Generic revision of the species formerly belonging to the genus Carangoides and its related genera (Carangiformes: Carangidae)

Abstract

The carangid genus Carangoides Bleeker 1851 has been considered to include 19 valid species, although the genus has been shown to be paraphyletic with molecular-phylogenetic methods. In this account, the previous Carangoides and its related genera (Alectis Rafinesque 1815, Atropus Oken 1817, Parastromateus Bleeker 1864, Selene Lacepède 1802, Ulua Jordan and Snyder 1908, and Uraspis Bleeker 1855) are reorganized into 15 (including five new and four resurrected) genera based on both molecular-phylogenetic results and morphological analyses. The members of the previously recognised Alectis are split into two genera, Alectis [Alectis ciliaris (Bloch 1787)] and Scyris Cuvier 1829 [Scyris alexandrina (Geoffroy St. Hilaire 1817) and Scyris indica Rüppell 1830]. Two new genera, Euprepocaranx and Paraselene are established for Carangoides dorsalis Gill 1863 [= Carangoides otrynter (Jordan and Gilbert 1883)] and Selene orstedii Lütken 1880, respectively. The genus Selene is valid, including six species except for Selene orstedii. Atropus atropos (Bloch and Schneider 1801), two species of Ulua and two species previously known in Carangoides [Carangoides armatus (Forsskål in Niebuhr 1775) and Carangoides hedlandensis (Whitley 1934)] composed the newly redefined Atropus. Craterognathus and Flavocaranx are new monotypic genera, consisting of the previous species Carangoides plagiotaenia Bleeker 1857 and Carangoides bajad (Fabricius in Niebuhr 1775), respectively. Platycaranx is also a new genus comprising three species, previous Carangoides chrysophrys (Cuvier in Cuvier and Valenciennes 1833), Carangoides malabaricus (Bloch and Schneider 1801) and Carangoides talamparoides Bleeker 1852. Resurrected genera, Carangichthys Bleeker 1853, Ferdauia Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927, and Turrum Whitley 1932 are composed of a few species previously assigned to Carangoides: Carangoides dinema Bleeker 1851, Carangoides humerosus (McCulloch 1915) and Carangoides oblongus (Cuvier in Cuvier and Valenciennes 1833); Carangoides ferdau (Fabricius in Niebuhr 1775) and Carangoides orthogrammus (Jordan and Gilbert 1882); and Carangoides coeruleopinnatus (Rüppell 1830), Carangoides fulvoguttatus (Forsskål in Niebuhr 1775) and Carangoides gymnostethus (Cuvier in Cuvier and Valenciennes 1833), respectively. Parastromateus and Uraspis continue to be valid genera, including Parastromateus niger (Bloch 1795), and Uraspis helvola (Forster in Bloch and Schneider 1801) and Uraspis uraspis (Günther 1860), respectively. The revised Carangoides consists of only two species, resurrected Carangoides ire (Cuvier in Cuvier and Valenciennes 1833) and Carangoides praeustus (Anonymous [Bennett] 1830). Diagnosis and brief description are provided for each genus.

Introduction

The members of the carangid genus Carangoides Bleeker 1851b are distributed widely in the tropical shallow waters of the Indo-Pacific and eastern Pacific (see Springer 1982). The genus includes 19 valid species (Fricke et al. 2021) that are all caught commercially and represent important food resources throughout the world. Although Bleeker (1851b) established the genus Carangoides, he neither gave a description of the genus nor designated the type species, only using this generic name for the following 13 nominal species: Carangoides praeustus (Anonymous [Bennett] 1830), Carangoides gymnostethoides Bleeker 1851b, Carangoides hemigymnostethus Bleeker 1851b, Carangoides coeruleopinnatus (Rüppel 1830), Carangoides oblongus (Cuvier in Cuvier and Valenciennes 1833), Carangoides dinema Bleeker 1851b, Carangoides chrysophryoides Bleeker 1851b, Carangoides talamparah Bleeker 1851a, Carangoides citula (Cuvier in Cuvier and Valenciennes 1833), Carangoides atropos (Bloch and Schneider 1801), Carangoides blepharis Bleeker 1851a and Carangoides gallichthys Bleeker 1851a. Subsequently Jordan and Evermann (1896) presented a brief description of Carangoides, and Jordan (1919) assigned Caranx praeustus Anonymous [Bennett] 1830 as the type species. According to Gushiken (1983), who completed a taxonomical study on the Japanese carangid fishes, the genus Carangoides was characterized by undeveloped adipose eyelids, cycloid scales along anterior straight part of lateral line, and anterior lobes of the second dorsal and anal fins sometimes elongate and falcate. Additionally, he stated that their body shapes were highly variable. Despite his statement, Carangoides praeustus, the type species of the genus, has a lower body than other species included in Carangoides and its second dorsal fin is not falcate, slightly higher than the first. Therefore, the morphology of Carangoides praeustus is considerably different from the other species of Carangoides.

Many previous authors have examined the morphology of carangids, with some proposing taxonomic changes. Suzuki (1962) studied the osteology of carangid fishes in detail, but he did not indicate interspecific differences in Carangoides. Smith-Vaniz (1984) presented phylogenetic relationships among the family Carangidae using morphological methods. Gushiken (1988) also showed the carangid phylogenetic relationships morphologically, and stated that Carangoides had no autapomorphies and was composed of some species groups (he described as subunits). Springer and Smith-Vaniz (2008) inspected the supraneural and pterygiophore insertion patterns in carangid fishes comprehensively, and stated that Ferdauia ferdau (Fabricius in Niebuhr 1775) [as Carangoides ferdau (Fabricius in Niebuhr 1775)] and Ferdauia orthogrammus (Jordan and Gilbert 1882a) [as Carangoides orthogrammus (Jordan and Gilbert 1882a)] were unique among Carangoides. However, no study has focused on the morphological differences among the species of Carangoides. Smith (1967) showed the naked thoracic areas of 11 species and indicated that the naked areas were considered diagnostic characters to Carangoides species.

With the onset of molecular phylogenetics, Kijima et al. (1986) constructed the phylogenetic tree of Carangidae using isozymes, and stated that Carangoides and Atropus Oken 1817 comprised a sister group and these two genera and Alectis Rafinesque 1815 showed a sister-group relationship. Reed et al. (2001, 2002) indicated that two species of Carangoides and two species of Alectis did not form their own monophyletic groups from their molecular analysis using the mitochondrial 16S ribosomal RNA (16S) locus. Persis et al. (2009) constructed a phylogenetic tree of the subfamily Caranginae based on 22 species, including three species of Carangoides, using the mitochondrial cytochrome c oxidase subunit I (COI) locus, and showed paraphyly in Carangoides. However, a detailed examination of relationships among the species belonging to Carangoides were not performed in Reed et al. (2001) or Persis et al. (2009). Santini and Carnevale (2014) constructed a time-calibrated phylogeny of Carangidae based on five mitochondrial and two nuclear loci, finding the non-monophyletic relationships of Carangoides species. Bogorodsky et al. (2017) reviewed Carangoides from the Red Sea and presented the molecular phylogeny of the species of Carangoides using the COI gene, suggesting that several species in the genus are composed of two species, e.g., Carangoides praeustus and Platycaranx chrysophrys (Cuvier in Cuvier and Valenciennes 1833) gen. nov. [as Carangoides chrysophrys (Cuvier in Cuvier and Valenciennes 1833)]. Recently, two studies using ultraconserved elements have also examined relationships among Carangidae. Glass (2019) found paraphyly of Carangoides and compare relationships within Carangidae using 18 species of Carangoides. Girard et al. (2020) examined the phylogenetic relationships among carangiform fishes and found paraphyly of Carangidae. However, they did not focus on the relationships among genera or species within Carangidae.

Using both morphological and molecular approaches, these studies on the phylogenetic relationships among the species belonging to Carangoides and its related genera have shown the non-monophyly of Carangoides multiple times. However, detailed phylogenetic relationships among the species of Carangoides have not been demonstrated. Here we construct the phylogenetic relationships among the species of the subfamily Caranginae to understand the phylogenetic positions of the species belonging to Carangoides, and relationships among the members of Carangoides and its related genera. Furthermore, we reconstruct them into 15 genera based on molecular data and present their morphological diagnoses, among them we establish five new genera and resurrect four genera.

