Osteology and Muscular Attachments of the Japanese Jack Mackerel, <I

BULLETIN OF MARINE SCIENCE. 58(2): 438-493. 1996

OSTEOLOGY AND MUSCULAR ATTACHMENTS OF THE JAPANESE JACK MACKEREL, TRACHURUS JAPONICUS

Yusuke Suda

ABSTRACT Detailed descriptions of the osteology, myology and arthrology of the Japanese jack mackerel Trachurus japonicus are presented. Several new characters of the family Carangidae identified are: fine ridges on the neural and hemal spines, branching of the lateral line scale canal, placement of the first hemal spine and the first anal proximal radial, presence of the anterior neural subzygapophyses, lateral keel on the fourth preural vertebra, and the arrangement of the ridges on the scales. Among these, fine ridges on the neural and hemal spines, plus the arrangements of ridges on the scales are considered to be important characters for future phylogenetic analyses of the Carangidae.

The Japanese jack mackerel, Trachurus japonicus (Temminck and Schlegel, 1844), is one of the most important food fishes in Japan (228, 894 ton catch in 199], FAa, ]993). However, species discrimination and confusion in the food industry has increased with the expansion of other jack mackerels imports. Even though this species is of commercial importance, detailed morphological descriptions of it as well as other Trachurus species are unavailable. Limited morphological information used to distinguish species of carangid fishes have been provided by many systematic workers, Starks (1911), Wakiya (1924), Oshima (1925), Suzuki (1962), Vergara (1974), Gushiken (1983), Smith- Vaniz (1984), and Gushiken (1988). Accordingly this study was initiated to try to provide complete osteological, myological and arthrological descriptions of T. japonicus. Analyses of external morphological differences and brief osteological description in this species were previously published (Suda, ]985, 1991, Suda et aI., 1987a, ]987b, ]987c). The data and observations from this study provide the bases for initial taxonomic, phylogenetic, paleontological and fisheries analyses.

MATERIAL AND METHODS

The following materials were used for dissection and/or study of the external morphology of T. japonicus. Measurements (given as mm SL unless otherwise indicated) and localities are in parentheses. The vertebral length is a length between the anterior tip of the atlas and the posterior margin of the 23rd vertebra. Comparative materials of familial species as well as others are listed in Appendix 2.

Trachurus japonicus (Temminck et Schlegel, 1844) NSMT-P 50363 (216, Nagasaki, Japan), NSMT-P 50366 (187, Sagami Bay, Japan), NSMT-P 50372 (I72, Nagasaki, Japan), NSMT-P 50379 (145, Sagami Bay, Japan), KMNH VR 100,099 (244, Kochi, Japan), KMNH VR 100,100 (228, Kochi, Japan), KMNH VR 100,101 (296, Kagoshima, Japan), KMNH VR 100,102 (221, Bungo Channel, Japan), KMNH VR 100,103 (53, Sagami Bay, Japan), KMNH VR 100,104 (268, Sagami Bay, Japan), KMNH VR 100,105 (262, Sagami Bay, Japan), KMNH VR 100,106 (292, Taiwan). Abbreviations of specimen catalogue numbers are: NSMT-P = Fish collection, Department of Zo- ology, National Science Museum, Tokyo; NSMT-PO = Skeletal specimens, Department of Paleontol- ogy, National Science Museum, Tokyo; NSMT-PV = Fossil collection, Department of Paleontology, National Science Museum, Tokyo; KMNH VR = Vertebrate collection, Kitakyushu Museum of Nat- ural History, Kitakyushu City, Japan; SKSK = Suruga-wan Kaiyo Seibutsu Kenkyukai (deposited in the Department of Zoology, National Science Museum, Tokyo). Bone was observed based on the stained and cleared or dissected materials, whereas muscles and ligaments based on the dissected materials without staining.

438 SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 439

Figure I. Left lateral view of the whole skeleton of Trachurus japonicus (NSMT-P 50372). SL 172 mm. Bar indicates 10 mm.

The drawings were made with the help of a camera lucida. Names of bones follow Uyeno (1975); bone parts (e.g., process, facet, condyle, foramen, etc.), mainly Barel et al. (1976), Potthoff and Kelley (1982), Potthof et al. (1988), Potthoff et al. (1987), Potthoff and Tellock (1993) and Tominaga (1968); cartilaginous elements in the caudal-fin, Fujita (1990); muscles and nerves, Winterbottom (1974) and Freihofer (1978), respectively. Abbreviations for bones, muscles and nerves are indicated in upper case, and for parts on bones in lower case.

MORPHOLOGICALDESCRIPTIONS Vertebra (Figs. 1-7).-Vertebral column total 24 elements, 10 abdominal (AVl- 10), 14 caudal vertebrae (CVI-14). Each centrum with neural arch (na) which ends with neural spine (ns) dorsally, and caudal vertebra with hemal arch (ha) ending with hemal spine (hs) ventrally. Basapophysis (ba) (basapophysis: Starck, 1979; Wake, 1979. basiventral: Goodrich, 1986. lateral process: Jollie, 1973. par- apophysis: Starks, 1901, 1911; Weitzman, 1962; Harder, 1975; Potthoff and Kelly, 1982; Collette and Russo, 1984. transverse process: Cole and Johnstone, 190 I; Stokely, 1952. ventral spinal process: Harder, 1975) projects ventrolaterally from anteroventral part of lateral side of centrum in AV4-10. Five kinds of articulation processes present (anterior neural zygapophysis: anz, anterior neural subzygapophysis: ansz, posterior neural zygapophysis: pnz, anterior hemal zygapophysis: ahz, posterior hemal zygapophysis: phz) on neural and hemal arches. BONYELEMENTS/SERIALTRANSFORMATIONS.Teardrop-like articulation fovea (af) and anterior face of centrum on first abdominal vertebra (or atlas) (AVI ) (Figs. 2, 3) (tripartite condyle, Potthoff and Tellock, 1993) articulate with exoccipital condyle (ec) and vertebral concavity (vc) on basioccipital (Figs. 22E, 26) respectively. Neural arch and spine of AVI autogenous. Base of both sides of neural arch divided into two projections: adaxially as arch pedicel (apd) and abaxially as transverse process (trp) (Fig. 3). Former articulates with dorsal face of centrum, and hollow of latter serves as articulation fovea for epipleural rib (fr', Fig. 2). Neural arch of AV2 projects laterally from centrum, and fovea for epipleural rib situated at lateral most part of its base (Fig. 2). In first six abdominal vertebrae, neural canal (neue) with rounded triangular shape in anterior view. Height of canal in abdominal vertebrae increases posteriorly, and decreases posteriorly in caudal vertebrae (Fig. 3). Groove present in anterior face of neural spines of AV2 to CV 12 (Fig. 3). Angle of inclination of neural spine to axis of centrum becomes smaller posteriorly to AV7, then larger to CV I, and again smaller posteriorly (Fig. 2). 440 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

AVI AV2 nf AV3 AV 4

AV5

Figure 2. Left lateral views of the vertebrae of Trachurus japonicus. NSMT-P 50366 for AVI and KMNH VR 100, 102 for remainder. See App. I for abbreviations.

Ventrally, origin of hemal arch moves from anterior position in eVI to extreme posterior position in eVl2 (Fig. 2). Hemal arches and spines on eV12, 13 autogenous. Shape of hemal canal (heme) is circular in AV9 and 10. Height of canal in caudal vertebrae becomes lower posteriorly (Fig. 3). Basapophyses on AV4-8 become larger posteriorly until fusing on AV9 and 10 (Figs. 2, 3) forming first hemal arches. Articulation fovea for rib (fr) present and well-defined on both sides of AV3-8, but poorly developed in AV9 and 10. SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 441

CYI2(PU3) CVt3(PU2) CY14IURSI

HYU30.4

HYUl'2

Iv' ridge ivf

Figure 2.-Continued.

Anterior neural zygapophysis (anz) presents on AV2-10 and eVl-13 (PU2), whereas posterior neural zygapophysis (pnz) on AVI-lO and eVl-1 I (Fig. 2). In AV7-10 and eVl-lO, small anterior neural subzygapophysis (ansz) beneath anz. Anterior hemal zygapophysis (ahz) originates from anterior part of hemal arch in eV2-II, while those of eV]2-14 arise from autogenous hemal arches on eV12, ]3 and parhypura] (PARH). Posterior hemal zygapophysis (phz) present from AV8 to eVlO. As shown in Figure 2, inferior vertebral foramen (ivf) (Starks, 1911; Suzuki, 1962; Gushiken, ]983; Smith-Vaniz, 1984. inferior foramen: Iwai et aI., 1965) appears at base of hemal arch in eV6-13, but in half of specimens, also found in eV5. Foramen very large anterior to eVIl, becoming quite small in eVIl 442 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

AVI AV2 AV3

Ava

eV4 eV5

eVil eV9 eVIO

Figure 3. Anterior views of the vertebrae of Trachurus japonicus. NSMT-P 50366 for AVI and KMNH VRIOO, 102 for remainder. See App. 1 for abbreviations. SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 443

ba ... ~;:~)::'i

. ,. ", '.L, '1. •.•• ".' ,\, ~" , AV2 AV3 A V4 "AV5 AV6

AV 7 AV8 AVlO CVl CV2

CV4 CV 3

I _~::::J (, . . .

~=~~~r~50CYIZ(PV31 CY1J(PV2) CY14(URSI

Figure 4. Ventral views of the vertebrae of Trachurus japonicus. NSMT-P 50366 for AVl and KMNH VR 100, 102 for remainder. See App. 1 for abbreviations. 444 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

Figure 5, Diagrammatic figures of the right lateral views of the ribs of Trachurus japonicus. A. pleural and epipleural rib on the 3rd-6th vertebrae; B. those on the 7-8th vertebrae; C. ligaments for the epipleural.

AV9

Figure 6. Diagrammatic figures of the ligaments of the vertebrae of Trachurus japonicus. A. left lateral view of the first two abdominal vertebrae; B. left lateral view of the 8-lOth abdominal vertebrae; C. left lateral view of the articulation part between the 2nd and 3rd caudal vertebrae; D. ventral view of the 8-lOth abdominal vertebrae. See App. I for abbreviations. SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 445

A 4 C ~ C 3 hSl 2 3 4

hSl

Figure 7. Air-bladder of Trachurus japonicus (KMNH VRlOO, 104). A, left lateral view; B. left lateral view of the posterior part; C, diagramatic figure of the anterodorsal view. Bars indicate 10 mm. See App. 1 for abbreviations. and posteriorly. Neural foramen (n1) (Potthoff and Kelley, 1982) appears on dorsolateral part of centra in AV2 to CV13. Prominent lateral keel (lk) protruding laterally from lateral side of centrum of eVil (Figs. 2-4). Fine ridges (Uyeno and Suda, 1991) run along longitudinal directions of lateral face of neural and hemal arches and spines (Figs. 2, 54A). They originate from both anterior and posterior portions of neural and hemal arches and join at basal part of each spine. Numbers of ridges at maximum at juncture and decrease distally on spine. Numbers of ridges 5 to 20 at juncture in specimens of 40 to 220 mm in vertebral lengths. Ridges clearly observed in dried material but may be difficult to see in wet condition. RIBS(Fig. 5). Pleural ribs (Weitzman, 1962; Whitehead and Teugels, 1985. haemal ribs: Phillips, 1942. ventral ribs: Harder, 1975) originate from fovea (fr) of AV3-1O. Epipleurals (Tominaga, 1968; Whitehead and Teugels, 1985. dorsal ribs: Harder, 1975. epicentrum: Phillips, 1942) present on AVI-lO and eVI-5, and lie in myoseptum intersections. Epipleurals on AVI and 2 articulate with fovea for epipleural (fr') (Fig. 2). Proximal tips of epipleurals on AV3-8 slightly flattened and articulate with furrow or facet on basal part of pleural rib (Fig. 5A, B). Epipleurals on AV9 and 10 attach to basal part of basapophyses, whereas those on eVI-5 attach to centra near base of hemal spines. Paired attaching ligaments are found only on epipleurals of AV3-8 (Fig. 5C): one from basapophysis and other from small projection on basal part of pleural rib. VERTEBRALLIGAMENTS(Fig. 6). Interspinal ligament (ISL) joins each neural spine to next one from AVI to eVll (Fig. 6A). Nuchal ligament (NL, Fig. 6A) connects posterior margin of rim of foramen magnum (fm, Fig. 22E) to antero- basal margin of neural spine of atlas. Interzygapophysis ligament (IL) connects adjoining vertebrae from: dorsal margin of posterior neural zygapophysis (pnz) 446 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

PAoe A

eV12 (PU3)

