. Body comparatively slender, its depth about
2.3 to 3 times in standard length; side of
snout and cheek often with vertically elon-
gate spots (Fig. 22) or wavy longitudinal blue
lines (Fig. 23) in fresh specimens longer Fig. 21 Gymnocranius griseus
than 20 to 25 cm standard length; smaller
specimens may have irregular dark bars on
side and dark subocular bar as in 6a . . . . . . . . . ® 7
7a. Body depth about 2.6 to 3 times in standard length; cheek often with vertically elongate
blue spots in fresh specimens (Fig. 22) . . . . . . . . . . . . . . . . . . . . Gymnocranius microdon
7b. Body depth about 2.3 to 2.5 (occasionally 2.6) times in standard length; juveniles below
about 200 mm standard length frequently with dark bar under eye and 5 or 6 irregular
dark bars across side of body; adults with wavy, blue lines on snout and cheek increasing
in number with growth (Fig. 23) . . . . . . . . . . . . . . . . . . . . . . Gymnocranius grandoculis
Fig. 22 Gymnocranius microdon Fig. 23 Gymnocranius grandoculis
Key to the species of Lethrinus occurring in the area

Remarks on key characters: 2 of the most useful characters in differentiating species of Lethrinus are the
presence or absence of scales in the axil of the pectoral fins and, the number of scale rows above the lateral
line to the middle spines of the dorsal fin. These characters are relatively constant for most species of
Lethrinus but in a few species they vary. Because of this, some previous keys have tended to de-emphasize
the use of these characters. After examining the variation of these characters in many specimens, however,
I found that the advantages of using them far outweighs the disadvantages of their variation. The strategy
to tolerate this variation has been to have species occur more than once in the key and to add redundant
diagnostic characters. Despite this strategy, the identification of preserved specimens of species of
Lethrinus, especially juveniles, can still be a formidable task.
1a. Second dorsal-fin spine distinctly longer than other dorsal-fin spines (Fig. 24); a pair of
canines in front of lower jaw substantially curved outward (Fig. 25a) . . . . . . Lethrinus genivittatus
1b. Third, fourth, or fifth dorsal-fin spine the longest; canines in front of lower jaw almost
straight or curved outward slightly (Fig. 25b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
canines
curved
a) Lethrinus b) other
genivittatus Lethrinus species
Fig. 24 Lethrinus genivittatus Fig. 25 anterior view of canines in lower jaw

(teeth in upper jaw not shown)
3010 Bony Fishes
2a. Inner surface of pectoral-fin base without scales (Fig. 26a ) . . . . . . . . . . . . . . . . . . . . ® 3

2b. Inner surface of pectoral-fin base covered mostly or partially with scales (Fig. 26b),
sometimes with only a fraction of lower part of inner base covered . . . . . . . . . . . . . . . . ® 17
pectoral fin pectoral fin scale rows above
folded forward folded forward lateral line
no scales on scales on
inner base inner base
a) b) scale rows below lateral line
Fig. 26 inner surface of pectoral fin Fig. 27 scale counts above and below lateral line
(do not include lateral-line scale or scale on ventral median)
villiform teeth
3a. Longitudinal scale rounded teeth
rows between lateral
line and base of mid- canines canines
dle dorsal-fin spines
4 ½ (Fig. 27); lateral
teeth in jaws conical posterolateral teeth conical b)
(Fig. 28a) . . . . . . . . . . . ® 4 a)
3b. Longitudinal scale tubercles
rows between lateral on molars
distinct molars
line and base of mid-
dle dorsal-fin spines
5 ½ (Fig. 29); lateral canines
teeth in jaws conical,
rounded, molars, or
molars with a tuber- conical teeth
cle (Fig. 28a-d) . . . . . . . . ® 15 c) d)
Fig. 28 types of teeth
4a. Body comparatively deep, its 4.5 scale
depth 2.2 to 2.75 times in stand- rows
ard length; head length usually
distinctly less than body depth,
sometimes head length almost
equal to body depth . . . . . . . . . . . . ® 5
4b. Body comparatively slender, its
depth 2.8 to 3.9 times in standard
length; head length almost always
distinctly greater than body depth,
head length rarely almost equal to Fig. 29 scale count above lateral line
body depth . . . . . . . . . . . . . . . . . ® 6
snout length
without lip
5a. Third dorsal-fin spine usually the longest; snout
length (measured without lip) 0.9 to 1 times in cheek
cheek height (Fig. 30); in life, red markings height
around eye (Fig. 31) . . . . . . . . . . . . Lethrinus miniatus
5b. Fourth dorsal-fin spine usually the longest; snout
length (measured without lip) usually 0.8 to 0.9
times in cheek height (Fig. 30); in life, blue mark-
ings around eye (Fig. 32) . . . . . . Lethrinus haematopterus preorbital
(reported thus far only from Hong Kong and Taiwan Province of China, length
although it may eventually be recorded from the area)
Fig. 30 measurement of cheek height
(lowermost point on orbit to furthest
point on angle of preopercle) and
snout (without lip)
red markings
blue markings
Fig. 31 Lethrinus miniatus Fig. 32 Lethrinus haematopterus
6a. Prominent scaleless patch above base of pectoral fins, bright red in life (Fig. 33)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lethrinus conchyliatus
6b. No prominent scaleless patch above base of pectoral fins . . . . . . . . . . . . . . . . . . . . .® 7
7a. Interorbital area distinctly or slightly concave (Fig. 34a) . . . . . . . . . . . . . . . . . . . . . . ® 8

7b. Interorbital area nearly flat or convex (Fig. 34b, c) . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
scaleless patch concave flat convex

(red in life)
lips red
a) b) c)
Fig. 33 Lethrinus conchyliatus Fig. 34 anterior view of head showing shape of

interorbital region
melanophores areas without

8a. Membrane between innermost rays of melanophores
pelvic fins covered with melanophores
(Fig. 35a); 5 to 8 scales in supratemporal
patch (Fig. 36); in life, lips yellowish and a
red spot on upper base of pectoral fins
(Fig. 36) . . . . . . . . . . . . Lethrinus xanthochilus
8b. Membrane between innermost rays of
pelvic fins not mostly covered with
melanophores (Fig. 35b); 7 to 10 scales in
supratemporal patch; in life, lips, upper
base of pectoral fins, upper edge of oper- a) b)
cle, sometimes posterior edge of preoper-
cle, and sometimes an indistinct band on Fig. 35 pelvic fin
snout, reddish (Fig. 37) . . . . . Lethrinus reticulatus
supratemporal
patch
lips yellowish reddish

red patch
Fig. 36 Lethrinus xanthochilus Fig. 37 Lethrinus reticulatus

3012 Bony Fishes
9a. Scale rows in transverse series below lateral line 13 or 14 (Fig. 38); posterior nostril a
vertical slit, often closer to anterior nostril than to orbit; body very slender, its depth 3.2
to 3.9 times in standard length (Fig. 38) . . . . . . . . . . . . . . . . . . . . . Lethrinus variegatus
9b. Scale rows in transverse series below lateral line 15 to 17; posterior nostril a longitudinal
slit closer to orbit than to anterior nostril, or about half-way between anterior nostril and
orbit; body depth 2.8 to 3.4 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . ®10
10a. Snout long, its length (measured without lip) 0.7 to 0.8 times in cheek height (Fig. 30); 3
dark streaks radiating forward from eye on snout usually visible (Fig. 39); inner surface
of pectoral-fin surface never red in life . . . . . . . . . . . . . . . . . . . . . . . Lethrinus microdon
10b. Snout length (measured without lip) 0.8 to 0.9 times in cheek height (Fig. 30); 3 distinct
dark streaks radiating from eye not usually apparent, but if they are apparent, inner
surface of pectoral-fin base is red in life . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
Fig. 38 Lethrinus variegatus Fig. 39 Lethrinus microdon
11a. Area on snout directly in front of eye with a distinct hump, snout profile concave (Fig. 40)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Lethrinus sp. 1
11b. Area on snout in front of eye without a distinct hump, profile of snout nearly straight or
concave . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
12a. Membranes between inner rays of pelvic fins mostly covered with melanophores, or with a
limited part of innermost ray not covered by melanophores (Figs 35a and 41) . . Lethrinus amboinensis
12b. Membranes between pelvic-fin rays closest to body without a dense covering of melano-
phores (Fig. 35b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Fig. 40 Lethrinus sp. 1 Fig. 41 Lethrinus amboinensis

13a. Wide scaleless area on upper posterior margin of opercle (Fig. 42a); 7 to 10 scales in
supratemporal patch (Fig. 43) . . . . . . . . . . . . . . . . . . . . . . . Lethrinus rubrioperculatus
13b. Upper posterior margin of opercle without wide scaleless area (Fig 42b); 4 to 8 scales in
supratemporal patch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
wide
scaleless
area
operculum operculum pectoral fin

pectoral fin
a) b)
Fig. 42 posterior margin of opercle Fig. 43 Lethrinus rubrioperculatus
14a. A large irregular black blotch on sides, bordering under lateral line and below soft-rayed
portion of dorsal fin (Fig. 44) . . . . . . . . . . . . . . . . . . . . . . . . . . . Lethrinus semicinctus
14b. No large black blotch on side (Fig. 45) . . . . . . . . . . . . . . . . . . . . . . . . . Lethrinus sp. 2
black blotch
Fig. 44 Lethrinus semicinctus Fig. 45 Lethrinus sp. 2
15a. Snout long, its length (measured without

lip) 0.6 to 0.8 times in cheek height
(Fig. 30); body relatively slender, its depth
2.8 to 3.4 times in standard length (for
specimens over 10 cm standard length)
(Fig. 46) . . . . . . . . . . . . . . Lethrinus olivaceus
dark
15b. Snout length (measured without lip) 0.8 to 1 streaks
times in cheek height (Fig. 30); body depth
Fig. 46 Lethrinus olivaceus
2.3 to 2.9 times in standard length . . . . . . . . ® 16
16a. In adults, lateral teeth in jaws with molars, molars with tubercles, or broadly rounded
(Fig. 28 b-d); body depth 2.5 to 2.9 times in standard length; posterior margin of opercle
red and base of pectoral fins often red in life (Fig. 47) . . . . . . . . . . . . . . . . Lethrinus lentjan
(widespread throughout the area)
16b. In adults, lateral teeth in jaws conical (Fig. 28a); body depth 2.25 to 2.6 times in standard
length; without red markings in life (Fig. 48) . . . . . . . . . . . . . . . . . . . . . . . Lethrinus sp. 3
(restricted to northwestern Australia)
red
Fig. 47 Lethrinus lentjan Fig. 48 Lethrinus sp. 3

17a. A large black blotch bordering below lateral line and centred on posterior tip of pectoral
fins, usually persistent in preserved specimens (Fig. 49); usually 13 or 14 scale rows in
lower series around caudal peduncle (Fig. 50); (longitudinal scale rows between lateral
line and base of fifth dorsal-fin spine usually 5 ½, sometimes 4 ½ scales) (Fig. 50) . Lethrinus harak
17b. No dark blotch below lateral line; usually 15 scale rows in lower series around caudal
peduncle (except in L. atkinsoni which also has 13 or 14 of these scale rows but can be
distinguished from L. harak in always having 4 ½ longitudinal scale rows between lateral
line and base of fifth dorsal-fin spine, whereas L. harak usually has 5 ½, but sometimes
4 ½, longitudinal scale rows between lateral line and base of fifth dorsal-fin spine) . . . . . . . ® 18
lateral-line
black blotch scales
lower
peduncular
scales
Fig. 49 Lethrinus harak Fig. 50 scale counts on caudal peduncle

(lower circumpeduncular scale count includes
lateral-line scales on both sides)
3014 Bony Fishes
18a. Longitudinal scale rows between lateral line and base of fifth dorsal-fin spine 4 ½
(Fig. 27) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
18b. Longitudinal scale rows between lateral line and base of fifth dorsal-fin spine 5 ½
(Fig. 29) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
19a. The third, fourth, or fifth anal-fin ray usually the long-
est, much longer than length of base of soft portion of
anal fin; length of longest anal-fin ray 0.8 to 1.1 times
in length of entire anal fin (Fig. 51) . . . . . . . . . . . . . . ® 20
soft
19b. The first or second anal-fin ray usually the longest, anal-fin
shorter than, approximately equal to, or slightly base
longer than length of soft portion of anal fin; length of
longest anal-fin ray 1.1 to 1.8 times in length of base
of entire anal fin (Fig. 51) . . . . . . . . . . . . . . . . . . . ® 21
anal-fin base
20a. Lateral teeth in jaws conical or rounded (Fig. 28a, b); Fig. 51 length of soft anal-fin base
lateral-line scales 46 to 48; membranes between in- and entire anal-fin base
ner rays of pelvic fins usually with densely distributed
melanophores; small orange spots on head often
visible in life (Fig. 52) . . . . . . . . . . Lethrinus erythracanthus
20b. Lateral teeth in jaws usually with distinct molars (Fig. 28c); lateral-line scales 44 to
46; membranes between inner rays of pelvic fins usually not densely populated with
melanophores; 2 light bars on caudal peduncle often visible (Fig. 53) . . . . Lethrinus erythropterus
44-46 lateral-line scales
46-48 lateral-line
scales
often 2 light
often orange bars
spots
Fig. 52 Lethrinus erythracanthus Fig. 53 Lethrinus erythropterus
21a. Angle of anterior snout relative to line of

upper jaw 52 to 62° (Fig. 54); lateral teeth
in jaws conical (Fig. 28a); in life, a bright
red spot on outer base of pectoral fins
(Fig. 55) . . . . . . . . . . . . . . . Lethrinus miniatus
21b. Angle of anterior snout relative to line of
upper jaw 65 to 80° (Fig. 54); lateral teeth
in jaws rounded or molars (Fig. 28b,c); no
red spot on outer base of pectoral fins
(Fig. 56) . . . . . . . . . . . . . . . Lethrinus atkinsoni
Fig. 54 angle of snout relative to upper jaw
red
Fig. 55 Lethrinus miniatus Fig. 56 Lethrinus atkinsoni
blue spots
22a. Melanophores covering most of pelvic-fin and lines
membranes, including near inner rays
(Fig. 35a); in life, blue spots and/or streaks
radiating forward from eye (Fig. 57) . Lethrinus nebulosus
22b. Melanophores absent from large portions of
membranes between inner rays of pelvic fins
(Fig. 35b) . . . . . . . . . . . . . . . . . . . . . . . ® 23
Fig. 57 Lethrinus nebulosus
23a. Maxilla with a knob (Figs 58a and 59a); angle
of anterior snout relative to line of upper jaw
50 to 60° (Fig. 54) (Fig. 59b) . . . . . Lethrinus obsoletus
23b. Maxilla smooth or with a ridge (Fig. 57b, c);
angle of anterior snout relative to line of the
upper jaw 59 to 73° (Fig. 54) . . . . . . . . . . . . . ® 24
smooth ridge
24a. Snout relatively short and blunt, the preorbital
knob
1.3 to 1.4 times in cheek height (Fig. 30);
angle of anterior snout relative to upper jaw
line 64° to 73° (Fig. 54); posterior edge of
preopercle and opercle red in life; profile of a) b) c)
head in front of eye usually convex (Fig. 60) Fig. 58 outer surface of left maxilla
. . . . . . . . . . . . . . . . . . . . . . Lethrinus ornatus
24b. Snout relatively pointed,
the preorbital 1 to 1.3 times
in cheek height (Fig. 30);
angle of anterior snout rela-
tive to upper jaw line 59° to
70° (Fig. 54); posterior knob
edge of preopercle not red,
but edge of opercle and
base of pectoral fins some- yellow
times red in L. lentjan; pro- stripe
a) b)
file of head in front of eye
approximately straight . . . . . . . ® 25 Fig. 59 Lethrinus obsoletus
25a. In adults, lateral teeth in jaws broadly rounded and sometimes with a pointed cusp,
molars, or molars with a tubercle (Fig. 28b-d); body depth 2.5 to 2.9 times in standard
length; 15 or 16 scale rows between origin of anal fin and lateral line; in life, posterior
margin of opercle and sometimes outer base of pectoral fins red (Fig. 61) . . . . . Lethrinus lentjan
(widely distributed in the area)
25b. Lateral teeth in jaws conical; body depth 2.3 to 2.6 times in standard length; 16 or 17
scale rows between origin of anal fin and lateral line; no red markings on body . . . . . . . . . ® 26
(restricted to Australia, southeastern Indonesia, Papua New Guinea, the Solomons, and New Caledonia)
usually yellow
red stripes red
Fig. 60 Lethrinus ornatus Fig. 61 Lethrinus lentjan
3016 Bony Fishes
26a. Centres of scales on upper sides anteriorly with distinct black marking; inner base of
pectoral fins usually more than half covered with scales (Figs 62a and 63) . . . Lethrinus laticaudis
(northern Australia, southeastern Indonesia, Papua New Guinea, the Solomons, and New Caledonia)
26b. Centres of scales on upper sides without distinct black marking, often pearly; inner base
of pectoral fins usually less than half covered with scales (Figs 62b and 48) . . . . . Lethrinus sp. 3
(restricted to northwestern Australia)
more than half less than half
covered with scales blue stripes
covered with scales
pectoral fin pectoral fin
folded forward folded forward
a) b)
Fig. 62 inner surface of pectoral fin Fig. 63 Lethrinus laticaudis
0
The symbol is given when species accounts are included.
0 Gnathodentex aurolineatus (Lacepède, 1802)
0
0 Gymnocranius audleyi Ogilby, 1916
0
0
Gymnocranius elongatus Senta, 1973
Gymnocranius euanus Günther, 1879
0
0
Gymnocranius frenatus Bleeker, 1873
Gymnocranius grandoculis (Valenciennes, 1830)
0 Gymnocranius griseus (Schlegel, 1844)
0 Gymnocranius sp.
Gymnocranius microdon (Bleeker, 1851)
0
0 Lethrinus amboinensis Bleeker, 1854
0
0
Lethrinus atkinsoni Seale, 1910
Lethrinus conchyliatus Smith, 1959
0 Lethrinus erythracanthus Valenciennes, 1830
0 Lethrinus erythropterus Valenciennes, 1830
Lethrinus genivittatus Valenciennes, 1830
1/
0
? Lethrinus haematopterus Temminck and Schlegel, 1844
0 Lethrinus harak (Forsskål, 1775)
0
0
Lethrinus laticaudis Alleyne and Macleay, 1877
Lethrinus lentjan (Lacepède, 1802)
0
0
Lethrinus microdon Valenciennes, 1830
Lethrinus miniatus (Schneider, 1801)
0
0
Lethrinus nebulosus (Forsskål, 1775)
Lethrinus obsoletus (Forsskål, 1775)
0
0
Lethrinus olivaceus Valenciennes, 1830
Lethrinus ornatus Valenciennes, 1830
0
0
Lethrinus reticulatus Valenciennes, 1830
Lethrinus rubrioperculatus Sato, 1978
0
0
Lethrinus semicinctus Valenciennes, 1830
Lethrinus sp. 1
0
0
Lethrinus sp. 2
Lethrinus sp. 3
0 Lethrinus variegatus Valenciennes, 1830
Lethrinus xanthochilus Klunzinger, 1870
0 Monotaxis grandoculis (Forsskål, 1775)
0 Wattsia mossambica (Smith, 1957)
Reference
Carpenter, K.C. and G.R. Allen. 1989. Emperor fishes and large-eye breams of the world (family Lethrinidae). An
annotated and illustrated catalogue of lethrinid species known to date. FAO Fish. Synop., (125)Vol. 9:118 p.
1/ Not yet reported from the area; reported from Hong Kong and Taiwan Province of China.
click for next page

Gnathodentex aurolineatus (Lacepède, 1802) (Plate XVII, 122)

FAO names: En - Striped large-eye bream; Fr - Empereur strié; Sp - Emperador estriado.
Diagnostic characters: Body oblong, its depth 2.3 to 2.8 times in standard length. Dorsal profile of head
moderately sloped; eye relatively large, its diameter about equal to length of snout. Jaw teeth in a narrow
villiform band bordered by an outer series of conical teeth; 4 moderate-sized canines at front of upper
jaw and 6 at front of lower jaw; maxilla (upper jaw) with a longitudinal denticulated ridge; a pair of
close-set, round nasal openings on each side of snout in front of eyes. Dorsal fin with X slender spines
and 10 soft rays, the first to fourth spines increasing in length, the remaining spines about equal; anal fin
with III slender spines and 8 or 9 (usually 9) soft rays; pectoral-fin rays 15 (including small splint-like
uppermost element); caudal fin strongly forked with pointed tips. Lateral-line scales about 68 to 74;
cheek with 4 to 6 rows of scales; 5 scale rows between lateral line and base of middle dorsal-fin spines;
inner surface of pectoral-fin axil scaleless. Colour: upper one-third of side (i.e. back region) dark brown
with narrow silvery stripes, corresponding with each horizontal scale row; remainder of head and body
mainly silver to grey except 4 or 5 brownish orange stripes on lower two-thirds of side, uppermost stripe
is the widest and originates at upper corner of gill flap (operculum); a prominent golden yellow to orange
blotch directly below posterior part of dorsal fin and just in front of caudal-fin base; snout and lips often
with a yellowish tinge; iris golden; a broad pearly stripe on preorbital region, continuing below eye to its
posterior margin; fins mainly clear, although dorsal, anal, and caudal fins sometimes light reddish;
yellow-orange spot at upper base of pectoral fins.
Habitat, biology, and fisheries: Generally found on coral reefs, sometimes in aggregations containing
100 or more individuals. The depth range is between about 3 and 20 m. Feeds chiefly on bottom-living
invertebrates. Frequently found in markets. Caught mainly with spears, traps, gill nets, and handlines.
Average quality flesh that is marketed mostly fresh.
the Indo-West and Central
Pacific (excluding Hawaii)
from the Tuamotu Islands to
the east coast of Africa, and
from Australia northwards to
Japan.
3018 Bony Fishes
Gymnocranius audleyi Ogilby, 1916 (Plate XVII, 123)

Frequent synonyms / misidentifications: Gymnocranius bitorquatus Cockerell, 1916 / None.
FAO names: En - Collared large-eye bream.
Diagnostic characters: Body oblong, its depth 2.2 to 2.4 times in standard length. Snout profile relatively
steep, a bony mid-dorsal ridge on nape of adults; lower edge of eye well above a line from snout tip to
middle of caudal-fin fork; eye relatively large, its diameter usually less than snout length; interorbital space
flattened to slightly convex, its width less than eye diameter; suborbital space about equal to eye diameter
or slightly less; jaws extending to a vertical through nostrils; outer surface of maxilla smooth; a pair of
small, slender canines at front of each jaw; lateral teeth villiform (small and slender) with a few larger outer
conical teeth. Dorsal fin with X slender spines and 10 soft rays, first 3 or 4 spines gradually increasing in
length, the remainder about equal, the middle soft rays of dorsal and anal fins slightly elevated; anal fin
with III slender spines and 9 or 10 soft rays; pectoral-fin rays 10; caudal fin only moderately forked with
pointed tips, the median rays longer than eye diameter. Lateral-line scales 48 plus 2 tubed scales extending
onto base of caudal fin; cheek with 4 or 5 transverse scale rows; 4 to 6 rows of scales on cheeks;
5 ½ scale rows between lateral line and base of middle dorsal-fin spines; inner surface of pectoral
fins scaleless. Colour: overall silver or whitish, often light brown or olive on upper sides; each scale on
sides with a brown spot forming longitudinal rows, those on back darkest; top of head dull brownish or
grey; cheek and opercle silvery, sometimes with a golden sheen; parietal scale patch (on side of nape,
above and behind eye) blackish surrounded by a brilliant white margin; fins uniformly clear to slightly
yellowish. Some individuals, especially those freshly dead, have scattered brown flecks on sides. Also a
blackish bar running through the eye may be evident.
Habitat, biology, and fisheries: Inhabits sand or rubble areas adjacent to reefs, usually at depths between
about 8 to 40 m. It feeds mainly on bottom-living invertebrates. Occasionally seen in markets, although not
a high-priority commercial fish. It sometimes has a distasteful iodine flavour that can actually be smelled
on freshly hooked fishes. However, “untainted” fishes are good eating.
Distribution: East coast of
southern Queensland, Aus-
tralia. Also occurs on the
southern half of the Great
Barrier Reef.
Gymnocranius elongatus Senta, 1973

FAO names: En - Forktail large-eye bream.
Diagnostic characters: Body oblong, its depth 2.2 to 2.4 times in standard length. Head profile evenly
rounded, snout somewhat pointed; lower edge of eye intersected by line from snout tip to middle of
caudal-fin fork; eye large, its diameter usually about equal to length of snout; interorbital space convex,
about equal to eye diameter; preorbital width less than eye diameter; mouth relatively small, posteriormost
part of jaws not reaching a vertical through front of eye; outer surface of maxilla smooth; 2 or 3 pair of
small, slender canines at front of each jaw, lateral teeth villiform to conical; a pair of close-set, round nasal
openings on each side of snout in front of eyes. Dorsal fin with X slender spines and 10 soft rays; anal fin
with III slender spines and 10 soft rays; pectoral-fin rays 14; caudal fin deeply forked with pointed tips,
the median rays shorter than eye diameter. Lateral-line scales 46 to 48 plus 2 to 4 tubed scales extending
onto base of caudal fin; cheek with 4 to 6 transverse scale rows; 5 ½ scale rows between lateral line
and base of middle dorsal-fin spines; inner surface of pectoral-fin axil scaleless. Colour: overall silvery,
sometimes slightly brownish dorsally; about 8 transverse brown bars on sides, the first crossing through
eye, the remainder below dorsal fin and across caudal peduncle; scattered blotches and speckling
sometimes evident on sides; fins clear to yellow-orange; caudal-fin margins and tips often deep red. There
is considerable variation in the intensity of the dark bars on the sides of the body.
Habitat, biology, and fisheries: Inhabits trawling grounds of the continental shelf, usually at depths
between about 50 and 100 m. Feeds mainly on bottom-living invertebrates. Occasionally seen in markets,
mainly fresh. Caught mostly with bottom trawls.
Distribution: Coastal and
shelf waters of the West Pa-
cific and eastern Indian
oceans from Japan to north-
ern Australia. It has also been
reported from the western In-
dian Ocean, but this needs to
be confirmed.
3020 Bony Fishes
Gymnocranius euanus Günther, 1879

Frequent synonyms / misidentifications: Gymnocranius japonicus Akazaki, 1961 / None.
FAO names: En - Japanese large-eye bream.
sloping steeply; lower edge of eye well above a line from tip of snout to middle of caudal-fin fork; eye
relatively large, although its diameter less than interorbital space and width of preorbital width (at least in
adults); mouth relatively small, posterior part of jaws reaching to about level of nostril openings; outer
surface of maxilla smooth; 2 or 3 pairs of small, slender canines at front of each jaw, other teeth villiform,
but well developed molars on lateral sections of jaws. Dorsal fin with X slender spines and 10 soft rays;
anal fin with III slender spines and 10 soft rays; pectoral-fin rays 14; caudal fin moderately forked, inner
edge of fork slightly convex, with blunt tips, the median rays slightly longer than eye diameter.
Lateral-line scales 48 or 49 plus 3 or 4 tubed scales extending onto base of caudal fin; cheek with 4 or 5
transverse scale rows; 4 ½ scale rows between lateral line and base of middle dorsal-fin spines;
inner surface of pectoral-fin axil scaleless. Colour: overall silvery white to greyish; scales of back and
sides with a small brownish basal spot; scattered prominent black blotches mainly on anterior half of body;
a very faint bar sometimes present below eye to lower edge of cheek; fins mainly clear to reddish, caudal
fin sometimes dusky reddish brown; faint spots may be evident on dorsal, caudal, and anal fins.
Habitat biology, and fisheries: Inhabits sand and rubble bottoms adjacent to rock and coral reefs, usually
at depths between 15 and 50 m. It feeds mainly on small bottom-living gastropod molluscs. Occasionally
seen in markets, usually fresh. Caught mainly with bottom longlines and handlines, sometimes speared.
Ocean including southern Ja-
pan, South China Sea, west-
ern Australia to the Great
Barrier Reef of Australia,
Coral Sea, New Caledonia,
and Tonga.
Gymnocranius frenatus Bleeker, 1873

FAO names: En - Yellowsnout large-eye bream.
Diagnostic characters: Body oblong, its depth 2.3 to 2.4 times in standard length. Dorsal profile of
head sloping steeply, the upper forehead profile somewhat angular, lower edge of eye well above a line
from tip of snout to middle of caudal-fin fork in adults; eye relatively large, its diameter slightly greater than
interorbital space and preorbital width (at least in adults); mouth relatively small, posterior part of jaws
reaching to about level of nostril openings; outer surface of maxilla smooth; 2 or 3 pair of small, slender
canines at front of upper jaw and an enlarged outward-projecting tusk on each side at front of lower jaw;
other teeth villiform, but well developed molars on lateral sections of jaws. Dorsal fin with X slender spines
and 10 soft rays; anal fin with III slender spines and 10 soft rays; pectoral-fin rays 14; caudal fin
moderately forked, inner edge of fork straight, with pointed tips, the median rays longer than eye
diameter. Lateral-line scales 48 or 49 plus 3 or 4 tubed scales extending onto base of caudal fin; cheek
with 4 or 5 transverse scale rows; 5 ½ scale rows between lateral line and base of middle dorsal-fin
spines; inner surface of pectoral-fin axil scaleless. Colour: overall silvery white, greyish on upper back;
about 5 to 7 narrow, irregular dark bars (may be very faint) on sides; a prominent blue-edged, yellow
band from front of eye to above upper lip, encompassing nostrils; cheek yellowish with 3 or 4
oblique, blue bands sloping upwards from above upper jaw towards rear part of eye; fins mainly clear or
whitish; a black spot at base of upper pectoral-fin rays and narrow brown bar across base of entire fin.
Habitat, biology, and fisheries: Inhabits sand, mud, and rubble bottoms, usually at depths between about
20 and 80 m. It feeds mainly on small bottom-living gastropod molluscs. Common in some markets of
Malaysia, Thailand, and Viet Nam, usually fresh or dried-salted. Caught mainly with bottom trawls, but
sometimes with bottom longlines and handlines.
Distribution: Indo-Malaysian
Archipelago and South China
Sea.
3022 Bony Fishes
Gymnocranius grandoculis (Valenciennes, 1830) (Plate XVII, 124)

Frequent synonyms / misidentifications: Gymnocranius robinsoni (Gilchrist and Thompson, 1909) /
None.
FAO names: En - Bluelined large-eye bream.
Diagnostic characters: Body oblong, its depth usually 2.4 to 2.5 times in standard length. Forehead
profile moderately steep; large adults developing a bony ridge on nape and bony shelf over front part of
eyes; lower edge of eye well above a line from tip of snout to middle of caudal-fin fork; eye relatively large,
about equal to or larger than interorbital space and preorbital width in small specimens (less than about
40 cm), but much smaller in comparison to interorbital and preorbital widths in large fishes; mouth relatively
small, posterior part of jaws not reaching a vertical through front of eye; outer surface of maxilla smooth;
2 or 3 pairs of slender canines at front of each jaw, although the outer canine on each side of lower jaw is
usually enlarged; other teeth villiform, becoming strongly conical on lateral sections of jaws. Dorsal fin with
X slender spines and 10 soft rays; anal fin with III slender spines and 10 soft rays; pectoral-fin rays 14;
caudal fin only moderately forked with pointed tips, the median rays much longer than eye diameter.
Lateral-line scales 48 plus 2 or 3 additional tubed scales extending onto base of caudal fin; cheek with 3
or 4 transverse scale rows; 5 ½ scale rows between lateral line and base of middle dorsal-fin spines;
inner surface of pectoral-fin axil scaleless. Colour: overall silvery with thin brown scale margins;
anterior half of head often brown; a series of narrow undulating, longitudinal lines on cheek and side of
snout; fins clear to yellow or orange; caudal fin frequently dusky grey-brown; a narrow brown bar across
base of pectoral fins. Juveniles under about 25 cm standard length often with 5 or 6 irregular dark bars
on side and dark bar below eye. The bar at the level of the pectoral-fin base is usually the widest and
the dark coloration extends onto the pelvic fins.
Habitat, biology, and fisheries: Inhabits trawling grounds of the continental shelf and offshore rocky
bottoms, usually at depths between about 50 and 100 m. It feeds mainly on bottom-living invertebrates
and small fishes. Occasionally seen in markets, mostly fresh. Caught mainly with bottom longlines and
bottom trawls. A good potential commercial fish if sufficient stocks can be located.
uted in the Indian Ocean and
western edge of the Pacific
Ocean, from East Africa to
southeastern Oceania and
Australia northward to Japan.
Gymnocranius griseus (Schlegel, 1844)

Frequent synonyms / misidentifications: None / Gymnocranius grandoculis (Valenciennes, 1830)
(juveniles).
FAO names: En - Grey large-eye bream; Fr - Empereur gris; Sp - Emperador gris.
juvenile
Diagnostic characters: Body oblong, its depth usually 1.9 to 2.3 times in standard length. Dorsal and
ventral profile of head evenly convex or ventral profile slightly straighter; lower edge of eye slightly above
a line from tip of snout to middle of caudal-fin fork; eye relatively large, its diameter about equal to or slightly
larger than preorbital and interorbital widths, mouth relatively small, posterior part of jaws reaching to about
level of anterior nostrils; outer surface of maxilla smooth; 2 or 3 pairs of small, slender canines at front
of each jaw, other teeth villiform, becoming conical on lateral sections. Dorsal fin with X slender spines
and 10 soft rays; anal fin with III slender spines and 10 soft rays; pectoral-fin rays 14; caudal fin
moderately forked with pointed tips, the median rays slightly longer than eye diameter. Lateral-line
scales 46 to 48 plus 2 or 3 additional tubed scales extending onto base of caudal fin; cheek with 4
inner surface of pectoral-fin axil scaleless. Colour: overall silvery, frequently with a diffuse to vivid
pattern of 5 to 8 narrow dark bars on side, including 1 through eye and across cheek; fins mainly
clear to yellowish, sometimes diffuse mottling or spotting on dorsal, caudal, and anal fins; often a narrow
brown bar across bases of pectoral fins. Some adults with a few scattered blue spots or scribbling on the
snout and cheek.
Size: Maximum total length reported as 80 cm, but it is doubtful that the species grows that large; maximum
reliable total length about 35 cm, commonly to 25 cm.
Habitat, biology, and fisheries: Inhabits coastal trawling grounds, usually at depths between about 20
and 80 m. It feeds mainly on bottom-living invertebrates. Sexual maturation occurs at a fairly small size,
about 15 to 17 cm standard length. Common in markets, mainly fresh, occasionally dried. Caught mainly
with bottom trawls, but sometimes with handlines.
Distribution: Southern
Japan to the Indo-Malaysian
region, western Australia, and
possibly eastward to India.
The species has often been
confused with the young of
Gymnocranius grandoculis,
hence the true distributional
limits are somewhat obscure.
Records of G. griseus from
the western Indian Ocean
p r o b a b l y p e r t a i n t o G.
grandoculis.
3024 Bony Fishes
Gymnocranius microdon (Bleeker, 1851) (Plate XVII, 125)

FAO names: En - Bluespotted large-eye bream.
Diagnostic characters: Body oblong, its depth 2.5 to 3 times in standard length. Forehead profile
moderately steep; lower edge of eye well above a line from tip of snout to middle of caudal-fin fork; eye
large, about equal to width of preorbital and interorbital spaces; mouth relatively small, posterior part of
jaws reaching to about level of nostril openings; outer surface of maxilla smooth; 2 or 3 pairs of small,
slender canines at front of each jaw, other teeth villiform becoming conical on lateral sections of jaws.
Dorsal fin with X slender spines and 10 soft rays; anal fin with III slender spines and 10 soft rays; pectoral-fin
rays 14; caudal fin moderately forked with pointed tips, median rays longer than eye diameter.
Lateral-line scales 48 or 49 plus 2 or 3 tubed scales extending onto base of caudal fin; cheek with 3 or 4
inner surface of pectoral-fin axil scaleless. Colour: overall silvery to brownish, scale margins with a
narrow brown border; numerous vertically elongate blue spots on snout and cheek, and sometimes
on operculum; fins clear to yellow or reddish, caudal fin sometimes dusky brown.
Habitat, biology, and fisheries: Inhabits sand and rubble bottoms adjacent to rock and coral reefs, usually
at depths between about 20 and 50 m. It feeds mainly on small bottom-living invertebrates. Occasionally
seen in markets, usually fresh. Caught mainly with bottom longlines and handlines.
Distribution: Known thus far
only from southern Japan, the
South China Sea to Scott reef
off northwestern Australia, the
Marshall Is lands, and off
Phuket, Thailand in the An-
daman Sea.
Gymnocranius sp.
Frequent synonyms / misidentifications: None / Gymnocranius lethrinoides (non Bleeker, 1850).
FAO names: En - Blacknape large-eye bream.
convex, the snout sloping steeply; lower edge of eye well above a line from tip of snout to middle of
caudal-fin fork; eye large, its diameter about equal to interorbital space and larger than width of preorbital;
mouth relatively small, posterior part of jaws reaching to about level of nostril openings; outer surface of
maxilla smooth; 2 or 3 pairs of small, slender canines at front of each jaw, other teeth villiform, becoming
conical on lateral sections of jaws. Dorsal fin with X slender spines and 10 soft rays; anal fin with III slender
spines and 10 soft rays; pectoral-fin rays 14; caudal fin moderately forked, inner edge of fork slightly
convex, with blunt tips, the median rays slightly longer than eye diameter. Lateral-line scales 47 or
48 plus 3 or 4 tubed scales extending onto base of caudal fin; cheek with 4 or 5 transverse scale rows;
5 ½ scale rows between lateral line and base of middle dorsal-fin spines; inner surface of pectoral-
fin axil scaleless. Colour: overall silvery or silvery white; scales of back (and sometimes lower on
sides) with a prominent blackish basal spot forming longitudinal rows; area immediately above eye
(supraorbital shelf) blackish; a blackish (may be very faint) bar below eye to lower edge of preopercle; fins
mainly clear to reddish, caudal fin sometimes dusky reddish brown.
Habitat, biology, and fisheries: Inhabits sand and rubble bottoms adjacent to rock and coral reefs, usually
at depths between 15 and 40 m. It feeds mainly on small bottom-living gastropod molluscs. Sexual
maturation is attained by a total length of at least 27 cm. Occasionally seen in markets, usually fresh.
Caught mainly with bottom longlines and handlines, sometimes speared.
from southern Japan, western
Australia to the Great Barrier
Reef of Australia, Coral Sea,
and New Caledonia.
3026 Bony Fishes
Lethrinus amboinensis Bleeker, 1854 (Plate XVII, 126)

Frequent synonyms / misidentifications: None / Lethrinus microdon non Valenciennes, 1830;
L. olivaceous non Valenciennes, 1830.
FAO names: En - Ambon emperor.
Diagnostic characters: Body moderately elongate, its depth 2.8 to 3.3 times in standard length. Head
length 1.1 to 1.3 times the body depth, 2.5 to 2.8 times in standard length, dorsal profile in front of eye
nearly straight; snout moderately long, its length about 1.7 to 1.9 times in head length, measured without
the lip the snout is 0.7 to 0.9 times in cheek height, its dorsal profile concave or nearly straight, snout
angle relative to upper jaw between 45° and 58°; interorbital space convex; posterior nostril an oblong
longitudinal opening, about half-way between orbit and anterior nostril; eye not situated close to dorsal
profile, its length 3.3 to 6.1 times in head length; cheek not very high, its height 2.5 to 4.0 times in head
length; lateral teeth in jaws conical; outer surface of maxilla smooth, or with a longitudinal ridge. Dorsal fin
with X spines and 9 soft rays, the third dorsal-fin spine usually the longest, its length 2.4 to 2.7 times in body
depth; anal fin with III spines and 8 soft rays, the first soft ray usually the longest, its length much less than
base of soft portion of anal fin and 1.5 to 1.9 times in length of entire anal-fin base; pectoral-fin rays 13;
pelvic-fin membranes between rays closest to body with dense melanophores. Lateral-line scales 46
to 48; cheek without scales; 4 ½ scale rows between lateral line and base of middle dorsal-fin spines;
usually 16 or 17 scale rows in transverse series between origin of anal fin and lateral line; 15 or 16
(usually 15) rows in lower series of scales around caudal peduncle; 5 to 11 scales in supratemporal patch;
inner surface of pectoral-fin base without scales; posterior angle of operculum fully scaly. Colour: body
yellowish with scattered indistinct dark blotches; head brown, sometimes with light streaks; lips reddish; axil
of pectoral fins orangish; basal portion of centre rays of pectoral fins white, the outer edges yellow; pelvic
and anal fins white or yellowish; dorsal and caudal fins mottled brown or yellow with and orange or reddish
edge.
Size: Maximum reported total length about 70 cm; commonly between 30 and 50 cm.
Habitat, biology, and fisheries: Inhabits deeper waters of coral reefs and adjacent sandy bottoms. Feeds
primarily on fishes and crustaceans. Very little is known of the biology of this species, perhaps because it
is easily mistaken for either
Lethrinus microdon or L. oli-
vaceus, particularly larger indi-
viduals. Caught primarily with
handlines; marketed fresh.
Distribution: I n d on es i a,
northwestern Australia, Philip-
pines to southern Japan, east-
ward through the Marshall
Islands, Solomons, Samoa,
and the Marquesas.
Lethrinus atkinsoni Seale, 1910 (Plate XVII, 127)

Frequent synonyms / misidentifications: None / Lethrinus mahsena (non Forsskål, 1775).
FAO names: En - Pacific yellowtail emperor.
Diagnostic characters: Body moderately deep, its depth 2.3 to 2.6 times in standard length. Head length
0.8 to 0.9 times in body depth, 2.7 to 2.9 times in standard length, dorsal profile near eye convex to
approximately straight; snout moderately short and blunt, its length about 1.9 to 2.6 times in head length,
measured without the lip the snout is 0.9 to 1.1 times in cheek height, its profile steep and nearly straight,
snout angle relative to upper jaw between 65° and 70°; interorbital space convex or slightly convex;
posterior nostril an oblong longitudinal opening, closer or slightly closer to orbit than to anterior nostril; eye
situated close to dorsal profile, its length 3.2 to 3.8 times in head length; cheek high, its height 2.1 to 2.8
times in head length; lateral teeth in jaws rounded or molars; outer surface of maxilla smooth, or with a
slight longitudinal ridge. Dorsal fin with X spines and 9 soft rays, the third or fourth dorsal-fin spine usually
the longest, its length 2.6 to 4 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray
usually the longest, its length equal to or longer than base of soft portion of anal fin and 1.1 to 1.4
times in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest
to body without dense melanophores. Lateral-line scales 46 to 47; cheek without scales; 4 ½ scale rows
between lateral line and base of middle dorsal-fin spines; 14 to 16 scale rows in transverse series
between origin of anal fin and lateral line; 13 to 15 (usually 14) rows in lower series of scales around
caudal peduncle; 4 to 8 scales in supratemporal patch; inner surface of pectoral-fin base densely
covered with scales; posterior angle of operculum fully scaly. Colour: upper sides bluish grey, tan or
yellowish, white ventrally, the caudal peduncle and sometimes a broad indistinct band on sides yellow; head
brown, lips reddish; fins pale yellowish, orange, or reddish; the base of pectoral fins, basal membranes of
anal fin and edges of pelvic, dorsal, anal, and caudal fins often reddish.
Size: Maximum size reported as about 40 to 50 cm total length; commonly between 30 and 35 cm.
Habitat, biology, and fisheries: Inhabits outer reef slopes, sandy areas in lagoons and seagrass beds to
depths of about 30 m. Reported as both typically solitary and swimming in schools. Feeds primarily on
crustaceans, molluscs, and fishes. Caught primarily by handline, trawl, shore seine, and gill net. Marketed
mostly fresh. Good eating but
not as sought after as other
emperors in some areas due to
its small size. Puts up a good
fight on a handline. Reported
to contribute about 4% to the
total commercial catch in New
Caledonia.
Distribution: Indonesia,
northern Australia, Philippines
to southern Japan, wide-
spread throughout West Pa-
cific to the Tuamotus.
3028 Bony Fishes
Lethrinus conchyliatus Smith, 1959 (Plate XVII, 128)

FAO names: En - Redaxil emperor; Fr - Empereur guele de vin; Sp - Emperador boca de vino.
Diagnostic characters: Body moderately elongate, its depth 3.2 to 3.4 times in standard length. Head length
about 1.1 to 1.3 times in body depth, about 2.5 to 2.8 times in standard length, dorsal profile near eye convex;
snout length about 1.7 to 1.9 times in head length, measured without the lip the snout is about 0.7 to 0.9
times in cheek height, its dorsal profile concave with a hump in front of eye, snout angle relative to upper
jaw between 45° and 60°; interorbital space convex; posterior nostril a longitudinal oblong opening, closer
to orbit than anterior nostril; eye not close to dorsal profile, its length about 4.0 to 6.0 times in head length;
cheek height about 2.6 to 3.2 times in head length; lateral teeth in jaws conical; outer surface of maxilla
smooth. Dorsal fin with X spines and 9 soft rays, the third or fourth dorsal-fin spine the longest; anal fin
with III spines and 8 soft rays, the first soft ray usually the longest, its length shorter than length of base of
soft-rayed portion of anal fin and about 1.7 to 1.9 times in length of entire anal-fin base; pectoral-fin rays 13;
pelvic-fin membranes between rays closest to body with or without dense melanophores. Lateral-line scales
47 or 48; cheek without scales; 4 ½ scale rows between lateral line and base of middle dorsal-fin
spines; usually 16 or 17 scale rows in transverse series between origin of anal fin and lateral line; usually
15 rows in lower series of scales around caudal peduncle; 9 to 11 scales in supratemporal patch; inner
surface of pectoral-fin base without scales; a large, distinct scaleless patch above base of pectoral
fins; posterior angle of operculum fully scaly. Colour: body and head brownish or grey, lighter ventrally;
lips, pectoral-fin base, patch above base of pectoral fins, and edge of operculum red; fins orangish, dusky,
or mottled.
Habitat, biology, and fisheries: Inhabits deep reefs to depths of 220 m. Feeds primarily on fishes and
crustaceans. Caught mostly by handline, traps, and trawl.
Distribution: Indian Ocean,
from Tanzania, northern
Madagascar, Chagos, An-
damans, and southwestern In-
donesia.
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Lethrinus erythracanthus Valenciennes, 1830 (Plate XVII, 129)

Frequent synonyms / misidentifications: Lethrinus kallopterus Bleeker, 1856 / None.
FAO names: En - Orangespotted emperor; Fr - Empereur capitaine; Sp - Emperador de manchas.
moderately deep, its depth 2.5
to 2.7 times in standard length.
Head length 0.9 to 1 times in
body depth, 2.5 to 2.8 times in
standard length, dorsal profile
near eye nearly straight or
slightly convex; snout moder-
ately short, its length about 1.8
to 2.4 times in head length,
measured without the lip the
snout is 0.8 to 1.1 times in cheek
height, its dorsal profile dis-
tinctly concave in large individu-
als a nd near ly s traight in
smaller individuals, snout angle
relative to upper jaw between
55° and 69°; interorbital space
convex; posterior nostril an ob-
long longitudinal opening, juvenile
closer to orbit than anterior nos-
tril; eye situated close to or far
removed from dorsal profile, its
length 2.7 to 5.2 times in head
length; cheek moderately high, subadult
its height 2 to 3.4 times in head
length; lateral teeth in jaws conical; outer surface of maxilla smooth or with a longitudinal ridge. Dorsal fin
with X spines and 9 soft rays, the fourth or fifth dorsal-fin spine the longest, its length 2.5 to 3.5 times
in body depth; anal fin with III spines and 8 soft rays, the third, fourth, or fifth soft ray usually the longest,
its length much longer than length of base of soft-rayed portion of anal fin and 0.8 to 1.1 times in
length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body
with or without dense melanophores. Lateral-line scales 46 to 48; cheek without scales; 4 ½ scale rows
between origin of anal fin and lateral line; usually 15 rows in lower series of scales around caudal peduncle;
5 to 8 scales in supratemporal patch; inner surface of pectoral-fin base densely covered with scales;
posterior angle of operculum fully scaly. Colour: body brown to dark grey, with indistinct scattered small
dark and light spots, with irregular light stripes sometimes on lower sides; head brown or grey, often with
many small orange spots on cheeks; pectoral and pelvic fins white to orangish, dorsal and anal fins mottled
orange and bluish; caudal fin often bright orange.
Habitat, biology, and fisheries: Inhabits deep lagoons and channels, and outer reef slopes and adjacent
soft-bottom areas at depth of 20 to 120 m. It feeds mostly on echinoderms, crustaceans, and molluscs. Caught
by handline, vertical longline, traps, and trawls. Marketed mostly fresh.
Distribution: Western Indian
Ocean to the Central Pacific,
from East Africa, Seychelles,
Chagos, and Maldives, to
T h a i l a n d, P h il i p pi ne s ,
Ryukyus, Micronesia, north-
western and northeastern
Australia, Samoa, Society Is-
lands, and Tuamotus.
3030 Bony Fishes
Lethrinus erythropterus Valenciennes, 1830 (Plate XVIII, 130)

Frequent synonyms / misidentifications: Lethrinus hypselopterus Bleeker, 1873 / None.
FAO names: En - Longfin emperor; Fr - Empereur békine.
0.8 to 1.0 times in body depth, 2.5 to 2.8 times in standard length, dorsal profile near eye nearly straight or
slightly convex; snout moderately short, its length about 1.9 to 2.2 times in head length, measured without
the lip the snout is 0.9 to 1 times in cheek height, its dorsal profile concave, snout angle relative to upper
jaw between 53° and 64°; interorbital space convex or almost flat; posterior nostril an oblong longitudinal
opening, closer to orbit than anterior nostril; eye situated close to or slightly removed from dorsal profile, its
length 3.3 to 4.6 times in head length; cheek moderately high, its height 2.1 to 2.7 times in head length;
lateral teeth in jaws of adults molars or rounded; outer surface of maxilla usually smooth. Dorsal fin with
X spines and 9 soft rays, the fourth or fifth dorsal-fin spine the longest, its length 2.7 to 3.3 times in body
depth; anal fin with III spines and 8 soft rays, the third, fourth, or fifth soft ray usually the longest, its
length much longer than length of base of soft-rayed portion of anal fin and 0.7 to 1 times in length
of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body without
dense melanophores. Lateral-line scales 44 to 46; cheek without scales; 4 ½ scale rows between lateral
line and base of middle dorsal-fin spines; 15 to 17 scale rows in transverse series between origin of anal
fin and lateral line; usually 15 rows in lower series of scales around caudal peduncle; 5 to 9 scales in
supratemporal patch; inner surface of pectoral-fin base densely covered with scales; posterior angle
of operculum fully scaly. Colour: head and body brown or rust red, lighter ventrally; sometimes 2 light bars
on caudal peduncle; area around eye, a broad streak from eye to tip of snout, lips, and base of pectoral fins
red; all fins reddish, often a bright red or orange.
Size: Maximum total length about 50 cm; commonly around 30 cm.
Habitat, biology, and fisheries: Mostly inhabits coral reefs and adjacent sandy areas. Occurs singly or in
small groups. Feeds on echinoderms, molluscs, crustaceans, and small fishes. Limited interest to fisheries
in the countries it occurs in. Caught mostly by handlines and traps. Marketed fresh.
Distribution: Western Indian
Ocean and West Pacific; Tan-
z a n i a an d M o z am bi qu e,
Chagos, Celebes, the Philip-
pines, northwestern Australia,
Papua New Guinea, Palau,
and Caroline Islands.
Lethrinus genivittatus Valenciennes, 1830 (Plate XVIII, 131)

Frequent synonyms / misidentifications: Lethrinus nematacanthus Bleeker, 1854 / None.
FAO names: En - Longspine emperor.
Diagnostic characters: Body moderately elongate, moderately slender, its depth 2.9 to 3.5 times in
standard length. Head length 1 to 1.2 times in body depth, 2.8 to 3 times in standard length, dorsal profile
near eye slightly convex; snout moderately short and blunt, its length about 2.2 to 2.5 times in head length,
measured without the lip the snout is 0.9 to 1 times in cheek height, its dorsal profile nearly straight, snout
angle relative to upper jaw between 60° and 74°; interorbital space convex to flat; posterior nostril an oblong
nearly vertical opening, about half-way between orbit and anterior nostril; eye situated close to dorsal profile,
its length 3.6 to 4 times in head length; cheek not high, its height 2.6 to 3.2 times in head length; lateral teeth
in jaws conical; a pair of teeth in lower jaw strongly curved outward; outer surface of maxilla with a
distinct knob. Dorsal fin with X spines and 9 soft rays, the second dorsal-fin spine the longest, sometimes
much longer than other dorsal-fin spines, its length 1.3 to 1.9 times in body depth; anal fin with III
spines and 8 soft rays, the first or second soft ray usually the longest, its length almost equal to or slightly
shorter than length of base of soft-rayed portion of anal fin and 1.2 to 1.7 times in length of entire anal-fin
base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body without dense melanophores.
Lateral-line scales 45 to 47; cheek without scales; 4 ½ scale rows between lateral line and base of middle
dorsal-fin spines; 15 or 16 scale rows in transverse series between origin of anal fin and lateral line; usually
15 rows in lower series of scales around caudal peduncle; 5 to 8 scales in supratemporal patch; inner surface
of pectoral-fin base without scales, partially covered with scales or, densely covered with scales; posterior
angle of operculum fully scaly. Colour: body tan or brown on upper sides, lower sides white with 3 brown
or tan stripes, sides often with scattered irregular black oblique bars and a square black blotch above pectoral
fins and bordering below lateral line; head brown or tan sometimes with several broad, somewhat indistinct
vertical and oblique bands (the bands sometimes composed of fine reticulations); fins pale, speckled with
small white blotches.
Size: Maximum total length about 25 cm; commonly to around 15 cm.
Habitat, biology, and fisheries: Primarily in shallow sandy and seagrass areas, but also in mangrove
swamps, lagoons, channels, and outer reefs slopes at depths of 5 to 25 m; penetrates into rivers in some
areas. Feeds mostly on crus-
taceans and small fishes.
Caught mostly by shore seine.
Minor importance in fisheries.
Marketed fresh.
Ocean and West Pacific, in-
cluding Indonesia, northern
Australia, the Philippines,
southern Japan, Papua New
Guinea, and the Caroline Is-
lands.
3032 Bony Fishes
Lethrinus harak (Forsskål, 1775) (Plate XVIII, 132)

Frequent synonyms / misidentifications: Lethrinus rhodopterus Bleeker, 1852 / None.
FAO names: En - Thumbprint emperor; Fr - Empereur Saint Pierre; Sp - Emperador San Pedro.
0.9 to 1 times in body depth, 2.7 to 3.1 times in standard length, dorsal profile near eye distinctly or slightly
convex; snout short and blunt, its length about 2.0 to 2.6 times in head length, measured without the lip the
snout is 0.9 to 1 times in cheek height, its dorsal profile nearly straight, snout angle relative to upper jaw
between 60° and 70°; interorbital space convex or almost flat; posterior nostril a narrow longitudinal slit,
closer to orbit than anterior nostril; eye situated close to dorsal profile, its length 3.4 to 4.2 times in head
length; cheek not very high, its height 2.3 to 3.1 times in head length; lateral teeth in jaws of adults molars
or rounded; outer surface of maxilla smooth or with a longitudinal ridge. Dorsal fin with X spines and 9 soft
rays, the fourth or fifth dorsal-fin spine the longest, its length 2.5 to 3.1 times in body depth; anal fin with III
spines and 8 soft rays, the first soft ray usually the longest, its length almost equal to or longer or shorter
than length of base of soft-rayed portion of anal fin and 1.2 to 1.6 times in length of entire anal-fin base;
pectoral-fin rays 13; pelvic-fin membranes between rays closest to body without dense melanophores.
Lateral-line scales usually 46 or 47; cheek without scales; usually 5 ½ but sometimes 4 ½ scale rows
between origin of anal fin and lateral line; 13 or 14 rows in lower series of scales around caudal peduncle;
posterior angle of operculum fully scaly. Colour: olive or grey above, shading to silvery white below; a large
elliptical black spot, often broadly edged in yellow, on side directly below lateral line and centred at
a vertical near the posterior tip of pectoral fins; sometimes light blue dots bordering lower rim of eye and
around nostrils; pectoral, pelvic, dorsal, and anal fins white to pinkish; caudal fin orange or reddish; vertical
fins sometimes lightly mottled or striped.
Size: Maximum total length about 50 cm (perhaps 60 cm); most commonly between 20 to 30 cm.
Habitat, biology, and fisheries: In shallow sandy, coral rubble, mangrove, lagoon, channel, and seagrass areas
inshore and adjacent to coral reefs. Feeds on polychaetes, crustaceans, molluscs, echinoderms, and small fishes.
Often observed solitary but also in small schools. Caught mostly by shore seine, gill nets, traps, and handline.
Distribution: Indian Ocean
and West Pacific, including
the Red Sea, East Africa, Sey-
chelles, Maldives, Sri Lanka,
Andamans, Indonesia, the
Philippines, southern Japan,
northeast Australia, Papua
New Guinea, the Caroline Is-
lands, the Marshall Islands,
Solomons, Vanuatu, Fiji, and
Samoa.
Lethrinus laticaudis Alleyne and Macleay, 1877

Frequent synonyms / misidentifications: Lethrinus fletus Whitley, 1943; L. anarhynchus Postel, 1965 /
Lethrinus fraenatus non Valenciennes, 1830.
FAO names: En - Grass emperor.
Diagnostic characters: Body relatively deep, its depth 2.3 to 2.5 times in standard length. Head length 0.8
to 0.9 times in body depth, 2.7 to 2.9 times in standard length, dorsal profile near eye nearly straight, or
concave in large individuals; snout length about 1.8 to 2.2 times in head length, measured without the lip
the snout is 0.9 to 1 times in cheek height, its dorsal profile concave, snout angle relative to upper jaw
between 58° and 66°; interorbital space convex; posterior nostril an oblong longitudinal opening, closer to
orbit than anterior nostril; eye removed from dorsal profile, its length 3.1 to 5.3 times in head length; cheek
moderately high, its height 2.2 to 2.9 times in head length; lateral teeth in jaws conical; outer surface of
maxilla with a longitudinal ridge. Dors al fin with X spines and 9 soft rays, the fourth dorsal-fin spine usually
the longest, its length 2.5 to 3.4 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray
usually the longest, its length almost equal to or slightly shorter than length of base of soft-rayed portion of
anal fin and 1.3 to 1.8 times in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes
between rays closest to body with or without dense melanophores. Lateral-line scales 46 to 48; cheek
without scales; 5 ½ scale rows between lateral line and base of middle dorsal-fin spines; 16 or 17
scale rows in transverse series between origin of anal fin and lateral line; usually 15 rows in lower series of
scales around caudal peduncle; 5 to 8 scales in supratemporal patch; inner surface of pectoral-fin base
more than half covered with scales; posterior angle of operculum fully scaly. Colour: body tan, brown, or
yellow with scattered irregular dark blotches; centres of scales on anterior upper sides with distinct
black markings; head brown or yellow with blue dots on cheeks and short blue stripes radiating in front and
behind eye, sometimes a number of blue cross stripes between the eyes; fins pale or yellow, the vertical
fins mottled.
Size: Maximum reported total length 56 cm; commonly between 30 and 40 cm.
Habitat, biology, and fisheries: Juveniles inhabit seagrass beds and mangrove swamps while adults are
found mostly on coral reefs. Feeds mostly on crustaceans and fishes. Significantly more and smaller females
than males have been con-
firmed in populations in Aus-
tralia. Caught primarily by
handlines. Mostly important
as a sport fish and in handline
fisheries in Australia. A good
food fish. Marketed fresh.
Distribution: Southern Indo-
nesia, nortwestern and north-
eastern Australia, Papua New
Guinea, and the Solomons.
3034 Bony Fishes
Lethrinus lentjan (Lacepède, 1802) (Plate XVIII, 133)

Frequent synonyms / misidentifications: Lethrinus opercularis Valenciennes, 1830 / None.
FAO names: En - Redspot emperor; Fr - Empereur lentilles; Sp - Emperador de lentejuelas.
Diagnostic characters: Body moderately deep, its depth 2.5 to 2.8 times in standard length. Head
length 0.9 to 1 times in body depth, 2.6 to 3 times in standard length, dorsal profile near eye nearly straight;
snout moderately short, its length about 1.9 to 2.4 times in head length, measured without the lip the snout
is 0.8 to 1 times in cheek height, its dorsal profile nearly straight, snout angle relative to upper jaw between
60° and 70°; interorbital space convex; posterior nostril an oblong longitudinal opening, closer to orbit than
anterior nostril; eye situated close to or far removed from dorsal profile, its length 3.3 to 4.8 times in head
length; cheek not high, its height 2.4 to 3.1 times in head length; lateral teeth in jaws rounded often with
conical tips, or molars often with tubercles; outer surface of maxilla with a longitudinal ridge. Dorsal
fin with X spines and 9 soft rays, the fourth dorsal-fin spine usually the longest, its length 2.4 to 3.4 times in
body depth; anal fin with III spines and 8 soft rays, the first soft ray usually the longest, its length almost
equal to or shorter than length of base of soft-rayed portion of anal fin and 1 to 1.2 times in length of entire
anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body without dense
melanophores. Lateral-line scales usually 46 or 47; cheek without scales; 5 ½ scale rows between lateral
line and base of middle dorsal-fin spines; 15 or 16 scale rows in transverse series between origin of
anal fin and lateral line; usually 15 rows in lower series of scales around caudal peduncle; 4 to 9 scales in
supratemporal patch; inner surface of pectoral-fin base densely covered with scales, with a few scales,
or naked; posterior angle of operculum fully scaly. Colour: body greenish or grey, shading to white below,
centres of scales on upper sides often white; posterior margin of opercle and sometimes base of pectoral
fins red; pectoral fins white, yellow, or pinkish; pelvic and anal fins white to orange; dorsal fin white and
orange mottled with a reddish margin; caudal fin mottled orange or reddish.
Size: Maximum size about 50 cm total length; commonly to around 30 cm.
Habitat, biology, and fisheries: Over sandy bottom in coastal areas, deep lagoons, and near coral reefs,
to depths of about 50 m. Juveniles and small adults often in aggregations over seagrass beds, mangrove
swamps and shallow sandy areas; adults are generally solitary and found in deeper waters. Feeds mostly
on crustaceans and molluscs,
b u t e c h i n o d e r m s , p ol y-
chaetes, and fishes are also
consumed. Caught primarily
by handline, traps, trawls,
beach seines, and gill nets.
Indo-West Pacfic, including
Red Sea, Persian Gulf, East
Africa to the Ryukus and
Tonga.
Lethrinus microdon Valenciennes, 1830 (Plate XVIII, 134)

Frequent synonyms / misidentifications: Lethrinus elongatus Valenciennes, 1830 / None.
FAO names: En - Smalltooth emperor; Fr - Empereur tidents; Sp - Emperador boquidulce.
relatively elongate, its depth 2.9
Head length 1 to 1.3 times in
standard length, dorsal profile near
eye nearly straight; snout moder-
ately long, its length about 1.8 to
2.3 times in head length, meas-
ured without the lip the snout is
0.7 to 0.8 times in cheek height,
its dorsal profile slightly concave,
snout angle relative to upper jaw
between 47° and 59°; interorbital mottled pattern
space convex to flat; posterior
nostril an oblong longitudinal open-
ing, closer to orbit than anterior
nostril; eye situated close to or far
removed from dorsal profile, its
length 3.4 to 5.7 times in head
length; cheek not high, its height
2.7 to 3.4 times in head length;
lateral teeth in jaws conical; outer
surface of maxilla smooth. Dorsal
fin with X spines and 9 soft rays, the
third or fourth dorsal-fin spine the
longest, its length 2.4 to 3 times in
body depth; anal fin with III spines
and 8 soft rays, the first soft ray silvery pattern
usually the longest, its length al-
most equal to or shorter than length
of base of soft-rayed portion of anal fin and 1.5 to 1.8 times in length of entire anal-fin base; pectoral-fin rays
13; pelvic-fin membranes between rays closest to body with or without dense melanophores. Lateral-line
scales 46 to 48; cheek without scales; 4 ½ scale rows between lateral line and base of middle dorsal-fin
spines; usually 16 or 17 scale rows in transverse series between origin of anal fin and lateral line;
usually 15 rows in lower series of scales around caudal peduncle; 7 to 11 scales in supratemporal patch;
inner surface of pectoral-fin base without scales; posterior angle of operculum fully scaly. Colour: body
grey or brown with scattered irregular dark blotches on sides; sometimes 3 dark streaks radiating forward
from eye; fins pale or orangish.
Size: Maximum total length about 70 cm; commonly between 30 and 50 cm.
Habitat, biology, and fisheries: Inhabits sandy areas near coral reefs to depths of about 80 m. Swims in
small schools, sometimes together with Lethrinus olivaceus. Feeds primarily on fishes, crustaceans,
cephalopods, and polychaetes during both night and day. Spawning in Palau occurs throughout most of the
year, during the first 5 days of
the lunar month, near the edge
of reefs. Caught mostly with
handlines, traps, gill nets, and
trawls. An excellent food fish.
Distribution: W i d e s pr e ad
Indo-West Pacific, including
Red Sea, Persian Gulf, East
Africa to Sri Lanka, to the
Ryukyu Islands, western Aus-
tralia, Palau, and Papua New
Guinea.
3036 Bony Fishes
Lethrinus miniatus (Schneider, 1801) (Plate XVIII, 135)

Frequent synonyms / misidentifications: Lethrinus chrysostomus Richardson, 1848; L. amamianus
Akazaki, 1962 / None.
FAO names: En - Trumpet emperor.
Diagnostic characters: Body moderately deep, its depth 2.4 to 2.8 times in standard length. Head
length 0.9 to 1 times in body depth, 2.5 to 2.9 times in standard length, dorsal profile near eye slightly convex;
snout moderately long, its length about 1.7 to 2.4 times in head length, measured without the lip the snout
is 0.9 to 1 times in cheek height, its dorsal profile slightly concave, snout angle relative to upper jaw
between 50° and 65°; interorbital space convex to flat; posterior nostril an oblong longitudinal opening, closer
to orbit than anterior nostril; eye situated close to dorsal profile, its length 3.4 to 5.4 times in head length;
cheek moderately high, its height 2.0 to 2.9 times in head length; lateral teeth in jaws conical; outer surface
of maxilla smooth or with a longitudinal ridge. Dorsal fin with X spines and 9 soft rays, the third dorsal-fin
spine usually the longest, its length 2.3 to 2.9 times in body depth; anal fin with III spines and 8 soft rays,
the first or second soft ray usually the longest, its length almost equal to or slightly longer than length of base
of soft-rayed portion of anal fin and 1.3 to 1.4 times in length of entire anal-fin base; pectoral-fin rays 13;
pelvic-fin membranes between rays closest to body usually with dense melanophores. Lateral-line scales
46 to 48; cheek without scales; 4 ½ scale rows between lateral line and base of middle dorsal-fin spines;
15 to 17 scale rows in transverse series between origin of anal fin and lateral line; usually 15 rows in lower
series of scales around caudal peduncle; 2 to 7 scales in supratemporal patch; inner surface of pectoral
fins partially or densely covered with scales; posterior angle of operculum fully scaly. Colour: body
silvery, tan, or yellowish, base of scales often black, often a series of 8 or 9 dark bars; base of pectoral fins
red; sometimes a red streak originating on upper operculum, passing beneath eye and onto snout;
2 red spots often on upper rim of eye; lips reddish; fins pale or reddish, sometimes brilliant red on
membranes near base of pelvic fins, and between spinous rays of dorsal and anal fin.
Size: Maximum size reported to about 90 cm total length and about 9 kg.
Habitat, biology, and fisheries: Feeds in sand around coral reefs in daytime; at nightime feeds mostly over
sandy sea floor near reefs; at depths between 5 and 30 m. Usually in small schools. Feeds mostly on
crustaceans, echinoderms, molluscs, and fishes, with crabs and sea urchins predominating. Caught primarily
by handline. One of the favourite food and sport fishes around the Great Barrier Reef. A major food fish in
New Caledonia. Occasionally
implicated in cases of fish
poisioning (ciguatoxin).
Distribution: The Ryukyu Is-
lands, eastern Philippines,
northern Australia, and New
Caledonia.
Lethrinus nebulosus (Forsskål, 1775) (Plate XVIII, 136)

Frequent synonyms / misidentifications: Lethrinus choerorynchus Schneider, 1801; L. fraenatus
Valenciennes, 1830 / None.
FAO names: En - Spangled emperor; Fr - Empereur moris; Sp - Emperador relámpago.
moderately deep, its depth 2.5
Head length 0.9 to 1 times in
standard length, dorsal profile
near eye nearly straight, or in
large individuals, distinctly
concave; snout moderately
long, its length about 1.8 to 2.4
times in head length, meas-
ured without the lip the snout is
0.8 to 1 times in cheek height,
its dorsal profile nearly straight
or concave, snout angle rela-
tive to upper jaw between 50°
and 68°; interorbital space
usually convex; posterior nos-
tril an oblong longitudinal
opening, closer to orbit than
anterior nostril or, about half-
way between orbit and anterior
nostril; eye not close to dorsal juvenile
profile except in small individu-
als, its length 3.6 to 5.9 times
in head length; cheek moderately high, its height 2.4 to 3 times in head length; lateral teeth in jaws rounded
with points or molars that often have tubercles; outer surface of maxilla smooth or with a longitudinal ridge.
Dorsal fin with X spines and 9 soft rays, the fourth or fifth dorsal-fin spine usually the longest, its length 2.7
to 3.6 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray usually the longest, its
length almost equal to or slightly shorter than length of base of soft-rayed portion of anal fin and 1.3 to 1.6
times in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest
to body usually with dense melanophores. Lateral-line scales 46 to 48; cheek without scales; 5 ½ scale
rows between lateral line and base of middle dorsal-fin spines; 16 or 17 scale rows in transverse series
between origin of anal fin and lateral line; usually 15 rows in lower series of scales around caudal peduncle;
posterior angle of operculum fully scaly. Colour: body yellowish or bronze, lighter below, centres of many
scales with a white or light blue spot, sometimes irregular dark indistinct bars on sides and a square black
blotch above pectoral fins bordering below lateral line; 3 blue streaks or series of blue spots radiating forward
and ventrally from eye; fins whitish or yellowish, pelvic fins dusky, edge of dorsal fin reddish.
Size: Maximum total length about 80 cm; commonly between 20 and 50 cm.
Habitat, biology, and fisheries: On nearshore and offshore coral reefs, coralline lagoons, seagrass beds,
mangrove swamps, and coastal sand and rock areas, to depths of 75 m. Adults alone or in small schools;
juveniles form large schools. Feeds mostly on echinoderms, molluscs, crustaceans, and to a lesser extent
on polychaetes and fishes.
Taken by handline, traps,
trawls, seines, and gill nets.
Marketed mostly fresh. Very
important commercial and
sport fish in some countries.
Indo-West Pacific including
Red Sea and Persian Gulf,
East Africa to southern Japan
and Samoa.
3038 Bony Fishes
Lethrinus obsoletus (Forsskål, 1775) (Plate XVIII, 137)

Frequent synonyms / misidentifications: Sciaena ramak (Forsskål, 1775) / None.
FAO names: En - Orangestriped emperor; Fr - Empereur bandes oranges; Sp - Emperador de bandes.
1 to 1.1 times in body depth, 2.6 to 2.9 times in standard length, dorsal profile near eye nearly straight or
slightly convex; snout length about 1.8 to 2.3 times in head length, measured without the lip the snout is 0.8
to 0.9 times in cheek height, its dorsal profile slightly concave, snout angle relative to upper jaw between
50° and 60°; interorbital space convex; posterior nostril a longitudinal oblong opening, closer to orbit than
anterior nostril; eye situated close to or not close to dorsal profile, its length 3.5 to 5 times in head length;
cheek height 2.5 to 3.2 times in head length; lateral teeth in jaws conical or rounded; outer surface of maxilla
with a distinct knob. Dorsal fin with X spines and 9 soft rays, the fourth or fifth dorsal-fin spine the longest,
its length 2.5 to 3.4 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray usually the
longest, its length almost equal to or slightly longer or shorter than length of base of soft-rayed portion of
between rays closest to body without dense melanophores. Lateral-line scales 45 to 48; cheek without
scales; 5 ½ scale rows between lateral line and base of middle dorsal-fin spines; 15 or 16 scale rows
in transverse series between origin of anal fin and lateral line; usually 15 rows in lower series of scales
around caudal peduncle; usually 4 to 9 scales in supratemporal patch; inner surface of pectoral-fin base
densely covered with scales; posterior angle of operculum fully scaly. Colour: body light tan to brown,
lighter below; centres of scales often lighter than background colour; an orange-yellow stripe on lower side
at level of pectoral-fin base, with 2 additional more faint orange-yellow stripes above and 1 below this stripe;
head often with several broad indistinct vertical and diagonal light and dark bands; sometimes white spots
below eye; posterior edge of operculum dark brown; fins whitish or tan, sometimes mottled.
Size: Maximum total length reported as 60 cm; commonly between 20 and 30 cm.
Habitat, biology, and fisheries: Inhabits seagrass beds, and sand and rubble areas of lagoons and reefs
to depths of about 30 m. Feeds mostly on crustaceans, molluscs, and echinoderms. Caught mostly by traps
and shore seines, occasionally by gill nets, handlines, and trawls. Marketed mostly fresh.
Indo-West Pacific including
the Red Sea, East Africa to the
Ryukyu Islands, Tonga, and
Samoa.
Lethrinus olivaceus Valenciennes, 1830 (Plate XIX, 138)

Frequent synonyms / misidentifications: None / Lethrinus elongatus non Valenciennes, 1830; L. miniatus
(non Schneider, 1801).
FAO names: En - Longface emperor; Fr - Empereur guele longue; Sp - Emperador trompudo.
Diagnostic characters: Body moderately slender, its depth 2.9 to 3.4 times in standard length. Head
length 1.1 to 1.3 times in body depth, 2.4 to 2.9 times in standard length, dorsal profile near eye nearly
straight or with small bump; snout length about 1.7 to 2.2 times in head length, measured without the lip
the snout is 0.6 to 0.8 times in cheek height, its dorsal profile slightly concave, snout angle relative to
upper jaw between 40° and 50°; interorbital space convex to flat; posterior nostril a longitudinal oblong
opening, closer to orbit than anterior nostril; eye situated close to or not close to dorsal profile, its length 3.4
to 6.2 times in head length; cheek height 3 to 3.8 times in head length; lateral teeth in jaws conical; outer
surface of maxilla smooth. Dorsal fin with X spines and 9 soft rays, the third or fourth dorsal-fin spine the
longest, its length 2.4 to 2.8 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray
usually the longest, its length almost equal to or slightly shorter than length of base of soft-rayed portion of
between rays closest to body with dense melanophores. Lateral-line scales 46 to 48; cheek without scales;
5 ½ scale rows between lateral line and base of middle dorsal-fin spines; usually 16 or 17 scale rows
in transverse series between origin of anal fin and lateral line; 15 rows in lower series of scales around caudal
peduncle; 6 to 9 scales in supratemporal patch; inner surface of pectoral-fin base without scales;
posterior angle of operculum fully scaly. Colour: body grey, lighter ventrally, often with scattered irregular
dark blotches; snout with wavy dark streaks, upper jaw, especially near corner of mouth sometimes edged
behind with red.
Habitat, biology, and fisheries: Inhabits sandy coastal areas, lagoons, and reef slopes, occurring to depths
of 185 m; juveniles found in shallow sandy areas. Often occurs in large schools. Feeds mostly on fishes,
crustaceans, and cephalopods. Caught mostly with handline and traps, occasionally by trawls and gill nets.
Large individuals often ciguatoxic in New Caledonia and possibly elsewhere in Oceania. Very important in
Palau where they are report-
edly overfished. In both Palau
and Tahiti this species is ac-
tively fished in locations and
times known to have large
spawning aggregations.
the Red Sea, East Africa to the
Ryukyu Islands, to Samoa and
Polynesia.
3040 Bony Fishes
Lethrinus ornatus Valenciennes, 1830 (Plate XIX, 139)

FAO names: En - Ornate emperor; Fr - Empereur orn; Sp - Emperador ornado.
Diagnostic characters: Body relatively deep, its depth 2.3 to 2.6 times in standard length. Head length 0.8
to 0.9 times in body depth, 2.7 to 3 times in standard length, dorsal profile near eye convex; snout length
about 2 to 2.5 times in head length, measured without the lip the snout is 0.9 to 1.1 times in cheek height,
its dorsal profile nearly straight or slightly concave, snout angle relative to upper jaw between 65° and
75°; interorbital space convex; posterior nostril a longitudinal oblong opening, closer to orbit than anterior
nostril; eye situated close to dorsal profile, its length 3.3 to 4.1 times in head length; cheek height 2.2 to 2.8
times in head length; lateral teeth in jaws rounded with points or molars; outer surface of maxilla usually
smooth, sometimes with a longitudinal ridge. Dorsal fin with X spines and 9 soft rays, the fourth or fifth
dorsal-fin spine the longest, its length 2.7 to 3.3 times in body depth; anal fin with III spines and 8 soft rays,
the first soft ray usually the longest, its length longer than length of base of soft-rayed portion of anal fin and
1.1 to 1.5 times in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays
closest to body without dense melanophores. Lateral-line scales usually 46 or 47; cheek without scales;
5 ½ scale rows between lateral line and base of middle dorsal-fin spines; 15 or 16 scale rows in
transverse series between origin of anal fin and lateral line; 13 to 15 rows in lower series of scales around
caudal peduncle; usually 3 to 8 scales in supratemporal patch; inner surface of pectoral-fin base densely
covered with scales; posterior angle of operculum fully scaly. Colour: body dusky whitish, lighter below,
with 5 or 6 orange stripes; posterior edge of opercle and preopercle bright red (the former more
conspicuous); head brown or tan, sometimes a red spot on lower front edge of eye; pectoral fins orangish;
pelvic and anal fins, and most of dorsal fin whitish; edge of dorsal and caudal fins reddish.
Habitat, biology, and fisheries: Inhabits sandy and soft bottom inshore bays, seagrass beds, lagoons, and
areas adjacent to reefs. Feeds on crustaceans, molluscs, echinoderms, polychaetes, and small fishes.
Caught by trap, handline,
shore seine, trawl, and han-
dline. Minor importance in
fisheries where it occurs.
Distribution: East Indian
Ocean and West Pacific, from
Sri Lanka to the Ryukyu Is-
lands, Papua New Guinea,
and northeastern Australia.
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Lethrinus reticulatus Valenciennes, 1830

FAO names: En - Redsnout emperor.
length 1.1 to 1.2 times in body depth, 2.5 to 2.8 times in standard length, dorsal profile near eye convex
or nearly straight; snout length about 1.9 to 2.4 times in head length, measured without the lip the snout is
0.8 to 0.9 times in cheek height, its dorsal profile concave, snout angle relative to upper jaw between 50°
and 60°; interorbital space flat or concave; posterior nostril a longitudinal oblong opening, closer to orbit
than anterior nostril; eye situated close to dorsal profile, its length 3.3 to 4.3 times in head length; cheek
height 2.7 to 3.4 times in head length; lateral teeth in jaws conical; outer surface of maxilla usually smooth.
Dorsal fin with X spines and 9 soft rays, the third dorsal-fin spine the longest, its length 2 to 2.8 times in body
depth; anal fin with III spines and 8 soft rays, the first soft ray usually the longest, its length almost equal to,
shorter, or slightly longer than length of base of soft-rayed portion of anal fin and 1.4 to 1.8 times in length
of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body without
dense melanophores. Lateral-line scales 46 to 48; cheek without scales; 4 ½ scale rows between lateral
line and base of middle dorsal-fin spines; 15 or 16 scale rows in transverse series between origin of anal
fin and lateral line; usually 15 rows in lower series of scales around caudal peduncle; 7 to 10 scales in
supratemporal patch; inner surface of pectoral-fin base without scales; posterior angle of operculum
fully scaly. Colour: body olive grey or tan, often with scattered irregular black blotches; base of pectoral,
upper edge of opercle, and sometimes posterior edge of preopercle red; head brown or olive with a somewhat
indistinct reddish band on snout, originating midway between eye and snout and terminating on tip of snout;
lips red; fins pale or orangish.
Habitat, biology, and fisheries: Nothing specific is reported on the habitat or biology of this species. It is
thought to occur on soft bottoms near reefs. The diet is presumably benthic invertebrates and fishes. Caught
mostly by trawl and handline.
Marketed fresh.
Distribution: Mauritius,
Chagos, southern Japan, the
Philippines to Irian Jaya, and
Samoa.
3042 Bony Fishes
Lethrinus rubrioperculatus Sato, 1978 (Plate XIX, 140)

Frequent synonyms / misidentifications: None / Lethrinus variegatus non Valenciennes, 1830.
FAO names: En - Spotcheek emperor; Fr - Empereur honteux; Sp - Emperador maquillado.
Diagnostic characters: Body moderately elongate, its depth 3 to 3.4 times in standard length. Head
length 1.1 to 1.3 times in body depth, 2.6 to 3 times in standard length, dorsal profile near eye convex
without a distinct hump; snout length 1.8 to 2.1 times in head length, measured without the lip the
snout is 0.7 to 0.9 times in cheek height, its dorsal profile nearly straight or slightly concave, snout
angle relative to upper jaw between 54° and 65°; interorbital space flat or slightly convex; posterior nostril
a longitudinal oblong opening, closer to orbit than anterior nostril; eye situated close or slightly removed from
dorsal profile, its length 3.9 to 4.7 times in head length; cheek height 2.4 to 3.3 times in head length; lateral
teeth in jaws conical; outer surface of maxilla smooth or with a longitudinal ridge. Dorsal fin with X spines
and 9 soft rays, the third dorsal-fin spine the longest, its length 2.4 to 2.7 times in body depth; anal fin with
III spines and 8 soft rays, the first soft ray usually the longest, its length shorter than length of base of
soft-rayed portion of anal fin and 1.4 to 1.8 times in length of entire anal-fin base; pectoral-fin rays 13;
pelvic-fin membranes between rays closest to body without dense melanophores. Lateral-line scales
47 to 49; cheek without scales; 4 ½ scale rows between lateral line and base of middle dorsal-fin
spines; 15 or 16 scale rows in transverse series between origin of anal fin and lateral line; usually 15
rows in lower series of scales around caudal peduncle; 7 to 10 scales in supratemporal patch; inner surface
of pectoral-fin base without scales; posterior angle of operculum with a wide scaleless area. Colour:
body olive grey or brown, with scattered irregular small black blotches; lips and a spot on upper edge of
operculum usually red; fins pale or pinkish.
Size: Maximum reported total length 50 cm; commonly to 30 cm.
Habitat, biology, and fisheries: Inhabits sand and rubble areas of outer reef slopes to depths of 160 m.
Feeds mostly on crustaceans, fishes, echinoderms, and molluscs. Caught mostly by handline, trap, and
trawl. Marketed mostly fresh. An excellent food fish. One of the most abundant species taken in bottom
fisheries in the Marianas.
East Africa to southern Japan
and the Marquesas.
Lethrinus semicinctus Valenciennes, 1830 (Plate XIX, 141)

FAO names: En - Blackblotch emperor; Fr - Empereur maill; Sp - Emperador malla.
but without a distinct hump; snout length about 1.9 to 2.1 times in head length, measured without the
lip the snout is 0.7 to 0.9 times in cheek height, its dorsal profile nearly straight, snout angle relative
to upper jaw between 55° and 67°; interorbital space flat or slightly convex; posterior nostril a longitudinal
oblong opening, closer to orbit than anterior nostril; eye situated close to dorsal profile , its length 3.6 to 4.3
times in head height; cheek length 2.5 to 3.0 times in head length; lateral teeth in jaws conical; outer
surface of maxilla smooth or with a longitudinal ridge. Dorsal fin with X spines and 9 soft rays, the third or
fourth dorsal-fin spine the longest, its length 2.3 to 2.9 times in body depth; anal fin with III spines and 8 soft
rays, the first soft ray usually the longest, its length aproximately equal to or shorter than length of base of
soft-rayed portion of anal fin and 1.3 to 1.7 times in length of entire anal-fin base; pectoral-fin rays 13;
pelvic-fin membranes between rays closest to body without dense melanophores. Lateral-line scales
46 to 48; cheek without scales; 4 ½ scale rows between lateral line and base of middle dorsal-fin
spines; usually 14 or 15 scale rows in transverse series between origin of anal fin and lateral line;
usually 15 rows in lower series of scales around caudal peduncle; 4 to 7 scales in supratemporal patch;
inner surface of pectoral-fin base without scales; posterior angle of operculum fully scaly. Colour:
body brown or tan, with scattered irregular small black blotches, a large oblong black blotch below
soft-rayed portion of dorsal fin and bordering below lateral line; fins pale or pinkish.
Habitat, biology, and fisheries: Inhabits shallow seagrass beds, reef flats, lagoons, and sandy areas near
coral reefs. Feeds on benthic invertebrates and small fishes. Caught by shore seine, trawl, trap, and handline.
Marketed fresh. Minor impor-
tance to fisheries.
Distribution: E a s t In di a n
Ocean and West Pacific, in-
cluding Sri Lanka, Indonesia,
northern Australia, the Ryukyu
Islands to the Marshall and
Solomon Islands.
3044 Bony Fishes
Lethrinus variegatus Valenciennes, 1830 (Plate XIX, 142)

Frequent synonyms / misidentifications: None / Lethrinus semicinctus non Valenciennes, 1830;
L. reticulatus non Valenciennes, 1830.
FAO names: En - Slender emperor; Fr - Empereur bas cou; Sp - Emperador moteado.
Diagnostic characters: Body slender, its depth 3.2 to 3.9 times in standard length. Head length 1.1
to 1.5 times in body depth, 2.6 to 2.9 times in standard length, dorsal profile near eye convex or nearly
straight; snout length 2.2 to 2.7 times in head length, measured without the lip the snout is 0.8 to 1 times
in cheek height, its dorsal profile nearly straight, snout angle relative to upper jaw between 50° and 65°;
interorbital space flat or convex; posterior nostril a vertical oblong opening, closer to anterior nostril than
orbit, or closer to orbit than anterior nostril; eye situated close to dorsal profile, its length 3.2 to 4.4 times in
head length; cheek height 2.9 to 3.8 times in head length; lateral teeth in jaws conical; outer surface of
maxilla smooth. Dorsal fin with X spines and 9 soft rays, the fourth dorsal-fin spine usually the longest, its
length 1.9 to 2.5 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray usually the
longest, its length aproximately equal to or shorter than length of base of soft-rayed portion of anal fin and
1.3 to 1.6 times in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays
closest to body without dense melanophores. Lateral-line scales 45 to 47; cheek without scales; 4 ½ scale
rows between lateral line and base of middle dorsal-fin spines; 13 or 14 scale rows in transverse
series between origin of anal fin and lateral line; 14 or 15 rows in lower series of scales around caudal
posterior angle of operculum fully scaly. Colour: body brown and grey, lighter ventrally, with scattered
irregular dark spots; often 2 dark bands below eye, 1 to lower rim of preopercle and 1 to corner of mouth,
another dark band across interorbital space; dorsal, anal, pectoral, and pelvic fins light or translucent; caudal
fin light and dark striped.
Habitat, biology, and fisheries: Inhabits sandy areas near coral reefs. Feeds on small benthic inverte-
brates. Caught mostly by shore seine and trawl. Of minor importance to fisheries where it occurs.
the Indo-West Pacific, includ-
ing the Red Sea, East Africa,
Sri Lanka, Chagos Archipel-
ago, Malaysia and Indonesia
to the Ryukyu Islands and
Palau, and to northwestern
Australia and New Caledonia.
Lethrinus xanthochilus Klunzinger, 1870 (Plate XIX, 143)

FAO names: En - Yellowlip emperor; Fr - Empereur bec de cane; Sp - Emperador trompa amarilla.
length 1 to 1.2 times in body depth, 2.6 to 3 times in standard length, dorsal profile near eye convex; snout
length about 1.7 to 2.5 times in head length, measured without the lip the snout is 0.8 to 0.9 times in
cheek height, its dorsal profile nearly straight, snout angle relative to upper jaw between 45° and 60°;
interorbital space distinctly concave; posterior nostril a longitudinal or vertical oblong opening, closer to
orbit than anterior nostril; eye situated close to dorsal profile, its length 2.8 to 5.1 times in head length; cheek
height 2.3 to 3.4 times in head length; lateral teeth in jaws conical; outer surface of maxilla smooth. Dorsal
fin with X spines and 9 soft rays, the third dorsal-fin spine the longest, its length 2 to 2.8 times in body depth;
anal fin with III spines and 8 soft rays, the first soft ray usually the longest, its length aproximately equal to
or shorter than length of base of soft-rayed portion of anal fin and 1.3 to 1.7 times in length of entire anal-fin
base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body with dense melanopho-
res. Lateral-line scales 47 or 48; cheek without scales; 4 ½ scale rows between lateral line and base
of middle dorsal-fin spines; 15 or 16 scale rows in transverse series between origin of anal fin and lateral
line; usually 15 rows in lower series of scales around caudal peduncle; 5 to 8 scales in supratemporal patch;
inner surface of pectoral fins without scales; posterior angle of operculum fully scaly. Colour: body yellowish
grey with scattered irregular dark spots; lips yellowish, upper lip more intense; a red spot at upper base of
pectoral fins; fins bluish grey and mottled, bases of fins lighter and edges of dorsal and caudal fins reddish.
Habitat, biology, and fisheries: Inhabits seagrass beds, sand and rubble areas of coral reefs, deep
channels, and lagoons. Often found in shallow water but recorded to depths of 150 m. Occurs usually in
small groups. Feeds primarily on crustaceans, fishes, and echinoderms. Caught mostly with handline, traps,
and trawl.
the Indo-West Pacific, includ-
ing the Red Sea, East Africa,
Central Indian Ocean, Indone-
sia to the Ryukyu Islands,
south to Queensland, and the
Caroline Islands to the Mar-
quesas.
3046 Bony Fishes
Lethrinus sp. 1
FAO names: En - Maldive emperor.
Diagnostic characters: Body moderately elongate, its depth 3 to 3.3 times in standard length. Head
with a hump in front of eye; snout length about 1.8 to 2 times in head length, measured without the lip
the snout is 0.8 to 0.9 times in cheek height, its dorsal profile concave, snout angle relative to upper
jaw between 50° and 60°; interorbital space flat or convex; posterior nostril a longitudinal oblong opening,
closer to orbit than anterior nostril; eye situated close to or not close to dorsal profile, its length 4.2 to 4.7
times in head length; cheek height 2.4 to 2.7 times in head length; lateral teeth in jaws conical; outer surface
of maxilla smooth. Dorsal fin with X spines and 9 soft rays, the third dorsal-fin spine the longest, its length
2.5 to 2.8 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray usually the longest,
its length approximately shorter than length of base of soft-rayed portion of anal fin and about 1.6 to 1.8
times in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to
body without dense melanophores. Lateral-line scales 47 or 48; cheek without scales; 4 ½ scale rows
between lateral line and base of middle dorsal-fin spines; usually 15 scale rows in transverse series
between origin of anal fin and lateral line; usually 15 rows in lower series of scales around caudal
posterior angle of operculum fully scaly. Colour: body light grey or tan, head darker; base of pectoral fins,
spot on upper posterior margin of opercle, posterior edge of preopercle, and indistinct area on snout
in front of eye usually reddish; fins pale or reddish.
Size: Maximum recorded total length about 32 cm, but presumably attains a much larger size.
Habitat, biology, and fisheries: Inhabits sandy areas near coral reefs. Presumably feeds mostly on
crustaceans and fishes. Occasionally caught by handline in the Maldives and by trawl in Indonesia.
Distribution: The Maldives
and southwestern Indonesia.
Lethrinus sp. 2
FAO names: En - Drab emperor.
length 1 to 1.2 times in body depth, 2.7 to 2.9 times in standard length, dorsal profile near eye convex,
without a distinct hump; snout length about 1.9 to 2 times in head length, measured without the lip the
snout is 0.8 to 0.9 times in cheek height, its dorsal profile nearly straight, snout angle relative to upper
jaw between 60° and 70°; interorbital space flat or convex; posterior nostril a longitudinal oblong opening,
closer to orbit than anterior nostril; eye situated close to dorsal profile, its length 3.6 to 4.1 times in head
length; cheek height 2.4 to 2.9 times in head length; lateral teeth in jaws conical; outer surface of
maxilla smooth or with a longitudinal ridge. Dorsal fin with X spines and 9 soft rays, the third dorsal-fin spine
the longest, its length 2.4 to 2.6 times in body depth; anal fin with III spines and 8 soft rays, the first soft ray
usually the longest, its length shorter than length of base of soft-rayed portion of anal fin and 1.5 to 1.7 times
in length of entire anal-fin base; pectoral-fin rays 13; pelvic-fin membranes between rays closest to body
without dense melanophores. Lateral-line scales 48 or 49; cheek without scales; 4 ½ scale rows
between lateral line and base of middle dorsal-fin spines; 15 or 16 scale rows in transverse series
between origin of anal fin and lateral line; usually 15 rows in lower series of scales around caudal
posterior angle of operculum fully scaly. Colour: body light grey or tan with scattered irregular dark blotches;
head brown or tan; fins pale.
Size: Maximum recorded total length about 27 cm, but probably attains a larger size.
Habitat, biology, and fisheries: Very little is recorded about the biology of this species. In the Philippines
it is caught together with smaller specimens of such species as Lethrinus olivaceus, L. reticulatus, and L.
semicinctus; it presumably has habitat characteristics similiar to these species. Caught by shore seine and
trawl.
Distribution: So far known
only from the Philippines, the
Ryukyu Islands, northwestern
to northeastern Australia,
New Caledonia, and the Loy-
alty Islands.
3048 Bony Fishes
Lethrinus sp. 3
Frequent synonyms / misidentifications: None / Lethrinus laticaudis non Alleyne and Macleay, 1877; L.
lentjan (non Lacepède, 1802); L. nebulosus (non Forsskål, 1775); L. punctulatus Macleay, 1878.
FAO names: En - Lesser spangled emperor.
Diagnostic characters: Body fairly deep, its depth 2.3 to 2.5 times in standard length. Head length 0.8
to 0.9 times in body depth, 2.6 to 2.9 times in standard length; dorsal profile near eye nearly straight; snout
moderately short, its length about 1.8 to 2 times in head length, measured without the lip the snout is 0.9
to 1 times in cheek height, its dorsal profile nearly straight to concave, snout angle relative to upper jaw
between 59° and 66°; interorbital space convex; posterior nostril an oblong longitudinal opening, closer to
orbit than anterior nostril; eye situated close to or not close to the dorsal profile, its length 3.7 to 5 times in
head length; cheek height 2 to 2.6 times in head length; lateral teeth in jaws conical, sometimes rounded
at base but still with conical tips; outer surface of maxilla smooth or with a longitudinal ridge. Dorsal
fin with X spines and 9 soft rays, the fourth or fifth dorsal-fin spine usually the longest, its length 3.1 to 3.8
times in body depth; anal fin with III spines and 8 soft rays, the first soft ray usually the longest, its length
almost equal to or shorter than length of base of soft-rayed portion of anal fin; pectoral-fin rays 13; pelvic-fin
membranes between rays closest to body without dense melanophores. Lateral-line scales usually 46 or
47; cheek without scales; 5 ½ scale rows between lateral line and base of middle dorsal-fin spines;
usually 16 or 17 scale rows in transverse series between origin of anal fin and lateral line; usually 15 rows
in lower series of scales around caudal peduncle; 5 to 8 scales in supratemporal patch; inner surface of
pectoral-fin base with usually less than 50% of lower part of base densely covered with scales, with
a few scales, or naked; posterior angle of operculum fully scaly. Colour: body and head brownish bronze
to yellowish, grading to light bluish on sides then white or just white ventrally, centres of some scales on
upper sides pearly white to light bluish, often a series of 9 or 10 broken indistinct bars on sides and caudal
peduncle; head with bluish or whitish streaks and spots around eye and on snout and cheeks, 1 distinct
streak emanating from eye and passing through or near nostrils and 2 less distinct streaks, sometimes
broken into spots, emanating from eye more ventrally; a broad, indistinct bar on both the preopercle and
opercle; fins hyaline to light brownish or yellowish, sometimes a tinge of reddish orange on fringe of soft
dorsal and caudal fins, upper rays of pectoral fins and outer rays of pelvic fins bright white to light bluish.
Size: Maximum total length about 40 cm; commonly to about 30 cm.
Habitat, biology, and fisheries: On coral
reefs and lagoons and adjacent sandy ar-
eas. Caught mostly by anglers.
Distribution: Apparently restricted to
Western Australia from around Exmouth
Gulf to Dampier Land and possibly to Dar-
win.
Remarks: This species may be unde-
scribed or it may turn out to be Lethrinus
punctulatus Macleay, 1878.
Monotaxis grandoculis (Forsskål, 1775) (Plate XIX, 144)

FAO names: En - Humpnose bigeye bream.
adult
Diagnostic characters: Body oblong, its depth 2.1 to 2 7 times

in standard length. Head profile strongly convex in front of eye,
the snout sloping steeply; eye large, its diameter about equal to
length of snout or 2.5 to 3.2 times in head length; interorbital
space wide and flat; hind margin of preopercle finely serrated;
posteriormost part of jaws reaching beyond vertical through
anterior margin of eye; a broad patch of small teeth in jaws
anteriorly, followed by a series of 6 or 7 round, flat molars
on each side and preceded by canines in front of each jaw; subadult
maxilla with a longitudinal denticulated ridge. Dorsal fin
with X slender spines and 10 soft rays, the first spine half the
length of second, otherwise remaining spines about equal; anal juvenile
fin with III spines and 9 soft rays; pectoral-fin rays 14 including
splint-like uppermost ray; caudal fin forked with pointed tips. Lateral-line scales 44 to 47, about 4 or 5
horizontal scale rows above lateral line; cheek with scales; inner surface of pectoral-fin base densely
scaly. Colour: ground colour bluish grey grading to whitish on ventral parts; lips yellow to pinkish; area
around eye often yellow or orange; fins without distinctive markings, the membranes clear or dusky, but
often reddish to yellow-orange; base of upper pectoral-fin rays and inner base of pectoral-fin axil black;
caudal fin frequently with blackish rays contrasted against paler membranes. Small juveniles whitish on
lower half with 3 prominent black saddles separated by narrower white bars on upper half; a prominent black
bar through eye; fins clear to white except the dorsal fin includes the continuation of 2 of the black saddles
of the upper side and the outer edges of the caudal-fin lobes yellow-brown. The juvenile pattern of bars
sometimes in relatively large (20 to 30 cm) individuals. Largest adults, although usually seen without
markings, are able to rapidly assume dark bars or saddles.
Habitat, biology, and fisheries: Usually on coral reefs, often on the sandy periphery or in sand and rubble
areas. Solitary fishes are often encountered, but large adults frequently form aggregations of up to about 50
individuals. The depth range is between about 3 and 60 m, although it appears to be most common between
about 5 and 30 m. Consumes mainly gastropod molluscs, ophiuroids, and echinoids, but also pagurid and
brachyuran crabs, polychaetes, tunicates,
and holothurians. Caught mainly with gill
nets, traps, spears, and handlines. Mar-
keted fresh. Sometimes ciguatoxic in the
Marshall Islands.
Distribution: Widespread in the Indo-West
and Central Pacific from Hawaii and south-
eastern Oceania to the east coast of Africa
and Red Sea and from Australia northwards
to Japan.
3050 Bony Fishes
Wattsia mossambica (Smith, 1957) (Plate XIX, 145)

FAO names: En - Mozambique large-eye bream.
Diagnostic characters: Body relatively deep, roughly rhomboid, its depth 1.9 to 2.2 times in standard length.
Head profile slightly convex; eye moderate in size, its diameter much shorter than snout length; jaw teeth
in a narrow, villiform band bordered by an outer series of conical teeth; anteriorly 4 moderate-sized
canines in upper jaw and 6 in lower jaw; maxilla with a longitudinal denticulated ridge; a pair of close-set
nasal openings, the posterior one slit-like, ending close to eye. Dorsal fin with X long, strong spines and 10
soft rays; first spine about 2/3 length of second, the second spine slightly longer than eye diameter, the fourth
to sixth spines longest; anal fin with III strong spines and 10 soft rays; third spine twice as long as eye
diameter; anal-fin base 1.4 to 1.6 times longer than longest anal-fin ray; caudal fin slightly forked with
broadly rounded lobes. Lateral-line scales 41 to 47; 5 scale rows between lateral line and base of middle
dorsal-fin spines; cheek with scales; inner surface of pectoral-fin axil scaleless. Colour: overall silvery
grey with yellowish suffusion; scale margins narrowly brownish; indistinct dark blotches or bars sometimes
apparent on body; lips whitish to yellow; fins yellow, faint brown spotting may be present on soft dorsal, anal,
and caudal fins; a narrow blackish bar across base of pectoral fins.
Habitat, biology, and fisheries: Inhabits the outer edge of the continental shelf at depths between 100 and
180 m. It feeds on bottom-liv-
ing invertebrates and small
fishes. Occasionally seen in
markets, mainly fresh. Caught
mainly with bottom longlines
and bottom trawls.
Distribution: Tropical Indian
Ocean and West Pacific from
Mozambique to southern Ja-
pan, the Marshall Islands,
northwestern Australia, and to
Queensland.
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Perciformes: Percoidei: Nemipteridae 3051
NEMIPTERIDAE
Threadfin breams (also whiptail breams, monocle breams,
dwarf monocle breams, and coral breams)
by B.C. Russell
D iagnostic characters: Elongate to moderately deep, compressed, small to medium-sized (to about
35 cm) sparoid fishes. Mouth terminal, small to moderate; premaxillaries moderately protrusible; teeth
in jaws conical, enlarged canines present anteriorly in Nemipterus and Pentapodus; vomer and palatine
without teeth. A single continuous dorsal fin, with X spines and 9 soft rays; anal fin with III spines
and 7 (except Nemipterus virgatus with 8) soft rays; caudal fin emarginate, forked, lunate or falcate; the
upper and/or lower tips of the fin with or without pointed or filamentous extensions; pectoral fins with 2
unbranched and 12 to 17 branched rays; pelvic fins thoracic, with I spine and 5 soft rays. Scales finely
ctenoid and moderate in size. Ascending premaxillary process never as long as the alveolar ramus; articular
process not fused to ascending process along anterior margin; postmaxillary process present, low,
broad-based; palato-premaxillary ligament well developed; ethmo-maxillary ligament well developed,
Y-shaped, with a ventrolateral branch inserting on the palatine anterior to the insertion of the palato-pre-
maxillary ligament. Gill membranes free from the isthmus; gill arches 4, a slit behind the fourth; pseudo-
branch well developed; gill rakers short, knob-like; 6 branchiostegal rays, the first 5 inserting on the
ceratohyal, the sixth inserting at the interspace between the ceratohyal and the epihyal; second and third
epibranchial tooth plates lacking. First and second infraorbitals deep; the third infraorbital never deep,
and the second infraorbital typically projecting backward below it (this posterior projection devel-
oped as an externally visible spine in Scolopsis); third infraorbital with a well-developed subocular shelf;
fourth infraorbital often with a very small shelf. Two predorsal bones present in the following configuration:
first predorsal, first neural spine, second predorsal, second neural spine, first pterygiophore supporting the
first 2 dorsal-fin spines plus the second pterygiophore supporting the third dorsal-fin spine, third neural
spine, third pterygiophore supporting the fourth dorsal-fin spine, fourth neural spine; the last pterygiophore
of the dorsal and anal fins trisegmental. Epipleural ribs 8 to 12; accessory subpelvic keel and post pelvic
process present; opisthotic well developed. Colour: extremely variable, but often pinkish or reddish with
red, yellow, or blue markings.
a single dorsal fin with
X spines and 9 soft rays
caudal fin
often with
filamentous
extension(s)

and usually 7 soft rays
Habitat, biology, and fisheries: Marine, bottom-living fishes. Species of the genus Nemipterus occur on
mud and sand bottoms in coastal inshore as well as offshore shelf waters and range in depths to about
300 m, although most species occur in much shallower water. Species of the genus Parascolopsis occur
on mud or sand bottoms mainly in offshore shelf waters in depths to about 400 m. Species of the genus
Pentapodus are benthic or free-swimming near the bottom, and usually occur on or close to coral reefs in
depths to 100 m. The monotypic Scaevius milii occurs on reef and shallow muddy or sand bottoms in inshore
areas. Species of the genus Scolopsis occur usually on coral reefs, or on sand or mud bottoms close to
reefs, in depths to about 60 m. Nemipterids can be solitary or schooling and do not appear to be territorial.
Carnivorous and feed mainly on other small fishes, cephalopods, crustaceans, and polychaetes. Many
nemipterids show size-related differences in sex ratios, with small fishes being mainly females and larger
fishes males. In some species this size-related skew in sex ratios appears to be due to higher growth rates
in males. However, some species of Pentapodus, Scolopsis, and Nemipterus appear to be protogynous
hermaphrodites. Spawning seasonality varies widely among different species and appears to vary also
3052 Bony Fishes
between localities. Mature ova are present in species of Nemipterus over a prolonged period, inferring a
protracted spawning season, with usually 1 or 2 periods of increased fecundity. Threadfin breams, and to a
much lesser extent monocle breams, are an important component of commercial and artisanal fisheries of
the Indo-West Pacific region. Whiptail breams are of artisanal fisheries importance in some areas and are
also taken occasionally by recreational fishermen. Dwarf monocle breams, because of their usually small
size and deeper-water habits are of little fisheries interest. Caught mainly by bottom trawl or by handline.
Other methods include longline, gill nets, lift nets, surrounding nets, drive-in nets, fish stakes, and traps.
Some species of Scolops is also are captured live by hand net for the aquarium trade. Nemipterids are
popular eating fishes and are marketed fresh, dry-salted, dry-smoked, fermented, and steamed. Trash fish
are made into fish balls, fish cakes, fish meal and surimi, or used as animal feed. In many areas, species of
Nemipterus are the second or third-most important catch species by weight. For 1995, the FAO Yearbook
of Fishery Statistics reports a total catch of around 134 200 t of nemipterids from the Western Central Pacific.
However, because of the largely artisanal nature of the fisheries of some areas and the paucity of landing
statistics, the available figures probably grossly underestimate the actual catch of nemipterids taken in the
area.
Similar families occuring in the area

Lutjanidae: dorsal fin with X to XII spines and 10 to 17 soft rays; anal fin with III spines and 7 to 11 soft rays;
teeth usually present on vomer and palatine.
Caesionidae: mouth small, protractile; dorsal fin with X to XV spines and 8 to 22 soft rays; anal fin with III
spines and 9 to 13 soft rays.
Lutjanidae Caesionidae
Lethrinidae: anal fin with III spines and 8 to 10 soft rays; no cheek scales in Lethrinus.
Haemulidae: dorsal fin with IX to XIV spines and 11 to 26 soft rays; anal fin with III spines and 6 to 18 soft
rays; scales present between eye and mouth; pores present on chin.
Lethrinidae Haemulidae
Key to the species of Nemipteridae occurring in the area

1a. Suborbital naked, with a large backwardly pointing spine and a series of smaller spines
or serrations on its posterior margin; posterior margin of preopercle coarsely denticulate
or serrate (Fig. 1); canine teeth absent . . . . . . . . . . . . . . . . . . . . . . . . . (Scolopsis) ® 2
1b. Suborbital scaly or naked, spine weak or absent; posterior margin of suborbital smooth,
finely serrate, or with a few small denticulations; posterior margin of preopercle finely
denticulate or smooth (Fig. 2); canine teeth in jaws absent, or present only anteriorly . .
. . . . . . . . . . . . . . . . . . . . . . . (Scaevius, Nemipterus, Pentapodus, Parascolopsis) ® 13
2a. Small forward pointing spine or bony ridge present below eye (not always well developed
in juveniles) (Fig. 3a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Small forward pointing spine or bony ridge below eye absent (Fig. 3b) . . . . . . . . . . . . . . . ® 6
eye
preopercle
coarsely antrorse
denticulate spine or
or serrate bony
ridge suborbital
spine
posterior margin a) b)
of suborbital
Fig. 1 Scolopsis Fig. 2 Fig. 3 suborbital bone
3a. Maxilla denticulate on its external edge (Fig. 4a) . . . . . . . . . . . . . . . . . . . Scolopsis ciliatus
(Andaman Sea, West Pacific)
3b. Maxilla smooth along its external edge (Fig. 4b) . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Scales on top of head extending forward to between level of snout and anterior nostril
(Fig. 5a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
4b. Scales on top of head not or only just extending forward to level of posterior nostril
(Fig. 5b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scolopsis xenochrous
(West Pacific and East Indian Ocean: Solomon Islands to Maldives)
5a. Pectoral fins reaching to level of anus; anterior part of anal fin black; body depth 2.5 to
3 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scolopsis bilineatus
(West Pacific and East Indian Ocean; Fiji to Laccadive Islands)
5b. Pectoral fins not reaching to level of anus; anterior part of anal fin not black; body depth
2 to 2.6 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . Scolopsis vosmeri
(Indo-West Pacific)
6a. Temporal region naked (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . Scolopsis temporalis

(Solomon Islands, New Guinea, Sulawesi)
6b. Temporal region scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
naked naked temporal
area region
denticulate smooth
scaly
area scaly
area
a) b)
a) b)
Fig. 5 top of head, showing
Fig. 4 outer surface of maxillary forward extent of scales Fig. 6 extent of scales on head
3054 Bony Fishes
7a. Head scales reaching forward only to level of

midpupil (Fig. 7b) . . . . . . . . . . . . . Scolopsis trilineatus
(West Pacific: Samoa to Philippines)
7b. Head scales reaching forward to or in front of level naked
of anterior margin of eye (Fig. 7a) . . . . . . . . . . . . . ® 8 area
8a. Lateral-line scales 37 to 39 . . . . . . . Scolopsis margaritifer

(West Pacific)
8b. Lateral-line scales 42 to 48 . . . . . . . . . . . . . . . . . ® 9
scaly
area scaly
9a. Three scale rows between lateral line and dorsal- area
fin origin; colour pattern consisting of 3 irregular
dark stripes horizontally on upper half of body; a) b)
total pectoral-fin rays 16 . . . . . . . . . . . Scolopsis lineatus
(West Pacific and Cocos-Keeling Islands) Fig. 7 top of head, showing forward
9b. Four to 5 scale rows between lateral line and extent of scales
dorsal-fin origin; colour pattern not as above; total
pectoral-fin rays 16 to 18 . . . . . . . . . . . . . . . . . ® 10
10a. Head scales reaching forward to or nearly to level of posterior nostril . . . . . . . . . . . . . . ® 11

10b. Head scales not reaching forward to level of posterior nostril . . . . . . . . . . . . . . . . . . . ® 12
11a. Bony opercular ridge naked or nearly so (Fig. 8a); lower limb of preopercle naked or
nearly so; red spot at base of pectoral fins . . . . . . . . . . . . . . . . . . . Scolopsis taeniopterus
(West Pacific: northern Australia to Philippines)
11b. Bony opercular ridge scaly (Fig. 8b); lower limb of preopercle with 1 or 2 rows of scales;
no red spot at base of pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . Scolopsis affinis
(West Pacific: northern Australia to Philippines)
12a. Four transverse scale rows between lateral line and first dorsal-fin spine; patch of scales
on top of head truncated anteriorly (Fig. 9a) . . . . . . . . . . . . . . . . . . . . . Scolopsis auratus
(southern Indonesia, Sri Lanka, Maldives)
12b. Five or 6 transverse scale rows between lateral line and first dorsal-fin spine; scales on
top of head not truncated anteriorly (Fig. 9b) . . . . . . . . . . . . . . . . . Scolopsis monogramma
(West Pacific and East Indian Ocean)
13a. Scales on top of head not reaching to level of eyes; temporal parts of head naked
(Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scaevius milii
13b. Scales on top of head reaching forward to or in front of middle of eyes; temporal parts of
head scaly (Fig. 2) . . . . . . . . . . . . . . . . . . (Nemipterus, Pentapodus, Parascolopsis) ® 14
naked forward extent

area of head scales
bony opercular
ridge naked bony opercular
ridge scaly naked area
lower lower scaly

limb of scaly area
limb of area
preopercle preopercle
naked scaly
a) b) a) b)
Fig. 8 scales on preopercle and opercle Fig. 9 shape of anterior margin Fig. 10 Scaevius milii
of head scales
14a. Suborbital spine weak or ab-

sent; 4 to 6 transverse scale
rows on preopercle (Fig. 11) . .
. . . . . . (Pentapodus, Parascolopsis) ® 38
14b. Suborbital spine absent; 3
transverse scale rows on pre-
opercle (Fig. 12) . . . . . (Nemipterus) ® 15
15a. Anal fin with III spines and 8 4-6 rows of scales
3 rows of scales
soft rays . . . . . . . . Nemipterus virgatus
(West Pacific) Fig. 11 Pentapodus Fig. 12 Nemipterus
15b. Anal fin with III spines and 7
soft rays . . . . . . . . . . . . . . . . . ® 16
16a. First 2 dorsal-fin spines close together, almost fused, produced to form a very long
filament (Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus nematophorus
(East Indian Ocean and West Pacific from South China Sea to Indonesia)
16b. First 2 dorsal-fin spines separated by a membrane, not produced into a long filament,
and shorter than the following spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
17a. Membrane between dorsal-fin spines deeply incised (Fig. 14) . . . . . . . . . . . Nemipterus peronii
(Indo-West Pacific)
17b. Membrane between dorsal-fin spines continuous or only slightly emarginate (Fig. 15) . . . . . ® 18
dorsal-fin filament dorsal-fin membrane deeply incised
Fig. 13 Nemipterus nematophorus Fig. 14 Nemipterus peronii
18a. Upper lobe of caudal fin produced to an elongate point, falcate, or extended into a narrow
filament (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
18b. Upper lobe of caudal fin pointed or rounded, not produced (Fig. 17) . . . . . . . . . . . . . . . ® 28
membranes not incised
a) elongate point b) falcate c) filamentous

a) b)
Fig. 15 dorsal fin Fig. 16 shapes of upper caudal-fin lobe Fig. 17 shapes of upper
caudal-fin lobe
19a. Upper lobe of caudal fin produced to an elongate point; dorsal fin elevated, first dorsal-fin
spine long, 1.1 to 1.4 times in length of longest dorsal-fin spine . . . . . . . . . . . . . . . . . ® 20
19b. Upper lobe of caudal fin falcate or extended into a short or long filament; dorsal fin not
notably elevated, first dorsal-fin spine short, 1.3 to 2.9 times in length of longest spine . . . . . ® 21
3056 Bony Fishes
20a. Pelvic fins reaching to or beyond level of origin of anal fin; pectoral fins reaching to or
just beyond level of anus, 1.1 to 1.3 times in head length; total gill rakers on first gill arch
11 to 13; 5 pale golden stripes on sides beneath the lateral line; anal fin with a narrow
golden stripe just above its base, this stripe extending out to tip of the last soft ray . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus sp. 1
(Indonesia)
20b. Pelvic fins reaching to or short of level of origin of anal fin; pectoral fins reaching to or
just short of level of origin of anal fin, 0.9 to 1.1 times in head length; total gill rakers on
first gill arch 13 to 16; 2 narrow golden stripes on sides beneath the lateral line; anal fin
without stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus thosaporni
(West Pacific from southern Japan to Indonesia)
21a. Pectoral fins reaching to or beyond level of origin of anal fin; upper lobe of caudal fin with
moderately long filament, about equal to head length . . . . . . . . . . . . . . Nemipterus japonicus
(Indo-West Pacific)
21b. Pectoral fins not reaching to level of origin of anal fin; upper lobe of caudal fin with short
or long filament or falcate, ribbon-like extension . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
22a. Pelvic fins reaching to or beyond level of origin of anal fin, pectoral fins moderately long,
reaching to beyond level of anus, but short of level of origin of anal fin; upper lobe of
caudal fin with short filament . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus marginatus
(West Pacific)
22b. Pelvic fins not reaching to level of origin of anal fin; pectoral fins short to very long; upper
lobe of caudal fin with long filament or falcate, ribbon-like extension . . . . . . . . . . . . . . . ® 23
23a. A line drawn upwards from posterior edge of the suborbital reaching the dorsal profile at
or behind origin of dorsal fin; total gill rakers on first gill arch 14 to 20 . . . . . Nemipterus aurifilum
(northeastern Australia)
23b. A line drawn upwards from posterior edge of the suborbital reaching the dorsal profile
before origin of dorsal fin; total gill rakers on first gill arch 10 to 16 . . . . . . . . . . . . . . . . ® 24
24a. Body elongate, its depth 3.8 to 4.6 times in standard length . . . . . . . . . . . . Nemipterus zysron
(Indo-West Pacific)
24b. Body moderately deep, its depth 2.9 to 4 times in standard length . . . . . . . . . . . . . . . . ® 25
25a. Lower margin of eye on or below a line from tip of snout to upper base of pectoral fins . . . . . ® 26
25b. Lower margin of eye on or above a line from tip of snout to upper base of pectoral fins . . . . . ® 27
26a. Upper lobe of caudal fin falcate or ribbon-like, uniformly yellow; sides of body with 2
yellow stripes, the lower stripe originating behind base of pectoral fins; dorsal fin with
undulating yellow lines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus bathybius
(West Pacific)
26b. Upper lobe of caudal fin filamentous, yellow with red lower margin; sides of body with 2
yellow stripes, the lower stripe originating above upper base of pectoral fins; dorsal fin
without undulating yellow lines . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus balinensis
27a. Total pectoral-fin rays 16; pectoral and pelvic fins reaching to or just short of level of
anus; sides of body with 2 yellow stripes; red spot between first 2 dorsal-fin spines . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus nemurus
(West Pacific)
27b. Total pectoral-fin rays 16 to 18 (usually 17 or 18); pectoral and pelvic fins reaching to or
beyond level of anus; sides of body with indistinct midlateral yellow stripe; red spot
between first 2 dorsal-fin spines absent . . . . . . . . . . . . . . . . . . . . . . Nemipterus vitiensis
(Fiji, Vanuatu)
28a. Lower margin of eye below a line drawn from tip of snout to upper base of pectoral fins;
suborbital narrow, 3.6 to 6.7 times in eye diameter; golden yellow spot, edged with red
above and below, above upper base of pectoral fins . . . . . . . . . . . . Nemipterus balinensoides
(West Pacific)
28b. Lower margin of eye on or above a line drawn from tip of snout to upper base of pectoral
fins; suborbital deeper, 0.9 to 2.8 times in eye diameter; no yellow spot above base of
pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 29
29a. Scales below lateral line in ascending rows anteriorly (Fig. 18) . . . . . . . . . . . . . . . . . . ® 30
29b. Scales below lateral line in more or less horizontal rows anteriorly (Fig. 19) . . . . . . . . . . . ® 32
scale rows
ascending scale rows more or less horizontal
Fig. 18 longitudinal scale rows Fig. 19 longitudinal scale rows
30a. Upper lobe of caudal fin pointed, tip black (pigment retained in preserved specimens)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus theodorei
(eastern Australia)
30b. Upper lobe of caudal fin rounded or slightly rounded, tip pink or same colour as rest of
fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 31
31a. Dorsal fin pale yellow, with a broad, bicoloured submedial stripe (yellow above, orange
below), yellow margin and pale mauve submarginal stripe; anal fin with a single, pale
lemon submedial stripe (sometimes broken) . . . . . . . . . . . . . . . . . . . . Nemipterus aurora
(West Pacific)
31b. Dorsal fin translucent pink, with 2 or 3 narrow yellow medial stripes, and reddish yellow
margin; anal fin with 2 narrow pale yellow medial stripes . . . . . . . . . . . . Nemipterus celebicus
(Indonesia, northern Australia)
32a. Pectoral fins reaching to or short of level of anus . . . . . . . . . . . . . . . . . . . . . . . . . ® 33

