Department of Land and
Natural Resources
Division of
Aquatic Resources
Technical Report
20-01
Areview of the biology of the
family Carangidae, with emphasis
on species found in Hawaiian waters
October 2000

Benjamin J. Cayetano
Governor
October 2000
DIVISION OF AQUATIC RESOURCES
Department of Land and Natural Resources
1151 Punchbowl Street, Room 330
Honolulu, HI 96813
Cover: Caranx sexfasciatus
photo by Michael R. Porter

Areview of the biology of the
family Carangidae, with emphasis
on species found in Hawaiian waters.
DAR Technical Report 20-01
Randy R. Honebrink
Division of Aquatic Resources
Department of Land and Natural Resources
1151 Punchbowl Street, Room 330
Honolulu, Hawaii 96813

Contents
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . .1
Taxonomic History . . . . . . . . . . . . . . . . . . . . .1
Diagnosis . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Distribution . . . . . . . . . . . . . . . . . . . . . . . . . . .5
Key to Hawaiian Genera . . . . . . . . . . . . . . . .9
Reproduction . . . . . . . . . . . . . . . . . . . . . . . .10
Early Life History . . . . . . . . . . . . . . . . . . . . .14
Age and Growth . . . . . . . . . . . . . . . . . . . . . .18
Feeding . . . . . . . . . . . . . . . . . . . . . . . . . . . .22
Behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . .27
Predation . . . . . . . . . . . . . . . . . . . . . . . . . . .29
Morbidity and Mortality . . . . . . . . . . . . . . . . .30
Ecologic Interactions . . . . . . . . . . . . . . . . . .30
Economic Importance . . . . . . . . . . . . . . . . .31
Literature Cited . . . . . . . . . . . . . . . . . . . . . .33
5

Introduction
The family Carangidae encompasses a diverse group of fishes known variously by such com-
mon names as jacks, trevallies (crevalles), amberjacks, pompanos, scads, kingfish, pilotfish,
rainbow runners, among others. Aconsiderable amount of the available literature deals with
species of greatest economic importance, especially pompanos of the genus Trachinotus, which
do not occur in Hawai‘i. The focus of this paper is on species found in Hawaiian waters,
although many of the studies referenced were conducted in other parts of the world.
Taxonomic History
The family is named for the genus Caranx, first described by Lacépède in 1801. Previous
descriptions of some carangid species, but assigned to other genera, include those by Artedi
(1738), Osbeck (1757), Linnaeus (1758), Forsskål (1775), and Bloch (1793). Linnaeus origi-
nally placed members of the carangid genera Naucratesand Trachinotusin the genus
Gasterosteus(the sticklebacks), and in 1766 described the related cobia (Rachycentron canad-
um) as Gasterosteus canadus. Forsskål and Bloch assigned various carangids to the genus
Scomber(the mackerels). Quoy and Gaimard (1824) described carangids captured during their
voyage around the world, and Cuvier and Valenciennes (1833) did early work on Indo-Pacific
species. Descriptions of carangids in Hawaiian waters were included in works by Jordan and
Everman (1903) and Fowler (1928).
Aconsiderable amount of revision in nomenclature has occurred over the years. For example,
Smith-Vaniz and Randall (1994) note that 16 junior names have been proposed for the white
trevally (Pseudocaranx dentex). Bleeker (1852) and Berg (1947) list several carangid genera as
belonging to the family Seriolidae, but that name is obsolete.
Carangids belong to the order Perciformes, suborder Percoidei, and superfamily Percoidea.
The family is divided into four subfamilies (tribes), originally described by Starks (1911):
Trachinotini, Scomberoidini, Naucratini, and Carangini. Eschmeyer (1990) lists 122 generic
names that have been ascribed to carangids at one time or another.There are now considered to
be about 32 genera and 140 species worldwide (Nelson 1994). 24 species from 13 genera occur
in Hawai‘i (Randall 1996, Suzumoto pers. comm.). In addition, the green jack, Caranx cabal-
lus, may be establishing itself in Hawaiian waters (Randall 1999), but is not included here.
Phylogenetic relationships within the suborder and family remain poorly defined (Laroche et
al. 1984). Freihofer (1978) noted that the families Nematistiidae (roosterfish), Echeneididae
(remoras), Rachycentridae (cobia), Coryphaenidae (dolphinfish or mahimahi), and Carangidae
possess an apparent specialization of the lateralis system in the form of one or two tubular ossi-
fications around the anterior extension of the nasal canal. This characteristic is rare in percoids,
and suggests these families form a monophyletic group, the carangoids (Figure 1).
1

Johnson (1984) further notes that these five families share small, adherent cycloid scales, and
lists three synapomorphies that suggest Carangidae, Coryphaenidae, Rachycentridae, and
Echeneididae are united as a monophyletic group: lack of the bony stay posterior to the ulti-
mate dorsal and anal pterygiophores found in nearly all other percoids, presence of two prenasal
canal units, and a lamellar expansion along the anterior margin of the coracoid. Hypothesized
relationships among these groups are shown in Figure 2, based on characteristics described by
Smith-Vaniz (1984a).
The roosterfish Nematistiushas sometimes been regarded as a member of Carangidae, but is
now considered a sister group of the other four families listed above. Nelson (1984) placed the
species Parastromateus nigerin its own family,Apolectidae (Formionidae), but in 1994 fol-
lowed Smith-Vaniz (1984a) and placed it in Carangidae.
Gushiken (1988) described a hypothetical phylogenetic tree for the carangids (Figure 3).
Presumed relationships were based on 25 characteristics that include gap between last two anal
spines, upper jaw, detached or semidetached finlets, scutes, adipose eyelids, pharyngeal and
premaxillary teeth, and a number of other muscular and skeletal features. As an example of the
taxonomic confusion in the family, Gushiken considered Kaiwarinusa sister genus to
Pseudocaranx, whereas Eschmeyer (1990) lists Kaiwarinusas a synonym of Carangoides.
Smith-Vaniz is currently revising the genera Pseudocaranx,Caranx, and Decapterus, which
will eventually result in further changes in nomenclature.
Diagnosis
Members of the family Carangidae are characterized by an anal fin with two anterior spines
(one spine in Elagatisand Seriolina) separated from the rest of the fin, but which often become
embedded with age (Figure 4). The caudal peduncle is very slender, and the caudal fin is
deeply forked. The dorsal fin is generally divided into an anterior portion with four to eight
2
Figure 1
Representatives of the five families of carangoid fishes. (a) Roosterfish - Nematistiidae,
(b) remora - Echeneididae, (c) cobia - Rachycentridae, (d) dolphinfish - Coryphaenidae,
(e) trevally - Carangidae. (a) After Eschmeyer and Herald (1983), (b) and (c) after McClane
(1974), (d) and (e) from Squire and Smith (1977)

3
Figure 2
Hypothesized cladogram of carangoid fishes including main groups of Carangidae. Numbers in
parentheses are estimated total number of species in taxon. After Smith-Vaniz (1984a).

4
Figure 3
Hypothetical phylogenetic tree of the family Carangidae. After Gushiken (1988).
Figure 4
Diagnostic characteristics typical of the family Carangidae. From Chan et al. (1984).
Selene
Alectis
Ulua
Atropus
Carangoides
Carangichthys
Gnathanodon
Caranx
Uraspis
Chloroscombrus
Kaiwarinus
Pseudocaranx
Hemicaranx
Alepes
Pantolabus
Selaroides
Atule
Selar
Decapterus
Trachurus
Megalaspis
Parastromateus
Seriolina
Seriola
Naucrates
Elegatis
Campogramma
Parona
Oligoplites
Scomberoides
Lichia
Trachinotus
Carangini
Naucratini
Scomberoidini
Trachinotini

5
spines and a posterior portion with one spine and 17 to 44 soft rays. In many carangids the last
rays of the dorsal and anal fins are detached and form one to nine small posterior finlets.
Pectoral fins are often long and falcate. The eye is usually protected by a transparent “adipose”
eyelid, which is immovable but not fatty.
Members of the family possess small cycloid scales, which in most species are modified into a
row of enlarged scutes along the posterior straight portion of the lateral line. In some
carangids, particularly the genera Carangoidesand Caranx, the breast is only partially scaled,
and the pattern of breast squammation is useful for species identification. Body shape is gener-
ally compressed but extremely variable, ranging from slender forms like Decapterusand
Elagatisto deep-bodied forms like Selene(Figure 5). The premaxilla is usually protrusible.
Teeth range from small and villiform to large and conical, and are located variously on the pre-
maxillae, dentary, vomer, palatines, tongue, and pharyngeals (Gunn 1990).
Distribution
Berg (1947) points out that fossil carangids are known from the Eocene, but further information
on origins and dispersal could not be located. Carangids are found in all tropical and subtropi-
cal marine waters of the world, and some occur in temperate regions. Table 1 lists the various
genera and number of species in each genus according to Smith-Vaniz (1984a), and the number
of species found in each major geographical area as indicated in more detail by Laroche et al.
(1984).
The species found in Hawai‘i are listed in Table 2. None are endemic, and all are probably
derived from the Indo-West Pacific. Nine of the 24 local species are circumtropical in distribu-
tion, as indicated in Table 3. Differences in local distribution of some species has been noted
anecdotally, especially between the Northwestern Hawaiian Islands (NWHI) and the main
islands. Pseudocaranx dentex and Carangoides ferdauseem to be more common in the NWHI,
and P. dentexis rarely seen in the main islands. Decapterus tablis also more common in the
Figure 5
Variation in body shape among carangids. (a) Elagatis bipinnulata, (b) Selene vomer.
(a) From Squire and Smith (1977), (b) after Böhlke and Chaplin (1993).

