MORPHOLOGY, SYSTEMATICS, AND BIOLOGY OF

THE SPANISH MACKERELS (SCOMBEROMORUS, SCOMBRIDAE)

1
BRUCE B. COLLE'ITE AND JOSEPH L. Russo 2

ABSTRACT

The Spanish mackerels and seerfishes of the genus Scomberomorus constitute the most speciose
group of the 44 genera in six families that comprise the suborder Scombroidei. As in higher scom-
brids, Scomberomorus, Acanthocybium, and Grammatorcynus have a well-developed median keel
on the caudal peduncle, but there is no bony support as is present in the Sardini and Thunnini.
Acanthocybium and Scomberomorus share 17 osteological characters and are considered sister-
groups. The relationships of Grammatorcynus are not clear but it is clearly more primitive than
Scomberomorus; therefore, we have used it as the outgroup for a cladistic analysis of Scomberomorus.
Scomberomorus differ from all other scombrids in having a spatulate anterior extension of the
vomer. There are 18 species in the genus, nearly 40% of the 49 species of scombrids: Eastern At-
lantic-tritor (Cuvier); western Atlantic-brasiliensis Collette, Russo and Zavala-Camin, cavalla
(Cuvier), maculatus (Mitchill), and regalis (Bloch); eastern Pacific-concolor Lockington and
sierra Jordan and Starks; and Indo-West Pacific-commerson (Lacepoo.e), guttatus (Bloch and
Schneider), koreanus (Kishinouye), lineolatus (Cuvier), munroi Collette and Russo, multiradiatus
Munro, niphonius (Cuvier), plurilineatus Fourmanoir, queenslandicus (Macleay), semifasciatus
(Macleay), and sinensis (Lacepede). A cladistic analysis of 58 characters shows six monophyletic
species-groups in Scomberomorus. The sinensis group is monotypic and is defined by the presence of
an abrupt downward curve in the lateral line under the first dorsal fin and by its retention of a swim
bladder. The commerson species-group contains commerson, niphonius, queenslandicus, and cavalla
and is defined by the presence of an intercalar spine of at least moderate length. Scomberomorus
cavalla and S. commerson share two additional specializations, the pterosphenoid bones are close
together and the lateral line curves abruptly downward under the second dorsal finlets. The munroi
species-group is monotypic and is defined by the loss of the anterior process on the outer surface of
the head of the maxilla. The semifasciatus species-group contains semifasciatus, plurilineatus, and
lineolatus, and is defined by the presence of a greatly expanded posterior end of the maxilla. Scom-
beromorus lineolatus and S. semifasciatus share an additional specialization, a wide parasphenoid,
but this character state appears independently in several other lines. The guttatus species-group
contains guttatus, multiradiatus, and koreanus and is defined by a high supraoccipital crest. Auxil-
iary branches extend off the anterior part of the lateral line in S. guttatus and S. koreanus. The
regalis species-group contains regalis, tritor, maculatus, concolor, sierra, and brasiliensis and is
defined by the presence of nasal denticles. All but the most primitive species in this group (S. tritor)
have an artery arising from the fourth left epibranchial artery. The four most advanced species (all
except tritor and maculatus) have developed a long posterior process on the pelvic girdle. The three
most advanced species (sierra, brasiliensis, and regalis) have a coeliaco-mesenteric shunt connect-
ing the fourth right epibranchial artery with the coeliaco-mesenteric artery.

The purposes of this paper are to define the 18 nus, and rely on previous work by Kishinouye
species of Scomberomorus, to clarify their rela- (1923), Munro (1943), Mago Leccia (1958), and
tionships, and to assess the systematic position Devaraj (1977).
of Scomberomorus within the Scombridae. The The Spanish mackerels have been placed by
methods used are similar to those of Collette and Collette and Chao (1975) and Collette and Russo
Chao (1975) in a revision of the bonitos and of (1979) in a tribe (the Scomberomorini) along with
Gibbs and Collette (1967) in a revision of Thun- Acanthocybium and Grammatorcynus, interme-
diate between the more primitive mackerels
1 National Marine Fisheries Service, Systematics Laboratory, (Scombrini) and the more advanced bonitos (Sar-
National Museum of Natural History, Washington, DC 20560. dini). Acanthocybium is clearly the specialized
2National Marine Fisheries Service, Systematics Laboratory,
National Museum of Natural History, Washington, DC 20560, sister group of Scomberomorus, but the phylo-
and Department of Biological Sciences, The George Washington genetic position of Grammatorcynus has been
University, Washington, DC 20006; present address: Office
of Information Research Resource Management, Smithsonian unclear.
Institution, Washington, DC 20560. Until recently, the number of valid species of
Manuscript accepted November 1983. 545
FISHERY BULLETIN: VOL. 82, NO.4, 1984.
 FISHERY BULLETIN: VOL. 82. NO. 4

Scomberomorus has been in doubt. In his revision cies of Scomberomorus based on a cladistic
of Australian species, Munro (1943) recognized 15 analysis of characters described in the first part,
species in the world (excluding Cybiosarda ele- using Grammatorcynus as the plesiomorphic
gans, a bonito, and Lepidocybium flauobrunneum, out-group.
a gempylid). Fraser-Brunner (1950) recognized
only nine species, placing five valid species in
synonymy. In the course of this revision, we have MATERIAL
discovered two previously undescribed species, S.
brasiliensis (Collette et al. 1978), which was con- The material examined is listed by general lo-
fused with S. maculatus, and S. munroi (Collette cality under four or five headings in the accounts
and Russo 1980), which was confused with S. for each of the 18 species of Scomberomorus.
niphonius. Comparative material of Acanthocybium and
Emphasis was placed on obtaining fresh or Grammatorcynus is listed at the end of this
frozen specimens for dissection from several pop- section. The numbers under these headings are
ulations of each species. Standard counts and not additive but are included to give some degree
measurements were taken, color pattern was of confidence in the morphological data presented
recorded, and a search made for parasitic cope- in the body of the paper. "Total specimens" is the
pods. Results of the copepod study have been total number of individuals examined whether
reported by Cressey and Cressey (1980), and preserved, dissected, or skeletonized. "Dissected"
analysis of these data from a host-parasite point are fresh or frozen specimens for which data on
of view has been completed (Cressey et al. 1983; the viscera and usually other characters were
Collette and Russo 1985). The viscera were ex- recorded. Specimens were subsequently made in-
amined and illustrated in situ following remov- to skeletons. "Measured and counted" includes
al of an oval portion of the ventral body wall. specimens that were subsequently dissected as
The viscera then were removed and drawings well as the preserved museum specimens used for
were made of the liver and other selected organs. detailed morphometric and meristic examination.
The kidneys and anterior parts of the arterial "Counts only" are additional museum specimens
system then were drawn. Counts of ribs and used only for meristic examination. "Skeletons"
intermuscular bones were made and the speci- refer to all the skeletal material examined, both
men was then skeletonized, facilitated by immer- specimens that were dissected and additional
sion in hot water. skeletal material already in museums. Asterisks
The base measurement for morphometric com- indicate type-specimens of nominal species.
parisons of fresh, frozen, and preserved specimens Material was examined from the following
was millimeters fork length (mm FL). institutions:
This paper is divided into three major parts.
The first part contains descriptions and illustra- AMNH American Museum of Natural His-
tions of morphometry, meristic characters, soft tory, New York
anatomy, and osteology of the species of Scom- AMS Australian Museum, Sydney
beromorus. Comparisons with Acanthocybium ANSP Academy of Natural Sciences,
solandri and Grammatorcynus bilineatus are in- Philadelphia
cluded. All references to Grammatorcynus in this BMNH British Museum (Natural History),
paper refer to G. bilineatus. The validity of the London
second species, G. bicarinatus, was only estab- CAS California Academy of Sciences, San
lished recently (Collette 1983). The second part Francisco
comprises separate species accounts including CSIRO CSIRO Marine Biological Laborato-
synonymy, types of nominal species, diagnosis ry, Cronulla, N.S.W, Australia
(based on characters from the first section), de- DASF Department of Agriculture, Stock,
scription, size, color pattern, summaries of pub- and Fisheries, Port Moresby, Pap-
lished information on biology and interest to ua New Guinea
fisheries, geographic distribution, and material FMNH Field Museum of Natural History,
examined. The most important references to each Chicago
species are marked with asterisks in the syn- HUMZ Laboratory of Marine Zoology,
onymies. The third part is an analysis of the Hokkaido University, Hakodate,
relationships of Acanthocybium and the spe- Hokkaido

546
COLLETTE and RUSSO: SPANISH MACKERELS

GCRL Gulf Coast Research Laboratory and (2); Australia (1); Marshall Is. (1); E
Museum, Ocean Springs, Miss. Pacific (2).
LACM Los Angeles County Museum of counts: 36.
Natural History, Los Angeles diss.: 11 (943-1,420): W Atlantic (7); Indian O.
MCZ Museum of Comparative Zoology, (3); Revillagigedos (1).
Harvard
MNHN Museum National d'Histoire Natu- Grammatorcynus bilineatus. -Total 52 (23.5-575
relle, Paris mmFL).
MPIP Museu de Pesca do Instituto de Pes- meas.: 34 (226-575): Red Sea (13, *Thynnus
ca, Santos bilineatus); Indian Ocean ? (1); Anda-
MSUF Museo de La Specola, Universita di man Sea (3); Celebes (1); New Guinea
Firenze, Florence (3); Australia (8); Philippine Is. (5,
MZUSP Museu de Zoologia da Universidade *Nesogrammus piersoni); Solomon Is.
de Sao Paulo, Sao Paulo (1); Caroline Is. (3); Marshall Is. (8);
NHMV Naturhistorisches Museum, Vienna Fiji (2).
NMC National Museum of Natural Sci- counts: 44.
ences, Ottawa diss.: 10 (382-453): Indian Ocean? (1); Timor
QM Queensland Museum, Brisbane Sea (2); Bismarck Arch. (1); Marshall Is.
RMNH Rijksmuseum van Natuurlijke His- (2); Queensland, Australia (4).
torie, Leiden
ROM Royal Ontario Museum, Toronto Grammatorcynus bicarinatus. -Total 9 (306-825
RUSI J. L. B. Smith Institute of Ichthyolo- mmFL).
gy, Rhodes University, Grahams- meas.: 9 (306-825): Western Australia (5);
town, South Africa Queensland (4).
SAM South African Museum, Capetow~ counts: 9.
SIO Scripps Institution of Oceanography, diss.: 2 (521 and 563): Queensland.
La Jolla, Calif.
TABL Miami Laboratory (formerly Tropi-
cal Atlantic Biological Laborato- KEY TO GRAMMATORCYNUS,
ry), NMFS, Miami, Fla. [Most ACANTHOCYBIUM, AND
specimens now at UE] SCOMBEROMORUS
UDONECI Universidad de Oriente, Nueva Es-
parta, Centro de Investigaciones, la. Two lateral lines, the lower joining the
Venezuela upper behind the pectoral fin base and
UF Florida State Museum, University at the caudal fin base; interpelvic pro-
ofFlorida, Gainesville cess single; teeth in jaws slender, coni-
UMMZ University of Michigan Museum of cal, not compressed; vertebrae 31 ....
Zoology, Ann Arbor .................... Grammatorcynus 2
USNM United States National Museum, lb. One lateral line; interpelvic process
Washington, D.C. double; teeth in jaws strong, com-
WAM Western Australia Museum, Perth pressed, almost triangular or knife-
ZMA Zoological Museum, Amsterdam like; vertebrae 39-64 3
ZMH Zoologisches Institut und Zoolog-
isches Museum, Hamburg 2a. Gill rakers 14-15; small eye, 3-4% FL;
ZMK Zoological Museum, Copenhagen frequently with small dark spots on
ZSI Zoological Survey of India, Calcutta lower sides of body .
..... G. bicarinatus (Quoy and Gaimard)
Acanthocybium solandri.-Total 47 (536-1,500 2b. Gill rakers 19-24; large eye, 7-9% FL;
mmFL). seldom with dark spots on sides of
meas.: 26 (536-1,500): W Atlantic (8); St. He- body G. bilineatus (Riippell)
lena (1); S. Africa (3); Indian Ocean (4);
Caroline Is. (6); 'fuamotu Is. (1); E Pa- 3a. Snout as long as rest of head; no gill
cific (3). rakers; 23-27 spines in first dorsal fin;
heads: 8 (202-380): Bahama Is. (1); St. Helena posterior end of maxilla concealed un-

547
 FISHERY BULLETIN: VOL. 82, NO.4

der preorbital bone; vertebrae 62-64 .. lOa. First dorsal fin black only on first 5-7
....... Acanthocybium solandri (Cuvier) interspinous membranes, white pos-
3b. Snout much shorter than rest of head; teriorly; intestine straight, with no
gill rakers 1-27; 12-22 spines in first folds; total vertebrae 48-50 .
dorsal fin; posterior end of maxilla · S. niphonius (Cuvier)
exposed; vertebrae 41-56 . lOb. First dorsal fin black to, or almost to,
...................... Scomberomorus 4 posterior end; intestine with 2 loops
and 3 limbs; total vertebrae 50-52 ....
4a. Lateral line abruptly curving down be- · S. munroi Collette and Russo
low first or second dorsal fin; verte-
brae 41-46 5 lIa. Lateral line with many small auxiliary
4b. Lateral line straight or descending gra- branches anteriorly 12
dually posteriorly; vertebrae 44-56 .... 7 lIb. Lateral line without auxiliary branches
or with only a few anteriorly 13
5a. Lateral line abruptly curving down be-
low first dorsal fin; total gill rakers 12a. Dorsal fin spines 15-18, usually 16 or
on first arch 12-15; caudal vertebrae more; intestine with 2 loops and 3
21-22 S. sinensis (Lacepede) limbs; total vertebrae 47-52, usually
5b. Lateral line abruptly curving down be- 48 or more; head longer, 20.2-21.5%
low second dorsal fin; total gill rakers FL; body depth less, 22.8-25.2% FL ...
on first arch 2-13; caudal vertebrae · S. guttatus (Bloch and Schneider)
23-27 .. , 6 12b. Dorsal fin spines 14-17, usually 15 or
fewer; intestine with 4 loops and 5
6a. Total gill rakers on first arch 7-13, usual- limbs; total vertebrae 46-47, usually
ly 9 or more; spines in first dorsal fin 46; head shorter, 19.7-20.4% FL; body
12-18, usually 15 or fewer; precaudal depth greater, 24.4-26.7% FL .
vertebrae 16-17 S. cavalla (Cuvier) · S. korean us (Kishinouye)
6b. Total gill rakers on first arch 3-8, usual-
ly 6 or fewer; spines in first dorsal fin 13a. Sides of body with spots and at least one
15-18, usually 16 or more; precaudal stripe, the stripes may be short, wavy
vertebrae 19-20 . or interrupted 14
............... S. commerson (Lacepede) 13b. Sides of body without any stripes, spots
usually present 16
7a. Total gill rakers on first arch 21-27; no
bars on body .... S. concolor Lockington 14a. One long stripe on sides with spots or
7b. Total gill rakers on first arch 1-18; spots, interrupted lines above and below the
bars, or other markings usually pres- stripe; total vertebrae 47-48, usually
ent on sides of body 8 48; total gill rakers on first arch 12-18,
usually 15 or more .... S. regalis (Bloch)
8a. Anal fin rays 25-29; second dorsal fin 14b. Sides with several short stripes; total
rays 21-25, usually 23 or more; gill vertebrae 44-47, usually 46; total gill
rakers on first arch 1-4; total verte- rakers on first arch 9-15, usually 14
brae 54-56; no pattern on body . or fewer 15
.. S. multiradiatus Munro
8b. Anal fin rays 15-24; second dorsal fin 15a. Sides with a series of short straight
rays 15-24; total gill rakers on first stripes and few if any spots; total gill
arch 3-18; total vertebrae 44-53; sides rakers on first arch usually 11 or
of body usually with spots or other fewer; second dorsal fin rays 15-19,
markings 9 usually 18 or fewer; distance from 2D
origin to caudal base 46.2-54.5% FL,
9a. Dorsal fin spines 19-22, usually 19 or x 50.0% S. lineolatus (Cuvier)
more · 10 15b. Sides with a series of short wavy mark-
9b. Dorsal fin spines 13-19, usually 18 or ings plus many small spots; total gill
fewer 11 rakers on first arch usually 12 or more;

548
COLLE'ITE and RUSSO: SPANISH MACKERELS

second dorsal fin rays 19-21, usually Lateral Line

20 or more; distance from 2D origin to
caudal base 51.8-57.5% FL, i 54.8% .. In most species of Scomberomorus, the lateral
............ S. plurilineatus Fourmanoir line runs posteriorly above the pectoral fin and
theh gradually descends to the middle ofthe body
16a. Sides with bars or large spots, larger at about the level of the second dorsal fin. Gram-
than the diameter of the eye ..... , ... 17 matorcynus differs from Bcomberomorus, Acan-
16b. Sides with small round spots, about the thocybium, and all other members of the family
diameter of the eye, orange colored by .having a second lateral line that joins' the.
in life 19 upper lateral line at a right angle behind the
pectoral fin base and then courses ventrally and
17a. Sides with large spots or blotches; total posteriorly along the ventral surface of the body
gill rakers on first arch 3-9, usually 7 to join the dorsal lateral line on the caudal
or fewer S. queenslandicus Munro peduncle. In Acanthocybium and three species of
17b. Sides plain or with bars; total gill rakers Scomberomorus, the lateral line moves abruptly
of first arch 6-15, usually 9 or more .. 18 downward under the first or second dorsal fin.
The abrupt downward curve is under the first
18a. First dorsal fin spines 13-15; second dor- dorsal fin in Acanthocybium and S. sinensis
sal fin rays 19-22, usually 20 or more; (see Figure 68); it is under the second dorsal in
total gill rakers on first arch 6-13, S. cavalla and S. commerson (see Figures 50
usually 11 or fewer; total vertebrae and 52).
44-46, usually 45; base first dorsal fin Scomberomorus guttatus and S. koreanus dif-
17.0-23.6% FL . fer from other members of the genus in having
.............. S. semifasciatus (Macleay) many fine branches from the anterior part of the
18b. First dorsal fin spines 15-18, usually 16 lateral line, both dorsally and ventrally (see
or more; second dorsal fin rays 16-19, Figures 54 and 56). Acanthocybium and S. ni-
usually 17; total gill rakers on first phonius (see Figure 62) may have branches from
arch 12-15; total vertebrae 46-47, usu- the lateral line but they are not as numerous or
ally 46; base first dorsal fin 23.8- distinct.
30.4% FL S. tritor (Cuvier)
Nasal Dentides
19a. Total vertebrae 51-53; second dorsal fin
rays 17-20, usually 18 or more . Nasal denticles (Fig. la, b) are small general-
................. S. maculatus (Mitchill) ized teeth found within the olfactory chamber
19b. Total vertebrae 46-49; second dorsal fin on the medial surface surrounding the posterior
rays 15-19, usually 18 or fewer ... " .. 20 nares and on the skin covering the anterior
surface of the lateral ethmoid. Nasal denticles
20a. Pectoral fin rays 21-24, usually 22 or are similar to the small villiform teeth present
more; pelvic fin short, 2.9-5.9% FL, i within the mouth cavity and adjoining regions of
4.5% .... S. brasiliensis Collette, Russo, stomadeal origin and on the skin covering the
and Zavalla-Camin cleithrum (Fig. le, d) and on the isthmus where
20b. Pectoral fin rays 20-24, usually 21 or they are contacted by the opercular membrane.
fewer; pelvic fin longer, 3.2-6.4% FL, These teeth typically fit into sockets in pads of
i 5.3% ..... S. sierra Jordan and Starks fine spongelike bone. They point posteriorly and
are aligned with presumed flow of water from the
anterior naris through the olfactory chamber and
COMPARATIVE MORPHOLOGY out the posterior naris. Nasal denticles were
found only in the six species of the Scomberom-
The morphological characters useful for distin- orus regalis species-group (brasiliensis, concolor,
guishing the species of Scomberomorus and for maculatus, regalis, sierra, and tritor). Nasal
evaluating their phylogenetic relationships are denticles are not present in Acanthocybium or
divided into six categories: lateral line, nasal Grammatorcynus. We do not know their function
denticles, morphometry, meristics, soft anatomy, and are not aware of such structures in other
and osteology. fishes.

549
01
01
o
a b

c d

i::l
til

i
CJ
r
.[15

FIGURE I.-Scanning electron photomicrographs of nasal dentides (a-b) and villiform teeth over the deithrum (c-d) in Scomberomorus sierra, GulfofCalifornia, 353 mm ...~
FL, USNM 217368. a, c. 50x. b, d. 250x.
COLLETTE and RUSSO: SPANISH MACKEREUl

Morphometric Characters ance (ANCOVA) was carried out on the regres-

sions of body parts on fork length. Results are
In addition to fork length, 26 measurements reported, under a section entitled Geographic
routinely were made on all specimens destined to Variation, in 11 of the 18 species accounts. Tests
be dissected, to insure that these data would of significance were made by Newman-Keuls
be available if needed. Preserved material also Multiple Range Test.
was measured until an adequate sample was
obtained. Measurements follow the methods of Meristic Characters
Marr and Schaefer (1949) as modified by Gibbs
and Collette (1967) and Collette and Chao (1975). Countable structures are of special, value sys-
Morphometric characters can be used to separate tematically because they are relatively easy to
species and populations within species. Tables record unambiguously and are easy to summarize
showing the 26 characters as thousandths of fork in tabular fashion. Meristic characters that have
length and 8 characters as thousandths of head proved valuable systematically in Scomberomo-
length are presented in the systematic section of rus include numbers of fin rays (first dorsal
the paper (see Tables 13-30). Most of the charac- spines, second dorsal rays, dorsal finlets, anal
ters are best used at the species level; therefore, rays, anal finlets, and pectoral rays), gill rakers,
only a summary table of the means of proportions teeth on the upper and lower jaws, vertebrae
(Table 1) is presented in this section. Where there (precaudal, caudal, and tota1), and lamellae in
was sufficient material from two or more poten- the olfactory rosettes. Olfactory lamellae are dis-
tially different populations, analysis of covari- cussed as the next to last section under soft

TABLE I.-Morphometric comparison of the species of Scomberomorus. Means as thousandths offork length or head length. Species
arranged alphabetically by the first three letters of their names. Ranges for the species given in Tables 13-30,
Character bra cay com con gut kor lin mac mul mun nip plu que reg sem sie sin tn Min. spp. Max. spp.
Fork length
Snoul-A 538 539 542 524 517 493 507 536 505 546 563 502 525 548 506 537 584 533 493kor 584 sin
Snout-20 511 506 510 507 481 467 501 503 477 528 536 473 501 521 472 510 559 513 467kor 559 sin
Snout-10 242 258 243 236 239 242 252 241 249 222 248 221 234 255 245 241 291 248 221 mun, 291 sin
plu
Snout-P2 253 258 257 242 251 248 245 257 243 249 263 233 251 265 250 252 290 266 233plu 290 sin
Snout-P, 219 232 237 209 209 210 212 217 213 201 225 193 229 234 219 221 258 222 193plu . 258sin
P,-P2 108 106 96 100 106 114 93 110 102 105 105 103 99 109 105 104 113 111 93 lin 114kor,
sin
Head length 213 223 229 202 205 208 206 212 208 198 216 193 220 223 213 212 255 217 193piu 255 sin
Max. body depth 198 191 187 187 209 237 181 197 229 190 172 206 188 197 211 190 218 206 172 nip 237kor
Max. body width 82 89 94 89 93 100 97 .91 95 100 84 97 101 91 94 84 102 90 82 bra 104 mun
P, length 123 129 122 125 109 133 139 129 131 109 111 123 120 126 147 123 158 134 109 gut, 158 sin
mun
P21ength 45 65 56 50 59 60 55 52 40 54 68 51 55 56 50 53 83 60 40mul 83 sin
Pz insertion-vent 273 271 273 261 251 227 241 263 247 281 285 243 254 267 237 267 273 250 237sem 285 nip
Pztip-vent 225 212 217 212 191 184 185 211 207 225 218 186 198 210 187 222 189 190 184 kor 225 bra,
mun
Base 10 263 245 261 254 235 218 231 256 216 307 282 240 263 257 210 260 260 262 210 sem 307 mun
Height 20 117 109 103 111 131 166 124 125 167 112 98 148 114 114 159 123 145 126 98 nip 167 kor,
mul
Base 20 118 106 104 127 141 160 114 128 178 115 113 128 113 114 138 120 121 122 104com 178mul
Height anal 114 106 100 107 127 160 117 118 164 108 97 135 112 112 156 117 145 125 97 nip 184mul
Base anal 113 108 100 134 133 154 122 123 216 105 107 125 108 110 145 119 122 120 100 com 216 mul
Snout (fleshy) 82 87 89 72 72 70 81 80 77 77 81 67 86 87 81 79 97 81 67plu 97 sin
Snout (bony) 72 79 81 63 84 62 74 70 67 70 75 59 80 79 72 70 91 72 59plu 91 sin
Maxilla length 123 132 131 113 108 111 113 119 125 104 120 96 125 124 119 121 147 123 96plu 147 sin
Postorb~al 95 98 104 96 96 101 91 96 86 90 102 94 102 98 95 98 117 96 86mul 117 sin
Orb~ (fleshy) 37 38 35 32 37 34 32 34 34 25 34 34 31 41 35 33 35 38 25mun 41 reg
OrM(bony) 54 51 49 48 53 50 48 51 52 39 47 45 49 56 51 49 52 53 37mun 56 reg
Interorb~al width 57 60 62 49 59 60 57 56 58 56 57 56 63 58 57 55 63 59 49 con 63 que,
sin
20-caudal 490 477 481 484 527 550 500 487 494 468 465 548 496 480 517 475 445 476 445 sin 550kor
Head length
Snout (fleshy) 386 392 390 353 351 339 395 376 372 386 376 348 391 390 378 371 382 376 339kor 395 lin
Snout (bony) 343 357 355 313 310 301 359 335 321 351 346 306 363 351 339 331 355 333 301 kor 363 que
Maxilla length 581 591 571 555 526 532 547 562 603 521 553 496 568 556 555 570 578 568 496 plu 603 mul
Postorb~a1 446 438 455 476 484 489 442 454 415 456 473 485 463 439 447 461 460 443 415 mul 489kor
Orb~ (fleshy) 175 168 147 159 174 157 156 160 165 134 150 179 142 178 162 158 138 173 129 mun 178 reg
Orb~ (bony) 249 229 211 226 252 238 231 242 252 199 215 232 223 247 238 235 202 245 191 mun 252 gut,
mul
Interorb~al width 270 268 270 241 284 292 276 266 280 282 264 290 286 262 267 253 249 272 241 con 292 kor

551
 FISHERY BULLETIN: VOL. 82, NO.4

anatomy. The other meristic characters are dis- 4 Australian species), Silas (1963, Gramma-
cussed in the relevant osteological sections of the torcynus), Mota Alves and Tome (l967a, S. caval-
paper. la), Mota Alves (1969, S. brasiliensis), Tongyai
(l971a, S. guttatus and S. commerson), and Col-
Soft Anatomy lette and Russo (1979, preliminary review of the
genus).
The relative position, shape, and size of the The anterior end of the liver abuts the trans-
various internal organs provide valuable diag- verse septum anteriorly in the body cavity. The
nostic characters. For purposes of discussion, the liver has three lobes. The left and right lobes are
characters in the soft anatomy are divided into longer than the middle lobe in all three genera
five sections: viscera, vascular system, urogenital (Fig. 4). The right lobe is longest in Scomberomo-
system, olfactory organ, and pharyngeal muscles. rus and Grammatorcynus. The left and right
lobes are about equal in length in Acanthocyb-
VISCERA ium. Two efferent (venous) vessels lead directly
from the anterior surface of the liver into the
Emphasis was placed on the appearance of the sinus venosus in all species. The short esophagus
viscera in ventral view, after removal of an oval leads into the stomach. The stomach is sometimes
segment of the belly wall (Figs. 2, 3). Previous visible in ventral view but this is dependant on
papers on the viscera include Kishinouye (1923, 5 the amount of food present, rather than showing
Japanese species of Scomberomorus, and Acan- differences between species. The pyloric portion
thocybium and Grammatorcynus), Munro (1943, of the intestine arises from the anterior end ofthe

·· ..
·· ..
···.
···
····
···
····

o ~ Q

-
a b c
LIVER I NTESTI N E

f~~~j(~Rj CAECAL MASS t::::::::J GONAD

FIGURE 2.-Viscera in ventral view. a. Scomberomo- W;EI:j:1~1 STOMACH URINARY BLADDER

rus maculatus, Georgia, 290 mm FL. b. Acanthocybium
solandri, Campeche Banks, Mexico, 1,280 mm FL. c.
Grammatorcynus bilineatus, Marshall Is., 424 mm FL. GALL BLADDER

552
COLLETrE and RUSSO: SPANISH MACKEREl.':)

stomach. At this point the main branches of the The efferent branchial (epibranchial) arteries
pyloric caeca join the intestine. The caeca branch and coeliaco-mesenteric artery form a unit at the
and form a dense dendritic conglomeration, the anterior end of the dorsal aorta (Figs. 6, 7). Two
caecal mass. Cells in the pyloric caeca are histo- anterior epibranchials on each side unite to form
logically similar to those in the intestine and a common trunk, and these trunks join as the "Y"
produce enzymes such as lipase, maltase, trypsin, of the aorta beneath the posterior part of the
and pepsin (Mota Alves and Tome 1970). The skull or the first or second vertebra. The posterior
intestine continues posteriorly and its course two epibranchials of each side unite immediately
appears to be species-specific. The intestine may before they join the aorta, usually ventral to the
be a simple straight tube from stomach to anus, second or third vertebra. As the aorta proceeds
have two descending and one ascending arm, or posteriorly, it gives rise to the large coeliaco-
have four bends with three descending and two mesenteric artery on the right side ventral to
ascending arms. The spleen is prominent in ven- the second to fourth vertebrae. The coeliaco-
tral view in most species but is hidden in others. mesenteric artery has two or three main branches
The gall bladder, an elongate tubular sac which which lead to the liver and other viscera.
is usually green, arises from the right lobe of the The postcardinal vein runs along the ventral
liver and usually lies along the first descending surface of the kidney (Fig. 8) from the vicinity
arm of the intestine on the right side. A swim of the first complete haemal arch anteriorly in
bladder is present in Grammatorcynus, Acantlw- the median line to the pectoral region. There it
cybium, and S. sinensis (Fig. 5) but is absent in curves to the right and discharges into the right
the other 17 species of Scomberomorus. Cuvierian duct. Posteriorly, the postcardinal re-
The Spanish mackerels can be divided into ceives a pair of small veins at the level of each
three groups based on the number of folds in vertebra. The postcardinal is composed of two
the intestine. Grammatorcynus, Acanthocybium main branches that join anterior to the Y of the
(Fig. 2b, c), and S. niphonius (Fig. 3k) have a ureter. The main branch leaves the haemal arch
straight gut not folded back on itself. Scomber- dorsally and the small branch runs under the
omorus koreanus (Fig. 3D has four folds and five surface of the kidney from the urogenital area.
distinct arms. The other species all have two folds Five species of Scomberomorus (brasiliensis,
and three long arms (Fig. 3). Collette and Russo concolor, maculatus, regalis , and sierra) have
(1980) used this character to differentiate S. unique specializations of the right and/or left
munroi from the North Pacific S. niphonius. fourth epibranchial arteries (Fig. 7c-g). Each of
The spleen is large and centrally located in these species has an artery arising from the
ventral view in four species: guttatus, koreanus, fourth left epibranchial artery. Other species of
munroi, and plurilineatus. The spleen is smaller thee genus (e.g., S. guttatus and S. tritor, Fig. 7a,
and distinctly on the left side in ventral view in b) lack these specializations. In S. concolor and
seven species: brasiliensis, commerson, lineola- S. brasiliensis this branch is small and goes into
tus, maculatus, multiradiatus, queenslandicus, the muscular tissue surrounding the left dorsal
and sinensis. It is not visible in ventral view in portion of the esophagus (Fig. 7d, D. In S. macu-
Grammatorcynus, Acanthocybium, and seven latus and S. sierra, this branch is large and
species of Scomberomorus: cavalla, concolor, ni- becomes the dorsal left gastric artery (Fig. 7c, e).
phonius, regalis, semifasciatus, sierra, and tri- In S. regalis this branch goes into the left lobe of
tor. the liver (Fig. 7g, hepatic branch). Scomberomo-
rus maculatus and S. sierra have lost the connec-
VASCULAR SYSTEM tion between the dorsal left gastric artery and the
coeliaco-mesenteric artery. It is replaced by a
The only published work on the vascular sys- connection to the fourth left epibranchial artery.
tem of the Spanish mackerels is on Japanese In S. regalis , the left dorsal gastric artery seems
species by Kishinouye (1923). No specialized sub- to have been reduced.
cutaneous vascular system and no cutaneous ar- Scomberomorus brasiliensis, S. sierra, and S.
teries or veins are present as they are in the regalis share a specialization of the right fourth
higher tunas, Thunnini, Auxis to Thunnus (Col- epibranchial artery. In these species an artery
lette 1979). Therefore, this description will be connects the fourth right epibranchial artery
confined to the anterior portion of the dorsal with a branch of the coeliaco-mesenteric artery
aorta and the postcardinal vein. (coeliaco-mesenteric shunt, Fig. 7e-g).

553
 FISHERY BULLETIN: VOL. 82, NO.4

b c
Q
Q

Q Q
FIGURE 3. -Viscera in ventral view of representative specimens of the 18 species of Scomberomo-
rus. a. S. brasiliensis, Belem Fish Market, Brazil, 556 mm FL. b. S. cavallo, off Miami, Fla., 797
mm FL. c. S. commerson, Gulf of Papua, 580 mm FL. d. S. concolor, Gulf of California, 495
mm FL. e. S. guttatus, Gulf of Mannar, 405 mm FL. f. S. koreanus, locality unknown, 812 mm
FL. g. S. lineolotus, Cochin, India, 786 mm FL. h. S. maculotus, St. Andrews Bay, Fla., 323

554
COLLETI'E and RUSSO; SPANISH MACKERELS

MIILIVER

[ill CAECAL MASS

~ INTESTINE

• SPLEEN

o ~ STOMACH

• GAll BLADDER

o E::::J GONAD

mi SWIM BLADDER

mIn FL. i. S. multiradiatus, Gulf of Papua, 272 mIn FL. j. S. munroi, Gulf of Papua, 512 mIn
FL, USNM 219374. k. S. niphonius, Korea, 235 mIn FL. 1. S. plurilineatus, Durban, S. Africa,
490 mIn FL. m. S. queenslandicus, Exmouth Gulf, Western Australia, 466 mm FL. n. S. regalis,
Bahamas, 456 mm FL. o. S. semifasciatus, Gulf of Papua, 715 mIn FL. p. S. sierra, Baja Cali·
fornia, 516 mID FL. q. S. sinensis, China, 711 mm FL. r. S. tritor, Gulf of Guinea, 415 mIn FL.

555
 FISHERY BULLETIN: VOL. 82, NO.4

r-.c--Iateral muscle

peri toneum
urinary bladder-H:..-

swim bladder

blood vessels

ureter - - - - \ f l .

urogenital opening

FIGURE 5.-Swim bladder and urinary bladder in ventral view of

Scomberomorus sinensis (body wall and viscera removed), off Zhoushan
Is., China, 714 mm FL, USNM 220856.

FIGURE 4.-Livers in ventral view. a.

Scomberomorus maculatus, Florida, 712
mm FL. b. Acanthocybium solandri,
Florida, 1,403 mm FL. c. Gramma-
torcynus bilineatus, Marshall Is., 444
mmFL.

556
COLLETIE and RUSSO: SPANISH MACKERELS

3
2
4
3
5 - --~ .
(::1' 4
6 - - --
5
COElIACO-
5 MESENTERIC

a b c
FIGURE 6.-Anterior arterial system in ventral view. Numbers indicate vertebral centra, stippled areas where pharyngeal
muscles originate. a. Scomberomorus multiradiatus, off the Fly River, Gulf of Papua, 272 mm FL. b. Acanthocybium
solandri, Revillagigedos Is., 1,068 mm FL. c. Grammatorcynus bilineatus, Timor Sea, 453 mm FL.

UROGENITAL SYSTEM mesenteries between the gonads in all species

except S. sinensis. Scomberomorus sinensis has a
The only reference to the anatomy of the uro- specialization of the urinary bladder unique to
genital system in Scomberomorus (other than scombrids and, so far as we know, vertebrates in
fishery biology studies of the gonads) is Kishi- general. In this species the urinary bladder has
nouye (1923) on Japanese species and Acantho- become hypertrophied and occupies the space
cybium. The paired gonads lie along the dorsolat- inside the swim bladder (Fig. 5). Acanthocybium
eral body wall and are visible in ventral view in (Fig. 2b) has an elongate urinary bladder that
mature adults. The kidney lies dorsal to the layer extends anteriorly one-third to two-thirds the
of fibrous connective tissue which forms the dor- length ofthe visceral cavity.
sal wall ofthe peritoneum. Anteriorly, the kidney
divides into a pair of narrow projections which OLFACTORY ORGAN
extend along the sides of the parasphenoid and
usually reach the posterior end of the "midridge" Kishinouye (1923) provided a generalized ac-
of the prootic. The anterior ends of the kidney count of the olfactory organ of several scombrids.
surround the origins of the pharyngeal muscles More detailed studies have been made on Scom-
on the vertebral column and usually separate ber scombrus (Burne 1909), Sarda sarda (Treti-
along the middle of the vertebral column. In the akov 1939), Allothunnus {allai (Nakamura and
vicinity of the esophagus, the kidney expands Mori 1966), Katsuwonus pelamis (Gooding 1963),
laterally and forms two projections which may Thunnus (Iwai and Nakamura 1964a; Gibbs and
extend anteriorly to the upper end of the gill slits. Collette 1967), and the bonitos, Sardini (Collette
Posteriorly, near the posterior fifth of the body and Chao 1975). As in other scombrids, the olfac-
cavity, the kidney narrows to an elongate trian- tory cavity in Scomberomorus has a small ante-
gle (Fig. 8). The branches of the "ureter" (meso- rior naris and a slitlike posterior naris. No infor-
nephric ducts) join to form a common trunk just mation on the supplementary sacs, or accessory
before entering the urinary bladder. The ureters olfactory cavity (Iwai and Nakamura 1964a), was
enter the urinary bladder either at its anterior obtained from the present study comparable with
end or on its dorsal surface. The urinary bladder that of Tretiakov (1939), who described three
(Figs. 9, 10) is either ovoid or elongate, depending supplementary sacs (middle, maxillary, and ros-
on degree of inflation, and is located in the tral sacs) in Sarda sarda. The central axis of the

557
 FISHERY BULLETIN: VOL. 82, NO.4

epibranchials

artery

a b c d

coeli acO- me se n te ri c
sh u n t

coeliaco-mesenteric shunt
left gastric artery

e f 9
FIGURE 7.-Anterior arterial system in dorsal view of seven species of Scomberomorus. a. S. guttatus, Pakistan, 545 mm FL. b. S.
tritor, Gulf of Guinea, 494 mm FL. c. S. maculatus, Chesapeake Bay, 312 mm FL. d. S. concolor, Gulf of California, 455 mm FL. e.
S. sierra, Ecuador, 512 mm FL. f. S. brasiliensis, Belem market, Brazil, 588 mm FL, USNM 217557, paratype. g. S. regalis, Bahama
Is., 490 mm FL.

olfactory rosette is located beneath the anterior laminae. Most species of bonitos have 21-39 la-
naris. Leaflike lamellae radiate from the central mellae in each nasal rosette but Gymnosarda
axis and occupy the anterior dorsal third of the unicolor is distinct in the group in having 48-56
olfactory cavity. Gooding (1963) studied the mor- (Collette and Chao 1975:532).
phology and histology of the olfactory organ of The number of olfactory lamellae was counted
Katsuwonus pelamis and found olfactory cells on on both sides in Scomberomorus and a wide
the olfactory epithelium of the lamellae. Iwai and range of variation was observed, 24-76 (Table 2).
Nakamura (1964a) found that the number of In bonitos, the number oflamellae increases from
lamellae per rosette varies among specimens of small specimens to adults but does not appear to
species of Thunnus but that there were differ- change after a certain size is reached, as Collette
ences among species in the shape of the nasal and Chao (1975:532) showed for Gymnosarda

558
COLLETTE and RUSSO: SPANISH MACKERELS

vel n TABLE 2.-Number of lamella in nasal rosettes of species of

Scomberomorus.
Overall
Species Side Min. Max. x N x Rank
brasiliensis L 24 40 33.67 12 33.88 6
rig ht R 25 42 34.08 13
cavalla L 30 56 42.92 24 43.13 12
R 31 55 43.35 23
commerson L 42 58 48.92 12 49.32 13
vel n R 43 60 49.80 10
conc%r L 26 35 30.92 13 30.92
R 26 34 30.92 13
guttatus L 30 76 53.41 27 53.43 16
R 31 73 53.46 26
koreanus L 47 56 50.67 3 54.75 18
R 48 73 57.20 5
linea/stus L 30 35 32.50 4 32.18 3
R 30 34 32.00 7
maculatus L 25 38 33.43 14 33.44 5
R 30 37 33.45 11
mult/rad/atus L 32 40 36.75 4 36.00 10
R 25 44 34.50 2
munroi L 54 54 54.00 3 53.84 17
R 54 57 53.67 3
niphonius L 25 42 33.67 15 34.41 7
R 26 42 35.21 14
pluril/neatus L 45 53 49.50 4 50.50 14
R 44 56 51.50 4
queens/and/cus L 43 59 49.75 4 50.67 15
R 43 61 51.40 5
regalis L 28 41 34.00 9 35.11 9
R 30 43 36.22 9
semifasciatus L 31 37 34.00 3 34.78 8
R 31 38 35.17 6
sierra L 30 36 32.64 14 32.07 2
R 28 34 31.50 14
sinensis L 38 38 38.00 1 42.50 11
R 41 47 44.00 3
tritor L 27 48 33.40 10 32.57 4
R 24 37 31.83 11

unicolor and Orcynopsis unicolor. We have not

examined many small Scomberomorus nasal ro-
settes but did find 23 lamellae in an 80 mm FL S.
guttatus, a species for which the minimum count
of lamellae for specimens larger than 100 mm
was 30.
Three species of Scomberomorus (korean us ,
munroi, and guttatus) had high counts, overall
means 53.4-54.8. The highest counts per side
were for S. koreanus (73) and S. guttatus (76).
Ten species had low counts, overall means 31.0-
36.0. These 10 included all 6 species of the regalis
group as well as lineolatus, multiradiatus, ni-
phonius, and semifasciatus.

PHARYNGEAL MUSCLES
FIGURE B.-Kidney and postcardinal vein in ventral view
of Scomberomorus queenslandicus, Palm I., Queensland, 641 The paired pharyngeal (retractor dorsalis)
mmFL.
muscles originate on the ventral surface of one or
two vertebrae between the third and the sixth
abdominal vertebrae and insert on the upper
pharyngeal bones (Fig. 2). We did not find any
differences between species as Collette and Chao
(1975) did for the bonitos.

559
 FISHERY BULLETIN: VOL. 82, NO.4

~;.;.;.+-peritoneum

ti~f--urlnary bladder
ovary--~~
:I"~
,""'oi'r.';';';'''---- i n t est i n e

anal pore
FIGURE 9.- Urogenital system in ventral view of
Scomberomorus (body wall and viscera removed)..
Composite illustration.
-=-m--urogenital papilla
geAital pore

peritoneum ureter

'~!--_ g 0 n a d
urinary bladder
urinary bladder

large intestine

a nus

FIGURE to.-Urogenital system in ventral view of Scomberomorus queenslandicus, Palm I., Queensland, 641 mm
FL. a. With intestine opened. b. Urinary bladder and ureters.

Osteology and the two earlier papers of most importance to

the osteology of Scomberomorus: Mago Leccia
Osteological characters proved to be useful in (1958) on three western Atlantic species (cavalla,
determining relationships among the 18 species maculatus, and regalis) and Devaraj (1977) on
of Scomberomorus and between this genus and four Indian species (commerson, guttatus, ko-
its presumed closest relatives, Acanthocybium reanus, and lineolatus) and Acanthocybium.
and Grammatorcynus. The osteological portion of
the paper is divided into five sections: skull, axial
skeleton, dorsal and anal fins, pectoral girdle, SKULL
and pelvic girdle. Osteological terminology gen-
erally follows Gibbs and Collette (1967) and Col- Description of the skull is presented in two
lette and Chao (1975). Organization within sec- sections: neurocranium (Figs. 11-19) and bran-
tions is similar to that of Collette and Chao (1975) chiocranium.

560
COLLETTE and RUSSO: SPANISH MACKERELS

Neurocranium view, the neurocranium of Scomberomorus is

more or less trapezoidal in shape. It is elongate
Following a general description of the neuro- and flat, particularly at the anterior region and is
cranium, the four major regions are discussed: deepest at the hind end of the orbit. The dorsal
ethmoid, orbital, otic, and basicranial. surface is marked by a median ridge and three
grooves on each side: dilator, temporal, and su-
GENERAL CHARACTERISTICS.-In dorsal pratemporal (Allis 1903:49). These grooves are

SPHENOTIC

INTERCALAR

EXOCCIPITAL

VOMER FIRST VERTEBRA

LATERAL ETHMOID EPIOTIC

SUPRAOCCIPITAL

b
FIGURE n.-Skulls in dorsal view. a. Scomberomorus commerson, Coffs Harbour, New South Wales, 1,155 mm FL. b. Scom-
beromorus munroi, Cairns, Queensland, 800 mm FL, USNM 219372, paratype.

561
 FISHERY BULLETIN: VOL. 82. NO.4

separated from each other by ridges of bone. nasal, and it is not interrupted above the eyes by
Thus, there are six grooves and five ridges in all. any transverse ridge. Posteriorly, the ridge ends
The median ridge is carried forward on the fron- at the epiotic where the medial process of the
tals to the ethmoid and is prolonged posteriorly in posttemporal attaches.
a large supraoccipital crest. This crest extends The external or pterotic ridge extends forward
down over the exoccipital suture more broadly to the midlevel of the orbit and develops anterior-
than in any other genus of the Scombridae. lya small auxiliary ridge that extends laterally
The internal ridge or temporal ridge almost and posteriorly toward the temporal ridge.
reaches anteriorly to the posterior portion of the The dilator groove is shorter than the other two

b
FIGURE I2.-Skulls in dorsal view. a. Scomberomorus koreanus, Singapore, 480 nun FL. b. Scomberomorus concolor, Gulf of
California, 495 mm FL.

562
COLLE'ITE and RUSSO: SPANISH MACKERELS

and can be detected easily in lateral view. The The interorbital and otic regions are not as
temporal groove is the middle one and is deeper broad as in the more advanced genera of the
than either of the other two. The remaining Sardini (Collette and Chao 1975) and Thunnini
groove, the supratemporal, is the largest of the (Gibbs and Collette 1967). The median and tem-
three and opens posteriorly between the supra- poral crests are higher in Scomberomorus than in
occipital crest and the middle portion of the other scombrids. The bonitos, particularly Or-
epiotic. cynopsis unicolor (Collette and Chao 1975:fig. 21),

b
FIGURE I3.-Skulls in dorsal view. a. Acanthocybium solandri, Caribbean Sea, 1,240 mm FL. b. Grammatorcynus
bilineatus, Scott Reef, Timor Sea, 453 mm FL.

563
 FISHERY BULLETIN: VOL. 82, NO.4

EPIOTIC SUPRAOCCIPITAL CREST

ETHMOID

INTERCALAR

VOMER
FIRST VERTEBRA

PTEROSPHENOID PARASPHENOID

b
FIGURE 14.-Skulls in lateral view. a. Scomberomorus commerson, Coffs Harbour, New South Wales, 1,155 mm FL. b.
Scomberomorus munroi, Cairns, Queensland, 800 mm FL, USNM 219372, paratype.

have the next highest crests. premaxilla. At its anterolateral aspect, the eth-
moid bone supports the nasals.
ETHMOID REGION.-This region is composed In Scomberomorus, only the most anterior part
of the ethmoid, lateral ethmoid, and vomer. The of the ethmoid bone is exposed in dorsal view,
nasal bone lies lateral to the ethmoid and lateral while the rest of it is overlapped by the frontals.
ethmoid and, therefore, is included here. In Acanthocybium, only the lateral aspects of
the bone are overlapped by the frontals and a
Ethmoid.-The ethmoid (dermethmoid) is a V-shaped dorsal median portion is exposed. The
forked median bone overlapped by the frontals ethmoid bone is longer in A. solandri than in
above and bounded by the vomer and lateral Scomberomorus.
ethmoid ventrally. The concave anterior surface
articulates with the ascending process of the Lateral ethmoid. -The lateral ethmoids (par-

564
COLLETIE and RUSSO: SPANISH MACKERELS

ethmoids) are massive paired bones which form the skull and contact the anterior edge of the
the anterior margin of the orbit and the posterior parasphenoid. The median half of each lateral
and mesial walls of the nasal cavity. The lateral ethmoid extends downward about three-fourths
portion of each bone extends downward from the as far as the lateral portion and has a large round
middle region ofthe frontals. The ventral surface foramen for the olfactory nerve which is promi-
ofthis wall mesially bears an articulating surface nently seen on the anterior surface. On the dorsal
for the palatine and laterally another articulat- surface, they abut the nasals anteriorly, the fron-
ing surface for the first infraorbital (lachrymal). tals posteriorly, and articulate with the ethmoid
The inner walls of the lateral ethmoids come mesially. On the anterior surface, ventral to the
closest to each other at the ventral median line of foramen, each lateral ethmoid bears a process

b
FIGURE 15.-Skulls in lateral view. a. Scomberomorus koreanus, Singapore, 480 mm FL. b. Scomberomorus concolor, Gulf of
California, 495 mm FL.

565
 FISHERY BULLETIN: VOL. 82, NO.4

that extends anteriorly and mesially to contact ethmoid dorsally and lateral ethmoid dorsolater-
the dorsolateral surface of the spear-shaped pos- ally. The pointed posterior process is firmly anky-
terior portion of the vomer. No appreciable differ- losed dorsally with the parasphenoid. On each
ence was noted in the lateral ethmoids of the side of the vomer, dorsolaterally and behind the
different species. spatulate anterior process, is a prominent articu-
lar surface for a loose articulation with the head
Vomer.-The vomer is the most anteroventral- of the maxilla. Posterior to this articular surface,
ly located bone of the cranium. The spatula- facing ventrolaterally, is a prominent sulcus for a
shaped anterior process bears a large oval patch similar movable articulation with the ventral
of fine teeth on its ventral surface. The vomerine branch of the anterolateral fork of the palatine.
tooth patch extends posteriorly as a narrow ridge The spatulate anterior process of the vomer is
in some specimens of some species, e.g., S. con- very long and extends beyond the anterior mar-
color (Fig. I5b). The vomer articulates with the gins of the nasal and ethmoid bone in Scomber-

b
FIGURE 16.-Skulls in lateral view. a. Acanthocybium solandri, Caribbean Sea, 1,240 mm FL. b. Grammatorcynus
bilineatus, Scott Reef, Timor Sea, 453 mm FL.

566
COLLEITE and RUSSO: SPANISH MACKERELS

omorus. No other scombrid has such a spatulate Nasal.-The nasal bones (Fig. 20) are flat,
anterior extension of the vomer. In fact, the roughly triangular bones with thickened lateral
vomer is either not visible in dorsal view or edges. The mesial edges are irregular and almost
protrudes anteriorly slightly beyond the ethmoid serrate in some species to form a firm immovable
in other scombrid genera. articulation with the lateral edge of the frontals.

PTEROSPHENOID
PTEROTIC
FRONTAL

LATERAL ETHMOID INTERCALAR

ETHMOID --\~~" BASIOCC IPIT A L

FIRST VERTEBRA

PARASPHENOID
SPHENOTIC EXOCCIPITAL

b
FIGURE 17.-Skulls in ventral view.a. Scomberomorus commerson, Coffs Harbour, New South Wales, 1,155 mm FL. b. Scom-
beromorus munroi, Cairns, Queensland, 800 mm FL, USNM 219372, paratype.

567
 FISHERY BULLETIN: VOL. 82, NO.4

FIGURE 18. -Skulls in ventral view. a. Scomberomorus koreanus, Singapore, 480 mm FL. b. Scomberomorus concolor, Gulf of Cali-
fornia, 495 mID FL.

The anterior margins fit neatly beside the ante- cynus (2.8-3.4, Fig. 20d), S. cavalla (2.8-3.1, Fig.
rior branches of the forked ethmoid bone as can 20a), and S. regalis (2.8-3.0). The other 14 species
be seen in the dorsal views of the skulls (Figs. 11, of Scomberomorus are intermediate (2.0-2.9). The
12). They are nonprojecting in that their anterior anterior end of the nasal bone is rounded and
margin is at the level of the ethmoid bone except heavy in Scomberomorus and Acanthocybium
in Grammatorcynus where they project well be- (Fig. 20a-c). The anterior end has a short, slightly
yond the anterior end of the neurocranium (Fig. angled arm in Grammatorcynus (Fig. 20d).
13b). Length divided by width ranges from 2.0 to
4.2 in the three genera. The widest nasal bones ORBITAL REGION.-The orbit is surrounded
are in S. koreanus (2.0-2.1) and S. sinensis (2.0- by the posterior wall of the lateral ethmoid, the
2.3, Fig. 20b). The most elongate nasals are in ventral side of the frontal, the pterosphenoid,
Acanthocybium (3.1-4.2, Fig. 20c), Grammator- sphenotic, prootic, suborbital, and lachrymal

568
COILETI'E and RUSSO: SPANISH MACKEREUl

FIGURE 19.-Skulls in ventral view. a. Acanthocybium solandri, Caribbean Sea, 1,240 mm FL. b. Grammatorcynus
bilineatus, Scott Reef, Timor Sea, 453 mm FL.

bones. The left and right orbits are partially the dorsal surface of the lateral ethmoid. The
separated by the basisphenoid. The sclerotic midlateral aspect is thickened to form the orbital
bones enclose the eyeballs. roof. Posteriorly, they are bounded by the supra-
occipital and parietals. Posterolaterally, they
Frontal. -The frontals are paired bones that overlap the pterotics and just anterior to the
form the largest portion of the dorsal surface of pterotics, cover the sphenotics. Ventrally, each
the neurocranium. Anteriorly they are pointed, frontal bears a sheet of bone, the orbital lamella,
and posteriorly they become expanded. Anterior- which is bounded by the sphenotic posteriorly,
ly, the frontals overlap the dorsal surface of the lateral ethmoid anteriorly, and pterosphenoid
ethmoid bone, the inner edge of the nasals, and mesially. On the base of the orbital lamella may

569
 FISHERY BUlLETIN: VOL. 82, NO.4

be seen a number of small foramina for the Scomberomorus, the left and right frontals are
branches of the supraorbital nerve trunk.. The attached very closely to each other such that
laterosensory canals of the frontals are evident there is no gap between them. However, in S.
on the pterotic crests as a series of pores. commerson and S. cavalla, there is a long narrow
In Acanthocybium, the frontals are separated slit between the left and right frontals, but it is
from each other by the dorsomedian pineal fenes- not a fenestra in the true sense, as the lower parts
tra lying just in front of the supraoccipital at the of the bones are very closely approximated. A
level of the pterosphenoids and another anterior third condition is found in S. sinensis. Here the
fontanel just posterior to the ethmoid bone (Fig. anterior part ofthe median ridge is almost absent
I3a). A smaller, more oval pineal opening is and there is a wide gap between the left and right
present between the posterior ends of the frontals frontals. The interorbital commissures of the lat-
in Grammatorcynus (Fig. I3b). When viewed eralis system are developed a little anterior to the
through the pineal fenestra, a part of the dorsal middle of each frontal in the form of two pores at
surface of the parasphenoid is visible through the the margin of the median ridge which lead into
opening of the brain chamber between the ptero- oblique tubes downwards and posteriorly. An-
sphenoids. There is a deep depression on the other pair of commissures of the lateralis sys-
frontals mesially, just anterior to the pineal fe- tem is developed along the anterolateral margin
nestra. This depression becomes shallower ante- of the frontals. These sensory canals are not
riorly, becoming confluent with the dorsal surface developed in Acanthocybium and Grammatorcy-
of the frontals. In Scomberomorus, the frontals nus.
join mesially along the median line on the neuro-
cranium where they form the anterior half of the Pterosphenoid. -The pterosphenoids (alisphe-
median ridge whose posterior half is composed of noids) form the posterodorsal region of the orbit.
the supraoccipital crest. In all but three species of They abut the basisphenoid and prootics posteri-

a b

c d
FIGURE 20.-Left nasal bones in lateral view. a. &omberomorus cavalla, Miami, 797 mm FL, 2x. b. Scomberomorus sinensis,
Tokyo, 1,850 mm FL, 1x. c. Acanthocybium solandri, Revillagigedos Is., 1,068 mm FL, 1.5x. d. Grammatorcynus bilineatus,
Queensland, 521 mm FL, 3 x .

570
COLLE'ITE and RUSSO: SPANISH MACKERELS

orly and the frontals and sphenotics laterally. the relative degree of development of the lateral
There is a space between the left and right ridges. Both features are best developed in S.
pterosphenoids opening into the brain chamber commerson where the length of the anterior pro-
just anterior to the basisphenoid. In most species cess is greater than the height of the vertical axis
of Scomberomorus, there is an anterior medially ofthe bone.
directed lobe on each ptero!?phenoid. These lobes
meet along the median line or at least come very Infraorbitals. - The infraorbital (suborbital)
close to each other in adults of three species: series of Scomberomorus consists of from 9 to 13
commerson (over 1,000 mm FL), cavalla (over 550 elements which enclose the infraorbital branch of
mm FL), and lineolatus (over 750 mm FL). Small- the lateral sensory canal system (Fig. 21a). Only
er specimens of these 3 species and all sizes of the 9 elements were observed in S. munroi, S. sierra,
other 15 species have a wide gap or fenestra and S. sinensis, but 13 elements were observed in
between the left and right lobes. The gap is about S. brasiliensis. The canal enters the infraorbital
equal to the width of the parasphenoid or slightly series at what is usually considered the last
larger in three species: brasiliensis, koreanus element (dermosphenotic) and continues around
(Fig. 12a), and concolor (Fig. 12b). The gap is the orbit to terminate on the first infraorbital
largest in S. multiradiatus, so large that there is (lachrymal).
virtually no medially directed lobe. This causes The first infraorbital (lachrymal or 101) is the
the window into the brain chamber to be almost first and largest element in the infraorbital se-
rectangular in this species. ries. Anteriorly, several canal tubes open on the
laminar, platelike surface of the bone. Posterior-
Sclerotic. -The sclerotic bones consist of two ly, the canal tube continues directly to the second
thickened semicircular segments connected by infraorbital. The first infraorbital is an elongate
cartilage on the inner lateral surface and by bone (length/height = 2.8-3.5) that covers part of
corneal membranes on the outside. The inner rim the maxilla and is attached to the lateral ethmoid
of the sclerotic bones appears elliptical externally dorsally by a mesially directed articular process.
as in the bonitos (Collette and Chao 1975) and The anterior portion is forked with a thin ante-
Thunnus (e.g., T. atlanticus , de Sylva 1955:fig. rior process. This process is a point of attachment
7). The sclerotic bones of Grammatorcynus are for a ligament connected to the nasal. The projec-
relatively larger, thinner, and close to circular. In tion is present in all species of Scomberomorus
Acanthocybium, the sclerotic bones are ellipti- except S. lineolatus and S. tritor. The portion
cal as in Scomberomorus, but they are heavier posterior to the articular process is elongate,
and extend further medially. The only species of pointed, and longer than the anterior portion.
Scomberomorus that appeared to differ from the The general shape of the first infraorbital in
other species is S. sinensis. The sclerotics are Scomberomorus is similar to that in the bonitos
especially thick in this species and there is a (Collette and Chao 1975:fig. 28), particularly
thick bony lump in the middle of the posterior Cybiosarda elegans, except that the anterior pro-
surface of one of the two sclerotics. Other species cess is smaller and more dorsally directed than in
of Scombero""orus have a thickening of the bone Cybiosarda. Acanthocybium differs from Scom-
a
in the same region but it does not form distinct beromorus in having the posterior portion of the
protrusion as it does in S. sinensis. first infraorbital short and broad, shorter than
the anterior portion (Fig. 21b). Grammatorcynus
Basisphenoid.-The basisphenoid is a small, has a feebly forked anterior end (Fig. 21c), lack-
median, V-shaped bone that connects the para- ing a distinct anterior process such as is present
sphenoid, prootics, and pterosphenoids. The com- in Scomberomorus and Acanthocybium.
pressed median vertical base bears an anterior As Devaraj (1977) noted, the dorsal margin
median process but lacks a posterior process as is of the anterior part of the first infraorbital is
present in other scombrids such as Thunnus straight, or nearly so, in S. cavalla and S. com-
(Gibbs and Collette 1967) and most bonitos (Col- merson but clearly concave in the other species.
lette and Chao 1975). In most species of Scomber- Mago Leccia (1958:pl. 4, fig. 7) indicated that S.
omorus there is at least a trace of a lateral ridge cavalla lacked the characteristic anterior projec-
that extends laterally and posteriorly on each tion, but we have found it to be present in our
side of the anterior process. There is great varia- material. In other respects, there seems to be as
tion in the length of the anterior process and in much variation between individuals of a species
571
 FISHERY BUlLETIN: VOL. 82, NO.4

dermosphenotic--~

IOl(lachrymal)

FIGURE 21.-Left infraorbital bones in lateral view. a. Scomberomorus maculatus, Cape Hatteras, N.C., 534 nun FL. b. Acantha-
cybium solandri, Revillagigedos Is., 1,068 mm FL. c. Grammatorcynus bilineatus, Timor Sea, 453 nun FL.

as between species in the shape of the first reported by those authors, we have found this
infraorbital. shelf to occur in all species of Scomberomorus as
The second infraorbital (102) sits firmly on the well as in all other genera of scombrids. The
dorsal edge of the anterior portion of the first shape of this shelf varies among specimens of the
infraorbital. It is a flat, somewhat compressed same species as well as between right and left
bone. sides of a single specimen.
The third infraorbital (103) is an elongate, The fourth through the penultimate elements
tubular bone. It has no platelike extensions, but (postorbitals) usually are simple tubelike bones
has a large mesial shelflike extension (subocular which may have pores accommodating canal
shelf of Smith and Bailey 1962). Although not tubes to the skin and cheek scales. The fourth
572
COLLE'ITE and RUSSO: SPANISH MACKERELS

through about the seventh elements may be ex- beromorus in the attitude of the attachment area
panded laterally as laminar plates which cover and its roughness. In many species, the lateral
the anterior end of the cheek scales. There may crest continues posteriorly almost perpendicular
be 10-16 rows of specialized cheek scales posterior to the skull. In some species such as S. commer-
to the infraorbitals. These scales originate mesial son (Fig. lla) and S. queenslandicus, the area of
to the infraorbital canal tubes and extend poste- attachment is flatter. This area is flat and rough
riorly as flat, sometimes pointed, platelike ele- in Acanthocybium (Fig. 13a) and forms a separate
ments. These platelike scales may themselves be process in Grammatorcynus (Fig. 13b).
covered with more typical cycloid scales and
exhibit the same morphology as the corselet Supraoccipital. -The supraoccipital forms the
scales of higher scombrids. The cheek scales of dorsomedial portion of the posterior end of the
Scomberomorus may represent the primitive neurocranium and bears a well-developed crest
condition of the corselet. which continues anteriorly on the frontals and is
pronounced posteriorly as a strong supraoccipital
OTIC REGION.-This region encloses the otic crest. The supraoccipital can be divided into two
chamber inside the skull, and is formed by the parts: a thin, elongate triangular crest and a
parietal, epiotic, supraoccipital, prootic, pterotic, roughly hexagonal base. The crest extends down
sphenotic, and intercalar (opisthotic) bones. over the exoccipitals along the median line where
the dorsal walls of the exoccipitals suture with
Parietals. - The parietals articulate with the each other, but it is not interposed between the
frontals anteriorly, the supraoccipital mesially exoccipitals. The hexagonal base is bounded an-
and the pterotics laterally, sphenotics ventrally, teriorly by the frontals and laterally by the
and epiotics posteriorly. The inner lateral crest parietals and epiotics. The crest extends posteri-
that originates at the middle of the frontal bones orly over the first vertebral centrum usually to a
continues through the parietals to terminate at level past the posterior margin of the centrum
the epiotics. This crest is typical of scombrids and (Figs. 14-16). The height ofthe crest varies among
is particularly well developed in Scomberomorus. species of Scomberomorus and is highest in three
These crests originate on the parietals, instead of species, S. guttatus, S. korean us (Fig. 15a), and
the frontals, in Acanthocybium and Grammator- S. multiradiatus. Dividing the height of the su-
cynus and are not as high as in Scomberomorus. praoccipital crest (ventral margin ofsupraoccipital
The parietals of all the species of Scomberomorus to edge of crest) by skull length (tip of vomer to·
are similar. posteroventral margin of basioccipital) gives a
There is a gap or fenestra on the dorsal surface ratio of 0.46-0.57 for these three species, com-
of the skull where the parietal, epiotic, and pter- pared with 0.34-0.45 in the other 15 species. Low
otic bones come together. It varies in shape from ratios are found in S. cavalla and S. commerson
roughly triangular to rectangular in most spe- (0.35-0.40) and in all six species of the regalis
cies. There is wide variation from specimen to group (0.34-0.42).
specimen that tends to obscure potential inter-
specific differences. The gap is very small in some Prootics.-In ventral view, the prootics con-
specimens of eight species: commerson, concolor nect with all bones on the ventral side ofthe skull
(Fig. 12b), koreanus (Fig. 12a), maculatus, mun- which compose the posterior part of the neuro-
roi (Fig. lIb), plurilineatus, queenslandicus, and cranium (Figs. 17-19). Each prootic is bordered
sierra. It is usually larger in the other species ventrally by the parasphenoid; posteriorly by the
and in most specimens of S. commerson (Fig. basioccipital, exoccipital, and intercalar; lateral-
lla). ly by the pterotic and sphenotic; and anteriorly
by the pterosphenoid and basisphenoid. The pro-
Epiotics. -The epiotics are massive, irregular, otic bones are irregular in shape and meet each
and bounded by the parietals anteriorly, the other along the ventromedian line of the brain
supraoccipital mesially, the exoccipitals posteri- case to form the anterior portion of the poste-
orly, and the pterotics laterally. The inner lateral rior myodome. On the ventral surface, extending
crests terminate at the posterior end of the epi- from the lateral wing of the parasphenoid to the
otics. The medial process of the posttemporal sphenotic, the prootic forms a thick bridge which
bone attaches here on a rough process. There are strengthens the trigemino-facialis chamber (Allis
slight differences between the species of Scom- 1903). A prootic foramen is present anterolateral-

573
 FISHERY BULLETIN: VOL. 82, NO.4

ly between the tip of the parasphenoid wing and Sphenotics. -The sphenotics form the most
the sphenotic. There is no trace of the prootic pit posterior dorsolateral part of the roof of the orbit.
characteristic of the Thunnini and Allothunnus They continue the outer lateral shelf from the
(Gibbs and Collette 1967; Collette and Chao 1975). frontals and articulate with the pterosphenoid
Specimens differ in the number and arrangement medially and the prootic and pterotic posteriorly.
of foramina leading into the brain cavity from A segment of the articular fossa for the head of
inside the anterior opening of the trigemino- the hyomandibula is afforded by the lateral wall
facialis chamber, but these do not seem to be of the sphenotic on the ventral surface. The
useful interspecific differences. sphenotic is pierced by a foramen for the ramus
oticus nerve (Allis 1903). When viewed dorsally,
Pterotics. -The pterotics form the lateral pos- the sphenotics spread out on both sides more
terior corners of the neurocranium. Posteriorly, prominently in Scomberomorus than in Acan-
each pterotic is produced into a truncate process thocybium, as noted by Devaraj (1977). Devaraj
or pointed spine. The pterotics articulate with the stated that the «midlateral projection" was large
epiotics and parietals medially and with the in koreanus, guttatus, maculatus, and regalis;
exoccipitals and intercalars posteriorly. A ridge, small in lineolatus, cavalla, and commerson; and
the pterotic ridge, originates on the dorsal sur- absent in Acanthocybium, but we are not clear as
face of the posterior third of the frontal and to what he was referring.
continues posteriorly, diverging to the posterior
corner of the pterotic, just anterior to the pterotic Intercalars.-The intercalars (opisthotics) are
spine. In ventral view, the pterotics articulate flat bones that form part ofthe posterior border of
with the sphenotics anteriorly and the prootics the neurocranium interposed between the pter-
and intercalars medially. Two contiguous fossae, otics and exoccipitals. The anterior portion on the
one at the posterior half of the pterotic bone and dorsal surface is concealed by the overlapping
one at its joint with the sphenotic, seat the dor- pterotic, thus exposing the bone on the dorsal
sal and anterior condyles of the hyomandibula. surface less than on the ventral side. Each inter-
Three closely situated lateral sensory canal pores calar bears a protuberance on the dorsal surface
open on each pterotic at the posteriormost region to receive the lateral arm of the posttemporal.
of the pterotic crest. The largest pore is the most This protuberance is followed by a posterior
posterior and opens dorsally; lateral to this is the projection of the intercalars in some species of
next largest opening laterally on the outside of Scomberomorus but not in Acanthocybium or
the pterotic crest; the smallest is the most ante- Grammatorcynus.
rior ofthe three, lying along the crest and usually Species of Scomberomorus may be roughly di-
more elongate in shape. vided into three groups based on the size of
The lengths and widths of the pterotic spines the posterior projection from the intercalar as
vary among the species. In eight species (brasili- Devaraj (1977) noted for Indian species. Eight
ensis, guttatus, koreanus (Fig. 18a), multiradi- species lack any posterior projection or have only
atus, plurilineatus, regalis, semifasciatus, and an insignificant projection: guttatus, koreanus
tritor), there is essentially no pterotic spine, (Fig. 18a), lineolatus, multiradiatus, munroi
merely a rounded posterior area of the skull. In (Fig. 17b), plurilineatus, semifasciatus, and si-
six species (concolor (Fig. 18b), lineolatus, macu- nensis. In each of these species, except S. multi-
latus, munroi (Fig. 17b), niphonius, and sierra), radiatus, the pterotic spine protrudes further
there is a blunt posteriorly projecting spine. posteriorly than does the intercalar region. In S.
Scomberomorus sinensis is similar to this group, multiradiatus, the posterior corners of the skull
but the posterior projection is broader and less are rounded and there is no pterotic spine so the
like a spine. The pterotic spines are longest in intercalars project further posteriorly. Eight spe-
three species (cavalla, commerson (Fig. 17a), and cies have a distinct posterior projection from the
queenslandicus), all of which also have promi- intercalar: brasiliensis, cavalla, concolor (Fig.
nent posterior projections of the intercalars. 18b), maculatus, niphonius, regalis, sierra, and
Grammatoreynus (Fig. 19b) is similar to the lat- tritor. The posterior projection is smaller in some
ter group, but the spine is thinner and sharper. specimens of S. niphonius, placing it somewhat
Acanthocybium (Fig. 19a) has a longer and thin- between groups 1 and 2. The posterior projection
ner pterotic spine than do Grammatorcynus and is a little longer in S. cavalla, between groups
the species of Scomberomorus. 2 and 3. Two species, commerson (Fig. 17a)

574
COLLETTE and RUSSO: SPANISH MACKERELS

and queenslandicus, have a prominent truncate ly, the lateral flanges fuse to form a circular
process. margin in a slightly backward oblique position
and attach to the margin of the first vertebral
BASICRANIAL REGION.-This region con- centrum. There are a variable number of small
sists of the parasphenoid, basioccipital, and exoc- pores in a shallow depression on the lateral
cipital bones, and forms the posteroventral base surfaces of the basioccipital. This depression is
of the skull. deepest in S. sinensis but does not approach
the basioccipital depression characteristic of the
Parasphenoid. -The parasphenoid is a long bonitos, Sardini (Collette and Chao 1975).
cross-shaped bone (Figs. 17-19) which articulates
with the vomer anteriorly and forms the ventral Exoccipital.-The exoccipitals connect the
axis of the skull. The lateral wing of the para- skull to the first vertebra dorsally. The exoccipi-
sphenoid extends dorsolaterally along the ventral tal articulates with the epiotic and supraoccipital
ridge of the prootic bones on either side, and has a bones anterodorsally, the intercalar laterally,
pointed end which forms part ofthe anteroventral and with the other exoccipital posterodorsally. In
wall of the posterior myodome. Posteriorly, the ventral view, the exoccipital articulates with the
parasphenoid bifurcates into two lateral flanges prootic anteriorly, basioccipital medioventrally,
which attach dorsally to the corresponding pos- and intercalar laterally. In posterior view, the
teroventral flanges of the basioccipital bone and foramen magnum is framed by the exoccipitals.
surround the posterior opening of the posterior Laterally, there are two foramina. The small
myodome. A ventrally projecting median keel is anterior glossopharyngeal foramen (Allis 1903)
present in the area anterior to the origin of the lies close to the posterior border of the prootic.
lateral flanges. In ventral view, the general The large posterior vagal foramen lies just under
characteristic of the parasphenoid is a gradual the overhanging shelf formed by the posterior
narrowing of the bone from anterior to posterior. margin of the exoccipital. Dorsally, a small fora-
The broadest portion of the parasphenoid is lo- men which opens into the brain cavity is present
cated usually at or before the tip of the V-shaped at the medioposterior corner of the exoccipital.
joint with the vomer. Broad parasphenoids are
also present in Acanthocybium and the bonitos, Branchiocranium
Sardini (Collette and Chao 1975). In lateral view
(Figs. 14-16), the parasphenoid forms the ventral The branchiocranium is divided into five sec-
border to the orbits and connects with the lateral tions: mandibular arch, palatine arch, hyoid arch,
ethmoids, basisphenoid, prootics, and basioccipi- opercular apparatus, and branchial apparatus.
tal bones dorsally.
The shaft of the parasphenoid is distinctly MANDIBULAR ARCH.-The mandibular arch
wider in seven species: S. commerson (Fig. 17a), is composed of the upper jaw (premaxilla, maxil-
lineolatus, munroi (Fig. 17b), niphonius, queens- la, and supramaxilla) and the lower jaw (dentary,
landicus, semifasciatus, and sinensis. Devaraj angular, and retroarticular). Teeth are borne on
(1977) included S. caualla along with S. lineola- the premaxilla and dentary, and the number of
tus and S. commerson as having a broad para- teeth on these bones is a useful taxonomic char-
sphenoid, based on Mago Leccia (1958). We find S. acter (see Dentition section).
caualla to have a broader parasphenoid than some
members of the regalis species group but not as Dentition.-Large, triangular, laterally com-
broad as in the group ofseven species listed above. pressed teeth are present in the upper and lower
jaws of Scomberomorus. Acanthocybium has sim-
Basioccipital.-The basioccipital is the most ilar teeth that are a little blunter and more
posteroventrally located bone of the skull. It is tightly packed. Grammatorcynus has long thin
shaped like an inverted U with lateral flanges on teeth that are slightly compressed laterally. Bo-
either side of the skull and forms the roof and nitos have conical teeth that are larger than the
lateral walls of the posterior myodome. Anterior- conical teeth of the higher tunas (Thunnini).
ly, the basioccipital is attached to the prootic Tooth replacement in Scomberomorus caualla
bones and dorsally with the exoccipital bones. Its was studied by Morgan and King (1983). The
lateral flanges expand ventrally to meet the flat number of jaw teeth in Scomberomorus varies
posterior flanges of the parasphenoid. Posterior- widely with a range of 5-39 in the upper jaw, 4-37
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 FISHERY BULLETIN: VOL. 82, NO.4

in the lower jaw (Tables 3, 4). Two species of Premaxilla. -The premaxilla (Fig. 22) is a
Scomberomorus stand out from the rest, S. multi- long, curved bone with a stout, arrowhead-shaped,
radiatus with the fewest teeth (5-10, x 8.0 on the anterior end that extends dorsally and posteri-
upper jaw; 5-11, x 7.8 on the lower jaw) and S. orly as an ascending process. The posterior shank
concolor with the most teeth (13-37, x 22.2 on the of the premaxilla is elongate and bears a row of 5-
upper jaw; 12-34, x 19.7 on the lower jaw). The 18 39 compressed triangular teeth on its ventral
species can be ranked from lowest to highest margin. There are two articular facets for the
as follows (mean for upper jaw followed by mean overlying maxilla at the junction of the posterior
for lower jaw): 1) multiradiatus (8.0, 7.8); 2) margin of the ascending process with the shank.
queenslandicus (13.3, 10.6); 3) semifasciatus The ascending processes of both premaxillae are
(12.8, 11.2); 4) cavalla (14.0, 10.9); 5) koreanus closely approximated to each other mesially and
(13.7,11.2); 6) commerson (14.1,11.3); 7) sinensis fit into the median groove of the ethmoid bone.
(13.4,12.2); 8) brasiliensis (14.0,11.9); 9) lineo- The ascending process forms an angle of 32°_61°
latus (15.1, 12.9); 10) guttatus (16.9, 14.4); 11) with the shank, and this process is 31-48% of the
sierra (17.3, 14.1); 12) maculatus (16.8, 14.6); total length of the premaxilla. Devaraj (1977:22)
13) munroi (17.5, 15.0); 14) plurilineatus (17.9, noted that S. lineolatus had the sharpest angle
15.4); 15) tritor (18.6, 15.4); 16) regalis (19.3, among the Indian species that he studied (23 0
15.8); 17) niphonius (19.6, 15.9); and 18) con- as he measured it), and we find that it has the
color (22.2, 19.7). The species with the fewest sharpest angle (Fig. 22b) of any of the species
teeth, S. multiradiatus, also has the fewest gill in the genus, 32°-36° according to our measure-
rakers (usually 2 or 3, see Table 5), and the spe- ments. The species with the largest angle is S.
cies with the most teeth, S. concolor, has the most guttatus, 60°_61°. Devaraj included guttatus along
gill rakers (usually 23-25, see Table 5) but the with koreanus, regalis, and maculatus as species
correlation is not so good in the other 16 species with angles of 40°_43°. Our data for these other
(compare Tables 3 and 4 with Table 5). three species are 40°_54°. Scomberomorus com-

TABLE 3.-Number of teeth in upper jaw in species of TABLE 4.-Number of teeth in lower jaw in species of
Scomberomorus. Scomberomorus.