32b. Pectoral fins reaching to beyond level of anus . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 34
33a. Pelvic fins short, reaching to or short of anus; a line drawn upwards from posterior edge
of the suborbital reaching the dorsal profile at about origin of dorsal fin; body pink, with
darker saddles on back; caudal fin with lower margin white-edged; dorsal and anal fins
without stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus furcosus
(East Indian Ocean to West Pacific)
33b. Pelvic fins very long, reaching to or beyond level of origin of anal fin; a line drawn
upwards from posterior edge of the suborbital reaching the dorsal profile 2 or more scale
rows before origin of dorsal fin; body pinkish; indistinct narrow, pale yellow stripes on
sides, 2 above lateral line, 5 below; dorsal fin pale yellow, lacking stripes; anal fin with
faint, broken yellow stripe near base which extends out on posterior rays to tip of fin . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus gracilis
(southern Indonesia)
3058 Bony Fishes
34a. Caudal fin forked or lunate, upper lobe falcate; last anal-fin ray elongate, notably longer
than other rays; yellow teardrop-shaped bar beneath eye . . . . . . . . . . . Nemipterus isacanthus
(West Pacific, from Philippines to northern Australia)
34b. Caudal fin forked, upper lobe pointed or rounded; last anal-fin ray not notably longer than
other rays; yellow bar beneath eye absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 35
35a. Enlarged canines anteriorly in upper and lower jaws; red ovoid spot present below origin
of lateral line; dorsal fin with narrow yellow stripe bordered on either side by translucent
interspace that is edged by a narrow pale bluish white stripe, this stripe extending
obliquely upwards to just above midposterior margin of fin; body deep, its depth 2.6 to
3.4 times in standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus hexodon
(West Pacific, from Andaman Sea to Solomon Islands)
35b. Enlarged canines anteriorly in upper jaw only; red spot below origin of lateral line absent;
colour of dorsal fin not as above; body moderately elongate, its depth 3.1 to 4 times in
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 36
36a. Pelvic fins very long, reaching to or beyond level of origin of anal fin . . . . . Nemipterus nematopus
(West Pacific)
36b. Pelvic fins moderately long, not reaching to level of origin of anal fin . . . . . . . . . . . . . . . ® 37
37a. Width of naked area of preopercle 1.4 to 1.7 times in width of scaly area; pectoral-fin
length 1.1 to 1.5 times in head length; caudal fin pink, with upper tip same colour as rest
of fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus mesoprion
(Gulf of Thailand, Indonesia)
37b. Width of naked area of preopercle 1.8 to 2.4 times in width of scaly area; pectoral-fin
length 1 to 1.1 times in head length; caudal fin pink, with upper tip sulphur yellow . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nemipterus tambuloides
(West Pacific, from Andaman Sea to Indonesia)
38a. No canine teeth in jaws; second anal-fin spine usually longer and more robust than third
spine (Fig. 20b); body depth 2.5 to 3 times in standard length . . . . . . . . . . (Parascolopsis) ® 39
38b. Two or 3 pairs of small canines anteriorly in upper jaw, a single pair of larger, flared
canines anteriorly on either side of lower jaw (Fig. 21); second anal-fin spine shorter and
less robust than third (Fig. 20a); body depth 3 to 3.5 times in standard length . . (Pentapodus) ® 44
39a. Total gill rakers on first gill arch 17 to 19 . . . . . . . . . . . . . . . . . . . . Parascolopsis eriomma

(Indo-West Pacific)
39b. Total gill rakers on first gill arch 8 to 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 40
40a. Preopercle more or less entirely scaly or with an incomplete naked flange posteriorly on
its free margin (Fig. 22a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parascolopsis tosensis
(southern Japan, Philippines, Indonesia)
40b. Preopercle with a distinct broad naked flange bordering its free margin (Fig. 22b) . . . . . . . . ® 41
large, flared canine
broad,
a) b) naked
a) b) flange
Fig. 20 anal-fin spines Fig. 21 lower jaw (dentary) Fig. 22 Squamation on preopercle
of Pentapodus
41a. Pelvic fins not reaching to or near level of anus, length of pelvic fins 1.6 to 1.9 times in
head length; 4 or 5 transverse scale rows above lateral line; interorbital width 1 to 1.3
times in eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . Parascolopsis rufomaculatus
(northwestern Australia)
41b. Pelvic fins reaching close to or beyond level of anus, length of pelvic fins 1.2 to 1.5 times
in head length; 2 ½ or 3 ½ transverse scale rows above lateral line; interorbital width 1.3
to 2.2 times in eye diameter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 42
42a. Triangular black spot above and behind eye . . . . . . . . . . . . . . . . Parascolopsis melanophrys
(East Malaysia, East Indonesia)
42b. Triangular black spot above and behind eye absent . . . . . . . . . . . . . . . . . . . . . . . . ® 43
43a. Dusky band joining nostrils across snout; fourth or fifth dorsal-fin ray elongate in larger
specimens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parascolopsis tanyactis
(Philippines, Indonesia, northwestern Australia)
43b. Dusky band across snout absent; fourth or fifth dorsal-fin ray not notably elongate . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parascolopsis inermis
(West Pacific to East Indian Ocean)
44a. Caudal fin with upper rays or upper and lower rays produced into long trailing filaments . . . . ® 45
44b. Caudal fin without long filamentous extensions, lobes pointed, sharply pointed, or falcate . . . . ® 47
45a. Upper and lower lobes of caudal fin produced

into long trailing filaments; lower limb of preop-
ercle scaly, at least anteriorly (Fig. 23a); lat-
eral-line scales 50 to 56 (usually 52 to 54) . .
. . . . . . . . . . . . . . . . . . . . . Pentapodus emeryii
(Philippines, Indonesia, northwestern Australia)
45b. Upper lobe of caudal fin only produced into a) b)
long trailing filament; lower limb of preopercle
naked (Fig. 23b); lateral-line scales 46 to 50 . . . . . ® 46 Fig. 23 squamation on preopercle
46a. Three dusky stripes (blue in life) on snout; first from eye to tip of snout, second from eye
across snout in front of nostrils, third joining eyes behind nostrils . . . . . . . Pentapodus paradiseus
(northeastern Australia, Papua New Guinea, Solomon Islands)
46b. Two dusky stripes (blue in life) on snout; first from eye to middle of upper lip, second from
eye to tip of snout; no stripe joining eyes . . . . . . . . . . . . . . . . . . . . . . Pentapodus setosus
(Philippines, Indo-Malay Archipelago)
47a. Head scales reaching forward dorsally to or in front of posterior nostril . . . . . . . . . . . . . . ® 48

47b. Head scales reaching forward dorsally to anterior margin of eye or almost to posterior
nostril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 49
wedge-shaped notch
naked area
48a. Head scales reaching to between level of poste-
rior and anterior nostrils; scaly area between
nostrils rectilinear (Fig. 24a); caudal-fin lobes
pointed . . . . . . . . . . . . . . . . Pentapodus bifasciatus
(Philippines, Indo-Malay Archipelago) scaly scaly
48b. Head scales extending to or in front of level of area area
anterior nostril; scaly area between nostrils usu-
ally with a naked, wedge-shaped notch anteri-
orly (Fig. 24b); caudal-fin lobes falcate, upper
lobe usually longer than lower lobe . . . Pentapodus caninus a) b)
(Philippines, Indo-Malay Archipelago)
Fig. 24 top of head showing forward
extent of scales
3060 Bony Fishes
49a. Lower limb of preopercle naked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 50

49b. Lower limb of preopercle scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 51
50a. Snout length greater than eye diameter; body moderately deep, its depth 3 to 3.7 times
in standard length; black spot on caudal peduncle . . . . . . . . . . . . . . . . Pentapodus porosus
(northwestern Australia)
50b. Snout length equal to or less than eye diameter; body slender, its depth 3.9 to 4.5 times
in standard length; no spot on caudal peduncle . . . . . . . . . . . . . . Pentapodus nagasakiensis
(Japan, South China Sea, northern Australia)
51a. Body pale with 3 dusky stripes (brown in life) along sides; dark bar at base of pectoral
fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pentapodus trivittatus
(Indo-Malay Archipelago, New Guinea, Solomon Islands)
51b. Body colour not as above; no dark bar at base of pectoral fins . . . . . . . . . . . . Pentapodus sp.
(West Pacific, from Taiwan Province of China to Samoa)

0
0 Nemipterus aurifilum (Ogilby, 1910)
0 Nemipterus aurora Russell, 1993
0 Nemipterus balinensis (Bleeker, 1858-9)
0 Nemipterus balinensoides (Popta, 1918)
0 Nemipterus bathybius Snyder, 1911
0 Nemipterus celebicus (Bleeker, 1854)
0 Nemipterus furcosus (Valenciennes, 1830)
0 Nemipterus gracilis (Bleeker, 1873)
0 Nemipterus hexodon (Quoy and Gaimard, 1824)
0 Nemipterus isacanthus (Bleeker, 1873)
0 Nemipterus japonicus (Bloch, 1791)
0 Nemipterus marginatus (Valenciennes, 1830)
0 Nemipterus mesoprion (Bleeker, 1853)
0 Nemipterus nematophorus (Bleeker, 1853)
0 Nemipterus nematopus (Bleeker, 1851)
0 Nemipterus nemurus (Bleeker, 1857)
0 Nemipterus peronii (Valenciennes, 1830)
0 Nemipterus sp. 1
0 Nemipterus tambuloides (Bleeker, 1853)
0 Nemipterus theodorei Ogilby, 1916
0 Nemipterus thosaporni Russell, 1991
0 Nemipterus virgatus (Houttuyn, 1782)
0 Nemipterus vitiensis Russell, 1990
0 Nemipterus zysron (Bleeker, 1856-57)
0 Parascolopsis eriomma (Jordan and Richardson, 1909)
0 Parascolopsis inermis (Schlegel, 1843)
0 Parascolopsis melanophrys Russell and Chin, 1996
0 Parascolopsis rufomaculatus Russell, 1986b
0 Parascolopsis tanyactis Russell, 1986b
0 Parascolopsis tosensis (Kamohara, 1938)
0 Parascolopsis townsendi
0 Pentapodus bifasciatus (Bleeker, 1848)
0 Pentapodus caninus (Cuvier, 1830)
0 Pentapodus emeryii (Richardson, 1843)
0 Pentapodus nagasakiensis (Tanaka, 1915)
0 Pentapodus paradiseus (Günther, 1859)
0 Pentapodus porosus (Valenciennes, 1830)
0 Pentapodus setosus (Valenciennes, 1830)

0 Pentapodus sp.
0 Pentapodus trivittatus (Bloch, 1791)
0 Pentapodus vitta
0 Scaevius milii (Bory de Saint-Vincent, 1823)
0 Scolopsis affinis Peters, 1877
0 Scolopsis auratus (Park, 1797)
0 Scolopsis bilineatus (Bloch, 1793)
0 Scolopsis ciliatus (Lacepède, 1802)
0 Scolopsis lineatus Quoy and Gaimard, 1824
0 Scolopsis margaritifer
0 Scolopsis monogramma (Kuhl and Van Hasselt, 1830)
0 Scolopsis taeniopterus (Kuhl and van Hasselt, 1830)
0 Scolopsis temporalis (Cuvier, 1830)
0 Scolopsis trilineatus Kner, 1868
0 Scolopsis vosmeri (Bloch, 1792)
0 Scolopsis xenochrous Günther, 1872
References
Russell, B.C. 1990. FAO species catalogue. Vol. 12. Nemipterid fishes of the world (threadfin breams, whiptail breams,
monocle breams, dwarf monocle breams, and coral breams). Family Nemipteridae. An annotated and illustrated
catalogue of nemipterid species known to date. FAO Fish. Synop., (125)12. Rome, FAO, 149 p.
Russell, B.C. 1991. Description of a new species of Nemipterus (Pisces: Perciformes: Nemipteridae) from the western
Pacific, with re-descriptions of Nemipterus marginatus (Valenciennes), N. mesoprion (Bleeker) and N. nematopus
(Bleeker). J. Nat. Hist., 25:1379-1389.
Russell, B.C. 1993. A review of the threadfin breams of the genus Nemipterus (Nemipteridae) from Japan and Taiwan,
with description of a new species. Japan. J. Ichthyol., 39:295-310.
click for next page

3062 Bony Fishes
Nemipterus aurora Russell, 1993 (Plate XX, 146)

Frequent synonyms / misidentifications: None / Previously misidentified as Nemipterus delagoae Smith,
1941 [= N. bipunctatus (Ehrenberg, 1830)], a species which it closely resembles but which is restricted to
the Indian Ocean.
FAO names: En - Dawn threadfin bream.
Diagnostic characters: Body depth 3.2 to 3.9 times in standard length. Snout length equal to or a little more
than eye diameter. Three pairs of small recurved canines anteriorly in upper jaw. Eye diameter 2.9 to 3.7
times in head length; lower margin of eye on or above a line from tip of snout to upper base of pectoral
fins; interorbital width 1.2 to 1.8 times in eye diameter; suborbital depth 1.1 to 2 times in eye diameter; a
line drawn up from posterior edge of suborbital reaching the dorsal profile somewhere between 2
scale rows before origin of dorsal fin and third dorsal-fin spine. Width of naked area of preopercle 1.2
to 1.7 times in width of scaly area. Total gill rakers on first gill arch 11 to 14. Total pectoral-fin rays 15 to 17;
pectoral fins moderately long, 1.1 to 1.4 times in head length, reaching to or just beyond level of
anus; pelvic fins very long, 0.9 to 1.3 times in head length, reaching to just before level of origin of
anal fin and third anal-fin spine; caudal fin forked, tips of fin pointed or slightly rounded. Scale rows
on body below lateral line upward-curved anteriorly. Colour: body pinkish above, shading through pale
mauve to silvery on ventral half; sides with 4 or 5 longitudinal, upward-curved silvery white stripes
along the middle of each scale row below lateral line, from behind head to caudal peduncle; head
pinkish with golden yellow reflections on opercle; upper lip with yellow edge; dorsal fin pale translucent
yellow, with lemon yellow margin and pale mauve submarginal stripe; broad bicoloured submedial
stripe, yellow above orange below, becoming uniformly yellow posteriorly; anal fin transparent, with
pale lemon submedial stripe (sometimes broken) above base of fin; caudal fin yellowish pink, upper
tip rosy; pelvic and pectoral fins transparent.
Size: Maximum standard length 20 cm, commonly to 15 cm.
Habitat, biology, and fisheries: A benthic species, found on sand or mud bottoms in depths of 56 to 90 m.
Taken in commercial trawls in the South China Sea and southern Gulf of Thailand together with other species
of Nemipterus. No major fishery exists.
Distribution: West Pac ific
from the Ryukyu Islands to
southeastern Indonesia.
Nemipterus bathybius Snyder, 1911 (Plate XX, 148)

FAO names: En - Yellowbelly threadfin bream.
Diagnostic characters: Body depth 2.9 to 3.6 times in standard length. Snout length equal to or a little less
than eye diameter. Three or 4 pairs of small recurved canines anteriorly in upper jaw. Eye diameter 2.7 to
3.6 times in head length; lower margin of eye tangent to or just below a line from tip of snout to upper
base of pectoral fins; interorbital width 1.3 to 1.9 times in eye diameter; suborbital depth 1.9 to 4.7 times
in eye diameter; a line drawn upwards from posterior edge of suborbital reaching the dorsal profile
in front of origin of dorsal fin. Width of naked area of preopercle 1.6 to 2.2 times in width of scaly area.
Total gill rakers on first gill arch 13 to 16. Pectoral-fins 15 to 17; pectoral fins long, 1 to 1.4 times in head
length, reaching almost to level of origin of anal fin; pelvic fins moderately long, 1.1 to 1.6 times in
head length, reaching beyond anus; caudal fin forked, upper lobe falcate, usually long and ribbon-
like in adult specimens. Colour: upper part of body pinkish, silvery below; 2 yellow lateral stripes,
the first beneath lateral line from behind opercle to upper caudal-fin base, the second from behind
upper part of pectoral-fin base to middle of caudal-fin base; a pair of yellow stripes, united anteriorly,
from the isthmus to lower caudal-fin base, passing just outside bases of pelvic and anal fins; dorsal
fin pink, edged with yellow, with a median stripe of yellowish undulating lines; anal fin transparent, pinkish
near outer margin; caudal fin pink, upper lobe and filament yellow; other fins hyaline; bases of pelvic fins
bright sulphur yellow.
Habitat, biology, and fisheries: A benthic species, found on sand or mud bottoms at depths of 35 to 300 m,
but most common in depths between 45 to 90 m. Larger fishes generally occur in depths greater than 110 m.
This species is a rudimentary hermaphrodite, in which all males have functional testes with rudimentary
ovarian portions throughout their life. In the South China Sea, spawning is reported from June to October.
The diet of larger fishes consists mainly of crustaceans, fish, and cephalopods. Young fishes (120 mm) feed
on copepods, ostracods, and amphipods. Males grow more quickly and to a larger size than females.
Estimates of the parameters of the von Bertalanffy growth curve made for the population (males) from off
Hong Kong are: L¥ = 30.2 cm total length, K = 0.32, t0 = - 0.80. Taken in commercial quantities in the South
China Sea by longline and
bottom trawl. Catch rates are
higher in the day-time. Mar-
keted mainly fresh.
Distribution: West Pacific, in-
c l ud i ng s ou th er n J ap an,
South China Sea, Philippines,
Indonesia, and northwestern
Australia.
3064 Bony Fishes
Nemipterus celebicus (Bleeker, 1854) (Plate XX, 149)

FAO names: En - Celebes threadfin bream.
Diagnostic characters: Body depth 2.9 to 4.2 times in standard length. Snout length about equal to or
greater than eye diameter. Three pairs of small recurved canines anteriorly in upper jaw. Eye diameter 3 to
4 times in head length; lower margin of eye tangent to or just above a line from tip of snout to upper
base of pectoral fins; interorbital width 1.2 to 1.8 times in eye diameter; suborbital depth 1 to 3 times in
eye diameter; a line drawn up from posterior edge of suborbital reaching the dorsal profile about 2
to 4 scales before origin of dorsal fin. Width of naked area of preopercle 1.1 to 1.8 times in width of scaly
area. Total gill rakers on first gill arch 10 to 14. Total pectoral-fin rays 16 or 17; pectoral fins long, 1.1 to
1.4 times in head length, reaching to between level of anus and origin of anal fin; pelvic fins very
long, 0.9 to 1.4 times in head length, reaching to or beyond level of origin of anal fin; caudal fin
forked, upper lobe slightly rounded, without filament. Scale rows on body below lateral line upward-
curved anteriorly. Colour: upper part of head and body pinkish, paling to silvery white on ventral
surface; violet and reddish reflections on scales above lateral line; indefinite pale golden or yellowish
stripe along lateral line and narrow golden stripe on each of 4 scale rows below lateral line, these
stripes ascending slightly before level of anal fin; head with golden and mauve reflections; traces of 2
silvery mauve lines in front of and below eye; bluish mauve spot on upper part of opercle; upper lip yellow;
dorsal fin translucent pink, with narrow reddish yellow margin and 2 or 3 narrow yellow stripes along
median part of fin; anal fin translucent with 2 narrow pale yellow stripes along median part of fin;
caudal fin pale golden, upper tip rosy; pectoral fins pale pink; pelvic fins translucent white.
Habitat, biology, and fisheries: A benthic species, found on sand or mud bottoms in depths of 30 to 100 m.
Feeds mainly on small crustaceans. Taken by line and bottom trawl. Appears in local fish markets in
Indonesia; taken commercially by trawlers in northern Australian waters together with other species. No
major fishery exists.
Distribution: I n d o n es i a,
northern Australia.
Nemipterus furcosus (Valenciennes, 1830) (Plate XX, 150)

FAO names: En - Forktailed threadfin bream.
Diagnostic characters: Body depth 3 to 3.9 times in standard length. Snout length equal to or greater than
eye diameter. Two or 3 pairs of small recurved canines anteriorly in upper jaw. Eye diameter 2.8 to 4 times
in head length; lower margin of eye tangent to or above a line from tip of snout to upper base of
pectoral fins; interorbital width 1.1 to 1.7 times in eye diameter; suborbital depth 0.9 to 1.9 times in eye
diameter; a line drawn up from posterior edge of suborbital reaching the dorsal profile at about origin
of dorsal fin. Width of naked area of preopercle 1 to 1.4 times in width of scaly area. Total gill rakers on
first gill arch 9 to 12. Total pectoral-fin rays 15 to 17; pectoral fins moderately long, 0.8 to 1.1 times in
head length, reaching to or just short of level of anus; pelvic fins moderately long, 1.3 to 1.7 times
in head length, reaching to or just short of level of anus; caudal fin deeply forked. Colour: head and
body pale iridescent pink, paling on sides to silvery white below; back with 9 indistinct cross bars,
extending to just beneath lateral line, third bar somewhat darker and more distinct, forming a reddish
shoulder spot behind origin of lateral line in some specimens; traces of indistinct yellowish stripes
along body, one above and several below lateral line; cheeks and opercle silvery, upper jaw rosy, lower
jaw silvery; eye rosy; dorsal fin pale rosy, sometimes with yellowish tinge, outer margin darker pink;
anal fin bluish white, with row of transparent or faint yellowish spots near base; caudal fin pale rosy,
with yellow tinge, lower margin of fin white; pelvic fins and axillary scales white; pectoral fins rosy.
Size: Maximum standard length 22.5 cm, commonly to 18 cm.
Habitat, biology, and fisheries: A benthic species, inhabiting sand and mud bottoms in depths of 8 to
110 m. Females predominate at small size classes while males dominate the larger size classes, and there
is some evidence of this species being a sequential hermaphrodite. Ripe females occur all year round in the
population from the northwest shelf of Australia, but a higher proportion of ripe fishes is present in November
and December. Feeding occurs during the day, with crustaceans and small fishes predominating. Estimates
of the parameters of the von Bertalanffy growth curve made for the population from the northwest shelf of
Australia are: L¥ = 41.9 cm fork length, K = 0.25, t0 = 0.74. Taken by handline and bottom trawl. Appears in
local fish markets. Considered a good food fish and fished commercially by trawlers together with other
species. Nemipterus furcosus is the principal catch species in the trawl fishery off the coast of northwestern
Australia.
Distribution: West Pacific from southern
Japan to northeastern Australia, and Indian
Ocean including the Gulf of Mannar, Sri
Lanka, Andaman Sea, Strait of Malacca,
and northwestern Australia.
3066 Bony Fishes
Nemipterus hexodon (Quoy and Gaimard, 1824) (Plate XX, 152)

FAO names: En - Ornate threadfin bream.
Diagnostic characters: Body depth 2.6 to 3.4 times in standard length. Snout length equal to or greater
4.6 times in head length; lower margin of eye above a line from tip of snout to upper base of pectoral
fins; interorbital width 1.3 to 1.9 times in eye diameter; suborbital depth 0.9 to 1.8 times in eye diameter; a
line drawn up from posterior edge of suborbital reaching the dorsal profile about 2 to 6 scale rows before
origin of dorsal fin. Width of naked area of preopercle 1.2 to 1.9 times in width of scaly area. Total gill rakers
on first gill arch 11 to 17. Total pectoral-fin rays 15 to 18; pectoral fins long, 1 to 1.4 times in head length,
reaching to or beyond level of anus; pelvic fins long, 1.2 to 1.6 times in head length, reaching to or
just beyond level of anus; caudal fin forked, upper lobe slightly longer than lower lobe. Colour: upper
part of body pinkish, paling to silvery white on ventral surface; 6 to 8 pale yellow stripes on sides
from below lateral line; blood red, ovoid spot below origin of lateral line, bordered below by bright
yellow; yellow stripe on either side of ventral midline, from isthmus to lower caudal-fin base; golden
reflections behind eye, on cheeks and opercle; dorsal fin translucent whitish, with yellow margin; a
narrow yellow stripe beginning anteriorly near base of fin and extending backwards to just above
midposterior margin, this stripe bordered on either side by a translucent interspace that is edged by
a narrow pale bluish white stripe; caudal fin pinkish, upper lobe tipped with yellow; anal fin
translucent; a narrow yellow stripe running from near anterior base of fin to midposterior margin
(extending to tip of fin along posterior ray in larger specimens); base of fin beneath this stripe pale
bluish white; pectoral and pelvic fins pale translucent pink; base of pelvic fins and axillary scale lemon
yellow.
Habitat, biology, and fisheries: A benthic species, occurs on mud or sand bottoms in depths of 10 to 80
m, but is most abundant in depths of 20 to 50 m. Females predominate at small sizes and males at larger
sizes as a result of faster growth rates in males. In the Gulf of Thailand, spawning probably occurs between
February and March. Feeds
mainly on small shrimps,
squid, small fishes, and ben-
thic animals. Taken by han-
dline and bottom trawl
together with other demersal
fishes throughout the year.
Appears in local markets. No
cific, from the Andaman Sea to
the Solomon Islands.
Nemipterus isacanthus (Bleeker, 1873) (Plate XX, 153)

FAO names: En - Teardrop threadfin bream.
Diagnostic characters: Body depth 3 to 3.5 times in standard length. Snout length equal to or greater than
eye diameter. Three pairs of small recurved canines anteriorly in upper jaw. Eye diameter 3 to 3.7 times in
head length; lower margin of eye above a line from tip of snout to upper base of pectoral fins; interorbital
width 1.4 to 1.9 times in eye diameter; suborbital depth 1.1 to 2.1 times in eye diameter; a line drawn up
from posterior edge of suborbital reaching the dorsal profile about 2 to 7 scale rows before origin of
dorsal fin. Width of naked area of preopercle 1.7 to 2.7 times in width of scaly area. Total gill rakers on first
gill arch 11 to 14. Total pectoral-fin rays 16 or 17; pectoral fins long, 1 to 1.4 times in head length, reaching
to beyond level of anus; pelvic fins moderately long, 1.3 to 1.6 times in head length, reaching to
between just short of to just beyond level of anus; caudal fin deeply forked, upper lobe falcate.
Colour: body pinkish mauve, silvery on lower part of sides and ventral surface; top of head and back
yellow-tinged pink; 2 broad pale golden stripes along body, the first from beneath origin of lateral line
to upper caudal-fin base, the second from behind pectoral-fin base to lower caudal-fin base; pale
yellow stripe on either side of ventral midline; snout pinkish; yellow teardrop-shaped bar beneath eye,
extending obliquely forwards about halfway down suborbital; upper lip yellow; bluish patch on upper
part of opercle; dorsal fin translucent, with narrow, red-edged yellow margin, and narrow yellow
submedial stripe; caudal fin pink, tip of upper lobe bright yellow, lower lobe yellow-tinged; anal fin
and pelvic fins translucent whitish; pectoral fins translucent pinkish.
Size: Maximum standard length 21 m, commonly to 17 cm.
Habitat, biology, and fisheries: A benthic species, occurs on mud or sand bottoms in depths greater than
50 m. Taken by handline and bottom trawl together with other species. Appears occasionally in local markets.
No major fishery exists.
cluding the Philippines, Gulf of
Thailand, Strait of Malacca, In-
donesia, and northern Austra-
lia.
3068 Bony Fishes
Nemipterus japonicus (Bloch, 1791) (Plate XXI, 154)

FAO names: En - Japanese threadfin bream; Fr - Cohana japonaise; Sp - Baga japonesa.
than eye diameter. Four or 5 pairs of small recurved canines anteriorly in upper jaw. Eye diameter 3.2 to 4.4
times in head length; lower margin of eye above a line from tip of snout to upper base of pectoral fins;
interorbital width 1 to 1.9 times in eye diameter; suborbital depth 1 to 1.9 times in eye diameter; a line drawn
up from the posterior edge of suborbital reaching the dorsal profile 2 to 4 scale rows before origin
of dorsal fin. Width of naked area of preopercle 1.1 to 2 times in width of scaly area. Total gill rakers on
first gill arch 14 to 17. Total pectoral-fin rays 17 or 18; pectoral fins very long, 1 to 1.3 times in head
length, reaching to or just beyond level of origin of anal fin; pelvic fins moderately long, 1.2 to 1.6
times in head length, reaching to or just beyond anus; caudal fin moderately forked, upper lobe
slightly longer than lower lobe and produced into a short or moderately long filament. Colour: upper
part of body pinkish, becoming silvery below; top of head behind eye with a golden sheen; 11 or 12 pale
golden yellow stripes along body from behind head to base of caudal fin; a prominent red-suffused
yellow blotch below origin of lateral line; dorsal fin whitish, margin of fin yellow, edged with red; a
pale lemon stripe near base of dorsal fin, this stripe narrow anteriorly and widening on posterior part
of fin; anal fin whitish with pale lemon broken lines or scribblings over most of fin; pectoral fins
translucent pinkish; pelvic fins whitish with yellow axillary scale; caudal fin pink, upper tip and filament
yellow.
Habitat, biology, and fisheries: A benthic species, very abundant in coastal waters, found on mud or sand
bottoms in depths of 5 to 80 m, usually in schools. Small fishes are generally most abundant in depths less
than 27 m; only larger fishes occur in depths greater than 45 m. Females predominate at small sizes and
males at large sizes, due to faster growth rates in males. Spawning occurs over an extended period, and is
reported to take place from May to October in the South China Sea, and from November to February in East
Malaysian waters. The diet consists mainly of small fishes, crustaceans, molluscs (mainly cephalopods),
polychaetes, and echinoderms. Estimates of the parameters of the von Bertalanffy growth curve made for
the population off the Strait of Malacca (Kedah State, Malaysia) are: L¥ = 31.4 cm total length, K = 0.55.
Taken in commercial quantities by longlines, bottom trawls, and gill nets. Juveniles are also caught in
considerable quantities by shrimp trawlers using beam trawls. Nemipterus japonicus constitutes an impor-
tant part of the trawl catch in the South China Sea. Marketed mainly fresh, but also steamed, dried salted,
dry-smoked, fermented, or made into fish
balls and fish meal. In the Philippines, this
species is especially valued in a convales-
cent diet.
Distribution: Widespread throughout the
Indian Ocean and West Pacific. A report on
N. japonicus as a Red Sea immigrant into
the Mediterranean is unconfirmed. Simi-
larly, a record of this species from north-
western Australia is without basis.
Nemipterus nematophorus (Bleeker, 1853) (Plate XXI, 157)

Frequent synonyms / misidentifications: Nemipterus filamentosus (Valenciennes in Cuvier and Valen-
ciennes, 1830) (name preoccupied by Cantharus filamentosus Rüppell) / None.
FAO names: En - Doublewhip threadfin bream.
greater than eye diameter. Three to 5 pairs of small recurved canines anteriorly in upper jaw. Eye diameter
3 to 4 times in head length; lower margin of eye tangent to a line from tip of snout to upper base of
pectoral fins; interorbital width 0.8 to 1.8 times in eye diameter; suborbital depth 1.4 to 2.6 times in eye
diameter; a line drawn up from the posterior edge of suborbital reaching the dorsal profile about 2 to
5 scale rows before origin of dorsal fin. Width of naked area of preopercle 1.3 to 1.9 times in width of
scaly area. Total gill rakers on first gill arch 12 to 15. Dorsal fin with anterior pair of spinous rays close
together, almost fused and produced into a long, trailing filament. Total pectoral-fin rays 16 or 17;
pectoral and pelvic fins long, reaching to between level of anus and origin of anal fin; pectoral fins
0.9 to 1.1 times in head length, pelvic fins 0.9 to 1.5 times in head length; caudal fin forked, upper
lobe produced into a trailing filament. Colour: upper part of head and body pinkish, silvery white below;
broad golden yellow stripe beneath lateral line, with distinct gold patch anteriorly beneath origin of
lateral line; 3 narrow golden yellow stripes laterally along ventral half of the body; yellow stripe on
either side of ventral midline; dorsal fin translucent pinkish, anterior dorsal filament and margin of
fin yellow; anal fin translucent, faint yellow or orange stripe along fin extending from near base of
first spine to middle of last ray, basal part of fin pale bluish white; caudal fin pink, upper tip and
caudal filament yellow; pectoral fins pale pink; pelvic fins pink, axillary scale lemon yellow.
Habitat, biology, and fisheries: A benthic species, occurs on sand or mud bottoms in depths to 75 m.
Taken by gill net and bottom trawl. In East Malaysian waters highest catches occur in depths of more than
50 m. Appears occasionally in local markets. No major fishery exists.
cific, including the Bay of Ben-
gal, Andaman Sea, Strait of
Malacca, Philippines, South
China Sea, Gulf of Thailand,
and Indonesia. Record of this
species from northwestern
Australia is apparently without
basis.
3070 Bony Fishes
Nemipterus nemurus (Bleeker, 1857) (Plate XXI, 159)

FAO names: En - Redspine threadfin bream.
Diagnostic characters: Body depth 3.3 to 4 times in standard length. Snout length a little more than or a
little less than eye diameter. Two to 5 pairs of small recurved canines anteriorly in upper jaw. Eye diameter
2.9 to 3.7 times in head length; lower margin of eye tangent to or just above a line from tip of snout to
upper base of pectoral fins; interorbital width 1.4 to 1.9 times in eye diameter; suborbital depth 1.3 to 2.2
times in eye diameter; a line drawn up from the posterior edge of suborbital reaching the dorsal profile
about 3 to 7 scale rows before origin of dorsal fin. Width of naked area of preopercle 1.5 to 2.2 times in
width of scaly area. Total gill rakers on first gill arch 12 to 13. Total pectoral-fin rays 16; pectoral and pelvic
fins moderately long, reaching to or just short of level of anus; pectoral fins 1.1 to 1.3 times in head
length; pelvic fins 1.2 to 1.5 times in head length; caudal fin forked, the upper rays produced into a
trailing filament. Colour: body pinkish above, with indistinct, broad, pale yellow stripe from behind eye
to caudal-fin base; indistinct, narrow golden stripes along each of 3 scale rows below lateral line;
ventral half of body pearly-white, with indistinct, broad, pale yellow stripe from behind pectoral fins
to caudal-fin base; head pinkish, with pale golden reflections on cheeks and opercle; golden yellow stripe
from posterior nostril extending through eye, and similar stripe from anterior margin of upper lip to
lower margin of eye; iris pink; dorsal fin pale yellow, base of fin and submargin translucent pale mauve,
margin of fin yellow; interspinous membrane of first 2 dorsal-fin spines bright red superiorly; anal
fin white, with series of yellow spots or irregular yellow stripe submedially; caudal fin pinkish,
posterior margin red; lower lobe of fin suffused with yellow, upper rays and filament yellow; pectoral
fins pale yellowish pink; pelvic fins white.
Habitat, biology, and fisheries: A benthic species, occurs on mud or sand bottoms. Feeds on small fishes
and larger benthic invertebrates. Taken by handline and bottom trawl together with other species. The most
abundant species of Nemipterus in bottom trawl catches from the South China Sea. Appears in small
numbers in local markets. No
cluding the Philippines, Gulf of
Thailand, South China Sea,
Strait of Malacca, and Indone-
sia.
Nemipterus peronii (Valenciennes, 1830) (Plate XXI, 160)

Frequent synonyms / misidentifications: Nemipterus tolu (Valenciennes in Cuvier and Valenciennes,
1830) / The name N. peronii has been misapplied to N. furcosus.
FAO names: En - Notchedfin threadfin bream; Fr - Cohana tolu; Sp - Baga tolu.
Diagnostic characters: Body depth 3.1 to 4.1 times in standard length. Snout length about equal to eye
diameter. Three or 4 pairs of small recurved canines anteriorly in upper jaw. Eye diameter 2.8 to 3.6 times
in head length; lower margin of eye just above a line from tip of snout to upper base of pectoral fins;
interorbital width 1.4 to 1.9 times in eye diameter; suborbital depth 1.1 to 2.2 times in eye diameter; a line
drawn up from the posterior edge of suborbital reaching the dorsal profile at or just before origin of
dorsal fin. Width of naked area of preopercle 1.5 to 2.2 times in width of scaly area; dorsal-fin spines
elongate, interspinous membrane deeply incised. Total gill rakers on first gill arch 9 to 12 (usually 10 or
11). Total pectoral-fin rays 16 or 17; pectoral fins short, 1.1 to 1.7 times in head length, not reaching to
level of anus; pelvic fins moderately long, 1.1 to 1.5 times in head length, reaching to level of anus;
caudal fin forked, upper lobe pointed and slightly longer than lower lobe. Colour: upper part of body
pinkish, with 7 or 8 indistinct darker pink saddles reaching to or just below the lateral line; lower part
of body silvery, with faint golden lines following each scale row; a diffuse pale reddish spot below
and just behind origin of lateral line; a golden yellow stripe on snout in front of eye passing through
nostrils; upper lip yellow; suborbital silvery mauve; golden reflections on preopercle and opercle; dorsal
fin pale whitish pink, with a pale yellow line or series of spots just above base of fin; tips of spinous
part of fin reddish yellow; anal fin pale whitish pink, suffused pale yellowish over its middle part; caudal
fin pinkish; pelvic fins whitish, axillary scale at base yellow, pectoral fins translucent.
Habitat, biology, and fisheries: A benthic species, found on sand or mud bottoms in depths to about 100 m.
Size of the fishes appears to increase with depth and males are relatively larger than females. Two prolonged
spawning seasons are likely in the South China Sea off the Trengganu coast: from November to February,
and another starting in May or June. This species is an active carnivore, feeding during daylight on fish,
crustaceans, molluscs, and polychaetes. Taken by handline and bottom trawl along with other species of
Nemipterus. Appears in small
numbers in local markets.
Trawled in commercial quanti-
ties in the Straits of Malacca
and off the Trengganu coast,
South China Sea.
Distribution: W e s t P a c i f ic
from Taiwan Province of China
to northern Australia, and In-
dian Ocean including the An-
daman Sea, Bay of Bengal, Sri
Lanka, Arabian Sea, Persian
Gulf, and Red Sea.
3072 Bony Fishes
Nemipterus tambuloides (Bleeker, 1853) (Plate XXI, 161)

FAO names: En - Fivelined threadfin bream.
4.1 times in head length; lower margin of eye tangent to or just above a line from tip of snout to upper
base of pectoral fins; interorbital width 1.5 to 1.9 times in eye diameter; suborbital depth 1.3 to 1.8 times
in eye diameter; a line drawn up from the posterior edge of suborbital reaching the dorsal profile about
2 to 7 scale rows before origin of dorsal fin. Width of naked area of preopercle 1.8 to 2.4 times in width
of scaly area. Total gill rakers on first gill arch 12 to 14. Total pectoral-fin rays 16 or 17; pectoral and pelvic
fins long, reaching to between level of anus and origin of anal fin; pectoral fins 1 to 1.1 times in head
length, pelvic fins 1.1 to 1.4 times in head length; caudal fin forked, upper lobe pointed. Colour: upper
part of head and body rosy, paling to silvery white on ventral surface; 5 well-defined sulphur yellow stripes
along body, the first stripe beginning above lateral line and extending along scale row to posterior
of dorsal fin, the second stripe running from snout through eye, postorbital part of head, and then
extending as a more or less straight line to upper caudal-fin base; remaining stripes beginning behind
head and following each scale row, slightly ascending posteriorly; sulphur yellow stripe along either
side of ventral midline from isthmus to lower caudal-fin base; head with 2 yellow stripes beneath
eye; cheeks and opercle with golden and mauve reflections; dorsal fin translucent pink, with yellow
margin and bluish grey inframarginal stripe; narrow sulphur yellow stripe extending just above base
of dorsal fin; anal fin translucent bluish white with pale yellow stripe near base of fin, this stripe bent
posteriorly and extending out to tip of last anal-fin ray; caudal fin bright rosy, upper tip sulphur
yellow; pectoral fins pale pink; pelvic fins pale yellowish, spine and first ray bluish grey.
Habitat, biology, and fisher-
ies: A benthic species, occurs
on sand or mud bottoms in
depths of 50 to 70 m. Taken by
handline and bottom trawl. An
abundant species in Thailand,
landed in large quantities
throughout the year.
Distribution: Andaman Sea,
Strait of Malacca, Philippines,
Gulf of Thailand, South China
Sea, and Indonesia.
Nemipterus thosaporni Russell, 1991 (Plate XXII, 162)

Frequent synonyms / misidentifications: None / Previously misidentified as Nemipterus marginatus
(Valenciennes, 1830) by most authors.
FAO names: En - Palefin threadfin bream.
greater than eye diameter. Three or 4 pairs of small recurved canines anteriorly in upper jaw. Eye diameter
2.9 to 4.3 times in head length; lower margin of eye tangent to or just above a line from tip of snout to
upper base of pectoral fins; interorbital width 0.9 to 1.8 times in eye diameter; suborbital depth 1.1 to 2.1
times in eye diameter; a line drawn up from the posterior edge of suborbital reaching the dorsal profile
about 3 to 7 scale rows before origin of dorsal fin. Width of naked area of preopercle 1.1 to 1.6 times in
width of scaly area. Total gill rakers on first gill arch 13 to 16. Total pectoral-fin rays 16 or 17; pectoral and
pelvic fins long, reaching to or just short of level of origin of anal fin; pectoral fins 0.9 to 1.1 times in
head length, pelvic fins 1 to 1.2 times in head length; caudal fin forked, upper rays prolonged and pointed,
but not forming a filament. Colour: body pinkish, silvery below; 2 narrow golden yellow stripes on sides,
the first from above upper preopercle margin to upper caudal-fin base, the second from behind upper
opercle margin, curving downwards slightly and extending to middle of caudal-fin base; cheeks and
opercle tinged with bluish gold; dorsal fin pale pinkish yellow, with bright yellow margin and pale dusky
submarginal stripe; caudal pinkish, upper tip bright yellow; anal fin translucent whitish; pelvic fins
translucent whitish, yellow stripe extending down second soft ray, axillary scale yellow; pectoral fins pale
pink.
Habitat, biology, and fisheries: A benthic species, occurs on sand or mud bottoms in depths of about 60
to 80 m. Estimates of asymptotic length (L¥), asymptotic weight (W¥), coefficient of growth (K), and rate of
natural mortality made for the population off Kalimantan in the South China Sea are: L¥ = 24.5 cm total
length, W¥ = 210 g, K = 0.42, M = 1.73. Taken by handline and bottom trawl. This species constitutes about
20% of the total nemipterid
catch in the South China Sea
off Kalimantan, but catch rates
are low (about 3.5 kg/hour at a
depth of 45 m). No major fish-
ery exists.
cluding southern Japan, Gulf
of Thailand, South China Sea,
Strait of Malacca, and Indone-
sia.
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3074 Bony Fishes
Nemipterus virgatus (Houttuyn, 1782) (Plate XXII, 163)

FAO names: En - Golden threadfin bream.
Diagnostic characters: Body depth 3.2 to 4 times in standard length. Snout length greater than eye
diameter. Three or 4 pairs of small recurved canines anteriorly in upper jaw. Eye diameter 3.4 to 4.3 times
in head length; lower margin of eye above a line from tip of snout to upper base of pectoral fins;
interorbital width 1.4 to 2.1 times in eye diameter; suborbital depth 1 to 1.7 times in eye diameter; a line
drawn up from the posterior edge of suborbital reaching the dorsal profile 5 to 7 scale rows before
origin of dorsal fin. Width of naked area of preopercle 1.1 to 2 times in width of scaly area. Total gill rakers
on first gill arch 12 to 16. Total pectoral-fin rays 16 or 18, pectoral and pelvic fins long, reaching to
between level of anus and origin of anal fin; pectoral fins 1 to 1.2 times in head length, pelvic fins 1 to
2.5 times in head length; anal fin with 8 rays; caudal fin moderately forked, upper lobe produced into
trailing filament. Colour: body pink, paler below; yellow stripe on back above lateral line; 5 yellow
stripes on sides beneath lateral line, the uppermost of these stripes beginning just beneath origin
of lateral line and running horizontally to upper base of caudal peduncle, the remaining stripes
somewhat concave and descending anteriorly; ventral surface white; head pink, a yellow stripe from
upper lip extending to anteroventral margin of eye; 2 faint yellow bars across cheeks sometimes
present; eye pinkish; upper lip yellow; dorsal fin pale pink, with broad yellow margin, edged below by
red anteriorly; a yellow stripe just above base of dorsal fin; anal fin translucent pinkish with yellow
submarginal stripe, and narrow yellow stripe just above base of fin, this extending to tip of last
anal-fin ray; caudal fin pink, upper margin of fin and filament yellow; pelvic fins pinkish, with yellow
stripe along second and third rays; pectoral fins translucent pinkish.
Habitat, biology, and fisheries: A benthic species, occurs on mud or sand bottoms in depths to about
220 m. Very abundant in the northern part of the South China Sea and southern part of the East China Sea.
Young fishes generally occur in shallower water, between 18 to 33 m. Females predominate at small sizes
and males at larger sizes, due to faster growth rates in males. This species is a rudimentary hermaphrodite,
in which all males have functional testes with rudimentary ovarian portions throughout their life. Spawning
takes place from February to June with a peak from February to April in the South China Sea. The diet
consists of crustaceans, fishes, and cephalopods. Estimates of the parameters of the von Bertalanffy growth
curve made for the population from Hong Kong are: L¥ = 44.7 cm total length, K = 0.41, t0 = - 0.12 (males);
and L¥ = 36.2 cm total length, K = 0.35, t0 = - 0.71 (females). Taken in commercial quantities by handlines,
longlines, and bottom trawls. Juveniles are also caught in considerable quantities by shrimp trawlers using
beam trawls. N. virgatus is one
of the most important commer-
cial fishes in the East China Sea
and northern South China Sea.
Marketed mainly fresh, but also
steamed or made into fish balls.
Distribution: Southern Japan,
East China Sea, northern part
of South China Sea from Viet-
nam to Formosa Strait, north-
western Australia, and Arafura
Sea.
Pentapodus setosus (Valenciennes, 1830) (Plate XXIII, 173)

FAO names: En - Butterfly whiptail.
Diagnostic characters: Snout length greater than eye diameter; head scales reaching forward to
between level of anterior margin of eyes and posterior nostrils; suborbital naked; lower limb of
preopercle naked. Total pectoral-fin rays 16 or 17 (usually 16); pelvic fins short, not reaching level of anus;
caudal fin forked, upper lobe produced into a very long trailing filament. Lateral-line scales 46 to 48
(usually 46 or 47). Colour: pale brownish on back; lower part of body whitish; a blue stripe along base of
dorsal fin; a yellow stripe from behind eye, gradually arching on back and terminating in a black spot
on upper caudal peduncle; a narrow blue line running through yellow stripe, this line convergent
with a blue line from origin of anal fin, both lines meeting at an acute angle behind black spot on
caudal peduncle; 2 bluish stripes across snout, first from middle of eye to tip of snout, second from
upper lip to lower margin of eye; top of snout dusky; caudal fin pinkish, filament pinkish brown.
Habitat, biology, and fisheries: A benthic species, inhabiting offshore waters. Juveniles occur close to
reefs. Food consists of small crustaceans. Trawled in fair numbers in the South China Sea and outer Gulf
of Thailand. Also taken by bamboo stake trap (Thailand). Not highly rated as food and brings a low price.
No major fishery exists. Used in fish balls, in fish meal, fried, and as duck food.
Distribution: P h i l i p p i n e s,
South China Sea, Singapore,
and Indonesia.
3076 Bony Fishes
Scolopsis monogramma (Kuhl and Van Hasselt, 1830) (Plate XXIV, 181)
Frequent synonyms / misidentifications: Scolopsis regina Whitley (1937) / This species has been
previously misidentified as S. temporalis.
FAO names: En - Monogrammed monocle bream.
Diagnostic characters: Body depth 2.5 to 3 times in standard length. Snout length greater than eye
diameter; head scales reaching to between anterior margin of eyes and posterior nostrils; lower limb
of preopercle scaly; antrorse suborbital spine absent; suborbital depth 1.2 to 2.1 times in eye diameter.
Total pectoral-fin rays 17 to 19 (usually 18); pelvic fins long, reaching almost to level of origin of anal fin;
caudal fin forked or lunate, upper lobe a little longer than lower lobe (lobes produced to form short
filamentous extensions in larger specimens). Lateral-line scales 46 to 49 (usually 47 or 48). Colour:
greyish on back, white below; brown longitudinal steaks on back above lateral line, and oblique
yellow streaks below lateral line; a brown midlateral stripe, expanded in the middle; 3 blue stripes
on snout; uppermost joining eyes above nostrils, middle stripe joining eyes through nostrils, lower
stripe from eye to tip of snout; interspaces between stripes yellow; a blue stripe on preopercle behind
eye; a blue chevron-shaped stripe running upwards onto opercle from below eye and bending
downwards towards pectoral-fin base; space between stripes on preopercle and opercle yellow; a
brown bar at base of pectoral fins; unpaired fins pale yellow, edged with blue.
Size: Maximum standard length 26 cm, commonly to 18 cm standard length.
Habitat, biology, and fisheries: A benthic species found on sand bottoms close to reefs in depths to 50 m.
Occurs solitary or in small groups. Feeds on small fishes, crustaceans, molluscs, and polychaetes. A
protogynous hermaphrodite. Appears in small numbers in local markets. Caught by trawl and bamboo stake
trap (Thailand), and by handline. Sold fresh. Prepared fried and in fish balls or used as duck food (Thailand).
No major fishery exists.
Distribution: We st Pacific
from Ryukyu Islands to north-
eastern Australia, and Eastern
Indian Ocean from the An-
daman Sea to northwestern
Australia.
Scolopsis taeniopterus (Kuhl and Van Hasselt, 1830) (Plate XXIV, 182)
Frequent synonyms / misidentifications: Scolopsis siamensis Akazaki, 1962 / This species has been
misidentified as Scolopsis cancellatus (= S. lineatus).
FAO names: En - Lattice monocle bream.
Diagnostic characters: Body depth 2.7 to 3.1 times in standard length. Snout length about equal to eye
diameter; head scales reaching forward to between level of anterior margin of eyes and posterior
nostrils; lower limb of preopercle naked; antrorse suborbital spine absent; suborbital depth 1.1 to 3.5
times in eye diameter. Total pectoral-fin rays 17 or 18 (usually 17); pelvic fins long, reaching to or beyond
level of anus; caudal fin emarginate. Lateral-line scales 45 to 48. Colour: greyish yellow on upper part of
body, whitish below; sides of body with faint oblique blue and yellow lines, these horizontal on the
caudal peduncle; a narrow blue stripe joining eyes just behind nostrils; a blue stripe from middle of
upper lip to lower edge of eye; upper part of pectoral-fin base with a reddish orange spot; fins
yellowish; dorsal fin with a blue stripe along its middle area; upper tip of caudal fin bright yellow;
upper base of caudal fin with a blue spot.
Habitat, biology, and fisheries: A benthic species, found on sand or mud bottoms in offshore areas, in
depths to 50 m. Swims in small groups. Feeds on small crustaceans, molluscs and fishes. A protogynous
hermaphrodite. This is the most abundant species of Scolopsis seen in markets in Singapore, Malaysia, and
Thailand. Trawled in moderate numbers along with other species in the South China Sea and Gulf of
Thailand. Highly rated as a food fish, although it sells for a low price. Sold fresh. It is prepared steamed or
used for fish balls; sometimes salted or dried. No major fishery exists.
Distribution: West Pacific,
from Taiwan Province of
China to northeastern Austra-
lia, and northwestern Austra-
lia.
3078 Bony Fishes
Nemipterus aurifilum (Ogilby, 1910)

En - Yellowlip threadfin bream.
Maximum standard length 18.5 cm, commonly to 14 cm. A benthic species, found on sand or mud
bottoms in depths of 24 to 220 m. Feeds on small crustaceans. Taken by handline and bottom trawl
in deeper water off the Queensland coast. Considered a good food fish. No major fishery exists.
Eastern Australia, from southern Queensland to northern New South Wales.
Nemipterus balinensis (Bleeker, 1858-9) (Plate XX, 147)

En - Balinese threadfin bream.
Maximum standard length 18 cm, commonly to 12 cm. A benthic species, found on sand or mud
bottoms in depths of 50 to 150 m. Occasionally taken by trawling in inshore waters, but taken mainly
by handline. Appears in small numbers in local markets. No major fishery exists. Southern
Indonesia, Sumatra to Flores.
Nemipterus balinensoides (Popta, 1918)

En - Dwarf threadfin bream.
bottoms in depths between 30 to 80 m. Occasionally taken by trawling in shallow inshore waters.
Rarely seen in markets. No major fishery exists. West Pacific, including the Philippines, Indonesia,
New Caledonia, and Phuket Island, Andaman Sea.
Nemipterus gracilis (Bleeker, 1873) (Plate XX, 151)

En - Graceful threadfin bream.
Maximum standard length 18 cm, commonly to 15 cm. A benthic species, occurs on mud or sand
bottoms in depths of 30 to 90 m. Females with ripe ova were present in June in Flores. Taken by
handline. Appears in small quantities in local markets. No major fishery exists. Indonesia: Sumatra
to Flores.
Nemipterus marginatus (Valenciennes, 1830) (Plate XXI, 155)

En - Red filament threadfin bream.
bottoms in depths between 12 to 70 m. Occurs in groups and feeds on small bottom-living animals.
Commonly taken by trawlers in the South China Sea. This species is taken in small quantities
(usually less than 1 kg per haul in South China Sea) along with other species. Regarded as a trash
fish and used mainly for fish meal or animal feed. West Pacific, including the South China Sea, Strait
of Malacca, southern coasts of Sumatra and Java, northern Australia, and the Gulf of Papua to the
Solomon Islands.
Nemipterus mesoprion (Bleeker, 1853) (Plate XXI, 156)

En - Mauvelip threadfin bream.
Maximum standard length 14 cm, commonly to 13 cm. A benthic species. Little is known of the
habits of this species. Taken by handline and bottom trawl, together with other species. Known from
southern Indonesia and the Gulf of Thailand.
3080 Bony Fishes
Nemipterus nematopus (Bleeker, 1851) (Plate XXI, 158)

En - Yellowtipped threadfin bream.
Maximum standard length 17.5 cm, commonly to 15 cm. A benthic species, occurs on sand or mud
bottoms in depths of 30 to 102 m. Taken by handline and bottom trawl together with other species.
Appears in small numbers in local markets. No major fishery exists. Philippines, Indonesia, and
northern Australia from the Timor Sea to northern Queensland.
Nemipterus sp. 1
En - Bluecheek threadfin bream.
Maximum standard length 26 cm, commonly to 15 cm. A benthic species, occurs in depths of 65 to
125 m. Taken so far only by exploratory bottom trawling. Not seen in local markets. No major fishery
exists. So far known only from Indonesia: southern coasts of eastern Java, Bali, and Lombok.
Nemipterus theodorei Ogilby, 1916

En - Theodore’s threadfin bream.
Maximum standard length 20 cm, commonly to 18 cm. A benthic species, occurs on sand or mud
bottoms in depths of 19 to 410 m. Taken by handline and bottom trawl. No major fishery exists.
Eastern Australia from north Queensland to New South Wales.
Nemipterus vitiensis Russell, 1990 (Plate XXII, 164)

En - Fiji threadfin bream.
Maximum standard length 22 cm, commonly to 17 cm. A benthic species, occurs on sand or mud
bottoms close to reefs in depths of 27 to 273 m. Taken by handline. Appears in local fish markets.
No major fishery exists. Fiji and Vanuatu.
Nemipterus zysron (Bleeker, 1856-57) (Plate XXII, 165)

En - Slender threadfin bream; Fr - Cohana élégante; Sp - Baga elegante.
Maximum standard length 18.5 cm, commonly to 16 cm. A benthic species, found on sand bottoms
near rocks in depths between 10 to 125 m. It swims in groups and feeds on bottom organisms.
Taken by handline and bottom trawl along with other species of Nemipterus. Appears in small
numbers in local markets. No major fishery exists. Widespread in the Indo-West Pacific from
northwestern Australia, the Indo-Malay Archipelago, Andaman Sea, Red Sea, Seychelles, and
Zanzibar.
Parascolopsis eriomma (Jordan and Richardson, 1909) (Plate XXII, 166)

En - Rosy dwarf monocle bream; Fr - Mamila rosée; Sp - Besugato rosado.
bottoms in offshore waters in depths of 25 to 175 m. Feeds mainly on benthic invertebrates. Taken
by vertical long line and by bottom trawl. No major fishery exists. Indian Ocean, including the Gulf
of Oman, Red Sea, east Africa (south to Delagoa Bay), Sri Lanka, and Andaman Sea; West Pacific,
including southern Japan, Taiwan Province of China, South China Sea, Philippines, and Indonesia.
3082 Bony Fishes
Parascolopsis inermis (Schlegel, 1843) (Plate XXII, 167)

En - Unarmed dwarf monocle bream.
bottoms in offshore waters in depths of 60 to 131 m. Feeds mainly on benthic invertebrates. Taken
by bottom trawl, deep-sea shrimp trap and vertical long line. No major fishery exists. Eastern Indian
Ocean, including the Laccadive Islands, Sri Lanka, and Andaman Sea; West Pacific, including
southern Japan and the South China Sea.
Parascolopsis melanophyrs Russell and Chin, 1996 (Plate XXII, 168)

En - Black eyebrow monocle bream.
Maximum standard length 17 cm. A little known but distinctive species. No major fishery exists.
Known only from 2 collected specimens from fish markets in Sabah (Malaysia) and West Timor
(Indonesia).
Parascolopsis rufomaculatus Russell, 1986

En - Redspot dwarf monacle bream.
Maximum standard length 15.5 cm, commonly to 10 cm. A benthic species, occurring in offshore
waters in depths of 200 to 350 m. Feeds mainly on benthic invertebrates. Taken by bottom trawl
(scampi net). No fishery exists. Known so far only from northwestern Australia.
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Parascolopsis tanyactis Russell, 1986

En - Longrayed dwarf monocle bream.
Maximum standard length 20.5 cm, commonly to 15 cm. A benthic species, occurring in offshore
waters in depths of 40 to 200 m. Feeds mainly on benthic invertebrates. Taken by bottom trawl. No
fishery exists. Western Pacific, including the Philippines, eastern Indonesia and northwestern
Australia.
Parascolopsis tosensis (Kamohara, 1938)

En - Tosa dwarf monocle bream.
Maximum standard length 10 cm, commonly to 8 cm. A benthic species occurring in offshore waters
in depths of 150 to 300 m. Very rarely found in markets. No major fishery exists. West Pacific,
including southern Japan, Taiwan Province of China, the Philippines, and eastern Indonesia.
Pentapodus bifasciatus (Bleeker, 1848) (Plate XXII, 169)

En - Whiteshouldered whiptail.
Maximum standard length 15 cm, commonly to 10 cm. A benthic species inhabiting shallow coral
reef areas. Food consists of small fishes, crustaceans (shrimps, caprellid amphipods), and poly-
chaetes. Appears occasionally in small numbers in local markets. No major fishery exists. Philip-
pines, Indonesia, and Singapore.
3084 Bony Fishes
Pentapodus caninus (Cuvier, 1830) (Plate XXIII, 170)

En - Smalltoothed whiptail.
Maximum standard length 18.5 cm, commonly to 15 cm. A free-swimming species which occurs
solitary or in small groups. Usually found close to the bottom over coral reef areas in depths of 15 m
or more. Food consists of small fishes and larger zooplankton as well as bottom-living organisms.
Taken by handline. Appears occasionally in small numbers in local markets. No major fishery exists.
West Pacific, including New Caledonia, Vanuatu, Solomon Islands, Papua New Guinea, Marshall
Islands, Palau, southern Japan, Philippines, and Indonesia.
Pentapodus emeryii (Richardson, 1843) (Plate XXIII, 171)

En - Double whiptail.
Maximum standard length 24.5 cm, commonly to 18 cm. A benthic species found close to reef areas.
Food consists of small fishes, crustaceans, ophiuroids, and sipunculid worms. Appears occasionally
in small numbers in local markets. No major fishery exists. Northwestern Australia, Indonesia, and
Philippines.
Pentapodus nagasakiensis (Tanaka, 1915)

En - Japanese whiptail.
Maximum standard length 15 cm, commonly to 10 cm. Usually inhabits deeper offshore waters, in
depths of 40 to 100 m. Food consists of small crustaceans (shrimps). Rarely caught. No major
fishery exists. Southern Japan, South China Sea, Indonesia (Lombok), and northwestern Australia.
Pentapodus paradiseus (Günther, 1859) (Plate XXIII, 172)

En - Paradise whiptail.
Maximum standard length 20 cm, commonly to 15 cm. A benthic species found close to reefs. Food
consists of small fishes, crustaceans, and polychaetes. Appears occasionally in small numbers in
local markets. Taken by handline by recreational fishermen in southern Queensland, Australia. No
major fishery exists. Northeastern Australia, southern Papua New Guinea, Solomon Islands, and
Philippines.
Pentapodus porosus (Valenciennes, 1830)

En - Northwest Australian whiptail.
Maximum standard length 23 cm, commonly to 15 cm. A benthic species found close to reefs in
offshore waters. Trawled in small numbers in offshore water, or taken by handline close to reefs.
No major fishery exists. Northwestern Australia and Aru Islands.
Pentapodus sp.
En - Yellowstriped whiptail.
Maximum standard length 16 cm, commonly to 10 cm. A benthic species inhabiting sand bottoms
adjacent to coral reefs. Appears occasionally in small numbers in local markets. No major fishery
exists. Taiwan Province of China, Philippines, eastern Indonesia (Ambon), Papua New Guinea,
eastern Australia, New Caledonia, Fiji, Tonga, and Samoa.
3086 Bony Fishes
Pentapodus trivittatus (Bloch, 1791) (Plate XXIII, 174)

En - Threestriped whiptail.
Maximum standard length 19 cm, commonly to 15 cm. A benthic species, common in shallow sandy
rubble and coral reef areas, in depths to 15 m. Often occurs in aggregations. Food consists of small
fishes, crustaceans, and polychaetes. Appears in small numbers in local markets. No major fishery
exists. Solomon Islands, New Guinea, Western Caroline Islands (Palau, Yap), Philippines, East
Malaysia, Indonesia, and Singapore.
Scaevius milii (Bory de Saint-Vincent, 1823) (Plate XXIII, 175)

En - Greenstriped coral bream.
Maximum standard length 20 cm, commonly to 15 cm. A benthic species, very common on shallow
inshore reefs and adjacent sand or mud bottoms, in depths to 20 m. Feeds on small fishes and
benthic invertebrates. Unimportant as a food fish. No major fishery exists. Northwestern Australia:
Gulf of Carpentaria to Abrolhos Islands.
Scolopsis affinis Peters, 1877 (Plate XXIII, 176)

En - Peters’ monocle bream.
Maximum standard length 19.5 cm. A benthic species, found on sand or mud bottoms close to reefs,
in depths to 60 m. Occurs solitary or in small aggregations. Appears occasionally in small numbers
in local markets. No major fishery exists. Western Pacific Ocean, including Ryukyu Islands,
Philippines, South China Sea, Indonesia, New Guinea, Solomon Islands, northeastern Australia,
and Andaman Islands.
Scolopsis auratus (Park, 1797) (Plate XXIII, 177)

En - Yellowstripe monocle bream; Fr - Mamila à bande dorée; Sp - Besugato de banda dorada.
Maximum standard length 21 cm, commonly to 18 cm. A benthic species, found close to reefs.
Appears occasionally in small numbers in local markets. No major fishery exists. Eastern Indian
Ocean, including the Maldives, Sri Lanka, and southern Indonesia.
Scolopsis bilineatus (Bloch, 1793) (Plate XXIV, 178)

En - Twolined monocle bream; Fr - Mamila griffée; Sp - Besugato rayado.
Maximum standard length 15.5 cm, commonly to 13 cm. A benthic species, common on coral reefs,
in depths to 20 m. Occurs solitary or in small aggregations. Feeds on small fishes and benthic
invertebrates. A protogynous hermaphrodite. Juveniles appear to be Batesian mimics of poison-
fanged blenniids (Meiacanthus). Appears occasionally in small numbers in local markets. Caught
mainly by handline. Live specimens captured by hand net for the aquarium export trade in the
Philippines. No major fishery exists. Western Pacific from the Ryukyu Islands to eastern Australia
and eastwards to New Caledonia and Fiji; also Indian Ocean from the Laccadive Islands, Sri Lanka,
Andaman Sea, and Western Australia.
Scolopsis ciliatus (Lacepède, 1802) (Plate XXIV, 179)

En - Sawjawed monocle bream.
Maximum standard length 13.5 cm, commonly to 10 cm. A benthic species, found on sandy bottoms
close to coral reefs. Occurs often in small groups. Feeds on benthic invertebrates and small fishes.
Appears occasionally in small numbers in local markets. Also taken incidentally by trawlers in the
Gulf of Thailand. No major fishery exists. Andaman Sea and Western Pacific, including the Ryukyu
Islands, Philippines, Indonesia, Solomon Islands, and Vanuatu.
3088 Bony Fishes
Scolopsis lineatus Quoy and Gaimard (1824) (Plate XXIV, 180)

En - Striped monocle bream; Fr - Scolopsis rayé.
Maximum standard length 18 cm, commonly to 13 cm. A benthic species, common on coral reefs
usually associated with sandy areas, in depths to 20 m. Occurs in pairs or small groups. Feeds on
small fishes and crustaceans. Appears occasionally in small numbers in local markets. No major
fishery exists. West Pacific from the Ryukyu Islands to the Marshall Islands and Vanuatu, and the
southeastern Indian Ocean including the Cocos-Keeling Islands and northwestern Australia. Record
from the Andaman Islands is unconfirmed.
Scolopsis margaritifer (Cuvier, 1830)

En - Pearly monocle bream.
Maximum standard length 17.5 cm, commonly to 15 cm. A benthic species found on sand bottoms
close to reefs, in depths to 20 m. Usually solitary. Feeds on crustaceans, polychaetes, molluscs,
and small fishes. Juveniles appear to be Batesian mimics of poison-fanged blenniids (Meiacanthus).
Appears occasionally in small numbers in local markets. Caught incidentally by trawlers in the Gulf
of Thailand. No major fishery exists. West Pacific from the South China Sea to Vanuatu, and
northwestern Australia.
Scolopsis temporalis (Cuvier, 1830)

En - Baldspot monocle bream.
Maximum standard length 28 cm, commonly to 20 cm. A benthic species found on sand bottoms
close to reefs in depths to 30 m. No major fishery exists. Eastern Indonesia, Solomon Islands, Santa
Cruz Islands, and Fiji.
Scolopsis trilineatus Kner, 1868 (Plate XXIV, 183)

En - Threelined monocle bream; Fr - Scolopsis à trois lignes.
Maximum standard length 16.5 cm, commonly to 14 cm. A benthic species, found on sand bottoms
in lagoons and close to reefs, in depths to 20 m. Occurs solitary or in small aggregations. Appears
occasionally in small numbers in local markets. No major fishery exists. Western Pacific, including
the Philippines, Western Caroline Islands, South China Sea, eastern Indonesia, northern Australia,
New Guinea, New Caledonia, Fiji, Samoa, Tonga, and northwestern Australia.
Scolopsis vosmeri (Bloch, 1792) (Plate XXIV, 184)

En - Whitecheek monocle bream; Fr - Mamila; Sp - Besugato cariblanco.
Maximum standard length 16 cm, commonly to 15 cm. A benthic species, found in inshore waters
usually on sand or mud bottoms close to reefs, as well as in offshore areas. Taken by bottom trawls,
handlines, and traps. Caught in wire traps and trawls (Thailand). Appears occasionally in small
numbers in local markets. In Thailand, this species is steamed or made into fish balls. No major
fishery exists. Indo-West Pacific, from the east African coast, Red Sea, and Persian Gulf to northern
Australia.
Scolopsis xenochrous Günther, 1872 (Plate XXIV, 185)

En - Obliquebarred monocle bream.
Maximum standard length 17 cm, commonly to 14 cm. A benthic species found around rocky-rubble
and seaweed areas as well as coral reefs, in depths of 5 to 50 m. Feeds mainly on benthic
crustaceans. Appears occasionally in small numbers in local markets. No major fishery exists. Indian
Ocean, including the Maldives, Sri Lanka, Andaman Sea, and northwestern Australia. West Pacific,
including Taiwan Province of China, the Philippines, Indonesia, New Guinea, northeastern Australia,
and the Solomon Islands.
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3090 Bony Fishes
POLYNEMIDAE
Threadfins
by R.M. Feltes
D iagnostic characters: Perciform fishes with oblong, somewhat compressed body (size from 17 to
200 cm). Eye covered by adipose tissue, highly variable in size. Snout conical, protruding anterior
past mouth. Mouth large, subterminal, extending posterior to eye. Supramaxillae absent. Cardiform teeth
on premaxillae, palatines, and ectopterygoids. Presence of teeth on vomer, basibranchials and gill arches
variable. Branchiostegal rays 7. Two widely separated dorsal fins; second or third spines of first dorsal fin
longest; margins of second dorsal fin and anal fin variously concave, anterior rays longest; first dorsal fin
with VII or VIII spines, if VIII, then first spine very small; second dorsal fin with I spine and 11 to 18 soft
rays; anal-fin insertion ventral to a point 1/4 to 1/3 caudad on second dorsal-fin base, anal fin with II or III
spines (if III, then first spine very small) and 9 to 30 soft rays; caudal fin deeply forked with pointed lobes,
principal caudal-fin rays 17; 3 to 16 separate articulated pectoral filaments inferior to 12 to 19 rays of
pectoral fins; pelvic fins abdominal, inserted just behind pectoral-fin bases, with I spine and 5 branched
rays. Body, most of head, and much of fins covered with finely ctenoid scales; lateral line continuous, and
extending to caudal-fin margin; lateral-line scales 42 to 109; scale rows above lateral line 4 to 9; scale rows
below lateral line 7 to 15. Nasal bones anterior with lateral aspects surrounding anterior of nasal capsules.
Long posterior process of coracoid extends dorsally, medial to pectoral radials. Fourth pectoral radials
elongate. Basipterygia not in direct contact with cleithra, but in ligamentous contact with second post-
cleithra. Vertebrae 10+14 or 10+15. Colour: silvery, golden, or light brown; some with dark spot at anterior
of lateral line or with longitudinal lines; dark silvery spot on opercle; fins usually yellow, orange, or brown,
and often dusky.
2 dorsal fins, the 1st with VII-VIII spines,

the 2nd with I spine and 11-18 soft rays
lateral line extends to posterior

eye covered by margin of caudal fin
adipose tissue
lower pectoral-fin rays anal fin with II-III spines

filamentous, separated from fin and 9-30 soft rays
Habitat, biology, and fisheries: Often inhabit sand and mud flats. May enter estuaries or rivers.
Development without marked metamorphosis. No external sexual dimorphism. Some species hermaphro-
dites. Recorded life span from 1 to 20 years. Many species feed largely on crustaceans and predominantly
prawns. From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports a range of yearly catch of
around 20 700 to 34 700 t of Polynemidae from the Western Central Pacific. The swimbladders of
polynemids have been valued for isinglass.
Remarks: The genus Polynemus Linnaeus, 1758 included 3 quite divergent species that are today
recognized in 3 distinct genera: Polynemus, Polydactylus, and Pentanemus. The International Commission
for Zoological Nomenclature established Polynemus paradiseus as the type of that genus in 1926. In spite
of this ruling many species have continued to be indiscriminately assigned to Polynemus.
Perciformes: Percoidei: Polynemidae 3091
Similar families occurring in the area: Superficially, threadfins can resemble other silvery fishes in the
area (e.g. certain species of Sciaenidae) but the combination of 2 separate dorsal fins and separate pectoral
filaments easily distinguishes them. Triglidae and Dactylopteridae also possess free pectoral filaments but
they have strong bony armour on their heads and dissimilar body shapes.
Key to the species of Polynemidae occurring in the area
1a. Pectoral-fin insertion well below midline of body; eye diameter variable, 1.3 or less in
snout length (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Pectoral-fin insertion near midline of body; eye diameter 1.3 or more in snout length
(Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
pectoral-fin
insertion below pectoral-fin
midline of body insertion near
midline of body
Fig. 1 Polydactylus plebeius Fig. 2 Polynemus dubius
2a. Teeth extending labially on anterior two-thirds of dentary; primary tooth

tooth plates extending posteriorly from lateral margin of plate on
primary tooth plate on vomer (Fig. 3); free lower lip re- vomer
stricted to posterior third of lower jaw; total gill rakers on first
gill arch less than 20 (most gill rakers replaced during
growth by tooth plates); oral cavity largely covered by small
tooth plates; pectoral filaments 3 or 4 . . . . . . . (Eleutheronema) ® 3
2b. Teeth on dentary restricted to dorsal surface or extending
labially only along anterior third of lower jaw; tooth plates on
vomer simple or absent; total gill rakers on first gill arch 18
to 60; oral cavity not largely covered by small tooth plates;
pectoral filaments 4 to 10 (typically) . . . . . . . . . . . . . . . . . ® 4 posterior tooth plates
Fig. 3 tooth plates on

3a. Pectoral filaments 3 (Fig. 4) . . . . . . . . . Eleutheronema tridactylum
roof of mouth
3b. Pectoral filaments 4 (Fig. 5) . . . . . . . . Eleutheronema tetradactylum
3 pectoral filaments 4 pectoral filaments

Fig. 4 Eleutheronema tridactylum Fig. 5 Eleutheronema tetradactylum
4a. Width of tooth bands on upper and lower jaws 2 times or more in space separating band
of teeth on one premaxilla from other (Fig. 6a); tooth plate on palatine shorter than tooth
plate on ectopterygoid; basisphenoid does not contact prootic (Fig. 7a); pectoral fila-
ments 5 to 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Filimanus) ® 5
4b. Width of tooth bands on upper and lower jaws less than 2 times in space separating band
of teeth on one premaxilla from other (Fig. 6b); tooth plate on palatine equal or longer
than tooth plate on ectopterygoid; basisphenoid contacts prootic (Fig. 7b); pectoral
filaments 5 to 10 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Polydactylus) ® 9
3092 Bony Fishes
upper-jaw
tooth bands
teeth on
palatine
a) Filimanus prootic
basisphenoid
teeth on
ecto-
pterygoid
a) Filimanus b) Polydactylus
b) Polydactylus
Fig. 6 tooth bands on upper and lower jaws Fig. 7 lateral view of neurocranium
5a. Pectoral filaments extending well past midpoint of anal fin; anal-fin rays 13 to 15; depth
of posterior margin of maxilla greater than or equal to eye diameter; snout blunt (Fig. 8) . . . . . ® 6
5b. Pectoral filaments not extending to midpoint of anal fin; anal-fin rays 10 to 12; depth of
posterior margin of maxilla less than eye diameter; snout protruding (Fig. 9) . . . . . . . . . . . ® 7
snout
snout protruding
blunt
13-15 anal-fin rays 11-12 anal-fin rays
long pectoral filaments short or moderate

pectoral filaments
Fig. 8 Filimanus hexanema Fig. 9 Filimanus heptadactyla
6a. Pectoral filaments 7; body depth less than 3.2 times in standard length . . . . . Filimanus perplexa
6b. Pectoral filaments 6; body depth more than 3.2 times in standard length . . . . Filimanus hexanema
7a. Pectoral filaments 8 (rarely 7); gill rakers on first gill arch 40 to 48 (mean 46.2); body blue
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filimanus sealei
7b. Pectoral filaments 6 or 7 (rarely 5); gill rakers on first gill arch 35 to 49; body yellow,
green, or silver . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Pectoral filaments 6 (occasionally 5), or asymmetrically 5 and 6, or 6 and 7; gill rakers
on first gill arch 36 to 46 (mean 41.8); body usually dusky yellow or green dorsally, silver
ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Filimanus xanthonema
(eastern Indian and southwestern Pacific oceans)
8b. Pectoral filaments 7; gill rakers on first gill arch 35 to 41 (mean 38.6); body usually brown
dorsally, yellow or silver ventrally . . . . . . . . . . . . . . . . . . . . . . . Filimanus heptadactyla
preopercular margin
9a. Posterior margin of preopercle largely en- with less than 10 spines
tire and covered by scales except ven-
trally where there are less than 10 small
spines; vomer lacking tooth patch; pecto-
ral filaments 6 (Fig. 10) . . . . Polydactylus nigripinnis
9b. Posterior margin of preopercle with serra-
tions on most of posterior margin, usually
with more than 10 small spines not cov-
ered by scales; vomer with or without
tooth patch . . . . . . . . . . . . . . . . . . . . .® 10 Fig. 10 Polydactylus nigripinnis
10a. Upper pectoral filaments longer than standard length; lateral-line scales 87 to 92;
pectoral filaments 7 (Fig. 11) . . . . . . . . . . . . . . . . . . . . . . Polydactylus macrophthalmus
10b. Upper pectoral filaments much shorter than standard length; lateral-line scales less than
80 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
11a. Caudal-peduncle length less than length of anal-fin base; anal-fin rays 14 to 18; pectoral
filaments 6 or 7 (usually 7) (Fig. 12) . . . . . . . . . . . . . . . . . . . . Polydactylus multiradiatus
11b. Caudal-peduncle length equal to or greater than length of anal-fin base; anal-fin rays 9
to 13 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
87-92 lateral-line scales
Fig. 11 Polydactylus macrophthalmus Fig. 12 Polydactylus multiradiatus
12a. Pigmented spot over several anterior dark spot

lateral-line scales; pectoral-filament
length less than head length; pecto-
ral filaments 5 or 6 (Fig. 13) . . . . . . . . . ® 13
12b. No pigmented spot over several an-
terior lateral-line scales; pectoral-fila-
ment length usually greater than
head length . . . . . . . . . . . . . . . . . . ® 14
13a. Pectoral filaments 5 . . . Polydactylus microstoma Fig. 13 Polydactylus sextarius

13b. Pectoral filaments 6 . . . . Polydactylus sextarius
14a. Length of postorbital greater than 3/4 of body depth at origin of second dorsal fin;
diameter of eye less than snout length; 2 times length of anal-fin base is less than
distance from pelvic-fin origin to anal-fin origin; filaments may extend from caudal-fin
lobes; pectoral filaments 5 (Fig. 14); swimbladder with lateral appendages (Fig. 15) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus indicus
14b. Length of postorbital usually less than or equal to 3/4 of body depth at origin of second
dorsal fin; diameter of eye greater than or equal to snout length; 2 times length of anal-fin
base is greater than distance from pelvic-fin origin to anal-fin origin; no filaments
extending from caudal-fin lobes; swimbladder without lateral appendages . . . . . . . . . . . . ® 15
caudal-fin filaments
Fig. 14 Polydactylus indicus Fig. 15 swimbladder (Polydactylus indicus)

3094 Bony Fishes
15a. Body depth at second dorsal-fin origin less than distance from pelvic-fin origin to anal-fin
origin; length of postorbital greater than distance from anterior of snout to posterior
margin of preopercle; pectoral-fin rays 13 to 15; pectoral filaments 4 or 5; gill rakers on
first gill arch 31 to 37; adults without numerous darker longitudinal lines on body (Fig. 16)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus macrochir
15b. Body depth at second dorsal-fin origin greater than distance from pelvic-fin origin to
anal-fin origin; length of postorbital less than distance from anterior of snout to posterior
margin of preopercle; pectoral-fin rays 14 to 18; pectoral filaments 5 or 6; gill rakers on
first gill arch 19 to 32; adults with numerous darker longitudinal lines on body . . . . . . . . . . ® 16
16a. Usually 5 pectoral filaments; gill rakers on first gill arch 19 to 28 (mean 25.6); pectoral-fin
rays 14 to 18 (mode 17) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus plebeius
16b. Usually 6 pectoral filaments; gill rakers on first gill arch 27 to 32 (mean 29.3); pectoral-fin
rays 14 to 17 (mode 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polydactylus sexfilis
17a. Posterior margin of preopercle largely entire; pectoral-fin rays 12 to 14; procurrent
caudal-fin rays (in dorsal or ventral series) 12 to 14; pectoral filaments 6 or 7 (typically
7); pectoral-filament length less then 1½ total length (Fig. 17) . . . . . . . . Parapolynemus verekeri
17b. Posterior margin of preopercle at least partly serrate; pectoral-fin rays 14 to 19; procur-
rent caudal-fin rays (in dorsal or ventral series) 15 to 19; pectoral filaments 7 to 16
(typically 7); pectoral-filament length may exceed 1½ total length . . . . . . . . . (Polynemus) ® 18
12-14 pectoral-fin rays

12-14 procurrent
caudal-fin rays
Fig. 16 Polydactylus macrochir Fig. 17 Parapolynemus verekeri

18a. Pectoral filaments 14 to 16; upper-jaw length less than or equal to caudal-peduncle
depth (Fig. 18) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynemus multifilis
18b. Pectoral filaments 7; upper-jaw length greater than caudal-peduncle depth . . . . . . . . . . . ® 19
19a. Lateral-line scales more than 88; first dorsal fin with VII spines; anal-fin spines II
(Fig. 19) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynemus hornadayi
19b. Lateral-line scales less than 87; first dorsal fin with VIII spines; anal-fin spines III . . . . . . . . ® 20
more than 88 lateral-line scales
14-17
pectoral
filaments
Fig. 18 Polynemus multifilis Fig. 19 Polynemus hornadayi
20a. Vomer without tooth patch; usually more than 1/2 of posterior margin of pectoral fins
black (for greater than 3 cm), usually not including distal half of dorsalmost rays of fin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Polynemus melanochir
20b. Vomer with tooth patch; pectoral fins usually not black or less than 1/2 of posterior margin
of fin black, dorsalmost rays of fins may be black distally . . . . . . . . . . . . . Polynemus dubius

0 Eleutheronema tetradactylum (Shaw, 1804)
0 Eleutheronema tridactylum (Bleeker, 1849)
0 Filimanus heptadactyla (Cuvier, 1829)
0 Filimanus hexanema (Cuvier, 1829)
0 Filimanus perplexa Feltes, 1991
0 Filimanus sealei (Jordan and Richardson, 1910)
0 Filimanus xanthonema (Valenciennes, 1831)
0 Parapolynemus verekeri (Saville-Kent, 1889)
0 Polydactylus indicus (Shaw, 1804)
0 Polydactylus macrochir (Günther, 1867)
0 Polydactylus macrophthalmus (Bleeker, 1859)
0 Polydactylus microstoma (Bleeker, 1851)
0 Polydactylus multiradiatus (Günther, 1860)
0 Polydactylus nigripinnis Munro, 1964
0 Polydactylus plebeius (Broussonet, 1782)
0 Polydactylus sexfilis (Valenciennes, 1831)
0 Polydactylus sextarius (Bloch, 1801)
0 Polynemus dubius Bleeker, 1851
0 Polynemus hornadayi Myers, 1936
0 Polynemus melanochir Valenciennes, 1831
0 Polynemus multifilis Schlegel, 1843
References
Cantor, T. 1849. Catalogue of Malayan fishes. J. Asiatic Soc. Bengal., 18(2):983-1443.
Feltes, R.M. 1991. Revision of the polynemid fish genus Filimanus, with the description of two new species. Copeia,
1991(2):302-322.
Feltes, R.M. 1992. Filimanus Myers, 1936 (Osteichthyes, Perciformes): proposed designation of Filimanus perplexa
Feltes, 1991 as the type species. Bull. Zool. Nomencl., 49(3):209-210.
Feltes, R.M. 1993. Parapolynemus, a new genus for the polynemid fish previously known as Polynemus verekeri.
Copeia, 1993(1):207-215.
Gloerfelt-Tarp, T. and P.J. Kailola. 1984. Trawled fishes of southern Indonesia and northwestern Australia. Jakarta,
Kailola, P., M. Williams, P. Stewart, R. Reichhelt, A. McNee, and C. Grieve. 1993. Australian fisheries resources.
Australia. Canberra, Australia, Dept. Primary Industries and Energy. Bureau of Resources Sciences, Fisheries
Research and Development Corporation,(10):422 p.
Myers, G.S. 1936. A new polynemid fish collected in the Sadong River, Sarawak by Dr. William T. Hornaday. J. Wash.
Acad. Sci., 26(9):376-382.
Weber, M.W.C. and L.F. De Beaufort. 1922. Fishes of the Indo-Australian Archipelago. IV. Heteromi, Solenichthyes,
Synentognathi, Percesoces, Labyrinthici, Microcyprini. Leiden, Netherlands, E.J. Brill, 410 p.
3096 Bony Fishes
Eleutheronema tetradactylum (Shaw, 1804)

Frequent synonyms / misidentifications: None / Eleutheronema tridactylum (Bleeker, 1849).
FAO names: En - Fourfinger threadfin; Fr - Barbure mamalí; Sp - Barbudo mamalí.
Diagnostic characters:
A large, relatively elon-
gate species. Body
depth at first dorsal-fin
origin 2.8 to 5 times in
standard length. Head
length 2.9 to 3.8 times in
standard length. Eye di-
a m e te r g r e a t e r tha n
s n o u t l e n g t h . Dor-
soposterior margin of
maxilla not elevated
but nearly straight, pos-
terior margin rounded;
upper lip thin or absent; teeth extending well onto labial surface of anterior two-thirds of dentary, and
thick lower lip restricted to posterior third; head of vomer with tooth plate, with posteriorly-directed
smaller patches extending from each lateral tip of this main tooth plate; tooth plates on palatine and
ectopterygoid and band of teeth on premaxilla and dentary wide; width of band of teeth on premaxillae less
than 2 times in space separating band of teeth on one premaxilla from other; most of oral cavity covered
with small tooth plates. Posterior margin of preopercle with less than 76 serrations. Gill rakers on first
gill arch 7 to 18 (replaced by tooth plates in larger fish). First dorsal fin with VIII spines; second dorsal fin
with I spine and 12 to 15 soft rays (mean 14); anal fin with III spines and 13 to 17 (mean 15) soft rays;
anal-fin base longer than second dorsal-fin base; caudal-peduncle length greater than length of anal-fin
base; about 14 to 17 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 15 to 19 (mean
17) simple rays, the fins insert low on body, reaching to middle or end of pelvic fins; 4 short pectoral
filaments, third or fourth filament, from ventralmost, the longest, extending to middle or past end of
pelvic fins; pelvic fins seldom extend past anus. Lateral-line scales 59 to 91 (mean 77); lateral line
terminating between first and second or second and third lower caudal-fin rays, or bifurcates terminating
at both these points, some specimens with third dorsal branch occurring further anteriorly and terminating
between first and second upper rays; scale rows above lateral line 7 to 14 (mean 11); scale rows below
lateral line 12 to 18 (mean 15). Basisphenoid contacts prootic. Vertebrae 10+15. Hundreds of small
pyloric caeca arranged in tufts. Swimbladder not apparent in adults. Colour: back and head silvery green
or silvery blue; silvery white or yellowish white ventrally; cheeks and suborbital region golden; iris silvery
or yellow with a golden green tinge dorsally; fins grey or with yellow or orange tinge, dusky and darkened
marginally; pectoral filaments white or creamy.
Size: Maximum standard length at least 160 cm, commonly to 59 cm standard length. In Queensland, it
rarely exceeds 95 cm standard length, usually not over 60 cm fork length. Early growth is rapid. In Australia
at 1 year they average 30 cm fork length.
Habitat, biology, and fisheries: Along coasts and in estuaries. Taken at least to 30 m in Gulf of
Carpentaria. Diet of adults is largely made up of prawns and fish, with occasional polychaetes. Fishes
eaten are largely members of Mugilidae, Engraulidae, and Sciaenidae. The frequency of crustaceans to
fish in the diet changes seasonally. Apparently breeds in the sea off South Papua in August, though it may
have an extended breeding season. Juveniles are found in estuaries. Matures at about 22 cm standard
length in Papua and 28 cm standard length in India. Adults ascend the rivers during winter. Caught with
beach seines, longlines, traps, and trawls. Marketed fresh, frozen, and dried or salted. Makes up about
5% of total landings in the Gulf of Carpen-
taria.
Distribution: Persian Gulf eastward along
the coast of the Indian Ocean to the Malay
Peninsula, Gulf of Thailand, mouth of the
Mekong River delta, China (including Tai-
wan Province), Philippines, through Indo-
nesia to southern New Guinea, Ashburton
River (Western Australia), around the
north coast to Sandy Cape (Queensland);
in the north possibly to South Japan.
Eleutheronema tridactylum (Bleeker, 1849)

Frequent synonyms / misidentifications: None / Eleutheronema tetradactylum (Shaw, 1804).
FAO names: En - Threefinger threadfin.
Diagnostic characters: Very similar to Eleutheronema tetradactylum. A medium-sized species. Body

depth at first dorsal-fin origin 3.7 to 4.6 times in standard length. Head length 3.1 to 3.8 times in standard
length. Eye diameter greater than snout length. Dorsoposterior margin of maxilla not elevated but nearly
straight, posterior margin rounded; upper lip thin or absent; teeth extending well onto lateral surface of
anterior two-thirds of dentary and thick lower lip restricted to posterior third; head of vomer with tooth plate,
with posteriorly-directed smaller patches extending from each lateral tip of this main tooth plate; tooth plates
on palatine and ectopterygoid and band of teeth on premaxilla and dentary wide; width of band of teeth on
premaxillae less than 2 times in space separating band of teeth on one premaxilla from other; most of oral
cavity covered with small tooth plates. Posterior margin of preopercle with less than 76 serrations. Gill
rakers on first gill arch 5 to 13 (replaced by tooth plates in larger fish). First dorsal fin with VIII spines;
second dorsal fin with I spine and 12 to 15 soft rays (mean 13); anal fin with III spines and 12 to 14 (mean
15) soft rays; anal-fin base longer than second dorsal-fin base; caudal-peduncle length greater than length
of anal-fin base; about 14 to 17 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 16
to 18 (mean 17) simple rays, the fins insert low on body, reaching to middle or end of pelvic fins; 3 short
pectoral filaments; second or third filament, from ventralmost, the longest, extending to middle or past
end of pelvic fins; pelvic fins seldom extend past anus. Lateral-line scales 69 to 77 (mean 74); lateral line
terminating between first and second or second and third lower caudal-fin rays, or bifurcates terminating
at both these points, some specimens with third dorsal branch occurring further anteriorly and terminating
between the first and second or second and third upper rays; scale rows above lateral line 9 to 11 (mean
9); scale rows below lateral line 11 to 15 (13). Basisphenoid contacts prootic. Vertebrae 10+15. Colour:
back and body green dorsally; silvery white or yellowish white ventrally; pectoral, dorsal, and caudal fins
dark with black borders; golden hue over body and fins.
Habitat, biology, and fisheries: Occurs along coasts. Has been taken with E. tetradactylum.
Distribution: Gulf of Thailand
to Singapore, north along Ma-
lay Peninsula at least to Krabi,
Thailand; Sumatra, Java, Bor-
neo, and the Moluccas.
3098 Bony Fishes
Filimanus heptadactyla (Cuvier, 1829)

Frequent synonyms / misidentifications: None / Filimanus similis Feltes, 1991; Polydactylus sextarius
(Bloch, 1801); P. multiradiatus (Günther, 1860).
FAO names: En - Sevenfinger threadfin.
Diagnostic characters: A small species. Body depth at first dorsal-fin origin 3 to 3.6 times in standard
length. Head length 3 to 3.6 times in standard length. Eye diameter equal to or greater than snout length.
Snout protruding. Depth of maxilla at posterior margin less than eye diameter; upper lip thin, lower lip
fleshy; upper and lower jaws having narrow bands of teeth, bands of teeth on opposing premaxillae
separated by gap 2 or more times width of band of teeth; vomer with small inconspicuous tooth patch;
tooth patch on palatines narrow and shorter than tooth patch on ectopterygoid. Posterior margin of
preopercle with less than 35 serrations. Gill rakers on first gill arch 35 to 41 (mean 39). First dorsal fin
with VIII spines; second dorsal fin with I spine and 12 or 13 soft rays (mean 12); anal fin with III spines
and 11 or 12 (mean 11) soft rays; base of anal fin about equal to or greater than base of second dorsal fin;
caudal-peduncle length greater than length of anal-fin base; about 11 to 15 procurrent caudal-fin rays in
dorsal or ventral series; pectoral fins with 14 to 16 (mean 15) simple rays, the fins insert low on body,
extending to between posterior three-quarters and end of pelvic fins; 7 pectoral filaments, third or fourth
filament, from ventralmost, the longest, not reaching midpoint of anal fin; pelvic fins extend to or just
past anus. Lateral-line scales 46 to 49 (mean 48); lateral line terminating between centre of caudal-fin
fork and first lower caudal-fin ray, or occasionally between centre and first upper ray or first and second
upper rays; scale rows above lateral line 6 or 7 (mean 6); scale rows below lateral line 9 or 10 (mean 10).
Basisphenoid does not contact prootic. Anterior margin of ventral section of coracoid expanded. Vertebrae
10+14. Swimbladder simple and elongate. Colour: body brown above and golden below; fins yellow with
black at least at margins; pectoral fins often mostly black.
Size: M a x i m u m s t a n d a r d
length at least 13 cm.
ies: Taken in trawls. No other
data available.
Distribution: Eastern Thai-
land through Indonesia to
Papua New Guinea.
Filimanus hexanema (Cuvier, 1829)

FAO names: En - Javanese threadfin.
Diagnostic characters: A small species. Body depth at first dorsal-fin origin 3.2 to 3.4 times in standard
length; body compressed, greatest body width 2.7 to 3 in body depth at first dorsal-fin origin. Head length
3.3 to 3.6 times in standard length. Eye diameter greater than snout length. Snout blunt. Mouth oblique;
posterior margin of maxilla deep, greater than eye diameter; upper lip thin, lower lip fleshy; upper and
lower jaws having narrow bands of teeth, bands of teeth on opposing premaxillae separated by gap
2 or more times width of band of teeth; vomer with small inconspicuous tooth patch; tooth patch on
palatines narrow and shorter than tooth patch on ectopterygoid. Posterior margin of preopercle with less
than 20 serrations. Gill rakers on first gill arch 47 to 50 (mean 50). First dorsal fin with VIII spines;
longest spine of first dorsal fin longer than caudal peduncle; second dorsal fin with I spine and 11 or
12 soft rays; anal fin with III spines and 14 soft rays; anal-fin base longer than second dorsal-fin base;
caudal-peduncle length variable relative to length of anal-fin base; about 11 to 15 procurrent caudal-fin
rays in dorsal or ventral series; pectoral fins with 14 or 15 (mean 14) simple rays, the fins insert low on
body, extending past end of pelvic fins, sometimes reaching past anal-fin origin; 6 pectoral filaments,
fourth or third filament, from ventralmost, the longest, extending to caudal-fin base; pelvic fins seldom
extend past anus. Lateral-line scales 49 or 50 (mean 50); lateral line terminating between centre of
caudal-fin fork and first lower caudal-fin ray, or occasionally between centre and first upper ray or first and
second upper rays; scale rows above lateral line 6; scale rows below lateral line 9 or 10 (mean 10).
Basisphenoid does not contact prootic. Anterior margin of ventral section of coracoid expanded. Vertebrae
10+14. Colour: body and fins golden yellow, body darker dorsally; fins, especially first dorsal and pectoral
fins, may be dusky, most pigment on pectoral fins lateral and highly variable; pectoral filaments black,
especially distally.
data available.
Distribution: Known reliably
only from Djakarta (Jawa)
with a record for Labuhan
(Labuan) on the Sunda Strait.
3100 Bony Fishes
Filimanus perplexa Feltes, 1991

Frequent synonyms / misidentifications: None / Polynemus melanochir Valenciennes, 1831.
FAO names: En - Splendid threadfin.
Diagnostic characters: A small species. Body moderately deep, body depth at first dorsal-fin origin 2.9
to 3.3 times in standard length. Head length 2.9 to 3.3 times in standard length. Eye diameter greater than
snout length. Snout blunt. Mouth oblique; posterior margin of maxilla deep, greater than eye diameter;
region of frontals slightly concave; upper lip thin, lower lip fleshy; upper and lower jaws having narrow
bands of teeth, bands of teeth on opposing premaxillae separated by gap 2 or more times width of
band of teeth; vomer with small inconspicuous tooth patch; tooth patch on palatines narrow and shorter
than tooth patch on ectopterygoid. Posterior margin of preopercle with less than 26 serrations. Gill rakers
on first gill arch 47 to 50 (mean 50). First dorsal fin with VIII spines; longest spine of first dorsal fin
longer than caudal peduncle; second dorsal fin with I spine and 11 soft rays; length of spine of second
dorsal fin greater than caudal-peduncle depth; anal fin with III spines and 13 to 15 (mean 14) soft rays;
anal-fin base longer than second dorsal-fin base; caudal-peduncle length variable relative to length of
anal-fin base; about 11 to 15 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins insert low
on body; pectoral fins with 13 or 14 (mean 14) simple rays, extending past end of pelvic fins, sometimes
reaching past anal-fin origin; 7 long pectoral filaments, third or fourth filament, from ventralmost, the
longest, extending past caudal-fin fork; pelvic fins seldom extend past anus. Lateral-line scales 49 to
51 (mean 50); lateral line terminating between centre of caudal-fin fork and first lower caudal-fin ray,
occasionally between centre and first upper ray or first and second upper rays; scale rows above lateral
line 5 to 8 (mean 6); scale rows below lateral line 9 or 10 (mean 10). Basisphenoid does not contact prootic.
Anterior margin of ventral section of coracoid expanded. Vertebrae 10+14. Swimbladder simple and
elongate. Pyloric caeca about 6, short and thick. Colour: body and fins fawn to yellow, body darker dorsally;
fins, especially first dorsal and pectoral fins, dusky to black; most pigment on pectoral fins lateral and highly
variable; pectoral filaments black, especially distally.
Size: M a x i m u m s t a n da r d
data available.
Distribution: Indonesia
including Nias, Padang on
Sumatra, Java, and Bali.
Filimanus sealei (Jordan and Richardson, 1910)

Frequent synonyms / misidentifications: None / Filimanus heptadactyla (Cuvier, 1829); Polydactylus
multiradiatus (Günther, 1860).
FAO names: En - Eightfinger threadfin.
Diagnostic characters: Very similar to Filimanus heptadactyla. A small species. Body depth at first
dorsal-fin origin 2.9 to 3.5 times in standard length. Head length 3 to 3.4 times in standard length. Eye
diameter equal to or greater than snout length. Snout protruding. Depth of maxilla at posterior margin less
than eye diameter; upper lip thin, lower lip fleshy; upper and lower jaws having narrow bands of teeth,
bands of teeth on opposing premaxillae separated by gap 2 or more times width of band of teeth; vomer
with small inconspicuous tooth patch; tooth patch on palatines narrow and shorter than tooth patch on
ectopterygoid. Posterior margin of preopercle with less than 35 serrations. Gill rakers on first gill arch
40 to 48 (mean 46). First dorsal fin with VIII spines; second dorsal fin with I spine and 11 or 12 soft rays
(mean 12); anal fin with III spines and 11 soft rays; base of anal fin about equal to or greater than base of
second dorsal fin; caudal-peduncle length greater than length of anal-fin base; about 11 to 15 procurrent
caudal-fin rays in dorsal or ventral series; pectoral fins with 14 or 15 (mean 14) simple rays, the fins insert
low on body, extending to posterior three-quarters of pelvic fins; 8 (rarely 7) pectoral filaments, third or
fourth filament, from ventralmost, the longest, not reaching midpoint of anal fin; pelvic fins extend to or just
past anus. Lateral-line scales 46 to 50 (mean 49); scale rows above lateral line 6 to 7 (mean 7); scale
rows below lateral line 9 or 10 (mean 10). Basisphenoid does not contact prootic. Anterior margin of ventral
section of coracoid expanded. Vertebrae 10+14. Colour: body blue above and silvery below; pectoral and
pelvic fins yellow with black at least at margins.
Habitat, biology, and fisheries: Taken in trawls. No other data available.
Distribution: T h e P h i l i p-
pines, New Guinea, Bismarck
Archipelago, and the Solo-
mon Islands.
3102 Bony Fishes
Filimanus xanthonema (Valenciennes, 1831)

Frequent synonyms / misidentifications: Polynemus pfeifferi Bleeker, 1853 / Filimanus similis Feltes,
1991; Polydactylus multiradiatus (Günther, 1860); P. sexfilis (Valenciennes, 1831); P. sextarius (Bloch,
1801).
FAO names: En - Yellowthread threadfin.
Diagnostic characters: Very similar to Filimanus heptadactyla. A small species. Body depth at first
dorsal-fin origin 2.8 to 3.7 times in standard length. Head length 2.9 to 3.6 times in standard length. Eye
diameter equal to or greater than snout length. Snout protruding. Depth of maxilla at posterior margin less
than eye diameter; upper lip thin, lower lip fleshy; upper and lower jaws having narrow bands of teeth,
bands of teeth on opposing premaxillae separated by gap 2 or more times width of band of teeth; vomer
with small inconspicuous tooth patch; tooth patch on palatines narrow and shorter than tooth patch on
ectopterygoid. Posterior margin of preopercle with less than 35 serrations. Gill rakers on first gill arch
36 to 46 (mean 41.8 in Pacific Ocean). First dorsal fin with VIII spines; second dorsal fin with I spine and
11 to 13 soft rays (mean 12); anal fin with III spines and 10 to 12 (mean 11) soft rays; anal-fin base about
equal to or greater than second dorsal-fin base; caudal-peduncle length greater than length of anal-fin
base; about 11 to 15 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 13 to 15 (mean
15) simple rays, the fins insert low on body, extending to posterior three-quarters of pelvic fins; 5 to 7
(mean 6) pectoral filaments, may be asymmetric, third or fourth filament, from ventralmost, the longest,
not reaching midpoint of anal fin; pelvic fins extend to or just past anus. Lateral-line scales 43 to 52 (mean
47); lateral line terminating between centre of caudal-fin fork and first lower caudal-fin ray, or occasionally
between centre and first upper ray or first and second upper rays; scale rows above lateral line 5 to 8 (mean
6); scale rows below lateral line 9 to 12 (mean 10). Basisphenoid does not contact prootic. Anterior margin
of ventral section of coracoid expanded. Vertebrae 10+14. Swimbladder simple and elongate. Pyloric caeca
about 14, short and thick. Colour: body green, dusky yellow, or brown above and silvery below; fins yellow
with black at least at margins; pectoral fins often mostly black; free filaments yellow.
Size: M a x i m u m a t l e a s t
14 cm standard length.
data available.
Distribution: Coasts of the
Bay of Bengal and the eastern
Indian Ocean; Singapore to
Lombok.
Parapolynemus verekeri (Saville-Kent, 1889)

Frequent synonyms / misidentifications: Polynemus intermedius Nichols, 1954 / None.
FAO names: En - Dwarf paradise fish.
Diagnostic characters: The smallest polynemid species. Relatively elongate body, its depth at first
diameter 2.5 or less times in snout length. Mouth large, obliquely turned down posteriorly, upper jaw less
than 1.3 times in postorbital length; posterior margin of maxilla moderately deep, less than 4 times in
upper-jaw length; broad head, interorbital less than 2.5 times in postorbital length; no free upper lip, lower
lip absent anteriorly and moderate to thick posteriorly; vomer edentate; tooth patch on palatine slightly
wider and longer than tooth patch on ectopterygoid; band of teeth on maxilla narrow with anterior gap
separating opposing sides; teeth extending laterally on anterior third of dentary of larger specimens.
Posterior margin of preopercle lacks serrations. Gill rakers on first gill arch 30 to 43 (mean 40). First
dorsal fin with VIII spines; second dorsal fin with I spine and 11 to 14 soft rays (mean 13); anal fin with III
spines and 10 to 12 (mean 11) soft rays; anal-fin base shorter than second dorsal-fin base; caudal-peduncle
length greater than length of anal-fin base; about 12 to 14 procurrent caudal-fin rays in dorsal or ventral
series; caudal-fin lobes long, lower often longest; usually longer than distance from snout to origin of second
dorsal fin; pectoral fins with 12 to 14 (mean 13) simple rays, the fins insert near midline on side of body
and may reach past posterior end of anal-fin base; 6 or 7 (mean 7) pectoral filaments; fifth filament, from
ventralmost, the longest, may extend past end of caudal fin, but less than 1.5 times in total length; pelvic
fins extend past anus. Lateral-line scales 50 to 60 (mean 56); lateral line terminating between centre of
caudal-fin fork and first lower caudal-fin ray; scale rows above lateral line 4 to 6 (mean 5); scale rows below
lateral line 8 to 11 (mean 10). Vertebrae 10+14. Swimbladder not apparent. Pyloric caeca narrow, about
15 to 24. Colour: body and head yellow, shaded with black dorsally; fins bright orange, pectoral filaments
intense vermilion.
Habitat, biology, and fisheries: Occurs in muddy estuaries and lower portions of rivers. Hundreds of
adults approaching breeding
condition were taken in trawl
on Medusa Banks at a depth
of 9 m in January or February
1968.
Distribution: Merauke River
(New Guinea) through Gulf of
Papua; Cambridge Gulf
(Western Australia) to Port
Stuart (Northern Territory).
3104 Bony Fishes
Polydactylus indicus (Shaw, 1804)

Frequent synonyms / misidentifications: N o n e / Polydactylus macrochir (Günther, 1867);
P. microstoma (Bleeker, 1851).
FAO names: En - Indian threadfin; Fr - Barbure indien; Sp - Barbudo indio.
Diagnostic characters: A large, elongate, shallow-bodied species; body depth at first dorsal-fin origin 3.9
to 5.3 times in standard length. Head length 2.9 to 3.6 times in standard length. Eye diameter less than
snout length. Upper lip thin; lower lip medium anteriorly, thick posteriorly; tooth patch on vomer rounded
triangle; premaxillary, dentary, palatine, and ectopterygoid tooth patches all wide; width of band of teeth
on premaxillae less than 2 times in space separating band of teeth on one premaxilla from other; on larger
specimens teeth extend laterally about 1/4 back on dentary; postorbital length greater than 3/4 of body
depth at second dorsal-fin origin. Posterior margin of preopercle with less than 65 serrations. Gill rakers
on first gill arch 18 to 30 (mean 20). First dorsal fin with VIII spines; second dorsal fin with I spine and
12 or 13 (mean 13) soft rays; anal fin with III spines and 11 soft rays; anal-fin base shorter than second
dorsal-fin base, 2 times length of anal-fin base less than distance from pelvic-fin origin to anal-fin origin;
caudal-peduncle length greater than length of anal-fin base; caudal-fin lobes long, often with filaments;
about 11 to 16 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 12 to 15 (mean
14) simple rays, the fins insert low on body, reaching to middle of pelvic fins or 3/4 of their length; 5
pectoral filaments, fifth filament, from ventralmost, the longest, extending past end of pelvic fins; pelvic
fins reach near or past anus. Lateral-line scales 64 to 73 (mean 69); lateral line terminates between
second and third lower rays of caudal fin; scale rows above lateral line 6 to 8 (mean 7); scale rows below
lateral line 10 to 12 (mean 11). Basisphenoid contacts prootic. Ventral section of the coracoid with foramina
and anterior margin of this section greatly expanded. Vertebrae 10+14. Swimbladder structure is unique
in family with at least 24 appendages entering body wall, swimbladder elongate, running entire length of
body cavity; ending in a point penetrating the caudal musculature; membrane is thick and silvery. About
16 pyloric caeca in the grouping proximal to the cardiac portion of the stomach, 8 to 10 in the distal group.
Colour: back and head greyish green or purplish black; silvery white ventrally, with a very indistinct stripe
along centre of each line of abdominal scales; fins dusky and darkened marginally.
Size: Maximum standard length at least 113 cm, commonly to 63 cm standard length.
Habitat, biology, and fisheries: Occurs mainly near estuaries, occasionally entering rivers, taken to a
depth of 100 m, though rare below 55 m. Young feed almost exclusively on crustaceans. The percent of
fishes in the diet increases
with size. Taken with beach
seines, traps, longlines, and
trawls.
Distribution: D u r b a n a n d
Delagoa Bay, South Africa;
Maurice; Sind Province
(Pakistan); east around the
coast of India; along the
Malay Peninsula; northwest
coast of Sumatra; Gulf of
Thailand, and Borneo.
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Polydactylus macrochir (Günther, 1867)

Frequent synonyms / misidentifications: Polynemus sheridani Macleay, 1884 / Polydactylus indicus
(Shaw, 1804).
FAO names: En - King threadfin.
Diagnostic characters: A large, elongate spe-
cies; body depth at first dorsal-fin origin 3.6 to
4.7 times in standard length, body depth at sec-
ond dorsal-fin origin less than distance from
pelvic-fin origin to anal-fin origin. Head length 3.3
to 3.6 times in standard length. Eye diameter
about equal to or less than snout length. Snout
is not acutely pointed; upper lip thin; lower lip
thick; tooth patch on vomer rounded triangle;
palatine tooth patches are wide; width of band of
teeth on premaxillae less than 2 times in space separating band of teeth on one premaxilla from other;
postorbital length greater than distance from snout to posterior margin of preopercle, postorbital length less
than or equal to 3/4 of body depth at second dorsal-fin origin. Posterior margin of preopercle with less than 50
serrations. Gill rakers on first gill arch 31 to 37 (mean 33). First dorsal fin with VIII spines; second dorsal
fin with I spine and 11 or 12 (mean 12) soft rays; anal fin with III spines and 9 to 11 (mean 11) soft rays;
anal-fin base longer than or equal to second dorsal-fin base; 2 times length of anal-fin base greater than
distance from pelvic-fin origin to anal-fin origin; caudal-peduncle length greater than length of anal-fin base;
about 11 to 16 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 13 to 15 (mean 14)
simple rays, the fins insert low on body, reaching to posterior quarter to end of pelvic fins; 5 (rarely 4)
pectoral filaments, fourth filament, from ventralmost, usually longest, extending almost to anal-fin
origin or a little past anal-fin origin; pelvic fins reach near or past anus. Lateral-line scales 63 to 75
(mean 71); lateral line terminates between first and second lower rays of caudal fin; scale rows above
lateral line 8 to 10 (mean 9); scale rows below lateral line 10 to 13 (mean 12). Basisphenoid contacts
prootic. Ventral section of the coracoid with foramina and anterior margin of this section greatly expanded.
Vertebrae 10+14. Swimbladder simple, elongate, and large. Pyloric caeca about 25. Colour: body bluish
silvery dorsally, and whitish ventrally; each line of scales shows a more or less distinct longitudinal streak;
during some seasons ventral surface flushed with yellow; dorsal, anal, and caudal fins blackish; pectoral
fins, pelvic fins, and pectoral filaments whitish or yellow; fins sprinkled with melanophores.
Size: Specimens with a standard length over 48 cm were abundant seasonally in the tidal waters of the
Mary River, Australia, and this fish could attain a mass of 45 kg. Fish over 88 cm standard length were
common at Maryborough, where the record was 30 kg. Fish of a weight of 45 kg are no longer caught; fish
taken usually weigh 9 to 14 kg. Reported to attain a fork length of 140 cm and 32 kg in the Gulf of
Carpentaria, and over 150 cm fork length in Princess Charlotte Bay, Australia. Estimated maximum size
of 170 cm fork length and 40 kg.
Habitat, biology, and fisheries: Frequents rivers and estuaries. Often over sand or mud flats to 5 m. Both
juveniles and adults are taken in the Trinity Inlet system at Cairns, Queensland. Along the coast of New
Guinea, found near and in the Lower Purari River system in mangrove areas and around the Nypa palms
between the mangroves and the swamps. More abundant there than Eleutheronema tetradactylum and
second in commercial importance after Lates calcarifer. Polydactylus macrochir comprises about 30%
of gill-net landings, by weight, from the Gulf of Carpentaria. P. macrochir is more abundant in the Kikori
area than in the Purari delta where larger fish are taken. Juveniles were taken along the coast. Prawns,
crabs, and fish are the predominant food items. It is suggested the species migrates down the Purari River
to the sea to spawn. They are known to move at least 550 km along the Australian coast. Can live more
than 20 years. Valued as a food fish, it is marketed as fresh or frozen fillets or whole fish. This is an important
game fish in Queensland.
Distribution: Early reports of this species
were from the Mary (Burnett) and Fitzroy rivers
of eastern Australia; common along the coast
and delta areas of Papua, New Guinea; south
coast of New Guinea (Oetoemboewe to Gulf
of Papua); in Australia, probably from as far
west as Broome (Western Australia) north and
east along the coast to the Brisbane River,
Queensland, although Günther reports the
type locality as New South Wales.
3106 Bony Fishes
Polydactylus macrophthalmus (Bleeker, 1859)

FAO names: En - River threadfin.
Diagnostic characters: A medium- to large-sized species, moderately elongate; body depth at first
diameter variable, about equal to snout length; narrow interobital, 4.7 to 5.3 in head length; operculum
with angular posterior margin; upper lip medium to thick; lower lip thick; tooth patch on vomer a wide “v”
shape; tooth patch on palatine broad; width of band of teeth on premaxillae less than 2 times in space
separating band of teeth on one premaxilla from other. Posterior margin of preopercle with less than 53
serrations. Gill rakers on first gill arch 27 to 38 (mean 32). First dorsal fin with VIII spines; second dorsal
fin with I spine and 13 or 14 (mean 14) soft rays; anal fin with III spines and 11 soft rays; anal-fin base
shorter than second dorsal-fin base; caudal-peduncle length greater than length of anal-fin base; about 11
to 16 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 13 or 14 (mean 13) simple
rays, the fins insert low on body, reaching to middle or end of pelvic fins; 7 long pectoral filaments, fifth
filament, from ventralmost, usually the longest, extending well past end of caudal fin, may exceed twice
standard length; pelvic fins reach near or past anus. Lateral-line scales 87 to 92 (mean 90); lateral line
terminates between first and second lower rays of caudal fin, or rarely lateral line bifurcating on caudal fin
with termination between centre of fork and first lower ray, and first and second lower rays; scale rows
above lateral line 9 to 11 (mean 10); scale rows below lateral line 15 to 17 (mean 16). Basisphenoid contacts
Vertebrae 10+14. Swimbladder simple, elongate, moderate to large in size. Pyloric caeca 25 to 35. Colour:
body dull silvery or brown to golden orange; unpaired fins dusky; tip of first dorsal fin may be black; second
dorsal fin may be dark distally; pectoral fins dusky only dorsally; pectoral filaments with some melanopho-
res.
ies: Middle to lower sections
of rivers. Esteemed as a food
fish along the Kapuas River,
Borneo.
Distribution: Hari and Musi
rivers of Sumatra and Kapuas
River (Borneo), at least as far
up as Silat.
Polydactylus microstoma (Bleeker, 1851)

Frequent synonyms / misidentifications: Polydactylus zophomus Jordan and McGregor in Jordan and
Seale, 1907 / Polydactylus sextarius (Bloch, 1801); P. plebeius (Broussonet, 1782).
FAO names: En - Smallmouth threadfin.
Diagnostic characters: A small species; body deep, its depth at first dorsal-fin origin 2.5 to 3.7 times in
standard length. Head length 2.7 to 3.3 times in standard length. Eye diameter greater than snout length.
Snout pointed; posterior margin of maxilla rounded, not deep; upper lip thin; lower lip moderate; vomer
edentate; anterior of tooth patch on palatine with medial arm; ectopterygoid tooth patch small; width
of band of teeth on premaxillae less than 2 times in space separating band of teeth on one premaxilla from
other. Posterior margin of preopercle with less than 53 serrations. Gill rakers on first gill arch 25 to 35
(mean 29). First dorsal fin with VIII spines; second dorsal fin with I spine and 12 or 13 (mean 13) soft rays;
anal fin with III spines and 10 to 13 (mean 12) soft rays; anal-fin base about equal to second dorsal-fin
base; caudal-peduncle length equal to or greater than length of anal-fin base; caudal-peduncle length
greater than length of anal-fin base; about 11 to 16 procurrent caudal-fin rays in dorsal or ventral series;
pectoral fins insert low on body; pectoral fins with 12 to 15 (mean 13) rays; usually 2 dorsalmost
pectoral-fin rays simple, others branched; pectoral fins and pectoral filaments short, 5 pectoral
filaments, fifth pectoral filament, from ventralmost, the longest, less than head length, both pectoral fins
and filaments extend to 1/3 to 3/4 of pelvic fins. Pelvic fins reach near or past anus. Lateral-line scales
42 to 50 (mean 47); lateral line usually terminates between first and second lower rays of caudal fin; scale
rows above lateral line 6 to 8 (mean 6); scale rows below lateral line 9 to 11 (mean 10). Basisphenoid
contacts prootic. Ventral section of coracoid with foramina and anterior margin of this section greatly
expanded. Vertebrae 10+14. Swimbladder simple, elongate, and highly variable in size. Pyloric caeca
about 15. Colour: body white, with metallic golden and green reflections; black spot behind opercle
covers about second through eighth lateral-line scales and diffusing over 1 row above and below;
dorsal and caudal fins a pale dusky green; pectoral, pelvic, and anal fins dusky, with wash of yellow;
pectoral fins dark mostly on medial surface and dorsolaterally.
Habitat, biology, and fisheries: Catches
of juveniles from the Labu estuary near the
mouth of the Markham River in Papua New
Guinea were associated with lunar phases.
Thirty-five percent of the specimens taken
in a collection from Palk Bay and Gulf of
Mannar, Sri Lanka, were hermaphrodites.
Distribution: Southeast India, Sri Lanka,
and northeast Bay of Bengal; Taiwan Prov-
ince of China south through Philippines,
Borneo, eastern Indonesia to eastern New
Guinea.
3108 Bony Fishes
Polydactylus multiradiatus (Günther, 1860)

Frequent synonyms / misidentifications: Polynemus specularis De Vis, 1883; Polydactylus auratus
McKay, 1970 / Filimanus heptadactyla (Cuvier, 1829).
FAO names: En - Australian threadfin.
Diagnostic characters: A small species with a deep compressed body; body depth greater than head
length, at first dorsal-fin origin 2.6 to 3.5 times in standard length. Head length 2.9 to 3.8 times in standard
length. Eye diameter greater than snout length. Snout not markedly pointed. Mouth small; upper lip
moderately thin; lower lip moderate; tooth patch on vomer rounded, wide “v” shape; anterior of palatine
tooth patch expanded medially; width of band of teeth on premaxillae less than 2 times in space separating
band of teeth on one premaxilla from other. Posterior margin of preopercle with less than 45 serrations.
Gill rakers on first gill arch 26 to 32 (mean 30). First dorsal fin with VIII spines; second dorsal fin with I
spine and 13 or 15 (mean 14) soft rays; base of second dorsal fin long, 4.2 to 5.6 times in standard length;
base of anal fin long, 3.5 to 5 times in standard length; anal-fin ray count high, anal fin with III spines
and 14 to 18 (mean 17) soft rays; anal-fin base longer than second dorsal-fin base; caudal peduncle
short and deep, caudal-peduncle length less than length of anal-fin base; about 11 to 16 procurrent
caudal-fin rays in dorsal or ventral series; pectoral fins with 14 to 16 (mean 15) simple rays, the fins insert
low on body, reaching to posterior three-quarters of pelvic fins; 7 (rarely 6) pectoral filaments, seventh
filament, from ventralmost, the longest, extending to posterior three-quarters of pelvic fins or to anal
origin; pelvic fins reach near or past anus. Lateral-line scales 50 to 58 (mean 54); lateral line usually
terminates between first and second lower rays of caudal fin; scale rows above lateral line 6 to 9 (mean
8); scale rows below lateral line 13 to 16 (mean 14). Basisphenoid contacts prootic. Ventral section of the
coracoid with foramina and anterior margin of this section greatly expanded. Vertebrae 10+14. Swimblad-
der not apparent. Pyloric caeca about 35. Colour: dorsal part of body and head golden, abdomen silvery
pink, the 2 colours often sharply defined; dorsal fins dusky; anal and caudal fins greenish yellow or yellowish
orange; pectoral fins yellowish with the middle or whole of fin black or bluish black; pectoral filaments pale
yellow.
Habitat, biology, and fisheries: Taken to a depth of at least 50 m in Gulf of Carpentaria. Feeds on prawns.
Commonly taken in association with prawns of the genus Penaeus.
Distribution: South coast of
New Guinea from Samdoe-
mande through the Gulf of
Papua; Exmouth Gulf (West-
ern Australia) along north and
east coast of Australia to
Clarence River (New South
Wales).
Polydactylus nigripinnnis Munro, 1964

