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Perciformes: Percoidei: Carangidae 2659

CARANGIDAE
Jacks and scads (also trevallies, queenfishes, runners, amberjacks, pilotfishes, pampanos, etc.)
by W.F. Smith-Vaniz

D iagnostic characters: Body highly variable in shape, ranging from elongate and fusiform to deep and
strongly compressed; caudal peduncle of medium width to notably slender, in some species with a
moderate lateral keel, bilateral paired keels or dorsal and ventral grooves. Head varying from moderately
long and rounded to short, deep and very compressed. Eye small to large, with adipose eyelid negligible
to strongly developed. Snout pointed to blunt; lower jaw protruding to subtended (included). Teeth in jaws
in rows or bands, either small to minute or an enlarged row of recurved canines present; teeth on roof of
mouth (vomer, palatines) or tongue present or absent depending on species or developmental stage. Gill
openings large, gill membranes not united, free from isthmus; branchiostegal rays 6 to 10 (usually 7); gill
rakers moderate in length and number to long and numerous, their number decreasing with growth in some
species; opercular bones smooth (but with spines in larvae and small juveniles). Two dorsal fins that are
separate in small juveniles, the first of moderate height or very low, with IV to VIII spines (spines
obsolete or embedded in adults of some species), the second dorsal fin with I spine and 18 to 44
soft rays and the anterior lobe scarcely produced to extremely long; anal fin with II anterior spines (only
I spine in Elagatis and Seriolina) that are separate from rest of fin by a gap (becoming embedded
in adults of some species) followed by I spine and 15 to 39 soft rays, with the anterior lobe low to
elongate; pectoral fins with about 14 to 24 soft rays, either long and falcate or short and pointed or rounded;
pelvic fins with I spine and 5 soft rays, moderately long in some species to becoming rudimentary in others
(absent in Parastromateus); caudal fin forked, with the lobes equal in most species. Scales small,
sometimes difficult to see, and cycloid (smooth to touch), but strongly lanceolate to needle-like in a few
species, usually absent from some areas of head and covering body (but absent on certain body areas in
some species) and sometimes extending onto fins. Lateral line arched (curved) or elevated above
pectoral fins and straight posteriorly, extending onto caudal fin; scutes (enlarged, thickened, and
often pointed scales in lateral line) present and prominent, or reduced in some species and absent
in some genera. Vertebrae 10-11+14-16 (usually 10+14; total 24 to 26). Colour: darker above (green or
blue to blackish) and paler below (silvery to white or yellow-golden), some species almost entirely silvery
when alive, others with dark or coloured bars or stripes on head, body or fins, and some able to change
patterns; the young of many species with bars or spots.

Atropus Trachinotus

Decapterus

Caranx Scomberoides

examples of body shapes

2660 Bony Fishes

Habitat, biology, and fisheries: Mostly schooling species (but Alectis generally solitary); some species
have largely continental distributions and occur primarily in brackish environments (especially young),
others such as Elagatis and Naucrates are pelagic, usually found at or near the surface, mostly in oceanic
waters, often far offshore. This is one of the most important families of commercial fishes, and all species are
used for food. For 1995, FAO’s Yearbook of Fishery Statistics reports a total catch of around 959 300 t of
Carangidae from the Western Central Pacific. Caught commercially with trawls, also with purse seines, traps,
and on line gear. The larger species of Trachinotus, Seriola, and Caranx are highly regarded as sportfish.
Similar families occurring in the area
The Carangidae is distinguished from all similar families
in having the first 2 anal-fin spines detached from rest of
fin (caution: these spines sometimes partially or com-
pletely embedded in large adults of several genera, espe- anal fin
cially Seriola, Alectis, and Caranx; however, they can be
found, without much difficulty by dissection). The pres- II detached spines, Carangidae
ence of enlarged, thickened scutes in the straight part of often embedded in skin
lateral line in some genera easily distinguishes them from
other families. Additional distinguishing characters of
similar families (especially to those carangid genera lack-
more dorsal-fin series of
ing scutes on the lateral line), are the following: spines finlets
Scombridae: dorsal-fin spines IX to XXVII (IV to VIII in
Carangidae); no scutes developed along posterior part of
lateral line; series of finlets present behind dorsal and anal
fins (Scomberoides, the only carangid that lacks scutes
and at the same time has a series of finlets, is further
distinguished by having II detached, depressible spines
in front of anal fin). no scutes here
Scombridae
Stromateidae: Pampus, the only genus in the area, can
be distinguished from carangids as follows: IV blade-like anal-fin spines, not detached from fin (barely
visible in adults, may be embedded in skin); pelvic fins minute or absent in adults (readily apparent in
carangids, except in adults of Parastromateus). Pampus is very similar in body shape to Parastromateus,
but has the lateral line strongly arched anteriorly (weakly arched in Parastromateus) and the straight part
does not form a slight keel on the caudal peduncle or has 8 to 19 weak scutes as in Parastromateus.

no keel here
keel of
scutes
IV blade-like anal-fin spines
(barely visible in adults)
Pampus (Stromateidae) Parastromateus niger (Carangidae)
Gempylidae (especially Lepidocybium and Ruvettus species): base of first dorsal fin longer than that of
second excluding finlets (equal or shorter than second in Carangidae); a series of dorsal and anal finlets
present in Lepidocybium and Ruvettus.

Lepidocybium Ruvettus
Gempylidae
Perciformes: Percoidei: Carangidae 2661

Pomatomidae: both jaws with a series of strong com-

pressed teeth; no grooves on caudal peduncle (present
in Seriola which is superficially similar). dorsal view
no groove caudal-peduncle groove

Pomatomidae lateral view

Rachycentridae: head broad and depressed, lower jaw Seriola (Carangidae)

projecting; body more slender; first dorsal fin with VIII
or IX short, free spines, each depressible in a groove,
II or III anal-fin spines, none detached from fin.
Centrolophidae, particularly the genus Hyperoglyphe:
III anal-fin spines, none detached from fin; adults with
margin of preopercle usually moderately denticulate
(smooth in Carangidae); jaw teeth all conical; simple
caudal fin, not deeply forked.
Lactariidae: III anal-fin spines, none detached from fin; Rachycentridae
anal fin has more soft rays than dorsal fin; caudal
peduncle not exceptionally narrowed.

anal-fin rays outnumbering

denticulated III anal-fin spines, not dorsal-fin rays
detached from fin
Hyperoglyphe (Centrolophidae) Lactariidae

Identification note
Dentition: Dentition has traditionally been used by past workers to finely subdivide a number of presumably
monophyletic species-groups centered around Caranx under different generic or subgeneric designations.
One such group is the “catch-basket” category “Carangoides.” The Indo-Pacific species of Carangoides
(sensu lato) exhibit a wide range of dentition types, including some that appear to be morphologically
intermediate. In other cases, genera long recognized as valid appear to be most closely related to a
subgroup within Carangoides. It is tempting to recognize a more inclusive genus Caranx, with Caran-
goides recognized as one of several subgenera. However, Caranx would then probably become undefin-
able as a monophyletic group, thus the status quo is maintained for the present with some reluctance.
Two nominal genera, Pseudocaranx and Ulua, are especially likely candidates for synonymization.
Fin spines: The 2 detached anterior anal-fin spines and some of the anterior spines of the first dorsal fin
(especially the first spine) frequently become completely embedded in the skin in large individuals of many
carangids (all spines of the first dorsal fin become embedded in Alectis and Parastromateus at a relatively
small size). Even in those genera with a relatively high spinous dorsal fin, the first spine is usually small
and closely appressed to the second spine and thus can easily be overlooked.
Breast squamation: Many species of Carangoides, Caranx, and Uraspis have the breast only partially
scaly, and the pattern of breast squamation is an important character used to distinguish species. The
pattern of breast squamation is sometimes difficult to observe in fresh specimens; observation is facilitated
by gently scraping the breast with a knife to remove mucous and allowing the breast to partially dry,
hastened by blowing air on the area.
Gill-raker counts: In species with relatively numerous gill rakers (e.g. Decapterus and Trachurus) great
care must be taken not to overlook rakers at either end of the gill arch. It is suggested that a small knife
be used to free the upper limb of the gill arch where it joins the skull. With a little practice this can be done
2662 Bony Fishes