Figure 8, Left lateral view of the caudal-fin skeleton (A) and cartilaginous elements in the caudal- fin (B) of Trachurus japonicus (NSMT-P 50372). Bar indicates 2 mm. See App, I for abbreviations. to ventral margin of anterior neural zygapophysis (anz) of next vertebra (AY 1- CVlO) (Fig. 6A, B, C); ventral margin of pnz to either dorsal margin of anterior neural subzygapophysis (ansz) of next vertebra (AY7-CVlO) (Fig. 6B, C) or to ventrolateral part of next centrum (AVl, 2) (Fig. 6A). As shown in Fig. 6D, internal pleural rib--basapophysis ligament (lRBL) extends from corresponding posterior furrow on basapophysis (ba) to medial face of basal part of the pleural rib. External pleural rib-basapophysis ligament (ERBL) arises from corresponding anterior furrow on basapophysis and divides into flat, thin medial and lateral bands; former connects to basal part of pleural rib anterior to it, while latter attaches to medial part of pleural rib two before it (Fig. 6B, D). AIR-BLADDER(Fig. 7). Torpedo shaped air-bladder lies from AVI to CV4. Tra- churus japonicus one of very few carangid species in which hemal spines (hsl- 3 or hsl--4) penetrate air-bladder without any special modification of bones. As with all carangids, no otophysic connection between anterior part of air-bladder and neurocranium. Caudal-fin Skeleton (Figs. 2,4, 8-9).-Caudal-fin skeleton composed of folJowing elements: third preural vertebra (PU3 = CVI2), second pre ural (PU2 = CVI3), urostyle (URS = CV14 = PUI) fused with uroneural (URN), parhypural (PARH), two epurals (EPUI, EPU2), fused hypural 1 + 2 (HYUI + 2), fused hypural 3 + 4 (HYU3 + 4), the fifth hypural (HYU5), 17 principal caudal-fin rays (PCR) (15 branched and two, one outer dorsad and outer ventrad, unbranched: ub) and procurrent rays (PROC). Neural spine of preural centrum 3 (npu3) and autogenous hemal spine of preural centrum 3 (hpu3) extend posteriorly to urostyle. PU2 also has posteriorly elongated autogenous hemal spine (hpu2), but this spine exhibits small ridge anteriorly at its base (Fig. 2). Neural spine of second preural (npu2) is very reduced. Fan- shaped lateral process, hypurapophysis (hyp), found on both sides of parhypural. Dorsally, two epurals present between npu3 and URS. Procurrent spur and associated basal shortening of anteriorly preceding ray as described by Johnson (1975) not found in Trachurus japonicus. Three free cartilaginous elements, post-hypural cartilage (CPHY5) and two post-hemal spine cartilage (CPHPU2, 3), present in the posterior part of caudal- fin skeleton (Fig. 8B). MUSCLESOF CAUDAL-FIN(Fig. 9). Superficial muscles of caudal-fin: interradialis (INT), epaxialis (EPAX), hypaxialis (HYP), lateralis superficialis (LATS), supracarinalis posterior (SCARP) and infracarinalis posterior (ICARP) (Fig. 9A). Interradialis (INT) interconnects ventrolateral face of 1st to 6th branched cau- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 447

A ====:::;=~b --- 7 6 5 4 3 2 1 1

ICARP

Figure 9. Diagrammatic figures of the left lateral views of the muscles of the caudal-fin of Trachurus japonicus. A, superficial muscles; B, interior muscles. See App. I for abbreviations.

dal-fin rays of upper lobe to dorsolateral face of 1st to the 5th rays of lower lobe. Posterior parts of epaxialis (EPAX) and hypaxialis (HYP) attach to lateral face of branched principal caudal-fin rays. Sheet like lateralis superficialis (LATS) lies on epaxialis and hypaxialis. While supracarinalis posterior (SCARP) originates from anterior groove of npu3 and connects it to dorsal-fin, infracarinalis posterior (lCARP) arises from anterior groove of hpu3 and connects it to anal-fin. Under superficial muscles are flexor dorsalis (FD), flexor dorsalis superior (FDS), hypochordal longitudinalis (HYL), flexor ventralis (FV), flexor ventralis inferior (FVI), and flexor ventralis externus (FVE) (Fig. 9B). Flexor dorsalis (FD) is longitudinal muscle running along lateral side of caudal peduncle, originating from PU2-4 and inserting on fin rays 1-8 of the upper lobe; flexor ventralis externus (FVE) similarly configured, but connecting PU3-4 to lower lobe fin rays 2-5. Flexor dorsalis superior (FDS), slim bundle of fibers, 448 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

Figure 10. Left lateral view of the dorsal-fin pterygiophores of Trachurus japonicus (NSMT-P 50363). Roman and Arabic numerals indicate spines and soft rays respectively. Bar indicates 5 mm. See App. I for abbreviations.

passes posteriorly from npu3 to upper lobe's unbranched principal fin ray, whereas ventrally, flexor ventralis inferior (FVI) connects hpu3 to lower lobe's unbranched principal fin ray. Posterodorsal face of hypurapophysis (hyp) and anterior part of lower hypural plate (HYUl + 2) are origins of hypochordal longitudinalis (HYL) which inserts on 6-9th upper fin rays. Flexor ventralis (FV) develops from lateral face of PU2 and splits, inserting on 3-8th and 9-11 th caudal-fin rays of lower lobe. Dorsal-fin Skeleton (Figs. 1, 10-12).-Three supraneurals (SUPN) (Weitzman, ]962; Mabee, 1988. predorsal: Smith and Bailey, 1961; Kendall, 1976; Potthoff et aI., 1988; Potthoff and Tellock, 1993; Potthoff et aI., 1987) with knob-like dorsal portion present anterior to first dorsal-fin pterygiophores. First supraneural lies anterior to first neural spine (ns1) of AV], SUPN 2,3 insert between ns2 and ns3 (Fig. 10). In Fig. 11, dorsal-fin pterygiophores consist of paired radials, proximal radial (PR) and distal radial (DR). These pterygiophores divided into two segments: first dorsal-fin set supporting eight dorsal-fin spines (OS), and second dorsal-fin set supporting one spine and 27 to 35 soft rays. In T. japonicus, no stay or end piece (Weitzman, 1962) present after last pterygiophore. Each proximal radial has a lateral wing (lw) on its shaft portion. Shape of distal radial supporting spines differ from those supporting soft rays. Former wing-like (Fig. ] lA) bearing hook- shaped prolongation (h, Bridge, 1896), whereas latter paired and pebble-like (Fig. ] 1B). Distal radials in spine supporting portion possess shallow groove between both wings, mating with similar groove on proximal radial. Only first proximal radial has no groove. These grooves serve to house dorsal-fin spine when deflected and is character mainly found in carangid and scombrid fishes. First two dorsal-fin spines exhibit chain-link articulation with PR 1 as no distal radial present (Fig. 11A). All other spines articulate as shown in Fig. 11A (PR2), including hook-shaped prolongation (h) that penetrates fenestra of dorsal spine and articulates to osseous tubercle (t, Bridge, ]896) of next proximal radial. Posteriorly positioned proximal radials of soft rays act in pairs to firmly pinch SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 449

OS3

.~/:'.. '

h .

...... ,,' ...:.....:: ... , •... : ..•.•. ~.. .~l:. ..."~' . ~~;~~,•...<~;.:. ...~.. ~ A ,--,/ OR2