Materials and methods

Materials for morphological examination (asterisk indicates cleaned and stained or skeletal specimens). Alectis Rafinesque 1815. Alectis ciliaris (Bloch 1787) (21 specimens): CAS-SU 11563, holotype of Hynnis hopkinsi Jordan and Starks in Jordan 1895, 548 mm SL (standard length), Sinaloa, Mexico; FRLM 11002, 11003, 23649, 25965, 25973, 28539, 35432, 36570, 37281, 39157, 40247, 40257, 41537, 43676, 44489, 54883, 60255*, 54–231 mm SL, Mie, Japan; RMNH.PISC. 26963, lectotype of Carangoides blepharis Bleeker 1851a, 272 mm SL, Jakarta, Indonesia; one of RMNH.PISC 6117, largest paralectotype of Carangoides blepharis, 275 mm SL, Jakarta, Indonesia; USNM 50874, holotype of Carangoides ajax Snyder 1904, 814 mm SL, Honolulu, USA. Atropus Oken 1817. Atropus armatus (Forsskål in Niebuhr 1775) [new combination of species name, formerly known as Carangoides armatus (Forsskål in Niebuhr 1775); nine specimens]: FRLM 32079, 200 mm SL, Phuket Thailand; FRLM 43850, 43862, 43863, 97–109 mm SL, Johor Strait, Malaysia; MNHN A-6181, paratype of Caranx citula Cuvier in Cuvier and Valenciennes 1833, 167 mm SL, Red Sea; MNHN A-6182, holotype of Caranx citula, 162 mm SL, Red Sea; MUFS 15237, 125 mm SL, Bangsaphan, Thailand; NSMT-P 60798, 34 mm SL, Hainan I., China; NSMT-P 71463, 115 mm SL, Phuket, Thailand. Atropus atropos (Bloch and Schneider 1801) (14 specimens): FRLM 8897, 90 mm SL, Japan; FRLM 32414*–32416, 71–131 mm SL, Bintan I., Indonesia; FRLM 33659, 33999, 34003, 63–108 mm SL, Selangor, Malaysia; FRLM 35453, 180 mm SL, Do Son, Vietnam; FRLM 44040–44042, 133–180 mm SL, Port Dickson, Malaysia; MNHN A-5508 (dry), A-6133 (dry), syntypes of Caranx nigripes Cuvier in Cuvier and Valenciennes 1833, 178–179 mm SL, Puducherry, India; MUFS 39862, 136 mm SL, Karachi, Pakistan. Atropus aurochs (Ogilby 1915) [new combination, formerly known as Ulua aurochus (Ogilby 1915); one specimen]: FRLM 14575, 134 mm SL, Ambon, Indonesia. Atropus hedlandensis (Whitley 1934) [new combination, formerly known as Carangoides hedlandensis (Whitley 1934); 32 specimens]: FRLM 2279, 2649, 3876*, 4960, 5627, 20819–20821, 24426, 24463, 27228, 29129, 31421–31424, 32640, 32641, 35541, 36660, 40542, 60251*, 69–173 mm SL, Mie, Japan; FRLM 20395, 20397*, 88–91 mm SL, Lombok, Indonesia; FRLM 24423, 159 mm SL, Miyazaki, Japan; FRLM 30723, 33731, 33821, 62–170 mm SL, Phuket, Thailand; FRLM 31475, 107 mm SL, Nha Trang, Vietnam; KPM-NI 25034, 130 mm SL, Kanagawa, Japan; MNHN A-5547, holotype of Caranx ciliaris Cuvier in Cuvier and Valenciennes 1833, ca. 94 mm SL, Puducherry, India; MNHN A-6102, paratype of Caranx citula (Cuvier in Cuvier and Valenciennes 1833), 158 mm SL, New Guinea. Atropus mentalis (Cuvier in Cuvier and Valenciennes 1833) [new combination, formerly known as Ulua mentalis (Cuvier in Cuvier and Valenciennes 1833); 24 specimens]: FRLM 14505, 16990, 35759, 35805, 35806, 54–173 mm SL, Ambon, Indonesia; FRLM 20388, 94 mm SL, Lombok, Indonesia; FRLM 35194, 102 mm SL, Songkhla, Thailand; FRLM 39998, 39999, 158–168 mm SL, Bitung, Indonesia; FRLM 43858–43861, 101–108 mm SL, Johor Strait, Malaysia; FRLM 44046, 104 mm SL, Port Dickson, Malaysia; MNHN B-2881, paratype of Carangoides citula (Cuvier in Cuvier and Valenciennes 1833), 201 mm SL, Red Sea; NSMT-P 71458, 129 mm SL, Phuket, Thailand; RMNH 6091 (7), syntypes and Bleeker specimens of Leioglossus carangoides Bleeker 1851b, 73–286 mm SL, Indonesia; RMNH 27642, syntype of Leioglossus carangoides, 183 mm SL, Indonesia. Carangichthys Bleeker 1853. Carangichthys dinema (Bleeker 1851b) (formerly known as Carangoides dinema Bleeker 1851b; 14 specimens): FRLM 3875, 29158, 34073, 35050, 35068, 35585, 36756, 39358, 43488, 94–348 mm SL, Mie, Japan; FRLM 38625, 162 mm SL, Okinawa, Japan; FRLM 40027, 40170, 124–128 mm SL, Bitung, Indonesia; NMW 60929, holotype of Caranx longipes Steindachner 1906, 59 mm SL, Samoa; RMNH.PISC. 26973, holotype of Carangichthys typus Bleeker 1853, 86 mm SL, Kema, Indonesia. Carangichthys humerosus (McCulloch 1915) [formerly known as Carangoides humerosus (McCulloch 1915); five specimens]: FRLM 21441, 21452, 21462, 21463, 111–126 mm SL, Ambon, Indonesia; FRLM 25724, 140 mm SL, Bitung, Indonesia. Carangichthys oblongus (Cuvier in Cuvier and Valenciennes 1833) [formerly known as Carangoides oblongus (Cuvier in Cuvier and Valenciennes 1833); 15 specimens]: FRLM 4221, 34080, 35051, 36444, 36695, 36898, 37150, 37263, 78–213 mm SL, Mie, Japan; FRLM 24411, 201 mm SL, Miyazaki Japan; FRLM 30637, 173 mm SL, Trang, Thailand; FRLM 34889, 40026, 40074, 106–156 mm SL, Bitung, Indonesia; MNHN A-6172, holotype of Carnax oblongus Cuvier in Cuvier and Valenciennes 1833, 261 mm SL, Vanikoro, Solomon Islands; USNM 51951, holotype of Caranx deani Jordan and Seale 1905b, 111 mm SL, Negros, Philippines. Carangoides Bleeker 1851b. Carangoides ire (Cuvier in Cuvier and Valenciennes 1833) (one specimen): MNHN 0000-5847, holotype of Caranx ire Cuvier in Cuvier and Valenciennes 1833, 131 mm SL, Puducherry, India. Carangoides praeustus (Anonymous [Bennett] 1830) (11 specimens): FRLM 25775*, 87 mm SL, Bitung, Indonesia; FRLM 31444*, 95 mm SL, Nha Trang, Vietnam; FRLM 32157, 32158, 135–159 mm SL, Phuket, Thailand; FRLM 35478, 35479, 129–138 mm SL, Ha Long Bay, Vietnam; KAUM–I. 12028, 168 mm SL, Sabah, Malaysia; KAUM–I. 23911, 103 mm SL, Samut Sakhon, Thailand; KAUM–I. 32903, 148 mm SL, Chachoengsao, Thailand; NSMT-P 65953, 69491, 33–123 mm SL, Thailand. Craterognathus gen. nov. Craterognathus plagiotaenia (Bleeker 1857) (formerly known as Carangoides plagiotaenia Bleeker 1857; 14 specimens): FRLM 8895, 265 mm SL, Japan; FRLM 21219, 344 mm SL, Parau; FRLM 24694, 26335, 39995–39997, 114–214 mm SL, Bitung, Indonesia; FRLM 37736, 279 mm SL, Okinawa, Japan; MUFS 18597, 140 mm SL, Makassar, Indonesia; RMNH.PISC. 6096 (3), paralectotypes of Carangoides plagiotaenia, 70–78 mm SL, Ambon, Indonesia; RMNH.PISC. 26974, lectotype of Carangoides plagiotaenia, 66 mm SL, Ambon, Indonesia; ZIN 2662, syntype of Caranx brevicarinatus Klunzinger 1871, 264 mm SL, Red Sea. Euprepocaranx gen. nov. Euprepocaranx dorsalis (Gill 1863c) [formerly known as Carangoides otrynter (Jordan and Gilbert 1883); seven specimens]: CAS-ICH 213450, 86 mm SL, Azuero Peninsula, Pacific Panama; USNM 4957 (2), syntypes of Carangoides dorsalis Gill 1863c, 183–190 mm SL, western coast of Central America; USNM 28153, 177 mm SL, Mazatlán, Pacific Mexico; USNM 362543, 314 mm SL, Panama City, Pacific Panama; USNM 382342, 141 mm SL, Buenaventura, Colombia; USNM 396266, 105 mm SL, Azuero Peninsula, Pacific Panama. Ferdauia Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927. Ferdauia ferdau (Fabricius in Niebuhr 1775) [new combination, formerly known as Carangoides ferdau (Fabricius in Niebuhr 1775); 21 specimens]: FRLM 3177, 12149, 14393, 14394, 25887, 25888, 34763, 36780, 36899, 37031, 41784, 42170, 133–233 mm SL, Mie, Japan; FRLM 26281, 145 mm SL, Bitung, Indonesia; FRLM 30716, 30720, 32095, 149–188 mm SL, Phuket, Thailand; FRLM 42761, 308 mm SL, Yoron I., Japan; FRLM 43330, 234 mm SL, Okinawa, Japan; RMNH.PISC. 6101, syntype of Carangoides hemigymnostethus Bleeker 1851b, 230 mm SL, Jakarta, Indonesia; RMNH.PISC. 26969, syntype or Bleeker specimen of Carangoides hemigymnostethus, 202 mm SL, Jakarta, Indonesia (or Timor); USNM 51729, holotype of Caranx gilberti Jordan and Seale 1906, 157 mm SL, American Samoa. Ferdauia orthogrammus (Jordan and Gilbert 1882a) [new combination, formerly known as Carangoides orthogrammus (Jordan and Gilbert 1882a); six specimens]: FRLM 28945, 369 mm SL, Okinawa, Japan; FRLM 36777, 36778, 255–292 mm SL, Wakayama, Japan; FRLM 40007, 226 mm SL, Bitung, Indonesia; FRLM 41754, 248 mm SL, Mie, Japan; SAIAB 1, holotype of Carangoides nitidus Smith in Smith 1972, 207 mm SL, Shimoni, Kenya. Flavocaranx gen. nov. Flavocaranx bajad (Fabricius in Niebuhr 1775) [formerly known as Carangoides bajad (Fabricius in Niebuhr 1775); six specimens]: HUMZ 165337, 183 mm SL, Red Sea, Saudi Arabia; MNHN A5587, A5752, syntypes of Caranx auroguttatus Cuvier in Cuvier and Valenciennes 1833, 326–372 mm SL, Red Sea (dry specimens); MUFS 33644, 125 mm SL, Mutrah, Oman; USNM 325262, 372 mm SL, Red Sea, Eritrea; USNM 403266, 207 mm SL, Dumaguete, Philippines. Paraselene gen. nov. Paraselene orstedii (Lütken 1880) (formerly known as Selene orstedii Lütken 1880; six specimens): CAS-ICH 80194, 87889, 74–76 mm SL, Pacific Panama; USNM 79971, 219 mm SL, Panama Bay, Pacific Panama (X-ray photo only); USNM 79975, 166 mm SL, Chame Point, Pacific Panama (X-ray photo only); ZMUC 461508, 461509, syntypes of Selene orstedii Lütken 1880, 136–155 mm SL, Puntarenas, Costa Rica. Parastromateus Bleeker 1864. Parastromateus niger (Bloch 1795) (seven specimens): FRLM 20399, 156 mm SL, Lombok, Indonesia; FRLM 32100, 98 mm SL, Phuket, Thailand; FRLM 32764, 63 mm SL, Iloilo, Philippines; FRLM 40277, 245 mm SL, Kagoshima, Japan; FRLM 44047, 44048, 87–217 mm SL, Port Dickson, Malaysia; USNM 55914, holotype of Citula halli Evermann and Seale 1907, 49 mm SL, Luzon, Philippines (photograph only). Platycaranx gen. nov. Platycaranx chrysophrys (Cuvier in Cuvier and Valenciennes 1833) [formerly known as Carangoides chrysophrys (Cuvier in Cuvier and Valenciennes 1833); 12 specimens]: FRLM 20396, 100 mm SL, Lombok, Indonesia; FRLM 35764, 126 mm SL, Ambon, Indonesia; FRLM 44037, 296 mm SL, Port Dickson, Malaysia; KAUM–I. 20701, 215 mm SL, Kagoshima, Japan; MNHN A-560, holotype of Caranx chrysophys Cuvier in Cuvier and Valenciennes 1833, 246 mm SL, Seychelles; NSMT-P 57169, 71460 (2), 123–137 mm SL, Phuket, Thailand; NSMT-P 60814, 95 mm SL, Hainan I., China; NSMT-P 69024, 69212, 123–142 mm SL, Nha Trang, Vietnam; RMNH 6113, syntype of Carangoides chrysophryoides Bleeker 1851b, 270 mm SL, Jakarta, Indonesia. Platycaranx malabaricus (Bloch and Schneider 1801) [formerly known as Carangoides malabaricus (Bloch and Schneider 1801); 18 specimens]: FRLM 9378, 165 mm SL, Japan; FRLM 13216, 13217, 146–159 mm SL, Samut Prakan, Thailand; FRLM 14574, 14577, 35762, 35763, 110–143 mm SL, Ambon, Indonesia; FRLM 32798, 59 mm SL, Iloilo, Philippines; KAUM–I. 12151, 12178, 22096, 94–132 mm SL, Sabah, Malaysia; KAUM–I. 17250, 116 mm SL, Terengganu, Malaysia; NSMT-P 57168, 111 mm SL, Phuket, Thailand; RMNH.PISC. 6098, lectotype of Carangoides talamparah Bleeker 1851a, 137 mm SL, Jakarta, Indonesia; RMNH.PISC. 25872 (largest specimen), paralectotype of Carangoides talamparah, 128 mm SL, Jakarta, Indonesia; RMNH.PISC. 25873 (3), paralectotypes of Carangoides talamparah, 78–176 mm SL, Jakarta, Indonesia. Platycaranx talamparoides (Bleeker 1852) (formerly known as Carangoides talamparoides Bleeker 1852; 11 specimens): FRLM 13219, 123 mm SL, Samut Prakan, Thailand; FRLM 25723, 40171, 97–142 mm SL, Bitung, Indonesia; FRLM 30724, 152 mm SL, Phuket, Thailand; FRLM 35192, 35193, 101–149 mm SL, Songkhla Thailand; FRLM 35657, 35697, 97–166 mm SL, Ambon, Indonesia; KAUM–I. 22095, 111 mm SL, Sabah, Malaysia; RMNH.PISC. 6099, lectotype of Carangoides talamparoides Bleeker 1852, 198 mm SL, Sibogha, Indonesia; RMNH.PISC. 26968, paralectotype of Carangoides talamparoides, 179 mm SL, Sibogha, Indonesia. Scyris Cuvier 1829. Scyris alexandrina (Geoffroy St. Hilaire 1817) [formerly known as Alectis alexandrina (Geoffroy St. Hilaire 1817); two specimens]: CAS-ICH 79955 (2), 154–167 mm SL, Cameroon. Scyris indica Rüppell 1830 [formerly known as Alectis indica (Rüppell 1830); 13 specimens]: FRLM 21659, 35702, 199–266 mm SL, Ambon, Indonesia; FRLM 24227, 267 mm SL, Okinawa, Japan; FRLM 25789, 84 mm SL, Bitung, Indonesia; FRLM 29957, 29990, 86–107 mm SL, Phuket, Thailand; FRLM 32763, 61 mm SL, Iloilo, Philippines; FRLM 35144, 157 mm SL, Songkhla, Thailand; FRLM 43904, 43908, 46–88 mm SL, Johor Strait, Malaysia; RMNH.PISC. 26962, lectotype of Carangoides gallichthys Bleeker 1851a, 118 mm SL, Indonesia; one of RMNH.PISC. 6118, largest paralectotype of Carangoides gallichthys, 414 mm SL, Indonesia; SMNS 10578, paralectotype of Carangoides gallichthys, 187 mm SL, Indonesia. Selene Lacepède 1802. Selene brevoortii (Gill 1863b) (six specimens): CAS-ICH 67914 (2), 184–187 mm SL, Nayarit, Pacific Mexico; CAS-ICH 90546, 132 mm SL, Sonora, Pacific Mexico; CAS-ICH 213465, 54 mm SL, Pacific Panama; USNM 4438, holotype of Argyriosus brevoortii Gill 1863b, 48 mm SL, Baja California, Mexico; USNM 39896, paratype of Argyreiosus pacificus Lockington 1877, 216 mm SL, Baja California, Mexico. Selene brownii (Cuvier 1816) (four specimens): CAS-ICH 39042 (2), 75–81 mm SL, off Laguna de Caratasca, Atlantic Honduras; CAS-SU 67160, 75 mm SL, La Habana, Cuba; USNM 37732, 222 mm SL, Jamaica. Selene dorsalis (Gill 1863a) (two specimens): CAS-ICH 214640, 120 mm SL, Sao Tome and Principe; SAM 13725, holotype of Selene gibbiceps Gilchrist 1914, 158 mm SL, Walvis Bay, Namibia. Selene peruviana (Guichenot 1866) (three specimens): CAS-ICH 213461, 54 mm SL, Azuero Peninsula, Pacific Panama; USNM 81773, holotype of Vomer declivifrons Meek and Hildebrand 1925, 151 mm SL, Panama City, Pacific Panama; USNM 368344, off Golfo de Fonseca, El Salvador (X-ray photo only). Selene setapinnis (Mitchill 1815) (16 specimens): AMNH I-7148, holotype of Vomer setapinnis cubensis Nichols 1918, 117 mm SL, Cuba; CAS-ICH 16370 (2), 35–41 mm SL, Rhode Island, USA; NSMT-P 120012–120021, 101–141 mm SL, off Suriname; NSMT-P 41713, 41726, 41727, 30–107 mm SL, off Suriname and French Guiana. Selene vomer (Linnaeus 1758) (four specimens): FRLM 49239, 52296, 54866, 189–242 mm SL, reared specimens; NSMT-P 65358, 192 mm SL, locality unknown. Turrum Whitley 1932. Turrum coeruleopinnatum (Rüppell 1830) [new combination, formerly known as Carangoides coeruleopinnatus (Rüppell 1830); 45 specimens]; CAS-SU 7283, paratype of Caranx formosanus Jordan and Snyder 1908, 179 mm SL, Kaohsiung, Taiwan; CAS-SU 9066, holotype of Caranx altissimus Jordan and Seale 1905a, 98 mm SL, Hong Kong, China; FRLM 2280, 14351, 23280, 29156, 39643, 39645, 39647, 43194, 51147, 43–156 mm SL, Mie, Japan; FRLM 13212–13215, 13218, 134–162 mm SL, Samut Prakan, Thailand; FRLM 14560, 21432, 21664, 35760, 35761, 64–260 mm SL, Ambon, Indonesia; FRLM 14653, 14757, 14758, 14778, 54–86 mm SL, Lombok, Indonesia; FRLM 25775–25777, 26259, 87–111 mm SL, Bitung, Indonesia; FRLM 28742, 32094, 107–161 mm SL, Phuket, Thailand; FRLM 32755, 57 mm SL, Panay I., Philippines; FRLM 35138–35142, 35154, 96–167 mm SL, Songkhla, Thailand; FRLM 44039, 133 mm SL, Port Dickson, Malaysia; KAUM–I. 16838, 96 mm SL, Terengganu, Malaysia; MUFS 18467, 18594, 18795, 76–97 mm SL, Makassar, Indonesia; RMNH.PISC. 6112, 26972, 54–66 mm SL, Ambon, Saparua or Manado, Indonesia. Turrum fulvoguttatum (Forsskål in Niebuhr 1775) [new combination, formerly known as Carangoides fulvoguttatus (Forsskål in Niebuhr 1775); nine specimens]: FRLM 37735, 45538, 356–625 mm SL, Okinawa, Japan; FRLM 40025, 165 mm SL, Bitung, Indonesia; MUFS 30182, 124 mm SL, Miyazaki, Japan; NMW 11627 (3), ?syntypes of Caranx bleekeri Klunzinger 1871, 159–187 mm SL, Red Sea; RMNH.PISC. 27127, syntype of Carangoides ophthalmotaenia Bleeker 1852, 66 mm SL, Ambon, Indonesia; ZIN 2544, syntype of Caranx bleekeri, 166 mm SL, Red Sea. Turrum gymnostethus (Cuvier in Cuvier and Valenciennes 1833) [new combination, formerly known as Carangoides gymnostethus (Cuvier in Cuvier and Valenciennes 1833); five specimens]: FRLM 31558, 121 mm SL, Nha Trang, Vietnam; FRLM 35143, 196 mm SL, Songkhla, Thailand; FRLM 39984, 39994, 159–185 mm SL, Bitung, Indonesia; RMNH.PISC. 6081, holotype of Carangoides gymnostethoides Bleeker 1851b, 334 mm SL, Jakarta, Indonesia. Uraspis Bleeker 1855. Uraspis helvola (Forster in Bloch and Schneider 1801) (21 specimens): ANSP 68453, holotype of Uraspis heidi Fowler 1938b, 215 mm SL, New Jersey, USA; CAS-ICH 305, holotype of Leucoglossa albilinguis Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927, 263 mm SL, Hawaii, USA; CAS-ICH 307, holotype of Uraspis reversa Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927, 265 mm SL, Hawaii, USA; CAS-ICH 309, paratype of Uraspis reversa, 300 mm SL, Hawaii, USA; FRLM 29895, 29896, 101–111 mm SL, East China Sea; FRLM 6651, 31356, 36715, 36905, 36906, 37073, 37521, 39389, 42093, 42094, 58658*, 49–285 mm SL, Mie, Japan; FRLM 42760, 273 mm SL, Kagoshima, Japan; KAUM–I. 51207, 282 mm SL, Kagoshima, Japan; MCZ 16702, 16703, syntypes of Caranx secundus Poey 1860, 296–259 mm SL, Cuba. Uraspis uraspis (Günther 1860) (14 specimens): CAS-SU 27962, holotype of Leucoglossa herklotsi Herre 1932, 185 mm SL, Hong Kong, China; FRLM 12076, 99 mm SL, Palawan, Philippines; FRLM 30714, 32119, 33365, 119–195 mm SL, Phuket, Thailand; KAUM–I. 22154, 22155, 22159, 22160, 22161, 130–142 mm SL, Sabah, Malaysia; UPVMI 440, 441, 657, 144–195 mm SL, Panay I., Philippines; USNM 98820, holotype of Uraspis pectoralis Fowler 1938a, 168 mm SL, Manila, Philippines. Other species of Carangidae and species of other Caragiformes used as comparative materials are listed in Electronic Supplementary Material (ESM) S1.