without leaving any stub with rakers attached. Once this has been accomplished, the gill rakers are much
easier to see. In some genera (e.g. Caranx and Seriola ) the number of developed rakers decreases with
growth with a concomitant increase in the number of rudiments (tubercles or short rakers with the diameter
of their bases greater than their height). When rudimentary rakers are included in the gill-raker counts, and
large specimens are being examined, it is very important that all of the tubercles are counted. In all cases
the raker in the angle of the gill arch is included in the count of lower-limb rakers.
Lateral-line scutes: In many carangids, size and configuration of the scales and scutes on the lateral line
is variable and there may be a gradual transition from one type to another. Scutes are here defined as
modified scales that either have their posterior margin with a small to moderate projecting spine or the
scale has a raised horizontal ridge and ends in an apex not exceeding a 90° angle. All scutes should be
counted, including those extending onto the caudal-fin base. In order to observe and accurately count the
lateral-line scales and scutes, good lighting and some magnification is recommended. In some species it
may also be necessary to remove small body scales that tend to overgrow or otherwise obscure the lateral
line.
Measurements: The curved part of the lateral line is measured as a chord of the arch extending from the
upper edge of the opercle to its junction with the straight part. The straight part of the lateral line is measured
from its junction with the curved part to its termination on the caudal-fin base (end of the last scute). In
cases where the junction of the curved and straight parts is very gradual, the curved part is considered to
begin with the scale or scute that has 3/4 of its height above the central axis of the straight part.

Key to the genera and subgenera of Carangidae occurring in the area

1a. Posterior straight part of lateral
line with hardened scutes; in
scutes
adults, pectoral fins long and fal-
cate, in most genera longer than
head (Fig. 1) (about equal to head
length in Selar a n d Trachurus,
and shorter than head length in
some Decapterus spp.) . . . . . . . . . . . ® 2
1b. No scutes in lateral line (only
pored scales, not enlarged); pec-
toral fins relatively short, shorter
than head (about 50 to 90% of
head length) . . . . . . . . . . . . . . . . ® 19 Fig. 1 Caranx

2a. Pored scales in curved lateral line scute-like, expanded dorsoventrally (Fig. 2) (caution:
in large fish may be obscured by overgrowth of smaller scales); dorsal accessory lateral
line normally extends posteriorly at least to below origin of second dorsal fin, usually
farther posteriorly (Figs 2 and 3) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachurus
2b. No enlarged scute-like scales in curved lateral line; dorsal accessory lateral line
terminating before origin of spinous dorsal fin . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
accessory 1st dorsal-fin spine
lateral line

dorsal view accessory lateral line

Fig. 2 Trachurus Fig. 3 Trachurus

3a. Pelvic fins, if present (absent in specimens larger than about 10 cm fork length),
positioned distinctly anterior to a vertical line through pectoral-fin base; soft anal-fin rays
35 to 39 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Parastromateus
3b. Pelvic fins (always present) not positioned distinctly anterior to a vertical through
pectoral-fin base; soft anal-fin rays 15 to 31 . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 4
Perciformes: Percoidei: Carangidae 2663

4a. Scales on body minute, inconspicu-

ous and embedded giving the im-
pression of naked skin; in smaller filamentous
fish, anterior soft rays of dorsal and rays
anal fins filamentous (Fig. 4) . . . . . . . . Alectis
4b. Scales on body small but conspicu-
ous, not embedded; in smaller fish,
anterior soft rays of dorsal and anal
fins not filamentous . . . . . . . . . . . . . . ® 5

5a. Second dorsal and anal fins with 1

or more distinctly separate finlets . . . . . . ® 6
5b. Second dorsal and anal fins without
finlets . . . . . . . . . . . . . . . . . . . . . ® 7 Fig. 4 Alectis

6a. Single detached terminal 2-rayed single finlet

finlet in dorsal and anal fins (Fig. 5);
shoulder girdle (cleithrum) margin
with 2 papillae, the lower papilla
larger (Fig. 6); maximum scute
height smaller than eye diameter . . . Decapterus
6b. Posterior soft dorsal- and anal-fin
rays consisting of 6 to 10 detached
finlets; shoulder girdle margin Fig. 5 Decapterus
smooth; maximum scute height
larger than eye diameter (Fig. 7) . . . Megalaspis
large scutes
small finlets
papilla

cleithral
margin

large
papilla

Fig. 7 Megalaspis

gill chamber after lifting operculum

small
Fig. 6 Decapterus papilla

7a. Shoulder girdle (cleithrum) margin with a furrow

ventrally, a large papilla immediately above it and cleithral
a smaller papilla near upper edge (Fig. 8) . . . . . . . . . Selar margin
7b. Shoulder girdle margin smooth . . . . . . . . . . . . . . . ® 8

8a. Upper jaw without teeth . . . . . . . . . . . . . . . . . . . ® 9 large

8b. Upper jaw with 1 or 2 rows or a band of minute papilla
teeth (caution: teeth difficult to detect in some
Carangoides) . . . . . . . . . . . . . . . . . . . . . . . . ® 10 furrow
gill chamber after lifting operculum

Fig. 8 Selar
2664 Bony Fishes

9a. Lower jaw with a series of minute teeth; a prominent black

opercular spot encroaching on shoulder; adipose eyelid
well developed posteriorly . . . . . . . . . . . . . . . . . . . Selaroides
9b. Lower jaw with a few feeble teeth in young (smaller than
10 cm fork length), absent in adults; no black opercular
spot; adipose eyelid poorly developed . . . . . . . . . . . Gnathanodon

10a. Tongue, roof and floor of mouth white, the rest dark (Fig. 9);
anal-fin spines reabsorbed or reduced and immovable; no
teeth on vomer or palatines . . . . . . . . . . . . . . . . . . . . Uraspis
10b. Lining of mouth not distinctly black and white; anal-fin
spines normal and movable; teeth present on vomer and
palatines . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 11
Fig. 9 Uraspis
11a. Fleshy adipose eyelid com-
pletely covering eye except for
a vertical slit centred on pupil
(Fig. 10a); terminal ray of dor-
sal and anal fins finlet-like, a
little more separated from
other rays but not detached,
and about twice length of pe-
nultimate ray . . . . . . . . . . . . . . Atule
11b. Fleshy adipose eyelid, if pre-
adipose
sent, not well developed ante- eyelid
riorly, most of anterior half of a) Atule b) Alepes
pupil exposed; terminal ray of
Fig. 10
dorsal and anal fins not finlet-
like (except terminal ray length
1.5 times the length of penulti-
m a t e r a y i n l a r g e Alepes
djedeba) . . . . . . . . . . . . . . . . ® 12

12a. Both jaws with a single row of numerous, comb-like teeth; adipose eyelid well developed
on posterior half of eye only (Fig. 10b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alepes
12b. Dentition not as above; adipose eyelid, if present, variously developed . . . . . . . . . . . . . ® 13