Overall Overall
Species Side Min. Max. i SD N x Rank Species Side Min. Max. x SD N x Rank
brasiliensis L 6 25 14.07 3.62 68 brasiliensis L 7 19 11.96 2.85 70
14.00 6 11.88 7
R 8 27 13.93 3.53 69 R 7 20 11.79 3.04 67
cavaJla L 8 29 14.24 5.82 50 cavalla L 6 24 10.94 3.64 50
14.00 7 10.92 3
R 6 28 13.74 5.23 46 R 7 22 10.90 3.78 48
commerson L 5 35 14.15 5.68 110 commerson L 5 29 11.37 4.40 108
11.27
14.06 8 6
R 7 38 13.96 5.28 109 R 4 27 11.17 3.84 106
conc%r L 15 35 22.15 4.94 26 conco/or L 13 30 19.46 3.96 26
22.20 18 19.71 18
R 13 37 22.26 5.77 23 R 12 34 19.96 4.96 25
guttatus L 12 36 16.78 4.13 89 guttatus L 10 25 14.49 3.06 98
16.68 11 14.42 11
R 11 35 16.97 4.25 93 R 9 23 14.34 2.70 97
koreanus L 9 19 14.17 2.76 24 koreanus L 8 17 11.25 2.19 24
13.71 5 11.21 4
R 10 16 13.25 2.11 24 R 9 15 11.17 1.31 24
lineo/atus L 10 27 15.28 3.94 29 linealatus L 7 28 12.72 3.69 29
15.07 9 12.93 9
R 9 28 14.86 4.19 29 R 9 26 13.14 3.20 29
maculatus L 10 32 17.04 4.06 55 maculatus L 10 30 14.89 3.70 55
16.82 10 14.64 12
R 7 30 16.57 3.80 49 R 8 26 14.37 3.02 52
multiradiatus L 5 10 7.88 1.24 26 multiradiatus L 6 11 8.00 1.20 26
8.04 7.75
R 6 10 8.19 1.20 26 R 5 9 7.50 0.95 26
munroi L 12 20 16.57 2.64 7 munroi L 11 29 15.88 5.62 8
17.50 13 15.01 13
R 12 23 18.22 3.90 9 R 11 19 14.13 2.75 8
niphonius L 12 26 19.53 2.71 32 niphonius L 12 20 15.55 2.05 33
19.56 17 15.93 17
R 14 26 19.58 2.75 33 R 12 20 16.30 2.05 33
plurilineatus L 16 22 18.25 1.70 24 piurilineatus L 12 22 15.83 2.08 23
17.92 14 15.37 14
R 12 23 17.58 2.52 24 R 12 20 14.92 1.89 24
queensla.ndicus L 8 17 13.33 2.43 30 queenslandicus L 6 14 10.59 1.79 32
13.29 3 10.61 2
R 10 18 13.24 2.31 29 R 7 14 10.64 1.99 28
regalis L 9 31 19.34 5.10 47 regalis L 10 24 15.72 4.17 46
19.29 16 15.80 16
R 10 30 19.25 4.74 48 R 8 23 15.87 4.10 47
semifasciatus L 10 23 13.03 3.07 33 semifasciatus L 7 18 11.24 2.98 33
12.76 2 11.23 5
R 8 21 12.48 2.92 33 R 7 18 11.21 2.93 33
sierra L 10 37 17.15 5.79 60 sierra L 7 37 13.90 5.29 61
17.32 12 14.05 10
R 7 39 17.48 7.34 62 R 7 32 14.19 5.30 62
sinensis L 10 16 13.64 1.69 14 sinensis L 10 15 12.43 1.87 14
13.43 4 12.22 8
R 10 17 13.21 1.67 14 R 10 15 12.00 1.41 14
tritor L 11 30 18.56 3.98 32 tritor L 10 21 15.24 2.95 33
18.58 15 15.40 15
R 11 28 18.59 4.38 32 R 10 23 15.56 3.19 33

576
COLLE'ITE and RUSSO: SPANISH MACKERELS

a
b

d
c
FIGURE 22.-Left premaxillae in lateral view. a. Scomberomorus semifasciatus, Port Moresby, New Guinea, 510 mm FL,
2 x. b. Scomberomorus lineolatus, Cochin, India, 786 mm FL, 2 x. c. Acanthocybium solandri, Miami, Fla., 1,403 mm FL,
1 x. d. Grammatorcynus bilineatus, Marshall Is., 424 mm FL, 2 x .

merson and S. cavalla also fall into this inter- S. cavalla, 31-32%. Acanthocybium has a longer
mediate group with angles of 41°_54°. Acantho- process than any of the species of Scomberomo-
cybium (Fig. 22c) has a sharp angle (34°-37°), like rus, 50% (according to our data and Devaraj 1977).
S. lineolatus. Grammatorcynus (Fig. 22d) has a
very large angle, 64°-67°, even greater than S. Maxilla.-The maxilla (Fig. 23) is a long,
guttatus. The ascending process is longest in S. curved bone surmounting the premaxilla dorso-
lineolatus, 46-48% ofthe length ofthepremaxilla laterally by means of an anterior head and ven-
(41-45% according to Devaraj) and S. sinensis, tral sulcus. The head consists of a thick massive
43-46%. The process is shortest in S. guttatus and inner condyle and a small lateral process (see S.

a b

c d
FIGURE 23. - Left maxillae in lateral view. a. Scomberomorus semifasciatus, Port Moresby, New Guinea, 510 mm FL, 2 x. b. Scom-
beromorus munroi, New Guinea, 512 mm FL, 2 x. c. Acanthocybium solandri, Miami, Fla., 1,403 mm FL, 1 x. d. Grammatorcynus
bilineatus, Timor Sea, 453 mm FL, 2 x .

577
 FISHERY BULLETIN: VOL. 82, NO.4

semifasciatus, Fig. 23a). The former possesses a Dentary.-The dentary (Fig. 24) is a large
prominent knob at its dorsolateral aspect that fits forked bone which forms the major part of the
into the articular surface of the vomer and an lower jaw. It is laterally flattened and bears a
anterior, deep concavity facing the inner wall of single row of 4-37 compressed triangular teeth on
the premaxilla. The head is 18-25% of the total the dorsal margin. Posteriorly, the dentary forms
length of the maxilla. Immediately posterior to two arms. The ventral arm is relatively narrow
the head is a shallow depression which receives and shorter than the dorsal arm, and its inferior
the anterior articulating process of the palatine. margin has a groove which accepts the angular
The shank of the maxilla is narrow and some- and the anterior end of Meckel's cartilage. The
what flattened. The posterior end expands into a base of the ventral arm has an external series of
thin, flat plate which is partially covered dorsally pores, which seem to be the preoperculomandibu-
by the supramaxilla. The height of the plate is 8- lar pores (Allis 1903; Mago Leccia 1958) of the
15% of the total length of the maxilla. Acantho- lateral line system. The length of the dentary
cybium (Fig. 23c) and Grammatorcynus (Fig. from its anterior margin to the tip of the lower
23d) lack any posterior expansion of the maxilla. arm is 86-97% of the length to the tip ofthe upper
In fact, in Acanthocybium there is a notch in the arm. The figures are similar for Acanthocybium
dorsal margin of the maxilla, and the posterior (91-96%). However, the lower margin is longer in
end is distinctly lower than the middle of the Grammatorcynus, 105-109% of the length of the
shaft of the bone. upper margin (Fig. 24c). The proportions are
Scomberomorus munroi is the only species in similar in all 18 species of Scomberomorus, with
the genus that is distinguishable from the others S. maculatus having the shortest lower margin
in characters of the maxilla; it totally lacks the (87-89%) and S. concolor the longest (92-97%).
anterior process on the outer surface of the head
of the maxilla (Fig. 23b). Devaraj (1977:23) stated
that the outer process was "flimsier" in S. com-
merson, but we find that the process varies from
small to moderate in our material of the species
and that S. commerson is not distinct in this
aspect.
The head ofthe maxilla is shorter than in most a
other species, relative to total length of the max-
illa, in the six species of the S. regalis group.
Starting with the shortest maxilla head length
(lowest mean percent), these six species (plus
koreanus and multiradiatus) rank as follows: 1)
concolor, 18.8; 2) brasiliensis, 19.0; 3) sierra,
19.8; 4) tritor, 19.9; 5) koreanus, 20.2; 6) macu-
latus,20.6; 7) multiradiatus, 21.0; and 8) tritor,
21.1. The longest heads are found in niphonius
(24.7), semifasciatus (24.1), and lineolatus (24.0).
The head of the maxilla is a little longer, relative
to total maxilla length, in Grammatorcynus
(26%) and much longer (33%) in Acanthocybium.
The posterior expansion of the maxilla is least
well-developed (lowest) relative to maxilla length
in S. multiradiatus (8-9%) and S. sinensis (9-
11%). The best-developed posterior expansion is in
S. plurilineatus (15%). The other 15 species range
from 11 to 14%. This range of variation is shown
in S. munroi but the specimen illustrated (Fig. c
23b) shows a relatively well-developed posterior
FIGURE 24. - Left dentaries in lateral view. a. Scomberomorus
expansion. The shape of the posterior expansion semifasciatus, Port Moresby, New Guinea, 510 mm FL, 2 x. b.
varies within and between species, but most of Acanthocybium solandri, Miami, Fla., 1,403 mm FL, 1 x. c.
the expansion is usually ventral. Grammatorcynus bilineatus, Marshall Is., 424 mm FL, 2 x .

578
COLLETTE and RUSSO: SPANISH MACKERELS

All the species of Scomberomorus and Gramma- Devaraj (1977) stated that the anterior notch was
torcynus have a notch on the anteroventral mar- distinct in S. cavalla and S. commerson. The
gin of the dentary. This notch is absent in Acan- notch may be a little more prominent in S.
thocybium. The notch seems to vary as much commerson than in the other species, but we
between specimens of a species of Scomberomo- cannot confirm this for S. cavalla.
rus as between species of the genus. Acanthocyb-
ium has a prominent notch on the anterior mar- Angular. -The triangular anterior end of the
gin of the dentary (Fig. 24b) which is indistinct or angular (frequently called articular) fits into the
absent in Scomberomorus and Grammatorcynus. dentary anteriorly (Fig. 25). The posterior end of

RETROARTICULAR

FIGURE 25.-Left angulars and retroarticulars in lateral view. a. Scomberomorus semifasciatus, Port
Moresby, New Guinea, 510 mm FL, 3.5x. b.Acanthocybium solandri, Miami, Fla., 1,403 mm FL, IX.
c. Grammatorcynus bilineatus, New Guinea, 382 mm FL, 4.5x.

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 FISHERY BULLETIN: VOL. 82, NO.4

the angular bears three large processes; the dor- Retroarticular.-The retroarticular bone (fre-
sal process directed forward and upward, the ven- quently called angular) is rhomboid and attached
tral process directed forward, and the posterior firmly, but not fused to the posteroventral mar-
process directed backward and upward. This pro- gin of the angular (Fig. 25). No differences were
cess is hooked and carries a transverse articular found among the retroarticulars of the species of
facet for the quadrate. Between the dorsal and Scomberomorus.
ventral processes is Meckel's cartilage which ex-
tends directly anterior into the space between the PALATINE ARCH. - The palatine arch con-
two arms of the dentary. The length of the angu- sists of four pairs of bones in the roof of the
lar to the tip of the dorsal process is 31-42% of the mouth: palatine, ectopterygoid, entopterygoid,
total length of the bone; the length to tip of the and metapterygoid.
ventral process is 42-53% of the total length. The
maximum width of the angular, measured from "
Palatine.-The palatine (Fig. 26) is forked
the tip of the dorsal process to the tip of the both posteriorly and anterolaterally. The dorsal
ventral process is 34-43% of the total length. branch of the anterolateral fork is hooked, and
Devaraj (1977) stated that the ventral process its anterior end articulates with a facet on the
was longer and narrower in S. commerson and maxilla, immediately ventral to the nasal. The
Acanthocybium than in other Indian species ventral branch is cone-shaped or pointed. The
and we confirm this. The ventral process is as exterior branch ofthe posterior fork carries on its
long or longer than the dorsal process in S. dorsal surface the shank ofthe ectopterygoid, and
commerson (ventral process 99-162% of the dorsal the inner, flat, thin branch is attached to the
process), Acanthocybium (99-148%), and also in anterior end of the entopterygoid. The lateral
S. queenslandicus (115-136%). The next longest aspect of the palatine is roughly triangular and
ventral processes are in S. caualla (80-104%) and concave', and closely attached to the mesial wall
S. sinensis (82-97%). The other 14 species of ofthe maxilla. Grammatorcynus (Fig. 26d) differs
Scomberomorus (and Grammatorcynus) have from Scomberomorus (Fig. 26a, b) and Acantho-
shorter ventral processes, 40-85% of the length of cybium (Fig. 26c) in almost lacking the anteriorly
the dorsal process. The shortest ventral process is directed ventral branch. Acanthocybium has a
in S. regalis, 40-44%. distinct ventral branch but it is shorter than the

a b

c d
FIGURE 26. - Left palatines in lateral view, slightly rotated to better show tooth patch.
a. Scomberomorus semifasciatus,
New Guinea, 740 mm FL, 2x. b. Scomberomorus commerson, New South Wales, 1,155 mm FL, IX. c. Acanthocybium
solandri, Miami, Fla., 1,403 mm FL, 1 x. d. Grammatorcynus bilineatus, Timor Sea, 453 mm FL, 2 x .

580
COLLE'ITE and RUSSO: SPANISH MACKERELS

dorsal branch, as pointed out by Devaraj (1977), widest tooth patch is in S. semifasciatus (Fig.
whereas the ventral branch is longer than the 26a), almost as wide as in Grammatorcynus but
dorsal branch in all species of Scomberomorus. with much finer teeth.
The distance from the anterior end of the ventral
branch to the end of the external branch divided Ectopterygoid. -The ectopterygoid (Fig. 27) is
by the distance from the tip of the dorsal hook to a T-shaped bone, the top of the T forming its
the end of the external branch is 120-123% in posterior end. It is joined with the entopterygoid
Grammatorcynus, 112-121% in Acanthocybium, dorsolaterally, the palatine laterally and anteri-
and only 87-107% in the species of Scomberomo- orly, and the quadrate and metapterygoid poste-
rus. Acanthocybium differs from both Scomber- riorly. The dorsal arm of the ectopterygoid is
omorus and Grammatorcynus in having the pos- shorter than the ventral arm in Scomberomorus
teriorly directed inner branch almost as long as and vice versa in Acanthocybium and Gramma-
the outer branch. The distance from the tip of the torcynus. This relationship can be expressed by
dorsal hook to the tip of the inner branch divided dividing the dorsal distance (from the anterior
by the distance to the tip of the outer branch is end ofthe bone to the tip of the dorsal arm) by the
97-99% in Acanthocybium and 54-84% in the ventral distance (from the anterior end to the tip
species of Scomberomorus and Grammatorcynus. of the ventral process). The range is 85-100% in
The tooth patch is long and narrow in Acantho- the species of Scomberomorus compared with
cybium (Fig. 26c), short and wide in Grammator- greater than 100% in Acanthocybium (103-109%)
cynus (Fig. 26d), and with the species of Scom- and Grammatorcynus (110-116%). The shank is
beromorus in between these extremes. The teeth longer in Acanthocybium than in the other two
are fine in all three genera, but a little larger in genera. The posterior edge of the ectopterygoid
Acanthocybium and Grammatorcynus than in (from the tip ofthe dorsal process to the tip of the
most species of Scomberomorus. ventral process) is shorter relative to the ventral
The species of Scomberomorus show some dif- distance in Acanthocybium (41-47%) than in the
ferences in the length of the ventral branch species of Scomberomorus (43-63%) and Gram-
relative to that of the length of the external matorcynus (64-68%).
branch, the relative length of the outer to the The ectopterygoids of the species of Scomber-
inner branch, the relative width of the tooth omorus are very similar. The shortest ventral
patch, and the size of the teeth in the tooth patch. distance is in S. sinensis, 85-88% of the dorsal
Dividing the length of the ventral margin, from distance, the longest in S. regalis, 99-100%. The
the anterior end of the ventral branch to the end shortest posterior edges are in S. niphonius and
of the external branch, by the length ofthe dorsal S. tritor (50-51% of the dorsal distance), the
margin, from the tip ofthe dorsal hook to the end longest posterior edges are in S. korean us (61-
of the ventral branch, shows three species of 63%), S. plurilineatus (60-63%), and S. semi-
Scomberomorus-sinensis (98-107%), tritor (100- fasciatus (59-62%);
102%), and commerson (94-102%)-to be most
similar to Acanthocybium (112-121%). The lowest Entopterygoid. - The entopterygoid is elongate
figures are for S. niphonius (87-88%). Dividing and o~al in shape (width 23-46% oflength) (Fig.
the length of the dorsal margin by the distance 28). The outer margin of the entopterygoid is the
from the tip of the dorsal hook to the end of the thickest part of the bone and is attached to the
inner branch shows four species of Scomberomo- inner margin ofthe ectopterygoid. The entoptery-
rus-plurilineatus (75-84%), munroi (77-79%), goid also connects with the palatine anteriorly
lineolatus (72-74%), and semifasciatus (70-73%) and the metapterygoid posterolaterally. The me-
-to resemble Grammatorcynus (71-75%). The sial and posterior borders are free from contacts
lowest figures are for S. multiradiatus (54-56%). with other bony elements. The dorsal surface is
The tooth patch is very narrow in S. commerson concave and the smooth convex ventral surface
(Fig. 26b), similar to the patch shape in Acantho- forms the major part of the buccal roof. The
cybium but with finer teeth. The tooth patch is anterior end is narrower than the posterior end in
also narrow in a 677 mm FL specimen of S. most species but a little wider in S. guttatus and
sinensis and reduced to only a single row of teeth S. koreanus. The entopterygoid is narrowest in S.
in a 1,082 mm specimen. The teeth in S. sinensis commerson (width 23-28% of length, Fig. 28a)
are larger than in other species of the genus, at and S. multiradiatus (29%). The shortest and
least the same size as in Acanthocybium. The widest entopterygoids are in sinensis (39-46%,
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 FISHERY BULLETIN: VOL. 82, NO.4

HYOMANDIBULA

QUADRATE

SYMPLECTIC

ECTOPTERYGOID

FIGURE 27.-Left suspensoria in mesial view. a. Scomberomorus semifasciatus,

Port Moresby, New Guinea, 510 mm FL, 2.5 x. b. Acanthocybium solandri, Revil-
lagigedos Is., 1,068 mm FL, 1.5x. c. Grammatorcynus bilineatus, Marshall Is.,
424 mm FL, 2x.

582
COLLE'ITE and RUSSO: SPANISH MACKERELS

a b

c d
FIGURE 28.-Left entopterygoids in dorsal view. a. Scomberomorus commerson, New South Wales, 1,155 mm FL, Ix. b. Scomber-
omorus sinensis, Hong Kong, 677 mm FL, 2 x. c. Acanthocybium solan.dri, Indian Ocean, 943 mm FL, 2x. d. Grammatorcynus
bilineatus, Marshall Is., 424 mm FL, 2.5 x .

Fig. 28b), maculatus (41-42%), and concolor with the dorsal end of the symplectic in Acantho-
(40-42%). Acanthocybium (Fig. 28c), Grammator- cybium and Grammatorcynus (Fig. 27b, c), but
cynus (Fig. 28d), and the other 13 species of Scom- not in most species of Scomberomorus (Fig. 27a).
beromorus are intermediate in width (30-40%). The posterior horizontal part of the ventral bor-
der is longer than the anterior oblique part in
Metapterygoid. - The metapterygoid (Fig. 27) Scomberomorus (anterior part 39-86% of poste-
is a flat, quadrangular or somewhat triangular rior part), but vice versa in Acanthocybium and
bone. The posterodorsal margin of this bone is Grammatorcynus (anterior part 132-218% of pos-
deeply grooved to receive the hyomandibula. The terior part).
dorsal portion is strongly ankylosed to the lamel- The anterior part of the ventral margin is
lar region of the hyomandibula. The ventroposte- relatively longer in S. multiradiatus (77-78% of
rior margin abuts the lowermost portion of the posterior part) and S. maculatus (65-86%), and
symplectic process of the hyomandibula, but does relatively shorter in S. plurilineatus (41-45%)
not touch the hyomandibula. There is a relatively and S. regalis (39-50%). Devaraj (1977) reported
long slit between the two bones, through which differences in the shape ofthe anterior free border
the hyoidean artery passes (Allis 1903). The ven- of the metapterygoid, as convex, nearly straight,
tral border is divided into two portions, the hori- or concave. We have found similar tendencies but
zontal portion in contact with the quadrate and it is difficult to place the species of Scomberomo-
the anterior oblique portion ankylosed to the rus in specific categories.
ectopterygoid. On the mesial surface, the meta-
pterygoid possesses a triangular-shaped area HYOID ARCH. - The hyoid arch is the chain of
which forms an interdigitating articulation with bones that connect the lower jaw and the opercu-
the upper arm of the ectopterygoid. The postero- lar apparatus with the skull. The arch is com-
ventral margin of the metapterygoid articulates posed of the hyomandibula, symplectic, quadrate,
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 FISHERY BULLETIN: VOL. 82, NO.4

hyoid complex (hypohyal, ceratohyal, epihyal, that fits into a groove on the inner surface of the
interhyal, and the seven branchiostegal rays), quadrate (Fig. 27). The symplectic is very narrow
and two median unpaired bones, the glossohyal in Scomberomorus, not filling the groove in the
and urohyal. quadrate (Fig. 27a). It is slightly wider in Gram-
matorcynus but the groove is narrower, the sym-
Hyomandibula. -The hyomandibula (Fig. 27) plectic more nearly filling the groove (Fig. 27c).
is an inverted L-shaped bone that connects the The symplectic is greatly expanded at its dorsal
mandibular suspensorium and opercular bones to end in Acanthocybium (Fig. 27b). In most species
the neurocranium. There are three prominent of Scomberomorus, the symplectic, like the poste-
condyles on the dorsal end of the hyomandibula. rior process ofthe quadrate, extends only a slight
The long dorsal condyle forms the base of the L distance beyond the dorsal margin of the quad-
and fits into the fossa at the junction of the rate. The symplectic is slightly longer than the
pterotic and sphenotic bones. The anterior con- posterior process in a species with a short process
dyle articulates with the ventral fossa of the (e.g., S. multiradiatus) and in one with a rela-
pterotic and the lateral process is attached to the tively long process (e.g., S. sinensis). No bony
inside of the opercle. Anterolaterally, the hyo- contact is present between the dorsal end of the
mandibula is drawn out into a lamellar region symplectic and either the metapterygoid or the
that joins the metapterygoid; posterolaterally, it hyomandibula in most species of Scomberomo-
has a long articulation with the preopercle. Ven- rus. The metapterygoid is in slight contact with
trally, the hyomandibula has a long symplectic the symplectic in S. sinensis and S. koreanus.
process; at the posterodorsal corner there is a Both Grammatorcynus and Acanthocybium have
small spine. A strong vertical ridge extends from much longer symplectics, extending well beyond
the ventral margin to a little below the dorsal the dorsal margin of the quadrate and even
border, thence it curves anteriorly to confluence beyond the dorsal end of the posterior process
with the anterior condyle. The portions lying to make firm contact with the metapterygoids.
anterior and posterior to this ridge are grooved Devaraj (1977:fig. 11) illustrated the symplectics
for articulation with the metapterygoid and pre- by themselves for the four Indian species (korea-
opercle respectively; in situ only the ridge and a nus, guttatus, lineolatus, and commerson) and
portion of the upper broader surface are visible Acanthocybium.
exteriorly. The upper surface of the symplectic is
connected to the ventral border of the hyoman- Quadrate.-The lower jaw is suspended from
dibula by way of a cartilage which is especially the cranium by means of the articulating facet of
well developed in Acanthocybium. There are two the ventral surface of the triangular quadrate.
deep fossae on the inner surface of the hyoman- The broad dorsal margin of the quadrate abuts
dibula of Acanthocybium but only one in Scom- the ventral border ofthe metapterygoid (Fig. 27).
beromorus and Grammatorcynus. The mesial surface of the quadrate bears a deep
The posterodorsal spine is best developed in groove which accepts the symplectic. There is a
Acanthocybium (Fig. 27b) and S. commerson, as strong process on the posterior margin of the
pointed out by Devaraj (1977). This spine is also quadrate that is attached along the lower ante-
well developed in S. queenslandicus and is pres- rior arm ofthe preopercle. The process is relative-
ent but small in the other 16 species of Scom- ly short in Scomberomorus, extending only a
beromorus (e.g., S. semifasciatus, Fig. 27a). No short distance beyond the dorsal margin of the
spine is present in Grammatorcynus (Fig. 27c). quadrate in most species (e.g., S. semifasciatus,
The total length of the hyomandibula (ventral tip Fig. 27a). The process is shortest in S. multiradia-
to dorsal margin of dorsal condyle) is greater tus, not reaching the dorsal margin. The process
relative to maximum width (tip of anterior con- is longest in S. commerson, S. lineolatus, and S.
dyle to outer margin of posterior condyle) in sinensis, but it is still shorter in these three
Grammatorcynus (width 35-36% of length) and species than in Acanthocybium (Fig. 27b) and
S. multiradiatus (36-39%). The hyomandibula is Grammatorcynus (Fig. 27c). An attempt was
shortest relative to width in S. sinensis (45-52%). made to quantify this by measuring from the
Acanthocybium (41-44%) is similar to the major- inside ofthe articular facet to the tip ofthe dorsal
ity of species of Scomberomorus (39-47%). process and to the tip of the anterior margin of
the quadrate. The short process in S. multiradia-
Symplectic.-The symplectic is a small bone tus is shown by the distance to the anterior
584
COLLETTE and RUSSO: SPANISH MACKERELS

margin being about equal (95-103%) to the dis- further posterior due to also having an anterior
tance to the tip of the process. In the other species pointed end to the hypohyals at the junction of
of Scomberomorus, the distance to the anterior the dorsal and ventral hypohyals. In addition,
margin is less than (76-96%) the distance to the Acanthocybium has a prominent anterolateral
tip of the process. This percent is low in S. process on the ventral hypohyal. The groove for
lineolatus (76%), indicative of a long process, but the hyoidean artery runs along the outer surface
the figures for S. commerson (80-83%) and S. of the epihyal, ceratohyal, and ventral portion of
sinensis (83-85%) are not much lower than those the dorsal hypohyal. The groove extends anteri-
for many other species with shorter processes. orly 29-54% of the length of the dorsal hypo-
The lowest figures are for Acanthocybium (72- hyal before becoming a covered tunnel in Scom-
80%) and Grammatorcynus (65-71%), indicative beromorus and Grammatorcynus or a foramen in
of the long process in these two genera. Acanthocybium leading to the inner side of the
dorsal hypohyal. The opening on the inner side
Hyoid complex.-This complex includes the appears as a small to moderate pit usually lo-
two hypohyals (= basihyal of Mago Leccia 1958), cated in the ventral portion of the dorsal hypo-
ceratohyal, epihyal, and interhyal bones, and the hyal in Scomberomorus and Grammatorcynus.
seven branchiostegal rays (Fig. 29). The hypo- The pit lies astride the junction of the dorsal and
hyals, ceratohyal, and epihyal are closely asso- ventral hypohyals in S. brasiliensis and extends
ciated and form a functional unit. slightly into the ventral hypohyal in S. macula-
tus and S. sierra. The pit also is larger in these
Hypohyals. -The hypohyals are composed of species.
separate dorsal and ventral elements joined lon-
gitudinally. In lateral view, the ventral hypohyal Ceratohyal. -The ceratohyal is a long flat
is clearly larger than the dorsal hypohyal in all bone, broadest at the posterior end and with an
species of Scomberomorus and in Grammator- anteroventral projection that articulates with the
cynus (Fig. 29a, c). The ventral hypohyal is about posteroventral notch of the ventral hypohyal. It is
three times larger than the dorsal in Acanthocyb- the largest bone of the hyoid complex. Posterior-
ium (Fig. 29b). Devaraj (1977:29) stated that the ly, the middle part of the ceratohyal interlocks
dorsal and ventral hypohyals were of equal size with the epihyal by means of odontoid processes
in S. commerson, but we find the ventral larger issuing from both elements (ceratohyal-epihyal
in lateral view, as in the other species of the suture of McAllister 1968), while the upper and
genus. In mesial view, the dorsal and ventral lower portions are joined by cartilage. Four acin-
hypohyals in S. commerson and the other 17 aciform branchiostegal rays are attached to the
species are about equal in size. The ventral hypo- respective articular surfaces along the concave
hyal is perhaps a little larger than the dorsal in middle portion of the ventral margin. In Scom-
mesial view in S. multiradiatus and S. queens- beromorus (Fig. 29a) the fifth branchiostegal ray
landicus. Laterally, the suture between the dor- usually is attached to the most posterior part of
sal and ventral hypohals runs almost horizontal- the ceratohyal or on the space between the cera-
ly in Acanthocybium but curves ventrally at tohyal and epihyal, not on the anterior part of the
various angles in Scomberomorus and Gramma- epihyal as stated by Devaraj (1977) and Mago
torcynus. Devaraj (1977) stated that it formed "an Leccia (l958:pl. 4). In Acanthocybium and Gram-
upward curve anteriorly in S. koreanus, S. lineo- matorcynus, the fifth ray is on the anterior part of
latus, S. regalis , and S. niphonius and runs the epihyal (Fig. 29b, c). The hyoidean groove
nearly straight in the other species including A. runs the length of the ceratohyal on its lateral
solandri." The specimen of S. commerson that he surface. The groove is so deep in some specimens
illustrated (figure 12D) does show a straight of some species that it forms a thin slit through
suture, but in our material a downward curve the bone, the ceratohyal window or beryciform
usually is present. Mesially, a pointed lateral foramen. Slits are common in 10 species of Scom-
process at the anterodorsal end of the dorsal beromorus: brasiliensis, commerson, concolor,
hypohyal forms a symphysis with the glossohyal, multiradiatus, munroi, niphonius, queensland-
urohyal, basibranchial, and the process of the icus, semifasciatus, sierra, and tritor; rare in
hypohyal from the opposite side in Scomberomo- four, cavalla, plurilineatus, maculatus, and si-
rus and Grammatorcynus. Acanthocybium also nensis plus Acanthocybium and Grammator-
has a pointed lateral process but it appears to be cynus; and occasional in the other four species:
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 FISHERY BULLETIN: VOL. 82, NO.4

CERATOHYAL EPIHYAL
INTERHYAL

HYPOHYALS

BRANCHIOSTEGAL RAYS
a
HYOIDEAN GROOVE

FIGURE 29.-Left hyoid complexes in lateral view. a. Scomberomorus commerson, New South Wales, 1,155 mm
FL, 1 x. b. Acanthocybium solandri, Miami, Fla., 1,403 mm FL, 1 x. c. Grammatorcynus bilineatus, Timor Sea,
453 mm FL, 2x.

guttatus, koreanus, lineolatus, and regalis. Both deeply concave and very much constricted in
large (S. commerson) and small (S. multiradia- Acanthocybium such that the dorsal margin of
tus) species have slits. Smaller specimens of a the bone comes closer to the groove for the hy-
species sometimes have slits (guttatus, pluriline- oidean artery. The margin is straight in Gramma-
atus, queenslandicus, regalis, and semifasciatus), torcynus and varies in Scomberomorus. Devaraj
while larger specimens lack them; sometimes the (1977:30-31) stated that the dorsal margin of the
situation is reversed (koreanus, lineolatus, and ceratohyal is convex in some species (koreanus
tritor). The dorsal margin of the ceratohyal is and lineolatus), almost straight in others (gut-
586
COLLE'ITE and RUSSO: SPANISH MACKERELS

tatus and niphonius), and slightly concave in

most (commerson, maculatus, regalis, and caval-
La). We find similar tendencies, but there is
extensive variation even in small samples. Six
species tend to have the dorsal margin convex:
guttatus, korean us , lineolatus, multiradiatus,
plurilineatus, and semifasciatus; seven species
tend to have the dorsal margin concave: cavalla,
commerson', maculatus, munroi, queenslandicus,
regalis , and sinensis; and five usually have the
dorsal margin nearly straight: brasiliensis, con-
color, niphonius, sierra, and tritor. a b
Epihyal. -The epihyal is a triangular bone
which interlocks anteriorly with the ceratohyal.
It has a posterior process which articulates with
the interhyal. In Scomberomorus, two branchios-
tegal rays are seated on the ventral portion of
the epihyal, not three as stated by Devaraj (1977)
or shown by Mago Leccia (1958). Three branchios-
tegal rays do articulate with the epihyal in
Acanthocybium and Grammatorcynus. The depth
of the epihyal is least in Acanthocybium, 58-62%
of the length from the smooth anterior margin of
the bone to the tip of the posterior process. Two
species of Scomberomorus (commerson and cav- c d
alta) have relatively low epihyals, 68-71% of FIGURE 30.-Glossohyals in dorsal view. a. Scomberomorus
length. Grammatorcynus also has a relatively plurilineatus, Natal, 910 mm FL, 4 x. b. Scomberomorus
low epihyal, 66-77% of length. The deepest epi- munroi, New Guinea,' 512 mm FL, 5 x. c. Acanthocybium
hyals are in four species of Scomberomorus: ko- solandri, Indian Ocean, 1,088 mm FL, 2 x. d. Grammatorcynus
reanus (90-98%), concolor (86-94%), plurilineatus bilineatus, Queensland, 521 mm FL, 4 x .
(87-91%), and guttatus (87-90%).

Interhyal. -The interhyal is a small flattened cynus differs in having a quadrangular to oval
bone that is attached to the epihyal dorsal to tooth plate fused to and covering the dorsal
the posterior process. The interhyal is directed surface of the bone (Fig. 30d). Two bonitos, Cyb-
obliquely upward and links the hyoid complex to iosarda and Orcynopsis, have a similar condition
the hyomandibula and symplectic. No differences but there are two separate oval tooth patches in
were noted among interhyals. these genera (Collette and Chao 1975:fig. 43a, b).
Another bonito, Gymnosarda, has what appears
Glossohyal. -The glossohyal (basihyal) (Fig. to be a single tooth plate on the glossohyal, but
30) is a median bone that supports the tongue and this plate is actually composed of left and right
overlies the first basibranchial bone at the ante- portions that fit over the bone rather than being
rior end of the branchial arch: In Scomberomo- fused to it (Collette and Chao 1975:fig. 43f). The
rus, the glossohyal is roughly rod-shaped or coni- glossohyal is a little wider in Grammatorcynus
cal in most species. Its width is 35-54% of its than in Acanthocybium or most species of Scom-
length. It generally has a flat or narrowed ante- beromorus, 47-55% oflength.
rior end and broadens posteriorly, but terminates The size of the ventral protrusion varies among
in a small posterior cone or flattened projection. the species of Scomberomorus. It is greatest in S.
The glossohyal protrudes ventrally adjacent to sinensis, commerson, and cavalta. The glossohyal
the posterior articulation. The glossohyal of is narrowest in S. multiradiatus and plurilinea-
Acanthocybium is flattened and spatulate with a tus (Fig. 30a), 35-36% of width. It is widest in
broad anterior end, a narrow posterior end, and S. sierra and munroi (Fig. 30b), 52-54%. The
no ventral protrusion (Fig. 30c). Grammator- anterior end is widest in S. niphonius and sinen-
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 FISHERY BULLETIN: VOL. 82, NO.4

sis, narrowest in brasiliensis, cavalla, and compared with 16-24%. Grammatorcynus also
commerson. has a low urohyal, depth 15-17% of length. The
length of the ventral margin is 68-91% of the
Urohyal. -The urohyal (Fig. 31) is a com- length of the dorsal margin. The ventral margin
pressed, median, unpaired bone. The anterior end of the urohyal does not extend as far posteriorly
of this element lies between, and is connected in Grammatorcynus, only 68-69% of the length
with, the hypohyals of the left and right sides. of the dorsal margin compared with 80-91% in
The dorsal and ventral margins are thickened. Acanthocybium and Scomberomorus. Both Mago
The anterior end has an articulation head and Leccia (1958:322) and Devaraj (1977:32) stated
the posterior end is deep. The maximum depth that the posterior end of the dorsal margin was
posteriorly is 13-24% of the length of the dorsal pointed but it ends in a distinct fork in all species
margin. The urohyal is not as deep in Acantho- of Scomberomorus and in Acanthocybium (Fig.
cybium as in the species of Scomberomorus, 31c). The major difference in Grammatorcynus is
depth 13-15% of the length of the dorsal margin that the shape of the posterior end of the dorsal

FIGURE 31.-Urohyals in left lateral view. a. Scomberomorus queenslandicus, Queens-

land, 641 mm FL, 2x. b. Scomberomorus munroi, New Guinea, 512 mm FL, 2x. c.
Acanthocybium solandri, Indian Ocean, 1,088 mm FL, 1 x. d. Grammatorcynus bilineatus,
New Guinea, 382 mm FL, 3 x. Inset to right is the posterior end of the dorsal margin, in
dorsal view.

588
COLLETrE and RUSSO: SPANISH MACKERELS

margin is tripartite (Fig. 31d) instead of forked. beromorus (caualla, regalis, and tritor), there is
Some specimens of Acanthocybium differ from also a weak anteroventral process. Instead of a
the other two genera by having a slight indenta- distinct process at this point, Acanthocybium and
tion in the anterior end of the urohyal. the other species of Scomberomorus have an
angle where the anterior margin meets the ante-
OPERCULAR APPARATUS.-Four wide flat roventral margin. The posterior margin and/or
bones (opercle, preopercle, subopercle, and inter- the posteroventral margin of the opercle are
opercle) fit together to form the gill cover which fimbriate in Acanthocybium and most species of
protects the underlying gill arches. Scomberomorus (brasiliensis, koreanus. lineola-
tus, maculatus, niphonius, queenslandicus, semi-
Opercle.-The opercle is broad (Fig. 32), and it fasciatus (Fig. 32a), sinensis, and tritor). Gram-
is overlapped laterally on its anterior margin by matorcynus (Fig. 32d) has a much narrower and
the posterior half of the preopercle. The narrow, more elongate opercle than do Acanthocybium
elongate, articular facet for the opercular process or the species of Scomberomorus. The most elon-
of the hyomandibula is located on the mesial gate opercle among Scomberomorus species is in
surface of the anterodorsal corner of the opercle. S. multiradiatus (Fig. 32b). The broadest is in
Grammatorcynus (Fig. 32d) and most species of Acanthocybium.
Scomberomorus have a weak process at the pos-
terodorsal corner. This process appears to be Preopercle. -The preopercle (Fig. 33) is a large
absent in Acanthocybium (Fig. 32c) and Scom- crescent-shaped flat bone, broadest at the lower
beromorus sinensis. In several species of Scom- posterior angle. The anterior portion of the bone

FIGURE 32.-Left opercles in lateral view.

a. Scomberomorus semifasciatus, New Guin-
ea, 510 mm FL. b. Scomberomorus multi-
radiatus, New Guinea, 294 mm FL. c. Acan-
thocybium solandri, Revillagigedos Is., 1,080
mm FL. d. Grammatorcynus bilineatus,
Marshall Is., 424 mm FL.

a b

d
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 FISHERY BULLETIN: VOL. 82. NO.4

Grammatorcynus and most species of Scomber-

omorus. Devaraj (1977:34) stated that the poste-
rior margin was convex in S. commerson and
Acanthocybium. We find it to be nearly straight
in Acanthocybium and very slightly concave in
S. commerson. The concave posterior border
makes the upper and lower parts appear as two
limbs, the lower of which is longer. As Devaraj
(1977:34) noted, the lower portion is longer in S.
guttatus than in the other species, the distance
a from the anterior margin of the bony ridge to the
posterior end of the lower lobe being 74-80% of
the height of the preopercle measured from the
ventral margin to the dorsal tip of the bone.
b Other species with long lower portions include S.
munroi (73-78%), S. plurilineatus (69-79%), S.
niphonius (73-75%), and Grammatorcynus (68-
75%). Devaraj (1977:34) stated that the anterior
ridge was forked at its upper part in all the
Indian species of Scomberomorus except S. com-
merson in which the fork is either indistinct
or absent, and that the fork was completely ab-
sent in Acanthocybium. We are unable to con-
firm this observation and find no differences
between Scomberomorus and Acanthocybium. In
these genera, and in Grammatorcynus, the an-
terodorsal margin terminates in a pore similar to
the preopercular lateral line canal pore at the
anteroventral margin of the bone.

c d Subopercle. - The subopercle is a flat triangu-

lar bone with a prominent anterior projection
FIGURE 33.-Left preopercles in lateral view. a. Scomberomo-
(Fig. 34). Two ridges converge posteriorly from
rus semifasciatus, New Guinea, 510 mm FL. b. Scomberomorus
multiradiatus, New Guinea, 294 mm FL. c. Acanthocybium the anterior projection on the lateral side of the
solandri, Revillagigedos Is., 1,068 mm FL. d. Grammatorcynus bone. The upper ridge articulates with the lower
bilineatus, Marshall Is., 424 mm FL. posterior projection of the opercle and the lower
ridge connects to the posterodorsal margin of the
interopercle. The dorsal ridge is much stronger
is thickened into a bony ridge. A series of 5-7 than the ventral ridge and extends over the main
pores along the lower margin of the ridge repre- part of the subopercle as a discrete shelf. The
sents the preopercular canal of the lateral line much weaker ventral ridge is difficult to detect in
system which continues into the dentary. On the some species. The angle between the anterior
mesial side, the ridge possesses a groove for projection and the anterior margin of the sub-
attachment to the hyomandibula and the quad- opercle varies from approximately a right angle
rate. There is a shelf mesial to the anteroventral in Acanthocybium (Fig. 34c) and most species of
end of the preopercle in Acanthocybium (Fig. Scomberomorus to acute in Grammatorcynus
33c) that is not present in Scomberomorus (Fig. (Fig. 34d) and Scomberomorus multiradiatus
33a, b) or Grammatorcynus (Fig. 30d). Devaraj (Fig. 34b). The length of the anterior projection
(1977:34) referred to this as a groove. The canals varies from 20 to 45% ofthe length ofthe anterior
leading to the preopercular pores are visible margin dorsal to the projection. The projection is
through the bone in Scomberomorus and Gram- longest in Acanthocybium (36-45%), S. sierra
matorcynus but cannot be seen in Acanthocybium (37-43%), and S. koreanus (33-41%). It is shortest
due to the thickness of the bone. The posterior in S. commerson (20-25%), S. semifasciatus
margin of the preopercle is distinctly concave in (21-23%, Fig. 34a), and S. queenslandicus (21-
590 1$
COLLEITE and RUSSO: SPANISH MACKERELS

a b

c d
FIGURE 34.-Left subopercles in lateral view. a. Scomberomorus semifasciatus, New
Guinea, 510 mm FL. b. Scomberomorus multiradiatus, New Guinea, 294 mm FL. c.
Acanthocybium solandri, Revillagigedos Is., 1,068 mm FL. d. Grammatorcynus bilinea-
tus, Marshall Is., 424 mm FL.

27%). It is also short, relative to the long, narrow 35d) and Acanthocybium (40-49%, Fig. 35c) have
subopercle, in Grammatorcynus (25-26%). Dev- lower interopercles than most species of Scom-
araj (1977:33) mentioned differences in the shape beromorus (Fig. 35a, b). The shallowest inter-
of the posteroventral margin and the dorsal edge opercles in this genus are in S. plurilineatus
of the subopercle, but we have not noted any (45-47%), S. munroi (47-49%), S. niphonius
consistent differences between species in these (47-49%), and S. semifasciatus (47-51%, Fig. 35a).
regions. A well-formed notch anterior to the crest on the
sloping anterior margin in Scomberomorus and
Interopercle.-The interopercle (Fig. 35) is Grammatorcynus is relatively poorly developed
roughly oval in shape with a crest on the superior in Acanthocybium, rendering the superior margin
margin. There is a well-developed facet on the nearly straight. The posterior margin is rounded
mesial side to receive the articular process of the in Scomberomorus and Grammatorcynus but di-
interhyal. The depth of the interopercle varies vided into two by a notch in Acanthocybium.
from 37 to 61% of the length of the bone. The
deepest interopercles are in Scomberomorus si- BRANCHIAL APPARATUS.-The branchial
nensis (54-61%, Fig. 35b) and S. sierra (57-58%). apparatus is composed of the five pairs of gill
The interopercles are moderately deep (50-58%) arches, gill filaments, gill rakers, pharyngeal
in seven species: brasiliensis, commerson, ko- tooth patches, and supporting bones. The general
reanus, lineolatus, multiradiatus, queenslandi- arrangement in the Scomberomorini (Fig. 36) is
cus, and tritor. Grammatorcynus (37-42%, Fig. similar to that found in other scombrids such as

591
 FISHERY BULLETIN: VOL. 82, NO.4

c d
FIGURE 35.-Left interopercles in lateral view. a. Scomberomorus semifasciatus, New Guinea, 510 mm FL. b. Scomber-
omorus sinensis, Hong Kong, 677 rom FL. c. Acanthocybium solandri, Revillagigedos Is., 1,068 mm FL. d. Grammator-
cynus bilineatus, Marshall Is., 424 rom FL.

the Sardini (Collette and Chao 1975), Thunnus branchial. The third basibranchial has an ex-
(Iwai and Nakamura 1964b:22, fig. 1; de Sylva panded anterior end at its junction with the
1955:21, fig. 40), Scomberomorus (Mago Leccia second basibranchial and then tapers posteriorly.
1958:327, pI. 12), and Rastrelliger (Gnanamuttu A prominent groove is present anteriorly which
1971:14, fig. 6). Within the Scomberomorini, the accepts the medial anterior end of the second
most useful differences are in the number of gill hypobranchial. A section of cartilage extends
rakers. Most of the branchial bones bear patches posteriorly to articulate with the fourth and fifth
oftiny teeth. ceratobranchials.

Basibranchials. - The three basibranchials Hypobranchials. - Three hypobranchials are

form an anteroposterior chain. The first and present. The first is interposed between the sec-
second are about the same size and considerably ond basibranchial and the first ceratobranchial.
shorter than the third. The first is covered dorsal- The second hypobranchial is about the same size
ly by the glossohyal. as the first, fits into a groove on the third basi-
In lateral view the first basibranchial is nar- branchial, and extends to the second cerato-
rowest in the middle. In Scomberomorus, it is branchial. The third hypobranchial is smaller
short with a wide base where it joins with the than the first or second, fits snugly against the
second basibranchial but it is much more elon- posterolateral margin of the third basibranchial
gate in Acanthocybium and Grammatorcynus. and its posterior end articulates with the third
The second basibranchial has a prominent notch ceratobranchial.
in the ventral margin and a distinct groove
laterally which extends from the anteroventral Ceratobranchials. - The five ceratobranchials
margin to the middorsal region of the bone. This are the longest bones in the branchial arches.
groove accepts the anterior end of the first hypo- They have a deep groove ventrally for the bran-
592
COLLETI'E and RUSSO: SPANISH MACKEREUl

chial arteries and veins. The ceratobranchials longest and bears two processes mesially. The
support most of the gill filaments and gill rakers. anterior process articulates with the first pharyn-
The first three are morphologically similar and gobranchial, and the posterior process attaches
articulate with the posterior ends of their respec- with the interarcual cartilage. The second epi-
tive hypobranchials. The fourth is more irregular branchial is similar to the first, but slightly
and attaches to a cartilage posterior to the third shorter. The anterior end is divided into two
basibranchial. The fifth ceratobranchial is also processes: the anterior process attaches to the
attached to the cartilage, has a dermal tooth second pharyngobranchial and the posterior pro-
plate fused to its dorsal surface, and the complex cess is coupled with the third pharyngobranchial
is termed the lower pharyngeal bone. It is covered by way of an elongate cartilage. This process is
with small conical teeth that are directed slightly much more elongate in Grammatorcynus than in
posteriad. Acanthocybium or Scomberomorus. The third
epibranchial is the shortest in the series. Lateral-
Epibranchials. - The posterolateral end of each ly, it is attached with the third ceratobranchial;
ofthe four epibranchials is attached to the ends of mesially, it is attached with the third pharyngo-
the first four ceratobranchials. Each epibranchial branchial. An elongate posterodorsal process is
bears a groove posterodorsally for the branchial present. This process joins with the fourth epi-
arteries and veins. The first epibranchial is the branchial. The fourth epibranchial is larger than

tt----IlASIIlRANCHIAL

----HYPOIlRANCHIAL

GILL RAKER~---

---CERATOIlRANCHIAL

UPPER
PHARYNGOIlRANCHIAL STAY

FIGURE 36.-Branchial apparatus of Scomberomorus semifasciatus, New Guinea, 510 mm FL. Dorsal view of the gill arches with the
dorsal region folded back to show their ventral aspect. Epidermis removed from right hand side to reveal underlying bones.

593
 FISHERY BULLETIN: VOL. 82, NO.4

the third and is interposed between the fourth between numbers of gill rakers, gap between gill
ceratobranchial and pharyngobranchial. It may rakers, and size of food items, as Magnuson and
be described as a curved bone with the angle Heitz (1971) have clearly shown for a number of
formed by the lateral and medial arms being species of Scombridae. The number of gill rakers
much more acute in Grammatorcynus than in is easily countable and is a useful taxonomic
Acanthocybium or Scomberomorus. A dorsal pro- character in Spanish mackerels as well as among
cess arises from the middle of the bone and other groups of the Scombridae.
attaches to the third epibranchial. Acantiwcybium differs from Grammatorcynus,
Scomberomorus, and the other genera of Scom-
Pharyngobranchials. -There are four pharyn- bridae by completely lacking gill rakers, Three
gobranchials attached basally to the epibranchial species of Scomberomorus have greatly reduced
of their respective gill arch. The first is long and numbers of gill rakers (Table 5): multiradiatus
slender, articulates dorsally with the prootic, and (1-4, sometimes only a single gill raker present,
is frequently called the suspensory pharyngeal at the junction of the upper and lower arches),
(Iwai and Nakamura 1964b). The elongate second commerson (1-8), and queenslandicus (3-9). One
pharyngobranchial bears a patch of teeth. The species, concolor, stands out from the rest of the
third is the largest element in the series; it has a genus in having many gill rakers, 21-27. Gram-
broad patch of teeth on its ventral surface, a matorcynus has more gill rakers (19"24) than 17
broad posterior end, and tapers to a narrow species of Scomberomorus but fewer than S.
anterior end. The third pharyngobranchial of concolor. There is a correlation between number
Scomberomorus is much more elongate than of gill rakers and number of jaw teeth (Tables 3,
those of Acanthocybiumand Grammatorcynus. 4) in Scomberomorus. The species with the fewest
The fourth pharyngobranchial also bears a ven- gill rakers, S. multiradiatus, also has the fewest
tral tooth plate, has a rounded posterior end, and jaw teeth (i 8.0 on the upper jaw, 7.8 on the lower
has an elongate strut (pharyngobranchial stay) jaw) and the species with the most gill rakers, S.
mesially which overlaps the third pharyngo- concolor, has the most teeth (i 22.2, 19.7).
branchial. This stay is much more elongate in
Scomberomorus than in Acanthocybium and AXIAL SKELETON
Grammatorcynus.
This section is divided into four parts: vertebral
Gill Rakers. -The hypobranchial, ceratobran- number, vertebral column, ribs and intermuscu-
chial, and epibranchial of the first gill arch sup- lar bones, and caudal complex.
port a series of slender, rigid gill rakers. The
longest gill raker is at or near the junction of the Vertebral Number
upper and lower arches, between the ceratobran-
chial and epibranchial. There is a correlation Vertebrae may be divided into precaudal (ab-

TABLE 5.-Total number of gill rakers on the first arch in Acantlwcybium, Grammatorcynus, and the species of Scomberomorus.
Species 0 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20. 21 22 23 24 25 26 27 N x
S. brasiliensis 9 49 37 22 11 129 13.8
S. cavalla 3 8 30 22 1 1 66 9.3
S. commerson 0 37 28 27 12 3 2 110 4.3
S. concolor 2 10 10 9 4 3 39 24.2
S. guttatus 3 7 24 53 25 9 2 123 11.1
S. koreanus 5 4 3 11 4 27 13.2
S. Iineolatus 0 3 9 12 3 1 29 10.5
S. maculatus 1 4 16 27 13 5 1 67 13.0
S. mulliradiatus 2 12 10 3 27 2.5
S. munroi 3 4 1 8 10.8
S. niphonius 3 18 14 2 38 12.5
S. plurilineatus 10 15 8 34 13.0
S. queenslandicus 2 3 3 11 13 1 34 6.2
S. regalis 1 2 4 19 13 6 46 15.4
S. samifasciatus 1 11 9 5 3 1 32 9.8
S. sierra 1 5 15 33 22 6 82 15.1
S. sinensis 7 6 1 1 16 12.6
S. tritor 7 18 11 5 41 13.3
Acanthocybium 30 30 0
Grammatorcynus 3 9 17 8 5 1 43 21.1

594
COLLETrE and RUSSO: SPANISH MACKERELS

dominal) and caudal (Tables 6-8). The first caudal Within Scomberomorus, S. multiradiatus has
vertebra is defined as the first vertebra that the most vertebrae (54-56), followed by S. macu-
bears a notably elongate haemal spine and lacks latus (51-53), S. munroi (50-52), and S. guttatus
pleural ribs. Vertebral counts include the uro- (47-52). The fewest vertebrae are found in S.
style which bears the hypural plate. Of the three cavalla (41-43) and S. sinensis (41-42). Vertebral
genera, Acanthocybium has the most vertebrae counts are useful in distinguishing species that
(62-64), Grammatorcynus the least (31), with the had previously been confused (Collette and Russo
species of Scomberomorus falling between (41- 1979); S. koreanus (46) from S. guttatus (usually
56). The same situation exists with precaudal 48-51) as shown by Devaraj (1976); S. brasiliensis
vertebrae (Acanthocybium 30-32, Scomberomo- (47-49) from S. maculatus (51-53) as shown by
rus 16-23, Grammatorcynus 12) and caudal ver- Collette et al. (1978); and S. munroi (50-52) from
tebrae (Acanthocybium 31-33, Scomberomorus S. niphonius (48-50) as shown by Collette and
20-36, Grammatorcynus 19). The presence of only Russo (1980). In general, low vertebral number is
31· vertebrae in Grammatorcynus is a primitive considered primitive in the genus, high vertebral
condition agreeing with Scomber and Rastrel- number advanced.
liger, the most primitive members of the Scom- Species with similar total numbers of vertebrae
brinae. The increased number of vertebrae in may differ in numbers of precaudal and caudal
Acanthocybium is clearly a specialization. vertebrae. Both S. cavalla and S. sinensis have

TABLE 6.-Number of precaudal vertebrae in Acanthocybium, Grammator-

cynus, and the species of Scomberomorus.

Species 12 I I 16 17 18 19 20 21 22 23 I I 30 31 32 N i1
S. brasiliensis 4 78 1 83 20.0
S. cavalla 1 28 29 17.0
S. commerson 41 69 110 19.6
S. concalor 1 14 5 20 19.2
S. guttatus 1 14 43 2 60 20.8
S. koreanus 24 24 20.0
S. lineolatus 2 13 1 16 18.9
S. maculatus 30 3 33 21.1
S. multiradiatus 9 16 25 20.6
S. munroi 2 10 12 21.8
S. niphonius 4 23 1 28 21.9
S. plurilineatus 1 12 13 19.9
S. queenslandicllS 1 13 14 19.9
S. rega/is 1 9 10 19.9
S. semifasciatus 1 21 22 19.0
S. sierra 3 45 3 51 20.0
S. sinensis 9 3 12 19.3
S. trilOr 2 24 26 18.9
Acanthocybium 2 4 2 8 31.0
GrammalOtcynus 14 14 12.0

TABLE 7.-Number of caudal vertebrae in Acanthocybium, Grammatorcynus, and the species of

Scomberomorus.

Species 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 N i1
S. brasiliensis 7 72 4 83 28.0
S. cava//a 1 27 1 29 25.0
S. commerson 11 33 49 7 9 109 24.7
S. concalor 2 13 5 20 28.2
S. guttatus 12 17 28 3 60 29.4
S. koreanus 22 2 24 26.1
S. lineo/atus 3 11 16 26.8
S. maculatus 15 18 33 30.5
S. multiradiatus 10 13 2 25 34.7
S. munroi 2 8 2 12 29.0
S. niphonius 23 5 28 27.2
S. plurilineatus 2 10 1 13 25.9
S. queenslandicus 10 4 14 28.3
S. regalis 10 10 28.0
S. semifasciatus 1 17 4 22 26.1
S. sierra 1 9 36 5 51 27.9
S. sinensis 2 9 11 21.8
S. tritor 23 3 26 27.1
Acanthocybium 4 2 7 32.1
Gramma/orcynus 14 14 19.0

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 FISHERY BULLETIN: VOL. 82, NO.4

TABLE 8. -Total number ofvertebrae in Acanthocybium, Grammatorcynus, and the species of Scomberomorus.
Species 31 /I 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 I I 62 63 64 N 51
S. brasiliensis 11 67 5 83 47.9
S. cava//a 2 28 1 31 42.0
S. commerson 7 16 34 39 15 111 44.4
S. concolor 1 11 8 20 47.4
S. guttatus 1 8 5 16 28 2 60 50.1
S. koreanus 23 2 25 46.1
S. linealatus 1 3 13 17 45.7
S. maculatus 13 19 2 34 51.7
S. multiradiatus 1 15 9 25 55.3
S. munroi 3 9 1 13 50.8
S. niphonius 3 20 6 29 49.1
S. plurilineatus 2 12 14 45.9
S. queenslandicus 12 3 15 48.2
S. rega/is 1 9 10 47.9
S. semifasciatus 1 23 3 27 45.1
S. sierra 1 8 38 4 51 47.9
S. sinensis 10 1 11 41.1
S. triWr 25 1 26 46.0
Acanthocybium 2 3 2 7 63.0
Grammatorcynus 16 16 31.0

41-43 vertebrae, but S. cavalla has 16-17 pre- Variable characters are found on the haemal
caudal and 24-26 caudal, while S. sinensis has 19 arches and haemapophyses. Laterally directed
or 20 precaudal and 21-22 caudal vertebrae (com- parapophyses, arising from the middle of the
pare Tables 6 and 7). centrum, appear on the fourth to sixth vertebrae,
where the intermuscular bones and pleural ribs
Vertebral Column are encountered (see section on Ribs and Inter-
muscular Bones). The parapophyses become
The neural arches and spines are stout and broader and longer posteriorly and gradually
compressed on the first to the fifth or sixth shift to the anteroventral portion ofthe centra. In
vertebrae in most species of Scomberomorus. lateral view, the first ventrally visible parapo-
Compressed neural spines extend to the seventh physes are found on the 7th-9th vertebra in
vertebra in S. commerson and Acanthocybium Scomberomorus, usually the 8th, on the 6th-7th
but only to the fourth vertebra in Grammator- in Grammatorcynus, and on the 14th-15th in
cynus. Posteriorly, toward the caudal peduncular Acanthocybium.
vertebrae and caudal complex, the neural spines Posteriorly, the distal ends of the paired para-
bend abruptly backward and cover most of the pophyses meet, forming the first closed haemal
neural groove; caudally they merge into the cau- arch. The first closed haemal arch is on the 8th
dal complex as in Thunnus (Kishinouye 1923; vertebra in Grammatorcynus (Fig. 37d), 10th-
Gibbs and Collette 1967) and the bonitos (Collette 16th in Scomberomorus (Fig. 37a, b), and 25th-
and Chao 1975). Neuropophyses are present on 28th in Acanthocybium (Fig. 37c). This location
all centra except the last one or two. The neural is correlated with the total number of vertebrae
prezygapophyses on the first vertebra are modi- (Table 8). Among the species of Scomberomorus,
fied to articulate with the exoccipital where the the first closed haemal arch is most anterior in S.
vertebral axis is firmly articulated with the cavalla (10th-11th vertebra, Fig. 37a) and S.
skull. They are stronger at the anterior portion of sinensis (12th), the two species with the few-
the vertebrae and are spurlike spines on the est vertebrae (40-43). The most posterior first
peduncular vertebrae and in the caudal complex. haemal arch is on the 15th-16th vertebra in S.
Neural postzygapophyses arise posterodorsally munroi and S. niphonius (Fig. 37b) and on the
from the centrum and overlap prezygapophyses 14th-15th in S. multiradiatus, species with many
posteriorly. The postzygapophyses progressively vertebrae (48-56). The other 13 species, including
merge into the neural spine in the peduncular S. guttatus and S. maculatus with high vertebral
region to disappear by the last 6-8 vertebrae. The counts (47-53), have the first haemal arch located
basic structure and elements of the neural arches at an intermediate position, 'on the 13th-14th
and neurapophyses are similar to those of other vertebra. The haemal spines become elongate
scombrids (Kishinouye 1923; Conrad 1938; Mago and point posteriorly until they abruptly become
Leccia 1958; Nakamura 1965; Gibbs and Collette more elongate on the first caudal vertebra. The
1967; Collette and Chao 1975; Potthoff 1975). paired pleural ribs (see section on Ribs and Inter-
596
COLLETTE and RUSSO: SPANISH MACKERELS

a b

c d
FIGURE 37. -Vertebra bearing first closed haemal arch in left lateral view (middle vertebra of each set of three). \ertebrae num-
bered from anterior. a. Scomberorrwrus cavalla, Chesapeake Bay, 672 mm FL, 1.5 x . b. Scomberorrwrus niphonius, Japan, 683
mm FL, 1.5 x. c. Acanthocybium solandri, Revillagigedos Is., 1,068 mm FL, Ix. d. Grommatorcynus bilineatus, Queensland, 521
mmFL, 1.5x.

muscular Bones} attach to the distal ends of the (Fig. 38). The haemal postzygapophyses fuse with
parapophyses and arches and extend posteriorly the haemal spine or disappear in the caudal
to the last precaudal vertebra. Symmetrically peduncle region.
with the neural arches and spines on the' caudal The haemal prezygapophyses arise from the
vertebrae, the haemal arches and spines bend anterior base of the haemal arches on the 8th-
posteriorly at the caudal peduncle and then 11th vertebra in Grammatorcynus, the lOth-22d
merge into the caudal complex. in Scomberomorus, and the 23d-25th in Acan-
Haemapophyses include pre- and postzygapo- thocybium. The most anteriorly located prezyga-
physes but their relative positions are different pophyses in Scomberomorus are in S. cavalla, on
from those of the neurapophyses, and they do not the 10th-12th vertebra. The most posteriorly lo-
overlap. The first haemal postzygapophyses arise cated are in S. sinensis (22d), S. queenslandicus
posteroventrally from the 6th-7th centrum in (18th-20th), S. multiradiatus (18th-19th), and S.
Grammatorcynus, the 6th-8th in Scomberomo- maculatus (17th-20th). The other 15 species have
rus, and the 9th-10th in Acanthocybium, and the first haemal prezygapophyses on the 13th-
they reach their maximum length at about the 19th vertebra. The data from Devaraj (1977)
junction of the precaudal and caudal vertebrae for the four Indian species fall in the interme-

597
 FISHERY BULLETIN: VOL. 82, NO.4

a b

c d
FIGURE 38. -Junction of precaudal and caudal vertebrae in left lateral view (middle vertebra of each set of three
is first caudal vertebra). Vertebrae numbered from anterior. a. Scomberomorus cavalla, Florida, 688 mm FL,
1 x. b. Scomberomorus munroi, Cairns, Queensland, 800 mm FL, 1 x. c. Acanthocybium solandri, Miami, Fla.,
1,242 mm FL, 1x . d. Grammatorcynus bilineatus, Timor Sea, 453 mm FL, 1.5 x .

diate group, but we found more variation, usu- the 56th-57th in Acanthocybium. Devaraj (1977)
ally a range of three or four vertebrae, than found only one inferior foramen in Acanthocyb-
Devaraj did. As do their counterpart neural pre- ium, on the 49th vertebra, but we found them
zygapophyses, the haemal prezygapophyses per- on 7-9 vertebrae in 10 specimens from the At-
sist symmetrically into the caudal complex. lantic, Indian, and Pacific Oceans. In Scom-
Struts between the haemal arch and the cen- beromorus, inferior foramina begin furthest an-
trum form the inferior foramina. Foramina are teriorly and extend furthest posteriorly in S.
present from the 18th-19th to the 27th-28th ver- multiradiatus, from the 21st-23d to the 51st-52d
tebra in Grammatorcynus, the 21st-33d to the vertebra. They begin furthest posteriorly in S.
35th-52d in Scomberomorus, and the 49th-51st to maculatus (29th-33d), S. niphonius (27th-33d),

598
COLLETI'E and RUSSO: SPANISH MACKERELS

and S. concolor (26th-38th). They extend posteri- vertebra: S. guttatus, S. munroi, and S. niphon-
orly only to the 35th-36th vertebra in S. cavalla. ius. The condition in the remaining 12 species
either varies or is based on only a single speci-
Ribs and Intermuscular Bones men. The greatest number of intermuscular
bones are found in S. guttatus, 26-30 pairs.
Pleural ribs are present from the 2d or 3d Counts as high as 27 are found in S. koreanus, S.
vertebra posterior to the 12th-31st vertebra, de- maculatus, and S. multiradiatus. Except for S.
pending on the species. Intermuscular bones start koreanus, the other three species with high num-
on the back of the skull or the first vertebra and bers of intermuscular bones also have high verte-
extend to the 10th-30th vertebra. bral counts. The fewest intermuscular bones in
Correlated with its high number of vertebrae, Scomberomorus are found in S. cavalla and S.
Acanthocybium has the most pleural ribs (30 sinensis, 20 pairs, and S.lineolatus, S. niphonius,
pairs) of the three genera. Similarly, Gramma- and S. semifasciatus, 20-23 pairs each. Gramma-
torcynus has the fewest pleural ribs (10 pairs) in torcynus has relatively few intermuscular bones
agreement with its low number of vertebrae. (19-21 pairs) and Acanthocybium, unexpectedly,
Species of Scomberomorus are intermediate in has the fewest (10 pairs) among the genera under
number of vertebrae and pleural ribs (15-21 discussion. This seems odd in view of its high
pairs). The first pleural rib articulates with the number of vertebrae and pleural ribs. Intermus-
centrum of the third vertebra in Grammator- cular bones extend back furthest in the four
cynus and most specimens of Scomberomorus. species with the highest number, S. guttatus (to
The first rib articulates with the centrum of the the 25th-29th), S. koreanus (24th-29th), S. macu-
second vertebra in Acanthocybium, as noted by latus (22d-27th), and S. multiradiatus (26th).
Devaraj (1977:44), and in one or two specimens of They extend back the shortest distance in S.
at least three species of Scomberomorus: com- cavalla (to the 19th), the species with the fewest
merson (1 of 5), maculatus (2 of 10), and sinensis intermuscular bones. Correlated with their low
(our only specimen). Pleural ribs extend posteri- number in Grammatorcynus and Acanthocybium,
orly usually to about the last precaudal vertebra. the bones extend back to the 19th-21st and to the
They extend to the 31st vertebra in Acantho- 10th vertebra respectively.
cybium, to the 17th-23d in Scomberomorus, and
only to the 12th in Grammatorcynus. Of the Caudal Complex
species of Scomberomorus, the most ribs are
found in S. munroi (20-21 pairs), S. guttatus (20), The supporting bones ofthe caudal fin (Fig. 39)
S. brasiliensis (18-20), and S. maculatus (18-20). consist of four or five preural centra in Scomber-
The fewest are in S. cavalla (15 pairs), S. semi- omorus. Having four preural centra supporting
fasciatus (15-17), and S. concolor (16-18). Ribs the caudal fin is not a diagnostic character of the
extend back furthest in the same four species family as stated by Potthoff (1975). Only three
with the most pleural ribs, S. munroi (to the 22d- preural centra support the caudal fin in Gram-
23d), S. guttatus (20th-22d), S. brasiliensis (20th- matorcynus, Scomber, and Rastrelliger. Five cen-
22d), and S. maculatus (17th-22d). They extend tra support the caudal fin in Acanthocybium. In
back the shortest distance in S. cavalla (to the Scomberomorus and Acanthocybium, preural
17th) and S. semifasciatus (17th-19th). As Dev- centra 4 and 3 bear stout haemal and neural
araj (1977:44) noted, the anterior ribs in Acan- spines. Preural centrum 2 has an epural. Preural
thocybium are very broad compared with those in centra 2 and 3 each have autogenous haemal
Scomberomorus. . spines. The urostyle represents a fusion of preural
Intermuscular bones start on the first vertebra centrum 1 and the ural centrum (Potthoff 1975).
in Acanthocybium, Grammatorcynus, and some The urostyle is fused with the triangular hypural
species of Scomberomorus. In some specimens of plate posteriorly and articulates with the uro-
at least 13 species of Scomberomorus, the first neural dorsally. Dorsally, the urostyle bears an
intermuscular bone is attached to the exoccipital autogenous epural and ventrally, the autogenous
on the skull. This appears to be the usual condi- parhypural. Preural centra 2-4 are compressed in
tion in three species, S. concolor, S. koreanus Scomberomorus and Acanthocybium but not so
(also noted by Devaraj 1977), and S. sierra. At much as in the bonitos and tunas (Collette and
least three other species usually appear to have Chao 1975; Gibbs and Collette 1967). Preural
the first intermuscular bone attached to the first centrum 4 is not at all shortened in Grammator-

599
 FISHERY BULLETIN: VOL. 82, NO.4

///)!:lk--- HY PUR A L 5

HYPURALS 3 + 4

PARHYPURAL

PU4 EPURALS 1 and 2

URONEURAL

b PARHYPURAPOPHYSIS

NEURAL SPINES

UROSTYLE

FIGURE 39.~Caudal complex in left lat- - - HYPURAL

eral view. a. Scomberomorus semifas-
ciatus, New Guinea, 510 mm FL, 3 x. b.
NOTCH
Acanthocybium solandri, Revillagigedos
Is., 1,068 mm FL, 2 x. c. Grammator-
cynus bilineatus, New Guinea, 382 mm HAEMAL SPINES
FL,4x. c
cynus and preural centrum 3 is only slightly the ventral part (1 and 2). The notch is absent in
shortened (Fig. 39c). In Scomberomorus and the more advanced bonitos and tunas (Collette
Acanthocybium, the posterior three neural and and Chao 1975). In two larger specimens of
three haemal spines bend abruptly away from the Grammatorcynus (453 and 521 mm FL), the fifth
vertebral axis and parallel the dorsal and ventral hypural is partially fused to the dorsal hypural
edges of the hypural plate. Only one neural and plate instead of being separate as in three small-
one haemal spine do so in Grammatorcynus. er specimens (382-410 mm FL, Fig. 39c). One of
The triangular hypural plate is composed of the diagnostic characters of the Scombridae is
4-5 fused hypural bones (Potthoff 1975). The that the bases of the caudal rays completely cover
dorsalmost (hypural 5) is not fused with the the hypural plate instead of only extending part
dorsal part of the hypural plate (hypurals 3 and way over the plate as is true of the Gempylidae
4). The primitive hypural notch is present on the and Trichiuridae with caudal fins.
middle of the posterior margin of the hypural The parhypural is separate from the ventral
plate (Fig. 39). This notch is a remnant of the hypural plate in Scomberomorus and Gramma-
fusion ofthe dorsal part of the hypural plate with torcynus but is fused with it in Acanthocybium
600
COLLETIE and RUSSO: SPANISH MACKERELS

(Fig. 39b). This fusion was also noted by Conrad been provided by several authors: S. sinensis and
(1938), Fierstine and Walters (1968), and Devaraj S. niphonius (Kishinouye 1923:pI. 23, fig. 40, pI.
(1977). The parhypural appears to be partially 24, fig. 41); S. caualla, S. maculatus, and S.
fused with the hypural plate in Scomberomorus regalis (Mago Leccia 1958:pI. 15, figs. 1-3); S.
niphonius (see Kishinouye 1923:figure 41) and S. tritor (Monod 1968:fig. 736); S. koreanus, S. gut-
plurilineatus. The two haemal arches preceding tatus, S. lineolatus, and S. commerson (Devaraj
the parhypural are autogenous in the three gen- 1977:fig. 15); S. semifasciatus (Collette and Russo
era although Devaraj (1977) stated that the two 1979:fig. 4B); and Acanthocybium (Kishinouye
haemal arches were fused with their centra in 1923:pI. 23, fig. 39; Conrad 1938:fig. 8; and Monod
Acanthocybium. 1968:fig. 737). There are problems with nomen-
The parhypural has a strongly hooked process, clature and labelling of various elements in these
the parhypurapophysis (or hypurapophysis), at papers, as discussed by Potthoff (1975).
its proximal end. The parhypurapophysis slopes
upwards in a similar manner in Scomberomorus DORSAL AND ANAL FINS
and Grammatorcynus but has a right angle and
then a level projection in Acanthocybium. Dev- Scombrids have two dorsal fins. The first dorsal
araj (1977:44) claimed that "the hypurapophysis fin is composed of stiff spines and is separated
is reduced to a small process" in Acanthocybium, from the second dorsal by a short distance, except
and his figure seems to show that. This conclu- in Rastrelliger, Scomber, and Auxis which have
sion must be based on a damaged specimen be- a greater distance between the fins. The second
cause the parhypurapophysis is well developed in dorsal fin is composed of soft rays and is followed
our specimens (Fig. 39b). The concentrations of by a series of free finlets, 6-11 in Scomberomorus.
tendons and muscular bands between the parhy- The anal fin is located approximately opposite
purapophysis and caudal rays in scombroids were the dorsal fin and is composed largely of soft rays
described by Fierstine and Walters (1968), but no followed by a series of anal finlets similar to the
specific study of this aspect was made during our dorsal finlets, 5-12 in Scomberomorus. Some
work. scombrids have a free or partially free spine
There are two epurals as in other scombrids preceding the anal fin, but in Scomberomorus it
(Potthoff 1975). In shape and size, the anterior is difficult to tell if the anterior elements are
epural (epural 1) resembles the neural spine of spiny or soft rays; therefore, all are included
adjacent preural centrum 3. The posterior epural as "anal rays". Numbers of fin rays are useful
(epural 2) is a free splint located between the characters in distinguishing groups of species in
anterior epural and the uroneural and fifth hy- Scomberomorus.
pural which are joined together. The range in number of spines in the first
Illustrations of the caudal complex of Acantho- dorsal fin is 11-27 among Scomberomorus, Acan-
cybium and 11 species of Scomberomorus have thocybium, and Grammatorcynus (Table 9). The

TABLE 9.-Number of spines in the first dorsal fin of Acanthocybium, Grammatorcynus, and the
species of Scomberomorus.
Species 11 12 13 14 15 16 17 18 19 20 21 2223 24 25 26 27 N ii
S. brasiliensis 61 67 2 130 17.5
S. cavalla 3 6 42 2 3 1 58 14.9
S. commerson 3 50 107 5 165 16.7
S. conc%r 1 3 20 7 31 17.1
S.guttatus 6 30 59 7 102 16.7
S. koreanus 2 20 5 1 28 15.2
S. linealatus 2 10 15 1 28 16.5
S. maculatus 10 40 7 57 17.9
S. multlradiatus 8 14 4 27 17.8
S. munroi 6 8 15 20.7
S. niphonius 4 29 5 38 20.0
S. plurilineatus 12 17 2 31 15.7
S. queenslandicus 8 24 2 34 16.8
S. regalis 9 31 6 46 16.9
S. semifasciatus 1 11 17 29 14.6
S. sierra 1 3 36 39 79 17.4
S. sinensis 1 7 6 14 16.4
S. tritar 3 7 19 11 40 17.0
Acanthocybium 2 2 7 19 4 34 25.6
Grammatarcynus 2 42 45 12.0

601
 FISHERY BULLETIN: VOL. 82, NO.4

usual variation for a species is 3 or 4 spines. (Table 10). The usual variation for a species is 3
Acanthocybium has the most dorsal spines, 23-27. or 4 finlets. The highest counts are in S. commer-
Of the species of Scomberomorus, munroi (20-22) son and S. queenslandicus, both usually 9 or 10
and niphonius (19-21) both usually have 20 or 21 finlets. The lowest counts, 6 or 7, are in S.
dorsal spines, more than the other 16 species in sinensis and Grammatorcynus. The next fewest
the genus, and this is one reason that S. munroi dorsal finlets, 7 or 8, rarely 9, are found in S.
was not described until recently (Collette and multiradiatus. The low number of finlets and
Russo 1980). Four species of Scomberomorus high number of second dorsal fin rays in this
have low counts: cavalla 12-18, usually 15; semi- species may indicate an extension of the fin at the
fasciatus 13-15; koreanus 14-17, usually 15; and expense of the number of finlets.
plurilineatus 15-17. Grammatorcynus has even Anal fin rays (Table 11) show a similar trend to
fewer dorsal spines, 11-13, usually 12. Dorsal that of dorsal fin rays. The range in the three
spine counts are roughly correlated with verte- genera is 11-29; the usual variation for a species
bral number (Table 8). Acanthocybium has the is 4-6 rays. Four of the five species of Scomber-
highest counts of precaudal and total vertebrae; omorus with high counts of second dorsal fin rays
S. munroi and S. niphonius have the highest also have high counts of anal fin rays: multira-
precaudal vertebral counts, but not highest total diatus 25-29; korean us 20-24,. usually 22 or 23;
vertebral number; S. cavalla has the lowest pre- guttatus 19-23, usually 20-22; and semifasciatus
caudal and total counts in the genus; and Gram- 19-22, usually 21 or 22. No species of Scomber-
matorcynus has the fewest precaudal, caudal, and omorus stands out with very low counts but six
total vertebrae. species usually have 17-19 anal fin rays, lower
The range in number of second dorsal fin rays than the other species of the genus: brasiliensis,
is 10-25 in the three genera (Table 10). The usual cavalla, commerson, munroi, niphonius, and si-
variation for a species is 4 or 5 rays. Five species nensis. Acanthocybium (11-14) and Grammator-
of Scomberomorus have high counts: multiradia- cynus (11-13, usually 12) again have the fewest
tus 21-25, usually 23 or 24; korean us 20-24, usu- rays in this fin.
ally 22 or 23; guttatus 18-24, usually 20-22; semi- Anal finlets range 5-12 in the three genera
fasciatus 19-22, usually 20; and plurilineatus (Table 11) with the usual variation for a species
19-21, usually 20. The lowest counts in Scomber- being 4 finlets. The most finlets are found in S.
omorus are in sinensis, 15-17 rays. Acanthocyb- queenslandicus (9-11, usually 10) followed by
ium (12-16) and Grammatorcynus (10-12, usually commerson and lineolatus usually having 9 or 10
11) have even fewer second dorsal fin rays. Verte- finlets, similar to the situation with dorsal fin-
bral counts (caudal and total) are highest in S. lets. The lowest counts, usually 6 finlets, are
multiradiatus and lowest in S. sinensis. Gram- found in S. multiradiatus, S. sinensis, and Gram-
matorcynus has the fewest vertebrae. matorcynus, just as with dorsal finlets. Again the
Dorsal finlets number 6-11 in the three genera anal fin of S. multiradiatus appears to have

TABLE 10. - Number ofsecond dorsal fin rays and dorsal finlets inAcanthocybium, Grammatorcynus, and the species of Scomberomorus.