Morphological examination. Counts and measurements followed Hubbs and Lagler (1947) and Springer and Smith-Vaniz (2008). The count of scutes terminated at the posterior margin of hypural plate. Body size was always expressed as standard length because some specimens had a damaged caudal fin. The relationships between standard length and fork length were provided in ESM S2. Suborbital depth, length of anal-fin base, and upper-lobe length of caudal fin were measured from the ventralmost point of dermal orbital rim to dorsoposterior corner of upper jaw, from the base of third spine to that of posteriormost soft ray, and from center of caudal-fin base to tip of upper lobe, respectively. All measurements were made with a digital caliper to the nearest 0.01 mm (≤ 180 mm), or a divider and scale to the nearest 0.1 mm (> 180 mm). Osteological observations were made from soft X-ray photographs, dissection and/or cleared and stained specimens by the method of Dingerkus and Uhler (1977). Cyanine blue was used to count lateral-line scales and observe scaly areas. Standard length, total length, snout length, suborbital depth, body depth and upper-lobe length of caudal fin are abbreviated as SL, TL, SnL, SOD, BD and UCF, respectively. Institutional codes follow Fricke and Eschmeyer (2021).

DNA preparation, amplification, and sequencing. The specimens used for the present molecular analyses are listed in ESM Table S1. The muscle tissues were stored in 99% ethanol at -18°C. Total DNA was extracted from preserved muscle tissue using the DNeasy Tissue Kit (Qiagen) according to the manufacturer’s protocol. The complete mitochondrial cytochrome b (cytb) gene, COI gene, NADH dehydrogenase subunit 5 (ND5) and 16S were amplified by polymerase chain reaction (PCR) using TaKaRa PCR Thermal Cycler Dice^TM mini (TAKARA BIO INC) with GoTaq^® Colorless Master Mix (Promega) and the primer pairs for cytb Simtcy-R [5’-TGA ARA ACC ACC GTT GTT WTT-3’] (Kuriiwa et al. 2007) and Epipro-F [5’-GTT TAA TTT AGA ATT CTA GCT TTG G-3’] (Kuriiwa, unpublished data), for COI Fish F1 [5’-TCA ACC AAC CAC AAA GAC ATT GGC AC-3’] (Ward et al. 2005) and Fish R1 [5’-TAG ACT TCT GGG TGG CCA AAG AAT CA-3’] (Ward et al. 2005), for ND5 L12321-Leu [5’-GGT CTT AGG AAC CAA AAA CTC TTG GTG CAA-3’] (Miya and Nishida 2000), L12329-Leu [5’-CTC TTG GTG CAA MTC CAA GT-3’] (Miya and Nishida 1999) and H13727-ND5 [5’-GCG ATK ATG CTT CCT CAG GC-3’] (Miya and Nishida 1999), for 16S Primer A [5’-GGT CCW RCC TGC CCA GTG A-3’] (Kurogi 2008) and Primer B [5’-CCG GTC TGR ACY AGA TCA CGT-3’] (Kurogi 2008). Cytb and 16S PCR thermal cycling conditions were as follows: pre-denaturation 94°C for 5 min, 35 cycles of denaturation at 94°C for 30 s, annealing at 57°C for 30 s, extension at 72°C for 1 min, and a final extension step at 68°C for 10 min. COI thermal cycling condition was as follows: pre-denaturation 95°C for 5 min, 35 cycles of denaturation at 95°C for 30 s, annealing at 58°C for 30 s, extension at 72°C for 30 s, and a final extension step at 72°C for 10 min. ND5 thermal cycling condition was as follows: 94°C for 5 min, 35 cycles of denaturation at 94°C for 15 s, annealing at 58°C for 30 s, extension at 72°C for 30 s, and a final extension step at 72°C for 10 min.

PCR products were purified using ExoSAP-IT^® (USB^®) or illustra^TM ExoStar^TM (GE Healthcare) according to the manufacturer’s protocols. The sequencing procedures were carried out in the Center for Molecular Biology and Genetics, Mie University (DNA sequencer: Applied Biosystems^® 3730 DNA Analyzer or Applied Biosystems^® 3130xl Genetic Analyzer). The sequences of cytb, COI, ND5 and/or 16S obtained here were deposited in DDBJ/EMBL/GenBank under the accession numbers shown in ESM Table S1. The DNA sequences were initially aligned using ATGC version 6.0.5 (GENETYX). In addition, the 16S were aligned using CLUSTAL X version 2.1 (Thompson et al. 1997) and then inspected and corrected by eye; after that, the region was aligned using the software ProAlign ver. 0.5a3 (Löytynoja and Milinkovitch 2003) in the 70% posterior probability.

Molecular phylogenetic analysis for the subfamily Caranginae. Maximum likelihood (ML) analysis for phylogenetic reconstruction was applied using RAxML version 8.2.10 (Stamatakis 2014) based on the dataset composed of a concatenated sequence of cytb, COI, ND5 and 16S. To conduct a more comprehensive investigation of the phylogenetic relationships among the species of Caranginae, we downloaded partial mitochondrial sequences of the subfamily from the databases (GenBank and BOLD; ESM Table S2). The GTR + Γ + I model (Yang 1994) was used as the best substitution model tested with MEGA X (Kumar et al. 2018), and rapid bootstrap (BS) analysis was conducted with 1,000 replications. Scomberoides lysan (Fabricius in Niebuhr 1775) and Trachinotus baillonii (Lacepède 1801) were used as the outgroup of Caranginae, and Kaiwarinus equula (Temminck and Schlegel 1844) [formerly known as Carangoides equula (Temminck and Schlegel 1844)] and Pseudocaranx dentex (Bloch and Schneider 1801) as the outgroup of the species formerly belonging to the genus Carangoides and their related species.