13a. Upper jaw anteriorly with 2 irregular rows of short conical teeth, posteriorly inner surface
of jaw paved with blunt teeth (Fig. 11a); snout shorter than eye diameter . . . . . . . . . . “Alepes”
13b. Dentition not as above; snout usually equal or larger than eye diameter . . . . . . . . . . . . ® 14

14a. Upper jaw with an outer series of moderate to strong canines and an inner band of fine
teeth (Fig. 11b); lower jaw with a single row of teeth . . . . . . . . . . . . . . . . . . . . . . . ® 15
14b. Dentition not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 16
2 rows of short
conical teeth
canines
conical
teeth

inner fine
surface teeth
of jaw

a) “Alepes” b) Caranx c) Pseudocaranx

Fig. 11 ventral view of teeth in upper jaw (teeth on roof of mouth not shown)
Perciformes: Percoidei: Carangidae 2665

15a. Body generally deep, with dorsal profile more convex than ventral; total gill rakers 20
to 31 on first gill arch; segmented dorsal- and anal-fin rays never produced as filaments;
2 to 4 canines anteriorly in each jaw . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx
15b. Body shallow with dorsal and ventral profiles equally convex; total gill rakers 34 to 39
on first gill arch; segmented dorsal- and anal-fin rays produced as filaments in adult
males; no canines anteriorly in either jaw . . . . . . . . . . . . . . . . . . . . . . . . . . Pantolabus

16a. Both jaws with single series of short, conical teeth (upper jaw sometimes with an inner
row of conical teeth anteriorly) (Fig. 11c); breast completely scaly . . . . . . . . . . . Pseudocaranx
16b. Both jaws with a band of teeth, at least anteriorly; breast naked ventrally (most species)
to completely scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 17

17a. Belly with a deep median groove, accommodating pelvic fins, anus, and anal-fin spines
(Fig. 12a); pelvic fins conspicuously long and black, tip of appressed fins extending
almost to origin of anal fin; curved lateral line short, chord of curved part of lateral line
contained 1.5 to 2 times in straight part (Fig. 12b) . . . . . . . . . . . . . . . . . . . . . . . Atropus
17b. Belly without median groove; pelvic fins not conspicuously long and black; curved lateral
line moderate in most species, with chord of curved part of lateral line contained less
than 1.5 times in straight part . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 18

median groove curved part of

lateral line

pelvic fins long,

black

a) underside of belly
b) lateral view

Fig. 12 Atropus

18a. Gill rakers long, feather-like, and project into mouth along side of tongue (Fig. 13); total
gill rakers 54 to 86 on first gill arch; lower jaw becoming prominent in large adults, with
the angle of “chin” projecting beyond upper jaw (Fig. 14) . . . . . . . . . . . . . . . . . . . . . Ulua
18b. Gill rakers of normal length and shape; total gill rakers 21 to 37 on first gill arch; shape
of lower jaw not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides

gill
rakers

tongue

Fig. 13 Ulua Fig. 14 Ulua

2666 Bony Fishes

19a. Bases of soft dorsal and anal fins unequal in length, anal-fin base shorter and only about
45 to 70% of dorsal-fin base length (Fig. 15); caudal-peduncle grooves present, dorsally
and ventrally (Fig. 16) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 20
19b. Base of soft anal fin as long as, or only slightly shorter than, base of dorsal fin; no
caudal-peduncle grooves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 23

spines
caudal-peduncle groove

dorsal view

lateral view

Fig. 15 Naucrates Fig. 16 Naucrates

20a. Terminal 2-rayed finlet present in 2-rayed finlet

dorsal and anal fins (Fig. 17); up-
per jaw ending distinctly before
eye (to below anterior margin of
eye in young) . . . . . . . . . . . . . Elagatis
20b. No finlets in dorsal and anal fins;
upper jaw ending below anterior
margin of eye to posterior margin
of eye . . . . . . . . . . . . . . . . . . . ® 21 Fig. 17 Elagatis

21a. Upper jaw broadly rounded posteriorly and usually terminating below posterior margin
of eye (Fig. 18a); gill rakers on first gill arch mostly consisting of rudiments, 4 to 10 total
elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriolina
21b. Upper jaw truncate or slightly rounded posteriorly and terminating below about anterior
margin of eye to middle of eye (Fig. 18b, c); gill rakers on first gill arch mostly well
developed, 11 to 29 total elements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 22

a) Seriolina b) Naucrates c) Seriola

Fig. 18
Perciformes: Percoidei: Carangidae 2667

22a. First dorsal fin with IV or V spines; fleshy keel

soft anal-fin rays 15 to 17; fleshy
keel laterally on caudal peduncle
well developed (Fig. 19) . . . . . . . Naucrates
22b. First dorsal fin with VII or VIII
spines (caution: anterior spines
may become completely embed-
ded in large individuals); soft
anal-fin rays 18 to 22 (except 15 groove
to 17 in S. hippos); fleshy keel on
caudal peduncle absent to mod- Fig. 19 Naucrates
erately developed (S. lalandi) . . . . . Seriola

23a. Posterior soft dorsal- and anal-fin rays consisting of semi-detached finlets (Fig. 20);
distal quarter to half of rays not connected by interradial membrane (unattached portion
of rays increasing with growth); lower jaw of adults with 2 rows of conical teeth separated
by a shallow groove; upper lip joined to snout at midline by a bridge of skin (frenum),
except crossed by a shallow groove in very young . . . . . . . . . . . . . . . . . . . Scomberoides
23b. Posterior soft dorsal- and anal-fin rays not consisting of semi-detached finlets (Fig. 21);
lower jaw without teeth or with band of small villiform teeth; upper lip separated from
snout at midline by a continuous deep groove . . . . . . . . . . . . . . . . . . . . . . . Trachinotus
semi-detached finlets

Fig. 20 Scomberoides Fig. 21 Trachinotus

Key to the species of Alectis occurring in the area

1a. Profile of nape and head
broadly rounded; suborbital
de pt h r el atively narrow,
contained 1.7 to 3 times in
upper jaw length (Fig. 22);
gill rakers (excluding
rudiments) on lower limb of
first arch 12 to 17 . . . . . Alectis ciliaris
1b. Profile of nape and head
somewhat angular; suborbi-
tal depth relatively broad,
contained 0.8 to 1 times in
upper jaw length (Fig. 23);
gill rakers (excluding rudi-
ments) on lower limb of first Fig. 22 Alectis ciliaris Fig. 23 Alectis indica
arch 21 to 26 . . . . . . . Alectis indica
2668 Bony Fishes

Key to the species of Alepes occurring in the area

1a. Interradial membranes of spinous dorsal fin black; total gill rakers 24 to 30 on first arch
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alepes melanoptera
1b. Interradial membranes of spinous dorsal fin transparent to dusky; total gill rakers 32 to
47 on first arch (except 27 to 30 in A. apercna) . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2

2a. Total gill rakers 27 to 30 on first gill arch; upper jaw with supramaxilla relatively small
and without an anterior spine-like projection (Fig. 24a) . . . . . . . . . . . . . . . . Alepes apercna
2b. Total gill rakers 32 to 47 on first gill arch; upper jaw with supramaxilla relatively large
and with an anterior spine-like projection (Fig. 24b, c) . . . . . . . . . . . . . . . . . . . . . . . ® 3

a) Alepes apercna b) Alepes djedaba c) Alepes vari

Fig. 24 upper jaw (supramaxilla shaded)