FAO names: En - Blackfin threadfin.
Diagnostic characters: A small species. Body depth at first dorsal-fin origin 2.8 to 3.8 times in standard
length; body wide, width above pectoral-fin insertion 5.7 to 8.1 times in standard length. Head length 2.8
to 3.2 times in standard length. Snout protruding. Eye diameter about equal to snout length. Head wide;
angle of posterior margin of opercle relatively ventral; upper lip very thin; lower lip thick; dentaries fit largely
within premaxillae when mouth completely closed; broad band of teeth cover ventral surface of
premaxilla; vomer edendate; palatine and ectopterygoid tooth plates narrow; width of band of teeth on
premaxillae less than 2 times in space separating band of teeth on one premaxilla from other. Posterior
margin of preopercle with less than 10 serrations. Gill rakers on first gill arch 24 to 27 (mean 26).
First dorsal fin with VIII spines; longest spine of first dorsal fin less than or equal to 2/3 of pectoral-fin
length; second dorsal fin with I spine and 12 or 13 (mean 13) soft rays; anal fin with III spines and 11 to
13 (mean 12) soft rays; anal-fin base shorter than or equal to second dorsal-fin base; caudal-peduncle
length greater than or equal to length of anal-fin base; about 12 to 14 procurrent caudal-fin rays in dorsal
or ventral series; pectoral fins with 16 to 18 (mean 17) simple rays; pectoral fins insert low on body, relatively
long, being about 1/3 standard length, reaching to end of pelvic fins; 6 pectoral filaments, sixth filament,
from ventralmost, the longest, extending to origin of anal fin; bases of second dorsal fin and anal fin of
equal length; pelvic fins reach near or past anus. Lateral-line scales 48 to 51 (mean 50); lateral line
terminates between first and second lower rays of caudal fin; scale rows above lateral line 6 to 8 (mean
7); scale rows below lateral line 11 or 12 (mean 11). Basisphenoid contacts prootic. Basisphenoid contacts
Vertebrae 10+14. Peritoneum with dark scattering of melanophores. No swimbladder apparent. Pyloric
caeca about 25. Colour: body dusky blue dorsally, silver golden below; spot immediately above origin of
pectoral fins; unpaired fins yellow with melanophores; pectoral fins, except uppermost ray and free
filaments, intensely black on lateral surface.
ies: Usually taken off mouths
of rivers. Has been taken at
150 m in the Gulf of Carpen-
taria, but typically at much
shallower dephs.
Distribution: Gulf of Papua
(New Guinea), specifically
Orokolo Bay; Cambridge Gulf
(Western Australia) to Van
Diemen Gulf (Northern Terri-
tory).
3110 Bony Fishes
Polydactylus plebeius (Broussonet, 1782)

Frequent synonyms / misidentifications: None / Polydactylus microstoma (Bleeker, 1851).
FAO names: En - Common threadfin; Fr - Barbure rayé; Sp - Barbudo rayado.
Diagnostic characters: M e-
dium-sized to large species.
Body depth at first dorsal-fin
origin 3.1 to 4.3 times in stand-
ard length, body depth at sec-
ond dorsal-fin origin greater
than distance from pelvic-fin
origin to anal-fin origin. Head
length 2.7 to 3.5 times in stand-
ard length. Eye diameter
greater than or about equal to
snout length. Snout moderately
conical; upper lip thin; lower lip
m o d e rate; tooth patch o n
vomer a wide “v” shape changing to a rounded triangle; premaxillary, dentary, palatine and ectopterygoid
tooth patches all wide; width of band of teeth on premaxillae less than 2 times in space separating band
of teeth on one premaxilla from other; postorbital length greater than distance from anterior of snout to
posterior margin of preopercle, postorbital length less than or equal to 3/4 of body depth at second dorsal-fin
origin. Posterior margin of preopercle with less than 70 serrations. Gill rakers on first gill arch 19 to 28
(mean 26). First dorsal fin with VIII spines; second dorsal fin with I spine and 12 to 14 (mean 13) soft rays;
anal fin with III spines and 9 to 12 (mean 11) soft rays; anal-fin base about equal to second dorsal-fin base,
2 times length of anal-fin base greater than distance from pelvic-fin origin to anal-fin origin; caudal-peduncle
series; pectoral fins with 14 to 18 (mean 17) simple rays, the fins insert low on body, reaching to middle
of pelvic fins or 3/4 of its length; 5 pectoral filaments, fifth filament, from ventralmost, the longest,
extending to 3/4 of pelvic fins or past origin of anal fin; pelvic fins reach near or past anus. Lateral-line
scales 59 to 69 (mean 62); lateral line terminates between first and second lower rays of caudal fin; scale
contacts prootic. Ventral section of coracoid with foramina and anterior margin of this section greatly
expanded. Vertebrae 10+14. Swimbladder simple, elongate, and variable in size. Colour: body brown or
dark olive dorsally, becoming lighter ventrally; adults with about 7 dark olive brown longitudinal lines above
lateral line and about 10 below lateral line which are more faintly developed; sometimes reddish towards
mouth and nose; dorsal, anal, caudal, and pectoral fins grey distally; pectoral fins often a dull yellowish;
pelvic fins lighter or whitish; smaller specimens usually lack horizontal stripes; may be lighter in coloration,
being more yellow or silvery; fish less than 8 cm may have a heavily pigmented first dorsal fin, pectoral
fins, pelvic fins, anterior bases of second dorsal and anal fins, and lobes of caudal fin; fish less than 5 cm
may have broad dark vertical bands passing through operculum, first dorsal fin and pelvic fins, caudal
peduncle, and caudal-fin lobes, in addition to dark base of second dorsal and anal fins.
Size: Maximum standard length at least 45 cm. An average large fish attains 26 cm standard length. Fishes
of 12 cm can be caught in large numbers.
Habitat, biology, and fisheries: Live off coasts over sand or mud flats. Caught in large numbers at mouths
of rivers. Frequently caught in the surf, commonly enters estuaries, and occurs at a depth of 20 to 122 m.
Specimens of 10 to 19 cm standard length feed mostly on crabs occurring on the muddy bottom near shore.
Fish are also part of the diet. Taken in beach seines and trawls.
Distribution: Indian Ocean from South Af-
rica, north along the east coast of Africa
including Madagascar and Mauritius; Mus-
cat, Oman, east along the coast of India;
islands of the northern Indian Ocean; Malay
Peninsula and Archipelago. Indonesia north
through the Philippines, Taiwan Province of
China to Korea and Mikowa Bay and Tokyo
in Japan; Geralton District (Western Austra-
lia) along the north coast to Port Stephens
(New South Wales); New Guinea eastwards
to Polynesia.
Polydactylus sexfilis (Valenciennes, 1831)

Frequent synonyms / misidentifications: Polynemus kuru Bleeker, 1853 / Filimanus xanthonema
(Valenciennes, 1831).
FAO names: En - Sixfinger threadfin.
Diagnostic characters: Medium-
sized to large species. Body depth
at first dorsal-fin origin 2.7 to 4.1
times in standard length, body
depth at second dorsal-fin origin
greater than distance from pelvic-
fin origin to anal-fin origin. Head
length 2.7 to 3.6 times in standard
length. Eye diameter greater than
snout length. Snout moderately
conical; upper lip thin; lower lip
moderate; tooth patch on vomer a
wide “v” shape changing to a
rounded triangle; premaxillary,
dentary, palatine and ectoptery-
goid tooth patches all wide; width of band of teeth on premaxillae less than 2 times in space separating
band of teeth on one premaxilla from other; postorbital length less than distance from anterior of snout to
posterior margin of preopercle, postorbital length less than or equal to 3/4 of body depth at second dorsal-fin
origin. Posterior margin of preopercle with less than 62 serrations. Gill rakers on first gill arch 27 to 32
(mean 29). First dorsal fin with VIII spines; second dorsal fin with I spine and 11 to 13 (mean 13) soft rays;
anal fin with III spines and 9 to 12 (mean 11) soft rays; anal-fin base about equal to second dorsal-fin base,
2 times length of anal-fin base greater than distance from pelvic-fin origin to anal-fin origin; caudal-peduncle
series; pectoral fins with 14 to 17 (mean 16) simple rays, the fins insert low on body, reaching to middle
of pelvic fins or 3/4 of their length; 6 pectoral filaments, fifth filament, from ventralmost, the longest,
extending to 3/4 of pelvic fins or past origin of anal fin; pelvic fins reach near or past anus. Lateral-line
scales 58 to 70 (mean 63); lateral line terminates between first and second lower rays of caudal fin; scale
contacts prootic. Ventral section of the coracoid with foramina and anterior margin of this section greatly
expanded. Vertebrae 10+14. Swimbladder simple, elongate, and variable in size. Colour: body brown or
dark olive dorsally, becoming lighter ventrally, adults with about 7 dark olive brown longitudinal lines
above lateral line and about 10 below lateral line which are more faintly developed; sometimes reddish
towards mouth and nose; dorsal, anal, caudal, and pectoral fins are grey or black distally; pectoral fins
sometimes appearing purplish; pelvic fins are lighter or whitish; smaller specimens usually lack the
horizontal stripes; may be lighter in coloration, being more yellow or silvery; or fish less than 8 cm may
have a heavily pigmented first dorsal fin, pectoral fins, pelvic fins, anterior bases of second dorsal and anal
fins, and lobes of caudal fin; or fish less than 5 cm may have broad dark vertical bands passing through
the operculum, the first dorsal fin and pelvic fins, the caudal peduncle, and the caudal-fin lobes, in addition
to the dark base of second dorsal and anal fins.
Size: Maximum at least 40 cm standard length.
Habitat, biology, and fisheries: Found near reef areas. Not taken over the reef, but found in schools in
lagoons and along sandy and rocky shores, often in zones of turbulence. Usually taken at depths of 20 to
50 m. A lunar spawning rhythm occurs close to shore. Sexual maturity as males occurs at 12 to 15 cm.
They subsequently undergo sex reversal passing through a hermaphroditic stage, and becoming females
between 18 and 24 cm. The potential of this species for aquaculture has been studied. The diet is
comprised mostly of crustaceans followed
by teleosts.
Distribution: Mauritius, Seychelles, Kenya,
and many of the islands of the northern
Indian Ocean; Indonesia north through the
Philippines, Ryukyu Islands to Ogasawara
and Yaeyama Islands east through the is-
lands of the Pacific to Hawaii, and from
Micronesia to Polynesia and Pitcairn Island.
3112 Bony Fishes
Polydactylus sextarius (Bloch, 1801)

Frequent synonyms / misidentifications: None / Filimanus heptadactyla (Cuvier, 1829).
FAO names: En - Blackspot threadfin; Fr - Barbure tache noire; Sp - Barbudo de mancha.
Diagnostic characters: A small species with a deep body; body depth at first dorsal-fin origin 2.7 to 3.5
times in standard length. Head length 2.6 to 3.4 times in standard length. Eye diameter greater than snout
length. Snout pointed; posterior margin of maxilla rounded and not deep; upper lip thin; lower lip
moderate; vomer edentate; anterior of tooth patch on palatine with medial arm; ectopterygoid tooth
patch small; width of band of teeth on premaxillae less than 2 times in space separating band of teeth on
one premaxilla from other. Posterior margin of preopercle with less than 50 serrations. Gill rakers on first
gill arch 25 to 33 (mean 28). First dorsal fin with VIII spines; second dorsal fin with I spine and 12 to 14
(mean 13) soft rays; anal fin with III spines and 11 to 13 (mean 12) soft rays; anal-fin base about equal to
second dorsal-fin base; caudal-peduncle length greater than length of anal-fin base; about 11 to 16
procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 13 to 15 (mean 14) rays, usually
2 dorsalmost pectoral-fin rays simple, others branched; pectoral fins short, insert low on body and
reach to 1/3 to 3/4 of pelvic fins; 6 short pectoral filaments, sixth filament, from ventralmost, the longest,
less than head length, extending midway along or beyond end of pelvic fins; pelvic fins reach to between
anus and anal-fin origin. Lateral-line scales 43 to 49 (mean 47); lateral line terminates between first and
second lower rays of caudal fin; scale rows above lateral line 5 to 7 (mean 6); scale rows below lateral line
8 to 12 (mean 11). Basisphenoid contacts prootic. Ventral section of the coracoid with foramina and anterior
margin of this section greatly expanded. Vertebrae 10+14. Swimbladder simple, elongate, and highly
variable in size. Pyloric caeca about 15. Colour: dorsally, body and head silvery or golden green to
olivaceous; becoming silvery white on sides and ventrally; dark spot present near origin of lateral line
covering about second through eighth lateral-line scale and diffusing over 1 row above and below;
fins dusky; dorsal fins, pectoral fins, and caudal fin light brown; first dorsal fin black distally, remainder
dusky; anal fin hyaline, becoming opaque medially; pelvic fins and pectoral filaments whitish.
Size: Maximum standard length at least 22 cm; an average large fish attains 14 cm standard length.
Habitat, biology, and fisheries: Frequently enters estuaries and is found over sand or sand-mud flats in
depths of 19 to 73 m. Stomach samples included primarily crustaceans, sponges, and fish scales. The
progression in this hermaphrodite is protandrous, changing from a juvenile to a hermaphrodite then a
female. Taken with beach seines and trawls.
Distribution: Algoa Bay (South Africa)
north along the east coast of Africa; east
coast of India and Bay of Bengal; Malay
Peninsula, coasts of Sumatra and Java,
and Borneo; from the Gulf of Thailand
northwards along the Asian coast, including
Taiwan Province of China, to the Yellow
Sea; possibly New Guinea.
Polynemus dubius Bleeker, 1851

Frequent synonyms / misidentifications: Polynemus longipectoralis Weber and de Beaufort, 1922 /
Polynemus paradiseus Linnaeus, 1758.
FAO names: En - Eastern paradise fish.
Diagnostic characters: A small, elongate species; body depth at first dorsal-fin origin 3.7 to 5.3 times in
standard length. Head length 3.5 to 4.2 times in standard length. Eye diameter 1.3 or more times in snout
length; small mouth, upper jaw length 2.1 to 2.6 times in head length, greater than caudal-peduncle depth;
upper lip thin; lower lip thick; tooth patch on vomer a small band or thin “v” shape. Posterior margin of
preopercle with serrations, but less than 25. Gill rakers on first gill arch 23 to 33 (mean 28). First dorsal
fin with VIII spines; second dorsal fin with I spine and 13 to 18 soft rays (mean 16); anal fin with III spines
and 10 to 13 (mean 12) soft rays; anal-fin base shorter than second dorsal-fin base; caudal-peduncle length
pectoral fins with 15 to 18 (mean 17) simple rays; pectoral fins insert about at midpoint on side of body,
reaching to origin or midpoint of anal fin; 7 long pectoral filaments, sixth or seventh filament, from
ventralmost, the longest, extending well past caudal fin, may exceed 1½ total length; pelvic fins seldom
extend past anus. Lateral-line scales 66 to 86 (mean 79); lateral line terminates between centre of
caudal-fin fork and first lower caudal-fin ray; scale rows above lateral line 6 to 7 (mean 7); scale rows below
lateral line 12 to 15 (mean 14). Vertebrae 10+15. No apparent swimbladder. Pyloric caeca about 9. Colour:
body golden yellow with green or grey dorsally and yellow ventrally; fins hyaline to yellowish; spot
immediately above origin of pectoral fins; finely scattered melanophores posterior to eyes; first dorsal,
second dorsal, and caudal fins with light dusky covering, first dorsal fin may be black distally, pectoral fins
usually not black or less than 1/2 of posterior margin of fin black.
Habitat, biology, and fisheries: Regularly enters fresh-water streams from estuaries during the spawning
season, and probably at other times. Sometimes found in abundance as much as 40 km up the Mae Nam
above Bangkok. Large numbers of adults and young caught during the rainy season in the Chao Phraya
River and other rivers of that system using finely meshed seines. Small fish may be used as duck feed and
fertilizer.
Distribution: Tonle Sap (Great Lake) in
Cambodia, the lower Bassac and Mekong
rivers; the Chao Phraya River system in
Thailand, at least to Ayutthaya; the mouth
of the Muar River, Malaysia, Bagan Siapi
Api, Sumatra, and Banjermasin in Borneo.
3114 Bony Fishes
Polynemus hornadayi Myers, 1936

Frequent synonyms / misidentifications: None / Polynemus paradiseus Linnaeus, 1758.
FAO names: En - Hornaday’s paradise fish.
Diagnostic characters: A small, elongate species; body depth at first dorsal-fin origin 3.8 to 4.2 times in
length; upper jaw length 1.4 to 2 times in head length, greater than caudal-peduncle depth; upper lip thin;
lower lip moderate to thick; tooth patch on vomer a small band or thin “v” shape. Posterior margin of
preopercle with serrations, but less than 25. Gill rakers on first gill arch 26 to 28 (mean 26). First dorsal
fin with VII spines; second dorsal fin with I spine and 13 to 16 soft rays (mean 15); anal fin with II spines
and 11 to 12 (mean 11) soft rays; anal-fin base shorter than second dorsal-fin base; caudal-peduncle length
pectoral fins with 17 to 19 (mean 17) simple rays; pectoral fins insert about at midpoint on side of body,
reaching to middle of anal fin; 7 long pectoral filaments, fifth or sixth filament, from ventralmost, the
longest, extending well past caudal fin, may exceed 1½ total length; pelvic fins seldom extend past anus.
Lateral-line scales 90 to 99 (mean 95); lateral line terminates between centre of caudal-fin fork and first
lower caudal-fin ray; scale rows above lateral line 9 to 11 (mean 10); scale rows below lateral line 13 to
18 (mean 16). Vertebrae 10+15. Swimbladder simple, elongate, and moderate in size. Pyloric caeca about
10. Colour: recently preserved in alcohol: yellowish grey or tan; dusky or grey dorsally; fine scattering of
melanophores posterior to eyes; first dorsal, second dorsal, and caudal fins lightly pigmented; pectoral
filaments white.
ies: Muddy, fast flowing riv-
ers, 12 m wide and 3 m deep.
Possibly in estuaries.
from Ensengi River of Sadong
River system below Simujan,
and Kuching market (south-
west Sarawak).
Polynemus melanochir Valenciennes, 1831

Frequent synonyms / misidentifications: Polynemus borneensis Bleeker, 1860; P. hilleri (Fowler,
1905) / Polynemus paradiseus Linnaeus, 1758.
FAO names: En - Blackhand paradise fish.
Diagnostic characters: A small, elongate species; body depth at first dorsal-fin origin 3.8 to 5.4 times
in standard length. Head length 3.2 to 4.1 times in standard length. Eye diameter 1.3 or more times in
snout length; upper jaw length 1.7 to 2.1 times in head length, greater than caudal-peduncle depth; upper
lip thin; lower lip thick; vomer edentate. Posterior margin of preopercle with serrations, but less than 25.
Gill rakers on first gill arch 27 to 34 (mean 30). First dorsal fin with VIII spines; second dorsal fin with I
spine and 14 to 17 soft rays (mean 16); anal fin with III spines and 11 to 13 (mean 12) soft rays; anal-fin
base shorter than second dorsal-fin base; caudal-peduncle length greater than length of anal-fin base;
about 15 to 19 procurrent caudal-fin rays in dorsal or ventral series; pectoral fins with 15 to 18 (mean 17)
simple rays; pectoral fins insert about at midpoint on side of body, reaching to origin or midpoint of
anal fin; 7 long pectoral filaments, sixth or seventh filament, from ventralmost, the longest, extending
well past caudal fin, may exceed 1½ total length; pelvic fins seldom extend past anus. Lateral-line scales
65 to 74 (mean 71); lateral line terminates between centre of caudal-fin fork and first lower caudal-fin ray;
scale rows above lateral line 6 to 8 (mean 7); scale rows below lateral line 12 to 15 (mean 14). Vertebrae
10+15. No apparent swimbladder. Pyloric caeca about 9. Colour: body greenish grey dorsally, ventrally
pale yellow; first dorsal fin with narrow black border, second dorsal fin and anal fin dusky to black; pectoral
fins blue grey or black with yellow or white bases; proximal end of pectoral fins and a few dorsal-fin
rays and connecting membranes, especially distally, not black; size and intensity of the black area variable,
but in adults usually dense black covering more than 1/2 pectoral fins.
Habitat, biology, and fisheries: Lower sections of rivers. Possibly in estuaries.
Distribution: T h e M e k o n g
River system at least as far as
Can Tho and the Bassac
River at least as far as My
Tho; the rivers of Borneo from
Sarawak south to Banjer-
masin.
3116 Bony Fishes
Polynemus multifilis Schlegel, 1843

Frequent synonyms / misidentifications: Polistonemus multifilis Schlegel in Temminck and Schlegel,
1843 / None.
FAO names: En - Elegant paradiseus fish.
Diagnostic characters: A small, elongate species, body depth at first dorsal-fin origin 3.9 to 4.8 times in
length; small mouth, upper-jaw length 2.2 to 2.8 times in head length, less than or equal to
caudal-peduncle depth; upper lip moderate to thick; lower lip thick; tooth patch on vomer a small band or
thin “v” shape; posterior margin of preopercle with less than 25 serrations. Gill rakers on first gill arch
24 to 29 (mean 27). First dorsal fin with VIII spines; second dorsal fin with I spine and 14 to 16 soft rays
(mean 15); anal fin with III spines and 11 to 14 (mean 12) soft rays; anal-fin base shorter than second
dorsal-fin base; caudal-peduncle length greater than length of anal-fin base; about 15 to 19 procurrent
caudal-fin rays in dorsal or ventral series; pectoral fins with 14 to 17 (mean 15) simple rays; pectoral fins
insert about at midpoint on side of body, reaching to origin of anal fin; 14 to 16 (mean 15) long pectoral
filaments, thirteenth or fourteenth filament, from ventralmost, the longest, extending well past caudal
fin, may exceed 1½ total length; pelvic fins seldom extend past anus, pelvic-fin origin to anal-fin origin
relatively long, 2.8 to 3.9 in standard length. Lateral-line scales 87 to 109 (mean 99); lateral line terminates
between centre of caudal-fin fork and first lower caudal-fin ray; scale rows above lateral line 7 to 10 (mean
9); scale rows below lateral line 15 to 22 (mean 18). Vertebrae 10+15. Swimbladder elongate and large.
Pyloric caeca about 12 to 14. Colour: recently preserved specimens (alcohol): yellow, yellowish grey, or
tan, dorsally duskier, grey; spot immediately above origin of pectoral fins; finely scattered melanophores
posterior and ventral to eyes, and over most of opercle; first dorsal, second dorsal, and caudal only lightly
pigmented; dorsomedial and dorsolateral area of pectoral fins sometimes with a small faint spot.
Habitat, biology, and fisheries: Middle and lower sections of larger rivers. Possibly in estuaries. A
specimen collected in the Kapuas River (Borneo), with a standard length of 13.2 cm had pectoral filaments
51 cm in length.
Distribution: Koke Tong Ca-
nal, a tributary of Chao
Phraya River (Thailand); Su-
matra, in the Musi River at
Moeara and the Hari River at
Djambi; Borneo in the Kapuas
River at least to Bunut and at
Banjermassin.
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Perciformes: Percoidei: Sciaenidae 3117
SCIAENIDAE
Croakers (drums)
by K. Sasaki
D iagnostic characters: Moderately elongate, moderately compressed, small to large (to 200 cm
standard length) perciform fishes. Head and body (occasionally also fins) completely scaly,
except tip of snout. Sensory pores often conspicuous on tip of snout (upper rostral pores), on lower edge
of snout (marginal rostral pores), and on chin (mental pores), usually 3 or 5 upper rostral pores, 5 marginal
rostral pores, and 3 pairs of mental pores; these pores usually distinct in bottom feeders with inferior to
subterminal mouth, whereas indistinct in midwater feeders with terminal to oblique mouth. A barbel
sometimes present on chin. Position and size of mouth variable from strongly inferior to oblique, larger in
species with oblique mouth, smaller in species with inferior mouth. Teeth differentiated into large and small
in both jaws or in upper jaw only; enlarged teeth always form outer series in upper jaw, inner series
in lower jaw; well-developed canines (more than twice as large as other teeth) may be present at front of
one or both jaws; vomer and palatine without teeth. Dorsal fin continuous, with deep notch between
anterior (spinous) and posterior (soft) portions; anterior portion with VIII to X slender spines (usually X),
and posterior portion with I spine and 21 to 44 soft rays; base of posterior portion elongate, much longer
than anal-fin base; anal fin with II spines and 6 to 12 (usually 7) soft rays; caudal fin emarginate to
pointed, never deeply forked, usually pointed in juveniles, rhomboidal in adults; pelvic fins with I spine and
5 soft rays, the first soft ray occasionally with a short filament. Scales cycloid (smooth) or ctenoid (rough);
lateral-line scales extending to hind margin of caudal fin. Dorsal side of head (skull) cavernous with
a series of bridge-like bony struts; infraorbitals and preopercle with variously developed ridges
which fold over canals of cephalic lateral-line system. Branchiostegal rays 7, of which 6 on ceratohyal
and 1 on epihyal. Swimbladder well developed with thick wall; carrot-shaped, or hammer-shaped,
with horn-like, tube-like, or arborescent appendages. Appendages may spread behind transverse
septum or entering head beyond the septum. Drumming muscles (sound-producing muscles on body
wall) usually developed in males, less so or absent in females. Sagitta (large earstone) thick, housed in
well-inflated auditory bulla (ear capsule), with a tadpole-shaped impression (or sulcus) on its inner surface.
Colour: highly variable from silvery to dark brown or black, either uniform or some species with spots and
dark bands; juveniles of many species have bands and/or spots on body.
notch
lateral line
swimbladder
II anal-fin
spines
upper rostral marginal
pores rostral pores
mental
lobe of pores
rostral
fold
emarginate truncate rounded
barbel with pore at tip infraorbitals preopercle

S-shaped rhomboidal pointed
ventral view of head bones of head (lateral view)
(Umbrina sp.) shapes of caudal fin
3118 Bony Fishes
Habitat, biology, and fisheries: Primarily inhabit coastal waters on continental shelves, but also occur in
estuaries and rivers, especially in breeding seasons and when juvenile and young. Never found in oceanic
island groups distantly separated from a continental shelf (such as Mariana Islands and Solomon Islands
in the area). A large majority of them are found over muddy or sandy bottoms. Some occur in large shoals
and are the object of sizable fisheries. From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports
a range of yearly catch of around 36 800 to 50 900 t of Sciaenidae from the Western Central Pacific. Actual
yearly catch is probably still higher, since available statistics do not cover small-scale fisheries. Most
croakers feed on small crustaceans, fishes, and benthic organisms. They are usually taken in bottom trawls
and bottom set gill nets.

Other perch-like fishes either with lateral line not extend-
ing to hind margin of caudal fin, or anal fin with III spines.
Centropomidae: also with lateral line extending to hind
margin of caudal fin, but anal fin with III spines.
Apogonidae and Sillaginidae: also with II anal-fin spines,
but lateral line terminated at the base of caudal fin (except
extending onto caudal fin in some apogonids). Centropomidae
Apogonidae Sillaginidae
Identification note
Correct identification of genera of this family is not possible without examination of the swimbladder and
the otoliths. General external features are highly variable even within a single genus, making it difficult to
utilize them as key characters for identification of genera. Users are strongly advised to examine
swimbladders and otoliths for reliable identification. The methods are described below:
1. The swimbladder is located between the viscera and the vertebral column, separated from the head
by a transverse membrane or septum. The body of the swimbladder is readily exposed after gutting
the fish. It becomes necessary to remove organs anteriorly, in order to check the condition of anterior
appendages.
2. Otoliths (earstones) are located in the ear capsules (auditory bulla) on each side of head; 1 pair
(sagitta, see figures below) is large, while the other 2 pairs are rudimentary (except in Johnius). To
examine the otoliths, it is necessary to remove them from the ear capsules by the following method:
while keeping the gill cover open wide, cut the lateral wall of the ear capsule and then extract otolith;
the lateral wall of ear capsule can be easily exposed by removing several muscle bundles connecting
skull and dorsal gill arches.
brain tadpole-shaped
impression
lapillus
sagitta
semi-circular
canals
inner view lateral view

dorsal view of head sagitta of Argyrosomus regius
Key to the genera of Sciaenidae occurring in the area

1a. Swimbladder with horn-like or tube-like appendages; their origins restricted to either
anterior or posterior portion of bladder (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Swimbladder with a number of branched appendages along its entire lateral side (Fig. 2) . . . . . ® 6
a) Atractoscion b) Otolithoides c) Boesemania d) Bahaba e) Panna

Fig. 1 swimbladders
supracleithrum
gill cover
exposed
portion of first
appendage
a) Protonibea b) Johnius c) Johnius cleithrum

Fig. 2 swimbladders
2a. Swimbladder with a pair of horn-like appendages at anterior portion of bladder (Fig. 1a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atractoscion
2b. Swimbladder with 1 to several pairs of tube-like appendages lying parallel to bladder
(Fig. 1b-e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Swimbladder appendages attached to posterior portion of bladder (Fig. 1b) . . . . . . . Otolithoides
3b. Swimbladder appendages attached to anterior portion of bladder (Fig. 1c-e) . . . . . . . . . . . ® 4
4a. Swimbladder with several pairs of tube-like appendages directed posteriorly (Fig. 1c) . . Boesemania
4b. Swimbladder with 1 pair of tube-like appendages directed posteriorly (Fig. 1d-e) . . . . . . . . . ® 5
5a. Swimbladder appendages simple, not divided, directed posteriorly (Fig. 1d) . . . . . . . . . Bahaba
5b. Swimbladder appendages dividing into a forward directed branch and a posteriorly
directed branch (Fig. 1e) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panna
6a. Swimbladder carrot-shaped (Fig. 2a); first appendage does not extend to lateral face of
pectoral arch; sulcus head of sagitta not very oblique, shape of sulcus tail variable but
not deepened as a hollow cone (Fig. 3a-c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6b. S w i m b l a d d e r h a m m e r -
shaped (Fig. 2b); a lateral sulcus
branch of first appendage head
extends to lateral face of
pectoral arch at junction of
cleithrum and supra-
cleithrum (visible under gill sulcus
cover) (Fig. 2c); sulcus tail
head of sagitta very obligue, a) Austronibea b) Atrobucca c) Larimichthys d) Johnius
sulcus tail deepened as a
hollow cone (Fig. 3d) . . . . . . . . ® 19 Fig. 3 sagittae
3120 Bony Fishes
7a. Anterior pair of swimbladder appendages branching behind posterior surface of trans-
verse septum and not entering head (Fig. 4a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7b. Anterior pair of swimbladder appendages entering into head beyond transverse septum
and branching between skull and upper gill arches (Fig. 4b) . . . . . . . . . . . . . . . . . . . ® 14
8a. Outer upper teeth enlarged and spaced, but without outstanding canines (i.e. less than
1/2 length of other teeth) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
8b. One or 2 pairs of outstanding canine teeth in upper jaw or both jaws (i.e. more than 1/2
length of other teeth) (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
transverse
septum
a) Argyrosomus b) Nibea a) Chrysochir b) Otolithes
Fig. 4 anterior portion of swimbladder Fig. 5
9a. First pair of mental pores close together behind tip of jaw and united by a groove
(Fig. 6a); lower fins dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Protonibea
9b. First pair of mental pores on front of chin, 1 on each side of tip of jaw, not united by a
groove (Fig. 6b); lower fins not dark . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Tail of tadpole-shaped impression of sagitta J-shaped (Fig. 7a) . . . . . . . . . . . . . Argyrosomus

10b. Tail of tadpole-shaped impression of sagitta only slightly curved (Fig. 7b) . . . . . . . . . . . ® 11
1st pair united 1st pair separated
tail
a) Protonibea b) Argyrosomus a) Argyrosomus b) Atrobucca
Fig. 6 ventral view of lower jaw (mental pores) Fig. 7 sagitta (inner surface)
dorsal
dorsal limb limb
11a. S w i m b l a d d er ap-
pe ndages without
well-developed dor-
sal limbs (Fig. 8a) . . . . Pennahia
11b. S w i m b l a d d er ap- ventral
pendages with well- limb
d e v e l o pe d d or s al ventral
limbs (Fig. 8b) . . . . . Atrobucca a) Pennahia b) Atrobucca limb
Fig. 8 swimbladder appendages
12a. Snout pointed; mouth inferior; large, outstanding

canines in upper jaw only (Fig. 5a) . . . . . . . . Chrysochir portion branching
on top of bladder
12b. Mouth terminal or lower jaw projecting; large,
outstanding canines on 1 or both jaws (Fig. 5b) . . . . . ® 13
13a. Swimbladder appendages without well-devel-

oped dorsal limbs wrapped around main bladder
(Fig. 9a); tail of tadpole-shaped impression of
sagitta only slightly curved (Fig. 10a) . . . . . . . . Otolithes
13b. Some or all swimbladder appendages with well-
developed dorsal limb branching on top of swim-
bladder (Fig. 9b); tail of tadpole-shaped
impression of sagitta J-shaped (Fig. 10b) . . . . Pterotolithus
a) Otolithes b) Pterotolithus
14a. Lower jaw with a single mental barbel (Fig. 11) . . Dendrophysa Fig. 9 swimbladder appendages
14b. No mental barbel . . . . . . . . . . . . . . . . . . . . . ® 15
15a. Scales on top of head, anterior part of back and belly extremely rough (with enlarged,
erect spinules) (Fig. 12) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aspericorvina
15b. No specialized scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
spinules
mental barbel
a) Otolithes b) Pterotolithus
Fig. 12 scale from top of head
Fig. 10 sagitta (inner surface) Fig. 11 Dendrophysa (Aspericorvina)
16a. Swimbladder appendages with well-devel-

oped dorsal and ventral limbs (Fig. 13a);
scales on anterior half of body cycloid
(smooth) . . . . . . . . . . . . . . . . . . . . . . . ® 17
dorsal
16b. Swimbladder appendages fan-like without limb
well-developed dorsal limbs (Fig. 13b);
scales on body ctenoid (rough to touch) . . . . . . .® 18
ventral
limb
17a. Dorsal-fin rays 28 to 29 . . . . . . . . . . . . . Sonorolux
17b. Dorsal-fin rays 33 to 36 . . . . . . . . . . . Larimichthys
18a. Second anal-fin spine length 23 to 41% of

head length . . . . . . . . . . . . . . . . . . Austronibea a) Larimichthys b) Austronibea
18b. Second anal-fin spine length 38 to 64% of Fig. 13 swimbladder appendages
head length (except for N. semifasciata, 34
to 44% of head length) . . . . . . . . . . . . . . . . Nibea
3122 Bony Fishes
19a. Lower jaw teeth uniform in size or in-

ner row of lower jaw teeth molariform
(Fig. 14a) . . . . . . . . . . . . Johnius (Johnius)
19b. Inner row of lower jaw teeth enlarged,
conical (Fig. 14b) . . . . . . . Johnius (Johnieops) row of
enlarged teeth
a) Johnius (Johnius) b) Johnius (Johnieops)
Fig. 14 dentition
Keys to the species of Sciaenidae

Remarks on key characters: a) a gill raker at the angle of the gill arch and denticulate flat plates are excluded
from the lower gill raker count; b) interorbital width is the least bony width measured above centres of eyes.
Key to the species of Argyrosomus occurring in the area

1a. Caudal fin rhomboidal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Argyrosomus amoyensis
1b. Caudal fin S-shaped or nearly truncate . . . . . . . . . . . . . . . . . . . . Argyrosomus japonicus
Key to the species of Atrobucca occurring in the area

1a. Dorsal-fin rays 23 or 24 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atrobucca adusta
1b. Dorsal-fin rays 27 to 33 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Membrane of gill cover with a distinct black blotch . . . . . . . . . . . . . . . . . Atrobucca kyushini
2b. Membrane of gill cover without a distinct black blotch . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Pectoral fins short, 20 to 24% of standard length; body brown or copper coloured above,
pale orange or white below when fresh . . . . . . . . . . . . . . . . . . . . . . . . Atrobucca brevis
3b. Pectoral fins long, 26 to 31% of standard length; body silvery, whitish below when fresh
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atrobucca nibe
Key to the species of Johnius (Johnius) occurring in the area

1a. Chin with a barbel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Chin without a barbel . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Dorsal-fin rays 23 to 26; scales on body cycloid; caudal fin S-shaped or truncate . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius amblycephalus
2b. Dorsal-fin rays 29 to 34; scales on body ctenoid; caudal fin rhomboidal . . . . Johnius macropterus
3a. Anterior portion of dorsal fin with VIII or IX (usually IX) spines; anal-fin rays 6 or 7
(usually 6); scale rows above lateral line to origin of dorsal fin 10 to 15, scale rows below
lateral line to origin of anal fin 17 to 21; scales on body cycloid . . . . . . . . Johnius trachycephalus
3b. Anterior portion of dorsal fin with IX to XI (usually X) spines; anal-fin rays 7; scale rows
above lateral line to origin of dorsal fin 5 to 11, scale rows below lateral line to origin of
anal fin 7 to 17; scales on body cycloid or ctenoid . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Scales on upper part of body cycloid; caudal fin truncate; a yellow or white streak present
along lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius carutta
4b. Scales on upper part of body ctenoid; caudal fin rhomboidal or pointed (at least in
specimens smaller than 15 cm standard length); a yellow or white streak absent along
lateral line . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Scales on head all cycloid; ctenii on body scales poorly developed; body smooth to touch
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius laevis
5b. Scales on head (at least on interorbital space) ctenoid; ctenii on body scales well
developed; body rough to touch . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Lower jaw with inner short row of molariform teeth posteriorly . . . . . . . . . Johnius macrorhynus
6b. Lower jaw teeth all conical, uniform in size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ®7
7a. Scale rows above lateral line to origin of dorsal fin 5 or 6; scale rows below lateral line to
origin of anal fin 7 to 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7b. Scale rows above lateral line to origin of dorsal fin 6 to 11; scale rows below lateral line
to origin of anal fin 10 to 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
8a. Dorsal-fin rays 31 to 33 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius hypostoma

9a. First gill arch with 6 to 8 gill rakers on lower limb . . . . . . . . . . . . . . . . . . Johnius trewavasae
9b. First gill arch with 9 to 12 gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Second anal-fin spine length 41 to 58% of head length . . . . . . . . . . . . . . . Johnius carouna
10b. Second anal-fin spine length 25 to 34% of head length . . . . . . . . . . . . . . Johnius heterolepis
11a. Dorsal-fin rays 31 to 33; scale rows above lateral line to origin of dorsal fin 5 or 6; scale
rows below lateral line to origin of anal fin 9 or 10; eye diameter 19 to 23% of head length
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius hypostoma
11b. Dorsal-fin rays 25 to 31; scale rows above lateral line to origin of dorsal fin 6 to 11; scale
rows below lateral line to origin of anal fin 10 to 17; eye size variable . . . . . . . . . . . . . . . ® 12
12a. Body dark pigmented; lower fins black . . . . . . . . . . . . . . . . . . . . . . . Johnius belangerii

12b. Body not dark pigmented; lower fins pale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. First gill arch with 7 to 10 gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . ® 14

13b. First gill arch with 9 to 12 gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
14a. Snout well projecting in front of upper jaw; body depth 20 to 26% of standard length . . . Johnius weberi
14b. Snout slightly projecting in front of upper jaw; body depth 25 to 30% of standard length . . . . . ® 15
15a. Interorbital width 26 to 31% of head length; second anal-fin spine length 32 to 37% of
head length (around 7 to 11 cm in standard length), 26 to 32% (around 11 to 16 cm)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius latifrons
15b. Interorbital width 21 to 28% of standard length; second anal-fin spine length 38 to 51%
of head length (around 6 to 11 cm in standard length), 29 to 42% (around 11 to 20 cm)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius australis
16a. Snout bluntly rounded; eye diameter 21 to 28% of head length; vertebrae 25 . . . . Johnius carouna
16b. Snout well pointed; eye diameter 13 to 24% of head length; vertebrae 26 . . . . . . . Johnius coitor
Key to the species of Johnius (Johnieops) occurring in the area

1a. Mental pores in 1 pair only . . . . . . . . . . . . . . . . . . . . . . . . . . . Johnius novaeguineae
1b. Mental pores in 3 pairs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
3124 Bony Fishes
2a. First pair of mental pores separated by symphysis . . . . . . . . . . . . . . . . Johnius plagiostoma

2b. First pair of mental pores by a crescent-shaped groove . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Snout pointed; second anal-fin spine length 24 to 42% of head length . . . . . . Johnius borneensis
3b. Snout bluntly rounded; second anal-fin spine length 45 to 54% of head length . . . Johnius pacificus
Key to the species of Nibea occurring in the area

1a. Lower jaw teeth uniform in size . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Inner row teeth on lower jaw enlarged . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Lateral-line scales 57 to 60 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nibea squamosa

2b. Lateral-line scales 48 to 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nibea microgenys
3a. Snout pointed, projecting in front of upper jaw; second anal-fin spine length 34 to 44%
of head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nibea semifasciata
3b. Snout blunt, not projecting in front of upper jaw; second anal-fin spine length 39 to 59%
of head length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Scale rows above lateral line to origin of dorsal fin 8 to 13, scale rows below lateral line
to origin of anal fin 12 to 17, total scale rows 21 to 29 . . . . . . . . . . . . . . . . . . Nibea soldado
4b. Scale rows above lateral line to origin of dorsal fin 14 to 17, scale rows below lateral line
to origin of anal fin 17 to 23, total scale rows 31 to 39 . . . . . . . . . . . . . . . . . Nibea leptolepis
Key to the species of Panna occurring in the area

1a. Dorsal-fin rays 33 to 36 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panna microdon
1b. Dorsal-fin rays 42 to 44 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Panna perarmatus
Key to the species of Pennahia occurring in the area

1a. Dorsal-fin rays 26 to 29 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pennahia macrocephalus
2a. Mental pores in 3 pairs; caudal fin rhomboidal; a large black bloch on spinous dorsal fin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pennahia pawak
2b. Mental pores in 2 pairs; caudal fin truncate; no large black blotch on spinous dorsal fin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pennahia anea
Key to the species of Pterotolithus occurring in the area

1a. Dorsal-fin rays 24 to 27 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterotolithus lateoides
1b. Dorsal-fin rays 30 to 34 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pterotolithus maculatus

0
0 Argyrosomus amoyensis (Bleeker, 1863)
0 Argyrosomus japaonicus Temminck and Schlegel, 1843
0 Aspericorvina jubata (Bleeker, 1855)
0 Atractoscion aequidens (Cuvier, 1830)
0 Atrobucca adusta Sasaki and Kailola, 1988
0 Atrobucca brevis Sasaki and Kailola, 1988

0 Atrobucca kyushini Sasaki and Kailola, 1988
0 Atrobucca nibe (Jordan and Thompson, 1911)
0 Austronibea oedogenys Trewavas, 1977
0 Bahaba polykladiskos (Bleeker, 1852)
0 Boesemania microlepis (Bleeker, 1858)
0 Chrysochir aureus (Richardson, 1846)
0 Dendrophysa russelli (Cuvier, 1830)
0 Johnius (Johnius) amblycephalus (Bleeker, 1855)
0 Johnius (Johnius) australis (Günther, 1880)
0 Johnius (Johnius) belangerii (Cuvier, 1830)
0 Johnius (Johnius) carouna (Cuvier, 1830)
Johnius (Johnius) cantori (Bleeker, 1874) 1/
0 Johnius (Johnius) carutta Bloch, 1793

0 Johnius (Johnius) coitor (Hamilton, 1822)
0 Johnius (Johnius) heterolepis Bleeker, 1873
0 Johnius (Johnius) hypostoma (Bleeker, 1853)
0 Johnius (Johnius) laevis Sasaki and Kailola, 1991
0 Johnius (Johnius) latifrons Sasaki, 1992
0 Johnius (Johnius) macropterus (Bleeker, 1853)
0 Johnius (Johnius) macrorhynus (Mohan, 1976)
0 Johnius (Johnius) trachycephalus (Bleeker, 1850)
0 Johnius (Johnius) trewavasae Sasaki, 1992
0 Johnius (Johnius) weberi Hardenberg, 1936
0 Johnius (Johnieops) borneensis (Bleeker, 1850)
0 Johnius (Johnieops) novaeguineae (Nichols, 1950)
0 Johnius (Johnieops) pacificus Hardenberg, 1941
0 Johnius (Johnieops) plagiostoma (Bleeker, 1850)
0 Larimichthys pamoides (Munro, 1964)
0 Nibea leptolepis (Ogilby, 1918)
0 Nibea microgenys Sasaki, 1992
0 Nibea semifasciata Chu, Lo, and Wu, 1963
0 Nibea soldado (Lacepède, 1802
0 Nibea squamosa Sasaki, 1992
0 Otolithes ruber (Bloch and Schneider, 1801)
0 Otolithoides biauritus (Cantor, 1850)
0 Panna microdon (Bleeker, 1849)
0 Panna perarmatus (Chabanaud, 1926)
0 Pennahia anea (Bloch, 1793)
0 Pennahia macrocephalus (Tang, 1937)
0 Pennahia pawak (Lin, 1940)
0 Protonibea diacantha (Lacepède, 1802)
0 Pterotolithus lateoides (Bleeker, 1850)
0 Pterotolithus maculatus (Cuvier, 1830)
0 Sonorolux fluminis Trewavas, 1977
Reference
Trewavas, E. 1977. The sciaenid fishes (croakers or drums) of the Indo-West-Pacific. Trans. Zool. Soc. Lond.,
33:253-541.
1/ Known only from a single, half-skin specimen from Malaysia.
click for next page

3126 Bony Fishes
Argyrosomus amoyensis (Bleeker, 1863)

Frequent synonyms / misidentifications: Nibea miichthyoides Chu, Lo, and Wu, 1963 / None.
FAO names: En - Amoy croaker.
mental
pores
Diagnostic characters: A fairly large species with an elongate
body. Mouth large, terminal; upper jaw extending backward below
posterior half of eye; jaws meeting evenly in front; chin without
barbel; teeth well differentiated into large and small in both jaws,
but none canine-like; the large ones widely spaced, forming outer
series in upper jaw, inner series in lower jaw; upper rostral pores
3, marginal rostral pores 5; mental pores in 3 pairs, the first
small, rounded at front of chin, separated by symphysis, the ventral view
others small, slit-like. Gill rakers slender, slightly shorter than gill of head
filaments at angle of arch, 8 to 10 on lower limb of first gill arch.
Dorsal fin with X spines, followed by a notch, second part of fin
with I spine and 25 to 27 soft rays; anal fin with II spines and 7
soft rays, second spine short, slender, 23 to 30% of head
length; caudal fin rhomboidal in adults. Scales cycloid
(smooth) on snout and below eye, elsewhere ctenoid (rough to
touch); lateral-line scales reaching to tip of caudal fin. Swimblad-
der carrot-shaped, with 22 to 34 pairs of arborescent append- sagitta swimbladder
ages of approximately equal size, branching in rather
ragged-looking fan-shape, none entering head. Sagitta (large earstone) with a tadpole-shaped
impression, the tail of which J-shaped. Colour: greyish above and white silvery below; faint oblique
stripes along scale rows on upper half of body; a pale yellow longitudinal stripe above lateral line; a black
spot at pectoral-fin base and a dark blotch on gill cover; spinous dorsal fin dark distally.
Habitat, biology, and fisheries: Inhabits coastal waters over muddy bottoms to a depth of about 60 m.
Caught with bottom trawls, gill nets, and handlines. Marketed fresh; also dried salted; swimbladder dried.
Distribution: S o u t h C h i n a
Sea and off Java, Indonesia;
also in the Arabian Sea from
Oman to western coast of In-
dia.
Argyrosomus japonicus (Temminck and Schlegel, 1843)

Frequent synonyms / misidentifications: Sciaena antarctica Castelnau, 1872 / Argyrosomus
hololepidotus (Lacepède, 1802).
FAO names: En - Japanese meagre.
Diagnostic characters: A large species with elongate body.

Mouth large, terminal; upper jaw extending backward below
posterior half of eye; jaws meeting evenly in front or lower jaw
slightly projecting; teeth in both jaws well differentiated into
large and small, but none canine-like; the large ones widely
spaced, forming outer series in upper jaw, inner series in lower
jaw; upper rostral pores 3, the outer pair minute or obsolete in
larger specimens; marginal rostral pores 5; mental pores in
3 pairs, the first small, rounded at front of chin, separated
by symphysis, the others slit-like. Gill rakers slender, about
2/3 length of gill filaments at angle of arch, 8 to 11 on lower
limb of first gill arch. Dorsal fin with X spines, followed by a
notch, second part of fin with I spine and 25 to 30 soft rays; sagitta
anal fin with II spines and 7 soft rays, second spine short,
slender, 26 to 34% of head length; caudal fin S-shaped or almost truncate. Scales
cycloid (smooth) on snout and below eyes, elsewhere ctenoid (rough to touch); lateral-line swimbladder
scales reaching to hind margin of caudal fin. Swimbladder carrot-shaped, with 21 to 31 (half)
pairs of arborescent appendages, branching in a rather ragged-looking fan-shape,
none entering head. Sagitta (large earstone) with a tadpole-shaped impression, the tail of which
J-shaped. Colour: silvery blue or grey on back shading to whitish on flanks and belly; a series of pearly
spots along lateral line; fins grey or brown; pectoral-fin axil with black spot; edges of jaws and inside of gill
cover yellow.
Habitat, biology, and fisheries: Inhabits lower reaches of rivers, estuaries, rocky reefs, ocean beaches,
and continental shelves down to 150 m. Feeds on fishes, crabs. prawns, and worms. Matures at about 6
years of age and when about 75 cm total length. Annual catches in Australia were about 300 t in 1989-90.
Caught by bottom set gill nets, haul nets, beach seines, and handlines. Common in markets as fresh, gutted
fish in Australia.
Distribution: C e n t r a l a n d
southern, mainland Austra-
lian waters. Also found off
South Africa, southern
Mozambique, and off Oman to
east coasts of India. In the
northern West Pacific, it oc-
c urs from Hong Kong to
southern Korea and Japan.
3128 Bony Fishes
Aspericorvina jubata (Bleeker, 1855)

Frequent synonyms / misidentifications: Johnius melanobranchium Fowler, 1934 / None.
FAO names: En - Prickly croaker.
Diagnostic characters: A small species with a moder-

ately deep body. Snout bluntly rounded, slightly projecting spinules
in front of upper jaw; mouth small, horizontal; upper jaw
extending backward below middle to hind margin of pupil;
upper rostral pores 5, marginal rostral pores 5; mental
pores in 3 pairs, the first small, close together, united
by a crescent-shaped groove just behind symphysis; scale from top of head
teeth differentiated into large and small in upper jaw
only, none canine-like; the large ones forming outer series,
closely spaced, largest at front of jaw; lower jaw with a band
of villiform teeth. Gill rakers short, blunt, about 1/3 length
of gill filaments at angle of arch, 6 to 9 on lower limb of first
gill arch. Dorsal fin with X spines, followed by a notch,
second part of the fin with I spine and 22 to 25 soft rays;
anal fin with II spines and 7 to 9 soft rays, second spine
short, 20 to 28% of head length; caudal fin rhomboidal.
Scales on head, back (to end of spinous part of dorsal
fin), throat and belly ctenoid (rough to touch), with 5 or swimbladder
6 strong projecting spinules on hind margin; else- lateral view of head (half)
where, scales cycloid (smooth); lateral-line scales reach-
ing to tip of caudal fin. Swimbladder carrot-shaped with about 16 pairs of appendages along its sides,
the first pair entering head beyond transverse septum. Sagitta (large earstone) with a tadpole-
shaped impression, the tail of which sharply curved at about an angle of 90° and ending close to
the ventral edge. Colour: body silvery, darker on back; opercle with dark spot.
Size: Maximum standard length 16 cm, commonly from 10 to 15 cm.
Habitat, biology, and fisheries: Inhabits shallow coastal waters, estuaries, and rivers.
Distribution: Gulf of Thai-
land, Sumatra, and Borneo.
Atractoscion aequidens (Cuvier, 1830)

Frequent synonyms / misidentifications: None / Argyrosomus hololepidotus (Lacepède, 1802).
FAO names: En - Geelbeck croaker; Fr - Téraglin; Sp - Corvinata prieta.
Diagnostic characters: A large species with a slender,

elongate body. Snout acutely pointed; mouth large,
strongly oblique; upper jaw extending backward below
hind margin of eye or a little behind; lower jaw projecting
in front of upper jaw; upper and marginal rostral pores
and anterior pair of mental pores absent, second and
third pair represented by minute pores in the skin; all
teeth depressible, those in upper jaw differentiated
into large and small, those in lower jaw about equal
size; none of teeth canine-like. Gill rakers short, den-
tate, about 1/4 length of gill filaments at angle of arch,
6 to 9 on lower limb of first gill arch. Dorsal fin with IX
to X spines, followed by a notch, second part of fin with
I spine and 25 to 32 (28 to 32 in Australia) soft rays;
anal fin with II spines and 9 or 10 soft rays, second
spine very short, slender, 11 to 18% of head length; sagitta swimbladder
caudal fin emarginate to lunate. Scales cycloid
(smooth) on head, ctenoid (rough) on body; lateral-line scales very small, about 70, reaching to hind
margin of caudal fin. Swimbladder carrot-shaped, with a single pair of horn-like appendages curved
between transverse septum and anterior end of bladder, not entering head. Sagitta (large earstone)
with a tadpole-shaped impression, the tail of which J-shaped. Colour: body iridescent blue and purple,
often with faint oblique lines on back and sides; pectoral-fin axil with a black blotch; edges of jaws and
inside of gill cover bright yellow.
Habitat, biology, and fisheries: Inhabits coastal waters down to 200 m; juveniles enter estuaries. Stays
close to bottom during the day and rises to midwater or the surface to feed pelagic fishes during the night.
Caught by handlines and oc-
casionally by traps; also a tar-
get fish in recreation fisheries.
Annual catches in Australia
were 35 t in 1989-90. Mar-
keted fresh; flesh highly es-
teemed.
Distribution: Along eastern
coasts of Australia. Else-
where, off southern Mozam-
b ique, South Afric a, and
southeastern Atlantic north to
Mauritania.
3130 Bony Fishes
Atrobucca adusta Sasaki and Kailola, 1988

FAO names: En - Scorched croaker.
(after Sasaki and Kailola, 1988)
Diagnostic characters: A fairly large species, with a long, slender

caudal peduncle (27 to 30% of standard length). Mouth large, oblique;
upper jaw extending backward below middle to hind margin of eye; chin
angular, without barbel; upper rostral pores 3, marginal rostral pores 5;
mental pores in 3 pairs, the first small, at front of chin, separated by
symphysis; teeth differentiated into large and small in both jaws, but none
canine-like; the large one forming outer series in upper jaw, inner series sagitta
in lower jaw. Gill rakers slender, slightly longer than gill filaments at angle
of arch, 9 or 10 on lower limb of first gill arch. Dorsal fin with X spines,
followed by a notch, second part of fin with I spine and 23 or 24 soft rays;
anal fin with II spines and 7 soft rays, second spine short, slender, 19 to
24% of head length; caudal fin rhomboidal. Scales cycloid (smooth) on
snout, gill cover, and throat, but ctenoid (rough) on other parts of head and
body; lateral-line scales reaching to tip of caudal fin; pectoral fins long,
24 to 27% of standard length. Swimbladder carrot-shaped, with about
30 pairs of branched appendages enveloping the bladder, each with a
well-developed dorsal and ventral limb, regularly arranged so that
terminal branches of dorsal limb point backward, those near tip of
ventral limb forward; no appendages entering head. Sagitta (large swimbladder appendages
earstone) with a tadpole-shaped impression, the tail of which only
slightly curved. Colour: body silvery when alive, becoming chocolate brown immediately after death; in
preservative, scales of belly dark margined, particularly on flanks; posterior mouth lining, gill
chamber, and body cavity black.
Size: Known only from 2 species at 41 and 46 cm standard length.
ies: Inhabits shallow coastal
waters and estuaries.
Distribution: K n o w n f r o m
near Lae, Papua New Guinea.
Atrobucca brevis Sasaki and Kailola, 1988

Frequent synonyms / misidentifications: None / Argyrosomus hololepidotus (Lacepède, 1802).
FAO names: En - Orange croaker.
Diagnostic characters: A medium-sized species, with a moderately

long caudal peduncle (22 to 27% of standard length). Mouth large,
slightly oblique; upper jaw extending backward below hind margin of eye;
chin rounded, without barbel; upper rostral pores 3, marginal rostral pores
5; mental pores in 3 pairs, the first small, at front of chin, separated
by symphysis; teeth differentiated into large and small in both jaws, but
none canine-like; the large ones forming outer series in upper jaw, inner
series in lower jaw. Gill rakers slender, as long as gill filaments at angle of
arch, 10 or 11 on lower limb of first gill arch. Dorsal fin with X spines, sagitta
followed by a notch, second part of fin with I spine and 29 to 32 soft rays;
anal fin with II spines and 7 soft rays, second spine short,
slender, 21 to 28% of head length; caudal fin rhomboidal;
pectoral fins short, 20 to 24% of standard length. Scales
cycloid (smooth) on snout, gill cover, and throat, but ctenoid
(rough) on other parts of head and body; lateral-line scales
reaching to tip of caudal fin. Swimbladder carrot-shaped, with
27 to 30 pairs of branched appendages, each with a well-
developed dorsal and ventral limb, regularly arranged so
that terminal branches of dorsal limb point backward,
those near tip of ventral limb downward; no appendages swimbladder appendages
entering head. Sagitta (large earstone) with a tadpole-
shaped impression, the tail of which only slightly curved. Colour: brown or copper-coloured above,
pale orange or white below; mouth lining pale except grey speckled palate; gill chamber and body cavity
dark brown.
S i z e : Maximum standard
length 26 cm, commonly to
20 cm.
i e s : Inhabits continental
shelves down to 110 m.
Distribution: Along north-
western and northern coast of
Australia and southern Papua
New Guinea.
3132 Bony Fishes
Atrobucca kyushini Sasaki and Kailola, 1988

FAO names: En - Blackspot croaker.
Diagnostic characters: A medium-sized species, with a moder-

ately long caudal peduncle (24 to 28% of standard length).
Mouth large, oblique; upper jaw extending backward below middle
of eye; chin rounded, without barbel; upper rostral pores 3, marginal
rostral pores 5; mental pores in 3 pairs, the first small, at front
of chin, separated by symphysis; teeth differentiated into large
and small in both jaws, but none canine-like; the large ones forming
outer series in upper jaw, inner series in lower jaw. Gill rakers stiff,
equal to or slightly shorter than gill filaments at angle of arch, 9 to
12 on lower limb of first gill arch. Dorsal fin with X spines, followed
by a notch, second part of fin with I spine and 30 to 32 soft rays; sagitta
anal fin with II spines and 7 soft rays, second spine stiff, 30 to 33%
of head length; caudal fin rhomboidal; pectoral fins long, 23 to
27% of standard length. Scales cycloid (smooth) on snout, below
eye, and throat, but ctenoid (rough) on other parts of head and
body; lateral-line scales reaching to tip of caudal fin. Swimbladder
carrot-shaped, with 25 to 28 pairs of branched appendages, each
with a well-developed dorsal and ventral limb, terminal
branches of dorsal and ventral limbs point upward to back-
ward, those near tip of ventral limb forward; no appendages
entering head. Sagitta (large earstone) with a tadpole-shaped
impression, the tail of which slightly curved. Colour: silvery swimbladder appendages
grey above, silvery white below; membrane of gill cover with a
distinct black blotch; mouth
lining pale; pectoral and pel-
vic fins tinged orange.
20 cm.
ies: I n h a b i t s c o n t i n e n t a l
shelves down to 100 m.
Distribution: Off Borneo,
South China Sea.
Atrobucca nibe (Jordan and Thompson, 1911)

FAO names: En - Blackmouth croaker.
Diagnostic characters: A medium-sized species with a moderately

long caudal peduncle (24 to 27% of standard length). Mouth large,
oblique; upper jaw extending backward below hind margin of eye; chin
angular, without barbel; upper rostral pores 3, marginal rostral pores
5; mental pores in 3 pairs, the first small, at front of chin, separated
by symphysis; teeth differentiated into large and small in both jaws,
but none canine-like; the large ones forming outer series in upper jaw,
inner series in lower jaw. Gill rakers slender, as long as gill filaments
at angle of arch, 9 to 12 on lower limb of first gill arch. Dorsal fin with
X spines, followed by a notch, second part of fin with I spine and 27 to
33 soft rays; anal fin with II spines and 7 soft rays, second spine short,
slender, 26 to 31% of head length; caudal fin rhomboidal; pectoral fins
long, 26 to 31% of standard length. Scales cycloid (smooth) on ventral view lateral view
snout, cheek, gill cover, and throat, but ctenoid (rough) on other parts
of head and body; lateral-line scales reaching to tip of caudal fin. swimbladder
Swimbladder carrot-shaped, with 24 to 30 pairs of branched appendages, each with a developed dorsal
and ventral limb, regularly arranged so that terminal branches of dorsal limb point backward, those
near tip of ventral limb forward; no appendages entering head. Sagitta (large earstone) with a
tadpole-shaped impression, the tail of which only slightly curved. Colour: body silvery, whitish below;
lining of mouth, gill chamber, and body cavity black.
Habitat, biology, and fisheries: Usually inhabits continental shelves down to 200 m, but also occurs in
deeper water. In the East China Sea, 50% mature at 24 or 25 cm total length (3 years), 100% mature at
28 to 30 cm total length (4
years).
Distribution: Philippines, In-
donesia, and off northern
Australia; also Chinese and
Japanese waters and disjunc-
tively off South Africa. Re-
cords from the Bay of Bengal
are based on misidentification
o f Atrobucca antonbruun
Sasaki, 1995.
3134 Bony Fishes
Austronibea oedogenys Trewavas, 1977

Frequent synonyms / misidentifications: None / Johnius spp.
FAO names: En - Yellowtailed croaker.
Diagnostic characters: A small species with a slender

body. Snout pointed, swollen, projecting well in front of
upper jaw; mouth inferior; upper jaw extending backward
below hind margin of eye, and extending anteriorly beyond
lower jaw; upper rostral pores 3, marginal rostral pores 5;
mental pores in 3 pairs, the first small, close together,
united by a crescent-shaped groove just behind sym- ventral view of head
physis; teeth differentiated into large and small in both jaws,
but none canine-like; the large ones forming outer series in
upper jaw, inner series in lower jaw. Gill rakers stiff, about
2/3 length of gill filaments at angle of arch, 7 to 12 on lower
limb of first gill arch. Dorsal fin with IX to X (most frequently
X) spines, followed by a notch, second part of fin with I spine
and 27 to 30 soft rays; anal fin with II spines and 7 soft
rays, second spine short, slender, 23 to 41% of head
length; caudal fin rhomboidal. Scales cycloid (smooth) on swimbladder
head and throat, ctenoid (rough) on other parts of body; sagitta (half)
small scales present on soft parts of dorsal fin; lateral-line
scales reaching to tip of caudal fin. Swimbladder carrot-
shaped, with about 20 pairs of fan-like appendages along its sides, the first pair entering head, spread
beyond transverse septum. Sagitta (large earstone) with a tadpole-shaped impression, the tail of
which sharply curved at about an angle of 90° and ending close to the ventral edge. Colour: body
dusky mauve, silvery white below; scales on flanks with orange margins; bluish green blotch on gill cover;
caudal fin yellow with fine
brown stippling.
15 cm.
Distribution: Northern and
northwestern Australia and
southern New Guinea.
Bahaba polykladiskos (Bleeker, 1852)

Frequent synonyms / misidentifications: None / Nibea spp.
FAO names: En - Spine bahaba.
(after Trewavas, 1977)
Diagnostic characters: A fairly large species with a slender caudal

peduncle (26 to 28% of standard length). Mouth large, terminal; upper
jaw extending backward below middle to hind margin of eye; lower jaw
slightly projecting; upper rostral pores 5, marginal rostral pores 5; mental
pores in 3 pairs, the first small, at front of chin, separated by sym-
physis; teeth differentiated into large and small in both jaws, but none
canine-like; the large ones forming outer series in upper jaw, inner series
in lower jaw. Gill rakers stiff, less than 1/2 length of gill filaments at angle
of arch, 7 or 8 on lower limb of first gill arch. Dorsal fin with X spines, followed
by a notch, second part of fin with I spine and 27 to 30 soft rays; anal fin
with II spines and 7 soft rays, second spine long, massive, 55 to 63%
of head length; caudal fin rhomboidal. Scales cycloid (smooth) on snout
and cheek, elsewhere ctenoid (rough to touch); lateral-line scales reaching
to tip of caudal fin; caudal fin thickly covered with small scales. Swim-
bladder carrot-shaped, with only 1 pair of appendages arising at anterior
end and extending, embedded in muscle of abdominal wall, to level of
pelvic-fin tips. Sagitta (large earstone) with a tadpole-shaped impression,
the tail of which sharply curved at about an angle of 90°. Colour: body swimbladder
brown, paler to whitish below.
Size: Maximum standard length to 40 cm, commonly to 30 cm.
Distribution: V i e t N a m ,
Cambodia, and Borneo.
3136 Bony Fishes
Boesemania microlepis (Bleeker, 1858)

Frequent synonyms / misidentifications: None / Nibea soldado (Lacepède, 1802).
FAO names: En - Boeseman croaker.
(after Sasaki, 1991)
Diagnostic characters: A large species with a slender

caudal peduncle (26 to 28% of standard length).
Mouth terminal, slightly oblique; upper jaw extending
backward below hind margin of eye; upper rostral pores
3, marginal rostral pores 5; mental pores in 3 pairs, the
first small, rounded, at front of chin, separated by
symphysis; teeth differentiated into large and small in
both jaws, but none canine-like; the large ones in upper
jaw forming outer series, inner series in lower jaw. Gill
rakers stiff, 2/3 length of gill filaments at angle of arch,
7 to 12 on lower limb of first gill arch. Dorsal fin with IX
to X (most frequently X) spines, followed by a notch, sagitta swimbladder
second part of fin with I spine and 27 to 34 (27 to 30 in
Mekong River Basin, 31 to 34 in Me Nam River Basin) soft rays; anal fin with II spines and 7 soft rays,
second spine long, massive, 44 to 53% of head length; caudal fin pointed. Scales cycloid (smooth) on
snout and cheek, elsewhere ctenoid (rough to touch); lateral-line scales reaching to tip of caudal fin; caudal
fin thickly covered with small scales. Swimbladder carrot-shaped, with a pair of forward-directed
appendages and 5 or 6 pairs of backward-directed appendages, both tube-like and arising at
anterior portion of bladder; former entering head beyond transverse septum, and latter extending
3/5 length of bladder. Sagitta (large earstone) with a tadpole-shaped impression, the tail of which
sharply curved at about an angle of 90°. Colour: greyish brown dorsally, paler below; faint dark oblique
lines on back; fins yellow, dorsal and caudal fin lightly mottled dusky.
S iz e : Maximum standard
length about 100 cm, com-
monly to 50 cm.
ies: Known only from rivers
and estuaries.
Distribution: Thailand, Viet
Nam, Cambodia, and Suma-
tra.
Chrysochir aureus (Richardson, 1846)

Frequent synonyms / misidentifications: Nibea acuta (Tang, 1937) / None.
FAO names: En - Reeve’s croaker; Fr - Courbine dorée; Sp - Corvina dorado.
mental
Diagnostic characters: A medium-sized species with a slen- pores
der body. Snout acutely pointed, projecting in front of
upper jaw; mouth large, nearly horizontal; upper jaw extend-
ing backward below hind margin of eye, and extending ante-
riorly beyond lower jaw; upper rostral pores 3, marginal rostral
pores 5; mental pores in 3 pairs, the first slit-like on either
side of symphysis; teeth differentiated into large and small
in both jaws; large teeth forming outer series in upper jaw, and ventral view
including 1 or 2 pairs of canines more than twice as long of head
as the other enlarged teeth at tip of jaw; large teeth in lower
jaw forming inner series, but not including any canines. Gill
rakers short, slender, about 1/2 length of gill filaments at angle
of gill arch, 8 or 9 on lower limb of first gill arch. Dorsal fin with
X spines, followed by a notch, second part of fin with I spine
and 25 to 28 soft rays; anal fin with II spines and 6 or 7 (most
frequently 7) soft rays, second spine short, slender, 17 to
31% of head length; caudal fin rhomboidal. Scales cycloid
(smooth) on snout and below eye, elsewhere finely ctenoid
(slightly rough to touch); lateral-line scales reaching to tip of
caudal fin. Swimbladder carrot-shaped, with 27 to 30 pairs of
fan-like appendages along its sides, none entering head. sagitta swimbladder
Sagitta (large earstone) with a tadpole-shaped impression, the tail of which only slightly curved,
ending in a disc at the posterior end of sagitta. Colour: metallic blue above, shading to silvery below;
oblique streaks along scale rows on back; pectoral fins yellow, other fins grey suffused with orange.
Habitat, biology, and fisheries: Inhabits shallow coastal waters.
cific, west to the Bay of Ben-
gal, east to southern China
and Indonesia.
3138 Bony Fishes
Dendrophysa russelli (Cuvier, 1830)

Frequent synonyms / misidentifications: Sciaena russelli (Cuvier, 1829) / Johnius amblycephalus
(Bleeker, 1855).
FAO names: En - Goatee croaker; Fr - Bourrugue chévre; Sp - Lambe chivato.
Diagnostic characters: A fairly small species with an oblong

body. Snout steeply rounded, projecting slightly in front of upper
jaw; mouth inferior, upper jaw extending backward below pos-
terior half of eye; a single, pointed barbel on chin; upper
rostral pores 3, marginal rostral pores 5; mental pores in 3 pairs,
the first close together, opening in a common pit at base of
barbel; teeth weakly differentiated into large and small in upper
jaw, large ones not widely spaced, forming outer series; lower
jaw teeth uniformly small. Gill rakers stiff, less than 1/2 length
of gill filaments at angle of arch, 8 to 10 on lower limb of first gill
arch. Dorsal fin with X spines, followed by a notch, second part
of fin with I spine and 25 to 28 soft rays; anal fin with II spines
and 7 soft rays, second spine long, stiff, its length 38 to 47%
of head length; caudal fin rhomboidal. Scales cycloid (smooth) sagitta swimbladder
on snout and below eye, elsewhere ctenoid (rough to touch);
lateral-line scales reaching to tip of caudal fin. Swimbladder carrot-shaped, with about 14 to 17 pairs
of fan-like appendages along its sides, the first pair entering head beyond transverse septum.
Sagitta (large earstone) with a tadpole-shaped impressions, the tail of which bent at a sharp angle and
terminally cutting into the ventral edge. Colour: back grey, shading to white on belly; a dark brown broad
band on nape; opercle with a deep blue blotch; upper edge of spiny part of dorsal fin dark.
Size: M a x i m u m s ta n d a r d
15 cm.
ies: Inhabits coastal waters
and estuaries.
to southern China, Philip-
pines, and Indonesia.
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Johnius (Johnius) amblycephalus (Bleeker, 1855)

Frequent synonyms / misidentifications: Johnius dussumieri (Valenciennes, 1833) / Johnius
macropterus (Bleeker, 1853); Dendrophysa russelli (Cuvier, 1830).
FAO names: En - Bearded croaker; Fr - Courbine barbiche; Sp - Corvina barba.
Diagnostic characters: A small to medium-sized species with

a moderately deep body. Snout steep, bluntly rounded, project-
ing in front of upper jaw; mouth inferior; a stiff, blunt barbel on
chin; upper rostral pores 5, marginal rostral pores 5; mental
pores in 3 pairs, the first open at base of the barbel in a common
pit; teeth differentiated into large and small in upper jaw
only, the large ones close-set, not canine-like, and forming
outer series; lower jaw with a band of villiform teeth. Gill
rakers stiff, less than 1/2 length of gill filaments at angle of arch,
6 to 9 on lower limb of first gill arch. Dorsal fin with X spines,
followed by a notch, second part of fin with I spine and 23 to 26
soft rays, second to fifth spines prolonged; anal fin with II sagitta swimbladder
spines and 7 soft rays, second spine moderately short, 24
to 40% of head length; caudal fin slightly rhomboidal, S-shaped or truncate. Scales small, 7 to 12
scale rows above lateral line to origin of dorsal fin, 13 to 18 scale rows below lateral line to origin
of anal fin; scales on head and body cycloid (smooth to touch); small scales present on soft parts
of dorsal and anal fins; lateral-line scales reaching to hind margin of caudal fin. Swimbladder hammer-
shaped, with 14 or 15 pairs of arborescent appendages along its sides, the first pair entering head
beyond transverse septum and sending a palmate branch to the front of pectoral arch. Sagitta (large
earstone) with a tadpole-shaped impression, the head of which has its long axis lying obliquely to
that of sagitta and the tail expanded and deepened as a hollow cone, connected with the head by
a narrow groove. Colour: back and flanks black or dark brown, belly whitish or cream yellow; upper part
of the rather high spinous part of dorsal fin black.
monly to 15 cm.
cific, west to Pakistan, east to
southern China and north-
eastern Australia.
3140 Bony Fishes
Johnius (Johnius) australis (Günther, 1880)

Frequent synonyms / misidentifications: None / Johnius belangerii (Cuvier, 1830); J. carouna (Cuvier,
1830).
FAO names: En - Bottlenose jewfish.
Diagnostic characters: A small to medium-sized species with a moderately deep

body; body depth 25 to 30% of standard length. Eye rather small, 18 to 24% of head
length. Interorbital width 21 to 28% of head length. Snout steep, bluntly rounded,
slightly projecting in front of upper jaw; mouth small, inferior; upper jaw extending
backward below middle of eye; no barbel on chin; upper rostral pores 5, marginal
rostral pores 5; mental pores in 3 pairs, the first open close together behind symphysis
in a common pit; teeth differentiated into large and small in upper jaw only, the
large ones close-set, not canine-like, and forming outer series; lower jaw with a band
of villiform teeth. Gill rakers blunt, about 1/2 length of gill filaments at angle of
arch, 7 to 10 on lower limb of first arch. Dorsal fin with IX to XI (usually X) spines,
followed by a notch, second part of fin with I spine and 26 to 31 soft rays; anal fin with
swimbladder
II spines and 7 soft rays, second spine long, stiff, 38 to 51% of head length (around
6 to 11 cm in standard length), 29 to 42% (aroud 11 to 20 cm); caudal fin rhomboidal. Scales moderately
small, 8 to 10 scale rows above lateral line to origin of dorsal fin, 11 to 16 scale rows below lateral
line to origin of anal fin; scales cycloid (smooth) on snout, below eye, and throat, ctenoid (rough to
touch) elsewhere; small scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to
tip of caudal fin. Swimbladder hammer-shaped, with about 16 pairs of arborescent appendages along
its sides, the first pair entering head beyond transverse septum and sending a palmate branch to
the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped impression, the head of
which has its long axis lying obliquely to that of sagitta and the tail expanded and deepened as a
hollow cone, connected with the head by a narrow groove. Vertebrae 25. Colour: body silvery,
everywhere so clouded with brown dots as to quite obscure the ground colour; pectoral and pelvic fins
pale.
18 cm.
waters, estuaries, and rivers.
Distribution: Along coasts of
northern and northeastern
Australia and New Guinea.
Johnius (Johnius) belangerii (Cuvier, 1830)

Frequent synonyms / misidentifications: Johnius fasciatus Chu, Lo, and Wu, 1963 / Johnius carouna
(Cuvier, 1830); J. australis (Günther, 1880); J. weberi Hardenberg, 1936.
FAO names: En - Belanger’s croaker; Fr - Courbine de Belanger; Sp - Corvina de Belanger.
Diagnostic characters: A small to medium-sized species with a

moderately deep body; body depth 26 to 32% of standard length. Eye
large, 23 to 28% of head length. Interorbital width 18.5 to 28% of
head length. Snout steep, obtusely rounded, slightly projecting in
front of upper jaw; mouth small, inferior; upper jaw extending back-
ward below middle of eye; no barbel on chin; upper rostral pores 5,
marginal rostral pores 5; mental pores in 3 pairs, the first open close
behind symphysis in a common pit; teeth differentiated into large
and small in upper jaw only, the large ones close-set, not canine-
like, forming outer series; lower jaw with a band of villiform teeth.
Gill rakers very short, club-shaped, less than 1/2 length of gill
filaments at angle of arch, 8 to 10 on lower limb of first gill arch.
Dorsal fin with IX or X (most frequently X) spines, followed by a
notch, second part of fin with I spine and 27 to 31 soft rays; anal fin sagitta swimbladder
with II spines and 7 soft rays, second spine long, stiff, 38 to 49%
of head length; caudal fin rhomboidal. Scales moderately small, 7 to 11 scale rows above lateral line
to origin of dorsal fin, 12 to 17 scale rows below lateral line to origin of anal fin; scales cycloid
(smooth to touch) on snout, strongly ctenoid (very rough to touch) on other parts of head and body;
small scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to tip of caudal
fin. Swimbladder hammer-shaped, with 11 to 15 pairs of arborescent appendages along its sides, the
first pair entering head beyond transverse septum and sending a palmate branch to the front of
pectoral arch. Sagitta (large earstone) with a tadpole-shaped impression, the head of which has its
long axis lying obliquely to that of sagitta and the tail expanded and deepened as a hollow cone,
connected with the head by a narrow groove. Vertebrae 25. Colour: dark pigmented, but pigments
sometimes irregular and irregularly concentrated into short, dark bars along back or on dorsal fin; spinous
part of dorsal fin black, lower fins also black in many cases; dark blotch shows through gill cover.
length 20 cm, commonly
15 cm.
waters down to 40 m.
cific, west to the Persian Gulf,
east to southern China and
Indonesia.
3142 Bony Fishes
Johnius (Johnius) carouna (Cuvier, 1830)

Frequent synonyms / misidentifications: None / Johnius australis (Günther, 1880); J. belangerii
(Cuvier, 1830); J. weberi Hardenberg, 1936.
FAO names: En - Caroun croaker; Fr - Courbine caroune; Sp - Corvina caruna.
Diagnostic characters: A small to medium-sized species with a moderately deep

body; body depth 26 to 33% of standard length. Eye large, 21 to 28% of head length.
Interorbital width 22 to 28% of head length. Snout bluntly rounded, slightly projecting
in front of upper jaw; mouth small, inferior; upper jaw extending backward below
middle to hind margin of eye; no barbel on chin; upper rostral pores 5, marginal rostral
pores 5; mental pores in 3 pairs, the first open close behind symphysis in a common
pit; teeth differentiated into large and small in upper jaw only, the large ones
close-set, not canine-like, forming outer series; lower jaw with a band of villiform
teeth. Gill rakers short, stiff, about 1/2 length of gill filaments at angle of arch, 10
to 12 on lower limb of first gill arch. Dorsal fin with X spines, followed by a deep
notch, second part of fin with I spine and 26 to 30 soft rays; anal fin with II spines and
7 soft rays, second spine long, stiff, 41 to 58% of head length; caudal fin rhomboidal.
Scales moderately large, 6 to 9 scale rows above lateral line to origin of dorsal swimbladder
fin, 10 to 14 scale rows below lateral line to origin of anal fin; scales cycloid
(smooth) on snout, cheek, and throat, but ctenoid (rough to touch) on other parts of head and body;
fin. Swimbladder hammer-shaped, with 14 or 15 pairs of arborescent appendages along its sides, the
long axis lying obliquely to that of sagitta and the tail expanded and deepened as a hollow cone
connected with the head by a narrow groove. Vertebrae 25. Colour: upper 2/3 of body light grey or with
a white sheen, lower 1/3 yellowish; pectoral, pelvic, and anal fins and lower part of caudal fin with
yellow tinge.
16 cm.
waters, entering estuaries
and mangrove swamps.
land, Sumatra, and Borneo;
north to southern China, west
to India.
Johnius (Johnius) carutta Bloch, 1793

FAO names: En - Karut croaker; Fr - Courbine carutte; Sp - Corvina carota.

a moderately deep body; body depth 28 to 32% of standard
length. Eye small to moderate sized, 19 to 24% of head
length. Interorbital width 25 to 31% of head length. Snout
bluntly rounded, slightly projecting in front of upper jaw;
mouth small, inferior; upper jaw extending backward below
middle of eye; no barbel on chin; upper rostral pores 5,
marginal rostral pores 5; mental pores in 3 pairs, the first open sagitta
close behind symphysis in a common pit; teeth differentiated
into large and small in upper jaw only, the large ones
close-set, not canine-like, forming outer series; lower jaw with
a band of villiform teeth. Gill rakers short, stiff, less than
1/2 length of gill filaments at angle of arch, 8 or 9 on lower
limb of first gill arch. Dorsal fin with X spines, followed by a
notch, second part of fin with I spine and 26 to 29 soft rays; anal
fin with II spines and 7 soft rays, second spine short, 25 to first gill arch swimbladder
35% of head length; caudal fin truncate or weakly S-
shaped. Scales small, 7 to 9 scale rows above lateral line to origin of dorsal fin, 12 to 16 scale rows below
lateral line to origin of anal fin; scales cycloid (smooth) on head and upper part of body, weakly ctenoid
(slightly rough to touch) on other parts of body; small scales present on soft parts of dorsal and anal
fins; lateral-line scales reaching to hind margin of caudal fin. Swimbladder hammer-shaped, with about
16 pairs of arborescent appendages along its sides, the first pair entering head beyond transverse
septum and sending a palmate branch to the front of pectoral arch. Sagitta (large earstone) with a
tadpole-shaped impression, the head of which has its long axis lying obliquely to that of sagitta
and the tail expanded and deepened as a hollow cone connected with the head by a narrow groove.
Vertebrae 25. Colour: upper two-thirds of body dark grey or brown with a yellow or white streak along
lateral line; belly whitish; up-
per two-thirds of first dorsal fin
black.
20 cm.
Distribution: West coasts of
Malay Peninsula to Pakistan.
3144 Bony Fishes
Johnius (Johnius) coitor (Hamilton, 1822)

Frequent synonyms / misidentifications: None / Johnius laevis Sasaki and Kailola, 1991.
FAO names: En - Coitor croaker.
(after Bleeker, 1950)
Diagnostic characters: A small species with a slender to moderately deep body; body
depth 24 to 30% of standard length. Eye small, 13 to 24% of head length. Interorbital
width 22 to 29% of head length. Snout pointed, projecting in front of upper jaw; mouth
small, inferior; upper jaw extending backward below middle of eye; no barbel on chin;
upper rostral pores 5, marginal rostral pores 5; mental pores in 3 pairs, the first open
close behind symphysis in a common pit; teeth differentiated into large and small in
upper jaw only, the large ones close-set, not canine-like, forming outer series; lower
jaw with a band of villiform teeth. Gill rakers slender, 1/2 to 2/3 length of gill
filaments at angle of arch, 9 to 12 on lower limb of first gill arch. Dorsal fin with IX
to XI (most frequently X) spines, followed by a deep notch, second part of fin with I spine
and 26 to 29 soft rays; anal fin with II spines and 6 to 8 (most frequently 7) soft rays,
second spine long, stiff, 37 to 59% of head length; caudal fin rhomboidal. Scales
small, 7 to 10 scale rows above lateral line to origin of dorsal fin, 11 to 16 scale rows
below lateral line to origin of anal fin; scales cycloid (smooth) on snout, ctenoid (rough
to touch) on other parts of head and body; small scales present on soft parts of dorsal
and anal fins; lateral-line scales reaching to tip of caudal fin. Swimbladder hammer- swimbladder
shaped, with 14 or 15 pairs of arborescent appendages along its sides, the first
pair entering head beyond transverse septum and sending a palmate branch to
the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped impres-
sion, the head of which has its long axis lying obliquely to that of sagitta and the
tail expanded and deepened as a hollow cone connected with the head by a narrow
groove. Vertebrae 26. Colour: light golden yellow with a light purple or blue sheen; spinous part of dorsal
fin with dusky to black border; soft parts of dorsal, anal and caudal fins with a dull green or grey border.
Distribution: Borneo and
Malay Peninsula, east to In-
dia.
Johnius (Johnius) heterolepis Bleeker, 1873

FAO names: En - Largescale croaker.
(after Sasaki., 1992)
Diagnostic characters: A small species with a moderately

deep body; body depth 25 to 27% of standard length. Eye small
to moderately large, 19 to 27% of head length. Interorbital width
23 to 28% of head length. Snout pointed, projecting slightly in
front of upper jaw; mouth small, inferior; upper jaw extending
backward below hind margin of pupil; no barbel on chin; upper
rostral pores 5, marginal rostral pores 5; mental pores in 3 pairs,
the first open close behind symphysis in a common pit; teeth
differentiated into large and small in upper jaw only, the
large ones close-set, not canine-like, forming outer series;
first gill arch
lower jaw with a band of villiform teeth. Gill rakers slender,
about 1/2 length of gill filaments at angle of arch, 9 to 11 on
lower limb of first gill arch. Dorsal fin with X or XI (usually X) swimbladder
spines, followed by a deep notch, second part of fin with I spine and 25 to 28 soft rays; anal fin with II
spines and 7 soft rays, second spine short, slender, 25 to 34% of head length; caudal fin rhomboidal.
Scales large, those on flanks much larger than those on lateral line; 5 scale rows above lateral line
to origin of dorsal fin, 10 or 11 scale rows below lateral line to origin of anal fin; scales cycloid
(smooth) on snout, cheek, and anterior half of gill cover, ctenoid (rough to touch) on other parts of head
and body; small scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to tip
of caudal fin. Swimbladder hammer-shaped, with about 14 pairs of arborescent appendages along
the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped impression, the tail of
hollow cone connected with the head by a narrow groove. Vertebrae 25. Colour: greenish or bluish
grey above, silvery below;
fins yellowish.
Size: Maximum standard
12 cm.
Habitat, biology, and fish-
eries: I n h a b i t s s h a l l o w
coastal waters.
Distribution: Malaysia and
Indonesia.
3146 Bony Fishes
Johnius (Johnius) hypostoma (Bleeker, 1853)

FAO names: En - Smallmouth croaker.
Diagnostic characters: A small species with a moderately deep body; body depth 24
to 34% of standard length. Eye small, 19 to 23% of head length. Interorbital width 28
to 34% of head length. Snout bluntly rounded, projecting slightly in front of upper jaw;
mouth small, inferior; upper jaw extending backward below posterior half of eye; no
barbel on chin; upper rostral pores 5, marginal rostral pores 5; mental pores in 3 pairs,
the first open close behind symphysis in a common pit; teeth differentiated into large
and small in upper jaw only, the large ones close-set, not canine-like, forming outer
series; lower jaw with a band of villiform teeth. Gill rakers slender, about 1/2 length
of gill filaments at angle of arch, 11 or 12 on lower limb of first gill arch. Dorsal fin
with IX to XI (most frequently X) spines and 31 to 33 soft rays; anal fin with II spines
and 7 soft rays, second spine moderately long, stiff, 30 to 48% of head length; caudal
fin rhomboidal. Scales large, those on flanks much larger than those on lateral line;
5 or 6 scale rows above lateral line to origin of dorsal fin, 9 or 10 scale rows below
lateral line to origin of anal fin; scales cycloid (smooth) on snout and cheek, ctenoid
(rough to touch) on other parts of head and body; small scales present on soft parts swimbladder
of dorsal and anal fins; lateral-line scales reaching to tip of caudal fin. Swimbladder
hammer-shaped, with about 10 pairs of arborescent appendages along its sides, the first pair
entering head beyond transverse septum and sending a palmate branch to the front of pectoral
arch. Sagitta (large earstone) with a tadpole-shaped impression, the head of which has its long axis
lying obliquely to that of sagitta and the tail expanded and deepened as a hollow cone connected
with the head by a narrow groove. Vertebrae 25. Colour: greenish above, silvery below; fins yellowish.
waters.
from Indonesia.
Johnius (Johnius) laevis Sasaki and Kailola, 1991

Frequent synonyms / misidentifications: None / Johnius coitor (Hamilton, 1822).
FAO names: En - Smooth croaker.
Diagnostic features: A small species with a deep body;

body depth 27 to 34% of standard length. Eye large, 22
to 30% of head length. Interorbital width 25 to 30% of
head length. Snout pointed, projecting well in front of
upper jaw; mouth small, inferior; upper jaw extending
backward below middle to posterior margin of eye; no
barbel on chin; upper rostral pores 5, marginal rostral
pores 5; mental pores in 3 pairs, the first open close
behind symphysis in a common pit; teeth differentiated first gill arch scales
into large and small in upper jaw only, the large ones
close-set, not canine-like, forming outer series; lower jaw with a band of villiform teeth. Gill rakers
slender, about 1/2 length of gill filaments at angle of arch, 10 to 12 on lower limb of first gill arch.
Dorsal fin with X to XI (usually X) spines, followed by a notch, second part of fin with I spine and 29 to
34 soft rays; anal fin with II spines and 7 soft rays, second spine moderately long, stiff, 29 to 52%
of head length; caudal fin rhomboidal. Scales large, those on flanks much larger than those on lateral
line, 5 or 6 scale rows above lateral line to origin of dorsal fin, 8 to 10 scale rows below lateral line
to origin of anal fin; scales cycloid (smooth) on head and throat, weakly ctenoid on other parts of
body, ctenii (small spines along posterior margin of scales) poorly developed, smooth to touch;
fin. Swimbladder hammer-shaped, with 11 to 14 pairs of arborescent appendages along its sides, the
pectoral arch. Sagitta (large earstone) with a tadpole-like impression, the head of which has its long
axis lying obliquely to that of sagitta and the tail expanded as a hollow cone connected with the
head by a narrow groove. Vertebrae 25. Colour: head and upper body iridescent mauve or bronze, lower
part of body silvery white; fins
yellow, black margins to dor-
sal and caudal fins.
11 cm.
ies: Inhabits coastal waters
down to 60 m.
Distribution: Northern Aus-
tralia and southern Papua
New Guinea.
3148 Bony Fishes
Johnius (Johnius) latifrons Sasaki, 1992

FAO names: En - Broadhead croaker.

deep body; body depth 25 to 30% of standard length. Eye
small to moderately large, 17 to 26% of head length.
Interorbital width 26 to 31% of head length. Snout
steeply rounded, projecting slightly in front of upper jaw;
mouth small, inferior, upper jaw extending backward below
middle of eye; no barbel on chin; upper rostral pores 5,
marginal rostral pores 5; mental pores in 3 pairs, the first
open close behind symphysis in a common pit; teeth dif-
ferentiated into large and small in upper jaw only, the
large ones close-set, not canine-like, forming outer series; first gill arch swimbladder
lower jaw with a band of villiform teeth. Gill rakers stiff, about 1/3 length of gill filaments at angle
of arch, 7 to 9 on lower limb of first gill arch. Dorsal fin with X spines, followed by a deep notch, second
part of fin with I spine and 25 to 29 soft rays; anal fin with II spines and 7 soft rays, second spine short,
stiff, 32 to 37% of head length (around 7 to 11 cm in standard length), 26 to 32% (around 11 to 16 cm);
caudal fin rhomboidal. Scales moderate in size, 7 to 9 scale rows above lateral line to origin of dorsal
fin, 11 to 14 scale rows below lateral line to origin of anal fin; scales cycloid (smooth) on snout, ctenoid
(rough to touch) on other parts of head and body; small scales present on soft parts of dorsal and anal
fins; lateral-line scales reaching to tip of caudal fin. Swimbladder hammer-shaped, with about 14 pairs
of arborescent appendages along its sides, the first pair entering head beyond transverse septum
and sending a palmate branch to the front of pectoral arch. Sagitta (large earstone) with a
Vertebrae 25. Colour: light
brown dorsally, creamy white
ventrally.
12 cm.
waters.
Distribution: Thailand and
Indonesia.
Johnius (Johnius) macropterus (Bleeker, 1853)

Frequent synonyms / misidentifications: Sciaena macroptera (Bleeker, 1853) / Johnius amblycephalus
(Bleeker, 1855).
FAO names: En - Largefin croaker; Fr - Courbine à grande nagoire; Sp - Corvina aletona.

deep body. Eye small to moderately large, 18 to 26% of head
length. Snout rounded, slightly projecting in front of upper jaw;
mouth small, inferior; upper jaw extending backward below
middle of eye; upper rostral pores 5, marginal rostral pores 5;
mental pores in 3 pairs, the first open close behind symphysis
in a common pit; a short, stiff barbel at chin; teeth differen-
tiated into large and small in upper jaw only, the large ones
close-set, not canine-like, forming outer series; lower jaw
with a band of villiform teeth. Gill rakers stiff, less than 1/2
length of gill filaments at angle of arch, 7 to 12 on lower limb
of first gill arch. Dorsal fin with X spines, followed by a notch, sagitta swimbladder
second part of fin with I spine and 29 to 34 soft rays; anal fin
with II spines and 7 soft rays, second spine stiff, 32 to 56% of head length; caudal fin rhomboidal.
Scales large, those on flanks much larger than those on lateral line, 4 to 7 scale rows above lateral
line to origin of dorsal fin, 9 to 12 scale rows below lateral line to origin of anal fin; scales cycloid
(smooth) on head, ctenoid (rough to touch) on body; small scales present on soft parts of dorsal
and anal fins; lateral-line scales reaching to tip of caudal fin. Swimbladder hammer-shaped, with 14 or
15 pairs of arborescent appendages along its sides, the first pair entering head beyond transverse
septum and sending a palmate branch to the front of pectoral arch. Sagitta (large earstone) with a
Colour: back dark brown, flanks and belly whitish with silvery reflections; spinous part of dorsal fin dark;
pectoral fins hyaline.
Size: M a x i m u m s ta n d a r d
10 cm.
waters.
cific, west to India and Sri
Lanka, east to New Guinea.
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3150 Bony Fishes
Johnius (Johnius) macrorhynus (Mohan, 1976)

Frequent synonyms / misidentifications: Johnieops macrorhynus Mohan, 1976 / Johnius weberi
Hardenberg, 1836.
FAO names: En - Bigsnout croaker; Fr - Courbine longnez; Sp - Corvina hocicona.
Diagnostic characters: A medium-sized species with an elongate body;

body depth 24 to 32% of standard length. Eye small to moderately large,
18 to 26% of head length. Interorbital width 20 to 25% of head length.
Snout bluntly rounded, projecting slightly in front of upper jaw; mouth
small, inferior; upper jaw extending backward below middle of eye; no
barbel on chin; upper rostral pores 5, marginal rostral pores 5; mental
pores in 3 pairs, the first open close behind symphysis in a common pit;
teeth differentiated into large and small in upper jaw, the large ones
close-set, not canine-like, forming outer series in upper jaw; lower jaw
with an outer villiform teeth band and an inner, short row of molari-
form teeth posteriorly. Gill rakers short, stumpy, about 1/4 length of sagitta
gill filaments at angle of arch, 5 to 9 on lower limb of first gill arch. Dorsal fin with X spines, followed
by a notch, second part of fin with I spine and 27 to 30 soft rays; anal fin with II spines and 7 soft rays,
second spine rather short, slender, 23 to 39% of head length; caudal fin rhomboidal, becoming
truncate or S-shaped in specimens over 15 cm standard length. Scales moderately large, 6 to 9 scale
rows above lateral line to origin of dorsal fin, 10 to 14 scale rows below lateral line to origin of anal fin;
scales cycloid (smooth) on snout and below eyes, ctenoid (rough to touch) on other parts of head and
body; small scales present on soft parts of dorsal and anal fins; lateral-line scales extending to hind
margin of caudal fin. Swimbladder hammer-shaped, with 13 or 14 pairs of arborescent appendages
along its sides, the first pair entering head beyond transverse septum and sending a palmate branch
to the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped impression, the head
of which has its long axis lying obliquely to that of sagitta and the tail expanded and deepened as
a hollow cone connected with the head by a narrow groove. Vertebrae 25. Colour: lower part of body
with a golden tinge; pectoral, pelvic, and anal fins yellow; a faint steel blue blotch on gill cover.
waters.
Distribution: Malay Penin-
sula, South China Sea, Bor-
neo, east to India and
Pakistan.
Johnius (Johnius) trachycephalus (Bleeker, 1850)

Frequent synonyms / misidentifications: Otolithoides siamensis Fowler, 1934 / Johnius coitor
(Hamilton, 1822).
FAO names: En - Leaftail croaker.
Diagnostic characters: A small species with a slender body; body depth 21 to

23% of standard length. Eye small, 18 to 21% of head length. Snout prominent,
swollen, projecting well in front of upper jaw; mouth small, inferior; upper jaw
extending backward below middle of eye; no barbel on chin; upper rostral pores
5, marginal rostral pores 5; mental pores in 3 pairs, the first open close behind
symphysis in a common pit; teeth differentiated into large and small in upper
jaw only, the large ones close-set, not canine-like, forming outer series; lower jaw
with a band of villiform teeth. Gill rakers slender, about 1/2 length of gill
filaments at angle of arch, 11 to 13 on lower limb of first gill arch. Dorsal fin
with VIII to IX (most frequently IX) spines, followed by a notch, second part of
fin with I spine and 24 to 27 soft rays; anal fin with II spines and 6 or 7 (most
frequently 6) soft rays, second spine stiff, 31 to 45% of head length; caudal
fin pointed. Scales small, 10 to 15 scale rows above lateral line to origin of
dorsal fin, 17 to 21 scale rows below lateral line to origin of anal fin; scales
on head and body cycloid (smooth); small scales present on soft parts of swimbladder
dorsal and anal fins; lateral-line scales reaching to tip of caudal fin. Swimbladder
hammer-shaped, with 11 to 13 pairs of arborescent appendages along its sides,
the first pair entering head beyond transverse septum and sending a palmate branch to the front
of pectoral arch. Sagitta (large earstone) with a tadpole-shaped impression, the head of which has
its long axis lying obliquely to that of sagitta and the tail expanded and deepened as a hollow cone
connected with the head by a narrow groove. Vertebrae 25. Colour: back grey, flanks lighter, belly
silver; fins yellowish.
Size: Maximum standard length 13 cm, commonly to about 10 cm.
Distribution: Sumatra, Bor-
neo, and Thailand.
3152 Bony Fishes
Johnius (Johnius) trewavasae Sasaki, 1992

Frequent synonyms / misidentifications: None / Johnius macrorhynus (Mohan, 1976).
FAO names: En - Trewavas’ croaker.
Diagnostic characters: Small species with a moderately

elongate body; body depth 24 to 30% of standard length. Eye
moderately large, 22 to 27% of head length. Interorbital width
22 to 24% of head length. Snout steep, obtusely rounded,
projecting slightly in front of upper jaw; mouth small, inferior;
upper jaw extending backward below front margin of pupil; no
barbel on chin; upper rostral pores 5, marginal rostral pores
5; mental pores in 3 pairs, the first open close behind sym-
physis in a common pit; teeth differentiated into large and
small in upper jaw only, the large ones close-set, not canine- first gill arch
like, forming outer series; lower jaw with a band of villiform swimbladder
teeth. Gill rakers short, obtuse, about 1/5 length of gill
filaments at angle of arch, 6 to 8 on lower limb of first gill arch. Dorsal fin with X to XI (most frequently
X) spines, followed by a notch, second part of fin with I spine and 24 to 27 soft rays; anal fin with II
spines and 7 or 8 (most frequently 7) soft rays, second spine rather short, slender, 25 to 37% of head
length; caudal fin rhomboidal. Scales large, those on flanks much larger than those on lateral line, 5
or 6 scale rows above lateral line to origin of dorsal fin, 7 to 10 scale rows below lateral line to origin
of anal fin; scales cycloid (smooth) on head and throat, ctenoid (rough to touch) on other parts of head
and body; small scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to tip
of caudal fin. Swimbladder hammer-shaped, with about 14 pairs of arborescent appendages along
the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped impression, the head of
hollow cone connected with the head by a narrow groove. Vertebrae 25. Colour: dark mottled pattern
dorsally, creamy white ventrally; scale pockets on dorsal side of head and body with broad, dark margins;
spinous dorsal fin black, pectoral fins grey, pelvic fins pale.
14 cm.
Distribution: S i n g a p o r e ,
north to Taiwan Province of
China, Hong Kong, and
Shanghai.
Johnius (Johnius) weberi Hardenberg, 1936

Frequent synonyms / misidentifications: None / Johnius macrorhynus (Mohan, 1976).
FAO names: En - Weber’s croaker.
Diagnostic characters: A small species with a slender, elongate body; body

depth 20 to 26% of standard length. Eye moderately large, 21 to 28% of head
length. Interorbital width 20 to 25% of head length. Snout swollen, projecting
well in front of upper jaw; mouth small, inferior; upper jaw extending backward
below middle of eye; no barbel on chin; upper rostral pores 5, marginal rostral
pores 5; mental pores in 3 pairs, the first open close behind symphysis in a
common pit; teeth differentiated into large and small in upper jaw only, the
large ones close-set, not canine-like, forming outer series; lower jaw with a band
of villiform teeth. Gill rakers stiff, 1/3 to 1/2 length of gill filaments at angle
of arch, 7 to 9 on lower limb of first gill arch. Dorsal fin with X spines, followed
by a notch, second part of fin with I spine and 26 to 29 soft rays; anal fin with II
spines and 7 soft rays, second spine moderately short to moderately long,
slender, 33 to 41% of head length; caudal fin rhomboidal. Scales moderately
large, 7 to 10 scale rows above lateral line to origin of dorsal fin, 11 to 14 scale
rows below lateral line to origin of anal fin; scales cycloid (smooth) on snout,
swimbladder
below eye, cheek, gill cover, and throat, but ctenoid (rough to touch) on other
parts of head and body; small scales present on soft parts of dorsal and anal fins; lateral-line scales
reaching to tip of caudal fin. Swimbladder hammer-shaped, with about 15 pairs of arborescent
appendages along its sides, the first pair entering head beyond transverse septum and sending a
palmate branch to the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped
impression, the head of which has its long axis lying obliquely to that of sagitta and the tail
expanded and deepened as a hollow cone connected with the head by a narrow groove. Vertebrae
25. Colour: body silvery; spinous dorsal fin dusky, pectoral and pelvic fins pale.
Habitat, biology, and fisheries: Inhabits shallow coastal waters and estuaries.
land, Sumatra, and Borneo.
3154 Bony Fishes
Johnius (Johnieops) borneensis (Bleeker, 1850)

Frequent synonyms / misidentifications: Johnius vogleri (Bleeker, 1853) / Johnius sina (non Cuvier,
1830).
FAO names: En - Sharptooth hammer croaker; Fr - Courbine dentue; Sp - Corvina dientusa.
Diagnostic characters: A medium-sized species with a deep

body. Eye moderately large, 20 to 25% of head length. Snout
pointed, but not swollen or projecting; mouth large,
oblique; upper jaw extending backward below posterior
half of eye; no barbel on chin; upper rostral pores 3, mar- sagitta jaws
ginal rostral pores 5; mental pores in 3 pairs, the first open
close behind symphysis connected by a crescent-shaped groove; teeth in narrow bands, well differen-
tiated into large and small in both jaws, but none canine-like; the large ones widely spaced, forming
outer series in upper jaw, inner series in lower jaw. Gill rakers slender, about 1/2 length of gill filaments
at angle of arch, 9 to 15 on lower limb of first gill arch. Dorsal fin with IX to XI (most frequently X)
spines, followed by a notch, second part of fin with I spine and 26 to 32 soft rays; anal fin with II spines
and 7 or 8 (most frequently 7) soft rays, second spine moderately long, 24 to 42% of head length;
caudal fin rhomboidal. Scales on head cycloid (smooth), those on body ctenoid (rough to touch); small
scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to tip of caudal fin.
Swimbladder hammer-shaped, with 14 to 17 pairs of arborescent appendages along its sides, the
long axis lying obliquely to that of sagitta and the tail expanded and deepened as a hollow cone
connected with the head by a narrow groove. Colour: head and body dusky mauve above, silver-white
below; 2 dusky streaks usually present along mid-sides; fins pale yellow or cream, outer two-thirds of
spinous dorsal fin black, sometimes a dusky streak along soft dorsal fin.
Habitat, biology, and fisheries: Inhabits shallow coastal waters, estuaries, and rivers. In India, it matures
at the size of 14 to 16 cm total length; attains an average size of 15.8 cm, 20.6 cm, and 24 cm total length,
at the end of 1 year, 1 ½ years,
and 2 years, respectively.
Crustaceans are the preferred
diet of juveniles; fishes show
piscivorous tendency as they
grow.
east to southern China, Tai-
wan Province of China, and
northern and northeastern
Australia and New Guinea.
Johnius (Johnieops) novaeguineae (Nichols, 1950)

FAO names: En - Paperhead croaker.
Diagnostic characters: A small species with a deep, swollen head. Eye very
small, 11 to 16% of head length. Snout very steeply rounded, but not swollen
or projecting; mouth very large, strongly oblique; upper jaw extending back-
ward well beyond hind margin of eye; chin angular, prominent; no barbel; no upper
rostral pores, marginal rostral pores 5; mental pores in 1 pair only, open behind
symphysis; teeth well differentiated into large and small in both jaws, but none
canine-like; the large ones widely spaced, forming outer series in upper jaw, inner
series in lower jaw. Gill rakers long, slender, equal to or longer than gill
filaments at angle of arch, 16 to 18 on lower limb of first gill arch. Dorsal fin
with VIII to X (most frequently IX) spines, followed by a notch, second part of
fin with I spine and 25 to 28 soft rays; anal fin with II spines and 6 to 8 (usually
7) soft rays, second spine long, slender, 31 to 44% of head length; caudal fin
acutely pointed. Scales cycloid (smooth) on head and throat, ctenoid (rough to
touch) on other parts of body; small scales present on soft parts of dorsal and
anal fins; lateral-line scales reaching to tip of caudal fin. Swimbladder hammer-
shaped, with 11 pairs of arborescent appendages along its sides, the first very
short, the second sending a palmate branch to the front of pectoral arch.
Sagitta (large earstone) with a tadpole-shaped impression, the head of which
has its long axis lying obliquely to that of sagitta and the tail expanded and
deepened as a hollow cone connected with the head by a narrow groove.
Colour (in preservative): pale pinkish brown, paler below, without markings, fins swimbladder
pale. (lateral view)
ies: Inhabits estuaries and
rivers.
Distribution: New Guinea
and northern Australia.
3156 Bony Fishes
Johnius (Johnieops) pacificus Hardenberg, 1941

FAO names: En - Pacific croaker.
Diagnostic characters: A small species with a moderately deep body. Eye

small, 16 to 23% of head length. Snout bluntly rounded; mouth horizontal,
slightly inferior; upper jaw extending backward below hind margin of eye; no
barbel on chin; upper rostral pores 5, marginal rostral pores 5; mental pores
in 3 pairs, the first open close behind symphysis connected by a crescent-
shaped groove; teeth differentiated into large and small in both jaws, the
large ones close-set, not canine-like, forming outer series in upper jaw, inner
series in lower jaw. Gill rakers slender, about 1/2 length of gill filaments at
angle of arch, 10 to 13 on lower limb of first gill arch. Dorsal fin with X
spines, followed by a deep notch, second part of fin with I spine and 25 to 29
soft rays; anal fin with II spines and 7 soft rays, second spine long, stiff, 45
to 54% of head length; caudal fin pointed. Scales on head and throat cycloid
(smooth), those on other parts of body ctenoid (rough to touch); small scales
present on soft parts of dorsal and anal fins; lateral-line scales reaching to
tip of caudal fin. Swimbladder hammer-shaped, with about 12 pairs of
arborescent appendages along its sides, the first pair entering head and
sending a palmate branch to the front of pectoral arch. Sagitta (large
earstone) with a tadpole-shaped impression, the head of which has its
long axis lying obliquely to that of sagitta and the tail expanded and
deepened as a hollow cone connected with the head by a narrow groove. swimbladder
Colour: body silvery, with faint dark oblique streaks on back; upper half of
spinous dorsal fin dusky, pectoral and pelvic fins pale.
Habitat, biology, and fisheries: Inhabits estuaries and rivers.
Distribution: Northern coasts
of New Guinea.
Johnius (Johnieops) plagiostoma (Bleeker, 1850)

FAO names: En - Large-eye croaker.
Diagnostic characters: A small species with a moderately deep body. Eye large,
about 30% of head length. Snout rounded, but not swollen or projecting; mouth
large, strongly oblique; upper jaw extending backward below hind margin of eye;
chin angular, prominent, without barbel; upper rostral pores 3, marginal rostral
pores 5; mental pores in 3 pairs, the first open at front of chin, separated by
symphysis; teeth well differentiated into large and small in both jaws, but
none canine-like; the large ones widely spaced, forming outer series in upper jaw,
inner series in lower jaw. Gill rakers slender, equal to or slightly shorter than
gill filaments at angle of arch, 14 to 17 on lower limb of first gill arch. Dorsal
fin with IX to XI (most frequently X) spines, followed by a notch, second part of fin
with I spine and 28 to 31 soft rays; anal fin with II spines and 6 to 8 (most
frequently 7) soft rays, second spine long, stiff, 40 to 50% of head length;
caudal fin rhomboidal. Scales on head cycloid (smooth), those on body ctenoid
(rough to touch); small scales present on soft parts of dorsal and anal fins;
lateral-line scales reaching to tip of caudal fin. Swimbladder hammer-shaped,
with about 13 pairs of arborescent appendages along its sides, the first pair
entering head beyond transverse septum and sending a palmate branch to swimbladder
the front of pectoral arch. Sagitta (large earstone) with a tadpole-shaped
impression, the head of which has its long axis lying obliquely to that of sagitta and the tail
expanded and deepened as a hollow cone connected with the head by a narrow groove. Colour:
bluish or greenish grey, flanks and belly silvery; fins yellowish.
to Borneo and Java, north to
Viet Nam.
3158 Bony Fishes
Larimichthys pamoides (Munro, 1964)

Frequent synonyms / misidentifications: Pseudotosciaena pamoides Munro, 1964 / None.
FAO names: En - Southern yellow croaker.
dorsal limb
Diagnostic characters: A small species with a moderately elongate body.
Snout rounded, but not swollen or projecting; mouth strongly oblique; upper
jaw extending backward below hind margin of pupil; chin without barbel;
upper rostral pore median one only, marginal rostral pores 5; mental pores
in 3 pairs, the first open at front of chin, separated by symphysis; teeth
differentiated into large and small in both jaws, the large ones widely spaced,
not canine-like, forming outer series in upper jaw, inner series in lower jaw.
Gill rakers long, slender, about 1.5 times longer than gill filaments at
angle of arch, 13 to 17 on lower limb of first arch. Dorsal fin with IX
spines, followed by a notch, second part of fin with I spine and 33 to 36
soft rays; anal fin with II spines and 7 soft rays, second spine short,
slender, 23 to 33% of head length; pectoral fins long, 25 to 27% of
standard length; caudal fin pointed. Scales on head and body cycloid
(smooth) except on belly and under side of caudal peduncle where ventral limb
weakly ctenoid scales cover (slightly rough to touch); small scales
present on a basal quarter of soft parts of dorsal and anal fins; lateral-line swimbladder appendages
scales reaching to tip of caudal fin. Swimbladder carrot-shaped, with about
26 pairs of arborescent appendages along its sides, each appendage
with a well-developed dorsal and ventral limb, the first pair entering head
beyond transverse septum. Colour (in preservative): body light brown;
inside of mouth, gill cavity, and body cavity dark brown.
Size: Known only from 2 specimens, 8.2 and 13.5 cm standard length.
Habitat, biology, and fisheries: Inhabits shallow coastal waters and estuaries to depths of 50 m.
northwestern Australia and
southern Papua New Guinea.
Nibea leptolepis (Ogilby, 1918)

FAO names: En - Smallscale croaker.
(after Ogilby, 1918)
septum

an arched back and deep body. Snout pointed, but not swollen
or projecting; mouth large, slightly oblique; upper jaw extending
backward below hind margin of pupil, upper jaw length 37 to
44% of head length, lower jaw length 47 to 52% of head
length; no barbel on chin; upper rostral pores 3, marginal rostral
pores 5; mental pores in 3 pairs, the first close together,
united by a crescent-shaped groove just behind symphysis;
teeth differentiated into large and small in both jaws, the
large ones widely spaced, not canine-like, forming outer series
in upper jaw, inner series in lower jaw. Gill rakers slender, about
2/3 length of gill filaments at angle of arch, 9 to 11 on lower limb
of first gill arch. Dorsal fin with X spines, followed by a notch,
second part of fin with I spine and 28 to 31 soft rays; anal fin
with II spines and 7 or 8 (most frequently 7) soft rays; second
spine long, stiff, 42 to 53% of head length; caudal fin rhom- sagitta swimbladder
boidal. Scales in lateral line 48 to 50, 14 to 17 scale rows above
lateral line to origin of dorsal fin, 17 to 23 scale rows below lateral line to origin of anal fin (total 31
to 39 scale rows); scales cycloid (smooth) on head and throat, ctenoid (rough to touch) on other parts of
body; small scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to tip of
caudal fin. Swimbladder carrot-shaped, with about 22 pairs of arborescent appendages along its
sides, the first pair entering head beyond transverse septum. Sagitta (large earstone) with a
tadpole-shaped impression, the tail of which J-shaped. Colour: silvery, flanks and belly deeply washed
with gold; spinose dorsal dark edged.
17 cm.
northern Australia.
3160 Bony Fishes
Nibea microgenys Sasaki, 1992

FAO names: En - Smalljaw croaker.
Diagnostic characters: A small to medium-sized species with an arched back

and deep body. Snout pointed, projecting in front of upper jaw; mouth
small, inferior, slightly oblique; upper jaw extending backward below hind
margin of pupil, upper jaw length 32 to 38% of head length, lower jaw length
34 to 40% of head length; no barbel on chin; upper rostral pores 3, marginal
rostral pores 5; mental pores in 3 pairs, the first close together, united by
a crescent-shaped groove just behind symphysis; teeth differentiated into
large and small in upper jaw only; large teeth in upper jaw closely spaced,
not canine-like, forming outer series; a band of villiform teeth in lower jaw.
Gill rakers slender, about 2/3 length of gill filaments at angle of arch, 5 to 8 on
lower limb of first gill arch. Dorsal fin with X spines, followed by a notch, second
part of fin with I spine and 29 to 31 soft rays; anal fin with II spines and 7 soft sagitta
rays; second spine long, stiff, 44 to 57% of head length; caudal fin rhomboi-
dal. Scales in lateral line 48 to 50, 9 to 14 scale rows above lateral line to origin of dorsal fin, 11 to 17 scale
rows below lateral line to origin of anal fin (total 22 to 29 scale rows); scales cycloid (smooth) on snout,
cheek, and throat, ctenoid (rough to touch) on other parts of head and body; small scales present on soft
parts of dorsal and anal fins; lateral-line scales reaching to tip of caudal fin. Swimbladder carrot-shaped,
with 19 to 21 pairs of arborescent appendages along its sides, the first pair entering head beyond
transverse septum. Sagitta (large earstone) with a tadpole-shaped impression, the tail of which
J-shaped. Colour (in preservative): brownish grey on back and flanks, whitish below; gill cover grey due
to blackish gill cavity.
monly to 19 cm.
northwestern and northern
Australia.
Nibea semifasciata Chu, Lo, and Wu, 1963

FAO names: En - Sharpnose croaker.
septum
Diagnostic characters: A small to medium-sized species with an arched back and
deep body. Snout acutely pointed, projecting in front of upper jaw; mouth large,
slightly inferior; upper jaw extending backward below hind margin of eye, upper
jaw length 41 to 43% of head length, lower jaw length 46 to 49% of head length;
no barbel on chin; upper rostral pores 3, marginal rostral pores 5; mental pores in
3 pairs, the first close together, united by a crescent-shaped groove just
behind symphysis; teeth differentiated into large and small in both jaws, the
large ones widely spaced, not canine-like, forming outer series in upper jaw, inner
series in lower jaw. Gill rakers stiff, about 1/2 length of gill filaments at angle of arch,
9 or 10 on lower limb of first gill arch. Dorsal fin with X spines, followed by a deep
notch, second part of fin with I spine and 26 to 29 soft rays; anal fin with II spines
and 7 soft rays, second spine stiff, 34 to 44% of head length; caudal fin
rhomboidal. Scales in lateral line 50 to 52, 11 or 12 scale rows above lateral line to
origin of dorsal fin, 15 to 17 scale rows below lateral line to origin of anal fin (total
26 to 29 scale rows); scales on head cycloid (smooth), those on body ctenoid (rough
to touch); small scales present on soft parts of dorsal and anal fins; lateral-line
scales reaching to tip of caudal fin. Swimbladder carrot-shaped, with 17 to 20
pairs of aborescent appendages along its sides, the first pair entering head
beyond transverse septum. Sagitta (large earstone) with a tadpole-shaped
impression, the tail of which J-shaped. Colour: body brownish grey, dark oblique swimbladder
lines on anterior part of back; margin of spinous dorsal fin darker.
Habitat, biology, and fisheries: Inhabits coastal waters.
Distribution: East China Sea
and Bangkok.
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3162 Bony Fishes
Nibea soldado (Lacepède, 1802)

Frequent synonyms / misidentifications: None / Nibea leptolepis (Ogilby, 1918).
FAO names: En - Soldier croaker; Fr - Courbine sodat; Sp - Corvina soldado.
septum
Diagnostic characters: A fairly large species with an arched
back and deep body. Snout rounded, but not swollen or
projecting; mouth large, terminal; upper jaw extending back-
ward below hind margin of eye, upper jaw length 38 to 44% of
head length, lower jaw length 44 to 53% of head length; no
barbel on chin; upper rostral pores 3, marginal rostral pores 5;
mental pores in 3 pairs, the first close together, united by a
crescent-shaped groove just behind symphysis; teeth dif-
ferentiated into large and small in both jaws, the large ones
widely spaced, not canine-like, forming outer series in upper jaw,
inner series in lower jaw. Gill rakers slender, equal to or slightly
shorter than gill filaments at angle of arch, 7 to 12 on lower limb
of first gill arch. Dorsal fin with IX to X (most frequently X) spines,
followed by a notch, second part of fin with I spines and 27 to 33
soft rays; anal fin with II spines and 7 soft rays, second spine
long, stiff, 39 to 59% of head length; caudal fin rhomboidal. sagitta
Scales in lateral line 48 to 51, 8 to 13 scale rows above lateral
swimbladder
line to origin of dorsal fin, 12 to 17 scale rows below lateral
line to origin of anal fin (total 21 to 29 scale rows); scales on head cycloid (smooth), those on body
ctenoid (rough to touch); small scales present on soft parts of dorsal and anal fins; lateral-line scales
reaching to tip of caudal fin; swimbladder carrot-shaped, with 18 to 22 pairs of aborescent appendages
along its sides, the first pair entering head beyond transverse septum. Sagitta (large earstone) with
a tadpole-shaped impression, the tail of which J-shaped. Colour: silvery with faint series of oblique
stripes along scale rows; margin of soft part of dorsal fin dark, pectoral and pelvic fins with yellow tinge.
cific, west to India, east to
Queensland coasts of Austra-
lia.
Nibea squamosa Sasaki, 1992

Frequent synonyms / misidentifications: None / Nibea semifasciata Chu, Lo, and Wu, 1963.
FAO names: En - Scale croaker.
Diagnostic characters: A fairly large species with a moderately elongate body.