6
Table 1
Worldwide distribution of carangid genera. After Laroche et al. (1984), Smith-Vaniz (1984a), and
Randall (1996).
No. of Ind. West Cent. East West East
SpeciesOcean Pac. Pac. Pac. Atl. Atl.Hawai‘i
Trachinotini
Lichia 1 1 1
Trachinotus 20 5 6 2 4 5 4
Scomberoidini
Oligoplites 5 3 3
Parona 1 1
Scomeroides 4 4 4 1 1
Naucratini
Campogramma 1 1
Elagatis 1 1 1 1 1 1 1 1
Naucrates 1 1 1 1 1 1 1 1
Seriola 9 3 5 4 3 5 4 3
Seriolina 1 1 1
Carangini
Alectis 3 2 2 1 1 1 2 1
Alepes 4 3 4
Atropus 1 1 1
Atule 1 1 1 1 1
Carangoides 22 17 18 4 3 2 3
Caranx 14 7 8 5 5 4 5 4
Chloroscombrus2 1 1 1
Decapterus 10 5 8 5 3 3 4 4
Gnathanodon 1 1 1 1 1 1
Hemicaranx 4 2 1 1
Megalaspis 1 1 1
Pantolabus 1 1
Parastromateus1 1 1
Pseudocaranx 3 1 2 1 1 1 1 1
Selar 2 2 2 1 1 1 2 1
Selaroides 1 1 1
Selene 7 3 3 1
Trachurus 12 3 3 2 1 4
Ulua 2 1 2
Uraspis 3 3 3 2 2 1 2 2

7
Table 2
Carangids found in Hawaiian waters. Hawaiian names from Pukui and Elbert (1986) and Titcomb
(1972). Species names, synonyms, and other valid names according to Eschmeyer (1997). Common
names based on Myers (1991), Randall (1996), and Randall et al. (1990).
Hawaiian name, Species, Synonyms*, other valid names Common name
Originally described as
Scomberoidini
Lai, Scomberoides lysan Chorinemus sanctipetri* Leatherback
Scomber lysanForsskål, 1775
Naucratini
Kamanu, Elagatis bipinnulata Rainbow runner
Seriola bipinnulataQuoy & Gaimard, 1825
Naucrates ductor Pilotfish
Gasterosteus ductorLinnaeus, 1758
Kähala,Seriola dumerili Greater amberjack
Caranx dumeriliRisso, 1810
Kähala,Seriola lalandiValenciennes, 1883 S. zonata*, S. aureovittata* Yellowtail
Kähala,Seriola rivolianaValenciennes, 1833 Almaco jack
Carangini
Ulua kihikihi, Alectis ciliaris Carangoides ajax* African pompano
Zeus ciliarisBloch, 1787
‘Ömaka,Atule mate Alepes mate Yellowtail scad
Caranx mateCuvier, 1833
Ulua, Carangoides equula Kaiwarinus equula Whitefin trevally
Caranx equulaTemminck & Schlegel, 1844
Ulua, Carangoides ferdau Barred jack
Scomber ferdauForsskål, 1775
Ulua (papa), Carangoides orthogrammus Island jack
Caranx orthogrammusJordan & Gilbert, 1882
Ulua aukea, Caranx ignobilis Giant trevally
Scomber ignobilisForsskål, 1775
Ulua lä‘uli,Caranx lugubrisPoey, 1860 Black trevally
‘Ömilu,Caranx melampygusCuvier, 1833 Bluefin trevally
Pake ulua, Caranx sexfasciatus Quoy & Gaimard, 1825 Bigeye trevally
‘Öpelu,Decapterus macarellus Caranx pinnulatus* Mackerel scad
Caranx macarellusCuvier, 1833
Decapterus macrosomaBleeker, 1851 Slender scad
Decapterus muroadsi Amberstripe scad
Caranx muroadsiTemmick & Schlegel, 1844
Decapterus tablBerry, 1968 Redtail scad
Ulua pa‘opa‘o, Gnathanodon speciosus Scomber rim Golden trevally
Scomber speciosusForsskål, 1775
Pseudocaranx dentex Charangus cheilio*, Caranx dentexWhite trevally
Scomber dentexBloch & Schneider, 1801
Akule, Selar crumenophthalmus Trachurops crumenophthalmus* Bigeye scad
Scomber crumenophthalmus
Uraspis helvola Whitemouth jack
Scomber helvolusForster, 1801
Uraspis secunda U. reversa* Cottonmouth jack
Caranx secundusPoey, 1860

NWHI, especially around seamounts (Seki pers. comm.). It is unclear to what extent these vari-
ations in abundance are due to differences in fishing pressure.
Akey to Hawaiian genera is given on the following page. Gosline and Brock (1960) present a
key to the Hawaiian species of carangids, but it is of limited usefulness since some of the
species listed have been determined not to occur in Hawaiian waters, a few of the more recently
identified species are not included, and many of the generic and specific names are outdated.
Meristic values for Hawaiian carangids are given in Table 4.
8
Table 3
Worldwide distribution of carangids occurring in Hawaiian waters. After Laroche et al. (1984).
Species Distribution
Scomberoidini
Scomberoides lysan Indian Ocean, Western and Central Pacific
Naucratini
Elagatis bipinnulata Circumtropical
Naucrates ductor Circumtropical
Seriola dumerili Indian Ocean, Western and Central Pacific, Atlantic
Seriola lalandi Circumtropical
Seriola rivoliana Circumtropical
Carangini
Alectis ciliaris Circumtropical
Atule mate Indian Ocean, Western and Central Pacific
Carangoides equula Indian Ocean, Western and Central Pacific
Carangoides ferdau Indian Ocean, Western and Central Pacific
Carangoides orthogrammusIndian and Pacific Oceans
Caranx ignobilis Indian Ocean, Western and Central Pacific
Caranx lugubris Circumtropical
Caranx melampygus Indian and Pacific Oceans
Caranx sexfasciatus Indian and Pacific Oceans
Decapterus macarellus Circumtropical
Decapterus macrosoma Indian and Pacific Oceans
Decapterus muroadsi Western and Central Pacific
Decapterus tabl Indian Ocean, Western and Central Pacific, Atlantic
Gnathanodon speciosus Indian and Pacific Oceans
Pseudocaranx dentex Indian Ocean, Western and Central Pacific, Atlantic
Selar crumenophthalmus Circumtropical
Uraspis helvola Indian and Pacific Oceans, Eastern Atlantic
Uraspis secunda Circumtropical