Dorsal rays Finlels

Species 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 N x 6 7 8 9 10 11 N x
S. brasiliensis 1 11 52 48 13 125 17.5 29 89 12 130 8.9
S. cavalla 4 12 25 19 60 17.0 1 15 35 6 57 8.8
S. commerson 1 21 61 67 11 1 162 17.4 12 76 72 4 164 9.4
S. cone%r 1 2 10 16 2 31 18.5 7 21 1 30 7.7
S. guttatus 3 5 24 40 24 7 104 21.0 6 52 43 4 105 8.4
S. koreanUs 1 4 8 13 27 22.3 7 17 4 28 7.9
S. lineaiatus 4 9 11 2 0 1 1 29 17.6 2 7 18 2 29 8.7
S. maculatus 5 19 25 7 56 18.6 2 29 25 56 8.4
S. multiradiatus 3 4 10 8 2 27 23.1 15 10 2 27 7.5
S. munroi 3 8 3 15 18.1 11 4 15 9.3
S. niphonius 1 12 12 12 1 38 17.0 9 22 7 38 7.9
S. plurilineatus 6 22 8 36 20.1 12 20 3 35 8.7
S. queens/andicus 10 11 10 31 18.0 12 17 2 31 9.7
S. regaiis 2 12 28 5 47 17.8 2 39 6 47 8.1
S. semifasciatus 6 18 5 3 32 20.2 9 19 3 31 8.8
S. sierra 2 29 41 6 78 17.7 2 43 33 1 79 8.4
S. sinensis 4 9 1 14 15.8 8 6 14 6.4
S. tritar 6 25 9 41 17.1 1 33 7 41 8.1
Acanthacybium 1 3 21 7 2 35 13.3 6 16 11 3 36 8.3
Grammatorcynus 5 37 1 43 10.9 40 5 45 6.1

602
COLLETIE and RUSSO: SPANISH MACKERELS

TABLE n.-Number of anal fin rays and anal finlets in Acanthocybium, Grammatorcynus, and the species of Scomberomorus.
Anal rays Finlets
Species 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 N x 5 6 7 8 9 10 11 12 N x
S. brasiliensis 6 35 30 44 9 124 18.1 1 41 84 2 128 8.7
S. cavalla 1 9 27 21 1 59 18.2 4 33 19 4 60 8.4
S. commerson 8 29 72 43 12 1 165 18.2 1 17 84 57 4 2 165 9.3
S. concolor 1 17 9 2 1 30 20.5 1 15 15 31 7.5
S. guttatus 2 23 49 26 3 103 21.0 20 63 17 4 104 8.0
S. koreanus 1 3 13 11 29 22.3 17 11 1 29 7.4
S. Iineolatus 2 0 7 17 2 1 29 19.7 2 2 14 11 29 9.2
S. maculatus 4 11 24 18 57 19.0 1 33 21 1 56 8.4
S. multiradiatus 3 7 8 8 1 27 26.9 18 8 1 27 6.4
S. munroi 8 5 2 15 17.6 2
10 3 15 9.1
S. niphonius 1 12 20 3 1 37 17.8 7 20
10 38 8.0
S. plurilineatus 4 16 13 34 20.3 1 15
17 2 35 8.6
S. queenslandicus 2 1 6 14 8 31 18.8 8 21 2 31 9.8
S. regalis 1 4 15 24 1 46 18.4 3 34 8 1 46 8.2
S. semifasciatus 1 4 17 11 33 21.2 1 11 19 2 33 8.7
S. sierra 3 6 19 33 17 1 79 18.7 6 48 23 1 78 8.2
S. sinensis 1 3 9 1 14 17.7 2 9 3 14 6.1
S. tritor 5 20 14 2 41 18.3 6 27 8 41 8.0
Acanthocybium 1 11 14 8 34 12.9 13 12 7 2 34 7.9
Grammatorcynus 5 22 17 44 12.3 1 39 4 44 6.1

extended posh:dorly to reduce the number of anal Posttemporal

finlets.
The posttemporal (Fig. 40) is a flat elliptical
bone with two sturdy anterior processes that
PECTORAL GIRDLE attach the pectoral girdle to the neurocranium.
The median (dorsal) process is concave at its
The pectoral girdle consists of the girdle itself dorsal surface and articulates with the dorsal
(cleithrum, coracoid, and scapula), the radials to surface of the epiotic. The lateral (ventral) pro-
which the pectoral fin rays attach, and a chain of cess is shorter, round in cross section, and its
bones that connect the girdle to the rear of the hollow anterior end articulates with the dorsal
skull (posttemporal, supracleithrum, supratem- protuberance of the intercalar. There is a thin
poral, and two postcleithra). shelf between the median and lateral processes in

c
FIGURE 40.-Left posttemporals in lateral view. a. Scomberomorus cavalla, Chesapeake Bay, 672 mm FL, 1.5x. b. Scomber-
omorus plurilineatus, South Africa, 910 mm FL, Ix. c. Acanthocybium solandri, Revillagigedos Is., 1,068 mm FL, Ix. d.
Grammatorcynus bilineatus, Queensland, 521 mm FL, 1.5x.

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Scomberomorus (Fig. 40a, b) but this shelf is phonius, and semifasciatus. Grammatorcynus
absent in Acanthocybium (Fig. 40c) and Gram- also has a short median process (66-71%). Mea-
matorcynus (Fig. 40d). A variably sized notch is sured this way, the longest lateral processes are in
present at the middle of the posterior edge of the koreanus (53-55%), sierra (52%), plurilineatus
flat body of the bone. Grammatorcynus has a and regalis (50-51%), guttatus (49-55%), and
prominent, anteriorly directed spine on the ven- Grammatorcynus (48-52%). The shortest lateral
tral margin of the median process about one-third processes are in munroi (37-40%), tritor (41-42%),
of the distance from the body of the bone to the and cavalla and niphonius (41-44%). Acantho-
anterior tip of the process. In Acanthocybium, cybium also has a relatively short lateral process
. there is a separate process extending anteriorly (42-51%) .
from the ventral wall of the median process. This Still another way of comparing relative lengths
auxiliary process (Kishinouye 1923) is as long or of the processes among species is to divide the
almost as long as the median process itself. It length of the lateral process by the length of the
ends in a series of several pointed processes. median process, both measured on the inner
(Both Conrad 1938 and Devaraj 1977 referred to surface of the posttemporal. By this technique,
the auxiliary process as the median process.) relatively greater proportional measurements of
The lengths of the median and lateral processes the lateral processes are found in S. koreanus
vary among· the species under discussion. To (75-77% of median process), S. semifasciatus
compare the species quantitatively, we made two (76%), Grammatorcynus (71-74%), S. multiradia-
sets of measurements and divided them by the tus (70-73%), and S. guttatus (69-71%). Relatively
total length of the posttemporal, from the ante- shorter proportional measurements of these lat-
rior tip of the median process to the posterior eral processes (55-63%) are found in Acantho-
margin of the bone. We measured to the tips of cybium and five species of Scomberomorus: ca-
the median and lateral processes from the most valla, munroi, niphonius, queenslandicus, and
posterior point on the shelf between the two tritor.
processes. Largely because of the lack of a shelf Another difference lies in the presence and, if
between the processes in Acanthocybium and present, in the shape of a spine or process at the
Grammatorcynus, both processes appear to com- base of the lateral process on the inner surface of
prise a larger proportion of total posttemporal the posttemporal. It appears to be absent in seven
length than they do in the species of Scombero- species of Scomberomorus: cavalla, guttatus,
morus, 53-65% vs. 36-51% for the median process maculatus, queenslandicus, regalis, semifascia-
and 27-40% vs. 15-36% for the lateral process. tus, and tritor. It is small and inconspicuous in
The median process is longer in Acanthocybium six species: brasiliensis, concolor, korean us , line-
than in Grammatorcynus, 56-65% vs. 53-60%, olatus, multiradiatus, and sierra. It is broader,
but the lateral process is slightly longer in Gram- usually shaped more like a shelf with a point in
matorcynus than in Acanthocybium, 35-40% the remaining five species of the genus: commer-
vs. 27-37%. Among the species of Scomberomo- son, munroi, niphonius, plurilineatus, and some-
rus, the longest median processes (48-51% total times in sinensis. The process has the form of
length) are found in S. commerson, plurilineatus, a wide flap in Grammatorcynus and of a long
and sierra; the shortest (36-40%) in cavalla, blunt process in Acanthocybium. Devaraj's (1977)
semifasciatus, and sinensis. The longest lateral data for Indian species correspond well with
processes are in koreanus (36%) and plurilinea- ours.
tus (30-31%); the shortest (15-19%) in cavalla,
munroi, niphonius, queenslandicus, and tritor. Supracleithrum
To eliminate the confounding factor ofthe shelf
between the median and lateral processes, mea- The supracleithrum (Fig. 41) is an ovate bone,
surements also were made on the inner surface of overlapped dorsolaterally by the posttemporal
the bone, from the point where the two processes and overlapping the anterior part of the dorsal
diverge to the tips of the processes: Measured this winglike extension of the cleithrum. The anterior
way, the longest median processes (74-79% of border of the bone on the mesial side is thickened
total length of the posttemporal) are in Acantho- into a ridge. Dorsally there is a small handle-
cybium and six species of Scomberomorus. The shaped process which curves into the posterior
shortest median processes (63-68%) are in three margin to end in a notch at the posterodorsal
species of Scomberomorus: multiradiatus, ni- aspect. A branch of the lateralis system extends
604
COLLETTE and RUSSO: SPANISH MACKERELS

from the posterior notch of the posttemporal onto factor because the smallest species of 8comber-
the supracleithrum. This short canal lies ventral omorus (8. multiradiatus) and small specimens
to the dQrsal process of the supracleithrum and of large species tend to have narrower supra-
extends to the posterior edge ofthe bone. cleithra than large species and large specimens.
The maximum width of the supracleithrum For example, the percentages for a series of five
varies from 42 to 75% of the total length of the 8.commerson are as follows: 354-364 mm FL,
bone in the three genera. The supracleithrum is 38-43%; 493 mm, 42%; 1,052 mm, 39-44%; 1,155
widest in Grammatorcynus (72-75% of length), mm,47%.
8comberomorus niphonius (55-62%), and 8. line- The dorsal process is prominent in Acanthocyb-
olatus (53-57%). It is narrowest in 8. multiradia- ium (Fig. 41c), Grammatorcynus (Fig. 41d), 8.
tus (43-53%), sinensis (45-46%), semifasciatus cavalla, commerson, and lineolatus (Fig. 41a). It
(46-51%), and sierra (45-49%). Specimen size is a is small but distinct in 8. multiradiatus (Fig.
41b). In most of the other species of 8comberomo-
rus,' it tends to be less sharply set off from the
main body of the supracleithrum.

Supratemporal

The supratemporal (Fig. 42) is a thin flat

triangular bone lying just underneath the skin
where its lateral process articulates with a dorsal
articular surface on the pterotic. Mago Leccia
(1958:324) failed to find the supratemporal in his
specimens. The anterior margin is concave and
the convex posterior margin slightly overlaps the
dorsal arm of the posttemporal. The supratem-
poral is deeper (from the tip of the median ante-
rior arm to the base) than wide (tip of lateral
a b anterior arm to tip of posterior arm), width
49-84% of depth in 8comberomorus and Acan-
thocybium. However, the supratemporal is wider
than deep in Grammatorcynus (Fig. 42d), width
101-113% oflength. Acanthocybium (Fig. 42c) has
a wider supratemporal (84-93% of depth) than do
the species of 8comberomorus (49-79%). The wid-
est supratemporals in 8comberomorus are in
niphonius (73%), guttatus (67-79%), sierra (69-
74%), and semifasciatus (63-72%). The narrowest
supratemporals are in multiradiatus (49-50%),
koreanus (54%), brasiliensis (53-59%), queens-
landicus (55-60%), and sinensis (54-62%).
The supratemporal bears a prominent lateral
line canal that extends out almost to the tips of
all three arms. Devaraj (1977:45) did not specifi-
cally mention the presence ofthis canal. In 8com-
beromorus, the canal along the anterior margin
of the bone is the longest and best developed, and
the canal along the lateral side the next longest.
c d In most species of the genus, the first canal has
three or four posteriorly directed branches. Speci-
FIGURE 41.-Left supracleithra in lateral view. a. Scomber-
mens of 8. niphonius and 8. semifasciatus (Fig.
omorus lineolatus, Cochin, India, 786 mm FL, 1.5 x. b. Scom-
beromorus multiradiatus, New Guinea, 294 mm FL, 3 x. c.
42a) had five or six branches. Two specimens of
Acanthocybium solandri, Caribbean Sea, 1,240 mm FL, 1 x. d. 8. multiradiatus had only a single posterior
Grammatorcynus bilineatus, Marshall Is., 424 mm FL, 2 x . branch. A specimen of 8. munroi had no posterior
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a b

FIGURE 42.-Left supratemporals in lateral

view. a. Scomberomorus semifasciatus, New
Guinea?, 740 mm FL, 1.5x. b. Scomberomo-
rus multiradiatus, New Guinea, 224 mm FL,
5 x. c. Acanthocybium solandri, Caribbean
Sea, 1,240 mm FL, 1 x. d. Grammatorcynus
bilineatus, New Guinea, 382 mm FL, 3 x . c d
branches, but it did have four pores along the becomes narrower posteriorly in most species of
main part of the canal. Acanthocybium has a Scomberomorus and in Grammatorcynus. The
single short posterior branch that opens into a posterior plate is longer and of uniform width in
very large pore (Fig. 42c). Grammatorcynus (Fig. Acanthocybium (Fig. 43c).
42d) lacks a distinct posterior branch but has a The lower part of the cleithrum is large and
relatively longer canal on the lateral side of the folded back upon itself as two walls: one lateral
bone. and the other mesial, which meet at their ante-
rior margins and run parallel to each other. The
Cleithrum mesial wall of the cleithrum forms a large tri-
angular slit with the coracoid. As Devaraj (1977:
The main body of the cleithrum is crescent- 46) pointed out, this slit is hidden in lateral view
shaped with an anterodorsal spine and a poste- in the species of Scomberomorus by the great
riorly projecting plate at the upper end, as in width of the lateral wall of the cleithrum. This
other scombrids (Fig. 43). The angle between the portion of the cleithrum is narrower in Acantho-
spine and the plate is wider in Acanthocybium cybium and Grammatorcynus, and consequently
(Fig. 43c) than in Grammatorcynus (Fig. 43d) the upper part of the slit is visible in lateral view.
and the species of Scomberomorus. The bonitos
have wider angles (Collette and Chao 1975:fig. Coracoid
61), except for Gymnosarda. The spine extends
about as far dorsally as the plate does in Acan- The coracoid is elongate and more or less tri-
thocybium and all the species of Scomberomorus, angular in shape (Fig. 43). It connects with the
except S. sinensis in which the spine extends well scapula along its dorsal edge and with the mesial
past the dorsal margin of the plate. In Gramma- shelf of the cleithrum anterodorsally and antero-
torcynus, the spine does not extend all the way to ventrally. There is a prominent elongate slit
the margin of the plate (Fig. 43d). The plate between the cleithrum and the coracoid that is
606
COLLETI'E and RUSSO: SPANISH MACKERELS

CORACOID

POSTERIOR
PLA TE

SCAPULAR
FORAMEN
e

c
ANTERODORSAL
SPI NE

FIGURE 43.-Left pectoral girdles in lateral view. a. Scomberomorus semifasciatus, New

Guinea, 510 mm FL. b. Scomberomorus sinensis, Hong Kong, 677 mm FL. c. Acanthocybium
solandri, Revillagigedos Is., 1,086 mm FL. d. Grammatorcynus bilineatus, Marshall Is., 424
mm FL. e. Scomberomorus koreanus , Indonesia, 480 mm FL, inset of scapular and interradial
foramina.

visible laterally in Acanthocybium and Gramma- Scapula

torcynus but is concealed by the lateral shelf of
the cleithrum in Scomberomorus. The coracoid is The anterior margin of the scapula connects to
relatively narrower in Acanthocybium than in the mesial shelf of the cleithrum (Fig. 43). This
Grammatorcynus and the species of Scombero- attachment extends to the posterior projecting
morus. We did not find the coracoid to be signif- plate anterodorsally. The scapula is attached to
icantly narrower in S. commerson, as reported by the coracoid posteriorly and with the first two
Devaraj (1977:47). and part of the third upper radials posterodorsal-
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 FISHERY BULLETIN: VOL. 82, NO.4

ly. The posterodorsal margin of the scapula is bers of the family to the more advanced: Scom-
drawn out into a facet which accepts the most brini 18-21, Scomberomorini 19-26, Sardini 21-28,
anterior ray of the pectoral fin. The scapula is Thunnini (except for Thunnus) 22-29, Thunnus
pierced by a large, usually round foramen near 30-36.
the lateral margin with the inner shelf of the
cleithrum. A prominent suture leads from the Radials
scapular foramen to the ventral margin of the
scapula. The foramen is largest in Acanthocyb- The four radials differ in size and shape and are
ium (Fig. 43c), Scomberomorus brasiliensis, and attached directly to the thickened posterior edges
S. regalis. It is smallest in S. guttatus and S. of the scapula and coracoid (Fig. 43). The size of
niphonius. We did not find it very large in S. the radials increases posteroventrally. Small for-
korean us (Fig. 43e), as stated by Devaraj (1977: amina are located between the second and third,
47). It is intermediate in size in Grammatorcynus and the third and fourth radials counting poste-
(Fig. 43d) and the other species of Scomberomo- riorly. In Scomberomorus and Acanthocybium
rus (e.g., S. semifasciatus and S. sinensis, Fig. (Fig. 43a-c), the first two radials and the upper
43a, b). third of the third radial attach to the scapula; the
ventral third of the third plus the fourth radial
Pectoral Fin Rays attach to the coracoid. In Grammatorcynus the
first two radials attach to the scapula, the second
The first (uppermost and largest) pectoral fin two to the coracoid (Fig. 43d). A much larger
ray articulates directly with a posterior process of foramen is present between the largest (fourth)
the scapula. The other rays attach to the radials. radial and the coracoid. Posteriorly, this foramen
The number of pectoral rays ranges from 19 to 26 is framed by a posterior process of the upper part
in the three genera (Table 12). Most species of of the fourth radial meeting an anterior process
Scomberomorus usually have 22 or 23 rays. Five from the posterior margin of the coracoid. The
species average fewer, with a mode of 21 rays: process on the fourth radial is only slightly devel-
concolor, guttatus, maculatus, sierra, and tritor. oped in Grammatorcynus (Fig. 43d). The foramen
The two species in the genus with the most is considerably larger than the scapular foramen
pectoral rays are S. plurilineatus (21-26, x 23.1) in five species of Scomberomorus: guttatus, ko-
and S. semifasciatus (22-25, x 23.3). Acanthocyb- reanus (Fig. 43e), lineolatus, niphonius, and plu-
ium and Grammatorcynus have slightly higher rilineatus. It is slightly larger than the scapular
counts than do the species of Scomberomorus, 22- foramen in seven species: commerson, concolor,
26, mostly 24 or 25. maculatus, multiradiatus, munroi, queenslandi-
Within the Scombridae, the number of pectoral cus, and tritor. The two foramina are about equal
fin rays increases from the more primitive mem- in size in six species of Scomberomorus (brasil-
iensis, caualla, regalis, semifasciatus (Fig. 43a),
sierra, and sinensis (Fig. 43b» and Grammator-
cynus (Fig. 43d). The scapular foramen is much
TABLE 12.-Number of pectoral fin rays in Acanthocybium,
Grammatorcynus, and the species of Scomberomorus.
larger than the foramen following the fourth
radial in Acanthocybium (Fig. 43c).
Species 19 20 21 22 23 24 25 26 N x
S. brasiliensis 9 38 21 69 22.2
S. cavalla 6 27 18 51 22.2 Postcleithra
S. commerson 18 52 32 8 110 22.3
S. concolor 8 21 4 34 20.8
S. guttatus 20 57 11 1 89 20.9 The posterior projecting plate of the cleithrum
S. koreanus 1 4 13 7 3 28 22.3
S. lineolatus 4 2 6 10 3 25 22.2
has its posterior end attached to the first post-
S. maculatus 8 33 14 1 56 21.1 cleithrum which connects ventrally to the second
S. multiradiatus 1 10 10 6 27 21.8
S. munroi 4 4 1 9 21.7 postcleithrum. The lamellar first postcleithrum
S. niphonius 9 22 5 36 21.9 (Fig. 44) is kidney-shaped with a narrow upper
S. plurilineatus 1 9 14 5 3 33 23.1
S. queenslandicus 5 19 5 0 1 30 22.1 end, rounded lower margin, concave anterior bor-
S. regalis 1 17 21 4 1 44 21.7 der and convex posterior margin. In Grammator-
S. semifasciatus 4 15 13 33 23.3
S. sierra 16 38 17 1 1 73 21.1 cynus (Fig. 44d), the first postcleithrum is very
S. sinensis 2 8 3 13 22.1
S. tritor 3 22 13 38 21.3
wide and short with a notch in the dorsal margin
Acanthocybium 3 4 17 12 1 37 24.1 instead of a pointed end, width/maximum length
Grammatorcynus 1 8 13 17 3 42 24.3
= 55-62%. It is wider (47-48%) in Acanthocybium

608
COLLETIE and RUSSO: SPANISH MACKERELS

a b

FIGURE 44.-Left first postcleithra in lateral

view. a. Scomberomorus sinensis, Hong Kong,
677 mm FL, 1 x. b. Scomberomorus koreanus,
Indonesia, 480 mm FL, 2 x. c. Acanthocyb-
ium solandri, Revillagigedos Is., 1,068 mm FL,
IX. d. Grammatorcynus bilineatus, Queens-
land, 521 mm FL, 2 x .

c d

(Fig. 44c) than in the species of Scomberomorus width of the bone makes the second postcleithrum
(24-41%). Three species of Scomberomorus have appear much wider in Grammatorcynus, 37-42%
wide first postcleithra (37-41%): commerson, si- of total length compared with 16-27% in the other
nensis (Fig. 44a), and tritor. Three species have two genera. The widest second postcleithra in
narrow first postcleithra: koreanus (24-26%, Fig. Scomberomorus, 22-27% of total length, are in
44b), lineolatus (28-29%), and guttatus (28-31%). lineolatus, maculatus, plurilineatus, and queens-
The other 12 species have moderately wide first landicus (Fig. 45a). The narrowest ones are in
postcleithra, 30-39%. Devaraj (1977:48) reported guttatus and koreanus (15-20%, Fig. 45b), and
long, narrow first postcleithra in the same three cavalla, sierra, and sinensis (19-20%). Acantho-
species (plus S. maculatus and S. regalis from cybium (Fig. 45c) and the other nine species of
Mago Leccia's 1958 work) and wider ones in S. Scomberomorus are intermediate, 20-24%. The
commerson and S. cavalla. ascending process appears longer in 6 species of
The second postcleithrum (Fig. 45) is broad and Scomberomorus (brasiliensis, cavalla, macula-
lamellar at the upper part with a short pointed tus, queenslandicus, regalis, and tritor) than in
ascending process and a long styliform descend- Acanthocybium, Grammatorcynus, and the other
ing process. Grammatorcynus (Fig. 45d) differs 12 species of Scomberomorus.
strikingly from Acanthocybium and Scombero-
morus in having a sharp process extending anteri- PELVIC GIRDLE
orly from the broad lamellar portion of the bone.
Inclusion of this process in measurements of the The pelvic fin rays (I, 5) attach directly to the
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 FISHERY BULLETIN: VOL. 82. NO.4

paired basipterygia which make up the pelvic basipterygium is composed of three main parts
girdle. The bones are united along the midline (Fig. 46): a wide anterodorsal plate; a thin, flat
and are imbedded in the ventral abdominal wall anterior process (anterior xiphoid process of
free from contact with any other bones. Each de Sylva 1955, anteromesial process of Devaraj
1977); and a strong posterior process (posterior
xiphoid process of de Sylva 1955). There are three
wings to the anterodorsal plate (Kishinouye
1923): lateral (external), mesial (internal), and
v~) ventral (vertical). Anteriorly, the lateral wing
turns into the same vertical plane and merges
into the ventral wing. The mesial wing and the

r lateral wing meet in one plane posteriorly along

a ridge.
To compare the pelvic girdles, the lengths of all
three parts were measured from their bases to
their tips. The anterior process comprised 15-52%
of the length of the anterodorsal plate. The long-
a b est anterior processes were in Grammatorcynus
(46-51%, Fig. 46d), Acanthocybium (35-47%, Fig.
46c), and seven species of Scomberomorus: sierra
(38-52%), concolor (36-44%), regalis (31-44%,
~
\'9: Fig. 46a), semifasciatus (35-36%), sinensis (35%),
tritor (32-36%), and maculatus (28-36%). The
shortest anterior processes were in three species
of Scomberomorus: koreanus (15-30%), multi-
radiatus (21-26%), and lineolatus (23-33%, Fig.
46b), but there is a large range of variation
within species. The posterior process comprised
20-85% of the length of the anterodorsal plate.
c d The longest posterior processes were in four
American species of Scomberomorus: regalis (78-
FIGURE 45. - Left second postcleithra in lateral view. a. Scom- 90%), brasiliensis (81%), sierra (62-85%), and
beromorus queenslandicus, Great Barrier Reef, 641 mm FL, concolor (67-68%). The other two species that
1 x. b. Scomberomorus koreanus, Indonesia, 480 mm FL,
1.5 x. c. Acanthocybium solandri, Revillagigedos Is., 1,068
belong to this group have shorter posterior pro-
mm FL, 1 x . d. Grammatorcynus bilineatus, New Guinea, 382 cesses: maculatus (38-48%) and tritor (36-50%).
mmFL,2x. The shortest posterior processes were in five

c d
FIGURE 46.-Right basipterygia of the pelvic girdle in mesial view. a. Scomberomorus regalis, Miami, Fla., 469 mm FL, 1.5x. b.
Scomberomorus lineolatus, Palk Strait, India, 428 mm FL, 2 x . c. Acanthocybium solandri, Miami, Fla., 1,403 mm FL, 1 x. d. Gram-
matorcynus bilineatus, Queensland, 521 mm FL, 1.5 x .

610
COLLETTE and RUSSO: SPANISH MACKERELS

species of Scomberomorus: guttatus (20-41%), ko- Polipturus Rafinesque 1815:84 (replacement

reanus (25-29%), lineolatus (29-35%), multi- name for Scomberomorus Lacepede, therefore,
radiatus (27-33%), and munroi (28-44%), plus takes the same type-species, Scomberomorus
Grammatorcynus (29-33%) and Acanthocybium plumierii Lacepede 1801).
(30-39%). Cybium Cuvier 1829:199 (type-species: Scomber
Grammatorcynus and some individuals of commerson Lacepede 1800 by subsequent des-
Acanthocybium and at least seven species of ignation of Gill 1862:126).
Scomberomorus have longer anterior processes Apolectus Bennett 1831:146 (type-species: Apolec-
than posterior processes. The lengths of the ante- tus immunis Bennett 1831 by monotypy, =
rior process as a percentage of the posterior Scomberomorus tritor (Cuvier in Cuvier and
process are Grammatorcynus (154-158%), Acan- Valenciennes 1831) ).
thocybium (91-156%), and the seven species of Apodontis Bennett 1832:169 (replacement name
Scomberomorus: sinensis (121%), semifasciatus for Apolectus Bennett, preoccupied by Apolec-
(111-116%), munroi (84-120%),guttatus (80-116%), tus Cuvier in Cuvier and Valenciennes 1831,
plurilineatus (89-113%), koreanus (57-105%), and Pisces).
tritor (66-100%). The shortest anterior processes Chriomitra Lockington 1879a:133 (type-species:
were in brasiliensis (42%), concolor (52-65%), Chriomitra concolor Lockington 1879a by
and sierra (56-62%). monotypy).
Devaraj (1977:48) alluded to differences in the Sierra Fowler 1905:766 (type-species: Cybium
relative depth of the anterior end of the antero- cavalla Cuvier 1829 by original designation
dorsal plate, but we have found this very difficult and monotypy).
to assess owing to different sizes and conditions of Sawara Jordan and Hubbs 1925:214 (type-species:
our material. Devaraj appears to be correct in Cybium niphonium Cuvier in Cuvier and
stating that the anterior end is particularly nar- Valenciennes 1831 by original designation and
row in S. lineolatus. The broadest anterior end is monotypy).
certainly in Grammatorcynus (Fig. 43d), which Pseudosawara Munro 1943:68 (type-species: Cyb-
Devaraj did not study. ium kuhlii Valenciennes 1831 by original des-
As Devaraj (1977:48) pointed out, a notch is ignation, = Scomberomorus guttatus (Bloch
present on the ventral wing of the anterolateral and Schneider 1801».
plate before it joins the other wings in Acantho- Indocybium Munro 1943:68-69 (type-species:
cybium (Fig. 46c) but is absent in Scomberomo- Cybium semifasciatum Macleay 1884a by orig-
rus (and also in Grammatorcynus). inal designation).
Except for Grammatorcynus, no differences
were found among the three genera in the fleshy Diagnosis. -Scomberomorus differs from all
bifid interpelvic process that is ventral to the other scombrids in possessing a spatulate vomer
paired posterior processes of the basipterygia. that projects anteriorly well beyond the anterior
Grammatorcynus differs from Scomberomorus margin of the neurocranium.
and Acanthocybium in having a single interpel- Scomberomorus differs from both Acanthocyb-
vic process. Auxis and Gymnosarda also have a ium and Grammatorcynus in a series of 12 osteo-
single interpelvic process, the former very large, logical characters: 1) posterior horizontal edge
the latter of moderate size. However, there is a of metapterygoid longer than anterior oblique
posterior process from each basipterygium re- edge (anterior oblique edge longer in Gramma-
gardless of whether the fleshy interpelvic process torcynus and Acanthocybium); 2) dorsal arm of
is single or bifid. ectopterygoid shorter than ventral arm (dorsal
arm longer or equal); 3) lateral wall of cleith-
rum wide, space between cleithrum and coracoid
SPECIES ACCOUNTS not visible in lateral view (narrow, space visible
in lateral view); 4) epiotic crests originate on
Scomberomorus Lacepede anterior part of frontal bones (originate behind
midfrontal region); 5) many (more than 11) ver-
Scomberomorus Lacepede 1801:292 (type-species: tebrae with inferior foramina (few, less than
Scomberomorus plumierii Lacepede 1801 by 11); 6) first basibranchial short (long); 7) strut
monotypy, = Scomberomorus regalis (Bloch on fourth pharyngobranchial elongate (not elon-
1793) ). gate); 8) symplectic short, not in contact with

611
 FISHERY BULLETIN: VOL. 82, NO.4

metapterygoid (long, in contact); 9) ventral lateral and medial arms of fourth epibranchial
hypohyal at least three times larger than dorsal less acute (more acute); 8) anterior process of
hypohyal (less than three times larger); 10) fifth second epibranchial not elongate (elongate); 9)
branchiostegal rayon suture between epihyal four or five vertebrae supporting caudal fin rays
and ceratohyal (on epihyal); 11) no shelf present (three); 10) no anterior process on second post-
between dorsal and ventral arms of posttemporal cleithrum (prominent spinelike process present);
(shelf present); and 12) epihyal much longer 11) anterior end of first postcleithrum pointed
than deep, depth 58-62% oflength (depth 66-98% (notched); 12) .base of third pectoral radial on
of length). suture between coracoid and scapula (completely
In three additional characters, Scomberomorus on coracoid); 13) jaw teeth compressed and tri-
differs from Acanthocybium and Grammator- angular (conical); 14) ventral surface of para-
cynus but is closer to the former than the latter: sphenoid convex (concave); 15) upper margin of
ventral branch of palatine equal to or longer than dentary longer than lower margin (lower longer);
(87-107%) dorsal branch (slightly shorter, 112-' and 16) posterior edge of ectopterygoid short,
121%, in Acanthocybium; much shorter, 120- 41-63% of ventral distance (long, 64-68%).
123%, in Grammatorcynus); supratemporal much
deeper than wide, 49-79% (deeper, 84-93%; wider
than deep, 101-113%); and first postcleithrum Scomberomorus brasiliensis Collette,
very narrow, 24-41% of length (narrow, 47-48%; Russo, and Zavalla-Camin
wide, 55-62%). Scomberomorus has a deep uro- Serra Spanish Mackerel
hyal; it is moderately deep in Grammatorcynus Figure 47
and shallow in Acanthocybium. Scomberomorus
has a moderate to high number of vertebrae (40- Scomberomorus maculatus. Not of Mitchill
56) compared with other members of the family, 1815. Ribeiro 1915:134-135 (Brazil). Lowe
more than Grammatorcynus (31), but less than 1962:679-686 (British Guiana continental
Acanthocybium (62-64). shelf). Cervigon 1966:720-721 (description,
Scomberomorus and Acanthocybium agree fishery; Venezuela), fig. 303. Bastos 1966:
with each other but differ from Grammatorcynus 113-117 (counts and measurements). Nomura
in a series of16 osteological characters: 1) supra- 1967:29-39 (biology; Ceara,' Brazil). Mota
cleithrum narrow, 42-62% of length (wide, 72- Alves and Tome 1968a:25-30 (sexual develop-
75% in Grammatorcynus); 2) pores present on ment). Mota Alves and Tome 1968b:139-140
dorsal arm of supratemporal (absent); 3) nasals (sperm). Fonteles Filho 1968:133-137 (fishery;
do not protrude far beyond ethmoid region (pro- Ceara, Brazil). Nomura and Costa 1968:95-99
trude far beyond); 4) posterior ~nd of dorsal (length-weight relationship). Costa and Paiva
margin of urohyal forked (tripartite); 5) glosso- 1969:89-95 (maximum size 125 cm FL; Ceara,
hyal without teeth fused to bone (large tooth Brazil). Mota Alves 1969:167-171 (digestive
patch fused to bone); 6) hyomandibula wide, 36- tract). *Menezes 1970:171-176 (food). Dahl
52% of length (narrow, 35-36%); 7) angle of 1971:278-279 (Colombia), photograph. Alcan-

FIGURE 47.-Scomberomorus brasiliensis. Belem market, Brazil, 502 mm FL, USNM 217550, holotype.

612
COLLETTE and RUSSO: SPANISH MACKERELS

tara Filho 1972a (gill net fishery; Ceara, Bra- Diagnosis. -This species possesses nasal denti-
zil). * Gesteira 1972:117-122 (reproduction and cles as do the other five species of the regalis
fecundity). Menezes 1972:86-88 (number of group (concolor, maculatus, regalis, sierra, and
gill rakers). Bastos et al. 1973 (canning, Bra- tritor) , has the artery that branches from the
zil). Costa and Almeida 1974:115-122 (length fourth left epibranchial artery as do all the spe-
frequencies). Menezes 1976:45-48 (size, sex- cies in the group except S. tritor, and shares a
ratio; NE Brazil). Fonteles-Filho and Alcan- specialization of the fourth right epibranchial
tara-Filho 1977 (gill net mesh selectivity curve; artery (Fig. 7f) with S. sierra and S. regalis. In
Ceara, Brazil). *Sturm 1978:155-172 (biology, these three species an artery connects the fourth
Trinidad). Ximenes 1983 (age and growth; right epibranchial with a branch of the coeliaco-
Ceara, Brazil). mesenteric artery. Scomberomorus brasiliensis
Scomberomorus brasiliensis Collette, Russo, and has shorter pelvic fins than do the other members
Zavalla-Camin 1978:273-279 (original descrip- of the regalis group (Fig. 48), 3.6-5.9% FL com-
tion; Brazil). Manooch et al. 1978 (annotated pared with 4.7-6.4 in S. sierra and 4.4-6.3 in S.
bibliography). Collette 1979:29 (characters). regalis. Together with three other species of the
Collette and Russo 1979:8-11 (diagnostic char- regalis group (concolor, regalis, and sierra), S.
acters, range). Cressey et al. 1983:264 (host- brasiliensis has a long posterior process on the
parasite list, 4 copepod species). Collette and pelvic girdle, 62-90% ofthe length ofthe anterior
Nauen 1983:60-61 (description, range, fig.). plate. Differs from S. sierra by essentially lack-
ing pterotic spines. Intercalar spine absent as in
Types.-Holotype: USNM 217550 (502 mm FL); the other five species of the regalis group and S.
BeH~m market; 22 May 1975; B. B. Collette niphonius.
1642. D XVIII + 17+ X; A 19+IX; PI 22; RGR I
3 + 1 + 10 = 14; vertebrae 19 + 28 = 47. Paratypes: Description. -Lateral line gradually descending
103 specimens (110-630 mm FL) from 54 Brazilian to midline on caudal peduncle. Intestine with two
collections (see Collette et al. 1978:276-278). folds and three limbs (Fig. 3a). Spines in first

40

35
•
•
E 30
E •
:I:
G25
Z
w
..... 20
Z
... 15
U
>
.....
w 10
Q..

5
o brasiliensis
• maculatus, sierra,
tritor, regalis
0
0 75 150 225 300 375 450 525 600 675
FORK LENGTH (mm)

FIGURE 48.-Regression of pelvic fin length on fork length in five species of Scomberomorus. The regression line for S. brasiliensis is
significantly different from those for S. maculatus, S. sierra, S. tritor, and S. regalis. The regression lines for the latter four species do
not differ significantly from each other. Therefore, the same symbol is used for plotting specimens of these four species. (From Collette
et al. 1978:fig. 1.)

613
 FISHERY BULLETIN: VOL. 82, NO.4

dorsal fin 17 or 18, rarely 19 (Table 9); second ured in Ceara from 1962 to 1966, 9 exceeded 95.0
dorsal fin rays 15-19, usually 17 or 18 (Table 10); cm FL, more than 60% each year from 1962 to
finlets 8-10, usually 9 (Table 10); anal fin rays 16- 1968 were in the size range 40-65 cm (Brazilian
20, usually 17-19 (Table 11); finlets 7-10, usually 9 records summarized by Collette et al. 1978). Sex-
(Table 11); pectoral fin rays 21-24, usually 22 or ual maturity is reached at age III or IV, 46 cm FL
23 (Table 12). For a sample of 90 Brazilian S. in Ceara (Gesteira 1972). The shortest mature
brasiliensis, Bastos (1966) found the following male in Trinidad was 38 em, the shortest ripe
numbers of fin rays to be most common: dorsal female 45 cm (Sturm 1978). The length-weight
spines 18 (86.6%), rays 18 (76.6%), finlets 9 relationship for the Brazilian population was
(75.3%); anal rays 18 (100%), finlets 9 (79.8%), given by Nomura (1967). Males and females grew
pectoral rays 22 (98.9%). Precaudal vertebrae 19- at roughly equal rates up to 4 yr of age but then
21, usually 20 (Table 6); caudal vertebrae 27-29, females grew faster on to age XIV (Ximenes
usually 28 (Table 7); total vertebrae 47-49, usu- 1983).
ally 48 (Table 8). The counts of 46 or 47 reported
by Bastos (1966) presumably exclude the hypural Color pattern. -Sides with several rows of round
plate which we include in our counts. Gill rakers yellowish-bronze (in life) spots (Fig. 47) similar to
on first areh (1-3) + (9-13) = 11-16, usually 2 + (11- S. maculatus and S. sierra but without any lines
12) = 13-15 (Table 5). For a large sample from or streaks such as are present in S. regalis.
Brazil (225 males, 275 females), Menezes (1972) Number of yellowish-bronze spots on sides of
found a similar range, 11-17, and a "typical" count body increases with size of fish; young specimens
of 3 + 1 + 11 = 15. Morphometric characters are (200 mm) have about 30 spots; adults more, 45
given in Table 13. spots (422 mm), 47(455), 46(470), 45(516), and
58(530) (Collette et al. 1978). Spots arranged in
Size. -Maximum size 125 cm FL (Costa and three or four rows (sometimes in two rows). The
Paiva 1969, Ceara, Brazil). Of 16,170 fish meas- rows are not very well defined but it is possible to
recognize them. First dorsal fin black in the
anterior half (first seven membranes), posterior
TABLE I3.-Summary of morphometric data of Scom- half white with upper edge black. Pectoral fin
beromorus brasiliensis. FL = fork length, HL = head dusky; pelvic and anal fins white.
length. There is a black and white photograph of a
Character N Min. Max. Mean SO specimen from Colombia in Dahl (1971:278) and a
Fork length 69 111 710 317 136 drawing of a Venezuelan specimen in Cervig6n
Snout-A roo FL 68 508 692 538 . 24
Snout-20 roo FL 68 483 672 511 23
(1966:fig. 303).
Snout-lO roo FL 68 196 337 242 20
Snout-P, roo FL 69 217 359 253 22
Snout-P, roo FL 69 191 317 220 20 Biology. -No extensive migrations are known
P,-P, roo FL 68 83 159 108 11 for S. brasiliensis, and it is available to the
Head length roo FL 69 121 309 213 22
Max. body depth roo FL 69 164 263 197 15 fishery in northeastern Brazil all year round.
Max. body width roo FL 67 54 114 81 11 There does appear to be some seasonal movement
P, length roo FL 68 97 143 123 9
P,length roo FL 66 29 59 45 5 around Trinidad (Sturm 1978). There is a spawn-
P2 insertion-vent roo FL 66 234 349 272 18
P, tip-vent roo FL 62 193 298 226 17
ing peak in the Gulf of Paria, Venezuela, in
Base 10 roo FL 68 232 360 263 18 October-April followed by a postspawning feed-
Height 20 roo FL 62 92 139 118 11
Base 20 roo FL 69 93 153 118 10 ing migration away from Venezuela with a period
Height anal roo FL 61 83 149 115 12 of maximum abmidance in Trinidad waters May-
Base anal roo FL 68 97 142 113 10
Snoul (fleshy) roo FL 69 69 120 82 8 September. Some spawning takes place in the
Snout (bony) roo FL 65 59 102 73 7 Gulf of Paria throughout the year with a peak
Maxilla length roo FL 68 104 188 124 13
Postorbital roo FL 69 84 127 95 6 in October-April (Sturm 1978). Ripe fish are
Orbital (fleshy) roo FL 69 27 57 37 6
Orbital (bony) roo FL 69 33 77 53 8
taken on the Guyana continental shelf in Sep-
Interorbital roo FL 69 48 107 57 7 tember (Lowe 1962). Spawning takes place all
20-caudal roo FL 67 427 594 490 31
Head length 69 33 140 66 25 year round off northeastern Brazil with a peak in
Snout (fleshy) roo HL 69 354 596 386 29 the third trimester, July-September (Gesteira
Snout (bony) roo HL 65 300 546 344 31
Maxilla length roo HL 68 541 937 581 47 1972). Spawning probably takes place mostly
Postorbital roo HL 69 387 758 446 43 offshore beyond the main fishing areas. There
Orbit (fleshy) roo HL 69 136 286 175 24
Orbit (bony) roo HL 69 172 408 249 31 appear to be no references to eggs or larvae of S.
Interorbital roo HL 69 240 564 269 44
brasiliensis. As with other species in the genus,
614
COLLETIE and RUSSO: SPANISH MACKERELS

food consists largely of fishes with smaller quan- Venezuela is S. brasiliensis as is also a large
tities of penaeoid shrimps and loliginid cephalo- proportion of the Brazilian catch of Scomberomo-
pods. The most important component of the food rus spp. In Trinidad, it is taken by drift gill nets
ofl,020 individuals (17.5-87.5 cm FL) from north- that are fished overnight and with beach seines
eastern Brazil was the thread herring, Opistho- (Sturm 1978). There are two major fisheries in
nema oglinum, (more than 25%) followed by Brazil. One employs gill nets (rede-de-pesca) from
Engraulidae, Carangidae, Hemiramphidae, and wooden boats not over 10 m long powered by gaso-
Pomadasyidae (Menezes 1970). line engines (Fonteles Filho 1968; Alcantara Filho
1972a). The other method is trolling from rafts
Interest to fisheries. -This is an important food (Fonteles Filho 1968; Costa and Almeida 1974).
fish throughout its range-Colombia (Dahl 1971), Most of the catch is consumed fresh, but in Brazil
Venezuela (Cervigon 1966), Trinidad (Sturm some has been salted (Paiva and Costa 1966) and
1978), the Guianas (Gines and Cervigon 1968), some has been canned (Bastos et al. 1973).
and especially in northeastern Brazil. The fish-
ery is concentrated in June-August in Trinidad Distribution. -Caribbean and Atlantic coasts of
(Sturm 1978) but is conducted year round in Central and South America from Belize at least
northeastern Brazil (Alcantara Filho 1972a). The as far south as Lagoa Tramandai, Rio Grande do
fishing grounds are 5-16 mi offshore in Brazil SuI, Brazil (Fig. 49). Previously confused with,
(Fonteles Filho 1968; Alcantara Filho 1972a). but not known to overlap the range of, S. macula-
Most of the catch previously reported as S. macu- tus which occurs in the Gulf of Mexico and along
latus from Fishing Area 31 (Western Central the Atlantic coast of the United States. Replaced
Atlantic) for Colombia, Trinidad and Tobago, and in the West Indies by S. regalis.

60

. i
.L I
oI I
i~-cI'~-+--+- +-+--j--+-
t"
of---+--f--4-,:-t-

cif-,-f--+--+- ,J'f--t-+----+-+----+-t--1-+--+-+--+----j40
, I

j , I tit'
o ---"----:':-::o---'-------'-~.---'--'---.L._-l-----'--l607.---'---L--'--f·--'--
30
ci -'----'-..-..L--'---"UJ

FIGURE 49.-Ranges of the regalis-group of Scomberomorus: S. tritor, S. maculatus, S. regalis, S. brasiliensis,

S. sierra, and S. concolor. (Range of S. regalis more extensive, see text.)

615
 FISHERY BULLETIN: VOL. 82. NO.4

Geographic variation. -Morphometric data from Cybium acervum Cuvier in Cuvier and Valen-
two populations of S. brasiliensis were compared ciennes 1831:186 (original description, Mar-
by ANCOVA: Central America (n = 9-11) and tinique, Santo Domingo, Cuba). Poey 1865:
Brazil (n = 39-44). Null hypotheses that the 2 322 (Cuba; color pattern of juveniles). Poey
sets of regressions are coincident were accepted 1868:362 (description; Cuba). Poey 1875:147
for 24 of 26 regressions. The two populations were (after Cuvier in Cuvier and Valenciennes
different in Sn-P2 and 2D-C. Comparison of mer- 1831). Poey 1878:4 (unable to find this species
istic characters for central and northern South in Cuba).
America versus Brazil did not reveal any differ- ?Cybium clupeoidum Cuvier in Cuvier and Va-
ences (Collette et aI. 1978:tables 1-3). lenciennes 1831:178 (original description, "tIe
de Norfolk, Nouvelle Hollande").
Material examined. - Total 146 (89-710 mm FL). Scomberomorus caballa. Jordan and Gilbert
1882:427 (synonymy, range). Goode 1884:316
meas.: 69 (111-710): Belize (2); Honduras (1); (range, size), pI. 94.
Costa Rica (3); Panama (5); Colombia Scomberomorus cavalla. Meek and Newland
(1); Venezuela (3); Trinidad (2); Guyana 1884:233, 235 (description, synonymy, range).
(2); Surinam (4); French Guiana (2); Dresslar and Fesler 1889:442 (in key), 444-445
Brazil (44, *S. brasiliensis). (synonymy, range), pI. 11 (specimen from
counts: 146. Woods Hole). Jordan and Evermann 1896b:
diss.: 6 (363-639): French Guiana (2); Belem, 875-876 (description, synonymy). Evermann
Brazil (4). and Marsh 1902:124 (description, synonymy;
Puerto Rico). Jordan and Evermann 1902:
287-288 (description, range), photograph.
Scomberomorus cavalla (Cuvier) Bean 1903:400-401 (synonymy, description,
King Mackerel range). Fowler 1905:766-767 (placed in new
Figure 50 subgenus Sierra; description, Santo Domingo
and St. Martins). Smith 1907:193-194 (diag-
Guarapucu. Marcgrave 1648:178 (Brazil). nosis, range; few or no records from North
Cybium cavalla Cuvier 1829:200 (original de- Carolina). Sumner et aI. 1913:750 (references,
scription after Marcgrave's Guarapucu; Brazil). occurrence; Menemsha Bight and Quisset Har-
Cybium caballa. Cuvier in Cuvier and Valen- bor, Mass.). Ribeiro 1915:135-136 (description;
ciennes 1831:187-190 (description; Brazil, West range S to Angra dos Reis, Brazil). Meek and
Indies). Gunther 1860:373 (synonymy, de- Hildebrand 1923:322-323 (description, synon-
scription; West Indies). Poey 1865:322 (Bra- ymy). Schroeder 1924:7 (maximum weight 75
zil, Puerto Rico; C. caballa is the juvenile of lb; Fla. Keys), fig. 5. Nichols and Breder 1927:
C. acervum). Poey 1875:147 (description; 124 (description, range), fig. 172. Nichols
Cuba). Poey 1878:3-4 (synonymy, charact~rs). 1929:230-231 (range, description), fig. 84. Bee-
Cybium immaculatum Cuvier in Cuvier and Va- be and Hollister 1935:213 (Union I., Grena-
lenciennes 1831:191 (original description, no dines). Baughman 1941:16-17 (Texas records).
locality). Gunther 1860:370. Poey 1878:5 Munro 1943:69, 71-72 (placed in subgenus Sier-
(after Cuvier). ra). La Monte 1945:26 (description, range),

FIGURE 50.-Scomberomoru8 cavalla. Woods Hole, Mass., 1,000 mm FL, USNM 19418. (From Goode 1884:pl. 94.)

616
COLLE'ITE and RUSSO: SPANISH MACKERELS

color pI. 11. Breder 1948:127 (range), fig. Erd- velopment, pigmentation, counts, and measure-
man 1949:301 (West Indies). Fraser-Brunner ments; 49 larvae and juveniles (3.3-31.0 mm
1950:160-161 (range), fig. 33. Baughman 1950: SL), figs. 4, 5, 6B (larvae and juveniles, 3.3-23
243-244 (previous Texas records). Knapp . mm SL». Dahl 1971:277 (uncommon in Co-
1950:141-142 (food in Texas, shrimps, squids, lombia), fig. Ivo 1972:27-29 (gonadal stages of
fishes). Rivas 1951:224-225 (synonymy, diag- 4,346 females; Ceara, Brazil). Moe 1972:16-17
nosis, range). Taylor 1951:270 (popular an- (migrations; Florida). Alcantara Filho 1972b
glers' fish taken by trolling; North Carolina). (gill net and trolling fisheries; NE Brazil).
La Monte 1952:50 (description, range). Bige- Richards and Klawe 1972:13 (range), 89 (refer-
low and Schroeder 1953:349 (description, range; ence to Wollam 1970). Miyake and Hayasi
Gulf of Maine record, N Truro, Cape Cod), fig. 1972:111:3 (in key), IV:11 (common names).
184. Pew 1954:26 (description, range, habits), Dwinell and Futch 1973 (139 larvae and juve-
fig. 22. Mather 1954:292 (13 specimens, about niles, 2.8-13.5 mm SL, all months; NE Gulf of
70 cm FL, in trap; Quisset, Mass.). Mather Mexico). Bastos et aI. 1973 (canning; NE Bra-
and Gibbs 1957:243 (9 specimens, 600-700 mm zil). *Beaumariage 1973 (age, growth, food,
FL; Buzzards Bay, Mass.). Briggs 1958:287 reproduction; Florida). Ivo 1974 (fecundity;
(range). *Mago Leccia 1958 (osteology, com- Ceara, Brazil). Berrien and Finan 1977a (spe-
parisons with S. maculatus and S. regalis), cies synopsis). Erdmann 1977:150 (in spawn-
figs. Butz and Mansueti 1962:130-135 (de- ing condition mainly in July and Aug.; NE
scription; N Chesapeake Bay; comparison with Caribbean). Klawe 1977:2 (common name,
specimens from Mass. and Fla.), fig. 2 (head). range). DeVane 1978 (stomach contents;
Moe 1963:108-109 (most sought fish in Fla. North Carolina). Fritzsche 1978:121-125 (de-
charter boat fishery). Collette 1966:365-367 scription, larval development), figs. 66-69 (lar-
(types of C. aeervum and C. immaculatum; vae). Collette 1978: Scombm 4 (description,
both names synonyms of S. cavalla). Nomura range), figs. Manooch et aI. 1978 (annotated
and Costa 1966:11-13 (length-weight of666 spec- bibliography). Lima and Oliveira 1978:6, 23
imens; Ceara, Brazil). Cervigon 1966:718-719 (common name "cavalla" in Brazil). Collette
(description; Venezuela). Randall 1967:753- 1979:29 (characters, range). Collette and
754 (food of 22 West Indian specimens, 92.3% Russo 1979:9 (diagnostic characters, range).
fishes). Nomura and Rodriguez 1967:79-85 Manooch 1979 (commercial U.S. catches aver-
(age and growth, condition factor, 1,504 speci- aged 2,541 t/yr over last 17 yr, recreational
mens, 30-120 em FL; Ceara, Brazil), fig. 1 catch statistics are inadequate). Meaburn
(sagitta). Mota Alves and Tome 1967a:103- 1979 (heavy metal contamination). McEach-
108 (anatomy and histology of the digestive ran et aI. 1980 (larvae off Texas coast). Fisch-
tract), figs. 1, 2 (arrangement of viscera), figs. er 1980:1-21 (size, length-weight, sex ratio;
3-7 (histology of gut). Mota Alves and Tome Louisiana). Sutherland and Fable 1980 (an-
1967b:1-9 (histology of gonads), figs. 1-11 (photo- nual migration from S Florida N to NE Gulf
micrographs). Mota Alves and Tome 1967c: of Mexico and W to S Texas in the spring).
173-175 (anatomy and histology of the liver and MacGregor et aI. 1981 (significant correlation
gall bladder). Mota Alves and Tome 1968c: found between gonadosomatic indices and se-
31-32 (sperm). Fonteles Filho 1968 (fishery; rum estrogens in females and with serum an-
NE Brazil). Nomura and Costa 1968:95-99 drogens in males). Fable et aI. 1981 (tempera-
(length-weight relationship, 104 males and 90 ture effects on catches; NW Florida). Lubbock
females; Ceara, Brazil). Randall 1968:119 (de- and Edwards 1981:150 (Saint Paul's Rocks).
scription, range, habits), fig. 136. Lyles 1969: Richardson and McEachran 1981 (larvae 2.0-
16-21 (summary of U.S. landings, 1880-1967). 2.9 mm SL, pigment characters, measurements;
Menezes 1969a:15-20 (food of 798 specimens; Gulf of Mexico), fig. 2A (2.3 mm larva). Naugh-
Ceara, Brazil; fishes compose main diet). ton and Saloman 1981 (stomach contents of 139
Menezes 1969b:175-178 (meristic characters, juveniles, 103-309 mm FL; Cape Canaveral,
osteology; NE Brazil). Mota Alves and Tome Fla.; diet mainly clupeoids). Sacchi et aI.
1970:181-184 (histology and enzymes of pylor- 1981:3 (French Antilles). Trent et al. 1981
ic caeca). Beardsley and Richards 1970:5 (size composition and sex ratio; SE U.S.).
(length-weight o£197 specimens, 585-1,500 mm Ximenes et aI. 1981 (age and growth; NE Bra-
FL, 1.47-32.09 kg; Florida). Wollam 1970 (de- zil). Morgan and King 1983 (tooth replace-
617
 FISHERY BULLETIN: VOL. 82, NO.4

ment). Johnson et aI. 1983 (age, growth, and curve in the lateral line under the first dorsal fin
mortality; SE U.s.). Cressey et aI. 1983:264 but the lateral line gradually descends in the
(host-parasite list, 4 copepod species). Col- other 15 species. Scomberomorus caualla differs
lette and Nauen 1983:61-62 (description, range), from S. commerson in having fewer vertebrae
fig. Saloman and Naughton 1983a (food in (41-43, usually 42 or fewer compared with 42-46,
SE U.S.). usually 43 or more) and more gill rakers (7-13,
usually 8 or more compared with 1-8, usually 7
Types of nominal species. -Cybium caualla Cu- or fewer). Ventral process of angular moderate,
vier, 1829 is based on Marcgrave's description 87-93% of dorsal process as in S. sinensis. Ascend-
and figure (1648:179) of the "Guarapucu"; there ing process of premaxilla short as in S. gutta-
are no extant types for this name. tus. Anterior ends ofpterosphenoid close together
Cybium aceruum Cuvier in Cuvier and Valen- as in S. commerson. Intercalar spine well devel-
ciennes, 1831. Lectotype: MNHN A.5781; Santo oped as in S. commerson and S. queenslandi-
Domingo; Ricord; 130 mm FL; selected by Collette cus.
(1966:365); D XV + 17 + VIII; A 18+ VIII; RGR 1
1 + 1 + 7 = 9; vertebrae 17 + 25 = 42; upper jaw Description. -Intestine with two folds and three
teeth 8-11; lower jaw teeth 7-8. Paralectotypes: limbs (Fig. 3b). Spines in first dorsal fin 12-18,
MNHN B.2508, out of A.5781; Santo Domingo; usually 15 (Table 9); second dorsal fin rays 15-18,
Ricord; 2(133-138 mm FL). A photograph of one usually 17 or 18 (Table 10); dorsal finlets 7-10,
of the syntypes was published by Blanc and usually 9 (Table 10); anal fin rays 16-20, usually
Bauchot (1964: pI. 1, fig. 1, upper fig.). 18 or 19 (Table 11); anal finlets 7-10, usually 8
Cybium immaculatum Cuvier in Cuvier and (Table 11); pectoral fin rays 21-23, usually 22 or
Valenciennes, 1831. Lectotype: MNHN A.5720; 23 (Table 12). Precaudal vertebrae 16 or 17, usu-
Martinique; Plee; 157 mm FL; selected by Col- ally 17 (Table 6); caudal vertebrae 24-26, usually
lette (1966:366); D XV + 17 + IX; A 17 + IX; P 1 23; 25 (Table 7); total vertebrae 41-43, usually 42
RGR 1 1 + 1 + 7 = 9; vertebrae 17 + 25 = 42; upper (Table 8). Gill rakers on first arch (1-2) + (6-11) =
jaw teeth 9-11; lower jaw teeth 9-12. Paralecto- 7-13, usually 1 + (8-9) = 9-10 (Table 5). Counts for
types: MNHN B.2509; out of A.5720; Martinique; a large Brazilian sample (353 individuals, Me-
Plee; 147 mm FL; and MNHN A.5780; Martin- nezes 1969b), were (0-2) + 1 + (5-9) = 6-11, usually
ique; Plee; 164 mm FL. Photographs of two of 1 + 1 + 7 = 9. Morphometric characters are given
the syntypes were published by Blanc and Bau- in Table 14.
chot (1964:pI. 2, fig. 12).
Cybium clupeoidum Cuvier in Cuvier and Va- Size. -Maximum size 172.5 em FL (female, 37.2
lenciennes,1831. Holotype: MNHN A.5784; "lIe kg; Beaumariage 1973); common to 70 em. The
de Norfolk, a l'oeust de la Nouvelle-Hollande"; all-tackle angling record is a 40.8 kg fish taken
Broussonet collection; 302 mm FL; D XV + 17 + off Key West, Fla., in 1976. In Florida, females
IX; A 18 + VIII; RGR 1 1 + 1 + 7 = 9; vertebrae 17 + usually mature in their fourth summer at a mean
25 = 42; upper jaw teeth 13-13; lower jaw teeth 11- length of 83.7 em, males in the third summer at
12. A photograph of the type was published by 73 em (Beaumariage 1973). In Brazil, females
Blanc and Bauchot (1964:pI. 1, fig. 3). The high mature at age V-VI, about 77 em according to Ivo
gill raker count and low vertebral number show (1972), at age IV and 63 em according to Gesteira
the type to be a specimen of the western Atlantic and Mesquita (1976). Males and females grow at
S. caualla as supposed by Bauchot and Blanc roughly equal rates up to age V but then females
(1961) and Blanc and Bauchot (1964) rather than grow faster (Ximenes et aI. 1981). They reach an
S. commerson as presumed by Collette (1966) age of at least XIV (Ximenes et aI. 1981; Johnson
based on geography. The locality has been sup- et aI. 1983). Length-weight relationships have
ported by Bauchot (1969), but the data or the been published for Brazil (Nomura and Costa
specimen must have been mixed with the western 1968; Ximenes et aI. 1981), Florida (Beaumariage
Atlantic species sometime in the past. 1973), and Louisiana (Fischer 1980).