Snout blunt, obtusely rounded, slightly projecting in front of upper jaw;
mouth small, inferior; upper jaw extending backward below hind margin of
pupil, upper jaw length 31 to 38% of head length, lower jaw length 34 to 40%
of head length; no barbel on chin; upper rostral pores 5, marginal rostral pores
5; mental pores in 3 pairs, the first close together, united by a crescent-
shaped groove just behind symphysis; teeth differentiated into large and
small in upper jaw only; the large ones closely spaced, not canine-like,
forming outer series; a band of villiform teeth on lower jaw. Gill rakers stiff,
about 1/4 length of gill filaments at angle of arch, 6 to 9 on lower limb of first gill
arch. Dorsal fin with X spines, followed by a notch, second part of fin with I spine
and 27 to 31 soft rays; bony lumps generally present on dorsal and caudal fins sagitta
in specimens longer than 30 cm standard length; anal fin with II spines and 7
soft rays, second spine long, stiff, 38 to 64% of head length, progressively thicker with age and size;
caudal fin rhomboidal. Lateral-line scales numerous, 57 to 60, scale rows above lateral line to origin of
dorsal fin 9 to 13, scale rows below lateral line to origin of anal fin 12 to 18 (total 22 to 30 scale rows);
scales on snout, cheek, and throat cycloid (smooth), those on other parts of head and body ctenoid (rough
to touch); small scales present on soft parts of dorsal and anal fins; lateral-line scales reaching to tip
of caudal fin. Swimbladder carrot-shaped, with about 18 pairs of aborescent appendages along its
sides, the first pair entering head beyond transverse septum. Sagitta with a tadpole-shaped
impression, the tail of which J-shaped. Colour (in preservative): uniformly brownish yellow, slightly paler
ventrally; a series of oblique dark spots along scale rows on anterior part of body below spinous dorsal fin.
Habitat, biology, and fisheries: Shallow coastal waters, estuaries, and rivers; known also from fresh
water up to about 500 km away from sea.
Distribution: N e w G u i n e a
and northern Australia.
3164 Bony Fishes
Otolithes ruber (Bloch and Schneider, 1801)

Frequent synonyms / misidentifications: Otolithes argenteus (Cuvier, 1830) / None.
FAO names: En - Tigertooth croaker; Fr - Grande verrue tigre; Sp - Bombache tigre mayor.
Diagnostic characters: Medium-sized to large species

with a slender, cylindrical body. Snout not swollen or
projecting; its dorsal profile rising evenly to origin of
dorsal fin; mouth large, strongly oblique; upper jaw extend-
ing backward below hind margin of eye; chin without barbel;
upper rostral pores absent, marginal rostral pores 3,
head
mental pores indistinct; teeth differentiated into large and
small in both jaws, with 1 or 2 pairs of strong canines at
front of one or both jaws. Gill rakers slender, about equal
to or 2/3 length of gill filaments at angle of arch, 8 to 11 on
lower limb of first gill arch. Dorsal fin with IX to X spines
(most frequently IX in the western Indian Ocean, X in the
Bay of Bengal and West Pacific), followed by a notch,
second part of fin with I spine and 26 to 30 soft rays; anal
fin with II spines and 7 soft rays, second spine short,
slender, 14 to 23% of head length; caudal fin rhomboidal.
Scales cycloid (smooth), but a few ctenoid (rough to sagitta swimbladder
touch) on lower part of caudal peduncle; lateral-line
scales reaching to tip of caudal fin. Swimbladder carrot-shaped with 32 to 36 pairs of fan-like
appendages along sides, each appendage lodged beside bladder and none widely lapping dorsal
surface of bladder wall, the first pair not entering head. Sagitta (large earstone) with a tadpole-
shaped impression, the tail of which only slightly curved, ending in a disc near the posterior edge.
Colour: brownish above, silvery with a golden sheen on flanks and belly, often with oblique dark streaks
dorsally; pectoral, pelvic, and anal fins reddish or light brown.
waters down to 40 m. In India,
the majority are mature at 20 cm
total length, all are ripe at 26 cm
total length and above. Feeds
mainly on fishes and prawns.
Distribution: Widespread along
coasts of the Indian Ocean and
West Pacific, west to South Af-
rica, east to southern China and
Queensland, Australia.
Otolithoides biauritus (Cantor, 1850)

Frequent synonyms / misidentifications: Otolithoides brunneus (Day, 1873) / None.
FAO names: En - Bronze croaker; Fr - Verrue bronzée; Sp - Bombache bronzeado.
Diagnostic characters: A large species with a slender, cy-

lindrical body. Eye small, 13 to 15% of head length. Snout
pointed, but not projecting in front of upper jaw; mouth large,
oblique; upper jaw extending backward well beyond hind mar-
gin of eye; chin without barbel; upper rostral pores 3, marginal
rostral pores 5; mental pores in 2 pairs, the first small, at
front of chin, separated by symphysis; teeth differentiated
into large and small in both jaws; the large ones widely spaced,
not canine-like, forming outer series in upper jaw, inner series
in lower jaw. Gill rakers slender, about 2/3 length of gill fila-
ments at angle of arch, 10 to 12 on lower limb of first gill arch.
Dorsal fin with IIX to IX (most frequently IX) spines,
followed by a notch, second part of fin with I spine and 27 to
32 soft rays; anal fin with II spines and 6 or 7 (most frequently
7) soft rays, second spine short, slender, 16 to 20% of head
length; caudal fin pointed. Scales cycloid (smooth) on head
and back, elsewhere ctenoid (rough to touch); lateral-line sagitta swimbladder
scales reaching to tip of caudal fin. Swimbladder carrot-
shaped, with a single pair of tube-like appendages arising from posterior end of bladder and running
forward beside main body of bladder and in front of it into head beyond transverse septum, where
they branch under skull. Sagitta (large earstone) with a tadpole-shaped impression, the tail of which
only slightly curved and ending in a disc. Colour: head and back green or grey, flanks gold or orange,
paler toward belly, minutely dotted with brown. Lateral line golden yellow; dorsal, anal, and caudal fins
brown or yellow to pale orange, pelvic fins pale orange, pectoral fins brown with black spot at base.
Habitat, biology, and fisheries: Inhabits coastal and inshore waters.
Distribution: India to Malay
Peninsula, Sumatra, Borneo,
and Viet Nam.
3166 Bony Fishes
Panna microdon (Bleeker, 1849)

FAO names: En - Panna croaker; Fr - Courbine ragane; Sp - Corvina ragana.
Diagnostic characters: A medium-sized species, with a slen-

der, cylindrical body. Eye small, 13 to 20% of head length.
Snout pointed, but not swollen or projecting in front of upper jaw;
mouth large, terminal; upper jaw extending backward below hind
margin of eye; chin without barbel; upper rostral pores 3, marginal
rostral pores 5; mental pores in 3 pairs, the first small, at front
of chin, separated by symphysis; teeth differentiated into large
and small in both jaws; the large ones widely spaced, not canine-
like, forming outer series in upper jaw, inner series in lower jaw.
Gill rakers slender, 2/3 to equal length of gill filaments at angle of
arch, 9 to 11 on lower limb of first gill arch. Dorsal fin with IX
to X (most frequently X) spines and 33 to 36 soft rays, the fin
not incised between the 2 parts; anal fin with II spines and 6 or sagitta
7 (most frequently 7) soft rays, second spine short, slender, 16
to 35% of head length; caudal fin pointed. Scales on head and throat cycloid
(smooth), those on body finely ctenoid (rough to touch); lateral-line scales, with
a coating of small secondary scales, reaching to tip of caudal fin; scales absent
on soft parts of dorsal and anal fins. Swimbladder carrot-shaped, with 1 pair of
tube-like appendages arising at anterior end and immediately dividing into an
anterior branch entering head, and ramifying under skull and a posterior tube
lying beside the main bladder to its posterior end. Sagitta (large earstone) with
a tadpole-shaped impression, the tail of which broad and deep with a sharp
bend between shorter proximal part and a longer proximal part. Colour: greyish swimbladder
above, becoming silvery on flanks and belly; pectoral and pelvic fins yellow; dorsal
and anal fins dusky with dark margin.
waters and estuaries; young
and juveniles occur in man-
grove swamps. Caught with
bottom-trawls, gill nets, and
handlines. Marketed fresh;
also dried salted.
Distribution: Thailand, Ma-
laysia, Viet Nam, and Indone-
sia.
Panna perarmatus (Chabanaud, 1926)

Frequent synonyms / misidentifications: Otolithoides perarmatus (Chabanaud, 1926) / None.
FAO names: En - Armour croaker.
adult (47 cm standard length)
Diagnostic characters: A
large species, with an elon-
gate, cylindrical body. Eye
small, 14 to 24% of head
length. Snout very steeply
rounded, but not projecting
in front of upper jaw; mouth
large, oblique; upper jaw ex- young specimen (16 cm standard length)
tending backward below (after Sasaki, 1995)
posterior margin of eye; chin
without barbel; upper rostral pore median one only, marginal rostral pores 5; mental
pores in 2 pairs, the first small, rounded, at front of chin, separated by symphysis;
teeth differentiated into large and small in both jaws, the large ones widely spaced, not
canine-like, forming outer series in upper jaw, inner series in lower jaw. Gill rakers
long, slender, equal to or slightly longer than gill filaments at angle of arch, 12
to 14 on lower limb of first gill arch. Dorsal fin with VI to VII spines and 42 to 44
soft rays, not incised between the 2 parts; anal fin with II spines and 7 or 8 soft
rays, second spine long, stiff, its length 37 to 46% of head length; the last dorsal,
second anal, and pelvic-fin spines progressively thicker with age or size; caudal
fin rhomboidal. Scales cycloid (smooth) on head, ctenoid (rough to touch) on body;
lateral-line scales reaching to tip of caudal fin; small scales present on soft parts of
dorsal and anal fins. Swimbladder carrot-shaped, with 1 pair of tube-like append-
ages arising at anterior end and immediately dividing into an anterior branch
entering head and ramifying under skull, and a short posterior tube. Colour: body
brownish or dark greenish; opercular membrane and fins blackish.
Size: Maximum standard length about 50 cm, commonly to 30 cm. swimbladder
Distribution: Thailand, Viet
Nam, and Borneo.
3168 Bony Fishes
Pennahia anea (Bloch, 1793)

Frequent synonyms / misidentifications: Pennahia macrophthalmus (Bleeker, 1850) / None.
FAO names: En - Donkey croaker.
mental pores
Diagnostic characters: A fairly small, moderately deep-

bodied species. Snout pointed, but not swollen or project-
ing; mouth large, oblique; upper jaw extending backward
below hind margin of eye; chin angular, without barbel;
upper rostral pores absent or minute, marginal rostral
pores 5; mental pores in 2 pairs, both small, the first
pair at front of chin, separated by symphysis; teeth well
differentiated into large and small in both jaws, but none
canine-like, the large ones widely spaced, forming outer ventral view
series in upper jaw, inner series in lower jaw. Gill rakers of head
slender, as long as gill filaments at angle of arch, 9 to
12 on lower limb of first gill arch. Dorsal fin with IX to
X (most frequently X) spines, followed by a notch, second
part of fin with I spine and 22 to 24 soft rays; anal fin with
II spines and 7 soft rays, second spine short, slender,
22 to 36% of head length; caudal fin truncate. Scales
cycloid (smooth) on head and flanks, elsewhere ctenoid
(rough to touch); lateral-line scales reaching to hind margin
of caudal fin. Swimbladder carrot-shaped, with 17 to 22 swimbladder
pairs of fan-like appendages along its sides, the first not sagitta
entering head. Sagitta (large earstone) with a tadpole-
shaped impression, the tail of which only slightly curved. Colour: head and body greyish light brown
on back, becoming paler ventrally with increasing silvery reflection to whitish abdomen; gill cover with
diffused dark blotch; upper two-thirds of spinous dorsal fin dusky.
Habitat, biology, and fisheries: Inhabits coastal waters down to 60 m. Caught with bottom trawls and
seine nets. Marketed fresh and dried salted.
east to Indonesia.
Pennahia macrocephalus (Tang, 1937)

Frequent synonyms / misidentifications: Argyrosomus macrocephalus (Tang, 1937) / None.
FAO names: En - Bighead pennah croaker.
Diagnostic characters: A fairly small, deep-bodied species. Snout pointed,

but not projecting or swollen; mouth large, oblique; upper jaw extending
backward below middle to hind margin of eye; chin angular, without barbel;
upper rostral pores 3, marginal rostral pores 5; mental pores in 3 pairs, the
first small, at front of chin, separated by symphysis; teeth well differenti-
ated into large and small in both jaws, but none canine-like; the large ones
widely spaced, forming outer series in upper jaw, inner series in lower jaw. Gill
rakers slender, as long as gill filaments at angle of arch, 10 to 13 on lower
limb of first gill arch. Dorsal fin with X spines, followed by a deep notch,
second part of fin with I spine and 26 to 29 soft rays; anal fin with II spines
and 7 soft rays, second spine moderately long, stiff, 28 to 39% of head
length; caudal fin rhomboidal. Scales on head and anterior part of body
cycloid (smooth), ctenoid (rough) posteriorly; lateral-line scales reaching to
tip of caudal fin. Swimbladder carrot-shaped, with about 18 pairs of fan-like
appendages along its sides, none entering head. Sagitta (large earstone)
with a tadpole-shaped impression, the tail of which only slightly curved.
Colour: counter-shaded from metallic blue above to white below; lining of
mouth and gill cover dark grey; a black spot at tip of lower jaw and one above
base of pectoral fins; pectoral, pelvic, and anal fins yellowish. swimbladder
Habitat, biology, and fisheries: Inhabits coastal water on continental shelf down to 100 m.
Distribution: East and South
China seas and Java, Indone-
sia. Records from the Bay of
Bengal are based on misiden-
tifaction of Pennahia ovata
Sasaki, 1996.
3170 Bony Fishes
Pennahia pawak (Lin, 1940)

Frequent synonyms / misidentifications: Argyrosomus pawak Lin, 1940 / None.
FAO names: En - Pawak croaker.
Diagnostic characters: A small to medium-sized, fairly deep-bodied species.

Snout pointed, but not swollen or projecting; mouth large, slightly oblique; upper
jaw extending backward below middle of eye; chin rounded, without barbel;
upper rostral pores 3, marginal rostral pores 5; mental pores in 3 pairs, the
first small, rounded, at front of chin, separated by symphysis; teeth differ-
entiated into large and small in both jaws, but none canine-like; the large ones
widely spaced, forming outer series in upper jaw, inner series in lower jaw. Gill
rakers slender, 2/3 to equal length of gill filaments at angle of arch, 7 to 9 on
lower limb of first gill arch. Dorsal fin with X spines, followed by a notch, second
part of fin with I spine and 23 to 25 soft rays; anal fin with II spines and 7 soft
rays, second spine short, slender, its length 17 to 31% of head length;
caudal fin rhomboidal. Scales on head cycloid (smooth), those on body
ctenoid (rough to touch); lateral-line scales reaching to tip of caudal fin. Swim-
bladder carrot-shaped, with 24 to 27 pairs of fan-like appendages along its
sides, none entering head. Sagitta (large earstone) with a tadpole-shaped
impression, the tail of which only slightly curved. Colour: body bronze,
greenish brown above; a large, characteristic blotch on spinous dorsal fin;
black lining of gill cover shows through conspicuously; a pale band between swimbladder
basal and distal dark zones in soft dorsal fin; caudal fin dark at margin.
Habitat, biology, and fisheries: Inhabits coastal waters on continental shelves.
Distribution: South China
Sea and southern Indonesia.
Protonibea diacantha (Lacepède, 1802)

Frequent synonyms / misidentifications: Nibea diacanthus (Lacepède, 1802) / None.
FAO names: En - Spotted croaker; Fr - Courbine pintade; Sp - Corvina pintade.
Diagnostic characters: A large species with a slender body.

Eye small, 16 to 21% of head length. Snout acutely pointed, but
not swollen or projecting; mouth large, slightly oblique; upper
jaw extending backward below hind margin of eye; chin without
barbel; upper rostral pores 3, marginal rostral pores 5; mental
pores in 3 pairs, the first small, close together, united by a
crescent-shaped groove just behind symphysis; teeth dif-
ferentiated into large and small in both jaws, but none canine-
like; the large ones well spaced, forming outer series in upper
jaw, inner series in lower jaw. Gill rakers stiff, about 1/2 length
of gill filaments at angle of arch, 7 or 8 on lower limb of first
arch. Dorsal fin with IX or X (most frequently X) spines,
followed by a notch, second part of fin with I spine and 22 to 25
soft rays; anal fin with II spines and 7 soft rays; second
spine short, slender, 27 to 33% of head length; caudal fin sagitta swimbladder
rhomboidal. Scales cycloid (smooth) on snout and below eye, ctenoid (rough to touch) elsewhere;
lateral-line scales reaching to tip of caudal fin. Swimbladder carrot-shaped, with 16 to 20 pairs of
arborescent appendages along its sides, none entering head. Sagitta (large earstone) with a
tadpole-shaped impression, the tail of which bents sharply at about an angle of 90°, very narrowly
separated from the ventral edge. Colour: 3 to 5 dark bars along back and many small black spots
(about size of pupil) on top of head, upper half of body and caudal fin; pectoral, pelvic, and anal
fins and lower part of caudal fin black; in larger fishes, bars and spots obscure or absent.
Habitat, biology, and fisheries: Inhabits coastal waters down to 60 m. Also ascends tidal rivers and
estuaries. In India, matures at around 85 cm standard length (3 or 4 years). Feeds mainly on prawns and
fishes. Caught with bottom
trawls and gill nets. Marketed
fresh and dried-salted.
east to northern Australia and
southern China, and Taiwan
Province of China.
3172 Bony Fishes
Pterotolithus lateoides (Bleeker, 1850)

Frequent synonyms / misidentifications: Otolithes lateoides Bleeker, 1850 / None.
FAO names: En - Bigmouth croaker.
Diagnostic characters: A fairly large species, with a slender, cylindrical

body. Eye small, 16 to 19% of head length. Snout pointed, its upper
profile rising evenly and almost horizontally to origin of dorsal fin; mouth
large, strongly oblique; upper jaw extending backward below hind margin
of pupil; chin prominent, without barbel; upper rostral pores absent,
marginal rostral pores 5; mental pores inconspicuous; teeth in a single
series in both jaws, with 1 or 2 pairs of strong canines at tip of upper
jaw and 1 pair at tip of lower jaw. Gill rakers slender, about 2/3 length of
gill filaments at angle of arch, 8 to 12 on lower limb of first gill arch. Dorsal
fin with X spines, followed by a notch, second part of fin with I spine and portion
24 to 27 soft rays; anal fin with II spines and 7 or 8 soft rays, second branching
spine short, slender, 23 to 31% of head length; caudal fin pointed. on top of
bladder
Scales cycloid (smooth) on head, elsewhere ctenoid (rough to touch);
lateral-line scales reaching to tip of caudal fin. Swimbladder carrot-
shaped, with about 40 pairs of arborescent appendages, some of the
dorsal limbs long and lie on the dorsolateral surface of the bladder,
none entering head. Sagitta (large earstone) with a tadpole-shaped
impression, the tail of which J-shaped. Colour: greyish above, silvery
flanks and belly; fins yellow, brown spots between soft dorsal-fin rays. swimbladder appendage
Habitat, biology, and fisheries: Inhabits coastal waters and estuaries. Caught with bottom trawls, gill
nets, and handlines. Marketed fresh; also dried-salted; swimbladder dried.
Distribution: Malay Penin-
sula, Java, and Borneo.
Pterotolithus maculatus (Cuvier, 1830)

Frequent synonyms / misidentifications: Otolithes maculatus Cuvier, 1830 / None.
FAO names: En - Blotched tigertoothed croaker.
Diagnostic characters: A fairly large species with a slender, cylindrical

body. Eye small, 21 to 22% of head length. Snout pointed, its upper
profile rising evenly and almost horizontally to origin of dorsal fin;
mouth large, strongly oblique; upper jaw extending backward below
middle of eye; chin prominent, without barbel; upper rostral pores portion
branching
absent, marginal rostral pores 5, mental pores inconspicuous; teeth on top of
differentiated into large and small in both jaws, a pair of strong canines bladder
at front of both jaws. Gill rakers slender, about 2/3 length of gill
filaments at angle of arch, 8 to 11 on lower limb of first gill arch. Dorsal
fin with IX spines, followed by a notch, second part of fin with I spine
and 30 to 34 soft rays; anal fin with II spines and 10 to 12 soft rays,
second spine short, slender, 23 to 30% of head length; caudal fin
rhomboidal. Scales on head and body cycloid (smooth); lateral-line
scales extending to tip of caudal fin. Swimbladder carrot-shaped, with
numerous (50 or more) pairs of branched appendages, the stems
of the ventral limbs forming a parallel series of tubes, those of the
dorsal limbs roofing the swimbladder with filigree of silvery
branched tubules, none entering head. Sagitta (large earstone)
with a tadpole-shaped impression, the tail of which J-shaped.
Colour: upper part of body marked with numerous black blotches,
about 1/2 size of eye or larger; fins yellow. swimbladder appendage
Habitat, biology, and fisheries: Inhabits shallow coastal waters and estuaries. Caught with bottom trawls,
gill nets, and handlines. Marketed fresh; also dried-salted; swimbladder dried.
Distribution: Bay of Bengal to
Borneo.
3174 Bony Fishes
Sonorolux fluminis Trewavas, 1977

FAO names: En - Estuary croaker.
(after Trewavas, 1977)
Diagnostic characters: A small species with a deep,

rounded body. Eye moderately large, 22 to 23% of
head length. Snout steeply rounded, but not inflated or
projecting; mouth large, oblique, upper jaw extending
backward slightly beyond hind margin of eye; lower jaw
upturned, chin prominent, without barbel; upper rostral
pores absent, marginal rostral pore median one
only; 2 or 3 pairs of mental pores, the first small,
rounded, at front of chin, separated by symphysis;
teeth differentiated into large and small in both jaws, but
none canine-like; the large ones widely spaced, forming
outer series in upper jaw, inner series in lower jaw. Gill
rakers long, slender, about 1.5 times as long as gill
filaments at angle of arch, 16 or 17 on lower limb of
first gill arch. Dorsal fin with X or XI spines and 28
to 29 soft rays, the fin not deeply incised into 2 sagitta
parts; anal fin with II spines and 7 or 8 soft rays, the swimbladder
second spine moderately long, slender, 33 to 37% (right side)
of head length; caudal fin rhomboidal. Scales on head
and anterior part of body cycloid (smooth), elsewhere ctenoid (rough to touch); lateral-line scales
reaching to tip of caudal fin; small scales present on soft parts of dorsal fins; many of ventral scales
covering patches of yellow, luminous tissue. Swimbladder carrot-shaped, with about 20 pairs of
arborescent appendages along its side, each with a well-developed dorsal and ventral limb, the first
pair entering head beyond transverse septum. Sagitta (large earstone) with a tadpole-shaped
impression, the tail of which nearly straight, only slightly curved, ending in a rounded disc. Colour:
unknown.
Size: K n o w n o n l y f r om 2
specimens, both about 8 cm
standard length.
ies: Inhabits estuaries.
Distribution: K n o w n f r o m
Sarawak, Borneo.
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Perciformes: Percoidei: Mullidae 3175
MULLIDAE
Goatfishes (surmullets)
by J.E. Randall
D iagnostic characters: Body moderately elongate and somewhat compressed (size to 50 cm). Two
long unbranched barbels on chin; mouth low on head, the lower jaw inferior, the cleft slightly oblique;
dentition variable but teeth conical, either in villiform bands or in 1 or 2 rows, never as enlarged canines
(except in adult males of western Atlantic and eastern Pacific species of Pseudupeneus, the teeth of which
are slightly enlarged). A single flat spine posteriorly on opercle (a second less developed spine may be
present); margin of preopercle smooth. Two well-separated dorsal fins, the first with VII or VIII
(usually VIII) slender spines (first spine often very small), the second fin with 9 soft rays (first
unbranched); anal fin with I spine and 6 or 7 soft rays; caudal fin deeply forked, with 13 branched
rays; pelvic fins with I spine and 5 soft rays; pectoral fins with 13 to 18 rays. Scales finely ctenoid; head
and body completely scaly (except preorbital region of some species of Upeneus). Lateral line complete,
following contour of back, the pored scales to base of caudal fin 27 to 38. Colour: ground colour in
preservative usually pale, in life often whitish to light red; most species with distinctive black, brown, red,
or yellow markings; median fins often with stripes or oblique bands.
2 dorsal fins, 1st with VII-VIII
spines, 2nd with 9 soft rays
2 barbels on chin
Habitat, biology, and fisheries: Most goatfishes inhabit shallow seas. They are usually found on open
sand or mud bottoms, at least for feeding (though the species of Parupeneus and Mulloidichthys are often
seen on coral reefs or rocky substrata). The goatfish barbels, with their chemosensory receptors, are
actively moved over or into the sediment to find food organisms. Once the prey is located, the fish often
roots into the sediment with its snout for the prey. Mullid fishes are carnivorous; they feed on a wide variety
of small animals, especially crustaceans and various worms. A few species feed in part on small fishes.
The flesh of goatfishes is of good quality and that of some species highly esteemed in certain countries.
Most of the catch is marketed fresh. Catch statistics are usually not recorded by individual species. For
1995, the FAO Yearbook of Fishery Statistics reports a total catch of around 44 250 t of Mullidae from the
Western Central Pacific.
IV-VI graduated
dorsal-fin spines
Polymixiidae: the only other family with a pair of long
barbels on chin; the species occur in deep water
(about 200 to 400 m). They and are distinguished
from the Mullidae by the following combination of
characters: a single, unnotched dorsal fin with IV to
VI spines; anal fin with III or IV spines; barbels
inserted well behind tip of lower jaw.
barbels inserted well III-IV anal-fin spines
behind tip of lower jaw Polymixiidae
3176 Bony Fishes
Key to the genera of Mullidae occurring in the area teeth on

vomer
1a. No teeth on vomer or palatines . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Teeth present on vomer and palatines (Fig. 1) or only on vomer . . . . . ® 3
2a. Teeth in jaws small, in 2 to several rows; lateral-line scales 33
to 39; 5 scale rows between dorsal fins . . . . . . . . . . . Mulloidichthys
2b. Teeth in jaws in 1 row, moderately large, bluntly conical, and
well spaced; lateral-line scales 27 to 29; 2 or 3 scale rows
between dorsal fins . . . . . . . . . . . . . . . . . . . . . . . Parupeneus teeth on
palatine
3a. Teeth on vomer, but none on palatines; no scales basally on
second dorsal and anal fins; lateral-line scales 26 to 28 . . . Upeneichthys Fig. 1
3b. Teeth on vomer and palatines; small scales basally on second
dorsal and anal fins; lateral-line scales 28 to 38 . . . . . . . . . . Upeneus
Key to the species of Mulloidichthys occurring in the area

1a. Peritoneum transparent to silvery white; head length 3.2 to 3.6 times in standard length;
colour, including fins, red, without any dark or light markings, shading to pale red or whitish
ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mulloidichthys pflugeri
(Hawaii and Society Islands to Ryukyu and Mascarene islands - all localities insular)
1b. Peritoneum dark brown to black; head length 2.95 to 3.45 times in standard length; body
with 1 or more blue or yellow stripes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Lateral-line scales 38 to 39; body depth 3.1 to 3.5 times in standard length; colour yellow,
including fins, the body with 5 narrow blue stripes, the upper 4 continuing anteriorly onto
head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mulloidichthys mimicus
(Marquesas Islands and Line Islands)
2b. Lateral-line scales 33 to 36; body depth 3.4 to 4.8 times in standard length; colour whitish
with a yellow stripe on upper side . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. First gill arch with 18 to 22 gill rakers on lower limb; body depth 3.7 to 4.8 times in standard
length; snout length 1.9 to 2.2 times in head length; barbel length 1.4 to 1.65 times in head
length; pectoral-fin length 1.45 to 1.7 times in head length; a blackish spot usually present on
yellow stripe of body below first dorsal fin; fins whitish to pale yellowish . . . Mulloidichthys flavolineatus
(Indo-Pacific)
3b. First gill arch with 23 to 26 gill rakers on lower limb (for West Pacific specimens; 20 to 25 in
Indian Ocean); body depth 3.4 to 4.5 times in standard length; snout length 2 to 2.45 times in
head length; barbel length 1.2 to 1.5 times in head length; pectoral-fin length 1.35 to 1.55 times
in head length; no black spot in yellow stripe on body; fins yellow . . . . . . Mulloidichthys vanicolensis
(Indo-Pacific)
Key to the species of Parupeneus occurring in the area
1a. A dark brown to dark reddish stripe on upper side of body ending anteriorly on caudal
peduncle; a dark spot larger than eye on side of caudal peduncle . . . . . . . . . . . . . . . . . . ® 2
1b. No dark stripe on upper side of body; no dark spot on side of peduncle, or if present, not
preceded by a dark stripe on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Pectoral-fin rays 16 to 18 (usually 17); last dorsal- and anal-fin ray equal to or slightly
longer than penultimate ray; barbels equal to or shorter than snout plus eye; peduncular
dark spot round, centred on lateral line at base of caudal fin; no dark band at base of
second dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus barberinus
(East Africa to French Polynesia)
2b. Pectoral-fin rays 15 to 17 (usually 16, rarely 17); last dorsal- and anal-fin ray distinctly
longer than penultimate ray; barbels very long, much longer than snout plus eye; pedun-
cular dark spot not round, anterior to base of caudal fin, and more above than below lateral
line; a blackish band at base of second dorsal fin, continuing to distal end of last ray . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus macronemus
(Red Sea and coast of East Africa to Indonesia)
3a. Posterior end of maxilla evenly convex (Fig. 2a) . . . . . ® 4

3b. Posterior end of maxilla oblique, the dorsoposte-
rior corner more acute (Fig. 2b) . . . . . . . . . . . . . . ® 5
evenly oblique
4a. Body moderately elongate, its depth 3.6 to 4.4 convex
times in standard length; pectoral-fin rays 15 or
16 (rarely 16); colour pink with a yellow stripe
a) b)
following lateral line anteriorly and passing
above and adjacent to lateral line posteriorly; no Fig. 2
dark spot on side of body . . . . . . . . Parupeneus jansenii
(Seychelles and India to Indonesia and Philippines)
4b. Body less elongate, its depth 3 to 3.55 times in standard length; pectoral-fin rays 15 to 17
(usually 16, rarely 15); a dark reddish spot about the size of pupil usually present just
below lateral line above outer third of pectoral fin; edges of scales dull orange-red on about
upper three-fourths of body, white below, the scales of upper 4 rows with a pale bluish
spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus heptacanthus
(Red Sea south to South Africa, east to western Pacific from the Ryukyu Islands to Lord Howe Island and New
Caledonia)
5a. Two or more dark bars extending ventrally from dorsal margin of body; first gill arch with
27 to 33 gill rakers on lower limb . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5b. At most 1 dark bar extending ventrally from dorsal margin of body (then only on caudal
peduncle); first gill arch with 18 to 26 gill rakers on lower limb (except 28 to 30 in P. moffitti) . . . . . ® 7
6a. A broad dark bar below each dorsal fin, a faint third bar sometimes present on caudal
peduncle, and a large dark area on head enclosing eye; barbels short, 1.65 to 1.9 times
in head length; body depth 2.5 to 3.4 times in standard length; dorsal profile of snout
distinctly concave; last 2 rays of second dorsal and anal fins subequal . . . . Parupeneus bifasciatus
(Indo-Pacific)
6b. Four or 5 dark bars on body, only the last 2 (below second dorsal fin and on caudal
peduncle) heavily pigmented, the one below interdorsal space narrowest; barbels long,
1.1 to 1.35 times in head length; body depth 3.15 to 3.65 times in standard length; dorsal
profile of snout straight to slightly concave; last ray of second dorsal and anal fins clearly
longer than penultimate ray . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus multifasciatus
(Central and western Pacific)
7a. First gill arch with 28 to 30 gill rakers on lower limb; colour red with a lighter red stripe on
back below dorsal fins, bordered above and below by a row of indistinct small dark brown
spots . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus moffitti
(Guam, from depths of 120 to 230 m)
7b. First gill arch with 18 to 26 gill rakers on lower limb; colour not as above . . . . . . . . . . . . . . ® 8
8a. First gill arch with 18 to 21 gill rakers on lower limb; a round black spot as large or larger
than eye posteriorly on side of peduncle, more above than below lateral line (but not
reaching dorsal edge of peduncle); a large horizontally-elliptical yellow spot centred on
lateral line above tip of pectoral fin . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus indicus
(East Africa to Samoa and the Caroline Islands)
8b. First gill arch with 21 to 26 (rarely 21) gill rakers on lower limb; colour not as above (a black
spot, if present on caudal peduncle, small or saddle-like) . . . . . . . . . . . . . . . . . . . . . . ® 9
3178 Bony Fishes
9a. Barbels very long, their length from longer than head to 1.2 times in head length; snout
long, 1.6 to 1.9 times in head length; 2 colour phases, one bluish grey with a saddle-like
yellow spot on caudal peduncle, the other yellow with a brighter yellow spot dorsally on
peduncle; attains 50 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus cyclostomus
(Indo-Pacific)
9b. Barbels not very long, their length 1.3 to 1.9 times in head length; snout not as long, 1.7
to 2.15 times in head length; colour not as above; maximum length 25 to 38 cm . . . . . . . . . ® 10
10a. Pectoral-fin rays 15 to 17 (usually 16, rarely 15); last dorsal- and anal-fin ray of adults
distinctly longer than penultimate ray of these fins; body depth 3.45 to 3.95 times in
standard length; colour whitish to pink with a black spot about 4 scales in width on lateral
line below rear of first dorsal fin, followed by a large oval white spot; base of second dorsal
fin broadly blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus pleurostigma
(Indo-Pacific)
10b. Pectoral-fin rays 14 to 16 (usually 15); last dorsal- and anal-fin ray subequal to penultimate
ray of these fins; body depth 2.95 to 3.55 times in standard length; colour not as above . . . . . ® 11
11a. Head and anterior half of body dark reddish brown, abruptly white and yellow on posterior
half; 2 diagonal white bands in dark part of head and body, the uppermost running from
front of snout, through upper part of eye and along back; an irregular black spot a little
smaller than eye on lateral line just behind rear base of second dorsal fin . . Parupeneus barberinoides
(western Pacific to Samoa and islands of Micronesia)
11b. Body not divided into very dark anterior and abruptly pale posterior zones, but 2 diagonal
whitish bands present from snout through eye (1 through upper part, the other through
lower), parallel with dorsal contour of body, ending below second dorsal fin . . . . . . . . . . . ® 12
12a. Barbels short, 1.55 to 1.9 times in head length; first gill arch with 24 to 27 gill rakers on
lower limb; colour light red, the scale edges reddish brown to dark brown except ventrally;
a slight intensification of dark pigment to form a blotch dorsally on caudal peduncle,
preceded by a narrow pale blotch . . . . . . . . . . . . . . . . . . . . . . . . . . Parupeneus ciliatus
(Society Islands and Rapa to New South Wales, north to southern Japan and west to Seychelles and Réunion)
12b. Barbels not short, 1.3 to 1.6 times in head length; first gill arch with 21 to 24 gill rakers on
lower limb; colour similar, with or without a black spot dorsally on caudal peduncle . . . . . . . . ® 13
13a. A black spot as large or larger than eye dorsally on caudal peduncle, its lower edge resting
on lateral line, its upper edge near top of peduncle, usually not meeting spot of other side
(or if it does, the intervening pigment is generally less intense) . . . . . . . . . . Parupeneus spilurus
(southern Japan to New South Wales, New Caledonia, and northern New Zealand; also Western Australia south to
Rottnest Island)
13b. No black spot on caudal peduncle (a faint dusky spot may be present dorsoposteriorly); 3
brown and 2 whitish stripes on upper half of body distinct, the lowermost brown stripe
midlateral from opercle at level of lower part of eye . . . . . . . . . . . . . . Parupeneus biaculeatus
(southern Japan to Viet Nam)
Key to the species of Upeneus occurring in the area

1a. First dorsal fin with VII spines, the first longest; pectoral-fin rays 13 or 14 (usually 13); total
gill rakers on first gill arch 27 to 29; upper lobe of caudal fin with 3 narrow oblique dark
bands, the lower lobe with 6 or 7; a midlateral dark stripe; a dark brown saddle-like spot
on caudal peduncle just posterior to second dorsal fin . . . . . . . . . . . . . Upeneus asymmetricus
(Philippines to Indonesia and northwestern Australia)
1b. First dorsal fin with VIII spines, the first minute (may be adherent to second spine);
pectoral-fin rays 13 to 17; total gill rakers on first gill arch 19 to 30; colour not entirely as
above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Second dorsal-fin spine very prolonged, extending when depressed beyond base of
second dorsal fin; pectoral-fin rays 13; light red, shading through pink on sides to white
ventrally; cheek and operculum suffused with yellow; barbels yellow . . . . . . . . . . Upeneus filifer
(southern Queensland and Chesterfield Islands)
2b. Second dorsal-fin spine about equal to or slightly longer than third; pectoral-fin rays 13 to
17; colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Second dorsal-fin spine long, 1.1 to 1.3 times in head length; total gill rakers on first gill
arch 19 to 22; lateral-line scales 31 or 32; a narrow brown to brownish yellow stripe on
body at level of upper part of eye containing 3 darker zones, 1 under each dorsal fin and
1 below anterior caudal peduncle; upper lobe of caudal fin with 3 or 4 dark cross bands
(may be faint), the lower lobe with a broad dusky posterior border, with or without faint
dark cross bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Upeneus sundaicus
(Philippines to northern Australia and Persian Gulf)
3b. Second dorsal-fin spine not long, 1.3 to 1.65 times in head length; total gill rakers on first
gill arch 20 to 30; lateral-line scales 29 to 38; colour not entirely as above . . . . . . . . . . . . . ® 4
4a. Lobes of caudal fin without dark cross bands; bronze dorsally, silvery on sides with 2
narrow brassy yellow stripes; first dorsal fin with 2 broad dusky yellowish stripes separated
by narrow white stripes, the tip broadly black; lateral-line scales usually 34 or 35; total gill
rakers on first gill arch 26 to 30 . . . . . . . . . . . . . . . . . . . . . . . . . . . Upeneus sulphureus
(Red Sea south to Mozambique and Madagascar, east to Fiji)
4b. One or both lobes of caudal fin with dark cross bands; colour not as above; lateral-line
scales 28 to 38; total gill rakers on first gill arch 20 to 30 . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Dorsal lobe of caudal fin with 6 or 7 dusky orange oblique bands, the lower lobe without
dark cross bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5b. Both lobes of caudal fin with dark cross bands . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
6a. A golden yellow stripe as wide as pupil from behind eye to upper base of caudal fin; back
above stripe iridescent silvery pink, below silvery; total gill rakers on first gill arch 26 to 30
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Upeneus moluccensis
(Red Sea to Madagascar east to the western Pacific from southern Japan to northern Australia)
6b. Silvery with 3 close-set iridescent pink stripes on back and 2 narrow brassy yellow stripes
on side; total gill rakers on first gill arch 25 to 29 . . . . . . . . . . . . . . . Upeneus quadrilineatus
(southern Japan to Indonesia)
7a. Pectoral-fin rays 15 to 17 (usually 16); total gill rakers on first gill arch 25 to 30; peritoneum
blackish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7b. Pectoral-fin rays 12 to 15 (usually 13 or 14); total gill rakers on first gill arch 21 to 25;
peritoneum whitish . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
8a. Silvery with 4 distinct narrow brassy yellow stripes on upper two-thirds of body; lower lobe
of caudal fin with 3 dark cross bands (not counting tip, sometimes black) the distal band
nearly twice as broad and more intensely black than middle band; first dorsal fin broadly
tipped with black; total gill rakers on first gill arch 27 to 31 . . . . . . . . . . . . . . . Upeneus vittatus
(Indo-Pacific)
8b. Silvery without distinct yellow stripes on body; lower lobe of caudal fin with 4 dark cross
bands of about equal width; tip of first dorsal fin narrowly black; total gill rakers on first gill
arch 25 to 29 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Upeneus subvittatus
(southern Japan to Indonesia and Red Sea to Mozambique)
3180 Bony Fishes
9a. No scales on side of snout; lateral-line scales 35 to 37; silvery, the scales dorsally on body
edged with green or orange-pink; 2 narrow orange-yellow stripes on side of body; upper
lobe of caudal fin of adults with 6 to 8 and the lower lobe with 4 to 6 dusky orange to black
cross bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Upeneus arge
(central and western Pacific to Seychelles and Mozambique)
9b. Scales on side of snout; lateral-line scales 28 to 30; a dark reddish brown to blackish stripe
from behind eye to base of caudal fin; body above stripe brownish to greenish grey,
flecked with small dark reddish brown to blackish spots; upper lobe of caudal fin with 4 to
6 blackish cross bands, and lower lobe with 5 to 7; outer third to half of first dorsal fin with
an irregular dark red to black spot containing 2 to several small yellow spots . . . . . Upeneus tragula
(Indo-Pacific)

0 Mulloidichthys flavolineatus (Lacepède, 1801)
0 Mulloidichthys pflugeri (Steindachner, 1900)
Mulloidichthys mimicus Randall and Guézé, 1980
0 Mulloidichthys vanicolensis (Valenciennes, 1831)

0 Parupeneus barberinoides (Lacepède, 1801)
0 Parupeneus barberinus (Bleeker, 1852)
0 Parupeneus bifasciatus (Lacepède, 1801)
Parupeneus biaculeatus (Richardson, 1846)
0 Parupeneus ciliatus (Lacepède, 1802)

0 Parupeneus cyclostomus (Lacepède, 1801)
0 Parupeneus heptacanthus (Lacepède, 1802)
0 Parupeneus indicus (Shaw, 1803)
0 Parupeneus macronemus (Lacepède, 1801)
Parupeneus jansenii (Bleeker, 1856)
0 Parupeneus multifasciatus (Quoy and Gaimard, 1824)

Parupeneus moffitti Randall and Myers, 1993
0 Parupeneus pleurostigma (Bennett, 1831)

0 Parupeneus spilurus (Bleeker, 1854)
Upeneichthys lineatus (Bloch and Schneider, 1801)
0 Upeneus arge (Jordan and Evermann, 1902)
Upeneus asymmetricus Lachner, 1954
0
Upeneus filifer (Ogilby, 1910)
Upeneus moluccensis (Bleeker, 1855)
Upeneus quadrilineatus Cheng and Wang, 1968
0
Upeneus subvittatus (Temminck and Schlegel, 1843)
0
Upeneus sulphureus Cuvier, 1829
0
Upeneus sundaicus (Bleeker, 1855)
0
Upeneus tragula (Richardson, 1846)
Upeneus vittatus (Forsskål, 1775)
References
Masuda, H., K. Amaoka, C. Araga, T. Uyeno, and T. Yoshino (eds). 1984. The fishes of the Japanese Archipelago.
Tokyo, Tokai University Press, 456 p.
Randall, J.E. and P. Guézé. 1992. Upeneus francisi, a new goatfish (Perciformes: Mullidae) from Norfolk Island and
New Zealand. Cybium, 16(1):21-29.
Randall, J.E., G.R. Allen, and R.C. Steene. 1990. Fishes of the Great Barrier Reef and Coral Sea, 506 p.
Stepien, C.A., J.E. Randall, and R.H. Rosenblatt. 1994. Genetic and morphological divergence of a circumtropical
complex of goatfishes: Mulloidichthys vanicolensis, M. dentatus, and M. martinicus. Pac. Sci., 48(1):44-56.
Mulloidichthys flavolineatus (Lacepède, 1801) (Plate XXV, 186)

Frequent synonyms / misidentifications: Mulloidichthys samoensis (Günther, 1802) / None.
FAO names: En - Yellowstripe goatfish; Fr - Capuciva à bande jaune; Sp - Salmonete de banda amarilla.
Diagnostic characters: Body elongate, its depth 3.7 to 4.8 times in standard length, and moderately
compressed; head length 3.1 to 3.45 times in standard length. Chin with a pair of slender barbels which
usually do not reach posterior margin of preopercle, their length 1.4 to 1.65 times in head length;
snout long, its length in adults 1.9 to 2.2 (to 2.4 in young) times in head length; mouth small, maxilla not
reaching front edge of eye; villiform teeth in a band in jaws; no teeth on roof of mouth. First gill arch with
8 or 9 (rarely 7 or 10) gill rakers on upper limb, 18 to 22 on lower limb (total 25 to 30). Two
well-separated dorsal fins, the first with VIII slender spines (initial spine very small), and the second with
9 soft rays; anal fin with I spine and 7 soft rays; pectoral-fin rays 16 to 18 (very rarely 15 or 19). Lateral-line
scales 33 to 36; 5 rows of scales between dorsal fins, and 11 or 12 rows along upper part of caudal
peduncle. Colour: silvery white with a yellow stripe on side of body at level of eye, usually containing a
blackish spot above posterior part of pectoral fins; barbels white; fins whitish to pale yellowish; peritoneum
blackish.
Size: Maximum reported total length 40 cm, commonly to 28 cm.
Habitat, biology, and fisheries: A shallow-water species usually found in lagoons and bays. Tends to
occur in aggregations by day over coral reefs or at times resting on sand bottom. Although observed to
feed by day, most of the feeding appears to commence with the onset of darkness (then as individual fish).
Feeds on a wide variety of small invertebrates such as small crabs, shrimps, polychaetes, sipunculids,
small pelecypods and gastropods, hermit crabs, heart urchins, and foraminifera; occasionally captures
small fishes. Caught by nets, traps, hook-and-line, and by spearing. The young are taken by seines or cast
nets; they are consumed fresh, dried, or used as bait.
Distribution: Occurs through-
out the Indo-Pacific region; in
the western Pacific from south-
ern Japan to Montague Island,
New South Wales (36°S); in
Oceania to Hawaii and Pitcairn
Group.
3182 Bony Fishes
Mulloidichthys pflugeri (Steindachner, 1900) (Plate XXV, 187)

FAO names: En - Red goatfish.
Diagnostic characters: Body moderately elongate, its depth 3.5 to 4.25 times in standard length, and
moderately compressed; head length 3.2 to 3.6 times in standard length. Chin with a pair of slender
barbels which usually do not reach rear margin of preopercle, their length 1.45 to 1.6 times in head
length; snout moderately long, its length in adults 1.85 to 2.2 times in head length; mouth small, maxilla
not reaching front edge of eye; villiform teeth in a band in jaws; no teeth on roof of mouth. First gill arch
with 7 or 8 gill rakers on upper limb, 19 to 22 on lower limb (total 27 to 30). Two well-separated dorsal
fins, the first with VIII slender spines (initial spine very small), and the second with 9 soft rays; anal fin with
I spine and 7 soft rays; pectoral-fin rays 17 or 18. Lateral-line scales 34 to 37; 5 rows of scales between
dorsal fins, and 11 or 12 rows along upper part of caudal peduncle. Colour: orange-red, shading to pale
orange-red or whitish ventrally; barbels white; fins light orange-red; peritoneum transparent to silvery
white.
Size: Largest specimen examined 48 cm, commonly to 35 cm.
Habitat, biology, and fisheries: Tends to occur deeper than other species of the genus (to at least 110 m),
but may be found in as little as 15 m. Usually encountered as solitary individuals or in pairs over sand flats
near coral reefs. When feeding, it may take on a strong pattern of 4 broad dark bars (darker red than rest
of body, not with black pigment). Not commonly seen in fish markets.
Distribution: Indo-Pacific, but
distribution spotty, and all
known localities insular: Ha-
waii, Society Islands, Marque-
sas, Tonga, Marshall Islands,
Mariana Islands, Ryukyu Is-
lands, Indonesia, Mauritius,
and Réunion.
Mulloidichthys vanicolensis (Valenciennes, 1831) (Plate XXV, 188)

Frequent synonyms / misidentifications: Mulloides vanicolensis (Valenciennes, 1831); M. auriflamma (non
Forsskål, 1775); M. erythrinus Klunzinger, 1884 / None.
FAO names: En - Yellowfin goatfish; Fr - Capucin de Vanicolo; Sp - Salmonete vanicolense.
moderately compressed; head length 2.95 to 3.4 times in standard length. Chin with a pair of slender
barbels which reach posterior to rear margin of preopercle, their length 1.2 to 1.5 times in head
length; snout not long, its length in adults 2 to 2.45 times in head length; mouth small, maxilla not reaching
front edge of eye; villiform teeth in a band in jaws; no teeth on roof of mouth. First gill arch with 8 to 10
gill rakers on upper limb, 23 to 26 on lower limb (total 32 to 36). Two well-separated dorsal fins, the
first with VIII slender spines (initial spine very small), and the second with 9 soft rays; anal fin with I spine
and 7 soft rays; pectoral-fin rays 16 to 18. Lateral-line scales 33 to 36; 5 rows of scales between dorsal
fins, and 11 or 12 rows along upper part of caudal peduncle. Colour: whitish to pink with a yellow stripe
edged in pale blue on body at level of upper end of gill opening; no black spot within yellow stripe;
barbels white to yellowish; fins yellow.
Size: Maximum total length, 38 cm, commonly to 25 cm.
Habitat, biology, and fisheries: A shallow-water species usually found in lagoons and bays, but generally
in deeper water than Mulloidichthys flavolineatus. Tends to occur in aggregations by day on coral reefs,
but disperses individually at night to feed on sand flats. Food habits similar to those of M. flavolineatus:
small crabs, shrimps, and other crustaceans, polychaetes and other worms, ophiuroids, small molluscs,
heart urchins, and foraminifera. Able to take on a disruptive pattern of broad irregular red bars, especially
at night.
Distribution: Widespread in the Indo-Pacific region; in the Indian Ocean from the Red Sea south to
southern Mozambique and Mauritius; in the western Pacific from Kyushu to central New South Wales; in
Oceania east to Hawaii, Tuamotu Archipelago, and Marquesas. The population in the Indian Ocean has
fewer gill rakers (29 to 34) from
that of the Pacific, and fish
there are able to change colour
to yellow with blue stripes,
hence a mimic of the snapper
Lutjanus kasmira (with which it
may school). Comparison of
the Indian Ocean and Pacific
populations of M. vanicolensis
by enzyme elecrophoresis
failed to show a genetic diver-
gence comparable to the gill-
raker difference.
3184 Bony Fishes
Parupeneus barberinoides (Bleeker, 1852) (Plate XXV, 189)

FAO names: En - Bicolour goatfish.
moderately compressed. Chin with 2 slender barbels usually reaching posterior to rear edge of
preopercle, their length 1.25 to 1.5 times in head length; head length 2.75 to 3.2 times in standard
length; snout moderate, its length 1.75 to 2 times in head length; dorsal profile of snout straight to
slightly concave; mouth small, maxilla reaching from 2/3 to 3/4 distance to anterior edge of eye; a single
row of well-spaced, stout, conical teeth in jaws; no teeth on roof of mouth. First gill arch with 6 or 7 gill
rakers on upper limb, 21 to 25 on lower limb (total 28 to 33). Two well-separated dorsal fins, the first
with VIII spines, the second fin with 9 soft rays; first dorsal-fin spine very short; longest dorsal-fin spine
1.4 to 1.8 times in head length; last ray of second dorsal and anal fins distinctly longer than penultimate
ray; anal fin with I spine and 6 soft rays; pectoral-fin rays 15 or 16 (usually 15). Lateral-line scales 27 or
28; 3 rows of scales between dorsal fins; 9 rows of scales along upper part of caudal peduncle. Colour:
head and anterior half of body dark reddish brown, abruptly white and yellow on posterior half; 2
diagonal white bands in dark part of head and body, the uppermost running from front of snout, through
upper part of eye and along back; an irregular black spot a little smaller than eye on lateral line just
behind rear base of second dorsal fin; barbels red.
Habitat, biology, and fisheries: Occurs on coral reefs and adjacent habitats, generally in protected areas
like bays or lagoons; juveniles often seen in seagrass beds. Adults are usually solitary, but juveniles
generally form small schools.
Distribution: W e s te rn P a-
cific from New South Wales to
southern Japan, east to Sa-
moa and the Marshall Islands.
Parupeneus barberinus (Lacepède, 1801) (Plate XXV, 190)

FAO names: En - Dash-and-dot goatfish; Fr - Rouget-barbet barberin; Sp - Salmonete.
moderately compressed. Chin with 2 slender barbels which usually reach posterior to rear margin of
preopercle (barbels equal to or longer than snout plus eye), their length 1.4 to 1.6 times in head
length; snout long, its length 1.5 to 2.1 times in head length (snout relatively longer with growth); dorsal
profile of snout straight; mouth small, maxilla reaching only about 1/2 distance to front edge of eye; a single
row of well-spaced, stout, conical teeth in jaws; no teeth on roof of mouth. First gill arch with 6 or 7 gill
rakers on upper limb, 20 to 25 on lower limb (total 26 to 32). Two well-separated dorsal fins, the first
with VIII spines, the second fin with 9 soft rays; first dorsal-fin spine very short; longest dorsal-fin spine
1.15 to 1.7 times in head length (relatively longer in larger fish; third and fourth spines prolonged in large
adults); last ray of second dorsal and anal fins equal to or a little longer than penultimate ray; anal fin with
I spine and 6 soft rays; pectoral-fin rays 16 to 18 (usually 17). Lateral-line scales 27 or 28; 3 rows of
scales between dorsal fins; 8 or 9 rows of scales along upper part of caudal peduncle. Colour: a black
stripe from snout through lower part of eye and along back to below rear base of second dorsal fin;
back above stripe light grey-brown to yellow, below whitish, the scale edges narrowly grey to reddish brown;
a round black spot, larger than eye in adults, posteriorly on midside of caudal peduncle; diagonal
pale blue lines alternating with brownish yellow extending dorsoposteriorly and ventroanteriorly from eye,
with 1 or 2 parallel blue lines on cheek. Fish taken from deeper water are more red than those from the
shallows; the dark stripe on body and spot on peduncle of the deep-dwelling fish may be red.
Size: Maximum reported total length about 50 cm, commonly to 30 cm.
Habitat, biology, and fisheries: This large goatfish is primarily a shallow-water species, but it is known
to reach a depth of 100 m. It is generally found in the vicinity of coral reefs, but feeds over sand bottoms,
sometimes excavating deeply into the sand. Stomach contents of 19 specimens examined by the present
author consisted of crabs (including portunids, xanthids, and raninids), shrimps, polychaetes, sipunculids,
pelecypods, small gastropods, isopods, brachiopods, and one had eaten a small eel.
out most of the Indo-Pacific re-
gion; unknown from the Red
Sea, Gulf of Oman, Persian
Gulf, and Hawaii. Ranges in
the western Pacific from south-
ern Japan to the southern
Great Barrier Reef.
3186 Bony Fishes
Parupeneus bifasciatus (Lacepède, 1801) (Plate XXV, 191 and 192)

Frequent synonyms / misidentifications: Parupeneus trifasciatus (Lacepède, 1801); P. crassilabris
(Valenciennes, 1831) / None.
FAO names: En - Doublebar goatfish; Fr - Rouget-barbet double-tauche; Sp - Salmonete de dos manchas.
Diagnostic characters: Body not elongate (for a mullid), its depth 2.65 to 3.4 times in standard length
(larger individuals deeper bodied), and moderately compressed. Chin with 2 slender barbels which
usually do not reach rear margin of preopercle, their length 1.65 to 1.9 times in head length; snout
not long, its length 1.7 to 2 times in head length; dorsal profile of snout concave; mouth small, maxilla
reaching at most a vertical through posterior nostril; a single row of well-spaced, stout, conical teeth in
jaws; no teeth on roof of mouth. First gill arch with 7 to 9 (usually 8) gill rakers on upper limb, 27 to
31 on lower limb (total 35). Two well-separated dorsal fins, the first with VIII spines, the second fin with
9 soft rays; first dorsal-fin spine very short; longest dorsal-fin spine 1.45 to 1.6 times in head length; last
2 rays of second dorsal and anal fins subequal; anal fin with I spine and 6 soft rays; pectoral-fin rays 15
to 17 (usually 16). Lateral-line scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of scales
along upper part of caudal peduncle. Colour: yellowish grey to brownish red, the centres of scales paler
than edges, shading to whitish ventrally, with a broad saddle-like dark brown bar centred below anterior
part of first dorsal fin, a second broad bar below second dorsal fin, and often a third faint bar dorsally on
caudal peduncle (bars reaching slightly below midside of body, those of juveniles more strongly marked
and those of large adults may be obscure); a large dark brown area on side of head, enclosing eye, and
extending onto side of snout (obscure in large adults); irregular blue lines and spots usually present on
side of head (including within dark area); second dorsal and anal fins often with oblique yellow and purple
or blue lines. Individuals from deeper water more red than those from the shallows.
Habitat, biology, and fisheries: A coral-reef species known from a depth range of a few to at least 60 m.
Analysis of stomach contents of 17 specimens revealed feeding mainly on crabs (44% of contents),
shrimps, octopuses, mantis shrimps, amphipods, other crustaceans, fishes, and polychaetes.
Distribution: Gulf of Aden and Gulf of Oman south to Natal, east throughout Oceania, including Hawaii
and Pitcairn Group. In the western Pacific from the Ryukyu Islands to the Great Barrier Reef. Differences
in colour, particularly with re-
spect to the dark bars, may be
seen among individuals from
the western Indian Ocean,
western Pacific, and Hawaii, in-
dicating population differentia-
tion. This merits further study.
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Parupeneus ciliatus (Lacepède, 1802) (Plate XXV, 193)

Frequent synonyms / misidentifications: Parupeneus fraterculus (Valenciennes, 1831); P. pleurotaenia
(Playfair and Günther, 1867) / Parupeneus porphyreus (non Jenkins, 1903); P. rubescens (non Lacepède,
1801).
FAO names: En - Diamondscaled goatfish.
Diagnostic characters: Body slightly elongate, its depth 2.95 to 3.4 times in standard length, and
moderately compressed. Chin with 2 slender barbels which do not reach rear margin of preopercle,
their length 1.55 to 1.9 times in head length; snout short, its length 1.85 to 2.1 times in head length;
dorsal profile of snout straight to slightly concave; mouth small, maxilla reaching a vertical through posterior
nostril; a single row of well-spaced, stout, conical teeth in jaws; no teeth on roof of mouth. First gill arch
with 6 to 8 gill rakers on upper limb, 24 to 27 (rarely 27) on lower limb (total 31 to 34). Two
well-separated dorsal fins, the first with VIII spines, the second fin with 9 soft rays; first dorsal-fin spine
very short; longest dorsal-fin spine 1.6 to 1.9 times in head length; last 2 rays of second dorsal and anal
fins subequal; anal fin with I spine and 6 soft rays; pectoral-fin rays 15 (rarely 14). Lateral-line scales 27
or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of scales along upper part of caudal peduncle.
Colour: light red, the scale edges reddish brown to dark brown except ventrally; a slight intensification of
dark pigment to form a blotch dorsally on caudal peduncle, preceded by a narrow pale blotch; 2 parallel
whitish to pale red bands from side of snout through eye (1 across upper part, 1 along lower edge) along
back to end beneath front of second dorsal fin; fins light red without markings except small pale spots on
dorsal and anal fins of some individuals.
Size: Largest specimen examined 38 cm total length, commonly to 28 cm.
Habitat, biology, and fisheries: A coral-reef species of shallow to moderate depths of lagoons and outer
reef zones; may also be seen in seagrass beds. Often observed at rest on reef substratum, suggesting
that it feeds nocturnally.
Distribution: Indo-Pacific ex-
cept the Red Sea to Persian
Gulf and Hawaii; in the west-
ern Pacific from southern Ja-
pan to New South Wales and
Lord Howe Island; east in
Oceania to the Marshall Is-
lands and French Polynesia
(including Rapa).
3188 Bony Fishes
Parupeneus cyclostomus (Lacepède, 1801) (Plate XXVI, 194 and 195)

Frequent synonyms / misidentifications: Parupeneus chryserydros (Lacepède, 1801); P. luteus
(Valenciennes, 1831) / None.
FAO names: En - Goldsaddle goatfish; Fr - Rouget-barbet doré; Sp - Salmonete dorado.
moderately compressed. Chin with 2 very long slender barbels, reaching to or slightly posterior to
end of opercle, their length 0.95 to 1.2 times in head length. Snout long, its length 1.6 to 1.9 times
in head length; dorsal profile of snout straight to slightly concave; mouth small, maxilla reaching little more
than 1/2 distance to anterior edge of eye; a single row of well-spaced, stout, conical teeth in jaws; no teeth
on roof of mouth. First gill arch with 6 or 7 gill rakers on upper limb, 22 to 26 on lower limb (total 29
to 33). Two well-separated dorsal fins, the first with VIII spines, the second fin with 9 soft rays; first dorsal-fin
spine very short; longest dorsal-fin spine 1.45 to 1.75 times in head length; last ray of second dorsal and
anal fins slightly longer than penultimate ray; anal fin with I spine and 6 soft rays; pectoral-fin rays 15 to
17 (usually 16). Lateral-line scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of scales
along upper part of caudal peduncle. Colour: 2 colour phases, the first yellowish grey with blue markings
on the scales and a broad saddle-like yellow spot dorsally on caudal peduncle; irregular blue lines radiating
from eye and below eye; second dorsal and anal fins with oblique blue lines; caudal fin with blue lines
paralleling rays; other phase entirely yellow with a brighter yellow saddle-like spot dorsally on caudal
peduncle.
Habitat, biology, and fisheries: This large goatfish is more closely associated with coral reefs than other
species. It is unusual for a mullid in its heavy feeding on fishes (about 70% of the diet). It has been observed
to use its barbels to frighten its prey from holes in the reef. Like other goatfishes while feeding, it is often
accompanied by other fishes such as jacks and wrasses which try to capture prey animals fleeing from its
intrusions in the reef or sand.
the Indo-Pacific from the Red
Sea to Natal and east to the
islands of Oceania, including
Hawaii (which lacks the yellow
p h a s e ) , an d th e P i tc a i r n
Group. Ranges in the western
Pacific from the Ryukyu Is-
lands to central New South
Wales.
Parupeneus heptacanthus (Lacepède, 1802) (Plate XXVI, 196)

Frequent synonyms / misidentifications: Parupeneus cinnabarinus (Cuvier, 1801); P. pleurospilos
(Bleeker, 1853) / None.
FAO names: En - Cinnabar goatfish; Fr - Barbet à point rouge.
Diagnostic characters: Body slightly elongate, its depth 3 to 3.55 times in standard length, and moderately
compressed. Chin with 2 moderately long slender barbels, reaching beyond rear edge of preopercle,
their length 1.15 to 1.35 times in head length; snout moderate, its length 1.75 to 2.1 times in head
length; dorsal profile of head convex; mouth small, maxilla nearly or just reaching a vertical at posterior
nostril; posterior end of maxilla evenly convex (other species of the genus except Parupeneus jansenii
with a dorsoposterior lobe on maxilla); a single row of well-spaced, stout, conical teeth in jaws; no teeth
to 30). Two well-separated dorsal fins, the first with VIII spines, the second fin with 9 soft rays; first
dorsal-fin spine very short; longest dorsal-fin spine 1.45 to 1.75 times in head length; last ray of second
dorsal and anal fins slightly longer than penultimate ray; anal fin with I spine and 6 soft rays; pectoral-fin
rays 15 to 17 (usually 16). Lateral-line scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows
of scales along upper part of caudal peduncle. Colour: brownish yellow to light red (deeper-dwelling fish
more red), the edges of the scales darker, shading to silvery white below; centres of scales of back with a
pale blue to pearly spot (more evident on some individuals than others); a small reddish brown spot on
upper side of body of adults just below seventh and eighth lateral-line scales; faint iridescent blue
lines extending dorsoposteriorly and ventroanteriorly from eye, and often a parallel one on cheek and snout
below eye; second dorsal and anal fins with faint pale blue to pink lines, alternating with yellowish.
Size: Largest of 184 specimens examined, 37 cm total length; commonly to 25 cm.
Habitat, biology, and fisheries: Associated more with somewhat turbid water and silty sand or weedy
bottoms than other species of the genus; not often seen on coral reefs; usually found at depths greater
than 15 m. Occurs singly or in
small groups.
Distribution: Red Sea and
coast of East Africa to 34°S,
east to Samoa and the Mar-
shall Islands; in the western
Pacific from southern Japan to
southern Queensland and
Lord Howe Island.
3190 Bony Fishes
Parupeneus indicus (Shaw, 1803) (Plate XXVI, 197)

FAO names: En - Indian goatfish; Fr - Rouget-barbet indien; Sp - Salmonete índico.
moderately compressed. Chin with 2 slender barbels which reach to or slightly beyond rear edge of
preopercle, their length 1.3 to 1.5 times in head length; snout moderate, its length 1.65 to 1.95 times
in head length; dorsal profile of snout straight, becoming convex anterior to eye; mouth small, maxilla
reaching about 1/2 distance to front edge of eye; a single row of well-spaced, stout, conical teeth in jaws;
no teeth on roof of mouth. First gill arch with 5 to 7 gill rakers on upper limb, 18 to 21 on lower limb
(total 24 to 27). Two well-separated dorsal fins, the first with VIII spines, the second fin with 9 soft rays;
first dorsal-fin spine very short; longest dorsal-fin spine 1.5 to 1.8 times in head length; last ray of second
dorsal and anal fin longer than penultimate ray; anal fin with I spine and 6 soft rays; pectoral-fin rays
15 to 17 (usually 16). Lateral-line scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of
scales along upper part of caudal peduncle. Colour: greenish brown to reddish brown dorsally, the scale
edges narrowly dark, shading to whitish or pale pink ventrally, with a roundish black spot larger then
eye posteriorly on side of caudal peduncle, 2/3 of which lies above lateral line; a large, horizontally-
elongate, yellow spot (sometimes partly white) centred on lateral line below posterior part of first
dorsal fin and interdorsal space; irregular pale blue lines radiating from eye; irregular oblique pale blue
lines on second dorsal and anal fins; caudal fin with pale blue lines parallel with rays.
Habitat, biology, and fisheries: A coastal species generally found on sand, silty sand or seagrass
substrata, generally in the vicinity of rocky bottom or reefs; feeds mainly on crabs, shrimps, amphipods,
small octopuses, polychaetes, and small fishes. Caught in trawls, fish traps, and by hook-and-line. Highly
esteemed as a food fish.
Distribution: Southern Ara-
bian Peninsula south to Port
Alfred, South Africa, east to
Samoa and the Caroline Is-
lands; in the western Pacific
from southern Japan to the
southern Great Barrier Reef.
Parupeneus macronemus (Lacepède, 1801) (Plate XXVI, 198)

FAO names: En - Longbarbel goatfish; Fr - Rouget-barbé badeau; Sp - Salmonete bandón.
moderately compressed. Chin with 2 very long slender barbels which reach or nearly reach rear edge
of opercle, their length 1.1 to 1.25 times in head length; snout moderate, its length 1.7 to 1.9 times
in head length; dorsal profile of snout straight; mouth small, maxilla not reaching a vertical at posterior
nostril; a single row of well-spaced, stout, conical teeth in jaws; no teeth on roof of mouth. First gill arch
with 7 to 9 gill rakers on upper limb, 27 to 30 on lower limb (total 34 to 39). Two well-separated dorsal
fins, the first with VIII spines, the second fin with 9 soft rays; first dorsal-fin spine very short; longest
dorsal-fin spine 1.45 to 1.75 times in head length; last ray of second dorsal and anal fins much longer
than penultimate ray; anal fin with I spine and 6 soft rays; pectoral-fin rays 15 to 17 (usually 16).
Lateral-line scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of scales along upper part
of caudal peduncle. Colour: reddish brown, paler posteriorly and ventrally (sometimes pale magenta or
lavender), with a dark brown band from front of snout, through eye, continuing as a black stripe
across upper end of gill opening, and ending beneath rear base of second dorsal fin; a roundish
black spot (usually larger than eye) on posterior third of caudal peduncle, the lateral line passing
through its lower third; a vertical pale blue line or spot on scales of body below dark stripe, and light blue
lines on margin of dark band on head; basal third of second dorsal fin blackish, with a narrow extension
to posterior tip of fin; lateral edge of pelvic fins broadly blackish.
Habitat, biology, and fisheries: An inshore species generally found on sand and weedy substrata; rarely
taken in depths of more than 40 m. Occurs as individuals or in small aggregations. Feeds like other
goatfishes by using its long barbels to probe into the sediment to find prey which consists mainly of
crustaceans and worms.
Distribution: Ranges from
the Red Sea and Persian Gulf
south to Natal and east to In-
donesia and the Philippines.
?
3192 Bony Fishes
Parupeneus multifasciatus (Quoy and Gaimard, 1824) (Plate XXVI, 199 and 200)
Frequent synonyms / misidentifications: None / Parupeneus trifasciatus (non Lacepède, 1801).
FAO names: En - Manybar goatfish; Fr - Rouget à trois bandes.
moderately compressed. Chin with 2 moderately long slender barbels which reach posterior to rear
edge of preopercle and exceptionally to posterior edge of opercle, their length 1.1 to 1.35 times in
head length; snout moderately long, its length 1.7 to 1.85 times in head length; dorsal profile of snout
straight to slightly concave; mouth small, maxilla reaching little more than 1/2 distance to anterior edge of
eye; a single row of well-spaced, stout, conical teeth in jaws; no teeth on roof of mouth. First gill arch with
7 to 9 gill rakers on upper limb, 29 to 32 on lower limb (total 36 to 41). Two well-separated dorsal fins,
the first with VIII spines, the second fin with 9 soft rays; first dorsal-fin spine very short; longest dorsal-fin
spine 1.45 to 1.85 times in head length; last ray of second dorsal and anal fins much longer than
penultimate ray; anal fin with I spine and 6 soft rays; pectoral-fin rays 15 to 17 (usually 16). Lateral-line
scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of scales along upper part of caudal
peduncle. Colour: light grey to light brownish red with a broad blackish bar below second dorsal fin,
an oblique elliptical black bar nearly crossing caudal peduncle, a narrow dusky bar below interdor-
sal space, a diffuse broad faint bar usually present below first dorsal fin, and another sometimes
evident on nape; area on side between bars paler than rest of body; a dusky band from front of snout
through eye, more darkly pigmented on postorbital head; basal half of second dorsal fin blackish,
broadening posteriorly to distal part of fin; anal fin and outer half of second dorsal fin yellow with dark-edged
blue (sometimes pink) lines; caudal fin with blue lines paralleling rays.
Habitat, biology, and fisheries: The most common goatfish found on coral reefs and adjacent sand and
rubble in the central and western Pacific. Although usually encountered inshore, it is reported to depths as
great as 140 m. Solitary and diurnal, feeding on crabs (about 40% of diet), shrimps (30%), other
crustaceans, small fishes, octopuses, pelecypods, gastropods, and foraminifera. Will also feed on the
larger animals of the zooplankton, such as crab megalops, when abundant. Females may be sexually
mature as small as 18 cm. Because of its relatively small size, of less commercial value than the larger
goatfishes.
out the islands of Oceania and
the western Pacific, including
Hawaii and the Pitcairn Group.
Not known from the Indian
Ocean except Christmas Island
and Cocos-Keeling Islands.
Ranges in the western Pacific
from southern Japan to central
New South Wales.
Parupeneus pleurostigma (Bennett, 1831) (PLate XXVII, 201)

FAO names: En - Sidespot goatfish; Fr - Rouget-barbet pastille; Sp - Salmonete pastilla.
Diagnostic characters: Body elongate, its depth 3.45 to 3.95 times in standard length, and moderately
compressed. Chin with 2 slender barbels which usually reach to or posterior to rear edge of
preopercle, their length 1.25 to 1.55 times in head length; snout moderate, its length 1.7 to 2.1 times
in head length; dorsal profile of snout straight to before eye; mouth small, maxilla reaching about 1/2
distance to anterior edge of eye; a single row of well-spaced, stout, conical teeth in jaws; no teeth on roof
of mouth. First gill arch with 6 to 8 (rarely 8) gill rakers on upper limb, 21 to 25 on lower limb (total
28 to 32). Two well-separated dorsal fins, the first with VIII spines, the second fin with 9 soft rays; first
dorsal-fin spine very short; longest dorsal-fin spine 1.4 to 1.8 times in head length; last ray of second
dorsal and anal fins distinctly longer than penultimate ray; anal fin with I spine and 6 soft rays;
pectoral-fin rays 15 to 17 (usually 16, one of 56 specimens examined with 15 on one side). Lateral-line
scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of scales along upper part of caudal
peduncle. Colour: yellowish grey to light red, often with a pale blue to whitish spot on scales of body and
irregular blue lines around eye; a black spot usually much larger than eye centred on lateral line below
rear base of first dorsal fin, with a large oval white spot posterior and adjacent to it; base of second
dorsal fin blackish, the outer part yellow with dark-edged blue lines.
Habitat, biology, and fisheries: Generally found foraging by day over sand adjacent to inshore coral
reefs; feeds on a wide variety of sand-dwelling invertebrates such as crabs (with crab larvae 36% of
stomach contents of 15 specimens), polychaetes (20%), shrimps, heart urchins, sipunculids, foraminifera,
ophiuroids, other crustaceans, and small gastropods; occasionally takes small fishes.
the Indo-Pacific region; in the
western Indian Ocean from
southern Oman to southern
Mozambique; in the western
Pacific from the Ryukyu Is-
lands to Montague Island,
New South Wales. East in
Oceania to Hawaii and the
Pitcairn Group.
3194 Bony Fishes
Parupeneus spilurus (Bleeker, 1854) (Plate XXVII, 202)

Frequent synonyms / misidentifications: Parupeneus signatus (Günther, 1867) / Parupeneus
fraterculus (non Valenciennes, 1831).
FAO names: En - Blackspot goatfish; Fr - Rouget-barbet à tache noire.
Diagnostic characters: Body slightly elongate, its depth 3.1 to 3.5 times in standard length, and
moderately compressed. Chin with 2 slender barbels which usually reach to below rear edge of
preopercle, their length 1.35 to 1.65 times in head length; snout short, its length 1.8 to 2.15 times
in head length; dorsal profile of snout before eye slightly concave to straight; mouth small, maxilla reaching
about 2/3 distance to front edge of eye; a single row of well-spaced, stout, conical teeth in jaws; no teeth
to 32). Two well-separated dorsal fins, the first with VIII spines, the second fin with 9 soft rays; first
dorsal-fin spine very short; longest dorsal-fin spine 1.4 to 1.8 times in head length; last ray of second dorsal
and anal fins slightly longer than penultimate ray; anal fin with I spine and 6 soft rays; pectoral-fin rays
15 or 16 (usually 16). Lateral-line scales 27 or 28; 3 rows of scales between dorsal fins; 8 or 9 rows of
scales along upper part of caudal peduncle. Colour: orange-red to reddish brown, the scale edges narrowly
darker, shading to pale red or white ventrally; a black spot as large or larger than eye dorsally at
posterior end of caudal peduncle, its lower edge resting on lateral line, its upper edge near top of
peduncle, usually not meeting spot of other side (or if it does, the intervening pigment is generally less
intense); an elongate pale yellowish blotch adjacent and anterior to black spot, extending to below rear
base of second dorsal fin; 3 brown stripes, separated by 2 pale red to whitish stripes, passing from snout
along anterior part of body parallel with dorsal contour, the middle brown stripe passing through eye, the
lower just above pectoral-fin base (brown stripes merge and white stripes disappear below anterior part of
second dorsal fin); a faint reddish to yellow stripe along lower side of body; fins orange-red; barbels white.
Size: Maximum reported total length 50 cm; largest of 74 specimens examined 35.5 cm total length.
Habitat, biology, and fisheries: An inshore species generally found on rocky substrata or coral reefs.
Often seen by day at rest on the bottom during daylight hours, suggesting possible nocturnal feeding.
Adults solitary or in small aggregations; juveniles tend to form small schools.
Distribution: Known only from the western Pacific from Japan (collected by the present author at
Shirahama in southern Honshu) to New South Wales, Lord Howe Island, Norfolk Island, northern New
Zealand, and New Caledonia;
also in Western Australia
south to Rottnest Island. Indi-
viduals collected in the Philip-
pines and Indonesia appear to
be the same. Unusual in its
broad range of latitude and
hence its tolerance of a wide
range of temperature.
Upeneus arge Jordan and Evermann, 1902

Frequent synonyms / misidentifications: None / Upeneus taeniopterus (non Cuvier, 1829).
FAO names: En - Bandtail goatfish.
Diagnostic characters: Body elongate, its depth 3.65 to 4.75 times in standard length; snout length 2.25
to 2.45 times in head length. Chin with 2 slender barbels which nearly or just reach or extend slightly
posterior to rear margin of preopercle, their length 1.3 to 1.6 times in head length. Mouth small;
villiform teeth in a narrow band in jaws, on palatines, and in 2 small patches on vomer. First gill arch with
5 or 6 gill rakers on upper limb, 15 to 17 on lower limb (total 20 to 23). Two well-separated dorsal fins,
the first with VIII spines (first spine minute), and the second with 9 soft rays; second and third dorsal-fin
spines longest, 1.25 to 1.6 times in head length; anal fin with I spine and 7 soft rays; pectoral-fin
rays 13 to 15 (usually 14). Lateral-line scales 35 to 37; no scales on side of snout; small scales
basally on second dorsal and anal fins; 5½ scales between dorsal fins; 12 or 13 scales along dorsal
edge of caudal peduncle. Colour: silvery, the scales dorsally on body edged with orange-pink; 2 narrow
orange-yellow stripes on side of body (the upper more orange, the lower more yellow); upper lobe of
caudal fin of adults with 6 to 8, and the lower lobe with 4 to 6 dusky orange to black cross bands;
first dorsal fin with oblique rows of reddish spots; second dorsal fin with oblique dull orange-red bands
which become blackish distally and separated at outer margin by white; remaining fins whitish; barbels
white to pale yellow; peritoneum whitish.
Size: Largest specimen examined 36.5 cm total length, commonly to 25 cm.
Habitat, biology, and fisheries: Generally found on sand bottoms near rocky substrata or coral reefs,
often in very shallow water. Ten specimens were collected in Aldabra on sand in less than 1.3 m. Usually
encountered in small groups.
Distribution: This species has been found mainly at relatively small islands. It is known in the Pacific from
Hawaii (type locality) and Tuamotu Archipelago west to the islands of Micronesia, Fiji, and the Ryukyu
Islands. In the Indian Ocean
from the Maldives, Chagos Ar-
chipelago, Seychelles, and a
single collection from Mozam-
bique. It remains unknown
from the coast of Asia and Aus-
tralia, Indonesia, Philippines,
and Taiwan Province of China.
Remarks: Upeneus arge has
been regarded in recent publi-
cations as a synonym of U.
taeniophorus Cuvier, following
the advice of Paul Guézé.
However, the latter species is
distinct in having a deeper body, slightly lower lateral-line scale counts, and conspicuously dark-striped
dorsal fins, the first dorsal fin tipped with black. It was described from Sri Lanka and may be confined to
southern Asian waters.
3196 Bony Fishes
Upeneus moluccensis (Bleeker, 1855) (Plate XXVII, 203)

FAO names: En - Goldband goatfish; Fr - Rouget-souris bande or; Sp - Salmonete de bande dorada.
Diagnostic characters: Body moderately elongate, its depth 3.5 to 4.05 in standard length (body tends
to become relatively deeper with growth); snout length 2.45 to 2.75 times in head length. Chin with 2
slender short barbels which do not reach posterior margin of preopercle, their length 1.55 to 2.1
times in head length. Mouth small; villiform teeth in a narrow band in jaws, on palatines, and in 2 small
patches on vomer. First gill arch with 7 to 9 gill rakers on upper limb, 19 to 22 on lower limb (total
26 to 30). Two well-separated dorsal fins, the first with VIII spines (first spine minute), and the second with
9 soft rays; second and third dorsal-fin spines longest, 1.45 to 1.65 times in head length; anal fin with I
spine and 7 soft rays; pectoral-fin rays 15 to 17 (nearly always 16). Lateral-line scales 33 to 35; no
scales on side of snout; small scales basally on second dorsal and anal fins; 5½ scales between dorsal
fins; 12 or 13 scales along dorsal edge of caudal peduncle. Colour: a golden yellow stripe about as
broad as pupil from behind upper part of eye to upper base of caudal fin; back above stripe silvery
pink, below silvery; snout silvery pink; upper lobe of caudal fin with 6 or 7 oblique red cross bars, the
lower lobe red or longitudinally streaked with red; dorsal fins with red stripes; barbels white to pink;
peritoneum brown to dark brown.
Habitat, biology, and fisheries: A species of continental or large insular shelves, generally on mud
substrata; known from a depth range of 10 to 80 m. Usually occurs in schools. Caught mainly by trawling.
Of limited commercial importance due to small size, but occasionally taken in large numbers.
Distribution: Red Sea (and
Mediterranean Sea as an im-
migrant via the Suez Canal)
south to Mozambique and
Madagascar (also an unex-
pected record from Réunion),
east along the southern shore
of Asia (but not the Persian
Gulf) to the western Pacific
where it ranges from Kyushu
to northern Australia.
Upeneus sulphureus Cuvier, 1829 (Plate XXVII, 204)

FAO names: En - Sulphur goatfish; Fr - Rouget-souris aurore; Fr - Salmonete aurora.
Diagnostic characters: Body slightly elongate, its depth 3.25 to 3.85 times in standard length; snout length
2.4 to 2.7 times in head length. Chin with 2 slender barbels which usually reach or extend posterior
to rear margin of preopercle, their length 1.25 to 1.7 times in head length (barbels relatively longer in
adults than young). Mouth small; villiform teeth in a narrow band in jaws, on palatines, and in 2 small
patches on vomer. First gill arch with 7 to 9 gill rakers on upper limb, 18 to 22 on lower limb (total
26 to 30). Two well-separated dorsal fins, the first with VIII spines (first spine minute), and the second with
9 soft rays; second and third dorsal-fin spines longest, 1.45 to 1.65 times in head length; anal fin with I
spine and 7 soft rays; pectoral-fin rays 14 to 17 (rarely 14 or 17). Lateral-line scales 33 to 36 (rarely
33 or 36); no scales on side of snout; small scales basally on second dorsal and anal fins; 5½ scales
between dorsal fins; 12 or 13 scales along dorsal edge of caudal peduncle. Colour: silvery greenish or
pink dorsally, shading to silvery on sides and ventrally, with 2 narrow brassy yellow stripes on side of
body; caudal fin without dark cross bands; first dorsal fin broadly tipped with black; rest of fin with
2 broad dusky yellow stripes, the narrow interspaces white; remaining fins dusky yellowish; barbels white;
peritoneum brown to dark brown.
Habitat, biology, and fisheries: Known from coastal waters of continents and large islands; generally
found on mud substrata in the depth range of 20 to 60 m; often enters estuaries. Caught mainly by trawling,
sometimes in large numbers.
Distribution: Southern Red
Sea to southern Mozambique
and Madagascar (also re-
corded from Seychelles), east
along continental shores to In-
donesia and northern Austra-
lia, north to southern Japan.
East in Oceania only to Fiji.
Absent from the Persian Gulf;
previous records of the spe-
cies from the Gulf are misi-
d e nt i fi c at i on s of Upeneus
doriae.
3198 Bony Fishes
Upeneus sundaicus (Bleeker, 1855) (Plate XXVII, 205)

Frequent synonyms / misidentifications: Upeneus luzonius Jordan and Seale, 1907 / None.
FAO names: En - Ochreband goatfish; Fr - Rouget-souris ocre; Sp - Salmonete de banda ocre.
(based on a photo of a very small specimen, 11 mm total length)
Diagnostic characters: Body moderately elongate, its depth 3.55 to 4 times in standard length; snout
length 2.05 to 2.5 times in head length. Chin with 2 slender barbels which usually reach or extend
slightly posterior to rear margin of preopercle, their length 1.3 to 1.7 times in head length. Mouth
small; villiform teeth in a narrow band in jaws, on palatines, and in 2 small patches on vomer. First gill
arch with 4 to 6 gill rakers on upper limb, 14 to 16 on lower limb (total 19 to 22). Two well-separated
dorsal fins, the first with VIII spines (first spine minute), and the second with 9 soft rays; second
dorsal-fin spine longest (but third nearly as long), 1.1 to 1.3 times in head length; anal fin with I spine
and 7 soft rays; pectoral-fin rays 13 to 15 (usually 14). Lateral-line scales 31 to 32; no scales on side
of snout; small scales basally on second dorsal and anal fins; 4½ scales between dorsal fins; 10 or 11
scales along dorsal edge of caudal peduncle. Colour: bronze to silvery green dorsally, shading to silvery
white on sides and ventrally, with a narrow brownish yellow stripe on body at level of upper part of
eye containing 3 darker zones, 1 under each dorsal fin and 1 below anterior caudal peduncle; upper lobe
of caudal fin with 3 or 4 dark cross bands (may be faint); lower lobe of caudal fin with a broad dusky
posterior border, with or without faint dark cross bands; fins yellowish; barbels yellow; peritoneum
pale.
Size: Largest specimen examined 17.2 cm, commonly to 13 cm.
Habitat, biology, and fisheries: Little is reported on the habitat or biology of this species. It is presumed
to live on mud or silty sand substrata, since it is usually taken by trawls or seines. Depth records of museum
specimens vary from 3 to 20 m.
Distribution: P h i l i p p i n e s ,
Papua New Guinea, Indone-
sia, Queensland, northwest-
ern Australia, Viet Nam, and
Thailand to Persian Gulf.
?
Upeneus tragula (Richardson, 1846) (Plate XXVII, 206)

Frequent synonyms / misidentifications: Upeneus oligospilus Lachner, 1954 / None.
FAO names: En - Freckled goatfish; Fr - Rouget-souris à bande sombre; Sp - Salmonete barborín.
Diagnostic characters: Body elongate, its depth 3.9 to 4.25 times in standard length; snout length 2.25
to 2.65 times in head length. Chin with 2 slender barbels, usually not reaching rear margin of
preopercle, their length 1.4 to 1.85 times in head length. Mouth small; villiform teeth in a narrow band
in jaws, on palatines, and in 2 small patches on vomer. First gill arch with 5 to 7 gill rakers on upper
limb, 15 to 18 (rarely 18) on lower limb (total 21 to 25). Two well-separated dorsal fins, the first with VIII
spines (first spine minute), and the second with 9 soft rays; second and third dorsal-fin spines longest,
1.35 to 1.65 times in head length; anal fin with I spine and 7 soft rays; pectoral-fin rays 13 or 14 (more
often 13). Lateral-line scales 28 to 30; scales present on side of snout; small scales basally on second
dorsal and anal fins; 4½ scales between dorsal fins; 10 or 11 scales along dorsal edge of caudal peduncle.
Colour: a dark reddish brown to blackish stripe from front of snout through eye, along upper side,
to base of caudal fin; head and body above stripe brownish to greenish grey, flecked with small
dark reddish brown or blackish spots; body below stripe white with rows of small dark spots; upper
lobe of caudal fin with 4 to 6 blackish cross bands, and lower lobe with 5 to 7; first dorsal fin whitish
with a large dark red to black blotch containing 2 to several small yellow spots on outer third to
half of fin; a second irregular dark blotch in lower part of fin; second dorsal with 3 dark reddish or blackish
bands; barbels usually yellow.
Size: Reported to 33 cm total length, commonly to 25 cm.
Habitat, biology, and fisheries: A shallow-water species of protected waters of bays and lagoons; often
found on sand or silty sand substrata near coral reefs; also common in estuaries. Caught by seines, trawls,
gill nets, and traps.
Distribution: P e r si a n G ulf
and southern Red Sea to
Mozambique, east to the
w e s t e r n P a c i fi c w h e r e i t
ranges from southern Japan
to central New South Wales,
east to New Caledonia, Vanu-
atu, and Palau.
3200 Bony Fishes
Upeneus vittatus (Forsskål, 1775) (Plate XXVII, 207)

FAO names: En - Striped goatfish; Fr - Rouget-souris orangé; Sp - Salmonete naranjero.
Diagnostic characters: Body moderately elongate, its depth 3.5 to 4 times in standard length; snout length
2.3 to 2.7 times in head length. Chin with 2 slender barbels, usually not reaching rear margin of
preopercle, their length 1.4 to 1.85 times in head length. Mouth small; villiform teeth in a narrow band
in jaws, on palatines, and in 2 small patches on vomer. First gill arch with 7 to 9 gill rakers on upper
limb, 19 to 21 (rarely 18) on lower limb (total 21 to 25). Two well-separated dorsal fins, the first with VIII
spines (first spine minute), and the second with 9 soft rays; second and third dorsal-fin spines longest, 1.3
to 1.55 times in head length; anal fin with I spine and 7 soft rays; pectoral-fin rays 15 to 17 (usually 16).
Lateral-line scales 34 to 36; scales present on side of snout; small scales basally on second dorsal
and anal fins; 5½ scales between dorsal fins; 11 or 12 scales along dorsal edge of caudal peduncle. Colour:
silvery greenish on back, silvery on sides, with 4 distinct narrow brassy yellow stripes on upper
two-thirds of body; ventral part of body often yellow; upper lobe of caudal fin with 4 or 5 dark cross bands;
lower lobe of caudal fin with 3 dark cross bands, the distal band nearly twice as broad and more
intensely black than middle band; first dorsal fin broadly tipped with black, with 2 broad dusky yellow
stripes below; second dorsal fin with 3 dusky yellow stripes; remaining fins whitish; barbels usually white;
peritoneum dark brown to blackish.
Size: Reported to 28 cm total length, commonly to 22 cm.
Habitat, biology, and fisheries: This species is usually found on mud bottoms from shallow estuarine
areas to depths of at least 100 m. It is caught mainly in seines, trawls, or traps.
Distribution: The most wide-
ranging species of the genus,
occurring from the Red Sea
south to East London, South
Africa, east to the Society Is-
lands and Marquesas; in the
western Pacific from Shikoku
to Queensland. Inadvertently
introduced to Hawaii with a
shipment of Marquesan sar-
dines.
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Perciformes: Percoidei: Pempheridae 3201
PEMPHERIDAE
Sweepers (bullseyes)
by R.D. Mooi
D iagnostic characters: Small to medium-sized fishes (up to 22 cm), elongate to deep-bodied and
compressed. Eye large, without adipose lid. Snout short. Opercular spines short and blunt. Gill rakers
on first arch long and numerous (more than 20). Branchiostegal rays 7. Dorsal fin very short, usually
higher than long, much shorter than anal-fin base and in advance of anal-fin origin, with V to VII
narrow spines and 7 to 13 soft rays; anal-fin base long, the fin with III spines (first spine short and often
difficult to see) and 18 to 45 soft rays; caudal fin shallowly forked; pelvic fins with I spine and 5 soft rays;
pectoral fins with 16 to 19 rays. At least some scales ctenoid in all species, in some species cycloid scales
dominate anterodorsally and laterally; lateral-line scales 44 to more than 80; lateral line complete,
extending far onto caudal fin; lateral-line scales exposed or covered. Coracoid expanded posteriorly.
Some species bioluminescent, with light organs associated with the pyloric caeca and/or rectum. Colour:
variable, but often pinkish or reddish, flanks silvery, iridescent; dorsal, anal, and caudal fins often bordered
or tipped by black.
short dorsal fin, in

advance of anal fin
large eye
lateral
line
extending
onto
caudal fin
long anal fin
Habitat, biology, and fisheries: Found mainly in tropical and subtropical areas in shallow water (less than
100 m), common on coral reefs. During the day, often found in large schools in caves or protected areas
from which they disperse at night to feed on crustaceans and polychaete worms. Many species have
bioluminescent organs associated with the pyloric caeca and sometimes the rectum; in at least Parapria-
canthus the bioluminescent chemistry appears to be obtained from food items. Family of no commercial
value, but their local abundance results in capture by trap nets and shrimp trawls, and explains their
appearance in fish markets. Used as fish paste non-commercially in at least Japan. Meat of some species
reported as tasty.
Remarks: Species generally poorly described with many taxonomic problems. The most obvious of these
is among the cycloid-scaled, keeled Pempheris. Four species of this group are recognized here (P. otaiten-
sis, P. oualensis, P. schwenkii, P. vanicolensis), although a further 4 nominal taxa are reported for the area
(P. adusta Bleeker, 1877; P. itoi Fowler, 1931; P. mangula Cuvier, 1829; P. molucca Cuvier, 1829). A
thorough revision of the family is required. As a result, distributions of individual species are not well known
and individual species maps are not provided. Examples of Pempheris are reported as far east as Pitcairn
Island and are expected throughout the covered area. Parapriacanthus has been reported from the
Phoenix Islands and westward in the WCP area.

Berycidae: similar in having a short dorsal fin in advance of a
long-based anal fin and the lateral line extending well onto the
caudal fin; easily distinguished by having IV anal-fin spines,
pelvic fins with I spine and 7 to 13 soft rays, head ornamenta-
tion consisting of serrate ridges and spines, very rough and
almost keeled scales, and deeply forked caudal fin.
Berycidae
3202 Bony Fishes
Leptobramidae: similar in having a short dorsal fin and long-

based anal fin; easily distinguished by the dorsal-fin origin
being posterior to anal-fin origin and closer to caudal-fin base
than snout tip, eye with adipose lid, maxilla extending beyond
posterior margin of eye, fewer than 15 gill rakers on first gill
arch, steel blue back.
Leptobramidae
Key to the species of Pempheridae occurring in the area
1a. Body elongate, its depth less than or equal to head length; anal fin naked or scaly only
basally; 28 or fewer soft rays in anal fin; lateral line extending about half-way along
middle caudal-fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Parapriacanthus) ® 2
1b. Body deep, its depth greater than head length; anal fin scaly over 1/2 of ray length; 30
or more soft rays in anal fin; lateral line extending to distal tips of middle caudal-fin rays
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Pempheris) ® 4
2a. Dorsal fin with VI spines and 10 or 11 soft
rays; anal-fin with 26 to 28 soft rays; first gill
arch with 7 or 8 gill rakers on upper limb, 19
to 22 on lower limb (Fig. 1) . . . . Parapriacanthus marei
2b. Dorsal fin with V or VI spines and 8 or 9 soft
rays; anal fin with 24 or fewer soft rays; first
gill arch with 4 to 7 gill rakers on upper limb,
14 to 19 on lower limb . . . . . . . . . . . . . . . . . ® 3
Fig. 1 Parapriacanthus marei
3a. Lateral-line scales 54 to 64 (usually 56 to 61); dorsal fin with VI spines (sometimes V);
anal fin with 22 to 24 soft rays (usually 23); deeper-bodied, with 2 or more scales
between midventral edge of coracoid and midlateral edge of pelvic bone (Fig. 2a) . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapriacanthus dispar
3b. Lateral-line scales 60 to 79 (usually 62 to 75); dorsal fin with V spines (sometimes VI);
anal fin with 18 to 23 soft rays (usually 19 or 20); more elongate body with 1 or 12 scales
between coracoid and midlateral edge of coracoid and midlateral edge of pelvic bone
(Figs 2b and 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parapriacanthus ransonneti
2 or more 1-1½ scale

scale rows rows
a) Parapriacanthus. b) Parapriacanthus
dispar Fig. 2 ransonneti
Fig. 3 Parapriacanthus ransonneti
4a. Scales on flank ctenoid and adherent; lat- sensory

tube
eral-line scales equal in size or slightly
larger than surrounding scales, sensory
tubes wider than long (Fig. 4a); prepelvic
area broad and unkeeled . . . . . . . . . . . . . . ® 5
4b. Scales on flank cycloid and deciduous; lat-
eral-line scales smaller than (and partly or
mostly concealed by) surrounding scales,
a) b)
sensory tubes longer than wide or wider
than long (Fig. 4b); prepelvic area narrow Fig. 4 lateral-line scales
and keeled or broad and unkeeled . . . . . . . . . ® 8
Perciformes: Percoidei: Pempheridae 3203
5a. Dark Y-shaped organ

visible just anterior to
anus (Fig. 5); axillary
scale present (shown
black in Fig. 2); first gill
arch with 5 to 7 (rarely 9)
gill rakers on upper limb
and 19 to 21 (rarely 22 or
23) on lower limb (total
24 to 29, but usually 26 or
27); lateral-line scales 62 Fig. 5 Y-shaped organ Fig. 6 Pempheris ypsilychnus
to 74, usually 64 to 69 between pelvic fins and anus
(Fig. 6) . . . Pempheris ypsilychnus in Pempheris ypsilychnus
(Australia only)
5b. No Y-shaped organ visible anterior to anus; no axillary scale; first arch with 9 to 13 gill rakers
on upper limb and 21 to 29 on lower limb (total 30 to 42); lateral-line scales 63 to 84 . . . . . . . . . ® 6
6a. Variably bronze or silvery brown with dark dorsal-fin tips and anterior anal-fin tips;
lateral-line scales 63 to 77 (usually less than 73); anal-fin rays 30 to 36 (usually 34 or
35); first gill arch with 32 to 36 gill rakers (Fig. 7) . . . . . . . . . . . . . . . . . . Pempheris analis
(in the area, known from eastern Australia)
6b. Coloration as above, but lateral-line scales 71 to 84; anal-fin rays 33 to 38 (usually 34 or
more); first gill arch with 30 to 42 gill rakers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Lateral-line scales 72 to 77; first gill arch with 30 to 34 gill rakers (usually 32 to 33) . Pempheris japonica
(Philippines northward)
7b. Lateral-line scales 72 to 84; usually more than 76; first gill arch with 37 to 42 gill rakers . . Pemperis rapa
(French Polynesia)
8a. Dorsal fin with V spines and 10 to 12 soft rays; prepelvic area broad and unkeeled;
lateral-line sensory tubes wider than long (Fig. 4a); scales above lateral line 6 to 8;
anterior luminous organ present and associated with pyloric caeca; yellowish brown to
brownish orange (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pempheris affinis
(southeastern Australia)
8b. Dorsal fin with VI spines and 8 to 10 soft rays; prepelvic area narrow and keeled;
lateral-line sensory tubes longer than wide (Fig. 4b); scales above lateral line 3 to 6;
anterior luminous organ absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Lateral-line scales 46 to 54; 3 or 4 scales between lateral line and dorsal-fin origin; 9 to 13
scales between lateral line and anal-fin origin; deep purplish dorsally, almost black posterior
to dorsal fin, shading to silvery flanks with pinkish or lavender iridescence, leading edge of
dorsal fin dark, base of anal fin dark with remainder of fin pale or lightly pigmented, caudal
fin dusky or yellow; usually less than 12.5 cm standard length (Fig. 9) . . . . . . . Pempheris schwenkii
9b. Lateral-line scales more than 53; 5 or 6 scales between lateral line and dorsal-fin origin; 12
to 16 scales between lateral line and anal-fin origin; body generally coppery or silver
iridescent, dorsal fin with dark leading edge and black distal tips to anterior soft rays, anal fin
with or without dark base but remainder of fin entirely pigmented or pale with black distal
margin, caudal fin often with distinct dark posterior margin; often larger than 12.5 cm
standard length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
Fig. 7 Pempheris analis Fig. 8 Pempheris affinis Fig. 9 Pempheris schwenkii

3204 Bony Fishes
10a. Anal fin completely dusky, or mostly dusky except for a pale distal margin; dorsal fin with
a dark leading edge that expands distally to produce dark tips on the first 2 or 3 soft rays;
pectoral fins usually with distinctly dark base; caudal fin usually without a dark posterior
margin (Fig. 10) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pempheris oualensis
10b. Anal fin pale or lightly pigmented with a dark distal margin; dorsal fin with a dark leading
edge with or without dark tips on the first 5 or 6 soft rays; pectoral fins with or without
distinctly dark base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
11a. Dorsal fin with dark leading edge, not expanded to produce dark tips on soft rays;
pectoral fins usually with a distinct dark base . . . . . . . . . . . . . . . . . . . Pempheris otaitensis
11b. Dorsal fin with dark leading edge expanding distally to produce dark tips on the first 5 or
6 soft rays; pectoral fin usually without a distinct dark base (Fig. 11) . . . . . Pempheris vanicolensis
Fig. 10 Pempheris oualensis Fig. 11 Pempheris vanicolensis

Parapriacanthus dispar (Herre, 1935)
Parapriacanthus marei Fourmanoir, 1971
Parapriacanthus ransonneti Steindachner, 1870
Pempheris affinis McCulloch, 1911
Pempheris analis Waite, 1910
Pempheris japonica Döderlein, 1883
Pempheris otaitensis Cuvier, 1831
Pempheris oualensis Cuvier, 1831
Pempheris rapa Mooi, 1998
Pempheris schwenkii Bleeker, 1855
Pempheris vanicolensis Cuvier, 1831
Pempheris ypsilychnus Mooi and Jubb, 1996
References
Mooi, R.D. and R.N. Jubb. 1996. Descriptions of two new species of the genus Pempheris (Pisces: Pempherididae)
from Australia, with a provisional key to Australian species. Rec. Aust. Mus., 48(2):117-130.
Tominaga, Y. 1963. A revision of the fishes of the family Pempheridae of Japan. J. Fac. Sci., Tokyo, Sect.
4,10(1):269-297.
Tominaga, Y. 1968. Internal morphology, mutual relationships and systematic position of the fishes belonging to the
family Pempheridae. Japan. J. Ichthyol., 15(2):43-95.
The present contribution is based upon work supported by the National Science Foundation under Grant No.
DEB-9317695.
Perciformes: Percoidei: Glaucosomatidae 3205
GLAUCOSOMATIDAE
Pearl perches
by R.J. McKay
D iagnostic characters: Robust to deeply ovate, compressed, marine coastal perch-like fishes (size to
about 70 cm). Head large; scales present on entire head except tip of snout, lips, and chin. Preopercle
with blunt spines at angle or entire; opercle with 1 blunt spine. Mouth large, oblique, terminal, and
protractile; lips thin; maxilla broad, scaly, with a narrow supplemental (supramaxillary) bone above,
scarcely slipping below preorbital; lower jaw protruding. Teeth pointed or conical, in narrow bands in jaws,
usually without canines; teeth on vomer, palatines, and tongue. Dorsal fin single, with VIII graduated
spines and 12 to 14 soft rays (much higher than spines); anal fin with III short graduated spines and
9 to 12 soft rays; caudal fin lunate or emarginate, frequently with tips produced or filamentous; pectoral
fins short and blunt; pelvic fins small, below base of pectoral fins, with I spine and 5 soft rays. Scales
ctenoid, small or moderate, a basal sheath present; lateral line continuous, tubes simple, continued onto
tail. Haemapophyses of fifth to tenth abdominal vertebrae modified, the sixth to eighth forming flat plates
to which the swimbladder is firmly bound. Colour: silvery grey to reddish brown with distinct or indistinct
stripes following scale rows; usually with 1 or more dark bands on head; a dark or silvery blotch at posterior
base of dorsal fin.
dorsal fin with VIII spines

and 12-14 soft rays
distinct, thin supplemental

bone (supramaxilla) on upper anal fin with III spines
end of maxilla and 26-30 soft rays
Habitat, biology, and fisheries: Generally frequenting submerged reefs, pinnacles, and rough rocky
bottoms in moderately deep water, but moving into shallow water at times. Usually strictly bottom feeders
but may move up into the water column to take a bait on occasions. These fishes feed mostly at dawn or
dusk and may be taken throughout the day and night in deep water. All are superior table fishes with white
flesh of excellent texture and superb flavour. Recognized in Australia to be some of the finest food fishes
available.
Lutjanidae: somewhat similar in shape to pearl perches, but have strong canine teeth on jaws, and differ
in lacking a distinct thin supplemental (supramaxillary) bone on the upper edge of the maxilla.
Pempheridae: related to the Glaucosomatidae in having the same structure of the swimbladder, and the
modification of the haemapophyses of the fifth to tenth abdominal vertebrae, but differ in having more
anal-fin elements (i.e. spines plus soft rays: 17 to 45 versus 9 to 12), less dorsal-fin spines (IV to VI versus
VIII or IX), and lack a supplemental (supramaxillary) bone.
Lutjanidae Pempheridae
3206 Bony Fishes
Key to the species of Glaucosomatidae occurring in the area supraclavicle covered with black membrane,
1a. Three dark vertical bands from nape, the first through “pearl-like” when exposed
eye, the second along edge of preopercle and the
third down to pectoral-fin base; dorsal, caudal, and
anal fins with some filamentous rays; dorsal-fin rays
14; anal-fin rays 12 . . . . . . . . . . . Glaucosoma magnificum
(northern Australia and southern New Guinea)
1b. A single dark vertical band through eye that may
disappear with increasing size; dorsal-fin rays 11;
anal-fin rays 9 . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Supraclavicle (shoulder bone) developed into a

prominent bony shield (Fig. 1) . . . . . . . Glaucosoma scapulare
(eastern Australia) Fig. 1
2b. Supraclavicle not developed into a bony shield; peri-
toneum and gill rakers black; lateral line with 49 to 51
pored scales . . . . . . . . . . . . . . . . . Glaucosoma buergeri

0
0
Glaucosoma buergeri Richardson, 1845
0
Glaucosoma magnificum (Ogilby, 1915)
Glaucosoma scapulare Ramsay, 1881
Reference
McKay, R.J. 1997. FAO species catalogue. Vol. 17. Pearl perches of the world (family Glaucosomatidae). An annotated
and illustrated catalogue of the pearl perches known to date. FAO Fish. Synop., (125)17:26 p.
Glaucosoma buergeri Richardson, 1845

Frequent synonyms / misidentifications: Glaucosoma fauvelii Sauvage, 1881; G. taeniatus Fowler,
1934; G. bürgeri Richardson, 1845 / Glaucosoma hebraicum non Richardson, 1845.
FAO names: En - Grey bigmouth bream.
adult
Diagnostic characters: Body robust, moderately compressed,

its depth 1.9 to 2.3 times in standard length. Head almost entirely
scaled, profile with a slight concavity before eye. Eye large, its
diameter 2.9 to 3.7 times in head length; interorbital space
convex, its width 3.9 to 4.8 times in head length; suborbital width
less than eye diameter (1.9 to 2.7 times in eye diameter) and 6.6 juvenile
to 7.8 times in head length. Snout short, its length 3.5 to 4.4 times
in head length. Mouth large, oblique, reaching to about posterior margin of eye; maxilla scaly, with thin
supplemental (supramaxillary) bone above, its width at the end 4.7 to 5.5 times in head length and 1.4 to
1.8 times in eye diameter; teeth small and canine-like (no enlarged canines), in several rows in upper jaw
and in a single series at side of lower jaw; a narrow V-shaped band of fine teeth on vomer. First gill arch
with 6 to 9 gill rakers on upper limb, 13 to 15 on lower limb. Dorsal-fin spines VIII, increasing in length from
a small anterior spine to a long posterior one; soft dorsal-fin rays 11, soft-rayed part of fin much higher
than spinous part with anterior ray often produced into a filament; anal fin with III slender spines and 9 soft
rays; caudal fin slightly emarginate with pointed tips; pectoral-fin rays 16. Lateral line almost straight, with
49 to 51 (usually 50) tubed scales extending onto caudal-fin base; 12 scale rows between lateral line and
origin of dorsal fin. Colour: body silvery grey; about 10 longitudinal thin bands along body, their widths
much less than diameter of pupil and less than their interspaces (bands become indistinct or
completely disappear with age); a vertical broad dark band from nape, curving through eye to lower
opercular angle and continued on subopercle; back at base of last dorsal-fin ray usually with a dark blotch;
roof of mouth with black mottling, coalescing lines, or completely black; tongue with black mottling;
peritoneum black (in dead specimens the black peritoneum may be slightly everted through the
anus); fins greyish. Juveniles with narrow dark bars and eye-stripe well defined.
Size: Maximum total length about 45 cm, commonly to 35 cm; maximum weight 2.5 kg.
Habitat, biology, and fisheries: Coastal waters in moderate depths on the continental shelf, generally
close to reefs or rough bottoms. Taken by bottom trawls and line. Marketed fresh, a small quantity is salted.
Distribution: Western Aus-
tralia from Koks Island (Shark
Bay) northwards to Cape Tal-
bot; Japan from Kochi Prefec-
ture (Ryukyu Islands)
southwards and along the
north China coast to Taiwan
Province of China and Viet
Nam; probably does not occur
in the Philippines.
3208 Bony Fishes
Glaucosoma magnificum (Ogilby, 1915)

FAO names: En - Threadfin pearl perch.
r o b u s t , m o d e r a t e l y c o m-
pressed, its depth 1.6 to 1.8
times in standard length. Head
almost entirely scaled, profile
evenly rounded. Eye large, its
diameter 2.3 to 2.6 times in
head length; interorbital space
convex, its width 2.9 to 3.2
times in head length; preorbital
width less than eye diameter
and 7.8 to 9.6 times in head
length. Snout short, its length 5
to 5.7 times head length.
Mouth large, oblique, reaching
to about middle of eye; maxilla
scaly, with thin supplemental
(supramaxilly) bone above, its
width at the end 4.9 to 5.4
times in head length and 1.9 to
2.3 times in eye diameter;
teeth small and canine-like (no enlarged canines), in several rows in upper jaw and in a single series at
side of lower jaw; a narrow V-shaped band of fine teeth on vomer. First gill arch with 5 to 10 gill rakers on
upper limb, 17 to 19 on lower limb. Dorsal-fin spines VIII, increasing in length from a small anterior spine
to a long posterior one; soft dorsal-fin rays 14, soft-rayed part of fin much higher than spinous part
with anterior ray produced into a short filament, the third ray prolonged into a long simple filament
which extends past caudal fin, and the fifth ray also extended to beyond hypural joint; anal fin with
III slender spines and 12 soft rays, the second ray prolonged beyond caudal-fin base; pectoral fins
with 15 or 16 rays, the supraclavicle not noticeably enlarged, covered with silvery membrane; caudal fin
slightly emarginate with long filamentous tips. Lateral line almost straight, with 46 to 48 tubed scales
extending onto caudal-fin base; 14 to 16 scale rows between lateral line and dorsal-fin origin; dorsal
and anal fins with bases scaly and small interradial scales extending onto proximal half of fins. Colour:
body silvery grey to reddish brown with middle of scales golden to yellowish; upper surface of head
lavender; a broad dark brown band running through eye to lower margin of opercle and onto subopercle;
a second narrower band from nape down posterior edge of preopercle; a third narrow band from
nape behind second band and curving on upper sides of back down the posterior margin of opercle
to the pectoral fins; in life, with a dark or lavender edged bright silvery spot on each side of base of last
dorsal-fin ray and another on posterior dorsal surface of caudal peduncle; roof of mouth with dark bluish
to black shading on posterior part; tongue pale to dusky; gill rakers pale, dusky, or black with pale tips;
peritoneum brown; fins greyish, filaments dark.
Habitat, biology, and fisheries: Coastal waters in moderate depths on the continental shelf, generally
close to submerged reefs or rough bottom. Feeds on crustaceans; small cuttlefish are also taken. Taken
by bottom trawls; the larger specimens are considered good eating.
Distribution: From Exmouth
Gulf in Western Australia
across northern Australia to
Cape York in Queensland. Al-
most certainly also in New
Guinean waters.
Glaucosoma scapulare Ramsay, 1881

FAO names: En - Pearl perch.
Diagnostic characters: Body robust, moderately compressed, its depth 2.4 times in standard length. Head
almost entirely scaled, profile with a slight concavity before eye, its length 2.6 times in standard length.
Eye large, its diameter 3.2 times in head length; interorbital space convex, its width 4 times in head length;
preorbital width less than eye diameter, 7.7 times in head length and 2.4 times in eye diameter. Snout
short, its length 4 times in head length. Mouth large, oblique, reaching to about posterior margin of eye;
maxilla scaly, with thin supplemental (supramaxillary) bone above, its width at the end 5.3 times in head
length and 1.7 times in eye diameter; teeth small and canine-like (no enlarged canines), in several rows
in the upper jaw and in a single series at side of lower jaw; a narrow V-shaped band of fine teeth on vomer.
First gill arch with 6 gill rakers on upper limb, 15 to 17 on lower limb. Dorsal-fin spines VIII, increasing in
length from a small anterior spine to a long posterior one; soft dorsal-fin rays 11, soft-rayed part of fin much
higher than spinous part; anal fin with III slender spines and 9 soft rays; pectoral-fin rays 16, the
supraclavicle (shoulder bone or suprascapulary, see Fig. 1 in key) large, dome-shaped and covered
with a black membrane when alive, but exposed as a pearly-white smooth bone in the market; caudal
fin slightly emarginate with pointed tips. Lateral line almost straight with 49 to 50 tubed scales extending
onto base of caudal fin; 10 scale rows between lateral line and dorsal-fin origin. Colour: body silvery grey,
scales with a small indistinct golden brown spot at base; head and back with lavender reflections; in
juveniles only, an indistinct curved brown band from nape passing through eye to lower opercular angle
and continued on subopercle; supraclavicle deep iridescent blue-black, becoming pearl white when the
thin membrane is removed; scales of back and upper sides with a small indistinct golden brown to dark
fawn spot; dorsal surface at base of last dorsal-fin ray usually with a dark blotch; pectoral fins hyaline with
a small brown or black spot on inner base (axillary spot); membranes of dorsal and caudal fins pale, rays
white; anal fin hyaline, rays silvery or pale fawn; pelvic fins hyaline, rays milk-white.
Habitat, biology, and fisheries: Coastal waters in moderate depths to 90 m on the continental shelf,
generally close to submerged reefs or rough bottom. The species is stated to be a midwater-feeder moving
well up from the bottom in a
shoal to take a bait during the
night until about 10.00 p.m.
Most fishes are taken on the
bottom during the day. Taken
by handline. The pearl perch
is highly regarded as a food-
fish in Queensland and New
South Wales.
Distribution: F r o m R o c k-
hampton in Queensland
southwards to Port Jackson in
New South Wales.
3210 Bony Fishes
LEPTOBRAMIDAE
Beach salmon
by R.D. Mooi
Leptobrama muelleri Steindachner, 1878

FAO names: En - Beach salmon. posteriorly placed dorsal fin
Diagnostic characters: Body mod- adipose eye lid
erately slender, compressed. Head
somewhat short. Eye with adipose
lid. Maxilla extending beyond pos-
terior margin of eye. Teeth small
and pointed, forming relatively broad
toothed surfaces on vomer, pala-
tines, meso- and ectopterygoids, and
jaws. Branchiostegal rays 6. Few (9 long-based anal fin
to 11), short gill rakers on first gill
arch, absent on remaining arches.
Dorsal-fin origin far back on body, closer to caudal-fin base than snout tip; dorsal fin with IV spines
and 16 to 18 soft rays; anal-fin origin in advance of dorsal-fin origin, and fin long based with III spines,
26 to 30 soft rays. Scales ctenoid, median fins in scale sheaths; lateral-line scales 75 to 77. Unusual
internal features include 9 spineless pterygiophores anterior to dorsal fin (some of these may be
supraneurals); cranium with 5 well-developed longitudinal ridges. Colour: back steel blue; sides and belly
silvery white; dorsal fin with black tip anteriorly (at least in males).
large eye without short dorsal fin in
Similar families occurring in the area adipose lid advance of anal fin
Pempheridae: similar in having a short dorsal fin and long-based
anal fin, but easily distinguished by having origin of dorsal fin in
advance of anal-fin origin; dorsal-fin height greater than length of
dorsal-fin base, or fin almost as high as long; not more than 13 soft
rays in dorsal fin; large eye without adipose lid; short maxilla not
reaching beyond eye; numerous elongate gill rakers (usually more
than 20, always more than 15).
Carangidae: similar in having an adipose lid over eye, a relatively Pempheridae
long-based anal fin, and some species elongate; in addition, if
the inconspicuous first dorsal-fin spines of some species were spiny 1st dorsal fin
overlooked, it would appear that they have a posteriorly posi-
tioned dorsal fin; most similar is Scomberoides. However,
carangids are very different upon close examination by having 2
dorsal fins, the first anterior with IV to VIII spines; usually more
than 18 soft rays in dorsal fin; anal fin with III spines, the first 2
separate and more anteriorly positioned; scales small and cy-
cloid; narrow caudal peduncle; caudal fin widely forked.
narrow caudal
separate anal-fin spines peduncle
Size: Maximum total length 43 cm at a maximum weight of 900 g. Carangidae
Habitat, biology, and fisheries: Marine and brackish water,
occasionally entering rivers. Feeds on small fishes of inshore waters, as well as on shrimps. No commercial
catch, although apparently provides good angling.
Distribution: Southern Papua New Guinea and north-
ern Australia, including Queensland, Northern Territory,
and Western Australia.
Reference
Tominaga, Y. 1965. The internal morphology and systematic
position of Leptobrama mülleri, formerly included in the
family Pempheridae. Japan. J. Ichthyol., 12(3/6):33-56.
Perciformes: Percoidei: Bathyclupeidae 3211
BATHYCLUPEIDAE
Bathyclupeids
by J.R. Paxton
D iagnostic characters: Moderate-sized (to 30 cm) perciform fishes, body moderately to distinctly
elongate, very compressed. Head moderate to large, dorsal profile horizontal. Eye very large, its
diameter greater than snout length. Mouth large, oblique to almost vertical, jaws not reaching level
of anterior margin of pupil. Small teeth in bands on jaws and palatine, inconspicuous V-shaped patch on
vomer. Gill rakers lath-like, 15 to 19 on first gill arch. Fin spines very weakly developed; a single
short-based dorsal fin near middle of body and over middle of anal fin, with I spine and 8 to 10 soft
rays; anal fin with I spine and 24 to 39 soft rays; pelvic fins subjugular, anterior to level of pectoral-fin
base, very short, with I spine and 5 soft rays; pectoral fins very large, reaching level of dorsal-fin
origin, with 26 to 30 rays. Scales large, cycloid on body and nape, head naked; lateral-line scales with
several small pores. Total vertebrae 31 (Bathyclupea gracilis). Colour: dorsal dark, ventral silvery.
Habitat, biology, and fisheries: Deep-sea fishes of slope and oceanic waters, meso- and bathy-, or
benthopelagic; mature adults unknown. Carnivores, mostly feeding on small crustaceans. Rare deep-sea
fishes of no commercial importance.
Remarks: One genus with 7 nominal species restricted to tropical and subtropical latitudes in the world
ocean, except the eastern Pacific and northeastern Atlantic. The family requires revision; some of the
species listed below may be synonyms.
Clupeidae: no fin spines; pelvic fins behind level of pectoral-fin base.
Pempheridae: pelvic fins moderate in length, behind level of pectoral-fin base; dorsal-fin origin anterior to
anal-fin origin.
Clupeidae

Pempheridae
Bathyclupea gracilis Fowler, 1938
Bathyclupea malayana Weber, 1913
Bathyclupea megaceps Fowler, 1938
Reference
Dick, M.M. 1972. A review of the fishes of the family Bathyclupeidae. J. Mar. Biol. Ass. India, 14(2):539-544.
3212 Bony Fishes
TOXOTIDAE
Archerfishes
by G.R. Allen
D iagnostic characters: Oval or rhomboidal-shaped, moderately compressed, small to medium-sized

(to about 50 cm) percoid fishes; longitudinal axis from tip of snout to middle of caudal fin passing
through centre of eye. Eye large, about equal to snout length. Mouth moderately large, protractile, with
lower jaw protruding; angle of jaw oblique, about 35° to 40° to horizontal; maxilla slender, scaly and without
a supplemental bone. Villiform teeth on jaws, vomer, and palatines; a deep longitudinal groove on roof
of mouth, which is converted to a tube when tongue is pressed against it. First gill arch with 2 to 8
gill rakers on lower limb. A single dorsal fin with IV to VI stout spines and 15 to 18 soft rays; anal fin
with III spines and 15 to 18 soft rays; pelvic fins with I spine and 5 soft rays; pectoral fins with 11 to 15
rays; caudal fin truncate to slightly emarginate. Branchiostegal rays 7. Scales moderate to relatively large
and ctenoid, extending onto head and median fins; cheeks and operculum scaly; lateral-line scales 25 to
47; scale rows on body running horizontally. Colour: most species silvery white with a pattern of dark bars,
large spots, or irregular stripes on sides.
dorsal-fin origin dorsal fin with IV-V spines

far back and 11-14 soft rays
mouth
terminal

and 15-18 soft rays
Habitat, biology, and fisheries: Mangrove shores, brackish estuaries, and fresh waters, always in shallow
depths. Usually seen in small aggregations, but sometimes alone, close to the surface. These fishes exhibit
one of nature’s most remarkable feeding adaptations. When suitable prey is sighted, usually a small insect,
the fish rises to the surface and squirts a jet of water by forcefully compressing the gill cover. The water is
propelled through the tubular structure formed when the tongue is pressed tightly against a deep groove
on the roof of the mouth. The aim is uncannily accurate over a distance of at least 2 m. The victim is knocked
into the water from overhanging vegetation and summarily devoured. Archerfishes are sometimes seen in
markets and are an important component of artisanal fisheries in many areas, particularly where man-
groves are plentiful.