Molecular phylogenetic analysis for the genus Uraspis. To check how many species the genus is composed of, ML analysis for phylogenetic relationship of the species of Uraspis was applied using MEGA X. A sequence was prepared as a dataset for analysis and set three codon positions for COI. The Kimura 2-parameter model (Kimura 1980) was used as the best substitution model tested with MEGA X, and BS analysis was conducted with 500 replications. Thirty COI sequences of Uraspis were downloaded from GenBank and BOLD (ESM Table S2) and included in the present analysis.

Results of molecular phylogeny and morphological characteristics of the clades

The complete DNA segments of 1,033–1,141 base pairs (bp) of the cytb, DNA segments of 551–652 bp of COI, DNA segments of 1,166–1,343 bp of ND5, and DNA segments of 467–545 bp of 16S were successfully sequenced and aligned (ESM Table S1).

Molecular phylogeny of the subfamily Caranginae. The present phylogenetic tree of Caranginae shows that the subfamily can be divided broadly into three clades (Fig. 1). Clade 1 consists of majority of the species formerly belonging to Carangoides and its related genera: Alectis, Atropus, Parastromateus, Scyris, Selene and Uraspis. Clade 2 is sister to Clade 1, and is composed of all species of Alepes Swainson 1839, Atule Jordan and Jordan 1922, Caranx Lacepède 1801, Chloroscombrus Girard 1858, Gnathanodon Bleeker 1850, Hemicaranx Bleeker 1862, Megalaspis Bleeker 1851b and Selaroides Bleeker 1851b. The type species of Carangoides, Carangoides praeustus, is nested in the Clade 2 and is distant phylogenetically from the species fomerly considered as congeners. Although Caranx vinctus Jordan and Gilbert 1882b is often considered a member of Carangoides (see Fricke et al. 2021), the species also belongs to Clade 2. Clade 3 is a sister to Clades 1 and 2, consisting of Decapterus Bleeker 1851b, Kaiwarinus Suzuki 1962, Pseudocaranx Bleeker 1863, Selar Bleeker 1851b and Trachurus Rafinesque 1810. Although K. equula is often considered a species of Carangoides, the species is distantly related to the species formerly belonging to Carangoides and its type species, Carangoides praeustus.

Fig. 1

Maximum likelihood tree of the subfamily Caranginae based on the concatenated sequences of the cytb, COI, ND 5 and 16S using the GTR + Γ + I model. Figures beside the internal branches bootstrap values. Species in red the former Carangoides (see Fricke et al. 2021). *Resurrected genus. **New genus

The phylogenetic tree of Caranginae obtained here (Fig. 1) is very similar to that of Carangini given by Santini and Carnevale (2014). Their result also indicates that the subfamily Caranginae consist of three clades, the first includes Decapterus and Trachurus, the second the former Carangoides and Selene, and the third Caranx and Alepes.

Morphological characteristics in the Clades 1–3. The members of Clade 1 have an obvious inferior foramen in the posterior two to four precaudal vertebrae (Starks 1911; Suzuki 1962; Fig. 2) and most of them have strongly compressed bodies. Moreover, they have feebly developed or undeveloped eyelids and comparatively small and weak scutes along the straight portion of lateral line (except for Uraspis species having developed scutes along entire length of the straight portion of lateral line). The species of Clade 2 possess an obvious inferior vertebral foramen in two to five posterior precaudal vertebrae and generally strongly compressed and deep bodies like those of Clade 1 although Atule mate (Cuvier in Cuvier and Valenciennes 1833), Carangoides praeustus, Caranx caballus Günther 1868 and Selaroides leptolepis (Cuvier in Cuvier and Valenciennes 1833) have rather slender bodies. With some exceptions, most of the members belonging to Clade 2 have developed eyelids and strong scutes along the entire straight portion of the lateral line. In all species of Clade 3, no inferior vertebral foramen is found in precaudal vertebrae with only exception of Decapterus tabl Berry 1968 having the foramen in last two precaudal vertebrae as previously stated by Gushiken (1983). Clade 3 consists of comparatively slender-bodied or cylindrical species such as Decapterus, Selar and Trachurus. The deep-bodied species Kaiwarinus equula is, however, included in Clade 3.

Fig. 2

Vertebral columns of six carangin fishes. a Alectis ciliaris, FRLM 60255, 203 mm SL; b Atropus hedlandensis, FRLM 60251, 154 mm SL; c Uraspis helvola, FRLM 58658, 285 mm SL; d Caranx ignobilis, FRLM 60254, 220 mm SL; e Decapterus macarellus, FRLM 60633, 144 mm SL; f Kaiwarinus equula, FRLM 58651, 264 mm SL. All specimens collected from Mie, Japan. Bars 10 mm

Molecular phylogeny of the species formerly included in Carangoides and its related genera. The more comprehensive phylogenetic relationship among the species formerly belonging to the genus Carangoides and their related species (Clade 1 in Fig. 1) is shown in Fig. 3. This relationship divides into 14 subclades (Subclades A–N). The species of previous Carangoides are included in eight subclades (Subclades C, F, G, H, I, J, L and N). Subclades A and B consist of the species formerly belonging to Alectis, but the genus clearly separated into Alectis and Scyris. Subclade C is composed of a single species of Euprepocaranx gen. nov., previously known as Carangoides otrynter (Jordan and Gilbert 1883), synonym of Euprepocaranx dorsalis. Subclade D also consists of a single species of Paraselene gen. nov., previously known as Selene orstedii. The remaining six species of Selene comprise Subclade E, which is sister to Paraselene (Subclade D), and Selene + Paraselene is sister to Euprepocaranx (Subclade C). Subclades A to E make up one lineage, and all members have remarkably deep and compressed bodies. The majority of previous Carangoides species except for E. dorsalis, Ferdauia ferdau, Ferdauia orthogrammus, Carangichthys dinema and Carangichthys oblongus comprise one lineage which divides into five subclades (Subclades F–J). Subclade H includes two previous Carangoides species (Atropus armatus and Atropus hedlandensis), Atropus atropos, and previous Ulua species (Atropus mentalis). Of these, Atropus atropos and the previous Ulua species are the most closely related. Subclade G is sister to Subclade H, including a single previous Carangoides species, Craterognathus plagiotaenia gen. nov. Subclade F is composed of three species of Turrum, previously known as Carangoides coeruleopinnatus, Carangoides fulvoguttatus and Carangoides gymnostethus. Of these, the latter two species form a sister pair. Flavocaranx gen. nov. includes a single species formerly known as Carangoides bajad, which is an independent subclade (Subclade I) sister to Subclade J. Subclade J consists of three species of Platycaranx gen. nov., previously known as Carangoides chrysophrys, Carangoides malabaricus and Carangoides talamparoides, showing a close relationship between the latter two species. Subclades M and N are composed of Uraspis and Carangichthys species, respectively. These two genera show a sister relationship although they have their own morphological characteristics. Subclade L consists of Ferdauia ferdau and Ferdauia orthogrammus, both previously known as Carangoides species. Parastromateus niger has unique morphological characteristics, composing an independent subclade (Subclade K).

Fig. 3

The more comprehensive maximum likelihood tree of the species formerly belonging to the genus Carangoides and their related species based on the concatenated sequences of the cytb, COI, ND 5 and 16S using the GTR + Γ + I model. Figures beside the internal branches bootstrap values. Species in red the former Carangoides (see Fricke et al. 2021). *Resurrected genus. **New genus

Morphological characteristics in the Subclades A–N and Carangoides

To clarify the morphological characteristics of the above-mentioned subclades and Carangoides praeustus which is separated phylogenetically from those Subclades, detailed observations were made on the following 13 characters.

Dorsal contour of upper lip around symphysis. Three types are found in dorsal contour of upper lip around symphysis. Steep type—dorsal edge of upper lip steeply protrudes upward—includes the species of Subclades B, D and E (Fig. 4b, d, e). Steep type is shared in a group Subclade D + Subclade E, but also found in Subclade B. Round chevron type—dorsal edge of upper lip around symphysis roundly protrudes upward—includes the species of Subclades A, C, F, H–J and L–N (Fig. 4a, c, f, h–j and l–n). Of these, Subclades C and J have narrower protrusions than other Subclades. Round chevron type is the most common. Arcuate type—dorsal edge of upper lip around symphysis gradually ascends showing smooth arc without distinct projection—includes the species of Subclades G and K (Fig. 4g, k). Carangoides praeustus (Clade 2 in Fig. 1) has Round chevron type of upper lip (Fig. 4o).

Fig. 4

Upper-jaw symphysis of 15 species belonging to Subclades A to N and Carangoides. a Alectis ciliaris (Subclade A), FRLM 23649, 141 mm SL; b Scyris indica (Subclade B), FRLM 21659, 199 mm SL; c Euprepocaranx dorsalis gen. nov. (Subclade C), USNM 382342, 141 mm SL; d Paraselene orstedii gen. nov. (Subclade D), CAS-ICH 80194, 76 mm SL; e Selene brevoortii (Subclade E), smaller specimen of CAS-ICH 67914, 184 mm SL; f Turrum coeruleopinnatum (Subclade F), FRLM 39643, 167 mm SL; g Craterognathus plagiotaenia gen. nov. (Subclade G), FRLM 39997, 149 mm SL; h Atropus armatus (Subclade H), FRLM 43850, 109 mm SL; i Flavocaranx bajad gen. nov. (Subclade I), HUMZ 165337, 183 mm SL; j Platycaranx malabaricus gen. nov. (Subclade J), FRLM 13216, 158 mm SL; k Parastromateus niger (Subclade K), FRLM 20399, 156 mm SL; l Ferdauia orthogrammus (Subclade L), FRLM 41754, 248 mm SL; m Uraspis helvola (Subclade M), FRLM 6651, 195 mm SL; n Carangichthys oblongus (Subclade N), FRLM 37263, 213 mm SL; o Carangoides praeustus, FRLM 35478, 129 mm SL

Tooth on jaws. Species of Subclades A–G, I, J and L have villiform teeth forming narrow bands on both jaws. In Clade H, species-specific variations of detition are found: slightly large conical teeth arranged uniserially or biserially, or villiform teeth forming bands. Subclade K, Parastromateus niger, has posteriorly curved long conical teeth arranged uniserially on both jaws. Species of Subclade M have small but stout conical teeth uniserially or biserially arranged on both jaws. In Subclades N, upper jaw has a conical tooth band, and lower jaw has an outer row of slightly large conical teeth and an inner band of villiform teeth. Carangoides praeustus has a band of small conical teeth on upper jaw, and has conical teeth forming a band anteriorly and uniserially arranged posteriorly on lower jaw.

Vomerine tooth patch. Species of Subclades A, B, F–J, L and N have chevron-shaped (Fig. 5a) or triangle (Fig. 5f) vomerine tooth patch. These shapes are the most common. The tooth patch of Subclade C is unique, diamond-shape with a rounded posterior corner (Fig. 5b). That of Subclade D is also unique, four-pointed-star-shape (Fig. 5c). These are considered autapomorphic characteristics of each Subclade. Subclade E has two types of the tooth patches: cross-shape with a long posterior extension (Fig. 5d)—Selene brownii, and diamond-shape with a long posterior extension (Fig. 5e)—Selene brevoortii, Selene dorsalis, Selene peruviana, Selene setapinnis and Selene vomer. The tooth patch with a long posterior extension is also considered an autapomorphy of Subclade E and a synapomorphy of the included species. Carangoides has a unique anchor-shaped vomerine tooth patch composed of chevron-shaped and posterior detached long slender tooth patches (Fig. 5g). Subclades K and M lack a vomerine tooth patch.