9
Key to Hawaiian genera of Carangidae
1a.Posterior straight part of LLwith scutes; in adults, Plong and falcate, in most
genera longer than head (except about equal for Selarand shorter in some
Decapterusspp.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
1b.No scutes on LL; Pusually shorter than head (ca. 0.9-2.0 in HL) . . . . . . .10
2a.Second D and Awith one or more distincly separate finlets . . . .Decapterus
2b.Second D and Awithout finlets . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
3a.Shoulder girdle margin with a furrow ventrally, a large papilla immediately
above it and a smaller papilla near upper edge . . . . . . . . . . . . . . . . . .Selar
3b.Shoulder girdle margin smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4a.In adults, spinous D entirely buried or spines short and disconnected, height
of longest spine less than pupil diameter; body superficially naked, scales
(where present) minute or embedded; opercle scaleless . . . . . . . . . .Alectis
4b.In adults, height of erect D usually longer than eye diameter and at least ante-
rior spines united by inter-radial membrane; small scales present over most
of body; opercle at least partially scaled . . . . . . . . . . . . . . . . . . . . . . . . . . .5
5a.Tongue, floor, and roof of mouth white, the rest of mouth dark . . . .Uraspis
5b.Tongue and mouth pigmentation not as above . . . . . . . . . . . . . . . . . . . . . .6
6a.Upper and lower jaws without teeth, except a few feeble teeth in lower jaw in
young . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Gnathanodon
6b.Jaw teeth always present, varying from 1 or 2 rows to a band of minute teeth
(difficult to detect in some species of Carangoides) . . . . . . . . . . . . . . . . . . .7
7a.Fleshy adipose eyelid completely covering eye except for vertical slit cen-
tered on pupil; terminal ray of D and Afinlet-like, a little more separated from
other rays, but not detached, and about twice the length of penultimate ray
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Atule
7b.Fleshy adipose eyelid, if present, not as developed as above, terminal ray of
D and Anot finlet-like . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8a.Upper jaw with an outer series of moderate to strong canines and an inner
band of fine teeth; lower jaw with a single row of teeth . . . . . . . . . . .Caranx
8b.Dentition not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
9a.Both jaws with a band of teeth, at least anteriorly; breast naked ventrally
(most species) to completely scaled . . . . . . . . . . . . . . . . . . . . .Carangoides
9b.Both jaws with single series of short, conical teeth (upper jaw sometimes with
an inner row of conical teeth anteriorly); breast completely scaled . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pseudocaranx
10a.Base of Aas long as, or only slightly shorter than, base of soft D; no C pedun-
cle grooves, posterior D and Arays consisting of semi-detached finlets, distal
1/4 to 1/2 of rays not connected by membrane; upper lip joined to snout at
midline by a bridge of skin, but crossed by a shallow groove in very young .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Scomberoides
10b.Base of Aabout 45-70% of D base length; C peduncle grooves present, dor-
sally and ventrally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11
11a.Single 2-rayed finlet behind end of D and A; maxilla ends distinctly before eye
(to below front margin of eye in young) . . . . . . . . . . . . . . . . . . . . . .Elagatis
11b.No finlets behind D and A; maxilla ends below eye . . . . . . . . . . . . . . . . . .12
12a.First D spines 4 or 5; Arays 15-17, cutaneous keel laterally on C peduncle
well developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Naucrates
12b.First D spines 7 or 8 (anterior spines may become entirely buried in large
fish); Arays 18-22; cutaneous keel laterally on C peduncle absent to moder-
ately developed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Seriola
Modified after Myers (1991) and Smith-Vaniz (1986a);
illustrations from Myers (1991), reprinted with permission.

Reproduction
Carangids are gonochoristic, and for the most part there is no apparent difference between the
sexes. However, sexual dichromaticism has been described by Talbot and Williams (1956) and
von Westernhagen (1974) in Caranx ignobilis, in which males appeared to be darker than
females. Clarke and Privitera (1995) noted the soft portion of the anal fin of male Selar cru-
menophthalmuswas dark black during spawning season, but white in females. Clarke (1996)
observed size differences between male and female Atule mate, as described below.
Male carangids have no intromittent organ, and fertilization is external. Females are oviparous
and iteroparous. Some species spawn pelagically, whereas others spawn close to shore.
Spawning seasons for most species are fairly long, generally peaking during summer months.
10
Table 4
Meristic values for Hawaiian carangids. Based on Masuda et al. (1984), Smith-Vaniz (1984b),
and Myers (1991).
Species Dorsal fin (D) Anal fin (A) Gill Rakers (GR)
Scomberoidini
Scomberoides lysan VI-VII+I,19-21 II+1,17-19 3-8+15-20
Naucratini
Elagatis bipinnulata VI+1,25-28+2 I+I,18-20+2 9-10+25-26
Naucrates ductor IV-V+I,25-29 II+I,15-17 5-7+16-19
Seriola dumerili VII+I,29-35 II+1,18-22 5-7+14-16
Seriola lalandi VII+I,30-35 II+I,19-22 8-9+18-21
Seriola rivoliana VII+I,27-33 II+I,18-22 6-9+17-20
Carangini
Alectis ciliaris VI+I,18-20 II+I,15-17 4-6+12-17
Atule mate VIII+I,22-25 II+I,18-21 10-13+26-31
Carangoides equula VIII+I,23-25 II+I,21-24 (not available)
Carangoides ferdau VIII+I,26-34 II+I,21-26 7-10+17-20
Carangoides orthogrammus VIII+I,28-31 II-1,24-26 8-10+20-23
Caranx ignobilis VIII+I,18-21 II+I,15-17 5-7+15-17
Caranx lugubris VIII+I,20-22 II+I,16-19 6-8+17-22
Caranx melampygus VIII+I,21-24 II+I,17-20 5-9+17-21
Caranx sexfasciatus VIII+I,19-22 II+I,14-17 6-8+15-19
Decapterus macarellus VIII+I,31-36+1 II+I,27-30+1 10-13+34-38
Decapterus macrosoma VIII+I,33-38+1 II+I,27-30+1 10-12+34-38
Decapterus muroadsi VIII+I,29-33+1 II+I,24-28+1 12-14+37-39
Decapterus tabl VIII+I,30-31+1 II+I,24-25+1 11-12+30-32
Gnathanodon speciosus VII+I,18-20 II+I,15-17 8-9+19-22+27-30
Pseudocaranx dentex VIII+I,25-26 II+I,21-22 11-14+24-26
Selar crumenophthalmus VIII+I,24-27 II+I,21-23 9-12+27-31
Uraspis helvola VIII+I,27-30 II+I,20-22 6-7+14-15
Uraspis secunda VIII+I,27-32 II+I,19-23 (not available)

Actual spawning in the wild has been described for only a few species, but seems to occur
repeatedly and periodically (Thresher 1984).
Caranx
Munro et al. (1973) report capturing ripe black trevallies (Caranx lugubris) in the Caribbean
from February through September. Sudekum et al. (1991) estimated sex ratio, spawning sea-
son, and size at first reproduction for the bluefin trevally, or ‘ömilu(C. melampygus), and the
giant trevally, or ulua aukea (C. ignobilis), in relatively unexploited waters of the Northwestern
Hawaiian Islands. In both species, the sex ratio was slightly skewed to females, with male:
female ratios of 1:1.148 for C. melampygusand 1:1.39 for C. ignobilis. Spawning season was
defined as the period of the year in which a significant proportion of fish had high gonadoso-
matic index (GSI) values, where GSI = (gonad wet weight/whole body wet weight) x 100.
In C. melampygus, gravid females were found from April to November, and from May through
July more than half were gravid, with June and July being peak months. Asummer peak was
also observed in C. ignobilis. Female C. melampygusappeared to reach sexual maturity at
about 350 mm standard length (SL), and female C. ignobilisat about 600 mm SL. From the
derived growth curves (see below), this corresponds to an age at first reproduction of about two
years for C. melampygusand four years for C. ignobilis. (Note that in Hawaiian waters the cur-
rent minimum size for all species of ulua and päpio(generally defined as ulua under ten
pounds) is seven inches total length, or 178 mm, for home consumption.) Fecundity was esti-
mated for C. melampygusand ranged from 49,700 mature ova for a fish of 760 g (328 mm SL)
to 4,270,000 for a fish of 6,490 g (640 mm SL). The authors suggest both species may repro-
duce for at least four to six years.
Williams (1965) noted that during the summer spawning season he sampled shoals of C. igno-
bilisin which either only males or females were caught, and suggested the sexes segregate dur-
ing the prespawning period. The catch data by Sudekum et al. (1991) supported this to some
extent, and was interpreted to mean that spawning is coordinated in space and time, and that
large numbers of males and females aggregate to spawn. These authors further suggest that
multiple clutches may be developed over the course of a year.
von Westernhagen (1974) described spawning in C. ignobilis. Groups of sometimes more than
100 individuals would gather close to shore, then break up into smaller groups of three to four,
and descend to two or three meters above the bottom. Males were easily recognized by their
black and white head region and black dorsal surface. Eventually one male would pair up with
a female, the two would sink to just above the bottom, and slowly swim around each other.
While circling, they were observed to release eggs and sperm into the water, during which time
they could be easily approached.
11

Thresher (1984) notes that the only other species of Caranxfor which spawning has been
observed is the bigeye trevally (C. sexfasciatus). Apair of fish in a 22.5 m by 7.6 m enclosure
were seen in late afternoon to swim faster than other fish in the school, approach each other
closely, and press their ventral surfaces together for some time, with one fish maintaining an
almost horizontal position.
Atule
Watson and Leis (1974) showed that yellowtail scad, or ‘ömaka(Atule mate), spawn in
Hawaiian waters between late spring and early fall, and spawning primarily occurs within par-
tially enclosed bays. Clarke (1996) determined the spawning season in Kane‘ohe Bay, O‘ahu to
extend from March through October, with some eggs present as late as December. He found
newly spawned ‘ömakaeggs in samples collected as early as 0800, and present until 1300, indi-
cating ‘ömakaspawn in the morning. These eggs were mostly found in open areas of the bay
where bottom depth was at least 10 m. Size at maturity was estimated at 150 to 160 mm.
Batch fecundity of females 188 to 232 mm SLranged from 63,000 to 161,000 ova, and for a
female of average weight (155 g) was estimated at about 115,000 eggs.
Sex ratios in this study were skewed, with males making up over two-thirds of the fish collect-
ed, which was interpreted to suggest segregation by sex and differential movement into the bay.
In addition, captured males were routinely considerably larger than females, which the authors
interpreted as evidence that males may have higher growth rates or lower mortality rates than
females.
Gnathanodon
Alimited amount of information regarding the golden trevally, or ulua pa‘opa‘o (Gnathanodon
speciosus), was obtained by Watson and Leis (1974) from studies in Kane‘ohe Bay. Newly
spawned eggs were collected in the bay from late February through mid-October, with a peak
from late April through early September. Five peaks of egg abundance were noted at approxi-
mately one-month intervals, and peaks seemed to coincide with the first and third quarters of
the moon. Pa‘opa‘o appeared to begin spawning in the early evening hours and continue for
some hours into the night.
Decapterus
Of the four species of Decapterusfound in Hawaiian waters, the mackerel scad, or ‘öpelu(D.
macarellus), is by far the most common. Spawning takes place in shallow (20 to 100 m)
inshore waters from April through August, peaking in May through July. Females have been
reported to spawn only once a season (DLNR 1979); however, that would not seem to be con-
sistent with what is known about spawning frequency in most of the rest of the family.
Size at maturity is estimated at 245 mm SLfor both males and females. Batch fecundity of
females 248 to 300 mm SLranged from 13,000 to 236,000 ova (Clarke and Privitera 1995).
12