Pempheridae: snout blunt, dorsal-fin base short,
positioned above pectoral fins; anal fin long, usually
with more than 30 soft rays; occur in marine rather
than estuarine and fresh-water habitats.
anal fin very long based,

with more than 30 soft rays
Pempheridae
Perciformes: Percoidei: Toxotidae 3213
Key to the species of Toxotidae occurring in the area

1a. Dorsal-fin spines IV; 4 or 5 black bars on upper sides (Fig. 1) . . . . . . . . . . . Toxotes jaculatrix
(India to Vanuatu)
1b. Dorsal-fin spines usually V; colour variable with either bars, spots, or irregular stripes on
sides, or coloration uniform without markings . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Lateral-line scales usually less than 38 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3

2b. Lateral-line scales usually 30 to 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
3a. General coloration uniform brown or bronze to silvery grey without markings (may have
about 10 faint, narrow bars on upper sides in life); pectoral-fin rays usually 14 (occasion-
ally 15); lateral line running in a straight course; gill rakers on lower limb of first gill arch
2 to 4 (usually 3) (Fig. 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxotes lorentzi
(fresh water only; northern Australia and southern New Guinea)
3b. General coloration consisting of dark bars or relatively large spots on a light background;
pectoral-fin rays usually 13 (rarely 11, 12, or 14); lateral line arched over pectoral-fin
region; gill rakers on lower limb of first gill arch 5 to 8 . . . . . . . . . . . . . . . . . . . . . . . . ® 4
Fig. 1 Toxotes jaculatrix Fig. 2 Toxotes lorentzi
4a. Colour pattern consisting of 6 or 7, alternating large and small black spots (Fig. 3) . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxotes chatareus
(widespread, India to New Guinea and northern Australia)
4b. Colour pattern consisting of 4 or 5, wedge-shaped black bars or saddles without
intervening small spots (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxotes oligolepis
(fresh water only; Molucca Islands, Kimberley region of Western Australia, and possibly Philippines)
Fig. 3 Toxotes chatareus Fig. 4 Toxotes oligolepis

3214 Bony Fishes
5a. Colour pattern consisting of irregular horizontal stripes on a light background (Fig. 5)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Toxotes blythi
(fresh water only; Burma)
5b. Colour pattern consisting of 3 or 4 large dark spots or vertical bars along back with a
small spot on upper back between each one (Fig. 6) . . . . . . . . . . . . . . . . Toxotes microlepis
(fresh water only; Thailand, Sumatra, and Borneo)
Fig. 5 Toxotes blythi Fig. 6 Toxotes microlepis
List of marine-estuarine species occurring in the area

0
0
Toxotes chatareus (Hamilton, 1822)
Toxotes jaculatrix (Pallas, 1767)
Reference
Allen, G.R. 1978. A review of the archerfishes (family Toxidae). Rec. Aust. Mus., 6(4):355-378.
Perciformes: Percoidei: Toxotidae 3215
Toxotes chatareus (Hamilton, 1822)

En - Spotted archerfish; Fr - Poisson archer tacheté; Sp - Arquero manchado.
Maximum total length about 50 cm, very rarely exceeding 35 cm. Mangrove shores, brackish
estuaries, and fresh waters, always in shallow depths. More common in fresh water. In groups or
solitary. Feeds on terrestrial insects. Moderately important food fish in some areas; taken by
handline, block nets, and gill nets; marketed mostly fresh. Continental shores and archipelagos of
the tropical eastern Indian Ocean and western Pacific, from Sri Lanka and India to New Guinea and
northern Australia.
Toxotes jaculatrix (Pallas, 1767)

En - Banded archerfish; Fr - Poisson archer sellé; Sp - Arquero ensillado.
Maximum total length about 30 cm, commonly to about 20 cm. Mangrove shores and brackish
estuaries, always in shallow depths. In groups or solitary. Feeds on terrestrial insects. Moderately
important food fish in some areas where there are abundant mangroves; taken by handline, block
nets, and gill nets; marketed mostly fresh. Continental shores and archipelagos of the tropical
eastern Indian Ocean and western Pacific, from India to northern Australia, New Guinea, Solomon
Islands, and Vanuatu.
click for next page

3216 Bony Fishes
MONODACTYLIDAE
Moonfishes
by M. Kottelat
D iagnostic characters: Body oval,

deep, strongly compressed (size to
about 25 cm). Eye moderately large, its
diameter longer than snout length. Mouth
small and oblique. Jaws with bands of
small conical teeth; vomer, palatines, and
tongue with granular teeth. Gill mem-
branes free from isthmus, separate. Bran-
chiostegals 6. Dorsal fin with V to VIII
spines and 26 to 31 soft rays; anterior
soft dorsal-fin rays elongated, situated
over midlength of body or somewhat in
front, in a single species at least as long
as 1/2 body depth. Anal fin with III spines
and 26 to 31 soft rays; anterior soft anal-
fin rays elongated, situated below
midlength of body or somewhat in front,
longer than elongated soft dorsal-fin rays.
Caudal fin truncate to forked. Body, head,
and unpaired fins covered by small, de-
ciduous scales. Colour: body and head
silvery, in most species with 1 or more
black vertical bar through eye, in front of
pectoral-fin base (usually continuous with
black margin along anterior edge of anal
fin) and on body; tip of dorsal and anal fins
black in most species.
Habitat, biology, and fisheries: Occur in shoals in estuaries, lagoons, and over shallow coral reefs, in
brackish and sea water, sometimes in fresh water. Minor commercial importance, occasionally sold fresh
in local markets. Some are caught for the aquarium-fish trade.
Remarks: Some authors place Schuetta in the Monodactylidae, but Tominaga (1968) considered it a
separate family. A single species has classically been recognized in the area, Monodactylus argenteus.
M. kottelati has recently been discovered in Sri Lanka and Laccadive Archipelago (India); this species
appears to be much rarer. In Sri Lanka, both species occur in sympatry, with M. kottelati occupying deeper
areas with higher tidal flow. More than 1 species apparently occur around Sumatra and one of them has
been identified as M. kottelati on the basis of juveniles; this should be confirmed with examination of adult
specimens.
Drepanidae, Ephippidae: pelvic fins well developed; gill membrane attached to isthmus.
Drepanidae Ephippidae
Perciformes: Percoidei: Monodactylidae 3217
Chaetodontidae, Scatophagidae: pelvic fins well developed; dorsal fin with X to XVII spines; spinous part
of dorsal fin well separated from soft-rayed part.
Chaetodontidae Scatophagidae
Carangidae: caudal fin forked; anal fin with II spines.

Menidae: ventral profile conspicuously more convex than dorsal profile; caudal fin forked; dorsal fin without
spines in adults; first 2 pelvic-fin rays elongated.
Carangidae Menidae
Key to the species of Monodactylidae occurring in the area

1a. Pelvic fins normally developed; caudal fin forked . . . . . . . . . . . . . . . Schuettea scalaripinnis
1b. Pelvic fins rudimentary or absent in adults; caudal fin truncate to slightly emarginate . . . . . . . ® 2
2a. Body depth 1.2 to 1.6 times in standard length; distance between tips of dorsal and anal
fins 0.8 to 0.9 times in standard length; caudal fin emarginate; posterior edge of dorsal
and anal fins distinctly concave . . . . . . . . . . . . . . . . . . . . . . . . Monodactylus argenteus
2b. Body depth 1 to 1.1 times in standard length; distance between tips of dorsal and anal
fins 0.6 to 0.7 times in standard length; caudal fin truncate to very slightly emarginate;
posterior edge of dorsal and anal fins almost straight . . . . . . . . . . . . . Monodactylus kottelati

0
The symbol is given when species accounts are include.
0 Monodactylus argenteus (Linnaeus, 1758)
0 Monodactylus kottelati Pethiyagoda, 1991
0 Schuettea scalaripinnis Steindachner, 1866
References
Pethiyagoda, R. 1991. Monodactylus kottelati, ein neues Flossenblatt aus Sri Lanka (Pisces: Monodactylidae). Aquar.
Terrar. Zeitschr., 44:162-167.
Tominaga, Y. 1968. Internal morphology, mutual relationships and systematic position of the fishes belonging to the
family Pempheridae. Japan. J. Ichthyol., 15:43-95.
3218 Bony Fishes
Monodactylus argenteus (Linnaeus, 1758)

FAO names: En - Silver moony; Fr - Lune d’argent; Sp - Ramball plateado.
(after Pethiyagoda, 1991)
Diagnostic characters: Body oval, deep, strongly compressed. Body depth 1.2 to 1.6 times in
standard length; distance between tips of dorsal and anal fins 0.8 to 0.9 times in standard length.
Eye moderately large, its diameter longer than snout length. Mouth small and oblique. Jaws with bands of
small conical teeth. Gill membranes free from isthmus, separate. Dorsal fin with VII or VIII spines and 26
to 31 soft rays; anterior soft dorsal-fin rays elongated, situated over midlength of body. Anal fin with III
spines and 26 to 31 soft rays; anterior soft anal-fin rays elongated, situated below midlength of body,
longer than elongated soft dorsal-fin rays. Posterior edge of dorsal and anal fins distinctly concave.
Caudal fin slightly emarginate. Pelvic fins rudimentary or absent in adults. Body, head, and unpaired
fins covered by small, deciduous scales. Colour: adults silvery, tip of dorsal and anal fins dusky; juveniles
silvery, with a curved black vertical bar through eye and 1 in front of pectoral-fin base; tip of dorsal and
anal fins black; anterior edge of anal fin with broad black margin.
Habitat, biology, and fisheries: Inhabits estuaries and lagoons, usually in shallow water, often in schools,
with a preference for around and under floating or sunken logs and branches. Juveniles occasionally enter
rivers but are rarely recorded from fresh waters. Sold fresh in local markets. Juveniles are caught for the
aquarium-fish trade.
the tropical Indo-West Pacific,
from South Africa and Red
Sea to Australia, Samoa, and
southern Ryukyu Islands.
Perciformes: Percoidei: Monodactylidae 3219
Monodactylus kottelati Pethiyagoda, 1991

Frequent synonyms / misidentifications: None / Monodactylus argenteus (Linnaeus, 1758).
FAO names: En - Angel mono; Fr - Monodactyle voile.
Diagnostic characters: Body oval, very
deep, strongly compressed. Body depth
1 to 1.1 times in standard length; dis-
tance between tips of dorsal and anal
fins 0.6 to 0.7 times in standard length.
Eye moderately large, its diameter longer
than snout length. Mouth small and oblique.
Jaws with bands of small conical teeth. Gill
membranes free from isthmus, separate.
Dorsal fin with VIII spines and 28 to 30 soft
rays; anterior soft dorsal-fin rays very
elongated, situated over midlength of
body. Anal fin with III spines and 28 to 30
soft rays; anterior soft anal-fin rays very
elongated, situated below midlength of
body, longer than elongated soft dorsal-fin
rays. Posterior edge of dorsal and anal
fins almost straight. Caudal fin truncate
to very slightly emarginate. Pelvic fins
rudimentary or absent in adults. Body,
head, and unpaired fins covered by small,
deciduous scales. Colour: adults silvery,
dorsal and anal fins dusky, blackish at the
tip; juveniles silvery, with a curved black
vertical bar through eye and one in front of
pectoral-fin base; dorsal and anal fins
blackish, denser black at the tip; anterior
edge of anal fin with broad black margin.
Size: Largest known specimen 10 cm total
length, but probably reaches a larger size.
Habitat, biology, and fisheries: Inhabits
estuaries and lagoons, usually in water
deeper than 5 m, often in schools, often
around sunken logs and branches; not
known to tolerate fresh waters. Apparently
unknown in local markets. Juveniles are (after Pethiyagoda, 1991)
caught for the aquarium-fish trade.
Distribution: Definitively
known only from a single lo-
cality in Sri Lanka and the Lac-
cadive Archipelago (India).
Juveniles specimens from
Medan (Sumatra) are tenta-
tively referred to this species.
3220 Bony Fishes
Schuettea scalaripinnis Steindachner, 1866

FAO names: En - Eastern pomfred.
(after Steindachner, 1866)
Diagnostic characters: Body oval, moderately deep, strongly compressed. Body depth about 2
times in standard length. Eye moderately large, its diameter longer than snout length. Mouth small and
oblique. Jaws with bands of small conical teeth. Gill membranes free from isthmus, separate. Dorsal fin
with V spines and 28 to 30 soft rays; anterior soft dorsal-fin rays elongated, situated in front of midlength
of body. Anal fin with III spines and 28 to 30 soft rays; anterior soft anal-fin rays elongated, situated in
front of midlength of body. Caudal fin forked. Pelvic fins normally developed, with I spine and 5 soft
rays. Body, head, and unpaired fins covered by small, ctenoid scales. Colour: adults silvery, yellow along
back; juveniles silver; fins yellow.
Habitat, biology, and fisheries: Inhabits large estuaries close to the sea and coastal bays. Schooling.
Distribution: Along the
coasts of New South Wales
and southern Queensland,
Australia.
Perciformes: Percoidei: Drepanidae 3221
DREPANIDAE
Sicklefishes
by P.C. Heemstra
D iagnostic characters: Body oval and strongly

compressed (size to about 50 cm). Interorbital,
preorbital, and broad preopercular flange naked; oper-
cle scaly dorsally; ventral edge of preopercle serrate in
juveniles. Mouth highly protrusible, forming a
downward-pointing tube when protruded; jaws with
bands of setiform teeth; no teeth on roof of mouth;
maxilla exposed posteriorly, no supramaxilla. Bran-
chiostegal membranes joined to isthmus; bran-
chiostegal rays 6. A single dorsal fin, with VIII to X
spines (third or fourth spines longest) and 19 to 22
soft rays; anal fin with III spines and 16 to 19 soft
rays; caudal fin rounded or bluntly wedge-shaped (al-
most truncate in large adults); pectoral fins elongate,
falciform, reaching caudal peduncle, with 16 to 18
rays; pelvic fins with I spine, 5 soft rays, and a fleshy
axillary process; first pelvic-fin ray elongate. Scales
small, finely ctenoid, extending onto top of head and
base of median fins; lateral line complete, strongly
curved over pectoral fins; lateral-line scales 48 to 55.
Vertebrae 10+14. Swimbladder large. Pyloric caeca 2
or 3. Large adults with a bump or bony knob on interorbital region, a result of hyperostosis of the frontal
bones. Colour: head and body silvery; sides of body with vertical rows of dark spots or lines.
Habitat, biology, and fisheries: Frequents a variety of habitats (sand or mud bottoms and coral reefs) in
shallow water, including estuaries and harbours. Feeds on benthic invertebrates (mainly crustaceans and
worms). Mostly caught with trawls.
Remarks: Three species are known for this family; 2 species occur in the area.
Chaetodontidae and Pentacerotidae: mouth not highly protrusible; pectoral fins not
much longer than head and well short of caudal peduncle; no notch between
spinous and soft-rayed parts of dorsal fin.
Ephippidae and Scatophagidae: mouth not highly protrusible; pectoral fins not
reaching past anal-fin base.
Monodactylidae: mouth not highly protrusible; pectoral fins shorter than head; eye
centred on horizontal axis through mouth. Chaetodontidae
Pentacerotidae Ephippidae Scatophagidae Monodactylidae

0
0 Drepane longimana (Bloch and Schneider, 1801)
0 Drepane punctata (Linnaeus, 1758)
Reference
Lloris, D. and J. Rucabado. 1987. Revisión sistematica y distribución de la familia Drepanidae (Pisces, Osteichthyes).
Misc. Zool., 11:277-288.
3222 Bony Fishes
Drepane longimana (Bloch and Schneider, 1801)

Frequent synonyms / misidentifications: None / Drepane punctata (non Linnaeus, 1758).
FAO names: En - Sicklefish.
o v a l a n d s t r o n g l y c o m-
times in standard length and 2
to 3 times in head length. Snout
profile straight or concave; pre-
orbital deep, the eye well
above horizontal line through
upper jaw symphysis. Interor-
bital, preorbital and broad pre-
opercular flange naked;
opercle scaly dorsally; ventral
edge of preopercle serrate in
juveniles; a fringe of cirri just
behind chin. Mouth highly
protrusible, forming a down-
ward-pointing tube when
protruded; jaws with bands of
setiform teeth; no teeth on roof
of mouth; maxilla exposed pos-
teriorly, no supramaxilla. First
gill arch with 6 to 8 gill rakers
on upper limb, 10 to 12 on
lower limb. A single dorsal fin,
with VIII or IX (usually VIII)
spines (fourth spine longest)
and 19 to 23 soft rays; anal
fin with III spines and 17 to
1 9 so f t r a ys; c a u d a l f in
rounded or bluntly wedge-
shaped (almost truncate in
large adults); pectoral fins
elongate, falciform, reaching caudal peduncle, with 16 to 18 rays. Lateral-line scales 46 to 55. Large
adults with a bump or bony knob on interorbital region, a result of hyperostosis of the frontal bones. Colour:
head and body silvery; 4 to 10 subvertical dark bars usually visible on dorsal part of body from head
to caudal-fin base (bars faint or absent in some specimens); median fins dusky, darker distally; 2 or 3
longitudinal rows of small dark spots (one on each interradial membrane) on soft dorsal fin; some specimens
with front part of head (interorbital, snout, cheeks, and lower jaw) dark, and lips white.
Habitat, biology, and fisheries: Occurs in a variety of inshore habitats, such as sand or mud bottoms,
reefs, estuaries, and harbours. Biology little known. Mostly caught with trawls, occasionally with hook-and-
line.
Distribution: Temperate and
tropical waters of the Indo-
West Pacific region from
South Africa to Red Sea,
Oman, Arabian Sea, India to
n o r t h e r n A u st r al i a , N e w
Guinea, Indonesia, Philip-
pines, Taiwan Province of
China, and Japan.
Perciformes: Percoidei: Drepanidae 3223
Drepane punctata (Linnaeus, 1758) SPS

Frequent synonyms / misidentifications: None / Drepane longimana (non Bloch and Schneider, 1801) .
FAO names: En - Spotted sicklefish; Fr - Forgeron tacheté; Sp - Catemo manchado.
o v a l a n d s t r o n g ly c o m-
times in standard length and 2 to
3 times in head length. Snout
profile straight or concave; pre-
orbital deep, the eye well above
horizontal line through upper jaw
symphysis. Interorbital, preorbi-
tal, and broad preopercular
flange naked; opercle scaly dor-
sally; ventral edge of preopercle
serrate in juveniles; a fringe of
cirri just behind chin. Mouth
highly protrusible, forming a
d o w n w a rd - p o i n t i n g t u b e
when protruded; j a w s w ith
bands of setiform teeth; no teeth
on roof of mouth; maxilla ex-
posed posteriorly, no supramax-
illa. First gill arch with 5 gill
rakers on upper limb, 10 or 11 on
lower limb. A single dorsal fin,
with VIII to X (usually IX)
spines (third or fourth spines
longest) and 20 to 22 soft rays;
anal fin with III spines and 16
to 19 soft rays; caudal fin
rounded or bluntly wedge-
shaped (almost truncate in large
adults); pectoral fins elongate,
falciform, reaching caudal
peduncle, with 16 to 18 rays. Lateral-line scales 46 to 50. Large adults with a bump or bony knob on
interorbital region, a result of hyperostosis of frontal bones. Colour: head and body silvery; dorsal part of
body from below dorsal fin to caudal peduncle with 5 to 10 series of black spots arranged in vertical
lines; fins dusky yellow; dorsal and caudal fins darker distally; 2 or 3 longitudinal rows of small dark spots
(1 on each interradial membrane) on soft dorsal fin.
Habitat, biology, and fisheries: Occurs in a variety of inshore habitats, such as sand or mud bottoms,
reefs, estuaries, and harbours. Biology little known. Mostly caught with trawls, occasionally with hook-and-
line. From 1990 to 1995, the FAO Yearbook of Fishery Statistics reports a range of yearly catch of around
580 to 950 t of Drepane punc-
tata from the Western Central
P a c ific (Malaysia, Philip-
pines).
tropical waters from India to
northern Australia, Indonesia,
New Guinea, Philippines, Tai-
wan Province of China, and
Japan.
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3224 Bony Fishes
CHAETODONTIDAE
Butterflyfishes
by R. Pyle
D iagnostic characters:
Oval-shaped (excluding
fins), deep-bodied, highly
compressed percoid fishes
(size to about 30 cm). Head
length roughly equal to head
height; preopercle smooth,
without prominent spines.
Eye moderately small, located
just on or slighly above lon-
gitudinal axis from tip of
snout to middle of caudal fin.
Snout length highly variable,
ranging from very short to ex-
tremely elongate (Forcipiger).
Mouth small, terminal, protrac-
tile, gape not extending to an-
terior rim of orbit. Teeth
bristle-like, curved, arranged in
rows or bands across jaws;
vomer and palatines without
teeth. Pseudobranchiae pre-
sent; 6 or 7 branchiostegal
rays. Gill rakers short, ranging
from 9 to 25 in number. A sin-
gle dorsal fin, continuous
and relatively smooth in most species; usually with X to XIV strong, stout spines (but as few as VI in
Parachaetodon and as many as XV or XVI in some species of Chaetodon and Hemitaurichthys), first few
to several interspinous membranes deeply incised, species of Heniochus with extremely elongate and
filamentous fourth dorsal-fin spine; no notch between spinous and soft dorsal fin; soft dorsal fin with 15 to
30 branched rays, some species with short horizontal filamentous extension of 1 or more soft dorsal-fin
rays at dorsoposterior margin of fin. Anal fin with III to V strong, stout spines (usually III), interspinous
membranes deeply incised; soft anal fin with 14 to 23 branched rays, margin usually rounded but
sometimes angular. Pectoral fins transparent with 13 to 15 soft rays. Pelvic fins with I stout spine and 5
branched rays; scaly axillary process at upper base of pelvic fins. Caudal fin rounded to slightly
emarginate, usually with 15 branched rays. Scales ctenoid, covering head, body, and median fins; largest
in centre of body, smaller on head, thorax, belly, caudal peduncle, and median fins; number of lateral-line
scales variable, ranging from 20 to 90 (depending on the genus). Vertebrae 11+13. Supraoccipital crest,
predorsal bones, and first dorsal pterygiophore articulated. Colour: most species very brightly coloured
with complex and varied colour patterns; most frequent colours include shades of yellow, orange, black,
and white; patterns usually include broad vertical bars, thin vertical stripes, cross-hatching patterns, spots,
and dark soft dorsal- and anal-fin margins; almost without exception, the eye is obscured within a darkly
coloured vertical ocular bar or dark region on the head; many species with a dark occellate
“false-eye” spot on the posterior portion of the body.
Habitat, biology, and fisheries: Mostly on shallow coral reefs, but also common in lagoons and outer reef
slopes to depths of 60 m or more. Species of the genera Roa and Prognathodes, and of the Chaetodon
subgenus Roaops usually restricted to depths in excess of 60 m. Juveniles of many species frequently
found in estuaries. Most species closely associated with the substratum, feeding on corals and other marine
invertebrates; some species (particularly species of Hemitaurichthys) aggregate in midwater and feed on
plankton. Although sometimes harvested as food fishes, the primary fishery value of chaetodontids is
through the ornamental marine aquarium trade, where they are the third most frequently exported fish by
number, and second highest in total value of all families of aquarium fishes in trade.
Remarks: Dichotomous keys are seldom used for identifying species of chaetodontids, as species in this
familiy exhibit highly conspicuous and distinctive colour patterns (the primary components of which usually
remain evident in preservative). Nevertheless, keys to genera and species of chaetodontids are included
here for purposes of consistency and clarity. Keys are more reliable when used for adult specimens.
Perciformes: Percoidei: Chaetodontidae 3225

Pomacanthidae: similar in general body shape and colour pattern to Chaetodontidae, but easily distin-
guished in having at least 1 prominent spine at the angle of preopercle. Also, Pomacanthidae lack the scaly
axillary process at the base of pelvic fins, lack the ocular bands and false-eye spots (except for a few
species), do not have produced snouts, and have 16 to 21 pectoral-fin rays.
Acanthuridae: similar general body shape and some species with bright colours, but can easily be
distinguished by the presence of a fixed or retractable sharp spine on the caudal peduncle, and typically
have fewer dorsal-fin spines (IV to IX) than most species of Chaetodontidae.
Acanthuridae
Pomacanthidae
Scatophagidae: distinguished from Chaetodontidae

by a deep notch between the spinous and soft
portion of the dorsal fin, and in having IV anal-fin
spines.
Zanclidae: the monotypic Zanclus cornutus closely
resembles some species of Heniochus, but is easily
distinguished by a strongly produced snout, bony
supraocular projections (in adults), and only VII
dorsal-fin spines.
Microcanthidae: a single species, Microcanthus
strigatus, likely to be confused with Chaetodonti-
dae; it is most easily distinguished by its distinctive
horizontally striped colour pattern.
Zanclidae
Scatophagidae Microcanthidae
3226 Bony Fishes
Key to the genera of Chaetodontidae occurring in the area

Note: species names are given when a genus is represented by a single species in the area.
1a. Lateral line incomplete, ending in vicinity of last rays of dorsal fin . . . . . . . . . . . . . . . . . ® 2
1b. Lateral line complete, ending at base of caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
2a. Dorsal-fin spines VI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parachaetodon ocellatus

2b. Dorsal-fin spines X to XVI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Third to fifth dorsal-fin spines not distinctly longer than others; body lacking vertical bars,
more than 2 vertical bars on body, or bars not extending entire depth of body . . . . . . . Chaetodon
3b. Third to fifth dorsal-fin spines distinctly longer than others; body colour pale to white with
2 broad dark vertical bars extending entire depth of body . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Snout length 2.7 to 3.7 times in head length; dark bars on body orangish brown;
prominent black ocellate spot on soft dorsal fin; ocular bar continuing ventral to eye . . Roa modestus
4b. Snout length 2.5 times in head length; dark bars on body black; no ocellate spot on soft
dorsal fin; ocular bar not continuing ventral to eye, instead extending anteriorly along
snout . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Prognathodes guezei
5a. Fourth dorsal-fin spine elongate to filamentous; supraorbital of adults with spines or
horns; nape of adults usually with hump or strong bony projection . . . . . . . . . . . . . Heniochus
5b. Fourth dorsal-fin spine normal or only slightly elongate; no horns on supraorbital;
normally no hump or projection at nape (some species with slight prominence or hump
on nape but fourth dorsal-fin spine normal) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. More than 65 scales in lateral line; pectoral fins elongate, falcate . . . . . . . . . . . . . . . . . ® 7
6b. Less than 60 scales in lateral line; pectoral fins not elongate, rounded . . . . . . . . . . . . . . ® 8
7a. Snout not elongate, not tubular . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemitaurichthys

7b. Snout elongate, tubular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forcipiger
8a. Normally XII dorsal-fin spines . . . . . . . . . . . . . . . . . . . . . . . Amphichaetodon howensis

8b. Normally VIII to X dorsal-fin spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
9a. Snout elongate, tubular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelmon

9b. Snout short, not tubular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coradion
Key to the species of Chaetodon occurring in the area

1a. Anal-fin spines IV (rarely V) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Anal-fin spines III . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Lateral-line scales 22 to 29; gill rakers 23 to 27; body elongate (depth 1.7 to 2.2 times in
standard length), pale with dark forward-facing chevron markings; dorsal and anal fins
orangish yellow; soft dorsal fin pointed at dorsoposterior margin . . . . . . . . Chaetodon trifascialis
2b. Lateral-line scales 36 to 41; gill rakers 19 to 24; body oval (depth 1.6 to 1.7 times in
standard length), yellow with longitudinal orange line; prominent black ocellate spot at
caudal peduncle; prominent longitudinal blue blotch just dorsal to centre of body . Chaetodon plebius
3a. Dorsal-fin spines XI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4

3b. Dorsal-fin spines XII or more . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
4a. Dorsal-fin rays 24 or 25; anal-fin rays 20 to 22 . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5

4b. Dorsal-fin rays 19 to 22; anal-fin rays 16 to 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5a. White or off-white chevron marking on caudal fin . . . . . . . . . . . . . . . . Chaetodon triangulum

5b. No such marking on caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon baronessa
6a. Dorsal-fin rays 19; anal-fin rays 15 to 17; 8 narrow black bars on head, body, and fins
(usually in sets of 2) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon octofasciatus
6b. Dorsal-fin rays 21 or 22; anal-fin rays 17 or 18; body with some bar patterns, but without
narrow black bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Body with 2 prominent orange-margined bars in addition to a single narrow orange bar
just posterior to operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon rainfordi
7b. Body mostly uniform in colour except for ocular bar and a single narrow orange bar just
posterior to operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon aureofasciatus
8a. Body yellow or whitish yellow with a single prominent dark spot or blotch just dorsal to
centre of body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9
8b. Body not yellow or whitish yellow, or lacking a dark spot, single prominent dark spot, or
blotch just dorsal to centre of body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
9a. Dorsal-fin rays 21 to 23; anal-fin rays 19 . . . . . . . . . . . . . . . . . . . Chaetodon unimaculatus

9b. Dorsal-fin rays 16 or 17; anal-fin rays 15 or 16 . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Prominent spot on side of body ocellate; 2 pale blue curved lines on side of body, 1 above
pectoral fins, 1 below, converging at or near origin of lateral line; ocular bar dark orange
with pale blue margins; usually 15 (rarely 14 or 16) pectoral-fin rays . . . . . . . Chaetodon bennetti
10b. Prominent spot not ocellate; no pale lines on side of body; ocular bar dark brownish black
and without margins; usually 14 (rarely 13) pectoral-fin rays . . . . . . . . . . Chaetodon speculum
11a. Snout not produced; head profile around mouth smooth and/or rounded . . . . . . . . . . . . . ® 12
11b. Snout slightly to moderately produced; head profile around mouth angular . . . . . . . . . . . ® 15
12a. Dorsal-fin spines XIII, dorsal-fin rays 20 to 22 (usually 21); lateral-line scales 30 to 39;
scale rows above lateral line to base of dorsal fin 4 to 6 . . . . . . . . . . . . Chaetodon trifasciatus
12b. Dorsal-fin spines XII, dorsal-fin rays 23 to 28; lateral-line scales 45 to 55; scale rows
above lateral line to base of dorsal fin 7 to 11 . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 13
13a. Body colour pattern reticulated, a large whitish spot on each scale; pelvic fins black
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon reticulatus
13b. Body colour pattern consisting of lines or stripes, not reticulated or spotted; pelvic fins
light . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
14a. Dorsal-fin rays 23 or 24; anal-fin rays 19 or 20 . . . . . . . . . . . . . . . . . . . Chaetodon meyeri

14b. Dorsal-fin rays 26 to 28; anal-fin rays 21 to 23 . . . . . . . . . . . . . . . . Chaetodon ornatissimus
15a. Body with a relatively straight, sharp colour demarcation extending diagonally from near
origin of dorsal fin ventroposteriorly to middle of soft anal fin; colour ventroanterior to
demarcation mostly white or very pale yellow, usually with black spot on most of scales;
colour dorsoposterior to demarcation black or brownish orange . . . . . . . . . . . . . . . . . ® 16
3228 Bony Fishes
16a. Body colour dorsoposterior to demarcation brownish orange . . . . . . . . . . . Chaetodon declivis

16b. Body colour dorsoposterior to demarcation black . . . . . . . . . . . . . . . . . . . . . . . . . ® 17
17a. No prominent band across nape . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon tinkeri

17b. Prominent black or yellow band across nape . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18
18a. Nape band black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon burgessi

18b. Nape band yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon flavocoronatus
19a. Scale rows below lateral line to origin of anal fin 24 to 30 . . . . . . . . . . . . . . Chaetodon smithi
19b. Scale rows below lateral line to origin of anal fin 11 to 21 . . . . . . . . . . . . . . . . . . . . . ® 20
20a. Prominent bump on nape; body colour uniformly dark . . . . . . . . . . . . . Chaetodon flavirostris
20b. No prominent bump on nape or slight bump, but body colour not uniformly dark . . . . . . . . . ® 21
21a. Most of body colour grey except for the dorsoposterior quadrant and soft dorsal fin, which
is black, black region separated from grey region by a broad white margin; snout, head
beneath eye, and thorax bright orange in life . . . . . . . . . . . . . . . . . . Chaetodon ephippium
21b. Colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
22a. No distinct ocular band (except in very small juveniles) . . . . . . . . . . . . . . Chaetodon nippon
22b. Distinct ocular band, either partial or complete . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23
23a. Proximal portion of caudal fin and posterior portion of caudal peduncle white . . . . . . . . . . ® 24
23b. Caudal fin colour not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 26
24a. Distal portion of caudal fin black and transparent white; broad black bar on posterior
portion of body; ocular band reduced to a small black spot above the eye . . . Chaetodon argentatus
24b. Distal portion of caudal fin orange (except for a thin transparent margin); broad orange
bar on posterior portion of body; ocular band extending ventral to the eye . . . . . . . . . . . . ® 25
25a. Scales on body white with black margins; distinct black spot with white margin on nape;
broad orange bar on posterior portion of body not reaching spinous portion of dorsal fin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon xanthurus
25b. Body white with black chevrons; spot on nape absent or faint with no distinct white
margin; broad orange bar on posterior portion of body reaches spinous portion of dorsal
fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon mertensii
26a. Ocular bar orange with black margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 27

26b. Ocular bar black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 30
27a. Dorsal half of body dark brown with 2 prominent white spots . . . . . . Chaetodon quadrimaculatus
27b. Body colour pale with dark spots or bands . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 28
28a. Caudal peduncle entirely orange; a dark spot at centre of each scale, but no obvious
vertical or diagonal bands on body, ocular bar extending vertically to origin of dorsal fin
and joined across nape . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon guttatissimus
28b. Caudal peduncle not entirely orange; body with vertical or diagonal parallel bands, ocular
bar not joined across nape and not reaching dark spot at origin of dorsal fin . . . . . . . . . . . ® 29
29a. Dark parallel bands on body diagonal, extending dorsoposteriorly from midbody to base
of soft dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon pelewensis
29b. Dark bands on body vertical . . . . . . . . . . . . . . . . . . . . . . . Chaetodon punctatofasciatus
30a. Thin white band on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 31

30b. No thin white band on caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 32
31a. Pectoral-fin rays 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon kleinii

31b. Pectoral-fin rays 15 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon trichrous
32a. Caudal peduncle black, or with black spots or markings . . . . . . . . . . . . . . . . . . . . . . ® 33

32b. Cadual peduncle orange or yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 41
33a. Body ground colour mostly brownish yellow . . . . . . . . . . . . . . . . . . . . . Chaetodon lunula

33b. Body ground colour mostly white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 34
34a. Body with a series of faint yellow dots in parallel diagonal rows extending dorsoposteri-
orly across body; no vertical or diagonal black lines on body . . . . . . . . . . . . Chaetodon selene
34b. No series of faint yellow dots in parallel diagonal rows; distinct vertical or diagonal lines
on body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 35
35a. Dorsal-fin rays 18 to 20, anal-fin rays 17 or 18 . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 36

35b. Dorsal-fin rays 22 to 27, anal-fin rays 18 to 22 . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 37
36a. Black diagonal lines on body joining a prominent black crescent band at base of soft
dorsal fin that extends onto caudal peduncle; lateral-line scales 33 to 39; scale rows
above lateral line to origin of dorsal fin 6 or 7; scale rows below lateral line to origin of
anal fin 15 to 17 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon melannotus
36b. No prominent black crescent at base of soft dorsal fin, a distinct black (usually ocellate)
spot on caudal peduncle; lateral-line scales 29 to 34; scale rows above lateral line to
origin of dorsal fin 5 or 6; scale rows below lateral line to origin of anal fin 13 or 14 . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon ocellicaudus
37a. Dorsal-fin spines XIII; black lines on body diagonal in 2 sets perpendicular to each other . . . . ® 38
37b. Dorsal-fin spines XII; black lines on body vertical in a single set . . . . . . . . . . . . . . . . . ® 39
38a. Soft portions of dorsal and anal fins mostly yellow (with some thin black lines) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon vagabundus
38b. Soft portions of dorsal and anal fins mostly black (with some thin yellow lines) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Chaetodon decussatus
39a. Two large black saddle-shape markings on side of body . . . . . . . . . . . . Chaetodon ulietensis
39b. A single large black crescent or diagonal band at base of soft dorsal fin . . . . . . . . . . . . . ® 40
40a. Large black band at base of soft dorsal fin extends well into base of soft anal fin; ocular
bar continuous and connected across nape, a white spot within it at nape . . . Chaetodon lineolatus
40b. Large black band at base of soft dorsal fin does not extend below caudal peduncle;
ocular bar discontinuous, a separate prominent black spot on nape . . . . . Chaetodon oxycephalus
41a. Dorsal-fin spines XII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon collare

41b. Dorsal-fin spines XIII or XIV (rarely XII) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 45
3230 Bony Fishes
42a. Body ground colour white or dirty white . . . . . . . . . . . . . . . . . . . . Chaetodon adiergastos

42b. Body ground colour brown or brownish yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 43
43a. Body colour dark brown with light brown spots on each scale; caudal fin bright orange;
pelvic fins black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon collare
43b. Body colour brownish yellow with dull orange horizontal or diagonal lines; caudal fin
brownish yellow with or without a prominent black bar; pelvic fins brownish yellow . . . . . . . ® 44
44a. Lines on body diagonal, extending dorsoposteriorly across body; ocular bar broad and
short, separated from a prominent black nape marking by a broad white area; prominent
black bar on caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon weibeli
44a. Lines on body horizontal, reduced to rows of ascending spots anteriorly above lateral
line; ocular bar vertical, relatively narrow, and complete; no prominent black bar on
caudal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon auripes
46a. Body white with 2 sets of diagonal black lines perpendicular to each other, ocular bar
broad ventral to eye and faint or absent dorsal to eye; prominent black spot on soft dorsal
fin of juveniles and adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodon auriga
46b. Body yellow or pale yellow with black dots or dark scale margins; ocular bar narrow both
dorsal and ventral to eye; no black spot on soft dorsal fin of adults . . . . . . . . . . . . . . . . ® 47
47a. Prominent black bar on caudal fin; scales on body with dark margins; scale rows above
lateral line to origin of dorsal fin 4 or 5 . . . . . . . . . . . . . . . . . . . . . . . Chaetodon rafflesii
47b. No black bar on caudal fin; a single dark spot in the centre of each body scale (but no
dark margins); scale rows above lateral line to origin of dorsal fin 6 to 8 . . . . . . . . . . . . . ® 48
48a. Broad black crescent bands at base of soft dorsal and soft anal fins; nape blue; a long
filament extending posteriorly from soft dorsal fin of adults . . . . . . . . . . . . Chaetodon semeion
48b. No broad black crescent bands at base of soft dorsal or soft anal fins; no blue on nape;
no filament on soft dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 49
49a. Dorsal-fin spines XIII, thin black submarginal line on entire edge of anal fin . . Chaetodon guentheri
49b. Dorsal-fin spines XIV, broad black margin on ventral edge of anal fin . . . . . Chaetodon citrinellus
Key to the species of Chelmon occurring in the area

1a. Snout short, 2.6 to 3 times in head length, 4.8 to 4.9 times in body depth; central body
bar broad, brownish in preservative; pelvic fins black; dorsal fin with IX spines and 26
soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelmon muelleri
1b. Snout longer, 1.7 to 2.1 times in head length, 2.4 to 3.4 times in body depth; central body
bar not broad, light with dark borders or absent; barred pattern continuing onto pelvic
fins, part greyish to white, part yellowish or hyaline . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Central body bar absent; spot in soft portion of dorsal fin normally faint, not ocellate;
snout length 3.1 to 3.4 times in body depth; interorbital width 4.4 to 4.8 times in head
length; caudal-peduncle depth 8.5 to 9.1 times in standard length; dorsal fin with IX
spines and 30 or 31 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelmon marginalis
2b. Central body bar evident; spot in dorsal fin obvious, ocellate; snout length 2.4 to 3.1 times
in body depth; interorbital width 4.4 to 4.8 times in head length; caudal-peduncle depth
9.1 to 11.3 times in standard length; dorsal fin with IX spines and 28 or 29 soft rays . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chelmon rostratus
Key to the species of Coradion occurring in the area

1a. Dorsal fin with VIII spines and 31 or 32 soft rays; soft dorsal fin elevated; median snout
stripe ending at upper lip; eyelids not continuous on isthmus; dorsal-fin ocellus absent
(sometimes faded spot evident); never an ocellus in anal fin . . . . . . . . . . . . Coradion altivelis
1b. Dorsal fin with IX or X spines and less than 30 soft rays . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Dorsal fin with IX spines and 28 or 29 soft rays; soft dorsal fin rounded, not elevated;
median snout stripe not continuous on chin as stripe (although spot or blotch may be
present); eyebands joined at isthmus and extended toward, but do not reach, insertion
of pelvic fins; dorsal fins always with ocellated spot; anal fin without ocellated spot . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coradion chrysozonus
2b. Dorsal fin with X spines and 24 to 27 (usually 25) soft rays; soft dorsal fin squarish in
shape; median snout stripe continuous on chin; eyebands not continuous on isthmus;
both dorsal and anal fins with well-developed ocellus . . . . . . . . . . . . . . Coradion melanopus
Key to the species of Forcipiger occurring in the area

1a. Dorsal fin with XII spines; lateral-line scales 74 to 80; snout length 1.6 to 2.1 times in
body depth; mouth opening slightly gaping; ventroposterior margin of operculum curved
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forcipiger flavissimus
1b. Dorsal-fin spines XI; lateral-line scales 66 to 75; snout length 1.1 to 1.5 times in body
depth; mouth opening very small; ventroposterior margin of operculum angular . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Forcipiger longirostris
Key to the species of Hemitaurichthys occurring in the area

1a. Dorsal and anal fins orange, centre of body and caudal fin white . . . . . Hemitaurichthys polylepis
1b. Body colour uniformly dark brownish, centre of body white . . . . . . . . . . . . . . . . . . . . ® 2
2a. Dorsal fin with XVI spines and 18 to 20 soft rays; anal fin with V spines and 15 soft rays
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemitaurichthys multispinosus
2b. Dorsal fin with XII spines and 26 soft rays; anal fin with III spines and 20 or 21 soft rays
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemitaurichthys thompsoni
Key to the species of Heniochus occurring in the area

1a. First dark band includes eyes as well as pectoral and pelvic fins . . . . . . Heniochus chrysostomus
1b. First dark band does not include eyes but does include pectoral and pelvic fins . . . . . . . . . . ® 2
2a. Dark band includes pectoral and pelvic fins and first 3 spines of dorsal fins . . . . . . . . . . . . ® 3
2b. Dark band which includes pectoral and pelvic fins can include 1 or more dorsal-fin spines
but always posterior to first 3 dorsal-fin spines . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Dark band which passes through eye extends below to edge of interopercle; a black
band encircles snout; a white streak crosses under chin . . . . . . . . . . . . Heniochus singularis
3b. Dark band which passes through eye does not continue below it to interopercle; no black
band encircling snout; no white streak crossing under chin . . . . . . . . . . Heniochus acuminatus
4a. Dark band which includes pectoral and pelvic fins does not include elongate (fourth)
dorsal-fin spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Heniochus monoceros
4b. Dark band which includes pectoral and pelvic fins includes elongate (fourth) dorsal-fin
spine . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
3232 Bony Fishes
5a. Body bands merge at centre part of body so that entire body from insertion of pelvic fins
to end of anal fin is uniformly dark coloured . . . . . . . . . . . . . . . . . . . . . Heniochus varius
5b. Body bands merging only in upper part of body, leaving a white triangular space in
midventral portion of body . . . . . . . . . . . . . . . . . . . . . . . . . . . Heniochus pleurotaenia

0
0 Amphichaetodon howensis (Waite, 1903)
0 Chaetodon adiergastos Seale, 1910
0 Chaetodon argentatus Smith and Radcliffe, 1911
0 Chaetodon aureofasciatus Macleay, 1878
0 Chaetodon auriga Forsskål, 1775
0 Chaetodon auripes Jordan and Snyder, 1901
0 Chaetodon baronessa Cuvier, 1831
0 Chaetodon bennetti Cuvier, 1831
0 Chaetodon citrinellus Cuvier, 1831
Chaetodon burgessi Allen and Starck, 1973
0 Chaetodon collare Bloch, 1787

0 Chaetodon declivis Randall, 1975
0 Chaetodon decussatus Cuvier, 1831
0 Chaetodon ephippium Cuvier, 1831
0 Chaetodon flavirostris Günther, 1873
0 Chaetodon guentheri Ahl, 1913
Chaetodon flavocoronatus Myers, 1980
0 Chaetodon guttatissimus Bennett, 1832

0 Chaetodon kleinii Bloch, 1790
0 Chaetodon lineolatus Cuvier, 1831
0 Chaetodon lunula (Lacepède, 1803)
0 Chaetodon melannotus Bloch and Schneider, 1801
0 Chaetodon mertensii Cuvier, 1831
0 Chaetodon meyeri Bloch and Schneider, 1801
0 Chaetodon nippon Steindachner and Döderlein, 1884
0 Chaetodon ocellicaudus Cuvier, 1831
0 Chaetodon octofasciatus Bloch, 1787
0 Chaetodon ornatissimus Cuvier, 1831
0 Chaetodon pelewensis Kner, 1868
Chaetodon oxycephalus Bleeker, 1853
0 Chaetodon plebeius Cuvier, 1831

0 Chaetodon punctatofasciatus Cuvier, 1831
0 Chaetodon quadrimaculatus Gray, 1831
0 Chaetodon rafflesii Bennett, 1830
0 Chaetodon rainfordi McCulloch, 1923
0 Chaetodon reticulatus Cuvier, 1831
0 Chaetodon selene Bleeker, 1853
0 Chaetodon semion Bleeker, 1855
0 Chaetodon smithi Randall, 1975
0 Chaetodon speculum Cuvier, 1831
0 Chaetodon tinkeri Schultz, 1951
0 Chaetodon trichrous Günther, 1874
Chaetodon triangulum Cuvier, 1831
0 Chaetodon trifascialis Quoy and Gaimard, 1824

0 Chaetodon trifasciatus Park, 1797
0 Chaetodon ulietensis Cuvier, 1831
0 Chaetodon unimaculatus Bloch, 1787
0 Chaetodon vagabundus Linnaeus, 1758
0 Chaetodon wiebeli Kaup, 1863
0 Chaetodon xanthurus Bleeker, 1857
0 Chelmon marginalis Richardson, 1842

0 Chelmon muelleri (Klunzinger, 1879)
0 Chelmon rostratus (Linnaeus, 1758)
0 Coradion altivelis McCulloch, 1916
0 Coradion chrysozonus (Kuhl and Van Hasselt in Cuvier, 1831)
0 Coradion melanopus (Cuvier, 1831)
0 Forcipiger flavissimus Jordan and McGregor, 1898
0 Forcipiger longirostris (Broussonet, 1782)
0 Hemitaurichthys multispinosus Randall, 1975
0 Hemitaurichthys polylepis (Bleeker, 1857)
Hemitaurichthys thompsoni Fowler, 1923
0 Heniochus acuminatus (Linnaeus, 1758)
0 Heniochus chrysostomus Cuvier, 1831
0 Heniochus monoceros Cuvier, 1831
0 Heniochus pleurotaenia Ahl, 1923
0 Heniochus singularis Smith and Radcliffe, 1911
0 Heniochus varius (Cuvier, 1829)
0 Parachaetodon ocellatus (Cuvier, 1831)
Prognathodes guezei (Maugé and Bauchot, 1976)
Roa modestus (Temminck and Schlegel, 1842)
References
Allen, G.R. 1985. Butterfly and angelfishes of the world. Volume 2. Melle, Germany, Mergus Publishers.
Allen, G.R., R.C. Steene, and M. Allen. 1998. A guide to angelfishes and butterflyfishes. Odyssey Publishing/Tropical
Reef Research, 250 p.
Burgess, W.E. 1978. Butterflyfishes of the world. A monograph of the Family Chaetodontidae. Neptune City, New
Jersey, T.F.H. Publications, 832 p.
Steene, R.C. 1977. Butterfly and angelfishes of the world. Volume 1. Melle, Germany, Mergus Publishes, 144 p.
3234 Bony Fishes
Amphichaetodon howensis (Waite, 1903)

En - Lord Howe coralfish.
Maximum total length about 20 cm. Inhabits coral reefs at depths of 10 to 50 m. Omnivorous; forms
pairs. Almost never exported through the aquarium trade. Distributed in the southwestern Pacific
Ocean, from New Caledonia to southern Australia and New Zealand.
Chaetodon adiergastos Seale, 1910

En - Philippine butterflyfish.
Maximum total length about 15 cm. Inhabits coral reefs and reef slopes at depths of 3 to 30 m.
Feeds on corals, crabs, worms, and other invertebrates; forms pairs or small groups. Rarely
exported through the aquarium trade. Distributed from northwestern Australia to Java, Philippines,
Taiwan Province of China, and the Ryukyu Islands.
Chaetodon argentatus Smith and Radcliffe, 1911

En - Asian butterflyfish.
Maximum total length about 20 cm. Inhabits outer reef slopes and drop-offs at depths of 5 to 20 m.
Omnivorous; forms pairs or small groups. Occasionally exported through the aquarium trade.
Distributed from the northern Philippines and Taiwan Province of China to southern Japan.
Chaetodon aureofasciatus Macleay, 1878

En - Goldenstriped butterflyfish.
Maximum total length about 13 cm. Inhabits coral reefs at depths of 5 to 15 m. Feeds on corals;
solitary or in pairs. Occasionally exported through the aquarium trade. Distributed from eastern
Australia and Great Barrier Reef to Papua New Guinea.
3236 Bony Fishes
Chaetodon auriga Forsskål, 1775

En - Threadfin butterflyfish; Fr - Chétodon cocher.
Maximum total length about 23 cm. Inhabits a wide variety of habitats at depths of 3 to 20 m. Feeds
on crabs, worms, and other invertebrates; solitary or in pairs. Frequently exported through the
aquarium trade. Distributed throughout the tropical Indo-Pacific region.
Chaetodon auripes Jordan and Snyder, 1901

En - Oriental butterflyfish.
Maximum total length about 20 cm. Natural diet unknown. Rarely exported through the aquarium
trade. Inhabits rocky reefs at depths of 4 to 25 m. Distributed from the northern Philippines and
Taiwan Province of China to southern Japan.
Chaetodon baronessa Cuvier, 1831

En - Baroness butterflyfish; Fr - Chétodon baronne.
forms pairs. Occasionally exported through the aquarium trade. Usually starves when kept in
captivity. Distributed from Cocos-Keeling in the eastern Indian Ocean through Indonesia, New
Guinea, and Philippines north to southern Japan, east to Fiji.
Chaetodon bennetti Cuvier, 1831

En - Bennett’s butterflyfish; Fr - Chétodon de Bennett.
solitary or in pairs. Occasionally exported through the aquarium trade. Usually starves when kept
in captivity. Distributed throughout the Indian Ocean and western and southern Pacific Ocean.
3238 Bony Fishes
Chaetodon citrinellus Cuvier, 1831

En - Speckled butterflyfish; Fr - Chétodon citron.
Maximum total length about 13 cm. Inhabits lagoons and coral reefs at depths of 2 to 15 m.
Omnivorous; forms pairs. Occasionally exported through the aquarium trade. Distributed throughout
the Indo-Pacific, except the southeastern Pacific Ocean.
Chaetodon collare Bloch, 1787

En - Collare butterflyfish; Fr - Chétodon à collier.
Maximum total length about 16 cm. Inhabits coral reefs and aggregates in midwater at depths of 3
to 15 m. Feeds on corals, crabs, worms, and other invertebrates; forms pairs or small to large
groups. Occasionally exported through the aquarium trade. Distributed from the Maldive Islands to
the Indo-Malayan region and the Philippines.
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Chaetodon declivis Randall, 1975

En - Marquesan butterflyfish.
Feeds on crabs, worms, and other invertebrates; forms pairs. Rarely exported through the aquarium
trade. Part of a group of butterflyfishes (subgenus Roaops) for which species limits are not well
defined. Known only from the Line and Marquesas islands.
Chaetodon decussatus Cuvier, 1831

En - Indian vagabond butterflyfish.
pairs. Rarely exported through the aquarium trade. Distributed from the central and eastern Indian
Ocean to Thailand and Indonesia.
3240 Bony Fishes
Chaetodon ephippium Cuvier, 1831

En - Saddleback butterflyfish; Fr - Chétodon à selle noire.
Maximum total length about 20 cm. Inhabits a wide variety of habitats at depths of 5 to 15 m.
Omnivorous; forms pairs. Frequently exported through the aquarium trade. Distributed from the
Indo-Malayan region throughout the tropical insular Pacific Ocean.
Chaetodon flavirostris Günther, 1873

En - Black butterflyfish; Fr - Chétodon à nez jaune.
Omnivorous; solitary or in pairs. Occasionally exported through the aquarium trade. Distributed
throughout the southern tropical Pacific Ocean from Australia to Rapa; also recorded from Easter
Island.
Chaetodon guentheri Ahl, 1913

En - Günther’s butterflyfish.
to 40 m. Feeds on plankton; forms small to large aggregations. Rarely exported through the
aquarium trade. Antitropical in distribution; from Taiwan Province of China to southern Japan in the
North, and from Papua New Guinea to southern Australia in the south.
Chaetodon guttatissimus Bennett, 1832

En - Spotted butterflyfish; Fr - Chétodon moucheté.
pairs. Occasionally exported through the aquarium trade. Distributed throughout the Indian Ocean,
eastward to Thailand and parts of Indonesia.
3242 Bony Fishes
Chaetodon kleinii Bloch, 1790

En - Klein’s butterflyfish; Fr - Chétodon de Klein.
to 60 m. Feeds on plankton; forms small to large aggregations. Frequently exported through the
aquarium trade. Distributed throughout most of the Indo-Pacific region.
Chaetodon lineolatus Cuvier, 1831

En - Lined butterflyfish; Fr - Chétodon strié.
pairs. Frequently exported through the aquarium trade. Distributed throughout the Indo-Pacific
region, except the southeastern Pacific Ocean.
Chaetodon lunula (Lacepède, 1803)

En - Raccoon butterflyfish; Fr - Chétodon à croissant.
small to large aggregations. Frequently exported through the aquarium trade. Distributed throughout
most of the Indo-Pacific region.
Chaetodon melannotus Bloch and Schneider, 1801

En - Blackbacked butterflyfish; Fr - Chétodon à dos noir.
forms pairs. Frequently exported through the aquarium trade. Distributed throughout the Indian
Ocean, Indo-Malayan region, and tropical western and southern Pacific Ocean.
3244 Bony Fishes
Chaetodon mertensii Cuvier, 1831

En - Merten’s butterflyfish; Fr - Chétodon de Mertens.
Omnivorous; forms pairs. Frequently exported through the aquarium trade. Distributed throughout
the tropical western and southern Pacific Ocean; also recorded from Easter Island.
Chaetodon meyeri Bloch and Schneider, 1801

En - Meyer’s butterflyfish; Fr - Chétodon de Meyer.
captivity. Occassionally observed as an unusal colour variety lacking the dark diagonal bands; may
represent hybrids with Chaetodon reticulatus or C. ornatissimus. Distributed throughout the Indian
Ocean, Indo-Malayan region to the Central Pacific Ocean.
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Chaetodon nippon Steindachner and Döderlein, 1884

En - Japanese butterflyfish.
Maximum total length about 15 cm. Inhabits rocky reefs at depths of 5 to 20 m. Feeds on crabs,
worms, and other invertebrates; forms small to large aggregations. Almost never exported through
the aquarium trade. Distributed from the northern Philippines to southern Japan.
Chaetodon ocellicaudus Cuvier, 1831

En - Spottail butterflyfish.
forms pairs. Occasionally exported through the aquarium trade. Distributed throughout the Indo-
Malayan region, eastward to Palau and the Solomon Islands; also reported from Zanzibar, but this
record is doubtful.
3246 Bony Fishes
Chaetodon octofasciatus Bloch, 1787

En - Eightbanded butterflyfish.
forms pairs or small groups. Occasionally exported through the aquarium trade. Often starves when
kept in captivity. An unusal colour variety in the Solomon Islands and Papua New Guinea exhibits
alternating yellow and orange bars. Distributed from the central Indian Ocean eastward to the
western Pacific Ocean, northward to the Ryukyu Islands.
Chaetodon ornatissimus Cuvier, 1831

En - Ornate butterflyfish; Fr - Chétodon à bandes orange.
captivity. Distributed from Sri Lanka in the Indian Ocean, eastward throughout the Indo-Malayan
region and tropical Pacific Ocean, northward to southern Japan.
Chaetodon pelewensis Kner, 1868

En - Dot-and-dash butterflyfish; Fr - Chétodon de Palau.
pairs. Frequently exported through the aquarium trade. Readily hybridizes with Chaetodon
punctatofasciatus in areas where the 2 species are sympatric. Distributed from the southwestern
Pacific Ocean northward to Papua New Guinea, eastward to Pitcairn; also reported from Easter
Island.
Chaetodon plebeius Cuvier, 1831

En - Bluespot butterflyfish; Fr - Chétodon à deux taches.
forms pairs. Occasionally exported through the aquarium trade. Distributed from the Andaman Sea
eastward throughout the Indo-Malayan Archipelago to Fiji, northward to southern Japan, southward
to Lord Howe Island.
3248 Bony Fishes
Chaetodon punctatofasciatus Cuvier, 1831

En - Spotbanded butterflyfish.
pairs. Frequently exported through the aquarium trade. Readily hybridizes with Chaetodon
pelewensis in areas where the 2 species are sympatric. Distributed from the western Indian Ocean
throughout the Indo-Malayan region, eastward to the Marshall Islands, northward to the Ryukyu
Islands.
Chaetodon quadrimaculatus Gray, 1831

En - Fourspot butterflyfish.
forms pairs or small groups. Frequently exported through the aquarium trade. Distributed throughout
the tropical northwestern and Central Pacific Ocean.
Chaetodon rafflesii Bennett, 1830

En - Latticed butterflyfish.
Maximum total length about 15 cm. Inhabits coral reefs at depths of 3 to 20 m. Feeds on corals,
crabs, worms, and other invertebrates; forms pairs. Frequently exported through the aquarium
trade. Distributed from Sri Lanka and the Andaman Sea throughout the Indo-Malayan region
northward to southern Japan, eastward across the tropical southern Pacific Ocean to the Tuamotu
Archipelago.
Chaetodon rainfordi McCulloch, 1923

En - Rainford’s butterflyfish.
Maximum total length about 13 cm. Inhabits lagoons and coral reefs at depths of 3 to 15 m. Feeds
on corals; solitary or in pairs. Rarely exported through the aquarium trade. Distributed from Papua
New Guinea southward to Lord Howe Island.
3250 Bony Fishes
Chaetodon reticulatus Cuvier, 1831

En - Reticulated butterflyfish; Fr - Chétodon maillé.
solitary or in pairs. Occasionally exported through the aquarium trade. Often starves when kept in
captivity. Distributed throughout the tropical western and Central Pacific Ocean.
Chaetodon selene Bleeker, 1853

En - Yellowdotted butterflyfish.
Maximum total length about 10 cm. Inhabits coral reefs at depths of 10 to 50 m. Omnivorous. Almost
never exported through the aquarium trade. Distributed throughout the Indo-Malayan region and
western New Guinea, northward to the Ryukyu Islands.
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Chaetodon semion Bleeker, 1855

En - Dotted butterflyfish.
Maximum total length about 26 cm. Inhabits coral reefs at depths of 5 to 25 m. Omnivorous; solitary
or in pairs. Occasionally exported through the aquarium trade. Distributed from the Maldive Islands
eastward throughout the Indo-Malayan region and tropical western and southern Pacific Ocean.
Chaetodon smithi Randall, 1975

En - Smith’s butterflyfish.
to 30 m. Feeds on plankton; forms small to large aggregations. Almost never exported through the
aquarium trade. Known only from Rapa to Pitcairn, and a single specimen from Easter Island.
3252 Bony Fishes
Chaetodon speculum Cuvier, 1831

En - Ovalspot butterflyfish; Fr - Chétodon à miroir.
solitary or in pairs. Occasionally exported through the aquarium trade. Distributed from the eastern
Indian Ocean eastward throughout the Indo-Malayan region to Tonga, northward to southern Japan,
southward to Lord Howe Island.
Chaetodon tinkeri Schultz, 1951

En - Tinker’s butterflyfish.
Feeds on crabs, worms, and other invertebrates; forms pairs. Rarely exported through the aquarium
trade. Part of a group of butterflyfishes (subgenus Roaops) for which species limits are not well
defined. Presently known from the main Hawaiian Islands, Johnston Atoll, and the Marshall Islands;
however, unidentified specimens resembling this species have been collected at Tarawa and
Rarotonga.
Chaetodon trichrous Günther, 1874

En - Tahiti butterflyfish.
to 25 m. Feeds on plankton; forms small to large aggregations. Almost never exported through the
aquarium trade. Known only from the Society and Marquesas islands.
Chaetodon trifascialis Quoy and Gaimard, 1824

En - Chevroned butterflyfish; Fr - Chétodon tête de flèche.
solitary. Occasionally exported through the aquarium trade. Usually starves when kept in captivity.
Distributed throughout most of the Indo-Pacifc region, excluding the main Hawaiian Islands,
Marquesas Islands, and Easter Island.
3254 Bony Fishes
Chaetodon trifasciatus Park, 1797

En - Oval butterflyfish; Fr - Chétodon à nageoire rouge orange.
on corals; forms pairs. Frequently exported through the aquarium trade. Usually starves when kept
in captivity. Distributed throughout most of the Indo-Pacifc region, excluding the Red Sea, Marque-
sas Islands, and Easter Island.
Chaetodon ulietensis Cuvier, 1831

En - Pacific doublesaddle butterflyfish; Fr - Chétodon d’Uliétéa.
pairs or small groups. Frequently exported through the aquarium trade. Similar in colour to
Chaetodon falcula of the Indian Ocean; the latter species has been reported from parts of Indonesia,
but this remains to be confirmed. Distributed from Western Australia through the Indo-Malayan
region eastward to the Tuamotu Archipelago, northward to southern Japan, southward to Lord Howe
Island.
Chaetodon unimaculatus Bloch, 1787

En - Teardrop butterflyfish; Fr - Chétodon ocellé.
on corals; forms pairs. Frequently exported through the aquarium trade. Distributed throughout most
of the tropical Indo-Pacific region.
Chaetodon vagabundus Linnaeus, 1758

En - Vagabond butterflyfish; Fr - Chétodon à vagabond.
pairs. Frequently exported through the aquarium trade. Distributed throughout much of the tropical
Indo-Pacific region, excluding Western Australia, Hawaii, and the Marquesas Islands.
3256 Bony Fishes
Chaetodon wiebeli Kaup, 1863

En - Wiebel’s butterflyfish.
Maximum total length about 18 cm. Inhabits coral reefs at depths of 4 to 25 m. Natural diet unknown.
Rarely exported through the aquarium trade. Distributed from the Indo-Malayan region northward
to southern Japan.
Chaetodon xanthurus Bleeker, 1857

En - Yellowtail butterflyfish.
Omnivorous; forms pairs. Frequently exported through the aquarium trade. Distributed from the
Indo-Malayan region northward to southern Japan.
Chelmon marginalis Rishardson, 1842

En - Margined coralfish.
Feeds on crabs, worms, and other invertebrates; usually in pairs. Almost never exported through
the aquarium trade. Distributed throughout the western coast of Australia and the Great Barrier
Reef.
Chelmon muelleri (Klunzinger, 1879

En - Müller’s coralfish.
on crabs, worms, and other invertebrates; usually in pairs. Almost never exported through the
aquarium trade. Known only from northern Australia.
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3258 Bony Fishes
Chelmon rostratus (Linnaeus, 1758)

En - Copperbanded butterflyfish.
Maximum total length about 20 cm. Inhabits coral reefs at depths of 3 to 20 m. Feeds on crabs,
worms, and other invertebrates; usually in pairs. Frequently exported through the aquarium trade.
Distributed from the Andaman Sea eastward throughout the Indo-Malayan region, northward to
southern Japan and the Great Barrier Reef.
Coradion altivelis McCulloch, 1916

En - Highfin coralfish; Fr - Coradion à grande voile.
Omnivorous; usually in pairs. Rarely exported through the aquarium trade. Distributed from the
Andaman Sea eastward throughout the Indo-Malayan region, northward to southern Japan and the
Great Barrier Reef.
Coradion chrysozonus (Kuhl and van Hasselt in Cuvier, 1831)

En - Orangebanded coralfish.
Omnivorous; usually in pairs. Rarely exported through the aquarium trade. Distributed from the
Andaman Sea eastward throughout the Indo-Malayan region, northward to southern Japan and the
Great Barrier Reef.
Coradion melanopus (Cuvier, 1831)

En - Two-eyed coralfish.
Omnivorous; usually in pairs. Rarely exported through the aquarium trade. Distributed throughout
the Indo-Malayan region eastward to Papua New Guinea.
3260 Bony Fishes
Forcipiger flavissimus Jordan and McGregor, 1898

En - Forcepsfish; Fr - Chelmon à long bec.
worms, and other invertebrates; forms pairs or small groups. Frequently exported through the
aquarium trade. Distributed throughout the entire tropical Indo-Pacific region to the eastern Pacific
Ocean.
Forcipiger longirostris (Broussonet, 1782)

En - Longnosed butterflyfish; Fr - Chelmon à très long bec.
worms, and other invertebrates; forms pairs. Rarely exported through the aquarium trade. Occa-
sionally exhibits an entirely dark (melanistic) colour phase. Distributed throughout most of the
tropical Indo-Pacific region, excluding Easter Island.
Hemitaurichthys multispinosus Randall, 1975

En - Manyspined butterflyfish.
to 50 m. Feeds on plankton; forms small to large aggregations. Never exported through the
aquarium trade. Known only from Pitcairn Island.
Hemitaurichthys polylepis (Bleeker, 1857)

En - Pyramid butterflyfish; Fr - Papillon à rideaux jaunes.
to 40 m. Feeds on plankton; forms small to large aggregations. Frequently exported through the
aquarium trade. Distributed from the Andaman Sea eastward throughout the Indo-Malayan region
to Hawaii and the Society Islands, northward to southern Japan.
3262 Bony Fishes
Heniochus acuminatus (Linnaeus, 1758)

En - Longfin bannerfish; Fr - Pavillon.
Maximum total length about 20 cm. Inhabits coral reefs at depths of 2 to 40 m. Omnivorous; usually
found in pairs, but rarely in aggregations. Frequently exported through the aquarium trade.
Distributed throughout much of the tropical Indo-Pacific region, excluding Hawaii and the Marquesas
Islands.
Heniochus chrysostomus Cuvier, 1831

En - Pennant bannerfish; Fr - Hénioche renversé.
forms small aggregations. Frequently exported through the aquarium trade. Distributed from the
tropical eastern Indian Ocean throughout the Indo-Malayan region eastward across the western
and Central Pacific Ocean to the Tuamotu Archipelago, northward to southern Japan.
Heniochus monoceros Cuvier, 1831

En - Masked bannerfish; Fr - Hénioche cornu.
worms, and other invertebrates; solitary or in pairs. Occasionally exported through the aquarium
trade. Distributed throughout much of the tropical Indo-Pacific region, excluding the Red Sea,
Western Australia, Hawaii, and the Marquesas Islands.
Heniochus pleurotaenia Ahl, 1923

En - Indian bannerfish; Fr - Hénioche singulier.
worms, and other invertebrates; forms small aggregations. Occasionally exported through the
aquarium trade. Distributed from the Maldive Islands and Sri Lanka to Java and the Andaman Sea.
3264 Bony Fishes
Heniochus singularis Smith and Radcliffe, 1911

En - Singular bannerfish; Fr - Hénioche bossu.
worms, and other invertebrates; solitary or in pairs. Occasionally exported through the aquarium
trade. Distributed from the Maldive Islands eastward throughout the Indo-Malayan region and
tropical western Pacific Ocean to Samoa, northward to southern Japan.
Heniochus varius (Cuvier, 1829)

En - Humphead bannerfish; Fr - Barbet souffré.
worms, and other invertebrates; solitary, pairs, or small groups. Frequently exported through the
aquarium trade. Distributed from Western Australia northward throughout the Indo-Malayan region
to southern Japan, eastward across the tropical western and southern Pacific Ocean.
Parachaetodon ocellatus (Cuvier, 1831)

En - Ocellate coralfish.
Omnivorous. Almost never exported through the aquarium trade. Distributed from Sri Lanka
eastward throughout the Indo-Malayan region northward to the Ryukyu Islands, southward to the
Great Barrier Reef.
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3266 Bony Fishes
POMACANTHIDAE
Angelfishes
by R. Pyle
D iagnostic characters: Oblong to oval-

shaped, deep-bodied, highly compressed
percoid fishes (size to about 50 cm). Head
length roughly equal to head height; 1 or more
prominent spines at angle of preopercle.
Eye moderately small, located above longitu-
dinal axis from tip of snout to middle of
caudal fin. Snout short, not produced.
Mouth small, terminal, protractile, gape not ex-
tending to anterior rim or orbit. Teeth bristle-
like, usually tricuspid, arranged in rows or
bands across jaws; vomer and palatines with-
out teeth. Gill rakers short, ranging from 9 to 25
in number. A single dorsal fin, continuous
and relatively smooth; IX to XV strong, stout
spines, first few to several interspinous mem-
branes deeply incised; no notch between spi-
nous and soft dorsal fin; soft dorsal fin with 15
to 33 branched rays, some species with fila-
mentous extension of 1 or more soft dorsal-fin
rays at dorsoposterior margin of fin. Anal fin
with III strong, stout spines, interspinous mem-
branes deeply incised; soft anal fin with 14 to
25 branched rays, some species with filamen-
tous extension of 1 or more soft dorsal-fin rays
at ventroposterior margin of fin. Caudal fin
rounded to slightly emarginate (except Genicanthus, which has a forked caudal fin with long filamentous
extensions). Pectoral fins transparent with 16 to 21 soft rays. Pelvic fins with a I stout spine and 5 branched
rays. Scales coarsly ctenoid, covering head, body, and median fins; largest in centre of body, smaller on
head, thorax, belly, caudal peduncle, and median fins; number of lateral-line scales variable, ranging from
30 to 90 (depending on genus). Vertebrae 10+14. Colour: almost all species very brightly coloured
with complex and varied colour patterns; juveniles of Pomacanthus with alternating black, blue, and
white vertical bands, strikingly different in colour from adults, which vary from species to species; strong
sexual dichromatism in species of Genicanthus, and lesser dichromatism in some species of Centropyge.
Habitat, biology, and fisheries: Mostly found on outer coral reefs at depths of 5 to 60 m (some species
restricted to greater depths), but some species occur in lagoons. Most species closely associated with the
substratum, feeding on sponges and other marine invertebrates; most species of Genicanthus usually found
in midwater and feed on plankton. Although sometimes harvested as food fish, the primary fishery value of
pomacanthids is through the ornamental marine aquarium trade, where they are the second most-frequently
exported fish by number, and highest in total value of all families of aquarium fishes in trade.
Remarks: Dichotomous keys are seldom used for identifying species of pomacanthids, as species in this
familiy exhibit highly conspicuous and distinctive colour patterns (the primary components of which usually
remain evident in preservative). Nevertheless, keys to genera and species of pomacanthids are included
here for purposes of consistency and clarity. Keys are more reliable when used for adult specimens.
Similar families occuring in the area

Chaetodontidae: similar in general body shape and colour pattern
to Pomacanthidae, but easily distinguished in lacking the spine on
margin of preopercle. Also, species of Chaetodontidae possess a
scaly axillary process at base of pelvic fins, usually have dark ocular
bands and false-eye spots, and often have produced snouts.
Chaetodontidae
Perciformes: Percoidei: Pomacanthidae 3267
Acanthuridae: similar general body shape and some species with

bright colours; however, species of Acanthuridae can easily be dis-
tinguished by the presence of a fixed or retractable sharp spine on
the caudal peduncle, and lack of spines at angle of preopercle. Also,
Acanthuridae typically have fewer dorsal-fin spines (IV to IX) than
most species of Pomacanthidae.
Scatophagidae: distinguished from Pomacanthidae by a deep notch
between spinous and soft portion of dorsal fin, lacking spines at angle
of preopercle, and in having IV anal-fin spines.
Ephippidae: lack spines at angle of preopercle; generally less colour-
ful as adults, and distinguished by greatly enlarged dorsal and anal Acanthuridae
fins as adults.
Pomacentridae: superficially resemble species of Centropyge, but lack spines at angle of preopercle.
Scatophagidae Ephippidae Pomacentridae
Key to the genera of Pomacanthidae occurring in the area

1a. Scale focus (convergence of radii) exposed, not hidden by cteni area . . . . . . . . . . . . . . . ® 2
1b. Scale focus (convergence of radii) not exposed, hidden by cteni area . . . . . . . . . . . . . . . ® 3
2a. Scale length 1 to 1.3 times in scale depth, 20 to 33 times in standard length . . . . . . Pomacanthus
2b. Scale length 1.5 to 1.7 times in scale depth, 35 to 51 times in standard length . . . Chaetodontoplus
3a. Caudal fin lunate, usually with long filamentous extensions on upper and lower lobes . .Genicanthus
3b. Caudal fin rounded or truncate, without filamentous extensions on upper or lower lobes . . . . . ® 4
4a. Suprecleithrum ovate, conspicuous; subopercle smooth . . . . . . . . . . . . . . . . Apolemichthys
4b. Supracleithrum elongate, well hidden by scales; subopercle with spines . . . . . . . . . . . . . . ® 5
4a. Interopercle spiny; no naked band of skin below and behind eye . . . . . . . . . . . . . . Centropyge
4b. Interopercle smooth; a naked band of skin from rear margin of orbit extending down and
forward below eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pygoplites
(a single species, P. diacanthus, in the genus)
Key to the species of Apolemichthys occurring in the area

1a. Colour of dorsal fin yellow . . . . . . . . . . . . . . . . . . . . . . . . . Apolemichthys trimaculatus
1b. Colour of dorsal fin mostly black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Colour of caudal and anal fins black; body colour greenish with an irregular bright yellow
spot on each scale . . . . . . . . . . . . . . . . . . . . . . . . . . . Apolemichthys xanthopunctatus
2b. Colour of caudal and anal fins mostly whitish grey; body colour grey with a few black
spots near dorsal-fin origin, dorsoposterior portion of body black with a distinct, broad
white band extending from about fourth dorsal-fin spine along base of dorsal fin to and
including caudal peduncle . . . . . . . . . . . . . . . . . . . . . . . . . . . . Apolemichthys griffisi
3268 Bony Fishes
Key to the species of Centropyge occurring in the area

1a. Posttemporal with spinules; 15 to 19 gill rakers on first gill arch . . . . . . . . . . . . . . . . . . ® 2
1b. Posttemporal without spinules; 21 to 25 gill rakers on first gill arch . . . . . . . . . . . . . . . . ® 13
2a. Dorsal-fin spines XIII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Dorsal-fin spines XIV or XV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Body with 8 to 12 vertical dark bars on pale background . . . . . . . . . . Centropyge multifasciata
3b. Body with 4 or 5 vertical white bars on bright red background . . . . . . . . . . . . Centropyge boylei
4a. Anterior third of body yellow, except for a dark blue nape spot just dorsal to eye; posterior
two-thirds of body and median fins dark blue; caudal fin yellow . . . . . . . . . . Centropyge bicolor
4b. Colour not as described above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Body colour dominated by yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5b. Body colour mostly black, brown, grey, or dark orange . . . . . . . . . . . . . . . . . . . . . . . ® 9
6a. Body, head, and fins yellow, without large blue or black markings . . . . . . . . . . . . . . . . . ® 7
6b. Body, head, or fins with large prominent blue or black markings . . . . . . . . . . . . . . . . . . ® 8
7a. Blue ring around eye, blue margins on median fins, and blue posterior edge of opercu-
lum; soft dorsal and anal fins rounded . . . . . . . . . . . . . . . . . . . . . . Centropyge flavissima
7b. No blue markings at all . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge heraldi
8a. Dorsal fin and dorsal portion of body bluish purple; body entirely yellow . . . . . . Centropyge colini
8b. Dorsal fin yellow; a prominent black spot just dorsal to centre of body (not to be confused
with juvenile C. flavissimus, which may have a prominent blue-margined black ocellate
spot at centre of body) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge narcosis
9a. Body and median fins dull orange or chocolate brown with many thin vertical pale
vermiculations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge aurantia
9b. Body and median fins primarily black or grey . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 10
10a. Body and median fins mostly black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
10b. Body and median fins mostly grey, black posteriorly . . . . . . . . . . . . . . . . . . . . . . . . ® 12
11a. A prominent white vertically oblong marking on centre of body; pelvic-fin spines and
margin of anal fin yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge tibicen
11b. Body and fins entirely black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge nox
12a. Posterior third of body, caudal fin, and soft dorsal and anal fins black; thorax grey; no
thin vertical orange vermiculations on body . . . . . . . . . . . . . . . . . . . . . Centropyge vroliki
12b. Black restricted to caudal fin, caudal peduncle, and posterior portion of soft dorsal fin;
thorax orange; numerous thin vertical orange vermiculations on body . . . . . . . . Centropyge eibli
13a. Dorsal-fin spines XIII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge nigriocella
13b. Dorsal-fin spines XIV to XV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
14a. Caudal fin opaque blue or purple with black markings . . . . . . . . . . . . . . . . . . . . . . . ® 15
14b. Caudal fin transparent yellow or orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
15a. Dorsoposterior portion of body blue, a large triangular blue patch extending from nape
and interorbital region to a point just posterior to opercular margin, just dorsal to insertion
of pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge venusta
15b. Body colour variable, but typically purple around margins and median fins, and orange
with thin vertical striations in centre of body . . . . . . . . . . . . . . . . . . . Centropyge bispinosa
16a. Two or 3 very prominent posteriorly projecting spines on preorbital . . . . . . Centropyge flavicauda
16b. No prominent posteriorly projecting spines on preorbital . . . . . . . . . . . . . . . . . . . . . ® 17
17a. Body with several to many distinct black or brown vertical bars . . . . . . . . . . . . . . . . . . ® 18
17b. Body without vertical bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
18a. Several broad black bars on bright reddish orange background, most bars at least 2/3
body depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge loricula
18b. Many thin greenish brown bars on a dull orange background, most bars less than 1/2
body depth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge shepardi
19a. Dorsal half of body white, ventral half yellow anteriorly (including thorax and pelvic fins)
and brownish blue posteriorly; a blue patch with black striations on nape above eye
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Centropyge multicolor
19b. Head and anterior abdomen orange, most of body orangish brown . . . . . . Centropyge hotumatua
Key to the species of Chaetodontoplus occurring in the area

1a. Head, body, and median fins except for caudal fin completely black with no markings;
caudal fin white; pelvic fins yellow . . . . . . . . . . . . . . . . . . . . . . . . Chaetodontoplus niger
1b. Colour not as described above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
2a. Dorsal-fin spines XI or XII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Dorsal-fin spines XIII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Scale rows about 80 to 90, each with 2 auxilliaries; fifth dorsal-fin spine longer than dorsal-fin
rays; ocular band black, narrow, and sharply defined; body colour light anteriorly, dark
posteriorly, longitudinal rows of white dots on side of body . . . . . . . . . Chaetodontoplus mesoleucus
3b. Scale rows more than 100, most with only 1 auxillary; fifth dorsal-fin spine longer than
dorsal-fin rays; ocular band broad, bluish black, and often diffuse at margins; a broad
white and yellow band extending from first dorsal-fin spine to pelvic fins . Chaetodontoplus duboulayi
4a. Body orangish brown with blue longitudinal stripes . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
4b. Body mostly black or dark blue, lacking longitudinal stripes . . . . . . . . . . . . . . . . . . . . . ® 6
5a. About 10 black-edged blue stripes on side of body and median fins . . Chaetodontoplus septentrionalis
5b. Blue stripes narrower, more numerous, irregular, confined to anterior portion of body
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodontoplus chrysocephalus
6a. Caudal fin with vertical black band, or black with yellow margin . . . . . . . . . . . . . . . . . . ® 7
6b. Caudal fin entirely yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
7a. Caudal fin black with yellow margin; posterior margins of soft dorsal and anal fins yellow;
pelvic fins black . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodontoplus melanosoma
7b. Caudal fin yellow with broad black posterior margin; soft dorsal- and anal-fin margins
bluish white; pelvic fins white . . . . . . . . . . . . . . . . . . . . . . Chaetodontoplus conspicilatus
8a. Body covered with tiny blue dots; no colour demarcation at head . Chaetodontoplus caeruleopunctatus
8b. Body without tiny blue dots; a sharp colour demarcation just posterior to head; yellow on
nape and thorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chaetodontoplus meredethi
Key to the species of Genicanthus occurring in the area

1a. Scale rows from upper end of gill opening to caudal-fin base 49 to 54; only first 2 dorsal-
fin interspinous membranes incised 1/2 or more length of spines . . . . . . . . . Genicanthus spinus
1b. Scale rows from upper end of gill opening to caudal-fin base 45 to 48; first 3 or 4 dorsal-
fin interspinous membranes incised 1/2 or more length of spines . . . . . . . . . . . . . . . . . ® 2
3270 Bony Fishes
2a. Anal-fin rays modally 18; males with numerous dark vertical bars, which extend to ventral
part of body, and a prominent black spot midventrally on thorax in front of base of pelvic
fins; females without dark markings on body . . . . . . . . . . . . . . . . Genicanthus melanospilos
2b. Anal-fin rays modally 16 or 17; males with black or orange-yellow horizontal stripes on
body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Basal one-half to three-fourths of upper lip scaly; first 4 dorsal-fin interspinous mem-
branes incised 1/2 or more length of spines . . . . . . . . . . . . . . . . . . . Genicanthus lamarck
3b. Lips not scales; first 3 dorsal-fin interspinous membranes incised 1/2 or more length of
spines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Males with 8 to 13 black horizontal stripes on about lower two-thirds of body; females
without dark markings on body but with black transverse bands on interorbital and
dorsally on snout and a black band in caudal-fin lobes . . . . . . . . . . . . Genicanthus watanabei
4b. Males lavender grey with a midlateral orange-yellow stripe (pale in alcohol) and an
orange-yellow band along back at base of dorsal fin; females with 3 diagonal dark bands
on body (lowermost blue in life), the middle one continuous with black band of lower
caudal-fin lobe, and 2 black bars on head, the posterior one curving on nape to join broad
black band at dorsal-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . Genicanthus bellus
Key to the species of Pomacanthus occurring in the area

Note: juveniles of P. sextriatus, P. navarchus, and P. xanthometopon are difficult to distinguish on the basis
of colour.
1a. Scales small or moderate, 70 or more scale rows in lateral line . . . . . . . . . . . . . . . . . . ® 2
1b. Scales large, fewer than 60 scale rows in lateral line . . . . . . . . . . . . . . . . . . . . . . . . ® 4
2a. Dorsal-fin spines XIV; adults bright blue with many longitudinal yellow lines; juveniles
with distinct white margin on complete length of dorsal fin, usually with a circular white
ring just anterior to caudal peduncle . . . . . . . . . . . . . . . . . . . . . . Pomacanthus imperator
2b. Dorsal-fin spines XIII; adult colour not as described above, juveniles without distinct
white margin on dorsal fin, white bars on body vertical or curved but none forming a
complete circle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
3a. Caudal fin dark with blue and white markings; adults greenish grey with a blue spot on
each body scale and filamentous extensions on soft dorsal and soft anal fins . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomacanthus semicirculatus
3b. Caudal fin transparent white; adults brown with several distinct diagonal blue lines and
a rounded soft anal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pomacanthus annularis
4a. Dorsal-fin rays 18 to 20; anal-fin rays 18 or 19; adults with 6 distinct dark vertical bars on
body and a broad white bar on head; caudal fin dark with blue dots . . . . . Pomacanthus sextriatus
4b. Dorsal-fin rays 16 to 18; anal-fin rays 16 to 18; no distinct vertical bars on adults; caudal
fin yellow or orange . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Eye within a bright orange patch across interorbital; a distinct dark spot at posterior base
of soft dorsal fin; caudal peduncle yellowish (adults) . . . . . . . . . . Pomacanthus xanthometapon
5b. Eye within a large blue patch covering most of head (except for thorax and region ventral
to eye); a broad dark blue band extending from soft anal fin dorsally across caudal
peduncle to posterior base of soft dorsal fin . . . . . . . . . . . . . . . . . Pomacanthus navarchus
List of species occuring in the area

0
0 Apolemichthys griffisi Carlson and Taylor, 1981
0 Apolemichthys trimaculatus (Lacepède in Cuvier, 1831)
0 Apolemichthys xanthopunctatus Burgess, 1973
0 Centropyge aurantia Randall and Wass, 1974
0 Centropyge bicolor (Bloch, 1787)
0 Centropyge bispinosa (Günther, 1860)
0 Centropyge boylei Pyle and Randall, 1993
0 Centropyge colini Smith-Vaniz and Randall, 1974
0 Centropyge eibli Klausewitz, 1963
0 Centropyge flavicauda Fraser-Brunner, 1933
0 Centropyge flavissima (Cuvier, 1831)
0 Centropyge heraldi Woods and Schultz, 1853
0 Centropyge hotumatua Randall and Caldwell, 1973
0 Centropyge loricula (Günther, 1860)
0 Centropyge multicolor Randall and Wass, 1974
0 Centropyge multifasciata (Smith and Radcliffe, 1911)
0 Centropyge narcosis Pyle and Randall, 1993
0 Centropyge nigriocella Woods and Schultz, 1953
0 Centropyge nox (Bleeker, 1853)
0 Centropyge shepardi Randall and Yasuda, 1979
0 Centropyge tibicen (Cuvier, 1831)
0 Centropyge venusta (Yasuda and Tominaga, 1969)
0 Centropyge vroliki (Bleeker, 1853)
0 Chaetodontoplus caeruleopunctatus Yasuda and Tominaga, 1976
0 Chaetodontoplus chrysocephalus Bleeker, 1854
0 Chaetodontoplus conspicillatus (Waite, 1900)
0 Chaetodontoplus duboulayi (Günther, 1867)
0 Chaetodontoplus melanosoma (Bleeker, 1853)
0 Chaetodontoplus meredithi Kuiter, 1990
0 Chaetodontoplus mesoleucus (Bloch, 1787)
0 Chaetodontoplus septentrionalis (Temminck and Schlegel, 1844)
Chaetodontoplus niger Chan, 1965
0 Genicanthus bellus Randall, 1975

0 Genicanthus lamarck (Lacepède, 1802)
0 Genicanthus melanospilos (Bleeker, 1857)
0 Genicanthus spinus Randall, 1975
0 Genicanthus watanabei (Yasuda and Tominaga, 1970)
0 Pomacanthus annularis (Bloch, 1787)
0 Pomacanthus imperator (Bloch, 1787)
0 Pomacanthus navarchus (Cuvier, 1831)
0 Pomacanthus semicirculatus (Cuvier, 1831)
0 Pomacanthus sextriatus (Cuvier, 1831)
0 Pomacanthus xanthometapon (Bleeker, 1853)
0 Pygoplites diacanthus (Boddaert, 1772)
References
Allen, G.R. 1985. Butterfly and angelfishes of the world. Volume 2. Melle, Germany, Mergus Publishers, pp. 145-352.
Allen, G.R., R.C. Steene, and M. Allen. 1998. A guide to angelfishes and butterflyfishes. Odyssey Publishing/Tropical
Reef Research, 250 p.
Steene, R.C. 1977. Butterfly and angelfishes of the world. Volume 1. Melle, Germany, Mergus Publishers., 144 p.
3272 Bony Fishes
Apolemichthys griffisi Carlson and Taylor, 1981

En - Griffis’ angelfish.
Feeds on sponges and tunicates; solitary, pairs, or small groups. Rarely exported through the
aquarium trade. Distributed from the Indo-Malayan region eastward to the Line Islands.
Apolemichthys trimaculatus (Lacepède in Cuvier, 1831)

En - Threespot angelfish; Fr - Poisson-ange à trois taches.
Feeds on sponges and tunicates; solitary, pairs, or small groups. Frequently exported through the
aquarium trade. Distributed throughout the Indian and tropical western Pacific oceans.
Apolemichthys xanthopunctatus Burgess, 1973

En - Goldflake angelfish.
Feeds on sponges and tunicates; solitary, pairs, or small groups. Occasionally exported through
the aquarium trade. Distributed from the Gilbert Islands to the Line Islands.
Centropyge aurantia Randall and Wass, 1974

En - Golden angelfish.
Maximum total length about 10 cm. Inhabits rubble and thick coral areas at depths of 3 to 20 m.
Natural diet and social structure unknown. This species is extremely cryptic and is seldom seen by
divers. Rarely exported through the aquarium trade. Known from Indonesia, Great Barrier Reef,
Solomon Islands, Pohnpei, and Samoa, and probably occurs elsewhere in the area.
Centropyge bicolor (Bloch, 1787)

En - Bicolor angelfish; Fr - Poisson ange jaune et bleu.
on algae; forms harems of 3 to 7 individuals. Frequently exported through the aquarium trade.
Distributed from the Indo-Malayan region eastward across the tropical Pacific Ocean to the Line
Islands, northward to southern Japan.
Centropyge bispinosa (Günther, 1860)

En - Coral beauty; Fr - Poisson ange à deux épines.
Maximum total length about 8 cm. Inhabits coral reefs and rubble areas at depths of 10 to 50 m.
Frequently exported through the aquarium trade. Feeds on algae; forms harems of 3 to 7 individuals.
Distributed throughout most of the tropical Indo-Pacific region, excluding the Red Sea, Hawaii, and
southeastern Pacific Ocean.
3274 Bony Fishes
Centropyge boylei Pyle and Randall, 1993

En - Peppermint angelfish.
Maximum total length about 7 cm. Inhabits ledges, caves, and rubble areas at depths of 55 to 110 m.
Natural diet unknown; forms pairs or small groups. Almost never exported through the aquarium
trade. Usually starves when kept in captivity. Known only from the Cook Islands.
Centropyge colini Smith-Vaniz and Randall, 1974

En - Colin’s angelfish.
Maximum total length about 8 cm. Inhabits ledges and caves at depths of 20 to 100 m. Natural diet
unknown; forms harems of 3 to 7 individuals. Almost never exported through the aquarium trade.
Known from Cocos-Keeling Islands, parts of Indonesia, Palau, Guam, Fiji, and the Marshall Islands.
Centropyge eibli Klausewitz, 1963

En - Eibl’s angelfish.
Maximum total length about 15 cm. Inhabits coral reefs at depths of 10 to 30 m. Feeds on algae;
forms harems of 3 to 7 individuals. Frequently exported through the aquarium trade. Readily forms
hybrids with Centropyge vroliki in areas where the 2 species are sympatric, and hybrids with
C. flavissima have been recorded from Christmas Island and Cocos-Keeling Islands in the eastern
Indian Ocean. This species is mimicked by juveniles of the acanthurid Acanthurus tristis. Distributed
from Sri Lanka to the eastern Indo-Malayan region.
Centropyge flavicauda Fraser-Brunner, 1933

En - Whitetail angelfish.
Maximum total length about 6 cm. Inhabits rubble areas at depths of 10 to 60 m. Feeds on algae;
forms aggregations. Occasionally exported through the aquarium trade. Probably synonymous with
Centropyge fisheri of Hawaii and Johnston Atoll. Distributed throughout much of the tropical
Indo-Pacific region, excluding the Red Sea, Hawaii, and southeastern Pacific Ocean.
Centropyge flavissima (Cuvier, 1831)

En - Lemonpeel angelfish; Fr - Centropyge tout jaune.
Readily forms hybrids with Centropyge vroliki in areas where the 2 species are sympatric, and
hybrids with C. eibli have been recorded from Christmas Island and Cocos-Keeling Islands in the
eastern Indian Ocean. This species is mimicked by juveniles of the acanthurid Acanthurus pyro-
ferus. Distributed throughout most of the Central Pacific Ocean from the Ogasawara Islands to the
Tuamotu Archipelago, excluding Hawaii and Johnston Atoll, as well as an unusual population in the
insular eastern Indian Ocean; also reported from the Coral Sea, Great Barrier Reef, and Ryukyu
Islands.
Centropyge heraldi Woods and Schultz, 1853

En - Herald’s angelfish.
forms harems of 2 to 4 individuals. Frequently exported through the aquarium trade. Distributed
from southern Japan southward to Taiwan Province of China and across the tropical western and
southern Pacific Ocean.
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3276 Bony Fishes
Centropyge hotumatua Randall and Caldwell, 1973

En - Hotumatua angelfish.
Maximum total length about 8 cm. Inhabits coral reefs at depths of 14 to 50 m. Natural diet unknown;
forms harems of 3 to 7 individuals. Rarely exported through the aquarium trade. Distributed from
the Austral Islands eastward to Easter Island.
Centropyge loricula (Günther, 1860)

En - Japanese pygmy angelfish.
forms harems of 3 to 7 individuals. Frequently exported through the aquarium trade. Specimens
from the Marquesas Islands lack the vertical black bars. Distributed throughout most of the Central
Pacific Ocean.
Centropyge multicolor Randall and Wass, 1974

En - Multicolor angelfish.
Distributed throughout the Central Pacific Ocean from Palau, Guam, and the Marshall Islands
eastward and southward to Fiji and the Society Islands. Natural diet unknown; feeds readily in
aquaria; forms harems of 3 to 7 individuals. Rarely exported through the aquarium trade.
Centropyge multifasciata (Smith and Radcliffe, 1911)

En - Multibarred angelfish.
unknown; forms pairs or small groups. Occasionally exported through the aquarium trade. Usually
starves when kept in captivity. Distributed from the eastern Indian Ocean across most of the Pacific
Ocean to the Society Islands, northward to the Ryukyu Islands.
Centropyge narcosis Pyle and Randall, 1993

En - Narc angelfish.
Maximum total length about 7 cm. Inhabits ledges and caves at depths of 100 to 130 m. Natural
diet unknown; feeds readily in aquaria; forms pairs or small groups. Almost never exported through
the aquarium trade. Known only from the Cook Islands.
Centropyge nigriocella Woods and Schultz, 1953

En - Blackspot angelfish.
Maximum total length about 6 cm. Inhabits coral reefs at depths of 4 to 15 m. Feeds on algae. Never
exported through the aquarium trade. Extremely cryptic; never seen by divers except at poison
stations. Patchy distribution; from Johnston Atoll to the Society Islands, and also reported from the
Mariana and Admiralty Islands; likely occurs in other areas within its range.
3278 Bony Fishes
Centropyge nox (Bleeker, 1853)

En - Midnight angelfish.
Maximum total length about 9 cm. Inhabits outer reef slopes, ledges, and caves at depths of 10 to
40 m. Feeds on algae; forms harems of 3 to 7 individuals. Frequently exported through the aquarium
trade. Distributed from the Ryukyu Islands southward to Indonesia, the Solomon Islands, Great
Barrier Reef, and New Caledonia.
Centropyge shepardi Randall and Yasuda, 1979

En - Shepard’s angelfish.
forms harems of 3 to 7 individuals. Occasionally exported through the aquarium trade. Known only
from the Mariana and Ogasawara Islands, and possibly a population southwest of Palau.
Centropyge tibicen (Cuvier, 1831)

En - Keyhole angelfish; Fr - Centropyge joueur de flûte.
Distributed throughout the Indo-Malayan region and western Pacific Ocean, northward to southern
Japan; southward to Lord Howe Island.
Centropyge venusta (Yasuda and Tominaga, 1969)

En - Purplemask angelfish.
unknown; forms pairs or small groups. Occasionally exported through the aquarium trade. Occa-
sionally forms hybrids with Centropyge multifasciata. The generic placement of this species is in
question. Distributed from the northern Philippines to southern Japan.
Centropyge vroliki (Bleeker, 1853)

En - Pearlscaled angelfish.
forms harems of 3 to 7 individuals. Frequently exported through the aquarium trade. Readily
hybridizes with both Centropyge eibli and C. flavissima at areas of sympatry. This species is
mimicked by juveniles of the acanthurid Acanthurus pyroferus. Distributed throughout the Indo-Ma-
layan region and in the Eastern Pacific Ocean to the Marshall Islands, northward to southern Japan,
southward to Lorde Howe Island.
Chaetodontoplus caeruleopunctatus Yasuda and Tominaga, 1976

En - Bluespotted angelfish.
Maximum total length about 14 cm. Depth distribution and habitat not known. Feeds on sponges
and tunicates. Rarely exported through the aquarium trade. Very little is known about this species.
Presently known only from the central Philippines.
3280 Bony Fishes
Chaetodontoplus chrysocephalus Bleeker, 1854

En - Orangefaced angelfish.
Maximum total length about 22 cm. Depth distribution and habitat not known. Feeds on sponges
and tunicates. Rarely exported through the aquarium trade. Specimens of this species may be the
adult male form of Chaetodontoplus septentrionalis, or might possibly represent hybrids. Distributed
from the Indo-Malayan region northward to southern Japan.
Chaetodontoplus conspicillatus (Waite, 1900)

En - Conspicuous angelfish; Fr -Chétodon armé à monocle.
Maximum total length about 34 cm. Inhabits coral reefs at depths of 20 to 40 m. Feeds on sponges
and tunicates; forms pairs. Rarely exported through the aquarium trade. Distributed from the
southern Great Barrier Reef and New Caledonia southward to Lord Howe Island.
Chaetodontoplus duboulayi (Günther, 1867)

En - Scribbled angelfish.
and tunicates; forms pairs or small groups. Occasionally exported through the aquarium trade.
Distributed from northern Australia and parts of Indonesia southward to Lord Howe Island.
Chaetodontoplus melanosoma (Bleeker, 1853)

En - Blackvelvet angelfish.
Maximum total length about 17 cm. Inhabits rocky and coral reefs at depths of 5 to 25 m. Feeds on
sponges and tunicates; solitary or in pairs. Frequently exported through the aquarium trade.
Distributed from the Indo-Malayan region and New Guinea northward to southern Japan.
Chaetodontoplus meredithi Kuiter, 1990

En - Meredith’s angelfish.
on sponges and tunicates; solitary, pairs, or small groups. Occasionally exported through the
aquarium trade. Distributed along the eastern coast of Australia southward to Lord Howe Island.
Chaetodontoplus mesoleucus (Bloch, 1787)

En - Vermiculated angelfish.
on sponges and tunicates; forms small groups. Frequently exported through the aquarium trade.
May consist of 2 separate species, 1 with an all-yellow tail, and 1 with a mostly grey tail. Distributed
throughout the Indo-Malayan region and western Pacific Ocean, northward to southern Japan,
southward to the east and west coasts of Australia.
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3282 Bony Fishes
Chaetodontoplus septentrionalis (Temminck and Schlegel, 1844)

En - Bluestripe angelfish.
Maximum total length about 20 cm. Inhabits rocky reefs at depths of 5 to 20 m. Feeds on sponges
and tunicates; solitary. Rarely exported through the aquarium trade. Distributed from the Malay
Peninsula northward to southern Japan.
Genicanthus bellus Randall, 1975

En - Ornate angelfish.
Feeds on plankton; forms harems of 3 to 7 individuals. Rarely exported through the aquarium trade.
The distribution of this species may be expanded with more deep reef exploration. Presently known
from Cocos-Keeling Island in the eastern Indian Ocean, the Philippines, Society Islands, Cook
Islands, Palau Islands, and a single specimen from Guam.
MALE
FEMALE
Genicanthus lamarck (Lacepède, 1802)

En - Lamarck’s angelfish.
Feeds on plankton; forms harems of 3 to 7 individuals. Frequently exported through the aquarium
trade. Distributed throughout the Indo-Malayan region eastward to Vanuatu, northward to southern
Japan, southward to the Great Barrier Reef.
MALE
FEMALE
Genicanthus melanospilos (Bleeker, 1857)

En - Swallowtail angelfish.
Feeds on plankton; forms harems of 3 to 7 individuals. Frequently exported through the aquarium
trade. Distributed throughout the Indo-Malayan region and southwestern Pacific Ocean to Fiji,
northward to the Ryukyu Islands.
3284 Bony Fishes
Genicanthus spinus Randall, 1975

En - Pitcairn angelfish.
Feeds on plankton; forms harems of 2 to 5 individuals. Almost never exported through the aquarium
trade. Distributed in southeastern Oceania from the Cook Islands to Ducie Atoll.
Genicanthus watanabei (Yasuda and Tominaga, 1970)

En - Watanabe’s angelfish.
Feeds on plankton; forms harems of 2 to 5 individuals. Occasionally exported through the aquarium
trade. Distributed in the western Pacific Ocean from the Ryukyu Islands southward to New
Caledonia and the Great Barrier Reef; also from the Cook Islands, Tuamotu Archipelago, and
Pitcairn.
Pomacanthus annularis (Bloch, 1787)

En - Blueringed angelfish.
on sponges and tunicates; forms pairs. Occasionally exported through the aquarium trade. Distrib-
uted from the east coast of Africa throughout Indonesia and New Guinea to New Caledonia,
northward to southern Japan.
Pomacanthus imperator (Bloch, 1787)

En - Emperor angelfish; Fr - Poisson ange impérial.
Maximum total length about 30 cm. Inhabits a wide variety of habitats at depths of 1 to 100 m. Feeds
on sponges and tunicates; forms pairs. Frequently exported through the aquarium trade. Juveniles
distinguished by a white dorsal-fin margin. Indian Ocean and Pacific Ocean populations may
represent separate species. Distributed throughout the Indo-Pacific region, excluding Easter Island,
Rapa, and the Marquesas Island; very rare in Hawaii.
Pomacanthus navarchus (Cuvier, 1831)

En - Bluegirdled angelfish.
and tunicates; solitary or in pairs. Occasionally exported through the aquarium trade. Distributed
throughout the Indo-Malayan region to the Solomon Islands and Great Barrier Reef.
Pomacanthus semicirculatus (Cuvier, 1831)

En - Koran angelfish; Fr - Poisson ange bleu.
and tunicates; solitary or in pairs. Frequently exported through the aquarium trade. Distributed
throughout the Indian Ocean and western Pacific Ocean, northward to southern Japan, eastward
to Samoa, and southward along the east coast of Australia.
3286 Bony Fishes
Pomacanthus sextriatus (Cuvier, 1831)

En - Sixbanded angelfish; Fr - Poisson ange à six bandes.
throughout the Indo-Malayan region to the Solomon Islands and east and west coasts of Australia,
northward to the Ryukyu Islands.
Pomacanthus xanthometapon (Bleeker, 1853)

En - Bluefaced angelfish.
throughout the central and eastern Indian Ocean and Indo-Malayan region, eastward to the Solomon
Islands, northward to the Ryukyu Islands, southward to New Caledonia.
Pygoplites diacanthus (Boddaert, 1772)

En - Regal angelfish; Fr - Poisson-ange duc.
and tunicates; solitary, pairs, or small groups. Frequently exported through the aquarium trade.
Distributed from the Red Sea throughout the Indian Ocean, eastward across the southern Pacific
Ocean to the Tuamotu Archipelago, northward to the Ryukyu Islands.
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Perciformes: Percoidei: Enoplosidae 3287
ENOPLOSIDAE
Old wife
by J.B. Hutchins
Enoplosus armatus (Shaw in White, 1790)

FAO names: En - Old wife.
D iagnostic characters: Body small to medium

sized (size to 31 cm), deep, and prominently
compressed. Head with obvious concavity over
eye, profile rising steeply to origin of dorsal fin. Eye
moderate in size. Preopercle serrated, with a
pair of enlarged spines at lower angle. Mouth
small, opening obliquely upwards, with small, fine
teeth in both jaws. Two completely separated
and highly elevated dorsal fins, first consisting
of VIII stout spines, second of I spine and 14
or 15 soft rays; anal fin similar in shape to second
dorsal fin, but not as elevated, consisting of III
spines and 14 or 15 soft rays; caudal fin large,
truncate to slightly lunate; pectoral fins moderately
long; pelvic fins large. Scales cycloid, very small;
lateral line prominently elevated anteriorly, de-
creasing in height posteriorly to midside of caudal
peduncle. Colour: head and body silvery white to
cream, with 7 or 8 black to dark brown cross-bands, bands 4 and 6 noticeably narrower, bands continued
onto dorsal, anal, and caudal fins; pelvic fins mostly dark brown with whitish leading edge.
Chaetodontidae: a single dorsal fin only (some members of Heniochus with a prominent notch between
spiny and rayed portions of fin); no strong spines at angle of preopercle.
Pomacanthidae: a single dorsal fin only; 1 strong spine at angle of operculum.
Scatophagidae: no strong spines at angle of preopercle; anal-fin spines IV.
notch
IV spines
Chaetodontidae Pomacanthidae Scatophagidae
Habitat, biology, and fisheries: Shallow coastal rocky reefs and seagrass beds; also found to depths of
100 m. Swims in small to large schools, but also occurs as solitary individuals. Feeds on crustaceans and
worms. Generally good to eat, but not marketed.
Distribution: Found only in the southern half of Austra-
lia, from Wide Bay in southern Queensland to Shark Bay
in Western Australia.
Reference
Gomon, M.F., J.C.M. Glover, and R.H. Kuiter. 1994. The fishes
of Australia’s south coast. Adelaide, State Print, 992 p.
3288 Bony Fishes
PENTACEROTIDAE
Armourheads
by P.C. Heemstra
A single species occurring in the area.
Histiopterus typus Temminck and Schlegel, 1844

Frequent synonyms / misidentifications: Histiopterus spinifer Gilchrist, 1904 / None.
FAO names: En - Sailfin armourhead; Fr - Matode voile; Sp - Botellón velero.
juvenile
adult
(after Heemstra in Smith and Heemstra, 1986)
Diagnostic characters: Body oblong-oval and moderately compressed, its depth greater than head
length. Head mostly naked, the bones exposed and rugose; a patch of small embedded scales below
eye; head of small juveniles with several small spines and knobs; preopercle edge very spiny in young,
entire or with small spines in adults; opercle without spines in adults. Snout profile concave; eye well above
horizontal line through upper jaw symphysis. Mouth small, terminal upper jaw slightly protrusible; maxilla
naked, mostly covered by preorbital; jaws with bands of small conical teeth; no supramaxilla; vomer with
or without teeth, no teeth on palatines. Branchiostegal membranes separate, free from isthmus; bran-
chiostegal rays 7. Gill slits 4, a slit behind the last; gill rakers short, first gill arch with 3 to 6 rakers on upper
limb, 14 to 16 on lower limb. Dorsal fin high, sail-like; dorsal-fin spines IV, the first 2 short, the third
and fourth much enlarged, longer than head, subequal to first soft ray; dorsal-fin rays 23 to 29; anal fin
with III spines (second longer than third) and 8 to 10 soft rays; caudal fin slightly emarginate; pectoral-fin
rays 16 to 18; pelvic fins with I strong spine and 5 soft rays, but no scaly axillary process. Scales small to
moderate, ctenoid; lateral line complete, strongly curved over pectoral fins; lateral-line pores 58 to 70.
Vertebrae 13+12. Swimbladder large. Pyloric caeca numerous. Colour: adult body pale, with dark brown
bars: first from dorsal-fin origin through pectoral-fin base to pelvic-fin base, second bar from anterior
dorsal-fin rays to belly, and third from base of fifth to seventh dorsal-fin rays to base of anal-fin
spines. Juveniles pale; large dark brown blotches over head, body, and dorsal, anal, and pelvic fins;
pectoral and caudal fins pale, without spots.
Perciformes: Percoidei: Pentacerotidae 3289

Chaetodontidae, Pomacanthidae, Drepanidae, Ephippidae, and Scatophagidae: head covered with scales;
anal-fin rays 14 or more (most pentacerotids with fewer than 14 anal-fin rays).
Chaetodontidae Pomacanthidae Drepanidae
Ephippidae Scatophagidae
Size: Maximum standard length 35 cm; commonly to 20 cm.
Habitat, biology and fisheries: Usually found inshore on reefs or on rough bottom of the outer continental
shelf and upper slope at depths of 40 to 400 m. Biology little known. Mostly caught with trawls, occasionally
with hook-and-line.
tropical waters of the Indo-
West Pacific region from
South Africa to Red Sea,
Oman, Arabian Sea, north-
western Australia, Philip-
pines, Taiwan Province of
China, and Japan.
Reference
Hardy, G.S. 1983. A revision of the fishes of the family Pentacerotidae. New Zeal. J. Zool., 10:177-220.
3290 Bony Fishes
KYPHOSIDAE
Sea chubs
by K. Sakai
D iagnostic characters: Medium-sized fishes (to 75 cm); body oblong or elliptical, moderately com-
pressed. Head small. Snout short. Posteroventral corner of preopercle serrate. Opercle with 2 small,
weak retrorse spines. Eye moderately small, its diameter shorter than snout length. Mouth small and
terminal, single row of lanceolate incisor-like teeth on both jaws; minute canine-like teeth in 2 or 3
rows medial to incisor-like teeth; palatines, vomer, and tongue with a band of villiform teeth. Preorbital
narrow, covering little of maxilla; maxilla barely reaching eye. First gill arch with 5 to 10 gill rakers on
upper limb, 14 to 24 on lower limb (total 20 to 34). Dorsal fin continuous, beginning above origin of
pelvic fins, with XI spines and 12 to 15 soft rays; anal fin beginning slightly behind middle of body, with III
spines and 11 to 14 soft rays; pectoral fins bluntly pointed posteriorly, with 16 to 20 rays, slightly longer
than pelvic fins; pelvic fins beginning a little behind pectoral-fin base; caudal fin more or less forked, with
pointed lobes. Scales ctenoid and not deciduous, extending onto most of soft portions of dorsal and
anal fins and proximal part of caudal fin. Pored scales in lateral line 49 to 55; longitudinal scale
rows 50 to 71. Scales above lateral line to origin of dorsal fin 11 to 15, scales below lateral line to origin
of anal fin 18 to 24. Colour: body bluish, bluish brown, or silvery grey dorsally and dusky grey or silvery
ventrally; distinct longitudinal lines on sides.
dorsal fin with XI spines and 12-15 soft rays
pored scales in
lateral line 49-55
single row of
lanceolate
incisor-like teeth
on both jaws
caudal fin
more or less
forked
longitudinal scale
rows 50-71
anal fin with III spines and
11-14 soft rays
Habitat, biology, and fisheries: Occur on rocky and coral reefs in tropical and temperate waters.
Herbivorous, feeding primarily on benthic algae. Schooling, sometimes in groups composed of several
species of kyphosids. Juveniles often occur far out at sea beneath floating debris and seaweeds. Highly
valued as food. Taken by handline, gill net, and spear.

Scorpididae: small lanceolate conical teeth in both jaws; usually smaller and weaker ctenoid scales; more
soft rays in dorsal and anal fins (dorsal- and anal-fin rays more than 24 versus less than 16 in kyphosids);
body laterally compressed, caudal peduncle low and narrow; caudal fin well forked.
Girellidae: outer teeth tricuspid in both jaws; more dorsal-fin spines (XIII to XVI dorsal-fin spines versus
XI in kyphosids); caudal peduncle deeper; caudal fin slightly emarginate.
Scorpididae Girellidae
Perciformes: Percoidei: Kyphosidae 3291
Key to the species of Kyphosidae occurring in the area

1a. Dorsal fin with 15 and anal fin with 14 soft rays; body depth 30 to 35% of standard length;
caudal fin distinctly forked (Fig. 1) . . . . . . . . . . . . . . . . . . . . . . . . . . Sectator ocyurus
1b. Dorsal fin with less than 15 and anal fin with less than 14 soft rays; body depth 35 to 50%
of standard length; caudal fin moderately forked . . . . . . . . . . . . . . . . . . . . (Kyphosus) ® 2
2a. Dorsal fin with 14 and anal fin with 13 soft rays (Fig. 2) . . . . . . . . . . . . . . Kyphosus vaigiensis
2b. Dorsal fin with 12 and anal fin with 11 soft rays . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
Fig. 1 Sectator ocyurus Fig. 2 Kyphosus vaigiensis
3a. Anterior soft portion of dorsal and anal fin elevated (Fig. 3a); total gill rakers on first gill
arch 26 to 31 (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kyphosus cinerascens
3b. Anterior soft portion of dorsal and anal fin not elevated (Fig. 3b); total gill rakers on first
gill arch less than 25 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
a) b)
Fig. 3 Fig. 4 Kyphosus cinerascens
4a. Longitudinal scale rows 52 to 58; total gill rakers on first gill arch 20 or 21; pectoral-fin
rays usually 18 (rarely 16 or 17) (Fig. 5) . . . . . . . . . . . . . . . . . . . . . Kyphosus sydneyanus
4b. Longitudinal scale rows 58 to 69; total gill rakers on first gill arch 21 to 24; pectoral-fin
rays usually 19 (rarely 18 or 20) (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . Kyphosus bigibbus
Fig. 5 Kyphosus sydneyanus Fig. 6 Kyphosus bigibbus

0 Kyphosus bigibbus Lacepède, 1801
0 Kyphosus cinerascens (Forsskål, 1775)
0 Kyphosus sydneyanus (Günther, 1886)
0 Kyphosus vaigiensis (Quoy and Gaimard, 1825)
0 Sectator ocyurus (Jordan and Gilbert, 1882)
Reference
Sakai, K. and T. Nakabo. 1995. Taxonomic review of the Indo-Pacific kyphosid fishes, Kyphosus vaigiensis (Quoy and
Gaimard). Japan. J. Ichthyol., 42(1):61-70.
3292 Bony Fishes
Kyphosus bigibbus Lacepède, 1801

Frequent synonyms / misidentifications: Kyphosus fuscus Lacepède, 1801 / None.
FAO names: En - Brown chub.
Diagnostic characters: Body depth 39 to 49%, its width 17 to 21% of standard length; caudal peduncle
slightly deep. Head length 25 to 31% of standard length, dorsal contour of snout slightly steep. Snout short,
its length longer than eye diameter. Mouth terminal, number of lanceolate incisor-like teeth about 34 to 38
in adults (over 20 cm standard length). First gill arch with 5 to 7 (usually 6) gill rakers on upper limb,
15 to 17 (usually 16 or 17) on lower limb (total 21 to 24, usually 22 or 23). Dorsal fin with XI spines,
the sixth or seventh spine longest, and 12 (rarely 11 or 13) soft rays; base of spinous portion longer
than base of soft portion; soft portion of dorsal fin low, the margin straight; anterior soft rays of dorsal fin
a little shorter than longest dorsal-fin spine. Anal fin with III spines, the third spine longest, with 11
(rarely 10 or 12) soft rays, anterior soft portion not elevated; longest anal-fin ray less than 2 times
length of third anal-fin spine. Pectoral fins with 19 (rarely 18 or 20) rays. Pelvic fins as long as pectoral
fins. Caudal fin shallowly forked with pointed lobes. Pored scales in lateral line 51 to 55 (usually 53);
longitudinal scale rows 58 to 69 (usually 63 to 65); scales above lateral line 12 to 15 (usually 13 or
14), scales below lateral line 20 to 25 (usually 21 or 22). Colour: body bluish brown dorsally, dusky
grey ventrally, with several brown or bluish brown longitudinal lines on flank, large specimens (about 50 cm
standard length) uniformly dusky blue or dark brown dorsally; head with 2 oblique brown or bluish brown
bands, one through eye, the other one below eye; dorsal and anal fins dark blue, the margin of soft portion
with a darker band; pectoral fins silver brown near base, the distal half slightly darker.
Habitat, biology, and fisheries: Inhabits coastal areas, primarily in coral and rocky reefs, often occurring
in the inner reefs. Herbivorous. Occasionally schools in groups with other kyphosid fishes. Juveniles often
occur far out at sea beneath floating debris and seaweed. Caught by gill nets, handlines, and spears.
the tropical Indo-West Pacific
from South Africa, including
the Red Sea, to Australia;
north to southern Japan.
Kyphosus cinerascens (Forsskål, 1775)

FAO names: En - Blue sea chub; Fr - Calicagère bleue; Sp - Chopa axul.
deep. Head length 26 to 34% of standard length, dorsal contour of snout slightly steep. Snout increasing
with growth, its length as long as or shorter than eye diameter to about 15 cm standard length, thereafter,
longer; number of lanceolate incisor-like teeth about 40 to 45 in adults (over 20 cm standard length). First
gill arch with 7 to 9 (usually 8 or 9) gill rakers on upper limb, 19 to 22 (usually 20 or 21) on lower
limb (total 26 to 31, usually 28 to 30). Dorsal fin with XI spines, the sixth or seventh spine longest,
and 12 (rarely 11 or 13) soft rays; base of spinous portion of dorsal fin longer than base of soft portion;
soft portion of dorsal fin well elevated, anterior soft rays clearly longer than longest dorsal-fin spine;
anal fin with III spines, the third spine longest, with 11 (rarely 10 or 12) soft rays, anterior soft
portion well elevated, similar to soft portion of dorsal fin; longest anal-fin ray 2.5 times longer than
third anal-fin spine. Pectoral fins with 18 (rarely 19) soft rays. Pelvic fins as long as pectoral fins. Caudal
fin shallowly forked with pointed lobes. Pored scales in lateral line 49 to 52 (usually 50 or 51);
longitudinal scale rows 50 to 57 (usually 53 to 56); scales above lateral line 9 to 12 (usually 10 or
11) scales below lateral line 17 to 21 (usually 18 to 20). Colour: body bluish dorsally, dusky greyish
ventrally, with several bluish brown longitudinal lines on flank, large specimens (about 40 cm standard
length) uniformly dusky or dark blue dorsally; head with 2 oblique brown or bluish brown bands; one through
eye, the other one below eye; dorsal and anal fins dark blue, margin of soft portion with darker band;
pectoral fins silver blue near base, distal half slightly darker.
Habitat, biology, and fisheries: Inhabits coastal areas, primarily coral and rocky reefs, often occurring
on inner reefs. Herbivorous. Occasionally schools in groups with other kyphosid fishes. Juveniles often
occur far out at sea beneath floating debris and seaweed. Caught by gill nets, handlines, and spears.
South Africa, including the
Red Sea, to Australia; north to
southern Japan.
3294 Bony Fishes
Kyphosus sydneyanus (Günther, 1886)

FAO names: En - Silver drummer.
slightly deep. Head length 24 to 30% of standard length, dorsal contour of snout before eye steep. Snout
short, its length longer than eye diameter. Mouth terminal, number of lanceolate incisor-like teeth about
40 to 50 in adults (over 20 cm standard length). Gill rakers on first gill arch 5 or 6 on upper limb, 14 or
15 on lower limb (total 20 or 21). Dorsal fin with XI (rarely X) spines, the sixth or seventh spine
longest, and 12 soft rays; base of spinous portion of dorsal fin longer than base of soft portion; soft portion
of dorsal fin low, the margin straight, anterior soft rays as long as or a little shorter than longest dorsal-fin
spine; anal fin with III spines, third spine longest, with 11 soft rays, anterior soft portion not
elevated; longest anal-fin ray less than 2 times length of third anal-fin spine. Pectoral fins with 17 or 18
soft rays. Pelvic fins as long as pectoral fins. Caudal fin shallowly forked with bluntly pointed lobes. Pored
scales in lateral line 49 to 53 (usually 50 or 51); longitudinal scale rows 52 to 58; scales above lateral
line 10 to 12; scales below lateral line 17 to 19. Colour: body dark silvery grey or dusky grey dorsally,
silver ventrally, with several olive brown or brown longitudinal lines on flank; head with 2 oblique olive
brown or brown bands; one before or behind eye, and the other one below eye; dorsal and anal fins dark
grey, the margin of soft portion with a darker band; pectoral fins silver near base, distal half slightly darker.
Habitat, biology, and fisheries: Inhabits coastal areas, primarily in shallow rocky reefs, often occurring
in bays and harbours. Herbivorous. May school with other kyphosid fishes in the northern part of its range.
Juveniles often occur far out at sea beneath floating debris and seaweeds. Caught by gill nets, handlines,
and spears.
Distribution: Southern part
of Australia; reasonably com-
mon in New South Wales,
South Australia, and Western
Australia, also found in
Queensland, Victoria, Tas-
mania, northern New Zea-
land, Lord Howe Island, and
Norfolk Islands.
Kyphosus vaigiensis (Quoy and Gaimard, 1825)

Frequent synonyms / misidentifications: Kyphosus lembus (Cuvier, 1831); K. gibsoni Ogilby, 1912 /
None.
FAO names: En - Brassy chub.
low. Head length 24 to 35% of standard length, dorsal contour of snout steep. Snout short, its length as
long as or slightly longer than eye diameter. Mouth terminal, number of lanceolate incisor-like teeth about
30 to 34 in adults (over 20 cm standard length). Gill rakers on first gill arch 8 to 10 (usually 9) on upper
limb, 21 to 24 (usually 22 or 23) on lower limb (total 29 to 34, usually 31 or 32). Dorsal fin with XI
(rarely X) spines, the sixth or seventh spine longest, and with 14 (rarely 13 or 15) soft rays; base of
spinous portion of dorsal fin longer than base of soft portion; anterior soft rays of dorsal fin a little shorter
or as long as longest dorsal-fin spine. Anal fin with III spines, third spine longest, with 13 (rarely 12)
soft rays, anterior soft portion not elevated, the longest soft ray 2 times longer than third anal-fin spine.
Pectoral fins with 19 (rarely 18 or 20) soft rays. Pelvic fins slightly shorter than pectoral fins. Caudal fin
shallowly forked with pointed lobes. Pored scales in lateral line 51 to 55 (usually 52 or 53); longitudinal
scale rows 56 to 64; scales above lateral line 11 to 14 (usually 12 or 13); scales below lateral line
18 to 21 (usually 18 or 19). Colour: body bluish brown dorsally, silver ventrally, with several olive brown
or yellow longitudinal lines on flank, large specimens (about 50 cm standard length) uniformly dark brown
dorsally; head with 2 oblique olive brown or yellow bands; one before or behind eye, and the other one
below eye; dorsal and anal fins dark blue, margin of soft portion with darker band; pectoral fins silver near
base, distal half slightly darker.
Habitat, biology, and fisheries: Inhabits coastal areas, primarily in coral and rocky reefs, often occurring
in inner reefs. Herbivorous. Occasionally schools with other kyphosid fishes. Juveniles often occur far out
at sea beneath floating debris and seaweed. Caught by gill nets, handlines, and spears.
South Africa, including the
Red Sea, to Australia; north to
southern Japan.
3296 Bony Fishes
Sectator ocyurus (Jordan and Gilbert, 1882)

FAO names: En - Bluestriped chub.
Diagnostic characters: Body oblong and moderately compressed, its depth 33 to 35%, its width 12
to 15% of standard length; caudal peduncle low, its depth 8 to 9% of standard length. Head small,
25 to 27% of standard length, dorsal contour of head prominently convex, interorbital space broad. Eye
small. Snout short, its length slightly longer than eye diameter. Mouth small and terminal; posterior end
of maxilla not reaching eye; single row of small lanceolate incisor-like teeth. Preorbital narrow,
covering little of maxilla. Gill rakers on first gill arch usually 9 on upper limb, 20 or 21 on lower limb
(total 29 or 30). Dorsal fin with XI (rarely X) spines, the sixth or seventh spine longest, with 15 (rarely
16) soft rays, base of spinous portion shorter than base of soft portion; soft portion of dorsal fin low and
straight; anterior soft rays clearly shorter than longest dorsal-fin spine. Anal fin with III spines, third
spine longest and with 14 soft rays, anterior part of soft portion not elevated, its height about equal
to soft portion of dorsal fin. Pectoral fins small, bluntly pointed posteriorly, with 19 or 20 soft rays. Pelvic
fins shorter than pectoral fins. Caudal fin well forked with pointed lobes. Pored scales in lateral line 57
to 61; longitudinal scale rows 64 to 73; scales above lateral line 14 to 16; scales below lateral line
22 to 24. Colour: a longitudinal cobalt blue stripe from slightly before dorsal-fin origin to upper part of
caudal-fin base, a distinct broad cobalt blue stripe from area above base of pectoral fins straight to base
of caudal fin, area between these 2 blue stripes dark grey; a golden yellow stripe below broad cobalt blue
stripe; head with 2 oblique cobalt blue bands; 1 from snout through upper margin of eye to upper part of
gill opening; the other one from angle of mouth through lower margin of eye to posterior end of opercle;
interspace and space below blue bands of head golden; anal and pelvic fins golden yellow; dorsal, pectoral,
and caudal fins blackish golden yellow.
Habitat, biology, and fisheries: A rare species in the area. Juveniles often occur far out at sea beneath
floating debris and seaweed.
Caught by gill nets or handli-
nes.
Distribution: Scattered in the
Pacific: Hawaii and Society Is-
lands; common in the eastern
Pacific from southern Califor-
nia to Ecuador; north to the Izu
Islands in Japan. Known in the
Central and West Pacific only
from Hawaii, Izu Islands, Ja-
pan, and a single specimen
from Moorea, Society Islands.
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Perciformes: Percoidei: Girellidae 3297
GIRELLIDAE
Nibblers
by K.E. Carpenter
D iagnostic characters (for species occurring in the area): Medium-sized fishes (to 76 cm); body oblong.
Mouth small, not extending posteriorly beyond eye; maxilla covered by suborbital; outer teeth in
jaws mostly tricupsid incisors. Single dorsal fin with XIII to XVI spines and 11 to 14 soft rays. Anal
fin with III spines and 11 or 12 soft rays. Pelvic fins inserted just under or distinctly just posterior to
pectoral-fin base. Caudal fin truncate to broadly forked. Scales small, ctenoid, extending onto cheeks and
upper part of opercles, and bases of dorsal and anal fins but not covering most of soft portion of dorsal
and anal fins. Lateral line continuous, following upper profile of body. Colour: uniformly dark bluish grey
to greenish grey above and silvery below, sometimes with thin dark bars on upper sides.
Habitat, biology, and fisheries: Coastal and estuarine, mostly on hard substrate bottoms that support
algal growth, from the intertidal zone to a depth of about 25 m. Nibblers feed on encrusting and floating
algae and incidentally on small crustaceans. Caught primarily by anglers. Girrelids are warm-temperate
and subtropical fishes that are mostly excluded from the area except at the southernmost fringe in Australia.
Girella punctata is a northwestern Pacific subtropical species that occurs as far south as Batan Island in
the Philippines, which is just north of the area.
Remarks: Girellids are combined under the Kyphosidae by some authors, sometimes together with
Microcanthidae and Scorpididae.
Kyphosidae: rear tip of maxilla exposed (covered by suborbital
in Girrellidae); dorsal fin with XI spines (XII to XVI in Girrellidae);
scales extend onto most of soft portion of dorsal and anal fins
(only basally in Girrellidae).
Scorpididae: no incisiform teeth (outer teeth mostly incisiform in
Girrellidae); spinous part of dorsal fin much lower than soft part
of dorsal fin; dorsal fin with IX or X spines (XII to XVI in
Girrellidae); scales extend onto most of soft portion of dorsal and
anal fins (only basally in Girrellidae). Kyphosidae
Microcanthidae: close-set brush-like teeth; dorsal fin with XI or
XII spines and 16 to 18 soft rays (XII to XVI spines and 11 to 14
soft rays in Girrellidae); anal fin with 16 to 19 soft rays (11 or 12
soft rays in Girrellidae).
Scorpididae Microcanthidae
3298 Bony Fishes
Key to the species of Girellidae occurring in the area

1a. Dorsal fin with XIII spines and 14 soft rays; pelvic fins inserted nearly just below bases
of pectoral fins; body uniformly dark . . . . . . . . . . . . . . . . . . . . . . . . . . . Girella elevata
1b. Dorsal fin with XIV to XVI spines and 11 or 12 soft rays; pelvic fins inserted distinctly
posterior to bases of pectoral fins; around 11 narrow bars along upper sides . . .Girella tricuspidata

0
0 Girella elevata Macleay, 1881
0 Girella tricuspidata (Quoy and Gaimard, 1824)
References
Gomon, M.F., J.C.M. Glover, and R.H. Kuiter (eds). 1994. The fishes of Australia’s south coast. Adelaide, State Print,
992 p.
Kuiter, R.H. 1993. Coastal fishes of south-eastern Australia. Honolulu, University of Hawaii Press, 437 p.
Girella elevata Macleay, 1881

En - Black drummer.
Maximum total length 76 cm. Typically schooling on rocky reefs from exposed shorelines to a depth
of around 25 m. Confined to the eastern coast of Australia from southern Queensland to Wilsons
Promontory in Victoria.
(after Hutchins and Swainston, 1986)
Girella tricuspidata (Quoy and Gaimard, 1824)

En - Luderick.
Maximum total length 62 cm. Schooling on rocky reefs and around hard structures in estuaries and
coastal areas to a depth of around 20 m. In Australia from southern Queensland to central southern
Australia, northern Tasmania, and northern New Zealand.

Perciformes: Percoidei: Scorpididae 3299
SCORPIDIDAE
Halfmoons
by K.E. Carpenter
Scorpis lineolata Kner, 1865

Frequent synonyms / misidentifications: Scorpis lineolatus Kner, 1865 / Scorpis aequipinnis
Richardson, 1848.
FAO names: En - Silver sweep.
(after Kuiter, 1993)
Diagnostic character: Body compressed and deep, its depth 1.8 to 2 times in standard length. Head
length 3.1 to 3.7 times in standard length; eyes 2.9 to 3.7 times in head length. Mouth terminal, oblique,
and small, its posterior end just reaching anterior margin of eye; maxilla exposed; strong conical teeth
in bands, the outer row enlarged and recurved. Single dorsal fin with IX or X spines and 26 to 28 soft
rays, the spines about 1/2 height of soft rays. Anal fin with III spines and 27 or 28 soft rays. Caudal
fin broadly forked. Pelvic fins inserted distinctly just posterior to pectoral-fin base. Scales small, ctenoid,
extending onto and covering most of soft portion of dorsal and anal fins. Lateral line continuous, gently
curved. Colour: mostly greyish, darker above and whitish below; upper opercular margin with a thin black
edge; base of pectoral fins black; upper and lower margins of caudal-fin lobes blackish.

Kyphosidae: incisiform teeth in jaws (no incisiform teeth in Scorpididae); dorsal fin with 12 to 15 soft rays
(26 to 28 in Sorpididae), dorsal-fin spines not noticeably shorter than soft rays (noticeably shorter in
Scorpididae); anal fin with 11 to 14 soft rays (27 or 28 in Scorpididae).
Girellidae: outer teeth mostly incisiform (no incisiform teeth in Scorpididae); spinous part of dorsal fin not
much lower than soft part of dorsal fin; dorsal fin with XII to XVI spines and 11 to 14 soft rays (26 to 28 in
Scorpididae); anal fin with 11 or 12 soft rays (27 or 28 in Scorpididae); scales not extending onto most of
soft portion of dorsal and anal fins (extend onto most of soft portion of dorsal and anal fins in Scorpididae).
Kyphosidae Girrellidae
3300 Bony Fishes
Microcanthidae: close-set brush-like teeth (strong conical

teeth in Scorpididae); spinous part of dorsal fin generally
longer than soft part of dorsal fin (spinous portion notice-
ably shorter in Scorpididae); dorsal fin with 16 to 18 soft
rays (26 to 28 in Scorpididae); anal fin with 16 to 19 soft
rays (27 or 28 in Scorpididae); scales not extending onto
most of soft portion of dorsal and anal fins (extend onto
most of soft portion of dorsal and anal fins in Scorpididae).
Size: Maximum total length to 37 cm.
Habitat, biology, and fisheries: Coastal and estuarine,
mostly on rocky reefs to a depth of about 30 m. Occurs in
large schools. Feeds on zooplankton and algae in midwa- Microcanthidae
ter. Caught mostly by anglers.
Distribution: S o u t h e a s t e r n
Australia, Lord Howe Island,
and northern New Zealand.
Remarks: Scorpidids are com-
bined under the Kyphosidae by
some authors, sometimes to-
gether with Microcanthidae
and Girellidae.
References
992 p.
Perciformes: Percoidei: Microcanthidae 3301
MICROCANTHIDAE
Microcanthids
by K.E. Carpenter
D iagnostic character: Moderate-sized fishes (to 25 cm); body ovate to high. Mouth terminal, small,
not extending posteriorly beyond anterior margin of eye; teeth small, close-set, and pointed, in a
broad band or a single row. Single dorsal fin with XI or XII spines and 16 to 18 soft rays. Anal fin with
III spines and 16 to 19 soft rays. Pelvic fins inserted distinctly just posterior to pectoral-fin base. Caudal
fin emarginate to forked. Scales small, ctenoid, extending onto cheeks and upper part of opercles, and
forming a scaly sheath at bases of dorsal and anal fins and extending onto soft portion of dorsal and
anal fins. Lateral line continuous, smoothly curved. Colour: yellowish or whitish with dark brown or
blackish nearly horizontal or oblique stripes.
Habitat, biology, and fisheries: Shallow coastal and rocky estuarine, mostly on hard substrate, from the
intertidal zone to a depth of about 30 m. Feed on small invertebrates and algae. Of little interest to fisheries
except in the aquarium trade. Microcanthids are warm-temperate and subtropical fishes that are mostly
excluded from the area except at the southernmost fringe in Australia.
Remarks: Microcanthids are combined under the Kyphosidae by some authors, sometimes together with
Microcanthidae and Scorpididae; microcanthids are also sometimes included under the Scorpididae.

Kyphosidae: teeth incisor-like (close-set pointed teeth in Microcanthidae); dorsal fin with 12 to 15 soft rays
(16 to 18 in Microcanthidae); anal fin with 11 to 14 soft rays (16 to 19 in Microcanthidae).
Girellidae: outer teeth mostly tricuspid incisors (close-set pointed teeth in Microcanthidae); dorsal fin with
XIII to XVI spines and 11 to 14 soft rays (XI or XII spines and 16 to 18 soft rays in Microcanthidae); anal
fin with 11 or 12 soft rays (16 to 19 soft rays in Microcanthidae).
Kyphosidae Girellidae
3302 Bony Fishes
Scorpididae: spinous part of dorsal fin much lower than soft part of dorsal fin; dorsal fin with IX or X spines
and 26 to 28 soft rays (XI or XII spines and 16 to 18 soft rays in Microcanthidae); anal fin with 27 or 28
soft rays (16 to 19 soft rays in Microcanthidae).
Chaetodontidae: caudal fin typically truncate or rounded (emarginate or forked in Microcanthidae); pelvic
fins inserted just below or nearly just below pectoral-fin base (distinctly behind pectoral-fin base in
Microcanthidae).
Scorpididae Chaetodontidae
Key to the species of Microcanthidae occurring in the area

1a. Lateral stripes nearly horizontal; lateral-line scales 70 to 78 . . . . . . . . . . . Atypichthys strigatus
1b. Lateral stripes distinctly oblique; lateral-line scales 56 to 60 . . . . . . . . . . Microcanthus strigatus

0 Atypichthys strigatus (Günther, 1860)
0 Microcanthus strigatus (Cuvier, 1831)
References
992 p.
Perciformes: Percoidei: Microcanthidae 3303
Atypichthys strigatus (Günther, 1860)

En - Australian Mado.
Maximum total length about 25 cm. Schooling in coastal waters and estuaries typically over hard
substrate, from the intertidal zone to a depth of about 30 m. Southern Queensland to Tasmania and
Lord Howe Island.
Microcanthus strigatus (Cuvier, 1831)

En - Stripey.
Maximum total length 16 cm. In small to large aggregations in shallow rocky coastal and estuarine
areas. Japan to Taiwan Province of China and Hawaii in the northern hemisphere and in the
southern hemisphere in southeastern Australia from the Capricorn Group to southern New South
Wales.
(after Randall, Allen, and Steene, 1990)

3304 Bony Fishes
ARRIPIDAE
Australian salmons
by J.B. Hutchins
Arripis trutta (Forster in Bloch and Schneider, 1801) ASA

Frequent synonyms / misidentifications: None / Arripis georgianus (Valenciennes, 1831).
FAO names: En - Australian salmon; Fr - Salmon australien; Sp - Salmón de Australia.
Diagnostic characters: Body medium sized (to about 1 m), robust and moderately elongate (streamlined);
caudal peduncle long and slender. Head conical with oblique terminal mouth, maxilla reaching below
centre of eye. Eye small with prominent adipose tissue along front and rear margins in large fish. Teeth
small, pointed, narrow bands in each jaw. Scales moderately small, very finely ctenoid, smooth to touch,
covering body and head except lower jaw, snout, and above eye; lateral line nearly straight. Single dorsal
fin with long base and low notch between spinous and rayed protions; spinous portion with IX spines,
noticeably higher than soft portion with 15 to 18 soft rays; anal fin small, with III spines and 9 or
10 soft rays; caudal fin deeply forked; pelvic fins relatively short, but slightly longer than pectoral fins.
Total gill rakers on first gill arch 33 or 40. Colour: upper portions of head and body bluish green, shading
to silvery white below; juveniles with series of vertical
golden bars or rows of spots on body; spots in adult
relatively smaller, darker, more irregularly placed, and
confined to upper sides; pectoral fins bright yellow, black-
ish at base, other fins clear, caudal fin with blackish
posterior border.
Caesionidae: dorsal-fin spines X or more; no distinct
notch between spinous and soft-rayed portions of dorsal Caesionidae
fin.
Lutjanidae: dorsal-fin spines X or more.
Size: Maximum total length about 1 m.
Habitat, biology, and fisheries: Shallow coastal wa-
ters, often entering estuaries. Swims in large schools
close to the shoreline, feeding on zooplankton and small
fishes. Preyed upon by larger carnivores especially
sharks. Popular recreational fishes that are commercially
important in temperate areas, but not marketed in the Lutjanidae
Western Central Pacific.
Distribution: Southern Australia, ranging as far north on
the east coast as southern Queensland. Also found in
New Zealand.
Reference
Paulin, C.D. 1993. Review of the Australian fish family
Arripididae (Percomorpha). Aust. J. Mar. Freshwater
Res., 44:459-471.
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Perciformes: Percoidei: Terapontidae 3305
TERAPONTIDAE
(= Therapontidae, Theraponidae, Teraponidae)
Terapon-perches (terapon-grunters)
by R.P. Vari
D iagnostic characters: Small to medium-sized (to 35 cm) perch-like fishes; body oblong to oblong-
ovate, slightly to moderately compressed laterally. Mouth moderate, protractile, with gape often
oblique, sometimes horizontal; jaws typically equal, sometimes upper jaw longer, the upper jaw not
extending beyond centre of orbit; jaw teeth usually in villiform bands (reduced to 2 or 3 distinct rows in
some species), with teeth of outer row distinctly larger than those in remainder of band; shape of jaw teeth
conical, flattened, or tricuspidate; teeth also present on roof of mouth (vomer and palatines) in juveniles
of many species, lacking in adults of most species. Posterior and ventral margins of preopercle variably
serrate, serrations more prominent in juveniles, usually more developed on vertical margin; first infraorbital
(lacrimal) with serrations along ventral margin, serrations less developed with age in some species;
opercle with 2 spines posteriorly, the lower one larger and stronger, extending beyond margin of
opercular lobe in some species. Posttemporal bone exposed and expanded posteriorly in some species
with posterior margin serrate. Dorsal fin single, arched, with XI to XIV spines and 8 to 14 soft rays,
fourth to seventh dorsal-fin spines longest, those following decreasing in length to penultimate spine which
is much shorter than ultimate spine in some species, this resulting in a notched spinous dorsal fin; anal fin
with III variably strong spines and 7 to 12 soft rays; pelvic-fin base located behind vertical line through
base of pectoral fins; caudal fin usually emarginate (truncate or rounded in some entirely fresh-water
species). Lateral line single and complete. Scales adherent, finely ctenoid (rough to touch). Colour: body
tan or light grey, often silvery in life with various dark markings; most marine species with 3 or more dark,
straight or downwardly curved longitudinal stripes on body; many marine species with dark transverse
bands on lobes of caudal fin.
dorsal fin with XI to XIV spines
and 8 to 14 soft rays
2 opercular
spines

and 7-12 soft rays
3 opercular spines
Habitat, biology, and fisheries: Marine terapontids inhabit
inshore marine and brackish waters, with some species also
entering hypersaline and fresh waters. Many terapontid spe-
cies limited to fresh waters of Australia and New Guinea.
Marine members of family are good food fishes entering
catches largely by artisanal and other inshore fisheries. None
of the species is important enough to support a special fishery.
Serranidae
Similar families occurring in the area X dorsal-fin
spines
Serranidae: mouth large, with upper jaw typically reaching to
below vertical through posterior margin of eye (not extending
beyond vertical though centre of eye in Terapontidae); caudal
fin typically rounded (usually emarginate in marine Teraponti-
dae); 3 strong spines on opercle (2 spines in Terapontidae).
Kuhliidae: dorsal fin with X spines (XI to XIV in Terapontidae).
Kuhliidae
3306 Bony Fishes
Key to the species of Terapontidae occurring in the area

1a. Posttemporal bone covered with skin and scales, not expanded posteriorly, its posterior
margin not serrate (Fig. 1a) . . . . . . . . . . . . . . . . . . . . . . . . . (Amniataba, Pelates) ® 2
1b. Posttemporal bone expanded posteriorly with skin and scale covering reduced, posterior
margin serrate (Fig. 1b) . . . . . . . . . . . . . . . . . (Terapon, Rhyncopelates, Mesopristes) ® 4
2a Lateral-line scales 46 to 54; spinous portion of dorsal fin without distinct patch of black
pigmentation anteriorly; body lacking distinct, dark, longitudinal stripes; lobes of caudal
fin with prominent oblique transverse black pigmentation . . . . . . . . . . Amniataba caudavittatus
2b Lateral-line scales 66 to 87; distinct patch of black pigmentation on anterior portion of
spinous dorsal fin; body with distinct, dark, longitudinal stripes; lobes of caudal fin without
prominent oblique transverse black pigmentation . . . . . . . . . . . . . . . . . . . . . (Pelates) ® 3
3a. Teeth tricuspidate; gill rakers (including rudiments) on first gill arch 6 or 7 on upper limb,
14 or 15 on lower limb; lateral-line scales 76 to 87 . . . . . . . . . . . . . . . . . Pelates sexlineatus
3b. Teeth lacking side cusps; gill rakers (including rudiments) on first gill arch 16 to 18 on
upper limb, 22 to 27 on lower limb; lateral-line scales 66 to 75 . . . . . . . . . Pelates quadrilineatus
4a. Lower opercular spine greatly developed, extending beyond margin of opercular lobe
(Fig. 2a); lobes of caudal fin with distinct oblique stripes; spinous dorsal fin with large
black blotch on middle rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Terapon) ® 5
4b. Lower opercular spine not extending beyond margin of opercular lobe (Fig. 2b); lobes of
caudal fin lacking distinct oblique stripes; spinous dorsal fin lacking large black blotch on
middle rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Rhyncopelates, Mesopristes) ® 7
posttemporal bone posttemporal bone

not exposed exposed, serrate
opercular
spine long opercular
spine short
margin of
opercular lobe
a) Amniataba, Pelates b) Terapon, Mesopristes
quadrilineatus Rhyncopelates a) Terapon b) Mesopristes
Fig. 1 Fig. 2
5a. Lateral-line scales 46 to 56; scale rows above lateral line 6 to 8 . . . . . . . . . . . Terapon theraps
5b. Lateral-line scales 70 or more; scale rows above lateral line 10 to 17 . . . . . . . . . . . . . . . ® 6
6a. Two to 4 straight, dark, longitudinal stripes along side of body; 18 to 24 gill rakers
(including rudiments) on lower limb of first gill arch . . . . . . . . . . . . . . . . . . . . Terapon puta
6b. Three downward curved, dark, longitudinal stripes along side of body; 12 to 15 gill rakers
(including rudiments) on lower limb of first gill arch . . . . . . . . . . . . . . . . . . . Terapon jarbua
7a. Lateral-line scales 60 to 75 . . . . . . . . . . . . . . . . . . . . . . . . Rhyncopelates oxyrhynchus

7b. Lateral-line scales 48 to 58 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Mesopristes) ® 8
8a. Cheek scale rows 4 to 6; body with dark, vertical bars dorsally and horizontal stripes
ventrally in all but largest adults . . . . . . . . . . . . . . . . . . . . . . . . Mesopristes cancellatus
8b. Cheek scale rows 8 or 9; body without dark vertical bars dorsally at any age; juveniles
with horizontal stripes on lateral and dorsolateral surfaces of body, stripes absent in
adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Mesopristes argenteus

Note: entirely fresh-water species from Australia and New Guinea are not included here.
0
0 Amniataba caudavittatus (Richardson, 1844)
0 Mesopristes argenteus (Cuvier, 1829)
0 Mesopristes cancellatus (Cuvier, 1829)
0 Pelates quadrilineatus (Bloch, 1790)
0 Pelates sexlineatus (Quoy and Gaimard, 1824)
0 Rhyncopelates oxyrhynchus (Temminck and Schlegel, 1842)
0 Terapon jarbua (Forsskål, 1775)
0 Terapon puta (Cuvier, 1829)
0 Terapon theraps (Cuvier, 1829)
Reference
Vari. R.P. 1978. The Terapon perches (Percoidei, Teraponidae), a cladistic analysis and taxonomic revision. Bull. Am.
Mus. Natl. Hist., 159(5):175-340.
3308 Bony Fishes
Amniataba caudavittatus (Richardson, 1844)

Frequent synonyms / misidentifications: Terapon caudavittatus (Richardson, 1844); Amphitherapon
caudavittatus (Richardson, 1844) / None.
FAO names: En - Flagtailed terapon.
(after Vari, 1978)
Diagnostic characters: A moderate-sized species; body moderately deep and compressed laterally.
Upper jaw slightly longer than lower jaw, gape oblique; posterior margin of upper jaw extending approxi-
mately to a vertical line through anterior margin of eye in juveniles, only to a vertical line in the region
between posterior nostril and eye in adults. Teeth conical, outer row enlarged and followed by a villiform
band in each jaw, vomer and palatines (on roof of mouth) toothless. Preopercle slightly serrate, serrations
largest along angle; lower opercular spine stronger and longer, not extending beyond margin of
opercular lobe. Posttemporal bone not exposed, covered with skin and scales. First gill arch with 6
to 8 gill rakers on upper limb, 12 or 13 on lower limb. Dorsal fin with XII or XIII spines and 8 to 10 soft
rays, the spinous part of dorsal fin arched, the fifth spine longest, and ultimate spine shortest; anal fin
with III spines and 8 or 9 soft rays, second anal-fin spine longer than third, shorter than longest anal-fin
rays. Pored scales in lateral line 46 to 54; 7 to 9 rows of scales above lateral line and 17 to 19 below it.
Colour: dorsal portions of body grey, ventral part of body lightly pigmented; dorsal half of body with
randomly dispersed spots somewhat smaller than pupil; some individuals with 5 or 6 incomplete vertical
bars extending from dorsal surface of body down to level of pectoral fins; spinous dorsal fin with irregular
spotting and faint duskiness distally, but without a distinct patch of dark pigmentation anteriorly; soft
dorsal fin somewhat dusky basally and distally; spinous portion of anal fin slightly dusky; soft portion of
anal fin slightly dusky anteriorly; caudal fin spotted basally, with distinct, black blotch extending
obliquely across each lobe.
Habitat, biology, and fisheries: Occurs in a variety of habitats ranging from fresh to hypersaline waters.
Within its range it occurs in a variety of near shore and fresh-water habitats. Caught with handlines, seines,
and other inshore fishing gear.
Distribution: Western and
northern Australia to southern
Papua New Guinea.
Mesopristes argenteus (Cuvier, 1829)

FAO names: En - Silver terapon.
(after Vari, 1978)
Diagnostic characters: A moderate-sized species; body moderately deep, distinctly compressed laterally,
ventral profile of head and body nearly straight to anus, more so in adults. Upper jaw longer than lower,
gape nearly horizontal; posterior margin of upper jaw extending to vertical line through anterior margin of
eye in juveniles, only to vertical through posterior nostril in adults. Teeth villiform, in bands in each jaw,
outer row of teeth distinctly enlarged; vomer and palatines (on roof of mouth) toothless. Preopercle serrate,
serrations larger along angle and vertical border; lower opercular spine stronger and longer, not
extending beyond margin of opercular lobe. Posttemporal bone expanded and exposed posteriorly,
with serrate posterior margin. First gill arch with 7 to 9 gill rakers on upper limb, 15 to 17 on lower limb.
Dorsal fin with XII spines and 9 to 11 soft rays, the spinous part of fin arched, the fourth or fifth spines
longest, and penultimate and ultimate spines subequal; anal fin with III spines and 8 or 9 soft rays,
second and third anal-fin spines very long and strong, second longest and as long as or slightly longer
than longest anal-fin rays. Pored scales in lateral line 52 to 58; 6 to 8 rows of scales above lateral line
and 18 to 21 below it; cheek-scale rows 8 or 9. Colour: dorsal portions of body grey, ventral part of body
silvery to silvery white; juveniles with 5 longitudinal body stripes; coloration in adults uniform; spinous
and soft portions of dorsal fin slightly dusky with blackish pigmentation sometimes present distally, but
lacking large black blotch on middle rays; caudal fin dusky, without distinct, dark, oblique stripes.
Habitat, biology, and fisheries: Found in marine, brackish, and fresh waters. Caught with handlines,
seines, and other inshore fishing gear.
Distribution: From Sumatra
through rest of Indonesia, to
northern Australia, New
Guinea, Vanuatu, and the
Philippines.
3310 Bony Fishes
Mesopristes cancellatus (Cuvier, 1829)

Frequent synonyms / misidentifications: Terapon cancellatus (Cuvier, 1829) / None.
FAO names: En -Tapiroid terapon.
(after Vari, 1978)
Diagnostic characters: A moderate-sized species; body moderately deep, distinctly compressed later-
ally, ventral profile of head and body nearly straight to anus, more so in adults. Upper jaw longer than lower
jaw, increasingly so in larger specimens; upper jaw fleshy and forming a prominent fleshy pad with age,
gape horizontal; posterior margin of upper jaw extending approximately to vertical line through anterior
margin, typically somewhat more posteriorly in adults. Teeth conical, outer row enlarged and followed by
bands of villiform teeth in each jaw; vomer and palatines (on roof of mouth) toothless. Preopercle serrate,
serrations very large in juveniles, less developed in adults in which serrations are larger along vertical
border; lower opercular spine stronger and longer, not extending beyond margin of opercular lobe.
Posttemporal bone expanded and exposed posteriorly, with serrate posterior margin. First gill arch
with 9 to 11 gill rakers on upper limb, 15 to 18 on lower limb. Dorsal fin with XI or XII spines and 10 or 11
soft rays, the spinous part of dorsal fin markedly arched, the fifth or sixth spines longest, and penultimate
and ultimate spines subequal, but penultimate spine distinctly stronger; anal fin with III spines and
8 or 9 soft rays, second and third anal-fin spines strong, second spine nearly as long as longest anal-fin
rays. Pored scales in lateral line 48 to 58; 6 to 8 rows of scales above lateral line and 15 to 21 below it;
cheek-scale rows 4 to 6. Colour: dorsal portions of body grey, ventral part of body silvery to silvery white;
body with 3 dark longitudinal stripes and with 5 dark oblique cross bars dorsal to uppermost stripe;
stripes and bars much less obvious in larger specimens and fading into general background coloration in
largest individuals; spinous dorsal fin clear, lacking large black blotch on middle rays; soft dorsal fin
with some dusky basal markings; spinous portion of anal fin slightly pigmented between second and third
spines and between third spine and first anal-fin ray; soft portion of anal fin with basal blotch on first 3 rays;
caudal fin with some of the body stripes continued onto its base, but without dark oblique stripes across
lobes.
Habitat, biology, and fisheries: Found in marine, brackish, and fresh waters. Caught with handlines,
seines, and other inshore fishing gear.
Distribution: From Sumatra
eastward through rest of Indo-
n e s ia , N e w G u i ne a, an d
Vanuatu to the Solomon Is-
lands; in the north to the Phil-
ippines and Taiwan Province
of China.
Pelates quadrilineatus (Bloch, 1790)

FAO names: En - Fourlined terapon; Fr - Violon crépuscule; Sp - Baraonga aurora.
Diagnostic characters: A moderately small species; body oblong, compressed laterally. Jaws equal, gape
oblique; posterior margin of upper jaw extending to vertical line through posterior nostril; teeth brown-
tipped, 2 rows in lower jaw and 3 rows or a villiform band with outer row of upper jaw enlarged in upper
jaw; vomer and palatines (on roof of mouth) toothless. Preopercle serrate, serrations larger along vertical
border, particularly in adults; lower opercular spine stronger and longer, but not extending beyond
margin of opercular lobe. Posttemporal bone not expanded or exposed posteriorly, covered with
skin and scales. First gill arch with 16 to 18 gill rakers on upper limb, 22 to 27 on lower limb. Dorsal
fin with XII or XIII spines and 9 to 11 soft rays, spinous part of fin arched, fifth to seventh spines longest,
and last 2 spines of approximately same length; anal fin with III spines and 9 or 10 soft rays, second
anal-fin spine subequal to third spine and shorter than longest anal-fin rays. Pored scales in lateral
line 66 to 75; 9 to 11 rows of scales above lateral line and 19 to 23 below it. Colour: dorsal portions of
body silvery grey, ventral part of body silvery white; 4 to 6 narrow, dark brown or black horizontal stripes
on body, the middle stripe extending onto caudal-fin base; juveniles in addition have 6 or 7 greyish vertical
bars; spinous portion of dorsal fin with a black blotch dorsally on membranes between third to
seventh dorsal-fin spines; a blotch of variable intensity on side of body posterior to nape; lobes of caudal
fin without prominent transverse black stripes; mouth and gill cavity red in life.
Habitat, biology, and fisheries: Found in inshore waters, sometimes in brackish waters. Feeds on small
fishes and invertebrates. Caught with all types of inshore fishing gear, including gill nets, traps, handlines,
and bottom trawls. Marketed fresh and dried-salted.
the Indo-Pacific from East Af-
rica, including the Red Sea
and Persian Gulf, eastwards
to the Philippines, Japan, and
China to New South Wales
and the Solomon Islands.
3312 Bony Fishes
Pelates sexlineatus (Quoy and Gaimard, 1824)

Frequent synonyms / misidentifications: Helotes sexlineatus (Quoy and Gaimard, 1824) / None.
FAO names: En - Sixlined terapon.
(after Vari, 1978)
Diagnostic characters: A small species; body oblong, slightly compressed laterally. Jaws equal, gape
short and oblique; posterior margin of upper jaw extending to vertical line midway between vertical through
posterior nostril and anterior margin of eye; teeth tricuspidate, cusps of nearly equal size, teeth in bands
in each jaw, outer row of teeth enlarged; vomer and palatines (on roof of mouth) toothless. Preopercle
serrate, serrations larger along vertical border; lower opercular spine stronger and longer, extending
nearly to, but not surpassing, margin of opercular lobe in adults. Posttemporal bone not expanded
or exposed posteriorly, covered with skin and scales. First gill arch with 6 or 7 rakers on upper limb,
14 or 15 on lower limb. Dorsal fin with XI or XII spines and 9 to 11 soft rays, spinous part of fin arched,
fifth or sixth spines longest, and penultimate spine shorter than ultimate spine; anal fin with III spines
and 9 to 11 soft rays, second anal-fin spine subequal to third spine and much shorter than longest
anal-fin rays. Pored scales in lateral line 76 to 87; 11 to 13 rows of scales above lateral line and 22 or
24 below it. Colour: dorsal portion of body greyish or bluish, ventral part of body silvery to silvery white; 5
to 8 narrow, dark brown or black horizontal stripes on body, the midlateral stripe extending onto
caudal-fin base; spinous part of dorsal fin with black margin dorsally on membranes; a blotch of variable
intensity on side of body posterior to nape; caudal fin with dark border and several bands of faint spots
basally, but without prominent transverse black stripes.
Size: Maximum total length about 32 cm, commonly between 18 and 24 cm
Habitat, biology, and fisheries: Marine inshore waters. Feeds on invertebrates and small fishes. Caught
with handlines, seines, and other inshore fishing gear. Marketed mostly fresh.
Distribution: Singapore to
Australia in the east and
south; northwards to China
and Okinawa (Japan).
Rhyncopelates oxyrhynchus (Temminck and Schlegel, 1842)

Frequent synonyms / misidentifications: Pelates oxyrhynchus (Temminck and Schlegel, 1842); Terapon
oxyrhynchus (Temminck and Schlegel, 1842) / None.
FAO names: En - Blotched terapon.
(after Vari, 1978)
Diagnostic characters: A moderate-sized species; body oblong, moderately deep, compressed laterally,
more so in juveniles. Jaws equal, gape slightly oblique; posterior margin of upper jaw extending to vertical
line through posterior nostril. Teeth villiform, in bands in each jaw, outer row of teeth much enlarged; vomer
and palatines (on roof of mouth) toothless. Preopercle serrate, serrations larger along vertical border; lower
opercular spine stronger and longer, not extending beyond margin of opercular lobe. Posttemporal
bone expanded and exposed posteriorly, with serrate posterior margin. First gill arch with 7 or 8 gill
rakers on upper limb, 14 to 16 on lower limb. Dorsal fin with XII spines and 9 to 11 soft rays, spinous part
of fin arched, third to sixth spines longest, and penultimate and ultimate spines subequal; anal fin with
III spines and 7 to 9 soft rays, second anal-fin spine longest and shorter than longest anal-fin rays.
Pored scales in lateral line 60 to 75; 10 or 11 rows of scales above lateral line and 20 to 24 below it.
Colour: dorsal portions of body grey, ventral part of body silvery to silvery white; 4 somewhat blotchy dark
brown or black horizontal stripes on body; some individuals with irregular stripes between 2 dorsal and 2
ventral primary stripes; spinous part of dorsal fin dusky basally and distally; soft portion of dorsal fin with
a basal band anteriorly and pigmentation across posterior rays. Spinous portion of anal fin unpigmented;
caudal fin clear in juveniles, with several narrow, irregular, parallel stripes on each lobe in adults.
Size: Maximum total length about 33 cm, commonly between 18 and 23 cm.
Habitat, biology, and fisheries: Juveniles entering into fresh waters. Adults found in inshore marine and
brackish waters. Feeds on small fishes and invertebrates. Caught with traps, handlines, and other inshore
fishing gear.
Distribution: Philippines,
southern China, Taiwan Prov-
ince of China, and southern
Japan.
3314 Bony Fishes
Terapon jarbua (Forsskål, 1775)

Frequent synonyms / misidentifications: Therapon servus Bloch, 1790 / None.
FAO names: En - Jarbus terapon; Fr - Violon jarbua; Sp - Baraonga jarbua.
Diagnostic characters: A moderate-sized species. Body oblong moderately compressed laterally. Jaws
equal, gape oblique; rear end of upper jaw reaching to vertical through anterior margin of eye in juveniles,
reaching to vertical through centre of orbit in adults; teeth conical, slightly recurved, in villiform bands, the
outer row much enlarged; vomer and palatines (on roof of mouth) with teeth in juveniles, often toothless
in adults. Preopercle strongly serrate, particularly at angle; lower opercular spine very long and strong,
extending distinctly beyond margin of opercular lobe. Posttemporal bone expanded, exposed and
serrate posteriorly; cleithrum serrate posteriorly. First gill arch with 6 to 8 rakers on upper limb, 12 to
15 on lower limb. Dorsal fin with XI or XII spines and 9 to 11 rays, spinous part of fin strongly arched and
deeply notched, fourth to sixth spines longest, and penultimate spine about 1/2 length of ultimate spine;
margin of soft part of dorsal fin straight or emarginate; anal fin with III spines and 7 to 10 soft rays, second
anal-fin spine subequal to third spine and shorter than longest anal-fin rays, margin of soft part of
anal fin concave; caudal fin emarginate. Pored scales in lateral line 75 to 100; 13 to 17 rows of scales
above lateral line and 19 to 34 rows below it. Colour: body silvery greyish or dorsally and silvery white
ventrally; 3 or 4 dark brown or black downwardly curved longitudinal stripes on body; spinous
portion of dorsal fin with a blackish blotch dorsally on membranes between third and sixth spines;
soft portion of dorsal fin with membranes of first 3 rays tipped with black and membranes between fifth and
seventh rays entirely black; caudal fin with medial rays pigmented; both caudal-fin lobes with dark tips
and a transverse band.
Habitat, biology, and fisheries: Inhabits coastal, brackish, and fresh waters, but apparently reproduces
solely in marine waters. Feeds on smaller fishes, invertebrates, and also selectively on scales of various
groups of fishes. Caught on all types of inshore fishing gear including gill nets, traps, handlines, and bottom
trawls. Marketed fresh and dried-salted.
rica, Red Sea, and Persian
Gulf to Fiji and Samoa in the
east, and New South Wales
(Australia) in the south; north-
wards to Japan.
Terapon puta (Cuvier, 1829)

Frequent synonyms / misidentifications: Authistes puta Whitley, 1943 / None.
FAO names: En - Smallscaled terapon; Fr - Violon grogneur; Sp - Baraonga ronco.
Diagnostic characters: A small-sized species. Body somewhat elongate and laterally compressed. Jaws
equal, gape horizontal; rear of upper jaw reaching to vertical through posterior nostril; teeth conical, strong,
in villiform bands, outer row enlarged; vomer and palatines (on roof of mouth) toothless. Preopercle strongly
serrate, with large spines at angle; lower opercular spine very long and strong, extending distinctly
beyond margin of opercular lobe. Posttemporal bone expanded, exposed and serrate posteriorly;
cleithrum serrate posteriorly. First gill arch with 7 to 9 gill rakers on upper limb, 18 to 24 on lower limb.
Dorsal fin with XI or XII spines and 9 to 11 soft rays, spinous part of fin strongly arched and deeply notched,
fifth or sixth spines longest and penultimate spine about 1/2 length of ultimate spine; margin of soft
part of dorsal fin straight or emarginate; anal fin with III spines and 8 or 9 soft rays, second anal-fin spine
shorter than third spine and longest anal-fin rays, margin of soft part of anal fin emarginate; caudal fin
emarginate. Pored scales in lateral line 70 to 85; 10 to 13 rows of scales above lateral line and 22 to 34
rows below it. Colour: body light grey or brown dorsally, tan or silvery white ventrally; 4 straight, narrow,
dark brown longitudinal stripes on body; juveniles also have 6 or 7 light grey vertical bars; spinous
portion of dorsal fin with a blackish blotch dorsally on membranes between third or fourth and
seventh or eighth spines; soft portion of dorsal fin with a black blotch along top of anterior rays; caudal
fin with medial rays pigmented; both lobes of caudal fin with dark tips and a transverse band.
Habitat, biology, and fisheries: Found in inshore waters, sometimes entering brackish and fresh waters.
Feeds on smaller fishes and invertebrates. Caught with all types of inshore fishing gear including gill nets,
traps, handlines, and bottom trawls. Marketed fresh and dried-salted.
rica north of Zanzibar, Red
Sea and Persian Gulf to New
Guinea and Vanuatu, and
north to the Philippines.
3316 Bony Fishes
Terapon theraps (Cuvier, 1829)

Frequent synonyms / misidentifications: Eutherapon theraps Whitley, 1943 / None.
FAO names: En - Largescaled terapon; Fr - Violon tigre; Sp - Baraonga tigre.
Diagnostic characters: A moderate-sized species; body oblong to moderately compressed laterally. Jaws
equal, gape slightly oblique; rear end of upper jaw reaching to vertical through anterior margin of orbit in
juveniles, falling short of that line in adults and only reaching a vertical through margin of posterior nostril;
teeth conical, in villiform bands, outer row much enlarged; vomer and palatines (on roof of mouth) toothless.
Preopercle serrate, serrations particularly developed at angle, more so in adults; lower opercular spine
very long and strong, extending distinctly beyond margin of opercular lobe; posttemporal bone
expanded, exposed and serrate posteriorly; cleithrum serrate posteriorly. First gill arch with 6 to 8 gill
rakers on upper limb, 14 to 17 on lower limb. Dorsal fin with XI or XII spines and 9 to 11 soft rays; the
spinous part of the fin strongly arched and deeply notched; the third to sixth dorsal-fin spines longest and
the penultimate spine about 1/2 length of ultimate spine; margin of soft part of dorsal fin emarginate;
anal fin with III spines and 7 to 9 soft rays, third anal-fin spine longest but shorter than longest anal-fin
ray; margin of soft part of anal fin angular, with a straight to slightly emarginate border; caudal fin shallowly
forked with rounded lobes. Pored scales in lateral line 46 to 56; 6 to 8 rows of scales above lateral line
and 14 to 16 rows below it. Colour: body dusky dorsally, silvery white ventrally; 4 dark brown horizontal
stripes on body; juveniles also have 6 or 7 light grey vertical bars; spinous part of dorsal fin with a
black blotch dorsally on fin membranes between third and seventh spine; soft portion of dorsal fin
with a dark band along upper portions of anterior rays and a horizontal band on posterior rays; soft portion
of anal fin with distal portions black; caudal fin with medial rays pigmented; upper lobe of caudal fin with
dark tip; both lobes of caudal fin with a dark, transverse band.
Habitat, biology, and fisheries: Found in inshore waters, sometimes entering brackish waters. Feeds on
smaller fishes and invertebrates. Caught with all types of inshore fishing gear, including traps, handlines,
and bottom trawls. Marketed
mostly fresh, sometimes
dried salted.
rica, Red Sea, and Persian
Gulf to the Solomon Islands.
Perciformes: Percoidei: Kuhliidae 3317
KUHLIIDAE
Flagtails
K.E. Carpenter
D iagnostic characters (for species found in the area): Oblong, compressed, moderate-sized (to about
50 cm) perch-like fishes. Head length less than body depth; preorbital and preopercle margins serrate;
2 flat spines on opercle. Eye moderately large, nearly bisected by a horizontal from tip of snout to middle
of caudal-fin base. Mouth small, slightly protrusible; maxilla not covered by preorbital; bands of fine teeth
in jaws, vomer, and palatines. A single dorsal fin, deeply notched between spinous and soft-rayed
portions, with X spines and 9 to 12 soft rays. Anal fin with III spines and 9 to 13 soft rays. Caudal
fin emarginate to moderately forked. Scales ctenoid, moderately large; lateral line complete, with 40 to 56
scales; a well-developed scaly sheath at base of dorsal and anal fins. Colour: mostly silvery, some
with black spots on body; dorsal and anal fins sometimes with dark spots or blotches; caudal fins with or
without species-specific characteristic dark blotches and bands.
single, deeply notched dorsal fin

2 spines on opercle with X spines and 9-12 soft rays

and 9-13 soft rays
Habitat, biology, and fisheries: Shallow water coastal fishes, on coral reefs, in rocky surge areas, bays,
harbors, estuaries, and some entering fresh water; juveniles often very close to shore and in tide pools.
Feed mostly on planktonic crustaceans at night. Excellent food fishes of minor importance to fisheries
because of their generally small size.
Remarks: The species of Kuhlia are in need of revision and the limits and relationships of the family
Kuhliidae needs to be better determined. The family is presently recognized as restricted to a single genus,
Kuhlia. The fresh-water genera Edelia, Nannatherina, and Nannoperca were formerly included in the
Kuhliidae but more recently determined to belong to the Percichthyidae. The West African genus Parakuh-
lia is placed in the Kuhliidae by some authors but otherwise recognized as belonging to the Haemulidae.

Terapontidae: eye mostly above horizontal from tip of snout to middle of caudal-fin base (nearly bisected
in Kuhliidae); dorsal fin with XI to XIV spines (X spines in Kuhliidae); vomer and palatines usually without
teeth (with fine teeth in Kuhliidae).
Gerreidae: mouth highly protrusible and pointing downward when extended (slightly protrusible in Kuhlii-
dae); dorsal fin not usually deeply notched (deeply notched in Kuhliidae); caudal fin deeply forked, without
dark bands or markings (emarginate to forked often with dark bands in Kuhliidae).
Terapontidae Gerreidae
3318 Bony Fishes
Haemulidae (superficially similar to some Pomadasys):

eye mostly above horizontal from tip of snout to middle of
caudal-fin base (nearly bisected in Kuhliidae); anal fin with
III spines, the second enlarged, and 7 to 9 soft rays
(second dorsal-fin spine not enlarged and 9 to 13 soft rays
in Kuhliidae); caudal fin truncate to slightly emarginate
(strongly emarginate to forked in Kuhliidae).
Haemulidae
Key to the species of Kuhliidae occurring in the area

1a. Gill rakers on lower limb of first gill arch 15 to 19 (rarely 20); small black spots typically
scattered on caudal peduncle and sometimes on sides . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Gill rakers on lower limb of first gill arch 21 to 26; typically no small black sides on caudal
peduncle or body . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Caudal fin weakly emarginate, tips of lobes rounded; anal fin with 9 or 10 soft rays; 5 or
6 gill rakers on upper limb of first gill arch . . . . . . . . . . . . . . . . . . . . . . . Kuhlia rupestris
2b. Caudal fin deeply emarginate to forked, tips of lobes pointed; anal fin with 11 to 13 soft
rays; 7 to 9 gill rakers on upper limb of first gill arch . . . . . . . . . . . . . . . . . Kuhlia marginata
3a. Caudal fin with 5 broad black stripes, 1 central horizontal stripe along shortest rays and
2 oblique stripes on each lobe; base of caudal fin without a dark band . . . . . . . . . Kuhlia mugil
3b. Caudal fin with a broad black marginal band or with tips of lobes with a black blotch; base
of caudal fin with a dark band . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Caudal fin with a broad black marginal band . . . . . . . . . . . . . . . . . . . . . . . Kuhlia munda
4b. Tips of caudal-fin lobes with a black blotch . . . . . . . . . . . . . . . . . . . . . . Kuhlia boninensis

0
0
Kuhlia boninensis (Fowler, 1907)
0
Kuhlia marginata (Cuvier, 1829)
0
Kuhlia mugil (Forster in Bloch and Schneider, 1801)
0
Kuhlia munda (De Vis, 1884)
Kuhlia rupestris (Lacepède, 1802)
References
Masuda, U., C. Araga, and T. Yoshino. 1975. Coastal fishes of southern Japan . Tokyo, Tokai University Press, 379 p.
Munro, I.S.R. 1967. The fishes of New Guinea. Port Moresby, New Guinea, Department of Agriculture, Stock and
Fisheries, 650 p.
Weber, M. and L.F. de Beaufort (eds). 1929. The fishes of the Indo-Australian Archipelago. V. Anacanthini,
Allotriognathi, Heterosomata, Berycomorphi, Percomorphi (Families: Kuhliidae, Apogonidae, Plesiopidae,
Pseudoplesiopidae, Priacanthidae, Centropomidae). Leiden, E. J. Brill, 458 p.
Perciformes: Percoidei: Kuhliidae 3319
Kuhlia boninensis (Fowler, 1907)

En - Bonin Island flagtail.
Maximum total length 25 cm. In estuaries and fresh water. Found in the Ryukyu Islands, Bonin
Islands, Ogasawara Islands, Micronesia, and Tahiti. Specific reports of this species from Micronesia
need to be confirmed and are therefore not included on the map.
Kuhlia marginata (Cuvier, 1829)

En - Spotted flagtail.
Maximum total length about 20 cm. Mostly in fresh water and estuaries, but also reported from rocky
and reef coastal areas. Mostly western Pacific but also reported from Sri Lanka.
Kuhlia mugil (Forster in Bloch and Schneider, 1801)

En - Barred flagtail; Fr - Crocro drapeau; Sp - Dara bandera.
Maximum total length 25 cm. Typically on or near coralline or rocky reefs but also enters estuaries
and fresh water; juveniles use tidepools and nearshore surge areas as nurseries. Feeds mostly on
zooplankton and small fish at night, shoaling in rocky or coral reef areas during daytime. Used as
food, bait, and as an aquarium fish; caught by seines, gill nets, and hook-and-line. Widespread in
the Indo-Pacific from South Africa to Central America. Often incorrectly identified as Kuhlia
taeniura, which is a junior synonym of K. mugil.
?
3320 Bony Fishes
Kuhlia munda (De Vis, 1884)

En - Silver flagtail.
Maximum total length 24 cm. Coastal bays, estuaries, and entering fresh water. Western Pacific
from Queensland, New Caledonia, Vanuatu, Papua New Guinea, and Fiji.
(after Munro, 1967)
Kuhlia rupestris (Lacepède, 1802)

En - Rock flagtail; Fr - Crocro sauvage; Sp - Dara campestre.
Maximum total length 56 cm. Mostly in fresh water and estuaries. A good food fish, caught primarily
by small scale beach seines, gill nets, throw nets, and by anglers with hook-and-line. Widespread
in the Indo-Pacific from the Red Sea to South Africa, from Ryukyu Islands to Queensland and
eastward to the Society Islands.
adult
juvenile
click for next page

Perciformes: Percoidei: Cirrhitidae 3321
CIRRHITIDAE
Hawkfishes
by J.E. Randall
D iagnostic characters: Oblong fishes (size to about 30 cm), body depth 2 to 4.6 times in standard
length. A fringe of cirri on posterior edge of anterior nostril. Two indistinct spines on opercle. A row of
canine teeth in jaws, the longest usually anteriorly in upper jaw and half-way back on lower jaw; a band of
villiform teeth inside the canines, broader anteriorly (in lower jaw only anteriorly). One or more cirri
projecting from tips of interspinous membranes of dorsal fin. Dorsal fin continuous, with X spines
and 11 to 17 soft rays, notched between spinous and soft portions; anal fin with III spines and 5 to 7
(usually 6) soft rays; pectoral fins with 14 rays, the lower 5 to 7 rays unbranched and usually enlarged,
with the membranes deeply incised; pelvic fins with I spine and 5 soft rays. Principal caudal-fin rays 15.
Branchiostegal rays 6. Scales cycloid. Swimbladder absent. Vertebrae 26. Colour: variable with species.
cirri
lower pectoral-fin rays

thickened and unbranched
Remarks: The hawkfish family consists of 10 genera and 38 species, 33 of which occur in the Indo-Pacific
region; 19 species are found in the Western Central Pacific.
Habitat, biology, and fisheries: Cirrhitids are bottom-dwelling fishes of coral reefs or rocky substrata; the
majority occur in shallow water. They use their thickened lower pectoral-fin rays to wedge themselves in
position in areas subject to surge. All species are carnivorous, feeding mainly on benthic crustaceans.
Cyprinocirrhitus polyactis, however, is primarily a zooplankton-feeder (though still often seen at rest on
the bottom). Most hawkfishes are small; separate species accounts are given below for the 3 largest
hawkfishes that are occasionally used as food. Some of the smaller species, notably the bright red
Neocirrhites armatus and the long-snouted Oxycirrhites typus, are of value as aquarium fishes.
Similar families Cirrhitidae occurring in the area

Serranidae: lower 5 to 7 pectoral-fin rays not simple
and thickened (except for a few Plectranthias); no
fringe of cirri on posterior nostril; scales usually
ctenoid; usually III spines on opercle; vertebrae
typically 24; swimbladder present.
Serranidae (Plectranthias)
3322 Bony Fishes
Key to the genera of Cirrhitidae occurring in the area

1a. Snout elongate, its length about 2 times in head length (Fig. 1); body slender, its depth
4.4 to 4.6 times in standard length; canine teeth in jaws only slightly longer than inner
villiform teeth and nearly uniform in size . . . . . . . . . . . . . . . . . . . . . . . . . . Oxycirrhites
1b. Snout not elongate, its length 2.8 to 5 times in head length; body not slender, its depth
2 to 3.4 times in standard length; canine teeth in jaws markedly longer than inner villiform
teeth, those at front of upper jaw and side of lower jaw enlarged . . . . . . . . . . . . . . . . . . ® 2
2a. Caudal fin lunate, the lobes produced (Fig. 2); soft dorsal-fin rays 16 or 17 . . . . . Cyprinocirrhites
2b. Caudal fin rounded, truncate, or slightly emarginate; soft dorsal-fin rays 11 to 15 . . . . . . . . . ® 3
Fig. 1 Oxycirrhites Fig. 2 Cyprinocirrhites
3a. No large scales on cheek (small scales in more than 12 irregular rows) (Fig. 3) . . . . . . . . . . ® 4
3b. Four to 6 rows of large scales on cheek (small scales usually present as well) (Fig. 4) . . . . . . ® 5
Fig. 3 Fig. 4
4a. Body not deep and not very compressed, its depth 2.6 to 3.4 times in standard length;
body width 1.5 to 1.9 times in body depth; soft dorsal-fin rays 11 (rarely 12); palatine
teeth present; upper margin of preopercle finely serrate or smooth; lower 7 pectoral-fin
rays unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitus
4b. Body deep and compressed, its depth 2 to 2.4 times in standard length; body width 2.9
to 3.1 times in body depth; soft dorsal-fin rays 13; palatine teeth absent; upper margin of
preopercle coarsely serrate; lower 6 pectoral-fin rays unbranched . . . . . . . . . . . . Neocirrhites
5a. Five rows of large scales between lateral line and spinous portion of dorsal fin; a single
cirrus from membrane near tip of each dorsal-fin spine (Fig. 5a) membranes of spinous
portion of dorsal fin not deeply incised, those between longest dorsal-fi n spines extend-
ing 4/5 or more of distance from base to tip of spines; palatine teeth absent . . . . . . Paracirrhites
5b. Three or 4 rows of large scales between lateral line and base of spinous portion of dorsal
fin; a tuft of cirri from membrane near tip of each dorsal-fin spine (Fig. 5b); membranes
of spinous portion of dorsal fin deeply incised, those between longest spines less than
2/3 distance from base to tip of spines; palatine teeth present or absent . . . . . . . . . . . . . . ® 6
a) Paracirrhites b) Cirrhitops
Fig. 5 spinous part of dorsal fin
6a. Soft dorsal-fin rays 14 (rarely 15); first 2 pectoral-fin rays unbranched; preorbital with a
free posterior margin; interorbital not scaly; snout not pointed, the profile from interorbital
to upper lip convex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitops
6b. Soft dorsal-fin rays 11 or 12 (rarely 13); first pectoral-fin ray unbranched, the second
usually branched; preorbital with or without a free posterior margin; interorbital scaly or
not scaly; snout pointed or not pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Palatine teeth absent; longest dorsal-fin spine 3.5 to 4 times in body depth; snout not
pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Isocirrhitus
7b. Palatine teeth present; longest dorsal-fin spine 1.7 to 3.2 times in body depth; snout
pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .®8
8a. Preopercular margin finely serrate (Fig. 6); preorbital without a free hind margin; interor-
bital scaly; first soft dorsal-fin ray not produced to a filament; lower 5 (rarely 6) pectoral-
fin rays unbranched . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amblycirrhitus
8b. Preopercular margin coarsely serrate (Fig. 7); preorbital with hind margin free for about
1/4 to 1/2 distance from its lower edge to eye; interorbital not scaly; first soft dorsal-fin
ray usually produced into a filament; lower 6 or 7 pectoral-fin rays unbranched . . . . Cirrhitichthys
coarsely serrate
Fig. 6 Amblycirrhitus Fig. 7 Cirrhitichthys
Key to the species of Amblycirrhitus occurring in the area

1a. Lateral-line scales 48 to 50; tips of membranes of spinous portion of dorsal fin black
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Amblycirrhitus unimaculata
(Ryukyu Islands, Taiwan Province of China, and Samoa)
1b. Lateral-line scales 39 to 42; tips of membranes of spinous portion of dorsal fin not black . . . . . ® 2
2a. A large ocellated black spot on opercle; head not crossed by 3 dark bars . . Amblycirrhitus bimacula
(Indo-Pacific)
2b. No large black spot on opercle; head crossed by 3 dark bars, the first passing ventrally from
eye, the next 2 on operculum . . . . . . . . . . . . . . . . . . . . . . . . Amblycirrhitus oxyrhynchos
(Goram Islands, Indonesia)
Key to the species of Cirrhitichthys occurring in the area

1a. Body depth 2.4 to 2.7 times in standard length; bony interorbital space not narrow, its
least width in adults about 1.7 times in eye diameter; free edge of preorbital usually with
1 to a few small spines; large dark spots on body, often coalescing to form 6 irregular
bars; a pale-edged black spot larger than pupil on opercle at level of eye; 5 or 6 narrow
oblique to diagonal dark bands on lower half of head, 3 or 4 of which extend ventroante-
riorly from eye . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitichthys aprinus
(southern Japan to Great Barrier Reef)
1b. Body depth 2.7 to 3 times in standard length; bony interorbital space narrow, its least
width about 2 times in eye diameter; free hind edge of preorbital without spines; colour
not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
3324 Bony Fishes
2a. Rear edge of maxilla slightly anterior to a vertical at front edge of orbit; fourth dorsal-fin
spine longest; first soft dorsal-fin ray not decidedly longer than other rays; body with
numerous small red to reddish brown spots, many of which join to form about 7 bars; first
bar vertical, centred on origin of dorsal fin and containing dark brown spots in its upper
half, the other bars more oblique; 2 red to reddish brown bands extending ventroanteri-
orly from eye and a median band on snout . . . . . . . . . . . . . . . . . . . . . Cirrhitichthys falco
(western Pacific to Maldive Islands)
2b. Rear edge of maxilla reaching to or beyond a vertical at front edge of orbit; fifth dorsal-fin
spine longest; first soft dorsal-fin ray decidedly longer than more posterior rays (3.5 cm
standard length or larger); body with large close-set red to reddish brown blotches in 4
longitudinal rows and an additional row of smaller blotches along lateral line; head with
numerous small red to reddish brown spots . . . . . . . . . . . . . . . . . Cirrhitichthys oxycephalus
(Indo-Pacific and tropical eastern Pacific)
Key to the species of Cirrhitus occurring in the area

1a. Supraoccipital crest visible as a low ridge; no scales on interorbital space; no small white
spots on head, body, and fins (white blotches nearly as large as eye present on body) .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitus pinnulatus
(Indo-Pacific)
1b. Supraorbital crest not visible externally; a narrow median band of scales on interorbital
space; small white spots on head, body, and fins, those on body arranged in about 12
longitudinal rows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cirrhitus albopunctatus
(Niuafoo Island, near Tonga Islands)
Key to the species of Paracirrhites occurring in the area

1a. Second pectoral-fin ray branched, at least near tip; scales present on snout anterior to
nostrils; no colour mark extending diagonally upward from rear part of eye . . . . . . . . . . . . ® 2
1b. Second pectoral-fin ray unbranched; no scales on snout anterior to nostrils; a prominent
elongate solid or U-shaped dark mark extending diagonally upward from rear part of eye . . . . . . . ® 3
2a. Tenth dorsal-fin spine slightly longer than ninth; lateral-line scales 45 to 49; numerous
small dark spots on head and anterior ly on body . . . . . . . . . . . . . . . . . Paracirrhites forsteri
(Indo-Pacific)
2b. Tenth dorsal-fin spine notably longer than ninth; lateral-line scales 48 to 51; numerous
small dark spots on body but none on head . . . . . . . . . . . . . . . . . Paracirrhites hemistictus
(Oceania except Hawaii; Great Barrier Reef, Christmas Island, and Cocos-Keeling Islands)
3a. Three dark-edged pale (orange in life) transverse bands on interopercle; postocular mark not
black within U-shaped border (though may be darker than rest of head); 1 colour form with
a broad pale stripe, lacking a black border, posteriorly on upper side; lateral-line scales 45
to 50 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracirrhites arcatus
(Indo-Pacific)
3b. No transverse bands on interopercle; postocular mark black within its U-shaped border;
a broad pale stripe, if present on upper side, broadly bordered with black; lateral-line
scales 48 to 52 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Uniformly pale (bright yellow in life); black postocular mark narrower than pupil, originat-
ing at or below centre of rear edge of eye, its lower border not extending anterior to eye;
no dark-edged white spots anteriorly on snout; no markings on maxilla; no white spot at
anterior nostril . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracirrhites xanthus
(Tuamotu Archipelago, Society Islands, and Marquesas)
4b. Light to dark brown (not yellow in life); postocular mark broader than pupil, originating
above centre of eye, its lower border extending anterior to eye; a pair of dark-edged white
spots anteriorly on snout near upper lip; one more small marks on maxilla; anterior nostril
nearly enclosed in a white spot . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Light brown with a broad pale stripe, broadly bordered in black, posteriorly on upper side
of body; upper margin of preopercle smooth or with only a few small serrations . Paracirrhites nisus
(Tuamotu Archipelago and Pitcairn Group)
5b. Dark brown except caudal peduncle, caudal fin, and extreme ventral part of body posterior
to insertion of pelvic fins which are pale (orange in life); no pale stripe on body; upper margin
of preopercle finely serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Paracirrhites bicolor
(Caroline Atoll and Tuamotu Archipelago)
List of the species occurring in the area

Amblycirrhitus bimacula (Jenkins, 1903)
Amblycirrhitus oxyrhinchos (Bleeker, 1858)
Amblycirrhitus unimaculata (Kamohara, 1957)
Cirrhitichthys aprinus (Cuvier, 1829)
Cirrhitichthys falco Randall, 1963
Cirrhitichthys oxycephalus (Bleeker, 1855)
Cirrhitops hubbardi (Schultz, 1943)
0
Cirrhitus albopunctatus Schultz, 1950
Cirrhitus pinnulatus (Bloch and Schneider, 1801)
Cyprinocirrhites polyactis (Bleeker, 1875)
Isocirrhitus sexfasciatus (Schultz, 1960)
Neocirrhites armatus Castelnau, 1873
Oxycirrhites typus Bleeker, 1875
Paracirrhites arcatus (Cuvier, 1829)1/
0
Paracirrhites bicolor Randall, 1963
0
Paracirrhites forsteri (Bloch and Schneider, 1801)
Paracirrhites hemistictus (Günther, 1874)
Paracirrhites nisus Randall, 1963
Paracirrhites xanthus Randall, 1963
References
Randall, J.E. 1963. Review of the hawkfishes (family Cirrhitidae). Proc. U.S. Natl. Mus., 114:389-451.
Randall, J.E. 1997. The hawkfish Cirrhitichthys serratus Randall, a synonym of C. falco Randall. Micronesica,
30(1):199-203.
1/ Paracirrhites amblycephalus (Bleeker, 1857), known from a single specimen, 78 mm standard length, from the Sangi
Islands (Indonesia) is here treated as a probable synonym of P. arcatus.
3326 Bony Fishes
Cirrhites pinnulatus (Bloch and Schneider, 1801)

Frequent synoynms / misidentifications: Cirrhitus alternatus Gill, 1862; C. spilotoceps Schultz, 1950 /
None.
FAO names: En - Stocky hawkfish; Fr - Epervier de corail; Sp - Solosolo robusto.
Diagnostic characters: Body depth 2.7 to 3 times in standard length. Snout short and blunt. Supraorbital
ridge low. Mouth large, the maxilla nearly reaching a vertical at posterior edge of eye; upper margin of
preopercle finely serrate. A continuous dorsal fin of X spines and 11 soft rays, deeply notched
between spinous and soft portions; a tuft of cirri from membrane near tip of each dorsal-fin spine;
anal fin with III spines and 6 soft rays; caudal fin slightly rounded; pectoral fins not reaching a vertical at
tips of pelvic fins; pectoral-fin rays 14, the lower 7 unbranched and thickened. Lateral-line scales 39
to 44; 4 rows of large scales above lateral line in middle of body; scales on cheek much smaller than
scales on body. Colour: ground colour white but overlaid with squarish dark brown blotches and
numerous dark orangish to reddish brown spots, leaving as white 3 rows of 5 or 6 spots of about size of
eye on body; ventral thorax and abdomen white; head with reddish brown spots, sometimes conjoined to
form irregular bands; median fins with reddish brown spots.
Habitat, biology, and fisheries: This robust hawkfish, the largest of the family in the Indo-Pacific, lives
on rocky shores and reef fronts exposed to wave action. Nocturnal; feeds mainly on crabs. Generally caught
from shore, usually by hook-and-line. Marketed mostly fresh.
out the Indo-Pacific region
from the Red Sea south to Port
Alfred (South Africa) and east
to French Polynesia (including
Rapa) and the Pitcairn Group;
in the western Pacific from
southern Japan to the Great
Barrier Reef.
Remarks: T h i s s p e c i e s i s
slightly different in colour in
Hawaii and appears to have a
different count of gill rakers on
the upper limb of the first gill arch (5 or 6 in Hawaii compared to 6 or 7 elsewhere). The name Cirrhites
alternatus Gill is available in the event a subspecific name is desired for the Hawaiian population.
Paracirrhites forsteri (Bloch and Schneider, 1801)

Frequent synonyms / misidentifications: Paracirrhites typee Randall, 1963 / None.
FAO names: En - Blackside hawkfish; Fr - Epervier à bande noire; Sp - Solosolo de banda negra.
Diagnostic characters: Body depth 2.6 to 2.8 times in standard length. Snout short. Mouth large, the
maxilla reaching posterior to a vertical at centre of eye; upper margin of preopercle finely serrate. A
continuous dorsal fin of X spines and 11 soft rays, notched between spinous and soft portions; a
single cirrus from membrane near tip of each dorsal-fin spine; anal fin with III spines and 6 soft rays;
pectoral-fin rays 14, the lower 7 unbranched and thickened; caudal fin slightly rounded. Lateral-line
scales 45 to 49; 5 rows of large scales between lateral line and spinous portion of dorsal fin; 5 or 6 rows
of large scales on cheek, in addition to small scales. Colour: yellowish with a faint longitudinal dark
banding following scale rows; usually a broad blackish band (sometimes broken into conjoined spots) on
upper side from below middle of dorsal fin nearly to middle of caudal fin; region below black band usually
yellow (by virtue of reduced longitudinal banding on scale rows); head, nape, and thorax with small
brownish red spots; fins varying from yellowish to pink.
Habitat, biology, and fisheries: A common coral-reef species, often observed sitting on live coral. Feeds
mainly on small fishes. Occasionally caught throughout its range. Taken mainly by hook-and-line. Marketed
mostly fresh.
Distribution: In d o-Pacific,
but not known from the Per-
sian Gulf or Gulf of Oman.
Tropical and subtropical is-
lands of the Indian Ocean and
coast of Africa from the Red
Sea to Natal. In the Pacific
from southern Japan to New
South Wales, east to the is-
lands of Oceania, including
Hawaii and the Pitcairn
Group.
3328 Bony Fishes
Paracirrhites hemistictus (Günther, 1874)

Frequent synonyms / misidentifications: Paracirrhites polystictus (Günther, 1874) / None.
FAO names: En - Halfspotted hawkfish.
first phase
second phase
Diagnostic characters: Body elongate for the genus, its depth 2.8 to 3.2 in standard length. Mouth
large, maxilla reaching to or posterior to a vertical at centre of eye. Upper two-fifths of margin of preopercle
finely serrate, a slight concavity in margin below serrate portion. A continuous dorsal fin of X spines
and 11 soft rays, notched between spinous and soft portions; tenth dorsal-fin spine distinctly longer
than ninth; a single cirrus from membrane near tip of each dorsal-fin spine; anal fin with III spines
and 6 soft rays; pectoral-fin rays 14, the lower 7 unbranched and thickened; caudal fin slightly rounded,
the corners sharply angular; lateral-line scales 48 to 51; 5 rows of large scales between lateral line and
spinous portion of dorsal fin; snout almost entirely scaly; 5 oblique rows of large scales on cheek, in addition
to small scales. Colour: 2 colour phases, not related to sex, the first greenish on back, densely spotted
with black, pale yellow below, with series of dark yellow spots forming stripes following scale rows; an
irregular white stripe slightly below midside of body with a few black spots below and occasionally within
stripe; head greenish to pinkish grey; fins yellowish. Second (Paracirrhites polystictus) phase grey with
numerous close-set dark brown spots on body and a white or pale pink spot slightly smaller than eye on
upper side in middle of body; head reddish grey without markings.
Habitat, biology, and fisheries: A shallow-water, coral-reef species generally found at oceanic Islands.
Rests motionless on the substratum, often on live coral. Food habits not investigated; probably feeds
opportunistically on crustaceans and small fishes that venture near. Generally caught by hook-and-line.
Of limited commercial importance.
Distribution: Occurs on all the islands of
Oceania except Hawaii and Easter Island;
also found on the Great Barrier Reef and
Christmas Island and Cocos-Keeling Is-
lands in the eastern Indian Ocean. Curi-
ously, it remains unreported from any
continental shore or any of the large is-
lands of the Indo-Malayan region.
Perciformes: Percoidei: Cheilodactylidae 3329
CHEILODACTYLIDAE
Morwongs
by J.E. Randall
D iagnostic characters: Body oblong and moderately compressed (size to about 45 cm). A single flat
spine on opercle; margin of preopercle smooth. Mouth small, terminal to slightly inferior, the adults
with thick lips; maxilla largely exposed on cheek; no supramaxilla. Teeth small, villiform, in bands
anteriorly in jaws, none on vomer or palatines. Branchiostegal rays 6. A continuous dorsal fin with
XIV to XXII spines and 19 to 39 soft rays; anal fin with III spines and 7 to 19 soft rays; caudal fin
forked; lower 4 to 7 pectoral-fin rays unbranched, thickened, and prolonged, as much as the distal
50% free of membrane; pelvic fins with I spine and 5 soft rays, their base distinctly posterior to base of
pectoral fins. Scales cycloid and small, 45 to 85 in lateral line; dorsal and anal fins with a basal scaly sheath.
Vertebrae 13-14 + 21. Colour: most of the species have vertical to oblique dark bands.
Habitat, biology, and fisheries: Morwongs are bottom-dwelling fishes, mainly found inshore, though a
single species is taken commercially in southern Australia by trawling to depths of 400 m. They feed on a
wide variety of small invertebrates which are sorted from the mouthfuls of sand and detritus that they ingest
from the substratum. The postlarvae are very deep bodied and extremely compressed (sometimes referred
to as the ‘paper fish-stage’, and may attain large size. The late postlarval stage of Goniistius vittatus may
reach 6 cm total length. A specimen of this species from the northeastern Hawaiian Islands collected from
the nest of a white tern was mistakenly described as a new genus and new family, Gregoryinidae, by Fowler
and Ball (1924). Most of the species are valuable food fishes, but only a few occur marginally in the Western
Central Pacific.
Distribution: The Cheilodactylidae is primarily a temperate southern hemisphere family. Australia has the
largest number of species, 14; New Zealand has 6 (all shared with Australia), South Africa 5, Chile and
Peru 3, and Argentina 1. In the northern hemisphere, there are 3 species in Japan and China and 1 in
Hawaii. The group is clearly antitropical in its distribution.
Remarks: Randall (1983) classified 8 Indo-Pacific species of Cheilodactylus in the subgenus Goniistius.
In a current study of the Cheilodactylidae, Chris Burridge (pers. comm.) has concluded that Goniistius
should be elevated to a genus for these species except for Cheilodactylus nigripes that may warrant the
description of a new genus. He has reclassified C. ephippium and C. fuscus in the genus Morwong Whitley.
He wrote, “I believe that Cheilodactylus should be restricted to 2 South African species, C. fasciatus and
C. pixi.”
3330 Bony Fishes

Of the 4 families of fishes in the superfamily Cirrhitoidea,
only the hawkfish family (Cirrhitidae) occurs in the area
with the Cheilodactylidae. The species of this family are
easily separated by having X dorsal spines compared to
XIV to XXII for the cheilodactylids.
Cirrhitidae
Key to the species of Cheilodactylidae occurring in the area
1a. Anal-fin rays 16 or 17; dorsal-fin rays 27 or 28; pectoral-fin rays 14 or 15 (usually 15); no
bony protuberances anteriorly on head . . . . . . . . . . . . . . . . . . . . Nemadactylus douglasii
(southern Queensland to Tasmania and New Zealand)
1b. Anal-fin rays 8 or 9; dorsal-fin rays 29 to 35; pectoral-fin rays 13 or 14; adults with 2 pairs
of knob-like bony protuberances, 1 anterior to eyes and 1 just above upper lip . . . . . . . . . . ® 2
2a. Third and fourth dorsal-fin spines subequal; body grey-brown dorsally and on most of
caudal peduncle, shading to whitish below, with 2 narrow white bars on caudal peduncle
and 2 short oblique white bands below soft portion of dorsal fin . . . . . . . . . . . Morwong fuscus
(southern Queensland to Victoria, east to Lord Howe Island and New Zealand)
2b. Fourth dorsal-fin spine much longer than third spine, the third 3.75 to 5 times in fourth;
body whitish with 2 oblique black bands; head whitish, with 2 oblique black bands which
converge and join at pectoral-fin base . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
3a. Dorsal-fin rays 32 to 35; body depth 2.6 to 2.85 times in standard length; no oblique black
band from below eye to chest; pelvic fins not dark brown to black . . . . . . . . . Goniistius vestitus
(southern Queensland to central New South Wales, east to Lord Howe Island, Norfolk Island, and New Caledonia)
3b. Dorsal-fin rays 29 to 33; body depth 2.5 to 2.6 times in standard length; an oblique black
band from below eye to ventrally on chest; pelvic fins dark brown to black . . . . Goniistius vittatus
(Hawaii, Lord Howe Island, Kermadec Islands, and New Caledonia)

Goniistius vestitus (Castelnau, 1878)
Goniistius vittatus (Garrett, 1864)
Morwong fuscus (Castelnau, 1879)
Nemadactylus douglasii (Hector, 1875)
References
Gomon, M.J., G.C.M. Glover, and R.H. Kuiter (eds). 1994. The fishes of Australia’s south coast. Adelaide, State Print,
992 p.
Randall, J.E. 1983. A review of the fishes of the subgenus Goniistius, genus Cheilodactylus, with descriptions of a new
species form Easter Island and Rapa. Occ. Pap. B.P. Bishop Mus., 25(7):1-24.
Perciformes: Percoidei: Cepolidae 3331
CEPOLIDAE
Bandfishes
by W.F. Smith-Vaniz
D iagnostic characters: Small (to about 40 cm), moderately to noticeably elongate fishes with com-
pressed, tapering body and lanceolate caudal fin. Head short, with blunt snout. Eyes relatively
large and high on head. Mouth large, oblique; upper jaw broad at end, without supramaxilla, and extending
to below posterior margin of eye; a single row of slender, slightly curved teeth in jaws with an inner cluster
of teeth at symphysis in some species. Dorsal fin continuous, with 0 to IV flexible spines and 21 to 89
segmented rays; anal fin with 0 or I spine and 13 to 102 segmented rays; caudal fin lanceolate, middle
9 to 15 rays branched; pelvic fins positioned below or slightly anterior to pectoral fins, with I spine
and 5 segmented rays; outermost segmented pelvic-fin ray unbranched or weakly branched, 4 inner
rays distinctly branched. Lateral line high on body, close to dorsal-fin base, terminating posteriorly near
end of fin; lateral-line tubes or canals on scales not embedded in skin. Scales cycloid (smooth) or with
crenulate margins, relatively large to minute. Colour: in life red or pink; most species have a distinctive
dark stripe on the membrane (usually hidden) connecting the premaxillary and maxillary bones of the upper
jaw.
Sphenanthias macrophthalmus Owstonia maccullochi
Sphenanthias sibogae Acanthocepola limbata
Habitat, biology, and fisheries: Relatively uncommon fishes taken by trawls in shallow to deep depths (to
at least 475 m). Bandfishes occur on level bottom, sand or mud substrates where they live in burrows, which
they excavate themselves. Not of significant commercial importance, but consumed by the local population
in some areas.

The combination of a lanceolate tail, large oblique
mouth, and the arrangement of the pelvic-fin rays,
consisting of I spine and 5 segmented rays (the outer-
most ray unbranched or weakly branched and the
inner 4 branched), will distinguish the bandfishes from
all other families.
Additional distinguishing characters of the superficially
similar Opistognathidae are the following:
Opistognathidae: dorsal-fin spines X or XI (0 to IV in
Cepolidae); caudal fin rounded (moderately to strongly
lanceolate in Cepolidae); pelvic fins with 5 segmented Opisthognathidae
rays, the outer 2 unbranched.
3332 Bony Fishes
Partial key to the species of Cepolidae occurring in the area

Note: the taxonomy of this family is very poorly known and even generic definitions and limits are not well
established. Following traditional usage, Sphenanthias Weber, 1913 is tentatively recognized as distinct from
Owstonia Tanaka, 1908, solely on the basis of a different lateral line configuration (see key). However, some
species of “Sphenanthias” appear to be more closely related to species of Owstonia than to S. sibogae, the
type species. A number of undescribed species belonging to the subfamily Owstoniinae occur in the area
and at this time it is not possible to provide a reliable key for their identification.
1a. Total dorsal-fin elements (i.e. spines plus soft rays) 71 to 89; last soft ray of dorsal and
anal fins narrowly to broadly connected to caudal fin by a membrane . . (subfamily Cepolinae) ® 2
1b. Total dorsal-fin elements 23 to 27; last soft ray of dorsal and anal fins not connected to
caudal fin by a membrane . . . . . . . . . . . . . . . . . . . . . . . . (subfamily Owstoniinae) ® 6
2a. Preopercular margin with spines; cheeks scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3

2b. Preopercular margin without spines; cheeks naked . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
3a. Middle 8 to 10 caudal-fin rays branched (segmented caudal-fin rays 7 in upper lobe, 6 in
lower lobe); vertebrae 12 + 44-48 . . . . . . . . . . . . . . . . . . . . . . Acanthocepola abbreviata
3b. No branched caudal-fin rays; vertebrae 12 + 49-67 . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
4a. Segmented caudal-fin rays 6 or 7 in upper lobe, 6 or 7 in lower lobe (total 12 to 14);
vertebrae 12 + 49-52 . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthocepola krustensternii
4b. Segmented caudal-fin rays 5 in upper lobe, 5 in lower lobe (total 10); vertebrae 12 + 60-
67 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthocepola limbata
5a. Dorsal fin with III spines and 54 to 57 soft rays; anal fin with I spine and 50 to 53 soft
rays; vertebrae 14 + 45-48 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cepola australis
5b. Dorsal fin with III spines and 65 soft rays; anal fin with 62 soft rays, anal-fin spine absent;
vertebrae 16 + 53 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cepola schlegelii
6a. Lateral line continues anteriorly, forming loop in front of dorsal fin . . . . . . . . . . . Owstonia spp.
6b. Lateral line does not form loop in front of dorsal fin . . . . . . . . . . . . . . . . . Sphenanthias spp.

Acanthocepola abbreviata (Valenciennes in Cuvier and Valenciennes, 1835)
Acanthocepola krustensternii (Temminck and Schlegel, 1845)
Acanthocepola limbata (Valenciennes in Cuvier and Valenciennes, 1835)
(Cepola indica Day, 1888 and Acanthocepola cuneatus Smith, 1935 are both considered to be
synonyms)
Cepola australis Ogilby, 1899
Cepola schlegelii Bleeker, 1854
Owstonia dorypterus (Fowler, 1934)
Owstonia maccullochi Whitley, 1934 (may be a junior synonym of O. totomiensis)
Owstonia totomiensis Tanaka, 1908
Sphenanthias grammodon (Fowler, 1934)
Sphenanthias macrophthalmus Fourmanoir, 1985
Sphenanthias nigromarginatus Fourmanoir, 1985
Sphenanthias sibogae Weber, 1913 (S. pectinifer Myers, 1939 appears to be a synonym)
References
Mok, H.-K. 1988. Osteological evidence for the monophyly of Cepolidae and Owstoniidae. Japan. J. Ichthyol., 34
(4):507-508.
Smith, J.L.B. 1968. New and interesting fishes from deepish water off Durban, Natal and southern Mozambique. S. Afr.
Ass. Mar. Biol. Res. Invest. Rep., 19:1-30.
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Perciformes: Labroidei: Cichlidae 3333
Suborder LABROIDEI
CICHLIDAE
Cichlids
by K.E. Carpenter
D iagnostic characters (for brackish-water tolerant species introduced into the area): Medium-sized (to
about 60 cm) fishes with variable body shape, from deep bodied and compressed to perch-like. Head
with a single nostril on each side. A single dorsal fin with XIII to XIX spines and 10 to 16 soft rays;
anal fin with III (except XII or XIII in Etroplus) spines and 7 to 12 soft rays; caudal fin typically rounded,
truncate, or slightly emarginate (forked in Etroplus). Lateral line interrupted, with 26 to 40 (except 83 to
102 in Cichla ocellaris) scales. Colour: highly variable body colour from blue-grey, grey-green, olive
green, brownish, blackish, silvery grey, to pale dusky, often with bars or blotches on sides; scales
sometimes with individual dark markings; fins sometimes with spots, bars, and blotches, and sometimes
bordered with a band of red or pink; males often exhibit distinct breeding coloration.
a single dorsal fin with XIII-XIX spines
and 10-16 soft rays
lateral line interrupted
a single
nostril on
each side
of head
anal fin with III (XII-XIII

in Etroplus) spines and
7-12 soft rays
Habitat, biology, and fisheries: Primarily fresh-water fishes that tolerate but generally do not breed and
become established in brackish water; exceptions to this in the area are Etroplus suratensis which is a
brackish-water species that can only tolerate fresh or marine water for short periods and Oreochromis
mossambicus which is primarily fresh water but can breed and live in brackish water. All cichlids in the area
have been introduced and are native to Africa or south Asia. Many species have been introduced into the
wild by accidental release of aquaculture or aquarium fish specimens. Of the many cichlids reported to have
established wild populations in the area, only 9 species have tolerance to brackish water: Cichla ocellaris
is native to South America and introduced into Guam; Etroplus suratensis is native to India and Sri Lanka
and introduced into Sabah, Sarawak, and possibly Indonesia; Hemichromis bimaculatus is native to West
Africa and established in urban drains of Cairns, Australia; Oreochromis aureus is native to Africa and the
Middle East and introduced into Fiji, the Philippines, Singapore, and Thailand; O. mossambicus is native of
East Africa with widespread introductions in the area; O. niloticus niloticus is native of East Africa with
widespread introductions but mostly established in the wild in freshwater systems; O. urolepis is native of
East Africa and introduced into Fiji and Malaysia; Tilapia rendalli is native to southern and eastern Africa
and introduced into Thailand and Papua New Guinea; T. zilli is native to Africa and the Middle East and
introduced to New Caledonia, Guam, Fiji, the Philippines, Malaysia, and Northern Marianas. Cichla ocellaris
and Hemichromis bimaculatus are predators while the other species are plant and sediment feeders.
Breeding in cichlids typically involves pair-formation, nest-building, mouthbrooding, and parental care of
young. Cichlids include many very important aquarium and aquaculture species although mostly for
fresh-water culture. However, there is limited culture under brackish water conditions. Only Oreochromis
mossambicus is regularly captured and consumed from small-scale brackish water fisheries.
3334 Bony Fishes

Cichlids are easily distinguished from all other families of fishes
based on the normal perciform characteristics (e.g. spines in fins
and pelvic-fin formula of I spine and 5 soft rays) and the fact that
they have a single nostril on each side of the head and an
interrupted lateral line. The only other perciforms with these
characteristics are damselfishes (Pomacentridae).
Pomacentridae: differ from cichlids in having almost always II
anal-fin spines (usually III in cichlids); lateral line most often
incomplete, not extending onto caudal peduncle (interrupted in
cichlids); caudal fin most typically forked (typically rounded, trun-
cate, or emarginate in cichlids). Pomacentrids are coastal marine Pomacentridae
fishes only rarely found in brackish water (2 species of over 200
are found in brackish water).
Key to the species of Cichlidae occurring in the area

Note: the following key is relevant only to those species of cichlids tolerant of brackish water and currently
known to be introduced into the area.
1a. Anal fin with XII or XIII spines (Fig. 1)
. . . . . . . . . . . . . . . . . . . Etroplus suratensis
1b. Anal fin with III spines . . . . . . . . . . . . . . ® 2
2a. Moderately large conical teeth present in

jaws; juveniles without black spot on soft-
dorsal fin . . . . . . . . . . . . . . . . . . . . . . ® 3
2b. Teeth fine, close set; juveniles with black
spot on soft dorsal fin . . . . . . . . . . . . . . . ® 4
3a. Dorsal fin deeply incised, nearly dividing

spinous- and soft-rayed portions; a
prominent black spot on caudal fin near Fig. 1 Etroplus suratensis
upper base; around 83 to 102 lateral-line
scales (Fig. 2) . . . . . . . . . . . . . Cichla ocellaris
3b. Dorsal fin continuous although middle dorsal-fin soft rays elongate; no black spot on
caudal fin; 26 to 28 lateral-line scales (Fig. 3) . . . . . . . . . . . . . . . Hemichromis bimaculatus
Fig. 2 Cichla ocellaris Fig. 3 Hemichromis bimaculatus
4a. First gill arch with 8 to 12 gill rakers on lower limb; dark spot at base of soft dorsal fin in
adults and juveniles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Tilapia) ® 5
4b. First gill arch with 14 to 28 gill rakers on lower limb; dark spot at base of soft dorsal fin
in juveniles only . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Oreochromis) ® 6
Perciformes: Labroidei: Cichlidae 3335
5a. Dorsal fin and upper half of caudal fin with small spots; no bands along flank; bases of
scales on flanks dark (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tilapia rendalli
5b. Dorsal fin and upper half of caudal fin without small spots; 1 or more indistinct broad
bands along flank; bases of scales on flanks not darkened (Fig. 5) . . . . . . . . . . . . . Tilapia zilli
Fig. 4 Tilapia rendalli Fig. 5 Tilapia zilli
6a. Lower limb of first gill arch with 14 to 20 (modally 17 or 18) gill rakers; caudal fin without
distinct dark narrow bars (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . Oreochromis mossambicus
6b. Lower limb of first gill arch with 18 to 28 (modally greater than 20) gill rakers; caudal fin
with or without distinct narrow bars . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
7a. Caudal fin without prominent narrow dark bars, with a broad pink distal margin (Fig. 7)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oreochromis aureus
7b. Caudal fin with distinct narrow dark bars, without a broad pink distal margin . . . . . . . . . . . ® 8
Fig. 6 Oreochromis mossambicus Fig. 7 Oreochromis aureus

8a. Caudal fin mostly covered with narrow dark bars; sides without distinct marking or with
dark bars (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oreochromis niloticus
8b. Caudal fin with narrow dark bars on base and upper half; sides with 2 to 4 dark blotches
(Fig. 9) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Oreochromis urolepis
Fig. 8 Oreochromis niloticus Fig. 9 Oreochromis urolepis

3336 Bony Fishes

All species tolerant to brackish water that have been introduced into the area are listed below. A single
species, Oreochromis mossambicus, has become well established in brackish water environments on a
broad scale in the area.
Cichla ocellaris Bloch and Schneider, 1801
Etroplus suratensis (Bloch, 1790)
Hemichromis bimaculatus Gill, 1862
Oreochromis aureus (Steindachner, 1864)
Oreochromis mossambicus (Peters, 1852)
Oreochromis niloticus niloticus (Linnaeus, 1758)
Oreochromis urolepis honorum (Trewavas, 1966)
Oreochromis urolepis urolepis (Norman, 1922)
Tilapia rendalli (Boulenger, 1897)
Tilapia zilli (Gervais, 1848)
References
De Silva, S.S. (ed.) 1989. Exotic aquatic organisms in Asia. Asian Fish. Soc. Spec. Publ., 154 p.
Levêque, C.D. Paugy, and G.G. Teugels (eds). 1992. Faune des poissons d’eaux douces et saumâtres d’Afrique de
l’Ouest. Tome 2. Coll Faune Tropicale 28. Paris, Musée Royal de l’Afrique Centrale, Tervuren, Belgique and
O.R.S.T.O.M., 902 p.
Talwar, P.K. and A.G. Jhingran. 1992. Inland fishes of India and adjacent countries. Vol. 2. Rotterdam, A.A. Balkema.
Trewavas, E. 1983. Tilapiine fishes of the Genera Sarotherodon, Oreochromis, and Danakilia. Brit. Mus. Natl. Hist.,
583 p.
Welcomme, R.L. 1988. International introductions of inland aquatic species. FAO Fish. Tech. Pap., (294):318 p.
click for next page

Perciformes: Labroidei: Pomacentridae 3337
POMACENTRIDAE
Damselfishes (anemonefishes)
by J.R. Allen
D iagnostic characters: Small percoid fishes (to about 25 cm); body elongate to rounded in lateral
profile, and laterally compressed. Eye medium sized, generally exceeding snout length. Mouth small,
jaws equal and oblique. Small conical to incisiform teeth in jaws in a single row, sometimes with additional
row of slender teeth behind primary row; teeth absent on vomer and palatines. Total gill rakers on first gill
arch 10 to 85. A single dorsal fin with VII to XVII stout spines and 9 to 21 soft rays; anal fin with II
spines and 9 to 16 soft rays; pelvic fins with I spine and 5 soft rays; pectoral fins with 14 to 22 rays;
caudal fin rounded to forked. Scales comparatively large to moderate sized and ctenoid, extending onto
head and median fins; cheeks and operculum scaly; lateral-line scales 12 to 60. Colour: extremely variable
with many species exhibiting shades of brown, black, blue, green, red, and yellow; a dark spot or ocellus
often present on dorsal fin, particularly in juvenile stages.
dorsal fin with VII-XVII spines

and 9-21 soft rays
single nostril
on each side
anal fin with II spines

and 9-16 soft rays
Habitat, biology, and fisheries: Coral and rocky reefs, but also common in other shallow-water habitats
including sand and rubble patches, silty embayments, harbours, and seagrass beds. Pomacentrids are one
of the most numerous groups occurring on tropical reefs, in terms of both number of species and number
of individuals. Solitary fishes, pairs, or aggregations are equally common, depending on the species. Most
species are highly territorial, especially during reproductive periods. Spawning occurs throughout most of
the year at tropical localities. Eggs are deposited on the bottom and are guarded by the male. Hatching
occurs within 2 to 7 days. Most species have a pelagic larval stage lasting between about 10 to 50 days,
except Acanthochromis polyacanthus, which bypasses this stage entirely. Food habits are variable. The
larger species tend to feed mainly on algae, many others are omnivorous, consuming algae and small
invertebrates, while others rely on planktonic items. Larger species, such as various Abudefduf, are
sometimes seen in markets, but the family is not considered as an important food fish. Their most important
commercial use is as aquarium fishes. Anemone fishes are the leading group in this respect.
3338 Bony Fishes

Cichlidae: anal-fin spines III or more (only II in Pomacentridae); few species occurring in the area and all
of these introduced fresh-water forms, commonly referred to as “tilapia”.
Serranidae (genus Pseudanthias): generally resemble the pomacentrid genus Chromis, but easily distin-
guished by the presence of III anal-fin spines and smaller, more numerous scales (usually 35 to 65
lateral-line scales, compared to 12 to 20 in Chromis).
III spines
Cichlidae III or more spines Serranidae (Anthiinae)
Key to the genera of Pomacentridae occurring in the area

1a. Scales small, usually more than 50 in a longitudinal series from rear edge of opercle to
base of caudal fin; all the opercles usually serrate . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Scales comparatively large, usually less than 45 in a longitudinal series; all the opercles
not serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
2a. Suborbital usually armed with a pair of elongate spines (Fig. 1); scales small, about 68
to 76 in a longitudinal series from upper edge of opercle to caudal-fin base . . . . . . . . . Premnas
2b. Suborbital without pair of elongate spines, although serrae present (Fig. 2); scales
moderate sized, about 50 to 60 in a longitudinal series . . . . . . . . . . . . . . . . . . Amphiprion
Fig. 1 Premnas Fig. 2 Amphiprion Fig. 3
3a. Upper and lower edge of caudal-fin base usually with 2 or 3 projecting spiniform
procurrent rays (Fig. 3); teeth conical, either biserial or multiserial; head usually entirely
scaly except narrow region at front of snout and around each nostril . . . . . . . . . . . . . . . . ® 4
3b. Upper and lower edge of caudal-fin base without projecting spiniform rays; teeth conical
or incisiform, uniserial, or biserial (Fig. 4); head entirely scaly or frequently without scales
on snout, preorbital, and suborbital . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7
a) conical teeth (uniserial) b) incisiform teeth (uniserial) c) biserial teeth
Fig. 4
4a. Dorsal-fin spines usually XVII . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Acanthochromis

4b. Dorsal-fin spines XII to XV . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5
5a. Suborbital without scales (Fig. 5) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Altrichthys

5b. Suborbital scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
6a. Suborbital and hind margin of preopercle finely serrate (Fig. 6) . . . . . . . . . . . . . . . Dascyllus
6b. Suborbital entire or covered with scales; hind margin of preopercle usually smooth
(Fig. 7) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chromis
7a. Body very elongate, 2.9 to 3 times in standard length; small papilla-like structures on
inner edge of posterior circumorbitals (Fig. 8) . . . . . . . . . . . . . . . . . . . . . . . Lepidozygus
7b. Body orbiculate to elongate, usually 1.5 to 2.8 times in standard length; papilla-like
structures on posterior circumorbitals absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8
8a. Scales small, usually about 44 to 46 in longitudinal series from upper edge of gill opening
to caudal-fin base; body depth 2.4 to 2.8 times in standard length . . . . . . . . . . . Teixeirichthys
8b. Scales large, usually about 26 to 35 (except 30 to 46 in Parma, which has body depth
1.6 to 1.9 times in standard length) in longitudinal series from upper edge of gill opening
to caudal-fin base; body depth 1.5 to 3 times in standard length . . . . . . . . . . . . . . . . . . ® 9
Fig. 8 eye of Lepidozygus

Fig. 5 Altrichthys Fig. 6 Dascyllus Fig. 7 Chromis showing papillae
9a. Hind margin of preopercle crenulate or weakly to strongly serrate (Fig. 9a-c) . . . . . . . . . . ® 10
9b. Hind margin of preopercle smooth (Fig. 9d) . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 19
10a. Lips greatly thickened, fimbriate, and curled back over snout (Fig. 10) . . . . . . . . . . Cheiloprion
10b. Lips normal, not greatly thickened and curled back over snout . . . . . . . . . . . . . . . . . . ® 11
a) crenulate b) weakly c) strongly d) smooth

serrate serrate
Fig. 9 preopercular and suborbital serrations Fig. 10 Cheiloprion
3340 Bony Fishes
11a. Teeth of jaws uniserial (Fig. 11a) . . . . . . . . . . . . . . . . . ® 12

11b. Teeth of jaws biserial (at least anteriorly) (Fig. 10b, c) . . . . . . ® 15
a) uniserial
12a. Edge of subopercle serrate (Fig. 12) . . . . . . . . . . . . . Pristotis
12b. Edge of subopercle smooth . . . . . . . . . . . . . . . . . . . . ® 13
b) biserial
13a. Suborbital scaly; snout scaly to about level of nostrils
(Fig. 13) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Stegastes
13b. Suborbital and snout scaleless; predorsal scales extend-
ing to about level of anterior edge of eyes . . . . . . . . . . . . ® 14
c) partly biserial
14a. Body elongate, its depth 2.6 to 2.9 times in standard

length; dorsal-fin spines XIV (Fig. 14) . . . . . . . . . Pomachromis Fig. 11 pomacentrid dentition
14b. Body ovate, its depth 2.1 to 2.3 times in standard length; (viewed from inner side)
dorsal-fin spines XIII . . . . . . . . . . . . . . . Amblypomacentrus
suborbital suborbital
scaly naked
Fig. 12 Pristotis Fig. 13 Stegastes Fig. 14 Pomachromis
15a. Suborbital scaleless (at least anteriorly); hind margin of preopercle distinctly serrate
(Fig. 9c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
15b. Suborbital scaly; hind margin of preopercle crenulate or weakly serrate (Fig. 9a, b) . . . . . . . ® 17
16a. Snout mostly scaleless, predorsal scales

extending to about front of eyes or slightly
beyond; notch between preorbital and sub-
orbital absent (Fig. 15) . . . . . . . . . . . Dischistodus
16b. Snout scaly to about level of nostrils or be-
yond; usually a prominent notch between
preorbital and suborbital (much reduced in
certain species) (Fig. 16) . . . . . . . . . . Pomacentrus
17a. Body more or less elongate, its depth usu-

ally 2.3 to 2.8 times in standard length; outer
r o w o f t e e t h w i t h fl a tt e n e d ti p s
(Fig. 17); caudal fin lunate or strongly emar-
ginate; posterior portion of dorsal and anal Fig. 15 Dischistodus Fig. 16 Pomacentrus
fins and outer caudal-fin rays frequently
produced into filaments . . . . . . . . . Neopomacentrus
17b. Body ovate, its depth 2 to 2.3 times in stand-
ard length; outer row of teeth with conical
tips; caudal fin rounded or slightly emargi-
nate; outer caudal-fin rays and posterior
portion of dorsal and anal fins not produced
into filaments . . . . . . . . . . . . . . . . . . . . . ® 18 Fig. 17 teeth
18a. Dorsal- and anal-fin rays 14 or

15; pectoral-fin rays 18; slight
notch between preorbital and
suborbital (Fig. 18); spinous
dorsal fin without dark spot at
base of hindmost spines . . . Pomacentrus
(subgenus Lepidopomacentrus)
18b. Dorsal- and anal-fin rays 11 to
13; pectoral-fin rays 15 or 16;
notch between preorbital and Fig. 18 Pomacentrus Fig. 19 Abudefduf
suborbital absent; spinous (subgenus Lepidopomacentrus)
dorsal fin with prominent dark
spot at base of hindmost spine
. . . . . . . . . . . . . . . . . . Chrysiptera
(C. traceyi and C. talboti only)
19a. Vertical scale rows in excess of 30; horizontal scale rows below lateral line to origin of
anal fin more than 14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parma
19b. Vertical scale rows about 28; horizontal scale rows from lateral line to origin of anal fin
less than 12 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
20a. Horizontal scale rows above middle of lateral line usually 3 to 3 ½ (Fig. 19) . . . . . . . . Abudefduf
20b. Horizontal scale rows above middle of lateral line 1 to 2 ½ (usually 1 ½) . . . . . . . . . . . . . ® 21
21a. Gill rakers on first gill arch approximately 65 to 85 . . . . . . . . . . . . . . . . . . Hemiglyphidodon
21b. Gill rakers on first gill arch 10 to 35 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22
22a. Dorsal-fin spines XII . . . . . . . . Plectroglyphidodon
22b. Dorsal-fin spines XIII or XIV . . . . . . . . . . . . ® 23
23a. Body somewhat orbiculate, its depth 1.5 to
1.8 times in standard length; longest dor-
sal-fin spine about equal in length to dis-
tance from tip of snout to uppermost portion
of preopercle border (Fig. 20) . . . . Amblyglyphidodon
23b. Body moderately deep to elongate, its
depth usually 1.8 to 2.7 times in standard
length; longest dorsal-fin spine usually
much shorter than distance from tip of
snout to uppermost portion of preopercle
border . . . . . . . . . . . . . . . . . . . . . . . . ® 24
24a. Body moderately deep, its depth usually
less than 2 times in standard length; preor- Fig. 20 Amblyglyphidodon
bital and suborbital scaly (Fig. 21); teeth
biserial, at least anteriorly (Fig. 11c) . . Neoglyphidodon
24b. Body ovate to elon-
gate, its depth usu-
ally greater than 2
times in standard
length; preorbital
a n d s u b or b it al
scaly or naked
( F i g. 22 ) ; te et h
biserial or uniserial
(Fig. 11a, b) . . . . . Chrysiptera
a) b)
Fig. 21 Neoglyphidodon Fig. 22 Chrysiptera
3342 Bony Fishes

Abudefduf bengalensis (Bloch, 1787)
Abudefduf lorenzi Hensley and Allen, 1977
Abudefduf notatus (Day, 1869)
Abudefduf septemfasciatus (Cuvier, 1830)
Abudefduf sexfasciatus Lacepède, 1802
Abudefduf sordidus Forsskål, 1775
Abudefduf vaigiensis (Quoy and Gaimard, 1825)
Abudefduf whitleyi Allen and Robertson, 1974
Acanthochromis polyacantha (Bleeker, 1855)
Altrichthys azurelineatus (Fowler and Bean, 1928)
Altrichthys curatus Allen, 1999
Amblyglyphidodon aureus (Cuvier, 1830)
Amblyglyphidodon azurelineatus (Fowler and Bean, 1928)
Amblyglyphidodon batunai Allen, 1995
Amblyglyphidodon curacao (Bloch, 1787)
Amblyglyphidodon leucogaster (Bleeker, 1847)
Amblyglyphidodon ternatensis (Bleeker, 1853)
Amblypomacentrus breviceps (Schlegel and Müller, 1839-44)
Amblypomacentrus clarus Allen and Adrim, 2000
0
0
Amphiprion akallopisos Bleeker, 1853
0
Amphiprion akindynos Allen, 1972
Amphiprion chrysopterus Cuvier, 1830
0
Amphiprion clarkii (Bennett, 1830)
0
Amphiprion ephippium (Bloch, 1790)
0
Amphiprion frenatus Brevoort, 1856
0
Amphiprion latezonatus Waite, 1900
Amphiprion leucokranos Allen, 1973
0
Amphiprion mccullochi Whitley, 1929
0
Amphiprion melanopus Bleeker, 1852
0
Amphiprion ocellaris (Cuvier, 1830)
0
Amphiprion percula (Lacepède, 1802)
0
Amphiprion perideraion Bleeker, 1855
Amphiprion polymnus (Linnaeus, 1758)
0
Amphiprion rubrocinctus Richardson, 1842
Amphiprion sandaracinos Allen, 1972
Amphiprion sebae Bleeker, 1853
Amphiprion thiellei Burgess, 1981
Amphiprion tricinctus Schultz and Welander, 1953
Cheiloprion labiatus (Day, 1877)
Chromis abyssicola Allen and Randall, 1985
Chromis acares Randall and Swerdloff, 1973
Chromis agilis Smith, 1960
Chromis alleni Randall, Ida, and Moyer, 1981
Chromis alpha Randall, 1988
Chromis amboinensis (Bleeker, 1873)
Chromis analis (Cuvier, 1830)
Chromis atripectoralis Welander and Schultz, 1951
Chromis atripes Fowler and Bean, 1928
Chromis axillaris (Bennett, 1831)
Chromis caudalis Randall, 1988
Chromis chrysura (Bliss, 1883)
Chromis cinerascens (Cuvier, 1830)
Chromis delta Randall, 1988
Chromis dimidiata (Klunzinger, 1871)

Chromis elerae Fowler and Bean, 1928
Chromis flavipectoralis Randall, 1989
Chromis flavomaculata Kamohara, 1960
Chromis fumea (Tanaka, 1917)
Chromis hypsilepis (Günther, 1867)
Chromis iomelas Jordan and Seale, 1906
Chromis klunzingeri Whitley, 1929
Chromis lepidolepis Bleeker, 1877
Chromis leucura Gilbert, 1905
Chromis lineata Fowler and Bean, 1928
Chromis margaritifer Fowler, 1946
Chromis mirationis Tanaka, 1917
Chromis nigrura Smith, 1960
Chromis nitida (Whitley, 1928)
Chromis opercularis (Günther, 1867)
Chromis ovatiformes Fowler, 1946
Chromis retrofasciata Weber, 1913
Chromis scotochiloptera Fowler, 1918
Chromis ternatensis (Bleeker, 1856)
0
Chromis vanderbilti Fowler, 1941
Chromis viridis (Cuvier, 1830)
Chromis weberi Fowler and Bean, 1928
Chromis westaustralis Allen, 1976
Chromis xanthochira (Bleeker, 1851)
Chromis xanthura (Bleeker, 1854)
Chrysiptera biocellata (Quoy and Gaimard, 1824)
Chrysiptera bleekeri (Fowler and Bean, 1928)
Chrysiptera brownriggii (Bennett, 1828) [formerly C. leucopoma]
0
Chrysiptera caruleolineata (Allen, 1973)
Chrysiptera cyanea (Quoy and Gaimard, 1824)
Chrysiptera cymatilis Allen, 1999
Chrysiptera flavipinnis (Allen and Robertson, 1974)
Chrysiptera galba (Allen and Randall, 1974)
Chrysiptera glauca (Cuvier, 1830)
Chrysiptera hemicyanea (Weber, 1913)
Chrysiptera niger (Allen, 1975)
Chrysiptera notialis (Allen, 1975)
0
Chrysiptera oxycephala (Bleeker, 1877)
Chrysiptera parasema (Fowler, 1918)
Chrysiptera pricei Allen and Adrim, 1992
Chrysiptera rex (Snyder, 1909)
Chrysiptera rollandi (Whitley, 1961)
Chrysiptera sinclairi Allen, 1987
Chrysiptera springeri Allen and Lubbock, 1976
Chrysiptera starcki (Allen, 1973)
Chrysiptera talboti (Allen, 1975)
Chrysiptera taupou (Jordan and Seale, 1906)
Chrysiptera trayceyi (Woods and Schultz, 1960)
Chrysiptera tricincta (Allen and Randall, 1974)
Chrysiptera unimaculata (Cuvier, 1830)
0 Dascyllus aruanus (Linnaeus, 1758)
Dascyllus carneus Fischer, 1885
Dascyllus flavicaudus Randall and Allen, 1974
Dascyllus melanurus Bleeker, 1854
Dascyllus reticulatus (Richardson, 1846)
0
Dascyllus strasburgi Klausewitz, 1960
Dascyllus trimaculatus (Rüppell, 1928)
3344 Bony Fishes
Dischistodus chrysopoecilus (Schlegel and Müller, 1839)

Dischistodus fasciatus (Cuvier, 1830)
Dischistodus melanotus (Bleeker, 1858)
Dischistodus perspicillatus (Cuvier, 1830)
Dischistodus prosopotaenia (Bleeker, 1852)
Dischistodus pseudochrysopoecilus Allen and Robertson, 1974
Hemiglyphidodon plagiometopon (Bleeker, 1852)
Lepidozygus tapeinosoma (Bleeker, 1856)
Mecaenichthys immaculatus (Ogilby, 1885)
Neoglyphidodon bonang (Bleeker, 1853)
Neoglyphidodon carlsoni (Allen, 1975)
Neoglyphidodon crossi Allen, 1991
Neoglyphidodon melas (Cuvier, 1830)
Neoglyphidodon nigroris (Cuvier, 1830)
Neoglyphidodon oxyodon (Bleeker, 1857)
Neoglyphidodon polyacanthus (Ogilby, 1889)
Neoglyphidodon thoracotaeniatus (Fowler and Bean, 1928)
Neopomacentrus anabatoides (Bleeker, 1847)
Neopomacentrus azysron (Bleeker, 1877)
Neopomacentrus bankieri (Richardson, 1846)
Neopomacentrus cyanomos (Bleeker, 1856)
Neopomacentrus filamentosus (Macleay, 1833)
Neopomacentrus metallicus (Jordan and Seale, 1906)
Neopomacentrus nemurus (Bleeker, 1857)
Neopomacentrus taeniurus (Bleeker, 1856)
Neopomacentrus violascens (Bleeker, 1848)
Parma alboscapularis Allen and Hoese, 1975
Parma bicolor Allen and Larson, 1979
Parma mccullochi Whitley, 1929
Parma microlepis Günther, 1862
Parma occidentalis Allen and Hoese, 1975
Parma oligolepis Whitley, 1929
Parma polylepis Günther, 1862
Parma unifasciata (Steindachner, 1867)
Plectroglyphidodon dickii (Lienard, 1839)
Plectroglyphidodon flaviventris Allen and Randall, 1974
Plectroglyphidodon imparipennis (Vaillant and Sauvage, 1875)
Plectroglyphidodon johnstonianus Fowler and Ball, 1924
Plectroglyphidodon lacrymatus (Quoy and Gaimard, 1824)
Plectroglyphidodon leucozonus (Bleeker, 1859)
Plectroglyphidodon phoenixensis (Schultz, 1943)
Pomacentrus adelus Allen, 1991
Pomacentrus albimaculus Allen, 1975
Pomacentrus alexanderae Evermann and Seale, 1907
Pomacentrus alleni Burgess, 1981
Pomacentrus amboinensis Bleeker, 1868
Pomacentrus armillatus Allen, 1993
Pomacentrus auriventris Allen, 1991
Pomacentrus australis Allen and Robertson, 1973
Pomacentrus azuremaculatus Allen, 1991
Pomacentrus bankanensis Bleeker, 1853
Pomacentrur bintanensis Allen, 1999
Pomacentrus brachialis Cuvier, 1830
Pomacentrus burroughi Fowler, 1918
Pomacentrus chrysurus Cuvier, 1830
0 Pomacentrus coelestis Jordan and Starks, 1901

Pomacentrus colini Allen, 1991
Pomacentrus cuneatus Allen, 1991
Pomacentrus emarginatus Cuvier, 1830
Pomacentrus geminospilus Allen, 1993
Pomacentrus grammorhynchus Fowler, 1918
Pomacentrus imitator (Whitley, 1964)
Pomacentrus javanicus Allen, 1991
Pomacentrus komodoensis Allen, 1999
Pomacentrus lepidogenys Fowler and Bean, 1928
Pomacentrus limosus Allen, 1992
Pomacentrus littoralis Cuvier, 1830
Pomacentrus melanochir Bleeker, 1877
0
Pomacentrus milleri Taylor, 1964
Pomacentrus moluccensis Bleeker, 1853
Pomacentrus nagasakiensis Tanaka, 1917
Pomacentrus nigromanus Weber, 1913
Pomacentrus nigromarginatus Allen, 1973
Pomacentrus opisthostigma Fowler, 1918
Pomacentrus pavo (Bloch, 1878)
Pomacentrus philippinus Evermann and Seale, 1907
Pomacentrus polyspinus Allen, 1991
Pomacentrus reidi Fowler and Bean, 1928
Pomacentrus saksonoi Allen, 1995
Pomacentrus similis Allen, 1991
Pomacentrus simsiang Bleeker, 1856
Pomacentrus smithi Fowler and Bean, 1928
Pomacentrus stigma Fowler and Bean, 1928
Pomacentrus taeniometopon Bleeker, 1852
Pomacentrus tripunctatus Cuvier, 1830
Pomacentrus vaiuli Jordan and Seale, 1906
Pomacentrus wardi Whitley, 1927
Pomacentrus xanthosternus Allen, 1991
Pomachromis exilis Allen and Emery, 1972
Pomachromis fuscidorsalis Allen and Randall, 1974
Pomachromis guamensis Allen and Larson, 1975
Pomachromis richardsoni (Snyder, 1909)
0 Premnas biaculeatus (Bloch, 1790)
Pristotis jerdoni (Day, 1873)
Stegastes albifasciatus (Schlegel and Müller, 1839)
Stegastes apicalis (De Vis, 1885)
Stegastes aureus (Fowler, 1927)
Stegastes emeryi Allen and Randall, 1974
Stegastes fasciolatus (Ogilby, 1889)
Stegastes gascoynei (Whitley, 1964)
Stegastes insularis Allen and Emery, 1985
Stegastes lividus (Bloch and Schneider, 1801)
Stegastes nigricans (Lacepède, 1802)
Stegastes obreptus (Whitley, 1948)
References
Allen, G.R. 1972. Anemonefishes. Neptune City, New Jersey, T.F.H. Publications, Inc., 288 p.
Allen, G.R. 1975. Damselfishes of the south seas. Neptune City, New Jersey. T.F.H. Publications, Inc., 240 p.
Allen, G.R. 1975. The anemone fishes. Their classification and biology. Second edition. Neptune City, New Jersey,
T.F.H. Publications, Inc., 352 p.
Allen, G.R. 1991. Damselfishes of the world. Melle, Germany, Mergus Publishers, 271 p.
3346 Bony Fishes
Amphiprion akallopisos Bleeker, 1853

En - Skunk anemonefish.
Maximum total length about 10 to 11 cm. Coral reefs of lagoons and seaward reefs to depths of at
least 15 m. Associated with the large anemones; frequently an adult pair and several smaller fishes
with each anemone. Feeds on zooplankton, small benthic invertebrates, and algae. Used as an
aquarium fish; caught with hand nets. Widespread in the Indian Ocean, ranging to the west coast
of Thailand and southern coasts Sumatra and Java, also occurs in the Java Sea and around Bali.
Amphiprion akindynos Allen, 1972

En - Barrier reef anemonefish.
Maximum total length about 12 to 13 cm. Coral reefs of lagoons and seaward reefs to depths of at
least 20 m. Associated with the large anemones; frequently an adult pair and several smaller fish
with each anemone. Feeds on zooplankton, small benthic invertebrates, and algae. Used as an
aquarium fish; caught with hand nets. Great Barrier Reef and adjacent Coral Sea to New Caledonia
and the Loyalty Islands.
Amphiprion chrysopterus Cuvier, 1830

En - Orangefin anemonefish.
Maximum total length about 15 cm. Coral reefs of lagoons and seaward reefs to depths of at least
20 m. Associated with the large anemones; frequently an adult pair and several smaller fish with
each anemone. Feeds on zooplankton, small benthic invertebrates, and algae. Used as an aquarium
fish; caught with hand nets. Widespread in the western Pacific including New Guinea, Coral Sea,
New Britain, Solomon Islands, Vanuatu, Fiji, Caroline Islands, Mariana Islands, Gilbert Islands,
Samoa, Society Islands, and Tuamotu Islands.
dark variety
light variety Melanesia variety
Amphiprion ephippium (Bloch, 1790)

En - Red saddleback anemonefish.
12 m. Associated with the large anemones; frequently an adult pair and several smaller fish with
each anemone. Feeds on zooplankton, small benthic invertebrates, and algae. Used as an aquarium
fish; caught with hand nets. Andaman and Nicobar islands, Thailand, Malaysia, Sumatra, and Java.
3348 Bony Fishes
Amphiprion frenatus Brevoort, 1856

En - Tomato anemonefish.
12 m. Associated with the large anemones, frequently an adult pair and several smaller fish with
each anemone, male partner is considerably smaller than the female. Feeds on zooplankton, small
benthic invertebrates, and algae. An important aquarium fish; caught with hand nets. South China
Sea and adjacent areas, north to Japan.
Amphiprion latezonatus Waite, 1900

En - Wideband anemonefish.
Maximum total length about 14 cm. Coral and rocky reefs to depths of at least 40 m. Associated
with the large anemones, frequently an adult pair and several smaller fish with each anemone.
Feeds on zooplankton, small benthic invertebrates, and algae. Used as an aquarium fish; caught
with hand nets. Lord Howe Island off southeastern Australia and mainland reefs near the Queens-
land-New South Wales border.
Amphiprion leucokranos Allen, 1973

En - Whitebonnet anemonefish.
Maximum total length about 9 cm. Coral reefs of lagoons and seaward reefs to depths of at least 15 m.
Associated with the large anemones, usually solitary individuals or pairs. Feeds on zooplankton, small
benthic invertebrates, and algae. Sometimes seen in the aquarium trade; caught with hand nets.
Northern New Guinea, including Manus Island, also New Britain and the Solomon Islands.
Amphiprion melanopus Bleeker, 1852

En - Red-and-black anemonefish.
10 m. Usually associated with colonial anemones, frequently an adult pair and several smaller fish
with each cluster of anemones. Feeds on zooplankton, small benthic invertebrates, and algae. An
important aquarium fish; caught with hand nets. Western Pacific, including Indonesia (Bali east-
wards), Melanesia, Micronesia, southeastern Polynesia (to Society Islands), and from Great Barrier
Reef to Coral Sea. Fish from Fiji to the Society Islands have a uniform red ground colour and some
fish from the Coral Sea lack a white bar on the head.
Vanuatu variety
western Pacific variety
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3350 Bony Fishes
Amphiprion ocellaris (Cuvier, 1830)

En - False clown anemonefish.
Maximum total length about 8 cm. Coastal and lagoon coral reefs to depths of at least 12 m.
Associated with the large anemones, frequently an adult pair and several smaller fish with each
anemone. Feeds on zooplankton, small benthic invertebrates, and algae. An important aquarium
fish; caught with hand nets. Easternmost part of Indian Ocean and western edge of the Pacific,
including Andaman and Nicobar islands, Indo-Malayan Archipelago, northwestern Australia, Phil-
ippines, and northward to Japan. Very similar to Amphiprion percula, but usually has XI dorsal-fin
spines and a taller spiny dorsal fin.
light variety
dark variety
Amphiprion percula (Lacepède, 1802)

En - Clown anemonefish.
Maximum total length about 8 cm. Coastal and lagoon coral reefs to depths of at least 12 m.
anemone. Feeds on zooplankton, small benthic invertebrates, and algae. An important aquarium
fish; caught with hand nets. Southwestern Pacific, including northern Queensland, New Guinea,
New Britain, New Ireland, Solomon Islands, and Vanuatu.
Melanesia variety
Amphiprion perideraion Bleeker, 1855

En - Pink anemonefish.
Maximum total length about 10 cm. Coral reefs of lagoons and seaward reefs to depths of at least 20 m.
anemone. Feeds mainly on algae, but also consumes zooplankton. An important aquarium fish; caught
with hand nets. Easternmost Indian Ocean and western Pacific, including Cocos (Keeling) Islands,
Christmas Island, and Indo-Australian Archipelago northwards to Micronesia and Japan.
Amphiprion polymnus (Linnaeus, 175)

En - Saddleback anemonefish.
Maximum total length about 12 cm. Sand bottoms adjacent to coastal, lagoon, and seaward coral
reefs to depths of at least 30 m. Associated with the large anemones, frequently an adult pair and
several smaller fish with each anemone. Feeds on zooplankton, small benthic invertebrates, and
algae. Sometimes seen in the aquarium trade; caught with hand nets. Indo-Malayan Archipelago
northwards to the Ryukyu Islands; also reported from the Northern Territory, Australia.
3352 Bony Fishes
Amphiprion sandaracinos Allen, 1972

En - Orange anemonefish.
15 m. Associated with the large anemones, frequently an adult pair and several smaller fish with
each anemone. Feeds on zooplankton, small benthic invertebrates, and algae. An important
aquarium fish; caught with hand nets. Easternmost part of Indian Ocean at Christmas Island and
Western Australia, also Indonesia, Melanesia, Philippines, and northwards to the Ryukyu Islands.
Chromis viridis (Cuvier, 1830)

En - Bluegreen chromis.
Maximum total length about 7 cm. Blue to pale green, depending on viewing angle. Lagoons and
sheltered reefs to depths of about 12 m. Forms aggregations around coral heads, especially
branching Acropora. Feeds on zooplankton. Sometimes captured for the aquarium trade; caught
with hand nets and surround nets. Widespread in the Indo-West Pacific from East Africa and the
Red Sea to the Line Islands and southeastern Polynesia, northward to southern Japan.
Chrysiptera cyanea (Quoy and Gaimard, 1824)

En - Blue devil.
Maximum total length about 8 cm. Generally bright blue. Coral reefs of lagoons and protected
coastal reefs to a depth of about 10 m; forms aggregations with 1 male to several females. Feeds
on zooplankton, small benthic invertebrates, and algae. An important aquarium fish; caught with
hand nets and surround nets. At most localities mature males have an orange tail. Females have
a bluish to translucent tail and a black spot at the base of the last dorsal-fin rays. Western Pacific
and the eastern edge of the Indian Ocean, including Western Australia, Great Barrier Reef, New
Guinea, New Britain, Solomon Islands, Mariana and Caroline islands, Indo-Malayan Archipelago,
Philippines, Taiwan Province of China, and Ryukyu Islands. Chrysiptera taupou from the Coral Sea
to Samoa is similar but has a yellow belly and tail.
Chrysiptera parasema (Fowler, 1918)

En - Goldtail demoiselle.
Maximum total length about 6 cm. Generally bright blue, abruptly yellow on posteriormost part of
body including tail. Coral reefs of lagoons and sheltered inshore reefs to depths of at least 16 m.
Occurs in aggregation amongst rich coral growth. Feeds mainly on zooplankton. An important
aquarium fish; caught with hand nets and surround nets. Western Pacific including Solomon Islands,
northern New Guinea, Indonesia, Philippines, and Solomon Islands. Several similar species in the
area, including C. hemicyanea (yellow belly), C. springeri (entirely deep blue, usually with XII
dorsal-fin spines) and C. sinclairi (entirely blue, usually with XIII dorsal-fin spines).
3354 Bony Fishes
Dascyllus aruanus (Linnaeus, 1758)

En - Humbug dascyllus.
Maximum total length about 7 cm. Coral reefs of lagoons and sheltered inshore reefs to depths of
about 12 m. Forms aggregations around small coral heads. Feeds on zooplankton, small benthic
invertebrates, and algae. An important aquarium fish; caught with hand nets. Widespread in the
Indo-West Pacific from East Africa and the Red Sea to the Line Islands and southeastern Polynesia,
northward to southern Japan. Another species in the area, Dascyllus melanurus, is similar, but
differs in having the posterior half of the caudal fin black.
Dascyllus trimaculatus (Rüppell, 1928)

En - Threespot dascyllus.
55 m. Juveniles frequently associated with the large anemones with Amphiprion. Feeds on
zooplankton, small benthic invertebrates, and algae. An important aquarium fish; caught with hand
nets. Widespread in the Indo-West Pacific from East Africa and the Red Sea to Polynesia and
Australia, northwards to Japan.
Pomacentrus coelestis Jordan and Starks, 1901

En - Neon damselfish.
Maximum total length about 10 cm. Generally neon blue with whitish to clear (occasionally pale
yellow) anal fin and tail. Coral reefs of lagoons and seaward reefs to depths of about 12 m. Forms
aggregations on rubble bottoms. Feeds mainly on zooplankton. An important aquarium fish; caught
with hand nets. Widespread in the eastern Indian Ocean and western Pacific including the Andaman
Sea, Cocos (Keeling) Islands, Christmas Island, Indo-Australian Archipelago, and Oceania in
general as far east as the Line Islands and Tuamotus, and northward to Japan. Similar species in
the area include Pomacentrus alleni (yellow anal fin and black streak on lower edge of caudal fin)
and P. auriventris (yellow on lower half of body).
Pomacentrus moluccensis Bleeker, 1853

En - Lemon damselfish.
Maximum total length about 7.5 cm. Bright yellow with small black spot, at upper pectoral-fin base.
Coral reefs of lagoons and seaward reefs to depths of at least 12 m. Found in areas of rich coral
growth. Feeds mainly on zooplankton, small benthic invertebrates, and algae. Fiji fish are variably
dusky purplish. An important aquarium fish; caught with hand nets and surround nets. Eastern Indian
Ocean and western Pacific including the Andaman Sea, Indo-Australian Archipelago east to Fiji,
and northwards to the Ryukyu Islands.
3356 Bony Fishes
Premnas biaculeatus (Bloch, 1790)

En - Spinecheek anemonefish.
Maximum total length about 16 cm. Sheltered inshore reefs, lagoons, and seaward reefs to depths
of at least 52 m. Occurs in pairs associated with the large anemones, male fish usually less than
half as large as female. Feeds on zooplankton, small benthic invertebrates, and algae. An important
aquarium fish; caught with hand nets. Indo-Australian Archipelago.
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Index of Scientific and Vernacular Names 3357
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.
3358 The Living Marine Resources of the Western Central Pacific
A Anemones . . . . . . . . . .
Angel mono . . . . . . . . .
.
.
.
.
.
.
.
.
.
.
. . . . . . 3354
. . . . . . 3219
abbreviatus, Gerres . . . . . . . . . . . . . . . . 2950 Angelfishes . . . . . . . . . . . . . . . . . . . . . 3266
Abudefduf . . . . . . . . . . . . . . . . . . . . . 3337 annularis, Pomacanthus . . . . . . . . . . . . . 3284
Acanthocepola limbata . . . . . . . . . . . . . . 3331 antarctica, Sciaena . . . . . . . . . . . . . . . . 3127
Acanthochromis polyacanthus . . . . . . . . . . 3337 antonbruun, Atrobucca . . . . . . . . . . . . . . 3133
Acanthopagrus australis . . . . . . . . . . . . . 2995 Aphareus . . . . . . . . . . . . . . . 2840, 2843, 2991
Acanthopagrus berda . . . . . . . . . . . . . . . 2996 Aphareus furca . . . . . . . . . . . . . . . . . . 2854
Acanthopagrus latus . . . . . . . . . . . . . . . 2997 Aphareus furcatus . . . . . . . . . . . . . . . . . 2854
ACANTHURIDAE . . . . . . . . . . . . . . 3225, 3267 Aphareus rutilans . . . . . . . . . . . . . . . . . 2855
See also Vol. 6 APOGONIDAE . . . . . . . . . . . . . . . . . . . 3118
Acanthurus pyroferus . . . . . . . . . . . . 3275, 3279 See also Vol. 4
Acanthurus tristis . . . . . . . . . . . . . . . . . 3274 Apogonids . . . . . . . . . .
. . . . . . . . . . . 3118
achlamys, Gazza . . . . . . . . . . . . . . . . . . 2798 Apolemichthys . . . . . . . .
. . . . . . . . . . . 3267
acinaces, Gerres . . . . . . . . . . . . . . . . . . 2951 Apolemichthys griffisi . . .
. . . . . . . . . . . 3272
Acropora . . . . . . . . . . . . . . . . . . . . . . 3352 Apolemichthys trimaculatus . . . . . . . . . . . 3272
acuminatus, Heniochus . . . . . . . . . . . . . . 3262 Apolemichthys xanthopunctatus . . . . . . . . . 3272
acuta, Nibea . . . . . . . . . . . . . . . . . . . . 3137 Aprion virescens . . . . . . . . . . . . . . . . . 2856
adetii, Lutjanus . . . . . . . . . . . . . . . . . . 2861 Archerfishes . . . . . . . . . . . . . . . . . . . . 3212
adiergastos, Chaetodon . . . . . . . . . . . . . . 3234 arge, Upeneus . . . . . . . . . . . . . . . . . . . 3195
adusta, Atrobucca . . . . . . . . . . . . . . . . . 3130 argentatus, Chaetodon . . . . . . . . . . . . . . 3235
adusta, Pempheris . . . . . . . . . . . . . . . . . 3201 argenteus, Mesopristes . . . . . . . . . . . . . . 3309
aequidens, Atractoscion . . . . . . . . . . . . . 3129 argenteus, Monodactylus . . . . . . . 3216, 3218-3219
aequipinnis, Scorpis . . . . . . . . . . . . . . . . 3299 argenteus, Otolithes . . . . . . . . . . . . . . . . 3164
aesticola, Pteraclis . . . . . . . . . . . . . . . . 2834 argenteus, Pomadasys . . . . . . . . . . . . . . 2983
affinis, Scolopsis . . . . . . . . . . . . . . . . . 3086 argentimaculatus, Lutjanus . . . . . . . . . . . 2862
akallopisos, Amphiprion . . . . . . . . . . . . . 3346 argyreus, Gerres . . . . . . . . . . . . . . . 2951, 2956
akindynos, Amphiprion . . . . . . . . . . . . . . 3346 argyreus, Pomadasys . . . . . . . . . . . . . . . 2984
albacares, Thunnus . . . . . . . . . . . . . . . . 2832 argyrogrammicus, Pristipomoides . . . . . . . . 2908
albovittatus, Plectorhinchus . . . . . . . . . . . 2970 Argyrops spinifer . . . . . . . . . . . . . . . . . 2998
alleni, Pomacentrus . . . . . . . . . . . . . . . . 3355 Argyrosomus . . . . . . . . . . . . . . . . . . . 3122
alternatus, Cirrhites . . . . . . . . . . . . . . . . 3326 Argyrosomus amoyensis . . . . . . . . . . . . . 3126
alternatus, Cirrhitus . . . . . . . . . . . . . . . . 3326 Argyrosomus hololepidotus . . . . . 3127, 3129, 3131
altivelis, Coradion . . . . . . . . . . . . . . . . . 3258 Argyrosomus japonicus . . . . . . . . . . . . . . 3127
amamianus, Lethrinus . . . . . . . . . . . . . . . 3036 Argyrosomus macrocephalus . . . . . . . . . . . 3169
amblycephalus, Johnius . . . . . . . . . . 3138, 3149 Argyrosomus pawak . . . . . . . . . . . . . . . . 3170
amblycephalus, Johnius (Johnius) . . . . . . . . 3139 armatus, Enoplosus . . . . . . . . . . . . . . . . 3287
Amblycirrhitus . . . . . . . . . . . . . . . . . . 3323 armatus, Neocirrhites . . . . . . . . . . . . . . . 3321
amboinensis, Lethrinus . . . . . . . . . . . . . . 3026 Armour croaker . . . . . . . . . . . . . . . . . . . 3167
Ambon emperor . . . . . . . . . . . . . . . . . . 3026 Armourheads . . . . . . . . . . . . . . . . . . . . 3288
Amniataba caudavittatus . . . . . . . . . . . . . 3308 Arquero ensillado . . . . . . . . . . . . . . . . . 3215
amoenus, Pristipomoides . . . . . . . . . . . . . 2908 Arquero manchado . . . . . . . . . . . . . . . . . 3215
Amoy croaker . . . . . . . . . . . . . . . . . . . 3126 ARRIPIDAE . . . . . . . . . . . . . . . . . . . . 3304
amoyensis, Argyrosomus . . . . . . . . . . . . . 3126 Arripis georgianus . . . . . . . . . . . . . . . . 3304
Amphichaetodon howensis . . . . . . . . . . . . 3234 Arripis trutta . . . . . . . . . . . . . . . . . . . . 3304
Amphiprion . . . . . . . . . . . . . . . . . . . . 3354 aruanus, Dascyllus . . . . . . . . . . . . . . . . 3354
Amphiprion akallopisos . . . . . . . . . . . . . . 3346 Asian butterflyfish . . . . . . . . . . . . . . . . . 3235
Amphiprion akindynos . . . . . . . . . . . . . . 3346 asper, Taractes . . . . . . . . . . . . . . . . . . 2835
Amphiprion chrysopterus . . . . . . . . . . . . . 3347 Aspericorvina jubata . . . . . . . . . . . . . . . 3128
Amphiprion ephippium . . . . . . . . . . . . . . 3347 atkinsoni, Lethrinus . . . . . . . . . . . . . . . 3027
Amphiprion frenatus . . . . . . . . . . . . . . . 3348 Atractoscion aequidens . . . . . . . . . . . . . . 3129
Amphiprion latezonatus . . . . . . . . . . . . . 3348 Atrobucca . . . . . . . . . . . . . . . . . . . . . 3122
Amphiprion leucokranos . . . . . . . . . . . . . 3349 Atrobucca adusta . . . . . . . . . . . . . . . . . 3130
Amphiprion melanopus . . . . . . . . . . . . . . 3349 Atrobucca antonbruun . . . . . . . . . . . . . . 3133
Amphiprion ocellaris . . . . . . . . . . . . . . . 3350 Atrobucca brevis . . . . . . . . . . . . . . . . . 3131
Amphiprion percula . . . . . . . . . . . . . . . . 3350 Atrobucca kyushini . . . . . . . . . . . . . . . . 3132
Amphiprion perideraion . . . . . . . . . . . . . 3351 Atrobucca nibe . . . . . . . . . . . . . . . . . . 3133
Amphiprion polymnus . . . . . . . . . . . . . . 3351 Atypichthys strigatus . . . . . . . . . . . . . . . 3303
Amphiprion sandaracinos . . . . . . . . . . . . 3352 audleyi, Gymnocranius . . . . . . . . . . . . . . 3018
Amphitherapon caudavittatus . . . . . . . . . . . 3308 aurantius, Centropyge . . . . . . . . . . . . . . 3273
analis, Hapalogenys . . . . . . . . . . . . . . . . 2968 auratus, Chrysophrys . . . . . . . . . . . . . . . 3002
anarhynchus, Lethrinus . . . . . . . . . . . . . . 3033 auratus, Pagrus . . . . . . . . . . . . . . . . . . 3002
anea, Pennahia . . . . . . . . . . . . . . . . . . 3168 auratus, Polydactylus . . . . . . . . . . . . . . . 3108
Anemonefishes . . . . . . . . . . . . . . . . . . . 3337 auratus, Scolopsis . . . . . . . . . . . . . . . . . 3087
aureofasciatus, Chaetodon . . . . . . . . . . . . 3235 bathybius, Nemipterus . . . . . . . . . . . . . . 3063

aureus, Chrysochir . . . . . . . . . . . . . . . . 3137 Bathyclupea gracilis . . . . . . . . . . . . . . . 3211
aureus, Leiognathus . . . . . . . . . . . . . . . 2800 BATHYCLUPEIDAE . . . . . . . . . . . . . . . . 3211
aureus, Oreochromis . . . . . . . . . . . . . . . 3333 Bathyclupeids . . . . . . . . . . . . . . . . . . . 3211
auricilla, Pristipomoides . . . . . . . . . . . . . 2909 Beach salmon . . . . . . . . . . . . . . . . . . . 3210
aurifilum, Nemipterus . . . . . . . . . . . . . . 3078 Bearded croaker . . . . . . . . . . . . . . . . . . 3139
auriflamma, Mulloides . . . . . . . . . . . . . . 3183 Belanger’s croaker .. . . . . . . . . . . . . . . . 3141
auriga, Chaetodon . . . . . . . . . . . . . . . . 3236 belangerii, Johnius . . . . . . . . . . . . . 3140, 3142
auripes, Chaetodon . . . . . . . . . . . . . . . . 3236 belangerii, Johnius (Johnius) . . . . . . . . . . 3141
auritus, Pomadasys . . . . . . . . . . . . . . . . 2985 bellus, Genicanthus . . . . . . . . . . . . . . . . 3282
auriventris, Pomacentrus . . . . . . . . . . . . . 3355 Bengal snapper . . . . . . . . . . . . . . . . . . 2863
aurolineatus, Gnathodentex . . . . . . . . . . . 3017 bengalensis, Lutjanus . . . . . . . . . . . . . . . 2863
aurora, Nemipterus . . . . . . . . . . . . . . . . 3062 Bennett’s butterflyfish . . . . . . . . . . . . . . . 3237
Australian emperor . .. . . . . . . . . . . . . . . 3048 bennetti, Chaetodon . . . . . . . . . . . . . . . . 3237
Australian Mado . . .. . . . . . . . . . . . . . . 3303 Berber ponyfish . . . . . . . . . . . . . . . . . . 2801
Australian salmon . .. . . . . . . . . . . . . . . 3304 berbis, Leiognathus . . . . . . . . . . . . . . . . 2801
Australian salmons . .. . . . . . . . . . . . . . . 3304 berda, Acanthopagrus . . . . . . . . . . . . . . 2996
Australian seabream .. . . . . . . . . . . . . . . 2995 berda, Mylio . . . . . . . . . . . . . . . . . . . . 2996
Australian threadfin .. . . . . . . . . . . . . . . 3108 BERYCIDAE . . . . . . . . . . . . . . . . . . . . 3201
australis, Acanthopagrus . . . . . . . . . . . . . 2995 See also Vol. 4
australis, Brama . . . . . . . . . . . . . . . . . . 2833 Besugato cariblanco . . . . . . . . . . . . . . . . 3089
australis, Johnius . . . . . . . . . . . . . . 3141-3142 Besugato de banda dorada . . . . . . . . . . . . 3087
australis, Johnius (Johnius) . . . . . . . . . . . 3140 Besugato rayado . . . . . . . . . . . . . . . . . . 3087
Austronibea oedogenys . . . . . . . . . . . . . . 3134 Besugato rosado . . . . . . . . . . . . . . . . . . 3081
Authistes puta . . . . . . . . . . . . . . . . . . . 3315 biaculeatus, Premnas . . . . . . . . . . . . . . . 3356
biauritus, Otolithoides . . . . . . . . . . . . . . 3165
B Bicolor angelfish . . . . . .
bicolor, Centropyge . . . .
. .
. .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. . . 3273
. . . 3273
baconensis, Gerres . . . . . . . . . . . . . . . . 2959 Bicolour goatfish . . . . . . . . . . . . . . . . . . 3184
Baga elegante . . . . .. . . . . . . . . . . . . . 3081 bifasciatus, Parupeneus . . . . . . . . . . . . . 3186
Baga japonesa . . . . .. . . . . . . . . . . . . . 3068 bifasciatus, Pentapodus . . . . . . . . . . . . . . 3083
Baga tolu . . . . . . . .
. . . . . . . . . . . . . . 3071 Bigbelly pomfret . . . . . . . . . . . . . . . . . . 2832
Bahaba polykladiskos . . . . . . . . . . . . . . . 3135 Bigeye snapper . . . . . . . . . . . . . . . . . . 2882
Baldspot monocle bream . . . . . . . . . . . . . . 3088 Bighead pennah croaker . . . . . . . . . . . . . . 3169
balinensis, Nemipterus . . . . . . . . . . . . . . 3078 bigibbus, Kyphosus . . . . . . . . . . . . . . . . 3292
balinensoides, Nemipterus . . . . . . . . . . . . 3078 Bigmouth croaker . . . . . . . . . . . . . . . . . 3172
Balinese threadfin bream . . . . . . . . . . . . . 3078 Bigscale pomfret . . . . . . . . . . . . . . . . . . 2836
balteatus, Dipterygonotus . . . . 2838, 2919-2920, 2931 Bigscaled ponyfish . . . . . . . . . . . . . . . . . 2821
Banana fusilier . . . . . . . . . . . . . . . . . . . 2937 Bigsnout croaker . . . . . . . . . . . . . . . . . . 3150
Banded archerfish . . . . . . . . . . . . . . . . . 3215 biguttatus, Lutjanus . . . . . . . . . . . . . . . . 2864
Banded grunt . . . . . . . . . . . . . . . . . . . . 2986 bilineatus, Scolopsis . . . . . . . . . . . . . . . 3087
Bandfishes . . . . . . . . . . . . . . . . . . . . . 3331 bimaculatus, Hemichromis . . . . . . . . . . . . 3333
Bandtail goatfish . . . . . . . . . . . . . . . . . . 3195 bindus, Leiognathus . . . . . . . . . . . . 2802, 2815
Baraonga aurora . . . . . . . . . . . . . . . . . . 3311 bipunctatus, Nemipterus . . . . . . . . . . . . . 3062
Baraonga jarbua . . . . . . . . . . . . . . . . . . 3314 bispinosus, Centropyge . . . . . . . . . . . . . . 3273
Baraonga ronco . . . . . . . . . . . . . . . . . . 3315 bitaeniatus, Lutjanus . . . . . . . . . . . . . . . 2865
Baraonga tigre . . . . . . . . . . . . . . . . . . . 3316 bitorquatus, Gymnocranius . . . . . . . . . . . . 3018
barberinoides, Parupeneus . . . . . . . . . . . . 3184 Black butterflyfish . . . . . . . . . . . . . . . . . 3240
barberinus, Parupeneus . . . . . . . . . . . . . 3185 Black drummer . . . . . . . . . . . . . . . . . . . 3298
Barbet à point rouge . . . . . . . . . . . . . . . . 3189 Black eyebrow monocle bream . . . . . . . . . . 3082
Barbet souffré . . . . . . . . . . . . . . . . . . . 3264 Black-and-white snapper . . . . . . . . . . . . . . 2898
Barbudo de mancha . . . . . . . . . . . . . . . . 3112 Blackbacked butterflyfish . . . . . . . . . . . . . 3243
Barbudo indio . . . . . . . . . . . . . . . . . . . 3104 Blackbanded snapper . . . . . . . . . . . . . . . 2893
Barbudo mamalí . . . . . . . . . . . . . . . . . . 3096 Blackblotch emperor . . . . . . . . . . . . . . . . 3043
Barbudo rayado . . . . . . . . . . . . . . . . . . 3110 Blackeared grunt . . . . . . . . . . . . . . . . . . 2989
Barbure indien . . . . . . . . . . . . . . . . . . . 3104 Blackfin threadfin . . . . . . . . . . . . . . . . . . 3109
Barbure mamalí . . . . . . . . . . . . . . . . . . 3096 Blackhand paradise fish . . . . . . . . . . . . . . 3115
Barbure rayé . . . . . . . . . . . . . . . . . . . . 3110 Blackmouth croaker . . . . . . . . . . . . . . . . 3133
Barbure tache noire . . . . . . . . . . . . . . . . 3112 Blacknape large-eye bream . . . . . . . . . . . . 3025
Baroness butterflyfish . . . . . . . . . . . . . . . 3237 Blackside hawkfish . . . . . . . . . . . . . . . . . 3327
baronessa, Chaetodon . . . . . . . . . . . . . . 3237 Blackspot angelfish . . . . . . . . . . . . . . . . 3277
Barred flagtail . . . . . . . . . . . . . . . . . . . 3319 Blackspot croaker . . . . . . . . . . . . . . . . . 3132
Barred tigerfish . . . . . . . . . . . . . . . . . . . 2945 Blackspot goatfish . . . . . . . . . . . . . . . . . 3194
Barrier reef anemonefish . . . . . . . . . . . . . . 3346 Blackspot snapper . . . . . . . . . . . . . . . . . 2873
Blackspot threadfin . .. . . . . . . . . . . . . . . 3112 Brownstripe snapper . . . . . . . . . . . . . . . . 2896

Blacktail snapper . . .. . . . . . . . . . . . . . . 2874 brunneus, Otolithoides . . . . . . . . . . . . . . 3165
Blackvelvet angelfish .. . . . . . . . . . . . . . . 3281 buergeri, Glaucosoma . . . . . . . . . . . . . . 3207
Blanche à pagaies . .. . . . . . . . . . . . . . . 2958 Bullseyes . . . . . . . . . . . . . . . . . . . . . . 3201
Blanche armie . . . .. . . . . . . . . . . . . . . 2957 bürgeri, Glaucosoma . . . . . . . . . . . . . . . 3207
Blanche commune . .. . . . . . . . . . . . . . . 2956 Burro arlequin . . . . . . . . . . . . . . . . . . . 2971
Blanche élégante . . .. . . . . . . . . . . . . . . 2955 Burro limón . . . . . . . . . . . . . . . . . . . . . 2973
Blanche elévée . . . .. . . . . . . . . . . . . . . 2950 Burro listado . . . . . . . . . . . . . . . . . . . . 2980
Blanche fil . . . . . .. . . . . . . . . . . . . . . 2952 Burro negro . . . . . . . . . . . . . . . . . . . . . 2974
Blanche gouvernail . .. . . . . . . . . . . . . . . 2951 Burro oriental . . . . . . . . . . . . . . . . . . . . 2982
blochii, Leiognathus . . . . . . . . . . . . . . . 2803 Burro trovador . . . . . . . . . . . . . . . . . . . 2981
Blotched grunt . . . . . . . . . . . . . . . . . . . 2988 Burro trucha . . . . . . . . . . . . . . . . . . . . 2978
Blotched therapon . . . . . . . . . . . . . . . . . 3313 Burro velero . . . . . . . . . . . . . . . . . . . . 2967
Blotched tigertoothed croaker . . . . . . . . . . . 3173 Butterfly whiptail . . . . . . . . . . . . . . . . . . 3075
Blubberlip snapper . . . . . . . . . . . . . . . . . 2889 Butterflyfishes . . . . . . . . . . . . . . . . . . . 3224
Blue devil . . . . . . . . . . . . . . . . . . . . . . 3353
Blue sea chub . . . . . . . .
Blue-and-gold fusilier . . . . .
.
.
.
.
.
.
.
.
.
.
. . . . . . 3293
. . . . . . 2926
C
Bluecheek silver grunt . . . . . . . . . . . . . . . 2984 caerulaurea, Caesio . . . . . . . . . . . . . . . . 2926
Bluecheek threadfin bream . . . . . . . . . . . . 3080 caeruleopunctatus, Chaetodontoplus . . . . . . 3279
Bluefaced angelfish . . . . . . . . . . . . . . . . 3286 Caesio azuror . . . . . . . . . . . . . . . . . . . 2926
Bluegirdled angelfish . . . . . . . . . . . . . . . . 3285 Caesio caerulaurea . . . . . . . . . . . . . . . . 2926
Bluegreen chromis . . . . . . . . . . . . . . . . . 3352 Caesio ceinture d’or . . . . . . . . . . . . . . . . 2933
Bluelined large-eye bream . . . . . . . . . . . . . 3022 Caesio croissant . . . . . . . . . . . . . . . . . . 2928
Blueringed angelfish . . . . . . . . . . . . . . . . 3284 Caesio cuning . . . . . . . . . . . . . . . . 2927-2928
Bluespot butterflyfish . . . . . . . . . . . . . . . . 3247 Caesio erythrogaster . . . . . . . . . . . . . . . 2927
Bluespotted angelfish . . . . . . . . . . . . . . . 3279 Caesio lunaris . . . . . . . . . . . . . . . . . . . 2928
Bluespotted large-eye bream . . . . . . . . . . . 3024 Caesio striata . . . . . . . . . . . . . . . . . . . 2926
Bluestripe angelfish . . . . . . . . . . . . . . . . 3282 Caesio teres . . . . . . . . . . . . . . . . . 2929-2930
Bluestriped chub . . . . . . . . . . . . . . . . . . 3296 Caesio varilineata . . . . . . . . . . . . . . . . . 2926
Boeseman croaker . . . . . . . . . . . . . . . . . 3136 Caesio ventre rouge . . . . . . . . . . . . . . . . 2927
Boesemania microlepis . . . . . . . . . . . . . . 3136 Caesio xanthonota . . . . . . . . . . . . . 2929-2930
bohar, Lutjanus . . . . . . . . . . . . . . . . . . 2866 CAESIONIDAE . . . . . . 2838, 2841, 2919, 3052, 3304
Bombache bronzeado . . . . . . . . . . . . . . . 3165 calcarifer, Lates . . . . . . . . . . . . . . . . . . 3105
Bombache tigre mayor . . . . . . . . . . . . . . . 3164 Calicagère bleue . . . . . . . . . . . . . . . . . . 3293
Bonin Island flagtail . . . . . . . . . . . . . . . . 3319 campbelli, Coius . . . . . . . . . . . . . . . . . . 2945
boninensis, Kuhlia . . . . . . . . . . . . . . . . 3319 cancellatus, Mesopristes . . . . . . . . . . . . . 3310
borneensis, Johnius (Johnieops) . . . . . . . . . 3154 cancellatus, Scolopsis . . . . . . . . . . . . . . . 3077
borneensis, Polynemus . . . . . . . . . . . . . . 3115 cancellatus, Terapon . . . . . . . . . . . . . . . 3310
Botellón velero . . . . . . . . . . . . . . . . . . . 3288 caninus, Pentapodus . . . . . . . . . . . . . . . 3084
Bottlenose jewfish . . . . . . . . . . . . . . . . . 3140 cantharoides, Paracaesio . . . . . . . . . . . . . 2904
Bourrugue chévre . . . . . . . . . . . . . . . . . 3138 Cantharus filamentosus . . . . . . . . . . . . . . 3069
boutton, Lutjanus . . . . . . . . . . . . . . 2867, 2890 Capucin de Vanicolo . . . . . . . . . . . . . . . . 3183
boylei, Centropyge . . . . . . . . . . . . . . . . . 3274 Capuciva à bande jaune . . . . . . . . . . . . . . 3181
Brama australis . . . . . . . . . . . . . . . . . . 2833 CARANGIDAE . . . . . . . . . 2793, 2825, 3210, 3217
Brama brama . . . . . . . . . . . . . 2824, 2831, 2833 See also Vol. 3
Brama dussumieri . . . . . . . . . . . . . . 2831-2832 carbunculus, Etelis . . . . . . . . . . . . . . . . 2857
Brama japonica . . . . . . . . . . . . . . . 2824, 2831 CARISTIIDAE . . . . . . . . . . . . . . . . 2825, 2837
Brama leucotaenia . . . . . . . . . . . . . . . . . 2831 carnolabrum, Lipocheilus . . . . . . . . . . . . 2860
Brama myersi . . . . . . . . . . . . . . . . 2831-2833 Caroun croaker . . . . . . . . . . . . . . . . . . . 3142
Brama orcini . . . . . . . . . . . . . . . . 2831-2832 carouna, Johnius . . . . . . . . . . . . . . 3140-3141
Brama pauciradiata . . . . . . . . . . . . . . . . 2834 carouna, Johnius (Johnius) . . . . . . . . . . . 3142
brama, Brama . . . . . . . . . . . . . 2824, 2831, 2833 carponotatus, Lutjanus . . . . . . . . . . . . . . 2868
BRAMIDAE . . . . . . . . . . . . . . . . . 2824, 2837 carutta, Johnius (Johnius) . . . . . . . . . . . . 3143
Brassy chub . . . . . . . . . . . . . . . . . . . . 3295 Castagnole mignonne . . . . . . . . . . . . . . . 2831
brevirostris, Leiognathus . . . . . . . . . . . . . 2805 Catemo manchado . . . . . . . . . . . . . . . . . 3223
brevis, Atrobucca . . . . . . . . . . . . . . . . . 3131 caudavittatus, Amniataba . . . . . . . . . . . . . 3308
brighami, Rooseveltia . . . . . . . . . . . . . . . 2915 caudavittatus, Amphitherapon . . . . . . . . . . 3308
Brilliant pomfret . . . . . . . . . . . . . . . . . . . 2834 caudavittatus, Terapon . . . . . . . . . . . . . . 3308
Broadbanded velvetchin . . . . . . . . . . . . . . 2968 Celebes threadfin bream . . . . . . . . . . . . . . 3064
Broadhead croaker . . . . . . . . . . . . . . . . . 3148 celebicus, Nemipterus . . . . . . . . . . . . . . . 3064
Bronze croaker . . . . . . . . . . . . . . . . . . . 3165 celebicus, Plectorhinchus . . . . . . . . . . . . . 2972
Brown chub . . . . . . . . . . . . . . . . . . . . . 3292 CENTROPOMIDAE . . . . . . . . . . . . . . . . 3118
Brownstripe red snapper . . . . . . . . . . . . . . 2896 See also Vol. 4
Centropyge . . . . . . . . . . . . . . . . . . 3266-3268 Chaetodon trichrous . . . . . . . . . . . . . . . 3253

Centropyge aurantius . . . . . . . . . . . . . . . 3273 Chaetodon trifascialis . . . . . . . . . . . . . . . 3253
Centropyge bicolor . . . . . . . . . . . . . . . . 3273 Chaetodon trifasciatus . . . . . . . . . . . . . . 3254
Centropyge bispinosus . . . . . . . . . . . . . . 3273 Chaetodon ulietensis . . . . . . . . . . . . . . . 3254
Centropyge boylei . . . . . . . . . . . . . . . . . 3274 Chaetodon unimaculatus . . . . . . . . . . . . . 3255
Centropyge colini . . . . . . . . . . . . . . . . . 3274 Chaetodon vagabundus . . . . . . . . . . . . . . 3255
Centropyge eibli . . . . . . . . . . . . 3274-3275, 3279 Chaetodon wiebeli . . . . . . . . . . . . . . . . . 3256
Centropyge fisheri . . . . . . . . . . . . . . . . . 3275 Chaetodon xanthurus . . . . . . . . . . . . . . . 3256
Centropyge flavicauda . . . . . . . . . . . . . . 3275 chaetodonoides, Plectorhinchus . . . . . . . . . 2971
Centropyge flavissimus . . . . . . . . 3274-3275, 3279 CHAETODONTIDAE . . . 3217, 3221, 3224, 3266, 3287,
Centropyge heraldi . . . . . . . . . . . . . . . . 3275 3289, 3302
Centropyge hotumatua . . . . . . . . . . . . . . 3276 Chaetodontoplus . . . . . . . . . . . . . . . . . 3269
Centropyge joueur de flûte . . . . . . . . . . . . . 3278 Chaetodontoplus caeruleopunctatus . . . . . . . 3279
Centropyge loriculus . . . . . . . . . . . . . . . 3276 Chaetodontoplus chrysocephalus . . . . . . . . 3280
Centropyge multicolor . . . . . . . . . . . . . . 3276 Chaetodontoplus conspicillatus . . . . . . . . . 3280
Centropyge multifasciatus . . . . . . . . . 3277, 3279 Chaetodontoplus duboulayi . . . . . . . . . . . 3280
Centropyge narcosis . . . . . . . . . . . . . . . 3277 Chaetodontoplus melanosoma . . . . . . . . . . 3281
Centropyge nigriocellus . . . . . . . . . . . . . . 3277 Chaetodontoplus meredithi . . . . . . . . . . . . 3281
Centropyge nox . . . . . . . . . . . . . . . . . . 3278 Chaetodontoplus mesoleucus . . . . . . . . . . . 3281
Centropyge shepardi . . . . . . . . . . . . . . . 3278 Chaetodontoplus septentrionalis . . . . . . 3280, 3282
Centropyge tibicen . . . . . . . . . . . . . . . . 3278 chatareus, Toxotes . . . . . . . . . . . . . . . . 3215
Centropyge tout jaune .
. . . . . . . . . . . . . . 3275 Checkered snapper . . . . . . . . . . . . . . . . 2869
Centropyge venustus . . . . . . . . . . . . . . . 3279 CHEILODACTYLIDAE . . . . . . . . . . . . 2991, 3329
Centropyge vroliki . . .
. . . . . . . . 3274-3275, 3279 Cheilodactylus . . . . . . . . . . . . . . . . . . . 3329
CEPOLIDAE . . . . . . . . . . . . . . . . . . . . 3331 Cheilodactylus ephippium . . . . . . . . . . . . 3329
Chaetodon . . . . . . .
. . . . . . . . . . . 3224, 3226 Cheilodactylus fasciatus . . . . . . . . . . . . . 3329
Chaetodon adiergastos . . . . . . . . . . . . . . 3234 Cheilodactylus fuscus . . . . . . . . . . . . . . . 3329
Chaetodon argentatus . . . . . . . . . . . . . . 3235 Cheilodactylus nigripes . . . . . . . . . . . . . . 3329
Chaetodon aureofasciatus . . . . . . . . . . . . 3235 Cheilodactylus pixi . . . . . . . . . . . . . . . . 3329
Chaetodon auriga . . . . . . . . . . . . . . . . . 3236 Chelmon . . . . . . . . . . . . . . . . . . . . . . 3230
Chaetodon auripes . . . . . . . . . . . . . . . . 3236 Chelmon à long bec . . . . . . . . . . . . . . . . 3260
Chaetodon baronessa . . . . . . . . . . . . . . . 3237 Chelmon à très long bec . . . . . . . . . . . . . . 3260
Chaetodon bennetti . . . . . . . . . . . . . . . . 3237 Chelmon marginalis . . . . . . . . . . . . . . . 3257
Chaetodon citrinellus . . . . . . . . . . . . . . . 3238 Chelmon muelleri . . . . . . . . . . . . . . . . . 3257
Chaetodon collare . . . . . . . . . . . . . . . . . 3238 Chelmon rostratus . . . . . . . . . . . . . . . . 3258
Chaetodon declivis . . . . . . . . . . . . . . . . 3239 Chétodon à bandes orange . . . . . . . . . . . . 3246
Chaetodon decussatus . . . . . . . . . . . . . . 3239 Chétodon à collier . . . . . . . . . . . . . . . . . 3238
Chaetodon ephippium . . . . . . . . . . . . . . . 3240 Chétodon à croissant . . . . . . . . . . . . . . . 3243
Chaetodon falcula . . . . . . . . . . . . . . . . . 3254 Chétodon à deux taches . . . . . . . . . . . . . . 3247
Chaetodon flavirostris . . . . . . . . . . . . . . 3240 Chétodon à dos noir . . . . . . . . . . . . . . . . 3243
Chaetodon guentheri . . . . . . . . . . . . . . . 3241 Chétodon à miroir . . . . . . . . . . . . . . . . . 3252
Chaetodon guttatissimus . . . . . . . . . . . . . 3241 Chétodon à nageoire rouge orange . . . . . . . . 3254
Chaetodon kleinii . . . . . . . . . . . . . . . . . 3242 Chétodon à nez jaune . . . . . . . . . . . . . . . 3240
Chaetodon lineolatus . . . . . . . . . . . . . . . 3242 Chétodon à selle noire . . . . . . . . . . . . . . . 3240
Chaetodon lunula . . . . . . . . . . . . . . . . . 3243 Chétodon à vagabond . . . . . . . . . . . . . . . 3255
Chaetodon melannotus . . . . . . . . . . . . . . 3243 Chétodon armé à monocle . . . . . . . . . . . . . 3280
Chaetodon mertensii . . . . . . . . . . . . . . . 3244 Chétodon baronne . . . . . . . . . . . . . . . . . 3237
Chaetodon meyeri . . . . . . . . . . . . . . . . . 3244 Chétodon citron . . . . . . . . . . . . . . . . . . 3238
Chaetodon nippon . . . . . . . . . . . . . . . . . 3245 Chétodon cocher . . . . . . . . . . . . . . . . . . 3236
Chaetodon ocellicaudus . . . . . . . . . . . . . 3245 Chétodon d’Uliétéa . . . . . . . . . . . . . . . . . 3254
Chaetodon octofasciatus . . . . . . . . . . . . . 3246 Chétodon de Bennett . . . . . . . . . . . . . . . 3237
Chaetodon ornatissimus . . . . . . . . . . 3244, 3246 Chétodon de Klein . . . . . . . . . . . . . . . . . 3242
Chaetodon pelewensis . . . . . . . . . . . . 3247-3248 Chétodon de Mertens . . . . . . . . . . . . . . . 3244
Chaetodon plebeius . . . . . . . . . . . . . . . . 3247 Chétodon de Meyer . . . . . . . . . . . . . . . . 3244
Chaetodon punctatofasciatus . . . . . . . . 3247-3248 Chétodon de Palau . . . . . . . . . . . . . . . . . 3247
Chaetodon quadrimaculatus . . . . . . . . . . . 3248 Chétodon maillé . . . . . . . . . . . . . . . . . . 3250
Chaetodon rafflesii . . . . . . . . . . . . . . . . 3249 Chétodon moucheté . . . . . . . . . . . . . . . . 3241
Chaetodon rainfordi . . . . . . . . . . . . . . . 3249 Chétodon ocellé . . . . . . . . . . . . . . . . . . 3255
Chaetodon reticulatus . . . . . . . . . . . . 3244, 3250 Chétodon strié . . . . . . . . . . . . . . . . . . . 3242
Chaetodon selene . . . . . . . . . . . . . . . . . 3250 Chétodon tête de flèche . . . . . . . . . . . . . . 3253
Chaetodon semion . . . . . . . . . . . . . . . . 3251 Chevroned butterflyfish . . . . . . . . . . . . . . 3253
Chaetodon smithi . . . . . . . . . . . . . . . . . 3251 Chinamanfish . . . . . . . . . . . . . . . . . . . . 2918
Chaetodon speculum . . . . . . . . . . . . . . . 3252 choerorynchus, Lethrinus . . . . . . . . . . . . . 3037
Chaetodon tinkeri . . . . . . . . . . . . . . . . . 3252 Chopa axul . . . . . . . . . . . . . . . . . . . . . 3293
Chromis . . . . . . . . . . . . . . . . . . . . . . 3338 Colin’s angelfish . . . . . . . . . . . . . . . . . . 3274

Chromis viridis . . . . . . . . . . . . . . . . . . 3352 colini, Centropyge . . . . . . . . . . . . . . . . . 3274
chryserydros, Parupeneus . . . . . . . . . . . . . 3188 Collare butterflyfish . . . . . . . . . . . . . . . . . 3238
Chrysiptera cyanea . . . . . . . . . . . . . . . . 3353 collare, Chaetodon . . . . . . . . . . . . . . . . 3238
Chrysiptera hemicyanea . . . . . . . . . . . . . 3353 Collared large-eye bream . . . . . . . . . . . . . 3018
Chrysiptera parasema . . . . . . . . . . . . . . . 3353 Collybus drachme . . . . . . . . . . . . . . . . . 2832
Chrysiptera sinclairi . . . . . . . . . . . . . . . 3353 Common bluestripe snapper . . . . . . . . . . . . 2879
Chrysiptera springeri . . . . . . . . . . . . . . . 3353 Common ponyfish . . . . . . . . . . . . . . . . . 2808
Chrysiptera taupou . . . . . . . . . . . . . . . . 3353 Common silverbiddy . . . . . . . . . . . . . . . . 2956
chrysocephalus, Chaetodontoplus . . . . . . . . 3280 Common threadfin . . . . . . . . . . . . . . . . . 3110
Chrysochir aureus . . . . . . . . . . . . . . . . 3137 conchyliatus, Lethrinus . . . . . . . . . . . . . . 3028
Chrysophrys auratus . . . . . . . . . . . . . . . 3002 conspicillatus, Chaetodontoplus . . . . . . . . . 3280
Chrysophrys major . . . . . . . . . . . . . . . . 3002 Conspicuous angelfish . . .. . . . . . . . . . . . 3280
chrysopterus, Amphiprion . . . . . . . . . . . . 3347 Copperbanded butterflyfish . . . . . . . . . . . . 3258
chrysostomus, Heniochus . . . . . . . . . . . . . 3262 Coradion . . . . . . . . . .
. . . . . . . . . . . . 3231
chrysostomus, Lethrinus . . . . . . . . . . . . . . 3036 Coradion à grande voile . .. . . . . . . . . . . . 3258
chrysotaenia, Plectorhinchus . . . . . . . . . . . 2972 Coradion altivelis . . . . .
. . . . . . . . . . . . 3258
chrysozona, Pterocaesio . . . . . . . . . . 2933, 2935 Coradion chrysozonus . . . . . . . . . . . . . . 3259
chrysozonus, Coradion . . . . . . . . . . . . . . 3259 Coradion melanopus . . . . . . . . . . . . . . . 3259
Cichla ocellaris . . . . . . . . . . . . . . . . . . 3333 Coral beauty . . . . . . . .
. . . . . . . . . . . . 3273
CICHLIDAE . . . . . . . . . . . . . . . . . 3333, 3338 Coral breams . . . . . . . .
. . . . . . . . . . . . 3051
Cichlids . . . . . . . . .
. . . . . . . . . . . 3333-3334 cornutus, Zanclus . . . . .. . . . . . . . . . . . 3225
ciliatus, Parupeneus . . . . . . . . . . . . . . . 3187 Corocoro jabalina . . . . .. . . . . . . . . . . . 2987
ciliatus, Scolopsis . . .
. . . . . . . . . . . . . . 3087 Corocoro mejilla azul . . . .
. . . . . . . . . . . . 2984
cinctus, Plectorhinchus . . . . . . . . . . . . . . 2978 Corocoro montura . . . . . . . . . . . . . . . . . 2988
cinerascens, Kyphosus . . . . . . . . . . . . . . 3293 Corocoro plateado . . . . .. . . . . . . . . . . . 2983
Cinnabar goatfish . . . . . . . . . . . . . . . . . 3189 coruscans, Etelis . . . . .. . . . . . . . . . . . 2858
cinnabarinus, Parupeneus . . . . . . . . . . . . . 3189 Corvina aletona . . . . . .. . . . . . . . . . . . 3149
Cirrhites alternatus . . . . . . . . . . . . . . . . 3326 Corvina barba . . . . . . .. . . . . . . . . . . . 3139
Cirrhites pinnulatus . . . . . . . . . . . . . . . . 3326 Corvina carota . . . . . . .
. . . . . . . . . . . . 3143
Cirrhitichthys . . . . . . . . . . . . . . . . . . . 3323 Corvina caruna . . . . . . .
. . . . . . . . . . . . 3142
CIRRHITIDAE . .
. .
. . . . . . . . . 2991, 3321, 3330 Corvina de Belanger . . . .. . . . . . . . . . . . 3141
Cirrhitids . . . . .
. .
. . . . . . . . . . . . . . . 3321 Corvina dientusa . . . . . .
. . . . . . . . . . . . 3154
CIRRHITOIDEA .
. .
. . . . . . . . . . . . . . . 3330 Corvina dorado . . . . . . .
. . . . . . . . . . . . 3137
Cirrhitus . . . . .
. .
. . . . . . . . . . . . . . . 3324 Corvina hocicona . . . . . .
. . . . . . . . . . . . 3150
Cirrhitus alternatus . . . . . . . . . . . . . . . . 3326 Corvina pintade . . . . . .. . . . . . . . . . . . 3171
Cirrhitus spilotoceps . . . . . . . . . . . . . . . 3326 Corvina ragana . . . . . . .
. . . . . . . . . . . . 3166
citrinellus, Chaetodon . . . . . . . . . . . . . . 3238 Corvina soldado . . . . . .. . . . . . . . . . . . 3162
Clown anemonefish . . . . . . . . . . . . . . . . 3350 Corvinata prieta . . . . . .
. . . . . . . . . . . . 3129
CLUPEIDAE . . . . . . . . . . . . . . . . . . . . 3211 Cotton silverbiddy . . . . .
. . . . . . . . . . . . 2953
See also Vol. 3 Courbine à grande nageoire . . . . . . . . . . . . 3149
Cocoa snapper . . . . . . . . . . . . . . . . . . . 2903 Courbine barbiche . . . . .. . . . . . . . . . . . 3139
coelestis, Pomacentrus . . . . . . . . . . . . . . 3355 Courbine caroune . . . . . . . . . . . . . . . . . 3142
Cohana élégante . . . . . . . . . . . . . . . . . . 3081 Courbine carutte . . . . . .
. . . . . . . . . . . . 3143
Cohana japonaise . . . . . . . . . . . . . . . . . 3068 Courbine de Belanger . . . . . . . . . . . . . . . 3141
Cohana tolu . . . . . . . . . . . . . . . . . . . . 3071 Courbine dentue . . . . . .. . . . . . . . . . . . 3154
COIIDAE . . . . . . . . . . . . . . . . . . . . . . 2942 Courbine dorée . . . . . . .
. . . . . . . . . . . . 3137
Coitor croaker . . . . . . . . . . . . . . . . . . . 3144 Courbine longnez . . . . . . . . . . . . . . . . . 3150
coitor, Johnius . . . . . . . . . . . . . . . 3147, 3151 Courbine pintade . . . . . .
. . . . . . . . . . . . 3171
coitor, Johnius (Johnius) . . . . . . . . . . . . . 3144 Courbine ragane . . . . . .. . . . . . . . . . . . 3166
Coius . . . . . . . . . . . . . . . . . . 2942-2943, 2945 Courbine sodat . . . . . . .
. . . . . . . . . . . . 3162
Coius campbelli . . . . . . . . . . . . . . . . . . 2945 crassilabris, Parupeneus . . . . . . . . . . . . . 3186
Coius microlepis . . . . . . . . . . . . . . . . . . 2942 Crimson jobfish . . . . . . .
. . . . . . . . . . . . 2910
Coius quadrifasciatus . . . . . . . . . . . . . . . 2945 Crimson seabream . . . . . . . . . . . . . . . . . 3001
Coius undecimradiatus . . . . . . . . . . . . . . 2942 Crimson snapper . . . . . .. . . . . . . . . . . . 2872
Colas à bandes dorées . . . . . . . . . . . . . . 2912 Croakers . . . . . . . . . .
. . . . . . . . . . . . 3117
Colas bagnard . . . . . . . . . . . . . . . . . . . 2915 Crocro drapeau . . . . . . . . . . . . . . . . . . 3319
Colas dentu . . . . . . . . . . . . . . . . . . . . 2914 Crocro sauvage . . . . . . . . . . . . . . . . . . 3320
Colas drapeau . . . . . . . . . . . . . . . . . . . 2909 Croupia roche . . . . . . .. . . . . . . . . . . . 2944
Colas écailleuse . . . . . . . . . . . . . . . . . . 2905 cuning, Caesio . . . . . . .
. . . . . . . . . 2927-2928
Colas fil . . . . . . . . . . . . . . . . . . . . . . . 2910 cyanea, Chrysiptera . . . .. . . . . . . . . . . . 3353
Colas lavande . . . . . . . . . . . . . . . . . . . 2913 cyclostomus, Parupeneus . . . . . . . . . . . . . 3188
Colas oeil doré . . . . . . . . . . . . . . . . . . . 2911 Cyprinocirrhitus polyactis . . . . . . . . . . . . 3321
Colas orné . . . . . . . . . . . . . . . . . . . . . 2908
D Drepane longimana . . . . . . . . . . . . . 3222-3223

Drepane punctata . . . . . . . . . . . . . . 3222-3223
DACTYLOPTERIDAE . . . . . . . . . . . . . . . 3091 DREPANIDAE . . . . . . . . . . . . . 3216, 3221, 3289
See also Vol. 4 Drums . . . . . . . . . . . . . . . . . . . . . . . 3117
Damselfishes . . . . .
. . . . .
. . . . . . . 3334, 3337 dubius, Polynemus . . . . . . . . . . . . . . . . 3113
Dara bandera . . . . .
. . . . .
. . . . . . . . . . 3319 duboulayi, Chaetodontoplus . . . . . . . . . . 3280
Dara campestre . . .. . . . .
. . . . . . . . . . 3320 Dussumier’s ponyfish . . . . . . . . . . . . . . . 2806
Darkbanded fusilier . .
. . . . .
. . . . . . . . . . 2940 dussumieri, Brama . . . . . . . . . . . . . 2831-2832
Dascyllus aruanus . . . . . .
. . . . . . . . . . 3354 dussumieri, Johnius . . . . . . . . . . . . . . . . 3139
Dascyllus melanurus . . . . .
. . . . . . . . . . 3354 dussumieri, Leiognathus . . . . . . . . . . . . . 2806
Dascyllus trimaculatus . . . .
. . . . . . . . . . 3354 Dwarf monocle breams . . . . . . . . . . . 3051-3052
Dash-and-dot goatfish . . . . .
. . . . . . . . . . 3185 Dwarf paradise fish . . . . . . . . . . . . . . . . . 3103
Datnioides . . . . . . . . . . .
. . . . . . . . . . 2942 Dwarf threadfin bream . . . . . . . . . . . . . . . 3078
DATNIOIDIDAE . . . . . . . .
. . . . . . . . . . 2942
daura, Leiognathus . . . . . .
. . . . . . . . . . 2804
Dawn threadfin bream . . . . .
. . . . . . . . . . 3062 E
decacanthus, Gerres . . . . .
. . . . . . . . . . 2959 Eastern paradise fish .. . . . . . . . . . . . . . . 3113
declivis, Chaetodon . . . . . .
. . . . . . . . . . 3239 Eastern pomfred . . .. . . . . . . . . . . . . . . 3220
decorus, Leiognathus . . . . .
. . . . 2802, 2805, 2815 Edelia . . . . . . . .. . . . . . . . . . . . . . . 3317
decussatus, Chaetodon . . . .
. . . . . . . . . . 3239 Ehrenberg’s snapper .. . . . . . . . . . . . . . . 2871
decussatus, Lutjanus . . . . .
. . . . . . . . . . 2869 ehrenbergii, Lutjanus . . . . . . . . . . . . . . . 2871
Deep pugnose ponyfish . . . .
. . . . . . . . . . 2822 Eibl’s angelfish . . . . . . . . . . . . . . . . . . . 3274
Deepbody silverbiddy . . . . .
. . . . . . . . . . 2950 eibli, Centropyge . . . . . . . . . . . 3274-3275, 3279
delagoae, Nemipterus . . . . .
. . . . . . . . . . 3062 Eightbanded butterflyfish . . . . . . . . . . . . . . 3246
Dendrophysa russelli . . . . .
. . . . . . . 3138-3139 Eightfinger threadfin . . . . . . . . . . . . . . . . 3101
Dentex fourmanoiri . . . . . .
. . . . . . . . . . 2999 Elegant paradiseus fish . . . . . . . . . . . . . . 3116
Dentex spariformis . . . . . .
. . . . . . . . . . 3000 Eleutheronema tetradactylum . . . . 3096-3097, 3105
Dentex tumifrons . . . . . . .
. . . . . . . . . . 3000 Eleutheronema tridactylum . . . . . . . . 3096-3097
diacantha, Protonibea . . . .
. . . . . . . . . . 3171 elevata, Girella . . . . . . . . . . . . . . . . . . 3298
diacanthus, Nibea . . . . . . .
. . . . . . . . . . 3171 Elongate ponyfish . . . . . . . . . . . . . . . . . 2807
diacanthus, Pygoplites . . . .
. . . . . . . . . . 3286 elongatus, Gymnocranius . . . . . . . . . . . . 3019
Diagramma labiosum . . . . .
. . . . . . . . . . 2966 elongatus, Leiognathus . . . . . . . . . . . 2807, 2816
Diagramma pictum . . . . . .
. . . . . . . 2966-2967 elongatus, Lethrinus . . . . . . . . . . . . 3035, 3039
Diagramma punctatum . . . .
. . . . . . . . . . 2967 emeryii, Pentapodus . . . . . . . . . . . . . . . 3084
Diagramme arlequin . . . . . .
. . . . . . . . . . 2971 EMMELICHTHYIDAE . . . . . . . . . . . . 2838, 2920
Diagramme citron . . . . . . .
. . . . . . . . . . 2973 Emmelichthys karnellai . . . . . . . . . . . . . 2838
Diagramme ménestrel . . . . .
. . . . . . . . . . 2981 Emmelichthys strusakeri . . . . . . . . . . . . . 2838
Diagramme noire . . . . . . . .
. . . . . . . . . . 2974 Emperador boca de vino . . . . . . . . . . . . . . 3028
Diagramme oriental . . . . . .
. . . . . . . . . . 2982 Emperador boquidulce . . . . . . . . . . . . . . . 3035
Diagramme rubanne . . . . . .
. . . . . . . . . . 2980 Emperador de bandes . . . . . . . . . . . . . . . 3038
Diagramme truité . . . . . . . .
. . . . . . . . . . 2978 Emperador de lentejuelas . . . . . . . . . . . . . 3034
Diagramme voilier . . . . . . .
. . . . . . . . . . 2967 Emperador de manchas . . . . . . . . . . . . . . 3029
diagrammus, Plectorhinchus . . . . . . . . . . 2975
. Emperador estriado . . . . . . . . . . . . . . . . 3017
Diamondscaled goatfish . . . . . . . . . . . . . 3187
. Emperador gris . . . . . . . . . . . . . . . . . . . 3023
digramma, Pterocaesio . . . . . . . . . . 2934, 2936
. Emperador malla . . . . . . . . . . . . . . . . . . 3043
Dipterygonotus . . . . . . . . . . . . . . . . . 2919
. Emperador maquillado . . . . . . . . . . . . . . . 3042
Dipterygonotus balteatus . . . 2838, 2919-2920, 2931
. Emperador moteado . . . . . . . . . . . . . . . . 3044
Dipterygonotus leucogrammicus . . . . . . . . . 2931 Emperador ornado . . . . . . . . . . . . . . . . . 3040
DIRETMIDAE . . . . . . . . . . . . . . . . 2825, 2837 Emperador relámpago . . . . . . . . . . . . . . . 3037
See also Vol. 4 Emperador San Pedro . . . . . . . . . . . . . . . 3032
Dirty ordure snapper . . . . . . . . . . . . . . . . 2902 Emperador trompa amarilla . . . . . . . . . . . . 3045
dodecacanthoides, Lutjanus . . . . . . . . . . . 2870 Emperador trompudo . . . . . . . . . . . . . . . 3039
Donkey croaker . . . . . . . . . . . . . . . . . . 3168 Empereur bandes oranges . . . . . . . . . . . . . 3038
doriae, Upeneus . . . . . . . . . . . . . . . . . . 3197 Empereur bas cou . . . . . . . . . . . . . . . . . 3044
Dormilona . . . . . . . . . . . . . . . . . . . . . 2944 Empereur bec de cane . . . . . . . . . . . . . . . 3045
Dot-and-dash butterflyfish . . . . . . . . . . . . . 3247 Empereur békine . . . . . . . . . . . . . . . . . . 3030
Dots-and-dashes ponyfish . . . . . . . . . . . . . 2819 Empereur capitaine . . . . . . . . . . . . . . . . 3029
Dotted butterflyfish . . . . . . . . . . . . . . . . . 3251 Empereur gris . . . . . . . . . . . . . . . . . . . 3023
Double whiptail . . . . . . . . . . . . . . . . . . . 3084 Empereur guele de vin . . . . . . . . . . . . . . . 3028
Doublebar goatfish . . . . . . . . . . . . . . . . . 3186 Empereur guele longue . . . . . . . . . . . . . . 3039
Doublelined fusilier . . . . . . . . . . . . . . . . . 2934 Empereur honteux . . . . . . . . . . . . . . . . . 3042
Doublewhip threadfin bream . . . . . . . . . . . . 3069 Empereur lentilles . . . . . . . . . . . . . . . . . 3034
Drab emperor . . . . . . . . . . . . . . . . . . . 3047 Empereur maill . . . . . . . . . . . . . . . . . . . 3043
drachme, Collybus . . . . . . . . . . . . . . . . . 2832 Empereur moris . . . . . . . . . . . . . . . . . . 3037
Empereur orn . . . . . . . . . . . . . . . . . . . . 3040 Filimanus perplexa . . . . . . . . . . . . . . . . 3100

Empereur Saint Pierre . . . . . . . . . . . . . . . 3032 Filimanus sealei . . . . . . . . . . . . . . . . . . 3101
Empereur strié . . . . . . . . . . . . . . . . . . . 3017 Filimanus similis . . . . . . . . . . . . . . 3098, 3102
Empereur tidents . . . . . . . . . . . . . . . . . . 3035 Filimanus xanthonema . . . . . . . . . . . 3102, 3111
Emperor angelfish . . . . . . . . . . . . . . . . . 3285 fisheri, Centropyge . . . . . . . . . . . . . . . . 3275
Emperor red snapper . . . . . . . . . . . . . . . 2892 Fivelined snapper . . . . . . . . . . . . . . . . . 2888
Emperor snappers . . . . . . . . . . . . . . . . . 3004 Fivelined threadfin bream . . . . . . . . . . . . . 3072
Emperors . . . . . . . . . . . . . . . . . . . . . . 3004 Flagtailed terapon . . . . . . . . . . . . . . . . . 3308
ENOPLOSIDAE . . . . . . . . . . . . . . . . . . 3287 Flagtails . . . . . . . . . . . . . . . . . . . . . . 3317
Enoplosus armatus . . . . . . . . . . . . . . . . 3287 Flame snapper . . . . . . . . . . . . . . . . . . . 2858
Epervier à bande noire . . . . . . . . . . . . . . . 3327 Flathead pomfret . . . . . . . . . . . . . . . . . . 2835
Epervier de corail . . . . . . . . . . . . . . . . . . 3326 flavicauda, Centropyge . . . . . . . . . . . . . . 3275
EPHIPPIDAE . . . . . . . . . . 3216, 3221, 3267, 3289 flavipinnis, Pristipomoides . . . . . . . . . . . . 2911
See also Vol. 6 flavirostris, Chaetodon . . . . . . . . . . . . . . 3240
ephippium, Amphiprion . . . . . . . . . . . . . 3347 flavissimus, Centropyge . . . . . . . . 3274-3275, 3279
ephippium, Chaetodon . . . . . . . . . . . . . . 3240 flavissimus, Forcipiger . . . . . . . . . . . . . . 3260
ephippium, Cheilodactylus . . . . . . . . . . . . 3329 flavolineatus, Mulloidichthys . . . . . . . 3181, 3183
Equula nuchalis . . . . . . . . . . . . . . . . . . 2805 flavomaculatus, Plectorhinchus . . . . . . . . . 2973
Equulites hastatus . . . . . . . . . . . . . . . . . 2812 fletus, Lethrinus . . . . . . . . . . . . . . . . . . 3033
Equulites novaehollandiae . . . . . . . . . . . . 2810 fluminis, Sonorolux . . . . . . . . . . . . . . . . 3174
equulus, Leiognathus . . . . . . . . . . . . 2806, 2808 Forcepsfish . . . . . . . . . . . . . . . . . . . . . 3260
eriomma, Parascolopsis . . . . . . . . . . . . . . 3081 Forcipiger . . . . . . . . . . . . . . . . . . 3224, 3231
erythracanthus, Lethrinus . . . . . . . . . . . . 3029 Forcipiger flavissimus . . . . . . . . . . . . . . 3260
erythrinus, Mulloides . . . . . . . . . . . . . . . 3183 Forcipiger longirostris . . . . . . . . . . . . . . 3260
Erythrocles schlegeli . . . . . . . . . . . . . . . 2838 Forgeron tacheté . . . . . . . . . . . . . . . . . . 3223
Erythrocles scintillans . . . . . . . . . . . . . . 2838 Forktail large-eye bream . . . . . . . . . . . . . . 3019
Erythrocles taeniatus . . . . . . . . . . . . . . . 2838 Forktailed threadfin bream . . . . . . . . . . . . . 3065
erythrogaster, Caesio . . . . . . . . . . . . . . . 2927 forsteri, Paracirrhites . . . . . . . . . . . . . . . 3327
erythropterus, Lethrinus . . . . . . . . . . . . . 3030 Fourfinger threadfin . . . . . . . . . . . . . . . . 3096
erythropterus, Lutjanus . . . . . . . . . . . 2872, 2884 Fourlined terapon . . . . . . . . . . . . . . . . . 3311
Estuary croaker . . . . . . . . . . . . . . . . . . 3174 Fourmanoir’s seabream . . . . . . . . . . . . . . 2999
Etelis . . . . . . . . . . . . . . . 2843, 2857-2859, 2916 fourmanoiri, Dentex . . . . . . . . . . . . . . . 2999
Etelis carbunculus . . . . . . . . . . . . . . . . 2857 Fourspot butterflyfish . . . . . . . . . . . . . . . . 3248
Etelis coruscans . . . . . . . . . . . . . . . . . . 2858 fraenatus, Lethrinus . . . . . . . . . . . . 3033, 3037
Etelis filamentosus . . . . . . . . . . . . . . . . . 2916 fraterculus, Parupeneus . . . . . . . . . . 3187, 3194
Etelis marshi . . . . . . . . . . . . . . . . . . . . 2857 Freckled goatfish . . . . . . . . . . . . . . . . . . 3199
Etelis nudimaxillaris . . . . . . . . . . . . . . . . 2916 frenatus, Amphiprion . . . . . . . . . . . . . . . 3348
Etelis radiosus . . . . . . . . . . . . . . . . . . . 2859 frenatus, Gymnocranius . . . . . . . . . . . . . 3021
Etroplus . . . . . . . . . . . . . . . . . . . . . . 3333 Freshwater snapper . . . . . . . . . . . . . . . . 2875
Etroplus suratensis . . . . . . . . . . . . . . . . 3333 fulviflamma, Lutjanus . . . . . . . . . . . . . . 2873
euanus, Gymnocranius . . . . . . . . . . . . . . 3020 fulvus, Lutjanus . . . . . . . . . . . . . . . . . . 2874
Eumegistus . . . . . . . . . . . . . . . . . . . . 2824 furca, Aphareus . . . . . . . . . . . . . . . . . . 2854
Eumegistus illustris . . . . . . . . . . . . . . . . 2834 furcatum, Pomadasys . . . . . . . . . . . . . . . 2986
Eutherapon theraps . . . . . . . . . . . . . . . . 3316 furcatus, Aphareus . . . . . . . . . . . . . . . . . 2854
Evynnis japonica . . . . . . . . . . . . . . . . . 3001 furcatus, Rhonciscus . . . . . . . . . . . . . . . . 2986
furcosus, Nemipterus . . . . . . . . . . . . 3065, 3071
F fuscescens, Lutjanus . . .
fuscus, Cheilodactylus . . .
.
.
.
.
.
.
.
.
.
.
.
.
. . . . . . 2875
. . . . . . 3329
falcula, Chaetodon . . . . . . . . . . . . . . . . 3254 fuscus, Kyphosus . . . . . . . . . . . . . . . . . . 3292
False clown anemonefish . . . . . . . . . . . . . 3350 Fusilero aceitunero . . . . . . . . . . . . . . . . . 2940
fangi, Plectorhinchus . . . . . . . . . . . . . . . 2978 Fusilero amarillo azulado . . . . . . . . . . . . . 2929
fasciatus, Cheilodactylus . . . . . . . . . . . . . 3329 Fusilero azur . . . . . . . . . . . . . . . . . . . . 2926
fasciatus, Johnius . . . . . . . . . . . . . . . . . 3141 Fusilero banana . . . . . . . . . . . . . . . . . . 2937
fasciatus, Leiognathus . . . . . . . . . . . . . . 2809 Fusilero cinta dorada . . . . . . . . . . . . . . . . 2933
fauvelii, Glaucosoma . . . . . . . . . . . . . . . 3207 Fusilero de banda ancha . . . . . . . . . . . . . . 2935
Fiji threadfin bream . . . . . . . . . . . . . . . . . 3081 Fusilero de dos bandas . . . . . . . . . . . . . . 2934
filamentosus, Cantharus . . . . . . . . . . . . . . 3069 Fusilero de lomo amarillo . . . . . . . . . . . . . 2930
filamentosus, Etelis . . . . . . . . . . . . . . . . 2916 Fusilero de Marr . . . . . . . . . . . . . . . . . . 2936
filamentosus, Gerres . . . . . . . . . . . . . . . 2952 Fusilero de Randall . . . . . . . . . . . . . . . . 2938
filamentosus, Nemipterus . . . . . . . . . . . . . 3069 Fusilero de tres bandas . . . . . . . . . . . . . . 2941
filamentosus, Pristipomoides . . . . . . . . . . . 2910 Fusilero de una bande . . . . . . . . . . . . . . . 2939
filamentosus, Randallichthys . . . . . . . . . . . 2916 Fusilero elegante . . . . . . . . . . . . . . . . . . 2932
Filimanus heptadactyla . 3098, 3101-3102, 3108, 3112 Fusilero jaspeado . . . . . . . . . . . . . . . . . 2931
Filimanus hexanema . . . . . . . . . . . . . . . 3099 Fusilero luna . . . . . . . . . . . . . . . . . . . . 2928
Fusilero vientre colorado . . . . . . . . . . . . . . 2927 Glaucosoma hebraicum . . . . . . . . . . . . . 3207

Fusilier à bande large . . . . . . . . . . . . . . . 2935 Glaucosoma magnificum . . . . . . . . . . . . . 3208
Fusilier à dos jaune . . . . . . . . . . . . . . . . 2930 Glaucosoma scapulare . . . . . . . . . . . . . . 3209
Fusilier à dos jaune et bleu . . . . . . . . . . . . 2929 Glaucosoma taeniatus . . . . . . . . . . . . . . . 3207
Fusilier à trois bandes . . . . . . . . . . . . . . . 2941 GLAUCOSOMATIDAE . . . . . . . . . . . . . . 3205
Fusilier à une bande . . . . . . . . . . . . . . . . 2939 Gnathodentex . . . . . . . . . . . . . . . . . . . 2991
Fusilier banane . . . . . . . . . . . . . . . . . . . 2937 Gnathodentex aurolineatus . . . . . . . . . . . . 3017
Fusilier de Marr . . . . . . . . . . . . . . . . . . . 2936 Goatee croaker . . . . . . . . . . . . . . . . . . . 3138
Fusilier de Randall . . . . . . . . . . . . . . . . . 2938 Goatfishes . . . . . . . . . . . . . . . . . . . . . 3175
Fusilier deux bandes jaunes . . . . . . . . . . . . 2934 Goldband fusilier . . . . . . . . . . . . . . . . . . 2933
Fusilier élégant . . . . . . . . . . . . . . . . . . . 2932 Goldband goatfish . . . . . . . . . . . . . . . . . 3196
Fusilier ligne olive . . . . . . . . . . . . . . . . . 2940 Goldbanded jobfish . . . . . . . . . . . . . . . . 2912
Fusilier marbré . . . . . . . . . . . . . . . . . . . 2931 Goldbanded sweetlips . . . . . . . . . . . . . . . 2972
Fusiliers . . . . . . . . . . . . . . . . . . . . . . 2919 Golden angelfish . . . . . . . . . . . . . . . . . . 3273
Golden ponyfish . . . . . . . . . . . . . . . . . . 2800
G Golden threadfin bream . . .
Goldeneye jobfish . . . . . .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. . 3074
. . 2911
Gazza achlamys . . . . . . . . . . . . . . . . . . 2798 Goldenstriped butterflyfish . . . . . . . . . . . . . 3235
Gazza minuta . . . . . . . . . . . . . . . . . . . 2799 Goldflag jobfish . . . . . . . . . . . . . . . . . . . 2909
Gazza spp. . . . . . . . . . . . . . . . . . . 2792, 2800 Goldflake angelfish . . . . . . . . . . . . . . . . . 3272
Geelbeck croaker . . . . . . . . . . . . . . . . . 3129 goldiei, Lutjanus . . . . . . . . . . . . . . . . . 2877
Genicanthus . . . . . . . . . . . . . . . . . 3266, 3270 Goldlined seabream . . . . . . . . . . . . . . . . 3003
Genicanthus bellus . . . . . . . . . . . . . . . . 3282 goldmanni, Plectorhinchus . . . . . . . . . . . . 2976
Genicanthus lamarck . . . . . . . . . . . . . . . 3283 Goldsaddle goatfish . . . . . . . . . . . . . . . . 3188
Genicanthus melanospilos . . . . . . . . . . . . 3283 Goldstripe ponyfish . . . . . . . . . . . . . . . . . 2804
Genicanthus spinus . . . . . . . . . . . . . . . . 3284 Goldtail demoiselle . . . . . . . . . . . . . . . . . 3353
Genicanthus watanabei . . . . . . . . . . . . . . 3284 Goniistius . . . . . . . . . . . . . . . . . . . . . 3329
genivittatus, Lethrinus . . . . . . . . . . . . . . 3031 Goniistius vittatus . . . . . . . . . . . . . . . . . 3329
georgianus, Arripis . . . . . . . . . . . . . . . . 3304 gonzalesi, Paracaesio . . . . . . . . . . . . . . . 2899
GERREIDAE . . . . . . . . . . . . . . 2793, 2946, 3317 Goret à six bandes . . . . . . . . . . . . . . . . . 2986
Gerres . . . . . . . . . . . . . . . . . . . . . . . 2946 Graceful threadfin bream . . . . . . . . . . . . . 3079
Gerres abbreviatus . . . . . . . . . . . . . . . . 2950 gracilis, Bathyclupea . . . . . . . . . . . . . . . 3211
Gerres acinaces . . . . . . . . . . . . . . . . . . 2951 gracilis, Nemipterus . . . . . . . . . . . . . . . . 3079
Gerres argyreus . . . . . . . . . . . . . . . 2951, 2956 Grande verrue tigre . . . . . . . . . . . . . . . . 3164
Gerres baconensis . . . . . . . . . . . . . . . . . 2959 grandoculis, Gymnocranius . . . . . . . . 3022-3023
Gerres decacanthus . . . . . . . . . . . . . . . . 2959 grandoculis, Monotaxis . . . . . . . . . . . . . . 3049
Gerres filamentosus . . . . . . . . . . . . . . . . 2952 Grass emperor . . . . . .. . . . . . . . . . . . . 3033
Gerres gigas . . . . . . . . . . . . . . . . . . . . 2955 Green jobfish . . . . . . .
. . . . . . . . . . . . . 2856
Gerres infasciatus . . . . . . . . . . . . . . . . . 2952 Greenstriped coral bream . . . . . . . . . . . . . 3086
Gerres japonicus . . . . . . . . . . . . . . . . . 2959 GREGORYINIDAE . . . . . . . . . . . . . . . . . 3329
Gerres kapas . . . . . . . . . . . . . . . . . . . . 2953 Grey bigmouth bream . . . . . . . . . . . . . . . 3207
Gerres macracanthus . . . . . . . . . . . . 2952, 2960 Grey large-eye bream . . . . . . . . . . . . . . . 3023
Gerres macrosoma . . . . . . . . . . . . . . . . 2954 Grey sweetlips . . . . . .
. . . . . . . . . . . . . 2966
Gerres oblongus . . . . . . . . . . . . . . . . . . 2955 Griffis’ angelfish . . . . .
. . . . . . . . . . . . . 3272
Gerres oyena . . . . . . . . . . . . . . 2951, 2953, 2956 griffisi, Apolemichthys .. . . . . . . . . . . . . 3272
Gerres poeti . . . . . . . . . . . . . . . . . . . . 2957 griseus, Gymnocranius . . . . . . . . . . . . . . 3023
Gerres punctatus . . . . . . . . . . . . . . . . . . 2952 Grondeur à joues bleus . . . . . . . . . . . . . . 2984
Gerres subfasciatus . . . . . . . . . . . . . . . . 2960 Grondeur argenté . . . . . . . . . . . . . . . . . 2983
gerroides, Leiognathus . . . . . . . . . . . . . . 2804 Grondeur javelot . . . . .
. . . . . . . . . . . . . 2987
Giant sweetlips . . . . . . . . . . . . . . . . . . . 2970 Grondeur selle . . . . . .
. . . . . . . . . . . . . 2988
gibbosus, Plectorhinchus . . . . . . . . . . . . . 2974 Grunts . . . . . . . . . .
. . . . . . . . . . . . . 2961
gibbus, Lutjanus . . . . . . . . . . . . . . . . . 2876 guentheri, Chaetodon . . . . . . . . . . . . . . . 3241
gibsoni, Kyphosus . . . . . . . . . . . . . . . . . 3295 Günther’s butterflyfish . .
. . . . . . . . . . . . . 3241
giganteus, Plectorhinchus . . . . . . . . . . . . . 2970 guttatissimus, Chaetodon . . . . . . . . . . . . . 3241
gigas, Gerres . . . . . . . . . . . . . . . . . . . . 2955 Gymnocaesio gymnoptera . . . . . . . . . 2919, 2932
Girella elevata . . . . . . . . . . . . . . . . . . . 3298 Gymnocranius . . . . . . . . . . . . . . . . 2991, 3007
Girella punctata . . . . . . . . . . . . . . . . . . 3297 Gymnocranius audleyi . . . . . . . . . . . . . . 3018
Girella tricuspidata . . . . . . . . . . . . . . . . 3298 Gymnocranius bitorquatus . . . . . . . . . . . . 3018
GIRELLIDAE . . . . . . . . . . 3290, 3297, 3299, 3301 Gymnocranius elongatus . . . . . . . . . . . . . 3019
Girellids . . . . . . . . . . . . . . . . . . . . . . . 3297 Gymnocranius euanus . . . . . . . . . . . . . . 3020
Glabrilutjanus nematophorus . . . . . . . . . . . 2918 Gymnocranius frenatus . . . . . . . . . . . . . . 3021
Glaucosoma buergeri . . . . . . . . . . . . . . . 3207 Gymnocranius grandoculis . . . . . . . . . 3022-3023
Glaucosoma bürgeri . . . . . . . . . . . . . . . . 3207 Gymnocranius griseus . . . . . . . . . . . . . . 3023
Glaucosoma fauvelii . . . . . . . . . . . . . . . . 3207 Gymnocranius japonicus . . . . . . . . . . . . . 3020
Gymnocranius lethrinoides . . . . . . . . . . . . 3025 Humpback red snapper . . . . . . . . . . . . . . 2876

Gymnocranius microdon . . . . . . . . . . . . . 3024 Humphead bannerfish . . . . . . . . . . . . . . . 3264
Gymnocranius robinsoni . . . . . . . . . . . . . 3022 Humpnose bigeye bream . . . . . . . . . . . . . 3049
Gymnocranius sp. . . . . . . . . . . . . . . . . . 3025 hypostoma, Johnius (Johnius) . . . . . . . . . . 3146
gymnoptera, Gymnocaesio . . . . . . . . . 2919, 2932 hypselopterus, Lethrinus . . . . . . . . . . . . . 3030
H I
HAEMULIDAE . . . . . . 2841, 2942, 2961, 2991, 3005, illustris, Eumegistus . . . . . . . . . . . . . . . 2834
3052, 3317-3318 imperator, Pomacanthus . . . . . . . . . . . . . 3285
Halfmoons . . . . . . . . . . . . . . . . . . . . . 3299 Indian bannerfish . . . . . . . . . . . . . . . . . . 3263
Halfspotted hawkfish . . . . . . . . . . . . . . . . 3328 Indian goatfish . . . . . . . . . . . . . . . . . . . 3190
Haneda’s ponyfish . . . . . . . . . . . . . . . . . 2818 Indian snapper . . . . . . . . . . . . . . . . . . . 2883
hanedai, Secutor . . . . . . . . . . . . . . . . . 2818 Indian threadfin . . . . . . . . . . . . . . . . . . . 3104
Hapalogenys . . . . . . . . . . . . . . . . . . . . 2961 Indian vagabond butterflyfish . . . . . . . . . . . 3239
Hapalogenys analis . . . . . . . . . . . . . . . . 2968 indicius, Secutor . . . . . . . . . . . . . . . . . 2819
Hapalogenys kishinouyei . . . . . . . . . . . . . 2969 indicus, Parupeneus . . . . . . . . . . . . . . . 3190
Hapalogenys mucronatus . . . . . . . . . . . . . 2968 indicus, Polydactylus . . . . . . . . . . . . 3104-3105
harak, Lethrinus . . . . . . . . . . . . . . . . . 3032 Indonesian snapper . . . . . . . . . . . . . . . . 2865
Harlequin sweetlips . . . . . . . . . . . . . . . . 2971 inermis, Parascolopsis . . . . . . . . . . . . . . 3082
harrawayi, Plectorhinchus . . . . . . . . . . . . 2970 infasciatus, Gerres . . . . . . . . . . . . . . . . 2952
Harry hotlips . . . . . . . . . . . . . . . . . . . . 2974 insidiator, Leiognathus . . . . . . . . . . . . . . 2820
hasta, Pomadasys . . . . . . . . . . . . . . 2983, 2987 insidiator, Secutor . . . . . . . . . . . . . . 2818-2820
hastatus, Equulites . . . . . . . . . . . . . . . . . 2812 intermedius, Polynemus . . . . . . . . . . . . . . 3103
Hawkfish . . . . . . . . . . . . . . . . . . . . . . 3330 interruptus, Secutor . . . . . . . . . . . . . . . . 2822
Hawkfishes . . . . . . . . . . . . . . . . . . . . . 3321 isacanthus, Nemipterus . . . . . . . . . . . . . . 3067
hebraicum, Glaucosoma . . . . . . . . . . . . . 3207 itoi, Pempheris . . . . . . . . . . . . . . . . . . 3201
Helotes sexlineatus . . . . . . . . . . . . . . . . 3312
Hemichromis bimaculatus . . . . . . . . . . . . 3333
hemicyanea, Chrysiptera . . . . . . . . . . . . . 3353
J
hemistictus, Paracirrhites . . . . . . . . . . . . . 3328 jaculatrix, Toxotes . . . . . . . . . . . . . . . . 3215
Hemitaurichthys . . . . . . . . . . . . . . . 3224, 3231 jansenii, Parupeneus . . . . . . . . . . . . . . . 3189
Hemitaurichthys multispinosus . . . . . . . . . 3261 Japanese butterflyfish . . . .. . . . . . . . . . . 3245
Hemitaurichthys polylepis . . . . . . . . . . . . 3261 Japanese large-eye bream .. . . . . . . . . . . 3020
Hénioche bossu . . . . . . . . . . . . . . . . . . 3264 Japanese meagre . . . . . .. . . . . . . . . . . 3127
Hénioche cornu . . . . . . . . . . . . . . . . . . 3263 Japanese pygmy angelfish . .. . . . . . . . . . . 3276
Hénioche renversé . . . . . . . . . . . . . . . . . 3262 Japanese silverbiddy . . . . .. . . . . . . . . . . 2959
Hénioche singulier . . . . . . . . . . . . . . . . . 3263 Japanese threadfin bream . .. . . . . . . . . . . 3068
Heniochus . . . . . . . . . . . . 3224-3225, 3231, 3287 Japanese whiptail . . . . . .. . . . . . . . . . . 3084
Heniochus acuminatus . . . . . . . . . . . . . . 3262 japonica, Brama . . . . . . .. . . . . . . . 2824, 2831
Heniochus chrysostomus . . . . . . . . . . . . . 3262 japonica, Evynnis . . . . . .. . . . . . . . . . . 3001
Heniochus monoceros . . . . . . . . . . . . . . 3263 japonicus, Argyrosomus . .. . . . . . . . . . . 3127
Heniochus pleurotaenia . . . . . . . . . . . . . 3263 japonicus, Gerres . . . . . .. . . . . . . . . . . 2959
Heniochus singularis . . . . . . . . . . . . . . . 3264 japonicus, Gymnocranius . .. . . . . . . . . . . 3020
Heniochus varius . . . . . . . . . . . . . . . . . 3264 japonicus, Nemipterus . . .. . . . . . . . . . . 3068
heptacanthus, Parupeneus . . . . . . . . . . . . 3189 Japuta menuda . . . . . . . .. . . . . . . . . . . 2831
heptadactyla, Filimanus 3098, 3101-3102, 3108, 3112 jarbua, Terapon . . . . . . .. . . . . . . . . . . 3314
Herald’s angelfish . . . . . . . . . . . . . . . . . 3275 Jarbus terapon . . . . . . . .. . . . . . . . . . . 3314
heraldi, Centropyge . . . . . . . . . . . . . . . . 3275 Javanese threadfin . . . . . .. . . . . . . . . . . 3099
heterolepis, Johnius (Johnius) . . . . . . . . . . 3145 Javelin grunt . . . . . . . . .. . . . . . . . . . . 2987
hexanema, Filimanus . . . . . . . . . . . . . . . 3099 Jobfishes . . . . . . . . . . .. . . . . . . . . . . 2840
hexodon, Nemipterus . . . . . . . . . . . . . . . 3066 John’s snapper . . . . . . . .. . . . . . . . . . . 2878
Highfin coralfish . . . . . . . . . . . . . . . . . . 3258 Johnieops macrorhynus . . .. . . . . . . . . . . 3150
hilleri, Polynemus . . . . . . . . . . . . . . . . . 3115 johnii, Lutjanus . . . . . . .. . . . . . . . . . . 2878
Histiopterus spinifer . . . . . . . . . . . . . . . . 3288 Johnius . . . . . . . . . . .. . . . . . . . . . . 3118
Histiopterus typus . . . . . . . . . . . . . . . . . 3288 Johnius (Johnieops) . . . .. . . . . . . . . . . 3123
hololepidotus, Argyrosomus . . . . . 3127, 3129, 3131 Johnius (Johnieops) borneensis . . . . . . . . . 3154
Hornaday’s paradise fish . . . . . . . . . . . . . . 3114 Johnius (Johnieops) novaeguineae . . . . . . . 3155
hornadayi, Polynemus . . . . . . . . . . . . . . 3114 Johnius (Johnieops) pacificus . . . . . . . . . . 3156
Hotlips . . . . . . . . . . . . . . . . . . . . . . . 2961 Johnius (Johnieops) plagiostoma . . . . . . . . 3157
Hotumatua angelfish . . . . . . . . . . . . . . . . 3276 Johnius (Johnius) . . . . . . . . . . . . . . . . . 3122
hotumatua, Centropyge . . . . . . . . . . . . . . 3276 Johnius (Johnius) amblycephalus . . . . . . . . 3139
howensis, Amphichaetodon . . . . . . . . . . . . 3234 Johnius (Johnius) australis . . . . . . . . . . . 3140
Humbug dascyllus . . . . . . . . . . . . . . . . . 3354 Johnius (Johnius) belangerii . . . . . . . . . . . 3141
Johnius (Johnius) carouna . . . . . . . . . . . . 3142 Kyphosus vaigiensis . . . . . . . . . . . . . . . . 3295

Johnius (Johnius) carutta . . . . . . . . . . . . 3143 kyushini, Atrobucca . . . . . . . . . . . . . . . . 3132
Johnius (Johnius) coitor . . . . . . . . . . . . . 3144
Johnius (Johnius) heterolepis . .
Johnius (Johnius) hypostoma . .
.
.
.
.
. . . . . . 3145
. . . . . . 3146
L
Johnius (Johnius) laevis . . . . . . . . . . . . . 3147 labiosum, Diagramma . . . . . . . . . . . . . . 2966
Johnius (Johnius) latifrons . . . . . . . . . . . 3148 LABROIDEI . . . . . . . . . . . . . . . . . . . . 3333
Johnius (Johnius) macropterus . . . . . . . . . 3149 laevis, Johnius . . . . . . . . . . . . . . . . . . . 3144
Johnius (Johnius) macrorhynus . . . . . . . . . 3150 laevis, Johnius (Johnius) . . . . . . . . . . . . . 3147
Johnius (Johnius) trachycephalus . . . . . . . . 3151 lamarck, Genicanthus . . . . . . . . . . . . . . 3283
Johnius (Johnius) trewavasae . . . . . . . . . . 3152 Lamarck’s angelfish . . . . . . . . . . . . . . . . 3283
Johnius (Johnius) weberi . . . . . . . . . . . . . 3153 Lambe chivato . . . . . . . . . . . . . . . . . . . 3138
Johnius amblycephalus . . . . . . . . . . . 3138, 3149 LAMPRIDIDAE . . . . . . . . . . . . . . . . . . . 2825
Johnius australis . . . . . . . . . . . . . . 3141-3142 See also Vol. 3
Johnius belangerii . . . . . . . . . . . . . 3140, 3142 Large-eye croaker . . . .. . . . . . . . . . . . . 3157
Johnius carouna . . . . . . . . . . . . . . 3140-3141 Largebodied silverbiddy .. . . . . . . . . . . . . 2954
Johnius coitor . . . . . . . . . . . . . . . . 3147, 3151 Largefin croaker . . . . .. . . . . . . . . . . . . 3149
Johnius dussumieri . . . . . . . . . . . . . . . . 3139 Largescale croaker . . . .. . . . . . . . . . . . . 3145
Johnius fasciatus . . . . . . . . . . . . . . . . . . 3141 Largescaled terapon . . .. . . . . . . . . . . . . 3316
Johnius laevis . . . . . . . . . . . . . . . . . . . 3144 Larimichthys pamoides . . . . . . . . . . . . . . 3158
Johnius macropterus . . . . . . . . . . . . . . . 3139 lateoides, Otolithes . . . . . . . . . . . . . . . . 3172
Johnius macrorhynus . . . . . . . . . . . . 3152-3153 lateoides, Pterotolithus . . . . . . . . . . . . . . 3172
Johnius melanobranchium . . . . . . . . . . . . 3128 Lates calcarifer . . . . . . . . . . . . . . . . . . 3105
Johnius sina . . . . . . . . . . . . . . . . . . . . 3154 latezonatus, Amphiprion . . . . . . . . . . . . . 3348
Johnius spp. . . . . . . . . . . . . . . . . . . . . 3134 laticaudis, Lethrinus . . . . . . . . . . . . 3048, 3033
Johnius vogleri . . . . . . . . . . . . . . . . . . . 3154 latifrons, Johnius (Johnius) . . . . . . . . . . . 3148
Johnius weberi . . . . . . . . . . . . 3141-3142, 3150 lativittata, Pterocaesio . . . . . . . . . . . . . . 2935
jubata, Aspericorvina . . . . . . . . . . . . . . . 3128 Lattice monocle bream . . . . . . . . . . . . . . . 3077
Latticed butterflyfish . . . . . . . . . . . . . . . . 3249
latus, Acanthopagrus . . . . . . . . . . . . . . . 2997
K latus, Mylio . . . . . . . . . . . . . . . . . . . . 2997
kaakan, Pomadasys . . . . . . . . . . . . . . . . 2987 latus, Sparus . . . . . . . . . . . . . . . . . . . . 2997
kallopterus, Lethrinus . . . . . . . . . . . . . . . 3029 Lavender jobfish . . . . . . . . . . . . . . . . . . 2913
kapas, Gerres . . . . . . . . . . . . . . . . . . . 2953 Leaftail croaker . . . . . . . . . . . . . . . . . . . 3151
karnellai, Emmelichthys . . . . . . . . . . . . . 2838 LEIOGNATHIDAE . . . . . . . . . . . . . . 2792, 2946
Karut croaker . . . . . . . . . . . . . . . . . . . 3143
. Leiognathus . . . . . . . . . . . . . . . . . . . . 2792
kasmira, Lutjanus . . . . . . . . . . . . . 2879, 3183
. Leiognathus aureus . . . . . . . . . . . . . . . . 2800
Keyhole angelfish . . . . . . . . . . . . . . . . 3278
. Leiognathus berbis . . . . . . . . . . . . . . . . 2801
King soldierbream . . . . . . . . . . . . . . . . 2998
. Leiognathus bindus . . . . . . . . . . . . . 2802, 2815
King threadfin . . . . . . . . . . . . . . . . . . 3105
. Leiognathus blochii . . . . . . . . . . . . . . . . 2803
kishinouyei, Hapalogenys . . . . . . . . . . . . 2969 Leiognathus brevirostris . . . . . . . . . . . . . 2805
Klein’s butterflyfish . . . . . . . . . . . . . . . . . 3242 Leiognathus daura . . . . . . . . . . . . . . . . 2804
kleinii, Chaetodon . . . . . . . . . . . . . . . . . 3242 Leiognathus decorus . . . . . . . . . 2802, 2805, 2815
Knifetail pomfret . . . . . . . . . . . . . . . . . . 2836 Leiognathus dussumieri . . . . . . . . . . . . . 2806
kohleri, Pterocaesio . . . . . . . . . . . . . . . . 2936 Leiognathus elongatus . . . . . . . . . . . 2807, 2816
Koran angelfish . . . . . . . . . . . . . . . . . . . 3285 Leiognathus equulus . . . . . . . . . . . . 2806, 2808
kottelati, Monodactylus . . . . . . . . . . . 3216, 3219 Leiognathus fasciatus . . . . . . . . . . . . . . . 2809
Kuhlia . . . . . . . . . . . . . . . . . . . . . . . 3317 Leiognathus gerroides . . . . . . . . . . . . . . . 2804
Kuhlia boninensis . . . . . . . . . . . . . . . . . 3319 Leiognathus insidiator . . . . . . . . . . . . . . . 2820
Kuhlia marginata . . . . . . . . . . . . . . . . . 3319 Leiognathus leuciscus . . . . . . . . . . . . . . 2810
Kuhlia mugil . . . . . . . . . . . . . . . . . . . . 3319 Leiognathus lineolatus . . . . . . . . . . . . . . 2817
Kuhlia munda . . . . . . . . . . . . . . . . . . . 3320 Leiognathus longispinis . . . . . . . . . . . . . 2811
Kuhlia rupestris . . . . . . . . . . . . . . . . . . 3320 Leiognathus moretoniensis . . . . . . . . . . . . 2812
Kuhlia taeniura . . . . . . . . . . . . . . . . . . 3319 Leiognathus oblongus . . . . . . . . . . . . . . . 2801
KUHLIIDAE . . . . . . . . . . . . . . . . . 3305, 3317 Leiognathus pan . . . . . . . . . . . . . . . . . 2813
kuru, Polynemus . . . . . . . . . . . . . . . . . . 3111 Leiognathus rapsoni . . . . . . . . . . . . . . . 2814
kusakarii, Paracaesio . . . . . . . . . . . . . . . 2900 Leiognathus smithursti . . . . . . . . . . . . . . 2811
KYPHOSIDAE . . . . . . 2991, 3290, 3297, 3299, 3301 Leiognathus sp. 1 . . . . . . . . . . . . . . . . . 2817
Kyphosus bigibbus . . . . . . . . . . . . . . . . 3292 Leiognathus sp. 2 . . . . . . . . . . . . . . . . . 2823
Kyphosus cinerascens . . . . . . . . . . . . . . . 3293 Leiognathus splendens . . . . . . . . . . . 2802, 2815
Kyphosus fuscus . . . . . . . . . . . . . . . . . . 3292 Leiognathus spp. . . . . . . . . . . . . . . . . . 2792
Kyphosus gibsoni . . . . . . . . . . . . . . . . . 3295 Leiognathus stercorarius . . . . . . . . . . . . . 2816
Kyphosus lembus . . . . . . . . . . . . . . . . . 3295 Leiognathus virgatus . . . . . . . . . . . . . . . 2802
Kyphosus sydneyanus . . . . . . . . . . . . . . . 3294 lembus, Kyphosus . . . . . . . . . . . . . . . . . 3295
lemniscatus, Lutjanus . . . . . . . . . . . . . . . 2880 lineolatus, Lutjanus . . . .. . . . . . . . . . . . 2882

Lemon damselfish . . . .
. . . . . . . . . . . . . 3355 lineolatus, Scorpis . . . . .. . . . . . . . . . . . 3299
Lemon sweetlips . . . . .
. . . . . . . . . . . . . 2973 Lipocheilus carnolabrum .. . . . . . . . . . . . 2860
Lemonpeel angelfish . . .
. . . . . . . . . . . . . 3275 Lobotes . . . . . . . . . . .. . . . . . . . . . . . 2942
lentjan, Lethrinus . . . .
. . . . . . . . . . 3034, 3048 Lobotes surinamensis . . .. . . . . . . . . . . . 2944
Leptobrama muelleri . .
. . . . . . . . . . . . . 3210 LOBOTIDAE . . . . . . . . . . . . . . . . 2942, 2991
.
LEPTOBRAMIDAE . . .
. . . . . . . . . . 3202, 3210 Longbarbel goatfish . . . . . . . . . . . . . . . 3191
.
leptolepis, Nibea . . . . .
. . . . . . . . . . 3159, 3162 Longface emperor . . . . . . . . . . . . . . . . 3039
.
Lesser bream . . . . . . .
. . . . . . . . . . . . . 2831 Longfin bannerfish . . . . . . . . . . . . . . . . 3262
.
Lesson’s sweetlips . . . .
. . . . . . . . . . . . . 2975 Longfin emperor . . . . . . . . . . . . . . . . . 3030
.
lessonii, Plectorhinchus . . . . . . . . . . . . . 2975 Longfin silverbiddy . . . . . . . . . . . . . . . . 2958
.
LETHRINIDAE . . 2841, 2920, 2961, 2991, 3004, 3052 Longhead grunt . . . . . . . . . . . . . . . . . 2985
.
lethrinoides, Gymnocranius . . . . . . . . . . . 3025 longimana, Drepane . . . . . . . . . . . 3222-3223
.
Lethrinus . . . . . . . . 2961, 3004-3005, 3009, 3052 longimanus, Pentaprion .. . . . . . . . . . . . 2958
Lethrinus amamianus . . . . . . . . . . . . . . . 3036 longipectoralis, Polynemus . . . . . . . . . . . . 3113
Lethrinus amboinensis . . . . . . . . . . . . . . 3026 longipinnis, Taractichthys . . . . . . . . . . . . 2836
Lethrinus anarhynchus . . . . . . . . . . . . . . 3033 longirostris, Forcipiger . . . . . . . . . . . . . . 3260
Lethrinus atkinsoni . . . . . . . . . . . . . . . . 3027 longispinis, Leiognathus . . . . . . . . . . . . . 2811
Lethrinus choerorynchus . . . . . . . . . . . . . 3037 Longnosed butterflyfish . . . . . . . . . . . . . . 3260
Lethrinus chrysostomus . . . . . . . . . . . . . . 3036 Longrayed dwarf monocle bream . . . . . . . . . 3083
Lethrinus conchyliatus . . . . . . . . . . . . . . 3028 Longspine emperor . . . . . . . . . . . . . . . . 3031
Lethrinus elongatus . . . . . . . . . . . . . 3035, 3039 Longspined silverbiddy . . . . . . . . . . . . . . . 2960
Lethrinus erythracanthus . . . . . . . . . . . . . 3029 Longtail silverbiddy . . . . . . . . . . . . . . . . . 2951
Lethrinus erythropterus . . . . . . . . . . . . . . 3030 Lord Howe coralfish . . . . . . . . . . . . . . . . 3234
Lethrinus fletus . . . . . . . . . . . . . . . . . . 3033 loriculus, Centropyge . . . . . . . . . . . . . . . 3276
Lethrinus fraenatus . . . . . . . . . . . . . 3033, 3037 Luderick . . . . . . . . . . . . . . . . . . . . . . 3298
Lethrinus genivittatus . . . . . . . . . . . . . . . 3031 Lunar fusilier . . . . . . . . . . . . . . . . . . . . 2928
Lethrinus harak . . . . . . . . . . . . . . . . . . 3032 lunaris, Caesio . . . . . . . . . . . . . . . . . . 2928
Lethrinus hypselopterus . . . . . . . . . . . . . . 3030 Lunartail snapper . . . . . . . . . . . . . . . . . . 2881
Lethrinus kallopterus . . . . . . . . . . . . . . . 3029 Lune d’argent . . . . . . . . . . . . . . . . . . . 3218
Lethrinus laticaudis . . . . . . . . . . . . . 3048, 3033 Lunero . . . . . . . . . . . . . . . . . . . . . . . 2791
Lethrinus lentjan . . . . . . . . . . . . . . 3034, 3048 Luneur . . . . . . . . . . . . . . . . . . . . . . . 2791
Lethrinus mahsena . . . . . . . . . . . . . . . . 3027 lunula, Chaetodon . . . . . . . . . . . . . . . . 3243
Lethrinus microdon . . . . . . . . . . . . . 3026, 3035 lunulatus, Lutjanus . . . . . . . . . . . . . . . . 2881
Lethrinus miniatus . . . . . . . . . . . . . 3036, 3039 luteus, Parupeneus . . . . . . . . . . . . . . . . 3188
Lethrinus nebulosus . . . . . . . . . . . . 3037, 3048 LUTJANIDAE . . 2839-2840, 2920, 2943, 2961, 2991,
Lethrinus nematacanthus . . . . . . . . . . . . . 3031 3005, 3052, 3205, 3304
Lethrinus obsoletus . . . . . . . . . . . . . . . . 3038 Lutjanus . . . . . . . . . . . . . . . . . . . . . . 2843
Lethrinus olivaceus . . . . . . 3026, 3035, 3039, 3047 Lutjanus adetii . . . . . . . . . . . . . . . . . . 2861
Lethrinus opercularis . . . . . . . . . . . . . . . 3034 Lutjanus argentimaculatus . . . . . . . . . . . . 2862
Lethrinus ornatus . . . . . . . . . . . . . . . . . 3040 Lutjanus bengalensis . . . . . . . . . . . . . . . 2863
Lethrinus punctulatus . . . . . . . . . . . . . . 3048 Lutjanus biguttatus . . . . . . . . . . . . . . . . 2864
Lethrinus reticulatus . . . . . . . . . 3041, 3044, 3047 Lutjanus bitaeniatus . . . . . . . . . . . . . . . 2865
Lethrinus rhodopterus . . . . . . . . . . . . . . . 3032 Lutjanus bohar . . . . . . . . . . . . . . . . . . 2866
Lethrinus rubrioperculatus . . . . . . . . . . . . 3042 Lutjanus boutton . . . . . . . . . . . . . . 2867, 2890
Lethrinus semicinctus . . . . . . . . . 3043-3044, 3047 Lutjanus carponatatus . . . . . . . . . . . . . . 2868
Lethrinus sp. 1 . . . . . . . . . . . . . . . . . . 3046 Lutjanus decussatus . . . . . . . . . . . . . . . 2869
Lethrinus sp. 2 . . . . . . . . . . . . . . . . . . 3047 Lutjanus dodecacanthoides . . . . . . . . . . . . 2870
Lethrinus sp. 3 . . . . . . . . . . . . . . . . . . 3048 Lutjanus ehrenbergii . . . . . . . . . . . . . . . 2871
Lethrinus variegatus . . . . . 3042-3044-3045, 3047 Lutjanus erythropterus . . . . . . . . . . . 2872, 2884
Lethrinus xanthochilus . . . . . . . . . . . . . . 3045 Lutjanus fulviflamma . . . . . . . . . . . . . . . 2873
leuciscus, Leiognathus . . . . . . . . . . . . . . 2810 Lutjanus fulvus . . . . . . . . . . . . . . . . . . 2874
leucogrammicus, Dipterygonotus . . . . . . . . . 2931 Lutjanus fuscescens . . . . . . . . . . . . . . . . 2875
leucokranos, Amphiprion . . . . . . . . . . . . . 3349 Lutjanus gibbus . . . . . . . . . . . . . . . . . . 2876
leucotaenia, Brama . . . . . . . . . . . . . . . . 2831 Lutjanus goldiei . . . . . . . . . . . . . . . . . . 2877
lewisi, Pinjalo . . . . . . . . . . . . . . . . . . . 2906 Lutjanus johnii . . . . . . . . . . . . . . . . . . 2878
limbata, Acanthocepola . . . . . . . . . . . . . . 3331 Lutjanus kasmira . . . . . . . . . . . . . . 2879, 3183
lineatus, Plectorhinchus . . . . . . . . . . . . . 2976 Lutjanus lemniscatus . . . . . . . . . . . . . . . 2880
lineatus, Scolopsis . . . . . . . . . . . . . . 3077, 3088 Lutjanus lineolatus . . . . . . . . . . . . . . . . 2882
Lined butterflyfish . . . . . . . . . . . . . . . . . 3242 Lutjanus lunulatus . . . . . . . . . . . . . . . . 2881
Lined sweetlips . . . . . . . . . . . . . . . . . . . 2976 Lutjanus lutjanus . . . . . . . . . . . . . . 2882-2883
lineolata, Scorpis . . . . . . . . . . . . . . . . . 3299 Lutjanus madras . . . . . . . . . . . . . . . . . 2883
lineolatus, Chaetodon . . . . . . . . . . . . . . . 3242 Lutjanus malabaricus . . . . . . . . . . . . 2884, 2895
lineolatus, Leiognathus . . . . . . . . . . . . . . 2817 Lutjanus maxweberi . . . . . . . . . . . . . . . 2885
Lutjanus mizenkoi . . . . . . . . . . . . . . . . 2886 marginata, Kuhlia . . . . . . . . . . . . . . . . . 3319

Lutjanus monostigma . . . . . . . . . . . . . . . 2887 marginatus, Nemipterus . . . . . . . . . . 3073, 3079
Lutjanus quinquelineatus . . . . . . . . . . . . 2888 Margined coralfish . . . .. . . .
. . . . . . . . . 3257
Lutjanus rivulatus . . . . . . . . . . . . . . . . . 2889 Marquesan butterflyfish . . . . .
. . . . . . . . . 3239
Lutjanus rufolineatus . . . . . . . . . . . . 2867, 2890 Marr’s fusilier . . . . . . .
. . . .
. . . . . . . . . 2936
Lutjanus russelli . . . . . . . . . . . . . . . . . 2891 marri, Pterocaesio . . . .. . . .
. . . . . . . . . 2936
Lutjanus sebae . . . . . . . . . . . . . . . 2884, 2892 marshi, Etelis . . . . . .. . . .
. . . . . . . . . 2857
Lutjanus semicinctus . . . . . . . . . . . . . . . 2893 Masked bannerfish . . . . . . . .
. . . . . . . . . 3263
Lutjanus stellatus . . . . . . . . . . . . . . . . . 2894 Matode voile . . . . . . .. . . .
. . . . . . . . . 3288
Lutjanus timorensis . . . . . . . . . . . . . . . . 2895 Mauvelip threadfin bream . . . .
. . . . . . . . . 3079
Lutjanus vitta . . . . . . . . . . . . . . . . . . . 2896 maxweberi, Lutjanus . . . . . .
. . . . . . . . . 2885
lutjanus, Lutjanus . . . . . . . . . . . . . . 2882-2883 megalolepis, Secutor . . . . . .
. . . . . . . . . 2821
luzonius, Upeneus . . . . . . . . . . . . . . . . . 3198 Meiacanthus . . . . . . . . . . .
. . . . . . 3087-3088
melannotus, Chaetodon . . . . .
. . . . . . . . . 3243
M melanobranchium, Johnius .
melanochir, Polynemus . . .
.
.
.
. . . . . . . . . 3128
.
. . . . . . 3100, 3115
maccullochi, Owstonia . . . . . . . . . . . . . . 3331 melanophyrs, Parascolopsis . .
. . . . . . . . . 3082
Macolor . . . . . . . . . . . . . . . . . . . . . . 2849 melanopus, Amphiprion . . . .
. . . . . . . . . 3349
Macolor macolor . . . . . . . . . . . . . . . . . 2898 melanopus, Coradion . . . . . .
. . . . . . . . . 3259
Macolor macularis . . . . . . . . . . . . . 2897-2898 melanosoma, Chaetodontoplus . . . . . . . . . 3281
.
Macolor niger . . . . . . . . . . . . . . . . 2897-2898 melanospilos, Genicanthus . . . . . . . . . . . 3283
.
Macolor typus . . . . . . . . . . . . . . . . . . . 2898 melanurus, Dascyllus . . . . . . . . . . . . . . 3354
.
macolor, Macolor . . . . . . . . . . . . . . . . . 2898 Mene maculata . . . . . . . . . . . . . . . . . 2791
.
macracanthus, Gerres . . . . . . . . . . . 2952, 2960 MENIDAE . . . . . . . . . . . . . . . . . 2791, 3217
.
macrocephalus, Argyrosomus . . . . . . . . . . . 3169 Meredith’s angelfish . . . . . . . . . . . . . . . 3281
.
macrocephalus, Pennahia . . . . . . . . . . . . 3169 meredithi, Chaetodontoplus . . . . . . . . . . 3281
.
macrochir, Polydactylus . . . . . . . . . . 3104-3105 Merten’s butterflyfish . . . . . . . . . . . . . . . 3244
.
macronemus, Parupeneus . . . . . . . . . . . . 3191 mertensii, Chaetodon . . . . . . . . . . . . . . 3244
.
macrophthalmus, Pennahia . . . . . . . . . . . . 3168 mesoleucus, Chaetodontoplus .
. . . . . . . . . 3281
macrophthalmus, Polydactylus . . . . . . . . . . 3106 mesoprion, Nemipterus . . . . . . . . . . . . . 3079
.
macrophthalmus, Sphenanthias . . . . . . . . . 3331 Mesopristes argenteus . . . . . . . . . . . . . 3309
.
macroptera, Sciaena . . . . . . . . . . . . . . . . 3149 Mesopristes cancellatus . . . . . . . . . . . . . 3310
.
macropterus, Johnius . . . . . . . . . . . . . . . 3139 Meyer’s butterflyfish . . . . . . . . . . . . . . . 3244
.
macropterus, Johnius (Johnius) . . . . . . . . . 3149 meyeri, Chaetodon . . . . . . . . . . . . . . . 3244
.
macrorhynus, Johnieops . . . . . . . . . . . . . . 3150 MICROCANTHIDAE . . . . . . 3225, 3297, 3300-3301
.
macrorhynus, Johnius . . . . . . . . . . . 3152-3153 Microcanthids . . . . . . . . . . . . . . . . . . 3301
.
macrorhynus, Johnius (Johnius) . . . . . . . . 3150 Microcanthus strigatus . . . . . . . . . . 3225, 3303
.
macrosoma, Gerres . . . . . . . . . . . . . . . . 2954 microdon, Gymnocranius . . . . . . . . . . . . 3024
.
macularis, Macolor . . . . . . . . . . . . . 2897-2898 microdon, Lethrinus . . . . . . . . . . . 3026, 3035
.
maculata, Mene . . . . . . . . . . . . . . . . . . 2791 microdon, Panna . . . . . . . . . . . . . . . . 3166
.
maculatus, Otolithes . . . . . . . . . . . . . . . . 3173 microdon, Pristipomoides . . . . . . . . . . . . 2913
.
maculatus, Pomadasys . . . . . . . . . . . . . . 2988 microgenys, Nibea . . . . . . . . . . . . . . . 3160
.
maculatus, Pterotolithus . . . . . . . . . . . . . 3173 microlepis, Boesemania . . . . . . . . . . . . . 3136
.
madras, Lutjanus . . . . . . . . . . . . . . . . . 2883 microlepis, Coius . . . . . . . . . . . . . . . . 2942
.
magnificum, Glaucosoma . . . . . . . . . . . . 3208 microlepis, Pristipomoides . . . . . . . . . . . 2910
.
mahsena, Lethrinus . . . . . . . . . . . . . . . . 3027 microphthalmus, Pinjalo . . . . . . . . . 2906-2907
.
major, Chrysophrys . . . . . . . . . . . . . . . . 3002 microstoma, Polydactylus . . . . . . 3104, 3107, 3110
.
major, Pagrus . . . . . . . . . . . . . . . . . . . 3002 Midnight angelfish . . . . . . . . . . . . . . . . 3278
.
major, Sparus . . . . . . . . . . . . . . . . . . . 3002 Midnight snapper . . . . . . . . . . . . . . . . . 2897
.
Malabar blood snapper . . . . . . . . . . . . . . . 2884 miichthyoides, Nibea . . . . . . . . . . . . . . 3126
.
malabaricus, Lutjanus . . . . . . . . . . . 2884, 2895 milii, Scaevius . . . . . . . . . . . . . . . 3051, 3086
.
Maldive emperor . . . .
. . . . . . . . . . . . . . 3046 miniatus, Lethrinus . . . . . . . . . . . . 3036, 3039
.
Mamila . . . . . . . . .
. . . . . . . . . . . . . . 3089 Minstrel sweetlips . . . . . . . . . . . . . . . . 2981
.
Mamila à bande dorée .
. . . . . . . . . . . . . . 3087 minuta, Gazza . . . . . . . . . . . . . . . . . . 2799
.
Mamila griffée . . . . .
. . . . . . . . . . . . . . 3087 mizenkoi, Lutjanus . . . . . . . . . . . . . . . 2886
.
Mamila rosée . . . . . .
. . . . . . . . . . . . . . 3081 Mojarra alona . . . . . . . . . . . . . . . . . . 2958
.
Manefishes . . . . . . .
. . . . . . . . . . . . . . 2837 Mojarra chata . . . . . . . . . . . . . . . . . . 2950
.
Mangrove red snapper .
. . . . . . . . . . . . . . 2862 Mojarra común . . . . . . . . . . . . . . . . . . 2956
.
mangula, Pempheris . . . . . . . . . . . . . . . 3201 Mojarra de hebra . . . . . . . . . . . . . . . . . 2952
.
Manybanded sweetlips . . . . . . . . . . . . . . . 2977 Mojarra elegante . . . . . . . . . . . . . . . . . 2955
.
Manybar goatfish . . . .. . . . . . . . . . . . . . 3192 Mojarra espinuda . . . . . . . . . . . . . . . . . 2957
.
Manyspined butterflyfish. . . . . . . . . . . . . . 3261 Mojarra timonera . . . . . . . . . . . . . . . . . 2951
.
margaritifer, Scolopsis . . . . . . . . . . . . . . 3088 Mojarras . . . . . . . . . . . . . . . . . . . . . 2946
.
marginalis, Chelmon . . . . . . . . . . . . . . . 3257 molucca, Pempheris . . . . . . . . . . . . . . . 3201
.
Moluccan snapper . . . . . . . . . . . . . . . . . 2867 munda, Kuhlia . . . . . . . . . . . . . . . . . . 3320

moluccensis, Pomacentrus . . . . . . . . . . . . 3355 Myers pomfret . . . . . . . . . . . . . . . . . . . 2833
moluccensis, Upeneus . . . . . . . . . . . . . . 3196 myersi, Brama . . . . . . . . . . . . . . . . 2831-2833
monoceros, Heniochus . . . . . . . . . . . . . . 3263 Mylio berda . . . . . . . . . . . . . . . . . . . . 2996
Monocle breams . . . . . . . . . . . . . . . . . . 3051 Mylio latus . . . . . . . . . . . . . . . . . . . . . 2997
Monodactyle voile . . . . . . . . . . . . . . . . . 3219
MONODACTYLIDAE . . . . .
Monodactylus argenteus . . .
. . . . . . . 3216, 3221
. . . . 3216, 3218-3219
N
Monodactylus kottelati . . . . . . . . . . . 3216, 3219 nagasakiensis, Pentapodus . . . . . . . . . . . . 3084
monogramma, Scolopsis . . . . . . . . . . . . . 3076 Naked toothpony . . . . . . . . . . . . . . . . . . 2798
Monogrammed monocle bream . . . . . . . . . . 3076 Nannatherina . . . . . . . . . . . . . . . . . . . 3317
monostigma, Lutjanus . . . . . . . . . . . . . . 2887 Nannoperca . . . . . . . . . . . . . . . . . . . . 3317
Monotaxis . . . . . . . . . . . . . . . . . . . . . 2991 Narc angelfish . . . . . . . . .
. . . . . . . . . . 3277
Monotaxis grandoculis . . . . . . . . . . . . . . 3049 narcosis, Centropyge . . . . .
. . . . . . . . . . 3277
Moonfish . . . . . . . . . . . .. . . . . . . . . . 2791 navarchus, Pomacanthus . . .
. . . . . . . . . . 3285
Moonfishes . . . . . . . . . . .. . . . . . . . . . 3216 nebulosus, Lethrinus . . . . .
. . . . . . . 3037, 3048
moretoniensis, Leiognathus . . . . . . . . . . . 2812 nematacanthus, Lethrinus . . .
. . . . . . . . . . 3031
Morwong . . . . . . . . . . . . . . . . . . . . . . 3329 nematophorus, Glabrilutjanus . . . . . . . . . . 2918
Morwongs . . . . . . . . . . . . . . . . . . . . . 3329 nematophorus, Nemipterus . . . . . . . . . . . . 3069
mossambica, Wattsia . . . . . . . . . . . . . . . 3050 nematophorus, Symphorus . . . . . . . . . . . . 2918
mossambicus, Oreochromis . . . . . . . . 3333, 3336 nematopus, Nemipterus . . . . . . . . . . . . . . 3080
Motambo bérbero . . . . . . . . . . . . . . . . . 2801 NEMIPTERIDAE . . . . . 2841, 2920, 2962, 3005, 3051
Motambo boxeador . . . . . . . . . . . . . . . . . 2820 Nemipterids . . . . . . . . . . . . . . . . . . . . 3051
Motambo buldog . . . . . . . . . . . . . . . . . . 2822 Nemipterus . . . . 3051-3052, 3062, 3070-3071, 3081
Motambo común . . . . . . . . . . . . . . . . . . 2808 Nemipterus aurifilum . . . . . . . . . . . . . . . 3078
Motambo de dos manchas . . . . . . . . . . . . . 2803 Nemipterus aurora . . . . . . . . . . . . . . . . 3062
Motambo de hebra . . . . . . . . . . . . . . . . . 2810 Nemipterus balinensis . . . . . . . . . . . . . . 3078
Motambo de onda . . . . . . . . . . . . . . . . . 2806 Nemipterus balinensoides . . . . . . . . . . . . 3078
Motambo de penacho . . . . . . . . . . . . . . . 2811 Nemipterus bathybius . . . . . . . . . . . . . . . 3063
Motambo dentudo . . . . . . . . . . . . . . . . . 2799 Nemipterus bipunctatus . . . . . . . . . . . . . 3062
Motambo dorado . . . . . . . . . . . . . . . . . . 2804 Nemipterus celebicus . . . . . . . . . . . . . . . 3064
Motambo esplendor . . . . . . . . . . . . . . . . 2815 Nemipterus delagoae . . . . . . . . . . . . . . . 3062
Motambo liso . . . . . . . . . .. . . . . . . . . . 2798 Nemipterus filamentosus . . . . . . . . . . . . . 3069
Motambo naranjón . . . . . . . . . . . . . . . . . 2802 Nemipterus furcosus . . . . . . . . . . . . 3065, 3071
Motambo ornado . . . . . . . . . . . . . . . . . . 2817 Nemipterus gracilis . . . . . . . . . . . . . . . . 3079
Motambo rayado . . . . . . . . . . . . . . . . . . 2809 Nemipterus hexodon . . . . . . . . . . . . . . . 3066
Mottled fusilier . . . . . . . . .
. . . . . . . . . . 2931 Nemipterus isacanthus . . . . . . . . . . . . . . 3067
Mozambique large-eye bream . . . . . . . . . . . 3050 Nemipterus japonicus . . . . . . . . . . . . . . . 3068
mucronatus, Hapalogenys . . . . . . . . . . . . . 2968 Nemipterus marginatus . . . . . . . . . . . 3073, 3079
muelleri, Chelmon . . . . . . . . . . . . . . . . 3257 Nemipterus mesoprion . . . . . . . . . . . . . . 3079
muelleri, Leptobrama . . . . . . . . . . . . . . . 3210 Nemipterus nematophorus . . . . . . . . . . . . 3069
mugil, Kuhlia . . . . . . . . . . . . . . . . . . . 3319 Nemipterus nematopus . . . . . . . . . . . . . . 3080
Müller’s coralfish . . . . . . . .
. . . . . . . . . . 3257 Nemipterus nemurus . . . . . . . . . . . . . . . 3070
Mullid fishes . . . . . . . . . .
. . . . . . . . . . 3175 Nemipterus peronii . . . . . . . . . . . . . . . . 3071
MULLIDAE . . . . . . . . . . . . . . . . . . . . . 3175 Nemipterus sp. 1 . . . . . . . . . . . . . . . . . 3080
Mulloides auriflamma . . . . . . . . . . . . . . . 3183 Nemipterus tambuloides . . . . . . . . . . . . . 3072
Mulloides erythrinus . . . . . .. . . . . . . . . . 3183 Nemipterus theodorei . . . . . . . . . . . . . . . 3080
Mulloides vanicolensis . . . . . . . . . . . . . . 3183 Nemipterus thosaporni . . . . . . . . . . . . . . 3073
Mulloidichthys . . . . . . . . . . . . . . . 3175-3176 Nemipterus tolu . . . . . . . . . . . . . . . . . . 3071
Mulloidichthys flavolineatus . . . . . . . . 3181, 3183 Nemipterus virgatus . . . . . . . . . . 3005, 3051, 3074
Mulloidichthys pflugeri . . . . . . . . . . . . . . 3182 Nemipterus vitiensis . . . . . . . . . . . . . . . . 3081
Mulloidichthys samoensis . . . . . . . . . . . . . 3181 Nemipterus zysron . . . . . . . . . . . . . . . . 3081
Mulloidichthys vanicolensis . . . . . . . . . . . 3183 nemurus, Nemipterus . . . . . . . . . . . . . . . 3070
Multibarred angelfish . . . . . .
. . . . . . . . . . 3277 Neocirrhites armatus . . . . . . . . . . . . . . . 3321
Multicolor angelfish . . . . . . .
. . . . . . . . . . 3276 Neon damselfish . . . . . . . . . . . . . . . . . . 3355
multicolor, Centropyge . . . . . . . . . . . . . . 3276 New Guinean tigerperch . . . . . . . . . . . . . . 2945
multidens, Pristipomoides . . . . . . . . . . . . 2912 Nibblers . . . . . . . . . . . . . . . . . . . . . . 3297
multifasciatus, Centropyge . . . . . . . . . 3277, 3279 nibe, Atrobucca . . . . . . . . . . . . . . . . . . 3133
multifasciatus, Parupeneus . . . . . . . . . . . . 3192 Nibea . . . . . . . . . . . . . . . . . . . . . . . . 3124
multifilis, Polistonemus . . . .. . . . . . . . . . 3116 Nibea acuta . . . . . . . . . . . . . . . . . . . . 3137
multifilis, Polynemusm . . . . . . . . . . . . . . 3116 Nibea diacanthus . . . . . . . . . . . . . . . . . 3171
multiradiatus, Polydactylus . . . 3098, 3101-3102, 3108 Nibea leptolepis . . . . . . . . . . . . . . . 3159, 3162
multispinosus, Hemitaurichthys . . . . . . . . . 3261 Nibea microgenys . . . . . . . . . . . . . . . . . 3160
multivittatus, Plectorhinchus . . . . . . . . . . . 2977 Nibea miichthyoides . . . . . . . . . . . . . . . . 3126
Nibea semifasciata . . . . . . . . . . . . . 3161, 3163 orientalis, Plectorhinchus . . . . . . . . . . . . . 2982

Nibea soldado . . . . . . . . . 3136, 3156, 3159, 3162 Ornate angelfish . . . . . . . . . . . . . . . . . . 3282
Nibea spp. . . . . . . . . . . . . . . . . . . . . . 3135 Ornate butterflyfish . . . . . . . . . . . . . . . . . 3246
Nibea squamosa . . . . . . . . . . . . . . . . . . 3163 Ornate emperor . . . . . . . . . . . . . . . . . . 3040
niger, Macolor . . . . . . . . . . . . . . . . 2897-2898 Ornate jobfish . . . . . . . . . . . . . . . . . . . 2908
niger, Parastromateus . . . . . . . . . . . . . . 2825 Ornate ponyfish . . . . . . . . . . . . . . . . . . 2817
See also Vol. 4 Ornate threadfin bream . . . . . . . . . . . . . . 3066
nigriocellus, Centropyge . . . . . . . . . . . . . 3277 ornatissimus, Chaetodon . . . . . . . . . . 3244, 3246
nigripes, Cheilodactylus . . . . . . . . . . . . . 3329 ornatus, Lethrinus . . . . . . . . . . . . . . . . 3040
nigripinnnis, Polydactylus . . . . . . . . . . . . 3109 otaitensis, Pempheris . . . . . . . . . . . . . . . 3201
nigrus, Plectorhinchus . . . . . .. . . . . . . . . 2974 Otolithes argenteus . . . . . . . . . . . . . . . . 3164
niloticus niloticus, Oreochromis . . . . . . . . . 3333 Otolithes lateoides . . . . . . . . . . . . . . . . . 3172
niloticus, Oreochromis niloticus . . . . . . . . . 3333 Otolithes maculatus . . . . . . . . . . . . . . . . 3173
nippon, Chaetodon . . . . . . . . . . . . . . . . 3245 Otolithes ruber . . . . . . . . . . . . . . . . . . 3164
Northwest Australian whiptail . . . . . . . . . . . 3085 Otolithoides biauritus . . . . . . . . . . . . . . . 3165
Notchedfin threadfin bream . . . . . . . . . . . . 3071 Otolithoides brunneus . . . . . . . . . . . . . . . 3165
novaeguineae, Johnius (Johnieops) . . . . . . . 3155 Otolithoides perarmatus . . . . . . . . . . . . . . 3167
novaehollandiae, Equulites . . . . . . . . . . . . 2810 Otolithoides siamensis . . . . . . . . . . . . . . . 3151
nox, Centropyge . . . . . . . . . . . . . . . . . . 3278 oualensis, Pempheris . . . . . . . . . . . . . . . 3201
nuchalis, Equula . . . . . . . . . . . . . . . . . 2805 Oval butterflyfish . . . . . . . . . . . . . . . . . . 3254
nudimaxillaris, Etelis . . . . . . . . . . . . . . . 2916 Ovalspot butterflyfish . . . . . . . . . . . . . . . . 3252
ovata, Pennahia . . . . . . . . . . . . . . . . . . 3169
O Owstonia . . . . . . . . .
Owstonia maccullochi .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. . . . . . 3332
. . . . . . 3331
Obliquebanded snapper . . . . . . . . . . . . . . 2915 OWSTONIINAE . . . . . . . . . . . . . . . . . . 3332
Obliquebarred monocle bream . . . . . . . . . . . 3089 Oxycirrhites typus . . . . . . . . . . . . . . . . . 3321
oblongus, Gerres . . . . . . . . . . . . . . . . . 2955 oxyrhynchus, Pelates . . . . . . . . . . . . . . . 3313
oblongus, Leiognathus . . . . . . . . . . . . . . . 2801 oxyrhynchus, Rhyncopelates . . . . . . . . . . . 3313
obscurum, Plectorhinchus . . . . . . . . . . . . . 2970 oxyrhynchus, Terapon . . . . . . . . . . . . . . . 3313
obsoletus, Lethrinus . . . . . . . . . . . . . . . . 3038 oyena, Gerres . . . . . . . . . . . . . 2951, 2953, 2956
ocellaris, Amphiprion . . . . . . . . . . . . . . . 3350
ocellaris, Cichla . . . . . . . . . . . . . . . . . . 3333
Ocellate coralfish . . . . . . . . . . . . . . . . . . 3265 P
ocellatus, Parachaetodon . . . . . . . . . . . . . 3265 Pacific croaker . . . . . . . . . . . . . . . . . . . 3156
ocellicaudus, Chaetodon . . . . . . . . . . . . . 3245 Pacific doublesaddle butterflyfish . . . . . . . . . 3254
Ochreband goatfish . . . . . . . . . . . . . . . . 3198 Pacific fanfish . . . . . . . . . . . . . . . . . . . 2834
octofasciatus, Chaetodon . . . . . . . . . . . . . 3246 Pacific yellowtail emperor . . . . . . . . . . . . . 3027
ocyurus, Sectator . . . . . . . . . . . . . . . . . 3296 pacificus, Johnius (Johnieops) . . . . . . . . . . 3156
oedogenys, Austronibea . . . . . . . . . . . . . . 3134 Pagre à nageoires jaunes . . . . . . . . . . . . . 2997
Old wife . . . . . . . . . . . . . . . . . . . . . . . 3287 Pagre picnic . . . . . . . . . . . . . . . . . . . . 2996
oligospilus, Upeneus . . . . . . . . . . . . . . . . 3199 Pagrus auratus . . . . . . . . . . . . . . . . . . 3002
olivaceus, Lethrinus . . . . . . 3026, 3035, 3039, 3047 Pagrus major . . . . . . . . . . . . . . . . . . . 3002
Onespot snapper . . . . . . . . . . . . . . . . . . 2887 Painted sweetlips . . . . . . . . . . . . . . . . . 2967
Onestripe fusilier . . . . . . . . . . . . . . . . . . 2939 Palefin threadfin bream . . . . . . . . . . . . . . 3073
opercularis, Lethrinus . . . . . . . . . . . . . . . 3034 pamoides, Larimichthys . . . . . . . . . . . . . 3158
opercularis, Pomadasys . . . . . . . . . . . . . . 2983 pamoides, Pseudotosciaena . . . . . . . . . . . . 3158
OPISTOGNATHIDAE . . . . . . . . . . . . . . . 3331 Pampus . . . . . . . . . . . . . . . . . . . . . . 2825
See also Vol. 4 See also Vol. 6
Orange anemonefish . . . . . . . . . . . . . . . . 3352 pan, Leiognathus . . . . . . . . . . . . . . . . . 2813
Orange croaker . . . . . . . . . . . . . . . . . . . 3131 Panchito abanderado . . . . . . . . . . . . . . . 2909
Orangebanded coralfish . . . . . . . . . . . . . . 3259 Panchito adornado . . . . . . . . . . . . . . . . . 2908
Orangefaced angelfish . . . . . . . . . . . . . . . 3280 Panchito de bandas doradas . . . . . . . . . . . 2912
Orangefin anemonefish . . . . . . . . . . . . . . 3347 Panchito de ojo dorado . . . . . . . . . . . . . . 2911
Orangefin ponyfish . . . . . . . . . . . . . . . . . 2802 Panchito dentón . . . . . . . . . . . . . . . . . . 2914
Orangespotted emperor . . . . . . . . . . . . . . 3029 Panchito escamudo . . . . . . . . . . . . . . . . 2905
Orangestriped emperor . . . . . . . . . . . . . . 3038 Panchito hebra . . . . . . . . . . . . . . . . . . . 2910
orcini, Brama . . . . . . . . . . . . . . . . 2831-2832 Panchito lavanda . . . . . . . . . . . . . . . . . . 2913
ordinalis, Plectorhinchus . . . . . . . . . . . . . 2977 Panchito rayado . . . . . . . . . . . . . . . . . . 2915
Oreochromis aureus . . . . . . . . . . . . . . . 3333 Panchote cacao . . . . . . . . . . . . . . . . . . 2903
Oreochromis mossambicus . . . . . . . . . 3333, 3336 Panchote rabo amarillo . . . . . . . . . . . . . . 2904
Oreochromis niloticus niloticus . . . . . . . . . 3333 Panchote sillevo . . . . . . . . . . . . . . . . . . 2900
Oreochromis urolepis . . . . . . . . . . . . . . . 3333 Panchote vanuatú . . . . . . . . . . . . . . . . . 2899
Oriental butterflyfish . . . . . . . . . . . . . . . . 3236 Panna . . . . . . . . . . . . . . . . . . . . . . . 3124
Oriental sweetlips . . . . . . . . . . . . . . . . . 2982 Panna croaker . . . . . . . . . . . . . . . . . . . 3166
Panna microdon . . . . . . . . . . . . . . . . . . 3166 Pargode rayas amarillas . . . . . . . . . . . . . 2880

Panna perarmatus . . . . . . . . . . . . . . . . 3167 Pargode rayas azules . . . . . . . . . . . . . . . 2879
Paperhead croaker . . . .. . . .
. . . . . . . . . 3155 Pargode Samoa . . . . . . . . . . . . . . . . . . 2886
Papillon à rideaux jaunes. . . .
. . . . . . . . . 3261 Pargode Timor . . . . . . . . . . . . . . . . . . . 2895
Papuan black snapper . . . . . .
. . . . . . . . . 2877 Pargodiablo . . . . . . . . . . . . . . . . . . . . 2918
Papuan ponyfish . . . . .. . . .
. . . . . . . . . 2814 Pargoeglefino . . . . . . . . . . . . . . . . . . . 2887
Paracaesio . . . . . . . .
. . . 2849, 2902, 2904, 2920 Pargoenano . . . . . . . . . . . . . . . . . . . . 2885
Paracaesio cantharoides . . . . . . . . . . . . . 2904 Pargoescarlata . . . . . . . . . . . . . . . . . . 2859
Paracaesio gonzalesi . . . . . . . . . . . . . . . 2899 Pargoestrellado . . . . . . . . . . . . . . . . . . 2894
Paracaesio kusakarii . . . . . . . . . . . . . . . 2900 Pargoimperial . . . . . . . . . . . . . . . . . . . 2892
Paracaesio paragrapsimodon . . . . . . . . . . 2901 Pargojaspeado . . . . . . . . . . . . . . . . . . 2878
Paracaesio pedleyi . . . . . . . . . . . . . . . . 2904 Pargojorobado . . . . . . . . . . . . . . . . . . . 2876
Paracaesio sordida . . . . . . . . . . . . . . . . 2902 Pargomadraseño . . . . . . . . . . . . . . . . . 2883
Paracaesio stonei . . . . . . . . . . . . . . . . . 2903 Pargomalabárico . . . . . . . . . . . . . . . . . 2884
Paracaesio tumida . . . . . . . . . . . . . . . . . 2904 Pargomanchado . . . . . . . . . . . . . . . . . . 2871
Paracaesio xanthura . . . . . . . . . . . . 2899, 2904 Pargomaori . . . . . . . . . . . . . . . . . . . . 2889
Parachaetodon . . . . . . . . . . . . . . . . . . 3224 Pargomedianoche . . . . . . . . . . . . . . . . . 2897
Parachaetodon ocellatus . . . . . . . . . . . . . 3265 Pargonachero . . . . . . . . . . . . . . . . . . . 2861
Paracirrhites . . . . . . . . . . . . . . . . . . . . 3324 Pargoojo de buey . . . . . . . . . . . . . . . . . 2891
Paracirrhites forsteri . . . . . . . . . . . . . . . 3327 Pargorabo negro . . . . . . . . . . . . . . . . . 2874
Paracirrhites hemistictus . . . . . . . . . . . . . 3328 Pargoraboluna . . . . . . . . . . . . . . . . . . . 2881
Paracirrhites polystictus . . . . . . . . . . . . . 3328 Pargorubí . . . . . . . . . . . . . . . . . . . . . 2857
Paracirrhites typee . . . . . . . . . . . . . . . . 3327 Pargosolrayo . . . . . . . . . . . . . . . . . . . 2870
Paradicichthys venenatus . . . . . . . . . . . . . 2918 Pargotintero . . . . . . . . . . . . . . . . . . . . 2873
Paradise whiptail . . . . . . . . . . . . . . . . . . 3085 Pargotrompudo . . . . . . . . . . . . . . . . . . 2860
paradiseus, Pentapodus . . . . . . . . . . . . . . 3085 Pargovelero . . . . . . . . . . . . . . . . . . . . 2917
paradiseus, Polynemus . . . . . . . . 3090, 3113-3115 Pargoverde . . . . . . . . . . . . . . . . . . . . 2856
paragrapsimodon, Paracaesio . . . . . . . . . . 2901 Parupeneus . . . . . . . . . . . . . . . . . 3175-3176
Parakuhlia . . . . . . . . . . . . . . . . . . . . . 3317 Parupeneus barberinoides . . . . . . . . . . . . 3184
Parapolynemus verekeri . . . . . . . . . . . . . 3103 Parupeneus barberinus . . . . . . . . . . . . . . 3185
Parapriacanthus . . . . . . . . . . . . . . . . . 3201 Parupeneus bifasciatus . . . . . . . . . . . . . . 3186
Parapristipomoides . . . . . . . . . . . . . . . . 2840 Parupeneus chryserydros . . . . . . . . . . . . . 3188
Parapristipomoides squamimaxillaris . . . . . . 2905 Parupeneus ciliatus . . . . . . . . . . . . . . . . 3187
Parascolopsis . . . . . . . . . . . . . . . . . . . 3051 Parupeneus cinnabarinus . . . . . . . . . . . . . 3189
Parascolopsis eriomma . . . . . . . . . . . . . . 3081 Parupeneus crassilabris . . . . . . . . . . . . . . 3186
Parascolopsis inermis . . . . . . . . . . . . . . . 3082 Parupeneus cyclostomus . . . . . . . . . . . . . 3188
Parascolopsis melanophyrs . . . . . . . . . . . . 3082 Parupeneus fraterculus . . . . . . . . . . . 3187, 3194
Parascolopsis rufomaculatus . . . . . . . . . . . 3082 Parupeneus heptacanthus . . . . . . . . . . . . 3189
Parascolopsis tanyactis . . . . . . . . . . . . . . 3083 Parupeneus indicus . . . . . . . . . . . . . . . . 3190
Parascolopsis tosensis . . . . . . . . . . . . . . 3083 Parupeneus jansenii . . . . . . . . . . . . . . . 3189
parasema, Chrysiptera . . . . . . . . . . . . . . 3353 Parupeneus luteus . . . . . . . . . . . . . . . . . 3188
Parastromateus niger . . . . . . . . . . . . . . . 2825 Parupeneus macronemus . . . . . . . . . . . . . 3191
See also Vol. 3 Parupeneus multifasciatus . . . . . . . . . . . . 3192
Parequula . . . . . . . . . . . . . . . . . . . . . 2946 Parupeneus pleurospilos . . . . . . . . . . . . . 3189
Pargo abanderado . . . . . . . . . . . . . . . . . 2868 Parupeneus pleurostigma . . . . . . . . . . . . 3193
Pargo ajedrezado . . . . . . . . . . . . . . . . . 2869 Parupeneus pleurotaenia . . . . . . . . . . . . . 3187
Pargo bermellón . . . . . . . . . . . . . . . . . . 2855 Parupeneus porphyreus . . . . . . . . . . . . . 3187
Pargo bitilla . . . . . . . . . . . . . . . . . . . . . 2896 Parupeneus rubescens . . . . . . . . . . . . . . 3187
Pargo blanco y negro . . . . . . . . . . . . . . . 2898 Parupeneus signatus . . . . . . . . . . . . . . . . 3194
Pargo boquidulce . . . . . . . . . . . . . . . . . . 2854 Parupeneus spilurus . . . . . . . . . . . . . . . 3194
Pargo carmesi . . . . . . . . . . . . . . . . . . . 2872 Parupeneus trifasciatus . . . . . . . . . . . 3186, 3192
Pargo cinteado . . . . . . . . . . . . . . . . . . . 2893 pauciradiata, Brama . . . . . . . . . . . . . . . 2834
Pargo de agua dulce . . . . . . . . . . . . . . . . 2875 Pavillon . . . . . . . . . . . . . . . . . . . . . . . 3262
Pargo de banda bianca . . . . . . . . . . . . . . 2864 Pawak croaker . . . . . . . . . . . . . . . . . . . 3170
Pargo de Bengala . . . . . . . . . . . . . . . . . 2863 pawak, Argyrosomus . . . . . . . . . . . . . . . 3170
Pargo de cinco lineas . . . . . . . . . . . . . . . 2888 pawak, Pennahia . . . . . . . . . . . . . . . . . 3170
Pargo de dos manchas . . . . . . . . . . . . . . 2866 Pearl perch . . . . . . . . . . . . . . . . . . . . . 3209
Pargo de Indonesia . . . . . . . . . . . . . . . . 2865 Pearl perches . . . . . . . . . . . . . . . . . . . 3205
Pargo de llama . . . . . . . . . . . . . . . . . . . 2858 Pearlscaled angelfish . . . . . . . . . . . . . . . 3279
Pargo de Madras . . . . . . . . . . . . . . . . . . 2882 Pearly monocle bream . . . . . . . . . . . . . . . 3088
Pargo de manglar . . . . . . . . . . . . . . . . . 2862 pedleyi, Paracaesio . . . . . . . . . . . . . . . . 2904
Pargo de Molucas . . . . . . . . . . . . . . . . . 2867 Pelates oxyrhynchus . . . . . . . . . . . . . . . . 3313
Pargo de Papua . . . . . . . . . . . . . . . . . . 2877 Pelates quadrilineatus . . . . . . . . . . . . . . 3311
Pargo de Randall . . . . . . . . . . . . . . . . . . 2916 Pelates sexlineatus . . . . . . . . . . . . . . . . 3312
pelewensis, Chaetodon . . . . . . . . . . . 3247-3248 Plectorhinchus celebicus . . . . . . . . . . . . . 2972

PEMPHERIDAE . . . . . . . .
. 3201, 3205, 3210-3212 Plectorhinchus chaetodonoides . . . . . . . . . 2971
Pempheris . . . . .
. . . . . .
. . . . . . . . . . 3201 Plectorhinchus chrysotaenia . . . . . . . . . . . 2972
Pempheris adusta .. . . . . .
. . . . . . . . . . 3201 Plectorhinchus cinctus . . . . . . . . . . . . . . 2978
Pempheris itoi . . .
. . . . . .
. . . . . . . . . . 3201 Plectorhinchus diagrammus . . . . . . . . . . . 2975
Pempheris mangula . . . . . .
. . . . . . . . . . 3201 Plectorhinchus fangi . . . . . . . . . . . . . . . . 2978
Pempheris molucca . . . . . .
. . . . . . . . . . 3201 Plectorhinchus flavomaculatus . . . . . . . . . 2973
Pempheris otaitensis . . . . .
. . . . . . . . . . 3201 Plectorhinchus gibbosus . . . . . . . . . . . . . 2974
Pempheris oualensis . . . . .
. . . . . . . . . . 3201 Plectorhinchus giganteus . . . . . . . . . . . . . 2970
Pempheris schwenkii . . . . .
. . . . . . . . . . 3201 Plectorhinchus goldmanni . . . . . . . . . . . . 2976
Pempheris vanicolensis .
. . .
. . . . . . . . . . 3201 Plectorhinchus harrawayi . . . . . . . . . . . . . 2970
Pennahia . . . . . . . . .
. . .
. . . . . . . . . . 3124 Plectorhinchus lessonii . . . . . . . . . . . . . . 2975
Pennahia anea . . . . .. . .
. . . . . . . . . . 3168 Plectorhinchus lineatus . . . . . . . . . . . . . . 2976
Pennahia macrocephalus . . .
. . . . . . . . . . 3169 Plectorhinchus multivittatus . . . . . . . . . . . 2977
Pennahia macrophthalmus . .
. . . . . . . . . . 3168 Plectorhinchus nigrus . . . . . . . . . . . . . . . 2974
Pennahia ovata . . . . . . . .
. . . . . . . . . . 3169 Plectorhinchus obscurum . . . . . . . . . . . . . 2970
Pennahia pawak . . . . . . . .
. . . . . . . . . . 3170 Plectorhinchus ordinalis . . . . . . . . . . . . . 2977
Pennant bannerfish . . . . . .
. . . . . . . . . . 3262 Plectorhinchus orientalis . . . . . . . . . . . . . 2982
PENTACEROTIDAE . . . . . .
. . . . . . . 3221, 3288 Plectorhinchus pictus . . . . . . . . . . . . 2967, 2978
Pentanemus . . . . . . . . . .
. . . . . . . . . . 3090 Plectorhinchus picus . . . . . . . . . . . . . . . 2979
Pentapodus . . . . . . . . . .
. . . . . . . . . . 3051 Plectorhinchus polytaenia . . . . . . . . . . . . 2980
Pentapodus bifasciatus . . . .
. . . . . . . . . . 3083 Plectorhinchus punctatissimus . . . . . . . . . . 2979
Pentapodus caninus . . . . . .
. . . . . . . . . . 3084 Plectorhinchus reticulatus . . . . . . . . . . . . 2973
Pentapodus emeryii . . . . . .
. . . . . . . . . . 3084 Plectorhinchus schotaf . . . . . . . . . . . . . . 2981
Pentapodus nagasakiensis . .
. . . . . . . . . . 3084 Plectorhinchus vittatus . . . . . . . . . . . . . . 2982
Pentapodus paradiseus . . . .
. . . . . . . . . . 3085 Plectorhynchus sinensis . . . . . . . . . . . . . . 2978
Pentapodus porosus . . . . . .
. . . . . . . . . . 3085 Plectranthias . . . . . . . . . . . . . . . . . . . 3321
Pentapodus setosus . . . . . .
. . . . . . . . . . 3075 pleurospilos, Parupeneus . . . . . . . . . . . . . 3189
Pentapodus sp. . . . . . . . .
. . . . . . . . . . 3085 pleurostigma, Parupeneus . . . . . . . . . . . . 3193
Pentapodus trivittatus . . . . .
. . . . . . . . . . 3086 pleurotaenia, Heniochus . . . . . . . . . . . . . 3263
Pentaprion . . . . . . . . . . .
. . . . . . . . . . 2946 pleurotaenia, Parupeneus . . . . . . . . . . . . . 3187
Pentaprion longimanus . . . .
. . . . . . . . . . 2958 poeti, Gerres . . . . . . . . . . . . . . . . . . . . 2957
Peppermint angelfish . . . . . .
. . . . . . . . . . 3274 Poison-fanged blenniids . . . . . . . . . . . 3087-3088
perarmatus, Otolithoides . . .
. . . . . . . . . . 3167 Poisson ange à deux épines . . . . . . . . . . . . 3273
perarmatus, Panna . . . . . .
. . . . . . . . . . 3167 Poisson ange à six bandes . . . . . . . . . . . . 3286
PERCICHTHYIDAE . . . . . .
. . . . . . . . . . 3317 Poisson ange bleu . . . . . . . . . . . . . . . . . 3285
percula, Amphiprion . . . . . . .
. . . . . . . . . . 3350 Poisson ange impérial . . . . . . . . . . . . . . . 3285
perideraion, Amphiprion . . . . .
. . . . . . . . . . 3351 Poisson ange jaune et bleu . . . . . . . . . . . . 3273
peronii, Nemipterus . . . . . . .
. . . . . . . . . . 3071 Poisson archer sellé . . . . . . . . . . . . . . . . 3215
perplexa, Filimanus . . . . . . .
. . . . . . . . . . 3100 Poisson archer tacheté . . . . . . . . . . . . . . . 3215
Peters’ monocle bream . . . .
. . . . . . . . . . 3086 Poisson-ange à trois taches . . . . . . . . . . . . 3272
petersii, Pterycombus . . . . .
. . . . . . . . . . 2835 Poisson-ange duc . . . . . . . . . . . . . . . . . 3286
pfeifferi, Polynemus . . . . . .
. . . . . . . . . . 3102 Polistonemus multifilis . . . . . . . . . . . . . . 3116
pflugeri, Mulloidichthys . . .
. . . . . . . . . . 3182 polyacanthus, Acanthochromis . . . . . . . . . 3337
Philippine butterflyfish . . . . .
. . . . . . . . . . 3234 polyactis, Cyprinocirrhitus . . . . . . . . . . . . 3321
Picnic seabream . . . . . . . .
. . . . . . . . . . 2996 Polydactylus . . . . . . . . . . . . . . . . . . . . 3090
pictum, Diagramma . . . . . .
. . . . . . . 2966-2967 Polydactylus auratus . . . . . . . . . . . . . . . 3108
pictus, Plectorhinchus . . . .
. . . . . . . 2967, 2978 Polydactylus indicus . . . . . . . . . . . . 3104-3105
picus, Plectorhinchus . . . . .
. . . . . . . . . . 2979 Polydactylus macrochir . . . . . . . . . . . 3104-3105
Pinjalo . . . . . . . . . . . . .
. . . . 2840, 2850, 2907 Polydactylus macrophthalmus . . . . . . . . . . 3106
Pinjalo lewisi . . . . . . . . .
. . . . . . . . . . 2906 Polydactylus microstoma . . . . . . . 3104, 3107, 3110
Pinjalo microphthalmus . . . .
. . . . . . . 2906-2907 Polydactylus multiradiatus . . . 3098, 3101-3102, 3108
Pinjalo pinjalo . . . . . . . . .
. . . . . . . 2906-2907 Polydactylus nigripinnnis . . . . . . . . . . . . 3109
pinjalo, Pinjalo . . . . . . . .
. . . . . . . 2906-2907 Polydactylus plebeius . . . . . . . . . . . . 3107, 3110
Pink anemonefish . . . . . . .
. . . . . . . . . . 3351 Polydactylus sexfilis . . . . . . . . . . . . . 3102, 3111
pinnulatus, Cirrhites . . . . .
. . . . . . . . . . 3326 Polydactylus sextarius . . . . 3098, 3102, 3107, 3112
pisang, Pterocaesio . . . . . .
. . . . . . . . . . 2937 Polydactylus zophomus . . . . . . . . . . . . . . 3107
Pitcairn angelfish . . . . . . . .
. . . . . . . . . . 3284 polykladiskos, Bahaba . . . . . . . . . . . . . . 3135
pixi, Cheilodactylus . . . . . .
. . . . . . . . . . 3329 polylepis, Hemitaurichthys . . . . . . . . . . . . 3261
plagiostoma, Johnius (Johnieops) . . . . . . . . 3157 POLYMIXIIDAE . . . . . . . . . . . . . . . . . . 3175
plebeius, Chaetodon . . . . . . . . . . . . . . . . 3247 See also Vol. 3
plebeius, Polydactylus . . . . . . . . . . . . 3107, 3110 polymnus, Amphiprion . . . . . . . . . . . . . . 3351
Plectorhinchus . . . . . . . . . . . . . . . . . . 2961 POLYNEMIDAE . . . . . . . . . . . . . . . . . . 3090
Plectorhinchus albovittatus . . . . . . . . . . . . 2970 Polynemus . . . . . . . . . . . . . . . . . . . . . 3090
Polynemus borneensis . . . . . .. . . . . . . . . 3115 Pseudotosciaena pamoides . . . . . . . . . . . . 3158

Polynemus dubius . . . . . . . .. . . . . . . . . 3113 Pseudupeneus . . . . . . . . . . . . . . . . . . . 3175
Polynemus hilleri . . . . .. . .. . . . . . . . . 3115 Pteraclis . . . . . . . . . .
. . . . . . . . . 2824-2825
Polynemus hornadayi . . . . . .. . . . . . . . . 3114 Pteraclis aesticola . . . . .
. . . . . . . . . . . . 2834
Polynemus intermedius . . . . .. . . . . . . . . 3103 Pteraclis velifera . . . . .. . . . . . . . . . . . 2835
Polynemus kuru . . . . . . . . .. . . . . . . . . 3111 Pterocaesio . . . . . . . . . . . . . . . . . . . . 2932
Polynemus longipectoralis . . .. . . . . . . . . 3113 Pterocaesio chrysozona . . . . . . . . . . . 2933, 2935
Polynemus melanochir . . . . .. . . . . . 3100, 3115 Pterocaesio digramma . . . . . . . . . . . 2934, 2936
Polynemus multifilis . . . . . .. . . . . . . . . 3116 Pterocaesio kohleri . . . . . . . . . . . . . . . . 2936
Polynemus paradiseusm . . . .. . . 3090, 3113-3115 Pterocaesio lativittata . . .
. . . . . . . . . . . . 2935
Polynemus pfeifferi . . . .. . .. . . . . . . . . 3102 Pterocaesio marri . . . . . . . . . . . . . . . . . 2936
Polynemus sheridani . . . . . .. . . . . . . . . 3105 Pterocaesio pisang . . . . . . . . . . . . . . . . 2937
Polynemus specularis . . . . . .. . . . . . . . . 3108 Pterocaesio randalli . . . .. . . . . . . . . . . . 2938
polystictus, Paracirrhites .
. . .. . . . . . . . . 3328 Pterocaesio sp. . . . . . . . . . 2934-2935, 2937-2938
polytaenia, Plectorhinchus . . .. . . . . . . . . 2980 Pterocaesio tessellata . . .. . . . . . . . . . . . 2939
POMACANTHIDAE . . . . . . 3225, 3266, 3287, 3289 Pterocaesio tile . . . . . .. . . . . . . . . 2926, 2940
Pomacanthus . . . . . . . . . . . . . . . . 3266, 3270 Pterocaesio trilineata . . .. . . . . . . . . . . . 2941
Pomacanthus annularis . . . . . . . . . . . . . 3284 Pterotolithus . . . . . . . .
. . . . . . . . . . . . 3124
Pomacanthus imperator . . . . . . . . . . . . . 3285 Pterotolithus lateoides . . . . . . . . . . . . . . 3172
Pomacanthus navarchus . . . . . . . . . . . . . 3285 Pterotolithus maculatus . . . . . . . . . . . . . 3173
Pomacanthus semicirculatus . . . . . . . . . . . 3285 Pterycombus . . . . . . . . . . . . . . . . . 2824-2825
Pomacanthus sextriatus . . . . . . . . . . . . . 3286 Pterycombus petersii . . . . . . . . . . . . . . . 2835
Pomacanthus xanthometapon . . . . . . . . . . 3286 Pugnose ponyfish . . . . . . . . . . . . . . . . . 2820
POMACENTRIDAE . . . . . . . . . . 3267, 3334, 3337 punctata, Drepane . . . . . . . . . . . . . 3222-3223
Pomacentrids . . . . . . . . . . . . . . . . . . . 3334 punctata, Girella . . . . . . . . . . . . . . . . . 3297
Pomacentrus alleni . . . . . . . . . . . . . . . . 3355 punctatissimus, Plectorhinchus . . . . . . . . . . 2979
Pomacentrus auriventris . . . . . . . . . . . . . 3355 punctatofasciatus, Chaetodon . . . . . . . 3247-3248
Pomacentrus coelestis . . . . . . . . . . . . . . . 3355 punctatum, Diagramma . . . . . . . . . . . . . . 2967
Pomacentrus moluccensis . . . . . . . . . . . . 3355 punctatus, Gerres . . . . . . . . . . . . . . . . . 2952
POMADASYIDAE . . . . . . . . . . . . . . . . . 2961 punctulatus, Lethrinus . . . . . . . . . . . . . . 3048
Pomadasys . . . . . . . . . . . . . . . . . . 2961, 3318 Purplemask angelfish . . . . . . . . . . . . . . . 3279
Pomadasys argenteus . . . . . . . . . . . . . . . 2983 puta, Authistes . . . .
. . . . . . . . . . . . . . . 3315
Pomadasys argyreus . . . . . . . . . . . . . . . 2984 puta, Terapon . . . . . . . . . . . . . . . . . . . 3315
Pomadasys auritus . . . . . . . . . . . . . . . . 2985 Pygmy snapper . . . . . . . . . . . . . . . . . . . 2885
Pomadasys furcatum . . . . . . . . . . . . . . . 2986 Pygoplites diacanthus .
. . . . . . . . . . . . . . 3286
Pomadasys hasta . . . . . . . . . . . . . . 2983, 2987 Pyramid butterflyfish . .
. . . . . . . . . . . . . . 3261
Pomadasys kaakan . . . . . . . . . . . . . . . . 2987 pyroferus, Acanthurus . . . . . . . . . . . 3275, 3279
Pomadasys maculatus . . . . . . . . . . . . . . . 2988
Pomadasys opercularis . . . . . . . . . . . . . . 2983
Pomadasys trifasciatus . . . . . . . . . . . . . . 2989
Q
Pomfrets . . . . . . . . . . . . . . . . . . . . . . 2824 quadrifasciatus, Coius . . . . . . . . . . . . . . 2945
Ponyfishes . . . . . . . . . . . . . . . . . . . . . 2792 quadrilineatus, Pelates . . . . . . . . . . . . . . 3311
Porgies . . . . . . . . . . . . . . . . . . . . . . . 2990 quadrimaculatus, Chaetodon . . . . . . . . . . . 3248
porosus, Pentapodus . . . . . . . . . . . . . . . 3085 quinquelineatus, Lutjanus . . . . . . . . . . . . 2888
porphyreus, Parupeneus . . . . . . . . . . . . . 3187
Premnas biaculeatus . . . . . . . . . . . . . . . 3356 R
Prickly croaker . . . . . . . . . . . . . . . . . . . 3128
Raccoon butterflyfish . . . . . . . . . . . . . . . . 3243
Prickly fanfish . . . . . . . . . . . . . . . . . . . 2835
radiosus, Etelis . . . . . . . . . . . . . . . . . . 2859
Pristipomoides . . . . . . 2851, 2909-2911, 2913-2914 rafflesii, Chaetodon . . . . . . . . . . . . . . . . 3249
Pristipomoides amoenus . . . . . . . . . . . . . . 2908 Rainford’s butterflyfish . . . . . . . . . . . . . . . 3249
Pristipomoides argyrogrammicus . . . . . . . . 2908 rainfordi, Chaetodon . . . . . . . . . . . . . . . 3249
Pristipomoides auricilla . . . . . . . . . . . . . . 2909 ramak, Sciaena . . . . . . . . . . . . . . . . . . 3038
Pristipomoides filamentosus . . . . . . . . . . . 2910 Ramball plateado . . . . . . . . . . . . . . . . . 3218
Pristipomoides flavipinnis . . . . . . . . . . . . 2911 Randall’s fusilier . . . . . . . . . . . . . . . . . . 2938
Pristipomoides microdon . . . . . . . . . . . . . 2913 Randall’s snapper . . . . . . . . . . . . . . . . . 2916
Pristipomoides microlepis . . . . . . . . . . . . . 2910 randalli, Pterocaesio . . . . . . . . . . . . . . . 2938
Pristipomoides multidens . . . . . . . . . . . . . 2912 Randallichthys . . . . . . . . . . . . . . . . . . 2840
Pristipomoides sieboldii . . . . . . . . . . . . . . 2913 Randallichthys filamentosus . . . . . . . . . . . 2916
Pristipomoides typus . . . . . . . . . . . . 2912, 2914 rapsoni, Leiognathus . . . . . . . . . . . . . . . 2814
Pristipomoides zonatus . . . . . . . . . . . . . . 2915 Rays bream . . . . . . . . . . . . . . . . . . . . 2833
Prognathodes . . . . . . . . . . . . . . . . . . . 3224 Red filament threadfin bream . . . . . . . . . . . 3079
Protonibea diacantha . . . . . . . . . . . . . . . 3171 Red goatfish . . . . . . . . . . . . . . . . . . . . 3182
Pseudanthias . . . . . . . . . . . . . . . . . . . 3338 Red pinjalo . . . . . . . . . . . . . . . . . . . . . 2906
See also Vol. 4
Red saddleback anemonefish .

Red-and-black anemonefish . .
.
.
.
.
.
.
.
.
. . . . . . 3347
. . . . . . 3349
S
Redaxil emperor . . . . . . . . . . . . . . . . . . 3028 Saddle grunt . . . . . . . . . . . . . . . . . . . . 2988
Redbelly yellowtail fusilier . . . . . . . . . . . . . 2927 Saddleback anemonefish . . . . . . . . . . . . . 3351
Redsnout emperor . . . . . . . . . . . . . . . . . 3041 Saddleback butterflyfish . . . . . . . . . . . . . . 3240
Redspine threadfin bream . . . . . . . . . . . . . 3070 Saddleback snapper . . . . . . . . . . . . . . . . 2900
Redspot dwarf monacle bream . . . . . . . . . . 3082 Sailfin armourhead . . . . . . . . . . . . . . . . . 3288
Redspot emperor . . . . . . . . . . . . . . . . . . 3034 Sailfin snapper . . . . . . . . . . . . . . . . . . . 2917
Reeve’s croaker . . . . . . . . . . . . . . . . . . 3137 Salmon australien . . . . . . . . . . . . . . . . . 3304
Regal angelfish . . . . . . . . . . . . . . . . . . . 3286 Salmón de Australia . . . . . . . . . . . . . . . . 3304
regina, Scolopsis . . . . . . . . . . . . . . . . . . 3076 Salmonete . . . . . . . . . . . . . . . . . . . . . 3185
rendalli, Tilapia . . . . . . . . . . . . . . . . . . 3333 Salmonete aurora . . . . . . . . . . . . . . . . . 3197
Reticulated butterflyfish . . . . . . . . . . . . . . 3250 Salmonete bandón . . . . . . . . . . . . . . . . . 3191
reticulatus, Chaetodon . . . . . . . . . . . 3244, 3250 Salmonete barborín . . . . . . . . . . . . . . . . 3199
reticulatus, Lethrinus . . . . . . . . . 3041, 3044, 3047 Salmonete de banda amarilla . . . . . . . . . . . 3181
reticulatus, Plectorhinchus . . . . . . . . . . . . 2973 Salmonete de banda ocre . . . . . . . . . . . . . 3198
Rhabdosargus sarba . . . . . . . . . . . . . . . 3003 Salmonete de bande dorada . . . . . . . . . . . . 3196
rhodopterus, Lethrinus . . . . . . . . . . . . . . 3032 Salmonete de dos manchas . . . . . . . . . . . . 3186
Rhonciscus furcatus . . . . . . . . . . . . . . . . 2986 Salmonete dorado . . . . . . . . . . . . . . . . . 3188
Rhyncopelates oxyrhynchus . . . . . . . . . . . 3313 Salmonete índico . . . . . . . . . . . . . . . . . . 3190
Ribboned sweetlips . . . . . . . . . . . . . . . . 2980 Salmonete naranjero . . . . . . . . . . . . . . . . 3200
River threadfin . . . . . . . . . . . . . . . . . . . 3106 Salmonete pastilla . . . . . . . . . . . . . . . . . 3193
rivulatus, Lutjanus . . . . . . . . . . . . . . . . 2889 Salmonete vanicolense . . . . . . . . . . . . . . 3183
Roa . . . . . . . . . . . . . . . . . . . . . . . . . 3224 Samoan snapper . . . . . . . . . . . . . . . . . . 2886
Roaops . . . . . . . . . . . . . . . . . 3224, 3239, 3252 samoensis, Mulloidichthys . . . . . . . . . . . . 3181
robinsoni, Gymnocranius . . . . . . . . . . . . . 3022 sandaracinos, Amphiprion . . . . . . . . . . . . 3352
Rock flagtail . . . . . . . . . . . . . . . . . . . . 3320 Sapsap à deux taches . . . . . . . . . . . . . . . 2803
Ronco de seis bandas . . . . . . . . . . . . . . . 2986 Sapsap berbère . . . . . . . . . . . . . . . . . . 2801
Rooseveltia brighami . . . . . . . . . . . . . . . 2915 Sapsap bouledogue . . . . . . . . . . . . . . . . 2822
rostratus, Chelmon . . . . . . . . . . . . . . . . 3258 Sapsap boxeur . . . . . . . . . . . . . . . . . . . 2820
Rosy dwarf monocle bream . . . . . . . . . . . . 3081 Sapsap commun . . . . . . . . . . . . . . . . . . 2808
Rouget à trois bandes . . . . . . . . . . . . . . . 3192 Sapsap dentu . . . . . . . . . . . . . . . . . . . 2799
Rouget-barbé badeau . . . . . . . . . . . . . . . 3191 Sapsap doré . . . . . . . . . . . . . . . . . . . . 2804
Rouget-barbet à tache noire . . . . . . . . . . . . 3194 Sapsap écriture . . . . . . . . . . . . . . . . . . 2817
Rouget-barbet barberin . . . . . . . . . . . . . . 3185 Sapsap fil . . . . . . . . . . . . . . . . . . . . . . 2810
Rouget-barbet doré . . . . . . . . . . . . . . . . 3188 Sapsap nu . . . . . . . . . . . . . . . . . . . . . 2798
Rouget-barbet double-tauche . . . . . . . . . . . 3186 Sapsap panache . . . . . . . . . . . . . . . . . . 2811
Rouget-barbet indien . . . . . . . . . . . . . . . . 3190 Sapsap rayé . . . . . . . . . . . . . . . . . . . . 2809
Rouget-barbet pastille . . . . . . . . . . . . . . . 3193 Sapsap soleil . . . . . . . . . . . . . . . . . . . . 2815
Rouget-souris à bande sombre . . . . . . . . . . 3199 Sapsap vaguelette . . . . . . . . . . . . . . . . . 2806
Rouget-souris aurore . . . . . . . . . . . . . . . . 3197 Sapsap voile orange . . . . . . . . . . . . . . . . 2802
Rouget-souris bande or . . . . . . . . . . . . . . 3196 sarba, Rhabdosargus . . . . . . . . . . . . . . . 3003
Rouget-souris ocre . . . . . . . . . . . . . . . . . 3198 sarba, Sparus . . . . . . . . . . . . . . . . . . . 3003
Rouget-souris orangé . . . . . . . . . . . . . . . 3200 Sargo aleta amarilla . . . . . . . . . . . . . . . . 2997
Rovers . . . . . . . . . . . . . . . . . . . . . . . 2838 Sargo dorado . . . . . . . . . . . . . . . . . . . . 3003
Rubberlips . . . . . . . . . . . . . . . . . . . . . 2961 Sargo picnic . . . . . . . . . . . . . . . . . . . . 2996
ruber, Otolithes . . . . . . . . . . . . . . . . . . 3164 Sargo real . . . . . . . . . . . . . . . . . . . . . 2998
rubescens, Parupeneus . . . . . . . . . . . . . . 3187 Sargue doré . . . . . . . . . . . . . . . . . . . . 3003
rubescens, Taractes . . . . . . . . . . . . . . . . 2836 Sawjawed monocle bream . . . . . . . . . . . . . 3087
rubrioperculatus, Lethrinus . . . . . . . . . . . 3042 Scaevius milii . . . . . . . . . . . . . . . . 3051, 3086
Ruby snapper . . . . . . . . . . . . . . . . . . . 2857 scalaripinnis, Schuettea . . . . . . . . . . . . . 3220
Rubyfishes . . . . . . . . . . . . . . . . . . . . . 2838 Scale croaker . . . . . . . . . . . . . . . . . . . . 3163
ruconius, Secutor . . . . . . . . . . . . . . 2821-2822 Scalemouth jobfish . . . . . . . . . . . . . . . . . 2905
rufolineatus, Lutjanus . . . . . . . . . . . 2867, 2890 Scalynosed silverbiddy . . . . . . . . . . . . . . . 2959
rufomaculatus, Parascolopsis . . . . . . . . . . 3082 scapulare, Glaucosoma . . . . . . . . . . . . . . 3209
rupestris, Kuhlia . . . . . . . . . . . . . . . . . 3320 Scarlet snapper . . . . . . . . . . . . . . . . . . 2859
Russell’s snapper . . . . . . . . . . . . . . . . . 2891 SCATOPHAGIDAE 3217, 3221, 3225, 3267, 3287, 3289
russelli, Dendrophysa . . . . . . . . . . . . 3138-3139 schlegeli, Erythrocles . . . . . . . . . . . . . . . 2838
russelli, Lutjanus . . . . . . . . . . . . . . . . . 2891 schotaf, Plectorhinchus . . . . . . . . . . . . . . 2981
russelli, Sciaena . . . . . . . . . . . . . . . . . . 3138 Schuetta . . . . . . . . . . . . . . . . . . . . . . 3216
Rusty jobfish . . . . . . . . . . . . . . . . . . . . 2855 Schuettea scalaripinnis . . . . . . . . . . . . . . 3220
rutilans, Aphareus . . . . . . . . . . . . . . . . 2855 schwenkii, Pempheris . . . . . . . . . . . . . . . 3201
Sciaena antarctica . . . . . . . . . . . . . . . . . 3127
Sciaena macroptera . . . . . . . . . . . . . . . . 3149
Sciaena ramak . . . . . . . . . . . . . . . . . . . 3038 Shining ponyfish . . . . . . . . . . . . . . . . . . 2813

Sciaena russelli . . . . . . . . . . . . . . . . . . 3138 Shortfin pomfret . . . . . . . . . . . . . . . . . . 2834
SCIAENIDAE . . . . . . . . . . . . . . . . . 3091, 3117 siamensis, Otolithoides . . . . . . . . . . . . . . 3151
scintillans, Erythrocles . . . . . . . . . . . . . . 2838 siamensis, Scolopsis . . . . . . . . . . . . . . . . 3077
Scolopsis . . . . . . . . . . . . . . . . 3051-3052, 3077 sibogae, Sphenanthias . . . . . . . . . . . 3331-3332
Scolopsis à trois lignes
. .
. . . . . . . . . . . . . 3089 Sickle pomfret . . . . . . . . . . . . . . . . . . . 2836
Scolopsis affinis . . .. .
. . . . . . . . . . . . . 3086 Sicklefish . . . . . . . . . . . . . . . . . . . . . . 3222
Scolopsis auratus . . . .
. . . . . . . . . . . . . 3087 Sicklefishes . . . . . . . . . . . . . . . . . . . . . 3221
Scolopsis bilineatus . . .
. . . . . . . . . . . . . 3087 Sidespot goatfish . . . . . . . . . . . . . . . . . . 3193
Scolopsis cancellatus . .
. . . . . . . . . . . . . 3077 sieboldii, Pristipomoides . . . . . . . . . . . . . 2913
Scolopsis ciliatus . . . .
. . . . . . . . . . . . . 3087 signatus, Parupeneus . . . . . . . . . . . . . . . 3194
Scolopsis lineatus . . . .
. . . . . . . . . . 3077, 3088 SILLAGINIDAE . . . . . . . . . . . . . . . . . . 3118
Scolopsis margaritifer .
. . . . . . . . . . . . . 3088 See also Vol. 4
Scolopsis monogramma . . . . . . . . . . . . . 3076
. Silver drummer . . . . . . . . . . . . . . . . . . . 3294
Scolopsis rayé . . . . . . . . . . . . . . . . . . 3088
. Silver flagtail . . . . . . . . . . . . . . . . . . . . 3320
Scolopsis regina . . . . . . . . . . . . . . . . . 3076
. Silver grunt . . . . . . . . . . . . . . . . . . . . . 2983
Scolopsis siamensis . . . . . . . . . . . . . . . 3077
. Silver moony . . . . . . . . . . . . . . . . . . . . 3218
Scolopsis taeniopterus .
. . . . . . . . . . . . . 3077 Silver seabream . . . . . . . . . . . . . . . . . . 3002
Scolopsis temporalis . . . . . . . . . . . 3076, 3088
. Silver sweep . . . . . . . . . . . . . . . . . . . . 3299
Scolopsis trilineatus . . . . . . . . . . . . . . . 3089
. Silver terapon . . . . . . . . . . . . . . . . . . . 3309
Scolopsis vosmeri . . . . . . . . . . . . . . . . 3089
. Silverbiddies . . . . . . . . . . . . . . . . . . . . 2946
Scolopsis xenochrous . . . . . . . . . . . . . . 3089
. similis, Filimanus . . . . . . . . . . . . . . 3098, 3102
Scomberoides . . . . . . . . . . . . . . . . . . 3210
. sina, Johnius . . . . . . . . . . . . . . . . . . . 3154
Scorched croaker . . . . . . . . . . . . . . . . 3130
. sinclairi, Chrysiptera . . . . . . . . . . . . . . . 3353
SCORPIDIDAE . . . . . .3290, 3297, 3299, 3301-3302 sinensis, Plectorhynchus . . . . . . . . . . . . . 2978
Scorpis aequipinnis . . . . . . . . . . . . . . . 3299
. Singular bannerfish . . . . . . . . . . . . . . . . 3264
Scorpis lineolata . . . . . . . . . . . . . . . . . 3299
. singularis, Heniochus . . . . . . . . . . . . . . . 3264
Scorpis lineolatus . . . . . . . . . . . . . . . . 3299
. Sixbanded angelfish . . . . . . . . . . . . . . . . 3286
Scribbled angelfish . . . . . . . . . . . . . . . . 3280
. Sixfinger threadfin . . . . . . . . . . . . . . . . . 3111
Sea chubs . . . . . . . . . . . . . . . . . . . . 3290
. Sixlined terapon . . . . . . . . . . . . . . . . . . 3312
Seabreams . . . . . . . . . . . . . . . . . . . . 2990
. Skunk anemonefish . . . . . . . . . . . . . . . . 3346
sealei, Filimanus . . . . . . . . . . . . . . . . 3101
. Slender emperor . . . . . . . . . . . . . . . . . . 3044
sebae, Lutjanus . . . . . . . . . . . . . . 2884, 2892
. Slender fusilier . . . . . . . . . . . . . . . . . . . 2932
Sectator ocyurus . . . . . . . . . . . . . . . . . 3296
. Slender ponyfish . . . . . . . . . . . . . . . . . . 2816
Secutor hanedai . . . . . . . . . . . . . . . . . 2818
. Slender silverbiddy . . . . . . . . . . . . . . . . . 2955
Secutor indicius . . . . . . . . . . . . . . . . . 2819
. Slender threadfin bream . . . . . . . . . . . . . . 3081
Secutor insidiator . . . . . . . . . . . . . 2818-2820
. Slipmouths . . . . . . . . . . . . . . . . . . . . . 2792
Secutor interruptus . . . . . . . . . . . . . . . 2822
. Smalljaw croaker . . . . . . . . . . . . . . . . . . 3160
Secutor megalolepis . . . . . . . . . . . . . . . 2821
. Smallmouth croaker . . . . . . . . . . . . . . . . 3146
Secutor ruconius . . . . . . . . . . . . . 2821-2822
. Smallmouth threadfin . . . . . . . . . . . . . . . 3107
Secutor spp. . . . . . . . . . . . . . . . . . . . 2792
. Smallscale croaker . . . . . . . . . . . . . . . . . 3159
selene, Chaetodon . . . . . . . . . . . . . . . . 3250
. Smallscaled terapon . . . . . . . . . . . . . . . . 3315
semicinctus, Lethrinus .
. . . . . . . 3043-3044, 3047 Smalltooth emperor . . . . . . . . . . . . . . . . 3035
semicinctus, Lutjanus . . . . . . . . . . . . . . 2893
. Smalltoothed jobfish . . . . . . . . . . . . . . . . 2854
semicirculatus, Pomacanthus . . . . . . . . . . 3285 Smalltoothed whiptail . . . . . . . . . . . . . . . 3084
semifasciata, Nibea . . . . . . . . . . . . . 3161, 3163 Smith’s butterflyfish . . . . . . . . . . . . . . . . 3251
semion, Chaetodon . . . . . . . . . . . . . . . . 3251 Smithhurst’s ponyfish . . . . . . . . . . . . . . . 2811
septentrionalis, Chaetodontoplus . . . . . 3280, 3282 smithi, Chaetodon . . . . . . . . . . . . . . . . . 3251
SERRANIDAE . . . . . . 2942, 2991, 3305, 3321, 3338 smithursti, Leiognathus . . . . . . . . . . . . . . 2811
See also Vol. 4 Smooth croaker . . . . . . . . . . . . . . . . . . 3147
servus, Therapon . . . . . . . . . . . . . . . . . 3314 Snapper . . . . . . . . . . . . . . . . . . . . . . 3183
setosus, Pentapodus . . . . . . . . . . . . . . . . 3075 Snappers . . . . . . . . . . . . . . . . . . . . . . 2840
Sevenfinger threadfin . . . . . . . . . . . . . . . 3098 soldado, Nibea . . . . . . . . . 3136, 3156, 3159, 3162
sexfilis, Polydactylus . . . . . . . . . . . . 3102, 3111 Soldier croaker . . . . . . . . . . . . . . . . . . . 3162
sexlineatus, Helotes . . . . . . . . . . . . . . . . 3312 Solosolo de banda negra . . . . . . . . . . . . . 3327
sexlineatus, Pelates . . . . . . . . . . . . . . . . 3312 Solosolo robusto . . . . . . . . . . . . . . . . . . 3326
sextarius, Polydactylus . . . . 3098, 3102, 3107, 3112 Sonorolux fluminis . . . . . . . . . . . . . . . . 3174
sextriatus, Pomacanthus . . . . . . . . . . . . . 3286 sordida, Paracaesio . . . . . . . . . . . . . . . . 2902
Sharpnose croaker . . . . . . . . . . . . . . . . . 3161 Southern fanfish . . . . . . . . . . . . . . . . . . 2835
Sharptooth hammer croaker . . . . . . . . . . . . 3154 Southern rays bream . . . . . . . . . . . . . . . . 2833
Sharptooth jobfish . . . . . . . . . . . . . . . . . 2914 Southern silverbiddy . . . . . . . . . . . . . . . . 2960
Shepard’s angelfish . . . . . . . . . . . . . . . . 3278 Southern yellow croaker . . . . . . . . . . . . . . 3158
shepardi, Centropyge . . . . . . . . . . . . . . . 3278 Spangled emperor . . . . . . . . . . . . . . . . . 3037
sheridani, Polynemus . . . . . . . . . . . . . . . 3105 Spanish flag snapper . . . . . . . . . . . . . . . . 2868
Spare royal . . . . . .
. . . . . . . . . . . . . . . 2998 Sweetlips . . . . . . . . . . . . . . . . . . . . . . 2961
SPARIDAE . . . . . . . . 2841, 2946, 2962, 2990, 3005 sydneyanus, Kyphosus . . . . . . . . . . . . . . 3294
spariformis, Dentex . . . . . . . . . . . . . . . . 3000 Symphorichthys spilurus . . . . . . . . . . 2841, 2917
Sparus latus . . . . .. . . . . . . . . . . . . . . 2997 Symphorus nematophorus . . . . . . . . . . . . 2918
Sparus major . . . . .. . . . . . . . . . . . . . . 3002
Sparus sarba . . . . .
Speckled butterflyfish
.
.
.
.
. . . . . . . . . . . . . 3003
. . . . . . . . . . . . . 3238
T
specularis, Polynemus . . . . . . . . . . . . . . . 3108 taeniatus, Erythrocles . . . . . . . . . . . . . . . 2838
speculum, Chaetodon . . . . . . . . . . . . . . . 3252 taeniatus, Glaucosoma . . . . . . . . . . . . . . 3207
Sphenanthias . . . . . . . . . . . . . . . . . . . 3332 taeniopterus, Scolopsis . . . . . . . . . . . . . . 3077
Sphenanthias macrophthalmus . . . . . . . . . 3331 taeniopterus, Upeneus . . . . . . . . . . . . . . 3195
Sphenanthias sibogae . . . . . . . . . . . . 3331-3332 taeniura, Kuhlia . . . . . . . . . . . . . . . . . . 3319
spilotoceps, Cirrhitus . . . . . . . . . . . . . . . 3326 Tahiti butterflyfish . . . . . . . . . . . . . . . . . 3253
spilurus, Parupeneus . . . . . . . . . . . . . . . 3194 Taius tumifrons . . . . . . . . . . . . . . . . . . 3000
spilurus, Symphorichthys . . . . . . . . . . 2841, 2917 tambuloides, Nemipterus . . . . . . . . . . . . . 3072
Spine bahaba . . . . . . . . . . . . . . . . . . . 3135 Tang’s snapper . . . . . . . . . . . . . . . . . . . 2860
Spinecheek anemonefish . . . . . . . . . . . . . 3356 tanyactis, Parascolopsis . . . . . . . . . . . . . . 3083
spinifer, Argyrops . . . . . . . . . . . . . . . . . 2998 Tapiroid terapon . . . . . . . . . . . . . . . . . . 3310
spinifer, Histiopterus . . . . . . . . . . . . . . . 3288 Taractes . . . . . . . . . . . . . . . . . . . . . . 2824
spinus, Genicanthus . . . . . . . . . . . . . . . 3284 Taractes asper . . . . . . . . . . . . . . . . . . . 2835
splendens, Leiognathus . . . . . . . . . . . 2802, 2815 Taractes rubescens . . . . . . . . . . . . . . . . 2836
Splendid ponyfish . . . . . . . . . . . . . . . . . 2815 Taractichthys longipinnis . . . . . . . . . . . . 2836
Splendid threadfin . . . . . . . . . . . . . . . . . 3100 Taractichthys steindachneri . . . . . . . . . . . 2836
Spotbanded butterflyfish . . . . . . . . . . . . . . 3248 taupou, Chrysiptera . . . . . . . . . . . . . . . . 3353
Spotcheek emperor . . . . . . . . . . . . . . . . 3042 Teardrop butterflyfish . . . . . . . . . . . . . . . 3255
Spottail butterflyfish . . . . . . . . . . . . . . . . 3245 Teardrop threadfin bream . . . . . . . . . . . . . 3067
Spotted archerfish . . . . . . . . . . . . . . . . . 3215 temporalis, Scolopsis . . . . . . . . . . . . 3076, 3088
Spotted butterflyfish . . . . . . . . . . . . . . . . 3241 Tenspined silverbiddy . . . . . . . . . . . . . . . 2959
Spotted croaker . . . . . . . . . . . . . . . . . . 3171 Téraglin . . . . . . . . . . . . . . . . . . . . . . . 3129
Spotted flagtail . . . . . . . . . . . . . . . . . . . 3319 Terapon cancellatus . . . . . . . . . . . . . . . . 3310
Spotted sicklefish . . . . . . . . . . . . . . . . . . 3223 Terapon caudavittatus . . . . . . . . . . . . . . . 3308
Spotted sweetlips . . . . . . . . . . . . . . . . . 2979 Terapon jarbua . . . . . . . . . . . . . . . . . . 3314
springeri, Chrysiptera . . . . . . . . . . . . . . . 3353 Terapon oxyrhynchus . . . . . . . . . . . . . . . 3313
squamimaxillaris, Parapristipomoides . . . . . . 2905 Terapon puta . . . . . . . . . . . . . . . . . . . 3315
squamosa, Nibea . . . . . . . . . . . . . . . . . 3163 Terapon theraps . . . . . . . . . . . . . . . . . . 3316
Star snapper . . . . . . . . . . . . . . . . . . . . 2894 Terapon-grunters . . . . . . . . . . . . . . . . . . 3305
steindachneri, Taractichthys . . . . . . . . . . . 2836 Terapon-perches . . . . . . . . . . . . . . . . . . 3305
stellatus, Lutjanus . . . . . . . . . . . . . . . . . 2894 TERAPONIDAE . . . . . . . . . . . . . . . . . . 3305
stercorarius, Leiognathus . . . . . . . . . . . . 2816 TERAPONTIDAE . . . . . . . . . . . . . . 3305, 3317
Stocky hawkfish . . . . . . . . . . . . . . . . . . 3326 teres, Caesio . . . . . . . . . . . . . . . . . 2929-2930
stonei, Paracaesio . . . . . . . . . . . . . . . . . 2903 tessellata, Pterocaesio . . . . . . . . . . . . . . . 2939
striata, Caesio . . . . . . . . . . . . . . . . . . . 2926 tetradactylum, Eleutheronema . . . . 3096-3097, 3105
strigatus, Atypichthys . . . . . . . . . . . . . . . 3303 Theodore’s threadfin bream . . . . . . . . . . . . 3080
strigatus, Microcanthus . . . . . . . . . . . 3225, 3303 theodorei, Nemipterus . . . . . . . . . . . . . . 3080
Striped goatfish . . . . . . . . . . . . . . . . . . . 3200 Therapon servus . . . . . . . . . . . . . . . . . . 3314
Striped large-eye bream . . . . . . . . . . . . . . 3017 THERAPONIDAE . . . . . . . . . . . . . . . . . 3305
Striped monocle bream . . . . . . . . . . . . . . 3088 THERAPONTIDAE . . . . . . . . . . . . . . . . . 3305
Striped ponyfish . . . . . . . . . . . . . . . . . . 2809 theraps, Eutherapon . . . . . . . . . . . . . . . . 3316
Striped velvetchin . . . . . . . . . . . . . . . . . 2969 theraps, Terapon . . . . . . . . . . . . . . . . . 3316
Stripey . . . . . . . . . . . . . . . . . . . . . . . 3303 thosaporni, Nemipterus . . . . . . . . . . . . . . 3073
STROMATEIDAE . . . . . . . . . . . . . . . . . 2825 Threadfin breams . . . . . . . . . . . . . . . . . 3051
See also Vol. 6 Threadfin butterflyfish . . . . . . . . . . . . . . . 3236
Strongspine silverbiddy . . . . . . . . . . . . . . 2957 Threadfin pearl perch . . . . . . . . . . . . . . . 3208
strusakeri, Emmelichthys . . . . . . . . . . . . . 2838 Threadfins . . . . . . . . . . . . . . . . . . . . . 3090
subfasciatus, Gerres . . . . . . . . . . . . . . . . 2960 Threefinger threadfin . . . . . . . . . . . . . . . . 3097
Sulphur goatfish . . . . . . . . . . . . . . . . . . 3197 Threelined monocle bream . . . . . . . . . . . . 3089
sulphureus, Upeneus . . . . . . . . . . . . . . . 3197 Threespot angelfish . . . . . . . . . . . . . . . . 3272
Sunbeam snapper . . . . . . . . . . . . . . . . . 2870 Threespot dascyllus . . . . . . . . . . . . . . . . 3354
sundaicus, Upeneus . . . . . . . . . . . . . . . . 3198 Threestripe fusilier . . . . . . . . . . . . . . . . . 2941
suratensis, Etroplus . . . . . . . . . . . . . . . . 3333 Threestriped whiptail . . . . . . . . . . . . . . . . 3086
surinamensis, Lobotes . . . . . . . . . . . . . . 2944 Thumbprint emperor . . . . . . . . . . . . . . . . 3032
Surmullets . . . . . . . . . . . . . . . . . . . . . 3175 Thunnus albacares . . . . . . . . . . . . . . . . 2832
Swallowtail angelfish . . . . . . . . . . . . . . . . 3283 tibicen, Centropyge . . . . . . . . . . . . . . . . 3278
Sweepers . . . . . . . . . . . . . . . . . . . . . . 3201 Tigertooth croaker . . . . . . . . . . . . . . . . . 3164
Tilapia rendalli . . . . . . . . . . . . . . . . . . 3333 Upeneus moluccensis . . . . . . . . . . . . . . . 3196

Tilapia zilli . . . . . . . . . . . . . . . . . . . . . 3333 Upeneus oligospilus . . . . . . . . . . . . . . . . 3199
tile, Pterocaesio . . . . . . . . . . . . . . . 2926, 2940 Upeneus sulphureus . . . . . . . . . . . . . . . 3197
Timor snapper . . . . . . . . . . . . . . . . . . . 2895 Upeneus sundaicus . . . . . . . . . . . . . . . . 3198
timorensis, Lutjanus . . . . . . . . . . . . . . . 2895 Upeneus taeniopterus . . . . . . . . . . . . . . . 3195
Tinker’s butterflyfish . . . . . . . . . . . . . . . . 3252 Upeneus tragula . . . . . . . . . . . . . . . . . . 3199
tinkeri, Chaetodon . . . . . . . . . . . . . . . . 3252 Upeneus vittatus . . . . . . . . . . . . . . . . . . 3200
tolu, Nemipterus . . . . . . . . . . . . . . . . . . 3071 urolepis, Oreochromis . . . . . . . . . . . . . . 3333
Tomato anemonefish . . . . . . . . . . . . . . . . 3348
Toothpony . . . . . . . . .
Toothy . . . . . . . . . . .
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
. . 2799
. . 2901
V
Tosa dwarf monocle bream . . . . . . . . . . . . 3083 Vagabond butterflyfish . . . . . . . . . . . . . . . 3255
tosensis, Parascolopsis . . . . . . . . . . . . . . 3083 vagabundus, Chaetodon . . . . . . . . . . . . . 3255
Toxotes chatareus . . . . . . . . . . . . . . . . . 3215 vaigiensis, Kyphosus . . . . . . . . . . . . . . . 3295
Toxotes jaculatrix . . . . . . . . . . . . . . . . . 3215 vanicolensis, Mulloides . . . . . . . . . . . . . . 3183
TOXOTIDAE . . . . . . . . . . . . . . . . . . . . 3212 vanicolensis, Mulloidichthys . . . . . . . . . . . 3183
trachycephalus, Johnius (Johnius) . . . . . . . 3151 vanicolensis, Pempheris . . . . . . . . . . . . . 3201
tragula, Upeneus . . . . . . . . . . . . . . . . . 3199 Vanuatu snapper . . . . . . . . . . . . . . . . . . 2899
Trewavas’ croaker . . . . . . . . . . . . . . . . . 3152 variegatus, Lethrinus . . . . . . 3042-3044-3045, 3047
trewavasae, Johnius (Johnius) . . . . . . . . . . 3152 varilineata, Caesio . . . . . . . . . . . . . . . . 2926
trichrous, Chaetodon . . . . . . . . . . . . . . . 3253 varius, Heniochus . . . . . . . . . . . . . . . . . 3264
tricuspidata, Girella . . . . . . . . . . . . . . . . 3298 velifera, Pteraclis . . . . . . . . . . . . . . . . . 2835
tridactylum, Eleutheronema . . . . . . . . 3096-3097 Velvetchins . . . . . . . . . . . . . . . . . . . . . 2961
trifascialis, Chaetodon . . . . . . . . . . . . . . 3253 venenatus, Paradicichthys . . . . . . . . . . . . . 2918
trifasciatus, Chaetodon . . . . . . . . . . . . . . 3254 venustus, Centropyge . . . . . . . . . . . . . . . 3279
trifasciatus, Parupeneus . . . . . . . . . . 3186, 3192 verekeri, Parapolynemus . . . . . . . . . . . . . 3103
trifasciatus, Pomadasys . . . . . . . . . . . . . . 2989 Vermiculated angelfish . . . . . . . . . . . . . . . 3281
TRIGLIDAE . . . . . . . . . . . . . . . . . . . . 3091 Vermiculated ponyfish . . . . . . . . . . . . . . . 2823
See also Vol. 4 Verrue bronzée . . . . . . . . . . . . . . . . . . . 3165
trilineata, Pterocaesio . . . . . . . . . . . . . . . 2941 Violon crépuscule . . . . . . . . . . . . . . . . . 3311
trilineatus, Scolopsis . . . . . . . . . . . . . . . 3089 Violon grogneur . . . . . . . . . . . . . . . . . . 3315
trimaculatus, Apolemichthys . . . . . . . . . . . 3272 Violon jarbua . . . . . . . . . . . . . . . . . . . . 3314
trimaculatus, Dascyllus . . . . . . . . . . . . . . 3354 Violon tigre . . . . . . . . . . . . . . . . . . . . . 3316
Tripletail . . . . . . . . . . . . . . . . . . . . . . 2944 virescens, Aprion . . . . . . . . . . . . . . . . . 2856
Tripletails . . . . . . . . . . . . . . . . . . . . . . 2942 virgatus, Leiognathus . . . . . . . . . . . . . . . 2802
tristis, Acanthurus . . . . . . . . . . . . . . . . . 3274 virgatus, Nemipterus . . . . . . . . . 3005, 3051, 3074
trivittatus, Pentapodus . . . . . . . . . . . . . . 3086 viridis, Chromis . . . . . . . . . . . . . . . . . . 3352
Trout sweetlips . . . . . . . . . . . . . . . . . . . 2978 vitiensis, Nemipterus . . . . . . . . . . . . . . . 3081
Trumpet emperor . . . . . . . . . . . . . . . . . . 3036 vitta, Lutjanus . . . . . . . . . . . . . . . . . . . 2896
trutta, Arripis . . . . . . . . . . . . . . . . . . . 3304 vittatus, Goniistius . . . . . . . . . . . . . . . . 3329
tumida, Paracaesio . . . . . . . . . . . . . . . . 2904 vittatus, Plectorhinchus . . . . . . . . . . . . . . 2982
tumifrons, Dentex . . . . . . . . . . . . . . . . . 3000 vittatus, Upeneus . . . . . . . . . . . . . . . . . 3200
tumifrons, Taius . . . . . . . . . . . . . . . . . . 3000 Vivaneau à bande blanche . . . . . . . . . . . . . 2864
Two-eyed coralfish . . . . . . . . . . . . . . . . . 3259 Vivaneau à bande brune . . . . . . . . . . . . . . 2896
Twoblotch ponyfish . . . . . . . . . . . . . . . . . 2803 Vivaneau à bandes noires . . . . . . . . . . . . . 2893
Twolined monocle bream . . . . . . . . . . . . . 3087 Vivaneau à cinq bandes . . . . . . . . . . . . . . 2888
Twospot banded snapper . . . . . . . . . . . . . 2864 Vivaneau à raies bleues . . . . . . . . . . . . . . 2879
Twospot red snapper . . . . . . . . . . . . . . . . 2866 Vivaneau à raies jaunes . . . . . . . . . . . . . . 2880
typee, Paracirrhites . . . . . . . . . . . . . . . . 3327 Vivaneau bourgeois . . . . . . . . . . . . . . . . 2892
typus, Histiopterus . . . . . . . . . . . . . . . . 3288 Vivaneau chin rouge . . . . . . . . . . . . . . . . 2866
typus, Macolor . . . . . . . . . . . . . . . . . . . 2898 Vivaneau cramoisi . . . . . . . . . . . . . . . . . 2872
typus, Oxycirrhites . . . . . . . . . . . . . . . . 3321 Vivaneau d’eau douce . . . . . . . . . . . . . . . 2875
typus, Pristipomoides . . . . . . . . . . . . 2912, 2914 Vivaneau damier . . . . . . . . . . . . . . . . . . 2869
Vivaneau de l’Indonesia . . . . . . . . . . . . . . 2865
Vivaneau de nuit . . . . . . . . . . . . . . . . . . 2861
U Vivaneau de Papua . . . . . . . . . . . . . . . . 2877
ulietensis, Chaetodon . . . . . . . . . . . . . . . 3254 Vivaneau de Randall . . . . . . . . . . . . . . . . 2916
Unarmed dwarf monocle bream . . . . . . . . . . 3082 Vivaneau de Samoa . . . . . . . . . . . . . . . . 2886
undecimradiatus, Coius . . . . . . . . . . . . . . 2942 Vivaneau de Timor . . . . . . . . . . . . . . . . . 2895
unimaculatus, Chaetodon . . . . . . . . . . . . 3255 Vivaneau des mangroves . . . . . . . . . . . . . 2862
Upeneus . . . . . . . . . . . . . . . . . . . 3175, 3178 Vivaneau des Moluques . . . . . . . . . . . . . . 2867
Upeneus arge . . . . . . . . . . . . . . . . . . . 3195 Vivaneau diable . . . . . . . . . . . . . . . . . . 2918
Upeneus doriae . . . . . . . . . . . . . . . . . . 3197 Vivaneau drapeau . . . . . . . . . . . . . . . . . 2868
Upeneus luzonius . . . . . . . . . . . . . . . . . 3198 Vivaneau du Bengale . . . . . . . . . . . . . . . 2863
Vivaneau écarlate . . . . . . . . . . . . . . . . . 2859 Wideband fusilier . . . . . . . . . . . . . . . . . . 2935

Vivaneau eglefin . . . . . . . . . . . . . . . . . . 2887 Wiebel’s butterflyfish . . . . . . . . . . . . . . . . 3256
Vivaneau encrier . . . . . . . . . . . . . . . . . . 2871 wiebeli, Chaetodon . . . . . . . . . . . . . . . . 3256
Vivaneau étoié . . . . . . . . . . . . . . . . . . . 2894
Vivaneau gibelot . . .
Vivaneau gros yeux .
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2873
2882
X
Vivaneau hublot . . . . . . . . . . . . . . . . . . 2891 xanthochilus, Lethrinus . . . . . . . . . . . . . 3045
Vivaneau job . . . . . . . . . . . . . . . . . . . . 2856 xanthometapon, Pomacanthus . . . . . . . . . . 3286
Vivaneau la flamme . . . . . . . . . . . . . . . . 2858 xanthonema, Filimanus . . . . . . . . . . 3102, 3111
Vivaneau madras . . . . . . . . . . . . . . . . . . 2883 xanthonota, Caesio . . . . . . . . . . . . . 2929-2930
Vivaneau malabar . . . . . . . . . . . . . . . . . 2884 xanthopunctatus, Apolemichthys . . . . . . . . 3272
Vivaneau maori . . . . . . . . . . . . . . . . . . . 2889 xanthura, Paracaesio . . . . . . . . . . . . 2899, 2904
Vivaneau minuit . . . . . . . . . . . . . . . . . . 2897 xanthurus, Chaetodon . . . . . . . . . . . . . . 3256
Vivaneau nain . . . . . . . . . . . . . . . . . . . 2885 xenochrous, Scolopsis . . . . . . . . . . . . . . 3089
Vivaneau pagaie . . . . . . . . . . . . . . . . . . 2876
Vivaneau plate . . . . . . . . . . . . . . . . . . . 2898 Y
Vivaneau queue lune . . . . . . . . . . . . . . . . 2881
Yellow-and-blueback fusilier . . . . . . . . . . . . 2929
Vivaneau queue noire . . . . . . . . . . . . . . . 2874
Yellowback fusilier . . . . . . . . . . . . . . . . . 2930
Vivaneau rouillé . . . . . . . . . . . . . . . . . . 2855
Yellowback seabream . . . . . . . . . . . . . . . 3000
Vivaneau rubis . . . . . . . . . . . . . . . . . . . 2857
Yellowbanded snapper . . . . . . . . . . . . . . . 2861
Vivaneau soleil . . . . . . . . . . . . . . . . . . . 2870
Yellowbelly threadfin bream . . . . . . . . . . . . 3063
Vivaneau tidents . . . . . . . . . . . . . . . . . . 2854
Yellowdotted butterflyfish . . . . . . . . . . . . . 3250
Vivaneau verrue . . . . . . . . . . . . . . . . . . 2860
Yellowfin goatfish . . . . . . . . . . . . . . . . . 3183
Vivaneau voilier . . . . . . . . . . . . . . . . . . 2917
Yellowfin seabream . . . . . . . . . . . . . . . . 2997
Vivaneau ziebelo . . . . . . . . . . . . . . . . . . 2878
Yellowfinned ponyfish . . . . . . . . . . . . . . . 2805
Vivanette queue jaune . . . . . . . . . . . . . . . 2904
Yellowlined snapper . . . . . . . . . . . . . . . . 2890
Vivanette sellée . . . . . . . . . . . . . . . . . . 2900
Yellowlip emperor . . . . . . . . . . . . . . . . . 3045
Vivanette vanuatu . . . . . . . . . . . . . . . . . 2899
Yellowlip threadfin bream . . . . . . . . . . . . . 3078
Vivanettte cacao . . . . . . . . . . . . . . . . . . 2903
Yellowsnout large-eye bream . . . . . . . . . . . 3021
vogleri, Johnius . . . . . . . . . . . . . . . . . . 3154
Yellowstreaked snapper . . . . . . . . . . . . . . 2880
vosmeri, Scolopsis . . . . . . . . . . . . . . . . . 3089
Yellowstripe goatfish . . . . . . . . . . . . . . . . 3181
vroliki, Centropyge . . . . . . . . . . 3274-3275, 3279
Yellowstripe monocle bream . . . . . . . . . . . . 3087
Yellowstriped whiptail . . . . . . . . . . . . . . . 3085
W Yellowtail blue snapper . . . . . . . . . . . . . . 2904
Watanabe’s angelfish . . . . . . . . . . . . . . . 3284 Yellowtail butterflyfish . . . . . . . . . . . . . . . 3256
watanabei, Genicanthus . . . . . . . . . . . . . 3284 Yellowtailed croaker . . . . . . . . . . . . . . . . 3134
Wattsia . . . . . . . . . . . . . . . . . . . . . . . 2991 Yellowthread threadfin . . . . . . . . . . . . . . . 3102
Wattsia mossambica . . . . . . . . . . . . . . . 3050 Yellowtipped threadfin bream . . . . . . . . . . . 3080
Weber’s croaker . . . . . . . . . . . . . . . . . . 3153
weberi, Johnius . . . . . . . . . . . . 3141-3142, 3150 Z
weberi, Johnius (Johnius) . . . . . . . . . . . . 3153
ZANCLIDAE . . . . . . . . . . . . . . . . . . . . 3225
Whipfin ponyfish . . . . . . . . . . . . . . . . . . 2810 See also Vol. 6
Whipfin silverbiddy . . . . . . . . . . . . . . . . . 2952 Zanclus cornutus . . . . . . . . . . . . . . . . . 3225
Whiptail breams . . . . . . . . . . . . . . . . . . 3051 Zig-zag-ponyfish . .. . . . . . . . . . . . . . . . 2812
Whitebonnet anemonefish . . . . . . . . . . . . . 3349 zilli, Tilapia . . . .
. . . . . . . . . . . . . . . . 3333
Whitecheek monocle bream . . . . . . . . . . . . 3089 zonatus, Pristipomoides . . . . . . . . . . . . . . 2915
Whiteshouldered whiptail . . . . . . . . . . . . . 3083 zophomus, Polydactylus . . . . . . . . . . . . . . 3107
Whitetail angelfish . . . . . . . . . . . . . . . . . 3275 zysron, Nemipterus . . . . . . . . . . . . . . . . 3081
Wideband anemonefish . . . . . . . . . . . . . . 3348
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LIST OF COLOUR PLATES
PLATE I - LEIOGNATHIDAE PLATE V - LUTJANIDAE

1. Gazza achlamys 27. Aphareus furca
2. Gazza minuta 28. Aphareus rutilans
3. Leiognathus aureus 29. Aprion virescens
4. Leiognathus berbis 30. Etelis carbunculus
5. Leiognathus bindus 31. Etelis coruscans
6. Leiognathus blochii 32. Lipocheilus carnolabrum
7. Leiognathus daura 33. Lutjanus adetii
8. Leiognathus decants 34. Lutjanus argentimaculatus
PLATE II – LEIOGNATHIDAE PLATE VI - LUTJANIDAE

9. Leiognathus dussumieri 35. Lutjanus bengalensis
10. Leiognathus elongates 36. Lutjanus biguttatus
11. Leiognathus equulus 37. Lutjanus bohar
12. Leiognathus fasciatus 38. Lutjanus boutton
13. Leiognathus leuciscus 39. Lutjanus carponotatus
14. Leiognathus longispinis 40. Lutjanus decussatus
41. Lutjanus dodecacanthoides
42. Lutjanus ehrenbergii
PLATE III - LEIOGNATHIDAE
15. Leiognathus moretoniensis
16. Leiognathus pan PLATE VII - LUTJANIDAE
17. Leiognathus rapsoni 43. Lutjanus fulviflamma
18. Leiognathus splendens 44. Lutjanus fulvus
19. Leiognathus stercorarius 45. Lutjanus gibbus
20. Leiognathus sp. 1 46. Lutjanus johnii
47. Lutjanus kasmira
48. Lutjanus lemniscatus
PLATE IV - LEIOGNATHIDAE 49. Lutjanus lunulatus
21. Leiognathus sp. 2 50. Lutjanus lutjanus
22. Secutor hanedai
23. Secutor indicius
24. Secutor insidiator PLATE VIII - LUTJANIDAE
25. Secutor megalolepis 51. Lutjanus madras
26. Secutor ruconius 52. Lutjanus malabaricus
53. Lutjanus monostigma
54. Lutjanus quinquelineatus
55. Lutjanus rivulatus
56. Lutjanus russelli
57. Lutjanus semicinctus
58. Lutjanus stellatus
PLATE IX – LUTJANIDAE PLATE XIII - HAEMULIDAE
59. Lutjanm vitta 91. Diagramma pictum
60. Paracaesio kusakarii 92. Diagramma pictum
61. Paracaesio sordida 93. Plectorhinchus albovittatus
62. Paracaesio xanthura 94. Plectorhinchus albovittatus
63. Parapristipomoides squamimaxillaris 95. Plectorhinchus albovittatus
64. Pinjalo lewisi 96. Plectorhinchus chaetodonoides
65. Pinjalo pinjalo 97. Plectorhinchus chaetodonoides
66. Pristipomoides argyrogrammicus 98. Plectorhinchus chaetodonoides
PLATE X - LUTJANIDAE PLATE XIV - HAEMULIDAE

67. Pristipomoides auricilla 99. Plectorhinchus flavomaculatus
68. Pristipomoides fllamentosus 100. Plectorhinchus flavomaculatus
69. Pristipomoides flavipinnis 101. Plectorhinchus gibbosus
70. Pristipomoides multidens 102. Plectorhinchus lessonii
71. Pristipomoides sieboldii 103. Plectorhinchus lineatus
72. Pristipomoides zonatus 104. Plectorhinchus pictus
73. Randallichthys fllamentosus 105. Plectorhinchus picus
74. Symphorichthys spilurus 106. Plectorhinchus picus juv.
PLATE XI - CAESIONIDAE PLATE XV - HAEMULIDAE

75. Caesio caerulaurea 107. Plectorhinchus picus
76. Caesio cuning 108. Plectorhinchu spolytaenia
77. Caesio lunaris 109. Plectorhinchus schotaf
78. Caesio teres 110. Plectorhinchus vittatus
79. Caesio xanthonota 111. Pomadasys argenteus
80. Dipterygonotus balteatus 112. Pomadasys furcatum
81. Gymnocaesio gymnoptera 113. Pomadasys kaakan
82. Pterocaesio chrysozona 114. Pomadasys maculatus
PLATE XII - CAESIONIDAE PLATE XVI - SPARIDAE

83. Pterocaesio digramma 115. Acanthopagrus berda
84. Pterocaesio lativittata 116. Acanthopagrus latus
85. Pterocaesio marri 117. Argyrops spinifer
86. Pterocaesio pisang 118. Argyrops spinifer
87. Pterocaesio randatti 119. Dentex tumifrons
88. Pterocaesio tessellata 120. Pagrus auratus
89. Pterocaesio tile 121. Rhabdosargus sarba
90. Pterocaesio trilineata
PLATE XVII - LETHRINIDAE PLATE XXI - NEMIPTERIDAE
122. Gnathodentex aureolineatus 154. Nemipterus japonicus
123. Gymnocranius audleyi 155. Nemipterus marginatus
124. Gymnocramus grandoculis 156. Nemipterus mesoprion
125. Gymnocramus microdon 157. Nemipterus nematophorus
126. Lethrinus ambomensis 158. Nemipterus nematopus
127. Lethrinus atkinsoni 159. Nemipterus nemurus
128. Lethrinus conchyliatm 160. Nemipterus peronii
129. Lethrinus erythracanthus 161. Nemipterus tambuloides
PLATE XVIII - LETHRINIDAE PLATE XXII - NEMIPTERIDAE

130. Lethrinus erythropterus 162. Nemipterus thosaporni
131. Lethrinus genivittatus 163. Nemipterus virgatus
132. Lethrinus harak 164. Nemipterus vitiensis
133. Lethrinus lentjan 165. Nemipterus zysron
134. Lethrinus microdon 166. Parascolopsis eriomma
135. Lethrinus miniatus 167. Parascolopsis inermis
136. Lethrinus nebulosus 168. Parascolopsis melanophrys
137. Lethrinus obsoletus 169. Pentapodus bifasciatus
PLATE XIX - LETHRINIDAE PLATE XXIII - NEMIPTERIDAE

138. Lethrinus olivaceus 170. Pentapodm caninus
139. Lethrinus ornatus 171. Pentapodm emeryii
140. Lethrinus rubrioperculatus 172. Pentapodus paradiseus
141. Lethrinus semicinctus 173. Pentapodus setosus
142. Lethrinus variegatus 174. Pentapodus trivittatus
143. Lethrinus xanthochilus 175. Scaevius milii
144. Monotaxis grandoculis 176. Scolopsis affinis
145. Wattsia mossambica 177. Scolopsis auratus
PLATE XX - NEMIPTERIDAE PLATE XXIV - NEMIPTERIDAE

146. Nemipterus aurora 178. Scolopsis bilineatus
147. Nemipterus balinemis 179. Scolopsis ciliatus
148. Nemipterus bathybius 180. Scolopsis lineatus
149. Nemipterus celebicus 181. Scolopsis monogramma
150. Nemipterus furcosus 182. Scolopsis taeniopterus
151. Nemipterus gracilis 183. Scolopsis trilineatus
152. Nemipterus hexodon 184. Scolopsis vosmeri
153. Nemipterus isacanthus 185. Scolopsis xenochrous
PLATE XXV - MULLIDAE
186. Mulloidichthys flavolineatus
187. Mulloidichthys pflugeri
188. Mulloidichthys vanicolensis
189. Parupeneus barberinoides
190. Parupeneus barberinus
191. Parupeneus bifasciatus
192. Parupeneus bifasciatus
193. Parupeneus ciliatus
PLATE XXVI - MULLIDAE

194. Parupeneus cyclostomus
195. Parupeneus cyclostomus
196. Parupeneus heptacanthus
197. Parupeneus indicus
198. Parupeneus macronemus
199. Parupeneus multifasciatus
200. Parupeneus multifasciatus
PLATE XXVII - MULLIDAE

201. Parupeneus pleurostigma
202. Parupeneus spilurus
203. Upeneus moluccemis
204. Upeneus sulphureus
205. Upeneus sundaicus
206. Upeneus tragula
207. Upeneus vittatus
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PLATE I - LEIOGNATHIDAE
1. Gazza achlamys 2. Gazza minuta

(ill. by E. D'Antoni) (ill. by E. D'Antoni)
3. Leiognathus aureus 4. Leiognathus berbis

(ill. by E. D'Antoni) (iII. by E. D'Antoni)
5. Leiognathus bindus 6.Leiognathus blochii

7. Leiognathus daura 8.Leiognathus decants

PLATE II - LEIOGNATHIDAE
9. Leiognathus dussumieri 10. Leiognathus elongatus

11. Leiognathus equulus 12. Leiognathus fasciatus

13. Leiognathus leuciscus 14. Leiognathus longispinis

PLATE III - LEIOGNATHIDAE
16. Leiognathus pan

(ill. by E. D'Antoni)
15. Leiognathus moretoniensis

17. Leiognathus rapsoni 18. Leiognathus splendens

19. Leiognathus stercorarius

20. Leiognathus sp. 1
PLATE IV - LEIOGNATHIDAE
21. Leiognathus sp. 2 22. Secutor hanedai

23. Secutor indicius 24. Secutor insidiator

25. Secutor megalolepis 26. Secutor ruconius

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PLATE V - LUTJANIDAE
2 7. Aphareus furca 24.7 cm SL, Enewetak, Marshall Is. 28. Aphareus rutilans 33.5 cm SL, Guam
(photo by J.E. Randall) (photo by J.E. Randall)
29. Aprion virescens 33.9 cm SL, Natal, South Africa 30. Etelis carbunculus 24.5 cm SL, New Caledonia
31. Etelis coruscans 37 cm SL, Bane Rouge, Mauritius 32. Lipocheilus carnolabrum 38 cm SL, Cochin, India
33. Lutjanus adetii 17 cm SL, Noumea, New Caledonia 3 4. Lutjanus argentimaculatus 25.8 cm SL, Suakin,
(photo by J.E. Randall) Red Sea, Sudan (photo by J.E. Randall)
PLATE VI - LUTJANIDAE
35. Lutjanus bengalensis 1 0 . 8 cm SL, Mauritius 36. Lutjanus biguttatus 14.2 cm SL, Palau
37. Lutjanus bohar 52 cm SL, Ulithi, Caroline I. 38. Lutjanus boutton 1 2 . 5 cm SL, Maumere, Flores,
(photo by J.E. Randall) Indonesia (photo by J.E. Randall)
39. Lutjanus carponotatus 21.5 cm SL, One Tree I., 40. Lutjanus decussatus 16.2 cm SL, Batu Nampar,
Australia (photo by J.E. Randall) Lombok, Indonesia (photo by J.E. Randall)
41. Lutjanus dodecacanthoides 19.5 cm SL, Negros, 42. Lutjanus ehrenbergii 18.2 cm SL, Ambon,
Philippines (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
PLATE VII - LUTJANIDAE
43. Lutjanus fulviflamma 20.9 cm SL, Kendrew I., 44. Lutjanus fulvus 1 8 . 8 cm SL, Mafia I., Tanzania
Western Australia (photo by J.E. Randall) (photo by J.E. Randall)
45. Lutjanus gibbus 29.1 cm SL, Palmyra, 46. Lutjanus johnii 24.7 cm SL, Jakarta, Indonesia
US Minor Outlying Is. (photo by J.E. Randall) (photo by J.E. Randall)
47. Lutjanus kasmira 22.1 cm SL, Nuku Hiva, 48. Lutjanus lemniscatus 21 cm SL, Tuticorin, India
Marquesas Is. (photo by J.E. Randall) (photo by J.E. Randall)
49. Lutjanus lunulatus 21 cm SL, Hikkaduwa, 50. Lutjanus lutjanus 17.6 cm SL, Mutrah, Oman
Sri Lanka (photo by J.E. Randall) (photo by J.E. Randall)
PLATE VIII - LUTJANIDAE
5 1. Lutjanus madras 20.5 cm SL, Tuticorin, India 52. Lutjanus malabaricus 32.5 cm SL, Ambon,
(photo by J.E. Randall) Indonesia (photo by J.E. Randall)
53. Lutjanus monostigma 24.9 cm SL, Florida Island, 54. Lutjanus quinquelineatus 16.7 cm SL,
Solomon Is. (photo by J.E. Randall) Sri Lanka (photo by J.E. Randall)
55. Lutjanus rivulatus 63.7 cm SL, Atiha Pass, Moorea, 56. Lutjanus russelli 14.4 cm SL, Hong Kong
French Polynesia (photo by J.E. Randall) (photo by J.E. Randall)
57. Lutjanus semicinctus 11.2 cm SL, Guadalcanal, 58. Lutjanus stellatus 24.7 cm SL, Keelung,
Solomon Is. (photo by J.E. Randall) Taiwan Province of China (photo by J.E. Randall)
PLATE IX - LUTJANIDAE
5 9. Lutjanm vitta 19 cm SL, Enewetak, Marshall Is. 60. Paracaesio kusakarii 1 9 . 6 c m SL, Naha, Okinawa,
(photo by J.E. Randall) Ryukyu Is. (photo by J.E. Randall)
61. Paracaesio sordida 20.6 cm SL, Cebu, Philippines 62. Paracaesio xanthura 19.7 cm SL, Tutuila,
(photo by J.E. Randall) American Samoa (photo by J.E. Randall)
63. Parapristipomoides squamimaxillaris 33.9 cm SL, 64. Pinjalo lewisi 27.6 cm SL, Dumaguete, Negros,
Rapa, French Polynesia (photo by J.E. Randall) Philippines (photo by J.E. Randall)
65. Pinjalo pinjalo 20.5 cm SL, Bahrain 66. Pristipomoides argyrogrammicus 26.9 cm SL,
(photo by J.E. Randall) Okinawa, Ryukyu Is. (photo by J.E. Randall)
PLATE X - LUTJANIDAE
67. Pristipomoides auricilla 26.7 cm SL, Tutuila, 68. Pristipomoides fllamentosus 26.4 cm SL,
American Samoa (photo by J.E. Randall) Trincomalee, Sri Lanka (photo by J.E. Randall)
69. Pristipomoides flavipinnis 15.4 cm SL, Negros, 70. Pristipomoides multidens 46.8 cm SL, Jeddah,
Philippines (photo by J.E. Randall) Saudi Arabia (photo by J.E. Randall)
71. Pristipomoides sieboldii 23.5 cm SL, Guam 72. Pristipomoides zonatus 28.2 cm SL, Honolulu,
(photo by J.E. Randall) Hawaii (photo by J.E. Randall)
73. Randallichthys fllamentosus 48.2 cm SL, Oahu, 74. Symphorichthys spilurus 30.5 cm SL, Florida Island,
Hawaii (photo by J.E. Randall) Solomon Is. (photo by J.E. Randall)
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PLATE XI - CAESIONIDAE
75. Caesio caerulaurea 20.8 cm SL, Alite Reef, 76. Caesio cuning 15.7 cm SL, Hikkaduwa, Sri Lanka
Solomon Is. (photo by J.E. Randall) (photo by J.E. Randall)
77. Caesio lunaris 26 cm SL, Jana I., Persian Gulf, 78. Caesio teres 23.9 cm SL, Enewetak, Marshall Is.
Saudi Arabia (photo by J.E. Randall) (photo by J.E. Randall)
79. Caesio xanthonota 25.5 cm SL, Ave Maria Rocks, 80. Dipterygonotus balteatus 8,9 cm SL, Philippines,
Seychelles (photo by J.E. Randall) (photo by K.E. Carpenter)
81. Gymnocaesio gymnoptera 11.2 cm SL, Alite Reef, 82. Pterocaesio chrysozona 9.9 cm SL, Sharm el Moya,
Solomon Is. (photo by J.E. Randall) Red Sea, Egypt (photo by J.E. Randall)
PLATE XII - CAESIONIDAE
83. Pterocaesio digramma 23 cm TL, Philippines 84. Pterocaesio lativittata holotype, 10.45 cm SL,
(photo by J.E. Randall) Cocos (Keeling) Is. (photo by J.E. Randall)
85. Pterocaesio marri 1 9 . 2 cm SL, Enewetak, 86. Pterocaesio pisang 9.8 cm SL, Palau
Marshall Is. (photo by J.E. Randall) (photo by J.E. Randall)
87. Pterocaesio randatti 11.4 cm SL, Solpa I., 88. Pterocaesio tessellata holotype, 10.6 cm SL,
Philippines (photo by J.E. Randall) Philippines (photo by J.E. Randall)
89. Pterocaesio tile 17.5 cm SL, Enewetak, Marshall Is. 90. Pterocaesio trilineata 7.9 cm SL, Maumere Bay,
(photo by J.E. Randall) Flores, Indonesia (photo by J.E. Randall)
PLATE XIII - HAEMULIDAE
9 1. D i a g r a m m a pictum 21 cm SL, northern Taiwan, 92. Diagramma pictum 35.2 cm SL, Ampenan,
Province of China (photo by J.E. Randall) Lombok, Indonesia (photo by J.E. Randall)
93. Plectorhinchus albovittatus 20.7 cm SL, Negombo 94. Plectorhinchus albovittatus 34.5 cm SL, Maldives
Sri Lanka (photo by J.E. Randall) (photo by J.E. Randall)
95. Plectorhinchus albovittatus juv., 6.4 cm SL, Bali 96. Plectorhinchus chaetodonoides preadult,
I n d o n e s i a ( p h o t o by J.E. Randall) 1 8 . 5 c m SL, Iwayama Bay, Palau (photo by J.E. Randall)
97. Plectorhinchus chaetodonoides intermediate phase, 98. Plectorhinchus chaetodonoides juv., 3.7 cm SL,
13.7 cm SL, Chesterfield Is. (photo by J.E. Randall) Palau (photo by J.E. Randall)
PLATE X I V - HAEMULIDAE
99. Plectorhinchus flavomaculatus adult, 37.3 cm SL, 100. Plectorhinchus flavomaculatus intermediate phase,
Lizard I., Australia (photo by J.E. Randall) 16.4 cm SL, Mombasa, Kenya (photo by J.E. Randall)
1 0 1. Plectorhinchus gibbosus 27.7 cm SL, 102. Plectorhinchus lessonii 14 cm SL, Lombok,

La Digue, Seychelles (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
103. Plectorhinchus lineatus 40 cm SL, 104. Plectorhinchus pictus 32.2 cm SL, Bahrain
Maitre I., New Caledonia (photo by J.E. Randall) (photo by J.E. Randall)
105. Plectorhinchus picus 33.3 cm SL, Mauritius 106. Plectorhinchus picus juv, 1 5 . 2 c m SL,
(photo by J.E. Randall) Mauritius (photo by J.E. Randall)
PLATE XV - HAEMULIDAE
107. Plectorhinchus picus 54 cm SL, Lord Howe I., 108. Plectorhinchu spolytaenia 31.1 cm SL, Manado,
Australia (photo by J.E. Randall) Celebes, Indonesia (photo by J.E. Randall)
109. Plectorhinchus schotaf 25.5 cm SL, Port Sudan, 110. Plectorhinchus vittatus 21.6 cm SL,
Sudan (photo by J.E. Randall) Sri Lanka (photo by J.E. Randall)
111. Pomadasys argenteus 20.2 cm SL, Jambianom, 1 1 2 . Pomadasys furcatum 13.6 cm SL, Vizhingan,
Lombok, Indonesia (photo by J.E. Randall) India (photo by J.E. Randall)
113. Pomadasys kaakan 18.9 cm SL, Ambon, 114. Pomadasys maculatus 12.2 cm SL, Cochin, India
Indonesia (photo by J.E. Randall) (photo by J.E. Randall)
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PLATE XVI - SPARIDAE
1 1 5 . Acanthopagrus berda 20.9 cm SL, Buchanan I., 116. Acanthopagrus latus 11.3 cm SL, Half Moon Bay,
Northern Territory, Australia (photo by J.E. Randall) Saudi Arabia (photo by J.E. Randall)
1 1 7. Argyrops spinifer 11.5 cm SL, Madras, India 1 1 8 . Argyrops spinifer 1 9 . 3 cm SL, Salalah, Oman
1 19. Dentex tumifrons 22.2 cm SL, Naha, Okinawa, 120. Pagrus auratus 17.8 cm SL, Goat I., New Zealand
Ryukyu Is. (photo by J.E. Randall) (photo by J.E. Randall)
1 2 1 . Rhabdosargus sarba 20 cm SL, Vizhingan,

India (photo by J.E. Randall)
PLATE XVII - LETHRINIDAE
122. Gnathodentex aureolineatus 14.7 cm SL, 123. Gymnocranius audleyi 19.3 cm SL, Barrier Reef,
Fanning I., Kiribati (photo by J.E. Randall) Wistari Reef, Australia (photo by J.E. Randall)
124. Gymnocramus grandoculis 37.5 cm SL, 125. Gymnocramus microdon 22 cm SL, Lombok,
Mauritius (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
126. Lethrinus ambomensis 31.5 cm SL, Nuku Hiva 127. Lethrinus atkinsoni 16.6 cm SL, Lombok,
Marquesas Is. (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
128. Lethrinus conchyliatm 50 cm TL, Nosy Be, 129. Lethrinus erythracanthus 34.5 cm SL,
Madagascar (photo by G. Bianchi) Enewetak, Marshall Is. (photo by J.E. Randall)
PLATE XVIII - LETHRINIDAE
130. Lethrinus erythropterus 21.2 cm SL, Lombok 131. Lethrinus genivittatus 13.9 cm SL, Lizard I.,
Indonesia (photo by J.E. Randall) Great Barrier Reef, Australia (photo by J.E. Randall)
132. Lethrinus harak 23.9 cm SL, Mauritius 133. Lethrinus lentjan 14 cm SL, Bahrain
(photo by J. E. Randall) (photo by J.E. Randall)
134. Lethrinus microdon 25.9 cm SL, Bahrain 135. Lethrinus miniatus 35 cm SL, Heron I.,
(photo by J.E. Randall) Great Barrier Reef, Australia (photo by J.E. Randall)
136. Lethrinus nebulosus 23.4 cm SL, Tuticorin, India 137. Lethrinus obsoletus 2 9.5 cm SL, Enewetak,
(photo by J.E. Randall) Marshall Is. (photo by J.E. Randall)
PLATE XIX - LETHRINIDAE
138. Lethrinus olivaceus 43 cm SL, Enewetak, 139. Lethrinus ornatus 20.4 cm SL, Negombo,
Marshall Is. (photo by J.E. Randall) Sri Lanka (photo by J.E. Randall)
140. Lethrinus rubrioperculatus 33.7 cm SL, 141. Lethrinus semicinctus 20.2 cm SL, Naha,
Ogasawara I., Japan (photo by J.E. Randall) Okinawa, Ryukyu Is. (photo by J.E. Randall)
142. Lethrinus variegatus 14.7 cm SL, Gubal I., 143. Lethrinus xanthochilus 31.3 cm SL, Enewetak,
Red Sea, Egypt (photo by J.E. Randall) Marshall Is. (photo by J.E. Randall)
144. Monotaxis grandoculis 22 cm SL, Enewetak, 145. Wattsia mossambica 24.9 cm SL, Okinawa,
Marshall Is. (photo by J.E. Randall) Ryukyu Is. (photo by J.E. Randall)
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PLATE XX - NEMIPTERIDAE
146. Nemipterus aurora Lombok, Indonesia 147. Nemipterus balinemis Bali, Indonesia
(photo by B.C. Russell) (photo by B.C. Russell)
148. Nemipterus bathybius 149. Nemipterus celebicus Arafura Sea

(ill. by D. Eggleston) (photo by B.C. Russell)
150. Nemipterus furcosus 17.7 cm SL, Ambon, 151. Nemipterus gracilis Lombok, Indonesia
Indonesia (photo by J.E. Randall) (photo by B.C. Russell)
152. Nemipterus hexodon 17.2 cm SL, Ambon, 153. Nemipterus isacanthus Northwestern Australia
Indonesia (photo by J.E. Randall) (photo by B.C. Russell)
PLATE XXI - NEMIPTERIDAE
154. Nemipterus japonicus 18 cm SL, Cochin 155. Nemipterus marginatus

India (photo by J.E. Randall) (from Russell, 1990; courtesy CSIRO)
156. Nemipterus mesoprion 14.1 cm SL, Lombok 157. Nemipterus nematophorus 12.3 cm SL, Madras,
Indonesia (photo by J.E. Randall) India (photo by J.E. Randall)
158. Nemipterus nematopus 159. Nemipterus nemurus 17.4 cm SL, Lombok,

(from Russell, 1990; courtesy CSIRO) Indonesia (photo by J.E. Randall)
160. Nemipterus peronii 26.4 cm SL, Lombok, 161. Nemipterus tambuloides

Indonesia (photo by J.E. Randall) (ill. by D. Eggleston)
PLATE XXII - NEMIPTERIDAE
162. Nemipterus thosaporni 163. Nemipterus virgatus 15 cm SL, Shirahama,

(ill. by D. Eggleston) Japan (photo by J.E Randall)
164. Nemipterus vitiensis 165. Nemipterus zysron 15.4 cm SL, Negros,

(photo by A. Lewis) Philippines (photo by J.E Randall)
166. Parascolopsis eriomma 19.8 cm SL, Okinawa, 167. Parascolopsis inermis 18.7 cm TL, Okinawa,
Ryukyu Is. (photo by J.E Randall) Ryukyus Is. (photo by J.E Randall)
168. Parascolopsis melanophrys 169. Pentapodus bifasciatus 11.3 cm SL, Biola I.,
(photo by B.C. Russell) Singapore (photo by J.E Randall)
PLATE XXIII - NEMIPTERIDAE
170. Pentapodm caninus 15.5 cm SL, Tulai, Pulau, 171. Pentapodm emeryii 18.9 cm SL, Lombok,
Malaysia (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
172. Pentapodus paradiseus 12.3 cm SL, 173. Pentapodus setosus 11.8 cm SL, Salu I.,
Queensland, Australia (photo by J.E. Randall) Singapore (photo by J.E. Randall)
174. Pentapodus trivittatus 13.3 cm SL,New Guinea I., 175. Scaevius milii 12.3 cm SL, Dudley Point,
Papua New Guinea (photo by J.E. Randall) Northern Territory, Australia (photo by J.E. Randall)
176. Scolopsis affinis 19.6 cm SL, Lizard I., Australia 177. Scolopsis auratus 20.7 cm SL, Maldives
PLATE XXIV - NEMIPTERIDAE
178. Scolopsis bilineatus 14.6 cm SL, Lord Howe I., 179. Scolopsis ciliatus 11.3 cm SL, Port Moresby,
Australia (photo by J.E. Randall) Papua New Guinea (photo by J.E. Randall)
181. Scolopsis monogramma 21.6 cm SL, Lombok,

180. Scolopsis lineatus 12.3 cm SL, Guam
Indonesia (photo by J.E. Randall)
(photo by J.E. Randall)
182. Scolopsis taeniopterus 18.1 cm SL, Cebu, 183. Scolopsis trilineatus 13.7 cm SL, Vava'u, Tonga
Philippines (photo by J.E. Randall) (photo by J.E. Randall)
184. Scolopsis vosmeri 15.9 cm SL, Hikkaduwa, 185. Scolopsis xenochrous 13.3 cm SL, Ambon,
Sri Lanka (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
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PLATE XXV - MULLIDAE
186. Mulloidichthys flavolineatus 17.2 cm SL, 187. Mulloidichthys pflugeri 28.6 cm SL, Tahiti,
Johnston I. (photo by J.E. Randall) Society Is. (photo by J.E. Randall)
188. Mulloidichthys vanicolensis 16 cm SL, Mafia I., 189. Parupeneus barberinoides 10.4 cm SL,
Tanzania (photo by J.E. Randall) Kwajalein, Marshall Is. (photo by J.E. Randall)
190. Parupeneus barberinus 29.7 cm SL, Enewetak, 191. Parupeneus bifasciatus 20,5 cm SL, Alite Reef,
Marshall Is. (photo by J.E. Randall) Solomon Is. (photo by J.E. Randall)
192. Parupeneus bifasciatus 24.2 cm SL, Johnston I. 193. Parupeneus ciliatus 21,9 cm SL, Lord Howe I.,
(photo by J.E. Randall) Australia (photo by J.E. Randall)
PLATE XXVI - MULLIDAE
194. Parupeneus cyclostomus 22.5 cm SL, 195. Parupeneus cyclostomus 15,9 cm SL,
Kona, Hawaii (photo by J.E. Randall) Enewetak, Marshall Is. (photo by J.E. Randall)
196. Parupeneus heptacanthus 16.8 cm SL, 197. Parupeneus indicus 25.3 cm SL, Port Moresby,
New Caledonia (photo by J.E. Randall) Papua New Guinea (photo by J.E. Randall)
198. Parupeneus macronemus 24.5 cm SL, 199. Parupeneus multifasciatus 17.1 cm SL, Oahu,
Mutrah, Oman (photo by J.E. Randall) Hawaii (photo by J.E. Randall)
200. Parupeneus multifasciatus 16.1 cm SL,

Tahiti, Society Is. (photo by J.E. Randall)
PLATE XXVII - MULLIDAE
201. Parupeneus pleurostigma 16 cm SL, Enewetak, 202. Parupeneus spilurus 17.7 cm SL, Cebu,
Marshall Is. (photo by J.E. Randall) Philippines (photo by J.E. Randall)
203. Upeneus moluccemis 13.6 cm SL, Ambon, 204. Upeneus sulphureus 11.9 cm SL, Lombok,
Indonesia (photo by J.E. Randall) Indonesia (photo by J.E. Randall)
205. Upeneus sundaicus 12.2 cm SL, Ambon, 206. Upeneus tragula 21.6 cm SL, Palau
Indonesia (photo by J.E. Randall) (photo by J.E. Randall)
207. Upeneus vittatus 19.9 cm SL, Oahu,

Hawaiian Islands (photo by J.E. Randall)
FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES
ISSN 1020-6868
THE LIVING MARINE RESOURCES OF THE
WESTERN CENTRAL
PACIFIC
Volume 5 Bony fishes part 3 (Menidae to Pomacentridae)
FAO SPECIES IDENTIFICATION GUIDE FOR FISHERY PURPOSES
THE LIVING MARINE RESOURCES OF THE
WESTERN CENTRAL PACIFIC
VOLUME 5
Bony fishes part 3 (Menidae to Pomacentridae)
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, 2001
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 92-5-104587-9
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 2001
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 5. Bony fishes part 3 (Menidae to
Pomacentridae).
Rome, FAO. 2001. pp. 2791-3380.
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, sto-
matopods, shrimps, lobsters, crabs, holothurians, sharks, batoid fishes, chimaeras,
bony fishes, estuarine crocodiles, sea turtles, sea snakes, and marine mammals. The
introductory chapter outlines the environmental, 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 introduced 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 account summarizing family diagnostic characters, bio-
logical 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 important families are treated in detail (arranged
alphabetically by genus and species) and include the species name, frequent synonyms
and names of similar species, an illustration, FAO common name(s), diagnostic char-
acters, biology and fisheries information, notes on geographical distribution, and a
distribution map. For less important species, abbreviated accounts are used. Gener-
ally, this includes the species name, FAO common name(s), an illustration, a distribution
map, and notes on biology, fisheries, and distribution. 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:
Propject manager: P. Oliver.

Editorial assistance: N. DeAngelis and M. Kautenberger-Longo.
Desktop publishers: M. Kautenberger-Longo and N. DeAngelis.
Scientific illustrator: E. D’Antoni.
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FAO Species Identification Guide For Fishery Purposes Western Central Pacific Vol 5

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Perciformes: Percoidei: Menidae 2791

Perciformes: Percoidei: Menidae MENIDAE

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2792 Bony Fishes

Similar families occurring in the area

Key to the species of Leiognathidae occurring in the area

a) Secutor b) Gazza and Leiognathus c) Leiognathus a) Secutor b) Gazza and Leiognathus

2a. Canine teeth present anteriorly in jaws (Fig. 4) . . . . . . . . . (Gazza) ® 3

3a. Body scales absent anterior to a line running from base of

16a. Dark blotch astride nape . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17

List of species occurring in the area

Gazza achlamys Jordan and Starks, 1917 (Plate I, 1)

(after Jones, 1985)

Gazza minuta (Bloch, 1797) (Plate I, 2)

Leiognathus aureus Abe and Haneda, 1972 (Plate I, 3)

(after Jones, 1985)

Leiognathus berbis (Valenciennes, 1835) (Plate I, 4)

Leiognathus bindus (Valenciennes, 1835) (Plate I, 5)

Leiognathus blochii (Valenciennes, 1835) (Plate I, 6)

Leiognathus daura (Cuvier, 1829) (Plate I, 7)

Leiognathus decorus (de Vis, 1884) (Plate I, 8)

(after Randall, 1996)

Leiognathus dussumieri (Valenciennes, 1835) (Plate II, 9)

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Perciformes: Percoidei: Leiognathidae 2807

Leiognathus elongatus (Günther, 1874) (Plate II, 10)

Leiognathus equulus (Forsskål, 1775) (Plate II, 11)

Leiognathus fasciatus (Lacepède, 1803) (Plate II, 12)

Leiognathus leuciscus (Günther, 1860) (Plate II, 13)

(after Jones, 1985)

Leiognathus longispinis (Valenciennes, 1835) (Plate II, 14)

Leiognathus moretoniensis Ogilby, 1912 (Plate III, 15)

(after Jones, 1985)

Leiognathus pan Wongratana, 1988 (Plate III, 16)

(after Wongratana, 1988)

Leiognathus rapsoni Munro, 1964 (Plate III, 17)

(after Munro, 1964)

Leiognathus splendens (Cuvier, 1829) (Plate III, 18)

Leiognathus stercorarius Evermann and Seale, 1907 (Plate III, 19)

Leiognathus sp. 1 (Plate III, 20)

Secutor hanedai Mochizuki and Hayashi, 1989 (Plate IV, 22)

Secutor indicius Monkolprasit, 1973 (Plate IV, 23)

(after Monkolprasit, 1973)

Secutor insidiator (Bloch, 1787) (Plate IV, 24)

Secutor megalolepis Mochizuki and Hayashi, 1989 (Plate IV, 25)

Secutor ruconius (Hamilton-Buchanan, 1822) (Plate IV, 26)

Leiognathus sp. 2 (Plate IV, 21)

(after Jones, 1985)

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2824 Bony Fishes

single dorsal fin

Similar families occurring in the area

Stromateidae (Pampus sp.) Carangidae (Parastromateus niger)

Key to the species of Bramidae occurring in the area

2a. Dorsal-fin origin over or behind hind margin of eye

fin origin thickened

Fig. 4 Pteraclis velifera Fig. 5 Pteraclis aesticola

5a. Scales on caudal peduncle greatly en-

Fig. 7 Taractes rubescens Fig. 8 Taractes asper

upper lip not connected to upper lip joined to

a) Eumegistus ventral ridge b) Taractichthys ventral