OS1 OS2

pUr

PR2

~~~ Iw DR

Figure II. Anterodorsal views of the dorsal-fin skeleton of Trachurus japonicus (KMNH VRIOO, 102). A, first two elements; B, 26th soft ray and its supporting pterygiophores; C, the last element of the soft ray portion. Broken lines indicate connections between articulations. Bar indicates 2 mm. See App. I for abbreviations. 450 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

DS1 DS2

Figure 12. Diagrammatic figures of the anterodorsal views of the muscles of the first two elements of the dorsal-fin skeleton of Trachurus japonicus. Solid lines indicate ligaments. See App. 1 for abbreviations.

neural spine between them to give second dorsal-fin extra degree of rigidity (note ns13-18 in Fig. 10 and Suzuki, 1962, pp. 122-126), MUSCLESANDLIGAMENTSOFDORSAL-FIN(Fig. 12). Series of three muscles and number of ligaments control dorsal-fins. Except for first two dorsal-fin spines in which erectores dorsales (ERECD) arises from anterolateral face of first proximal radial (PRl), it originates from posterolateral face of preceding proximal radial. In both cases, insertion point is small process for insertion of erector muscle (ep) on basal part of each fin element. In opposition, depressores dorsales (DEPRD) originates from anterolateral face of proximal radial and inserts on small process for insertion of depressor muscle (dp), situated at posterolateral basal part of each fin element. Inclinatores dorsales (INCLD) arising from fascia between skin and insertion portion of epaxialis body muscle, divides in two on spines, and remains single on soft rays. With former, anterior section inserts on ventral face of wing-like process of distal radial (DR) and posterior section inserts on area between both processes for insertions of depressor and erector muscles. In soft ray supporting portion, insertion of inclinatores dorsales is only basal part of soft ray. Ligament connects attaching process for ligament of the fin ray (plfr) to attaching process for ligament of proximal radial (plpr). Anal-fin Skeleton (Figs. 1, 13-16).-Anal-fin pterygiophores consist of paired proximal and distal radials (PR, DR, Fig. 15). These pterygiophores support three anal-fin spines (AS) and 25 to 31 soft rays. Wide space present between anterior two spines and third spine (Figs. 13, 14). No stay (Weitzman, 1962) after last pterygiophore (Fig. 15B). In Figure 15, first proximal radial (PRl) is divided into shaft portion and articulation head. Posterior margin of former closely approximates to anterior face of first hemal spine (hsl, Figs. 13, 14), whereas latter expanded anteriorly serving as attachment for infracarinalis medius muscle (ICARM) (Fig. 16) and elongated posteriorly. Wide space between anterior two anal spines and third spine, syna- pomorphy of Carangidae (Smith-Vaniz, 1984), produced by posterior elongation of articulation head. Posterior end of elongated articulation head deeply forked into which anterior projection of distal radial (apdr) inserts. Hook-shaped prolon- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 451

gation (h, Bridge, 1896) extends posteriorly from DR I and penetrates fenestra of basal part of AS3 (Fig. 15A). Each proximal radial except first element has longitudinal lateral wing (lw) of shaft portion. Shape of distal radial supporting soft rays paired and pebble-like. In same manner as in dorsal-fin, first two anal spines exhibit chain-link articulation with PR 1. Many proximal radials of soft rays act in pairs to firmly pinch hemal spine between them to give anal-fin extra degree of rigidity (Fig. 13). MUSCLESOF ANAL-FIN(Fig. 16). Series of three muscles and several ligaments control anal-fin. Erectores anales (ERECA) originates from posterolateral face of preceding proximal radial (PR) except for anal-fin spines which arise from anterolateral face of proximal radials. For both cases, insertion point is small process for insertion of erector muscle (ep, Fig. 15A) on basal part of each fin element. Depressores anales (DEPRA) originates from anterolateral face of PR and inserts on small process for insertion of depressor muscle (dp, Fig. 15A) situated at posterolateral basal part of each fin ray. Inc1inatores anales (INCLA) arises from fascia between skin and epaxialis body muscle. It inserts on posterior elongation of articulation head of PR1 in spine supporting area, whereas in soft ray supporting portion, insertion for inclinatores anales is only basal part of soft ray. Infracarinalis medius (ICARM) arises from anterolateral face of anteriorly expanded part of articulation head of PRI and attaches to posterior xiphoid process (px) of pelvic girdle (Fig. 21B).

Pectoral Girdle (Figs. ]7-] 9).-Pectoral girdle comprises posttemporal (PT), su- prac1eithrum (SCL), cleithrum (CLE), scapula (SC), coracoid (CRA), 4 proximal pectoral radials (PPR), 20-21 cartilaginous distal pectoral radials (DPR), dorsal element of postcleithrum (DPCL), ventral element of postcleithrum (YPCL), 21- 22 fin rays, dorsal element of supratemporal-intertemporal (DST), and ventral element of supratemporal-intertemporal (YST). Posttemporal (Fig. 18A) has two arms, dorsal arm of posttemporal (da) and ventral arm of posttemporal (va). Former marked by its flattened termination articulates with triangular shaped dorsal face of epiotic (sulp, Fig. 22B). Latter stick-like and articulates with protuberance of intercalar (pIp, Fig. 26). Tubular structure for lateral line canal present at center part of bone. Lateral line canal extending from ventral element of supratemporal-intertemporal (YST, Fig. 17) enters anterior formamen of posttemporal (afpt), and emerges through posterior foramen of posttemporal (pfpt) to supracleithrum. Medial surface of posttemporal articulates with supracleithrum at three places (Fig. 18A): dorsal facet for supracleithrum (dsclf) connects with dorsal facet for posttemporal (dptf) at dorsal vertex of supracleithrum; other two articulations, mesial facet for supracleithrum (msclf) and ventral facet for supracleithrum (vsclf) articulate with mesial and ventral facet for posttemporal (mptf, vptf) respectively. Supracleithrum (Fig. 18A) thickened anteriorly and thin posteriorly. Lateral line canal from posttemporal enters anterior foramen of supracleithrum (afscl) and emerges through posterior foramen of supracleithrum (pfscl) to first lateral line scale. Baudelot's ligament (BL) arises from medial surface of bone and attaches to basioccipital (sBL, Fig. 26). Scapula (Fig. 18B) has large scapula foramen (sf) at its center. Posterior margin of bone forms articulations for proximal pectoral radials (PPRl-4). Saddle-like posterodorsal corner is articulation facet for first pectoral-fin ray. Scapula posteroventrally connected with coracoid by sutured joint. Proximal pectoral radials 452 BULLETIN OF MARINE SCIENCE. VOL. 58. NO.2. 1996

hsB

Figure 13. Left lateral view of the anal-fin pterygiophores of Trachurus japonicus (NSMT-P 50363). Roman and Arabic numerals indicate spines and soft rays respectively. Bar indicates 5 mm. See App. 1 for abbreviations. become larger ventrally and connected to scapula by ligaments. Distal pectoral radials (DPR) located at bases of pectoral-fin rays. Cleithrum (Fig. 18C) is largest bone in pectoral girdle, and forms main portion of anterior margin of girdle. It terminates dorsally and ventrally in pointed process. Bone divided into dorsal expanded region (posterior plate: Collette and Rus- so, 1984) and longitudinal shaft region with two flanges (lfc: lateral flange, mfc: medial flange). In adults, anterior margin of dorsal expanded region and/or ventral part of lateral flange often exhibit hyperostosis (Starks, 1911). Coracoid (Fig. 18C, D) anteriorly connected to cleithrum and dorsally to scapula. Two fenestrae, dorsal fenestra (dorf) and ventral fenestra (venf), present between coracoid and cleithrum. Ventral process of coracoid (vp) slender and forms posteroventral margin of ventral fenestra. As both ventral tips of coracoid and cleithrum not connected, ventral fenestra open ventrally. Dorsal element of postcleithrum (DPCL) bean-like in shape with slightly thickened anterior margin (Fig. 17), whereas ventral element (VPCL) divided into dorsal lamellar portion and ventral elongated portion. Ligament arising from part between lamellar and elongated portions of ventral element attaches to dorsal process of pelvic girdle (dpp, Fig. 20). Dorsal element of supratemporal-intertemporal (DST) cylindrical, whereas ventral element (YST) is trifurcate tubular bone (Fig. 17). Sensory canal enters ventral foramen of YST from sixth neurocraniallateral-line foramen (nlf6, Figs. 22A, B, 26) and bifurcates dorsally and posteriorly. Dorsal branch goes up to DST and posterior branch goes to posttemporal (PT) through posterior foramen of YST. MUSCLESOF PECTORALGIRDLE(Fig. 19). On lateral side of pectoral girdle, levator pectoralis (LPECT), sternohyoideus (STH), pharyngoclavicularis externus (PHCE), pharyngoclavicularis intern us (PHCI), hypaxialis (HYP) and abductor superficialis (ABS) are recognized superficially (Fig. 19A, B). The levator pectoralis (LPECT) arises from the lateral face of the ventral arm of the posttemporal (PT) and attaches to the pterotic (Fig. 27). The pharyngo- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 453

Figure 14. Left lateral view of the anterior part of the anal-fin skeleton of Trachurus japonicus (NSMT-P 50366). Bar indicates 5 mm. See App. 1 for abbreviations.

c1avicularis extemus (PHCE) and pharyngoclavicularis internus (PHCI) originate from the anterior part of the cleithrum (CLE) and attach to the ventral part of the fifth ceratobranchial (CBR5, Fig. 49B). The stemohyoideus (STH) also originates from the anterior face of the c1eithrum and attaches to the urohyal anteriorly (Fig. 45B, C) and hypaxialis (HYP) posteriorly. The abductor superficialis (ABS) originates from the posterior face of the c1eithrum and inserts on the lateral face of the pectoral-fin rays. After removing the abductor superficialis, the arrector ventralis (ARRV, ARRV'), abductor profundus (ABP) and coracoradialis (CRR) are exposed (Fig. 19C, 0). The arrector ventralis (ARRV) originates from the inside of the deep groove between two flanges (mfc, Ifc) of the cleithrum and inserts on the anterodorsal tip of the first pectoral-fin ray. A subdivision of the arrector ventralis (ARRV') arises from the lateral face of the coracoid (CRA) and inserts on the posterolateral face of the tip of the first pectoral-fin ray (Fig. 190). The origin of the abductor profundus (ABP) is the lateral face of the coracoid and the insertion includes the lateral face of the basal parts of the second to ventralmost fin rays. The coracoradialis (CRR) originates from the posterodorsal face of the posteriorly 454 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

Figure 15. Anal-fin skeleton of Trachurus japonicus (KMNH VRIOO. 102), A. anteroventral view of the left side of the first two elements; B. left lateral view of the last element. Broken lines indicate connections between articulations, Bar indicates 2 mm. See App, I for abbreviations,

protruded part of the coracoid and inserts on the ventral face of the fourth proximal pectoral radial. Muscles on medial side of pectoral girdle consist of adductor superficialis (ADS), adductor profundus (ADP), arrector dorsalis (ARRD) and adductor radialis (ADR) (Fig. 19E, F). Adductor superficialis (ADS) develops from medial face of cleithrum and inserts on medial faces of pectoral-fin rays 2-17. Origin of adductor profundus (ADP) is medial face of coracoid and insertions are lateral faces of pectoral-fin rays 2-16. After removing those muscles, arrector dorsalis (ARRD) and adductor radialis (ADR) exposed (Fig. 19F). Former originates from near anterior margin of cleithrum and inserts on anterior tip of first fin ray, while latter arises from medial faces of basal parts of third and fourth proximal pectoral radials and inserts on 17th to ventral most rays. Infracarinalis anterior (ICARA) (Figs. 19D, 21A) connects pectoral girdle to SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 455

DEPRA

Figure 16 (left). Diagrammatic figure of the left lateral view of the muscles of the first element of the anal-fin skeleton of Trachurus japonicus. Solid lines indicate ligaments. See App. 1 for abbreviations.

Figure 17 (right). Left pectoral girdle of Trachurus japonicus (NSMT-P 50371). Left, lateral view; right, medial view. Bar indicates 5 mm. See App. I for abbreviations. pelvic girdle. It attaches mainly to ventromedial face of cleithrum and partially to lateral face of ventral tip of same bone. Pelvic Girdle (Figs. 20, 21).-Two longitudinal keels, dorsally suprapelvic keel (spk) and ventrally subpelvic keel (sbk) develop from lateral margin of pelvis (P). Both keels originate from head part that articulates with pelvic spine and rays. Suprapelvic keel extends to anterior tip of pelvis, whereas subpelvic keel reaches anterior one-third of pelvis. Two projections, anterior and posterior xiphoid process (ax, px) (de Sylva, 1984; Potthoff et aI., 1987, 1988; Potthoff and Tellock, 1993), present on articulation head part. Both posterior xiphoid processes in firm contact with each other in anterior half and are deeply forked posteriorly. Small and deep hollow present between articulation head and posterodorsal face of pelvis. Pelvic girdle connects with pectoral girdle at two places. Anteriorly, lateral surface of pelvis closely attaches to medial face of pectoral girdle (Fig. 180). Posteriorly, ligament connects dorsal process of pelvic girdle (dpp) to ventral element of postcleithrum (VPCL, Fig. 17). MUSCLESOF PELVICGIRDLE(Fig. 21). Muscles of pelvic girdle consist of abductor superficialis pelvicus (ABSP), abductor profundus pelvicus (ABPP), arrector ventralis pelvicus (ARRVP), adductor superficialis pelvicus (AOSP), adductor profundus pelvicus (ADPP), arrector dorsalis pelvicus (ARRDP), infracarinalis anterior (ICARA) and infracarinalis medius (ICARM). Abductor superficialis pelvicus (ABSP) and abductor profundus pelvicus (ABPP) originate from ventral face of pelvis (P) and insert on bases of all soft rays. Origin of former just inside latter. Origin of arrector ventralis pelvicus (ARRVP) is ventrolateral face of pelvis and insertion is ventrolateral face of pelvic-fin spine. Adductor superficialis pelvicus (AOSP) and adductor profundus pelvicus (AOPP) develop from dorsal face of pelvis and insert on bases of all soft rays. Origin of arrector dorsalis pelvicus (ARROP) is ventrolateral face of pelvis and 456 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