Studies by Widodo (1991) on the locally occurring congener D. macrosomain the Java Sea
indicated a size at maturity of 148 mm for males and 155 mm for females. In the Philippines,
D. tablspawns somewhat later than D. macarellus, and batch fecundity ranges from 28,700 to
48,700 eggs (Shiota 1986b).
Selar
The spawning season for bigeye scad, or akule (Selar crumenophthalmus), in Hawaiian waters
has been given variously as April through November (Kawamoto 1973), February through
August (DLNR 1979), and April through October (Clarke and Privitera 1995). The latter
authors note that there appears to be some annual variation in season, and spawning seems to
take place between dawn and dusk. Akule form large schools in shallow, flat-bottomed waters
during the spawning season, and are believed to be Hawai‘i’s most abundant carangid (DLNR
1979).
Size at first maturity is estimated by Clarke and Privitera (1995) at 190 to 200 mm SL. Batch
fecundity of females 199 to 256 mm SLranged from 48,000 to 262,000 eggs. Individual
females apparently spawned about every three days.
Seriola
Kikkawa and Everson (1984) give the size at maturity in greater amberjack, or kähala(Seriola
dumerili), females as 72 cm FL, with a spawning season from February through June peaking
in March and April. Batch fecundity estimates ranged from 1.3 to 4.0 million eggs for fish 83.0
to 118.6 cm FL.
Marino et al. (1995) collected wild S. dumeriliin the South Mediterranean Sea during the 1990-
1992 spawning seasons. Maturing ovaries were present in 53.9% of four-year-old females, and
spermatozoa were found in deferent ducts in 80% of four-year-old males. They concluded that
the first reproductive season was four years of age for both sexes, although 40% of males were
sexually mature at three years. The median length at which 50% of fish attained sexual maturi-
ty was estimated at 109 cm SLin males and 113 cm SLin females. All fish at least 128 cm SL
were sexually mature.
The spawning period had been considered to peak between late May and mid-July in those
waters, but observations of mature ovaries supported a longer spawning season. Post-ovulatory
follicles were found in some maturing ovaries, and their presence was interpreted to indicate S.
dumerilimay release batches of eggs in several waves during the same spawning season.
Thompson and Munro (1974) note that ripe S. dumeriliand S. rivolianahave been collected in
the Caribbean as late as November.They also cite reports of ripe ovaries in large S. dumerili
that occupied over one-third of the abdominal cavity, and were estimated to contain over a mil-
lion eggs.
13

Elagatis
Rainbow runners, or kamanu (Elagatis bipinnulata), spawn throughout the year, and peak
around March. The larvae are described as the most abundant of the epipelagic carangids (Seki
1986a).
Early Life History
Carangid eggs range in diameter from about 0.7 to 1.3 mm, and are generally transparent and
spherical with a segmented yolk, narrow perivitelline space and one to several oil globules,
often with melanophores (Figure 6). They are difficult to distinguish from eggs of many other
families of marine fishes. Hatching commonly occurs 24 to 48 hours after spawning, depend-
ing on egg size and water temperature. However,Watson and Leis (1974) found that eggs of
ulua pa‘opa‘o (Gnathanodon speciosus) hatch after 18 hours.
At hatching, carangid larvae are usually 1.0 to 2.0 mm notochord length (NL), and the yolk sac
is fairly large. Notable characteristics of yolk sac larvae include an oil globule at the anterior
yolk sac ventral to the head, and the presence of 24 to 27 myomeres, but for the most part
newly hatched larvae are also difficult to identify and distinguish from those of many other
marine fishes (Leis and Trnski 1989). Within a day after hatching the gut develops as a straight
tube, and a single gut loop forms after about five days, when larvae are 3 to 4 mm NL. The
lengths and times at which this occurs vary somewhat with species (Figure 7).
The oil globule and yolk sac are absorbed by 72 hours after hatching (Venkataramanujam and
Ramamoorthi 1983). Following absorption, which occurs at 2.0 to 4.0 mm NL, larval carangids
vary in shape from relatively slender forms with body depth (BD) 20 to 27% SLto more deep
bodied forms with BD 32 to 59% SL. In general, carangid larvae have large heads and are
deeper bodied than those of many other marine fishes. They become more recognizable on the
basis of body shape, presence of 24 myomeres for all Hawaiian species (except Scomberoides,
14
Figure 6
(a) Generalized carangid egg. (b-d) Development of egg of ‘ömaka,Atule mate:(b) early mid-
dle stage, (c) advanced middle stage, (d) early late-stage. (a) From Watson and Leis (1974), (b-d)
from Miller and Sumida (1974).

which has 26), long preopercular spines (except Atule mate), dorsal crest (again except A.
mate), and midline melanophores. Identification of species is difficult, since major characteris-
tics are usually differences in pigmentation which change with development (Miller et al.
1979).
The formation of head spines occurs fairly early, at about the same time as yolk sac absorption.
The first to appear is a preopercular spine at the angle of the posterior margin of the preopercle,
and its size and shape are useful in distinguishing genus. Aseries of spines develops along the
posterior margin, and a second series develops along the anterior margin. The number of spines
increases through preflexion, flexion, and postflexion stages to a maximum of about 9 anterior
and 11 posterior, and does not appear to be constant for a particular species. Numbers of spines
decrease as the larva transforms into the early juvenile stage, and eventually spines become
overgrown by tissue and bone.
In many species a median supraoccipital crest develops on the head during preflexion. Its pres-
ence is useful in distinguishing carangid larvae from those of other fishes, since it doesn’t occur
in larvae of most other fish families. The crest persists until late in transformation when it also
becomes overgrown by bone and tissue. Supraocular spines and serrations develop in larvae
from a number of carangid genera, and supracleithral spines develop in all species (Figure 8).
At the time of hatching, dorsal, preanal, anal, and caudal finfolds are already present. The pec-
toral fin base and finfold develop in yolk-sac larvae. Fin formation generally proceeds sequen-
tially from caudal, pectoral, anal and soft dorsal, spinous dorsal, to pelvic, with the dorsal and
anal fins developing anterior to posterior. Completion of dorsal and anal fin rays occurs in con-
junction with or shortly after flexion.
15
Figure 7
Larval development in Atule mate.(a) just after hatching, 1.62 mm SL; (b) three days old, 2.26 mm
SL; (c) six days old, 3.06 mm SL; (d) twelve days old, 3.96 mm SL. From Miller and Sumida (1974).