Fig. 5

Vomerine tooth patches of the species formerly belonging to the genus Carangoides and its related genera. a Alectis ciliaris (Subclade A), FRLM 54883, 114 mm SL; b Euprepocaranx dorsalis (Subclade C), USNM 28153, 177 mm SL; c Paraselene orstedii (Subclade D), CAS-ICH 80194, 76 mm SL; d Selene brownii (Subclade E), USNM 37732, 222 mm SL; e S. vomer (Subclade E), FRLM 52296, 189 mm SL; f Atropus hedlandensis (Subclade H), FRLM 20397, 88 mm SL; g Carangoides praeustus, FRLM 35478, 129 mm SL. Bars 1 mm

Naked thoracic areas. Subclades A–D, F, H–J, L–N and Carangoides have naked area(s) on breast (naked thoracic area, Fig. 6a–d, f, h–j, l–o), whereas the breast is completely covered by scales except for pectoral-fin base in Subclades G and K (Fig. 6g, k). Subclade I has a very narrow naked area along the ventral mid-line anteriorly on the breast (Fig. 6i). The wide naked thoracic area continues to a naked pectoral-fin base in Subclades A–D, H, J and L (Fig. 6a–d, h, j, l). Subclades F, M and L include both species with continuous naked thoracic areas alike Subclade G, and those with separated naked thoracic areas. Subclade E has a variety of naked thoracic areas: no naked areas except for a naked pectoral-fin base (Fig. 6e₁), small naked areas anteriorly on the breast and around the pectoral-fin base (Fig. 6e₂), and narrow naked areas along the ventral midline on the breast extending to the anal-fin origin, along the shoulder girdle and the pectoral-fin base (Fig. 6e₃). Carangoides ire has an obvious naked area on the breast, extending to the pelvic-fin insertion (Fig. 6o₁), whereas the naked area is very small, and found along ventral midline on the jugular in Carangoides praeustus (Fig. 6o₂).

Fig. 6

Naked thoracic areas in Subclades A to N and Carangoides. a Alectis ciliaris (Subclade A), FRLM 25965, 176 mm SL; b Scyris alexandrina (Subclade B), smaller specimen of CAS-ICH 79955, 154 mm SL; c Euprepocaranx dorsalis (Subclade C), USNM 382342, 141 mm SL; d Paraselene orstedii (Subclade D), CAS-ICH 87889, 74 mm SL; e₁ Selene brownii (Subclade E), USNM 37732, 222 mm SL; e₂ Selene vomer (Subclade E), FRLM 54866, 242 mm SL; e₃ Selene brevoortii (Subclade E), smaller specimen of CAS-ICH 67914, 184 mm SL; f₁ Turrum coeruleopinnatum (Subclade F), FRLM 39643, 156 mm SL; f₂ Turrum fulvoguttatum (Subclade F), FRLM 37735, 165 mm SL; g Craterognathus plagiotaenia (Subclade G), FRLM 37736, 279 mm SL; h₁ Atropus armatus (Subclade H), FRLM 43850, 109 mm SL; h₂ Atropus atropos (Subclade H), FRLM 44041, 137 mm SL; h₃ Atropus hedlandensis (Subclade H), FRLM 33731, 170 mm SL; h₄ Atropus mentalis (Subclade H), FRLM 39999, 168 mm SL; i Flavocaranx bajad (Subclade I), USNM 325262, 372 mm SL; j₁ Platycaranx chrysophrys (Subclade J), FRLM 35764, 126 mm SL; j₂ Platycaranx malabaricus (Subclade J), FRLM 13216, 159 mm SL; j₃ Platycaranx talamparoides (Subclade J), FRLM 35697, 166 mm SL; k Parastromateus niger (Subclade K), FRLM 40277, 248 mm SL; l₁ Ferdauia ferdau (Subclade L), FRLM 37031, 206 mm SL; l₂ Ferdauia orthogrammus (Subclade L), FRLM 40007, 226 mm SL; m₁ Uraspis helvola (Subclade M), FRLM 36905, 178 mm SL; m₂ Uraspis uraspis (Subclade M), FRLM 30714, 195 mm SL; n₁ Carangichthys dinema (Subclade N), FRLM 36756, 150 mm SL; n₂ Carangichthys humerosus (Subclade N), FRLM 25724, 140 mm SL; n₃ Carangichthys oblongus (Subclade N), FRLM 34889, 156 mm SL; o₁ Carangoides ire, MNHN 0000-5847, 131 mm SL; o₂ Carangoides praeustus (Clade 2), FRLM 35478, 129 mm SL

Lateral-line scales. The lateral line of the subfamily Caranginae separates into anterior curved and posterior straight parts. The curved part in all Subclades and Carangoides is mainly covered by small cycloid scales, although a few small scutes are sometimes found at the posteriormost part in Subclades H, M and N. The strait part is covered usually by small cycloid scales anteriorly and scutes posteriorly in Subclades A–G and I–L, but usually by scutes only in Subclades M and N. In Subclade H, the strait part of Atropus atropos, Atropus aurochs and Atropus mentalis is usually covered by only scutes, whereas that of Atropus armatus and Atropus hedlandensis usually by both cycloid scales and scutes. The characteristic that whole straight part of lateral line covered by scutes is shared by a lineage Subclade M + Subclade N. The scutes of Subclades A, B, E and K are very small and undeveloped; in particular, scutes in Subclade E are minute thickened scales without obvious spines.

Fin membrane between dorsal-fin spines. Subclades A and B have no membrane between the dorsal-fin spines. Despite short and degenerated dorsal-fin spines in adults, Subclade E has interspinous membranes in the first dorsal fin when the spines are not embedded. Juvenile Parastromateus niger (Subclade K) has very short dorsal-fin spines without any interspinous membrane. In adults, dorsal-fin spines are embedded. Lacking of interspinous membranes in the first dorsal fin is shared in a lineage Subclade A + Subclade B, but found also in Subclade K.

Shapes of neural- and hemal-spine tips of ninth caudal vertebra. Three types of shapes of neural- and hemal-spine tips of ninth caudal vertebra are found. Oar-shape—the distal portions of neural and hemal spines are remarkably expanded posteriorly—is found in Subclades A and C–J (Fig. 7a). Blade-shape—the distal portions of the spines are flattened and slightly expanded—is found in Subclade L (Fig. 7b). Spine-shape—the distal portions of the spines are not expanded, the tips are pointed—is found in Subclades K, M, N and Carangoides praeustus (Fig. 7c). In Subclade B, Scyris alexandrina has the Blade-shape spines, whereas Scyris indica has Oar- or Blade-shape spines. Oar-shape is the most common and it is seen in the lineage consisting of Subclades A to J, except for Subclade B as stated above. The lineage composed of Subclades K to N, sister to the above lineage, has Blade-shape (Subclade L only) or Spine-shape.

Fig. 7

Shapes of neural- and hemal-spine tips of ninth caudal vertebra. a Platycaranx chrysophrys (Subclade J), FRLM 35764, 126 mm SL; b Ferdauia ferdau (Subclade L), FRLM 25888, 155 mm SL; c Carangichthys dinema (Subclade N), FRLM 35585, 147 mm SL. Arrow neural and hemal spines of ninth caudal vertebra

Shapes of first hemal spine. The first hemal spine of Subclade M (Uraspis) is distinctly crooked just below the centrum (Fig. 2c). In the remaining Subclades, the first hemal spine is almost linear or gently curved (Fig. 2a, b). The crooked first hemal spine is unique in Subclade M, therefore, this characteristic is considered an autapomorphy of the Subclade and a synapomorphy of the included species.

Adjacency of ultimate dorsal- and anal-fin pterygiophores with neural and hemal spines. The ultimate dorsal- and anal-fin pterygiophores connect the neural and hemal spines of tenth caudal vertebra, respectively, in Subclades D (Paraselene gen. nov., Fig. 8a) and E (Selene, Fig. 8b). Such pterygiophores of Subclade D are barely apart from the neural and hemal spines. Adjacency of ultimate dorsal- and anal-fin pterygiophores with neural and hemal spines is also found in Atropus atropos (Fig. 8c) and Atropus mentalis of Subclade H, all species of Platycaranx gen. nov (Subclade J, Fig. 8d). Although Alectis ciliaris (Subclade A) and two species of Scyris (Subclade B) have extremely deep and compressed bodies and are closely similar to Selene species in general body shape, the former three species have the ultimate dorsal- and anal-fin pterygiophores fully apart from neural and hemal spines, respectively.

Fig. 8

Adjacency of ultimate dorsal- and anal-fin pterygiophores with neural and hemal spines of tenth caudal vertebra (arrows). a Paraselene orstedii (Subclade D), CAS-ICH 80194, 76 mm SL; b Selene vomer (Subclade E), FRLM 54866, 242 mm SL; c Atropus atropos (Subclade H), FRLM 8897, 90 mm SL; d Platycaranx malabaricus (Subclade J), FRLM 13217, 146 mm SL

Numbers of dorsal- and anal-fin soft rays. Subclades A–E, G, H, J and N show fewer total numbers of dorsal- and anal-fin soft rays (31–44); whereas Subclades I, L and M have more total numbers of dorsal- and anal-fin soft rays (46–61; Fig. 9, Table 1). Subclade F indicates an exceptionally wide range of the total number, because one of the included species, Turrum coeruleopinnatum has much fewer total number of dorsal- and anal-fin soft rays (38–43) than the other two species, Turrum fulvoguttatum (48–51) and Turrum gymnostethus (55–58). Parastromateus niger (Subclade K) also possesses an exceptionally numerous total number of soft rays (78–83). Carangoides shows comparatively fewer total number of dorsal- and anal-fin soft rays (41–44).

Fig. 9

Total number of second dorsal- and anal-fin soft rays in Subclades A to N and Carangoides

Table 1 Number of fin rays in the species belonging to the subclade A to N and the genus Carangoides

Number of neural spines between pterygiophores supporting first and last dorsal-fin spines. Subclades A–J, N and Carangoides have six neural spines between pterygiophores supporting the first and last dorsal-fin spines. Subclades K, L and M have three to five, five and two to four spines, respectively.

Number of pterygiophores between first and second hemal spines. Subclades A–E, G, H, J, N and Carangoides have three pterygiophores between the first and second hemal spines. Of Subclade F, Turrum coeruleopinnatum has three pterygiophores, whereas the other two species Turrum fulvoguttatum and Turrum gymnostethus have four. Subclades I, K, L and M have four (or rarely five), eight to ten, five to six and three to six pterygiophores, respectively.

Proportion of snout length. Relative lengths of snout (SnL/SL) roughly divide into three groups (Fig. 10a). Subclades G, H, K–N and Carangoides have relatively shorter snouts (7.4–12.1%), whereas Subclades B and E have relatively longer snouts (14.2–23.4%) (Fig. 10a, Table 2). These two groups are clearly separated. However, the remaining Subclades A, C, F, I and J show the intermediate values (10.4–14.5%), and the values overlap with those of the former two groups.

Fig. 10

Ratios of a snout length and b suborbital depth to standard length (%) in Subclades A to N and Carangoides

Table 2 Proportional measurements of head parts and caudal fin in the species belonging to the subclade A to N and the genus Carangoides

Proportion of suborbital depth. Ratios of suborbital depth (SOD/SL) clearly divide into two groups (Fig. 10b). Subclades B, D and E have a broad suborbital (12.5–22.5%), whereas Subclades A, C, F–N and Carangoides have narrower suborbital (2.4–9.0%) (Fig. 10b, Table 2). In Subclades B, D and E, the suborbital depth is slightly shorter than the snout length (SOD 80.1–99.4% SnL). Of the remaining Subclades, Subclades A and C have relatively wide suborbital, 5.5–9.0% SL. In Subclade F, Turrum coeruleopinnatum has wider suborbital (5.3–8.6% SL) than Turrum fulvoguttatum (4.5–5.9%) or Turrum gymnostethus (4.2–4.6%).

Reconstitution of the former Carangoides and its related genera

The relationship between the above morphological features and each Subclade is summarized in Tables 3 and 4. Although no clear autapomorphy could be revealed for all subclades, they possess a unique combination of morphological characteristics, and they are certainly considered as independent genera. Therefore, the former Carangoides and its relatives are sorted and redefined as follows.

Table 3 Comparison of external morphology among Subclades and the genus Carangoides

Table 4 Comparison of internal morphology among Subclades and the genus Carangoides

Alectis Rafinesque 1815

(Subclade A, Fig. 11)

Fig. 11

Alectis ciliaris, the species of monotypic genus Alectis (Subclade A). a₁ FRLM 25965, 176 mm SL, Mie, Japan (fresh condition); a₂ RMNH.PISC. 26963, lectotype of Carangoides blepharis, 272 mm SL, Jakarta, Indonesia

Gallus Lacepède 1802: 583 (type species: Gallus virescens Lacepède 1802)

Alectis Rafinesque 1815: 84 (type species: Gallus virescens Lacepède 1802)

Blepharis Cuvier 1816: 322 (type species: Zeus ciliaris Bloch 1787)

Blepharichthys Gill 1861: 36 (type species: Zeus ciliaris Bloch 1787)

Type species. Gallus virescens Lacepède 1802, a junior synonym of Alectis ciliaris (Bloch 1787).