Diagnosis. -This species shares with S. commer- Color pattern. -Adults have plain silvery sides
son an abrupt downward curve in the lateral line without bars or spots, juveniles have bronze spots
under the second dorsal fin (Fig. 50). Scomber- smaller than the pupil of the eye in five or six
omorus sinensis also has an abrupt downward irregular rows (Randall 1968:119). Adults have no
618
COLLETIE and RUSSO: SPANISH MACKERELS

TABLE 14.-Summary of morphometric data of Scomberomorus cavalla. FL = fork length, HL = head length.
Unit.,p States West Indies South America Total

Character N Mi". Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO
Fork length 10 560 1,160 769 179 20 126 710 323 209 23 147 860 326 183 54 126 1,160 405 257
Snout-A %< FL 10 510 562 535 15 20 519 582 547 23 23 463 568 533 21 54 463 582 539 21
Snout-20 r. FL 10 474 521 499 14 20 486 534 512 12 23 430 524 504 18 54 430 534 505 16
Snout-1D %< FL 10 225 266 241 11 20 247 297 268 13 23 210 273 256 15 54 210 297 258 16
Snout-P, r. FL 8 223 260 236 11 20 239 290 268 14 23 205 293 256 21 52 205 293 257 20
Snout-P, %< FL 10 199 241 212 12 20 212 260 243 14 23 194 252 231 14 54 194 260 232 17
P,-P, %< FL 5 86 103 93 7 13 91 136 107 15 22 73 135 107 15 41 73 136 105 15
Head length %< FL 10 192 229 203 10 20 206 250 231 13 23 177 246 224 17 54 177 250 223 17
Max. body depth %< FL 7 149 173 159 10 10 153 258 198 37 22 147 237 197 25 40 147 258 190 30
Max. body width %< FL 8 95 109 102 5 11 78 101 89 9 21 70 126 85 13 41 70 126 89 12
P, length r. FL 10 114 138 124 7 15 108 143 129 12 23 111 142 130 7 49 108 143 129 9
P,length r. FL 10 53 62 58 3 12 54 95 68 10 21 36 85 66 12 44 36 95 64 11
P2 insertion-vent r. FL 8 264 294 281 9 15 245 309 277 17 22 211 298 266 18 46 211 309 272 18
P, tip-vent %< FL 8 205 237 222 11 12 187 236 214 14 21 162 289 208 26 42 162 289 212 21
Base 10 %< FL 9 221 255 242 11 19 224 267 246 11 23 207 269 247 14 52 207 269 245 13
Height 20 r. FL 9 85 108 95 7 14 76 127 108 13 20 79 128 114 12 44 76 128 108 13
Base 20 %< FL 10 78 103 93 7 14 82 124 105 12 23 81 135 112 15 48 78 135 106 14
Height anal r. FL 8 89 102 95 5 11 81 122 107 12 17 89 127 110 9 37 81 127 105 11
Base anal %< FL 10 87 111 98 9 17 92 123 107 10 23 83 129 112 12 51 83 129 108 12
Snout (fleshy) %< FL 10 74 103 84 8 17 83 98 89 4 23 72 92 86 5 51 72 103 87 8
Snout (bony) %< FL 10 68 96 78 8 17 73 86 81 4 22 68 85 78 4 50 68 96 79 5
Maxilla length r. FL 10 108 137 117 8 20 120 152 138 9 23 103 147 132 11 54 103 152 132 12
Postorbital r. FL 7 86 99 92 4 17 89 112 99 6 23 81 104 98 6 48 81 112 98 6
Orbital (fleshy) r. FL 10 24 43 30 5 17 33 58 41 8 23 27 54 39 6 51 24 58 38 7
Orbital (bony) r. FL 10 28 48 41 6 17 42 73 55 10 23 38 65 52 6 51 28 73 51 9
Interorbital %< FL 10 52 68 55 4 20 56 66 62 3 23 48 66 60 5 54 48 68 60 4
20-caudal %< FL 7 485 527 506 13 13 452 500 473 14 23 445 516 472 18 44 445 527 477 20
Head length 10 112 266 158 45 20 32 151 72 44 24 36 152 73 35 55 32 266 88 51
Snout (fleshy) r. HL 10 362 451 415 25 17 350 424 391 25 24 354 422 384 17 52 350 451 392 24
Snout (bony) %< HL 10 355 417 384 20 17 305 388 353 27 23 318 388 351 19 51 305 417 358 26
Maxilla length %< HL 10 546 604 576 18 20 577 613 '598 10 24 570 609 590 10 55 546 613 591 14
Postorbital %< HL 7 428 485 453 22 17 408 463 434 16 24 395 463 437 16 49 395 485 439 18
Orbit (fleshy) r. HL 10 123 210 147 24 17 152 233 176 24 24 147 220 171 18 52 123 233 168 24
Orbit (bony) %< HL 10 138 238 204 28 17 191 305 237 31 24 201 265 231 15 52 138 305 229 27
Interorbital %< HL 10 256 295 272 11 20 248 284 267 10 24 254 281 269 7 55 248 295 269 9

black area in the anterior part of the first dorsal San Blas in May (Sutherland and Fable 1980).
fin as do many species of Scomberomorus. The main run usually arrives in Panama City,
Black and white photographs are given by Fla., in late Mayor early June. The westward
Jordan and Evermann (1902) and Randall (1968: migration along the northern Gulf of Mexico ends
fig. 136). The drawing published by Goode (1884: off west Texas in June-July (Sutherland and
pI. 94) is'included here as Figure 50, Fable 1980). Return migration in the fall from
summer feeding grounds in the northwest Gulf to
Biology. -A summary of biological information winter feeding grounds off southern Florida has
has been presented by Berrien and Finan (1977a) been confirmed by recaptured tagged fish (Suth-
and there is also a useful annotated bibliography erland and Fable 1980), Based on gonad develop-
by Manooch et aI. (1978). King mackerel appear ment and larval distribution, spawning takes
to be present all year in Louisiana (Fischer 1980) place in the northeastern Gulf of Mexico and in
and in the state of Ceara in northeastern Brazil. the Atlantic offshore of Cape Kennedy, Fla., and
Some populations appear to be resident in south northward in late summer (Moe 1972). According
Florida waters as they are available to the recre- to Beaumariage (1973), spawning in Florida may
ational fishery throughout the year. However, be protracted as indicated by successive increase
the large schools that are found in south Florida in vitellogenic oocyte size during the summer.
waters during January and February move north Spawning takes place in May-September in the
along both coasts in the spring (Moe 1972). western Gulf of Mexico, especially in September
Schools that occur offshore of Palm Beach and in waters 35-183 m deep over the middle and
Martin Counties on the east coast of Florida in outer continental shelf (McEachran et al. 1980).
winter and early spring move north. They appear In the northeastern Caribbean, spawning peaked
off North Carolina in April and remain until fall in July and August (Erdman 1977). Spawning is
(DeVane 1978). On the west coast of Florida, king year round offshore of Ceara, northeastern Brazil
mackerel move north to the Naples-Ft. Myers or (Ivo 1972). Larvae and juveniles (139 specimens,
St. Petersburg-Tampa areas by April and Cape 2.8-28.8 mm SL) were taken off the northwest
619
 FISHERY BULLETIN: VOL. 82, NO.4

coast of Florida from June to October with larvae coast of northeastern Brazil where they are taken
< 3.1 mm taken in June, August, and September all year (Nomura and Rodrigues 1967). The main
(Dwinell and Futch 1973). Most of these larvae fishing grounds in northeastern Brazil are 6-16
and juveniles were taken in surface plankton nmi from the coastline (Fonteles Filho 1968). An
tows at surface temperatures of 26.3°-31.0°C and historical summary of the fishery in the United
salinities of 26.92-35.0%•. Larvae were taken in States has been presented by Lyles (1969). Com-
increasing numbers from May to September (35% mercial catches in the United States have aver-
or more of larvae in September of each year) in aged 2,541 t a year with a value of $1.3 million
the western Gulf of Mexico, particularly over the over 17 yr with a peak in 1974 of 4,764 t (Manooch
middle and outer continental shelf (McEachran 1979). The bulk of these landings were made in
et al. 1980). Larvae and juveniles have been Florida by hook and line and gill net fisheries
described and illustrated by Wollam (1970; 3 (Manooch 1979). Data on the large recreational
figures, 3.3-23 mm SL), Fritzsche (1978; 12 fig- catch are inadequate. The catch reported from
ures, 2.98-17 mm), and Richardson and McEach- Fishing Area 31 (Western Central Atlantic) to-
ran (1981, 2.3 mm SL). As with other members of talled 7,122 t in 1982 (FAO 1984) but is higher
the genus, food consists primarily of fishes with than this because much of the catch of 1,105 tons
smaller quantities ofpenaeoid shrimps and squids of unclassified Scomberomorus species is S. ca-
(Knapp 1950, Texas; Randall 1967, Caribbean; valla (or S. regalis). It is fished for with hook and
Menezes 1969a, northeastern Brazil; Beaumar- line in all the southeastern United States (Trent
iage 1973, Florida; DeVane 1978, North Carolina; et al. 1981). In addition, there is a commercial
Saloman and Naughton 1983a, United States). fishery using snapper hooks and line in Missis-
Clupeids such as Opisthonema, Harengula, Sar- sippi, a commercial gill net fishery in southern
dinella, and Brevoortia are particularly impor- Florida, and commercial hook and line fisheries
tant (Randall 1967; Menezes 1969a; Beaumariage in North Carolina and southern Florida (Trent et
1973; DeVane 1978; Saloman and Naughton al. 1981). The gill net fishery has employed power
1983a), even in juveniles 103-309 mm FL (Naugh- block retrieval since 1963 and aerial spotting is
ton and Saloman 1981). Other fishes commonly sometimes used (Beaumariage 1973). The king
consumed include Carangidae (particularly De- mackerel is the staple of the charter boat indus-
capterus), Lutjanidae, Pomadasyidae, and Hemi- try in Florida and is the most sought fish by
ramphidae (Randall 1967; Menezes 1969a; Beau- private boats (Moe 1963). In Florida it is most
mariage 1973; Saloman and Naughton 1983a). often fished at the surface with trolled lure or
small bait fish (Moe 1963). It is less commonly
Interest to fisheries. -The king mackerel is an caught than is S. brasiliensis across the northern
important species for recreational, commercial, coast of South America (Dahl 1971; Cervigon
or artisanal fisheries throughout its range from 1966; Gines and Cervigon 1968). Both gill nets
southeastern United States to northeastern Bra- and trolling are used in northeastern Bra~il, the
zil. North of southern Florida, the fishery is former catching 87.6% II-IV yr fish and the lat-
concentrated in the summer months. In North ter 78.2% IV-VI yr fishes (Alcantara Filho
Carolina, sport fisning is carried out from April 1972b). The Brazilian fishery is also carried out
to December (DeVane 1978) but is concentrated from rafts with hooks baited with thread herring
in spring and fall (Taylor 1951). In the Panama (Fonteles Filho 1968). Most of the catch is pro-
City area ofthe Florida panhandle, fish are taken cessed into steaks or sold fresh (Lyles 1969), but
from April to November and are most often it has been canned (Bastos et al. 1973) and
caught in August and September (Fable et al. salted (Paiva and Costa 1966) in northeastern
1981). From December to March the fishery along Brazil.
the east coast of Florida is concentrated from
Jupiter Inlet to Palm Beach Inlet, the rest of the Distribution. - Western Atlantic Ocean from
year the fishery is further north from Ft. Pierce Massachusetts to Rio de Janeiro, Brazil (Fig. 51).
to Sebastian Inlet (Beaumariage 1973). There is a There are several summer records from the south-
winter commercial fishery in the Florida Keys ern side of Cape Cod (Dresslar and Fesler 1889;
(Beaumariage 1973). King mackerel are taken all Sumner et al. 1913; Mather 1954; Mather and
year in Louisiana with a maximum in November- Gibbs 1957) but only one stray is known to have
January (Fischer 1980). King mackerel is the moved around to the north side of Cape Cod, to
main species of commercial interest along the North Truro in the Gulf of Maine (Bigelow and
620
FIGURE 51.-Ranges of Scomberomorus cavalla, S. commerson, and S. sinensis.
 FISHERY BULLETIN: VOL. 82, NO.4

Schroeder 1953:349; MCZ 37041, 560 mm FL). Scomberomorus commerson (Lacepede)

Abundant in the West Indies. The range extends N arrow-Barred King Mackerel
south to at least Rio de Janeiro (Angra dos Reis, Figure 52
Ribeiro 1915; Rio de Janeiro, MCZ 17269, BMNH
1903.6.9.79). Scomber commerson Lacepede 1800:598, 600-603
(original description after a figure from Com-
Geographic variation. -Samples were adequate merson's manuscripts), pI. 20, fig. 1.
to compare the morphometric data of three popu- Scomber Konam Russell 1803:27-28 (description;
lations of S. cavalla by ANCOVA· (Table 14): Vizigapatam, Coromandel coast of India), pI.
United States (n = 7-10), West Indies (n = 11-20), 135.
and South America (n = 17-23). Null hypotheses Scomber Commersonii. Shaw 1803:589 (descrip-
that the 3 sets of regressions are coincident were tion after Lacepede), pI. 85 (bottom fig.).
accepted for only 10 of 26 regressions, rejected for Scomber Maculosus Shaw 1803:592 (original de-
the other 16. For 12 regressions (Sn-A, Sn-2D, Sn- scription based on the Konam of Russell 1803,
1D, Sn-P 2 , Sn-PI, P I -P2 , Hd L, Sn (fleshy and pI. 135).
bony), maxilla L, postorbital, and interorbital), Cybium Commerson(i)(ii). Cuvier 1829:200
all three populations were significantly different (listed in footnote after Sc. Commersonii Lace-
from each other by the Newman-Keuls Multiple pede). Cuvier in Cuvier and Valenciennes
Range Test. For these 12 regressions,. there is a 1831:165-170 (description, earlier references;
cline from the United States to the West Indies to Pondichery and Malabar, India; Mauritius).
South America, a decreasing cline in slopes. The Richardson 1846:268 (range, references).
United States and West Indies populations dif- Bleeker 1853:42 (Malabar and Pondichery, In-
fered in one additional regression (2D-C) and the dia; Mauritius, Red Sea, China). Gunther
South America and the West Indies populations 1860:370 (synonymy, description; Malayan
differed in two additional regressions (maximum Peninsula and Cape Seas). Playfair and Gun-
depth and Ht 2D). No meristic differences were ther 1866:67 (Zanzibar and E coast of Africa).
found between the three populations, all usually Klunzinger 1871:494-495 (description, range).
had 15 spines in the first dorsal fin, 9 or 10 gill Bleeker 1873:131 (China, listed). Bleeker
rakers on the first arch, and 42 vertebrae. 1874:100 (Mauritius, listed). Day 1878:255-
256 (synonymy, description, range), pI. 56, fig.
Material examined. -Total 76 (126-1,160 mm FL). 5. Bleeker 1879:18 (Mauritius, listed). Cas-
telnau 1879:352 (Port Jackson, Australia;
meas.: 54 (126-1,160): E United States (3); Fla. listed). Kent 1893:229 (Great Barrier Reef,
(7); Veracruz (1); West Indies (20) (*C. Australia), pI. 46, fig. 1. Kishinouye 1923:416-
acervum Cuvier, *C. immaculatum Cu- 418 (synonymy, C. multifasciatum Kishinouye
vier), Trinidad (3); Guyana (10); Suri- a synonym of C. commerson; description, anat-
name (3); Brazil (7). omy; Japan, Taiwan, and S China), pI. 22, fig.
counts: 76. 36 (adult). Reeves 1927:8 (NE China and Ko-
diss.: 7 (557-909): Chesapeake Bay (1); Miami rea; listed). Umali 1936:98-99 (food fish; Phil-
(4); Panama City, Fla. (2). ippine Is.), fig. 59. Umali 1938:182 (fishery;

FIGURE 52.-Scomberomorus commerson. Queensland, 968 mm FL. (From Munro 1943:pl. 6B.)

622
COLLETIE and RUSSO: SPANISH MACKERELS

Ragay Gulf, Luzon, Philippine Is.). Domantay Herre 1953:245-246 (synonymy; Philippine rec-
1940:379 (important species; Margosatubig, ords). Ommanney 1953:66 (off Marie Louise
Zamboanga, Philippine Is.). Chevey and Du- I., S Amirante Is.). Devanesen and Chidam-
rand 1945:27 (description, food fish; Indochina), baram 1953:32-36 (names, description, fishery,
fig. Chacko 1949:89 (stomach contents of 12 economic importance), fig. 34. Tham 1953:49
specimens, 21-43 cm FL; Gulf of Mannar, India; (Singapore Straits). Fowler 1959:167 (descrip-
mostly clupeoids such as Stolephorus and Dus- tion, synonymy, locality records; Suva, Fiji),
sumiera). Chacko 1950:171 (characters of eggs 583 (additional references). Jones et al. 1960:
and larvae; Krusadai I., Gulf of Mannar, India). 136 (Andaman-Nicobar Is. waters). Jones
Mori 1952:136 (Fusan, Korea; listed). La 1962:113-117 (larvae and juveniles; S Kerala,
Monte 1952:51 (description, range), color pI. 19. India), figs. 9-14 (postlarvae and juveniles 14.4-
Gopalan Nayar 1958:49-51 (fishery; Vizhing- 278 mm). Bauchot and Blanc 1961:370 (de-
am, S India). Munro 1958b:262-263 (many scription of "neosyntypes"). Kaikini 1961:357
records; New Guinea region). Fourmanoir (largest species in the seerfish fishery at Mal-
and Crosnier 1964:386-387 (found along the wan, India, reaching 17.24 kg). Venkatara-
entire coast of Madagascar; one of most impor- man 1961:292 (teleosts in stomachs of 2 speci-
tant food fishes; occasional in lagoon at May- mens; Calicut, India). Kumaran 1964:586-587
otte, Comores Is.). Chacko et aI. 1967:1007- (stomach contents 283 specimens, 17-225 mm
1008 (drift net fishery; Madras State). FL; Vizhingam, W coast of India; 79% small
Cybium Konam Bleeker 1851a:357 (original de- fishes, 43% Anchoviella). Baissac 1964:186
scription, Batavia). Bleeker 1852:39-40 (de- (now scarce in Mascarene waters). Boeseman
scription; Batavia). Bleeker 1853:42 (Coro- 1964:467 (types of C. konam = S. commer-
mandel, India; East Indies). Kner 1865:144 sonii), pI. 4, fig. 16 (lectotype of S. ko-
(description; Manila). nam). Blanc and Bauchot 1964:444-445 Cneo-
Scomberomorus commerson(i)(ii). Jordan and syntypes" of C. commersonii), pI. 1-2, figs.
Seale 1906:228 (New Guinea, East Indies; 4-7. Gorbunova 1965a:53 (spawning sea-
listed). Jordan and Seale 1907:13 (description; son). George and Athanassiou 1965:1-4 (St.
Cavite, Luzon, Philippine Is.). Jordan and George Bay, Lebanon; first Mediterranean rec-
Dickerson 1908:610 (Suva market; Fiji). Fowl- ords; description), fig. 1 (49.0 and 60.3 cm TL
er 1918:63 (Philippine Is.; listed). Whitley specimens). Collette 1966:369 (Bauchot and
1927:5 (Fiji; listed). Herre 1931:33 (Philippine Blanc's "neosyntypes" invalid). Kamohara
localities). Whitley 1932:289 (Snapper I., 1967:44 (description; Japan), color pI. 22, fig.
Great Barrier Reef). Herre 1933:7 (Dumagu- 4. George and Athanassiou 1967:238 (listed
ete, Philippine Is.; listed). Hardenberg 1936: among species entering the Mediterranean
252 (mouth of Kapuas R., W Borneo). *Munro through the Suez Canal). Anonymous 1967:46
1942:33-48 (spawning, eggs, early larvae; N (off NW coast of Borneo). Mauge 1967:120
Queensland), pIs. 2-4, figs. 1-17 (eggs and early (listed from Smith's Fishes of South Africa).
larvae). *Munro 1943:67, 71-72 (placed in sub- Arnoult and Fourmanoir 1967:134, 139 (juve-
genus Cybium), 74-82 (description, anatomy, niles in mangrove swamp; Nossi-Be, Madagas-
synonymy, occurrence in Australia); pI. 6, fig. car). Ben-Tuvia 1968:35 (commercially impor-
B (968 mm FI specimen; Queensland); fig. 2.4 tant fish; several caught trolling in Dahlak
(viscera); pI. 8, fig. 3 (368 mm FL immature Archipelago; many from coast of Ethiopia, com-
specimen; N Queensland). Chapman 1946:169 mon in Eilat). Ben-Yami 1968:37 (caught by
(off New Caledonia). Herre and Urnali 1948 trolling and purse seine; Ethiopia). Wongra-
(common names in several languages and dia- tana 1968 (trawl survey; Thailand). Silas
lects; Philippine Is.). Barnard 1948:380 (49- 1967:1096 (leaping out of the water; Gulf of
in, 24-lb specimen; False Bay, South Africa). Mannar), 1,113-1,115 (length-weight). Merce-
Norman and Fraser 1949:153-154 (range). ron 1970:72-81 (length-weight, maturity, food
Fraser-Brunner 1950:161 (synonymy, range), mostly anchovies, movements; Cambodia).
fig. 34. Umali 1950:9 (found throughout the *Tongyai 1970 (distribution, peak fishing
Philippines in open sea, bays, and gulfs). months, migrations, food, fishery; Thailand).
Warfel and Manacop 1950:42 (in otter-trawl Collette 1970:3, 5 (Mediterranean coast of Is-
catches; Philippine Is.). Warfel 1950:2 (regu- rael). *Prado 1970:91-116 (synonymy; descrip-
larly found in fresh fish market, Philippine Is.). tion; biology, length-frequency, food, sex ratio,
623
 FISHERY BULLETIN: VOL. 82, NO.4

reproduction; Madagascar). Ben-Thvia 1971: Types of nominal species. -Scomber commerson

20-21 (3 specimens from Mediterranean coast Lacepede, 1800 is based on a figure from Com-
of Israel). Tongyai 1971a:9-13 (description), merson's manuscript; no types of this name are
pI. II (photograph), pI. III (viscera). Tongyai extant.
1971b:3 (economically important; Thailand), pI. Scomber Maculosus Shaw 1803 is based on the
7, 8, 10, 13 (photographs). Dhawan et aI. 1972: "konam" of Russell (1803:pI. 135); no types of this
183 (trolling line operations; Goa; feed on sar- name are extant.
dines). Nagabhushanam and Chandrasek- Cybium konam Bleeker 1851b. Lectotype:
hara Rao 1972:303 (Minimoy Atoll, Laccadive RMNH 6051; Batavia; P. Bleeker; 444 mm FL;
Archipelago). Shiino 1972:71 (common name). selected by Boeseman (1964:467); D XVII + 18 +
Richards and Klawe 1972:13 (range), 90-91 (ref- VIII; A 18 + IX; PI 22-22; RGR I 0 + 1 + 2 = 3; upper
erences to eggs, larvae, and juveniles). Mag- jaw teeth 15-16; lower jaw teeth 15-12. A photo-
nuson 1973:350 (short pectoral fin). Orsi 1974: graph of the lectotype was published by Boese-
175 (Vietnam; listed). Ronquillo 1974 (caught man (1964:pI. 4, fig. 16). Paralectotypes: RMNH
by light fishing; Philippine Is.). Lewis et aI. 24087; 12 specimens; in part. The original de-
1974:82-85 (93 specimens, 53.7-144.5 cm FL; scription was based on 12 specimens 90 to 490
ova diameters, maturity of ovaries; Bismark lines (= mm) long from Batavia. Boeseman (1964)
Archipelago, Papua New Guinea). Van der found more than 12 specimens in RMNH 6051,
Elst 1976:25 (important predator on Pomato- selected the largest specimen as lectotype, re-
mus saltatrix; Natal, South Africa). Baissac moved 2 specimens that were below the mini-
1976:216 (Mauritius). *Devaraj 1977 (osteol- mum size of the type-series, and recatalogued the
ogy). Klawe 1977:2 (common names, range). remainder of the material as RMNH 24087.
Randall et aI. 1978:166 (Persian Gulf: photo-
graph), 212 (color photograph 56). Uchida Cybium multifasciatum Kishinouye 1915. The
1978:13, 17, 20 (fishery resource; Cook Is., New original description was based on a specimen
Caledonia; Wallis and Futuna Is.). Collette from Yamaguchi Prefecture, Japan in 1914 and is
1979:29 (characters, range). Collette and Rus- probably no longer extant. Data from the original
so 1979:9, 13 (diagnostic characters, range). description show this name to be a junior syn-
Golani and Kredo 1981:41 (fishery; Mediterra- onym of S. commerson: D XVII + 15 + IX; A 14 +
nean coast of Israel). Hutchins 1979:83 (Rott- IX; GR 1+2=3; vertebrae 20+24=44; and lat-
nest I., off Perth, W Australia). Joubert 1981: eralline forming a deep bend. The author himself
5 (minor component of shore angler's catch; (Kishinouye 1923:416) subsequently placed mul-
Natal, South Africa). McPherson 1981 (biol- tifasciatum in synonymy.
ogy, migrations; Queensland). Van der Elst
1981:274 (photograph, description, natural his- Diagnosis. -This species shares with S. cavalla
tory, range). Kyushin et aI. 1982:227 (descrip- an abrupt downward curve in the lateral line
tion, photograph). * Devaraj 1982 (age and under the second dorsal fin (Fig. 52). One species,
. growth). Sivasubramaniam and Mohamed S. sinensis, has an abrupt downward curve in the
1982:65 (Qatar, Persian Gulf). Lewis and En- lateral line under the first dorsal fin but the
dean 1983 (presence of a ciguatoxin-like sub- lateral line descends gradually in the other 15
stance in Queensland specimens caught be- species. It differs from S. cavalla in having more
tween lat. 24°S and 26°S). Lewis et aI. 1983: vertebrae (42-46, usually 43 or more compared
14-21 (biology; Fiji). Cressey et aI. 1983:264 with 41-43, usually 42 or fewer) and fewer gill
(host-parasite list, 10 copepod species). Lee rakers (1-8, usually 7 or fewer compared with 7-
and Yang 1983:229-230 (Taiwan), fig. 19 (580 13, usually 8 or more). Posterodorsal spine of
mm FL). Collette and Nauen 1983:63-64 (de- hyomandibula large as in S. queenslandicus and
scription, range), fig. Jenkins et aI. 1984:348- Acanthocybium. Palatine tooth patch very nar-
351 (62 larvae, 3.5-9.3 mm SL; off Townsville, row (Fig. 23b) as in S. sinensis and Acanthocyb-
Qld.), fig. 3 (6 larvae, 3.7-9.1 mm SL). ium. Ventral process of angular long, 117-126% of
Cybium multifasciatum Kishinouye 1915:9 (orig- dorsal process, as in S. queenslandicus and Acan-
inal description; Yamaguchi Prefecture, Ja- thocybium. Anterior ends of pterosphenoid close
pan), pI. 1, fig. 3. together (Fig. 17a) as in S. cavalla. Intercalar
Scomberomorus konam. Herre 1953:246 (syn- spine well developed (Fig. 11a) as in S. cavalla
onymy). and S. queenslandicus.
624
COLLE'ITE and RUSSO: SPANISH MACKERELS

Description. -Intestine with two folds and three Color pattern. -Munro (1943:75) presented a
limbs (Fig. 3c). Spines in first dorsal fin 15-18, good description of Australian specimens. Sides
usually 17 (Table 9); second dorsal fin rays 15- pale silver gray marked with transverse vertical
20, usually 17 or 18 (Table 10); dorsal finlets bars of a darker gray. Bars narrow and slightly
8-11, usually 9 or 10 (Table 10); anal fin rays 16- wavy, sometimes breaking up into spots ventral-
21, usually 18 or 19 (Table 11); anal finlets 7-12, ly. Bars number 40-50 in adults but are usually
usually 9 or 10 (Table 11); pectoral fin rays 21-24, fewer than 20 in juveniles up to 450 mm FL.
usually 22 or 23 (Table 12). Precaudal vertebrae Munro reported the cranial regions and upper
19 or 20, usually 20 (Table 6); caudal verte- regions of the back to be mottled with iridescent
brae 23-27, usually 24 or 25 (Table 7); total blue and green. Cheeks, lower jaw, and belly
vertebrae 42-46, usually 44 or 45 (Table 8). Gill silvery white. First dorsal fin bright blue rapidly
rakers on first arch (0-2) + (1-8) = 1-8, usually fading to blackish blue. Pectoral fin light grey
(0-1)+ (3-4)= 3-5 (Table 5). Morphometric char- turning to blackish blue. Caudal fin lobes, second
acters given in Table 15. dorsal, anal, and dorsal and anal finlets pale
grayish white turning to dark gray. Juveniles
have the anterior membranes of the first dorsal
Size. -Maximum size 230 cm FL and 59 kg; jet black contrasting with pure white posteriorly
commonly 60-120 cm (Lewis 1981). The all-tackle (Munro 1943:pl. 8, fig. 3).
angling record is a 44.9 kg fish taken at Scott- There is an excellent illustration of an adult S.
burgh, Natal, South Africa, in 1982. Sexual ma- commerson from Japan in Kishinouye (1923:pl.
turity is attained at a length of 70-80 cm FL in 22), of an adult (968 mm FL, here reproduced as
Madagascar (Prado 1970), Papua New Guinea Figure 52), and a juvenile (368 mm) from Austra-
(Lewis et al. 1974), and Fiji (Lewis et al. 1983), lia in Munro (1943), and of an adult from India in
but not until 90-100 cm in South Africa (van der Jones and Silas (1962:fig. 2). There are color
Elst 1981). Females attain larger sizes than males paintings in La Monte (1952:pl. 19) and Grant
(Prado 1970; Lewis et al. 1974, 1983). (1982:627) and color photographs of a specimen

TABLE I5.-Summary of morphometric data of Scomberomorus commerson. FL = fork length, HL = head length.

Red Sea Indian Ocean East Indies Total

Character N Min. Max. Mean SD N Min. Max. Mean SD N Min. Max. Mean SD N Min. Max. Mean SD
Fork length 12 266 854 346 162 34 180 1.085 443 275 22 94 628 234 132 119 94 1,155 403 247
Snout-A %. FL 12 530 559 549 8 33 426 575 535 26 22 521 635 563 26 117 426 635 543 23
Snout-2D %. FL 12 491 525 506 8 34 405 538 501 22 22 500 545 522 13 118 405 625 509 20
Snout-1D %. FL 12 233 255 245 8 34 188 261 237 17 22 231 273 258 12 118 188 273 243 16
Snout-P, r. FL 12 247 269 259 6 33 201 321 256 21 22 229 299 269 18 117 201 321 257 18
Snout-P , %. FL 12 228 258 243 8 34 185 260 233 18 22 226 274 254 13 118 185 274 238 17
P,-P, %. FL 12 89 103 96 4 31 70 105 90 8 22 83 119 99 10 112 70 123 96 9
Head length %. FL 12 218 239 230 6 34 177 251 226 18 22 215 261 240 12 119 177 261 229 15
Max. body depth %. FL 12 174 202 188 9 32 141 215 177 15 22 165 234 193 19 115 141 235 187 18
Max. body width %. FL 12 76 99 87 6 31 74 118 95 10 22 64 142 85 17 109 64 142 93 12
P, length %. FL 11 118 148 124 8 33 103 137 124 7 22 86 135 108 11 114 86 153 122 12
P, length %. FL 12 47 67 57 5 33 46 71 56 6 22 38 127 61 16 115 38 127 56 10
P2 insertion-vent r. FL 12 264 285 275 7 33 207 386 270 27 22 269 320 288 14 117 207 386 273 19
P, tip-vent %. FL 12 203 233 220 10 33 170 238 210 15 22 209 278 231 17 115 170 278 217 16
Base 10 %. FL 12 248 276 261 7 33 210 286 257 14 22 243 285 265 11 117 210 286 261 13
Height 2D r. FL 12 98 125 108 7 31 91 119 104 8 21 76 108 91 9 102 76 135 103 11
Base 2D r. FL 12 87 123 104 14 33 81 130 99 11 22 78 171 106 18 117 78 171 104 14
Height anal %. FL 11 90 117 100 7 32 89 120 102 8 21 70 114 91 12 103 70 128 100 10
Base anal %. FL 12 89 108 98 6 33 81 113 97 8 22 80 119 98 10 116 80 164 100 11
Snout (fleshy) r. FL 12 80 91 88 3 33 66 99 88 7 22 84 100 93 5 117 66 100 89 6
Snout (bony) r. FL 12 72 82 77 3 33 61 136 82 12 22 77 91 84 4 117 61 136 81 7
Maxilla length %. FL 12 119 137 130 5 32 101 147 129 13 22 115 163 142 13 117 101 163 131 13
Postorbital %. FL 12 100 111 104 3 33 81 119 104 9 21 98 109 104 3 114 81 119 104 6
Orbital (fleshy) %. FL 12 28 38 34 3 32 21 47 33 6 22 28 50 40 6 115 21 50 34 6
Orbital (bony) r. FL 12 35 53 48 5 33 31 81 48 10 22 38 64 54 7 116 31 81 49 8
Interorbital %. FL 12 59 66 63 2 33 47 66 60 5 22 58 69 64 3 115 47 71 62 4
2D-caudal %. FL 12 458 520 474 20 33 380 540 491 31 21 420 489 454 18 109 380 540 480 27
Head length 12 61 186 79 34 34 41 214 96 50 22 24 135 55 28 119 24 242 89 48
Snout (fleshy) %. HL 12 351 402 381 14 33 326 408 388 15 22 356 412 389 12 117 326 424 390 15
Snout (bony) %. HL 12 324 372 338 14 33 333 571 362 39 22 329 371 349 10 117 316 571 354 25
Maxilla length %. HL 12 548 596 566 13 32 518 610 569 20 22 533 651 593 29 117 518 651 570 23
Postorbital r. HL .12 436 483 455 15 33 418 496 461 18 21 397 497 436 22 114 397 504 455 21
Orbit (fleshy) r. HL 12 128 167 147 10 32 111 189 144 17 22 129 200 166 18 115 101 224 147 21
Orbit (bony) %. HL 12 161 232 209 18 33 155 352 210 35 22 179 256 223 22 116 147 352 211 28
Interorbital %. HL 12 256 303 275 12 33 245 286 265 8 22 246 289 266 11 115 245 316 270 12

625
 FISHERY BULLETIN: VOL. 82, NO.4

from Kuwait in Kuronuma and Abe (1972:pl. 17), Merceron 1970). Other food items mentioned by
a Japanese specimen in Masuda et al. (1975:79), these authors include small carangids, Leiogna-
a Persian Gulf specimen in Randall et al. (1978: thus, squids such as Loligo, and penaeoid
212), a South African specimen in van der Elst shrimps. Feeding apparently takes place day and
(1981:274), a Queensland specimen in Grant night (Tongyai 1970).
(1982:pl. 325), and a 344 mm specimen from the
South China Sea in Kyushin et al. (1982:248). Interest to fisheries. - This species is taken
throughout its range by commercial, artisanal,
Biology. -Adults frequently undertake lengthy and recreational fisheries. Although it may be
seasonal longshore migrations (Lewis 1981). Mi- present the year round, e.g., in the coastal water
grations occur along the entire eastern coast of of Madras State (Chacko et al. 1967), fisheries are
Queensland (McPherson 1981). Tongyai (1970:fig. usually concentrated in some seasons, particu-
4) has mapped the migration route in the Gulf of larly those with the best weather conditions for
Thailand; from the Cambodian border in October fishing. Peak fishing seasons in' some areas are
to the northernmost part of the Gulf of Thailand as follows: Taiwan-spring (Kishinouye 1923);
in December to February, then south along the Great Barrier Reef-August to September (Grant
west coast of the gulf in April. At least some 1978); Cambodia-the dry season, October to
individuals are present year round in some areas, April (Merceron 1970); Gulf of Thailand-Octo-
e.g., Cambodia (Merceron 1970) and East Africa ber to May (Tongyai 1970); Waltair, northeastern
(Williams 1964). Spawning apparently occurs India-March-April, June-July, and December
over a long period in some regions, e.g., October (Venkata Subba Rao et al. 1981); Vizhingam,
to July in East Africa (Williams 1964), July to southeastern India-September to April (Gopa-
December in Papua New Guinea (Lewis et al. Ian Nayar 1958); and Malwan, south of Bombay
1974). Spawning times have been reported as -February to March and October to December
spring in Taiwan (Kishinouye 1923), October- (Kaikini 1961). There are important fisheries in
December on the Great Barrier Reef (Munro Fishing Areas 51, 57, and 71. The total catch
1942), October to February, peaking in December fluctuated between 63,290 and 79,047 t/yr in
and January in Fiji (Lewis et al. 1983), May 1979-82 (FAO 1984). The five countries with the
to July in the coastal waters of Madras State largest reported catch in this period were Indo-
(Chacko et al. 1967), and December-February in nesia, Philippines, Sri Lanka, Yemen, and Paki-
Madagascar (Fourmanoir and Crosnier 1964). stan. The landings in Queensland were around
Munro (1942) described and illustrated the devel- 1,000 tons/yr during the mid-1970's but have
opment of artificially fertilized eggs and early dropped to 730-770 tons in 1978-80 (McPherson
larvae from the Great Barrier Reef. Jones (1962) 1981). The 1982 catch in Fiji probably exceeded
described and illustrated five postlarvae and ju- 300 tons (Lewis et al. 1983). There is also an
veniles (14.4-54.4 mm) from Vizhingam along the important drift net fishery in India, but the catch
coast of southern Kerala taken in shore seines is not identified to species in the statistics. Drift
from February to June. The most complete larval nets (gill nets) that are usually fished over night
description is by Jenkins et al. (1984) of 62 larvae appear to be the most important gear used for
(3.5-9.3 mm SL) from the shelf waters of the S. commerson in Thailand, Malaysia, and India
Barrier Reef. Tongyai (1970) reported that juve- (Tongyai 1970; Pathansali 1968; Kaikini 1961;
niles 100-450 mm were taken in waters of high Chacko et al. 1967, respectively); other gear in-
turbidity and salinity in the Gulf of Thailand. cludes shore seines in Taiwan and India (Kishi-
Juveniles were caught with dip nets in Papua nouye 1923; Gopalan Nayar 1958), trolling lines
New Guinea waters in July, October, November, in Taiwan, Malaysia, India, and East Africa
and December (Lewis et al. 1974). Like other (Kishinouye 1923; Pathansali 1968; Dhawan et
species of the genus, S. commerson feeds primar- al. 1972; Williams 1964, respectively). Hand lines
ily on small fishes particularly anchovies such as (bett-tok) baited with mackerel (Rastrelliger)
Anchoviella and Stolephorus and clupeids such or squid (Loligo) and trotlines (bett-Iaak) with
as Sardinella (South Africa-van der Elst 1981; spoons are also employed in the Gulf of Thailand
Madagascar-Prado 1970; Madras-Chacko et (Tongyai 1970). It is taken fairly commonly in the
al. 1967; Waltair, east coast of India-Rao 1964; inshore fishery along the Mediterranean coast of
Vizhingam, southern India-Kumaran 1964; Israel with trammel nets and occasionally with
Gulf of Manaar-Chacko 1949; and Cambodia- purse seines (A. Ben-Tuvia 3 ). The yearly catch
626
COLLETIE and RUSSO: SPANISH MACKERELS

will be about 20 t out of the 2,000 t taken in Andamans and Nicobars (Jones et al. 1960), Lac-
the inshore fisheries according to A. Ben-Tuvia cadives (Nagabhushanam and Chandrasekhara
and D. Golani.4 It is a highly regarded species Rao 1972), Amirantes (Ommanney 1953), Com-
that commands a good price in the Philippine ores and Madagascar (Fourmanoir and Crosnier
Islands, Thailand, India, Madagascar, and East 1964), and Mauritius (Bleeker 1874; Baissac
Africa (Warfel 1950; Tongyai 1971b; Devanesen 1976). It has strayed into the South Atlantic
and Chidambaram 1953; Fourmanoir and Cros- because we have examined the head of a speci-
nier 1964; Williams 1964, respectively). It is a men (BMNH 1965.12.1.104) collected by Arthur
prime target of the Natal ski-boat fishermen and Loveridge from Egg Island, St. Helena. It has
is pursued by sport and commercial anglers in even traversed the Suez Canal and entered the
South Africa, using lures, feathers, clupeids, and eastern Mediterranean Sea where it is now
anchovies as bait (van der Elst 1981). It is mar- known from Lebanon (George and Athanassiou
keted fresh, on ice, or salted and dried (Gopalan 1965) and Israel (Collette 1970; USNM 226334;
Nayar 1958; Fourmanoir and Crosnier 1964; Wil- Golani and Kredo 1981).
liams 1964; Tongyai 1971b; McPherson 1981). A
lipid-soluble toxin similar to ciguatoxin has been Geographic variation. -Samples were adequate
found in the flesh of S. commerson between lat. to compare the morphometric data of three popu-
24 Sand 26 S along the east coast of Queensland
0 0
lations of S: commerson by ANCOVA (Table 15):
(Lewis and Endean 1983). From 1976 to 1980, at Red Sea (n = 12), Indian Ocean (n = 31-34), and
least 38 toxic S. commerson, resulting in 217 East Indies (n = 21-22). Null hypotheses that the
poisonings, came from this area. three sets of regressions are coincident were
accepted for 11 of 26 regressions, rejected for the
Distribution. -Widespread throughout the Indo- other 15. For one set, interorbital width, the
West Pacific from South Africa and the Red Sea regressions for all three populations differed sig-
east through the Indo-Australian Archipelago to nificantly in slope. The Red Sea population dif-
Australia and Fiji and north to Hong Kong, fers significantly from the Indian Ocean popula-
Formosa, and Japan (Fig. 51). The northernmost tion in six regressions: Sn-1D, Sn-PI , Ht 2D, Base
record is from the northern coast of Yamaguchi 2D, Ht A, and interorbital width. The Indian
Prefecture, southern Honshu, on the Sea of Japan Ocean population differs from the East Indies
(Kishinouye 1923:417). Its range extends farther population in eight regressions: P r P 2 , Hd L, P 2
out into the Pacific islands than any of the other tip-vent, Ht 2D, Sn (fleshy), Sn (bony), maxilla L,
species of Scomberomorus, throughout the Phil- and interorbital width.
ippine Islands, to New Caledonia (Chapman 1946; There are also geographic differences in meris-
Fourmanoir and Laboute 1976; Uchida 1978) and tic characters. Populations in the Red Sea and
Fiji (Jordan and Dickerson 1908; Whitley 1927; Persian Gulf tend to have fewer vertebrae (23-24
Fowler 1959). Records from Wallis and Futuna caudal, 43 total) and fewer rays in the second
Islands and Cook Islands (Uchida 1978) are doubt- dorsal and anal fins (usually 16-17 second dorsal
ful and need to be verified. In Australia the range and 17-18 anal) than other populations (24-27
extends south to Sydney (Castelnau 1879; AMS caudal, 44-46 total vertebrae; 17-18 second dorsal,
1.9693) and, rarely, even to Victoria and Tas- 18-19 anal rays). Populations in the East Indies
mania (Munro 1958a; Whitley 1964a) on the east '. and Gulf of Thailand tend to have more vertebrae
coast and to Rottnest Island off Perth, Western (25-27 caudal, 45-46 total) and anal finlets (mode
Australia (Hutchins 1979). From Australia and 10 rather than 9). Gill rakers tend to be fewer in
the East Indies, the range extends along the coast the East Indies, Gulf of Thailand, and South
of the Indian Ocean including the Persian Gulf China Sea (2-6, usually 3 or 4) compared with
and Red Sea to False Bay, Cape Town, South other populations (3-8, usually 4-6).
Africa (Barnard 1948). The range includes many
major offshore island groups in the Indian Ocean: Material examined. -Total 262 (94.2-1,155).

meas.: 120 (94.2-115): Israel (2); Red Sea (12);

3 A. Ben-Tuvia, Professor of Zoology, Zoology Department, The
Hebrew University, 91904 Jerusalem, Israel, pers. commun. Gulf of Aden (2); St. Helena (1); W
October 1982. Indian Ocean (14); Arabian Sea (14); Bay
4A. Ben-Tuvia, Professor of Zoology, and D. Golani, Zoology
Department, The Hebrew University, 91904 Jerusalem, Israel, of Bengal (5); Andaman Sea (7); Gulf of
pers. commun. October 1982. Thailand (14); East Indies (22, *C. ko-
627
 FISHERY BULLETIN: VOL. 82, NO.4

nam Bleeker), New Guinea (2); Austra- SL; from Gulf of California; measurements).
lia (5); Philippine Is. (6); South China Croker 1937:245-246 (Long Beach). Walford
Sea (8); Fiji (5). 1937:25-26 (description, occurrence). Roedel
counts: 262. 1939:341 (Long Beach; fifth record of recent
diss.: 14 (260-1,155): Israel (1); W Indian years). Munro 1943:69, 71-72 (placed in sub-
Ocean (2); Pakistan (1); New Guinea (2); genus Chriomitra). Fowler 1944:498 (listed;
New South Wales, Australia (2); Phil- Mexico; Panama Bay record probably S. sier-
ippine Islands (2); Hong Kong (4). ra). Fitch 1948:134 (Santa Monica Bay; sixth
specimen since 1880's). *Fitch and Flechsig
1949:275-280 (history of previous captures; de-
Scomberomorus concolor (Lockington) scription), fig. 75. Fraser-Brunner 1950:157-
Monterey Spanish Mackerel 158 (description), fig. 26. Clothier 1950:53
Figure 53 (47-48 vertebrae). Fitch 1950:70 (Newport
Harbor; seventh California record since 1880's;
Chriomitra concolor Lockington 1879a:134-136 comparison with S. sierra). Roedel 1951:510
(original description; Monterey Bay, Calif.). (Long Beach; 8th to 10th specimens since
Lockington 1879b:34 (uncommon; San Francis- 1880's; may have spots). Fitch 1952:560 (Los
co market). Angeles Harbor). Roedel 1953:85 (occasional
Scomberomorus concolor. Jordan and Gilbert in S California). Radovich 1961:21, 30 (years
1881a:456 (Monterey Bay; Chriomitra placed in of California captures). Collette et al. 1963:54
synonymy of Scomberomorus). Jordan and (compared with S. sierra; previous California
Jouy 1881:13 (specimens from Soquel, Calif.; records of S. sierra = S. concolor). Clemens
USNM 27205; distributed as duplicates). Jor- and Nowell 1963:260 (Gulf of California).
dan and Gilbert 1881b:45 (Monterey Bay). Jor- Fitch and Craig 1964:202, fig. 5 (otolith).
dan and Gilbert 1882:425-426 (description). Klawe 1966:445 (compared with S. sierra; more
Meek and Newland 1884:232-233 (synonymy, gill rakers on upper and lower arches). Fitch
description). Goode 1884:316 (Soquel, Monte- 1969:65 (jaw fragments and teeth; Chumash
rey Bay; occurrence, price). Dresslar and Fes- Indian village archaeological site; Ventura,
ler 1889:442-443 (synonymy, description). J or- Calif.). Castro-Aguirre et al. 1970:156-157
dan and Evermann 1896a:341 (listed). Jordan (abundant in Gulf of California). Fitch and
and Evermann 1896b:873-874 (description, syn- Lavenberg 1971:131, 168 (listed). Miller and
onymy). Jordan and Evermann 1902:284 (de- Lea 1972:192 (description; range Gulf of Cali-
scription). Starks 1918:121 (not reported from fornia to Soquel, Calif.), fig. Buen 1972:291
Monterey Bay in 40 yr). Meek and Hildebrand (Mexico). Bullis et al. 1972:75 (bionumeric
1923:325-326 (description; Soquel, Calif.). Jor- code number). Richards and Klawe 1972:13
dan et al. 1930:257 (listed). Phillips 1932:99 (range), 91 (references to juveniles). Magnu-
(Monterey Bay; first record in more than 40 yr). son 1973:350 (short pectoral fin). Sharp 1973:
Breder 1936:12 (2 specimens, 491-520 mm 384, fig. 3 (hemoglobin electrophoretic patterns

FIGURE 53.-Scomberomorus concolor. Gulf of California, 440 mm FL, USNM 233681.

628
COLLETTE and RUSSO: SPANISH MACKERELS

of S. sierra, S. concolor, and Acanthocybium usually 28 (Table 7); total vertebrae 46-48, usu-
identical or very similar). Johnson 1975:20 ally 47 or 48 (Table 8). Gill rakers on first arch (4-
(procurrent spur not present). Shiino 1976: 8) + (15-21)= 21-27, usually (6-7) + (17-18) = 23-25
231 (common name). Thomson and McKibbin (Table 5). Morphometric characters given in Ta-
1976:46 (description; Gulf of California). ble 16.
Klawe 1977:2 (common name, range). Fitch
and Schultz 1978:85, fig. 4G (otolith). Horn
TABLE 16.-Summary of morphometric data of Scom-
and Allen 1978:39 (range lat. 36°N to 32°N
beromorus concolor. FL = fork length, HL = head
along California coast). Collette 1979:29 length.
(characters, range). Collette and Russo 1979: Character N Min. Max. Mean SO
13 (diagnostic characters, range). Cressey et Fork length 34 134 685 401 139
al. 1983:264 (host-parasite list, 3 copepod spe- Snout-A %, FL 34 504 547 524 10
cies). Collette and Nauen 1983:64-65 (descrip- Snout-20 r. FL 34 488 534 506 12
Snout-lO %, FL 34 220 258 236 10
tion, range), fig. Snout-P, %, FL 34 192 287 242 18
Snout-P, %, FL 34 124 263 209 21
P,-P, %, FL 30 89 116 100 6
Types. -Chriomitra concolor Lockington 1879a. Head length r. FL 34 185 230 202 10
Max. body depth %, FL 29 164 220 187 16
Description based on a 21-in FL (533 mm FL) Max. body width %, FL 32 70 111 89 10
specimen obtained in the San Francisco market P, length %, FL 34 116 137 125 6
P,length r. FL 31 41 61 50 4
and probably originating in Monterey Bay. Lock- P2 insertion-vent %, FL 32 222 294 261 16
ington stated that the specimen was "in the P, tip-vent r. FL 29 173 248 212 18
Base 10 r. FL 34 210 292 254 15
possession of the Cal. Acad. of Sciences", but it is Height 20 r. FL 26 98 129 111 9
Base 20 %, FL 33 108 153 128 10
not now present in the CAS collection. Data from Height anal r. FL 25 26 137 108 20
the original description are "D XV + 17 + VII; A Base anal %, FL 33 107 171 135 14
Snout (fleshy) r. FL 33 53 83 72 5
18 + VIII. Body color dark steel blue above, be- Snout (bony) %, FL 33 57 76 64 3
coming silvery below; no streaks". Maxilla length %, FL 33 103 128 113 6
Postorbital %, FL 29 85 101 96 4
Orbital (fleshy) r. FL 33 25 47 32 7
Orbital (bony) %, FL 30 34 63 46 8
Diagnosis. -The species of Scomberomorus with Interorbital r. FL 34 44 55 49 3
the most gill rakers, a total of 21-27 on the first 20-caudal r. FL 30 413 524 484 24
Head length 34 31 134 80 27
arch, compared with 1-18 in the other 17 species. Snout (fleshy) %, HL 33 276 389 353 19
It possesses nasal denticles as do the other five Snout (bony) r. HL 33 273 355 314 16
Maxilla length %, HL 33 532 575 555 11
species of the regalis group (brasiliensis, macu- Postorbital r. HL 29 419 524 475 22
Orbit (fleshy) %, HL 33 125 212 158 26
latus, regalis, sierra, and tritor). Like S. macu- Orbit (bony) %, HL 30 175 285 226 32
latus, S. concolor lacks the artery that goes from Interorbital %, HL 34 220 274 242 12
the fourth right epibranchial artery to the coeli-
aco-mesenteric artery (Fig. 7d), but it has the
artery that comes off the fourth left epibranchial Size. -Maximum size 76.2 cm FL, 2.3-3.6 kg
artery as do all the species in the group except S. (Goode 1884).
tritor. Together with three other species of the
regalis group (brasiliensis, regalis, and sierra), Color pattern. -According to Walford (1937),
S. concolor has a long posterior process on the males are steel blue on the back, silvery on the
pelvic girdle, 62-90% of the length ofthe anterior sides and below, and are without streaks or spots..
plate. Intercalar spine absent as in the other five Females are darker, with two alternate series of
species of the regalis group and S. niphonius. brown spots on sides. The spots on the sides of the
females are gold in life (Fitch and Flechsig 1949).
Description. -Lateral line gradually descending A black and white photograph of S. concolor is
to midline on caudal peduncle. Intestine with two included in Fitch and Flechsig (1949:fig. 75).
folds and three limbs (Fig. 3d). Spines in first
dorsal fin 15-18, usually 17 (Table 9); second Biology. -Little is known about the biology of S.
dorsal fin rays 16-20, usually 18 or 19 (Table 10); concolor. In the 1880's, they appeared in Monte-
dorsal finlets 6-9, usually 8 (Table 10); anal fin rey Bay in September and disappeared in Novem-
rays 19-23, usually 20 (Table 11); anal finlets 6-8, ber (Goode 1884). There are no references to eggs,
usually 7 or 8 (Table 11); pectoral fin rays 19-22, larvae, or juveniles (Richards and Klawe 1972).
usually 21 (Table 12). Precaudal vertebrae 18-20,
usually 19 (Table 6); caudal vertebrae 27-29, Interest to fisheries. -Some accounts indicate
629
 FISHERY BULLETIN: VOL. 82, NO.4

that S. concalor was of considerable commercial description based on the wingeram of Russell
importance in Monterey Bay in the 1870's and 1803:pI. 134).
1880's, in great demand and at a high price, 30-50 Cybium guttatum. Cuvier 1829:200 (listed in
cents a pound according to Goode (1884). Accord- footnote from Sc. guttatus Bloch and Schnei-
ing to other authors, such as Lockington (1879a, der). Cuvier in Cuvier and Valenciennes
b), it was not abundant even then. No longer of 1831:173-176 (description). Richardson 1846:
any commercial significance. 268 (synonymy, range). Cantor 1849:1093-
1095 (synonymy, description, range; Pinang).
Distribution. -An eastern Pacific endemic origi- Bleeker 1852:38, 39 (synonymy, description;
nally described from Monterey Bay, Calif. (Lock- East Indies). Bleeker 1853:42 (India). Bleek-
ington 1897a). This apparently was the northern er 1860:13 (Borneo). Gunther 1860:371 (syn-
limit of the range, and there have been only onymy, description). Bleeker 1861a:52 (Singa-
about 10 recent records from the California coast pore; listed). Bleeker 1861b:74 (Pinang; listed).
(Long Beach, Santa Monica Bay, Newport Har- Kner 1865:143-144 (description). Day 1873:
bor; Fitch and Flechsig 1949; Radovich 1961). Its 225 (description, range). Bleeker 1873:131
present range is concentrated in the Gulf of (China; listed). Day 1878:255 (synonymy, de-
California (Castro-Aguirre et aI. 1970; Miller and scription, range), pI. 55, fig. 1 (young), pI.
Lea 1972; Collette and Russo 1979:13, fig. 8). 56, fig. 4 (adult). Tirant 1885:46 (Cambodia;
listed). Kishinouye 1923:419-420 (description,
Material examined. - Total 34 (134-685 mm FL). anatomy), pI. 34, fig. 61 (adult). Chabanaud
1926:22 (Cote d'Annam, Tonkin; listed). Har-
meas.: 34 (134-685): Soquel, Calif. (6); Gulf of denberg 1931:141 (Sumatra). Delsman 1931:
California (27). 402 (vertebrae 20 + 25 = 45), figs. 1-9 (eggs
counts: 30. and larvae). Morice 1953:37 (villiform tongue
diss.: 6 (420-495): Gulf of Calif. teeth present). Gopalan Nayar 1958:49-51
(fishery; Vizhingam, S India).
Cybium interruptum Cuvier in Cuvier and Va-
S comberomorus guttatus lenciennes 1831:172-173 (original description;
(Bloch and Schneider) Pondichery, India). Gunther 1860:371 (de-
Indo- Pacific King Mackerel scription after Cuvier). Day 1873:225 (de-
Figure 54 scription, range). Day 1878:254-255 (syn-
onymy, description), pI. 56, fig. 3.
Scomber guttatus Bloch and Schneider 1801:23- Cybium Kuhlii Cuvier in Cuvier and Valencien-
24 (original description; Tranquebar, India), nes 1831:178-179 (original description; Bom-
pI. 5. bay). Hardenberg 1931:140 (often found in
Scomber wingeram Russell 1803:26-27 (descrip- river mouths; Sumatra). Delsman 1931:402
tion; Corornandel coast of India), pI. 134. (vertebrae 20+ 25= 45), 407 (commonest spe-
Scomber leopardus Shaw 1803:591-592 (original cies of Cybium at Bagan Si Api Api).

FIGURE 54.-Scomberomorus guttatus. Gulf of Thailand, 459 mm FL, CAS GVF Reg. 1512.

630
COLLETTE and RUSSO: SPANISH MACKERELS

Cybium Croockewitii Bleeker 1851b:161 (original inate in food of juveniles and adults; Waltair
description; Banka). Bleeker 1852:37-38 (de- coast, India). Gorbunova 1965a:52-53 (spawn-
scription). Gunther 1860:372 (description ing). Gorbunova 1965b:174-175 (spawning;
after Bleeker). Gulf of Tonkin), fig. 6 (4.3 and 5.8 mm larvae).
Scomberomorus guttatus. Fowler 1905:766 (Su- Menon 1966:396 (Tranquebar, India). Thiem-
matra; ANSP 27490-91). Jordan and Richard- medh 1966:129, 140 (Thai names). Tongyai
son 1909:177 (Formosa; FMNH 59284). Reeves 1966a:7-13 (synonymy, occurrence in Thailand,
1927:8 (Swatow, China). Fowler 1928:109 biology), pi. 2C. Tongyai 1966b:3-17 (length
(Bombay). Chevey 1934:45-46 (Tirant's C. frequency; Andaman Sea). Collette 1966:368-
guttatum = S. guttatus). Delsman and Har- 369 (Cybium kuhlii a junior synonym of S.
denberg 1934:341-342 (description; East In- guttatus, lectotype of C. kuhlii s~lected). Jones
dies), fig. 247 (adult), fig. 248 (larva, with 1968:998 (seerfish fishery; India). Pathansali
myomeres 15+ 35= 50). Hardenberg 1934:. 1968:1001-1002 (fishery on east and west coasts
311 (Sumatra; listed). Hardenberg 1936:252 of Malaya). Tongyai 1970:559 (distribution;
(mouth of Kapuas R., Borneo). Hardenberg Thai waters), 561 (spawning), 561-562 (food).
1937:12 (mouth of Kumai R., Borneo). Herre Merceron 1970:75-81 (length-weight; Cambo-
and Myers 1937:21 (Singapore). Munro 1943: dia). Tongyai 1971a:13-16 (description), pi. I
68, 71 (placed in subgenus Indocybium). (viscera), pi. IV (photograph). Tongyai 1971b:
Quraishi 1945:28 (pyloric caeca arranged in 3 (undetermined economic potential; Thailand),
dendritic pattern). Norman and Fraser 1949: pi. 8,13 (photographs). Latiff1971:92 (descrip-
153 (Indo-Pacific species). Fraser-Brunner tion; Penang waters; photograph). Banerjee
1950:160 (synonymy in part, range), fig. 31. and Chakrubarty 1972 (drift gill netting; Lower
Tham 1950:21 (feeds largely on Stolephorus). Sundarbans, W Bengal). Fernando 1972:524,
de Beaufort 1951:232-234 (synonymy, descrip- 530 (incidental catches in trawls; Wadge Bank,
tion, range). Tham 1953:49 (Singapore Straits), Ceylon). Kuronuma and Abe 1972:105 (de-
50 (correlation of catch with physical factors scription; Kuwait), color pi. 17. Richards and
and presence of food fishes such as Stolepho- Klawe 1972:13-14 (range), 91-92 (references to
rus). Vijayaraghavan 1955:360-372 (com- eggs, larvae, and juveniles). Magnuson 1973:
monest species of genus in Madras; eggs, larval 350 (small pectoral fin). Banerji 1973:129-130
development). Krishnamoorthi 1957:236 (sec- (seerfish fishery; India). Orsi 1974:175 (listed;
ond in importance among fishes landed at Ra- Vietnam). Roy and Roy 1974:44, 51, 53 (a
meswaram I., Palk Bay), 239-242 (catch), 251 principal species in gill net fishery; Balashore,
(value of catch). Krishnamoorthi 1958:270- India). Shenoy and James 1974 (ice storage).
281 (spawning season and fisheries; Rame- Devaraj 1976:80-85 (distinguished from S.
swaram I., SE India). Venkataraman 1961: koreanus), fig. 4 (vertebrae), 5 (preopercle and
287, fig. 4C (food of 133 specimens; Calicut, liver). Rao 1976:63-78 (biometric compari-
India; mostly teleosts). Kaikini 1961:361 son of 5 Indian populations). Shiino 1976:
(seerfish fishery; Malwan, India). Jones and 231 (common names). *Devaraj 1977 (oste-
Silas 1962:195-197 (synonymy, description, ology). Rao and Ganapati 1977:107-111 (com-
range), fig. 3 (533 mm adult), fig. 5D (head, not parison with postlarvae and juveniles of S.
5C as labelled), fig. 6C (gill arch), fig. 7C lineolatus and S. commerson). Klawe 1977:2
(caudal peduncle keels). Jones 1962:107-113 (common name, range). Randall et ai. 1978:
(development, 14.8-239 mm), figs. 2-6 (speci- 167 (Persian Gulf; photograph). Collette 1979:
mens 14.8, 22.9, 41.2, 66.8, and 239 mm long). 29 (characters, range). Collette and Russo
Misra 1962:295-296 (description, distribution), 1979:13 (diagnostic characters, range). Zhang
fig. 181 (size given as "1828 mm"). Jones and and Zhang 1981:104 (range in part). Naka-
Kumaran 1964:344-346 (larval development), mura and Nakamura 1982:446 (3 specimens,
figs. 1-3 (from Jones 1962). Kumaran 1964: description; Sea of Japan), fig. 1B. * Devaraj
587-589 (postlarval and juvenile fishes form 1982 (age and growth). Sivasubramaniam
most of diet of juveniles; W coast of India). and Mohamed 1982:64 (Qatar, Persian Gulf).
Blanc and Bauchot 1964:449 (specimens exam- Cressey et al. 1983:264 (host-parasite list,
ined by Cuvier). Boeseman 1964:468 (syn- 4 copepod species). Lee and Yang 1983:230-
types of C. kuhlii), pi. V, fig. 18 (photograph of 231 (Taiwan), fig. 21 (353 mm FL). Collette
syntype). Rao 1964:592-597 (teleosts predom- and Nauen 1983:65-66 (description, range), fig.
631
 FISHERY BULLETIN: VOL. 82, NO.4

Scomberomorus guttatum. Malpas 1926:72-74 Collette (1966:368); D XVII + 21 + VIII; A 21 +

(87 specimens; length, weight, gonads, stomach VIII; PI 22; RGR I 2 + 1 + 9 = 12; vertebrae 21 + 30
contents; Ceylon). Scott 1959:113 (description; = 51. A photograph ofthe lectotype was published
Malaya), photograph. by Blanc and Bauchot (1964:pI. 3, fig. 14, upper
Scomberomorus kuhlii. Chevey 1934:20 (Ti- fish). Paralectotypes: RMNH 1239 (1, 190 mm
rant's C. kuhlii = S. kuhlii). Hardenberg FL) and 1241 (1, 108 mm FL); Java; Kuhl and van
1934:311 (listed; Sumatra). Hardenberg 1936: Hasselt; and MNHN A.5715 (1, 115 mm FL);
252 (mouth of Kapuas R., Borneo). Harden- Bombay; Dussumier. Photographs of RMNH 1239
berg 1937:12 (mouth of Kumai R., Borneo). and MNHN A. 5715 have been published by Boese-
Munro 1943:68, 71 (placed in subgenus Pseudo- man (1964:pI. 5, fig. 19) and Blanc and Bauchot
sawara). Herre and Herald 1951:339 (Sanda- (1964:pI. 3, fig. 14, lower fish), respectively.
kan market; N Borneo). Bauchot and Blanc Cybium Croockewitii Bleeker 1851. Holotype:
1961:372 (types of C. kuhlii; recognized as valid RMNH 6054; Banka, Strait near Muntok, East
species). Blanc and Bauchot 1964:447 (types Indies (= Indonesia); J. H. Croockewit; D XV + 24
of C. kuhlii), pI. III, fig. 14 (photograph of + VII; A 23 + VII; PI 21-21; RGR 1 2 + 1 + 9 = 12;
type-specimens). Orsi 1974:175 (listed; Viet- lateral line with fine branches anteriorly. A pho-
nam). tograph of the type was published by Boeseman
Scomberomorus croockewiti. de Beaufort 1951: (1964:pI. 5, fig. 18).
234-235 (description), fig. 40 (drawing made for
Bleeker). Boeseman 1964:467 (holotype), pI. Diagnosis. -This species shares with S. ko-
V, fig. 17 (photograph ofholotype). reanus the presence of numerous fine auxiliary
Indocybium guttatum. Munro 1955:221 (descrip- branches from the anterior part of the lateral line
tion; Ceylon), fig. 652. Chacko et aI. 1967: (Fig. 54). It differs from S. koreanus in having the
1006 (fishery; Madras). usual two loops and three limbs to the intestine
Scomberomorus lineolatus. Not of Cuvier, 1831. instead of four loops and five limbs. Anterior end
Bauchot and Blanc 1961:371 (type of C. inter- of premaxilla forms a blunt rather than inter-
ruptum). Blanc and Bauchot 1964:446-447 mediate or acute angle. Ascending process of
(type of C. interruptum), pI. III, fig. 13 (photo- premaxilla short as in S. caualla. Scapular fora-
graph ofholotype of C. interruptum). men small as in S. koreanus and S. niphonius.
Scomberomorus guttatus guttatus. Jones and Supraoccipital crest high as in S. koreanus and
Silas 1964:62-63 (synonymy, description, S. multiradiatus.
range), pI. VII, fig. B. Silas 1964:325-329
(synonymy, description, range; C. koreanum Description. -Lateral line gradually descending
considered a subspecies of S. guttatus). to midline on caudal peduncle. Intestine with two
folds and three limbs (Fig. 2e). Spines in first
Types of nominal species. -Scomber guttatus dorsal fin 15-18, usually 16 or 17 (Table 9); second
Bloch and Schneider 1801. The original descrip- dorsal fin rays 18-24, usually 20-22 (Table 10);
tion was based on a specimen from Tranquebar, dorsal finlets 7-10, usually 8 or 9 (Table 10); anal
India. No ~ypes are known to be extant. The fin rays 19-23, usually 20-22 (Table 11); anal
figure in the original description leaves little finlets 7-10, usually 8 (Table 11); pectoral fin rays
doubt as to the identity of the name. 20-23, usually 21 (Table 12). Precaudal vertebrae
Scomber leopardus Shaw 1803 was based on 19-22, usually 21 (Table 6); caudal vertebrae 28-
the "wingeram" of Russell (1803:pI. 134); no types 31, usually 29 or 30 (Table 7); total vertebrae 47-
are extant. 52, usually 50 or 51 (Table 8). Gill rakers on first
Cybium interruptum Cuvier in Cuvier and arch (1-2) + (7-12) = 8-14, usually 2 + (9-10) = 11-12
Valenciennes 1831. Holotype: MNHN A.5522; (Table 5). Morphometric characters given in Ta-
Pondichery, India; Leschenault; 375 mm FL; D ble 17.
? +? + IX; A ? + VII; lateral line branched anteri-
orly; dried, dorsal fin badly damaged. A photo- Size. -Maximum size 76 em FL. Size at first
graph of the type was published by Blanc and maturity 48-52 em TL in southern India (Krish-
Bauchot (1964:pI. 3, fig. 13). namoorthi 1958), 41-45 em TL in Thailand (Ton-
Cybium kuhlii Cuvier in Cuvier and Valen- gyai 1966b).
ciennes 1831. Lectotype: MNHN A.5771; Java;
Kuhl and van Hasselt; 108 mm FL; selected by Color pattern. -Nakamura and Nakamura (1982)
632
 ("J
0
t-<
t-<
l'J
::ll'J
":"-:l
:xl
c::
rn
rn
9
rn
FL = fork length, HL = head length.
~
TABLE 17.-Summary of morphometric data of Scomberomorus guttatus.
Indian Ocean East Indies Gulf of Thailand China Total OJ
~
Character N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO· N Min. Max. Mean SO is:
Fork iength 22 109 545 309 144 24 107 515 242 100 17 173 459 331 84 31 63 760 291 167 143 63 760 317 128 f.;
524 494 468 517 138 468 516 :>1
Snout-A %. FL 20 495 568 519 19 24 503 559 14 17 518 505 7 28 568 20 568 17 l'J
Snout-2D r. FL 22 452 509 483 14 24 454 504 481 12 17 459 494 479 9 28 463 510 481 10 140 451 512 481 11 ~
Snout-1D %. FL 22 224 273 242 12 22 225 260 241 10 17 219 248 232 9 29 220 275 238 13 123 219 275 238 11 f;.;
Snout-P, %. FL 19 228 285 253 17 22 231 278 255 12 17 225 358 248 31 26 225 296 248 20 116 224 358 250 19
Snout-PI %. FL 22 193 241 213 13 23 191 235 212 12 17 190 225 201 8 29 186 245 207 19 124 185 245 208 14
P,-P, %. FL 19 94 126 109 8 22 93 119 107 7 17 92 120 104 7 25 95 129 106 9 114 89 129 106 8
Head length r. FL 22 188 234 211 13 24 188 221 208 9 17 190 218 199 8 31 185 259 207 19 143 185 259 205 13
Max. body depth roo FL 19 171 326 218 30 19 190 229 208 10 16 99 226 202 29 30 191 236 209 9 129 99 326 209 18
Max. body width %. FL 15 69 125 92 16 15 69 106 88 10 16 85 104 93 6 22 71 106 91 8 92 69 146 93 12
P, length %. FL 17 92 130 113 9 20 89 130 107 11 11 93 122 107 7 23 89 133 108 9 99 89 133 109 9
P, Ien9th r. FL 13 47 70 63 6 16 47 69 59 5 13 47 62 57 4 20 48 74 59 5 88 41 74 59 6
P2 insertion~vent %. FL 14 224 270 250 16 22 240 279 257 11 15 231 268 245 12 23 218 269 248 13 101 218 288 251 13
P, tip-vent roo FL 10 158 208 187 17 14 178 223 196 13 12 174 209 187 12 22 167 211 189 13 81 158 229 191 15
Base 10 %. FL 22 203 253 232 13 21 210 258 234 13 17 208 256 240 12 29 216 246 232 8 122 203 270 234 12
Height 20 roo FL 18 113 153 138 11 20 103 167 124 16 10 114 169 132 15 22 117 155 132 10 113 103 254 131 17
Base 20 %. FL 22 125 161 146 11 23 121 156 136 10 17 113 157 136 13 29 100 170 144 13 124 100 170 141 12
Height anal %. FL 15 109 165 135 13 16 93 166 122 18 13 107 170 127 15 24 109 153 125 11 106 93 170 127 14
Base anal r. FL 22 115 165 138 13 23 83 154 127 13 17 111 147 130 10 29 116 146 134 7 124 83 171 133 12
Snout (fleshy) r. FL 22 60 81 72 5 24 64 80 72 3 17 61 75 69 4 29 61 170 75 19 125 60 170 72 10
Snout (bony) %. FL 22 56 73 64 4 24 57 69 64 3 17 54 68 61 4 29 48 161 66 19 125 47 161 63 10
Maxilla length roo FL 22 92 130 110 9 24 86 121 111 9 17 90 118 102 9 28 95 132 108 11 124 86 132 108 9
Postorbital %. FL 22 85 112 97 7 24 87 100 95 4 17 79 105 93 5 29 88 112 95 6 123 79 112 95 5
Orbital (fleshy) %. FL 22 25 50 36 7 24 28 56 40 6 17 29 41 34 3 29 23 52 35 8 125 23 56 36 7
Orbital (bony) %. FL 22 37 70 53 10 24 47 69 57 6 17 41 60 48 5 29 38 71 52 8 124 37 71 52 8
Interorbital %. FL 22 55 66 60 3 24 55 62 59 2 17 52 62 57 3 29 53 63 58 3 124 52 66 58 3
20-caudal r. FL 22 481 552 519 23 23 491 558 534 14 17 480 553 507 23 28 465 579 528 34 138 465 579 527 26
Head length 22 25 125 64 29 24 24 98 50 18 17 38 88 65 15 31 16 145 58 30 143 16 145 64 24
Snout (fleshy) %. HL 22 296 383 344 21 24 319 368 344 12 17 319 365 345 12 29 310 734 364 72 '125 296 734 351 38
Snout (bony) roo HL 22 265 328 303 15 24 281 334 306 12 17 282 320 304 12 29 245 694 320 74 125 232 694 309 38
Maxilla length %. HL 22 455 564 523 23 24 451 564 530 26 17 463 550 512 30 28 497 565 531 17 124 451 565 525 23
Postorbital roo HL 22 412 499 463 20 24 410 498 457 20 17 413 502 469 25 29 393 563 470 34 123 382 563 465 26
Orbit (fleshy) %. HL 22 118 217 173 30 24 148 253 193 24 17 151 198 168 13 29 121 220 169 26 125 118 253 174 27
Orbit (bony) %. HL 22 158 306 253 41 24 240 314 275 19 17 214 275 242 18 29 197 304 252 26 124 158 314 252 30
Interorbital roo HL 22 244 325 284 16 24 258 298 284 8 17 272 296 284 8 29 254 301 284 13 124 244 325 285 12
 FISHERY BULLETIN: VOL. 82, NO.4

described fresh specimens taken in Wakasa Bay Thailand might be deduced from seasonal changes
in the Sea of Japan. Body greyish blue dorsally, in peak fishing months along the coast of Thai-
silvery white laterally and ventrally. Several land. These peaks are November-December in
longitudinal rows of small brownish spots scat- eastern Thailand, late December-January in the
tered rather densely along lateral median line. northern part of the Gulf, and January-March in
First dorsal fin membrane black. Pectoral, second the western part of the Gulf (Tongyai 1970).
dorsal, and caudal fins dark brown. Pelvic and Based on occurrence of ripe females and size of
anal fins silvery white. maturing eggs, spawning probably occurs from
There are good illustrations of a specimen from April to July around Rameswaram Island be-
the North Pacific in Kishinouye (1923:fig. 61) and tween India and Sri Lanka (Krishnamoorthi
of one from India in Jones and Silas (1962:fig. 3). 1958). Ripe females 32.5-46.5 cm FL were taken
There are photographs of specimens of S. gut- in Thai waters in May. Larvae and juveniles have
tatus from India in Jones and Silas (1964:pI. 7) been reported from Indonesian and Indian waters
and Silas (1964:pI. 2), and there is a good photo- but apparently the only certain accounts are
graph of a specimen from the Sea of Japan in those of Jones (1962) and Jones and Kumaran
Nakamura and Nakamura (1982:fig. lB). A good (1964) who illustrated four postlarvae (14.8, 22.9,
color photograph of a specimen from the Persian 41.2, and 66.8 mm). As with other species of
Gulf is included in Kuronuma and Abe (1972: Scomberomorus, the food is primarily fishes. Ju-
pI. 17). veniles in India feed mainly on teleosts, particu-
larly clupeoids such as Anchoviella (Venkata-
Biology. -Little is reported in the literature raman 1961; Kumaran 1964; Rao 1964). Adults
about movements and migration of S. guttatus also feed mainly on teleosts with small quantities
but it appears to be less migratory than S. of crustaceans and squids (Thailand - Tongyai
commerson. Possible movements in the Gulf of 1970, India-Rao 1964). Anchovies are particu-

90

S. koreanus

--+-+-rl'--+---+-+---+--f-"i,--+--I20

o' o

~_hrt- -'h--+-+--+-"-420

\:" r.
S. queenslandicus FIGURE 55.-Ranges of five
Indo-West Pacific species of
Scomberomorus: S. guttatus,
S. koreanus, S. semifascia-
tus, S. queenslandicus, and
60' 90' 150· 180· 150. 6 S. multiradiatus.

634
COLLETTE and RUSSO: SPANISH MACKERELS

larly important: Stolephorus in Singapore Straits Amoy (BMNH 1860.7.20.110), and Swatow, China
(Tham 1950, 1953) and Anchoviella in Waltair, (Reeves 1927). There are many records and speci-
India (Rao 1964). mens from Indochina, the Gulf of Thailand, and
the East Indies. There are records and specimens
Interest to fisheries. -There are commercial or of S. guttatus from Borneo (Bleeker 1860; Har-
artisanal fisheries for S. guttatus in Cambodia denberg 1936; Herre and Herald 1951; ANSP
(Merceron 1970), Thailand (Tongyai 1971b), Ma- 72282) and Makassar, Celebes (RMNH 24096).
laysia (Pathansali 1968), and India, particularly The range extends further out in the East Indies
in the lower Sundarbans, West Bengal (Banerjee than that of either S.lineolatus or S. koreanus, at
and Chakrubarty 1972), the Ball;l.shore coast (Roy least to Bali. The report of S. guttatus from
and Roy 1974), around Madras (Vijayaraghavan Western Australia (McKay 1970) is based on a
1955), the Gulf of Mannar-Palk Bay area (Krish- specimen (HUMZ F-423) of S. queenslandicus.
namoorthi 1957), and Malwan, south of Bombay Earlier reports of S. guttatus from Australia
(Kaikini 1961). It is caught all year round in some (Macleay 1881; Stead 1906, 1908; Rendahl 1923)
areas (Cambodia-Merceron 1970; Ramaswaram are also based on S. queenslandicus (Munro 1943:
1., India-Krishnamoorthi 1957) but there are 86). Early reports from New Zealand are based on
peaks of abundance that differ from region to "a damaged specimen of a Cybium, probably
region. It is taken in the non-monsoon months C. guttatum, was obtained at the Chatham
(September-May) along the Balashore coast south Islands...." (Hutton 1895). This report has led to
of Calcutta with the catch increasing from Octo- subsequent records (Hutton 19Q4; Phillipps 1927;
ber to February (Roy and Roy 1974). Catches peak Whitley 1968). We concur with Whitley's conclu-
in September-October, December-February, and sion that this record is "very doubtful". The range
May in Waltair near Vishakhapatnam further extends west into the Persian Gulf (Kuronuma
south along the west coast of the Bay of Bengal and Abe 1972; ZMK 3-4).
(Venkata Subba Rao et al. 1981). The season
extends from September-October to March-April Geographic variation. -Morphometric data for
in Vizhingam, southern India, (Gopalan Nayar five populations of S. guttatus were compared
1958) with the peak catches usually in September with ANCOVA (Table 17): Arabian Sea (n = 7-
or October. It is one ofthe principal species in the 13), Bay of Bengal (n = 5-9), East Indies (n = 14-
drift net seerfish fishery in India, but the catch 24), Gulf of Thailand (n = 10-17), and China (n =
is not identified to species in the statistics. Indo- 22-31). Null hypotheses that the 5 sets of regres-
nesia reported the only catch identified as S. sion lines are coincident were accepted for 18 sets,
guttatus (4,254-5,249 t/yr) in 1979-82 (FAO 1984). rejected for 8 sets: Sn-lD, Sn-P1 , Head L, maxi-
The primary gear in most areas appears to be the mum depth, maxilla L, orbit (fleshy), interorbit,
drift gill net which is set overnight, but it is also and 2D-C. The five populations were arranged
taken in bamboo stake traps and with hand lines geographically from west to east as listed above.
in Thailand (Tongyai 1970) and by trolling or No significant differences were found between
with hook and line in India and Malaysia (Rao populations in the Arabian Sea and Bay of Ben-
1964; Jones 1968; Pathansali 1968). It is utilized gal, but there were significant differences be-
fresh or salted in most areas (India-Jones 1967; tween all other adjacent populations: Bay of Ben-
Cambodia-Merceron 1970; Thailand-Tongyai gal vs. East Indies (Sn-P1 ), East Indies vs. Gulf of
1971b). It can be stored on ice for 10-13 d (Shenoy Thailand (Interorbital), Gulf of Thailand vs. Chi-
and James 1974). Although less abundant than na (Sn-lD, interorbital, and 2D-C). The Arabian
the Indian mackerels (Rastrelliger spp.), it is Sea and Bay of Bengal populations were com-
highly esteemed for food and commands a high- bined, the regressions rerun, and compared with
er price in Thailand and India (Tongyai 1966a; the other three populations with ANCOVA. Null
Pathansali 1968). hypotheses that the 4 sets of regression lines are
coincident were accepted for 15 sets, rejected for
Distribution. -Indo-West Pacific from Taiwan to 11 sets: Sn-lD, Sn-P1 , Head L, maximum body
the Gulf of Thailand, Java, and Sumatra west depth, Base ID, Base 2D, Base A, maxilla L, orbit
around the Bay of Bengal and Arabian Sea into (fleshy), interorbital, and 2D-C. The Newman-
the Persian Gulf (Fig. 55). The northernmost Keuls Multiple Range Test was able to distin-
records are from Wakasa Bay, Japan (Nakamura guish populations that differed significantly for 7
and Nakamura 1982), Taiwan (FMNH 59284), sets of regressions but could not do so for 4:
635
 FISHERY BULLETIN: VOL. 82, NO.4

maximum body depth, Base ·ID, Base 2D, and 254 (description, synonymy), pI. 46, fig. 2.
Base A. Significant differences were found be- Delsman 1931:402,407 (vertebrae 20+ 25= 45).
tween the Indian Ocean population and that in Hardenberg 1931:140 (common, often found in
the East Indies and Gulf of Thailand population river mouths; Bagan Si Api Api, Sumatra).
in one (interorbital); and between the Gulf of Cybium koreanum Kishinouye 1915:11 (original
Thailand and China populations in two (Sn-ID description; Korea), pI. 1, fig. 6. Kishinouye
and maxilla L). 1923:420-421 (description), pI. 21, fig. 35. Mori
One meristic difference was found between 1928:5 (Fusan, Korea; listed). Morice 1953:37
populations of S. guttatus. The Indian Ocean (villiform teeth on tongue).
population has a mode of 50 vertebrae while Sawara koreanum. Soldatov and Lindberg
populations in the East Indies, Gulf of Thailand, 1930:112 (description after Kishinouye).
and China have modes of 51. Gill rakers were Cybium guttatum. Not of Bloch and Schneider
usually 11 and second dorsal rays 21 in all four 1801. Delsman 1931:402, 407 (vertebrae 20+
populations. 25 = 45). Hardenberg 1931:141 (Bagan Si Api
Api, Sumatra).
Material examined.-TotalI49 (63.3-760). Scomberomorus guttatus. Not of Bloch and
Schneider 1801. Hardenberg 1934:311 (Su-
meas.: 144 (80.0-760): Persian Gulf (2); N Ara- matra). Delsman and Hardenberg 1934:340-
bian Sea (6); Malabar coast of India 343 (in part, description, fishery), fig. 248 (in
(25); Gulf of Mannar (7); Coromandel part, myomeres 13 + 33 = 46).
Coast of India (6, *C. interruptum Scomberomorus koreanus. Munro 1943:68, 71
Cuvier); "India" (4); Burma (2); Anda- (placed in subgenus Pseudosawara Munro).
man Sea (3); East Indies (31, *C. crooc- Okada 1955:150 (description), fig. 137 (after
kewitii Bleeker); Gulf of Thailand (17); Kishinouye). Kamohara 1967:43-44 (descrip-
China (33). tion, range), color pI. 22, fig. 3. Shiino 1972:71
counts: 143. (common name). Magnuson -1973:350 (short
diss.: 14 (367-548): Karachi, Pakistan (6); pectoral fin). *Devaraj 1976:79-87 (descrip-
Cochin, India (1); Gulf of Mannar (4); tion, validation of species, comparison with S.
Hong Kong (2). guttatus and S. semifasciatus, synonymy), fig.
2 (745 mm adult; Palk Bay, India), fig. 3
(second dorsal and anal fins), fig. 4 (vertebral
Scomberomorus koreanus (Kishinouye) column), fig. 5 (preopercle and liver). Shiino
Korean Seerfish 1976:231 (common name). Klawe 1977:2 (com-
Figure 56 mon name, range). *Devaraj 1977 (osteology).
Collette 1979:24 (characters, range). Col-
Cybium kuhlii. Not of Cuvier, 1831. Day 1878: lette and Russo 1979:13 (diagnostic characters,

FIGURE 56.-Scomberomorus korean us. Ning Po, China, 525 mm FL, NHMVuncat.