Ossification of dorsal and anal fin spines occurs proximally from the distal tip in many species.
The two anteriormost anal fin spines are separated by a gap from the third spine once the fins
have formed, and this is an important distinguishing characteristic unique to carangids. Most
species develop an antrose spine on the anterodorsal margin of the first dorsal fin pterygio-
phore, which is obvious in larvae and juveniles but eventually becomes covered with skin.
Pigmentation is variable in carangid larvae, and its diagnostic usefulness is limited. In general,
rows of melanophores appear along the dorsal and ventral margins of the tail by late preflexion,
then on the head over the brain. Also during preflexion a row of small melanophores develops
along the lateral midline, which persists into the juvenile stage. This is sometimes referred to
as a “lateral line streak.”
Laroche et al. (1984) describe developmental osteology in some detail, and elaborate on distin-
guishing characters useful in identifying flexion and postflexion carangid larvae to the genus
level.
Scales begin to form during transformation to the juvenile stage, and in many species scutes
develop as modified scales along the posterior portion of the lateral line. Scale development
proceeds anteriorly, dorsally, and ventrally from the region of the lateral line anterior and adja-
cent to the caudal peduncle. Except as noted, most of the generalized information about early
life history in carangids presented above is based on Laroche et al. (1984).
Very little information is available concerning larval behavior. Cha et al. (1994) sampled fish
larvae off the Florida keys and found 92.4% of carangid larvae occurred in the upper 25 m of
the water column, and 100% occurred in the upper 50 m.
Development from larval through juvenile toward adult stages proceeds rather directly in
carangids, and adult characters are gradually acquired. There are no sudden developmental rate
16
Figure 8
Supraoccipital crest, and peropercular, supraocular, and supracleithral spines in carangid larvae.
(a) Selar crumenophthalmus, 5.6 mm; (b) Naucrates ductor, 4.7 mm. After Laroche et al. (1984).

changes between stages. By late postflexion, at a size of 8 to 9 mm, larvae are recognizeable as
small carangids (Figure 9a), and Berry (1959) considers transformation from larva to juvenile to
occur at about 8 mm SL. Juveniles develop spotted or barred pattems at 15 to 20 mm SL
(Figure 9b), and these pigmentation pattems persist until the fish is 100 to 200 mm SL, depend-
ing on species.
Leis (1991) indicates that carangid fishes do not settle. Young juveniles often associate with
floating or drifting objects, including jellyfish, clumps of algae, flotsam, etc. As they grow
larger, they tend to lose the banded pattern and move inshore, where they often take up a
schooling existence. Migration to inshore waters seems to occur when the fish are 21 to 50 mm
SL(Berry 1959). Some forms, such as the pilotfish (Naucratesductor) and golden trevally
(Gnathanodon speciosus) do not lose the banded pattern. Naucratesjuveniles apparently do not
move inshore; as they grow, they continue to accompany larger objects, including sharks and
other large carnivorous fishes (Böhlke and Chaplin 1993).
Anumber of authors note that the young of many carangid species are known to enter estuaries.
Blaber and Cyrus (1983) studied this phenomenon in estuaries of Natal, South Africa, in several
species which also occur in Hawai‘i. The following summaries are from their work.
Caranx ignobilisjuveniles were captured in estuaries at sizes of at least 40 mm. Juveniles
could tolerate salinities of 1.5 to 38o⁄
oo
, and subadults from 0.5 to 35o⁄
oo
. Juveniles below 200
mm would occur in highly turbid waters. Recruitment into estuaries occurred during the austral
summer, but larger fish were present throughout the year.
C. melampygusjuveniles were also captured at sizes of at least 40 mm. Those below 170 mm
tolerated salinity ranges from 6.0 to 35o⁄
oo
, and those above 170 mm were found in waters from
8.0 to 35o⁄
oo
. Juveniles were found only in clear waters, and recruitment occurred from October
through March.
17
Figure 9
(a) Late postflexion larva of Elagatis bipinnulata(11.4 mm), (b) small juvenile of Uraspis secunda
(25.6 mm). After Laroche et al. (1984).

Recruitment of C. sexfasciatusinto estuaries occurred from October through April, when fish
were from 30 to 60 mm. All sizes tolerated water from 0.5 to 40
o
⁄oo, and juveniles below 120
mm were found in waters of varying turbidity.
Scomberoides lysanfry entered estuaries during the late austral summer, from January through
April, at sizes of 20 to 30 mm. Juveniles tolerated salinity ranges of 6.5 to 35
o
⁄oo, and subadults
from 0.5 to 35
o⁄oo.All size classes were found in water of low turbidity. Gosline and Brock
(1960) note that in Hawai‘i, juveniles of one to four inches in length are often found in shallow
brackish water.
Recruitment of Atule matein Hawaiian waters may be a fairly straightforward process, since
the eggs are spawned in bays, which is also a preferred habitat of adults. Gosline and Brock
(1960) state that juveniles under one inch in length are often found under jellyfish.
Age and Growth
Carangids are noted for the changes they undergo with growth (Böhlke and Chaplin 1993), and
these changes have likely been responsible for misidentification of specimens and contributed
to some of the general taxonomic confusion that has occurred. In the case of Uraspis helvola,
for example, fish below 150 mm SLpossess both anterior and posterior retrose scutes and mid-
dle antrose scutes, but in larger fish the antrose scutes have gradually transformed and only ret-
rose scutes are present. Other changes that take place as the fish passes 150 mm SLare found
in dentition, pectoral fin length, and pelvic fin length and shape, in addition to changes in band-
ing pattern as described above (Reuben 1968).
An interesting example of change with growth occurs in juveniles of the African pompano, or
ulua kihikihi (Alectis ciliaris), which are easily recognized by the presence of long filaments
trailing from the first four or five rays of the dorsal and anal fins. These filaments may be
twice the length of the body, and as the fish gets larger they gradually shorten and eventually
disappear. It has not been determined what exactly happens to them, or their possible function.
Randall et al. (1990) and Myers (1991) speculate the filaments may serve to mimic the stinging
tentacles of jellyfishes for protection from predators.
‘Öpelu(Decapterus macarellus) grow to about 20.0 cm in 12 months, 27.1 cm in 24 months,
and 30.7 cm in 36 months (cited by Shiota 1986a). Akule (Selar crumenophthalmus) grow to
about 22.9 cm in 12 months and 30.5 cm in 24 months (Shiota 1986c).
von Bertalanffy growth equations could be found for a few local species, and are given in Table
5. Watarai (1973) describes the growth of juvenile ‘ömakaby the equation
Y= (2.29) X 6.61x10
-1
and of adult ‘ömakaby Y= (143.05) e 3.45x10
-4
X
,where Yis calculated
18

length in mm for a given day (X). He does not give a von Bertalanffy equation. Growth curves
for certain species are shown in Figure 10.
Length-weight relationships were also found for some local species, and are given in Table 6.
Watarai gives the equation for juvenile ‘ömakaas Y= (2.01xl0-5) X 3.02, and for adults as Y=
(7.68) e 1.43x10
-2
X, where Yis apparently wet weight in grams and X length in mm. Length-
weight curves for certain species are shown in Figure 11.
19
Table 5
Von Bertalanffy qrowth equations for some local carangids. From: 1 - Sudekum et al. (1991),
2 - Kawamoto (1973), 3 - Chiou and Chen (1993), 4 - Iwasaki (1995), 5 - Humphreys (1986).
Table 6
Length-weight relationships for some local carangids. From: 1 - Sudekum et al. (1991), 2 - Seki
(1986b), 3 - Kawamoto (1973), 4 - Seki (1986a), 5 - Humphries (1986).
Species Growth Equation
Caranx ignobilis1 Lt= 1838 (1 - e -0.111(t- 0.097))
Caranx melampygus1 Lt= 897 (1 - e -0.233(t+ 0.044))
Selar crumenophthalmus2Lt= 270 (1 - e -0.215(t+ 0.333))
Uraspis helvola3 Lt= 633 (1 - e -0.214(t+1.213))
Elagatis bipinnulata4 L
t= 930.2 (1 - e -0.214(t+ 0.449))
Seriola dumerili5 L
t= 149.3 (1 - e -0.314(t- 0.0420))
Species Length-Weight Relationship
Caranx ignobilis1 W = 2.30 x 10-5(SL) 2.977
Caranx melampygus1 W = 2.86 x 10-5(SL) 2.974
Pseudocarnax dentex2W = 1.70 x 10-8(FL) 3.0074
Selar crumenophthalmus3W = 1.80 x 10-6(FL) 3.397
Elagatis bipinnulata4 W = 2.32 x 10-4(FL) 2.24
Seriola dumerili5 W = 2.21 x 10-8(FL) 2.9412

20
Figure 10
Growth curves for six Hawaiian carangids. (a) From Kawamoto (1973), (b) from Dalzell and Peñaflor
(1989), (c) from Sudekum et al. (1991), (d) from Watarai (1973), (e) from Chiou and Chen (1993), (f) from
Iwasaki (1995).

Sudekum et al. (1991) conducted maximum feeding rate experiments on Caranx ignobilisand
C. melampygus, in which the fish were fed ad libitumat least three times per day during several
periods ranging from four to ten consecutive days each. C. melampygusgrew from an average
of 174 mm SLto 239 mm SLover 161 days, for an average growth rate of 0.40 mm/day, as
compared with an instantaneous growth rate of 0.45 mm/day calculated from the von
Bertalanffy growth curve. Mean weight increased from 124.5 g to 302.5 g over the same peri-
od. (Results for C. ignobiliswere not given, because the fish seemed to be in poor health dur-
ing the experiment.) These authors also provided the only set of otolith data (for the same two
species) that could be found.
Maximum sizes attained by species found in Hawaiian waters are given in Table 7.
21
Figure 11
Length-weight curves for two Hawaiian carangids. (a) From Kawamoto (1973), (b) from Watarai
(1973).