Included species. Alectis ciliaris (Bloch 1787).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body strongly compressed, very deep, disk-like (juveniles) or rhomboidal (adults); dorsal edge of upper lip around symphysis roundly protruding upward (Round chevron type; Fig. 4a); dorsal contour of head somewhat roundly convex; jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch chevron-shaped or triangular (Fig. 5a, f); adipose eyelid poorly developed; naked area on breast extending beyond pelvic-fin insertion and connected with naked pectoral-fin base (Fig. 6a); curved part and anterior straight part of lateral line with small cycloid scales (94–107 + 9–32), the posterior straight part with 7–25 small scutes; first dorsal fin without interspinous membranes, with V–VIII spines, second dorsal fin with 18–20 soft rays; anal-fin soft rays 15–17; some soft rays of dorsal and anal fins very long and filamentous in juveniles; first and second anal-fin spines embedded in juveniles and adults; no finlets; both neural and hemal spines of ninth caudal vertebra Oar-shape (Fig. 7a); total gill rakers 18–22; snout relatively short, 10.4–11.9% SL; suborbital moderately broad, 7.0–9.0% SL.

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth almost terminal, lower jaw slightly protruding. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 11th–13th dorsal-fin soft ray. First dorsal fin embedded in adults; second dorsal-fin base slightly longer than anal-fin base. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen in last two precaudal vertebrae.

Table 5 Number of scales and gill rakers in the species belonging to the subclade A to N and the genus Carangoides

Table 6 Proportional measurements of body parts in the species belonging to the subclade A to N and the genus Carangoides

Distribution. Distributed in tropical, subtropical and temperate seas circumglobally.

Gender. Feminine.

Comparison. Alectis and Scyris are distinguishable from former Carangoides and its relative genera except for Parastromateus by lack of membranes of first dorsal fin in both juveniles and adults, and the fin embedded in adults (vs. membrane present in first dorsal fin). Both genera can be distinguished from Parastromateus by having a naked area widely on breast (vs. breast completely scaly; Fig. 6). Alectis differs from Scyris by having a gently protruded dorsal edge of the upper lip around symphysis (Fig. 4a; vs. sharply protruded, Fig. 4b), slightly rounded dorsal contour of head (vs. distinctly concave), a fewer gill rakers (18–22 vs. 31–37, Table 5), a narrower suborbital (SOD 7.0–9.0% SL vs. 12.5–17.3% SL) and inferior vertebral foramen in posterior two precaudal vertebrae (vs. in posterior three precaudal vertebrae). Both genera are also distinguished from other carangid genera in having villiform tooth bands on both jaws, poorly developed eyelid, curved part of lateral line entirely covered with cycloid scales, scutes on posterior straight part of lateral line, and no finlets.

Remarks. The genus Alectis is a replaced name of Gallus Lacepède 1802 (see Daget and Smith-Vaniz 1986; Fricke et al. 2021). Gallus Lacepède 1802 is objectively invalid because this nominal genus was preoccupied by Brisson (1760) in Aves (Smith-Vaniz et al. 1979). The type species of Gallus Lacepède 1802 (and also Alectis) is Gallus virescens Lacepède 1802. The original description of G. virescens includes Zeus gallus Linnaeus 1758 as a synonym, and the former nominal species is considered a replacement of the latter (Smith-Vaniz et al. 1990; Fricke et al. 2021). Zeus gallus was established based on four studies (see Wheeler 1985; Kottelat, personal communication), and was characterized by having the tenth dorsal-fin and second anal-fin rays longer than the body. Linnaeus (1758) also stated that the habitat is America. Good and Bean (1885) and Wheeler (1985) considered Zeus gallus as a species of Selene. However, the anterior second dorsal- and anal-fin soft rays of Selene and Paraselene gen. nov. are not so long, shorter than the standard lengths except for juveniles. Whereas, the young Alectis ciliaris and two species of Scyris have very long filamentous dorsal- and anal-fin soft rays longer than their standard lengths. Of these three species, Alectis ciliaris is only distributed in America. From these facts, Zeus gallus is identifiable with Alectis ciliaris (= Zeus ciliaris Bloch 1787). Although Zeus gallus is a senior synonym of Zeus cilialis, Good and Bean (1885) was the last time the former was used as a valid name and has not been used as valid since 1900 as far as we know. On the other hand, Alectis ciliaris has been used as the presumed valid name by many authors over a long period of time (Smith-Vaniz et al. 1979, 1990; Gushiken 1983, 1984, 1988; Smith-Vaniz 1983, 1986, 1995, 1999, 2002, 2016; Gloerfelt-Tarp and Kailola 1984; Laroche et al. 1984; Sainsbury et al. 1985; Kijima et al. 1986; Allen and Swainston 1988; Randall et al. 1990; Mok 1993; Allen 1997; Rao et al. 2000; Reed et al. 2001, 2002; Senou 2002; Shao and Chen 2003; Rao 2004; Randall 2005; Hoese and Gates 2006; Springer and Smith-Vaniz 2008; Kimura 2009, 2011, 2013, 2017, 2018; White et al. 2013; Santini and Carnevale 2014; Hata 2019). Because Art. 23.9.1.1 and 23.9.1.2 of ICZN (1999) are satisfied, the younger name, Alectis ciliaris is valid for the species in accordance with Art. 23.9.2 of ICZN (1999).

The genus Blepharis established by Cuvier (1816) on the basis of Zeus ciliaris is a junior subjective synonym of Alectis Rafinesque 1815, because the type species of the latter genus G. virescens is a subjective synonym of Zeus ciliaris as stated above. The generic name Blepharis was also given to plants of the family Acanthaceae by Jussieu (1789). Because of this, Gill (1861) established Blepharichthys as a replace name for Blepharis. However, because Blepharis Cuvier 1816 is not a homonym of Blepharis Jussieu 1789 (Art. 52.7, ICZN 1999), this replacement is unneeded (Smith-Vaniz et al. 1990).

At the same time as the description of Zeus ciliaris, Bloch (1787: pl. 192, fig. 1) gave the redescription and drawing of Zeus gallus (Biodiversity Heritage Library: https://doi.org/10.5962/bhl.title.62139; accessed on 7 Mar 2021). This drawing is undoubtedly a species of Scyris because it has a concave dorsal contour of head as stated above.

Scyris Cuvier 1829

(Subclade B, Fig. 12)

Fig. 12

Two included species of the genus Scyris (Subclade B), a Scyris alexandrina, type species and b Scyris indica. a smaller specimen of CAS-ICH 79955, 154 mm SL, Cameroon; b₁ FRLM 35144, 157 mm SL, Songkhla, Thailand (fresh condition); b₂ RMNH.PISC. 26962, lectotype of Carangoides gallichthys, 118 mm SL, Indonesia

Scyris Cuvier 1829: 209 (type species: Gallus alexandrinus Geoffroy St. Hilaire 1817)

Gallichtys Cuvier in Cuvier and Valenciennes 1833: 168 (type species: Gallichtys major Cuvier in Cuvier and Valenciennes 1833)

Hynnis Cuvier in Cuvier and Valenciennes 1833:195 (type species Hynnis goreensis Cuvier in Cuvier and Valenciennes 1833)

Type species. Gallus alexandrinus [= Scyris alexandrina (Geoffroy St. Hilaire 1817)].

Included species. Scyris alexandrina (Geoffroy St. Hilaire 1817) and Scyris indica Rüppell 1830.

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body strongly compressed, very deep, rhomboidal; dorsal edge of upper lip around symphysis steeply protruding upward (Steep type; Fig. 4b); dorsal contour of head just anterior to eye distinctly concave; jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch chevron-shaped or triangular (Fig. 5a, f); adipose eyelid poorly developed; naked area on breast extending beyond pelvic-fin insertion and connected with naked pectoral-fin base (Fig. 6b); curved part and anterior straight part of lateral line with small cycloid scales (72–92 + 27–42), the posterior straight part with 8–20 small scutes; first dorsal fin without fin membranes, with VI or VII spines, second dorsal fin with 17–22 soft rays; anal-fin soft rays 15–20; some soft rays of dorsal, anal and pelvic fins elongate and filamentous in juvenile; first and second anal-fin spines embedded in juveniles and adults; no finlets; both neural and hemal spines of ninth caudal vertebra Oar- or Blade-shape (Fig. 7a, b); total gill rakers 29–39; snout long, its length 15.2–20.0% SL (Fig. 10a, Table 2); suborbital broad, 12.5–17.3% SL (Fig. 10b, Table 2).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Lower jaw slightly protruding. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 11th–12th dorsal-fin soft ray. First dorsal fin embedded in adults; second dorsal-fin base slightly longer than anal-fin base. Six neural spines between pterygiophores of first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 16 in Scyris alexandrina and 10 + 14 in Scyris indica; inferior foramen in last three precaudal vertebrae.

Distribution. Scyris alexandrina is known from the eastern Atlantic Ocean from Morocco to southern Angola and the Mediterranean (Smith-Vaniz 2016); Scyris indica is known from the Indo-West Pacific from the Red Sea and the eastern coast of Africa to Japan and the eastern coast of Australia (Smith-Vaniz 1983, 1999).

Gender. Feminine.

Comparison. See “Comparison” of Alectis.

Remarks. Scyris alexandrina and Scyris indica have previously been congeneric with Alectis ciliaris. Because the former two species and the latter are genetically well separated and each of them have unique morphological characteristics, they are treated as independent genera. Scyris is the oldest generic name for those two species. Gallichtys major, the type species of Gallichtys, is identifiable with Scyris indica from the original drawing. Hynnis goreensis, the type species of Hynnis, can be also identified with Scyris alexandrina from its original drawing. Therefore, nominal genera of Gallichtys and Hynnis are the subjective synonyms of Scyris.

Scyris indica was originally described as Scyris indicus by Rüppell (1830). However, “indica” is the correct spelling of the specific name because the genus Scyris is feminine and “indicus” is a Latin adjective (ICZN 1999, Art. 31.2).

Euprepocaranx gen. nov.

(Subclade C, Fig. 13)

Fig. 13

Euprepocaranx dorsalis, the species of monotypic genus Euprepocaranx gen. nov. (Subclade C). a₁ larger specimen of USNM 4957, syntype of Carangoides dorsalis, 190 mm SL, western coast of Central America (photographed by Y. Hibino); a₂ USNM 396266, 105 mm SL, Azuero Peninsula, Pacific Panama (right side, mirror image)

Type species. Carangoides dorsalis Gill 1863c.

Included species. Euprepocaranx dorsalis (Gill 1863c).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body strongly compressed, very deep, disk-like (juveniles) or rhomboidal (adults); dorsal edge of upper lip around symphysis roundly protruding upward (Round chevron type), protrusion of upper-jaw symphysis narrow (Fig. 4c); dorsal contour of head straight or slightly concave; jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch diamond-shaped with round posterior corner (Fig. 5b); adipose eyelid poorly developed; naked area on breast extending beyond pelvic-fin insertion and connected with naked pectoral-fin base (Fig. 6c); curved part and anterior straight part of lateral line with cycloid scales (57–71 + 1–12), the posterior straight part with 43–56 very small scutes; first dorsal fin with interspinous membranes, with VII–VIII spines, second dorsal fin with 17–19 soft rays; anal-fin soft rays 15–17; anterior soft rays of second dorsal and anal fins elongate and filamentous; no finlets; both neural and hemal spines of ninth caudal vertebra Oar-shape (Fig. 7a), total gill rakers 21–24; snout relatively long, 11.7–13.6% SL (Table 2, Fig. 10a); suborbital depth 5.5–8.4% SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth terminal. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of ninth or tenth dorsal-fin soft ray. Second dorsal-fin base slightly longer than anal-fin base. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen on last three precaudal vertebrae.

Distribution. Euprepocaranx dorsalis is known from tropical eastern Pacific Ocean from Baja California to the northern Ecuador (Smith-Vaniz 1995).

Etymology. The generic name “Euprepocaranx” derived from “euprepes”, the Greek meaning “good-looking”, and “Caranx”, the type genus of Carangidae, in reference to beautiful body shape with long filamentous dorsal- and anal-fin soft rays of the type species.

Gender. Masculine.