636
COLLE'ITE and RUSSO: SPANISH MACKERELS

range). Nakamura and Nakamura 1982:445- Color pattern. -Nakamura and Nakamura (1982)
446 (3 specimens; Wakasa Bay, Sea of Japan; described fresh specimens taken in Wakasa Bay
description), figs. lA, 2A. Kyushin et al.1982: in the Sea of Japan. Body greyish blue dorsally,
249 (description, photograph). Cressey et al. silvery white laterally and ventrally. Several
1983:264 (host-parasite list, 3 copepod species). longitudinal rows of small brownish spots rather
Lee and Yang 1983:231 (Taiwan), fig. 22 (550 sparsely scattered along lateral median line. First
mm FL). Collette and Nauen 1983:66-67 (de- dorsal fin membrane black. Pectoral, second dor-
scription, range), fig. sal, and caudal fins dark brown. Pelvic and anal
Scomberomorus semifasciatus. Not of Macleay fins silvery white.
1884. Fraser-Brunner 1950:159 (C. koreanus There are good drawings of S. koreanus from
placed in synonymy of S. semifasciatus). Japan in Kishinouye (1923:pl. 21) and from India
Sawara koreana. Mori 1952:136 (listed; Fusan in Devaraj (1976:fig. 2), and there is a good
and Chinnampo, Korea). photograph of a specimen from the Sea of Japan
Scomberomorus guttatus koreanus. Silas 1964: in Nakamura and Nakamura (1982:fig. lA). There
313-314, 325-326, 328-329 (description and is a good color illustration of S. korean us in
range in part). Kamohara (1967:pl. 22) and a color photograph of
a 411 mm specimen from the South China Sea in
Types. -Cybium koreanum Kishinouye 1915 was Kyushin et al. (1982:249).
based on a specimen collected by Yojiro Wakiya
on the west coast of Korea in 1913. There is no Biology.-Little is known of the migrations or
evidence to indicate that the specimen is still movements of S. koreanus. Kishinouye (1923)
extant. Data from the original description: D reported that it spawns at the mouth of Daidoko,
XIV+18-21+IX; A 18-21+VIII; GR 3+10=13; near Chinnanpo, Korea, in July. Feeds on sar-
vertebrae 20 + 26 = 46. dines, anchovies, and shrimps (Kishinouye 1923).

Diagnosis. -The only species of Scomberomorus

with four loops and five limbs to the intestine
(Fig. 3D. Other species have two loops and three TABLE lB.-Summary of morphometric data of Scom-
limbs or a straight intestine. It shares with S. FL = fork length, HI.. = head
beromorus koreanus.
guttatus the presence of numerous fine auxiliary length.

branches that branch from the anterior part of Character N Min. Max. Mean SO

the lateral line on the body (Fig. 56). Scapular Fork length 30 160 812 386 201
Snout-A %. FL 30 443 547 493 20
foramen small (Fig. 43e) as in S. guttatus and S. Snout-20 %. FL 30 427 512 467 15
niphonius. Supraoccipital crest high (Fig. 15a) as Snout-10 %. FL 30 215 254 241 9
Snout-P2 %. FL 29 224 267 247 11
in S. guttatus and S. multiradiatus. Snout-P, %. FL 29 189 223 209 9
P,-P2 %. FL 28 99 127 114 7
Head length %. FL 30 187 224 207 11
Description. -Lateral line gradually descending Max. body depth %. FL 30 185 263 236 16
Max. body width %. FL 26 83 115 101 9
to midline on caudal peduncle. Spines in first P, length %. FL 30 112 157 134 11
dorsal fin 14-17, usually 15 (Table 9); second P21ength %. FL 26 44 67 60 7
P2 insertion-vent %. FL 29 192 265 227 19
dorsal fin rays 20-24, usually 22 or 23 (Table 10); P2 tip-vent %. FL 26 135 197 165 19
dorsal finlets 7-9, usually 8 (Table 10); anal fin Base 10 %. FL 29 198 270 217 14
Height 20 %. FL 26 112 190 165 21
rays 20-24, usually 22 or 23 (Table 11); anal Base 20 %. FL 30 120 189 161 14
finlets 7-9, usually 7 or 8 (Table 11); pectoral fin Height anal %. FL 26 96 184 160 19
Base anal %. FL 30 100 171 154 15
rays 20-24, usually 22 or 23 (Table 12). Precaudal Snout (fleshy) %. FL 30 63 77 70 3
Snout (bony) %. FL 30 56 68 62 3
vertebrae 20 (Table 6); caudal vertebrae 26 or Maxilla length %. FL 30 93 120 110 8
27, usually 26 (Table 7); total vertebrae 46 or Postorbital %. FL 29 94 114 101 5
Orbital (fleshy) %. FL 30 21 42 33 7
47, usually 46 (Table 8). Gill rakers on first Orbital (bony) %. FL 30 32 61 50 9
arch (1-2)+(9-12)=11-15, usually 2+(11-12)=13- Interorbital %. FL 30 52 69 61 4
20-caudal %. FL 29 508 586 550 21
14 (Table 5). Morphometric characters given in Head length 30 35 152 78 37
Table 18. Snout (fleshy) %. HL 30 310 367 340 13
Snout (bony) %. HL 30 271 335 300 12
Maxilla length %. HL 30 484 565 531 17
Postorbital %. HL 29 459 536 488 18
Size. -Maximum size 150 cm FL and 15 kg in Orbit (fleshy) %. HL 30 110 197 157 27
weight; matures at 75 cm and 2.25 kg (Kishinouye Orbit (bony) %. HL 30 170 286 238 33
Interorbital %. HL 30 259 350 293 22
1923); common to 60 cm.
637
 FISHERY BULLETIN: VOL. 82, NO.4

Interest to fisheries. -The fishery for this species S. koreanus: India (n = 5), East Indies (n = 9),
was begun in Daidoko, Korea, by Japanese fisher- and Japan and China (n = 8-12). Null hypotheses
men in 1917; it is caught in summer and autumn that the 3 sets of regression lines are coincident
with drift nets and pound nets (Kishinouye 1923). were accepted for 25 sets, rejected only for body
It is usually not distinguished from other species width. The Newman-Keuls Multiple Range Test
of seerflshes but comprises an important part of showed that the population from Japan and Chi-
the drift net fishery in Palk Bay and the Gulf na differed significantly in slope from that in
of Mannar between southeastern India and Sri the East Indies. The population in the East Indies
Lanka (Devaraj 1976). did not differ significantly from that in India so
these two populations were combined and retested.
Distribution. -Continental Indo-West Pacific The only significant difference was again maxi-
from Japan, Korea, and China south to Singapore mum body width and the combined India-East
and Sumatra and west to Bombay, India (Fig. 55). Indies population differed significantly from the
The northern limit ofthe range is Wakasa Bay in Japan-China population (slopes 0.090,0.123, Q =
the Sea of Japan (Nakamura and Nakamura 5.987**). No meristic differences were found be-
1982). This species usually does not occur north of tween populations.
the west and south coasts of Korea (K{shinouye
1923). Specimens obtained in the Tokyo markets Material examined. - Total 30 (160-812 mm FL).
apparently are usually imported from Korea
(Okada 1955). There are museum specimens from meas.: 30 (160-812) Tokyo market (4); Hong
Ningpo (MNHN 5513), Swatow, and Hong Kong Kong (4); Swatow and Ning-Po (4), Chi-
(BMNH 1939.1.17.48) along the coast of China. na; Indochina (1); Sumatra (8); Indone-
There appear to be few specimens or records from sia (1); India (5).
the coast of Indochina or the Gulf of Thailand, but counts: 30.
we have examined specimens from "Cochinchine" diss.: 6 (420-812): Tokyo market, probably
(MNHN A.6827). There are several reports and spec- Korean fish (4); Indonesia (1); Hong
imens from Sumatra (Bagan Api Api, Hardenberg Kong? (1).
1931 as Cybium kuhlii; Delsman and Hardenberg
1934 as S. guttatus; ZMA 114.593), but the range Scomberomorus lineolatus (euvier)
apparently does not extend out further into the Streaked Seerfish
East Indies. Dependable Indian records are from Figure 57
Pondicherry (USNM 216698), Palk Bay, and the
Gulf of Mannar (Devaraj 1976) on the east coast, Cybium lineolatum Cuvier in Cuvier and Valen-
and Bombay (ANSP 88360) on the west coast. ciennes 1831:170-172 (original description; Mal-
abar, India). Cantor 1849:1092-1093 (descrip-
Geographic variation. -Morphometric data were tion, range; Pinang). Bleeker 1852:40-41
compared by ANCOVA for three small samples of (description, synonymy; East Indies). Bleeker

FIGURE 57.-Scomberomorus lineolatus. Cochin, India, 588 mm FL, USNM223538.

638
COLLETTE and RUSSO: SPANISH MACKERELS

1853:42 (India). Gunther 1860:370 (descrip- phorus). Shiino 1976:231 (common name).
tion; Malaya). Bleeker 1861a:52 (Singapore; *Rao and Ganapati 1977:101-111 (postlarvae
listed). Bleeker 1861b:74 (Pinang, Malaya; and juveniles; India). Klawe 1977:2 (common
listed). Day 1873:225 (description, range). name, range). * Devaraj 1977 (osteology).
Day 1878:256 (description). Collette 1979:29 (characters). Collette and
Scomberomorus lineolatum. Malpas 1926:74 (3 Russo 1979:13 (diagnostic characters, range).
males, 54.5-82.5 cm TL, 1.1-3.4 kg,Ceylon). *Devaraj 1982 (age and growth). Cressey et
Frost 1928:329 (otolith similar to that of S. ai. 1983:264 (host-parasite list, 3copepod spe-
regalis). cies). Collette and Nauen 1983:68 (description,
Scomberomorus lineolatus. Munro 1943:68; 70 range), fig.
(placed in subgenus Indocybium; vertebral Scomberomorus guttatus. Not of Bloch and
count of 21 + 29 = 50 pertains to another species Schneider 1801. Fraser-Brunner 1950:160 (Cyb-
of Scomberomorus, such as S. guttatus). De ium lineolatum placed in synonymy of Scomber-
Beaufort 1951:235-236 (synonymy, description, omorus guttatus).
range). Tham 1953:49 (Singapore Straits), 50 Indocybium lineolatum. Munro 1955:221 (de-
(correlation of catch with physical factors and scription; Ceylon); fig. 651. Chacko et ai. 1968:
presence of food fishes such as Stolephorus and 1006 (fishery; Madras).
Clupea). Scott 1959:114 (description; Malaya),
photograph. Bauchot and Blanc 1961:372 Types. -Holotype: MNHN A.6866; Malabar coast
(types of Cybium lineolatum). Jones 1962:117- of India; Dussumier; 707 mm FL; D about XVII +
119 (eggs, larvae, and juveniles). Jones and 17 + IX;AI9+ X; P 1 21;RGR1 2+ 1+ 8= 11; pattern
Silas 1961:197-198 (description, range), fig. 4 of three rows of elongate streaks still visible on
(680 mm adult), fig. 5C (not 5D as legend reads, type in 1975. Photograph of type published by
lateral view of head), fig. 6D (gill arch), fig. 7E Blanc and Bauchot (1964:pI. 3, fig. 15). Paratype:
(caudal peduncle keels). Jones and Silas 1964: MNHN 6357; Mahe (Malabar Coast), Belenger;
58-61 (synonymy, description, range), pI. 7, fig. only head and tail of a fish about 710 mm FL
A (photograph of 740 mm specimen). Silas (judging from head length of145 mm). Photograph
1964:317, 323-324 (specimens from India only). of paratype published by Blanc and Bauchot
Jones and Kumaran 1964:347 (larvae as yet (1964:pI. 3, fig. 16).
undescribed). Blanc and Bauchot 1964:447
(types of Cybium lineolatum), pI. III, figs. 15, 16 Diagnosis.-The only species of Scomberomorus
(photographs of type-specimens). Rao 1964: that has a pattern of short lines on its sides (Fig.
592-594 (teleosts constitute 97% of diet of ju- 57). Other species have some spots, blotches, or
veniles, Waltair coast of India). Thiemmedh bars, or are plain. Posterior end of maxilla greatly
1966:140 (common names). Collette 1966:367- expanded as in S. plurilineatus and S. semifascia-
368 (type of Cybium lineolatum). Tongyai tus. Anterior end of premaxilla forms an acute
1966a:7-1O (synonymy, occurrence, Thailand), angle (Fig. 22b). Ascending process of premaxilla
pI. 2D. Tongyai 1966b:3-15 (5 specimens, 46.5- very long as in S. sinensis and Acanthocybium.
76.5 cm FL; Terutao Is., Andaman Sea). Pa- Supracleithrum wide (Fig. 41a), 53-57% oflength,
thansali 1968:1002-1003 (fishery; Malaya). Si- as in S. niphonius. Foramen between last pectoral
las 1968:1114 (fishery; Gulf of Mannar), pI. 3A radial and coracoid larger than in any other
(photograph of adult). Rajan et al. 1969:90 species of Scomberomorus.
(outer channel; Chilka Lake, India). Tongyai
1970:559-561 (found in areas of low turbidity Description. -Lateral line gradually descending
and high salinity offshore; Thailand). Merce- to midline on caudal peduncle. Intestine with two
ron 1970:72 (specimens from near Sihanouk- folds and three limbs (Fig. 3g). Spines in first
ville, Cambodia). Tongyai 1971a:16-18 (Thai- dorsal fin 15-18, usually 16 or 17 (Table 9); second
land). Fernando 1972:524, 530 (incidental dorsal fin rays 15-22, usually 17 or 18 (Table 10);
catches in trawls; Wadge Bank, Ceylon). Rich- dorsal finlets 7-10, usually 9 (Table 10); anal fin
ards and Klawe 1972:14 (range), 92 (references rays 17-22, usually 20 (Table 11); anal finlets 7-10,
to larvae and juveniles). Banerji 1973:129-130 usually 9 or 10 (Table 11); pectoral fin rays 20-24,
(seerfish fishery; India). Magnuson 1973:350 usually 23 (Table 12). Precaudal vertebrae 18-20,
(short pectoral fin). Orsi 1974:175 (listed; Viet- usually 19 (Table 6); caudal vertebrae 25-28,
nam). Tham 1974 (possible predator of Stole- usually 27 (Table 7); total vertebrae 44-46, usually
639
 FISHERY BULLETIN: VOL. 82, NO.4

46 (Table 8). Gill rakers on first arch 0-2) + (6-ll) were taken in shore seines in February-April,
= 7-13, usually 2+ (8-9)= 10-11 (Table 5). Morpho- more advanced stages from boat seine catches in
metric characters given in Table 19. July-September. Juveniles feed on teleosts in
India (Venkataraman 1961; Rao 1964).
TABLE 19.-Summary of morphometric data of &om-
beromorus lineolatus. FL = fork length, HL = head Interest to fisheries. -There are small fisheries for
length. S. lineolatus in the waters around Thailand,
Character N Min. Max. Mean SO Malaysia, and India. It is taken from October to
Fork length 31 144 786 434 191 November in Thai waters of the Indian Ocean
Snout-A %. FL 30 464 540 507 16
5ooul-20 %. FL 29 453 525 502 17 (Tongyai 1970). It is less abundant than either S.
Snout-1D %. FL 30 214 264 253 17 commerson or S. guttatus in the Gulf of Thailand
Snout'P2 %. FL 27 209 266 245 13
Snout-P, %. FL 30 175 237 213 14 and along the Thai coast of the Bay of Bengal
P,-P2 %. FL 27 81 104 93 6 being found in areas of lower turbidity and higher
Head length %. FL 31 174 231 207 13
Max. body depth %. FL 27 153 211 181 11 salinity than the other two species (Tongyai 1970).
Max. body width %. FL 25 65 117 96 13
P, length %. FL 28 117 156 140 11
Fished for on both coasts of Malaysia, on the west
P21englh %. FL 20 45 70 56 7 coast from November to February in the north and
P2 insertion-vent %. FL 24 216 276 240 14
P2tip·vent %0 FL 20 156 223 184 15 March to July in the south, and on the east coast
Base 10 %. FL 24 198 270 231 15 from February to March and August to November
Height 20 %. FL 23 87 166 124 19
Base 20 %0 FL 29 89 155 113 15 (Pathansali 1968). Species of Scomberomorus are
Heighl anal %. FL 22 80 148 118 15 taken on both coasts of Malaysia mainly by gill
Base anal %. FL 28 100 150 123 14
Snout (fleshy) %. FL 30 64 88 82 6 nets, but hand lines and trolling lines are also
Snout (bony) %. FL 30 55 83 74 6
Maxilla length %0 FL 30 93 136 113 11
important on the east coast (Pathansali 1968). In
Postorbital %. FL 30 75 108 92 8 India, there is an important coastal fishery for the
Orb~al (fleshy) %0 FL 30 23 43 33 6
Orb~al (bony) %0 FL 29 34 64 48 9 three species of seerfishes of which S.lineolatus is
Interorbital %0 FL 29 47 61 57 3 the least common (Silas 1968). Small individuals,
20-caudal %. FL 29 462 545 500 23
Head length 31 33 147 88 35 up to 50 cm, are taken, together with S. commer-
Snout (fleshy) %. HL 30 325 438 395 24 son and S. guttatus, during the multiple troll
Snout (bony) %. HL 30 276 398 358 28
Maxilla length %0 HL 30 473 596 547 24 fishery season (May-September) in gill nets 5-12 .
Postorb~al %0 HL 30 404 512 443 24
Orbit (fleshy) %. HL 30 131 215 157 24
mi offTuticorin in the GulfofMannar, India (Silas
Orbit (bony) %. HL 29 188 284 231 30 1968). Gill nets, hook and line, and trolling are the
Interorbital %. HL 29 255 292 275 10
most important gear types in India (Silas 1968).
Scomberomorus spp., or pIa in-see in Thai, are
highly esteemed foodfishes in Thailand and are
Size.-Maximum size 80 cm FL. consumed as spicy fish-burgers (tad-muD. pIa in-
see) or high-quality salted fish (Tongyai 1966a). A
Color pattern. -Body dark bluish dorsally, sil- monthly average of about 100 t, fresh or salted, is
very white ventrally, marked with several rows of consumed in Bangkok alone (Tongyai 1966a).
elongate lines (Fig. 57). First dorsal fin black Seerfishes form a much smaller proportion of the
anteriorly, white posteriorly. catch in India than mackerels (Rastrelliger spp.),
There is a good drawing of a specimen from but are much in demand both fresh and salt-cured
India in Jones and Silas (1962:fig. 4). There are (Jones 1968).
also poor photographs in Jones and Silas (1964:pl.
7) and Silas (1968:pl. 3). Distribution. -Gulf of Thailand and Java west
around India at least to Bombay (Fig. 58). There
Biology. -Little has been reported in the litera- are records and specimens from Cambodia (Merce-
ture on the biology of S. lineolatus. A ripe male ron 1970) and Thailand (Tongyai 1966b, 1971a;
(82.5 cm TL, 3.4 kg) was taken on Wadge Bank, off CAS-GVF 60-286) in the Gulf of Thailand and
southern India on 2 October (Malpas 1926). A from both coasts of Malaysia (Cantor 1849; Bleek-
running ripe female (76.5 cm, 4.4 kg) was caught er 1861b; Scott 1959; CAS SU 14100; BMNH
in January in the Bay of Bengal off Satun, 1860.3.19.215), Singapore Straits (Tham 1953),
Thailand, near the border with Malaysia (Tongyai Java (USNM 72632), and Sumatra (NHMV 1874.1;
1966b). Postlarvae and juveniles (18.4-99.5 mm) ZMA 114.595). Central Indian Ocean reports and
were described from Waltair on the east coast of specimens are from Madras (ZSI 2156-7), Palk Bay
India by Rao and Ganapati (1977). Early stages (Devaraj 1977; dissections), Sri Lanka (Fernando
640
COLLE'ITE and RUSSO: SPANISH MACKERELS

60' 90'

FIGURE 58.-Ranges of four Indo-West Pacific species of Scomberomorus: S. lineolatus, S. plurilin-

eatus, S. niphonius, and S. munroi.

1972), Wadge Bank south of India (Malpas 1926), Andaman Sea (4); Gulf of Thailand (5);
the Malabar coast (MNHN A.6866, holotype of C. East Indies (7).
lineolatum), Cochin (5 specimens measured, 1 counts: 31.
dissected), and Bombay (MNHN A.5783). Records diss.: 5 (417-786): Palk Strait (4); Cochin (1).
of S. lineolatus from East Mrica (Williams 1960)
are referrable to S. plurilineatus. The report of S.
lineolatus from Western Australia (McKay 1970) Scomberomorus maculatus (Mitchill)
was based on a specimen (HUMZ F-422) of S. Spanish Mackerel
munroi. Figure 59

Geographic variation. -Comparisons of morpho- Scomber maculatus Mitchill 1815:426-427 (orig-

metric data by ANCOVA were made for four small inal description; New York), pI. 6, fig. 8.
samples of S. lineolatus: Arabian Sea (n = 7-12), Cybium maculatum. Cuvier 1829:200 (listed in
Bay of BengalCn = 4-6), Gulf of Thailand (n = 3- footnote after Se. maculatus Mitchill). Cuvier
5), and East Indies (n = 4-6). No significant in Cuvier and Valenciennes 1831:181 (descrip-
differences were found. No meristic differences tion; New York). Storer 1855:146-147 (synon-
were found between populations. ymy, description; exceedingly rare in Massa-
chusetts, 1 from Lynn and 4 from Provincetown),
Material examined.-TotaI31 (160-786 mm FL). pI. 13, fig. 1. Holbrook 1860:68-72 (synonymy,
description, color, anatomy, range in part), pI. 9,
meas.: 31 (160-786): India, Arabian Sea (8, *C. fig. 1. Giinther 1860:372 (synonymy, descrip-
lineolatum Cuvier), Bay of Bengal (7); tion). Poey 1878:4 (synonymy, description).
641
 FISHERY BULLETIN: VOL. 82, NO.4

FIGURE 59.-Scomberomorus maculatus. New York market, 270 mm FL, USNM 15582. (From Goode 1884:pl. 93.)

Smiley 1881 (distribution, fishery). Ryder 1882 1937-42; Texas). La Monte 1945:26, 28 (de-
(gonads, embryology, development), pI. 1-4 scription, range), color pI. 11. Breder 1948:127
(eggs, embryos, and larvae). Earll 1883 (name, (range in part, biology), fig. Erdman 1949:301
description, distribution, movements, reproduc- (range confined to coastal America and the
tion, fishery, artificial propagation), pI. 1. N coast of Cuba; other West Indian records
Scomberomorus maculatus. Jordan and Gilbert are misidentifications). Fraser-Brunner 1950:
1882:426 (synonymy, range in part). Goode 159 (synonymy and range in part), fig. 29.
1884:307-315 (range, fishery, reproduction), pI. Baughmall- 1950:244 (numerous Texas records
93. Meek and Newland 1884:232-234 (descrip- throughout the year). Knapp 1950:142 (458
tion, synonymy, range in part). Dresslar and stomach contents, Texas). Rivas 1951:225-226
Fesler 1889:442 (in key), 443 (synonymy and (synonymy in part, diagnosis, range). Taylor
range in part), pI. 9. Jordan and Evermann 1951:116-118 (biology, occurrence in North Caro-
1896b:874 (description, synonymy in part). lina), 271-272 (angling in North Carolina). La
Jordan and Evermann 1900:pI. 134, fig. 368 Monte 1952:50 (description, range), color pI.
(specimen from New York market). Jordan 18. Bigelow and Schroeder 1953:347-348 (de-
and Evermann 1902:285-286 (description, scription, range in part, only a stray in the Gulf
range), fig., photograph. Bean 1903:396-398 of Maine), fig. 182. Pew 1954:26 (description,
(synonymy, description, range in part, occur- range, habits), fig. 23. Mather and Day 1954:
rence in New York). Smith 1907:190-192 (diag- 185 (comparison of Wand E Atlantic speci-
nosis, size, range, catch, price; North Carolina), mens; S. tritor at best racially distinct from
fig. 77. Sumner et aI. 1913:750 (references, S. maculatus). Briggs 1958:286 (range in
occurrence, parasites; Buzzards Bay and Vine- part). Springer and Pirson 1958:175 (catch in
yard Sound, Mass.). Schroeder 1924:6-7 (most Texas). *Mago Leccia 1958 (osteology, com-
valuable food fish in the Florida Keys, range in parisons with S. regalis and S. cavalla), figs.
part), fig. 3. Nichols and Breder 1927:123-124 *Klima 1959 (distribution, biology). Moe 1963:
(description, range, biology), fig. 170. Frost 109 (second most fished for species by private
1928 (sagitta similar to that ofS. regalis). Hil- boats in Florida). Jorgensen and Miller 1968:
debrand and Schroeder 1928:203-205 (descrip- 9,13 (SL-FL-TL conversions). *Mendoza 1968
tion, range, biology, synonymy; Chesapeake (biology; Veracruz). Lyles 1969:1-15 (landings,
Bay), fig. 115. Nichols 1929:229-230 (synon- in part). Wollam 1970 (development, pigmen-
ymy, range, description), fig. 82. Hildebrand tation, counts, and measurements; 175 larvae
and Cable 1938:508-518 (development of larvae and juveniles, 3.1-25 mm SL, from the Gulf of
and postlarvae; Beaufort, N.C.), figs. 2-10 (lar- Mexico), figs. 2,3, 6A (larvae and juveniles, 3.1-
vae and juveniles, 2.75-97 mm). Baughman 25 mm SL). Beardsley and Richards 1970:5
1941:17 (migratory off Texas coast). Munro (length-weight of35 specimens from SE Florida,
1943:67, 71-72 (placed in subgenus Scomber- 330-770 mm FL, 0.45-4.76 kg). Farragut 1972
omorus). Fowler 1945:185-186 (synonymy, de- (use of antioxidants to prevent rancidity during
scription; South Carolina). Gunter 1945:55 frozen storage). Richards and Klawe 1972:14
(occurrence, Texas), 145 (monthly production, (range in part), 92-93 (references to eggs and
642
COLLE'ITE and RUSSO: SPANISH MACKERELS

larvae). Miyake and Hiyasi 1972:III-3 (in key); rate sides; more than half anterior part of first
IV-11 (common names). Dwinell and Futch dorsal fin black, remainder white) leave no doubt
1973 (188 larvae and juveniles, 2.8-42.2 mm SL, as to identity of name.
caught in June, Aug., and Sept., NE Gulf of
Mexico). Marquez 1973 (distribution, biology, Diagnosis. -Scomberomorus maculatus pos-
fishery). * Powell 1975 (age, growth, reproduc- sesses nasal denticles as do the other five species of
tion; Florida). * Berrien and Finan 1977b (in the regalis group (brasiliensis, concolor, regalis,
part; species synopsis). Klawe 1977:2 (com- sierra, and tritor) and has an artery branching
mon names, range). Fritzsche 1978:126-132 from the fourth left epibranchial artery as do all
(description, larval development), figs. 70-74 the species in the group except S. tritor. Like S.
(eggs, larvae, and juveniles). Collette 1978: concolor, S. maculatus lacks the shunt from the
Scombm 5 (description, range), figs. Collette fourth right epibranchial artery to the coeliaco-
et aI. 1978:274-275 (comparison with other mesenteric artery (Fig. 7c). It also has more
American species of Scomberomorus). Ma- vertebrae (51-53) than any ofthe other five species
nooch et aI. 1978 (annotated bibliography). in the group (46-49). Intercalar spine absent as in
Pristas and Trent 1978:582-588 (most abundant the other five species of the regalis group and S.
spring-fall; St. Andrew Bay, Fla.). Collette niphonius.
1979:29 (characters, range). Collette and
Russo 1979:13 (diagnostic characters, range). Description. - Lateral line gradually descending
Trent and Anthony 1979 (commercial and rec- to midline on caudal peduncle. Intestine with two
reational fisheries in U.S.). Doi and Men- folds and three limbs (Fig. 3h). Spines in first
dizabal 1979 (Mexican catch). Meaburn 1979 dorsal fin 17-19, usually 18 (Table 9); second dorsal
(heavy metal contamination). Hale 1979 (pres- fin rays 17-20, usually 18 or 19 (Table 10); dorsal
ervation technology). Amezcua-Linares and finlets 7-9, usually 8 or 9 (Table 10); anal fin rays
Yaiiez-Arancibia 1980:86-90 (Campeche, Mex- 17-20, usually 19 or 20 (Table 11); anal finlets 7-10,
ico). McEachran et aI. 1980 (larvae off Texas usually 8 or 9 (Table 11); pectoral fin rays 20-23,
coast). Sutherland and Fable 1980 (annual usually 21 (Table 12). Precaudal vertebrae 21 or
migration from wintering grounds off S Florida 22, usually 21 (Table 6); caudal vertebrae 30 or 31
and Campeche to summer grounds along the N (Table 7); total vertebrae 51-53, usually 51 or 52
coast of the Gulf of Mexico, return migration in (Table 8). Gill rakers on first arch (1-4)+ (8-13)=
fall). Deardorff and Overstreet 1981 (larvae of 10-16, usually 2 + (10-11) = 12-14 (Table 5). Mor-
4 forms of the anisakid nematode Hysterothy- phometric characters given in Table 20.
lacium found in mesentary of specimens from
the Gulf of Mexico). Johnson 1981 (electro- Size. - Maximum size 77 cm FL, 4.8 kg (Beardsley
phoresis; NW Florida). Skow and Chittenden arid Richards 1970). Sexual maturity in Florida is
1981 (differences between Atlantic coast and attained by age II, at 25-32 cm FL for females, 28-
Gulf of Mexico populations by hemoglobin elec- 34 cm for males (Klima 1959). Length-weight
trophoresis). Richardson and McEachran 1981 equations have been presented for populations in
(larvae 1.8-2.9 mm SL, pigment characters, Florida (Powell 1975) and Veracruz (Doi and
measurements; Gulf of Mexico), fig.lB (2.1 mm MendizabaI1979).
larva). Naughton and Saloman 1981 (stomach
contents of 344 juveniles, 117-432 mm FL; Cape Color pattern. -Dark bluish above, silvery below,
Canaveral, Fla., and Galveston, Tex.; diet main- . sides marked with about three rows of round to
ly clupeoids). Adkins and Bourgeois 1982:12- elliptical dark spots (Fig. 59), orange in life. First
13, 32-35, 48 (gill net; Louisiana). Cressey et dorsal fin black anteriorly and at distal margin
aI. 1983:264 (host-parasite list, 4 copepod spe- posteriorly, basal part of posterior membranes
cies). Collette and Nauen 1983:69-70 (descrip- white.
tion, range), fig. Saloman and Naughton 1983b There is a color painting of an S. maculatus in
(food in U.S. waters). La Monte (l945:pI. 11, 1952:pI. 18), and a black and
white photograph of one in Jordan and Evermann
Types. -Scomber maculatus Mitchill 1817 was (1902). The drawing published by Goode (1884:pI.
based on a 19-in (482.6 mm) fish from New York. 93) is included here as Figure 59.
No types known to be extant. The figure (pI. 6, fig.
8) and description (about 20 yellowish spots deco- Biology. -Summaries of biological information

643
 FISHERY BULLETIN: VOL. 82, NO.4

have been presented by Mendoza (1968), Marquez support the Panama City to Mobile and Port
(1973), and Berrien and Finan (1977b). There is Aransas to Veracruz migrations (Sutherland and
also a useful annotated bibliography by Manooch Fable 1980). Spawning takes place in New York-
et al. (1978). The Spanish mackerel is clearly a New Jersey late August-late September, in Chesa-
migratory species that moves north from Florida peake Bay mid-June to the end of summer, and in
along the Atlantic coast of the United States and the Carolinas starting in April (Earll 1883). Ripe
north and wes~ along the coast of the Gulf of females were found in Florida from July to Sep-
Mexico in the spring and returns in the fall, but tember by Klima (1959) and from April to Septem-
the details of the migration are not completely ber by Powell (1975). Powell felt that individuals
known. There are large concentrations in the spawned repeatedly in a prolonged spawning
winter in Florida and the Florida Keys (Beau- season in Florida. Spanish mackerel spawn from
mariage 1970) which move north to reach Charles- May to September in waters < 50 m over the inner
ton, S.C., in late March, North Carolina in April, continental shelfofTexas (McEachran et al. 1980).
Chesapeake Bay in May, and Sandy Hook, N.J., Spawning in Veracruz takes place in July-Sep-
to Narragansett Bay, R.I., by late July (Earll tember (Mendoza 1968). Early studies on develop-
1883; Beaumariage 1970). Schools also move north ing eggs and larvae (to 6 d old) were carried out by
along the Gulf coast of Florida in the spring (Moe " Ryder (1882) in North Carolina. Larvae have been
1972), and west across the northern Gulf from described from North Carolina (14-20 mm, Hilde-
Panama City, Fla., to Mobile, Ala. (Sutherland brand and Cable 1938; some misidentified, see
and Fable 1980), and possibly on into Texas reach- Wollam 1970), the west coast of Florida (3.1-35.0
ing Galveston in early March and Port Aransas in mm, June-September, Wollam 1970), the north-
late March (Baughman 1941). There is also north- eastern Gulf of Mexico (2.8-42.2 mm SL, June-
south migration along the Mexican coast, from September, Dwinell and Futch 1973), and Texas
south to north in March-April, north to south in (1.8-11.5 mm SL, May-September, McEachran et
August-November (Mendoza 1968). Tag returns al. 1980). Most were taken over the middle and

TABLE 20.-Summary of morphometric data of Scomberomorus maculatus. FL = fork length, HL =

head length.

AUantic GuU of Mexico Total

Character N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO
Fork length 24 163 712 330 137 36 152 593 307 95 60 152 712 316 113
Snout-A %. FL 24 515 570 538 14 36 507 565 534 14 60 507 570 536 14
Snout-20 %. FL 24 478 540 505 14 36 466 543 502 16 60 466 543 503 15
Snout· 10 %. FL 24 227 260 240 10 36 226 258 242 7 60 226 260 241 8
Snout-P2 %. FL 22 216 279 249 14 32 228 300 263 17 54 216 300 257 17
Snout-P, %. FL 24 199 247 217 14 36 193 261 217 12 60 193 261 217 13
P,-P2 %. FL 21 95 121 108 8 34 97 132 111 9 55 95 132 110 9
Head length %. FL 24 195 281 213 18 36 195 227 211 9 60 195 281 212 13
Max. body depth %. FL 18 168 232 194 19 33 178 228 198 12 51 168 232 197 15
Max. body width %. FL 18 70 110 89 12 36 65 137 92 15 54 65 137 91 14
P, length %. FL 24 114 146 131 7 35 107 140 129 8 59 107 146 129 8
P21ength %. FL 21 43 66 55 6 33 35 68 50 9 54 35 68 52 8
P2 insertion-vent %. FL 22 241 305 270 16 33 223 299 259 19 55 223 305 263 19
P2tip-vent %. FL 21 188 246 215 18 31 175 233 208 13 52 175 246 211 15
Base 10 %. FL 24 236 288 258 13 36 222 286 254 14 60 222 288 256 13
Height 20 %. FL 21 92 153 124 14 28 108 141 125 10 49 92 153 125 12
Base 20 %. FL 24 109 150 128 10 35 110 158 128 11 59 109 158 128 11
Height anal %. FL 23 97 140 117 13 31 88 133 119 12 54 88 140 118 12
Base anal %. FL 24 100 157 122 14 36 101 147 124 10 60 100 157 123 12
Snout (fleshy) %. FL 24 73 86 78 3 36 74 90 80 4 60 73 90 80 4
Snout (bony) %. FL 24 59 78 69 4 23 63 84 72 5 47 59 84 70 5
Maxilla length %. FL 24 108 134 117 7 36 110 136 120 6 60 108 136 119 6
Postorbital %. FL 23 87 108 98 5 36 84 101 95 4 59 84 108 96 4
Orbital (fleshy) %. FL 24 24 47 34 7 35 26 47 34 4 59 24 47 34 5
Orbital (bony) %. FL 24 41 62 50 7 36 40 65 52 6 60 40 65 51 7
Interorbital %. FL 24 51 60 56 2 36 52 63 57 3 60 51 63 56 3
20-caudal %. FL 23 462 521 493 19 36 438 524 483 22 59 438 524 487 21
Head length 24 38 147 69 27 36 34 129 65 20 60 34 147 67 23
Snout (fleshy) %. HL 24 271 401 369 25 36 346 416 381 16 60 271 416 376 21
Snout (bony) %. HL 24 237 360 327 25 23 325 385 343 15 47 237 385 335 22
Maxilla length %. HL 24 401 572 552 34 36 546 628 568 15 60 401 628 562 25
Postorbllal %. HL 23 317 496 461 37 36 407 486 450 16 59 317 496 454 27
Orbit (fleshy) %. HL 24 103 204 159 27 35 121 216 160 17 59 103 216 160 21
Orbit (bony) %. HL 24 153 284 234 29 36 186 308 246 27 60 153 308 242 28
Interorbital %. HL 24 204 296 284 20 36 234 296 268 13 60 204 296 266 16

644
COLLETIE and RUSSO: SPANISH MACKERELS

inner continental shelf, over depths from 12 to Anthony 1979). Since 1950, over 92% of the total
50 m, off Texas at surface water temperatures of U.S. catch has been landed in Florida (Trent and
19.6°-29.8° C and salinities of28.3-37.4%. (McEach- Anthony 1979). In 1976 about 18 million pounds
ran et al. 1980). Summaries of previous larval valued at about $3.2 million were landed by com-
work and illustrations of larvae 2.6-13.5 mm and mercial fishermen in the United States; in 1970 an
. of juveniles 14-97 mm are contained in Fritzsche estimated 23 million pounds were landed by rec-
(1978). As with other members of the genus, food reational fishermen (Trent and Anthony 1979).
consists chiefly of small fishes with lesser quanti- Spanish mackerel is second in volume among
ties of penaeoid shrimps and cephalopods. Clupe- Mexico's Gulf of Mexico fisheries with an average
oids such as menhaden (Brevoortia), alewives annual production from 1968 to 1976 of4,900 t (Doi
(Alosa), thread herring (Opisthonema), Spanish and Mendizabal 1979). Most of this (80%) is
sardine (Sardinella), and anchovies (Anchoa) are produced in the state of Veracruz with lesser
particularly important in North Carolina, Flor- amounts from Campeche (15%) and Yucatan (5%).
ida, Texas, and Veracruz (Earll 1883; Knapp 1950; The early fishery in the United States utilized
Miles and Simmons 1951; Klima 1959; Mendoza trolling lines, gill nets, and pound nets (Earll
1968; Naughton and Saloman 1981; Saloman and 1883). The commercial fishery in Florida utilizes
Naughton 1983b). Juveniles (100-400 mm FL) ate stab or floating gill nets, which capture fish of age
more anchovies (Naughton and Saloman 1981; II-III, 30-65 cm FL (52% 36-41 cm), and hook and
Saloman and Naughton 1983b) than adults did. line, which captures smaller fish, age I-II, 21-69
Other fishes commonly consumed include Caran- cm FL (38% 33-35 cm) (Klima 1959). Larger
gidae, Mugilidae, and Trichiuridae (Klima 1959; vessels now entering the fishery have power-
Saloman and Naughton 1983b). rollers to retrieve the nets which are mostly nylon;
airplane spotter pilots locate the fish (Trent and
Interest to fisheries. - The Spanish mackerel is a Anthony 1979). Recreational anglers catch Span-
valued fish to recreational or commercial fisheries ish mackerel from boats while trolling or drifting
throughout its range. The fishery along the Atlan- and from boats, piers, jetties, and beaches by
tic coast of the United States north of southern casting, livebait fishing, jigging, and drift fishing
Florida is seasonal (Klima 1959): late July to (Trent and Anthony 1979). Fishermen in Veracruz
September from Rhode Island to New Jersey employ beach seines (chinchorros playeros), gill
(Earll 1883), Mayor June to September in Chesa- nets (redes agalleras), trolling spoons (curricanes),
peake Bay (Hildebrand and Schroeder 1928), April and trap nets (almadrabas) (Doi and Mendizabal
to June 'on the way north, and September-October 1979). Nearly all the catch is consumed fresh,
on the way south through the Carolinas (Smith frozen, or smoked (Lyles 1969). A few attempts
1907; Taylor 1951). The fishery in southern Florida have been made at canning Spanish mackerel but
is concentrated in the winter months, October- the product has not been widely accepted (Earll
February or March (Klima 1959; Beaumariage 1883; Lyles 1969). Frozen fish begin to show signs
1970). In northwest Florida, the fishery peaks of rancidity after as little as 3 mo time in frozen
March-April (Beaumariage 1970); in Louisiana in storage, a problem which has been treated with
June and October (Adkins and Bourgeois 1982); antioxidants and EDTA (Farragut 1972; Hale
and in Texas March-April and July-September 1979).
(Springer and Pirson 1958). As in the Carolinas,
there are two major capture seasons in Veracruz: Distribution. - Western Atlantic Ocean from
45% of the annual production is taken March- Massachusetts south along the Atlantic coast of
April during the northward migration and 30% the United States and the coast of the Gulf of
October-December in the southward migration Mexico from Florida to Yucatan, Mexico (Fig. 49).
(Doi and MendizabaI1979). The beginnings ofthe There are several summer records from the south-
Spanish mackerel fishery in the United States ern side of Cape Cod, Buzzards Bay, Woods Hole,
were discussed by Earll (1883) and a historical and Vineyard Sound (Sumner et al. 1913; Bigelow
summary of the U.S. catch from 1887 to 1967 was and Schroeder 1953), but only strays are known
provided by Lyles (1969). The commercial fishery from further north. Storer (1855) recorded the
began along the middle Atlantic and Chesapeake capture of an individual at Lynn in Massachu~etts
Bay areas before 1850, and by 1880 about 86% of Bay and stated that individuals had been obtained
the total U.S. catch of 1.9 million pounds was at Provincetown, at the tip of Cape Cod, and at
landed in the Chesapeake Bay area (Trent and Monhegan Island in Maine. There do not appear to
645
 FISHERY BULLETIN: VOL. 82, NO.4

be any extant specimens to verify these records; Gulf of Mexico coast of U.S. (35); Yuca-
there is one specimen labelled only as "Cape Cod" tan (1).
(MCZ 23929). The known southern limit of the counts: 67.
range is Progreso, Yucatan (MCZ 32894); it is diss.: 16 (281-712): Va.-N.C. (3); Ga.-Fla. (7); St.
replaced by S. brasiliensis from Belize to Rio de Andrews Bay, Gulf of Mexico, Fla. (6).
Janeiro and by S. regalis in the Bahamas and
West Indies. Reports of S. maculatus from the
West Indies (except for th,e north coast of Cuba) Scomberomorus multiradiatus Munro
are referrable to S. regalis (Erdman 1949), those Papuan Seerfish
from the eastern Pacific are based on S. sierra, and Figure 60
those from the eastern Atlantic on S. tritor:
Scomberomorus multiradiatus Munro 1964:168-
Geographic variation. - Morphometric characters 169 (original description; Gulf of Papua off
were compared for two populations of S. macula- mouth of Fly R.), fig. 12 (holotype). Munro
tus by ANCOVA (Table 20): Atlantic coast of the 1967:200 (description), fig. 339. Magnuson
United States (n = 18-24) and Gulf of Mexico (n = 1973:350 (short pectoral fin). Kailola 1975:235
28-36). Null hypotheses that the 2 sets of regres- (4 specimens listed; Orokolo Bay, Tureture, 17
sion lines are coincident were accepted for 20 sets mi W of Fly R.). Klawe 1977:2 (range, common
of regressions and rejected for 6: Sn-lD, Sn-P2 , name). Kailola and Wilson 1978:34 (trawled
P1L, Snout (fleshy), Snout (bony), and maxilla in Gulf of Papua), 60 (number of fin rays). Col-
length. lette 1979:29 (diagnostic characters). Col-
There is also a difference in vertebral numbers lette and Russo 1979:13 (diagnostic characters,
between the eastern U.S. and Gulf of Mexico range). Lewis 1981:16 (photograph, biology).
populations. The eastern U.S. population usually Cressey et al. 1983:264 (host-parasite list, 2
has 31 caudal and 52 total vertebrae ex 30.4, 51.5), copepod species). Collette and Nauen 1983:
the gulfpopulation has 30 or 31 caudal and 51 or 52 70-71 (description, range), fig.
total vertebrae (x 30.9, 52.1).
The Gulf ofMexico and eastern U.S. populations Types. -Holotype: CSIRO C.3172; off northern
also differ electrophoretically. Skow and Chitten- mouth of Fly River, Papua New Guinea; MV Fciir-
den (1981) found differences in hemoglobin pheno- wind; 1948-50; 232 mm FL; D XVII + 23 + VIII; A
types between samples from Port Aransas, Tex., 28+ VI; P 1 23-23; RGR 1 0+ 1 + 2= 3; vertebrae
and Beaufort, N.C. 20 + 35 = 55; no spots or vertical bars. Holotype
illustrated by Munro (1964:fig. 12, 1967:fig. 339).
Material examined. -Total 69 (152-712 mm FL).
Diagnosis. -The species of Scomberomorus with
meas.: 60 (152-712): Atlantic coast of U.S. (24); the most vertebrae (54-56) and the fewest gill

FIGURE 60. -Scomberomorus multiradiatus. Gulf of Papua, 312 mm FL, USNM 233696.

646
COLLETIE and RUSSO: SPANISH MACKERELS

rakers (1-4) compared with the other 17 species Size. -Maximum size 35 cm FL, 0.5 kg, the
(vertebrae 41-53, gill rakers 1-27). It has the most smallest species in the genus, sexually mature at
rays in the anal fin (25-29, compared with 15-24 < 30 cm (Lewis 1981).
in the other 17 species). Posterior end of maxilla
only slightly expanded as in 8. sinensis. Supra- Color pattern. -Dark bluish black dorsally, sil-
occipital crest high as in 8. guttatus and S. very white ventrally, with no spots, blotches, or
koreanus. Supracleithrum narrow (Fig. 41b), 43- bars (Fig. 60). First dorsal fin black anteriorly and
53% of length as in 8. sinensis, semifasciatus, and along distal edge posteriorly with some white at
sierra. Foramen between last pectoral radial and base of fin posteriorly. The only previously pub-
coracoid smaller than in any other species of lished figures were of the holotype by Munro
Scomberomorus. (1964:fig. 12, 1967:fig. 339) and a photograph by
Lewis (1981:16).
Description. -Lateral-line gradually descending
to midline on caudal peduncle. Intestine with two Biology. -Schooling and other behavior unknown
folds and three limbs (Fig. 3i). Spines in first (Lewis 1981).
dorsal fin 16-19, usually 17 or 18 (Table 9); second
dorsal fin rays 21-25, usually 23 or 24 (Table 10); Interest to fisheries. -Trawled in the Gulf of
dorsal finlets 7-9, usually 7 or 8 (Table 10); anal fin Papua but too small to be of any commercial
rays 25-29, usually 26-28 (Table 11); anal finlets 6- significance.
8, usually 6 (Table 11); pectoral fin rays 20-23,
usually 21 or 22 (Table 12). Precaudal vertebrae 20 Distribution. -Restricted to shallow turbid wa-
or 21 (Table 6); caudal vertebrae 34-36, usually 34 ters of the Gulf of Papua off the mouth ofthe Fly
or 35 (Table 7); total vertebrae 54-56, usually 55 or River (Fig. 55). The western known limit is Ture-
56 (Table 8). Gill rakers on first arch 0 + (1-4) = 1-4, ture village, 12 mi west of Daru (Kailola 1975;
usually 0+ (2-3)= 2-3 (Table 5). Morphometric DASF FO 2851), and the eastern limit Freshwater
characters given in Table 21. Bay at Kerema (lat. 8°12' S, long. 145°59'E; Kai-
lola 1975; several dissected specimens, USNM).
TABLE 21.-Summary of morphometric data of Scom-
beromorus multiradiatus. FL = fork length, HL = head Material examined. -Total 28 (203-350 mm FL).
length.

Character N Min. Max. Mean SO meas.: 27 (203-350): off Fly R., Gulf of Papua
Fork length 27 203 350 253 39 (*8. multiradiatus).
Snout-A roo FL 27 471 540 505 15
Snout-20 roo . FL 27 448 508 477 15
counts: 28.
Snout-10 %0 FL 27 230 268 249 9 diss.: 4 (224-323).
Snout-P, %0 FL 27 228 255 243 7
Snout-P, roo FL 27 201 229 213 7
P,·P2 %0 FL 26 93 118 102 6
Head length %0 FL 27 198 247 208 8
Max. body depth %0 FL 27 205 265 228 16 Scomberomorus munroi Collette and Russo
Max. body width %0 FL 27 81 113 95 9 Australian Spotted Mackerel
P, length %0 FL 22 121 143 131 5
P,length %0 FL 25 29 56 40 6 Figure 61
P2 insertion-vent %0 FL 26 204 299 247 19
P2 tip-vent %0 FL 24 160 269 207 24
Base 10 %0 FL 25 188 243 216 13 8comberomorus niphonius. Not of Cuvier 1831.
Height 20 %0 FL 17 144 186 167 12
Base 20 %0 FL 27 154 198 177 13 Munro 1943:86-90 (first Australian record, de-
Height anal %0 FL 20 151 178 164 8 scription, range) pI. vii, fig. A. Serventy 1950:
Base anal %0 FL 27 176 268 216 18
Snout (fleshy) %0 FL 27 70 84 77 3 19 (throughout N Australia, S to Kimberly).
Snout (bony) roo FL 27 59 74 67 4
Maxilla length %0 FL 27 117 134 125 3
Roughley 1951:110 (description), pI. 45.cfig. after
Postorbital roo FL 27 77 96 86 6 Munro 1943). Taylor 1964:282 (Arnhem Land;
Orbital (fleshy) roo FL 27 30 41 34 3
Orbital (bony) roo FL 27 43 60 52 4 listed after Munro 1958a). Marshall 1964:364-
Interorbital %0 FL 27 51 71 58 5 365 (description after Munro 1943), pI. 50, fig.
20-caudal %0 FL 24 470 522 494 15
Head length 27 42 73 53 9 351. Marshall 1966:205 (description), pI. 50,
Snout (fleshy) roo HL 27 317 407 372 18 fig. 351. Kailola 1974:72 (description, first rec-
Snout (bony) roo HL 27 278 349 321 19
Maxilla length roo HL 27 514 630 603 21 ord from Papua New Guinea). Kailola 1975:
Postorbital roo HL 27 339 469 415 35
Orbit (fleshy) roo HL 27 121 195 165 16
237 (listed, fish. reference collection). Zhang
Orbit (bony) %0 HL 27 190 287 252 23 and Zhang 1981:104 (Australia). -
Interorbital roo HL 27 257 342 280 22
Sawara niphonia. Not of Cuvier 1831. Munro
647
 FISHERY BULLETIN: VOL. 82, NO.4

FIGURE 61.-Scomberomorus munroi. Deception Bay, Queensland, 740 mm FL, USNM 218387, holotype.

1958a:20 (description; new record for W Austra- straight intestine, and in having more vertebrae
lia); fig. Whitley 1964a:240 (fig. 5, range), (50-52, usually 51 vs. 48-50, usually 49).
252 (range, size). Whitley 1964b:48 (listed).
Grant 1965:175 and 1972:104 (description after Description. - Lateral line gradually descending
Munro 1958a), fig. Grant 1975:162 (descrip- to midline on caudal peduncle. Intestine with two
tion), 163 (color pI. 41). Grant 1978:190 (de- folds and three limbs (Fig. 3j). Spines in first
scription), 191 (color pI. 73). dorsal fin 20-22, usually 20 or 21 (Table 9); second
Sawara niphonius. Not of Cuvier 1831. Rohde dorsal fin rays 17-20, usually 18 (Table 10); dorsal
1976 (no monogenes found on 1 specimen from finlets 9 or 10, usually 9 (Table 10); anal fin rays
Coffs Harbour, N.s.W). 17-19, usually 17 or 18 (Table 11); anal finlets 8-10,
Scomberomorus sp. Collette 1979:29 and Collette usually 9 (Table 11); pectoral fin rays 21-23,
and Russo 1979:13 (Australian population re- usually 21 or 22 (Table 12). Precaudal vertebrae 21
ferred to S. niphonius actually an undescribed or 22, usually 22 (Table 6); caudal vertebrae 28-30,
species). usually 29 (Table 7); total vertebrae 50-52, usu-
Scomberomorus munroi Collette and Russo 1980: ally 51 (Table 8). Gill rakers on first arch 2 +
243-248 (original description; Australia, Papua (8-10)=10-12, usually 8+(8-9)=10-11 (Table 5).
New Guinea), fig. 1 (holotype). Lewis 1981: Morphometric characters given in Table 22.
18 (photograph, biology). Grant 1982:622 (de-
scription), 623 (color pI. 323). Cressey et Size. -Maximum size 100 cm FL, 8 kg; more
al. 1983:264 (host-parasite list, 4 copepod spe- commonly 50-80 cm and 4.5 kg in weight (Lewis
cies). Collette and Nauen 1983:71-72 (descrip- 1981); size at first maturity 50-55 cm (A. D.
tion, range), fig. Lewis 5 ). A 9.1 kg fish was caught near Rock-
hampton, Queensland, in 1976 (G. McPherson 6 ).
Types. -Holotype: AMS 1.21029-001; Deception
Bay N of Brisbane, Queensland; M. Dredge; 15 Color pattern. -Sides with several poorly defined
May 1975; 705 mm FL; D XXI+18+X; A 18+X; rows of round spots, larger than pupil but smaller
P 1 22; RGR 1 2 + 1 + 7 = 10; vertebrae 22 + 30 = than diameter of eye (Fig. 61). Scomberomorus
52. Paratypes: 10 (373-820 mm FL) from Queens- niphonius has more numerous, smaller spots,
land and southern coast of Papua New Guinea (see usually about size of pupil. Munro (1943:87) re-
Collette and Russo 1980:247). ported sides of freshly caught specimens light
silvery grey, upper part of back and inner surface
Diagnosis. - This species differs from all other of pectoral fin dark blue, cheeks and belly silvery
species of Scomberomorus in lacking an anterior
process on the outer surface of the head of the
maxilla (Fig. 23b). It is superficially similar to S.
5 A. D. Lewis, Principal Fisheries Officer, Department of
niphonius in being spotted and having many Agriculture & Fisheries, P.O. Box 358, Suva, Fiji, pers. commun.
spines (20-22) in the first dorsal fin. It differs from February 1983.
"G. McPherson, Fishery Biologist, Northern Fisheries Ser-
S. niphonius in having the usual two loops and vice, Queensland Fisheries Service, % Post Office, Bungalow,
three limbs to the intestine instead of having a Cairns, Qld. 4870, Australia, pers. commun. February 1983.

648
COLLETTE and RUSSO: SPANISH MACKERELS

white, anal fin light silvery grey, and anal finlets coast of Queensland from Double Island Point to
silvery grey. First dorsal fin black (bright steely Southport (Grant 1982). Other biological informa-
blue in fresh specimens according to Munro) with tion is lacking on this species.
blotches of white toward bases of more posterior
membranes in some specimens. First dorsal fin Interest to fisheries. -Together with Grammator-
membranes entirely black in holotype (Fig. 61). cynus and three other species of 8comberomorus,
Most other species of 8comberomorus with more mackerel fishing is Queensland's second major
extensive white areas on posterior half or middle finfishery with an annual output of about 1,000
third of dorsal fin. tons of whole and filleted fish (Anonymous 1978).
An excellent black and white illustration of 8. The best catches are made by drifting or anchoring
munroi drawn by Munro has appeared several over inshore Queensland reefs and fishing with
times in the literature (Munro 1943, 1958a; lines baited with small fish on a gang of three or
Roughley 1951; Marshall 1964, 1966) and another four linked hooks (Grant 1982). Also taken by
drawing by George Coates has been published by trawlers in the Gulf of Papua.
Grant (1965 and subsequent editions). We (Col-
lette and Russo 1980) illustrated the holotype with Distribution. -Inshore coastal waters of the
the same figure that is included here. Grant has northern coast of Australia (Fig. 58) from Abrol-
included a color photograph of a freshly caught hos Islands region in Western Australia to Coffs
specimen in the last three editions of his book Harbour and Kempsey in northern and central
(1972:pI. 41, 1978:pI. 73, 1982:pI. 323). A photo- New South Wales (Munro 1943, 1958a; Serventy
graph was included by Lewis (1981:18). 1950; Whitley 1964a; Lewis 1981) and Gulf of
Papua along southern coast of Papua New Guinea
Biology. -At the end of summer in the Southern from Kerema to Port Moresby (Kailola 1974,
Hemisphere (December to April or May), large 1975). Previously referred to as 8. niphonius in
schools of 8. munroi move close inshore along the Australia (Munro 1943).

Material examined. - Total 19 (296-820 mm FL).

TABLE 22.-Summary of morphometric data of Scom-
beromorus munroi. FL = fork length, HL = head meas.: 19 (296-820): Papua New Guinea (12);
length. Australia: N. Territory (3); Queensland
Character N Min. Max. Mean SO (3, *8. munroi); W. Australia (1).
Fork length 19 296 820 517 177 counts: 19.
Snout-A %. FL 17 521 572 547 12
Snout-20 r. FL 17 507 545 527 10 diss.: 4 (373-800): New Guinea (3); Queens-
Snout-10 %. FL 17 196 237 222 10 land (1).
Snout-P2 %. FL 17 228 267 249 10
Snout-P, %. FL 17 178 222 202 11
P,-P2 %. FL 18 95 126 105 7
Head length %. FL 18 176 213 198 8
Max. body depth r. FL 19 169 203 189 9 Scomberomorus niphonius (Cuvier)
Max. body width r. FL 15 85 122 100 9 Japanese Spanish Mackerel
P, length %. FL 14 100 126 108 7
P21ength %. FL 15 48 59 54 4 Figure 62
P2 insertion-vent %. FL 16 261 308 282 14
P2 tip-vent r. FL 13 200 244 225 12
Base 10 r. FL 17 281 322 307 12 Cybium niphonium Cuvier in Cuvier and Valen-
Height 20 %. FL 12 99 125 111 7
Base 20 r. FL 17 106 128 116 8 ciennes 1831:180-181 (original description based
Height anal r. FL 12 93 120 108 8 on a figure of a specimen from Japan). Tem-
Base anal r. FL 18 87 125 105 9
Snout (fleshy) %. FL 18 70 85 76 4 minck and Schlegel 1844:101-102 (description),
Snout (bony) %. FL 18 63 77 70 4
Maxilla length r. FL 18 91 115 103 5
pI. 53, fig. 2 (color painting of adult). Richard-
Postorbital r. FL 19 73 100 90 6 son 1846:268 (Sea ofJapan). GUnther 1860:371
Orbital (fleshy) r. FL 19 20 36 26 4
(description after Cuvier and Temminck and
Orbital (bony) r. FL 19 30 50 39 6
Interorbital r. FL 19 52 63 56 3 Schlegel). Gunther 1880:66 (Inland Sea, Ja-
2D-caudal r. FL 19 435 521 468 25
Head length 18 62 151 98 29 pan). Kitahara 1897:3 (description), fig. 11.
Snout (fleshy) %. HL 18 354 426 386 19 Kishinouye 1915:10 (description), pI. 1, fig. 4.
Snout (bony) %. HL 18 314 389 351 19
Maxilla length r. HL 18 486 544 521 16 *Kishinouye 1923:421-424 (description, biolo-
Postorbital %. HL 18 390 496 456 29
Orbit (fleshy) r. HL 18 113 174 134 18
gy), pI. 15, fig. 6 (soft anatomy in color); pI. 16,
Orbit (bony) r. HL 18 163 242 199 23 fig. 9 (transverse section of vertebrae); pI. 20,
Interorbital %. HL 18 256 318 282 16
fig. 32 (adult); pI. 24, fig. 41 (skull and vertebral
649
 FISHERY BULLETIN: VOL. 82, NO.4

FIGURE 62.-Scomberomorus niphonius. Japan, 719 mm FL, USNM 268909.

column). Boeseman 1947:95-96 (specimens in pectoral fin). Kusaka 1974:146 (urohyaD, fig.
Burger's collection). 269. Anonymous 1975:184 (description, range),
Cybium gracile Gunther 1873:378-379 (original map of fishery, color fig. Masuda et al. 1975:79
description; Cheefo, China). Morice 1953:37 (fig. G, color photograph), 256 (range). Uyeno
(tongue smooth). and Fuji 1975:14 (characters ofcaudal complex).
Scomberomorus niphonius. Steindachner and Shiino 1976:231 (common name). Klawe 1977:
Doderlein 1884:180-181 (description; Japan). 2 (common name, range). Collette 1979:24
Jordan et a1. 1913:121 (Japanese common (characters, range). Collette and Russo 1979:
names). Kamiya 1922:14-15, 26-32 (eggs and 13 (diagnostic characters, range). Liu 1981:
larvae; Ja,pan), pI. IV-V, figs. 1-24 (develop- 129-137 (age and growth). Zhang and Zhang
mental sequence of eggs and larvae). Kamiya 1981:104 (range in part). *Liu et al. 1982:170-
1924:35 (eggs). Tanaka 1927:154-157 (descrip- 178 (age and growth; Yellow Sea). *Wang
tion from 530 mm specimen from Tokyo mar- 1982:51-55 (catch, length-weight, management;
ket), pI. 42, figs. 163 and 164, pI. 44, fig. 173. Yellow Sea). Cressey et al. 1983:264 (host-
Soldatov 1929:5 (listed). Munro 1943:67-68, 71 parasite list, 4 copepod species). Lee and Yang
(subgenus Sawara). Fraser-Brunner 1950: 1983:230 (Taiwan), fig. 20 (619 mm FL). Col-
157-158 (description, range; "Scomber" gracile a lette and Nauen 1983:72-73 (description, range),
synonym of S. niphonius), fig. Okada 1955: fig. Ye and Zhu 1984 (bioeconomics).
149 (description, range, biology), fig. 136 (after Sawara niphonia. Jordan and Hubbs 1925:214
Kishinouye). Mori 1956:23 (Kasumi, Hamada; (new genus Sawara; specimen from Kobe mar-
S. Japan Sea). Mito 1960:79, 93 (eggs compared ket). Reeves 1927:9 (Ningpo, Peitaiho, China;
with those of S. commerson). Mito 1961:457 listed). Mori 1928:5 (Fusan, Korea). Solda-
(eggs and larvae). Tominaga 1964:vol. 1, pI. tov and Lindberg 1930:112 (synonymy, descrip-
198 (figure; anatomy), vol. 3:256-257 (descrip- tion, range). Suyehiro 1942:123-124 (intestine
tion, habits, distribution). Jones and Silas straight), fig. 78 (pyloric caeca). Honma 1952:
1964:52-54 (description, synonymy, range; in 143 (Echigo Provo = Niigata Pref., Japan).
part, only Chinese and Japanese specimens). Mori 1952:136 (Fusan, Masan, Quelpart I., Ko-
Gorbunova 1965a:53 (spawning season). Sha rea; listed). Nalbant 1970:58 (Kamchatka?).
et a1. 1966:1-12 (eggs and larvae), figs. Mito Scomberomorus gracileus (sic). Reeves 1927:8
1966:22-23 (fig. 15a, egg), 46-47 (fig. 26, lar- (Cheefo, Chinwangtao, China; listed).
va). Mito 1967:41 (vertical distribution of lar-
vae). *Hamada and Iwai 1967:1013-1020 (fish- Types of nominal species. -Cybium niphonium
ing seasons, length-weight, growth; Inland Sea Cuvier in Cuvier and Valenciennes 1831 was
of Japan), pI. 1 (otoliths). Kamohara 1967:43 based on a figure of a specimen from Japan,
(description, range in part), color pI. 22, fig. no type-specimens extant (Blanc and Bauchot
2. Tokida and Kobayashi 1967:158 (identifica- 1964:449).
tion of C. niphonium from Uchimura's unpub- Cybium gracile Gunther 1873. Holotype:
lished 1884 manuscript). Kim 1970:37-40 (age BMNH 1873.9.23.4; Cheefoo, N China; R. Swin-
determination; Korea). Uyeno 1971:79 (Japan hoe; 547 mm FL; D XX+16+IX; A 18+ VIII; P 1
Sea). Richards and Klawe 1972:14 (range), 93 22-23; RGR 1 2 + 1 + 10 = 13; vertebrae 22 + 27 = 49.
(references to eggs and larvae). Shiino 1972:71
(common name). Magnuson 1973:350 (short Diagnosis.-The only species of Scomberomorus
650
COLLETIE and RUSSO: SPANISH MACKERELS

with a straight intestine (Fig. 3k). The other Color pattern. -Kishinouye (1923:422) described
species have two or four loops. Scapular foramen S. niphonius as shining with a metallic lustre.
small as in S. guttatus and S. korean us. Intercalar Dorsum light greyish blue washed with green,
spine absent as in the six species of the regalis belly silvery. Seven or more rows of longitudinal
group. Supracleithrum wide, 55-62% oflength, as spots on the sides. Some spots connected together
in S. lineolatus. (Fig. 62). There are more numerous, smaller spots
than in S. munroi, about pupil size (Collette and
Description. -Lateral line gradually descending Russo 1980). Anterior quarter of first dorsal fin
to midline on caudal peduncle. Spines in first and a narrow distal margin ofthe rest ofthe dorsal
dorsal fin 19-21, usually 20 (Table 9); second dorsal fin black, most of basal membranes of posterior
fin rays 15-19, usually 16-18 (Table 10); dorsal three-quarters of fin white.
finlets 7-9, usually 8 (Table 10); anal fin rays 16- There are color paintings of S. niphonius in
20, usually 17 or 18 (Table 11); anal finlets 6-9, Kamohara (1967:pl. 22) and Anonymous (1975:pl.
usually 8 or 9 (Table 11); pectoral fin rays 21-23, 184), a color photograph in Masuda et al. (197q:79),
usually 22 (Table 12). Precaudal vertebrae 21-23, and a good black and white illustration in Kishi-
usually 22 (Table 6); caudal vertebrae 27 or 28, nouye (1923:fig. 32). We (Collette and Russo 1980:
usually 27 (Table 7); total vertebrae 48-50, usually fig. 1b) included the drawing that is presented
49 (Table 8). Gill rakers on first arch (2-3) + (9- here in the paper describing S. munroi.
12)=11-15, usually 2+(10-11)=12-13 (Table 5).
Morphometric characters given in Table 23. Biology.-There are two migrations in the Inland
Sea of Japan, a spawning migration in the spring
Size. -Maximum size 100 cm FL, 4.5 kg in weight (March to June) and feeding migration in the fall
(Kishinouye 1923). Age and growth studies have (September to November) according to Hamada
been published by H~mada and Iwai (1967), Kim and Iwai (1967). The spawning season in Japan is
(1970), and Liu et al. (1982). from April to May (Kishinouye 1923). The ripe

TABLE 23.-Summary of morphometric data of Scomberomorus niphonius. FL = fork length, HL =

head length.

Japan and Korea China Total

Character N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO
Fork length 13 157 788 439 200 16 105 623 280 167 32 105 788 351 197
Snout-A %. FL 11 539 574 553 10 16 558 605 573 14 30 484 605 563 22
Snout-2D %. FL 12 505 563 535 17 16 523 576 542 13 31 427 576 536 25
Snout-lO r. FL 12 202 271 239 21 16 224 278 256 16 31 202 278 248 20
Snout-P, %. FL 12 227 307 255 23 15 232 305 269 20 29 227 307 263 22
Snout-P, %. FL 12 184 252 216 20 16 197 264 232 18 31 184 264 225 20
P,-P, %. FL 12 89 125 100 11 13 95 129 109 12 26 89 129 105 12
Head length %. FL 13 180 249 207 18 16 192 252 223 18 32 180 252 215 19
Max. body depth r. FL 11 148 203 167 17 11 152 190 172 13 23 148 211 171 17
Max. body width r. FL 10 77 95 88 6 12 71 92 80 7 25 71 103 84 8
P, length r. FL 12 87 122 105 9 16 104 125 115 6 31 87 125 111 9
P,length r. FL 11 51 76 62 7 15 56 127 74 16 29 45 127 68 14
P2 insertion-vent %. FL 12 243 318 283 25 13 258 321 290 18 27 232 321 285 23
P, tip-vent %. FL 11 186 256 221 23 12 188 245 216 17 25 186 256 218 20
Base 10 %. FL 12 271 320 293 17 16 259 310 280 14 31 219 320 282 20
Height 20 r. FL 12 66 110 92 10 13 95 112 101 5 28 66 124 98 10
Base 20 r. FL 12 90 131 104 12 16 94 132 113 11 31 90 189 113 19
Height anal %. FL 9 84 108 92 7 15 63 106 97 11 27 63 124 97 11
Base anal %. FL 11 92 117 105 9 16 93 135 106 11 30 92 147 107 12
Snout (fleshy) r. FL 13 66 88 78 7 16 70 95 84 7 32 66 95 81 8
Snout (bony) r. FL 13 59 83 72 7 16 62 89 77 7 32 56 89 75 8
Maxilla length %. FL 13 94 136 114 13 16 103 145 125 11 32 93 145 119 14
Postorbital %. FL 13 91 104 98 4 16 89 115 104 7 31 89 115 102 7
Orbital (fleshy) %. FL 13 19 41 28 6 16 25 51 36 7 32 19 51 33 8
Orbital (bony) r. FL 13 29 59 41 9 16 36 65 51 8 32 29 65 47 10
Interorbital r. FL 13 51 63 56 4 16 54 61 57 2 32 51 63 57 3
2D-caudal r. FL 12 412 490 459 27 15 420 488 468 16 29 412 490 465 21
Head length 13 39 142 88 34 16 27 120 60 30 32 27 142 72 35
Snout (fleshy) r. HL 13 351 401 377 16 16 362 393 378 11 32 345 401 376 14
Snout (bony) %. HL 13 314 373 346 15 16 325 365 348 12 32 294 373 348 16
Maxilla length %. HL 13 504 582 549 23 16 538 582 560 14 32 484 582 553 22
Postorbital r. HL 13 417 511 477 26 16 405 518 470 26 31 405 518 473 26
Orbit (fieshy) %. HL 13 103 170 136 19 16 122 204 158 21 32 103 204 150 23
Orbit (bony) %. HL 13 161 250 194 28 16 188 257 230 23 32 161 257 215 30
Interorbital r. HL 13 248 294 271 14 16 226 285 258 18 32 226 303 264 18

651
 FISHERY BULLETIN: VOL. 82, NO.4

eggs are large, about 1.5 mm in diameter and recently described S. munroi (Collette and Russo
number about 550,000-870,000 (Kishinouye 1923). 1979).
Immature fish of about 30 mm are found in April
and May (Kishinouye 1923). Eggs and larvae up to Geographic variation. -Morphometric characters
35 mm TL are described by Sha et al. (1966) from were compared for two populations of S. niphonius
plankton net samples from Kiaochow Bay, Tsing- by ANCOVA (Table 23): Japan and Korea (n = 9-
tao, China. Although it feeds on small fishes 13) and China (n = 11-16), Null hypotheses that
(Kishinouye 1923), no detailed food studies seem the 2 sets of regression lines are coincident were
to have been published. accepted for 21 regressions and rejected for 5 sets:
Sn-A, Sn-1D, maximum body width, PlL, and
Interest to fisheries. -In the Inland Sea of Japan, orbit (bony). No meristic differences were found
the main fishing seasons are from March to June between the two populations.
and from September to November (Hamada and
Iwai 1967). Angling and gill nets are important Material examined. -Total 38 (86.5-788 mm FL).
gear in this region. There are also important
fisheries in the Huanghai Sea (Yellow Sea) and meas.: 31 (97.5-705): Japan (8); Korea (5); China
Bohai Sea (Liu et al. 1982). Ye and Zhu (1984) have (16); unknown locality (2).
developed a bioeconomic model for this fishery, counts: 38.
estimating maximum revenue, optimum econom- diss.: 2 (683-788): Japan.
ic effort, and optimum energy consumption. The
annual catch reported by China, Japan, and Korea
varied from 60,733 to 77,356 t between 1979 and Scomberomorus plurilineatus Fourmanoir
1982 (FAO 1984). Queen Mackerel or Kanadi Kingfish
Figure ?3
Distribution. -Confined to temperate and sub-
tropical waters of the western North Pacific, Cybium lineolatum. Not of Cuvier 1831. Gil.-
Japan, Korea, and northern China (Fig. 58). The christ and Thompson 1911:41 (description;
northernmost locality is Vladivostok, U.S.S.R., in Durban).
the Sea of Japan (BMNH 1893.1.27.10-12). In Scomberomorus lineolatum. Not of Cuvier 1831.
Japan, it is found from southern Hokkaido to Gilchrist and Thompson 1917:395 (Natal).
Honshu, Shikoku, and Kyushu, west to Pusan, Scomberomorus lineolatus. Not of Cuvier 1831.
Korea (CAS SU 31263), and Ningpo, Peitaiho Barnard 1927:803 (description; Natal). Fowler
(Reeves 1927; ZMA 114.597), Cheefo (= Yentai) 1934:441 (Durban). Smith 1935:210-211 (de-
(Gunther 1873; BMNH 1873:9.23.40; UMMZ scription; Port Alfred, South Africa). *Wil-
167374), and Tsingtao (USNM 130474) on the liams 1960:183-192 (description, synonymy,
Shantung Peninsula of northern China. Records range), pI. 2. *Williams 1964:151-154 (distri-
of S. niphonius from northern Australia and bution, fishery biology). Merrett and Thorpe
southern Papua New Guinea are referrable to the 1966:371-372 (references, range, size, biology).

FIGURE 63.-Scomberomorus plurilineatus. Durban, South Africa, 598 mm FL, USNM 264809.

652
COLLE'ITE and RUSSO: SPANISH MACKERELS

Scomberomorus leopardus. Not of Shaw 1803. Description. -Lateral line gradually descending
Fowler 1929:254 (description; Natal). Smith to midline on caudal peduncle. Intestine with two
1949, 1953, 1961:301 (description, range), fig. folds and three limbs (Fig. 31). Spines in first
841, pI. 64. Morrow 1954:815 (near Shimoni dorsal fin 15-17, usually 15 or 16 (Table 9); second
and Pemba I., East Africa). Talbot 1965:469 dorsal fin rays 19-21, usually 20 (Table 10); dorsal
(Mafia area, Tanganyika). Mauge 1967:234 finlets 8-10, usually 9 (Table 10); anal fin rays 19-
(Anakao, Thlear region, Madagascar). Shiino 22, usually 20 or 21 (Table 11); anal finlets 7-10,
1976:231 (common name). usually 8 or 9 (Table 11); pectoral fin rays 21-26,
Scomberomorus sp. Williams 1956:44 (Kenya). usually 22 or 23 (Table 12). Precaudal vertebrae 19
Scomberomorus guttatus. Not of Bloch and or 20, usually 20 (Table 6); caudal vertebrae 25-27,
Schneider 1801. Smith 1956:722 (Aldabra). usually 26 (Table 7); total vertebrae 45 or 46,
Smith and Smith 1963:43 (Seychelles), pI. usually 46 (Table 8). Gill rakers on first arch (2-
30B. Smith 1964:176-177 (description; Durban 3)+(9-13)=12-15, usually 2+(10-11)=12-13 (Ta-
and Delagoa Bay), pI. 8, figs. 3-5. Silas 1964: ble 5). Morphometric characters given in Table 24.
314-328 (western Indian Ocean population
only). Smith and Smith 1966:72 (Natal), color TABLE 24.-Summary of morphometric data of Scom-
pI. 841. beromorus plurilineatus. FL = fork length, HL = head
Cybium leopardus. Not of Shaw 1803. Four- length.
manoir 1957:227 (description; Mozambique Character N Min. Max. Mean SO
Channel). Fork length 37 144 910 547 211
Cybium lineolatus. Not of Cuvier 1831. Four- Snout-A %. FL 36 478 614 502 22
Snout-20 r. FL 36 446 622 473 28
manoir and Crosnier 1964:387-388 (Mada- Snout-1D %. FL 36 202 247 221 12
gascar). Snout-P, r. FL 36 207 261 232 12
Snout-P, %. FL 37 176 228 192 11
Scomberomorus plurilineatus Fourmanoir 1966: P,-P, r. FL 26 96 117 103 5
Head length %. FL 37 175 222 193 11
223-226 (original description; Madagascar), fig. Max. body depth %. FL 32 184 225 205 11
1. Klawe 1977:2 (range, common name). Col- Max. body width %. FL 20 76 123 97 13
P, length r. FL 31 98 140 123 10
lette 1979:29 (characters). Collette and Russo P, iength %. FL 35 44 66 51 6
1979:13 (diagnostic characters, range). * Van P2 insertion-vent r. FL 24 223 257 244 9
P, tip-vent r. FL 23 158 207 187 15
der Elst 1981:275 (description, natural history, Base 10 %. FL 23 219 256 240 11
Height 20 %. FL 29 122 166 148 11
range, photograph). Joubert 1981:5 (minor Base 20 %. FL 27 101 156 128 11
component of shore angler's catches; Natal, Height anal r. FL 27 91 155 135 14
South Africa). Cressey et aI. 1983:264 (host-
Base anal r. FL 27 111 143 126 9
Snout (fleshy) %. FL 35 61 75 67 3
parasite list, 5 copepod species). Collette and Snout (bony) %. FL 25 48 67 58 4
Maxilla length %0 FL 36 83 113 96 7
Nauen 1983:73-74 (description, range), fig. Postorbital %. FL 36 81 107 94 6
Orbital (fleshy) r. FL 37 22 139 34 19
Orbital (bony) r. FL 27 33 60 45 8
Types. -Scomberomorus plurilineatus Fourma- Interorbital r. FL 36 46 67 56 4
20-caudal %. FL 25 518 575 549 15
noir 1966 was based on a 740 mm specimen Head length 37 31 159 104 37
collected near Nossi-Be, Madagascar, in 1965. The Snout (fleshy) r. HL 35 309 371 348 13
type was supposed to be transferred from the
Snout (bony) r. HL 25 256 327 305 15
Maxilla length r. HL 36 469 529 496 14
O.R.S.T.O.M. collections to the MNHN collec- Postorbital r. HL 36 413 527 485 23
Orbit (fleshy) r. HL 37 121 769 179 104
tion but apparently was inadvertently discarded Orbit (bony) %. HL 27 178 276 231 27
(M.-L. Bauchot and P. Fourmanoir 7 ). Interorbital %. HL 36 253 352 290 21

Diagnosis. -The only species of Scomberomorus

that has a pattern of short wavy lines and spots on Size. -Maximum size 120 cm FL, South African
its sides (Fig. 63). Other species have straight angling record 10.0 kg, sexual maturity attained
lines, spots, blotches, or bars on the side or are at about 80 cm FL (van der Elst 1981).
plain. Posterior end of maxilla greatly expanded
as in S. lineolatus and S. semifasciatus. Color pattern. -Williams (1960) published a good
description of fresh specimens from Zanzibar.
7M._L. Bauchot, Curator of Fishes, Laboratoire d'Ichtyologie
Generale et Appliquee, Museum National d'Histoire Naturelle, Head blue-grey above, silvery white below, except
43 Rue Cuvier, 75231 Paris Cedex 05, France, and P. Fourmanoir, for lower jaw tip, preorbital area and maxillary
Office de la Recherche Scientifique et Technique Outre-Mer,
Institut Francais d'Oceanie, Noumea, B.P. 4, Nouvelle-Cale-
groove dusky to black. Pupil of eye black, rest
donie, pers. commun. December 1974. silvery. Body iridescent blue-grey above lateral
653
 FISHERY BULLETIN: VOL. 82, NO.4

line, silver below becoming whitish ventrally. A gill net prevails (Williams 1964). On the west
series of about six to eight interrupted horizontal coast of Zanzibar a trap net called the mensab is
black lines on sides of body much narrower than used to intercept fish on their projected paths of
interspaces. Anteriorly, usually only one of these movements (Williams 1964). More recently, tuna
lines above lateral line; replaced posteriorly by a purse seines are used in Zanzibar with catches of
number of short oblique black lines becoming several tons reported off the northwest coast
somewhat confused, and only two or three con- (Merrett and Thorp 1966). In Natal, South Africa,
tinue through to caudal peduncle. Horizontal it is a popular gamefish with ski-boat fishermen
black lines on body interrupted to varying degrees, and also with spearfishermen (van der Elst 1981).
beginning almost intact in places, but broken up
into a series of small rectangular "spots" in others. Distribution. -Common in coastal waters, espe-
Juveniles have spots but develop adult pattern of cially near rocky and coral reefs. Western Indian
interrupted lines by the time they reach a length Ocean along the coast of East Africa from Kenya
of 400 mm (Smith 1964:177). Upper areas of caudal (lat. 1 30' S) and Zanzibar (Williams 1964) to
0

peduncle and median keel black, lower areas Natal, South Africa (Fig. 58). The southernmost
dusky. First dorsal fin black except lower areas of records are from Algoa Bay (Smith and Smith
membrane may be pale posteriorly. Second dorsal 1966). Also found in the Seychelles Islands (Smith
fin with leading edge and tips of rays dusky, rest and Smith 1963) and along the west coast of
silver to pale; finlets dusky with a silver area at Madagascar.
center. Anal fin, leading edges and tips of rays
dusky, rest silvery; finlets white with a dusky Material examined. -Total 37 (165-910 mm FL).
central area. Pectoral fins black inside, as is axil;
dusky outside with edges black; pelvic fins pale meas.: 37 (165-910): Natal, South Africa (25);
whitish with outside of midrays dusky, groove on Mozambique (1); Kenya (1); Zanzibar (10,
body a little dusky. Caudal fin basally pale, rest of F. Williams' data).
fin dusky to black. counts: 37.
Black and white photographs of S. plurilineatus diss.: 5 (490-910): South Africa (4); Kenya (1).
have been published by Williams (1960:pI. 2, 640
mm Zanzibar specimen) and Fourmanoir (1966:
fig. 1, 740 mm holotype from Madagascar). Illus- Scomberomorus queenslandicus Munro
trations of a spotted 300 mm juvenile and two Queensland School Mackerel
adults over 1 m long were presented by Smith Figure 64
(1964:pI. 8). A colored figure of the juvenile is
included in Smith and Smith (1966:fig. 841). Cybium guttatum. Not of Bloch and Schneider
1801. Macleay 1880:559 (description; Port
Biology. -Large schools are present in the Zanzi- Jackson, Australia). Ogilby 1887:30 (listed;
bar Channel from March-April until August-Sep- Port Jackson).
tember, average weight 3.2-3.5 kg (Williams 1960). Scomberomorus guttatus. Not of Bloch and
Angling statistics point to a peak abundance in Schneider 1801. Waite 1904:42 (New South
Natal, South Africa, during May (van der Elst Wales). Stead 1906:165-166 (N.SW.). Stead
1981). Spawning probably takes place in August- 1908:98 (description; N .S.W). McCulloch
September in the Zanzibar Channel (Williams 1922:105 (N.S.W). McCulloch 1929:264-265
1964). There do not appear to be any published (range in part; Queensland, N.S.W).
references to eggs or larvae of S. plurilineatus. Scomberomorus (CybiumJ queenslandicus Munro
This species feeds mainly on anchovies (Ancho- 1943:82-86 (original description; Queensland
viella sp.), clupeids (Amblygaster sp., Sardinella and west Australia), pI. 7, fig. B, pI. 8, fig.
fimbrata, S. perforata), other small fishes, squids, 1. Coates 1950:24 (description), fig. Roughley
and mantis shrimps (Williams 1964; Merrett and 1951:110 (description), pI. 45, top fig. (after
Thorp 1966; van der Elst 1981). Munro). Jones and Silas 1962:202 (may turn
up in Indian waters), fig. 8 (after Munro).
Interest to fisheries. -In the Malindi area of Jones and Silas 1964:61-62 (description, range),
Kenya, catches of S. plurilineatus are mainly fig. 11 (after Munro). Taylor 1964:282 (listed
made by trolling and hand lines, while in the after Whitley 1954). Marshall 1964:363-364
Zanzibar Channel all methods are used but the (description; Qld.), pI. 49, fig. 350 A and B (after
654
COLLETTE and RUSSO: SPANISH MACKERELS

FIGURE 64.-Scomberomorus queenslandicus. Exmouth Gulf, Western Australia, 635 mm FL, USNM 268910.

Munro). Marshall 1966:205 (Qld.), pI. 49, fig. spots (larger than the diameter of the eye) on its
350 A and B (after Munro). Richards and sides (Fig. 64). In having few gill rakers (3-9), it is
Klawe 1972:14 (range), 94 (references to juve- superficially similar to S. commerson but differs
niles). Magnuson 1973:350 (short pectoral in lacking an abrupt downward curve in the
fin). Kailola 1974:71 (description; Gulf of lateral line under the second dorsal fin and in hav-
Papua; range extension). Kailola 1975:237 ing more vertebrae (48 or 49 vs. 42-46). Postero-
(specimen in Kanudi Fisheries collection). dorsal spine of hyomandibula large as in S. com-
Shiino 1976:231 (common name). Klawe 1977: merson and Acanthocybium. Ventral process of
2 (common name; range). Kailola and Wilson angular long, 117-126% of dorsal process, as in
1978:35 (trawled in Gulf of Papua), 60 (number S. commerson and Acanthocybium. Intercalar
of fin rays). Collette 1979:29 (characters, spine well developed as in S. cavalla and S.
range). Collette and Russo 1979:13 (diagnostic commerson.
characters, range). Grant 1982:624 (descrip-
tion, fishery in S Qld.), 625 (color pI. 324). Description. -Lateral line gradually descending
Rainer and Munro 1982:1046 (inshore group, to midline on caudal peduncle. Intestine with two
Gulf of Carpentaria), 1050-1051 (avoids low folds and three limbs (Fig. 3m). Spines in first
salinity areas in the southern Gulf). Cressey dorsal fin 16-18, usually 17 (Table 9); second dorsal
et aI. 1983:264 (host-parasite list, 5 copepod fin rays 17-19 (Table 10); dorsal finlets 9-11, usu-
species). Collette and Nauen 1983:74-75 (de- ally 9 or 10 (Table 10); anal fin rays 16-20, usually
scription, range), fig. Jenkins et aI. 1984:348- 19 (Table 11); anal finlets 9-11, usually 10 (Table
351 (193 larvae, 3.5-9.9 mm SL; off Townsville, 11); pectoral fin rays 21-23, rarely 25 (Table 12).
Qld.), fig. 4 (6 larvae, 3.6-9.5 mm SL). Precaudal vertebrae 19 or 20, usually 20 (Table 6);
Cybium queenslandicum. Whitley 1947:129 (W caudal vertebrae 28 or 29, usually 28 (Table 7);
Australia). Whitley 1948:24 (W Australia). total vertebrae 48 or 49, usually 48 (Table 8). Gill
Whitley 1954:27 (Parry Shoal, N. Territory). rakers on first arch (0-2) + (3-8) = 3-9, usually
Whitley 1964a:251-252 (description; W Aus-· 1+(5-6)=6-7 (Table 5). Morphometric characters
tralia and N. Territory). Whitley 1964b:48 given in Table 25.
(listed).
Cybium queenslandicus. Munro 1958a:112 (de- Size. -Maximum size 100 cm FL, 8 kg in weight,
scription, range), fig. 750 (after Munro). Grant commonly 50-80 cm (Lewis 1981).
1965:174 (description after Munro; Moreton Bay,
Queensland), fig. Grant 1972:103, 1975:161, Color pattern. -In his original description of the
1978:194 (description after Munro; fishery in S species, Munro (1943) provided a good description
Qld.), fig. of freshly caught specimens from Queensland.
Cranial regions and upper part of back iridescent
Types. -Holotype: QM 1.6588; Cape Cleveland, N bluish green, cheeks and belly silvery white. In
Queensland, Australia; G. Coates; 463 mm FL; D adult fish, sides marked with about three indefi-
XVII + 18+IX;A20+IX;P1 23;RGR1 1+ 1+4=6. nite rows of indistinct bronze-grey blotches, each a
little larger than orbit. Membrane of first dorsal
Diagnosis. -This species has relatively few large fin jet black with large contrasting areas of

655
 FISHERY BULLETIN: VOL. 82, NO.4

TABLE 25.-Summary of morphometric data of Scom- were described and illustrated by Jenkins et al.
beromorus queenslandicus. FL = fork length, HL = (1984).
head length.

Character N Min. Max. Mean SO Interest to fisheries. -Caught by recreational and

Fork length 28 156 641 392 123
Snout-A %0 FL 28 502 558 525 12
commercial line-fishermen trolling with lures and
Snout-20 r. FL 28 470 518 502 11 spoons, and using cut baits along the coast of
Snout-10 %0 FL 28 219 254 234 10
Snout-P, r. FL 27 234 273 252 11
Queensland (Grant 1982). They also form the basis
Snout-P, %0 FL 27 211 252 228 10 of a substantial net fishery, using set nets by day
P,-P, %0 FL 27 85 118 99 7
Head length %0 FL 28 203 245 220 10 or night. Considerable quantities of juveniles are
Max. body depth r. FL 25 161 207 188 11 trawled in parts of Moreton Bay during autumn
Max. body width %0 FL 26 81 118 101 11
P, length %0 FL 24 103 135 119 8 (March-May). Together with Grammatorcynus
P, length %0 FL 23 44 62 55 5
P2 insertion-vent %0 FL 24 233 275 254 12
and three other species of Scomberomorus, mack-
P, tip-vent r. FL 22 178 222 198 13 erel fishery is Queensland's second major finfish-
Base 10 r. FL 28 239 283 263 11
ery with an annual output of about 1,000 tons of
Height 20 %0 FL 18 89 136 114 12
Base 20 %0 FL 28 80 135 113 11 whole and filleted fish (Anonymous 1978). It was
Height anal r. FL 17 94 157 112 15
Base anal r. FL 28 89 141 108 12 known to south Queensland fishermen as school
Snout (fleshy) %. FL 28 78 97 86 4 mackerel for over 60 yr prior to its formal descrip-
Snout (bony) %0 FL 27 73 125 80 9
Maxilla length %0 FL 27 109 146 125 8 tion by Munro (1943). Trawled in the Gulf of
Postorbital r. FL 28 90 110 102 5 Papua.
Orbital (fleshy) %0 FL 28 23 41 31 5
Orbital (bony) %0 FL 27 39 66 49 8
Interorbital %0 FL 28 56 74 63 5
20-caudal r. FL 17 440 532 496 22
Distribution. -Confined to inshore coastal waters
Head length 28 36 141 85 25 of southern Papua New Guinea and the northern
Snout (fleshy) %0 HL 28 357 423 390 17
Snout (bony) %0 HL 27 331 554 363 41 three quarters of Australia (Fig. 55). The western-
Maxilla length %0 HL 27 526 597 568 17 most records are from Shark Bay (Munro 1943)
Postorbital r. HL 28 410 502 463 21
Orbit (fleshy) %0 HL 28 112 171 141 16 and Onslow (AMS IB.1576, WAM P. 8669-8678),
Orbit (bony) r. HL 27 183 283 222 29
Western Australia. The range extends south to
Interorbital %0 HL 28 259 330 287 17
Port Jackson (USNM 4795, AMS 1.15026) and
Botany Bay (USNM 47948) in the Sydney area. A
tentative record from Fiji (Collette and Russo
intense white between sixth and last spines. 1979) was based on small specimens of S. commer-
Second dorsal fin, finlets, and caudal fin pearly son (USNM 227183, 203-242 mm FL) with less dip
grey with darker margins. Pelvic fins, anal fin, in the lateral line than is usual in the species. The
and anal finlets white. Pectoral fins greyish, origin of a specimen of S. queenslandicus (USNM
darkest on inner surface. Munro also noted the 213539, 215 mm FL) that was purchased in the
absence of characteristic blotches in a 95 mm Anbon market in the Moluccas is unknown but
juvenile. joint venture trawlers which fish in the Arafua
Munro (1943:pl. 7, 8) provided excellent illustra- Sea unload at Anbon (Lewis footnote 5).
tions of adult (545 mm FL) and juvenile (140 mm
FL) specimens from Queensland. There is also a Geographic variation. -Comparisons were made
figure drawn by George Coates in Grant (1982 and ofmorphometric characters ofthree small samples
previous editions). A photograph of two juveniles of S. queenslandicus by ANCOVA: Western Aus-
(about 300 mm FL) was included in Lewis (1981: tralia (n = 5-9), eastern Australia (n = 9-13), and
15). A color photograph of a 230 mm specimen was southern New Guinea (n = 3-5). Null hypotheses
added to the fifth edition of Grant (1982:pl. 324). that the 3 sets of regression lines are coincident
were accepted for 24 sets of regressions and
Biology. -Schools move into bays and estuaries rejected for 2 sets: orbit (fleshy) and orbit (bony).
and inshore coastal waters the length of the entire In both cases, the Western Australian population
coast of Queensland during midwinter and early was significantly different by the Newman-Keuls
spring in the Southern Hemisphere (Grant 1982). Multiple Range Test from the population in east-
A female 450 mm FL with mature ovaries was ern Australia (intercepts 5.140, 2.847, Q = 5.630**
collected in Moreton Bay, Queensland, in January for fleshy orbit; intercepts 8.380, 1.951, Q =
1980 (Lewis, footnote 5). Information on eggs 4.562** for bony orbit) and the population in
(Richards and Klawe 1972) and on spawning and eastern Australia was not significantly different
food habits is lacking. Larvae (3.5-9.5 mm SL) from that in New Guinea. No significant meristic
656
COLLE'ITE and RUSSO: SPANISH MACKERELS

differences were found between populations. Seomberomorus regalis. Jordan and Gilbert
1882:426 (description, synonymy). Jordan and
Material examined. - Total 35 (156-641 mm FL). Gilbert 1883c:573 (Seomberomorus plumierii
identical with Scomber regalis). Goode 1884:
meas.: 28 (156-641): New South Wales, Austra- 316 (size; Fla. Keys), pI. 94. Meek and Newland
lia (5); Queensland (8, *S. queensland- 1884:233-234 (description, synonymy, range).
icus); W Australia (9); New Guinea (5); Jordan 1884:120 (description; Key West).
Anbon market (1). Dresslar and Fesler 1889:442 (in key), 444
counts: 35. (synonymy, range Cape Cod to Brazil), pI. 10.
diss.: 6 (354-641): Queensland (4); W Austra- Jordan and Evermann 1896b:875 (description,
lia (1); New Guinea (1). synonymy). Jordan and Evermann 1900:pl.
135, fig. 369 (Key West specimen). Evermann
and Marsh 1902:124 (description, synonymy in
Scomberomorus regalis (Bloch) part; Puerto Rico), fig. 28. Jordan and Ever-
Cero mann 1902:286-287 (description, range), fig.
Figure 65 Bean 1903:398-400 (synonymy, description,
range). Fowler 1905:766 (description; Santo
Scomber regalis Bloch 1793:38-43 (original de- Domingo). Smith 1907:192 (diagnosis; North
scription, after a drawing by Plumier; Martin- Carolina record from Yarrow 1877), fig. 78.
ique), color pI. 333. Bloch 1797:31-34 (French Sumner et aI. 1913:750 (references, occurrence;
translation of original description), color pI. Buzzards Bay, Mass.). Ribeiro 1915:135 (de-
333. Bloch and Schneider 1801:22 (description scription, range S to Angra dos Reis, Brazil).
after Bloch). Shaw 1803:583 (after Bloch). Meek and Hildebrand 1923:323-324 (descrip-
Lacepede 1803:711 (Scomberomore plumier is tion, synonymy). Schroeder 1924:7 (Fla. Keys),
the same as Scomber regalis). fig. 4. Nichols and Breder 1927:124 (descrip-
Seomberomorus Plumierii Lacepede 1802:292-294 tion, distribution, biology), fig. 171. Frost
(original description after Plumier's drawing; 1928:329-330 (sagittae), pI. 12, fig. 7 (sagitta).
Martinique). Beebe and Tee-Van 1928:97 (description; Port-
Cybium regale. Cuvier 1829:200 (listed in foot- au-Prince Bay, Haiti), fig. Hildebrand and
note from Se. regalis Bloch; Seomberomorus Schroeder 1928:205 (description, range in part,
plumieri a synonym). Cuvier in Cuvier and biology; Chesapeake Bay), fig. 116. Nichols
Valenciennes 1831:184-185 (description; Santo 1929:230 (range, description; Puerto Rico), fig.
Domingo). Castelnau 1855:23 (Bahia, Brazil). 83. Beebe and Hollister 1935:213 (Union I.,
Gunther 1860:372-373 (synonymy, description; Grenadines). Hubbs 1936:253 (description of 4
Jamaica). Poey 1865:322 (description; Santo juveniles, 36-50 mm, from Yucatan). Baugh-
Domingo). Kner 1865:144 (description; Rio de man 1941:17-18 (Texas records). Munro 1943:
Janeiro). Poey 1868:362 (description; Cuba). 67, 71-72 (placed in subgenus Seomberomo-
Poey 1875:147 (description; Cuba). Poey 1878: rus). Fowler 1945:186 (synonymy, description;
4 (synonymy, characters). Charleston, S.C.), 290 (Florida). La Monte

FIGURE 65.-Scomberomorus regalis. Key West, Fla., 625 mm FL, USNM 12527. (From Goode 1884:pl. 94.)

657
 FISHERY BULLETIN, VOL. 82, NO.4

1945:28 (description, range). Breder 1948: Bloch (color pI. 333) leaves no doubt as to the
127 (range), fig. Erdman 1949:301 (frequently identity ofthe species.
found in the West Indies). Fraser-Brunner
1950:160 (synonymy in part, range), fig. 32. Diagnosis.-The only species of Scomberomorus
Baughman 1950:244 (previous Texas records). that has a stripe on its sides (which may be broken
Rivas 1951:225 (synonymy, diagnosis, range). into several segments) with small dots above and
La Monte 1952:50-51 (description, range). below the stripe (Fig. 65). Other species have
Bigelow and Schroeder 1953:348 (description; lines, spots, blotches, or bars, or are plain. Scom-
one record from Gulf of Maine, Monomoy, Cape beromorus regalis possesses nasal denticles as do
Cod), fig. 183. Pew 1954:26, 28 (description, the other five species of the regalis group (brasil-
range, habits; 3-ft specimen from Port Aransas, iensis, concolor, maculatus, sierra, and tritor),
Tex.), fig. 24. * Erdman 1956:317 (range, has an artery that comes off the fourth left
spawning periods; Puerto Rico). Briggs 1958: epibranchial artery as do all the species in the
286 (range). * Mago Leccia 1958 (osteology, group except S. tritor, and shares a specialization
comparisons with S. maculatus and S. cavalla), of the fourth right epibranchial artery (Fig. 7g)
figs. Cervigon 1966:720-721 (description; Ven- with S. brasiliensis and S. sierra. In these three
ezuela). *Randall 1967:754 (food of 116 West species an artery connects the fourth right epi-
Indian specimens, 96.1% fishes). Bohlke and branchial with a branch ofthe coeliaco-mesenteric
Chaplin 1968:573 (description, range), fig. artery. Together with three other species of the
Randall 1968:117-118 (description, range, hab- regalis group (brasiliensis, concolor, and sierra),
its), fig. 134 (photograph of Virgin Is. specimen). S. brasiliensis has a long posterior process on the
Beardsley and Richards 1970:5 (length-weight pelvic girdle (Fig. 46a), 62-90% of the length ofthe
of 58 specimens, 213-835 mm FL, 0.13-4.88 kg; anterior plate. Intercalar spine absent as in the
Florida). Dahl 1971:278 (common in Colom- other five species of the regalis group and S.
bia), fig. Richards and Klawe 1972:14 (range), niphonius. Pectoral fin covered with scales.
94 (reference to Hubbs 1936). Miyake and
Hayasi 1972:111-3 (in key), IV-ll (common Description. -Lateral line gradually descending
names). Klawe 1977:2 (common name, range). to midline on caudal peduncle. Intestine with two
Erdman 1977:150 (spawn virtually all year; NE folds and three limbs (Fig. 3n). Spines in first
Caribbean). Collette et al. 1978:274-275 (com- dorsal fin 16-18, usually 17 (Table 9); second dorsal
parison with other American species of Scom- fin rays 16-19, usually 17 or 18 (Table 10); dorsal
beromorus ). Fritzsche 1978:133-135 (descrip- finlets 7-9, usually 8 (Table 10); anal fin rays 15-
tion, larval development), figs. 75-76 (larvae). 20, usually 18 or 19 (Table 11); anal finlets 7-10,
Collette 1978:Scombm 6 (description, range), usually 8 (Table 11); pectoral fin rays 20-24,
figs. Lima and Oliveira 1978:13, 24 (common usually 21 or 22 (Table 12). Precaudal vertebrae 19
name "sierra-penincho" in Brazil). Manooch or 20, usually 20 (Table 6); caudal vertebrae 28
et al. 1978 (annotated bibliography). Collette (Table 7); total vertebrae 47 or 48, usually 48
1979:29 (characters, range). Collette and (Table 8). Gill rakers on first arch (2-4) + (10-
Russo 1979:13 (diagnostic characters, range). 14) = 12-18, usually 3 + (12-13) = 15-16 (Table 5).
Sacchi et al. 1981:3 (French Antilles). Koster Morphometric characters given in Table 26.
1981:55 (Isla Rosario, Colombia). Cooper 1982
(fishery in Jamaica). Cressey et aI. 1983:264 Size. -Maximum size 83.5 em FL, 4.9 kg (Beards-
(host-parasite list, 7 copepod species). Garzon ley and Richards 1970). Sexual maturity in Flor-
and Acero 1983:18 (Isla Providencia, Colombia). ida is attained at about 350 mm FL for males, 380
Collette and Nauen 1983:75-76 (description, mm for females (Finucane and Collins 1984).
range), fig. Finucane and Collins 1984 (repro-
ductive biology, Fla.). Color pattern. -Bluish green on back, sides sil-
very, with a midlateral row of yellow streaks of
Types of nominal species. -Both Scomber regalis variable length and small yellow spots above and
Bloch 1793 and Scomberomorus plumierii La- below this row (Randall 1968). Distal part of
cepede 1802 are based on Plumier's drawing of a anterior lobe of first dorsal fin black, rest of fin
specimen from Martinique, and there are no white.
known type-specimens extant. The pattern of Bloch (1793:pl. 333) included a color plate in his
spots and lines in the color plate published by original description of Scomber regalis. There is a
658
COLLETTE and RUSSO: SPANISH MACKERELS

TABLE 26.-Summary of morphometric data of Scom- Interest to fisheries. -Taken commercially with
beromorus regalis. FL = fork length, HL = head gill nets and on lines in Florida, the West Indies,
length.
and the Bahama Islands. Also a valued spo~tfish
Character N Min. Max. Mean SO
taken by trolling with cut bait. It is taken by
Fork length 53 77 544 329 121
Snout-A %., FL 53 509 583 548 14
trolling and with hooks baited with live bait in
Snout-20 r. FL 53 489 560 520 14 Jamaica (Cooper 1982). Only 76-106 t identified as
Snout-1D %., FL 53 226 285 254 11
Snout-P, r. FL 49 210 306 264 16 S. regalis were reported from Fishing Area 31
Snout-P, r. FL 53 195 286 233 16 (Western Central Atlantic) in 1979-82, all from the
P,-P, %., FL 50 96 128 109 7
Head length %., FL 53 190 258 222 11 Dominican Republic (FAO 1984), but the actual
Max. body depth %., FL 49 113 231 197 18
Max. body width r. FL 45 56 176 91 18
catch is higher because an additional 985-1,108 t of
P, length %., FL 51 87 166 125 12 unidentified Scomberomorus was also reported
P,length %., FL 49 40 63 55 5
P2 insertion-vent %., FL 50 238 304 267 16
from this area and this is S. caualla and S. regalis.
P, tip-vent r. FL 48 187 252 210 17
Base 10 %., FL 53 227 283 258 12
Height 20 %., FL 45 75 145 115 11 Distribution. ~Most abundant in clear waters
Base 20 %., FL 53 94 135 116 9 around reefs in southern Florida, the Bahamas,
Height anal r. FL 44 84 138 112 10
Base anal %., FL 53 89 130 111 10 and West Indies (Fig. 49), but there are scattered
Snout (fleshy) %., FL 53 72 98 86 5 records from Cape Cod to Brazil. The northern-
Snout (bony) r. FL 52 64 88 78 5
Maxilla length %., FL 53 105 145 123 7 most records appear to be Monomoy at the south-
Postorbital r. FL 52 87 128 97 6
ern elbow of Cape Cod (Bigelow and Schroeder
Orbital (fleshy) %., FL 53 31 63 40 6
Orbital (bony) %., FL 53 40 73 55 6 1953:348) and Buzzards Bay on the south shore of
Interorbital %., FL 53 49 64 58 3
20-caudal %., FL 51 416 509 480 23 Cape Cod (Sumner et al. 1913). There are also
Head length 53 20 119 72 25 records from Chesapeake Bay (Hildebrand and
Snout (fleshy) r. HL 53 357 426 390 16
Snout (bony) %., HL 52 316 383 351 17 Schroeder 1928) and further south along the
Maxilla length roo HL 53 528 606 556 15 Atlantic coast of the United States. Several au-
Postorbital %., HL 52 376 569 440 26
Orbit (fleshy) roo HL 53 150 248 178 20 thors have reported occurrences in Texas (Baugh-
Orbit (bony) r. HL 53 199 292 247 21
man 1941, 1950; Pew 1954). Reports are scattered
Interorbital roo HL 53 220 286 262 13
from the northern coast of South America-Isla
Providencia, Colombia (Garzon and Acero 1983);
good black and white photograph of a 400 mm Colombia (Dahl 1971; USNM 94766), Venezuela
specimen from the Virgin Islands in Randall (Cervigon 1966; USNM 123081). The southern end
(1968:fig. 134) and a small color photograph in of the range is apparently about at Rio de Janeiro
Walls (1975:fig; 410). The drawing published by (Ribeiro 1915; BMNH 1903.6.9.80).
Goode (1884:pl. 94) is included here as Figure 65.
Material examined. -Total 60 (76.5-544 mm FL).
Biology. -Little is known about migrations or
movements of S. regalis. Young adults are taken meas.: 53 (76.5-544): Florida (3); Bahamas (6);
throughout the year in small numbers over the Cuba (5); Hispaniola (3); Jamaica (4);
Jamaica shelf (Cooper 1982). Around Puerto Rico, Puerto Rico (2); Virgin Is. (13); Antigua
spawning takes place virtually all year (Erdman (1); St. Eustatius (2); Martinique (1);
1977). Spawning takes place from April to October Barbados (5); Colombia (1); Curacao (1);
at California Bank, south of Jamaica (Cooper Venezuela (1); Brazil (3); Mexico (1).
1982). The majority offish appeared to be sexually counts: 60.
mature during most of the period between August diss.: 5 (456-544): Florida (2); Bahamas (3).
and October in the coastal waters of southern
Florida (Finucane and Collins 1984). Fecundity
estimates for 20 females 380-800 mm FL ranged Scomberomorus semifasciatus (Madeay)
from 161,000-2,234,000 (Finucane and Collins Broad-barred Spanish Mackerel
1984). The only published reference to eggs and Figure 66
larvae is by Fritzsche (1978:133-135) based on C.
A. Mayo's Ph.D. thesis. Food in the West Indies is Cybium semifasciatum Macleay 1884a:205-206
96% fishes, particularly small schooling clupeoids (original description; Burdekin R., Queensland).
(Harengula, Jenkinsia, and Opisthonema) and Macleay 1884b:28 (Burdekin R.). Whitley
atherinids (Allanetta) but also including squids 1936:40-42 (description in part; AMS IA.1598
and shrimps (Randall 1967). and IA.6573 = S. queenslandicus (Munro 1943:
659
 FISHERY BULLETIN: VOL. 82, NO.4

FIGURE 66.-Scomberomorus semifasciatus. Queensland, 451 mm FL. (From Munro 1943:pl. 6A.)

95); C. tigris a synonym of S. semifasciatum); vae, 3.3-10.5 mm SL; off Townsville, Qld.), fig. 2
fig. 3 (holotype of C. semifasciatum), fig. 4 (6 larvae, 3.8-10.5 mm SL).
(holotype of C. tigris). Indocybium semifasciatus. Fraser 1953:6 (abun-
Cybium tigris De Vis 1884:545 (original descrip- dant along NW coast of Western Australia).
tion; Cape York). Indocybium semifasciatum. Whitley 1947:129
Scomberomorus semifasciatum. McCulloch and (Western Australia). Whitley 1948:24 (W.
Whitley 1925:142 (Burdekin R.; after Macleay Australia). Whitley 1954:27 (between Darwin
1884a, b). and Pt. Charles, Northern Territory). Munro
Scomberomorus tigris. McCulloch and Whitley 1958a:112 (description, range), fig. 752. Whit-
1925:142 (Cape York; after De Vis 1884). ley 1964a:252 (description, range in part; ma-
McCulloch 1929:265 (Cape York; after De Vis terial from Qld., N. Territory, W. Australia).
1884). Whitley 1964b:48 (listed). Grant 1965:173
Scomberomorus semifasciatus. McCulloch 1929: (description after Munro), fig. Grant 1972:
264 (Burdekin R.; after Macleay 1884). *Mun- 106, 1975:164, 1978:194 (description after Mun-
ro 1943:91-95 (description, range, synonymy), ro; fishery in Qld.), fig.
pI. 6A (451 mm FL specimen from Mackay Dist.,
N Queensland), fig. 4 (45 and 58 mm juvs., Types of nominal species. -Cybium semifascia-
Townsville, N Qld.). Coates 1950:23 (descrip- tum Macleay 1884a. Holotype: AMS 1.18288;
tion), fig. Fraser-Brunner 1950:159 (descrip- lower Burdekin River, Queensland, Australia; A.
tion, range and synonymy in part). Roughley Morton; Aug. 1883; 285 mm FL; D ? + 20 + VIII; A
1951, 1953:110-111 (description), pI. 45, fig. C 21 + VI~I; P l 24-24; RGR l 2 + 1 + 6 = 9; vertebrae
(after Munro). Jones and Silas 1964:57-58 (de- 19+26=45.
scription, range), fig. 10 (after Munro). Taylor Cybium tigris De Vis 1884. Holotype: QM
1964:282 (description, references, specimens 1.119; Cape York, Queensland, Australia; K.
from Nightcliffe, N. Territory). Marshall Broadbent; 286 mm FL; D XIV + 19 + IX; A 21 +
1964:365-366 (description; Qld.), pI. 50, figs. 350 IX; P l 24-24; RGR l 2 + 1 + 7 = 10.
A & B (after Munro). Marshall 1966:205
(Qld.), pI. 50, figs. 350 A & B (after Mun- Diagnosis. -This species has wider lateral keels
ro). Richards and Klawe 1972:14 (range), 95 on the caudal peduncle than other species in the
(references to juveniles). Magnuson 1973:350 genus, but this is difficult to quantify and so is not
(short pectoral fin). Kailola 1975:237 (3 collec- a very useful diagnostic character. Superficially
tions from Gulf of Papua in Kanudi Fisheries similar to S. commerson in having prominent
collection). Shiino 1976:231 (common name). vertical bars on the sides (specimens over 700 mm
Klawe 1977:2 (common name, range). Collette FL lose their bars), but the bars are much wider
1979:29 (characters, range). Collette and (Fig. 66) than the many narrow bars of S. commer-
Russo 1979:13 (diagnostic characters, range). son. Also differs in lacking an abrupt downward
Grant 1982:630 (description, fishery in Qld.), curve in the lateral line under the second dorsal
631 (color pI. 328). Cressey et aI. 1983:264 fin, in having fewer spines in the first dorsal fin
(host-parasite list, 5 copepod species). Collette (15 or fewer vs. usually 16 or 17), and in having
and Nauen 1983:76-77 (description, range), more gill rakers on the first arch (6-13, usually 9
fig. Jenkins et aI. 1984:345, 348-351 (101 lar- or more vs. 1-8, usually 6 or less). Palatine tooth
660
COLLETfE and RUSSO: SPANISH MACKERELS

patch wider (Fig. 26a) than in any other species Queensland. Juveniles « 100 mm) with cranial
of Scomberomorus. Posterior end of maxilla ex- regions and upper regions of the back pale green
panded (Fig. 23a) as in S. lineolatus and S. with a bronze sheen and marked with 12-20 broad
plurilineatus. vertical dark grey bands. Bars confined to region
of body above lateral line, number increasing with
Description. -Lateral line gradually descending age. Cheeks and belly silver white. Snout dark
to midline on caudal peduncle. Intestine with two slate grey, patch of green above orbit. First dorsal
folds and three limbs (Fig. 30). Spines in first fin jet black with contrasting areas of white in
dorsal fin 13-15, usually 14 or 15 (Table 9); second central region. Second dorsal fin cream with
dorsal fin rays 19-22, usually 20 (Table 10); dorsal yellow anteriorly. Anal fin and finlets transparent
finlets 8-10, usually 9 (Table 10); anal fin rays 19- white. Caudal flukes creamy white at margins
22, usually 21 or 22 (Table 11); anal finlets 7-10, and dusky or blackish near hypuraI. Pectoral fins
usually 8 or 9 (Table 11); pectoral fin rays 22-25, dusky.
usually 23 or 24 (Table 12). Precaudal vertebrae With increase in size the bronze-green colora-
18 or 19, usually 19 (Table 6); caudal vertebrae 25- tion of the back turns greenish blue. The vertical
27, usually 26 (Table 7); total vertebrae 44-46, bands on the back are most marked in specimens
usually 45 (Table 8). Gill rakers on first arch < 500 mm and in larger fish there is a tendency for
(1-2) + (5-11) = 6-13, usually 2 + (7-9) = 9-10 (Ta- these markings to become less distinct, break into
ble 5). Morphometric characters given in Table 27. spots or fade out more or less completely. Above
700 mm, dead fish assume a drab greyish-yellow
Size.-Maximum size 120 cm FL, 10 kg (Lewis blotchy appearance with little or no evidence of
1981). markings. This uniform grey color apparently
accounts for the vernacular "grey mackerel" of
Color pattern. -Munro (1943) provided good de- Queensland fishermen as applied to older age
scriptions ofthe colors ofjuveniles and adults from groups of the species.
Munro included excellent illustrations of a 451
mm specimen and a 140 mm juvenile in his 1943
TABLE 27.-Summary of morphometric data of Scom- paper (pI. 6, 8). The illustration of the larger
beromorus semifasciatus. FL = fork length, HL = head individual has been reproduced in Grant (1982
length. and previous editions) and herein (Fig. 66). There
Character N Min. Max. Mean SD is a color photograph of a 470 mm specimen in
Fork length 31 142 840 344 217 Grant (1982:pI. 328).
Snout-A %. FL 31 477 533 505 16
Snout-2D %. FL 31 452 495 472 11
Snout-1D %. FL 30 212 261 245 12 Biology. - Little is known of the biology of this
Snout-P2 %. FL 29 195 273 249 18
Snout-P, %. FL 31 183 248 219 15 species other than it forms small schools. Juve-
P,-P2 %. FL 29 85 116 104 7 niles ranging in size from 45 to 100 mm are
Head length %. FL 31 180 236 212 14
Max. body depth %. FL 29 190 231 210 10 common along the beaches in the vicinity of
Max. bOdy width %. FL 29 71 121 94 13
P, length %. FL 30 132 166 147 10
Townsville, Queensland, during November and
P21ength %. FL 28 36 59 50 6 grow to twice this size by January (Munro 1943).
P2 insertion-vent %. FL 22 198 271 237 13
P2 tip-vent %. FL 19 168 210 187 10 Larvae (3.3-10.5 mm SL) were described and
Base 10 %. FL 30 170 236 210 14 illustrated by Jenkins et aI. (1984).
Height2D %. FL 25 112 177 160 13
Base2D %. FL 30 119 211 138 18
Height anal %. FL 23 135 220 156 17 Interest to fisheries. -Fish of 60-90 cm are caught
Base anal %. FL 29 124 168 145 11
Snout (fleshy) %. FL 30 72 89 81 4 on fishing grounds north of Yeppoon, Queensland,
Snout (bony) %. FL 30 63 80 72 4
Maxilla length %. FL 30 92 134 118 11
in November while smaller age groups are caught
Postorbital %. FL 29 84 109 95 6 in estuaries along the Queensland coast north of
Orbital (fleshy) %. FL 30 23 48 35 7
Orbital (bony) %. FL 31 34 63 51 8
Moreton Bay (Munro 1943). It is taken by setnet-
Interorbital %. FL 31 48 80 56 5 ting as well as by trolling and is popularly fished
2D-caudal %. FL 25 470 565 518 34
Head length 31 33 156 70 39 by Queensland anglers in small outboard-powered
Snout (fleshy) %. HL 30 355 409 379 17 boats trolling with small lures or cut bait (Grant
Snout (bony) %. HL 30 289 409 340 26
Maxilla length %. HL 30 496 596 554 21 1982). Together with Grammatorcynus and three
Postorbital %. HL 29 385 486 448 24
Orbit (fleshy) %. HL 30 122 202 162 24
other species of Scomberomorus, mackerel fishing
Orbit (bony) %. HL 31 176 269 237 24 is Queensland's second major finfishery with an
Interorbital %. HL 31 242 357 266 20
annual output of about 1,000 tons of whole and

661
 FISHERY BULLETIN: VOL. 82, NO.4

filleted fish (Anonymous 1978). Also trawled in Scomberomorus sierra Jordan and Starks
the Gulf of Papua. Sierra
Distribution. -This is the most estuarine of the Figure 67
species of Scomberomorus in Australia. It is
confined to estuarine and coastal waters of north- 8comberomorus maculatus. Not ofMitchill 1815.
ern Australia from Shark Bay, Western Australia, Jordan and Gilbert 1882:426 (in part; E. Pacific).
through the Northern Territory, and Queensland Jordan and Gilbert 1883a:106 (Mazatlan). Jor-
to northern New South Wales (Whitley 1964a:252, dan and Gilbert 1883b:110 (Panama). Meek
fig. 5f; Lewis 1981:17) and southern Papua New and Newland 1884:234 (synonymy in part; E.
Guinea (Fig. 55). Reports ofspecimens collected by Pacific). Dresslar and Fesler 1889:443 (synon-
D. G. Stead from Thailand and Malaya (Whitley ymy in part; E. Pacific). Evermann and Jen-
1964a:252) are based on misidentifications. kins 1891:128 (not previously reported N of
Mazatlan), 137 (important food fish; Guaymas,
Geographic variation. -Comparisons of morpho- Mexico). Meek and Hildebrand 1923:324-325
metric characters for three small samples of 8. (in part; Pacific coast of Panama; 8. sierra
semifasciatus were made with ANCOVA: Queens- considered a synonym of 8. maculatus). Herre
land (n = 3-4), Northern Territory (n = 3-5), and 1936:105 (description, several specimens; Albe-
New Guinea (n = 13-22). Null hypotheses that the marle 1., Galapagos; 8. sierra considered a
3 sets of regression lines are coincident were synonym of 8. maculatus). Fowler 1938:30-31
accepted for 23 sets, rejected for 3 sets: Base (synonymy in part, description; specimen from
1D, orbit (fleshy), and orbit (bony). The New- Charles 1., Galapagos). Hildebrand 1946:376-
man-Keuls Multiple Range Test showed that the 377 (synonymy in part; 14 specimens, 100-606
Queensland and New Guinea populations differed mm SL, from Gulf of Guayaquil; comparison of
significantly from each other in Base ill (slopes Atlantic and Pacific populations, "... at most
0.256 and 0.211, Q = 4.392**) and that the New different races for the opposite coasts"). Fra-
Guinea population differed from the Northern ser-Brunner 1950:159 (8. sierra considered a .
Territory population in fleshy orbit (slopes 0.020 synonym of8. maculatus). Bini and Tortonese
and 0.030, Q = 5.492**) and bony orbit (slopes 1955:32 (specimens from several localities in
0.034 and 0.046, Q = 7.523**). Peru). Ricker 1959:13 (Petalco Bay and San
BIas I., Mexico). Castro-Aguirre et a1. 1970:
Material examined.-TotaI34 (142-840 mm FL). 156 (Gulf of California). Sanchez T. and Lam
C. 1970:58-59 (length-weight; weights of body
meas.: 31 (142-840): Queensland, Australia (4, parts; diagram of vertebral column, 19 + 28 =
*C. semifasciatus, *8. tigris); N. Terri- 47); photograph. Leon 1973:22 (Puntarenas,
tory, Australia (5); New Guinea (22). Costa Rica). Amezcua-Linares 1977:10 (lagoon
counts: 33. system; Sinaloa, Mexico). Kong 1978:6-9 (An-
diss.: 6 (406-840): New Guinea. tofagasta, Chile). Yafiez-Arancibia 1980:111-

FIGURE 67. -Scomberomorus sierra. Gulf of California, 354 nun FL, USNM 217368.

662
COLLETIE and RUSSO: SPANISH MACKERELS

112 (Guerrero, Pacific coast of Mexico), pI. 29, 1957:306 (specimens taken with bait net and
fig.!. night light; Panama, Nicaragua). Collette et
Scomberomorus sierra Jordan and Starks in Jor- aI. 1963:53-54 (594 mm FL specimen, La Jolla,
dan 1895:428-429 (original description; Mazat- first California record, comparison with S. con-
lan, Mexico; also found in Panama). Jordan color), fig.!. Clemens and Nowell 1963:260 (19
and Evermann 1896a:341 (listed). Jordan and stations, Gulf of California, off Baja California,
Evermann 1896b:874-875 (description). Jor- Mexico, Costa Rica, Gulf of Panama, Gulf of
dan and Evermann 1902:286 (description). Gil- Guayaquil). Fitch and Craig 1964:202 (sagitta
bert and Starks 1904:68-69 (description; Pan- similar to that of S. concolor). Quiroga and
ama Bay; comparison with S. maculatus). Orbes 1964 (fishery; Esmeraldas Prov., Ecua-
Snodgrass and Heller 1905:360 (Albemarle I., dor). Klawe 1966:445-451 (occurrence ofyoung
Galapagos). Starks 1910:89-90 (osteology, ver- and spawning; E. Pacific), fig. 2 (11 mm speci-
tebrae 19 + 30 = 49). Kendall and Radcliffe mens). Fierstine and Walters 1968:4 (locali-
1912:96 (2 specimens from Panama Bay; de- ties), 12 (aspect ratio of caudal fin; vertebral
scription). Osburn and Nichols 1916:158 (Agua counts), fig. 12D (caudal fin complex). Lindsey
Verde Bay; Gulf of California). Evermann and 1968:1988-1990 (muscle temperature 1.15°C
Radcliffe 1917:55-56 (synonymy, description; higher than surface water temperature). Ma-
specimen from Paita, Peru). Starks 1918:120- tsumoto 1968:309-310 (jaw development com-
121 (description, occurrence), fig. 64. Higgins pared with that in Acanthocybium). Wollam
1920:33-34 (imported to San Diego from Mex- 1970:22 (larval differences between S. macula-
ico). Craig 1926:167 (used locally; Calexico, tus and S. sierra warrant recognition of S.
Mexico). Ulrey 1929:6 (Gulf of California). sierra). Erdman 1971:68 (gonads ripe late
Jordan et aI.1930:257 (listed). Croker 1933:14- Aug. to end of Nov., Gulf of Nicoya, Costa Rica;
15 (fishery), fig. 3, fig. 59 (yearly deliveries to S. sierra distinct from S. maculatus). * Artun-
San Diego and San Pedro by California boats duaga Pastrana 1976 (species synopsis, Colom-
fishing off Baja California). Breder 1936:11 bia). Miller and Lea 1972:192 (description,
(specimen taken by second "Pawnee" expedi- range), fig. Richards and Klawe 1972:14
tion). Walford 1937:24-25 (description, range, (range), 95 (references to larvae and juveniles).
angling notes), color pI. 39. Seale 1940:16-17 Magnuson 1973:350 (short pectoral fin). Sharp
(taken along the coast of Mexico and in the 1973:384 (electrophoretic patterns of hemoglo-
Galapagos Is.). Munro 1943:67, 71-72 (placed bin the same as in S. concolor). Shiino 1976:
in subgenus Scomberomorus). Fowler 1944: 231 (common name). Thomson and McKibbin
172-173 (synonymy, description, specimen from 1976:46 (description; Gulf of California), fig.
Balboa Harbor, Panama). Nichols and Mur- Klawe 1977:2 (common name, range). Collette
phy 1944:240 (Bay of Malaga, and Buenaven- et aI. 1978:274-275 (comparison with S. brasil-
tura, Colombia; more spots than in S. macu- iensis and other American species of Scornber-
latus). La Monte 1945:29 (common names, omorus). Horn and Allen 1978:41 (California
range). Eckles 1949:247-250 (description of 12 range lat. 33°N to 32°N). Collette 1979:29
juveniles, 21-71 mm FL, 9 from Costa Rica, 3 (characters, range). Collette and Russo 1979:
from the Gulf of California; vertebrae (19-20) + 13 (diagnostic characters, range). Phillips
28= (47-48), fig. 2 (21 mm specimen), fig. 3 (71 1981:54 (EI Salvador). Cressey et ai. 1983:264
mm specimen). Fitch and Flechsig 1949:278 (host-parasite list, 3 copepod species). Collette
(compared with S. concolor). Fitch 1950:70 and Nauen 1983:77-78 (description, range), fig.
(comparison of17 S. sierra with 30 S. concolor). Cybium concolor. Not of Lockington, 1889.
Clothier 1950:52-53 (vertebrae 47-49), pI. X Boulenger, 1899:3 (Golfe de Panama).
(outline of axial skeleton). Roedel 1951:510 Scomberomorus maculatus sierra. Chirichigno
(comparison with S. concolor). Mead 1951:121 1969:75 (common names, Ecuador, Peru, and
(2 ripe females, Gulf of California). Roedel Chile). Chirichigno 1974:325 (in key), fig. 641,
1953:85 (Mexican species; California records of 349 (range-S. Calif. to Bahia de Pisco, Peru
S. sierra probably refer to S. concolor). Clem- and Galapagos Is.).
ens 1956:76, 78 (14 postlarvae, 12-22 mm SL,
caught at lat. 08°06'N, long. 79°06'W; 2 sur- Types. -The original description was based on at
vived in shipboard aquaria for 13 d), fig. 2 least two specimens: "Types, 1720, L. S. Jr. Univ.
(postlarvae 16.5 and 54.0 mm SL). Clemens Mus.; the largest 24 inches long" (Jordan 1895:
663
 FISHERY BULLETIN: VOL. 82, NO.4

429). Bohlke (1953:105) considered SU 1720 to be ra has a longer pelvic fin (Fig. 48) than does S.
the holotype of the species but there is a specimen brasiliensis (4.7-6.4% FL vs. 3.6-5.9%) and lacks
in the British Museum (BMNH 1895.5.24.104) the lateral stripe that is the diagnostic feature of
which is also labelled as "type". To clarify the the pigment pattern of S. regalis. Together with
issue, we hereby select CAS SU 1720 as lectotype: three other members ofthe regalis group, S. sierra
Mazatlan, Mexico; D. S. Jordan; 332 mm FL; D has a long posterior process on the pelvic girdle,
XVIII+17+IX; A 19+VIII; PI 22; RGR I 4+1 62-90% of the length of the anterior plate. Dif-
+ 11 = 16; 3 rows of faint small spots visible on fers from S. brasiliensis by having distinct pterot-
both sides in 1962, size of spots about equal to ic spines. Intercalar spine absent as in the other
half diameter of eye. Paralectotype: BMNH five species of the regalis group and S. niphoni-
1895.5.24.104; Mazatlan, Mexico; D. S. Jordan; tin us.
tag with no. 1720 attached to specimen; 550 mm
FL; D XVII + 17+ IX; A 16+1X; PI 21-22; LGRI Description.-Lateralline gradually descending
3 + 1 + 10 = 14; 3 rows of spots on sides. to midline on caudal peduncle. Intestine with two
folds and three limbs (Fig. 3p). Spines in first
Diagnosis. -This species possesses nasal denti- dorsal fin 15-18, usually 17 or 18 (Table 9); second
cles (Fig. la, b) as do the other five species of the dorsal fin rays 16-19, usually 17 or 18 (Table 10);
regalis group (brasiliensis, concolor, maculatus, dorsal finlets 7-10, usually 8 or 9 (Table 10); anal
regalis, and tritor), has an artery branching off fin rays 16-21, usually 18-20 (Table 11); anal finlets
the fourth left epibranchial artery as do all the 7-10, usually 8 or 9 (Table 11); pectoral fin rays 20-
species in the group except S. tritor, and shares a 24, usually 21 (Table 12). Precaudal vertebrae 19-
specialization of the fourth right epibranchial 21, usually 20 (Table 6); caudal vertebrae 26-29,
artery (Fig. 7e) with S. brasiliensis and S. regalis. usually 28 (Table 7); total vertebrae 46-49, usu-
In these three species, an artery connects the ally 48 (Table 8). Gill rakers on first arch (2-4) +
fourth right epibranchial with a branch of the (9-14)= 12-17, usually 3+ (12-13)= 15-16 (Table 5).
coeliaco-mesenteric artery. Scomberomorus sier- Morphometric characters given in Table 28.

TABLE 28. -Summary of morphometric data of Scomberomorus sierra. FL = fork length, HL = head length.

Mexico Panama Colombia Total

Character N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO N Min. Max. Mean SO
Fork length 36 163 615 344 139 21 98 596 349 154 14 180 266 226 22 97 68 621 340 139
Snout-A %. FL 35 518 565 539 13 21 519 560 536 12 14 510 562 526 13 95 510 616 537 17
Snout-20 %. FL 35 496 534 513 10 21 466 529 507 14 14 489 511 504 6 95 466 586 510 14
Snout-l0 %. FL 36 222 268 245 13 21 221 266 242 12 14 216 251 237 11 97 213 276 241 14
Snout-P, %. FL 36 224 297 255 18 20 238 286 257 15 14 250 264 257 5 96 198 330 252 22
Snout-P, %. FL 36 195 249 222 14 21 205 245 220 11 14 211 227 219 5 97 181 260 220 15
P"P, %. FL 31 93 118 106 8 20 93 123 104 8 14 91 111 105 5 91 89 123 104 8
Head length %. FL 36 190 246 216 14 21 197 237 212 9 14 206 219 213 4 97 183 260 212 14
Max. body depth %. FL 34 157 205 191 12 19 164 210 184 14 14 190 221 205 9 93 157 221 190 15
Max. body width %. FL 32 53 111 84 13 19 63 104 86 11 14 64 86 74 8 90 53 115 84 12
P, length %. FL 36 110 141 123 7 21 102 140 123 11 14 117 135 127 6 97 78 145 123 9
P21ength %. FL 36 40 64 53 4 19 36 64 53 9 14 50 61 55 3 94 32 64 53 6
P2 insertion-vent %. FL 35 215 318 268 23 20 239 294 271 16 14 235 277 255 11 95 193 339 267 25
P2 tip-vent %. FL 35 162 264 216 20 19 194 262 221 18 14 183 216 200 10 93 162 284 221 22
Base 10 %. FL 35 232 286 261 11 20 206 280 253 18 14 238 288 260 12 95 206 306 259 16
Height 20 %. FL 35 103 139 120 8 18 109 139 124 8 13 127 141 135 5 88 103 146 123 10
Base 20 %. FL 36 69 142 118 13 21 105 140 120 10 14 109 144 125 8 97 69 184 120 14
Height anal %. FL 35 103 130 115 7 20 106 135 123 8 14 121 136 128 5 93 91 136 118 9
Base anal %. FL 36 97 163 121 12 21 103 157 121 12 14 115 131 122 5 97 88 163 119 12
Snout (fleshy) %. FL 36 69 91 81 6 21 73 89 80 4 14 73 78 76 2 97 62 92 79 6
Snout (bony) %. FL 33 43 82 72 7 19 65 78 71 4 9 62 69 66 2 85 43 150 70 11
Maxilla length %. FL 36 103 146 124 11 21 115 139 122 6 14 114 129 120 4 97 99 150 121 10
Postorbital %. FL 31 91 107 100 4 21 90 102 97 3 14 94 101 98 2 92 85 116 98 5
Orbital (fleshy) %. FL 36 25 49 34 7 21 23 48 34 7 14 34 43 39 2 97 2 53 34 7
Orbital (:>ony) %. FL 31 39 64 50 8 21 37 64 50 8 14 50 61 54 3 92 16 77 50 9
Interorbital %. FL 36 29 62 56 6 21 52 59 54 2 14 51 58 54 2 97 2 65 54 7
2D-caudal %. FL 30 430 520 476 25 20 436 509 471 18 14 443 511 467 18 89 234 580 475 38
Head length 36 40 135 73 26 21 23 122 73 31 14 39 57 48 5 97 18 135 71 26
Snout (fleshy) %. HL 36 350 405 375 13 21 345 434 379 23 14 344 370 358 7 97 338 434 371 16
Snout (bony) %. HL 33 182 375 332 30 19 310 373 336 21 9 299 320 308 8 85 182 576 332 37
Maxilla length %. HL 36 540 610 573 18 21 561 610 578 13 14 542 592 566 12 97 531 610 570 17
Postorbital %. HL 31 403 502 458 26 21 415 497 460 22 14 446 4n 460 8 92 388 509 461 26
Orbit (fleshy) %. HL 36 124 200 156 22 21 114 203 158 27 14 160 196 182 11 97 9 232 158 27
Orbit (bony) %. HL 31 184 278 226 25 21 173 292 234 33 14 228 280 256 15 92 81 337 235 35
Interorbital %. HL 36 143 284 257 22 21 220 288 257 14 14 241 270 254 8 97 11 288 253 33

664
COLLE'ITE and RUSSO: SPANISH MACKERELS

Size. -Maximum size 96.5 cm FL, 5.44 kg; of271 by artisanal fishermen for a total catch in 1971 of
sierra caught in the Gulf of Nicoya, Costa Rica, 50 127 tons (Artunduaga Pastrana 1976).
measured more than 63.5 cm and weighed more
than 5.4 kg (Erdman 1971). Size at first maturity Distribution.-Eastern Pacific (Fig. 49) from La
26-32 cm FL in Colombia (Artunduaga Pastrana Jolla, southern California (Collette et al. 1963)
1976). A length-}Veight regression curve has been south past Payta, Peru (Collette and Russo 1979:
published for 310 Colombian specimens 15-63 em, fig. 8) to Antofagasta, Chile (lat. 23°24'8, long.
0.03-2.4 kg (Artunduaga Pastrana 1976:fig. 6). 0
70 26'W, Kong 1978). Also found around the
Galapagos Islands.
Color pattern. - Bluish above, silvery white ven-
trally, sides with numerous round brownish Geographic variation. -Comparisons were made
(orange in life) spots, three rows below lateral line, of morphometric data for three populations of S.
one above (Fig. 67). First dorsal fin black distally, sierra by ANCOVA (Table 28): Mexico (n =
white at base. Second dorsal tinged with yellow, 31-36), Panama (n = 18-21), Colombia (n = 9-14).
margins black. Anal white. Null hypotheses that the 3 sets of regression lines
There is a color painting of S. sierra by Malm- are coincident were accepted for 18 sets, rejected
quist in Walford (l937:pl. 39), and there is a good for 8 sets: Sn-lD, Sn-P2 , Head L, maximum body
black and white underwater photograph ofseveral depth, Ht 2D, Ht A, Snout (fleshy), and maxilla L.
specimens in the Gulf of California in Thomson et The Newman-Keuls Multiple Range Test identi-
al. (1979:fig. 115). fied populations that were significantly different
for 6 sets of regressions. Five of these (Sn-lD,
Biology.-Spawning probably takes place near Sn-P2 , Head L, Ht A, and maxilla L) indicated that
the coast over most of its range (Klawe 1966). the populations from Mexico and Panama were
Spawning occurs off Mexico in July-September significantly different, one (maximum depth) that
(Klawe 1966). Ripe males and females were found the Panama and Colombia populations were sig-
from late August to the end of November in the nificantly different. The samples from Panama
Gulf of Nicoya, Costa Rica (Erdman 1971). The and Colombia were then combined and the com-
maximum incidence of ripe females extends from bined regressions were compared with those for
November to April in Colombia with a peak in Mexico. Null hypotheses were rejected for 7 ofthe
February-April (Artunduaga Pastrana 1976). 26 sets of regression lines; the same ones as were
Larvae and juveniles, 4.5-139 mm FL, of S. sierra rejected in the first test except for maximum body
have been taken from Baja California to Peru depth. No meristic differences were found between
duringJanuary-April and July-September (Klawe populations.
1966:fig. 1, table 1). The smallest larvae were
taken off Baja California July-September: 4.5-9.5 Material examined.-Total123 (68-621 mm FL).
mm FL, 13 September; 4.8 mm, 12 August; 8.4 mm,
9 July (Klawe 1966:table 1). Food of adults con- meas.: 97 (68-621): California (1); Mexico (39,
sists of small fishes (Walford 1937). In Colombia *S. sierra); EI Salvador (1); Costa Rica
(Artunduaga Pastrana 1976), the commonest (5); Panama (21); Colombia (14); Ecuador
fishes in stomach contents were anchovies (En- (7); Peru (6); Galapagos Is. (3).
graulidae, Anchoa and Cetengraulis) and clu- counts: 123.
peids (Odontognatus and Opisthonema). diss.: 13 (368-590): Mexico (2); Panama (5);
Ecuador (5).
Interest to fisheries. - According to Walford (1937),
S. sierra seems to be the most abundant game fish Scomberomorus sinensis (Lacepede)
along the Pacific coasts of Mexico and Central Chinese Seerfish
America. It is an excellent food fish frequently Figure 68
taken by anglers and abundant enough to support
a commercial fishery (Eckles 1949). Statistics are Scomber sinensis Lacepede 1800:599 (original
reported from Fishing Areas 77 and 87; the bulk of description). Lacepede 1802:23 (description
the catch is reported for Mexico 4,028-11,999 t/yr based on a Chinese drawing). GUnther 1860:
and for Peru 320-579 t/yr in 1979-82 (FAO 1984). 369 (footnoted as dubious species).
No specific commercial fishery exists for S. sierra Cybium chinense Cuvier in Cuvier and Valen-
in Colombia, but it is taken by the shrimp fleet and ciennes 1831:180 (original description based on
665
 FISHERY BULLETIN: VOL. 82, NO.4

FIGURE 68.-Scomberomorus sinensis. Shanghai, China, 714 mm FL, USNM 220856.

same figure used by Lacepede). Temminck range). Zama and Fujita 1977:118 (Ogasawara
and Schlegel 1844:100-101 (description), pI. 53, Is., listed after Sugiura 1970). Collette 1979:
fig. 1 (color painting of adult). Richardson 29 (characters, range). Collette and Russo
1846:268 (synonymy; seas of China and Japan). 1979:13 (diagnostic characters, range). Ohe et .
GUnther 1860:369 (footnoted as dubious spe- al. 1981:42-43 (comparison with Miocene Acan-
cies). Bleeker 1873:131 (China; listed). Kish- thocybium from Japan). Zhang and Zhang
inouye 1915:11 (description), pI. 1, fig. 5. * Kish- 1981:104 (range). Lee and Yang 1983:229 (Tai-
inouye 1923:418-419 (description, range), pI. 21, wan), fig. 18 (1,056 mm). Cressey et al. 1983:
fig. 34 (adult), pI. 23, fig. 40 (skull, vertebral 264 (host-parasite list, 3 copepod species). Col-
column). Kishinouye 1924:92 (26 mmjuvenile lette and Nauen 1983:78 (description, range),
from skipjick stomach). Boeseman 1947:95 fig.
(identification of Burger's plate). Morice 1953: Scomberomorus chinensis. Jordan et al. 1913:
37 (villiform tongue teeth present). 122 (Japanese common names). Norman and
Scomberomorus sinensis. Jordan and Snyder Fraser 1949:153 (China and Japan). Devaraj
1900:352 (Tokyo; listed). Jordan and Snyder 1977:56-57 (S. chinensis intermediate between
1901:64 (Japanese localities). Reeves 1927:8 Acanthocybium and other species of Scomber-
(Chefoo, China; listed). Mori 1928:5 (Fusan, omorus).
Korea; listed). Soldatov and Lindberg 1930:111 Cybium cambodgiense Durand 1940:37-38 (orig-
(synonymy, description, range). Munro 1943: inal description; Phnom Penh, Cambodia), pI. 6.
69, 71 (placed in subgenus Sierra; C. cambod- Scomberomorus caualla (not of Cuvier 1829).
giense Durand a junior synonym of S. sinensis). Fraser-Brunner 1950:160-161 (Cybium sinensis
Mori 1952:137 (Fusan, Korea; listed). Mori placed in synonymy ofS. caualla and considered
1956:23 (Kasumi and Hamada, S Japan Sea; a subspecies of it).
listed). Blanc et al. 1965:121-123 (C. cambod- Scomberomorus chinense. Richards and Klawe
giense Durand a junior synonym of S. sinensis). 1972:13 (range), 90 (reference to Kishinouye
*D'Aubenton and Blanc 1965:233-243 (descrip- 1924).
tion, range, biology), fig. 1 (191 mmjuvenile), fig. Scomberomorus sp. Kawamoto et al. 1972:49 (de-
2 (1,170 mm adult). Kamohara 1967:43 (com- scription; Mekong Delta, Vietnam), fig. 96.
parison with S. niphonius). Tokida and Koba- Scomberomorus cambodgiense. Orsi 1974:174
yashi 1967:158 (identification of C. chinense (Vietnam; listed).
of Uchimura's unpublished 1884 manuscript).
Sugiura 1970:205 (Bonin Is., listed). Richards Types of nominal species. -Both Scomber sinen-
and Klawe 1972:15 (range), 95-96 (reference to sis Lacepede 1800 and Cybium chinense Cuvier in
Blanc et al. 1965 and D'Aubenton and Blanc Cuvier and Valenciennes 1831 are based on a
1965). Shiino 1972:71 (common name). Mag- Chinese drawing; no types are extant (Blanc and
nuson 1973:350 (short pectoral fin). Orsi 1974: Bauchot 1964:449).
175 (Vietnam; listed). Shiino 1976:231 (com- Cybium cambodgiense Durand 1940. The orig-
mon name). Klawe 1977:2 (common name, inal description was based on a 215 mm specimen
666
COLLE'ITE and RUSSO: SPANISH MACKERELS

taken in Phnom Penh, Cambodia, 28 January middle of body (D'Aubenton and Blanc 1965:fig.
1939. Because this specimen is not known to still 2).
exist, data are presented from the original descrip- There is an excellent figure of S. sinensis in
tion: D XVI + 16 + VII; A 19 + VI; P l 22; GR 3 + 9 = Kishinouye (1923:fig. 34) and there are drawings
12. The figure (pI. 6) accompanying the original of ajuvenile (191 mm FL, fig. 1) and an adult (1,017
description clearly shows the deep dip in the mm FL, fig. 2) of S. sinensis from the Mekong
lateral line under the posterior part of the first River in Cambodia in D'Aubenton and Blanc
dorsal fin. (1965).

Diagnosis. -The only species of Scomberomorus Biology. -No information is available on the
that has a swim bladder and the only species with movements of S. sinensis. Although it penetrates
an abrupt downward curve in the lateral line great distances up the Mekong River, D'Aubenton
beneath the first dorsal fin (Fig. 68). Two spe- and Blanc (1965) reported that they failed to find
cies, S. cavalla and S. commerson, also have the slightest trace ofsexual activity in Cambodian
abrupt downward curves in the lateral line but freshwater specimens and so they concluded that
they are under the second dorsal fin. The lateral S. sinensis must reproduce exclusively in the sea.
line in the other 15 species descends gradually No information is available on eggs or larvae
without any prominent dips. The pectoral fins are (Richards and Klawe 1972), but D'Aubenton and
large and rounded rather than pointed as in the Blanc (1965) did report on juveniles as small as 166
other 17 species. Palatine tooth patch very narrow mm FL from TonIe Sap, Cambodia.
as in Scomberomorus commerson and Acantho-
cybium. Ventral process of angular moderate, 87- Interest to fisheries. - No catches were reported as
93% as long as the dorsal process, as in S. cavalla. S. sinensis by FAO for the period 1979-82 (FAO
Posterior end of maxilla only slightly expanded 1984). However, it is a prized food fish in Japan
as in S. multiradiatus. Ascending process of and probably in China as well. Kishinouye(1923)
premaxilla very long as in S. lineolatus and
Acanthocybium.
TABLE 29.-Summary of morphometric data of Scom-
Description. -Intestine with two folds and three beromorus sinensis. FL = fork length, HL = head
limbs (Fig. 3q). Spines in first dorsal fin 15-17, length.

usually 16 or 17 (Table 9); second dorsal fin rays Character N Min. Max. Mean SO
15-17, usually 15 or 16 (Table 10); dorsal finlets 6-7 Fork length 18 157 714 330 176
Snout-A %. FL 14 573 605 584 11
(Table 10); anal fin rays 16-19, usually 17 or 18 Snout-20 %. FL 14 526 571 558 13
(Table 11); anal finlets 5-7, usually 6 (Table 11); Snout-lO %. FL 16 268 309 292 9
Snout-P, %. FL 16 270 318 290 12
pectoral fin rays 21-23, usually 22 (Table 12). Snout-P, %. FL 16 234 279 259 11
Precaudal vertebrae 19 or 20 (Table 6); caudal P,-P, %. FL 16 101 128 113 7
Head length %. FL 18 230 264 255 8
vertebrae 21 or 22, usually 22 (Table 7); total Max. body depth %. FL 16 201 231 218 10
Max. body width %. FL 16 77 127 102 14
vertebrae 41 or 42, usually 41 (Table 8). Gill rakers P, length %. FL 18 133 186 157 14
on first arch (1-3)+ (10-12)= 11-15, usually 2+ p,length %. FL 16 75 89 83 5
P2 insertion-vent %. FL 15 254 295 273 13
(10-11)= 12-13 (Table 5). Morphometric characters P, tip-vent %. FL 15 161 216 189 17
given in Table 29. Base 10 %. FL 16 230 282 260 12
Height 20 %. FL 15 130 164 145 10
Base 20 %. FL 15 99 137 121 11
Size. - Maximum size 200 cm FL, 80 kg in weight Height anal %. FL 16 129 164 145 10
Base anal %. FL 16 99 149 122 12
(Kishinouye 1923). A length-weight graph for fish Snout (fleshy) . %. FL 16 86 106 97 5
Snout (bony) %. FL 16 77 99 90 5
up to 120 cm FL and 18 kg was published by Maxilla length %. FL 16 131 159 147 7
D'Aubenton and Blanc (1965:fig. 4). Postorbital %. FL 16 104 126 117 7
Orbital (fleshy) %. FL 16 25 40 35 5
Orbital (bony) %. FL 16 40 61 52 7
Color pattern_ -Back greenish blue, belly silvery, Interorbital %. FL 16 54 72 63 5
20-caudal %. FL 13 394 469 445 27
fins mostly blackish. Pelvic and anal fins with Head leng1h 18 41 173 83 41
blackish margins, anal finlets colorless (Kishi- Snout (fleshy) %. HL 16 360 414 382 14
Snout (bony) %. HL 16 335 387 355 14
nouye 1923). Large (larger than the diameter of Maxilla length %. HL 16 561 603 578 12
the eye), round, indistinct spots on sides in two Postorbital %. HL 16 404 482 460 24
Orbit (fleshy) %. HL 16 104 155 138 17
poorly defined rows in adults (Fig. 68). Juveniles Orbit (bony) %. HL 16 157 236 202 23
Interorbital %. HL 16 208 289 249 23
with saddlelike blotches extending down to about
667
 FISHERY BULLETIN: VOL. 82, NO.4

reported that 2 or 3 dozens of this species were 2 sets of regression lines are coincident were
often caught on an autumn day in pound nets on accepted for all but 1 set of the 26 sets, Sn-P2
the southern coast of Korea. It is caught in the (intercepts 7.552 and 8.385 respectively). The
Mekong River of Cambodia and commanded a Chinese sample had slightly fewer gill rakers on
high price in the Phnom Penh market in 1964 the first gill arch (11-14, mode 12, i 12.30) than the
(D'Aubenton and Blanc 1965:242). Mekong sample (12-15, mode 13, i 13.17). No other
meristic differences were found.
Distribution. -Western Pacific from Japan and
China south to Cambodia and Vietnam where it Material examined.-TotaI19 (157-714 mm FL).
enters the Mekong River (Fig. 51). The northern
limit is Akita, Honshu, in the Sea ofJapan and the meas.: 18 (157-714): China (10); Mekong R.,
Chiba-Tokyo area on the Pacific coast (Kishinouye Cambodia (6); Cochinchine (2).
1923:418). There are records or specimens from counts: 18.
Pusan, Korea (Mori 1928, 1952), Cheefo (= Yentai) diss.: 1 (plus 1 head, 431 mm).
on the Shantung Peninsula (Reeves 1927), the
Zhoushan Islands (lat. 30o N, long. 122°E, USNM
220856), Foochow (ZMH 11384), Amoy (USNM Scomberomorus tritor (Cuvier)
221277), and Hong Kong (CAS GVF HK 127). A West African Spanish Mackerel
record from the Bonin (or Ogasawara) Islands Figure 69
(Sugiura 1970; repeated by Zama and Fujita 1977)
has not been verified and seems very far offshore Cybium tritor Cuvier in Cuvier and Valenciennes
for this species. It has been taken on the coast of 1831:176-177 (original description; Goree, Sen-
Vietnam at Nha Trang (Blanc et aI. 1965). Scom- egal), pI. 218. GUnther 1860:372 (description
beromorus sinensis is the only species ofthe genus after Cuvier). Rochebrune 1882:96 (very com-
and of the family to move any significant distance mon; Goree, Dakar). Osorio 1898:197 (Prin-
into freshwater. It was described as a distinct cipe). Pellegrin 1908:75 (Dakar). Chabanaud
species, Cybium cambodgiense, by Durand (1940) and Monod 1927:278 (Port-Etienne), fig. 30 A.
from material from Phnom Penh, Cambodia, about Cadenat 1947:15 (common names; W Africa).
300 km up the Mekong River. Specimens (MNHN Postel 1954:357-358 (stomach contents of 286
1965-286-9) have come from TonIe Sap (or Grand specimens), 362 (gonosomatic index). A. Postel
Lac) which is even further lip the Mekong River 1955:60-61, fig. 3 (lower jaw teeth 10-21, upper
(Blanc et aI. 1965). jaw teeth 13-27; 190 specimens), 63, fig. 5
(number of jaw teeth in males and females).
Geographic variation. -Morphometric data for *Postel 1955a:4-158 (distribution, size, repro-
two small samples of S. sinensis were compared duction), pI. II, bottom figure. Postel 1955b:
with ANCOVA: China (n = 7-10) and the Mekong 31-32 (sex ratio, maximum size, number of
River, Cambodia (n = 6). Null hypotheses that the eggs). Frade and Postel 1955:35 (histology of

FIGURE 69.-Scomberomorus tritor. Liberia, 430 mm FL, USNM 193178.

668
COLLE'ITE and RUSSO: SPANISH MACKERELS

gonads; mature in June), fig. 7 (oocytes). Pos- Miyake and Hayasi 1972:I1I-3 (in key), IV-lO
tel 1958:107-111 (summary of Postel 1955a). (common names). Magnuson 1973:350 (short
Postel 1959:163 (listed). Postel 1960:257 (Cap pectoral fin). Klawe 1977:2 (common name;
Blanc to Senegambie). Marchal 1961:106 (lat. range). Penrith 1978:187 (Baia dos Tigres,
9°12'N, long. 13°42'W). Franca 1964:table 3 Angola). Collette et aI.I978:274-275 (compar-
(Angola). Daget and Iltis 1965:280-281 (Ebrie ison with W Atlantic species). Collette 1979:
and Abi lagoons, Ivory Coast), fig. 178. San- 29 (characters, range). Collette and Russo
ches 1966:146 (Angola). Blache et aI. 1970:375 1979:13 (diagnostic characters, range). Col-
(in key), fig. 960 (not fig. 961). Conand 1970:40 lette 1981:Scombm 7 (description, range), fig.
(distribution of larvae). Seret and Opic 1981:332-333 (description), fig.
Apolectus immunis Bennett 1831:146 (original Cressey et aI. 1983:264 (host-parasite list, 4
description; "Atlantic Coast of North Africa"). copepod species). Collette and Nauen 1983:79
Cybium maculatum. Not ofMitchill 1815. Stas- (description, range), fig.
sano 1890:44 (Spanish Sahara). Vinciguerra
1890:100-103 (synonymy). Osorio 1898:197 Types of nominal species. -Holotype: MNHN
(Sao Thome). A.6871; Goree, Senegal; Rang; 658 mm FL; D
Scomberomorus argyreus Fowler 1905:764-765 XV + 17 + VIII; A 17 + VIII; PI 21; RGR I 2+ 1+ 10
(original description, "West Africa"), pI. 51. = 13. Paratype: MNHN A.6868; Goree; Senegal;
Scomberomorus maculatus. Not of Mitchilll815. Rang; 505 mm FL. Photographs of the holotype
Fowler 1936:628-629. Cadenat 1937:482 (Da- and paratype were published by Blanc and Bau-
kar). Scaccini 1941:19 (synonymy in part; Mau- chot (1964:pI. 4, figs. 19, 20).
ritania). Navarro 1943:131 (Cabo Barbas and Scomberomorus argyreus Fowler 1905. Holo-
Blanco, Banco Arguin, Mauritania). Tortonese type: ANSP 11400; west coast of Africa; Dr. Sav-
1949:65 (accidental in Mediterranean Sea). age; 148mmFL; DXVII+ 17+ VIII;AI9+ VIII; PI
Sanz Echeverria 1950:1-2 (sagitta compared 22-22; RGRI 2+ 1 + 11= 14; vertebrae 18+28= 46.
with other scombrids), pI. 1, figs. 1-4 (photo-
graphs of sagittae). Mather and Day 1954:182, Diagnosis. -This species possesses nasal denti-
185 (Sierra Leone, Dakar, Canary Is.; S. tritor cles as do the other five species ofthe regalis group
not specifically distinct from S. maculatus). (brasiliensis, concolor, maculatus, regalis , and
Tortonese 1956:7 (accidental in Mediterranean sierra) but lacks the artery branching from the
Sea). Poll 1959:104-106 (description; S to Baie fourth left epibranchial artery that is present in
des Tigres, Angola), fig. 34. Maurin et aI. the other species (Fig. 7b). Intercalar spine absent
1970:19 (Nouakchott, NW Africa). Lozano as in the other five species ofthe regalis group and
Cabo 1970:158 (Sahara coast). Fagade and S. niphonius.
Olaniyan 1973:212, 220, 224 (piscivorous, feed-
ing largely on Ethmalosa fimbriata in Lagos). Description. -Lateral line gradually descending
Fagade and Olaniyan 1974:249 (caught in Lagos to midline on caudal peduncle. Intestine with two
Lagoon when water was brackish). Tortonese folds and three limbs (Fig. 3r). Spines in first
1975:354-355 (description, Italy), fig. 155. dorsal fin 15-18, usually 17 or 18 (Table 9); second
Scomberomorus tritor: Munro 1943:67-71 (placed dorsal fin rays 16-19, usually 17 (Table 10); dorsal
in subgenus Scomberomorus). Irvine 1947: finlets 7-9, usually 8 (Table 10); anal fin rays
186-187 (Accra, Ghana), fig. 108. Fraser- 17-20, usually 18 or 19 (Table 11); anal finlets
Brunner 1950:158 (synonymy in part), fig. 27. 7-9, usually 8 (Table 11); pectoral fin rays 20-22,
Chaine 1957:504-509 (otoliths), pI. IV (otoliths). usually 21 (Table 12). Precaudal vertebrae 18 or 19,
Gras 1961:583 (Lagune de Cotonou and Lac usually 19 (Table 6); caudal vertebrae 27 or 28,
Nokoue, Dahomey). Bauchot and Blanc 1961: usually 27 (Table 7); total vertebrae 46 or 47, •
372-373 (types). Blanc and Bauchot 1964:448 usually 46 (Table 8). Gill rakers on first arch
(types), pI. IV, figs. 19-20 (photographs oftypes). (1-3) + (10-13) = 12-15, usually 2 + (11-12) = 13-14
Gorbunova 1965a:54 (spawning season). Col- (Table 5). Postel (1955a) reported a range of 10-15
lette 1966:367 (types). Williams 1968:436, ta- gill rakers for 240 males and 520 females, 94% of
ble 593 (taken from Gambia to the Congo during both sexes 12-14. Morphometric characters given
the Guinean Trawling Survey). Collette 1970: in Table 30.
4-5 (in key; Mediterranean Sea). Richards and
Klawe 1972:15 (range), 96 (references to larvae). Size. -Maximum size of males 83.9 em FL, fe-

669
 FISHERY BULLETIN: VOL. 82, NO.4

males 97.5 cm FL; commonly 50-70 cm; age at first 8. tritor with food contained the clupeid Ethma-
maturity of both sexes 45 cm (Postel 1955a). losa fimbriata (Fagade and Olaniyan 1973). No
sexually mature stages of 8. tritor were found in
Color pattern. - Upper parts of body bluish, belly the lagoon so Fagade and Olaniyan concluded that
silvery, sides marked with several poorly defined the lagoon served as a feeding ground for this and
rows of elongate spots (Fig. 69). First dorsal fin other piscivorous fishes that could tolerate the
black anteriorly and along distal margin posteri- reduced salinity of the lagoon.
orly, white at base.
There are drawings of 8. tritor in Postel (1955a: Interest to fisheries. - Taken throughout the Gulf
pI. 2) and Poll (1959:fig. 34), Collette (1981: ofGuinea, but catches are reported only for Ghana
Scombm 7), and Seret and Opic (1981:333). and Angola in the period 1979-82. Most of the
catch is reported for GhaBa, 1,569 to 4,412 t/yr
Biology.-Sexual maturity in both sexes of 8. (FAO 1984).
tritor occurs from April to October in Senegal
(Postel 1955a). Postel (1955a) reported 1 million Distribution. -Eastern Atlantic, concentrated in
eggs in a 95 cm FL female. Juveniles have been the Gulf of Guinea from the Canary Islands
seined along the shore near Dakar in July (Postel (Mather and Day 1954; MCZ 26416), West Sahara
1955a). Seven larvae 3.5-8.1 mm were caught in (Stassano 1890), and Dakar, Senegal (original
September, December, February, and March, description), south to Baia dos Tigres, southern
south of the Ivory Coast at water temperatures of Angola (Fig. 49). Accidental and rare in the
23.2°-26°C and salinities of 34.38-35.45%0 (Zhu- Mediterranean Sea (Tortonese 1949, 1956), with
dova 1969). Three larvae identified as 8. tritor several extant specimens from Nice (NHMV
were reported from a station track from Dakar to 14599, MSUF M.1665), Villefranche, and Palermo
Recife (Zharov and Zhudova 1967), but this distri- (Tortonese 1975).
bution seems highly unlikely. In Lagos Lagoon,
Nigeria, stomach contents of 24 of 26 specimens of Material examined.-TotaI49 (69-658 mm FL).

meas.: 40 (102-658): N. Mediterranean (2); Ca-

TABLE 30.-Summary of morphometric data of Scom- nary Is. (1); Spanish Sahara (1); Senegal
beromorus tritor. FL = fork length, HL = head length. (4, *C. tritor), Guinea (1); Sierra Leone
Character N Min. Max. Mean SO (5); Liberia (5); Ivory Coast (10); Ghana
Fork length 40 102 658 347 151 (5); Nigeria (4); Angola (2); "West Afri-
Snout-A %. FL 40 506 567 533 15
Snout-20 roo FL 39 482 551 513 14
ca" (1, *8. argyreus Fowler).
Snout-1O %. FL 39 190 276 246 13 counts: 49.
Snout-P, %. FL 40 237 301 266 17
Snout-P, %. FL 40 197 247 222 13 diss.: 10 (394-600): Ivory Coast.
P,-P, %. FL 40 95 138 111 11
Head length roo FL 40 199 242 217 10
Max. body depth %. FL 38 166 240 206 21 RELATIONSHIPS
Max. body width %. FL 36 67 110 90 11
P, length %. FL 39 110 152 134 9
P,length %. FL 38 48 71 60 6 After comparing the species of 8comberomorus
P2 insertion-vent %. FL 39 213 304 250 18
P, tip-vent %. FL 38 146 243 190 17
with each other and with Grammatorcynus and
Base 10 roo FL 38 238 304 262 14 Acanthocybium (Comparative Morphology), all
Height 20 %. FL 31 97 161 126 14
Base 20 roo FL 40 101 144 122 10 the characters that differentiated species or
Height anal %. FL 36 92 159 125 13 genera were listed. Character polarities were
Base anal %. FL 40 93 139 119 10
Snout (fleshy) %. FL 40 73 93 81 5 determined by considering the character state
Snout (bony) %. FL 40 64 86 72 5 present in Grammatorcynus to represent the ple-
Maxilla length %. FL 40 110 143 123 8
Postorbital roo FL 39 89 107 96 4 siomorphous condition. 8comberomorus, Acan-
Orbital (fleshy) %. FL 40 26 52 38 6
Orbital (bony) roo FL 40 28 68 53 8
thocybium, the Sardini, and the Thunnini have 4
Interorbital %. FL 40 51 68 59 4 or 5 caudal vertebrae supporting the caudal fin
20-caudal roo FL 38 432 534 476 29
Head length 40 23 145 74 31 and 37 ·or more total vertebrae. These derived
Snout (fleshy) %. HL 40 332 409 376 18 characters indicate that these taxa form a mono-
Snout (bony) %. HL 40 290 388 333 21
Maxilla length' %. HL 40 541 608 568 14 phyletic group within the Scombridae. Gramma-
Postorbital %. HL 39 399 474 443 20 torcynus lies between the Scombrini and the
Orbit (fleshy) %. HL 40 124 233 173 23
Orbit (bony) %. HL 40 133 307 245 32 higher scombrids and is clearly more primitive
Interorbital %. HL 40 230 301 272 13
than 8comberomorus because it has, as in the
670
COLLETTE and RUSSO: SPANISH MACKERELS

Scombrini, only 3 vertebrae supporting the caudal semifasciatus from node 9; guttatus from node 8;
fin and only 31 total vertebrae. Therefore, we have and regalis from node 5.
used it as the outgroup for comparison with Scom- The sinensis group is monotypic. It is defined by
beromorus. Of the 72 characters that differenti- the presence of an abrupt downward curve in the
ated at least 1 taxon from the others, 14 were lateral line under the first dorsal fin (character 19,
autapomorphies of Acanthocybium. These cannot state 1). A similar abrupt downward curve is
contribute to an understanding of relationships present in two species ofthe commerson group but
within Scomberomorus and were omitted from the the curve is under the second dorsal in those two
analysis. The remaining 58 characters were em- species. Scomberomorus sinensis is the only spe-
ployed to generate a cladogram using a computer cies in the genus with a well-developed swim
program (WAGNER 78) written by J. S. Farris bladder (character 18, state 0), but this is a plesio-
(following Farris 1970 and Farris et al. 1970). The morphous character. This species is restricted to
order of the taxa was "shuffled" to determine if the northwestern Pacific from Cambodia to Ja-
another equally parsimonious tree would be gen- pan. There is no genus-group name available for
erated. Another cladogram was produced with the this group.
same total length, 112 steps. The first cladogram The commerson group contains four species:
has a deviation ratio (sum of the homoplasies niphonius, queenslandicus, cavalla, and commer-
between all pairwise combinations of terminal son. This group is defined by the presence of an
taxa divided by the sum of the character changes intercalar spine of at least moderate length (char-
between all pairwise combinations of terminal acter 17, state 1). Three species (queenslandicus,
taxa) of 0.24, the second 0.21. The difference is due cavalla, and commerson) have a long (state 2)
to characters 3 and 17. We feel that the first intercalar spine. Scomberomorus cavalla and S.
cladogram (Fig. 70) more reasonably reflects our commerson share two additional specializations:
concepts of evolution within the genus. A sum- the pterosphenoid bones are close together (char-
mary ofthe character states with references to the acter 13, state 1) and there is an abrupt downward
relevant figures is presented as Appendix 1. curve in the lateral line under the second dorsal
We recognize six species groups within Scom- fin (character 19). Three ofthese species are Indo-
beromorus (Fig. 70, Table 31):' sinensis from node West Pacific: niphonius from China, Korea, and
17; commerson from node 15; munroi from node 11; Japan; queenslandicus from off northern Austra-

,".'"
'> ~
~ ,~
-;;
~
,:: ,:: ,::" ,::;
;; ~
.~
,<:I
,'>
" ::" ,::" -;; ~
,
" ""
"V
'" . '" •."'" "'"
.;:: ,::
,'> 0 ~

..;: "- ~
• /:; ,~" :::'" ,::" '>
,
! '" '"
..:::
,, ,.-S" ~
co ~
'"
~ "

:J a"'"'"
c a a <:I
'" ~ i:: c -::: if
,"• •
<:I
~ ~
~ <:I
,~ a ~ '>" ~
~
i.:"
$
'"
'> (] ~
.0;
..::: ~
'>
.;; c a'" ,~
,~
<:I

G
<:I
'\' "i
'" " vi "i
Co
"0'
~

"0' "0'
"-
"0' "0'
" "0' "0' "0' '" "0 "0'~ "0'c "0'" "0''"
"0' "0'

FIGURE 70.-Cladogram of the Scomber-

omorini, node numbers refer to numbers
used in Table 31.

671
 FISHERY BULLETIN: VOL. 82, NO.4

TABLE 31.-Changes in character states based on the most lia and southern Papua New Guinea; and com-
parsimonious cladogram in Figure 70. Numbers under acquisi. merson widespread throughout the Indo-West Pa-
tion and reversal columns refer to nodes, three-letter mnemonics cific. The fourth, cavalla, is restricted to the
refer to species of Scomberomorus, and four-letter mnemonics
refer to genera, i.e., ACAN = Acanthocybium. Two or more
western Atlantic. The genus-group name Cybium
components per cell indicate independent acquisition or loss of a Cuvier (type-species S. commerson) is available
character state. Two or more states of the same character ac- for this group.
quired at a single node assume that the more primitive state was Scomberomorus niphonius is the only species in
transitional during the acquisition of the more advanced state.
the genus with a straight gut. This species has
Char- Acqui- Char- Acqui- very small scapular foramina (character 11, state
acter State sitian Reversal acter State sition
0 19 26
2), a character state also found, evidently homo-
0 19
1 18 1 18 plasiously, in S. guttatus and S. koreanus. It is re-
2 ACAN,13 CAY 2 17
2 0 19 27 0 19
stricted to the northwestern Pacific from China,
1 18 SEM 1 18 Korea, and Japan. The genus-group name Sawara
2 COM, 18 15 2 ACAN
3 0 8,17 11 28 0 ACAN Jordan and Hubbs is available for S. niphonius.
1 19 1 19 The munroi group is monotypic. It is defined by
2 ACAN 2 17
4 0 19 29 0 19 the loss ofthe anterior process on the outer surface
1 13, 18 15 1 17 of the head of the maxilla. It is restricted to
2 13, ACAN CAY 30 0 19
5 0 19 1 17 northern Australia and southern Papua New
1 17 31 0 19
2 15 MUL 1 17
Guinea. There is no genus-group name available
3 10 32 0 19 for this group.
6 0 8 GUT 1 17
1 17 NIp, 16 33 0 19 The semifasciatus group contains three species:
2 19 1 17 plurilineatus, lineolatus, and semifasciatus. This
3 ACAN 34 0 19
7 0 19 1 17 group is defined by the presence of a greatly
1 16,18 CAV,9 35 0 19 expanded posterior end of the maxilla (character
8 0 19 1 17
1 18 GUT 36 0 19 5, state 3). Two species, S. lineolatus and S.
2 ACAN, LIN 1 17
9 0 CAV,GUT 37 0 19
semifasciatus, share an additional specialization,
1 19 1 17 a wide parasphenoid (character 7, state 1). This
2 18, LIN 15 38 0 19
10 0 19 1 17 character state appears independently in several
1 5 BRA 39 0 19 other lines. All are Indo-West Pacific species,
11 0 ACAN 1 17
1 19 40 0 19 plurilineatus along the coast of East Africa plus
2 NIp, 7 1 17 Madagascar, lineolatus along the continental
12 Ii 19 41 0 19
1 17 1 18 coast from India to Indonesia, and semifasciatus
2 17 14 42 0 19
13 0 19 1 18
in northern Australia and southern Papua New
1 LIN, 14 43 0 19 Guinea. The genus-group name Indocybium Mun-
14 0 19 1 18
1 6 44 0 19 ro (type-species S. semifasciatus) is available for
15 0 19 1 18 this group.
1 3,9 LIN, 4 45 0 19
16 0 19 1 18 The guttatus group contains three species: mul-
l 1 46 0 19
17 0 19 1 18
tiradiatus, guttatus, and koreanus. This group is
1 5,12 47 0 19 defined by a high supraoccipital crest (character
2 13 1 18
18 0 19 48 0 19 14, state 1). Two species, guttatus and koreanus ,
1~
15 1 18 share the presence of auxiliary branches of the
19 0 19 49 0 19
1 14,18 15 1 18 lateral line (character 20, state 1). They also have
20 0 19 50 0 19 very small scapular foramina (character 11, state
1 7 1 18
21 0 19 51 0 19 2), a character state shared homoplasiously with
1 5 1 18
22 0 19 52 0 19
S. niphonius. All are Indo-West Pacific species,
1 4 1 18 guttatus and koreanus along the coast of Asia and
2 2 53 0 19
23 0 19 1 18 multiradiatus confined to a small section of the
1 17 MUL 54 0 19 Gulf of Papua off the mouth of the Fly River. The
2 LIN 1 18
24 0 19 55 0 19 genus-group name Pseudosawara Munro (type-
1 18 1 18 species S. guttatus) is. available for this group.
2 17 56 0 19
25 0 19 1 18 The regalis group contains six Atlantic and
1 18 57 0 19
2 17 1 18
eastern Pacific species: tritor, maculatus, con-
58 0 17 color, sierra, brasiliensis, and regalis. This group
1 19
is defined by the presence of nasal denticles (char-
672
COLLETTE and RUSSO: SPANISH MACKERELS

acter 21, state 1), a synapomorphy unique to the (formerly at DASF); Pearl Sonoda (CAS); Camm
group. All species also have a moderately long Swift (LACM); Frank Talbot (CAS, formerly at
intercalar spine (character 17), but this is also SAM and AMS); H. Wilkens (ZMH); Richard
present in S. niphonius. All species in the group Winterbottom (ROM, formerly at RUSI); and Luis
have a vomerine ridge (character 10, state 1) Alberto Zavalla-Camin (MPIP and MZUSP).
except for S. brasiliensis in which it has been Frozen material, vital to this project, was ob-
secondarily lost. The five most advanced species tained through the much appreciated efforts of
(all except S. tritor) have an artery arising from Tokiharu Abe (University of Tokyo); Adam Ben-
the fourth left epibranchial artery (character 22, Tuvia (Sea Fisheries Research Station, Haifa);
state 1). The four most advanced species (all except Frederick H. Berry and Mark D. Lange (TABL);
S. tritor and S. maculatus) have developed a long John Carleton (Queensland Fisheries Service); M.
posterior process on the pelvic girdle (character 16, Devaraj (formerly at Central Marine Fisheries
state 1). The three most advanced species (sierra, Research Institute, Mandapam Camp, India);
brasiliensis, and regalis) have a coeliaco-mesen- Jeffrey B. Graham (formerly at Smithsonian
teric shunt connecting the fourth right epibran- Tropical Research Institute, Balboa, Panama);
chial artery with the coeliaco-mesenteric artery Elwood K. Harry (International Game Fish Asso-
(character 22, state 2). The two most advanced ciation, Fort Lauderdale, Fla.); Barry Hutchins
species (brasiliensis and regalis ) have lost the (WAM); S. A. Jaleel (Marine Fisheries Depart-
pterotic spine (character 15, state 1) but this spine ment, Karachi, Pakistan); W. L. Klawe (Inter-
has also been independently lost in other lines. American Tropical Tuna Commission, La Jolla,
The genus-group name Scomberomorus Lacepede Calif.); Leslie W. Knapp (Smithsonian Oceano-
sensu stricto (type-species S. regalis ) applies to graphic Sorting Center); A. D. Lewis (formerly at
this group. DASF); Eugene L. Nakamura (NMFS, Panama
City, Fla.); the late Al Pflueger (Miami); Gary
ACKNOWLEDGMENTS Sharp (FAO, formerly at Inter-American Tropi-
cal Tuna Commission); J. M. Stretta (Centre de
For permission to examine specimens in their Recherches Oceanographiques, Abidjan, Ivory
institutions, or for donating specimens to the Coast); Camm Swift (LACM); Sen Min Tan (Ma-
USNM collections, we thank the following: Kunio rine Fisheries Research Department, SEAFDEC,
Amaoka (HUMZ); Maria Luisa Azzaroli (MSUF); Singapore); Rudy van der Elst (Oceanographic
Reeve M. Bailey (UMMZ); Marie-Louise Bauchot Research Institute, Durban); and Charles Wen-
(MNHN); Adam Ben-Tuvia (Sea Fisheries Re- ner (South Carolina Department of Marine
search Station, Haifa); M. Boeseman (RMNH); the Resources).
late James E. Bohlke (ANSP); Ian W. Brown Work at the Australian Museum in 1969-70 was
(formerly Senior Fisheries Officer, Fiji); Dan made possible through the National Marine Fish-
M. Carlsson (ZMK, formerly at Phuket Marine eries Service and the Trustees of the Australian
Biological Center, Thailand); F. Cervigon M. Museum, its Director at that time, Frank H.
(UDONECl); C. E. Dawson (GCRL); Alan R. Talbot and its Curator of Fishes, John R. Paxton.
Emery (ROM); William N. Eschmeyer (CAS); For assistance with computer processing we
William L. Fink (formerly at MCZ); Carter R. thank K. K. Beach and E. M. Hamilton of the
Gilbert (UF); C. G. Gruchy (NMC); Karsten Hartel George Washington University Center for Aca-
(MCZ); Philip C. Heemstra (RUSl); P. A. Hulley demic and Administrative Computing. L. C.
(SAM); Robert K. Johnson (FMNH); Paul Kahs- Hayek, Office of Computer Services, Smithsonian
bauer (formerly at NHMV); Robert J. Lavenberg Institution, provided consultation and suggestions
(LACM); Don E. McAllister (NMC); R. J. McKay regarding the statistical procedures used herein.
(formerly at WAM); Geoff McPherson (Queens- Ruth E. Gibbons and Gary A. Pettit assisted with
land Fisheries Service); Naercio A. Menezes data analysis.
(MZUSP); A. G. K. Menon (ZSI); Ian S. R. Munro The figures, which are an integral part of this
(CSIRO); Eugene L. Nakamura (NMFS, Panama paper, were drawn by Keiko Hiratsuka Moore.
City, Fla.); Js;>rgen G. Nielsen (ZMK); Han Nijssen Plates were prepared by Ruth E. Gibbons. Radio-
(ZMA); John R. Paxton (AMS); Thomas Potthoff graphs were taken by George Clipper and Ruth E.
(TABL); William J. Richards (TABL); C. Richard Gibbons. Jack Marquardt and his staff at the
Robins (University of Miami); Donn E. Rosen Smithsonian library were most helpful in finding
(AMNH); Richard Rosenblatt (SIO); B. R. Smith and obtaining early or obscure references. Typing,
673
 FISHERY BULLETIN: VOL. 82, NO.4

retyping, proofreading, xeroxing, and all the BANERJI, S. K.

other necessary clerical work was done by Arleen 1973. An assessment of the exploited pelagic fisheries of
the Indian Seas. In Proceedings of the Symposium on
McClain, Virginia R. Thomas, and Sara E. Col- Living Resources of the Seas Around India, Spec. Publ.,
lette. Robert H. Gibbs, Jr., Steven Gray, Melissa p.114-136. Cent. Mar. Fish. Res. Inst.
Lakich, Adrienne Mims, Linda Pushee Mercer, BARNARD, K. H.
and Frances Matthews Van Dolah participated in 1927. A monograph of the marine fishes of South Africa.
Ann. S. Afr. Mus. 21:419-1065.
some dissections. Drafts of the manuscript were
1948. Further notes on South African marine fishes. Ann.
read by A. D. Lewis, Eugene L. Nakamura, Izumi S. Afr. Mus. 36:341-406.
Nakamura, Thomas Potthoff, William J. Rich- BASTOS, J. R.
ards, James C. Tyler, and Austin B. Williams. 1966. Sobre a biometria da serra, Scomberomorus macu-
latus (Mitehill), da costa do Estado do Ceara. Arq. Estac.
BioI. Mar. Dniv. Fed. Ceara 6:113-117.
BASTOS, J. R., T. T. ALVES, C. A. ESTEVES ARARIPE, AND F. J.
LITERATURE CITED SIQUEIRA TELLES.
1973. SObre a elabora9iio de conservas de pescado em leite
ADKINS, G., AND M. J. BOURGEOIS. de coco e em oleos de algodao e de baba9u. Arq. Ci'mc.
1982. An evaluation of gill nets of various mesh sizes. La. Mar 13:25-29.
Dep. Wild. Fish. Tech. Bull. 36, 59 p. BAUCHOT, M. L.
ALCANTARA FILHO, P. DE. 1969. Les poissons de la collection de Broussonet au Mu-
1972a. Sobre a captura da serra, Scomberomorus macu- seum National d'Histoire Naturelle de Paris. Bull. Mus.
latus (Mitchill), com redes-de-espera, no Estado do Ceara. Nat. Hist. Nat. Paris, Ser. 2, 41:125-143.
Arq. Ciimc. Mar 12:77-84. BAUCHOT, M. L., AND M. BLANC.
1972b. Sobre a captura da cavala, Scomberomorus cavalla 1961. Catalogue des types de Scombroidei (poissons tele-
(Cuvier), com redes-de-espera, no Estado do Ceara. Arq. osteens Perciformes) des collections du Museum National
Ci'mc. Mar 12:133-138. d'Histoire Naturelle de Paris. Bull Mus. Nat. Hist. Nat.
ALLIS, E. P., JR. Paris, Ser. 2, 33:369-379.
1903. The skull, and the cranial and first spinal muscles
and nerves in Scomber scomber. J. Morphol. 18:45-328. BAUGHMAN, J. L.
AMEZCUA-LINARES, F. 1941. Scombriformes, new, rare, or little known in Texas
1977. Generalidades ictiologicas del sistema lagunar cos- waters with notes on their natural history or distribu-
tero de Huizache-Caimanero, Sinaloa, Mexico. An. Cen- tion. Trans. Tex. Acad. Sci. 24:14-26.
tro Ciime. Mar Limnol., Dniv. Nal. Auton. Mex. 4:1-26. 1950. Random notes on Texas fishes. Part II. Tex. J. Sci.
2;242~263.
AMEZCUA-LINARES, F., AND A. YAN'EZ-ARANCIBIA.
1980. Ecologia de los sistemas f1uvio-lagunares asociados BEAN, T. H.
a la Laguna de Terminos. El habitat y estructura de las 1903. Catalogue of the fishes of New York. N.Y. State
comunidades de peces. An. Centro Ci'mc. Mar Limnol., Mus. Bull. 60, 784 p.
Dniv. Nal. AutOn. Mex. 7:69-118. BEARDSLEY, G. L., JR., AND W. J. RICHARDS.
ANONYMOUS. 1970. Size, seasonal abundance, and length-weight rela-
1967. Fishes caught by trawling off the northwest coast of tion of some scombrid fishes from southeast Florida.
Borneo, Dec. 5-11, 1966. The "Oshoro Maru" Cruise 21. D.S. Fish Wildl. Serv., Spec. Sci. Rep. Fish. 595, 6 p.
Data Ree. Oceanogr. Obs. Explor. Fish., Hokkaido Dniv., BEAUMARIAGE, D. S.
No. 12, p_ 45-49. 1970. Current status of biological investigations of Flor-
1975. Pictoral catalogue of Chinese marine fishes. (In ida's mackerel fisheries. Proc. Gulf Caribb. Fish. Inst.
Chin.] Inst. Oceanol., Sinica Acad. and Shanghai Nat. 22d Annu. Meet. 1969:79-86.
Hist. Mus. Peoples Press, Shanghai, PRC., Vol. 1, 230 pI. 1973. Age, growth and reproduction of king mackerel,
1978. Major Queensland bid to track Spanish macker- Scomberomorus cavalla in Florida. Fla. Mar. Res. Publ.
el. Aust. Fish. 37(3):18-19. 1,45 p.
ARNOULT, A., AND P. FOURMANOIR. BEEBE, w., AND G. HOLLISTER.
1967. Sur une collection de poissons marins et de reptiles, 1935. The fishes of Dnion Island, Grenadines, British
provenant de l'Ile de Nossi-Be (Madagascar). Bull. Mus. West Indies, with the description of a new species of star-
Nat. Hist. Nat., Ser. 2, 39:134-142. gazer. Zoologica (N.Y.) 19:209-224.
ARTUNDUAGA PASTRANA, E. BEEBE, w., AND J. TEE-VAN.
1976. La sierra (Scomberomorus sierra Jordan y Starks) 1928. The fishes of Port-au-Prince Bay, Haiti, with a sum-
del Pacifico Colombiano. Divulg. Pesq., INDERENA, mary of the known species of marine fish of the island of
Bogota 8(4-5):1-72. Haiti and Santo Domingo. Zoologica (N.Y.) 10:1-279.
BAISSAC, J. DE B. BENNETT, E. T.
1964. The scombroid fishes of Mascarene waters. Proc. 1831. Characters of new genera and species of fishes from
Symp. Scombroid Fishes, Mar. BioI. A~soc. India, Symp. the Atlantic Coast of Northern Africa, presented by
Ser. I(Part 1):185-186. Captain Belcher, R.N. Proc. Zool. Soc. Lond., Part I:
1976. Poissons de mer des eaux de l'Ile Maurice. Proc. R. 145-148.
Soc. Arts Sci. Mauritius 3(2):191-266. 1832. Observations on a collection of fishes from the
BANERJEE, M. K., AND R. D. CHAKRUBARTY. Mauritius, presented by Mr. Telfair, with characters of
1972. Drift gill-netting in Lower Sundarbans, West Ben- new genera and species. Proc. Zool. Soc. Lond. 1830-
gal. Indian J. Fish. 16:75-81. 31:165-169.

674
COLLETTE and RUSSO: SPANISH MACKERELS
BEN-TuvIA, A. Akad. Kunst. J. Morino Comp., Ber!., Part 7, 144 p.
1968. Report on the fisheries investigations of the Israel 1797. Ichtyologie, ou histoire naturelle, generale et par-
South Red Sea Expedition, 1962. Bul!. Sea Fish. Res. ticuliere, des poissons. Dixieme Partie, 120 p. Berlin.
Stn. Haifa 52, p. 21-55. BLOCH, M. E., AND J. G. SCHNEIDER.
1971. Revised list of the Mediterranean fishes of Isra- 1801. Systema ichthyologiae. Berolini, 584 p.
e!. Isr. J. Zoo!' 20:1-39. BOESEMAN, M.
BEN-YAMI, M. 1947. Revision of the fishes collected by Burger and Von
1968. Observations on the distribution and behaviour of Siebold in Japan. Zoo!. Meded. Rijksmus. Nat. Rist.
pelagic schooling fish in the southern Red Sea. Bul!. Sea Leiden 28:1-242.
Fish. Res. Stn. Haifa 51, p. 31-46. 1964. Scombroid types in the Leidim Museum collection.
BERRIEN, P., AND D. FINAN. Proc. Symp. Scombroid Fishes, Mar. Bio!' Assoc. India,
1977a. Biological and fisheries data on king mackerel, Symp. Ser. I(Part 1):461-468.
Scomberomorus cavalla (Cuvier). Sandy Hook Lab. BOHLKE, J. E.
Tech. Ser. Rep. 8, 40 p. 1953. A catalogue of the type specimens of Recent fishes
1977b. Biological and fisheries data on Spanish mackerel, in the Natural History Museum of Stanford University.
Scomberomorus maculatus (Mitchill). Sandy Hook Lab. Stanford Ichthyo!. Bul!. 5:1-168.
Tech. Ser. Rep. 9, 52 p. BOHLKE, J. E., AND C. C. G. CHAPLIN.
BIGELOW, H. B., AND W. C. SCHROEDER. 1968. Fishes of the Bahamas and adjacent tropical wa-
1953. Fishes ofthe Gulf of Maine. U.S. Fish Wild!. Serv., ters. Livingstone Pub!. Co., Wynnewood, Pa., 771 p.
Fish. Bul!. 53:1-577. BOULENGER, G. A.
BINI, G., AND E. TORTONESE. 1899. Viaggio del Dott. Enrico Festa nel Darien e regioni
1955. Missione sperimentale di pesca nel Cile e nel Peru - vicine. Poissons de I'Amerique Centrale. Bol!. Mus.
pesci marini Peruviani. Bol!. Pesca Pisco Idrobio!., New Zoo!' Anat. Compo Torino 14(346):1-4.
Ser.,9(2):1-39. BREDER, C. M., JR.
BLACHE, J., J. CADENAT, AND A. STAUCH. 1936. Scientific results of the second oceanographic expe-
1970. Cles de determination des poissons de mer signales dition of the ·Pawnee" 1926. Heterosomata to Pediculati
dans l'atlantique oriental entre Ie 20e parallele Nord et from Panama to lower California. Bul!. Bingham Ocean-
Ie 15e paralleIe Sud. Faune Trop. XVIII, O.RS.T.O.M., ogr. Collect. Yale Univ. 2(3):1-56.
Paris, 480 p. 1948. Field book of marine fishes of the Atlantic coast from
BLANC, M., AND M. L. BAUCHOT. Labrador to Texas; being a short description of their
1964. Les Scombroidei (poissons tel{,osteens Perciformes) characteristics and habits with keys for their identifica-
du Museum National d'Histoire Naturelle de Paris. tion. G. P. Putnam's Sons, N.Y., 332 p.
Proc. Symp. Scombroid Fishes, Mar. Bio!' Assoc. India, BRIGGS,'J. C.
Symp. Ser. HPart 1):443-458. 1958. A list of Florida fishes and their distribution. Bul!.
BLANC, M., F. D'AUBENTON, AND P. FOURMANOIR. Fla. State Mus., Bio!. Sci. 2:223-318.
1965. A propos d'un Scombridae des eaux douces Cam- BUEN, F. DE.
bodgiennes: Scomberomorus sinensis (Lacepede, 1802). 1972. Clase V. Los peces tele6stomos (TeleostomiJ. In
Bul!. Mus. Hist. Nat., Ser. 2, 37:121-123. Luis Cendrero (editor), Zoologia Hispanoamericana,
BLEEKER,P. Vertebrados, p. 55-332. Editorial Porrua, Mexico,
1851a. Over eenige nieuwe geslachten en soorten van D.F.
Makreelachtige visschen van der Indischen Archipel. BULLIS, H. R, R B. ROE, AND J. C. GATLIN.
Nat. Tijdschr. Ned.-Indie 1:341-372. 1972. The Southeast Fisheries Center bionumeric code
185lb. Visschen van Banka. Nat. Tijdschr. Ned.-Indie Part I: Fishes. U.S. Dep. Commer.; NOAA Tech. Rep.
1:159-161. NMFS SSRF-659, 95 p.
1852. Bijdrage tot de kennis der Makreelachtige visschen BURNE, RH.
van den Soenda-Molukschen Archipe!. Verh. Bat. Gen. 1909. The anatomy of the olfactory organ of teleostean
24(5):1-93. fishes. Proc. Zoo!' Soc. Lond. 1909:610-663.
1853. Nalezingen op de icthyologische fauna van Bengalen BUTZ, G., AND R J. MANSUETI.
en Hindostan. Verh. Bat. Gen. 25:1-164. 1962. First record of the king mackerel, Scomberomorus
1860. Dertiende bijdrage tot de kennis der vischfauna van cavalLa, in northern Chesapeake Bay, Maryland. Chesa-
Borneo. Act. Soc. Sci. Indo-Neerl. 8:1-64. peake Sci. 3:130-135.
1861a. Mededeeling omtrent vischsoorten, nieuw voor de CADENAT,J.
kennis der fauna van Singapoera. Vers!. Akad. Arnst. 1937. Recherches systematiques sur les poissons littoraux
12:28-63. de la Cote Occidental d'Afrique, recoltes par Ie navire
1861b. lets over de vischfauna van het eiland Pinang. President Theodore-Tissier, au cours de sa 5" croisiere
Vers!. Akad. Arnst. 12:64-80. (1936). Rev. Trav. Off. Sci. Tech. pech. Marit. 10:
1873. Memoire sur la faune ichthyologique de Chine. 423-562.
Ned. Tijdschr. Dierk. 4:113-154. 1947. Noms vernaculaires des principales formes d'ani-
1874. Poissons de Madagascar et de l'Ile de la Reunion. maux marins des cotes de l'Afrique Occidentale Fran-
In Recherches sur la faune de Madagascar et de ses 9aise. Inst. Fr. Afr. Noire, Cat. II, 56 p.
dependances d'apres les decouvertes de F. P. L. Pollen CANTOR, T. E.
et D. C. van Dam. 4 Partie. Leiden, 104 p. 1849. Catalogue of Malayan fishes. J. Asiat. Soc., Ben-
1879. Contribution ilIa faune ichthyologique de l'Ile Mau- gal 18:983-1443.
rice. Verh. Akad. Arnst. 18:1-23. CASTELNAU, F. DE.
BLOCH,M.E. 1855. Poissons. In Animaux nouveaux ou rares recueillis
1793. Naturgeschichte der ausliindischen Fische. Konig!. pendant I'expedition dans les parties centrales de

675
 FISHERY BULLETIN: VOL. 82, NO.4

I'Amerique du Sud, de Rio de Janeiro a Lima, et de Lima COLLETTE, B. B.

au Para; executee par ordre du gouvernement Fran~ais 1966. Revue critiques des types de Scombridae des collec-
pendant les annees 1843 a 1847, Tome II, p. 1-112. P. tions du Museum National d'Histoire Naturelle de Paris.
Bertrand, Paris. Bull. Mus. Natl. Hist. Nat. Paris, Ser. 2, 38:362-375.
1879. Essay on the ichthyology of Port Jackson. Proc. 1970. Rastrelliger kanagurta, another Red Sea immigrant
Linn. Soc. NB.w. 3(4):347-402. into the Mediterranean Sea, with a key to the Mediter-
CASTRO-AGUIRRE, J. L., J. ARVIZU-MARTINEZ, AND J. PAEZ- ranean species of Scombridae. Bull. Sea Fish. Res. Stn.
BARRERA. <Haifa) 54:3-6.
1970. Contribucion al conocimiento de los peces del Golfo 1978. Scombridae. In W. Fischer (editor), FAO species
de California. Rev. Soc. Mex. Hist. Nat. 31:107-181. identification sheets for fishery purposes, Western Cen-
CERVIGON M., F. tral Atlantic, fishing area 31; Vol. IV, 34 p. FAO, Rome.
1966. Los peces marinos de Venezuela. Thmo II. Estac. 1979. Adaptations and systematics of the mackerels
Invest. Mar. Margarita, Fund. La Salle De Cienc. Nat., and tunas. In G. D. Sharp and A. E. Dizon (editors), The
Caracas. Monogr. 12, p. 449-951. physiological ecology of tunas, p. 7-39. Acad. Press, NY.
CHABANAUD,P. 1981. Scombridae. In W. Fischer, G. Bianchi, and W. B.
1926. Inventaire de la faune ichtyologique de I'Indo- Scott (editors), FAO species identification sheets for
chine. Servo Oceanogr. pech. Indochine, Stn. Mar. fishery purposes, Eastern Central Atlantic, fishing area
Cauda, Note 1, 26 p. 34; Vol. III, 32 p. FAO and Dep. Fish. Oceans, Ottawa.
CHABANAUD, P, AND T. MONOD. 1983. Recognition oftwo species of double-lined mackerels
1927. Les poissons de Port-Etienne. Bull. Comito Etud. (Grammatorcynus: Scombridae). Proc. BioI. Soc. Wash.
Hist. Sci. Afr. Occident. Fr. 1926:225-287. 96:715-718.
CHACKO, P 1. COLLETTE, B. B., AND L. N. CHAO.
1949. Food and feeding habits of the fishes of the Gulf of 1975. Systematics and morphology of the bonitos (Sarda)
Manaar. Proc. Indian Acad. Sci. Sec. B, 29:83-97. and their relatives (Scombridae, Sardini). Fish. Bull.,
1950. Marine plankton from waters around the Krusadai UB. 73:516-625.
Island. Proc. Indian Acad. Sci. Sec. B, 31:162-174. COLLETTE, B. B., AND C. E. NAUEN.
CHACKO, P 1., S. D. THOMAS, AND C. M. PILLAY. 1983. FAO species catalogue. Vol. 2. Scombrids of the
1968. Scombroid fisheries of Madras State, India. Proc. world. An annotated and illustrated catalogue of tunas,
Symp. Scombroid Fishes. Mar. BioI. Assoc. India, Symp. mackerels, bonitos and related species known to date.
Ser. HPart 3):1006-1008. FAO Fish. Synop. 125, 137 p.
CHAINE, J. COLLETTE, B. B., AND J. L. Russo.
1957. Recherches sur les otolithes des poisson". Etude 1979. An introduction to the Spanish mackerels, genus
descriptive et comparative de la sagitta des teleosteens Scomberomorus. In E. L. Nakamura and H. R. Bullis,
(VII). Bull. Cent. Etud. Rech. Sci. Biarritz 1:463-557. Jr. (editors), Proceedings of the Mackerel Colloquium, p.
CHAPMAN, W. M. 3-16. Gulf States Mar. Fish. Comm. No.4.
1946. Observations on tuna-like fishes in the tropical 1980. Scomberomorus munroi, a new species of Spanish
Pacific. Calif. Fish Game 32:165-170. mackerel from Australia and New Guinea. Aust. J. Mar.
CHEVEY,P Freshw. Res. 31(1):241-250.
1934. Revision synonymique de l'ouevre ichtyologique de 1985. Interrelationships of the Spanish mackerels (Pisces:
G. Tirant. In!'t. Oceanogr. Indochine, Stn. Mar. Cauda, Scombridae: Scomberomorus) and their copepod para-
Note 7, 291 p. sites. Cladistics 1:141-158.
CHEVEY, P., AND J. DURAND. COLLETTE, B. B., J. L. RUSSO, AND L. A. ZAVALLA-CAMIN.
1945. Principaux poissons comestibles d'Indochine. Inst. 1978. Scomberomorus brasiliensis, a new species of Span-
Oceanogr. Indochine, 42 p. ish mackerel from the western Atlantic. Fish. Bull.,
CHIRICHIGNO F., N. UB.76:273-280.
1969. Lista sistematica de los peces marinos comunes para COLLETTE, B. B., F. H. TALBOT, AND R. H. ROSENBLATT.
Ecuador, Peru, y Chile. Conf. Explot. Conserv. Riquezas 1963. The first California record of sierra, Scomberomorus
Marit. Pac. Surv., 108 p. sierra Jordan and Starks. Calif. Fish Game 49:53-54.
1974. Clave para identificar los peces marinos del Peru. CONAND,F.
Inst. Mar. Peru, Inf. 44:1-387. 1970. Distribution et abondance des larvaes de quelques
CLEMENS, H. B. famille et especes de poissons des cotes senegambiennes
1956. Rearing larval scombrid fishes in shipboard aquaria. en 1968. O.R.S.T.O.M. Dakar Doc. Sci. Provo No. 26, 52 p.
Calif. Fish Game 42:69-79. CONRAD,G.M.
1957. Fishes collected in the tropical eastern Pacific, 1954. 1938. The osteology and relationships of the wahoo (Acan-
Calif. Fish Game 43:299-307. thocybium solandri), a scombrid fish. Am. Mus. Nat.
CLEMENS, H. B., AND J. C. NOWELL. Hist. Novit. 1000, 32 p.
1963. Fishes collected in the eastern Pacific during COOPER,A.
tuna cruises, 1952 through 1959. Calif. Fish Game 1982. A preliminary study of the fishery for cero mackerel
49:240-264. (Scomberomorus regalis, Bloch) in Jamaican waters.
CLOTHIER, C. R. Proc. 34th GulfCaribb. Fish. Inst., p. 149-155.
1950. A key to some southern California fishes based on COSTA, R. S. DA, AND H. T. DE ALMEIDA.
vertebral characters. Calif. Div. Fish Game, Fish Bull. 1974. Notas sabre a pesca da cavala e da serra no Ceara -
79,83 p. Dados de 1971 a 1973. Arq. Cienc. Mar 14:115-122.
COATES,G. COSTA, R. S. DA, AND M. P PAIVA.
1950. Fishing on the Barrier Reef and inshore. T. WiII- 1969. Notas sabre a pesca da cavala e da serra no Ceara -
mett and Sons, Townsville, 72 p. Dados de 1968. Arq. Cienc. Mar 9:89-95.

676
COLLETTE and RUSSO: SPANISH MACKERELS
CRAIG, J. A. DEVARAJ,M.
1926. A new fishery in Mexico. Calif. Fish Game 12: 1976. Discovery ofthe scombrid Scomberomorus koreanus
166-169. (Kishinouye) in India, with taxonomic discussion on the
CRESSEY, R. F., B. B. COLLETTE, AND J. L. RussO. species. Jpn. J. Ichthyol. 23(2):79-87.
1983. Copepods and scombrid fishes: a study in host- 1977. Osteology and relationships of the Spanish macker-
parasite relationships. Fish. Bull., U.S. 81:227-265. els and seerfishes ofthe tribe Scomberomorini. IndianJ.
CRESSEY, R. F., AND H. B. CRESSEY. Fish. 22:1-67.
1980. The parasitic copepods of mackerel- and tuna-like 1982. Age and growth of three species of seerfishes
fishes (Scombridae) of the world. Smithson. Contrib. Scomberomorus commerson, S. guttatus and S. lineo-
Zool. 311:1-186. latus. Indian J. Fish. 28:104-127.
CROKER, R. S. DE VIS, C. W.
1933. The California mackerel fishery. Calif. Div. Fish 1884. New fishes in the Queensland Museum. No.3. Proc.
Game, Fish Bull. 40,149 p. Linn. Soc. N.S.W. 9:537-547.
1937. Monterey Spanish mackerel taken at Long Beach. DHAWAN, R. M., P. V. S. NAMBOOTHIRI, AND V. G. GOPINATHAN.
Calif. Fish Game 23:245-246. 1972. Results of trolling line operations in Goa waters
CUVIER, G. during 1965-68. Indian J. Fish. 14:181-187.
1829. Le regne animal. Nouv. ed. Tom. II. Deterville, DOl, T., AND D. MENDIZABAL.
Paris, 406 p. 1979. Evaluacion preliminar de la poblacion de sierra,
CUVIER, G., AND A. VALENCIENNES. Scomberomorus maculatus (Mitchill), frente a las costas
1831. Histoire naturelle des poissons. Vol. 8,375 p. F. G. de Veracruz. In E. L. Nakamura and H. R. Bullis, Jr.
Levrault, Paris. (editors); Proceedings ofthe Mackerel Colloquium, p. 43-
DAGET, J., AND A. ILTIS. 55. Gulf States Mar. Fish. Comm. No.4.
1965. Poissons de Cote d'Ivoire, eaux douces et saumatres. DOMANTAY, J. S.
Mem. Inst. Fr. Afr. Noire 74, 385 p. 1940. The fishing industry of Margosatubig. Philipp. J.
DAHL,G. Sci. 72:371-382.
1971. Los peces del norte de Colombia. INDERENA, DRESSLAR, F. B., AND B. FESLER.
Bogota, 391 p. 1889. A review of the mackerels (Scombrinae) of America
D'AUBENTON, F., AND M. BLANC. and Europe. Bull. U.S. Fish Comm. 7:429-446.
1965. Etude systematique et biologique de Scomber- DURAND,J.
omorus sinensis (Lacepede, 1802), poisson des eaux douces 1940. Notes sur quelques poissons d'especes nouvelles ou
du Cambodge. Bull. Mus. Natl. Hist. Nat., Ser. 2, 37: peu connus des eaux douces cambodgiennes. Notes Inst.
233-243. Oceanogr. Indochine 36:1-40.
DAY, F. DWINELL, S. E., AND C. R. FUTCH.
1873. Report on the sea fish and fisheries of India and 1973. Spanish and king mackerel larvae and juveniles in
Burma. Gov. Press, Calcutta, 86 + 332 p. the northeastern Gulf of Mexico, June through October
1878. The fishes of India; being a natural history of the 1969. Fla. Dep. Nat. Resour., Mar. Res. Lab., Leaf!. Ser.
fishes known to inhabit the seas and fresh waters ofIndia, 4, pt. 1, No. 24, p. 1-14.
Burma, and Ceylon. Vol. 1, p.1-778. Bernard Quaritch, EARLL, R. E.
Lond. 1883. The Spanish mackerel, Cybium maculatum (Mitch.)
DEARDORFF, T. L., AND R. M. OVERSTREET. Ag.; its natural history and artificial propagation, with
1981. Larval Hysterothylacium (= Thynnascaris) (Nema- an account of the origin and development of the fish-
toda: Anisakidae) from fishes and invertebrates in the ery. Rep. U.S. Comm. Fish Fish. 1880:395-426.
Gulf of Mexico. Proc. Helminthol. Soc. Wash. 48:113- ECKLES, H. H.
126. 1949. Observations on juvenile oceanic skipjack (Katsu-
DE BEAUFORT, L. F. wonus pelamis) from Hawaiian waters and sierra mack-
1951. Percomorphi (concluded). The fishes of the Indo- erel (Scomberomorus sierra) from the eastern Pacific.
.Australian Archipelago. Vol. 9, p. 1-242. E. J. Brill, U.S. Fish Wildl. Serv., Fish. Bull. 51:245-250.
Leiden. ERDMAN, D. S.
DELSMAN, H. C. 1949. Does the Spanish mackerel, Scomberomorus macu-
1931. Fish eggs and larvae from the Java Sea. 18. The latus (Mitchill), occur throughout the West Indies?
genus Cybium, with remarks on a few other Scom- Copeia 1949:301.
bridae. Treubia 13:401-410. 1956. Recent fish records from Puerto Rico. Bull. Mar.
DELSMAN, H. C., AND J. D. F. HARDENBERG. Sci. Gulf Caribb. 6:315-340.
1934. De· Indische Zeevisschen en Zeevisscherij. Bibl. 1971. Notes on fishes from the Gulf of Nicoya, Costa Rica.
Ned.-Ind. Naturhist. Ver., VI, Batavia-Centrum, 338 p. Rev. BioI. Trop. 19:59-71.
DE SYLVA, D. P. 1977. Spawning patterns of fish from the northeastern
1955. The osteology and phylogenetic relationships of the Caribbean. In Cooperative Investigations of the Carib-
blackfin tuna, Thunnus at/anticus (Lesson). Bull. Mar. bean and adjacent regions II, p. 145-169. FAO Fish.
Sci. Gulf Caribb. 5:1-41. Rep. 200.
DEVANE, J. C., JR. EVERMANN, B. w., AND O. P. JENKINS.
1978. Food of king mackerel, Scomberomorus caval/a, in 1891. Report upon a collection of fishes made at Guaymas,
Onslow Bay, North Carolina. Trans. Am. Fish. Soc. Sonora, Mexico, with descriptions of new species. Proc.
107:583-586. U.s. Natl. Mus. 14:121-165.
DEVANESEN, D. w., AND K. CHIDAMBARAM. EVERMANN, B. w., AND R. LEWIS.
1953. The common food-fishes of the Madras State. Gov. 1917. The fishes of the west coast ·of Peru and the Titicaca
Press, Madras, 79 p. Basin. Bull. U.S. Natl. Mus. 95, 166 p.

677
 FISHERY BULLETIN: VOL. 82, NO.4

EVERMANN, B. w., AND M. C. MARSH. Calif. Press, Berkeley, 179 p.

1902. The fishes of Porto Rico. Bull. U.S. Bur. Fish. FlTCH, J. E., AND S. A. SCHULTZ.
20:51-350. 1978. Some rare and unusual occurrences of fishes off
FABLE, W. A., JR., H. A. BRUSHER, L. TRENT, AND J. FlNNE- California and Baja California. Calif. Fish Game 64:
GAN,JR. 74-92.
1981. Possible temperature effects on charter boat catches FONTELES-FILHO, A. A.
of king mackerel and other coastal pelagic species in 1968. Sobre a captura e abundiincia da cavala e da serra
northwest Florida. Mar. Fish. Rev. 43(8):21-26. nos pesqueiros do Estado do Ceara. Arq. Est. BioI. Mar
FAGADE, S. 0., AND C. I. O. OLANIYAN. Univ. Fed. Ceara 8:133-137.
1973. The food and feeding interrelationship of the fishes FONTELES-FlLHO, A. A., AND P. DE ALCANTARA-FlLHO.
in the Lagos Lagoon. J. Fish BioI. 5:205-225. 1977. Curva de seletividade de redes-de-espera utilizadas
1974. Seasonal distribution of the fish fauna of the Lagos na captura de serra, Scomberomorus maculatus (Mitch-
Lagoon. I.F.A.N. Bull. 36A(1):244-252. iii). Arq. Ciemc. Mar 17:53-62.
FAO. FOURMANOIR, P.
1984. 1982 yearbook of fishery statistics. Catches and 1957. Poissons teleosteens des eaux malgaches du Canal
landings. Vol. 54, 393 p. FAO Fish. Ser. 21 and FAO de Mozambique. Mem. Inst. Sci. Madagascar, Ser. F,
Stat. Ser. 52, Rome, Italy. Oceanogr. 1:1-316.
FARRAGUT, R. N. 1966. Nouvelle denomination proposee pour un Scom-
1972. Effects of some antioxidants and EDTA on the bridae du canal de Mozambique: Scomberomorus plu-
development of rancidity in Spanish mackerel (Scom- rilineatus nov. sp. Bull. Mus. Hist. Nat. Paris, Ser. 2,
beromorus maculatus) during frozen storage. U.S. Dep. 38:223-226.
Commer., NOAA Tech. Rep. NMFS SSRF 650, 12 p. FOURMANOIR, P., AND A. CROSNIER.
FARRIS, J. S. 1964. Rapport sur la campagne de peche au thon a la
1970. Methods for computing Wagner trees. Syst. Zool. palangre derivante, faite par Ie "Maran Atha" dans les
19:83-92. environs de Majunga (Cote Nord-Ouest) du 21 Septembre
FARRIS, J. S., A. G. KLUGE, AND M. J. ECKARDT. au 15 October 1961. Proc. Symp. Scombroid Fishes, Mar.
1970. A numerical approach to phylogenetic systemat- BioI. Assoc. India, Symp. Ser. I(Part 1):379-388.
ics. Syst. Zool. 19:172-189. FOURMANOIR, P., AND P. LABOUTE.
FERNANDO, E. F W. 1976. Poissons de Nouvelle Caledonie et des Nouvelles
1972. Species composition of fish captured by trawlers in Hebrides. Les Editions Pacifique, Papeete, 376 p.
the Wadge Bank. Proc. Indo-Pac. Fish. Counc. 13(I1I): FOWLER, H. W.
521-531. 1905. New, rare or little-known scombroids. No. 1. Proc.
FlERSTINE, H. L., AND V. WALTERS. Acad. Nat. Sci. Phila. 56:757-771.
1968. Studies in locomotion and anatomy of scombroid 1918. A list of Philippine fishes. Copeia No. 58, p. 62-
fishes. Mem. South. Calif. Acad. Sci. 6:1-31. 65.
FlNUCANE, J. H., AND L. A. COLLINS. 1928. Further notes and descriptions of Bombay shore
1984. Reproductive biology of cero, Scomberomorus regal- fishes. J. Bombay Nat. Hist. Soc. 33:100-119.
is, from the coastal waters of south Florida. Northeast 1929. New and little-known fishes from the Natal coast.
Gulf Sci. 7(1):101-107. Ann. Natal Mus. 6:245-264.
FISCHER, M. 1934. Fishes obtained by Mr. H. W. Bell-Marley chiefly in
1980. Size distribution, length-weight relationships, sex Natal and Zululand in 1929 to 1932. Proc. Acad. Nat. Sci.
ratios, and seasonal occurrence of king mackerel (Scom- Phila. 86:405-514.
beromorus caualia) off the southeast Louisiana coast. 1936. The marine fishes of West Africa based on the
La. Dep. Wildl. Fish., Contrib. Mar. Res. Lab. Tech. Bull. collection of the American Museum Congo Expedition,
31, p. 1-21. 1909-1915. Bull. Am. Mus. Nat. Hist. 70:607-1493.
FITCH, J. E. 1938. The fishes of the George Vanderbilt South Pacific
1948. Some new and unusual fishes from southern Cali- Expedition, 1937. Acad. Nat. Sci. Phila. Monogr. 2,
fornia. Calif. Fish Game 34:133-135. 349p.
1950. Notes on some Pacific fishes. Calif. Fish Game 1944. The fishes. In Results of the fifth George Vander-
36:65-73. bilt Expedition (1941) (Bahamas, Caribbean Sea, Panama,
1952. Distributional notes on some Pacific Coast marine Galapagos Archipelago and Mexican Pacific Islands), p.
fishes. Calif. Fish Game 38:557-564. 57-529. Acad. Nat. Sci. Phila., Monogr. 6.
1969. Fish remains, primarily otoliths, from a Ventura, 1945. A study of the fishes of the southern piedmont and
California, Chumash village site (Ven-3). Mem. South. coastal plain. Acad. Nat. Sci. Phila. Monogr. 7, 408 p.
Calif. Acad. Sci. 8, App. A, p. 56-71. 1959. Fishes of Fiji. Gov. Fiji, Suva, 670 p.
FlTCH, J. E., AND W. L. CRAIG. FRADE, F., AND E. POSTEL.
1964. First records for the bigeye thresher (Alopias super- 1955. Contribution a l'etude de la reproduction des scom-
ciliosus) and slender tuna (Aliothunnus (aliai) from brides et thonides de I' Atlantique tropical. Rapp. P.-V.
California, with notes on eastern Pacific scombrid oto- Reun. Cons. Perm. Int. Explor. Mer 137:33-35.
liths. Calif. Fish Game 50:195-206. FRANCA, M. DE L. P. DA.
FlTCH, J. E., AND A. O. FLECHSIG. 1964. Peixes emalhados nas redes lagosteiras em agnas
1949. A brief account of the Monterey Spanish mack- costeiras de Angola. Notas Mimeogr. Cent. BioI. Piscat.
erel (Scomberomorus concolor). Calif. Fish Game 35: 40:1-13.
275-280. FRASER, A. J.
FlTCH, J. E., AND R. J. LAVENBERG. 1953. The fisheries of Western Australia. West. Aust.
1971. Marine food and game fishes of California. Univ. Fish. Dep., Fish. Bull. 4, 21 p.

678
COLLE'ITE and RUSSO: SPANISH MACKERELS

FRASER-BRUNNER, A. GOPALAN NAYAR, S.

1950. The fishes of the family Scombridae. Ann. Mag. 1958. A preliminary account of the fisheries of Vizh-
Nat. Hist., Ser. 12, 3:131-163. ingam. Indian J. Fish. 5:32-55.
FRITZSCHE, R. A. GORBUNOVA, N. N.
1978. Scombridae-mackerels and tunas. In Develop- 1965a. Seasons and conditions of spawning of the scom-
ment of fishes of the mid-Atlantic Bight. Vo!' V, Chaeto- broid fishes (Pisces, Scombroidei). [In Russ., Eng!.
dontidae through Ophidiidae, p. 61-167. U.S. Fish Wild!. summ.] Tr. Inst. Okeano!. Akad. Nauk SSSR 80:36-61.
Serv., Bio!' Servo Prog. FWS/OBS-78/12. 1965b. On spawning of scombroid fishes (Pisces, Scom-
FROST,G.A. broidei) in the Gulf of 'Ibnkin (South China Sea). [In
1928. A comparative study of the otoliths ofthe neopteryg- Russ., Eng!. summ.] Tr. Inst. Okeano!' Akad. Nauk
ian fishes-suborder Scombroidea. Ann. Mag. Nat. SSSR 80:167-176.
Hist., Ser. 10, 2:329-330. GRANT,E.M.
GARZON F., J., AND A. ACERO P. 1965. Guide to fishes. Queens!. Dep. Harbours Mar.,
1983. Notas sobre la pesca y los peces comerciales d la Isla Brisbane, 280 p.
de Providencia (Colombia), incluyendo nuevos registros 1972. Guide to fishes. 2d ed. Queens!. Dep. Primary
para el Caribe occidenta!' Caribb. J. Sci. 19(3-4):9-19. Ind., Brisbane, 472 p.
1975. Guide to fishes. 3d ed. Co-ord.-Gen. Dep., Bris-
GEORGE, C. J., AND V. ATHANASSIOU.
bane, 640 p.
1965. On the occurrence of Scomberomorus commersoni
1978. Guide to fishes. 4th ed. Dep. Harbours Mar.,
(Lacepooe) in St. George Bay, Lebanon. Doriana, Ann.
Brisbane, 768 p.
Mus. Civ. Stor. Nat. Genova 4(157):1-4.
1982. Guide to fishes. 5th ed. Dep. Harbours Mar.,
1967. A two-year study ofthe fishes appearing in the seine
Brisbane, 896 p.
fishery of St. George Bay, Lebanon. Ann. Mus. Civ. Stor.
GRAS, R.
Nat. Genova 76:237-294.
1961. Liste des poissons du Bas-Dahomey faisant partie de
GESTEIRA, T. C. V.
la collection du laboratoire d'Hydrobiologie du Service des
1972. Sobre a reproducao e fecundidade da serra, Scom-
Eaux, Forets et Chasses du Dahomey. Bul!. Inst. Fr. Afr.
beromorus maculatus (Mitchill), no Estado do Ceara.
Noire 23A:572-586.
Arq. Cilmc. Mar 12:117-122.
GUNTER,G.
GESTEIRA, T. C. v., AND A. L. LOBO DE MESQUITA.
1945. Studies on marine fishes of Texas. Pub!. Inst. Mar.
1976. Epoca de reproducao, tamanho e idade na primeira
Sci. Univ. Tex. 1:1-190.
desova de cavala e da serra, na costa do Estado do Ceara
GUNTHER, A.
(Brasil). Arq. Cilmc. Mar 16:83-86.
1860. Catalogue of the acanthopterygian fishes in the
GIBBS, R. H., JR., AND B. B. COLLE'ITE. collection of the British Museum. Vo!. 2, 548 p. Taylor
1967. Comparative anatomy and systematics of the tunas, and Francis, Lond.
genus Thunnus. U.S. Fish Wild!. Serv., Fish. Bul!. 1873. On a collection of fishes from Chefoo, North China.
66:65-130. Ann. Mag. Nat. Hist., Ser. 4, 12:377-380.
GILBERT, C. H., AND E. C. STARKS. 1880. Report on the shore fishes procured during the
1904. The fishes of Panama Bay. Mem. Calif. Acad. Sci. voyage of H.M.S. Challenger in the years 1873-1876.
4,304 p. Zoo!' Challenger Exped., Part 6, 82 p.
GILCHRIST, J. D. F., AND W W THOMPSON. HALE,M. B.
1911. Descriptions of fishes from the coast of Natal (Part 1979. Preservation technology for Spanish mackerel and
111.). Ann. S. Afr. Mus. 11:29-58. related species-a literature review. In E. L. Naka-
1917. A catalogue of the sea fishes recorded from Natal, mura and H. R. Bullis, Jr. (editors), Proceedings of the
part 2. Ann. Durban Mus. 1(4):291-431. Mackerel Colloquium, p. 73-77. Gulf States Mar. Fish.
GILL, T. N. Comm. No.4.
1862. On the limits and arrangement of the family of HAMADA, T., AND S. IWAI.
scombroids. Proc. Acad. Nat. Sci. Phila. 14:124-127. 1967. Biological studies on sawara resources in Harima-
GINES, H., AND F. CERVIGON M. Nada and adjacent waters-I. On some morphological
1968. Exploraci6n pesquera en las costas de Guayana characters and growth. [In Jpn., Eng!. abstr.] Bull.
y Surinam, ano 1967. Mem. Soc. Cienc. Nat. La Salle Jpn. Soc. Sci. Fish. 33:1013-1020.
28:5-96. HARDENBERG,J.D.F.
1931. The fishfauna of the Rokan Mouth. Treubia 13:
GNANAMU'ITU, J. C. 81-168.
1971. Osteology of the Indian mackerel, Rastrelliger kana- 1934. Additional notes to my paper "The fishfauna of the
gurta (Cuvier). Indian J. Fish. 13:1-26. Rokan Mouth." Treubia 14:299-312.
GOLANI, D., AND D. KREDO. 1936. On a collection of fishes from the estuary and the
1981. Hook-and-line and entangling net fisheries survey lower and middle course of the River Kapuas (W Borneo).
along the Mediterranean coast oflsrae!. [In Heb., Eng!. Treubia 15:225-254.
abstr.] Fish. Fish Breed. Isr. 16(1):31-42. 1937. Hydrological and ichthyological observations in the
GooDE,G.B. mouth of the Kumai River (S.W Borneo). Treubia 16:
1884. The food fishes of the United States. In The fish- 1-14.
eries and fishery industries of the United States; Sect. 1, HERRE,A. W
Part 3, p. 163-682. 1931. The fishes of the Herre 1931 Philippine expedition
GOODING, R. M. with descriptions of 17 new species. Newspaper Enter-
1963. The olfactory organ of the skipjack Katsuwonus prise Ltd., Hong Kong, 106 p. Reprinted by N. K. Gregg,
pelamis. FAO Fish. Rep. 6, p. 1621-1631. Kentfield, Calif.

679
 FISHERY BULLETIN: VOL. 82, NO.4

1933. A check list of fishes from Dumaguete, Oriental 1964b. Olfactory organs of tunas with special reference to
Negros, P. I., and its immediate vicinity. J. Pan-Pac. their systematic significance. Bull. Misaki Mar. Bio!.
Res. lnst. 8(4):6-11. Inst., Kyoto Univ:, No.7, p. 1-8.
1936. Fishes of the Crane Pacific Expedition. Field Mus. JENKINS, G. P., N. E. MILWARD, AND R. F. HARTWICK.·
Nat. Hist., Zoo!' Ser., 21:1-472. 1984. Identification· and description of larvae of Span-
1953. Check list of Philippine fishes. U.S. Fish Wildl. ish mackerels, genus Scomberomorus (Teleostei: Scom-
Serv.,Res. Rep. 20, 977 p. bridae), in shelf waters ofthe Great Barrier Reef. Aust.
HERRE, A. w., AND E. S. HERALD. J. Mar. Freshw. Res. 35:341-353.
1951. Noteworthy additions to the Philippine fish fauna JOHNSON, A. G.
with descriptions ofa new genus and species. Philipp. J. 1981. Electrophoretic patterns of proteins in Spanish
Sci. 79:309-340. mackerel (Scomberomorus maculatus). U.S. Dep. Com-
HERRE, A. w., AND G. S. MYERS. mer., NOAA Tech. Memo. NMFS-SEFC-76, 11 p.
1937. A contribution to the ichthyology of the Malay JOHNSON, A. G., W. A. FABLE, JR., M. L. WILLIAMS, AND L. E.
Peninsula. Bull. Raffles Mus. 13:5-75. BARGER.
HERRE, A. w., AND A. F. UMALI. 1983. Age, growth, and mortality of king mackerel,
1948. English and local common names of Philippine Scomberomorus cavalla, from the southeastern United
fishes. U.S. Fish Wild!. Serv., Circ. 14, 128 p. States. Fish. Bull., U.S. 81:97-106.
HIGGINS, E. JOHNSON, G. D.
1920. Mexican fishes in California ports. Calif. Fish 1975. The procurrent spur: An undescribed perciform
Game 6(1):33-34. caudal character and its phylogenetic implications. Oc-
HILDEBRAND, S. F. cas. Pap. Calif. Acad. Sci. 121, 23 p.
1946. A descriptive catalog of the shore fishes of Peru. JONES, S.
U.s. Natl. Mus. Bull. 189, 530 p. 1962. Notes on eggs, larvae and juveniles of fishes from
HILDEBRAND, S. F., AND L. E. CABLE. Indian waters. VIII. Scomberomorus guttatus (Bloch and
1938. Further notes on the development and life history Schneider), IX. Scomberomorus commerson (LacepMe)
of some teleosts at Beaufort, N.C. Bull. U.S. Bur. Fish. and X. Scomberomorus lineolatus (Cuvier). Indian J.
48: 505-642. Fish. 8:107-120.
HILDEBRAND, S. F., AND W. C. SCHROEDER. 1968. The scombroid fishery of India-present and fu-
1928. Fishes of Chesapeake Bay. Bull. U.S. Bur. Fish. ture. Proc. Symp. Scombroid Fishes, Mar. BioI. Assoc.
43:1-388. India, Symp. Ser. l(Part 3):994-1000.
HOLBROOK, J. E. JONES, S., AND M. KUMARAN.
1860. Ichthyology of South Carolina. 2d ed. Russell and 1964. Eggs, larvae and juveniles of Indian scombroid
Jones, Charleston, S.C., 205 p. fishes. Proc. Symp. Scombroid Fishes, Mar. BioI. Assoc.
HONMA,Y. India, Symp. Ser. l(Part 1):343-378.
1952. A list ofthe fishes collected in the Province ofEchigo, JONES, S., AND E. G. SILAS.
including Sado Island. [In Jpn.J Jpn. J. Ichthyol. 2: 1961. On fishes ofthe subfamily Scomberomorinae (family
138-146. Scombridae) from Indian waters. Indian J. Fish. 8:
HORN, M. H., AND L. G. ALLEN. 189-206.
1978. A distributional analysis of California coastal ma- 1964. A systematic review of the scombroid fishes of In-
rine fishes. J. Biogeogr. 5:23-42. dia. Proc. Symp. Scombroid Fishes, Mar. BioI. Assoc.
HUBBS, C. L. India, Symp. Ser. l(Part 1):1-105.
1936. Fishes of the Yucatan Peninsula. Publ. Carnegie JONES, S., E. G. SILAS, AND E. DAWSON.
lnst. Wash. 457, p. 157-287. 1960. New records of scombroid fishes from the Andaman-
HUTCHINS, B. Nicobar waters. J. Mar. BioI. Assoc. India 2:136-137.
1979. A guide to the marine fishes of Rottnest Island. JORDAN, D. S.
Creative Research, Perth, 103 p. 1884. List of fishes collected at Key West, Florida with
HUTTON, F. W. notes and descriptions. Proc. U.S. Natl. Mus. 7:103-150.
1895. Notes on some New Zealand fishes, with description 1895. The fishes of Sinaloa. Proc. Calif. Acad. Sci., Ser. 2,
of a new species. Trans. N. Z. Inst. 28:314-318. 5:377-514.
HUTTON, F. W. (EDITOR). JORDAN, D. S., AND M. C. DICKERSON.
1904. Index faunae Novae Zealandiae. Philos. Inst. Can- 1908. On a collection of fishes from Fiji, with notes on
terbury, N.Z., 372 p. certain Hawaiian fishes. Proc. U.S. Nat!. Mus. 34:
IRVINE, F. R. 603-617.
1947. The fishes and fisheries of the Gold Coast. Crown JORDAN, D. S., AND B. W. EVERMANN.
Agents for the Colonies, Lond., 352 p. 1896a. A check-list of the fishes and fish-like vertebrates
IVO,C. T. C. of North and Middle America. Rep. U.S. Comm. Fish
1972. Epoca de desova e idade na primeira matura~iio Fish. 1895, 21:207-584.
sexual da cavala, Scomberomorus cavalla (Cuvier), no 1896b. The fishes of North and Middle America. U.S.
Estado do Ceara. Arq. Cienc. Mar 12:27-29. Natl. Mus. Bull. 47:1-1240.
1974. Sabre a fecundidade da cavala, Scomberomorus 1900. The fishes of North and Middle America. U.S.
cavalla (Cuvier), em aguas costeiras do Estado do Ceara Natl. Mus. Bull. 47:3137-3313,392 pIs.
(Brasil). Arq. Cienc. Mar 14:87-89. 1902. American food and game fishes. Doubleday, Page
IWAI, T., AND 1. NAKAMURA. & Co., N.Y., 572 p.
1964a. Branchial skeleton ofthe bluefin tuna, with special JORDAN, D. S., B. W. EVERMANN, AND H. W. CLARK.
reference to the gill rays. Bull. Misaki Mar. BioI. lnst., 1930. Check list of the fishes and fishlike vertebrates of
Kyoto Univ., No.6, p. 21-25. North and Middle America north of the northern bound-

680
COLLE'ITE and RUSSO: SPANISH MACKERELS

ary of Venezuela and Colombia. [U.S.] Bur. Fish., Rep. KAMIYA, T.

U.S. Comm. Fish., 1928, append. 10, 670 p. 1922. Pelagic fish eggs and larvae in the Tateyama Bay.
JORDAN, D. S., AND C. H. GILBERT. Part 2: Pelagic fish eggs and larvae in the Seto Inland
1881a. List of the fishes of the Pacific coast of the United Sea. [In Jpn.] Rep. Fish. Inst. (Suisan Koshujo Shiken
States, with a table showing the distribution of the spe- Hokoku) 18(3):1-39.
cies. Proc. U.S. Nat!. Mus. 3:452-458. 1924. Key to the pelagic fish eggs of Japan. [In Jpn.] J.
1881b. Notes on the fishes of the Pacific coast of the United Fish. Res. (Suisan Kenkyu-Shi) Supp!. 19(6):33-40.
States. Proc. U.S. Nat!. Mus. 4:29-70. KAMOHARA, T.
1882. Synopsis of the fishes of North America. U.S. Nat!. 1967. Fishes of Japan in color. Hoikusha Pub!. Co., Osa-
Mus. Bul!. 16, 1018 p. ka, Jpn., 135 p.
·1883a. List of fishes collected at Mazatlan, Mexico, by KAWAMOTO, N., N. V. TRUONG, AND T. T. TuY-HOA.
Charles H. Gilbert. Bul!. U.S. Fish Comm. 2:105-108. 1972. Illustrations of some freshwater fishes of the Me-
1883b. List of fishes collected at Panama by Charles H. kong delta, Vietnam. Contrib. Fac. Agric., Univ. Cantho
Gilbert. Bul!. U.S. Fish Comm. 2:109-111. 1,49p.
1883c. On certain neglected generic names of La Ce- KENDALL, W. C., AND L. RADCLIFFE.
pede. Proc. U.S. Nat!. Mus. 5:570-576. 1912. The shore fishes. Reports on the scientific results of
JORDAN, D. S., AND C. L. HUBBS. the expedition to the eastern tropical Pacific, in charge of
1925. Record of fishes obtained by David Starr Jordan in Alexander Agassiz, by the U.S. Fish Commission Steamer
Japan, 1922. Mem. Carnegie Mus. 10:93-346. "Albatross;' from October, 1904, to March, 1905, Lieut.
JORDAN, D. S., AND P. L. JOUY. Commander L. M. Garrett, U.S.N., Commanding. XXv.
1881. Check-list of duplicates of fishes from the Pacific -Mem. Mus. Compo Zoo!' 35:77-172.
coast of North America, distributed by the Smithsonian KENT, W. S.
Institution in behalf of the United States National Mu- 1893. The Great Barrier ReefofAustralia; its products and
seum, 1881. Proc. U.S. Nat!. Mus. 4:1-18. potentialities. W. H. Allen and Co., Ltd., Lond., 387 p.
JORDAN, D. S., AND R. E. RICHARDSON. KIM, W. S.
1909. A catalog of the fishes of the Island of Formosa, 1970. Studies on the Spanish mackerel populations (1) age
, or Taiwan, based on the collections of Dr. Hans Sauter. determination. [In Korean, Eng!. abstr.] J. Oceano!.
Mem. Carnegie Mus. 4(4):159-204. Soc. Korea 5(1):37-40.
JORDAN, D. S., AND A. SEALE. KISHINOUYE, K.
1906. The fishes of Samoa. Description of the species found 1915. A study of the mackerels, cybiids, and tunas. [In
in the Archipelago, with a provisional check-list of the Jpn.] Suisan Gakkai Ho 1(1):1-24. [Eng!. trans!., U.S.
fishes of Oceania. Bul!. [U.S.] Bur. Fish. 25:173-455. Fish Wild!. Serv., Spec. Sci. Rep. Fish. 24,14 p.]
1907. Fishes of the islands of Luzon and Panay. Bull. 1923. Contributions to the comparative study of the so-
[U.S.] Bur. Fish. 26:1-48. called scombroid fishes. J. Col!. Agric., Imp. Univ.,
JORDAN, D. S., AND J. O. SNYDER. Tokyo 8:293-475.
1900. A list of fishes collected in Japan by Keinosuke 1924. Observations on the skipjack fishing grounds. [In
Otaki, and by the United States steamer Albatross, with Jpn.] Suisan Gakkai Ho 4(2):87-92. [Eng!. trans!.,
descriptions of fourteen new species. Proc. U.s. Nat!. U.S. Fish Wild!. Serv., Spec. Sci. Rep. Fish. 19, p.12-14.]
Mus. 23:335-380. KITAHARA, T.
1901. A preliminary check list of the fishes of Japan. An- 1897. Scombridae of Japan. [In Jpn., Eng!. summ.] J.
not. Zoo!. Jpn. 3(2-3):31-159. Imp. Fish. Bur., Tokyo, 6, 15 p.
JORDAN, D. S., S. TANAKA, AND J. O. SNYDER. KLAWE, W. L.
1913. A catalogue ofthe fishes ofJapan. J. Col!. Sci., Imp. 1966. Notes on occurrence of young and spawning of
Univ., Tokyo 33(1):1-497. Scomberomorus sierra in the eastern Pacific Ocean.
JORGENSON, S. C., AND G. L. MILLER. Pac. Sci. 20:445-451.
1968. Length relations of some marine fishes from coastal 1977. What is a tuna? Mar. Fish. Rev. 39(11):1-5.
Georgia. U.S. Fish Wild!. Serv., Spec. Sci. Rep. Fish. KLIMA, E. F.
575,16 p. 1959. Aspects of the biology and fishery for Spanish
JOUBERT, C. S. W. mackerel, Scomberomorus maculatus (Mitchi]]), of south-
1981. A survey of shore anglers' catches at selected sites on ern Florida. Fla. State Board Conserv., Tech. Ser. 27,
the Natal Coast, South Africa. S. Afr. Assoc. Mar. Biol. 39p.
Res., Oceanogr. Res. Inst., Invest. Rep. 52, p. 1-13. KLUNZINGER, C. B.
KAIKINI, A. S. 1871. Synopsis der Fische des Rothen Moores. II. Thei!.
1961. The fisheries of Malwan. Indian J. Fish. 7:348- Verh. Zoo!. Bot. Ges. Wien 21:441-668.
368. KNAPP, F. T.
KAILOLA, P. J. 1950. Menhaden utilization in relation to the conservation
1974. Additions to the fish fauna of Papua New Guinea of food and game fishes of the Texas Gulf coast. Trans.
III. Dep. Agric., Stock Fish. (Papua New Guinea), Res. Am. Fish. Soc. 79:137-144.
Bull. 12, p. 54-89. KNER, R.
1975. A catalogue of the fish reference collection at the 1865. Fische. Novara-Exped., Zoo!' Theil 1(2):110-272.
Kanudi Fisheries Research Laboratory, Port Moresby. KONG, I.
Dep. Agric., Stock Fish. (Papua New Guinea), Res. Bul!. 1978. Scomberomorus maculatus (Mitchill), 1815 en Anto-
16,277 p. fagasta, Chile. (Perciformes, Scombridae). Mus. Nac.
KAILOLA, P. J., AND M. A. WILSON. Hist. Nat. Bo!. (Santiago), 22(262):6-9.
1978. The trawl fishes ofthe Gulf of Papua. Dep. Primary KOSTER,F.
Ind., (Papua New Guinea), Res. Bul!. 20, 85 p. 1981. Observaciones de la ictiofauna de las Islas del

681
 FISHERY BULLETIN: VOL. 82, NO.4

Rosario. An. Inst. Invest. Mar. Punta Betln 11:49-57. Hai and Bo Hai. [In Chin., Engl. abstr.] Trans. Chin.
KRISHNAMOORTHI, B. khthyol. Soc. 2:129-137.
1957. Fishery resources of the Rameswaram Island. In- LIU, C., X. ZHANG, AND K. YANG.
dian J. Fish. 4:229-253. ' 1982. Studies on the growth of Spanish mackerel, Scom-
1958. Observations on the spawning season and the fish- beromorus niphoniw; in the Huanghai Sea and Bohai
eries of the spotted seer, Scomberomorus guttatus (Bloch Sea. [In Chin., Eng!. abstr.] Oceano!' Limnol. Sin.
& Schneider). Indian J. Fish. 5:270-281. 13(2):170-178.
KUMARAN,M. LoCKINGTON, W. N.
1964. Observations on the food of juveniles of Scomber- 1879a. On a new genus and species of Scombridae. Proc.
omorus commerSon (Lacep(!de) and S. guttatus (Bloch and Acad. Nat. Sci. Phila. 31:133-136.
Schneider) from Vizhingam, west coast of India. Proc. 1879b. Report upon the food fishes of San Francisco. Rep.
Symp. Scombroid Fishes, Mar. Bio!' Assoc. India, Symp. COl)llll. Fish., State of Calif. 1878-79:17-58.
Ser. I(Part 2):586-590. LoWE, R H. (MCCONNELL).
KURONUMA, K., AND Y. ABE. 1962. The fishes of the British Guiana continental shelf,
1972. Fishes of Kuwait. Kuwait Inst. Sci. Res., Kuwait Atlantic coast of South America, with notes on their
City, 123 p. natural history. J. Linn. Soc. Lond. Zool. 44:669-700.
KUSAKA, T. LoZANO CABO, F.
1974. The urohyal of fishes. Univ. Thkyo Press, Tokyo, 1970. Caracteristiques zoogeographiques de la faune ich-
320p. thyologique des cOtes des iIes Canaries, du Maroc, du
KYUSHIN, K., K. AMAOKA, K. NAKAYA, H. IDA, Y. TANINO, Sahara espagnol et de la Mauritanie avec une etude
AND T. SENTA. speciale des poissons ci>tiers. Rapp. P.-v. Reun., Cons.
1982. Fishes ofthe South China Sea. Jpn. Mar. Fish. Res. Perm. Int. Explor. Mer 159:152-164.
Cent., Thkyo, 333 p. LUBBOCK, R, AND A. EDWARDS.
LACEPEDE, B. G. E. 1981. The fishes of Saint Paul's Rocks. J. Fish BioI. 18:
1800-1803. Histoire naturelle des poissons. Vo!. 2, 1800, 135-157.
632 p.; Vol. 3,1801, 558 p. Plasson, Paris. LYLES,C. H.
LA MONTE,F. 1969. The Spanish mackerel and king mackerel fish-
1945. North American game fishes. Doubleday, Doran eries. U.S. Dep. Commer., NOAA, NMFS, Curro Fish.
and Co., Inc., Garden City, N.Y., 202 p. Stat. 4936, 21 p.
1952. Marine game fishes of the world. Doubleday and MAcGREGOR, R, 1lI, J. J. DINDO, AND J. H. FINUCANE.
Co., Inc., Garden City, N.Y., 190 p. 1981. Changes in serum androgens and estrogens during
LATIFF,M. spawning in bluefish, Pomatomus saltator, and king
1971. A guide to trawl species in Penang waters. Minist. mackerel, Scomberomorus cavalla. Can. J. Zoo!' 59:
Agric. Lands, Malaysia, 150 p. 1749-1754.
LEON, P. E. MACLEAY, W
1973. Ecologia de la ictiofauna del Golfo de Nicoya, Costa 1881. Descriptive catalogue ofthe fishes of Australia. Part
Rica, un estuario tropical. Rev. Bio!. Trop. 21:5-30. ll. Proc. Linn. Soc. N.S.w 5:510-629.
LEE, S., AND H. YANG. 1884a. Notes on a collection of fishes from the Burdekin
1983. Fishes ofthe suborder Scombroidei ofTaiwan. Bull. and Mary Rivers, Queensland. Proc. Linn. Soc. N.SW.
Inst. Zool. Acad. Sin. 22:217-242. 8:199-213.
LEWIS, A. D. 1884b. Supplement to the descriptive catalogue of the
1981. Population genetics, ecology and systematics orInda- fishes of Australia. Proc. Linn. Soc. N.S.W 9:2-64.
Australian scombrid fishes, with particular reference MAGNUSON, J. J.
to skipjack tuna (Katsuwonus pelamis). Ph.D. Thesis, 1973. Comparative study of adaptations for continuous
Australian National Univ., 314 p. swimming and hydrostatic equilibrium of scombroid and
LEWIS, A. D., L. B. CHAPMAN, AND A. SESEWA. xiphoid fishes. Fish. Bull., U.S. 71:337-356.
1983. Biological notes on coastal pelagic fishes in Fiji. MAGNUSON, J. J., AND J. G. HEITZ.
Fish. Div. (MAF) Fiji Tech. Rep. 4, p. 1-68. 1971. Gill raker apparatus and food selectivity among
LEWIS, A. D., B. R SMITH, AND R E. KEARNEY. mackerels, tunas, and dolphins. Fish. Bull., U.S. 69:
1974. Studies on tunas and baitfish in Papua New Guinea 361-370.
waters - II. Dep. Agric. Stock Fish., Res. Bul!. 11, 113 p. MAGO LECCIA, F.
LEWIS, R J., AND R ENDEAN. 1958. The comparative osteology of the scombroid fishes of
1983. Occurrence of a ciguatoxin-like substance in the the genus &omberomorus from Florida. Bull. Mar. Sci.
Spanish mackerel (&omberomorus commersoni). Thxi- Gulf Caribb. 8:299-34L
con 21:19-24. MALPAS, A. H.
LIMA, H. DE H., AND OLIVEIRA, A. M. E. DE. 1926. The marine biological survey of the littoral waters of
1978. Segunda contribui~o ao conhecimento dos nomes Ceylon. Bull. Ceylon Fish. 2:13-165.
vulgares de peixes marinhos do nordeste Brasileiro. Bol. MANOOCH, C. S., 1lI.
CiEmc. Mar 29:1-26. 1979. Recreational and commercial fisheries for king
LINDSEY, C. C. mackerel, Scomberomor.us cavalla, in the South Atlantic
1968. Temperatures of red and white muscle in recently Bight and Gulf of Mexico, U.S.A. In E. L. Nakamura
caught marlin and other large tropical fish. J. Fish. Res. and H. R Bullis, Jr. (editors), Proceedings ofthe Macker-
Board Can. 25:1987-1992. el Colloquium, p. 33-41. Gulf States Mar. Fish. Comm.
LIU,C. No.4.
1981. Age determination of Spanish mackerel, Scomber- MANOOCH, C. S., 1lI, E. L. NAKAMURA, AND A. B. HALL.
omorus niphonius (Cuvier and Valenciennes) in Huang 1978. Annotated bibliography of four Atlantic scombrids:

682
COLLE'ITE and RUSSO: SPANISH MACKERELS

Scomberomorus brasiliensis, S. cavalla, S. maculatus, of Mexico (Pisces: Scombridae). Northeast Gulf Sci. 4
and S. regalis. U.S. Dep. Commer., NOAA Tech. Rep. (1):1-16.
NMFS Circ. 418, 166 p. MCKAY, R. J.
MARCGRAVE, G. 1970. Additions to the fish fauna of Western Australia -
1648. Historiae rerum naturalium Brasiliae libri viii. 5. West. Aust. Fish. Bul!. 9:3-24.
Leyden and Amsterdam, 303 p. MCPHERSON, G. R.
MARCHAL,E. 1981. Preliminary report: Investigations of Spanish mack-
1961. Quelques observations complementaires sur les erel Scomberomorus commerson in Queensland wa-
Scombridae peches par Ie "Pyrrhus." Bull. mst. Fr. Afr. ters. In C. J. Grant and D. G. Walter (editors), Northern
Noire 23A:102-107. pelagic fish seminar, p. 51-58. Aust. Gov. Pub!. Serv.,
MARQUEZ M., R. Canberra.
1973. Informe sinoptico de la sierra, Scomberomorus mac- MEABURN, G. M.
ulatus (Mitchill), del Golfo de Mexico. Inst. Nac. Pesca, 1979. Heavy metal contamination of Spanish mackerel,
INP/SI: i 14, 19 p. Scomberomorus maculatus and king mackerel, S. caval-
MARR, J. C., AND M. B. SCHAEFER. lao In E. L. Nakamura and H. R. Bullis, Jr. (editors),
1949. Definitions of body dimensions used in describing Proceedings of the Mackerel Colloquium, p. 61-66. Gulf
tunas. U.S. Fish Wild!. Serv., Fish. Bul!. 51:241-244. States Mar. Fish. Comm. No.4.
MARSHALL, T. C. MEAD,G.W
1964. Fishes of the Great Barrier Reef and coastal wa- 1951. Postlarval Neothunnus macropterus, Auxis thazard,
ters of Queensland. Angus and Robertson, Sydney, 566 and Euthynnus lineatus from the Pacific coast of Cen-
p. (Also 1965, Livingston Pub!. Co., Narbeth.) tral America. U.S. Fish Wild!. Serv., Fish. Bul!. 52:121-
1966. Tropical fishes of the Great Barrier Reef. Angus 127.
and Robertson, Sydney, 239 p. MEEK, S. E., AND S. F. HILDEBRAND.
MASUDA, H., C. ARAGA, AND T. YOSHINO. 1923. The marine fishes of Panama. Field Mus. Nat.
1975. Coastal fishes of southern Japan. Tokai Univ. Rist., Pub!. 215, Zoo!' Ser. 15, Part 1, 330 p.
Press, Tokyo, 379 p. MEEK, S. E., AND R. G. NEWLAND.
MATHER, F. J., III. 1884. A review of the American species of Scomberomo-
1954. Northerly occurrences of warmwater fishes in the rus. Proc. Acad. Nat. Sci. Phila. 36:232-235.
western Atlantic. Copeia 1954:292-293. MENDOZA N., A.
MATHER, F.J., III, AND C. G. DAY. 1968. Consideraciones sobre la biologia pesquera de la
1954. Observations of pelagic fishes of the tropical Atlan- sierra, Scomberomorus maculatus (Mitchil!), en el Estado
tic. Copeia 1954:179-188. de Veracruz. Bios 1(2):11-22.
MATHER, F. J., III, AND R. H. GIBBS, JR. MENEZES, M. FERREIRA DE.
1957. Distributional records of fishes from waters off New 1969a. Alimenta~iio da cavala, Scomberomorus cavalla
England and the middle Atlantic states. Copeia 1957: (Cuvier), em aguas costeiras do Estado do Ceara. Arq.
242-244. Cienc. Mar 9:15-20.
MATSUMOTO, W M. 1969b. Algumas observa~oes osteologicas e merlsticas so-
1968. Morphology and distribution of larval wahoo Acan- bre a cavala, Scomberomorus cavalla (Cuvier), do nord-
thocybium solandri (Cuvier) in the central Pacific Ocean. este brasileiro. Arq. Cienc. Mar 9:175-178.
U.S. Fish Wild!. Serv., Fish. Bull. 66:299-322. 1970. Alimenta~iio da serra, Scomberomorus maculatus
MAUGE,L. A. (Mitchil!), em aguas costeiras do Estado do Ceara. Arq.
1967. Contribution preliminaire a l'inventaire ichtyolo- Ciimc. Mar 10:171-176.
gique de la region de Tulear. Ann. Univ. Madagascar, 1972. Numero de rastros da serra, Scomberomorus macu-
ser. Sci. Nat., Math. 5:215-246. (Also published in Re- latus (MitchiH), das aguas costeiras do Estado do Ce-
cueil Trav. Stn. Mar. Endoume-Marseille, Fasc. hors sar. ara. Arq. Cienc. Mar 12:86-88.
supp!. no. 7, p. 101-132.) 1976. Aspectos biol6gicos da serra, Scomberomorus macu-
MAURIN, C., F. LOZANO CABO, AND M. BONNET. latus (Mitchill), capturada por currais-de-pesca. Arq.
1970. Inventaire faunistique des principales especes ich- Cienc. Mar 16:45-48.
thyologiques frequentant les cotes nord-ouest africaines. MENON, A. G. K.
Rapp. P.-v. Reun. Cons. Int. Explor. Mer 159:15-21. 1966. On a collection of fish from the Coromandel Coast of
MCALLISTER, D. E. India including Pondichery and Karaikkal areas. Rec.
1968. The evolution of branchiostegals and associated Indian Mus. 59:369-404.
opercular, gular, and hyoid bones, and the classification of MERCERON, M.
teleostome fishes, living and fossi!. Bul!. Nat. Mus. Can. 1970. Note sur les tazars (Scomberomorus spp.) des cotes
221,239 p. cambodgiennes et leur peche. Univ. Rennes Trav. Fac.
MCCULLOCH, A. R. Sci. Ser. Oceanogr. Bio!' 3:69-90.
1922. Check list of the fish and fish-like animals of New MERRETT, N. R., AND C. H. THORP.
South Wales. Part 3. Aust. Zoo!' 2(3):86-130. 1966. A revised key to the scombroid fishes of East Africa
1929. A check-list of the fishes recorded from Australia. with new observations on their biology. Ann. Mag. Nat.
Aust. Mus. Syd. Mem. 5(1):1-144. Rist., Ser. 13, 8:367-384.
MCCULLOCH, A. R., AND G. P. WHITLEY. MILES, D. W, AND E. G. SIMMONS.
1925. A list of the fishes recorded from Queensland wa- 1951. The menhaden fishery. 'lex. Game, Fish, Oyster
ters. Mem. Queens!. Mus. 8(2):125-182. Comm., Bull. 39, 28 p.
McEACHRAN, J. D., J. H. FINUCANE, AND L. S. HALL. MILLER, D. J., AND R. N. LEA.
1980. Distribution, seasonality and abundance of king 1972. Guide to the coastal marine fishes of California.
and Spanish mackerel larvae in the northwestern Gulf Calif. Dep. Fish Game, Fish Bul!. 157, 235 p.

683
 FISHERV BULLETIN: VOL. 82, NO.4

MISRA,K. S. Univ. Fed. Ceara 7:173-175.

1962. An aid to the identification of the common commer- 1968a. Observa~Oes sabre 0 desenvolvimento maturativo
cial fishes of India and Pakistan. Rec. Indian Mus. 57: das ganadas da serra, Scomberomorus maculntus (Mitch-
1-320. ill, 1815). Arq. Estac. BioI. Mar. Univ. Fed. Ceara 8:
MITCHILL, S. L. 25-30.
1815. The fishes of New-York, described and arranged. 1968b. Algumas observa~aes sabre 0 semen da serra,
Trans. Lit. Philos. Soc. N.Y 1:355-492. Scomberomorus maculntus (Mitchill). Arq. Estac. BioI.
Mrro,S. Mar. Univ. Fed. Ceara 8:139-140.
1960. Keys to the pelagic fish eggs and hatched larvae 1968c. Considera~oes sabre 0 semen da cavala, Scomber-
found in the adjacent waters of Japan. [In Jpn., Engi. omorus cavalla (Cuvier). Arq. Estac. BioI. Mar. Univ.
summ.] Bull. Arts, Sci. Fac. Agric. Kyushu Univ. 18(1): Fed. Ceara 8:31-32.
71-94. 1970. On the pyloric caeca in fishes ofthe genus Scomber-
1961. Pelagic fish eggs and larvae from Japanese waters- omorus Lacepede. Arq. Cienc. Mar 10:181-184.
II. Lamprida, Zeida, Mugilina, Scombrina, Carangina MUNRO, I. S. R.
and Stromateina. [In Jpn., Engi. summ.] Bull. Arts, 1942. The eggs and early larvae of the Australian barred
Sci. Fac. Agric. Kyushu Univ. 18:451-466. Spanish mackerel, Scomberomorus commersoni (Lace-
1966. Nihon kaiyo plankton zukan - Dai 7 Kan-Gyoran - pede) with preliminary notes on the spawning of that
Chigyo (Japanese marine plankton picture book - Vol. species. Proc. R. Soc. Queens!. 54(4):33-48.
7 - Fish eggs -larvae). [In Jpn.] Soyosha, 'Ibkyo, 75 p. 1943. Revision of Australian species of Scomberomorus.
1967. Some ecological notes on the planktonic fish larvae. Mem. Queensi. Mus. 12:65-95.
[In Jpn., Engi. abstr.] Inf. Bull. Plankton Soc. Jpn. 14: 1955. The marine and fresh water fishes of Ceylon. Dep.
33-49. [Nihon Purankuton Kenkyu Renrakukaiho.] External Afr., Canberra, 351 p.
MIYAKE, M., AND S. HAvASI. 1958a. Families Polynemidae to Scomberomoridae.
1972. Field manual for statistics and sampling of Atlantic Handbook of Australian fishes, No. 28:113-116. Aust.
tunas and tuna-like fishes. Int. Comm. Conserv. Ati. Fish. Newsi. 17(10):17-20.
Tunas, Madrid, Spain, 95 p. 1958b. The fishes of the New Guinea region. Papua New
MOE, M. A., JR. Guinea Agric. J. 10(4):97-369.
1963. A survey of offshore fishing in Florida. Fla. State 1964. Additions to the fish fauna of New Guinea. Papua
Board Conserv. Mar. Res. Lab., Prof. Pap. Ser. 4, 115 p. New Guinea Agric. J. 16(4):141-186.
1972. Movement and migration of South Florida fishes. 1967. The fishes of New Guinea. Dep. Agric., Stock,
Fla. Dep. Nat. Resour. Mar. Res. Lab., Tech. Ser. 69, 25 p. Fish., Port Moresby, New Guinea, 650 p.
MONOD,T. NAGABHUSHANAM, A. K., AND G. CHANDRASEKHARA RAo.
1968. Le complexe urophore des poissons teIeosteens. 1972. An ecological survey of the marine fauna of Minicoy
Mem. Inst. Fond. Afr. Noire 81, 705 p. Atoll (Laccadive Archipelago, Arabian Sea). Mitteil.
MORGAN, E. C., AND W. K. KING. Zooi. Mus. Beri. 48(2):265-324.
1983. Tooth replacement in king mackerel, Scomberomo- NAKAMURA, I.
rus cavalln (Pisces: Scombridae). Southwest. Nat. 28(3): 1965. Relationships of fishes referable to the subfamily
261-269. Thunninae on the basis of the axial skeleton. Bull.
MORI, T. Misaki Mar. BioI. Inst., Kyoto Univ. 8:7-38.
1928. A catalogue of the fishes of Korea. J. Pan-Pac. Res. NAKAMURA, I., AND K. MORI.
Inst. 3(3):3-8. 1966. Morphological study of the slender tuna, Allothun-
1952. Check list of the fishes of Korea. Mem. Hyogo nus {allni Serventy obtained from the Tasman Sea. Rep.
Univ. Agric. 1(3):1-228. Nankai Reg. Fish. Res. Lab. 23:67-83.
1956. Fishes of San-in District including Oki Islands and NAKAMURA, I., AND R. NAKAMURA.
its adjacent waters (Southern Japan Sea). Mem. Hyogo 1982. New records of two species of Scomberomorus from
Univ. Agric. 2(3):1-62. Japan. Jpn. J. Ichthyoi. 28:445-449.
MORICE,J. NALBANT, T. T.
1953. Essai systematique sur les familles des Cybiidae, 1970. Noi contributii la studiul ihtiofaunei din Oceanul
Thunnidae et Katsuwonidae, poissons scombroides. Rev. Pacific de Nord. [In Romanian, Engi. summ.] Bui.
Trav. Inst. Sci. Tech. Peches Marit. 18:35-63. Cercet. Piscic. 29(1-2):57-64.
MORROW, J. E. NAUGHTON, S. P., AND C. H. SALOMAN.
1954. Fishes from East Africa, with new records and 1981. Stomach contents of juveniles of king mackerel
descriptions of two new species. Ann. Mag. Nat. Hist., (Scomberomorus cavalla) and Spanish mackerel (S. mac-
Ser. 12, 7:797-820. ulatus). Northeast Gulf Sci. 5(1):71-74.
MOTA ALVES, M. I. NAVARRO, F. DE P.
1969. SObre 0 trato digestivo da serra, Scomberomorus 1943. La pesca de arrastre en los fondos del Cabo Blanco
maculatus (Mitchill). Arq. Cienc. Mar 9:167-171. y del Banco Arguin (Africa Sahariana). Trab. Inst. Esp.
MOTA ALVES, M. I., AND G. DE SOUZA TOME. Oceanogr. 18, 225 p.
1967a. Anatomia e histologia do tubo digestivo de Scom- NICHOLS,J. T.
beromorus cavalla (Cuvier, 1829). Arq. Estac. BioI. Mar. 1929. The fishes of Porto Rico and the Virgin Islands,
Univ. Fed. Ceara 6:103-108. Branchiostomidae to Sciaenidae. Scientific Survey of
1967b. Alguns aspectos do desenvolvimento maturativo Porto Rico and the Virgin Islands. N.Y. Acad. Sci.
das ganadas das cavala, Scomberomorus cavalla (Cuvier, 10(2):169-295.
1829). Arq. Estac. BioI. Mar. Univ. Fed. Ceara 7:1-9. NICHOLS, J. T., AND C. M. BREDER, JR.
1967c. Notas sabre os anexos digestivos da cavala, Scom- 1927. The marine fishes of New York and southern New
beromorus cavalln (Cuvier 1829). Arq. Estac. BioI. Mar. England. Zoologica (N.Y) 9:1-192.

684
COLLE'ITE and RUSSO: SPANISH MACKERELS

NICHOLS, J. T., AND R C. MURPHY. PEW, P.

1944. A collection of fishes from the Panama Bight, Pacific 1954. Food and game fishes of the Texas coast. Tex.
Ocean. Bull. Am. Mus. Nat. Hist. 83:221-260. Game Fish Comm., Mar. Lab. Bull. 33, Ser. IV; 68 p.
NOMURA,H. PHILLIPPS, W. J.
1967. Dados biol6gicos sobre a serra, Scomberomorus mac- 1927. A check list of the fishes of New Zealand. J. Pan-
ulatus (Mitchill), das aguas cearenses. Arq. Estac. BioI. Pac. Res. Inst. 2(1):9-15.
Mar. Univ. Fed. Ceara 7:29-39. PHILLIPS, J. B.
NOMURA,H., AND R S. DA COSTA. 1932. Monterey Spanish mackerel landed in Monterey.
1966. SObre 0 comprimento e 0 peso da cavala e da serra das Calif. Fish Game 18:99.
aguas cearenses. Arq. Estac. BioI. Mar. Univ. Fed. PHILLIPS, P. C.
Ceara 6:11-13. 1981. Annotated checklist of fishes at Jiquilisco Bay, EI
1968. Length-weight relationship of two species of Scom- Salvador.' Rev. BioI. Trop. 29(1):45-58.
bridae fishes from Northeastern Brazil. Arq. Estac. PLAYFAIR, R L., AND A. C. L. GUNTHER.
BioI. Mar. Univ. Fed. Ceara 8:95-99. 1866. The fishes of Zanzibar. John van Voorst, Lond., 153
NOMURA, H., AND M. S. R DE SOUSA. p. (Reprint, 1971, N. K. Gregg, Kentfield, Calif.)
1967. Biological notes on king mackerel, Scomberomorus POEY,F.
cavalla (Cuvier), from northeastern Brazil. Arq. Estac. 1865. Revista de los tipos Cuvierianos y Valencienniannos
BioI. Mar. Univ. Fed. Ceara 7:79-85. correspondientes a los peces de la isla de Cuba. Repert.
Fis.-Nat. Isla Cuba, 1:308-338.
NORMAN, J. R, AND F. C. FRASER.
1868. Synopsis piscium Cubensium. Repert. Fis.-Nat.
1949. Field book of giant fishes. G. P. Putnam's Sons,
Isla Cuba, 2:279-484.
N.Y.,376p.
1875. Enumeratio piscium Cubensium. Part 1. Anal. Soc.
OGILBY, J. D.
Espaii. Hist. Nat. 4 parts, 224 p.
1887. Catalogue of the fishes of New South Wales with
1878. Notes on the American species of the genus Cybium.
their principal synonyms. Fisheries of the Colony. Rep.
Proc. U.S. Natl. Mus. 1:3-5.
Comm. Fish. N.sW. 1886, Append. 6, p. 1-67.
POLL,M.
OHE, E, H. NISHIMOTO, Y. OKUMURA, AND Y. AzUMA.
1959. Expedition oceanographique beIge dans les eaux
1981. Fossil "Acanthocybium" sp. (Scombridae, Pisces) of
cotieres Africaines de I'Atlantique Sud (1948-1949). Vol.
the Miocene Mizunami group, Central Japan. [In Jpn.,
IV. Fasicule 3B. Poissons V- Teloosteens acanthoprery-
sect. in Engl.] Bull. Mizunami Fossil Mus. 8:25-46.
giens (deuxieme partie). Bruxelles, 417 p.
OKADA, Y.
POSTEL, A. E.
1955. Fishes of Japan; illustrations and descriptions of
1955. Considerations biometriques sur la denture des
fishes off Japan. Maruzen Co., Ltd., Tokyo, 434 p.
Cybiides (dents maxillaires). Bull. Stn. Oceanogr. Sal-
OMMANNEY, F. D.
ammbO 51:57-67.
1953. The pelagic fishes and a note on tow nettings:
POSTEL, E.
Distribution of macroplankton, fish eggs and young
1954. Contribution a l'etude des thonides de I'Atlantique
fish. In Report on the Mauritius-Seychelles fisheries
tropical. J. Cons. 19:356-362.
survey 1948-49. Part II. Vol. 1, p. 58-104. G.B. Colon.
1955a. Contribution a l'etude de la biologie de quelques
Off. Fish. Publ.
Scombridae de I'Atlantique tropico-oriental. Stn. Ocean-
ORSI,J. J.
ogr. SalammbO, 10, 167 p.
1974. A check list of the marine and freshwater fishes of
1955b. Contribution a l'etude des thonides de I'Atlantique
Vietnam. Publ. Seto Mar. BioI. Lab. 21:153-177.
tropical (deuxieme note). Rapp. P.-v. Reun. Cons. Perm.
OSBURN, R C., AND J. T. NICHOLS. Int. Explor. Mer 137:31-32.
1916. Shore fishes collected by the 'Albatross' expedition in 1958. Contributions a l'etude de la biologie de quelques
Lower California with descriptions of new species. Bull. Scombridae de I'Atlantique tropico-oriental. Bull. Soc.
Am. Mus. Nat. Hist. 35:139-181. Sci. Bretagne 31:107-111.
OSORIO, B. 1959. Liste commentee des poissons signales dans I'Atlan-
1898. Da distribui~ii.o geographico dos peixes e crustaceos tique tropico-oriental nord, du Cap Spartel au Cap Roxo,
colhidos nas possessoes Portuguezas d'Africa Occidental e suivie d'un bref aper~u sur leur repartition bathymet-
existentes no Museu Nacional de Lisboa. J. Sci. Math. rique et geographique. Bull. Soc. Sci. Bretagne 34:
Phys., Nat. Acad. Real Sci., Seg. Ser. 5(19):185-202. 129-170.
PAIVA, M. P., AND R S. DA COSTA. 1960. Liste commentee des poissons signales dans I'Atlan-
1966. Considera~oes sobre a produ~ii.o de pescado marinho tique tropico-oriental nord, du Cap Spartel au Cap Roxo,
salgado no Estado do Ceara. Bol. Estac. BioI. Mar. Univ. suivie d'un bref aper~u sur leur repartition bathymet-
Fed. Ceara, No. 15, 11 p. rique et geographique. Bull. Soc. Sci. Bretagne 34:
PATHANSALI, D. 241-281.
1968. Note on the scombroid fishery in Malaya. Proc. 1973. Scomberomoridae. CLOFNAM-Check-list of the
Symp. Scombroid Fishes, Mar. BioI. Assoc. India, Symp. fishes of the north-eastern Atlantic and of the Mediter-
Ser. l(Part 3):1001-1005. ranean. UNESCO 1:473-475.
PELLEGRIN, J. POTTHOFF, T.
1908. Mission des pecheries de la cote occidentale d'Af- 1975. Development and structure of the caudal com-
rique. Poissons (2" Memoire). Actes Soc. Linn. Bordeaux plex, the vertebral column, and the pterygiophores in
62:71-102. the blackfin tuna (Thunnus atlanticus, Pisces, Scom-
PENRITH, M. J. bridae). Bull. Mar. Sci. 25:205-231.
1978. An annotated check-list of the inshore fishes of POWELL,D.
southern Angola. Cimbebasia, ser. A, 4:179-190. 1975. Age, growth, and reproduction in Florida stocks

685
 FISHERY BULLETIN: VOL. 82, NO.4

of Spanish mackerel, Scomberomorus maculatus. Fla. 679p.

Dep. Mar. Res. Publ. 5, 21 p. RICHARDS, W. J., AND W. L. KLAWE.
PRADO, J. 1972. Indexed bibliography of the eggs and young of tu-
1970. Notes sur Scomberomorus commerson Lacepede nas and other scombrids (Pisces, Scombridae) 1880-1970.
1800 de la cote nord-ouest de Madagascar. Univ. Rennes U.S. Dep. Commer., NOAA Tech. Rep. NMFS SSRF-652,
Trav. Fac. Sci. ser. Oceanogr. BioI. 3:91-116. 107p.
PRISTAS, P. J., AND L. TRENT. RICHARDSON, J.
1978. Seasonal abundance, size, and sex ratio of fishes 1846. Report on the ichthyology of the seas of China and
caught with gill nets in St. Andrew Bay, Florida. Bull. Japan. Rep. Br. Assoc. Adv. Sci., 15th Meet., p. 187-320.
Mar. Sci. 28:581-589. RICHARDSON, S. L., AND J. D. MCEACHRAN.
QUIROGA, D., AND A. ORBES ARMAS. 1981. Identification of small « 3 mm) larvae of king and
1964. Apuntes e informaciones sobre las pesquerias en la Spanish mackerel, Scomberomorus cavalla and S. macu-
Provincia de Esmeraldas. Inst. Nac. Pesca Equador, Bol. latus. Northeast Gulf Sci. 5(1):75-79.
Inf. 1(6):1-26. RICKER, K. E.
QURAISHI, M. R. 1959. Mexican shore and pelagic fishes collected from
1945. A comparative study of the morphology, histology Acapulco to Cape San Lucas during the 1957 cruise of the
and probable functions of the pyloric caeca in Indian "Marijean:' Univ. B.C. Inst. Fish., Mus. Contrib. 3:1-18.
fishes, together with a discussion on their homology. RIVAS, L. R.
Proc. Indian Acad. Sci. 21B:1-37. 1951. A preliminary review of the western North Atlantic
RADOVICH, J. fishes of the family Scombridae. Bull. Mar. Sci. Gulf
1961. Relationships of some marine organisms of the Caribb. 1:209-230.
Northeast Pacific to water temperatures, particularly ROCHEBRUNE, A. T. DE.
during 1957 through 1959. Calif. Dep. Fish Game, Fish 1882. Faune de la Senegambie. Poissons. Actes Soc.
Bull. 112, 62 p. Linn. Bordeaux 36, Ser. 4, 6:23-188.
RAFINESQUE, C. S. ROEDEL, P. M.
1815. Analyse de la nature ou tableau de l'univers et des 1939. Another record of the Monterey Spanish macker-
corps organises. Palermo, 224 p. el. Calif. Fish Game 25(4):343.
RAINER, S. F., AND I. S. R. MUNRO. 1951. Noteworthy southern California records of four spe-
1982. Demersal fish and cephalopod communities of an cies of marine fishes. Calif. Fish Game 37:509-510.
unexploited coastal environment in northern Austra- 1953. Common ocean fishes ofthe California coast. Calif.
lia. Aust. J. Mar. Freshw. Res. 33:1039-1055. Dep. Fish Game, Fish Bull. 91, 184 p.
RAJAN, S., S. PATNAINK, AND N. C. BASU.. ROHDE,K.
1969. New records of fishes from the Chilka Lake. J. 1976. Monogenean gill parasites of Scomberomorus com-
Zool. Soc. India 20(1/2):80-93. mersoni Lacepede and other mackerel on the Australian
RANDALL, J. E. east coast. Z. Parasitenk. 51:49-69.
1967. Food habits of reef fishes of the West Indies. Stud. RONQUILLO, I. A.
Trop. Oceanogr. (Miami) 5:665-847. 1974. A review of the roundscad fishery in the Philip-
1968. Caribbean reef fishes. T. F. H. Publ. Inc., Jersey pines. Indo-Pac. Fish. Counc., Proc. 15th Sess. Sect. III:
City, N.J., 318 p. 351-375.
RANDALL, J. E., G. R. ALLEN, AND W. F. SMITH-VANIZ. ROUGHLEY, T. C.
1978. Illustrated identification guide to commercial fishes. 1951. Fish and fisheries of Australia. Angus and Robert-
FAO Reg. Fish. Dev. Proj., FI:DP/RAB/71/278/3, 221 p. son, Sydney, 343 p.
RAo,K. S. RoY, J. C., AND S. ROY.
1964. Observations on the food and feeding habits of 1974. Observations on the pelagic and semi-pelagic fishery
Scomberomorus guttatus (Bloch & Schneider) and juve- of the Balashore Coast, India. Indo-Pac. Fish. Counc.,
niles of S. lineolatus (Cuvier & Valenciennes) and S. Proc. 15th Sess. Sect. 111:40-55.
commerson (Lacepede) from the Waltair coast. Proc. RUSSELL, P.
Symp. Scombroid Fishes, Mar. BioI. Assoc. India, Symp. 1803. Descriptions and figures of two hundred fishes,
Ser. 1(Part 2):591-598. collected at Vizagapatam on the coast of Coromandel.
1976. Biometric comparison of the spotted seer, Scomber- Vol. 2, 85 p. W. Bulmer and Co., Lond.
omorus guttatus (Bloch and Schneider) from five localities RYDER,J.A.
along the Indian coast. Matsya 1:63-78. 1882. Development of the Spanish mackerel (Cybium'
RAO, K. S., AND P. N. GANAPATI. maculatum). Bdl. U.S. Fish. Comm. 1:135-173.
1977. Description of the post-larvae and juveniles of Scom- SACCHI, J., A. LAGIN, V CHAUDEMAR, AND C. LANGLAIS.
beromorus lineolatus (Cuvier & Valenciennes, 1831) from 1981. La pilche des especes pelagiques aux Antilles Fran-
Indian waters. J. Nat. Hist. 11:101-111. ~aises - Etat actuel et persPectives de developpement.
REEVES, C. D. Sci. Poche, Bull. Inf. Doc., Inst. Sci. Tech. Peches Mar.
1927. A catalogue of the fishes of northeastern China and 312:1-15.
Korea. J. Pan-Pac. Res. Inst. 2(3):3-16. SALOMAN, C. H., AND S. P. NAUGHTON.
RENDAHL,H. 1983a. Food of king mackerel, Scomberomorus cavalla,
1921. Results of Dr. E. Mjobergs Swedish Scientific Expe- from the southeastern United States including the Gulf
ditions to Australia 1910-13. XXVIII. Fische. Kungl. of Mexico. U.S. Dep. Commer., NOAA Tech. Memo.
Svenska Vet. Akad. Handl. 61(9):1-24. NMFS-SEFC-126, 25 p.
RIBEIRO, A. DE M. 1983b. Food of Spanish mackerel, Scomberomorus macu-
1915. Fauna Brasiliense. Peixes. V Eleutherobranchios, latus, from the Gulf of Mexico and southeastern seaboard
Aspirophoros (Physoclistil. Arch. Mus. Nac. Rio de J., of the United States. U.S. Dep. Commer., NOAA Tech.

686
COLLEITE and RUSSO: SPANISH MACKERELS

Memo. NMF8-SEFC-128, 22 p. 1968. Tuna fishery of the Tinnevelly Coast, Gulf of Man-
SANCHES, J. G. nar. Proc. Symp. Scombroid Fishes, Mar. BioI. Assoc.
1966. Peixes de Angola (teloosteos). Notas Mimeogr. India, Symp. Ser. 1(Part 3):1083-1118.
Centro BioI. Piscat., Lisboa, No. 46, 227 p. SIVASUBRAMANIAM, K., AND I. A. MOHAMED.
SANCHEZ T., J., AND R. LAM C. 1982. Common fishes of Qatar. Sci. Atlas Qatar 1:1-176.
1970. Algunas caracteristicas fisicas y quimicas de las SKOW, L. C., AND M. E. CHIITENDEN, JR.
principales especies para consumo humano y sus rendimi- 1981. Differences in hemoglobin phenotypes among Span-
entos en productos pesqueros, en el Peru. Inst. Mar ish mackerel, Scomberomorus maculatus. Northeast
Peru, Callao, Inf. 33, 92 p. Gulf Sci. 5(1):67-70.
SANZ ECHEVERRIA, J. SMILEY, C. W.
1950. Notas sobre otolitos de peces proeedentes de las 1881. The Spanish mackerel, and its artificial propaga-
costas del Sahara. Segunda parte. Bol. Inst. Espaii. tion. Proc. Am. Assoc. Adv. Sci. 29(1880):575-583.
Oceanogr. 27, 14 p. SMiTH, C. L., AND R. M. BAILEY.
SCACCINI,A. 1962. The subocular shelf of fishes. J. Morphol.110:1-17.
1941. Primo elenco di pesci raccolti in Atlantico neUe SMITH,H.M.
acque della Mauritania, del Sahara spagnolo e delle 1907. The fishes of North Carolina. N.C. Geol. Econ.
Canarie. Thalassia 4(10):1-49. Survey, Raleigh 2, 453 p.
SCHROEDER, W. C. SMITH, J. L. B.
1924. Fisheries of Key West and the clam industry of 1935. New and little known fishes from South Africa.
Southern Florida. Rep. U.S. Comm. Fish., 1923, App. Rec. Albany Mus. 5:169-235.
12:1-74. (Doc. 962.) 1949[1953, 1961, 4th ed.]. The sea fishes of Southern
SCOIT,J.S. Mrica. Central News Agency, Ltd., South Africa, 550 p.
1959. An introduction to the sea fishes of Malaya. Min- 1956. The fishes of Aldabra. Part V. Ann. Mag. Nat.
istry of Agriculture, Kuala Lumpur, 180 p. Hist., Ser.12, 9(106):721-729.
SEALE, A. 1964. Scombroid fishes of South Africa. Proc. Symp.
1940. Report on the fishes from Allan Hancock expeditions Scombroid Fishes, Mar. BioI. Assoc. India, Symp. Ser.
in the California Academy of Sciences. Allan Hancock l(Part 1):165-183.
Pac. Exped. 9:1-46. SMITH, J. L. B., AND M. M. SMITH.
SERET, B., AND P. OPIC. 1963. The fishes of Seychelles. Rhodes Univ., Dep. leh-
1981. Poissons de mer de l'ouest Africain tropical. thyol., Grahamstown, S.A., 215 p.
O.R.S.T.O.M., Collect. Initiat.-Doc. Tech. 49, 416 p. 1966. Fishes of the Tsitsikama Coastal National Park.
SERVENTY, D. L. Natl. Parks Board, S. Afr., 161 p.
1950. Tuna survey ofnorth Australia. Aust. Fish. Newsl. SNODGRASS, R. E., AND E. HELLER.
9:18-20. 1905. Papers from the Hopkins-Stanford Galapagos Expe-
SHA, X.-S., G.-F. HE, AND H.-W. CHANG. dition, 1898-1899. XVII. Shore fishes ofthe Revillagigedo,
1966. A description of the morphological characters of the Clipperton, Cocos and Galapagos Islands. Proc. Wash.
eggs and larvae of the blue spotted mackerel, Scom- Acad. Sci. 6:333-427.
beromorus niphonius (Cuvier and Valenciennes). [In SOLDATOV, V. K.
Chin., Engl. summ.] Oceanologia Limnol. Sin. 8(1):1-12. 1929. A check list of fishes recorded from Russian Pacific
SHARP, G. D. waters. J. Pan-Pac. Res. Inst. 4(1):3-7.
1973. An electrophoretic study of hemoglobins of some SOLDATOV, V. K., AND G. J. LINDBERG.
scomhroid fishes and related forms. Compo Biochem. 1930. A review ofthe fishes of the seas of the Far East.
Physiol. 44B:381-388. [In Russ.] Bull. Pac. Sci. Fish. Inst. 5, 576 p.
SHAW,G. SPRINGER, V. G., AND J. PiRSON.
1803. General zoology or systematic natural history. Pis- 1958. Fluctuations in the relative abundance of sport
ces 4(Part 2):187-632. Kearsley, Lond. fishes as indicated by the catch at Port Aransas, Texas.
SHENOY, A. v., AND M. A. JAMES. Publ.lnst. Mar. Sci., Univ. Tex. 5:169-185.
1974. Spoilage of spotted seer (Scomberomorus guttatus) STARKS, E. C.
during ice storage. Fish. Technol. 11:67-72. 1910. The osteology and mutual relationships of the fishes
SHIINO, S. M. belonging to'the family Scombridae. J. Morphol. 21:
1972. List of English names of Japanese fishes with prop- 77-99.
osition of new names. [In Jpn. and Engl.] Sci. Rep. 1918. The mackerel and mackerel-like fishes of Califor-
Shima Marineland, No.1, 210 p. nia. Calif. Fish Game 4:118-129.
1976. List of common names of fishes of the world, those STASSANO, E.
prevailing among English-speaking nations. Sci. Rep. 1890. La pesca sulle spiagge atlantiche del Sahara. An.
Shima Marineland, No.4, 252 p. Agr. Roma 1890:7-59.
SILAS, E. G. STEAD, D. G.
1963. Synopsis of biological data on double-lined mackerel 1906. Fishes ofAustralia: A popular and systematic guide
Grammatorcynus bicarinatus (Quoy and Gaimard) (Indo- to the study ofwealth within om: waters. Wm. Brooks &
Pacific). FAO Fish. Rep. 6:811-833. Co., Sydney, 278 p.
1964. Cybium croockewitii Bleeker (1850) and C. kore- 1908. The edible fishes of New South Wales: Their present
anum Kishinouye (1915) considered synonyms of Scom- importance and their potentialities. Gov. Printer, Syd-
beromorus guttatus (Bloch and Schneider) with a rede- ney, 123 p.
scription and annotated bibliography of S. guttatus. STEINDACHNER, F., AND L. OODERLEIN.
Proc. Symp. Scombroid Fishes, Mar. BioI. Assoc. India, 1884. Beitriige zur Kenntniss der Fische Japan's [III].
Symp. Ser. l<Part 1):309-342. Denkschr. Akad. WISS. Wien 49:171-212.

687
 FISHERY BULLETIN: YOLo 82, NO.4

STORER, D. H. (1884, unpublished). Bull. Fac. Fish., Hokkaido Univ.

1855. A history ofthe fishes ofMassachusetts. Mem. Am. 18:137-182.
Acad. Arts Sci. 5:122-168. ToMINAGA, S.
STuRM, M. G. DE L. 1964. Anatomical sketches of 500 fishes. Kadokawa
1978. Aspects of the biology of Scomberomorus maculatus Shoten, Tokyo, Japan, 3 vols.
(Mitchill) in Trinidad. J. Fish BioI. 13:155-172. TONGYAI, M. L. P.
SUIGIURA, H. 1966a. Background notes for Spanish mackerel, (Scom-
1970. Fishes. In T. Tuyama and S. Asami (compilers), beromorus spp.) from Thai waters, studies, 1954-1965.
Ogasawara no shizen (The nature in the Bonin Islands); Thai Mar. Fish. Lab., Scomberomorus Rep. Pap. 1, 25
Vol. 1, p. 197-210. p.
SUMNER, F. B., R. C. OSBURN, AND L. J. COLE. 1966b. Spanish mackerels from M. Y. "DhanaIjata" (trawl)
1913. Section Ill. A catalogue ofthe marine fauna. In A during January, February, and May, 1966. Thai Mar.
biological survey of the waters ofWoods Hole and vicinity; Fish. Lab., Scomberomorus Rep. Pap. 2, 17 p.
Vol. 33, p. 545-794. Bull. [U.S.] Bur. Fish. 31, part 2. 1970. Plah in-see, Scomberomorus spp., of Thailand,
SUTHERLAND, D. F., AND W. A FABLE, JR. 1967. In J. C. Marr (editor), The Kuroshio, a symposium
1980. Results ofa king mackerel (Scomberomorus cavalla) on the Japan Current, p. 557-564. East-West Center
and Atlantic Spanish mackerel (Scomberomorus macu- Press, Honolulu.
latus) migration study: 1975-79. U.S. Dep. Commer., 1971a. A taxonomic study of the pla in-si (Pisces: Scom-
NOAA Tech. Memo. NMFS-SEFC-12, 23 p. broidei) of Thailand. Div. Mar. Fish. Tech., Dep. Fish.,
SUYEHIRO, Y. Bangkok, Thailand, 36 p.
1942. A study ofthe digestive system and feeding habits of 1971b. An economic evaluation of the pIa in-si (Pisces:
fish. Jpn. J. Zool. 10:1-303. Scombroidei) fishery. Bangkok, Thailand, 17 p.
TALBOT, F. H. ToRTONESE,E.
1965. A description of the coral structure of Tutia Reef 1949. Identification di due Sgombroide (pesci) accidentali
(Tanganyika Territory, East Africa), and its fish fauna. nel Mediterraneo. Boll. Zool. Torino 16(1-3):61-66.
Proc. Zool. Soc. Lond.145:431-470. 1956. Brevi considerazioni sui pesci Mediterranei del Sot-
TANAKA,S. tordine Sgombroidi. Boll. Inst. Mus. Zool. Univ. Torino
1927. Figures and descriptions of the fishes of Japan 5(3):3-11.
including Riukiu Islands, Bonin Islands, Formosa, Kurile 1975. Osteichthyes (Pesci ossei). Parte seconda. Fauna
Islands, Korea, and Southern Sakhalin. Kazama Shobo, Italia 11:1-636, Edizioni Calderini, Bologna.
Tokyo 36:677-692. TRENT, L., AND E. A. ANTHONY.
TAYLOR, H. F., AND STAFF. 1979. Commercial and recreational fisheries for Spanish
1951. Survey of marine fisheries of North Carolina. Univ. mackerel, Scomberomorus maculatus. In E. L. Naka-
North Carolina Press, Chapel Hill, 555 p. mura and H. R. Bullis, Jr. (editors), Proceedings of the
TAYWR, w.R. Mackerel Colloquium, p. 17-32. Gulf States Mar. Fish.
1964. Fishes ofArnhem Land. Rec. Am.-Aust. Sci. Exped. Comm.No.4.
Arnhem Land 4:45-307. TRENT, L., R. O. WILLIAMS, R. G. TAYLOR, C. H. SAWMAN, AND
TEMMINCK, C. J., AND H. SCHLEGEL. C. S. MANOOCH 1lI.
1844. Pisces. In P. F. von Siebold, Fauna Japonica ... 1981. Size and sex ratio of king mackerel, Scomberomorus
1823-30. Lugduni Batavorum, Part 3, p. 73-112. cavalla, in the southeastern United States. U.S. Dep.
THAM,A.K. Commer., NOAA Tech. Memo. NMFS-SEFC 62, 59 p.
1950. The food and feeding relationships of the fishes TRETIAKOY, D. K.
of Singapore Straits. Colon. Off., Lond., Fish. Publ. 1939. Olfactory organ in Sarda sarda C.Y. Dokl. Akad.
1(1):1-35. Nauk SSSR 22(4):202-205.
1953. A preliminary study of the physical, chemical and UCHIDA, R. N.
biological characteristics of Singapore Straits. Colon. 1978. The fish resources of the western central Pacific
Off., Lond., Fish. Publ. 1(4):1-65. islands. FAO Fish. Circ. 712, 53 p.
1974. Stolephorus resources in the South China Sea. UENO,T.
Indo-Pac. Fish. Counc. Proc. 15 Sess., Sect. lll:182-191. 1971. List of the marine fishes from the waters ofHokkaido
THIEMMEDH, J. and its adjacent regions. [In Jpn., Engl. abstr.] Sci.
1966. Fishes of Thailand: their English, scientific, and Rep. Hokkaido Fish. Exp. Stn.13:61-102.
Thai names. [Bangkok] Kasetsart Univ. Fish. Res. ULREY,A.B.
Bull. 4:1-212. 1929. A check-list of the fishes of southern California and
THOMSON, D. A, L. T. FINDLEY, AND A. N. KERSTITCH. Lower California. J. Pan-Pac. Res. Inst. 4(4):2-11.
1979. Reef fishes ofthe Sea ofCortez. John Wiley & Sons, UMALI,A.F.
N.Y.,302p. 1936. Edible fishes of Manila. Philipp. Dep. Agric.
THOMSON, D. A, AND N. McKIBBIN. Comm., Pop. Bull. 6, 192 p.
1976. Gulf of California fishwatcher's guide. Golden 1938. The fishery industries of Ragay Gulf. Philipp. J.
Puffer Press, Tucson, Ariz., 75 p. Sci. 65:175-200.
TIRANT,G. 1950. Key to the families of common commercial fishes
1885. Notes sur les poissons de la Basse-Cochinchine et du in the Philippines. U.S. Fish Wildl. Serv., Res. Rep. 21,
Cambodge. Excur. Reconnais. Cochinchine Fr. 9:413- 47 p.
438 and 10:91-198. (Reprinted 1929, SerVo Oceanogr. UYENO, T., AND S. FUJII.
Pech Indochine, Stn. Mar. Cauda, Note 6, p. 43-163.) 1975. A fish fossil ofthe family Scombridae from a Miocene
TOKIDA, J., AND K. KOBAYASHI. bed in Toyama Prefecture, Japan. Bull. Nat. Sci. Mus.
1967. "Catalogue of Japanese Fishes" by Kanzo Uchimura Tokyo, Ser. C(Geol.) 1(1):11-16.

688
COLLETTE and RUSSO: SPANISH MACKERELS

VAN DER ELST, R. Zealand region. Aust. Zool. 15(1):1-102.

1976. Game fish of the east coast of southern Africa. I. The WILLIAMS, F.
biology of the elf, Pomatomus saltattix (Linnaeus), in the 1956. Preliminary survey of the pelagic fishes of East
coastal waters of Natal. S. Afr. Assoc. Mar. BioI. Res., Africa. Colon. Off. Fish. Publ. 8, 68 p.
Oceanogr. Res. Inst. Invest. Rep. 44, 59 p. 1960. On Scomberomorus lineolatus (C.v.) 1831, from Brit-
1981. A guide to the common sea fishes of southern Af- ish East African waters (Pisces, Scombridae). Ann.
rica. C. Struik, Capetown, [South Africa], 367 p. Mag. Nat Hist., Ser. 13, 3:183-192.
VENKATARAMAN, G. 1964. The scombroid fishes of East Africa. Proc. Symp.
1961. Studies on the food and feeding relationships of the Scombroid Fishes, Mar. BioI. Assoc. India, Symp. Ser.
inshore fishes off Calicut on the Malabar coast. Indian l(part 1):107-164.
J. Fish. 7:275-306. 1968. Report on the Guinean Trawling Survey. Vol. 1.
VENKATA SUBBA RAO, K., V. R. RAO, P. MOJUMDER, T. A. ROA, General Report. Org. Afr. Unity, Sci., Tech. Res. Comm.,
S. REUBEN, S. S. DAN, AND B. N. RAo. Lagos, Nigeria. OAU/STRC Publ. 99, 828 p.
1981. Pelagic fishery resources of Lawson's Bay, Wal- WOLLAM, M. B.
tair. Indian J. Fish. 27:35-53. 1970. Description and distribution of larvae and early
VIJAYARAGHAVAN, P. juveniles of king mackerel, Scomberomorus cavalla
1955. Life-history and feeding habits of the spotted seer (Cuvier), and Spanish mackerel, Scomberomorus macu-
Scomberomorus guttatus (Bloch & Schneider). Indian latus (Mitchill); (Pisces: Scombridae); in the western
J. Fish. 2:360-372. North Atlantic. Fla. Dep. Nat. Resour. Tech. Ser. 61:
VINCIGUERRA, D. 1-35.
1890. Intorno ad alcune specie di pesci raccolte dal dottore WONGRATANA, T.
Enrico Stassano presso la costa occidentale del Sahara. 1968. Check list of fishes caught during the trawl surveys
Ann. Agrar. Roma 1890:61-103. in the Gulf of Thailand and off the east coast ofthe Malay
WAITE, E. R. Peninsula. Contrib. Mar. Fish. Lab. Bangkok 13, 96 p.
1904. A synopsis of the fishes of New South Wales. Mem. XIMENES, M. O. C.
N.S.w. Nat. Club 2, 59 p. 1983. Idade e crescimento da serra, Scomberomorus bra-
WALFORD, L. A. siliensis, no Estado do Ceara (Brasil). Arq. Cienc. Mar
1937. Marine game fishes of the Pacific coast from Alaska 21:47-54.
to the equator. Univ. Calif. Press, Berkeley, 205 p. XIMENES, M. O. C., M. F. DE MENEZES, AND A. A. FONTELES-
WALLS,J. G. FILHO.
1975. Fishes of the northern Gulf of Mexico. T. F. H. 1981. Idade e crescimento da cavala, Scomberomorus ca-
Publ., Neptune City, N.J., 432 p. valla (Cuvier), no Estado do Ceara (Brasil). Arq. Cienc.
WANG,Q. Mar 18:73-81.
1982. Preliminary studies on the rational utilization ofthe YANEZ-ARANCIBIA, A.
resources of Japanese Spanish mackerel, Scomberomorus 1980. Taxonom ia, ecologia y estructura de las comunidades
niphonius (Cuvier et Valenciennes). [In Chin.] Mar. de peces en lagunas costeras con bocas efimeras del Pa-
Fish., Shanghai 4(2):51-55. cifico de Mexico. Centro Cienc. Mar Limnol. Univ. Nac.
WARFEL, H. E. Auton. Mexico, Pub!. Esp. 2:1-306.
1950. Outlook for development of a tuna industry in the YE, C., AND D. ZHU.
Philippines. U.S. Fish Wild!. Serv., Res. Rep. 28, 37 p. 1984. The best economic result for the fisheries of the
WARFEL, H. E., AND P. R. MANACOP. Spanish mackerel. [In Chin., Eng!. abstr.] J. Fish.
1950. Otter trawl explorations in Philippine waters. U.S. China 8(2):171-177.
Fish Wildl. Serv., Res. Rep. 25, 49 p. ZAMA, A., AND K. FuJITA.
WHITLEY, G. P 1977. An annotated list of fishes from the Ogasawara
1927. A check list of fishes recorded from Fijian waters. J. Islands. J. Tokyo Univ. Fish. 63:87-138.
Pan-Pac. Res. Inst. 2(1):3-8. ZHANG, Q., AND Y. ZHANG.
1932. Fishes. Great Barrier Reef Exped. 1928-29. Sci. 1981. On the fish fauna of the South-Fujian and Taiwan
Rep. 4(9):267-316. Bank. [In Chin., Eng!. abstr.] Trans. Chinese Ichthyol.
1936. More ichthyological miscellanea. Mem. Queensl. Soc. 2:91-109.
Mus. 11(1):23-51. ZHAROV, V. 1., AND A. M. ZHUDOVA.
1947. New sharks and fishes from Western Australia. 1967. Some data on occurrence of scombroid larvae (order
Part 3. Aust. Zool. 11:129-150. Perciformes, suborder Scombroideil in the open waters of
1948. A list of the fishes of Western Australia. West. the tropical Atlantic. Tr. AtiantNIRO 18:201-214. [In
Aust. Fish. Dep., Fish. Bull. 2, 35 p. Russ., translated by W 1. Klawe, 1969, 21 p., Inter-Am.
1954. New locality records for some Australian fishes. Trop. Tuna Comm., La Jolla, Calif.]
Proc. R. Zoo!' Soc. N.S.w. 1952-53:23-30. .ZHUDOVA, A. M.
1964a. Scombroid fishes of Australia and New Zealand. 1969. Materials on the study ofthe eggs and larvae of some
Proc. Symp. Scombroid Fishes, Mar. BioI. Assoc. India, species of fish from the Gulf of Guinea and the adjacent
Symp. Ser. l(Part 1):221-253. waters of the open ocean. Tr. AtlantNIRO 22:135-163.
1964b. A survey of Australian ichthyology. Proc. Lin- [In Russ., translated by W L. Klawe, Inter-Am. Trop.
nean Soc. N.S.W 89:11-127. Tuna Comm., La Jolla, Calif.]
1968. A check-list of the fishes recorded from the New

689
 APPENDIX L

Characters used in analysis of Scomberomorus relationships

1) Posterodorsal spine of hyomandibula (Fig. 11) Relative size of scapular foramen (Fig. 43).
27). Plesiomorphous condition: absent. Char- Plesiomorphous condition: medium-sized. Char-
acter states: o(absent) , l(present and small), 2 acter states: O(large), l(medium), 2(small).
(large).
12) Pineal foramen (Figs. 11-13). Plesiomor-
2) Palatine tooth patch (Fig. 26). Plesiomor- phous condition: present. Character states: 0
phous condition: wide. Character states: O(wide), (present), l(reduced), 2(absent).
l(narrow), 2(very narrow).
13) Anterior ends of pterosphenoids (Figs. 17-
3) Inner process of palatine bone. Plesiomor- 19). Plesiomorphous condition: far apart. Char-
phous condition: short. Character states: O(very acter states: O(far apart), l(close together).
short, distance from dorsal hook of palatine to end
of inner process 54-72% of length to end of outer 14) Height of supraoccipital crest (Figs. 14-16).
process), l(short, 70-84%), 2(long, 97-99%). Plesiomorphous condition: low. Character states:
O(low), l(high).
4) Ventral process of angular (Fig. 25). Ple-
siomorphous condition: short. Character states: 15) Pterotic spine (Figs. 11-13). Plesiomorphous
O(short, 42-80% of length of dorsal process), 1 condition: well developed. Character states: 0
(moderate, 87-93%), 2(long, 117-126%). (well developed), l(essentially absent).

5) Posterior expansion of maxilla (Fig. 23). Ple- 16) Pelvic girdle: relative length of posterior
siomorphous condition: no expansion. Character process (Fig. 46). Plesiomorphous condition:
states: O(no posterior expansion of maxilla), 1 short to moderate-sized posterior process. Char-
(slight expansion), 2(moderate expansion), 3 acter states: O(short to moderate posterior process,
(marked expansion). 20-50% of length of anterior plate), l(long poste-
rior process, 62-90% of length of anterior plate).
6) Length of head of maxilla. Plesiomorphous
condition: long. Character states: O(short), 1 17) Spine on intercalar (Figs. 17-19). Plesiomor-
(medium), 2(long), 3(very long). phous condition: absent. Character states: O(ab-
sent), l(moderate length), 2(long).
7) Width of parasphenoid. Plesiomorphous
condition: narrow. Character states: O(narrow), 18) Swim bladder. Plesiomorphous condition:
l(wide). present. Character states: O(present), l(absent).

8) Angle of margins of anterior end ofpremaxilla 19) Curvature of lateral line (Figs. 50, 52, 68).
(Fig. 22). Plesiomorphous condition: blunt. Plesiomorphous condition: no abrupt curve.
Character states: O(blunt), l(intermediate), 2 Character states: O(no abrupt curve), l(abrupt
(acute). downward curve).

9) Length of ascending process of premaxilla 20) Auxiliary branches offlateralli.ne (Figs. 54,
(Fig. 22). Plesiomorphous condition: moderately 56). Plesiomorphous condition: absent. Char-
long. Character states: O(short), l(moderately acter states: O(absent), l(present).
long), 2(very long).
21) Nasal denticles (Fig. 1). Plesiomorphous
10) Vomerine ridge (Figs. 17-19). Plesiomor- condition: nasal chamber without denticIes.
phous condition: absent. Character states: 0 Character states: O(denticles absent), l(nasal den-
(absent),I(present). ticles present).

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COLLE'ITE and RUSSO: SPANISH MACKERELS

22) Anterior epibranchial artery (Fig. 7). Ple- ulate), l(spatulate, extending beyond anterior
siomorphous condition: unmodified. Character margin of ethmoid complex).
states: O(unmodified), l(esophageal artery arises
from fourth epibranchial), 2(coeliaco-mesenteric 32) Width of lateral wall of cleithrum (Fig.
shunt arises from fourth epibranchial). 43). Plesiomorphous condition: lateral wall nar-
row. Character states: O(lateral wall narrow,
23) Relative size of foramen between last radial space between cleithrum and coracoid visible in
and coracoid (Fig. 43). Plesiomorphous condi- lateral view), l(lateral wall wide, space between
tion: small. Character states: o(small), l(large), cleithrum and coracoid not visible in lateral view).
2(very large).
33) Epiotic crests (Figs. 11-13). Plesiomorphous
24) Length ofbranches ofpalatine bone (Fig. 26). condition: originate behind midfrontal region.
Plesiomorphous condition: ventral branch much Character states: O(originate behind midfrontal
shorter than dorsal branch. Character states: region), l(originate on anterior part of frontals).
O(ventral branch much shorter than dorsal branch,
120-123%), l(ventral branch slightly shorter, 112- 34) Vertebrae with inferior foramina. Plesio-
121%), 2(ventral branch equal to or longer than morphous condition: few. Character states: 0
dorsal branch, 87-107%). (few, < 11), l(many, more than 11).

25) Width of supratemporal (Fig. 42). Plesio- 35) Size of first basibranchial. Plesiomorphous
morphous condition: wider than deep. Character (:ondition: elongate. Character states: O(elon-
states: O(wider than deep, 101-113%), l(deeper gate),l(short).
than wide, 84-93%), 2(much deeper than wide,
49-79%). 36) Strut on fourth pharyngobranchial. Plesio-
morphous condition: not elongate. Character
26) Width of first postcleithrum (Fig. 44). states: O(not elongate), l(elongate).
Plesiomorphous condition: wide. Character
states: O(wide, 55-62% of length), l(narrow, 47- 37) Length of symplectic (Fig. 27). Plesiomor-
48%), 2(very narrow, 24-41%). phous condition: long. Character states: O(long,
in contact with metapterygoid), l(short, not in
27) Total number of vertebrae (Table 8). Ple- contact with metapterygoid).
siomorphous condition: few (31). Character
states: o(few, 31), l(moderate number, 41-56), 38) Size of dorsal and ventral hypohyals (Fig.
2(many, 62-64). 29). Plesiomorphous condition: ventral < 3 times
larger than dorsal. Character states: O(ventral
28) Depth ofurohyal (Fig. 31). Plesiomorphous hypohyal < 3 times larger than dorsal hypohyal in
condition: moderately deep. Character states: lateral view), l(ventral hypohyal > 3 times larger
O(shallow), l(moderately deep), 2(deep). than dorsal).

29) Shape of metapterygoid (Fig. 27). Plesio- 39) Position of fifth branchiostegal ray (Fig.
morphous condition: anterior oblique edge longer 29). Plesiomorphous condition: located on epi-
than posterior horizontal edge. Character states: hyal. Character states: O(completely on epihyal),
O(anterior oblique edge longer than posterior l(on suture between epihyal and ceratohyal).
horizontal edge), l(posterior horizontal edge long-
er than anterior oblique edge). 40) Posttemporal shelf (Fig. 40). Plesiomor-
phous condition: no shelf present. Character
30) Length of arms of ectopterygoid (Fig. 27). states: O(no shelf present between dorsal and
Plesiomorphous condition: dorsal arm longer than ventral arms of posttemporal), l(shelf present).
or equal to ventral arm. Character states: O(dor-
sal arm longer than or equal to ventral arm), 41) Width of supracleithrum (Fig. 41). Plesio-
l(dorsal arm shorter than ventral arm). morphous condition: wide. Character states: 0
(wide, 72-75% oflength), l(narrow, 42-62%).
31) Vomer (Figs. 17-19). Plesiomorphous condi-
tion: not spatulate. Character states: O(not spat- 42) Supratemporal pores (Fig. 42). Plesiomor-
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 FISHE~Y BULLETIN: VOL. 82, NO.4

phous condition: no pores. Character states: O(no 51) Anterior process on second postcleithrum
pores), l(pores present on dorsal arm). (Fig. 45). Plesiomorphous condition: elongate
process present on anterior margin of second
43) Position of nasals (Figs. 11-13). Plesiomor- postcleithrum. Character states: O(process pres-
phous condition: protrude beyond ethmoid region. ent), l(process absent).
Character states: O(protrude far beyond ethmoid
region), l(do not protrude, located adjacent to 52) Anterior end of first postcleithrum (Fig.
ethmoid region). 44). Plesiomorphous condition: notched. Char-
acter states: O(notched), l(pointed).
44) Shape of posterior end of dorsal margin of
urohyal (Fig. 31). Plesiomorphous condition: 53) Position of third pectoral fin radial (Fig.
tripartite. Character states: O(tripartite), 1 43). Plesiomorphous condition: base of third
(forked). radial completely on coracoid. Character states:
O(completely on coracoid), l(on suture between
45) Glossohyal teeth (Fig. 30). Plesiomorphous coracoid and scapula).
condition: glossohyal teeth present. Character
states: O(patch of teeth fused to dorsal surface of 54) Tooth shape. Plesiomorphous condition:
glossohyal), l(no glossohyal teeth). conical. Character states: O(conical), l(triangu-
lar and compressed). .
46) Width of hyomandibula (Fig. 27). Plesio-
morphous condition: narrow. Character states: 55) Parasphenoid contour. Plesiomorphous
O(narrow, width 35-36% of length), l(wide, width condition: concave ventrally. Character states:
36-52% oflength). O(concave),l(convex).

47) Angle of lateral and medial arms of fourth 56) Relative length of arms of dentary (Fig.
epibranchial. Plesiomorphous condition: more 24). Plesiomorphous condition: lower arm long-
acute. Character states: O(more acute), l(less est. Character states: O(lower arm longer than
acute). upper arm), l(upper arm longer than lower arm).

48) Anterior process of second epibranchial. 57) Length of posterior edge of ectopterygoid
Plesiomorphous condition: elongate. Character (Fig. 27). Plesiomorphous condition: posterior
states: O(elongate), l(not elongate). edge long. Character states: O(posterior edge
long, 64-68% of ventral distance), l(posterior edge
49) Preural centra 2-4 (Fig. 39). Plesio- short, 41c 63%).
morpho us condition: not shortened and com-
pressed. Character states: O(not compressed), 58) Shape of epihyal (Fig. 29). Plesiomorphous
l(compressed). condition: not much longer than deep. Character
states: O(depth 68-98% of length), l(much longer
50) Number of vertebrae supporting caudal than deep, 58-62% oflength).
fin. Plesiomorphous condition: 3 vertebrae sup-
port caudal fin. Character states: 0(3 vertebrae
support caudal), 1(4 or 5 vertebrae).

692