opk

Figure 18, Bones which compose the pectoral girdle of Trachurus japonicus (NSMT-P 50363 for A, C, D; NSMT-P 50371 for B). A, posttemporal and suprac1eithrum (left, lateral view; right, medial view); B, left lateral view of basal part of the pectoral-fin; C, left lateral view of the cleithrum, scapula and coracoid; D, medial view of the lower part of the left pectoral girdle especially showing articulation with the pelvic girdle. Broken lines indicate connections between articulations. Bars indicate 2 mm for A, B, D and 5 mm for C. See App. I for abbreviations. insertion is base of pelvic-fin spine. Part of fibers of ARRDP continue anteriorly to aponeurosis which attaches to medial face of coracoid. Infracarinalis anterior (ICARA) connects pelvic girdle to pectoral girdle. It originates from cleithrum (Fig. 19D) and inserts on dorsal process (dpp) of pelvic girdle and partially on small process situated just anterior to base of pelvic-fin spine. Infracarinalis medius (ICARM) connects pelvic girdle to anal fin. It arises from posterior xiphoid process (px) and attaches to articulation head part of first proximal radial of anal-fin (PR1, Fig. 16). SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 457

B c

Figure 19. Diagrammatic figures of the muscles of the left pectoral girdle of Trachurus japonicus. A-D. lateral views; E-F, medial views.

Neurocranium (Figs. 22-28).-Dorsal part of neurocranium has three crests: continuous frontal and supraoccipital crests (frocr, spocr) run along mid-line of dorsal face of neurocranium; pterotic crest (ptocr) forms dorsolateral margin of neurocranium; frontal-parietal crest (fpcr) (temporal crest: Tominaga, 1968) runs between those two crests. ETHMOIDREGION(Fig. 23). Prevomer (PREV), mesethmoid (MESETH) and lateral ethmoids (ETH LAT) occupy anterior part of neurocranium. Prevomer connects with mesethmoid and lateral ethmoid dorsally and parasphenoid (PARA) posteroventrally. Dorsal mid-line of prevomer and mesethmoid keeled. Keel of prevomer capped with cartilage. Prevomer has row of tiny conical teeth along anteroventral margin of its head region (Figs. 23B, 24A) and low longitudinal crest with minute serration (serrate crest on prevomer: sere) on ventral part of shaft region (Figs. 23B, 24B). Concavity located just behind tooth bearing part of head region is attaching surface (sV) for posterior palato-vomerine ligament (V). Cartilaginous articulation facet for palatine (fp) present at junction between prevomer and lateral ethmoid. Lateral ethmoid characterized by having wing like projection that spreads out laterally. Large hollow for olfactory rosette, olfactory fossa (of), lies between 458 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

sbk

Figure 20. Pelvic girdle of Trachurus japonicus (KMNH VRlOO, 102). A, dorsal view; B, left posteroventral view of the head portion. Shaded areas indicate ligaments. Bars indicate 2 mm. See App. I for abbreviations. anterior faces of mesethmoid and lateral ethmoid. Foramen for olfactory nerve (fon) present on orbital side of lateral ethmoid (Fig. 23B). Two condyles capped with cartilage present on ventral face of wing like projection of lateral ethmoid. Outer one, articulation condyle for lachrymal (el), connects with dorsal surface of lachrymal (fnc, Fig. 30) and inner one, articulation condyle for ectopterygoid (ce), attaches to facet of ectopterygoid (lep, Fig. 35). An area surrounded by ventral margin of lateral ethmoid, prevomer and parasphenoid is ethmoid cartilage (EC) which is continuous with lamina orbitonasalis (LAMO, Stiassny, 1986). ORBITALREGION(Fig. 25). Dorsal and ventral parts of orbital region comprise the frontals (FRO), pterosphenoids (PTE), basisphenoid (BASP) and parasphenoid (PARA). Frontal crest (frocr) originates from anteriormost part of frontal (FRO) and continues to supraoccipital crest (spocr). Frontal-parietal crest (fpcr) arises from anterior one-third of frontal and runs parallel to frontal and supraoccipital crest. SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 459

Figure 21. Diagrammatic figures of the muscles of the pelvic girdle of Trachurus japonicus. A, ventral view; B, dorsal view. See App. I for abbreviations.

Four foramina for frontal sensory canals, neurocranial lateral line foramen (nlfO- 3) present on frontal (Fig. 22): nlfO opens dorsally at posterior part of frocr; nlfl opens at anteriormost part of bone; nlf2 is at anterolateral part of bone; and nlf3 positioned posterolaterally on frontal. Sensory canals run completely through bony tube. Nasal continuous with nlfl (Fig. 42B). Pterosphenoid (PTE, Fig. 25) located on both sides of anterior opening of cavum cranii (aocc). It connects with frontal anterolaterally, sphenotic (SPH) laterally, prootic (PRO) posteroventrally and basisphenoid (BASP) ventromedially. Basisphenoid (BASP, Fig. 25) composed of peduncle and wing-like portions (basisphenoid peduncle: bsp, basisphenoid wing: bsw) present at posterior corner of orbital cavity and situated in front of anterior opening of myodome (aom). Basisphenoid peduncle develops from basisphenoid wing vertically to parasphenoid (PARA). Anterior opening of cavum cranii divided into anterior and posterior portions by bsw. Parasphenoid (PARA, Figs. 23, 25, 26) is elongate bone that stretches under ethmoid, otic and occipital regions. At anterior part of otic region, parasphenoid wing (w) developed dorsally and connects with prootic (PRO) by suture. Internal carotid foramen (fica) present at posterobasal part of parasphenoid wing. Posterior part of bone extending to posterior opening of myodome (porn) deeply incised (Fig. 26). OTIC REGION(Figs. 25, 26). Composed of sphenotics (SPH), prootics (PRO), pterotics (PTO), epiotics (EPO) and intercalars (INCA). Sphenotic (SPH) positioned at anterodorsal part of otic region. It bears a tri- gonal postorbital process (Barel et aI., 1976) (pop) at its lateral most part. Neu- rocranium widest at this process. Pterotic (PTO) situated at dorsolateral portion of region. Pterotic crest (ptocr) is tubular structure with three neurocranial lateral-line foramina (nlf4-6) on its edge: largest nlf4 at junction between PTO and FRO; nlf5 and 6 located at posterior part of ptocr. Lateral line sensory canal running through ptocr bifurcated at 460 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

D lPocr

Figure 22. Neurocranium of Trachurus japonicus (NSMT-P 50366), A, left lateral view; B, dorsal view; C. ventral view; D, anterior view; E. posterior view. Bars indicate 5 mm, See App. I for abbreviations.

posterior part of crest: ventrally it emerges through nlf5 to preopercle; dorsally it emerges through nlf6 to ventral element of supratemporal-intertemporal. Three other accessory foramina present on crest. Posterolaterally, ptocr ends as back- wardly projecting pterotic spine (ps). Just posterior to postorbital process of sphenotic (pop), two concavities for articulation with hyomandibular present: anteriorly articulation facet for anterior condyle of hyomandibular (fach), and posteriorly articulation facet for posterior condyle of hyomandibular (fpch). Former occurs between prootic and sphenotic, whereas latter confined to pterotic. Dila- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON1CUS 461

A ", ~.'.~. .. ·frocr

.FRO

Figure 23. Ethmoid region of the neurocranium of Trachurus japonicus (NSMT-P 50363). A, left anterodorsal view; B, left posteroventral view. Dotted and shaded areas show cartilage and ligament respectively. Bar indicates 2 mm. See App. I for abbreviations. tator area (dar) is narrow space between pterotic crest and postorbital process from which dilatator operculi muscle (DO) originates (Figs. 27, 41 A). Prootic (PRO) occupies anterior part of otic bulla and surrounded by sphenotic anterodorsally, pterotic dorsally, exoccipital (EXO) and basioccipital (BAO) posteriorly, and parasphenoid ventrally. Cavity positioned at anterolateral corner of 462 BULLETIN OF MARINE SCIENCE. VOL. 58, NO.2, 1996

A -'(Lr'i ,-''~"~_ () . Vl.../- V--

B ---\-;v' .,". .••.

Figure 24 (left), Posterior view of teeth on the prevomer (A) and right view of teeth on the serrate crest of prevomer (B) of Trachurus japonicus (NSMT-P 50363). Bar indicates 0.5 mm.

Figure 25 (right). Anteroventral view of the left side of the otic and the trigemino facialis chamber (encircled by broken line) region of Trachurus japonicus (NSMT-P 50366). Bar indicates 2 mm. See App. I for abbreviations.

Figure 26. Posteroventral view of the left side of the neurocranium showing the otic and occipital region of Trachurus japonicus (NSMT-P 50366). Bar indicates 2 mm. See App. I for abbreviations, SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 463

LO LPOST LEXT

Figure 27. Diagrammatic figure of the left lateral view of the muscles of the neurocranium of Tra- churus japonicus. See App. I for abbreviations. LINT2, 3 and TDA are not drawn because they present behind LEXT. prootic just beneath sphenotic is trigemino-facialis chamber (Fig. 25). Chamber consists of lateral commissure (llc) laterally and two foramina (trigeminal foramen: tf, facial foramen: ff, Patterson, 1964) inside chamber. Five nerves (otic branch of facial nerve, superficial ophthalmic nerve, profundus nerve, trigeminal nerve, and buccal branch of facial nerve) and two blood vessels (jugular vein, orbital artery) emerge from trigeminal foramen. On other hand, hyomandibular trunk of facial nerve, jugular vein, and orbital artery emerge from facial foramen. Oculomotor nerve emerges from another foramen positioned just anterior to trigeminal foramen. Structure of chamber and arrangement of nerves and blood vessels shows "Perciformes" pattern (Patterson, 1964: 438). Epiotics (EPO) are separated by supraoccipital. Triangular shaped dorsal face of epiotic is articulation surface for upper limb of posttemporal (sulp). From posterior end of articulation surface, round bony pillar runs posteroventrally and joins to exoccipital by suture. Intercalar (INCA) is very thin bone lying on ventrolateral face of pillar part formed by pterotic and exoccipital. Small projection of intercalar, articulation protuberance for lower limb of posttemporal (pIp) located on its lateral edge.

A B~ ..... " ~ ~

transparent zone ~ opaque zone ----

Figure 28. Left sagitta of Trachurus japonicus (NSMT-P 50363). A, lateral view; B, dorsal view; C, medial view. Bar indicates I mm. 464 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

OCCIPITALREGION.Region occupies posterior part of neurocranium and consists of the parietals (PARI), supraoccipital (SUPO), exoccipital (EXO) and basioccipital (BAO). Parietals (PARI, Fig. 22) sit on both sides of supraoccipital on dorsolateral face of neurocranium and forms part of frontal-parietal crest (fpcr). Supraoccipital (SUPO, Fig. 22) occupies posterodorsal part of neurocranium and characterized by having dorsoposterior extension of supraoccipital crest (spocr). Crest has no ridge on its lateral face. In adults, the basal part of the crest often hyperossified (hyperostosis: Starks, 1911). Exoccipital (EXO, Figs. 22, 26) located at posterior center of neurocranium. Just below supraoccipital crest is foramen magnum (fm). Exoccipital condyles (ec) articulate with articulation fovea (af) of atlas (Figs. 2, 3). Bony pillar extends anteriorly from condyle and branches anterolaterally to pterotic and dorsally to epiotic. Adjoining two foramina for nerves present on ventral face of basal portion of bony pillar: anteriorly foramen for nervus glossopharyngeus (fng) and posteriorly foramen for nervus vagus (fnv) (Freihofer, 1978). Basioccipital (BAO) situated at posteroventral portion of neurocranium and makes up posterior half of otic bulla. Posterior end of bone contains vertebral concavity (vc) for first centrum. Large posterior opening of myodome (porn) opens just anteroventral to articulation concavity. Baudelot's ligament originates from attaching surface (sBL) situated just dorsal to opening of myodome. MUSCLESOF NEUROCRANIUM(Fig. 27). The posterior face of the postorbital process (pop) of the sphenotic is the origin of the levator arcus palatini (LAP) which inserts on the lateral face of the articulation head of the hyomandibular. The posterior part of the sphenotic and anterodorsal part of the lateral face of the prootic is the origin of the adductor arcus palatini (not figured). The dilatator operculi (DO) originates from the narrow space (dilatator area: dar, Fig. 26) between the pterotic crest and postorbital process and inserts on the dilatator process (dilp) of the operc1e (Figs. 39, 41A). The origin of the levator operculi (LO) is the posterior face of the pterotic crest and the insertion is the upper part of the operc1e (Fig. 41A). The posterior part of the pterotic and anterodorsal part of the exoccipital is the origin of the levator pectoralis (LPECT) which inserts on the posttemporal (Figs. 19A, 41A). The neurocranium is connected with the gill arch by four muscles (levatores externi: LEXT, levatores interni: LINT, levator posterior: LPOST, tranversus dorsalis anterior: TDA) (Fig. 49A). LEXT and LINT arise from the flat part of the prootic surrounded by the postorbital process, facial foramen of the trigemino- facialis chamber, and articulation facet for the anterior condyle of the hyomandibular. The former inserts on the epibranchials 1-4 (EPBI-4, Fig. 49A) and the latter inserts on the dorsal face of the pharyngobranchials 2 and 3 (PB2, 3, Fig, 49A). LPOST develops from the ventral face of the pillar part of the exoccipital and inserts on the dorsal face of the fourth epibranchial (EPB4, Fig. 49A). TDA originates from the parasphenoid and prootic, and inserts on the epibranchial 2 (EPB2). OTOLITHS(Fig. 28). Sagitta, largest otolith, has small anterior projection, irreg- ularly waved dorsal edge and smooth ventral margin. Degree of development of anterior projection differs among individuals. Prominent groove, sulcus (Chaine and Duvergier, 1934), present on medial side.

Sensory Canal Bones (Figs. 29-31).-Nasal elements consist of nasals (NAS) and fragmentary prenasals (PRENAS), situated superficially between neurocraniallat- eralline foramen (nlfl) of frontal and palatinal wing of maxilla (plw) (Fig. 42B). SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 465 \)~ vST

Figure 29. Outline of the left facial bones of Trachurus japonicus (NSMT-P 50363). Bar indicates 5 mm. Broken lines indicate overlapping bones. See App. 1 for abbreviations.

IFO 1 Figure 30. Lateral view of the left sensory canal bones of Trachurus japonicus (NSMT-P 50366). Bar indicates 5 mm. See App. 1 for abbreviations. 466 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2. 1996

asp

c~~ _ = ~-l"~~"'''.'' ..' ...., ....:...... " ,- -' •••••• -0, ,....: .• : J' " ~._-~.' " If05

Figure 31 (left). Right third infraorbital bone of Trachurus japonicus (KMNH VRlOO, 102). A, lateral view; B, dorsal view; C, ventral view. Bar indicates 2 mm. See App. 1 for abbreviations.

Figure 32 (right). Left premaxilla of Trachurus japonicus (NSMT-P 50379 for A. B, C; NSMT-P 50363 for D). A, lateral view; B, medial view; C, ventral view; D, medial view of teeth. Bars indicate 2 mm for A, B, C and 0.5 mm for D. See App. 1 for abbreviations.

Infraorbital sensory canal runs through bony tube of infraorbitals (IFOI-6), except sixth element in which canal runs through open groove (Fig. 30). Tube of each element has several foramina on lateral surface. Lachrymal (IFOI) is largest and only element that has articulation. Articulation facet for neurocranium (fnc) situated at dorsal part of bone articulates with condyle on wing part of lateral ethmoid (c1, Fig. 23). Third infraorbital (Fig. 31) has medially developed infraorbital shelf (ifos) from its dorsal margin and elongate projection at posterobasal part of infraorbital shelf. Latter connected with anterior face of preopercle by ligament (Fig. 4IA). Ventral surface of infraorbital shelf is origin of section Al of adductor mandibulae (Fig. 4IA). Oromandibular Area.-JAws (Figs. 29, 32-34). Premaxilla (PREM, Fig. 32) composed of two arms, vertical ascending arm and horizontal dentigerous arm. Former includes small ascending process (asp) laterally, articulation process for maxilla (pm) medially and rostral cartilage (RC, Fig. 42) anteriorly, while latter bears triangular postmaxillary process (pmp) dorsally and row of small conical teeth with medially bent tips (Fig. 32C, D). Depth of ascending arm about 45% of length of dentigerous arm. Ascending process reduced and shorter than articulation process. Posterior face of articulation process articulates with head of maxilla. Maxillo-rostroid ligament (VII) and palato-premaxillary ligament (XII) attach to anterior face of rostral cartilage (Fig. 42A, B). Maxilla (MAX, Fig. 33) consists of anterior articulation head and spatula-like handle portions. Articulation head has three processes protruding in various di- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 467

D ...... UPM

"';."." ", .. ~

Figure 33 (left). Left maxilla and supramaxilla of Trachurus japonicus (NSMT-P 50379). A, lateral view; B, dorsal view, C, medial view; D, lateral view of the supramaxilla. Bar indicates 2 mm. See App. I for abbreviations.

Figure 34 (right). Left lower jaw bones of Trachurus japonicus (NSMT-P 50379 for A, B, C; NSMT- P 50363 for D). A, lateral view; B. medial view; C, dorsal view of dentary; D, medial view of teeth. Bars indicate 2 mm for A, B, C and 0.5 mm for D. See App. I for abbreviations. rections: neurocranial condyle (nc) dorsally, premaxillary wing (pmw) medially and palatinal wing (plw) laterally. Neurocranial condyle is semicircular in shape and situated at dorsal part of articulation head. Anterodorsal part of condyle is attaching surface for maxillo-rostroid ligament (VII, Fig. 42A, B). Medial face of palatinal wing is attaching surface for anterior maxillo-premaxillary ligament (IX, Fig. 42C). Maxillary process of palatine (mxp) articulates with palatinal wing just anterior to nail-like attaching process for primordial ligament (ppl) (Figs. 41A, 42A). Round hollow formed by these three processes is articulation facet for the premaxilla (fpmx, Fig. 33C) which sits astride the articulation process for the maxilla (pm, Fig. 32). The medial surface just behind the neurocranial condyle is the insertion for the section Al of the adductor mandibulae (Fig. 4IA). The handle portion is posteriorly expanded. The posterodorsal corner is angular, whereas the posteroventral corner is rounded. A long supramaxilla (SUPM, Fig. 330) with an anterior falciform projection is present. Its length is about two-thirds of the maxilla. The lower jaw (Fig. 34) is interrupted by two fissures, the dorsal obturated foramen (dof) and the ventral obturated foramen (vof). The mandibular sensory canal runs through a bony tube along the ventral margin of the dentary (DEN) 468 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

Figure 35. Lateral view of the left suspensorium of Trachurus japonicus (NSMT-P 50366), Bar indicates 5 mm. See App. I for abbreviations. and angular (ANG) which have a series of mandibule sensory canal foramina (fmsc: 1-3 on ANG, 4-8 on DEN). The dentary has a dorsal limb referred as the coronoid wing (cw) with an expanded coronoid process (cp). The dentigerous area runs from the symphysis to the area below the anterior portion of the coronoid process and bears a row of small conical teeth with medially bent tips (Fig. 34C, D). The dental recess (dr) is a shallow hollow located on the anterior part of the lateral surface of the bone. The bifurcated area of the dentary is deeply penetrated anteriorly and forms the dentary fossa (dent) into which the anterior part of the angular inserts. The anterior part of the lateral surface of the coronoid process is an attachment for robust connective tissue from the medial surface of the maxilla. The angular is arrowhead shaped. A thumb-like dorsal projection is the primordial process (prmp), in which a low primordial ridge (pr) runs along posterior margin of primordial process. Saddle-like suspensorial articulation facet (sat) which connects with quadrate present on dorsal face of posterior area of bone. This facet bordered dorsolaterally by semicircular lateral facet rim (lfr). Short horizontal ridge at basal part of primordial process is attaching ridge for primordial ligament (cpl, Fig. 4IA). On medial face, Meckelian cartilage (MC) originates from Meckelian ridge (Mr) and extends anteriorly into dentary fossa. Retroarticular (RET) is at posteroventral corner of lower jaw. PALATINE-PTERYGOID-QUADRATE-HYOMANDIBULAR-SYMPLECTICCOMPLEX(Figs. 29, 35-39). Palatine (PAL, Fig. 36) situated at anteriormost part of complex and continues posteriorly to ectopterygoid. It has finger-like maxillary process (mxp) anteriorly which articulates with lateral surface of articulation head of maxilla (Figs. 41A, 42A). Ventral margin of bone bears irregular rows of minute teeth SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 469

Figure 36 (left). Right palatine of Trachurus japonicus (KMNH VRIOO, 102 for A, B, C; NSMT-P 50363 for D). Bar indicates 2 mm for A, B, C and 0.5 mm for D. See App. I for abbreviations.

Figure 37 (right). Medial view of the left quadrate, symplectic and metapterygoid of Trachurus japonicus (NSMT-P 50363). Bar indicates 2 mm. See App. 1 for abbreviations.

(Fig. 36D). On dorsomedial surface is articulation facet (fme) with mesethmoid (fp, Fig. 23) present. Just anterior to this facet is attaching furrow (fppl) for palato- premaxillary ligament (XII) which connects with rostral cartilage (RC, Fig. 42A, B). Below articulation facet for mesethmoid is attaching process (ppvl) for posterior palato-vomerine ligament (V, Fig. 42C). Ectopterygoid (ECP, Fig. 35) has lateral ethmoid process (lep) at anterodorsal part of bone that articulates with articulation condyle for ectopterygoid of lateral ethmoid (ce, Fig. 23). Bone has small bill-shaped projection on its posterodorsal corner, Endopterygoid (ENP, Fig. 35) is very thin bone next to ectopterygoid and metapterygoid. Its anterior part contacts orbital side of lateral ethmoid by sheet- like connective tissue. Dorsal part of metapterygoid (MET, Figs. 35, 37) connected to hyomandibular by suture. Delicate flat wing, metapterygoid process (mpp) (metapterygoid lamina: Katayama, 1959), develops from its posterior margin to anterior margin of arm rim of hyomandibular. Quadrate (QUA, Figs. 35, 37) is fan-shaped bone situated at anteroventral corner of suspensorium and articulates with angular by articulation facet for mandibule (fmd). Bone extends posterodorsally and ends as spinous projection. On medial surface, symplectic-quadrate groove (sqg) present along posteroventral margin of bone into which symplectic (SYM) inserts. Two interosseous spaces present around symplectic (Figs. 35, 37): dorsally 470 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

pen

' . ::;"':,:=...... '::,',:' )",:,' " .: ....

. ~"'. ~.. .~ . '- :~:}.!:i)~ . '::" ....~. ',: ...... " .. "~' ..;.....:. " . ':",.' .," .,' ..... :>" " • ;:',:·f . '" .,"I , '

Figure 38. Left hyomandibular of Trachurus japonicus (KMNH VRlOO, 102). Left, lateral view; right, medial view. Bar indicates 2 mm. See App. I for abbreviations. dorsal preopercle-symplectic interosseous space (dpss) among preopercle, posterior tip of symplectic, ventral tip of hyomandibular and metapterygoid process; posteriorly ventral preopercle-symplectic interosseous space (vpss) between posterior margin of symplectic and preopercle, Hyomandibular (HYO, Fig. 38) divided into three portions: articulation head, arm rim, and flange zone. Articulation head is dorsal part of bone that has three condyles: anterior condyle of hyomandibular for neurocranium (acn), posterior condyle of hyomandibular for neurocranium (pen), and posteriormost opercular condyle (oc). First two condyles articulate with lateral otic region of neurocranium (fach, fpch, Fig. 26) respectively and opercular condyle articulates with opercle SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 471

Figure 39 (left). Medial view of the articulation part of the left opercle and hyomandibular of Tra- churus japonicus (NSMT-P 50366). Bar indicates 2 mm. See App. I for abbreviations.

Figure 40 (right). Lateral view of the left interopercle of Trachurus japonicus (NSMT-P 50363). Shaded area shows ligament. Bar indicates 5 mm. See App. I for abbreviations.

(Fig. 39). Arm rim is vertical shaft part of bone. Transverse ridge that originates from place below posterior condyle of hyomandibular runs on rim. Flange zone situated at anterior part of bone. Ventral part of zone connects with metapterygoid by suture. Hyomandibular trunk of facial nerve enters dorsal foramen for hyomandibular trunk of facial nerve (df) on medial side of flange zone, runs through arm rim ventrally and emerges through ventral foramen for hyomandibular trunk of facial nerve (vf) near ventral tip of arm rim. Opercular Series (Figs. 29, 35, 39, 40).-Opercular series consists of opercle (OPE), preopercle (PREO), interopercle (INO) and subopercle (SUBOP). Opercle (OPE) articulates with hyomandibular at suspensorium articulation socket (sas) anteriorly (Fig. 39). Nail-like adductor process (ap) present on medial surface just posterior to articulation socket. Suspensorium ridge (sr) runs along anterior margin of bone and terminates as dilatator process (dilp) at dorsal end (Fig. 35). As posterior part of opercle largely indented, bone appears to have two posterior processes (Fig. 35). Dorsal process weakly ossified and fimbriates, whereas ventral one is triangular in shape. Preopercular sensory canal runs through tubular portion of preopercle (PREO, Fig. 35). Area has several foramina along this portion. Margin of flange area of PREO almost smooth. Conspicuous ligament connects anterior middle face of bone with spinous projection of third infraorbital (Figs. 31, 41A). Interopercle (INO) is thin lamellar bone with slightly thickened dorsal area (Fig. 40). Two ligaments originate from dorsal margin of bone. Anterior one attaches to ligament process on lateral side of posterior ceratohyal (pipl), whereas posterior one to spur of epihyal (se) (Figs. 43, 45). Ventral margin of bone smooth. Anterodorsal corner of subopercle (SUBOP) projects dorsally. Posterodorsal end of bone poorly ossified. MUSCLESOF CHEEK(Fig. 41). Most conspicuous muscle of cheek region is adductor mandibulae, which is primarily responsible for closing jaws; subdivided into section AI, A2a, A213 and AW. Section Al (AI' of Smith-Vaniz, 1984) originates from ventral face of infraorbital shelf (ifos, Fig. 31) of third infraorbital (IF03) and inserts on neurocranial condyle (nc) of maxilla (Fig. 33). This pattern 472 BULLETIN OF MARINE SCIENCE. VOL. 58, NO.2. 1996

Figure 41. Diagrammatic figures of the left cheek and jaw muscles of Trachurus japonicus. A, lateral view; B, medial view. Solid line, ligament; striped line, nerve. See App. I for abbreviations. corresponds to carangid type of adductor mandibulae (Gosline, 1986). Origin of A2a is dorsal half of the anterior face of preopercle and that of A213 includes metapterygoid, quadrate, symplectic and ectopterygoid. Both A2a and A213con- tinue anteriorly to primordial ligament (PL) and AW. Primordial ligament attaches to attaching process (ppl) on anterolateral face of maxilla. Insertion of AW is medial face of angular laterally and dentary fossa (denf) anteriorly (Fig. 41B). Ramus maxillaris inferioris (103) runs along dorsal face of the AI. Ramus mandibularis trigeminus (102) runs between layers of A2a and A213,and emerges externally near base of quadrate. It subsequently divides into two branches, of which lower branch goes anteriorly between angular and AW. Ramus mandibularis externus facialis (RMEF) emerges from ventral foramen for hyomandibular trunk of facial nerve (vf) on hyomandibular shaft (Fig. 38) and goes into ventral preopercle-symplectic interosseous space (vpss); subsequently runs along symplectic- quadrate groove (sqg, Fig. 37) and goes over medial surface of angular above suspensorial articulation facet (saf, Fig. 34B). Protractor hyoidei (PRHY) originates from anteromedial surface of dentary and inserts on anterior ceratohyal (AC, Fig. 45A). Intermandibularis (lMD) interconnects anteromedial part of dentary (DEN) of both sides. Buccal Ligaments (Fig. 42).-Buccalligament system comprises posterior palato- vomerine ligaments (V, Roman numerals adopted to Stiassny, 1986), maxillo- rostroid ligaments (VII), anterior maxillo-premaxillary ligaments (IX), and palato- premaxillary ligaments (XII). Stout posterior palato-vomerine ligament (V) originates from concavity on ventral surface of prevomer (sV, Figs. 23B, 42C) and inserts on attaching process for posterior palato-vomerine ligament (ppvl, Fig. 36B) on medial side of palatine (PAL). Maxillo-rostroid ligament (VII) connects medial part of neurocranial condyle of maxilla (nc, Fig. 33A, B) to rostral cartilage (RC). Short anterior maxillo- premaxillary ligament (IX) present between medial surface of rostral cartilage and premaxillary wing of maxilla (pmw, Fig. 33A, B, C). Palato-premaxillary ligament (XII) arises from furrow (fppl, Fig. 36B), crosses over VII and inserts on rostral cartilage. Generally, basal acanthomorph system involves 'V', anterior palato-vomerine ligaments (VI), VII, IX, ethmo-maxillary ligaments (XI) and XII (Stiassny, 1986). SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 473

FRO

Figure 42. Buccal ligaments of Trachurus japonicus (NSMT-P 50372). A, left lateral view; B, dorsal view; C, ventral view (left upper jaw is protruded). Dotted and shaded areas show cartilage and ligaments respectively. Bars indicate 2 mm. See App. I for abbreviations.

As Trachurus japonicus lacks VI and XI, it has more simple buccal ligament system than basal acanthomorphs. Hyoid Arch (Figs. 43--45).-Basihyal (BAH) spatula-shaped in dorsal view. Pos- teroventral part serves as attachment for anterodorsal corner of dorsal hypohyal. Dorsal face of bone bears minute conical teeth with posteriorly bent tips (Fig. 43C, D). 474 BULLETIN OF MARINE SCIENCE. VOL. 58, NO.2. 1996

BAH

fdh A

B c

Figure 43. Left hyoid arch of Trachurus japonicus (NSMT-P 50366). A, lateral view; B, medial view; C, dorsal view of the basihyal; D, lateral view of the teeth on the basihyal. Dotted area, cartilage. Bars indicate 5 mm for A and B, I mm for C, and 0.5 mm for D. See App. I for abbreviations.

Hypohyal consists of dorsal and ventral hypohyals (DHYPH, VHYPH); connected to each other by suture joint and cartilage. Ligament connects attaching process for dorsal hypohyal-hypobranchialligament (phhl) on anterodorsal corner of dorsal hypohyal to anterolateral part of first hypobranchial (HYBR1, Fig. 46). Another ligament from anteroventral surface of basibranchial attaches to lateral surface of ventral hypohyal just below suture. Anterior part of medial surface of ventral hypohyal and hyoid process of urohyal (hp, Fig. 44) connected to each other by robust ligament. Hyoidean artery passes through foramen and bone from medial to lateral on dorsal hypohyal (fdh) on lateral surface. Anterior ceratohyal (AC) (Potthoff and Tellock, 1993) connected with hypohyals by synchondrosis and with posterior ceratohyal (PC) (Potthoff and Tellock, 1993) by strongly jagged suture on medial surface. Groove for hyoidean artery (gha) which partially interrupted by Beryciform foramen (bf, McAllister, 1968) on anterior ceratohyal runs longitudinally on dorsolateral surface. Ventral margin of anterior ceratohyal notched where proximal tips of branchiostegals rays 2 and 3 (BRA2, 3) attach. Posteroventral part of lateral surface shallowly concave for attachment of branchiostegals 4 and 5 (BRA4, 5). SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 475

A Ir

HAD hp

Iw B hp

STH

Figure 44 (left). Urohyal of Trachurus japonicus (NSMT-P 50366). A, left lateral view; B, ventral view. Bar indicates 2 mm. See App. I for abbreviations.

Figure 45 (right). Diagrammatic figures of the muscles of the hyoid arch of Trachurus japonicus. A, left lateral view; B, left lateral view of the urohyal; C, ventral view of the urohyaL Shaded areas indicate ligaments. See App. I for abbreviations.

Posterior part of posterior ceratohyal is articulation facet for epihyal (fe). On lateral surface, just anteroventral to facet is process for interoperc1e-posterior ceratohyal ligament (pipl). Anteroventral part of lateral surface shallowly concave for attaching surfaces of branchiostegal rays 6 and 7. Distal tip of epihyal (EPH, Potthoff and Tellock, 1993) attaches to posterior tip of symplectic and ventral tip of hyomandibular by cartilage. Ligament originates from spur of epihyal (se) and attaches to interoperc1e (Fig. 40). Branchiostegal rays (BRA) become longer posteriorly, first five branchiostegals connect with anterior ceratohyal and remaining two with posterior ceratohyal. Proximal tips of BRAl-3 pointed and have spur proximally at posterior margin, whereas BRA4-7 expanded and have no projections. Urohyal (Fig. 44) is slender lamellar bone and positioned beneath branchial arch. Spine like attaching process (plb) for ligament from second basibranchial, is present at anterodorsal corner of urohyal. Pair of hyoid processes (hp) present on anterior end of bone and connected with ventral hypohyals by ligaments. Two lateral wings (Iw) present along ventral margin of bone. Low lateral ridge (Ir) runs along lamellar part of bone from anteroventral part to posterodorsal tip. MUSCLESOFHYOIDARCH(Fig. 45). Protractor hyoidei (PRHY) extends from medial surface of dentary (Fig. 41B), and attaches to anterolateral surface of anterior ceratohyal and proximal anterior face of branchiostegal rays 2 and 3. Hyohyoidei abductores (HAB) originates from ventromedial surface of ventral hypohyal and inserts on anterior face of branchiostegal ray ] of opposite side. Hyohyoidei adductores (HAD) lies distally between branchiostegal rays. 476 BULLETIN OF MARINE SCIENCE, VOL. 58. NO.2, 1996 BAH--1 A

HYSR2

HYSR3

GSR 1

GSR 2

GSR3

GSR4

GSR5 (LP)

UP4 , ~ EPB4~~J EPB3 :::itJ~::::~ ~ a u' PBl ---1J ~

Figure 46. Branchial arch of Trachurus japonicus (NSMT-P 50366). A, ventral view; B, dorsal view. Three tooth plate groups are encircled by broken lines respectively in B. Bar indicates 5 mm. See App. 1 for abbreviations.

Sternohyoideus (STH) connects urohyal to cleithrum (CLE, Fig. 19A). It arises from inner face of groove formed by two longitudinal wings and area between lateral ridge and wing. Gill Arch (Figs. 46-49),-Four basibranchials (BABI-4), 3 hypobranchials (HYBRI-3) and 5 ceratobranchials (CBRI-5) make lower half of the gill arch, and 4 epibranchials (EPB 1-4), 4 pharyngobranchials (PB 1-4) and interarcual cartilage (IC) form upper half. Plate-like basibranchial 1 (BAB 1) connects with anteroventral corner of hypobranchial 1 (HYBR1) on its posteroventral corner. Basibranchial 2 has two triangular flanges extending ventrally from its ventral surface. Bone attaches to HYBR3 on its lateral sides. Stick-like BAB3 longest element in basibranchial series and attaches to HYBR2, 3. Basibranchial 4 is pebble-like and cartilaginous. Hypobranchials (HYBR) attach to corresponding basibranchials anterolaterally and to ceratobranchials posteriorly. Hypobranchials 1 and 2 composed of head parts and ventrally grooved shank parts. Hypobranchial 3 is triangular in dorsal view, and has large flange on ventromedial margin. Ceratobranchials 1-4 (CBRI-4) stick-like in shape with grooved ventral surfaces. Last element (CBR5) is specialized lower pharyngeal bone (LP). Medial margin of CBR4 clearly notched at anterior one-third of bone. Lower pharyngeal SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 477

EPB4

PB3

Figure 47. Upper half of the left branchial arch of Trachurus japonicus (NSMT-P 50363). A. artic- ulated condition (dorsal view); B, medial view; C. disarticulated condition (dorsal view). Bars indicate 2 mm. See App. I for abbreviations.

(CBR5) marked by long triangular pharyngeal tooth plate bearing small conical teeth that become larger posteromedially (Fig. 48A). Epibranchials (EPB, Fig. 47) consist of axial parts and caudally situated uncinate processes (unc, Hosoya and Kawamura, 1993; Lauder and Liem, 1983; Rosen, 1973; Travers, 1981). Anterior tip of EPBl connects with pharyngobranchial 1 (PB 1). It also connects with PB2 by interarcual cartilage (IC, Hosoya and Kawamura, 1993; Lauder and Liem, 1983; Travers, 1981) which originates from basal part of uncinate process. Pharyngobranchiall (suspensory pharyngeal: Tominaga, 1968) attaches to otic region of neurocranium just behind internal carotid foramen (fica, Fig. 26). Anterior tip of EPB2 slightly expanded and attaches to posterior cartilaginous part of PB2 and centrally situated cartilaginous condyle of PB3. Epibranchial 3 articulates with cartilaginous condyle at posterolateral part of pharyngobranchial 3. Epibranchial 4 connects with cartilaginous and block-like PB4 situated at anteromedial part of upper pharyngeal tooth plate 4 (UP4). Bony components constituting the upper half of the gill arch, presence of a cartilaginous pharyngobranchial 4, and articulation pattern between epibranchial 2 and pharyngobranchials 2 and 3 correspond to primitive perciform and percoid conditions mentioned by Johnson (1986). 478 BULLETIN OF MARINE SCIENCE, VOL. 58. NO.2, 1996

, -.\ '1 c

Figure 48. Gill rakers on the lower half of the left gill arch and lower pharyngeal bone of Trachurus japonicus (NSMT-P 50363). A, dorsal view of the lower pharyngeal bone; B, dorsal view of the posterior area of the left lower gill arch; C, anterior view of the gill raker on the outer row of the first gill arch; D, dorsal view of the gill rakers of the second gill arch. Bar indicate I mm. See App. I for abbreviations.

BASI BRANCHIAL DENTITION (Fig. 46B). Various sizes of free tooth plates arranged along long axis of basibranchial series. These free plates divided into three groups: from anterior to posterior, tp (tooth plate group) 1, tp2 and tp3. Tooth plate group 1 consists of two small plates situated on junction between basi bran- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 479

A B ~

Figure 49. Diagrammatic figures of the gill arch muscles of Trachurus japonicus. A, dorsal view of the upper gill arch muscles; 8, ventral view of the lower gill arch muscles. See App. I for abbreviations. chials I and 2 (BAB I, 2) and hypobranchial 1 (HYBRl). Tooth plate 2 is group of plates scattered near junction between BAB3 and HYBR2. Tooth plate 3 group positioned between HYBR3 of both sides. GILLRAKERS(Fig. 48). Gill rakers arranged in two rows on each gill arch except on fifth arch that has lower pharyngeal tooth plate. Gill rakers on outer row of first arch are comb-like with minute spinelets on their medial faces (Fig. 48C), whereas other gill rakers tubercle-like in shape with minute spinelets on their dorsal faces (Fig. 48D). MUSCLESOF GILLARCH(Fig. 49). Upper gill arch includes: levatores externi (LEXTI-4), levatores interni (LINT2, 3), levator posterior (LPOST), transversus dorsalis anterior (TDA), obliqui dorsales (OBLD), obliquus posterior (OBLP), transversus dorsalis posterior (TDP), retractor dorsalis (RDORS), adductores (AD) and sphincter oesophagi (SPHO) (Fig. 49A). Levatores externi (LEXT), levatores interni (LINT), levator posterior (LPOST) and transversus dorsalis anterior (TDA) connect gill arch to neurocranium (Fig. 27). LEXT originate from lateral face of flat part of prootic (PRO, Fig. 26) and insert on uncinate processes (unc) of epibranchials (EPB). LINT originate from flat part of prootic just behind origin of LEXT and insert on dorsal faces of pharyngobranchials 2 and 3 (PB2, 3). LPOST originates from ventral face of pillar part of exoccipital (Fig. 27) and inserts on uncinate process of epibranchial 4 (EPB4). TDA arises from ventral part of parasphenoid and prootic, and inserts on dorsomedial face of epibranchial 2. Obliquus dorsalis (OBLD) interconnects uncinate process of EPB3 and PB3, 480 BULLETIN OF MARINE SCIENCE. VOL. 58. NO.2, 1996

Figure 50. Left lateral view of the scutes of Trachurus japonicus (KMNH VRlOO, 103 for A; KMNH VR100, 102 for B-E). A, whole length; B, the largest scute on the curved portion (left, lateral view; right, medial view); C, scute on the transition part; D, the largest scute on the straight portion; E, scute on the caudal peduncle. Bar indicates 5 mm for B-E.

4. Obliquus posterior (OBLP) connects posterior face of EPB4 to posterior tip of ceratobranchial 5 (LP). Transversus dorsalis posterior (TOP) interconnects dorsomedial faces of EPB3 of both sides. Origin of retractor dorsalis (ROORS) is vertebral column and insertion is dorsal face of upper pharyngeal 4 (UP4). Ad- ductores (AD) are small muscles interconnecting epibranchials and ceratobranchials. Sphincter oesophagi (SPHO) attaches to ventral face of upper pharyngeal 4 and ventral face of ceratobranchial 5 (LP). Muscles of lower gill arch include obliqui ventrales (OBLV), recti ventrales (RECTV), rectus communis (RCOMM), pharyngoclavicularis externus (PHCE), pharyngoclavicularis intemus (PHCI) and transversi ventrales (TRV) (Fig. 49B). Obliqui ventrales (OBLV) span joint between ventral surface of hypobranchials (HYBR) and ceratobranchials (CBR) of the first three arches. Recti ventrales (RECTV) connect posterior margin of triangular process on ventral surface of HYBR3 to ventral surface of CBR4. Rectus communis (RCOMM) connects ventral margin of triangular process of HYBR3 to ventrolateral part of CBR5 (LP). Transversi ventrales (TRV) interconnects ventrolateral part of CBR5 (LP) of both sides. Pharyngoclavicularis extemus (PHCE) arises from ventrolateral part of CBR5 just outside origin of transversi ventrales and attaches to anteroventral part of cleithrum (CLE, Fig. 19A). Just posterior to insertion of pharyngoclavicularis externus is insertion of pharyngoclavicularis intemus (PHC!) which originates from anteromesial face of cleithrum (Fig. 19A). Scales (Figs. 50-53).-Lateralline scales transformed into strongly armed scutes. Anterior half of lateral line curved, whereas posterior half is straight (Fig. 50A). Posterior margin of each scute on curved portion of lateral line slightly and round- ly projected (Fig. 50B), whereas those in transitional portion between curved and straight portions slightly pointed (Fig. 50C), and those in straight portion have strong spines (Fig. 500, E). Scutes number 32-43 (mean 37.7) in curved portion, 28-39 (mean 33.6) on straight portion and 66-77 (mean 71.3) total (Suda, 1985). While depths of scutes on curved portion are of almost equal size, deepest scute situated at anterior one-third of straight portion (Fig. 50D). Lateral line sensory canal enters anterior foramen on lateral face of each scute and branches into three directions (Fig. 51). Body scales (Fig. 52) small, deciduous and cycloid. Although size of scales SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 481

Figure 51. Diagrammatic figure of the flow of the body lateral line canal of Trachurus japonicus. Scutes in medial view. Solid line shows the body lateral line canal.

Figure 52. Enlarged left lateral views of the scales for the various parts of the body of Trachurus japonicus (KMNH VR100, 104). *, enlarged figure of the scale on the same area of KMNH VRI00, 102 specimen is shown in Figure 53. Bar indicates 2 mm for the scale. 482 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

groove

Figure 53. Body scale under the pectoral-fin of Trachurus japonicus (KMNH VRlOO, 102) (from same area as the scale with asterisk in Figure 52). Bar indicates I mm. differs on different body areas, their shapes are almost identical. Focus situated at center of scale and anterior field has two to four grooves or radii. Arrangement of ridges (circuli) on scale (Fig. 53) not concentric. In horizontal direction, ridges more densely spaced than those in vertical direction. Each ridge bent over on vertical midline of scale.

DISCUSSION

Remarks on Some Character States in the Family Carangidae Distributions of the newly described characters are: FINERIDGESONTHENEURAL ANDHEMALARCHES.Fine ridges were observed in following recent carangids: SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 483

Alectis cilia lis. Carangoides ferdau, Caranx sexfasciatus, Decapterus macarellus, D. macrosoma, D. muroadsi, D. punctatus, D. russelli, Elagatis bipinnulata, Hem- icaranx amblyrhynchus, Kaiwarinus equula, Naucrates ductor, Parastromateus niger, Pseudocaranx dentex, Scomberoides lysan, Selar crumenophthalmus, Ser- iola lalandi, S. quinqueradiata, Seriolina nigrofasciata, Trachinotus bail/onii, Trachurus capensis, T. declivis, T. japonicus, T. lathami, T. murphyi, T. novaezelandiae, Uraspis helvola, Chloroscombrus chrysurus (but weak); and the Mid- dle Miocene Scomberoides maruoi (Uyeno and Suda, 1991). Of the carangids studied, only Selaroides leptolepis did not exhibit such ridges. Phylogenetic relationships of the Carangidae have been discussed with and by: Bathyc\upeidae (Jordan, 1923), Coryphaenidae (Matsubara, 1963; Smith- Vaniz, 1984), Echeneididae (Smith-Vaniz, 1984), Lactariidae (Jordan, 1923; Matsubara, 1955), Leiognathidae (Jordan, 1923; Matsubara, 1955, ] 963; Deng and Zhan, 1986), Leptobramidae (Tominaga, 1965), Menidae (Jordan, 1923; Matsubara, 1955, 1963; Deng and Zhan, 1986), Nematistiidae (Jordan, 1923; Smith-Vaniz, ]984), Pomatomidae (Jordan, 1923; Matsubara, 1963), Rachycentridae (Jordan, 1923; Matsubara, 1955, ]963; Smith-Vaniz, 1984; Deng and Zhan, 1986), Lu- varoidei (Matsubara, 1963), Scombroidei (Starks, 1911; Matsubara, ] 963), and Stromateoidei (Matsubara, 1963). Of these mentioned families and suborders, only Coryphaenidae, Echeneididae, Lactariidae, Leiognathidae, Leptobramidae, Men- idae, Scombroidei, and Stromateoidei have been examined by the author, and found to have the fine ridges. Species that were examined and found to have such ridges in Percoidei similar to those of Trachurus are: Coryphaenidae, Coryphaena hippurus; Echeneididae, Echeneis naucrates; Lactariidae, Lactarius lactarius; Leptobramidae, Leptobrama mulleri; Leiognathidae, Gazza sp.; Menidae, Mene maculata. Somewhat different forms of ridges were found to be present in these Percoidei: Sciaenidae, Argyrosomus argenteus; Bramidae, Brama japonica, Tar- achtichthys longipinnis-these Scombroidei: Scombridae, Rastrelliger kanagur- ata, Sarda orientalis, Scomberomorus niphonius, Thunnus thynnus-and these Stromateoidei: Centrolophidae, Hyperoglyphe japonica, Psenopsis anomala; Stro- mateidae Pampus argenteus. These forms differ from those of the Carangidae in the density of ridges (number of ridges per mm of width of the lateral basal part of the neural spine) and the degree of expression (Fig. 54). The density in the Carangidae is higher than the stromateoidei ranging from 9 to 37.8 in the former and 2.7 to 12 in the latter. With the Scombroidei, however, the range (11.7 to 21.7) falls within that of the Carangidae, but the expression of the ridges is so weak that at first the surfaces look almost smooth. Of the eight species of Sciaen- idae examined, only Argyrosomus argenteus was found to possess ridges, however, the juncture of the ridges from both anterior and posterior arches is situated more dorsally than in the Carangidae. In the Bramidae, the density of the ridges is less than the Carangidae (6.8-10). BRANCHESOF THE LATERALLINE CANAL.The tribe Carangini (Smith-Vaniz, 1984) is characterized by having the posterior part of the lateral line scales mod- ified into scutes. In Trachurus japonicus, the latera] line canal has dorsal and ventral branches from central canal on the medial side of each scute. Kimura and Suzuki (1981) have shown in their figure that the carangine genus Decapterus also has a branched lateral line canal on the scutes, but did not mentioned this character in the body of the paper. Other carangid tribes examined (Naucratini: Seriola quinqueradiata, Seriolina nigrofasciata; Trachinotini: Trachinotus baillonii; Scomberoidini: Scomberoides lysan), exhibited straight lateral lines without branching. The Coryphaenidae (Coryphaena hippurus), one of the out-groups of the Carangidae (Smith- Vaniz, 1984), also has a non-branching lateral line. 484 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2. 1996

11 F ; : , !

Figure 54. Left lateral views of the fine ridges of the neural spine on the first caudal vertebrae, A, Trachurus japonicus (KMNH VRIOO, 102); B, Argyrosomus argentatus (NSM-PO 337); C, Brama japonica (NSM-PO 438); D, Rastrelliger kanagurta (NSM-PO 439); E, Thunnus thynnus (NSM-PO 22); F, Psenopsis anomala (NSM-PO 658). Bar indicate 1 mm.

CONNECTIONBETWEENTHE FIRST HEMAL SPINE AND FIRST ANAL PROXIMALRA- DIAL. In Trachurus japonieus (tribe Carangini), the first hemal spine and first anal proximal radial are strongly developed and closely approximates each other. The vertebral column and the combined hemal spine and proximal radial (when viewed laterally) form the· shape of the letter 1: This condition is also found in other tribes (Trachinotini and Scomberoidini), Development and connection of these parts are weak in the tribe Naucratini (Seriola quinqueradiata, Nauerates duetor, Seriolina nigrofasciata). They are also weak in the out-groups Nematis- tiidae (Rosenblatt and Bell, 1976) and Coryphaenidae. PRESENCE OF THE SUBZYGAPOPHYSIS.The anterior neural subzygapophysis is present below the anterior neural zygapophysis in Traehurus japonieus. All members of the Carangidae and the out-group Coryphaenidae have this character. LATERAL KEEL ON THE FOURTH PREURAL VERTEBRA. The anterior part of the lateral face of the 4th preural vertebra (= 11th caudal vertebra) is well keeled in SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 485

Trachurus japonicus. Other members of the Carangini exhibit the same character. Of the Naucratini examined, only Seriola minimally exhibits a lateral keel, none being present in other members (Elagatis bipinnulata, Naucrates ductor, Seriolina nigrofasciata); and none in Trachinotini (Trachinotus baillonii). The condition in Scomberoides lysan, a Scomberoidini, resembles that of Seriola. It is noteworthy that the lateral keel is present only on the 4th preural vertebra regardless of the total number of vertebrae. ARRANGEMENTOF THE RIDGES ON THE SCALES. In Trachurus japonicus, arrangement of the ridges on the scale is not in a concentric configuration. Spaces between ridges are dense in the horizontal direction and coarse in vertical direction. Each ridge is bent over on the vertical midline of the scale. Other members of Carangini (Caranx ignobilis, Decapterus muroadsi, Hemi- caranx amblyrhynchus, Kaiwarinus equula, Selar crumenophthalmus) exhibit the same character, whereas the Naucratini (Naucrates ductor, Seriola quinqueradiata), Trachinotini (Trachinotus baillonii), Scomberoidini (Scomberoides lysan) and the out-group Coryphaenidae (Coryphaena hippurus) do not have this feature.

ACKNOWLEDGMENTS I express my gratitude to Dr. T. Uyeno of the National Science Museum, Tokyo, Dr. W. F. Smith- Vaniz of Florida Museum of Natural History, University of Florida, and Dr. R. Winterbottom of the Royal Ontario Museum, Dr. B. B. Collette of the National Museum of Natural History, U.S. Depart- ment of Commerce and Dr. T. Potthoff of the Narragansett Laboratory, Northeast Fisheries Science Center, U.S. Department of Commerce for critical reading of the manuscript and valuable suggestions. For the loan or donation of specimens, I thank Dr. K. Matsuura of the National Science Museum, Tokyo, Dr. M. Shimizu of the University of Tokyo, Dr. Y. Yabumoto of the Kitakyushu Museum of Natural History, Dr. J. R. Paxton and Ms. S. Reader of the Australian Museum, Dr. S. Monkolprasit of Kasetsart University, Thailand, and Drs. U. Yamada and M. Tokimura of the Seikai Regional Fisheries Research Institute, Fishery Agency of Japan. Mr. N. M. Teitler of the Tokyo University of Fisheries kindly corrected the English of the manuscript.

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Gushiken, S. 1983. Revision of the carangid fishes of Japan. Ga]axea 2: 135-264. ---. 1988. Phylogenetic relationships of the perciform genera of the family Carangidae. Jpn. J. Ichthyol. 34: 443--461. Harder, W. 1975. Anatomy of fishes, part I and 2. E. Schweizerbart'sche Verlagsbuchhandlung, part I, xii + 612 p., part 2, 132 p. Hosoya, K. and K. Kawamura. 1993. Skeletal structure of the branchia] arches in Paralichthys oli- vaceus (Temminck et Schlegel). Bu]1. Natl. Res. Inst. Aquaculture, No. 22: ]-10. Iwai, T., I. Nakamura and K. Matsubara. 1965. Taxonomic study of the tunas. Misaki Mar. BioI. Inst., Kyoto Univ. Spec. Rep. No.2: 51 p. (in Japanese) Johnson, G. D. 1975. The procurrent spur: an undescribed perciform caudal character and its phylogenetic implications. Occ. Pap. Calif. Acad. Sci. No. ]21: 1-23. ---. 1986. Scombroid phylogeny: an alternative hypothesis. Bu]1. Mar. Sci. 39: ]--41. Jollie, M. 1973. Chordate morphology. Robert E. Krieger Publishing Company, xiv + 478 p. Jordan, D. S. ]923. A classification of fishes including families and genera as far as known. Stanford Univ. Publ., Univ. Ser., BioI. Sci. 3: 77-243. Katayama, M. 1959. Studies of the serranid fishes of Japan I. Bull. Fac. Edu. Yamaguchi Univ. 8: 103-181. Kendall, A. W. Jr. ]976. Predorsal and associated bones in serranid and granunistid fishes. Bull. Mar. Sci. 26: 585-592. Kimura, S. and K. Suzuki. ]98]. Taxonomica] consideration on Japanese carangid fishes of the genus Decapterus Bleeker. Bull. Fac. Fish., Mie Univ. No.8: 1-9. Lauder, G. V. and K. E Liem. 1983. The evolution and interrelationships of the actinopterygian fishes. Bull. Mus. Compo Zool. 159: 85-197. Mabee, PM. ]988. Supraneural and predorsa] bones in fishes: Development and homologies. Copeia 1988: 827-838. Matsubara, K. ]955. Fish morphology and hierarchy, part 1 and 2. Ishizaki Shoten, Tokyo. 1605 p. (in Japanese) ---. ]963. Pisces, Vertebrata (Ib), Anima] phylogenetic systematics 9. Nakayama Shoten, Tokyo. 5 + 197-531 p. (in Japanese) McAllister, D. E. 1968. Evolution of branchials and associated opercular, gular, and hyoid bones and the classification of teleostome fishes, living and fossils. Bull. Nat. Mus. Can. 22] (BioI. Ser. No. 77): 1-239. Oshima, M. 1925. A review of the carangoid fishes found in the waters of FornlOsa. Philip. J. Sci. 26: 345--413. Patterson, C. 1964. A review of Mesozoic acanthopterygian fishes, with special reference to those of the English Chalk. Phil. Trans. Roy. Soc. London, Ser. B, (739, 247): 2] 3--482. Phillips, J. B. 1942. Osteology of the sardine (Sardinops caerulea). J. Morph. 70: 463-500. Potthoff, T. and S. Kelley. 1982. Development of the vertebral column, fins and fin supports, bran- chiostega] rays, and squamation in the swordfish, Xiphias gladius. Fish. Bull., U.S. 80: 161-186. --- and J. A. Tellock. 1993. Osteological development of the snook, Celllropomus undecimalis (Teleostei, Centropomidae). Bull. Mar. Sci. 52: 669-7]6. ---, S. Kelly and L. A. Collins. 1988. Osteological development of the red snapper, Lwjanus campechanus (Lutjanidae). Bull. Mar. Sci. 43: 1--40. ---, ---, V. Saksena, M. Moe and E Young. 1987. Description of larval and juvenile yellow- tail damsel fish, Microspathodon chrysurus, Pomacentridae, and their osteological development. Bull. Mar. Sci. 40: 330-375. Rosen, D. E. 1973. Interrelationships of higher euteleostean fishes. Pages 397-513 In P. H. Green- wood, R. S. Miles and C. Patterson, eds. Interrelationships of fishes. Suppl. (I), Zool. J. Linn. Soc. 53, Supplement No. ]. Academic Press, London, New York. Rosenblatt, R. H. and M. A. Bell. 1976. Osteology and relationships of the roosterfish, Nematistius pectoralis Gill. Contrib. Sci., Natl. His!. Mus. Los Angeles County No. 279: 1-23. Smith-Vaniz, W. F. 1984. Carangidae: Relationships. Pages 522-530 in H. G. Moser, W. J. Richards, D. M. Cohen, M. P Fahay, A. W. Kendall, Jr. and S. L. Richardson, eds. Ontogeny and systematics of fishes. Spec. PubI. I. Am. Soc. Ichthyol. Herpetol. Smith, C. L. and R. M. Bai]ey. 1961. Evolution of the dorsal fin supports of percoid fishes. Michigan Acad. Sci. Arts. Letters 46: 345-363. Starck, D. 1979. Vergleichende Anatomie der Wirbeltiere, 2. Springer-Verlag, xii + 776 p. Starks, E. C. 1901. Synonomy of the fish skeleton. Proc. Wash. Acad. Sci. 3: 507-539. ---. 1911. The osteology and relationships of the fishes belonging to the family Carangidae. Leland Stanford Junior Univ., Publ., Univ. Ser. No.5: 27--49. Stiassny, M. L. J. 1986. The limits and relationships of the acanthomorph teleosts. J. Zool., Lond. (B) ]: 41 ]--460. Stokely, P S. ]952. The vertebral axis of two species of centrarchid fishes. Copeia 1952: 255-261. SUDA:OSTEOLOGYANDMYOLOGYOFTRACHURUS JAPON1CUS 487

Suda, y. 1985. Revision and intraspecific variations of Japanese jack mackerel, genus Trachurus Rafinesque. Dissertation, University of Tokyo, iii + 269 p. (in Japanese) ---. 1991. Skeletal features of the Japanese horse mackerel. Bull. Kitakyushu Mus. Nat. Hist. No. 10: 53-89. (in Japanese) ---, M. Shimizu and Y. Nose. 1987a. Morphological differences between cultivated and wild jack mackerel Trachurus japonicus. Nippon Suisan Gakkaishi 53: 59-61. ---, ---and ---. 1987b. Morphological variations of the Japanese jack mackerel Tra- churus japonicus. Nippon Suisan Gakkaishi 53: 1913-1919. ---, --- and ---. 1987c. Two types of Japanese jack mackerel Trachurus japonicus of summer spawning group in the Sagami Bay region. Nippon Suisan Gakkaishi 53: 1921-1923. Suzuki, K. 1962. Anatomical and taxonomical studies on the carangid fishes of Japan. Rep. Fac. Fish. Pref. Univ. Mie 4: 43-232. Tominaga, Y. 1965. The internal morphology and systematic position of Leptobrama mulleri, formely included in the family Pempheridae. Jpn. J. Ichthyol. 12: 33-56. ---. 1968. Internal morphology, mutual relationships and systematic position of the fishes belonging to the family Pempheridae. Jpn. J. Ichthyol. 15: 43-95. Travers, R. A. 1981. The interarcual cartilage; a review of its development, distribution and value as an indicator of phyletic relationships in euteleostean fishes. J. Nat. Hist. 15: 853-871. Uyeno, T 1975. Pisces. Pages 181-242 in T Shikama, ed. Paleontology III, new edition. Asakura Shoten, Tokyo. (in Japanese) --- and Y. Suda. 1991. A new Miocene queenfish of the genus Scomberoides (Pisces, Carangidae) from Tottori, Japan. Bull. Natn. Sci. Mus., Tokyo Ser. C 17: 41--48. Vergara, R. R. 1974. Estudio fenetico de los carangidos Cubanos (Pisces, Teleostei). Academia de Ciencias de Cuba Instituto de Oceanologia, Ser. Oceanogr. No. 23: 1-30. Wake, M. H. 1979. Hyman's comparative vertebrate anatomy, 3rd ed. The University of Chicago Press, Chicago. xi + 788 pp. Wakiya, Y. 1924. The carangoid fishes of Japan. Ann. Carnegie Mus. 15: 139-293. Weitzman, S. H. 1962. The osteology of Brycon meeki, a generalized characid fish, with an osteological definition of the family. Stanford Ichthyol. Bull. 8: 1-77. Whitehead, P. J. P. and G. G. Teugels. 1985. The West African pygmy herring Sierralhrissa leonensis: general features, visceral anatomy, and osteology. Am. Mus. Novit. No. 2835: 1-44. Winterbottom, R. 1974. A descriptive synonymy of the striated muscles of the Teleostei. Proc. Acad. Natr. Sci. Philadelphia 125: 225-317.

DATEACCEPTED:September 19, 1994.

ADDRESS:Laboratory of Fisheries Management, National Fisheries University, 2-7-1 Nagata-hon- machi, Shimonoseki-shi, 759-65 Japan.

ApPENDIX 1 Abbreviations Used in Text and Figures

V posterior palato-vomerine ligament VIl maxillo-rostroid ligament IX anterior maxillo-premaxillary ligament XII palato-premaxillary ligament Al section Al of adductor mandibulae A20t section A20t of adductor mandibulae A2~ section A213of adductor mandibulae ABP abductor profundus ABPP abductor profundus pelvicus ABS abductor superficialis ABSP abductor superficialis pelvicus AC anterior ceratohyal acn anterior condyle of hyomandibular for neurocranium AD adductor ADP adductor profundus ADPP adductor profundus pelvicus ADR adductor radialis ADS adductor superficialis ADSP adductor superficialis pelvicus af articulation fovea 488 BULLETIN OF MARINE SCIENCE, VOL 58, NO.2. 1996

afpt anterior foramen of posttemporal afscl anterior foramen of supracleithrum ahz anterior hemal zygapophysis ANG angular ansz anterior neural subzygapophysis anz anterior neural zygapophysis aocc anterior opening of cavum cranii aom anterior opening of myodome ap adductor process apd arch pedicel apdr anterior projection of distal radial ARRD arrector dorsalis ARRDP arrector dorsalis pelvicus ARRV arrector ventralis ARRV' subdivision of the arrector ventralis ARRVP arrector ventralis pelvicus AS anal-fin spine asp ascending process AV abdominal vertebra AW section AW of adductor mandibulae ax anterior xiphoid process ba basapophysis BAB basi branchial BAH basihyal BAO basioccipital BASP basisphenoid bf beryciform foramen BL Baudelot's ligament BRA branchiostegal bsp basisphenoid peduncle bsw basisphenoid wing CBR ceratobranchial ce articulation condyle for ectopterygoid cI articulation condyle for lachrymal CLE c1eithrum cp coronoid process CPHPU post-hemal spine cartilage CPHY post -hypural cartilage CRA coracoid CRR coracoradialis CV caudal vertebra cw coronoid wing da dorsal arm of the posttemporal dar dilatator area DEN dentary denf dentary fossa DEPRA depressores an ales DEPRD depressores dorsales df dorsal foramen for hyomandibular trunk of facial nerve DHYPH dorsal hypohyal dilp dilatator process DO dilatator operculi dof dorsal obturated foramen dorf dorsal fenestra dp process for insertion of depressor muscle DPCL dorsal element of postcleithrum dpp dorsal process of pelvic girdle DPR distal pectoral radial dpss dorsal preopercle-symplectic interosseous space dptf dorsal facet for posttemporal DR distal radial dr dental recess DS dorsal-fin spine SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 489 dsclf dorsal facet for suprac1eithrum DST dorsal element of the supratemporal-intertemporal EC ethmoid cartilage ec exoccipital condyle ECP ectopterygoid ENP endopterygoid ep process for insertion of erector muscle EPAX epaxialis EPB epibranchial EPH epihyal EPO epiotic EPU epural ERBL external pleural rib-basapophysis ligament ERECA erectores anales ERECD erectores dorsales ETH LAT lateral ethmoid EXO exoccipital fach articulation facet for anterior condyle of hyomandibular FD flexor dorsalis fdh foramen of dorsal hypohyal FDS flexor dorsalis superior fe articulation facet for epihyal ff facial foramen fica internal carotid foramen fm foramen magnum fmd articulation facet for mandibule fme articulation facet for mesethmoid fmsc mandibule sensory canal foramen fnc articulation facet for neurocranium fng foramen for nervus glossopharyngeus fnv foramen for nervus vagus fon foramen for olfactory nerve fp articulation facet for palatine fpch articulation facet for posterior condyle of hyomandibular fpcr frontal-parietal crest fpmx articulation facet for premaxilla fppl attaching furrow for palato-premaxillary ligament fr articulation fovea for rib fr' articulation fovea for epipleural FRO frontal frocr frontal crest FV flexor ventralis FVE flexor ventralis externus FVI flexor ventralis inferior gha groove for hyoidean artery h hook-shaped prolongation ha hemal arch HAB hyohyoidei abductores HAD hyohyoidei adductores heme hemal canal hp hyoid process hpu hemal spine of pre ural centrum hs hemal spine HYBR hypobranchial HYL hypochordal longitudinalis HYO hyomandibular HYU hypural hyp hypurapophysis HYP hypaxial is IC interarcual cartilage ICARA infracarinalis anterior ICARM infracarinalis medius ICARP infracarinalis posterior 490 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

IFO infraorbital ifos infraorbital shelf IL interzygapophysis ligament IMD intermandibularis INCA intercalar INCLA inclinatores anales INCLD inclinatores dorsales INO interopercle INT interradial is 102 ramus mandibularis trigeminus 103 ramus maxillaris inferioris IRBL internal pleural rib-basapophysis ligament ISL interspinal ligament ivf inferior vertebral foramen LAMO lamina orbitonasalis LAP levator arcus palatini LATS lateral is superficialis lep lateral ethmoid process LEXT levator externus Ifc lateral flange Ifr lateral facet rim LINT levator internus Ik lateral keel lIc lateral commissure LO levator operculi LP lower pharyngeal LPECT levator pectoralis LPOST levator posterior Ir lateral ridge Iw lateral wing MAX maxilla MC Meckelian cartilage MESETH mesethmoid MET metapterygoid mfc medial flange mpp meta pterygoid process mptf mesial facet for posttemporal Mr Meckelian ridge msclf mesial facet for supracleithrum mxp maxillary process na neural arch NAS nasal nc neurocranial condyle neuc neural canal nf neural foramen NL nuchal ligament nlf neurocranial lateral-line foramen npu neural spine of preural centrum ns neural spine OBLD obliquus dorsalis OBLP obliquus posterior OBLV obliquus ventralis oc opercular condyle of olfactory fossa OPE opercle P pelvis PAL palatine PARA parasphenoid PARR parhypural PARI parietal PB pharyngobranchial PC posterior ceratohyal pcn posterior condyle of hyomandibular for neurocranium SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPONICUS 491

PCR principal caudal-fin ray pfpt posterior foramen of posttemporal pfscl posterior foramen of supracleithrum PHCE pharyngoclavicularis externus PHCI pharyngoclavicularis internus phhl attaching process for dorsal hypohyal-hypobranchial ligament phz posterior hemal zygapophysis pipl process for the interopercle-posterior ceratohyal ligament PL primordial ligament plb attaching process for ligament from branchial arch plfr attaching process for ligament of fin ray pip articulation protuberance for lower limb of posttemporal plpr attaching process for ligament of proximal radial plw palatinal wing pm articulation process for maxilla pmp postmaxillary process pmw premaxillary wing pnz posterior neural zygapophysis porn posterior opening of myodome pop postorbital process ppJ attaching process for primordial ligament PPR proximal pectoral radial ppvl attaching process for posterior palato-vomerine ligament PR proximal radial pr primordial ridge PREM premaxilla PRENAS prenasal PREO preopercle PREY prevomer PRHY protractor hyoidei prmp primordial process PRO prootic PROC procurrent ray ps pterotic spine PT posttemporal PTE pterosphenoid PTO pterotic ptocr pterotic crest PU preural vertebra px posterior xiphoid process QUA quadrate RC rostral carti lage RCOMM rectus communis ROORS retractor dorsalis RECTY rectus ventralis RET retroarticular RMEF ramus mandibularis externus facialis rpl attaching ridge for primordial ligament saf suspensorial articulation facet sas suspensorium articulation socket sbk subpelvic keel sBL attaching surface for Baudelot's ligament SC scapula SCARP supracarinalis posterior SCL supracleithrum se spur of epihyal serc serrate crest on prevomer sf scapula foramen SPH sphenotic SPHO sphincter oesophagi spk suprapelvic keel spocr supraoccipital crest sqg symplectic-quadrate groove 492 BULLETIN OF MARINE SCIENCE, VOL. 58, NO.2, 1996

sr suspensorium ridge STH stemohyoideus SUBOP subopercle sulp articulation surface for upper limb of posttempora] SUPM supramaxilla SUPN supraneura] SUPO supraoccipital sV attaching surface for posterior pa]ato-vomerine ligament SYM symplectic t osseous tubercle TOA transversus dorsalis anterior TOP transversus dorsalis posterior tf trigeminal foramen tp tooth plate group trp transverse process TRV transversus ventralis ub unbranched principal caudal-fin ray unc uncinate process UP upper pharyngeal URN uroneural URS urostyle va ventral arm of posttempora] vc vertebral concavity venf ventral fenestra vf ventral foramen for hyomandibular trunk of facial nerve VHYPH ventral hypohyal vof ventral obturated foramen vp ventral process of coracoid VPCL ventral element of postcleithrum vpss ventral preopercle-symplectic interosseous space vptf ventral facet for posttemporal vsclf ventral facet for supracleithrum VST ventral element of the supratemporal-intertemporal w parasphenoid wing

ApPENDIX 2

Comparative Skeletal Materials

Cypriniformes: Catostomus commersoni commersoni (NSM-PO 312), Carassius auratus (NSM-PO 293), Hemibarbus barbus (NSM-PO 305), Beloniformes: Tylosurus crocodilus crocodilus (NSM-PO 77), Cypselurus atrisignis (NSM-PO 46), Hyporamphus sajori (NSM-PO 48), Gadiformes: Gadus macrocephalus (NSM-PO 138), Beryciformes: Beryx splendens (NSM-PO 181,273), Zeiformes: Zeus faber (NSM-PO 44), Percoidei Carangidae: Alectis ciliaris (uncat,), Carangoides ferdau (NSM-PO 353), Caranx sexfasciatus (NSM-PO 172; uncal.), Caranx sp. (uncat.), Chloroscombrus chrysurus (SKSK 11500), Decap- terus macarellus (NSMT-P 41222; uncal.), D. macrosoma (uncal.), D. muroadsi (uncal.), D. punctatus (NSMT-P 4]213), D. russelli (uncal.), Elagatis bipinnulata (NSM-PO 184), Hemicaranx amblyrhynchus (NSMT-P 41134), Kaiwarinus equula (uncat.), Naucrates ductor (uncal.), Parastro- mateus niger (uncal.), Pseudocaranx dentex (NSM-PO 434; uncal.), Scomberoides lysan (NSM-PO 164, 202), S, maruoi (NSM-PV 19631), Selar crumenophthalmus (NSMT-P 41326; uncal.), Sela- roides leptolepis (uncat.), Seriola lalandi (NSM-PO 173, 239), S. quinqueradiata (uncal.), Seriolina nigrofasciata (NSMT-P 50353), Trachinotus baillonii (uncal.), Trachurus capensis (NSMT-P 50377; uncal.), T. declivis (NSMT-P 50368; uncal.), T. lathami (NSMT-P 50519), T. murphyi (uncat.), T, novaezelandiae (NSMT-P 50375; uncal.), Uraspis helvola (NSM-PO 739; uncat.); Centropomidae: Psammoperca waigiensis (NSM-PO 129); Cheilodactylidae: Gonistius zonatus (NSM-PO III, 115); Cichlidae: Tilapia mossambica (NSM-PO 435); Coryphaenidae: Coryphaena hippurus (NSM-PO 118; uncal.); Echeneididae: Echeneis naucrates (NSMT-P 11627); Emmelichthyidae: Plagiogenion rubiginosus (NSM-PO 276); Ephippididae: Drepane punctata (NSM-PO 157); Gerreidae: Gerres oyena (uncal.), G. sp. (NSM-PO 104); Lactariidae: Lactarius lactarius (uncal.); Leiognathidae: Leiognathus nuchalis (NSMT-P 19220), Secutor ruconius (NSMT-P 30600); Leptobramidae: Lep- SUDA: OSTEOLOGY AND MYOLOGY OF TRACHURUS JAPON/CUS 493

tobrama mulleri (Australian Museum Catalogue Number I. 15557-188); Lobotidae: Lobores suri- namensis (NSM-PO 107); Lutjanidae: Aprion virescens (NSM-PO 277), Etelis coruscans (NSM- PO 109); Menidae: Mene maculata (NSMT-P 31434); Mullidae: Parupeneus indicus (NSM-PO 105, 106); Nemipteridae: Nemipterus virgatus (NSM-PO 108); Pentacerotidae: Histiopterus typus (NSM- PO 160); Percichthyidae: Doederleinia berycoides (NSM-PO 116), Lateolabrax japonicus (NSM- PO 189, 190), Stereolepis doederleini (NSM-PO 159); Pomacanthidae: Pomacanthus maculosus (NSM-PO 128); Pomadasyidae: Hapalogenys nigripinnis (NSM-PO 274), Parapristipoma trilinea- tus (NSM-PO 279), Pomadasys hasta (NSM-PO 124); Priacanthidae: Priacanthus hamrur (NSM- PO 281), P. sp. (NSM-PO 103); Sciaenidae: Argyrosomus argentatus (NSM-PO 337), Equetus lanceolatus (NSMT-P 40581), Larimus breviceps (NSMT-P 40590), Menticirrhus americanus (NSMT-P 40586), Paralonchurus elegans (NSMT-P 40610), Pareques acuminatus (NSMT-P 40582), Stellifer rastrifer (NSMT-P 40614,40615); Scombropidae: Scombrops boops (SKSK 15-5- 20); Serranidae: Epinephelus microdon (NSM-PO 112), Plectropomus leopardus (NSM-PO 123), Variola louti (NSM-PO 114); Sparidae: Pagrus major (NSM-PO 113). Labroidei: Cheilinus undulatus (NSM-PO 247), Scarns gibbus (NSM-PO 254). Scombroidei: Auxis thazard (NSM-PO 182), Euthynnus affinis (NSM-PO 38), Gymnosarda unicolor (NSM-PO 31), Katsuwonus pelamis (NSM-PO 32), Rastrelliger kanagurta (NSM-PO 439), Sarda orientalis (NSM-PO 28), Scomber japonicus (NSM-PO 176), Scomberomorus niphonius (NSM-PO 41), Thunnus albacares (NSM-PO 36), T. thynnus (NSM-PO 22). Stromateoidei: Cubiceps caeruleus (NSMT-P 41935), Hyperoglyphe japonica (NSM-PO 334), Psen- apsis anomala (NSM-PO 246, 658), Pampus argenteus (uncat.). Scorpaeniformes: Hexagrammos lagocephalus (NSM-PO 135), Sebastiscus marmoratus (NSM-PO 149). Pleuronectiformes: Limanda herzensteini (NSM-PO 336). Tetraodontifonnes: Odonus niger (NSM-PO 346), Stephanolepis cirrhifer (NSM-PO 343), Thamna- conus modestus (NSM-PO 340).

Osteology and Muscular Attachments of the Japanese Jack Mackerel, &lt;I

Osteology and Muscular Attachments of the Japanese Jack Mackerel, <I