3a. Total gill rakers on first gill arch 38 to 47, of which 10 to 14 on upper limb, and 27 to 33
on lower limb; lateral line with 31 to 36 scales and 39 to 51 scutes (total 77 to 85); scutes
larger (Fig. 25a); ultimate ray of dorsal and anal fins about 1.3 to 1.5 times the length
of penultimate ray . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Alepes djedaba
3b. Total gill rakers on first gill arch 32 to 38, of which 9 to 12 on upper limb, and 23 to 26
on lower limb; lateral line with 42 to 50 scales, 48 to 69 scutes (total 86 to 119); scutes
smaller (Fig. 25b); ultimate and penultimate rays of dorsal and anal fins of equal length . Alepes vari

a) Alepes djedaba b) Alepes vari

c) Alepes apercna d) Alepes melanoptera

Fig. 25 lateral line

Key to the species of Carangoides occurring in the area

Note: species of Carangoides that have variable patterns of breast squamation may key out under both
sections of couplets when this character is utilized.
1a. Breast completely scaly or with a small, median naked area ventrally, scarcely if at all
visible in lateral view (Fig. 26) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Breast partially to completely naked . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5

small naked area

small
naked area
lateral view ventral view (showing examples of variation)
Fig. 26
Perciformes: Percoidei: Carangidae 2669

2a. Second dorsal fin with a conspicuous black blotch or submarginal band; vomerine tooth
patch anchor-shaped, with a long posteromedian extension (Fig. 27a, b) . . . . . . . . . . . . ® 3
2b. Second dorsal fin without a conspicuous black blotch or submarginal band; vomerine
tooth patch without a distinct posteromedian extension (Fig. 27c) . . . . . . . . . . . . . . . . ® 4

vomerine
tooth patch

vomerine vomerine
tooth patch tooth patch
a) Carangoides praeustus b) Carangoides equula c) Carangoides plagiotaenia
Fig. 27 tooth patches on roof of mouth and upper jaw
3a. Second dorsal fin with a conspicuous black blotch anteriorly (Fig. 28); soft anal-fin rays
18 to 20; total gill rakers on first gill arch 32 to 37 . . . . . . . . . . . . . . . Carangoides praeustus
3b. Second dorsal fin with a submarginal black band (Fig. 29); soft anal-fin rays 21 to 24;
total gill rakers on first gill arch 27 to 32 . . . . . . . . . . . . . . . . . . . . . Carangoides equula

Fig. 28 Carangoides praeustus Fig. 29 Carangoides equula

4a. Soft anal-fin rays 18 to 20; poste-

rior margin of preopercle outlined
in black, at least in adults
(Fig. 30); scutes 11 to 18 . . . .
. . . . . . . . . . . . Carangoides plagiotaenia
4b. Soft anal-fin rays 21 to 24; poste-
rior margin of preopercle not
black; scutes 20 to 30 . . . Carangoides bajad Fig. 30 Carangoides plagiotaenia

5a. Naked area of breast separated from naked base of pectoral fins by a broad band of
scales (Fig. 31) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 6
5b. Naked area of breast uninterrupted to naked base of pectoral fins (Fig. 37) . . . . . . . . . . . ® 13

6a. Second dorsal fin with a conspicuous black blotch anteriorly (Fig. 28); vomerine tooth
patch anchor-shaped, with a long posteromedian extension (Fig. 27a) . . . Carangoides praeustus
6b. Second dorsal fin without a conspicuous black blotch; vomerine tooth patch without a
distinct posterior extension . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 7

7a. Soft dorsal-fin rays 25 to 34; soft anal-fin rays 21 to 26 . . . . . . . . . . . . . . . . . . . . . . ® 8

7b. Soft dorsal-fin rays 17 to 23; soft anal-fin rays 15 to 19 . . . . . . . . . . . . . . . . . . . . . . ® 10
2670 Bony Fishes

8a. Naked area of breast extends posteroventrally well beyond origin of pelvic fins (Fig. 31a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides fulvoguttatus
8b. Naked area of breast does not extend posteroventrally beyond origin of pelvic fins
(Fig. 31b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 9

naked area naked

long area short

a) Carangoides fulvoguttatus b) Carangoides ferdau, C. orthogrammus

Fig. 31

9a. In life, typically 5 or 6 distinct dusky bands on sides of adults and yellow or orange spots
on sides, if present, small, numerous and mostly above lateral line (Fig. 32); lips not
papillose in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides ferdau
9b. In life, dark bands usually absent on sides of adults and several relatively large, oblong
yellow spots with dark centres present mostly below lateral line (Fig. 33); lips finely
papillose in adults . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides orthogrammus

bands typically bands usually

present absent

oblong
yellow spots
Fig. 32 Carangoides ferdau Fig. 33 Carangoides orthogrammus

10a. Straight part of lateral line slightly longer than curved part (Fig. 34); scutes 37 to 45
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides oblongus
10b. Straight part of lateral line slightly shorter than curved part (Fig. 35); scutes 16 to 38 . . . . . ® 11

straight part longer straight part shorter

Fig. 34 Carangoides oblongus Fig. 35 Carangoides dinema

11a. No small dark blotches on dorsum between bases of dorsal-fin rays . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides caeruleopinnatus
11b. Small dark blotches (becoming larger posteriorly) on dorsum between bases of dorsal-
fin rays . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 12
Perciformes: Percoidei: Carangidae 2671

12a. Soft dorsal-fin rays 20 to 22 (rarely 19); large black spot usually present on shoulder;
naked area of breast extends posteroventrally well beyond origin of pelvic fins (Fig. 36a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides humerosus
12b. Soft dorsal-fin rays 17 to 19; no large black spot present on shoulder; naked area of
breast typically does not extend posteroventrally beyond origin of pelvic fins (Fig. 36b)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides dinema

naked area naked area

long shorter

a) Carangoides humerosus b) Carangoides dinema

Fig. 36
13a. Soft dorsaI-fin rays 25 to 32 (rarely 25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 14
13b. Soft dorsal-fin rays 17 to 23 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 15

14a. Profile of snout angular and, in specimens larger than about 30 cm fork length, horizontal
line from tip of snout distinctly below level of eye (Fig. 37a); soft anal-fin rays 21 to 26
(rarely 25 or 26); total gill rakers (including rudiments) on first gill arch 22 to 27, of which
6 to 8 on upper limb, and 17 to 21 on lower limb; vertebrae 10+14 . . . . Carangoides fulvoguttatus
14b. Profile of snout moderately rounded and, in specimens larger than about 30 cm fork
length, horizontal line from tip of snout at or through level of eye (Fig. 37b), soft anal-fin
rays 24 to 26 (usually 25); total gill rakers (including rudiments) on first gill arch 27 to
31, of which 7 to 9 on upper limb, and 19 to 22 on lower limb; vertebrae 10+15 . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides gymnostethus

a) Carangoides fulvoguttatus b) Carangoides gymnostethus

Fig. 37

15a. Small area naked of scales anteriorly just above pectoral-fin base (Fig. 38a) . . . . . . . . . . ® 16
15b. Area anteriorly just above pectoral-fin base completely scaly (Fig. 38b) . . . . . . . . . . . . . ® 17

no naked area

naked area here

a) Carangoides malabaricus b) Carangoides armatus

Fig. 38
2672 Bony Fishes

16a. Total gill rakers (including rudiments) on first gill arch 32 to 38, of which 8 to 12 on upper
limb, and 21 to 27 on lower limb; in life, tongue greyish brown to brown . . Carangoides malabaricus
16b. Total gill rakers (including rudiments) on first gill arch 27 to 31, of which 6 to 9 on upper
limb, and 19 to 22 on lower limb; in life, tongue white to pale grey . . . . Carangoides talamparoides