Feeding
Carangids are generally described as fast-swimming carnivores and pursuit predators. Randall
(1967) divides the family by diet into fish-feeders and plankton-feeders, although that is some-
what over-simplified. The planktivores include the genera Atule,Decapterus, and Selar.A.
matefeeds diurnally on small fish and crustaceans, and is described by Gosline and Brock
(1960) as a “voracious plankton feeder.” D. macarellusfeeds largely on hyperiid amphipods
and crab megalops, along with fish larvae. D. tablfeeds in the water column on copepods,
crustacean mysids, and small fishes (Shiota 1986a, 1986b).
The diet of S. crumenophthalmusincludes small fishes (esp. anchovies and holocentrids) and
crustaceans such as stomatopods, copepods, shrimps, and crab megalops taken mostly by night
22
Table 7
Maximum sizes of carangids found in Hawaiian waters. After Smith-Vaniz (1986a), Smith-Vaniz
et al. (1990), Myers (1991). Hawai‘i records from C. Johnston, Hawai‘i Fishing News (pers. comm.).
*Not identified to species, reported as kähalaand ‘öpeluonly.
Species Length, Weight Hawai‘i Record (kg)
Scomberoidini
Scomberoides lysan to 70 cm FL 2.4 (5 lbs 4.6 oz)
Naucratini
Elagatis bipinnulata to 115 cm FL, 15.3 kg 14.1 (30 lbs 15 oz)
Naucrates ductor to 70 cm TL
Seriola dumerili to 188 cm, 80.6 kg 55.0 (121 lbs*)
Seriola lalandi to 193 cm, 58 kg
Seriola rivoliana to 110 cm FL, 24 kg
Carangini
Alectis ciliaris to 80 cm FL, 18.8 kg (Atlantic)21.8 (48 lbs)
Atule mate to 34 cm FL
Carangoides equula to 37 cm
Carangoides ferdau to 70 cm, 8.0 kg (S. Africa)
Carangoides orthogrammusto at least 71 cm FL, 6.1 kg 7.7 (17 lbs 1.2 oz)
Caranx ignobilis to at least 165 cm FL, 68 kg 86.8 (191 lbs)
Caranx lugubris to 91 cm FL, 15.5 kg
Caranx melampygus to at least 80 cm FL, 7 kg 10.2 (22 lbs 6.5 oz)
Caranx sexfasciatus to 85 cm FL 7.1 (15 lbs 8.8 oz)
Decapterus macarellus to 35 cm SL 1.4 (3 lbs 2.6 oz*)
Decapterus macrosoma to 35 cm FL
Decapterus muroadsi to 50 cm FL
Decapterus tabl to 50 cm
Gnathanodon speciosus to 110 cm, 14.8 kg (S. Africa) 7.4 (16 lbs 3 oz)
Pseudocaranx dentex to 96 cm
Selar crumenophthalmus to 30 cm FL 0.9 (1 lb 15.5 oz)
Uraspis helvola to 46 cm FL
Uraspis secunda to 48 cm

from the water column. Juveniles apparently feed by day, at least to some extent (Shiota
1986c).
Diet and feeding behavior in Caranx ignobilishas been fairly well studied in Hawai‘i.
Sudekum et al. (1991) found their diet to consist of parrotfish, ‘öpelu, wrasses, bigeyes, eels,
cephalopods (both squid and octopus), and crustaceans (lobsters, crabs, and shrimp). The diet
indicates C. ignobilisfeeds nocturnally at least part of the time, and forages in both shallow
water reef areas and open water habitats. C. ignobilisis one of the few fishes known to eat
large lobsters, and it has been observed feeding on undersized and berried lobsters released
from commercial fishing vessels.
Major (1978) studied predation behavior by small (212 to 245 mm SL) C. ignobilison
Hawaiian anchovies, or nehu (Stolephorus purpureus), at Coconut Island, O‘ahu. He described
the jack as a facultative schooling species, schooling temporarily for feeding or reproduction
but not as a requirement for survival. He also considered it to be an opportunistic carnivore,
feeding by day and night on fish and crustaceans, with nocturnal feeding occurring mostly in
the early evening.
Major’s study involved interactions between young jacks (päpio) singly or in small groups with
nehu schools of various sizes. Single predators were most successful at capturing isolated prey,
and had difficulty capturing schooled prey, perhaps because of the amount of time required to
identify and align themselves with an individual prey item for a successful strike.
When päpiowere allowed to hunt as a small group (three to five individuals), they had more
success with schooled prey. If the group formed prior to interacting with the nehu, there would
usually be a lead päpiowith others following. The leader would often charge the school as if it
were hunting individually, and the following päpiowould seem to key in on the activity of the
leader.The leader may have been orientating to individual prey items that the other päpiodid
not see. Orientations to the prey were mostly directed at individuals that seemed to make mis-
takes, such as moving too far away from the school to snap at something in the water, turning
the wrong way when the school changed directions, or falling behind or moving too far ahead
of the school. In general, the marginal or peripheral nehu were the ones most likely to be
attacked.
In some instances the päpiogroups would bunch together and form an inverted “V” that would
rapidly penetrate the nehu school. The penetration would break up the school, usually causing
some prey to become isolated. Before these isolated nehu could rejoin the school, they would
be attacked. This ability to break up prey schools was considered to be a major benefit of
grouping by predators. Parrish (1993) also noted increased predatory success by jacks (C.
caballus) when hunting in groups of three to five, especially when breaking up schools and ori-
23

enting to stragglers and “pseudopods” (groups that briefly detach from the school and then rein-
tegrate).
Major also made observations regarding handling times by päpio. Often prey were engulfed
and swallowed nearly simultaneously. But sometimes they were caught by the head or tail, or
crossways between the päpio’sjaws, and had to be manipulated before swallowing. The päpio
would then spit the prey out, swim at it again, and engulf it. Sometimes a number of such
expulsions would be required before the päpiowas successful. While the prey was being
manipulated in this way, it was accessible to other predators. If other päpiowere present, they
would often break off their own interactions with prey and chase and snap at portions of the
nehu sticking out of the mouth of the other päpio, usually with limited success.
Potts (1980) studied predatory behavior in C. melampygusin the Indian Ocean. He described
the species as a diurnally active predator with increased activity at dawn and dusk when small
midwater planktivorous fishes are moving to or from the shelter of the reef. From his observa-
tions, it appeared C. melampygususually hunted singly or in pairs, but sometimes in small
groups. In Hawaiian waters, single or small groups of ‘ömiluare often seen following closely
behind groups of the blue goatfish, or moano kea (Parupeneus cyclostomus), perhaps to capture
prey driven out by the goatfish (Hobson 1974, pers. obs.).
Sudekum et al. (1991) note some apparent ontogenetic shift in the diet of ‘ ömilu. Smaller indi-
viduals seem to take a higher incidence of crustaceans. Larger fishes feed on wrasses, goatfish,
damselfish, parrotfish, and filefish. The lack of deep-water species suggests C. melampygus
feeds mostly in reef areas, and the low incidence of nocturnally active prey indicates it is proba-
bly a diurnal or crepuscular predator.
Blaber and Cyrus (1983) found that juvenile C. sexfasciatusbelow 200 mm were primarily pis-
civorous on fry of estuarine species, but a large part of the diet consisted of penaeid shrimps.
C. sexfasciatusabove 200 mm were almost entirely piscivorous, but Myers (1991) describes
them as nocturnal feeders on small fishes and crustaceans.
C. lugubrisfeeds primarily on other fishes (Smith-Vaniz 1986b), but little else is known about
its feeding behavior. Longhurst and Pauly (1987) state that shoals of Caranxoften feed at the
surface and form “feeding flurries” by forcing shoals of small fish to the surface, but they do
not elaborate about the species and no other reference to that effect could be found.
The island jack (Carangoides orthogrammus) is sometimes seen “rooting” in the sand for crus-
taceans and fishes (Myers 1991). No other information regarding feeding in this or the other
two locally occurring Carangoidesspecies could be located.
24

The white trevally (Pseudocaranx dentex) is described as an opportunistic carnivore. Its diet
includes mainly fishes like conger eels, bigeyes, and groupers, but also octopuses, and crus-
taceans such as crabs and shrimps. This indicates a range of foraging grounds, including near
the bottom in deeper offshore waters (Seki 1984, 1986d).
Adult kamanu (Elagatis bipinnulata) feed on invertebrates, especially crustaceans, and small
fishes. Postlarvae and juveniles eat mostly calanoid copepods and Corycaeus(Seki 1986a)
The pilotfish (Naucrates ductor) is well known for its commensal relationship with sharks,
rays, and other large fishes. It feeds on scraps of the host’s food, along with small fishes and
invertebrates, and may eat ectoparasites from the host’s body (Smith-Vaniz 1986b).
Humphreys (1986) notes that as kähala(Seriola dumerili) increase in size their diet consists of a
larger proportion of Decapterus. He also mentions dietary differences between the Northwest-
ern Hawaiian Islands (NWHI) and the main islands. In the NWHI the main dietary components
are octopus and other bottom-associated prey (Humphreys and Kramer 1984), but in the main
islands the diet is predominantly Decapterusand other prey from the water column.
Badalamenti et al. (1995) describe ontogenetic shifts in the diet of S. dumeriliin the
Mediterranean. Individuals up to 80 mm SLfeed mostly on copepods and crustacean larvae.
Fish from 80 to 120 mm SLrepresent a transition stage in which zooplankton are still eaten,
along with some benthic and nectonic organisms like pipefishes and seahorses associated with
seagrass systems. Individuals above 120 mm SLfeed mainly on nectonic and nectobenthic
items such as goatfishes and porgies.
Grau et al. (1992) point out that the large number of pyloric caeca between the stomach and
large intestine of S. dumeriliis consistent with a diet of large prey. Sanderson et al. (1996) ana-
lyzed a videotape ofS. dumerilidisplaying what appeared to be ram suspension feeding behav-
ior in the field. An individual was observed swimming in a tight circular pattem around a reef
with an open mouth and abducted opercula, and periodically closing its mouth in a manner sim-
ilar to that of suspension-feeding fish. Because the fish was not captured, its stomach contents
could not be analyzed to confirm this behavior.Although long, closely spaced gill rakers typi-
cal of suspension feeders are absent in this species, scanning electron microscopy showed denti-
cles on the branchial surfaces which could trap particles.
Schmitt (1982) observed cooperative feeding behavior by yellowtail (S. lalandi) in two loca-
tions. At Santa Catalina Island, CA, seven yellowtail swam parallel to shore along the seaward
flank of a school of over 2,000 jack mackerel (Trachurus symmetricus). The yellowtail split a
small group of prey from the school, herded it shoreward toward a rocky coast, and surrounded
25

it. When the isolated prey then formed a dense aggregation, one yellowtail rushed through and
scattered it toward the surrounding predators.
At Danzante Island, Mexico, in the Gulf of California, Schmitt observed three episodes in
which groups of 8 to 15 yellowtail formed a line parallel to the reef and approached schools of
700 to 1,000 Cortez grunts (Lythrulon flaviguttatum), then broke into two lines and separated
small groups of about 15 grunts from the school. The yellowtail drove the small groups away
from the reef, presumably to keep them from taking cover in crevices, then encircled and
attacked them. Schmitt points out that in addition to demonstrating cooperative feeding, these
episodes show that yellowtail’s foraging behavior is plastic enough to respond to different prey
species and habitats, and provides convincing evidence that yellowtail forage diurnally since
these encounters took place in mid-day.
S. rivolianais described by Myers (1991) as a roving predator of small fishes, but no other
information on feeding could be found.
Leatherbacks, or lai (Scomberoides lysan) show an interesting ontogenetic shift in diet. Major
(1973) observed individuals in Kane‘ohe Bay from 27 to 52 mm SLswimming alongside and
almost in contact with silversides (Pranesus insularum) of 35 to 75 mm SLfor several minutes,
then striking the silversides along the dorsal surface and sides. In the lab, lai below 150 mm SL
demonstrated the same behavior with mullet (Mugil cephalus) from 40 to 150 mm SL. The lai
were immediately sacrificed and their stomach contents examined, which were found to contain
scales and epidermal tissue. Further examination of stomach contents from preserved speci-
mens also resulted in the presence of scales in lai below 150 mm SL.
The diet of lai below 50 mm SLconsists of scales and epidermal tissue of schooling fishes. At
50 mm SLthe outer dentary teeth of juvenile lai begins to be replaced, and the fish shifts its
diet to include whole schooling fishes and crustaceans in addition to scales. By 150 mm SLthe
lai’s dentition is completely replaced by adult teeth, and the fish becomes fully carnivorous.
Gerking (1994) notes that scale eaters generally do no particular damage to their host, but
Major found that lepidophagous lai often struck the same spot on mullet and silversides repeat-
edly and caused bleeding.
26

Behavior
Carangids exhibit the carangiform mode of swimming (Figure 12), obviously named for this
group of fishes. The anterior one-half to two-thirds of the body bends only slightly while
swimming, and most of the thrust is generated in the rear third. The characteristic narrow cau-
dal peduncle and high aspect ratio forked caudal fin serve to increase the efficiency of the tail
(Bond 1996). The scutes, when present, probably help reinforce the narrow peduncle (Randall
1983). Because of the caudal fin’s stiffness, small changes in direction must be made by other
fins.
Holland et al. (1996) studied fine scale movement of ‘ömilu(Caranx melampygus) in Kane‘ohe
Bay.They found ‘ömiluutilized different parts of the reef at different times, often staying near
a particular reef during the day and moving to an adjacent reef at night. Most fish showed
strong site fidelity, and didn’t move far from the point of release.
Sub-adult akule (Selar crumenophthalmus), known as halalü, form large schools in shallow
water from about July through December. Eventually they join up with adult schools, which
apparently do not make any large scale movements, even around the same island (DLNR 1979).
As has been mentioned previously, some carangids form associations with other fishes.
Examples include ‘ömilufollowing blue goatfish, and the relationship between pilotfish and
other large carnivores.
Schooling behavior and habitat preferences for Hawaiian carangids are summarized in Table 8.
27
Figure 12
The carangiform mode of swimming. After Bond (1996).

28
Table 8
Schooling behavior and habitat for carangids in Hawaiian waters. After Smith-Vaniz (1986b),
Smith-Vaniz et al. (1990), Myers (1991).
S p e c i e s Schooling; habitat
S c o m b e r o i d i n i
Scomberoides lysan Often schools near surface; clear lagoons to offshore reef areas, from sur-
face to 100 m
N a u c r a t i n i
Elagatis bipinnulata May form large schools; from surface to depths of 150 m or more in clear
o ffshore waters, occasionally in reef areas as shallow as 3 m
Naucrates ductor Small groups, pelagic, in open waters, commensal relationship with sharks
and other large fishes
Seriola dumerili Solitary or small to moderate schools; reefs or at dropoffs, from 8 to at
least 100 m
Seriola lalandi Large shoals; pelagic and epibenthic, to 50 m
Seriola rivoliana Outer reef slopes to depths of 160 m or more, perhaps more oceanic than
other S e r i o l as p p .
C a r a n g i n i
Alectis ciliaris Adults usually solitary; near bottom in shallow coastal waters to 60 m or
more; juveniles pelagic and drifting
Atule mate Schools; inshore waters to 50 m
Carangoides equula Benthic, 100 to 200 m
Carangoides ferdau Coastal waters adjacent to sandy beaches and reefs, to 60 m
Carangoides orthogrammusSmall schools; sandy channels of lagoons, seaward reefs from 3 to 168 m
Caranx ignobilis Juveniles in small schools over inshore sandy bottoms, adults usually soli-
tary and range widely over reefs to depths of 80 m
Caranx lugubris Singly or small groups; along outer reef slopes and offshore banks at 12 to
354 m
Caranx melampygus Singly or small groups; enter channels and inshore reefs to feed
Caranx sexfasciatus Singly or small groups; in deep channels and outer reefs to 96 m
Decapterus macarellus Schooling; open water and insular habitats, sometimes at surface, occa-
sionally over outer reefs, usually in waters 40-200 m
Decapterus macrosoma Schooling; in waters 30-170 m
Decapterus muroadsi U n c e r t a i n
Decapterus tabl Schooling; midwater or near bottom, in waters 200-360 m
Gnathanodon speciosus Small schools; adults in seaward reefs, young accompany sharks and
other large fish
Pseudocaranx dentex Schools; banks and inshore slopes in waters 80-200 m
Selar crumenophthalmus Small to large schools; adults generally offshore in waters to 170 m
Uraspis helvola Schools; dropoff areas from 30 to 65 m
Uraspis secunda Small schools; surface, benthic, and pelagic

Predation
Berry (1959) notes that young jack crevalles are food for a number of surface-feeding carni-
vores. Juveniles of varying sizes were found in the stomachs of fishes as indicated in Table 9.
The prey items were apparently not identified more specifically than “jack crevalle.”
Nakamura (1972) makes reference to studies which identified unspecified carangids in the
stomach contents of black marlin (Makaira indica) in the equatorial Pacific, and Decapterusin
the same predators off Japan.
In Hawai‘i, halalü(Selar crumenophthalmus) ranging in size from four to seven inches are sub-
ject to predation by ulua and kawakawa (Euthynnus affinis) when they school in shallow water
(Gosline and Brock 1960). As mentioned previously, ‘öpelu(Decapterus macarellus) have
been found in the stomachs of Caranx ignobilisand Seriola dumeriliin the Northwestern
Hawaiian Islands (Sudekum et al. 1991, Humphreys 1986).
Smith-Vaniz and Staiger (1973) describe the venom apparatus of the leatherback (Scomberoides
lysan). The first seven dorsal fin spines and first two anal spines are associated with venom
glands, with the anal spines being the most venomous. The anal spines can be locked outward
and inflict painful stings. This is an obvious antipredatory trait, although the identity of the
leatherback’s potential predators is not specified.
Hobson (1972, 1974) states that schools of nocturnal carangids, which spend the day hovering
in exposed areas near the reef, are targets of larger crepuscular predators, but does not elabo-
rate. Presumably, they may include larger carangids. In general, it’s probable that carangids
are potential prey for any piscivore large and fast enough to capture them.
29
Table 9
Predators on carangids, and maximum sizes of jacks found in stomach contents.
After Berry (1959).
Predator Max. prey size (mm)
Barracuda, Sphyraena barracuda 167
Greater amberjack, Seriola dumerili 90
Dolphinfish, Coryphaena hippurus 140
Tripletail, Lobotes surinamensis 65
Skipjack tuna, Katsuwonus pelamis 77
Little tunny,Euthrynnus alletteratus28
Blackfin tuna, Thunnus atlanticus 29
Yellowfin tuna, Thunnus albacares 90

Morbidity and Mortality
Thompson and Munro (1974) state that there is “no evidence” regarding factors causing mor-
bidity and mortality in carangids, and their major predators are unknown. The flesh of large
kähala(Seriola dumerili) is often infested with larval tapeworms, a condition that occurs in
other geographic areas as well. It is uncertain what effects these parasites have on the fish.
Kähalaand a number of other carangids are also prone to ciguatoxicity here and in other parts
of the world, but the effects of the toxin on the fish is also unknown.
Kawamoto (1973) estimated natural mortality rate of akule (Selar crumenophthalmus). The
mean monthly survival rate was calculated to be 66.3%, so mortality was 33.7%. Estimated
annual survival was then 0.7%, and annual mortality 99.3%. No other data on morbidity or
mortality in carangids could be found.
Ecologic Interactions
Since many carangids obtain their food from reef areas, they would be expected to compete
with other reef carnivores (Thompson and Munro 1974). Sudekum et al. (1991) estimated aver-
age daily consumption rates for Caranx ignobilisand C. melampygus, then extrapolated to cal-
culate individual food rations for each species per year. Individual C. ignobiliswere estimated
to consume an average of 150.69 kg/yr, and C. melampygusan average of 47.82 kg/yr. Based
on population estimates at French Frigate Shoals of 130,000 and 230,000, respectively, this
resulted in a total consumption of 30,600 tons per year by just these two species.
The estimates, however crude they may be, emphasize the importance of these two species as
apex predators both to their own populations and those of their prey.Alow overlap of specific
diet items indicates they do not compete much with each other.The authors suggest the two
species represent one of the most important top-level trophic paths in the French Frigate Shoals
ecosystem, and probably many others. Their combined predation pressure exceeds the com-
bined estimate for the three major shark species in the area (gray reef (Carcharhinus ambly-
rhynchos), Galapagos (C. galapagensis), and tiger (Galeocerdo cuvier), as determined by De
Crosta et al. (1984)) by a factor of about 40.
No further information could be found on the ecology of local carangids. All provide food for
other fishes at some point in their lives, and the smaller schooling species are probably impor-
tant food items for larger carnivores, as noted earlier. It is likely that predation by reef-associat-
ed carangids upon new recruits, juveniles, and adult prey items plays an important role as
described by Hixon (1991) for piscivores in general structuring abundance, distribution, and
local diversity in reef fish communities, and reducing intra- and inter-specific competition as a
result of postsettlement mortality.
30

Economic Importance
Carangids have received attention in various parts of the world during the past 30 years or so as
potential aquaculture candidates. Most work in the United States has focused on the Florida
pompano (Trachinotus carolinus), permit (Trachinotus falcatus), and palometa (Trachinotus
goodei). The white trevally (Pseudocaranx dentex) is extensively cultured in net cages in
Japan, and 1990 production was 959 metric tons (Ogawa 1992). In this species, parasitic and
viral infections present problems that must be overcome (Ogawa 1992, Nguyen et al. 1996).
Marino et al. (1995) note that the greater amberjack (Seriola dumerili) has considerable aqua-
culture potential due to its rapid growth rate and commercial value, but neither sexual maturity
nor gamete release has yet been achieved in captivity.
In Hawai‘i, spawning of akule (Selar crumenophthalmus) has been accomplished in captivity,
but larval survival to day 75 was less than 3% (Iwai et al. 1996). The Oceanic Institute on
O‘ahu has recently developed techniques for culture of ‘ömilu(Caranx melampygus)
(Ostrowski pers. comm.).
As indicated earlier, state law prohibits the taking of ulua and papio under seven inches in
length for personal consumption. Sale of ulua and päpiois prohibited for fishes under a pound.
There is, therefore, no aquarium fishery for carangids in Hawai‘i, and the small specimens seen
in some local pet stores are imported.
Anumber of carangids have important commercial fisheries in Hawai‘i. Akule are generally
taken by handline and surround net. The efficiency of the latter method increased considerably
after World War II with the introduction of spotter aircraft, a technique still in use. ‘Öpelu
(Decapterus macarellus) are considered an excellent food fish, and are taken by handline and
hoop nets (Shiota 1986a).
Commercial fishing for ulua aukea (Caranx ignobilis), ‘ömilu, and Pseudocaranx dentexis
done mostly by handlines and traps. Commercial catches of carangids for 1998 as reported to
the Division of Aquatic Resources are summarized in Table 10.
The palatability of carangids in general ranges from excellent to poor, and some species caught
by recreational fishers are not eaten (McClane 1974, Porter pers. comm.). Most local species
are considered good eating, although ciguatera poisoning is a concern, especially among the
larger reef-associated species. Kähala(Seriola dumerili) are excellent fighters, but are invari-
ably released due to their reputation as a species particularly prone to ciguatoxin. The follow-
ing information on recreational fishing comes from Rizzuto (1977, 1983), Sakamoto (1985,
1988), and personal observations.
31

Kähalaand kamanu (Elagatis bipinnulata) are often caught with shorecasting gear, especially in
areas where the bottom drops off rapidly into deep water.They are occasionally caught by
trolling, but mostly by bottom fishing from boats.
Lai (Scomberoides lysan) are popular sportfish on ultralight tackle, whipping lures or baits just
below the surface. They are not highly prized as food fish, but their tough skin (hence the
name “leatherback”) is often stripped off, dried, and used for trolling lures.
As mentioned previously, young akule, or halalü, frequently form large schools close to shore
from July through December, often entering bays and harbors. During the peak months of July
and August recreational anglers are out in force with fiberglass or bamboo handpoles. Adult
akule are caught handlining from boats at night, using lights to attract plankton and eventually
the fish. ‘Öpeluare usually taken by sportfishers jigging from boats. Both akule and ‘öpeluare
good eating, but are also used as bait, either cut or whole, for larger fishes.
Ulua and päpioare highly sought after by recreational fishermen, and ulua are considered by
many to be the ultimate shoreline sportfish. Ulua are caught with slide-bait from rocky cliffs,
by whipping plugs and bait from boats or shore, and by bottom fishing from boats. They are
also taken by spearing, as was the case with the Hawai‘i record 191-pounder.Päpioare taken
32
Table 10
Commercial catches and sale of carangids from Hawaiian waters, calendar year 1998, sorted
by value. From Division of Aquatic Resources (1998).
Pounds Pounds
Species Landed Sold Value ($)
Akule and halalü,Selar crumenophthalmus1,311,201 1,177,018 1,619,594
‘Öpelu,Decapterus macarellus 234,376 221,678 375,971
Ulua/päpio(misc.) 39,770 31,020 62,010
“Buta” ulua, Pseudocaranx dentex 40,522 37,953 53,643
Ulua aukea, Caranx ignobilis 9,280 8,614 12,291
Ulua “papa,” Carangoides orthogrammus 3,878 3,438 6,923
“Dobe” ulua, Uraspis helvola 4,118 4,118 6,903
Kamanu, Elagatis bipinnulata 3,394 2,727 3,836
Ulua kihikihi, Alectis ciliaris 2,052 1,191 2,343
‘Ömilu,Caranx melampygus 1,737 1,125 2,139
‘Ömaka,Atule mate 237 220 743
Pake ulua, Caranx sexfasciatus 297 278 345
‘Öpelu“mama,” Decapterus muroadsi 32 26 51

with lighter tackle, mostly by whipping plugs, soft plastics, and bait, but also by dunking bait,
trolling, and spearing.
In part because Hawai‘i does not require a marine recreational fishing license, there is no direct
data on the economic value of carangids in the state’s recreational fishery. Most sportfishers
target them at one time or another.The casting gear required for ulua fishing is expensive, and
considering the number of ulua fishermen seen on shorelines around the state, their contribution
to the local economy is significant. Gaffney (pers. comm.) estimates the value of the recre-
ational ulua fishery on Hawai‘i’s economy at over $31 million annually, far exceeding the eco-
nomic impact contributed by commercial fishing of carangids.
p
33
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37
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