Comparison. Euprepocaranx is distinguishable from the former Carangoides and other genera by a unique shape of vomerine tooth patch, diamond-shaped with round posterior corner (Fig. 5b). Euprepocaranx dorsalis shares the disc-like or rhomboid body and elongate dorsal- and anal-fin soft rays with species of Alectis and Scyris, but the former has the first dorsal fin not embedded, with interspinous membranes (vs. embedded, without interspinous membranes). The former species, moreover, differs from Scyris species by having a round dorsal edge of upper jaw symphysis (Fig. 4c; vs. sharp chevron-shaped, Fig. 4b). Euprepocaranx dorsalis is similar to Carangichthys species in having long anterior second dorsal- and anal-fin soft rays and small dark blotches along the second dorsal-fin base, but differs from the later species in having cycloid scales in the anterior straight part of the lateral line (vs. whole straight part of the lateral line covered by scutes) and both neural and hemal spines of ninth caudal vertebra Oar-shape (vs. Spine-shape, Fig. 7).

Remarks. Euprepocaranx dorsalis has been previously known as Carangoides otrynter. Jordan and Gilbert (1883) described the present species as a new species Caranx otrynter without any types, and they listed “Carangoides dorsalis Gill, Proc. Ac. Nat. Sci. Phila., 1863, 166 (Panama; not Vomer dorsalis Gill)” as a synonym. From this, they perceived that their new species and Gill’s Carangoides dorsalis were conspecific. Perhaps they described this new species to replace the specific name of Carangoides dorsalis to avoid confusion or homonymy with Vomer dorsalis Gill 1863a [= Selene dorsalis (Gill 1863a)]. However, they did not use “replacement” or its same meaning in the original description of Caranx otrynter. Since Caranx otrynter is not a replacement name of Carangoides dorsalis superficially, the former is a junior synonym of the latter.

Paraselene gen. nov.

(Subclade D, Fig. 14)

Fig. 14

Paraselene orstedii, the species of monotypic genus Paraselene gen. nov. (Subclade D). ZMUC P461509, syntype of Selene orstedii, 136 mm SL, Puntarenas, Costa Rica (photographed by M. Krag)

Type species. Selene orstedii Lütken 1880 [= Paraselene orstedii (Lütken 1880)].

Included species. Paraselene orstedii (Lütken 1880).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body strongly compressed, very deep, disk-like; dorsal edge of upper lip around symphysis steeply protruding upward (Steep type; Fig. 4d); dorsal contour of snout distinctly concave; jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch shaped four-pointed-star (Fig. 5c); adipose eyelid poorly developed; naked area on breast extending beyond pelvic-fin insertion and connected with naked pectoral-fin base (Fig. 6d); curved part and anterior straight part of lateral line with cycloid scales (45–51 + 18–24), the posterior straight part with 18–22 scutes; first dorsal fin with interspinous membranes, with IV–VIII spines, second dorsal fin with 16–18 soft rays; anal-fin soft rays 14–16; no finlets; both neural and hemal spines of ninth caudal vertebra Oar-shape (Fig. 7a); ultimate neural and hemal pterygiophores connecting neural and hemal spines of tenth caudal vertebra, respectively (Fig. 8a); total gill rakers 19–23; snout long, its length 17.9–20.6% SL (Fig. 10a, Table 2); suborbital broad, 15.9–17.5% SL (Fig. 10b, Table 2).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Lower jaw protruded. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of seventh dorsal-fin soft ray. Second dorsal-fin base slightly longer than anal-fin base. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen on last three precaudal vertebrae.

Distribution. Paraselene orstedii is known from the eastern Pacific Ocean from Mexico to Colombia (Smith-Vaniz 1995).

Etymology. The generic name “Paraselene” derived from “para”, the Greek meaning “near” or “beside”, and “selene”, the Greek meaning “moon”, and the generic name to which the present species has belonged before, in reference to a close relationship with the genus Selene.

Gender. Feminine.

Comparison. Paraselene is distinguishable from the former Carangoides and other genera by having a vomerine tooth patch shaped four-pointed-star (Fig. 5c). Although Paraselene is very similar to Selene in general appearance, the former is distinguishable from the latter by having a wide thoracic naked area extending to pelvic-fin insertion and connected with the naked pectoral-fin base (vs. naked area absent or very small area, Fig. 6) and fewer total gill rakers (19–23 vs. 30–40).

Remarks. Paraselene shows sister relationship to Selene in our result of molecular phylogeny (Fig. 3). The genetic difference between them is rather large and enough to separate them into different genera. In the phylogenetic tree from maximum-parsimony analyses by Reed et al. (2001), Paraselene orstedii (as Selene orstedii) was nested outside of the species of Selene + Alectis and Scyris (as Alectis) + Euprepocaranx (as Carangoides otrynter). In the phylogenetic tree from maximum-likelihood analysis by Santini and Carnevale (2014), Paraselene orstedii (as Selene orstedii) was also located outside of Selene and Euprepocaranx (as Carangoides otrynter). Those results support that Paraselene orstedii belongs to an independent genus which is different from Selene species.

Selene Lacepède 1802

(Subclade E, Fig. 15)

Fig. 15

Six included species of the genus Selene (Subclade E), a Selene brevoortii, b Selene brownii, c Selene dorsalis, d Selene peruviana, e Selene setapinnis and f Selene vomer, type species. a₁ USNM 4438, holotype of Argyriosus brevoortii, 48 mm SL, Baja California, Mexico; a₂ USNM 39896, paratype of Argyreiosus pacificus, 216 mm SL, Baja California, Mexico; b CAS-SU 67160, 75 mm SL, La Habana, Cuba; c SAM 13725, holotype of Selene gibbiceps, 158 mm SL, Walvis Bay, Namibia (photographed by H. Hata); d USNM 81773, holotype of Vomer declivifrons, 151 mm SL, Panama City, Pacific Panama; e AMNH I-7148, holotype of Vomer setipinnis cubensis, 117 mm SL, Cuba; f FRLM 49239, 206 mm SL, reared specimen (fresh condition, right side, mirror image)

Selene Lacepède 1802: 560 (type species: Selene argentea Lacepède 1802)

Argyreiosus Lacepède 1802: 566 (type species: Zeus vomer Linnaeus 1758)

Vomer Cuvier 1816: 316 (type species: Vomer brownii Cuvier 1816)

Platysomus Swainson 1839: 176 (type species: Vomer brownii Cuvier 1816)

Type species. Selene argentea Lacepède 1802, a junior synonym of Selene vomer (Linnaeus 1758).

Included species. Selene brevoortii (Gill 1863b), Selene brownii (Cuvier 1816), Selene dorsalis (Gill 1863a), Selene peruviana (Guichenot 1866), Selene setapinnis (Mitchill 1815) and Selene vomer (Linnaeus 1758).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body strongly compressed, very deep, disk-like; dorsal edge of upper lip around symphysis steeply protruding upward (Steep type; Fig. 4e); dorsal contour of head just anterior to eye distinctly concave; jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch diamond- or cross-shaped with a long posterior extension (Fig. 5d, e); adipose eyelid poorly developed; breast completely scaly, or narrow naked area present behind operculum but not connected to naked pectoral-fin base (Fig. 6e_1–3); curved part and usually anterior straight part of lateral line with cycloid scales (52–94 + 0–67), the posterior straight part with 6–71 scutes; first dorsal fin with interspinous membranes, with VIII (rarely VII) spines, second dorsal fin with 19–24 soft rays; anal-fin soft rays 16–20; no finlets; both neural and hemal spines of ninth caudal vertebra Oar-shape (Fig. 7a); ultimate neural and hemal pterygiophores connecting neural and hemal spines of tenth caudal vertebra, respectively (Fig. 8b); total gill rakers 30–44; snout long, its length 14.2–23.4% SL (Fig. 10a, Table 2); suborbital very broad, 13.1–22.5% SL (Fig. 10b, Table 2).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Lower jaw protruded. Scales on body small, cycloid, except for posterior lateral line; naked thoracic area variable: no naked area—Selene brownii and Selene setapinnis (Fig. 6e₁), small naked area just posterior to ventroposterior edge of operculum—Selene peruviana and S. vomer (Fig. 6e₂), narrow naked areas just posterior to operculum and along ventral midline at breast extending to origin of soft anal fin (Fig. 6e₃)—Selene brevoortii and Selene dorsalis; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of tenth dorsal-fin soft ray. Second dorsal-fin base almost equal to anal-fin base or slightly shorter. Six neural spines between pterygiophores supporting first and last dorsal-fin spine; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen on last three precaudal vertebrae.

Distribution. Eastern Pacific Ocean from Baja California to the northern Ecuador (Selene brevoortii and Selene peruviana), western Atlantic Ocean from Nova Scotia to the northern Argentina (Selene brownii, Selene setapinnis and Selene vomer), and eastern Atlantic Ocean, Madeira, Canarias, Cabo Verde and from Senegal to Namibia (Selene dorsalis) (Smith-Vaniz 2016).

Gender. Feminine.

Comparison. Selene is distinguishable from the former Carangoides and other genera by having a diamond- or cross-shaped vomerine tooth patch with a long posterior extension (Fig. 5d, e), and an elongated snout (its length 14.2–23.4% SL; Fig. 10a, Table 2) as well as a broad suborbital, 13.1–22.5% SL (Fig. 10b, Table 2). Selene shares a sharp chevron type dorsal edge of upper lip around symphysis, a longer snout and a broader suborbital with Scyris (Figs. 4, 10a, b), but the former is distinguishable from the latter by having interspinous membranes of first dorsal fin (vs. no interspinous membranes) and ultimate neural and hemal pterygiophores closely adjacent to the neural and hemal spines of the tenth caudal vertebra, respectively (Fig. 8b; vs. separated). Paraselene also has a sharp-chevron type of dorsal edge of the upper lip around symphysis, a long snout and a broad suborbital (Figs. 4, 10a, b), morphological differences between Selene and Paraselene are described in “Comparison” of the later genus.

Remarks. Because Zeus vomer, the type species of Argyreiosus Lacepède 1802, is a member of Selene (Selene vomer), both nominal generic names are synonymous. Although Argyreiosus and Selene were published at same time, Selene was given precedence (ICZN 1959).

Turrum Whitley 1932

(Subclade F, Figs. 16, 17)

Fig. 16

Turrum coeruleopinnatum of the genus Turrum (Subclade F). a₁ FRLM 35139, 154 mm SL, Songkhla, Thailand (fresh condition); a₂ RMNH.PISC. 6112, 54 mm SL, Ambon, Saparua or Manado, Indonesia; a₃ RMNH.PISC. 26972, 66 mm SL, Ambon, Saparua or Manado, Indonesia; a₄ FRLM 40025, 165 mm SL, Songkhla, Thailand (fresh condition); a₅ FRLM 37735, 356 mm SL, Okinawa, Japan (fresh condition); a₆ FRLM51147, 129 mm SL, Mie, Japan

Fig. 17

Turrum fulvoguttatum, type species (a) and Turrum gymnostethus (b) of the genus Turrum (Subclade F). a₁ FRLM 40025, 165 mm SL, Songkhla, Thailand (fresh condition); a₂ FRLM 37735, 356 mm SL, Okinawa, Japan (fresh condition); a₃ largest specimen of NMW 11627, syntype? of Caranx bleekeri, 187 mm SL, Red Sea (© Naturhistorisches Museum, Wien, photographed by A. Naseka); a₄ RMNH.PISC. 27127, syntype of Carangoides ophthalmotaenia, 66 mm SL, Ambon, Indonesia; a₅ ZIN 2544, syntype of Caranx bleekeri, 166 mm SL, Red Sea (photographed by M. Matsunuma); b₁ FRLM 31558, 121 mm SL, Nha Trang, Vietnam (fresh condition); b₂ FRLM 39984, 185 mm SL, Bitung, Indonesia (fresh condition); b₃ RMNH.PISC. 6081, holotype of Carangoides gymnostethoides, 334 mm SL, Jakarta, Indonesia

Turrum Whitley 1932: 337 (Type species: Turrum emburyi Whitley 1932)

Type species. Turrum emburyi Whitley 1932, a junior synonym of Turrum fulvoguttatum (Forsskål in Niebuhr 1775).

Included species. Turrum coeruleopinnatum (Rüppell 1830), Turrum fulvoguttatum (Forsskål in Niebuhr 1775), and Turrum gymnostethus (Cuvier in Cuvier and Valenciennes 1833).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body either compressed, deep, oval in Turrum coeruleopinnatum or somewhat slender, oblong in adult Turrum fulvoguttatum and Turrum gymnostethus; dorsal contour of upper lip around symphysis roundly convex (Round chevron type; Fig. 4f); jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch chevron-shaped or triangular (Fig. 5a, f); adipose eyelid poorly developed; naked area on breast extending beyond base of last pelvic-fin soft ray (Fig. 6f_{1, 2}); curved part and anterior straight part of lateral line with cycloid scales (66–110 + 1–34), the posterior straight part with 13–36 scutes; first dorsal fin with interspinous membranes, with VIII (rarely VII) spines, second dorsal fin with 20–32 soft rays; anal-fin soft rays 16–26; anterior second dorsal and anal fins falcate; no finlets; both neural and hemal spines of ninth caudal vertebra expanding, Oar-shape (Fig. 7a); total gill rakers 21–31; snout longer than eye diameter, SnL/SL 10.9–14.3% (Fig. 10a, Table 2); suborbital depth 4.2–8.6% SL (Fig. 10b, Table 2).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth almost terminal, or lower jaw slightly protruding. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, straight part beginning below base of 12th–18th dorsal-fin soft ray; naked area on breast connecting naked area on pectoral-fin base in Turrum coeruleopinnatum (Fig. 6f₁), but not connecting in Turrum fulvoguttatum (Fig. 6f₂) and Turrum gymnostethus. Second dorsal-fin base slightly longer than soft portion of anal-fin base; first and second anal-fin spines small, separated from the third spine, and not embedded in adults. Six neural spines between pterygiophores supporting first and last dorsal-fin spine; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14 (14–15 in Turrum gymnostethus); inferior foramen on last three (four in some Turrum coeruleopinnatum) precaudal vertebrae.

Distribution. Indo-Pacific from the Red Sea and the estern coast of Africa to Samoa (Smith-Vaniz 1983, 1999; Randall 2005).

Gender. Neuter.

Comparison. The genus Turrum is distinguished from Alectis, Scyris and Parastromateus by having interspinous membranes in the first dorsal fin (vs. lacking the membranes), from Euprepocaranx, Paraselene and Selene by having a chevron-shaped or triangular vomerine tooth band (vs. diamond-, cross- or four-pointed-star-shaped; Fig. 5), from Craterognathus gen. nov., Parastromateus, Flavocaranx gen. nov., Ferdauia, Uraspis, Carangichthys and Carangoides by having a thoracic naked area extending posteriorly beyond pelvic-fin insertion (vs. thoracic naked area absent or not extending beyond pelvic-fin insertion; Fig. 6), from Atropus by having a longer snout (10.9–14.3% SL vs. 7.4–10.7% SL; Fig. 10b, Table 2), and Platycaranx gen. nov. by having wide protrusion of upper-jaw symphysis (vs. narrow protrusion; Fig. 4). Turrum coeruleopinnatum apparently resembles species of Platycaranx in having a deep, compressed and oval body. The former species differs from Platycaranx malabaricus and Platycaranx talamparoides in lacking a small naked area anterodorsally to pectoral-fin base (vs. naked area present; Fig. 6) and from P. chrysophrys in having 20–23 second dorsal-fin soft rays and somewhat pointed snout (vs. 18–21 rays and truncate snout) in addition to the morphological difference of upper-jaw symphysis. Moreover, Turrum fulvoguttatum and Turrum gymnostethus are different from Flavocaranx in number of pterygiophores between the first and second hemal spines (three vs. four or five), and from two species of Ferdauia in shape of neural and hemal spines of ninth caudal vertebra (Oar-shape vs. Blade-shape; Fig. 7).

Remarks. The genus Turrum was established by Whitley (1932) on the basis of Turrum emburyi Whitley 1932 as a monotypic genus. Turrum emburyi has been synonymized under Turrum fulvoguttatum (as Carangoides fulvoguttatus) by Dor (1984), Paxton et al. (1989), Gunn (1990) and Smith-Vaniz (1999). We here reconfirmed this synonymy because important characteristics of Turrum emburyi described by Whitley (1932), i.e., second dorsal fin with 29 soft rays, anal fin with 24 soft rays, anterior second dorsal and anal fins forming moderate lobes, naked area on breast extending behind the pelvic fin and does not continue to the naked area on pectoral-fin base, and lower margin of eye located above a horizontal line through snout tip, and dorsal profile of snout almost straight (based on Whitley 1932: pl. 38, fig. 4), agree well with those of Turrum fulvoguttatum. As diagnoses of the genus, Whitley (1932) stated the following characters: villiform tooth bands on jaws, vomer and palatines; lips not sharp-edged; gill rakers not extending forward into mouth; body compressed, dorsal contour of body more convex than the ventral contour; naked area on breast; straight part of lateral line beginning well posterior to verticals through second dorsal- and anal-fin origins; scutes well developed posteriorly but not hooked forward; second dorsal and anal fins lobed anteriorly without convex margins, produced rays, or finlets; and no transverse bands across head or body. These characters, excluding transverse bands, agree well with those of Turrum coeruleopinnatum, Turrum fulvoguttatum and Turrum gymnostethus. Juveniles of these three species have an oblique dark band through eye on head.

Of the included three species, Turrum fulvoguttatum and Turrum gymnostethus are genetically divergent from Turrum coeruleopinnatum as shown in Fig. 3. Turrum fulvoguttatum and Turrum gymnostethus morphologically resemble each other in having somewhat slender body, especially in adults (BD/SL 31.7–54.1% and 30.4–48.4%, respectively) and comparatively numerous soft rays of dorsal (25–30 and 28–32, respectively) and anal fins (21–26 and 24–26, respectively). In contrast, Turrum coeruleopinnatum has considerably deep body (BD/SL 45.8–69.3%) and fewer soft rays of dorsal and anal fins (20–23 and 16–20, respectively), showing morphological difference from the former two congeners.

Although Carangoides ophthalmotaenia was described by Bleeker (1852) based on two syntypes, 86 and 126 mm in body length, collected from Ambon, Indonesia, there are three questionable syntypes or lectotype and paralectotypes as RMNH.PISC. 6112 (Fig. 16a₂), 26972 (Fig. 16a₃) and 27127 (Fig. 17a₄). The label on the bottles of RMNH.PISC. 6112 said “Caranx caeruleopinnatus Rüppell possibly a paralectotype, 1 ex. rem to RMNH 56972, 1 ex. to RMNH. 27127. Amboina, Saparua or Celebes (Manado), E. Indies, 1851–57. Coll. Dr. P. Bleeker, 1879.” That of RMNH.PISC. 26972 and 27127 said “Caranx opththalmotaenia (Bleeker) lectotype, 1 ex. from RMNH. 6112. G. J. van Thienen & Dr. J. Hartzfeld, Coll. Dr. P. Bleeker, 1879. Amboina, E. Indies, ca 1851.” and “Caranx fluvoguttatus (Forsskål) possibly a paralectotype, 1 ex. from RMNH. 6112. Coll. Dr. P. Bleeker, 1879. Amboina, Saparua or Celebes (Manado), E. Indies, 1851-57.”, respectively. From the original description, the syntypes have 27 or 28 dorsal-fin and 23 anal-fin soft rays, whereas the present specimens have 23 and 18 in RMNH.PISC. 6112, 22 and 17 in RMNH.PISC. 26972, and 28 and 23 in RMNH.PISC. 27127. On the basis of those fin-ray counts, RMNH.PISC. 6112 and 26972 clearly differ from the original description, and they are not included the syntypes of Carangoides ophthalmotaenia. Despite the damaged tip of caudal fin, the body length of RMNH.PISC. 27127 is estimated as 81 mm. Although this body length is rather smaller than that in the original description, the fin-ray counts and other characteristics of RMNH.PISC. 27127 agree well with the original description. From these facts, it is highly possible that RMNH.PISC. 27127 is the smaller syntype of Carangoides ophthalmotaenia. In our identification, RMNH.PISC. 27127 is Turrum fulvoguttatum and RMNH.PISC. 6112 and 26972 are Turrum coeruleopinnatum. Therefore, Carangoides ophthalmotaenia is a junior synonym of Turrum fulvoguttatum.

Figure 3 shows intraspecific differentiation in Turrum coeruleopinnatum. The genetic difference between two specimens, FRLM 35716 and 44039, indicates ca. 3.5% base substitution. Those specimens indicate some morphological differences as follows: the dorsal contour of head from tip of upper jaw to just above the eye roundly convex in FRLM 35716 (Fig. 16a₄, 128 mm SL) vs. almost straight in FRLM 44039 (Fig. 16a₅, 133 mm SL); dorsal and ventral contours of body almost the same convexity in FRLM 35716 vs. the dorsal contour more convex than the ventral in FRLM 44039; body slightly deeper (BD 54.3% SL) in FRLM 35716 than FRLM 44039 (BD 51.8% SL); and distance between pelvic-fin insertion and base of the first anal-fin spine barely shorter (18.4% SL) in FRLM 35716 than FRLM 44039 (19.4% SL). Gushiken (1983, 1984) and Senou (2002, 2013) presented a species of the former Carangoides closely similar to Turrum coeruleopinnatum as Carangoides uii (Wakiya 1924). They stated about differences between their Carangoides uii and Turrum coeruleopinnatum [as Carangoides caeruleopinnatus in Gushiken (1983, 1984) and Carangoides coeruleopinnatus in Senou (2002, 2013)] that the narrow scaly area just anterior to pectoral-fin base is short, not reaching to a horizontal through base of ventralmost pectoral-fin ray in their Carangoides uii, whereas such scaly area is rather long, reaching to the horizontal in Turrum coeruleopinnatum. Additionally, Gushiken (1983, 1984) mentioned that the first soft ray of second dorsal fin is distinctly longer than the first anal-fin soft ray in his Carangoides uii in the range of 100 to 200 mm SL, whereas the former is clearly shorter than the latter in T. coeruleopinnatum. Senou (2002, 2013) also indicate that the anteriormost part of second dorsal fin is elongated and longer than head, and such elongated part is longer than the elongated part of anal fin in juvenile his Carangoides uii; whereas the anteriormost part of second dorsal fin is shorter than head and such elongated part is shorter than the elongated part of anal fin in juvenile T. coeruleopinnatum. The specimen of FRLM 35716 is apparently similar to the drawings and photograph of Gushiken’s (1983: fig. 64, 1984: pl. 141H) and Senou’s (2002: 808, 2013: 899) Carangoides uii in round forehead and equally convex dorsal and ventral contours of body. However, the scaly area just anterior to pectoral-fin base in FRLM 35716 is long, extending downward beyond a horizontal line through base of ventralmost pectoral-fin ray, so the specimen disagrees with their Carangoides uii. Carangoides uii was established by Wakiya (1924) as Caranx (Citula) uii Wakiya 1924 on the basis of two specimens collected from Wakayama, Japan. He stated that the first soft rays of dorsal and anal fins are almost equally prolonged, and the original figure of the holotype (Wakiya 1924: pl. 22, fig. 1) shows almost straight contour of forehead. Moreover, the image of the holotype [(c) Field Museum of Natural History. CC BY-NC. https://collections-zoology.fieldmuseum.org/catalogue/638496 (accessed on 14 June 2021)] indicates the long scaly area just anterior to pectoral-fin base, extending below a horizontal through the base of ventralmost pectoral-fin ray, and more convex dorsal contour of body than the ventral. The specimens similar to Caranx (Citula) uii were often collected and identified as Turrum coeruleopinnatum (Fig. 16f). Therefore, it is appropriate that Caranx (Citula) uii is a junior synonym of Turrum coeruleopinnatum at present. In addition, the drawings of Carangoides uii given by Allen and Swainston (1988), Randall et al. (1990) and Allen (1997) show a long filamentous anterior dorsal-fin soft ray, a short anterior anal-fin soft ray and almost straight forehead. The fish in the underwater photograph of Carangoides uii published by Kuiter (1997) is similar to the present specimen of FRLM 51147 (Fig. 16f) in having equally long filamentous soft rays of second dorsal and anal fins and almost straight forehead. On the other hand, the present Turrum coeruleopinnatum includes two or more groups which are genetically and morphologically different. However, in the present stage, it is very difficult to divide them clearly and to give them correct species names.

Because the original specific names of “coeruleopinnatus” and “fulvoguttatus” are Latin masculine adjectives, these specific names must agree in gender with the generic name (ICZN 1999, