Editorial Type:
Article Category: Article Commentary
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Online Publication Date: 20 May 2010

Osteology and Systematics of Parastromateus niger (Perciformes: Carangidae), with Comments on the Carangid Dorsal Gill-Arch Skeleton

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Page Range: 312 – 333
DOI: 10.1643/CI-09-118
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Abstract

The monotypic Indo-Pacific genus Parastromateus Bleeker, 1864 is morphologically peculiar among carangid fishes in its overall body form and has been difficult to place phylogenetically. This has been partly the result of a lack of detailed morphological data for this and other carangid fishes. Here, we describe and analyze the osteology of Parastromateus niger (Bloch, 1795), review its taxonomic history, and describe characters that are suggestive of its phylogenetic affinities. In addition, we made a broad survey of gill-arch skeletal characters for carangids. Previous studies have noted that a combination of features from the skeletal anatomy of Parastromateus firmly places it within the Carangidae (presence of a gap between the second and third anal-fin spine) and within the tribe Carangini (presence of scutes, inferior vertebral formina). Within the Carangini, however, its phylogenetic placement is less clear. In our survey of the gill arches of Carangidae, we found that only Parastromateus and Hemicaranx have an enlarged toothplate that bridges the epibranchial 4–ceratobranchial 4 joint. Additionally, these two genera have a superficially similar form of teeth on the pharyngeal toothplates (elongate and filamentous vs. conical and stout in other carangins). Although outside the scope of our study, these and other morphological aspects of Parastromateus need to be brought into a broad-based cladistic analysis of the Carangidae.

Copyright: 2010 by the American Society of Ichthyologists and Herpetologists
Fig. 1
Fig. 1

Parastromateus niger. (A) Fresh juvenile specimen (ASIZP 66167, 88 mm SL, Taiwan). (B) Fresh adult specimen (ASIZP 65880, 261 mm SL, Taiwan). (C) Cleared-and-stained larva (USNM 388226, 11 mm SL). (D) Cleared-and-stained skeleton (ANSP 62088, 100 mm SL). Anterior facing left. (A) and (B) from K. T. Shao. The Fish Database of Taiwan. WWW Web electronic publication, version 2005/5 http://fishdb.sinica.edu.tw.

Fig. 2
Fig. 2

Photographs of the skull roof and neurocranium of Parastromateus niger in (A) dorsal, (B) ventral, and (C) left lateral views (AMNH 98949, est. 185 mm SL). (D) Line drawing of skull roof and neurocranium in lateral view (ANSP 62088, 120 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Abbreviations: boc, basioccipital; bsp, basisphenoid; epo, epioccipital; exo, exoccipital; fr, frontal; fX, foramen for the vagus nerve; hyfa, anterior facet for hyomandibula; hyfp, posterior facet for hyomandibula; ic, intercalar; let, lateral ethmoid; met, mesethmoid; n, nasal; pa, parietal; pnc, prenasal canal bone; pro, prootic; pto, pterotic; pts, pterosphenoid; soc, supraoccipital; spo, sphenotic; v, vomer.

Fig. 3
Fig. 3

Infraorbital series of Parastromateus niger (ANSP 62088; 120 mm SL). (A) Left infraorbital bones in lateral view. (B) Medial view of io2 and io3 showing subocular shelf. Anterior facing left (in B, image flipped). Dashed line indicates course of sensory canals. Abbreviations: dsp, dermosphenotic; io, infraorbital; socs, subocular shelf.

Fig. 4
Fig. 4

Suspensorium, opercular bones, and oral jaws of Parastromateus niger (ANSP 62088, 120 mm SL). (A) Lateral view of suspensorium and opercular bones; anterior facing left. (B) Medial view of suspensorium and opercular bones; anterior facing right. (C) Lateral view of oral jaws; anterior facing left. (D) Medial view of oral jaws; anterior facing right. Bone in stipple, cartilage in black; dashed line indicates course of sensory canals; gray in (D) indicates portion of Meckel's cartilage inside dentary. Abbreviations: a, anguloarticular; cm, coronomeckelian; d, dentary; ecp, ectopterygoid; enp, endopterygoid; h, hyomandibula; iop, interopercle; mc, Meckel's cartilage; mpt, metapterygoid; mx, maxilla; op, opercle; pal, palatine; pmx, premaxilla; pop, preopercle; q, quadrate; rar, retroarticular; rc, rostral cartilage; smx, supramaxilla; sym, symplectic.

Fig. 5
Fig. 5

Portions of the ventral hyoid arch of Parastromateus niger (ANSP 62088, 120 mm SL). (A) Lateral and (B) medial views of the ventral hyoid arch, with the exception of the basihyal (see Figs. 6, 7). (C) Urohyal in lateral view. Anterior facing left in (A, C); anterior facing right in (B). Bone in stipple, cartilage in black. Abbreviations: br, branchiostegal; cha, anterior ceratohyal; chp, posterior ceratohyal; dhh, dorsal hypohyal; ih, interhyal; uh, urohyal; vhh, ventral hypohyal.

Fig. 6
Fig. 6

Gill arches of Parastromateus niger (ANSP 62088, 120 mm SL). (A) Dorsal gill arches from left side in dorsal view. (B) Dorsal portions of first gill arch in anterolateral view. (C) Ventral gill arches in ventral view. Anterior facing left. Bone in stipple, cartilage in black. Gill rakers omitted. Abbreviations: bb, basibranchial; bh, basihyal; cb, ceratobranchial; eb, epibranchial; eb1-m, medial cartilaginous division of eb1; hb, hypobranchial; iac, interarcual cartilage; pb, pharyngobranchial; tp-pb4, tooth plate of pb4; unp(a), anterior fork of uncinate process of eb1; unp(d), dorsal fork of uncinate process of eb1.

Fig. 7
Fig. 7

Cleared-and-stained specimens of Parastromateus niger (ANSP 62088, 120 mm SL). (A) Lateral view of skull, intact. (B) Lateral view of skull with jaws, suspensorium, and infraorbital bones removed to show position of gill arches in situ (right side, image reversed). (C) Ventral gill arches in dorsal view. (D) Dorsal gill arches in ventral view; arrow indicates enlarged tooth plate positioned between the anterior and posterior rows of gill rakers and bridging the eb4–cb4 joint. Anterior facing left. Bone stained red, cartilage stained blue.

Fig. 8
Fig. 8

Vertebrae of Parastromateus niger (ANSP 62088, 120 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Abbreviations: en, epineural; hs, haemal spine; ns, neural spine; r, rib.

Fig. 9
Fig. 9

Caudal skeleton of Parastromateus niger. (A) Photograph of cleared-and-stained larval specimen; bone stained red, cartilage stained blue (USNM 388226, 11 mm SL). (B) Line drawing of large juvenile specimen (ANSP 62088, 120 mm SL); bone in stipple, cartilage in black. Proximal portions of fin rays shown in light gray; arrows indicate the longest dorsal and ventral unbranched caudal-fin ray ( = dorsalmost and ventralmost principal caudal-fin rays). Anterior facing left. Abbreviations: dcr, distal caudal radial; ep, epural; hspu, haemal spine of preural centrum; hy, hypural; nspu, neural spine of preural centrum; phy, parhypural; pu, preural centrum; un, uroneural; ust, urostylar centrum.

Fig. 10
Fig. 10

Median fin supports of Parastromateus niger (ANSP 62088, 120 mm SL). (A) Supports of entire dorsal fin showing pattern of intercalation of proximal-middle radials and neural spines. Arrow indicates longest unbranched ( = first principal) dorsal-fin ray. (B) Close-up of articulation of spines with proximal-middle radials. (C) Close-up of articulation of dorsal-fin rays and proximal-middle radials. (D, E) Articulation of dorsal-fin ray and distal radial in anterior (D) and posterior (E) views. (F) Supports of entire anal fin showing pattern of intercalation of proximal-middle radials and haemal spines. Arrow indicates longest unbranched ( = first principal) anal-fin ray. (G) Close-up of articulation of anal-fin spines with proximal-middle radials. Anterior facing left in A–C, F, and G. Bone in stipple, cartilage in black, and proximal portions of fin rays shown in light gray. Abbreviations: afs, anal-fin spine; dfr, dorsal-fin ray; dfs, dorsal-fin spine; dr, distal radial; hs, haemal spine; ns, neural spine; pm, proximal-middle radial; sn, supraneural.

Fig. 11
Fig. 11

Pectoral and pelvic girdles of Parastromateus niger (ANSP 62088, 120 mm SL). (A) Lateral view; anterior facing left. (B) Medial view; anterior facing right. (C) Detail of pectoral radials and fin rays; anterior facing left; proximal portions of fin rays shown in light gray. Bone in stipple, cartilage in black; dashed line indicates course of sensory canals. Abbreviations: cl, cleithrum; co, coracoid; dpr, distal pectoral radials; pcl, postcleithrum; plvb, pelvic bone; pt, posttemporal; ra, pectoral radials ( = actinosts); sc, scapula; scl, supracleithrum.

Fig. 12
Fig. 12

Pelvic fin and girdle of Parastromateus niger in lateral view at two stages of development. (A) ANSP 150531 (73.6 mm SL). (B) ANSP 62088 (120 mm SL). Anterior facing left. Bone in stipple, cartilage in black; fin rays shown in light gray. Only left fin is shown and note that the hemilepidotrichia of each fin ray are disassociated from each other in B. Abbreviation: plvb, pelvic bone.

Fig. 13
Fig. 13

Scutes of Parastromateus niger (ANSP 62088, 120 mm SL). (A, B) Isolated scute from the middle of the series in external (A) and internal (B) views (image flipped). (C) Semi-schematic drawing of a series of four scutes showing the relationship of scutes with each other. The position of the external anterior pore of the lateral line canal has been omitted for clarity of the figure, but this opening receives the lateral line from the middle pore of the lateral line. Anterior facing left. Dashed line indicates course of the lateral line. Abbreviations: allp, anterior external lateral line pore of a scute; mllp, middle internal lateral line pore of scute; pllp, posterior external lateral line pores of scute.

Fig. 14
Fig. 14

Dorsal gill arches of selected non-carangid carangoids in dorsal view; from left side. (A) Nematistius pectoralis (ANSP 148654, est. 220 mm SL). (B) Rachycentron canadum (MCZ 157656, 157 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Gill rakers and toothplate of eb2 omitted. Abbreviations: acc, accessory cartilage; eb, epibranchial; eb1-m, medial cartilaginous division of epibranchial 1; iac, interarcual cartilage; pb, pharyngobranchial; tp-eb3, toothplate of eb3; tp-pb4, tooth plate of pb4; unp, uncinate process of eb1.

Fig. 15
Fig. 15

Dorsal gill arches of Scomberoidini in dorsal view; from left side. (A) Parona signata (ANSP 148655, 184 mm SL). (B) Oligoplites saurus (USNM 383079, 37 mm SL). (C) Scomberoides tala (USNM 36080, 135 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Gill rakers and toothplate of eb2 omitted. Abbreviations: acc, accessory cartilage; eb, epibranchial; eb1-m, medial cartilaginous division of eb1; iac, interarcual cartilage; pb, pharyngobranchial; tp-eb3, toothplate of eb3; tp-pb4, tooth plate of pb4; unp, uncinate process of eb1.

Fig. 16
Fig. 16

Dorsal gill arches of Trachinotini in dorsal view; from left side. (A) Trachinotus carolinus (USNM 383083, 51 mm SL). (B) Trachinotus ovatus (USNM 319754, 70 mm SL). (C) Lichia amia (USNM 383085, 78 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Gill rakers and toothplate of eb2 omitted. Abbreviations: acc, accessory cartilage; eb, epibranchial; eb1-m, medial cartilaginous division of eb1; iac, interarcual cartilage; pb, pharyngobranchial; tp-eb3, toothplate of eb3; tp-pb4, tooth plate of pb4; unp, uncinate process of eb1.

Fig. 17
Fig. 17

Dorsal gill arches of Naucratini in dorsal view; from left side. (A) Seriola fasciata (MCZ 157674, 109 mm SL). (B) Seriolina nigrofasciata (ANSP 140622, est. 132 mm SL); note that tp-eb3 is present but is hidden in this view. (C) Elagatis bipinulata (MCZ 163480, 79 mm SL). (D) Campogramma glaycos (ANSP 135406, 166 mm SL). (E) Naucrates ductor (MCZ 164201, 37 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Gill rakers and toothplate of eb2 omitted. Abbreviations: acc, accessory cartilage; eb, epibranchial; eb1-m, medial cartilaginous division of eb1; iac, interarcual cartilage; pb, pharyngobranchial; tp-eb3, toothplate of eb3; tp-pb4, tooth plate of pb4; unp, uncinate process of eb1.

Fig. 18
Fig. 18

Dorsal gill arches of Carangini in dorsal view; from left side unless otherwise noted. (A) Alectis alexandrinus (USNM 383089, 92 mm SL). (B) Alepes djedaba (USNM 347148, 110 mm SL); right gill arches, image reversed. (C) Atropus atropos (ANSP 144403, 95 mm SL). (D) Atule mate (USNM 325271, 110 mm SL). (E) Carangoides plagiotaenia (USNM 362714, 129 mm SL). (F) Caranx crysos (USNM 158866, 98 mm SL). (G) Chloroscombrus chrysurus (USNM 383077, 64 mm SL). (H) Decapterus punctatus (USNM 383086, 108 mm SL). (I) Gnathanodon speciosus (USNM 383087, 87 mm SL). Anterior facing left. Bone in stipple, cartilage in black. Gill rakers and toothplate of eb2 omitted. Abbreviations: acc, accessory cartilage; eb, epibranchial; eb1-m, medial cartilaginous division of eb1; iac, interarcual cartilage; pb, pharyngobranchial; tp-eb3, toothplate of eb3; tp-pb4, tooth plate of pb4; unp, uncinate process of eb1.

Fig. 19
Fig. 19

Dorsal gill arches of Carangini in dorsal view; from left side unless otherwise noted. (A) Hemicaranx amblyrhynchus (ANSP 139101a, 106 mm SL). (B) Megalaspis cordyla (USNM 347146, 130 mm SL). (C) Pantolabus radiatus (ANSP 147709, 122 mm SL). (D) Parastromateus niger (ANSP 62088, 120 mm SL). (E) Pseudocaranx dentex (USNM 222104, 127 mm SL). (F) Selar crumenophthalmus (MCZ 149687, 56 mm SL). (G) Selaroides leptolepis (MCZ 59266, 65 mm SL); right gill arches, image flipped. (H) Selene vomer (USNM 383081, 54 mm SL). (I) Trachurus mediterraneus (USNM 383084, 81 mm SL). (J) Ulua mentalis (USNM 347149, 100 mm SL). (K) Uraspis sp. (USNM 362716, 100 mm SL); right gill arches, image flipped. Anterior facing left. Bone in stipple, cartilage in black. Gill rakers and toothplate of eb2 omitted. Abbreviations: acc, accessory cartilage; eb, epibranchial; eb1-m, medial cartilaginous division of eb1; iac, interarcual cartilage; pb, pharyngobranchial; pb2-d, dorsal extension of pb2; tp-eb3, toothplate of eb3; tp-pb4, tooth plate of pb4; unp, uncinate process of eb1.

Fig. 20
Fig. 20

Left cb5 in dorsal view showing the pharyngeal dentition and elements of the left fourth branchial arch in medial view (images reversed) showing the toothplate bridging the eb4–cb4 joint in Hemicaranx and Parastromateus. (A) Atropus atropos (ANSP 159005, 91 mm SL). (B) Hemicaranx amblyrhynchus (ANSP 139101, 85 mm SL). (C) Parastromateus niger (ANSP 62088, 100 mm SL). Boxes indicate area shown in close-up photos in D–F. Close-up photos showing elements of the left fourth branchial arch in medial view (images reversed) in (D) Atropus, (E) Hemicaranx, and (F) Parastromateus. Anterior facing left. Bone stained red, cartilage stained blue. Arrows indicate eb4–cb4 toothplate in Hemicaranx and Parastromateus. Abbreviations: acc, accessory cartilage; cb, ceratobranchial; eb, epibranchial; tp-pb4, toothplate of pb4.

Contributor Notes

Associate Editor: D. Buth.

1 Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, Virginia 23062, E-mail: ehilton@vims.edu. Send reprint requests to this address.
2 Department of Zoology, Division of Fishes, Smithsonian Institution, P.O. Box 37012, National Museum of Natural History, MRC 0159, Washington, D.C. 20013-7012, E-mail: johnsond@si.edu.
3 Florida Museum of Natural History, Dickinson Hall, Museum Road, University of Florida, Gainesville, Florida 32611-7800, E-mail: smithvaniz@gmail.com.
Received: 24 Jun 2009
Accepted: 03 Feb 2010
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