17a. Small dark blotches (becoming larger posteriorly) on back between bases of dorsal-fin
rays; naked area of breast typically does not extend posteroventrally beyond origin of
pelvic fins (Fig. 36b); soft dorsal-fin rays 17 to 19 . . . . . . . . . . . . . . . . . Carangoides dinema
17b. Colour pattern not as above; naked area of breast extends posteroventrally well beyond
origin of pelvic fins (Fig. 38b); soft dorsal-fin rays 18 to 23 . . . . . . . . . . . . . . . . . . . . ® 18

18a. Total gill rakers (including rudiments) on first gill arch 31 to 37 . . . . . . . . . Carangoides armatus
18b. Total gill rakers (including rudiments) on first gill arch 20 to 27 . . . . . . . . . . . . . . . . . . ® 19

19a. Adults with a steep dorsal head profile, and with a distinct break (“bump”) in profile in the
interorbital region (Fig. 39a); 3 to 8 (usually 5 to 7) of central soft rays of dorsal and anal
fins elongated in mature males (about 17 cm fork length); eye diameter about equal to,
or larger than snout length . . . . . . . . . . . . . . . . . . . . . . . . . . Carangoides hedlandensis
19b. Dorsal head profile not as steep in adults, and no distinct break (“bump”) in profile in the
interorbital region (Fig. 39b, c); central soft rays of dorsal and anal fins not elongated in
mature males; eye diameter slightly to much smaller than snout length . . . . . . . . . . . . . ® 20

20a. Dorsal profile of snout gently sloped, then abruptly vertical just above mouth cleft
(Fig. 39b); soft dorsal-fin rays 18 to 20; soft anal-fin rays 15 to 17 . . . . . Carangoides chrysophrys
20b. Dorsal profile of snout not as above (Fig. 39c), soft dorsal-fin rays 20 to 23 (usually 22
or 23); soft anal-fin rays 16 to 20 (usually 18 or 19) . . . . . . . . . . Carangoides caeruleopinnatus

bump

a) Carangoides hedlandensis b) Carangoides chrysophrys c) Carangoides caeruleopinnatus

Fig. 39

Key to the species of Caranx occurring in the area

Note: species of Caranx that have variable patterns of breast squamation will key out under both sections
of a couplet when this character is utilized.
1a. Breast completely scaly . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2
1b. Breast naked ventrally, frequently with small patch of prepelvic scales (Fig. 40) . . . . . . . . . ® 6

2a. In life, body coloration essentially uniform grey to brown; lobe of dorsal fin relatively long,
contained 2.3 to 5.3 times in fork length; profile of head relatively steep and angular
(Fig. 41) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx lugubris
2b. Body coloration not as above; lobe of dorsal fin contained 4.2 to 8.8 times in fork length;
profile of head not noticeably steep and angular . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3
Perciformes: Percoidei: Carangidae 2673

prepelvic
scale patch naked
area

pelvic fins
Fig. 40 ventral view Fig. 41 Caranx lugubris

3a. Small black spots scattered on head and body (forming at about 16 to 22 cm fork length);
snout length contained 9.2 to 12.7 times in fork length; total gill rakers (including
rudiments) on first gill arch 25 to 29 (usually 26 or 27), and total soft dorsal- and anal-fin
rays 39 to 44 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx melampygus
3b. No small black spots scattered on head and body; snout length contained 13.1 to 18.4
times in fork length; total gill rakers (including rudiments) on first gill arch 22 to 25, except
24 to 27 in C. heberi which has 34 to 38 total soft dorsal- and anal-fin rays . . . . . . . . . . . . ® 4

4a. No small black spot present on upper margin of opercle; upper lobe of caudal fin
frequently with distal half noticeably dark or black, especially in juveniles; in adults,
adipose eyelid only slightly developed (Fig. 42a) . . . . . . . . . . . . . . . . . . . . Caranx heberi
4b. A small, black spot present on upper margin of opercle; upper lobe of caudal fin usually
uniformly pigmented; in adults, adipose eyelid well developed, especially posteriorly
(Fig. 42b, c) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5

5a. In adults, dorsal-fin lobe without white tip; dorsal profile of head strongly convex and a
black spot on upper margin of opercle, in adults, at least 1/2 the diameter of pupil
(Fig. 42b); in specimens larger than 15 cm fork length, postorbital head length longer,
contained 5.7 to 7.3 times in fork length, and dorsal-fin lobe shorter, contained 5.7 to 8.8
times in fork length; vertebrae 10+14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx tille
5b. In adults, dorsal-fin lobe with white tip; dorsal profile of head moderately convex and
black spot on upper margin of opercle, its size in adults no larger than twice the diameter
of pupil (Fig. 42c); in specimens larger than 15 cm fork length, postorbital head length
shorter, contained 6.4 to 8.2 times in fork length, and dorsal-fin lobe longer, contained 5
to 6.6 times in fork length; vertebrae 10+15 . . . . . . . . . . . . . . . . . . . . Caranx sexfasciatus

opercular opercular
adipose eyelid spot spot

a) Caranx heberi b) Caranx tille c) Caranx sexfasciatus

Fig. 42
2674 Bony Fishes

6a. Naked area of breast uninterrupted to naked base of pectoral fins (Fig. 43); curved part
of lateral line short, chord of curved part contained 2.5 to 3.3 times in straight part; scutes
larger (Fig. 44a) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx bucculentus
6b. Naked area of breast separated from naked base of pectoral fins by a broad band of
scales; curved part of lateral line moderate, with chord of curved part contained less than
1.5 times in straight part; scutes smaller (Fig. 44b) . . . . . . . . . . . . . . . . . . . . . . . . . ® 7

curved
part very
short
a) Caranx bucculentus

naked area b) Caranx ignobilis

Fig. 43 Caranx bucculentus Fig. 44 lateral line

7a. Total gill rakers (including rudiments) 20 to 24 on first gill arch; general body colour
silvery to black; in specimens larger than 15 cm fork length, body depth contained 2.5 to
3.2 times in fork length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Caranx ignobilis
7b. Total gill rakers (including rudiments) 23 to 30 (rarely 23) on first gill arch; general body
colour bronze to yellow-green; in specimens larger than 15 cm fork length, body depth
contained 2.7 to 3.8 times in fork length . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 8

8a. Total soft dorsal- and anal-fin rays 34 to 38 (rarely

38); upper lobe of caudal fin frequently with distal brilliant white spot
half noticeably dark or black (especially in juve-
niles), and posterior margin of lower lobe without
a narrow white border; no pale spot on shoulder
just behind posterodorsal margin of opercle;
adults without small black spots . . . . . . . . Caranx heberi
8b. Total soft dorsal- and anal-fin rays 37 to 41 (rarely
37); upper lobe of caudal fin usually uniformly
pigmented, and posterior margin of lower lobe
with a narrow white border; in life a conspicuous
pale spot, approximately the diameter of pupil, on
shoulder just behind posterodorsal margin of op-
ercle (Fig. 45); adults with small black spots on
body above lateral line (forming at about 25 cm
fork length) . . . . . . . . . . . . . . . . . . Caranx papuensis
Fig. 45 Caranx papuensis
Perciformes: Percoidei: Carangidae 2675

Key to the species of Decapterus occurring in the area

1a. Posterior end of upper jaw concave above, rounded and produced below (Fig. 46a);
straight part of lateral line with 14 to 29 scales (Fig. 47a) . . . . . . . . . . . Decapterus macrosoma
1b. Posterior end of upper jaw straight above, straight to slightly concave below (Fig. 46b-e);
straight part of lateral line with 0 to 15 scales (except 18 to 39 scales in D. macarellus) . . . . . ® 2

a) Decapterus macrosoma b) Decapterus macarellus

c) Decapterus tabl d) Decapterus russelli e) Decapterus kurroides

Fig. 46

2a. Posterior end of upper jaw noticeably slanted anteroventrally (Fig. 46b); straight part of
lateral line with 18 to 39 scales and 24 to 40 scutes = 52 to 67 total (Fig. 47b); pectoral
fins usually shorter in adults, 58 to 72% head length; oral valve (membrane) at symphysis
of upper jaw conspicuously white (Fig. 48) . . . . . . . . . . . . . . . . . . . Decapterus macarellus
2b. Posterior end of upper jaw not as noticeably slanted anteroventrally (Fig. 46c-e); straight
part of lateral line with 0 to 10 scales and 30 to 40 scutes = 30 to 49 total (except 5 to 15
scales and 32 to 42 scutes = 41 to 50 total in D. muroadsi; Fig. 47c); pectoral fins usually
longer in adults, 71 to 105% head length (except 62 to 83% in D. tabl); oral valve
(membrane) at symphysis of upper jaw dusky or transparent (except white in
D. muroadsi) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 3

curved part of lateral line straight part

14-29 scutes
scales 18-39
scales
a) Decapterus macrosoma b) Decapterus macarellus

5-15 scales 4-10 scales

c) Decapterus muroadsi d) Decapterus tabl

0-4 scales no scales

e) Decapterus russelli f) Decapterus kurroides

Fig. 47 lateral line

2676 Bony Fishes
oral valve
3a. In life, caudal fin with upper lobe greenish yellow (or membrane)
and lower lobe grey; oral valve (membrane) at white
symphysis of upper jaw conspicuously white in
adults (Fig. 48); gill rakers on lower limb of first gill
arch 36 to 42 . . . . . . . . . . . . . . . Decapterus muroadsi
3b. In life, caudal fin with upper and lower lobes both
hyaline, brownish or red; oral valve (membrane)
at symphysis of upper jaw transparent or dusky;
gill rakers on lower limb of first gill arch 26 to 33 Fig. 48 ventral view of upper
(except 30 to 39 in D. russelli) . . . . . . . . . . . . . . . ® 4 jaw and roof of mouth

4a. Scutes relatively small (Fig. 47d); curved part of

lateral line with 61 to 73 scales; straight part of
lateral line with 4 to 12 anterior scales; total lat-
eral-line scales and scutes (excluding scales on
caudal fin) 103 to 118; in life, caudal fin red;
posterodorsal margin of opercular membrane mi-
nutely serrated in large adults (Fig. 49) . . . . Decapterus tabl
4b. Scutes relatively large (Fig. 47e, f); curved part of
lateral line with 42 to 62 scales; straight part of
lateral line with 0 to 4 anterior scales; total lateral-
line scales and scutes (excluding scales on cau-
dal fin) 77 to 102; in life, caudal fin dusky or red; Fig. 49 Decapterus tabl
posterodorsal margin of opercular membrane
smooth in adults . . . . . . . . . . . . . . . . . . . . . . . ® 5

5a. In life, caudal fin hyaline to dusky; gill rakers on lower limb of first gill arch usually 32 to
39; soft anal-fin rays (including finlet) 25 to 29 (rarely 25); eye usually smaller . . Decapterus russelli
5b. In life, caudal fin red; gill rakers on lower limb of first gill arch usually 26 to 32; soft anal-fin
rays (including finlet) 22 to 26 (rarely 26); eye usually larger . . . . . . . . . . Decapterus kurroides

Key to the species of Pseudocaranx occurring in the area

1a. Scales in curved part of lateral line
57 to 78; no scales on preorbital
bone below and in front of eye and
on expanded part of maxilla; pos-
terior end of upper jaw essentially
vertical (Fig. 50a); total gill rakers
(including rudiments) 29 to 35 on
first gill arch . . . . . . . Pseudocaranx dentex
1b. Scales in curved part of lateral line
37 to 48; scales present on preorbi-
tal bone below and in front of eye
and on expanded part of maxilla;
posterior end of upper jaw slanted a) Pseudocaranx dentex b) Pseudocaranx wrighti
posteroventrally (Fig. 50b); total gill
rakers (including rudiments) 36 to Fig. 50
44 on first gill arch . . . . Pseudocaranx wrighti
(western and southern Australia; not yet recorded
from the area, but easily confused with P. dentex and
therefore included here)
Perciformes: Percoidei: Carangidae 2677

Key to the species of Scomberoides occurring in the area

1a. Total gill rakers 8 to 15 on first gill
arch; scales on midbody below
lateral line broadly oblong or
blunty lanceolate (Fig. 51a, b);
lobe of dorsal fin uniformly pig-
mented (Figs 52 and 53) . . . . . . . . . ® 2
1b. Total gill rakers 21 to 27 on first
gill arch; scales on midbody be-
low lateral line sharply lanceo-
a) Scomberoides commersonnianus b) Scomberoides tala
late or needle-like (Fig. 51c, d);
distal half of dorsal-fin lobe
abruptly and heavily pigmented
(Figs 55 and 56) . . . . . . . . . . . . . . ® 3

2a. Upper jaw extends well beyond

posterior margin of eye, espe-
cially in adults (Fig. 52); in adults,
teeth of inner and outer rows in c) Scomberoides lysan d) Scomberoides tol
lower jaw subequal in size
(Fig. 54a); in life, large oval Fig. 51 scales on midbody below lateral line
blotches above or touching lat-
eral line . . Scomberoides commersonnianus
2b. Upper jaw extends slightly beyond posterior margin of eye (Fig. 53); in adults, inner row
of teeth in lower jaw distinctly larger than those in outer row (Fig. 54b); in life, vertically
elongate blotches intersecting lateral line . . . . . . . . . . . . . . . . . . . . . . Scomberoides tala

Fig. 52 Scomberoides commersonnianus Fig. 53 Scomberoides tala

a) Scomberoides commersonnianus

b) Scomberoides tala

Fig. 54 teeth in lower jaw (teeth in inner row black)

3a. Scales on midbody below lateral line sharply lanceolate (Fig. 51c); in adults, upper jaw
extends to or slightly beyond posterior margin of eye (Fig. 55); in life, a double series of
6 to 8 dusky, roundish blotches above and below lateral line, occasionally connected by
narrow isthmus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Scomberoides lysan
3b. Scales on midbody below lateral line slender, needle-like (Fig. 51d); in adults, upper jaw
does not extend to posterior margin of eye (Fig. 56); in life, oval or vertically oblong dark
blotches, the first 4 or 5 intersecting lateral line . . . . . . . . . . . . . . . . . . . . Scomberoides tol
2678 Bony Fishes

Fig. 55 Scomberoides lysan Fig. 56 Scomberoides tol

Key to the species of Selar occurring in the area

1a. Curved part of lateral line with 48 to 56 scales; curved part of lateral line moderate, with
chord of curved part contained 0.7 to 1.2 times in straight part; scutes smaller (Fig. 57a)
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Selar crumenophthalmus
1b. Curved part of lateral line with 21 to 24 scales; curved part of lateral line short, chord of
curved part contained 2.1 to 3 times in straight part; scutes larger (Fig. 57b) . . . . . . . Selar boops
48-56 scales

21-24
scales

a) Selar crumenophthalmus b) Selar boops

Fig. 57 lateral line

Key to the species of Seriola occurring in the area

1a. Soft dorsal- and anal-fin rays 22 to 25 and 15 to 17 (rarely 17), respectively; in specimens
larger than about 20 cm fork length, total gill rakers (excluding rudiments) on first gill arch
10 to 12; in life, papillae surrounding broad band of teeth in both jaws engorged with
blood giving teeth a red appearance . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola hippos
1b. Soft dorsal- and anal-fin rays 27 to 35 and 18 to 22, respectively; in specimens larger
than about 20 cm fork length, total gill rakers (excluding rudiments) on first gill arch 11
to 29; in life, papillae surrounding broad band of teeth in both jaws not engorged with
blood, appearance of teeth white . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 2

2a. Upper jaw moderately slender posteriorly, with slender supramaxilla (Fig. 58a); caudal
fin yellowish; in adults, a moderate cutaneous keel laterally on caudal peduncle; verte-
brae 11+14 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola lalandi
2b. Upper jaw relatively broad posteriorly, with broad supramaxilla (Fig. 58b-d); caudal fin
dark to dusky sometimes with a lighter posterior margin; in adults, cutaneous keel on
caudal peduncle absent to slightly developed; vertebrae 10+14 . . . . . . . . . . . . . . . . . . ® 3

supramaxilla supramaxilla supramaxilla supramaxilla

narrow broad broad broad
a) Seriola lalandi b) Seriola rivoliana c) Seriola dumerili d) Seriola hippos
Fig. 58 lateral view of head (supramaxilla shaded)
Perciformes: Percoidei: Carangidae 2679

3a. In adults, length of dorsal-fin lobe about 1.3 to 1.6 times longer than pectoral fins
(Fig. 59a) and 18 to 22% of fork length; in specimens larger than about 20 cm fork length,
total gill rakers (excluding rudiments) 22 to 26 on first gill arch; first pterygiophore of anal
fin straight in specimens larger than about 10 cm fork length (Fig. 60a) . . . . . . . Seriola rivoliana
3b. In adults, length of dorsal-fin lobe about equal to, or slightly longer than pectoral fins
(Fig. 59b) and 13 to 18% of fork length; in specimens larger than about 20 cm fork length,
total gill rakers (excluding rudiments) 11 to 19 on first gill arch; first pterygiophore of anal
fin moderately curved (Fig. 60b) . . . . . . . . . . . . . . . . . . . . . . . . . . . . Seriola dumerili

vertebrae

a) Seriola rivoliana

pterygiophore
pterygiophore curved
straight ventral
margin
of belly
anterior
margin of
first 2 spines detached from anal fin anal fin
(embedded in skin in large individuals)
b) Seriola dumerili
a) Seriola rivoliana b) Seriola dumerili
Fig. 59 Fig. 60

Key to the species of Trachinotus occurring in the area

1a. One to 7 spots in a longitudinal row on or
near lateral line (spots absent on speci-
mens smaller than about 10 to 13 cm
fork length); soft dorsal-fin rays 21 to 25 . . . . . ® 2
1b. No spots in a longitudinal row on or near
lateral line; soft dorsal-fin rays 18 to 20
(except 21 to 23 in T. africanus) . . . . . . . . . ® 4
2a. In adults, all spots equal to, or smaller
than eye diameter, and with about half of
spot below lateral line (Fig. 61); gill rak-
ers (including rudiments) on lower limb Fig. 61 Trachinotus baillonii
of first gill arch 15 to 19; vomerine and
palatine tooth patches typically type “A”
(Fig. 62a) . . . . . . . . . . . . Trachinotus baillonii
2b. In adults, anterior 2 spots larger than eye diameter and at least two-thirds of spot above
lateral line (Figs 63 and 64); gill rakers (including rudiments) on lower limb of first gill arch
11 to 15; vomerine and palatine tooth patches usually type “B” or “C” (Fig. 62b, c) . . . . . . . . ® 3
premaxilla
vomer

palatines
vomer
palatines

a) type “A” b) type “B” c) type “C”

(Trachinotus baillonii) (Trachinotus botla) (Trachinotus copperingi )
Fig. 62 tooth patches on roof of mouth and upper jaw
2680 Bony Fishes

3a. Soft anal-fin rays 19 to 21; in adults only 1 dark spot above pectoral fins, and in large
specimens spots oval-shaped (Fig. 63); dorsal-fin lobe usually longer than anal-fin lobe
in specimens larger than about 25 cm fork length; pelvic fins long, their length contained
1.5 to 1.7 times in pectoral-fin length in specimens larger than about 25 cm fork length;
vomerine tooth patch usually type “B” (Fig. 62b) . . . . . . . . . . . . . . . . . . . Trachinotus botla
3b. Soft anal-fin rays 22 to 24; in adults 2 dark spots above pectoral fins, and in large
specimens spots more vertically elongate (Fig. 64); dorsal-fin lobe consistently shorter
than anal-fin lobe in specimens larger than about 25 cm fork length; pelvic fins relatively
short, their length contained 1.7 to 2.2 times in pectoral-fin length in specimens larger
than about 25 cm fork length; vomerine tooth patch usually type “C” (Fig. 62c) . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachinotus copperingi

Fig. 63 Trachinotus bolta Fig. 64 Trachinotus copperingi

4a. Soft dorsal- and anal-fin rays 21 to 23 and 19 to 21, respectively; body often with a black
oval-shaped blotch in axillary base under pectoral fins . . . . . . . . . . . . . Trachinotus africanus
4b. Soft dorsal- and anal-fin rays 18 to 20 and 16 to 18, respectively; body uniformly
pigmented in axillary base under pectoral fins . . . . . . . . . . . . . . . . . . . . . . . . . . . . ® 5

5a. First predorsal bone inverted teardrop- or oval-shaped (Fig. 65a, this character is easily
observed by simple dissection along midline of nape); height of dorsal-fin lobe longer,
35 to 60% of fork length in specimens 10 to 40 cm fork length . . . . . . . . . . Trachinotus blochii
5b. First predorsal bone shaped like an inverted “L” with the arm projecting anteriorly
(Fig. 65b, c); anal-fin lobe yellow without a brownish anterior margin; height of dorsal-fin
lobe moderate, 24 to 37% of fork length in specimens 10 to 40 cm fork length . . . . . . . . . . ® 6

teardrop- or oval-shaped inverted L-shape inverted L-shape

a) Trachinotus blochii b) Trachinotus mookalee c) Trachinotus anak

Fig. 65 anterior part of skeleton in lateral view

(hyperossified bones shown in black)
Perciformes: Percoidei: Carangidae 2681

6a. T e e t h in a na r r o w ba n d on
tongue, persisting to about 50 cm
fork length; supraoccipital bone of
skull becoming broad and sau-
sage-shaped (Fig. 66a) in speci-
mens larger than about 30 cm fork dorsal
length; profile of snout evenly supraoccipital bone view
rounded in specimens larger than supraoccipital bone
about 20 cm fork length; preorbi-
tal and nasal bones never exhibit-
ing hyperostosis . . . . Trachinotus mookalee
6b. No teeth on tongue; supraoccipi-
tal bone of skull thin and blade-
like in adults (Fig. 66b); profile of
lateral
snout essentially straight immedi- view
ately above upper lip in speci-
mens larger than about 20 cm fork a) Trachinotus mookalee b) Trachinotus anak
length; preorbital and nasal bones Fig. 66 skull
hyperossified in specimens larger Note: the skull of Trachinotus africanus is used here for
than about 30 cm fork length T. anak; the supraoccipital bone shape is nearly identical in
(Fig. 65c) . . . . . . . . . . Trachinotus anak both species

Key to the species of Trachurus occurring in the area

1a. Dorsal accessory lateral line immediately below base of dorsal fin terminates below fifth
to eleventh (usually seventh to ninth) soft dorsal-fin ray; total scales and scutes in lateral
line 71 to 89 (usually more than 75); curved part of lateral line essentially parallel to axis
of body for most of its length (Fig. 67) . . . . . . . . . . . . . . . . . . . . . . . . Trachurus declivis
1b. Dorsal accessory lateral line immediately below base of dorsal fin terminates below first
to fifth (usually first or second) soft dorsal-fin ray; total scales and scutes in lateral line
67 to 81 (rarely more than 76); curved part of lateral line slanted for most of its length
(Fig. 68) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Trachurus novaezelandia

dorsal accessory
lateral line

Fig. 67 Trachurus declivis Fig. 68 Trachurus novaezelandiae

Key to the species of Ulua occurring in the area

1a. Total gill rakers on first gill arch 54 to 61, of which 16 to 21 on upper limb, and 37 to 41
on lower limb; tongue with central band of villiform teeth; in specimens smaller than about
15 cm fork length, first anal-fin ray produced but not filamentous, not extending beyond
caudal peducle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulua aurochs
1b. Total gill rakers on first gill arch 74 to 86, of which 23 to 27 on upper limb, and 51 to 61
on lower limb; tongue without central band of villiform teeth; in specimens smaller than
about 15 cm fork length, first anal-fin ray filamentous, extending beyond caudal peduncle
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Ulua mentalis
2682 Bony Fishes

Key to the species of Uraspis occurring in the area

1a. Naked area of breast extends
uninterrupted to naked base
of pectoral fins (Fig. 69a);
scales in curved part of lateral
line 61 to 82 . . . . . . . . Uraspis uraspis
1b. Naked area of breast sepa- continuous
rated from naked base of pec- continuous broad band
naked area naked area
toral fins by a broad band of of scales
scales (Fig. 69b); scales in
curved part of lateral line 48 to a) Uraspis uraspis b) Uraspis helvola and U. secunda
66 . . . . . Uraspis helvola and U. secunda
Fig. 69
(no verified records from the area; adults of these
2 species are virtually impossible to distinguish)

List of species occurring in the area

The generic limits and intrafamilial relationships of carangids are not well established; a few of the generic
units recognized here are subject to change and are used for convenience. At least 5 additional species,
“Caranx” koheru Hector, Pseudocaranx wrighti (Whitley), Seriola quinqueradiata Temminck and
Schlegel, Trachurus japonicus (Temminck and Schlegel), and T. murphyi Nichols, occur in the western
Pacific Ocean outside of the area of coverage.
The symbol 0 is given when species accounts are included. Species with a question mark have not yet
been recorded from the area but should be watched for.
0 Alectis ciliaris (Bloch, 1788)
0 Alectis indica (Rüppell, 1830)
0 Alepes apercna Grant, 1987
0 Alepes djedaba (Forsskål, 1775)
0 “Alepes” kleinii (Bloch, 1793)
0 Alepes melanoptera Swainson, 1839
1/

0 Alepes vari (Cuvier, 1833)

0 Atropus atropos (Schneider, 1801)
0 Atule mate (Cuvier, 1833)
0 Carangoides armatus (Rüppell, 1830)
0 Carangoides bajad (Forsskål, 1775)
0 Carangoides caeruleopinnatus (Rüppell, 1830)
0 Carangoides chrysophrys (Cuvier, 1833)
0 Carangoides dinema Bleeker, 1851
0 Carangoides equula (Temminck and Schlegel, 1844)
0 Carangoides ferdau (Forsskål, 1775)
0 Carangoides fulvoguttatus (Forsskål, 1775)
0 Carangoides gymnostethus (Cuvier, 1833)
0 Carangoides hedlandensis (Whitley, 1933)
0 Carangoides humerosus (McCulloch, 1915)
0 Carangoides malabaricus (Bloch and Schneider, 1801)
0 Carangoides oblongus (Cuvier, 1833)
0 Carangoides orthogrammus (Jordan and Gilbert, 1882)
0 Carangoides plagiotaenia Bleeker, 1857
0 Carangoides praeustus (Bennett, 1830)
0 Carangoides talamparoides Bleeker, 1852
0 Caranx bucculentus Alleyne and Macleay, 1877
0 Caranx ignobilis (Forsskål, 1775)
0 Caranx heberi (Bennett, 1830)
0 Caranx lugubris Poey, 1860
1/ Generic allocation of this species uncertain; might eventually be assigned to a separate, monotypic genus.
Perciformes: Percoidei: Carangidae 2683

0 Caranx melampygus Cuvier, 1833

0 Caranx papuensis Alleyne and Macleay, 1877
0 Caranx sexfasciatus Quoy and Gaimard, 1825
0 Caranx tille Cuvier, 1833
0 Decapterus kurroides Bleeker, 1855
0 Decapterus macarellus (Cuvier, 1833)
0 Decapterus macrosoma Bleeker, 1851
0 Decapterus muroadsi (Temminck and Schlegel, 1844)
0 Decapterus russelli (Rüppell, 1830)
0 Decapterus tabl Berry, 1967
0 Elagatis bipinnulata (Quoy and Gaimard, 1825)
0 Gnathanodon speciosus (Forsskål, 1775)
0 Megalaspis cordyla (Linnaeus, 1758)
0 Naucrates ductor (Linnaeus, 1758)
0 Pantolabus radiatus (Macleay, 1881)
0 Parastromateus niger (Bloch, 1795) 2/

0 Pseudocaranx dentex (Bloch and Schneider, 1801)

0 Scomberoides commersonnianus Lacepède, 1801
0 Scomberoides lysan (Forsskål, 1775)
0 Scomberoides tala (Cuvier, 1832)
0 Scomberoides tol (Cuvier, 1832)
0 Selar boops (Cuvier, 1833)
0 Selar crumenophthalmus (Bloch, 1793)
0 Selaroides leptolepis (Cuvier, 1833)
0 Seriola dumerili (Risso, 1810)
0 Seriola hippos Günther, 1876
0 Seriola lalandi Valenciennes, 1833
0 Seriola rivoliana Valenciennes, 1833
0 Seriolina nigrofasciata (Rüppell, 1829)
0 Trachinotus africanus Smith, 1967
0 Trachinotus anak Ogilby, 1909
0 Trachinotus baillonii (Lacepède, 1801)
0 Trachinotus blochii (Lacepède, 1801)
0 Trachinotus botla (Shaw, 1803)
0 Trachinotus copperingi (Günther, 1884)
0 Trachinotus mookalee Cuvier, 1832
0 Trachurus declivis (Jenyns, 1841)
0 Trachurus novaezelandiae Richardson, 1843
0 Ulua aurochs (Ogilby, 1915)
0 Ulua mentalis (Cuvier, 1833)
0? Uraspis helvola (Forster, 1801)
0? Uraspis secunda (Poey, 1860)
0 Uraspis uraspis (Günther, 1860)
References
Gunn, J.S. 1990. A revision of selected genera of the family Carangidae (Pisces) from Australia. Rec. Aust. Mus., Supp.
(12):77 p.
Gushiken, S. 1983. Revision of the carangid fishes of Japan. Galaxea, 2:135-264.
Lin, P.-L. and K.-T. Shao. 1999. A review of the carangid fishes (family Carangidae) from Taiwan with descriptions of
five new records. Zool. Stud. , 38(1):33-68.

2/ Many previous authors have assigned this species to a separate family, Apolectidae (or Formionidae)

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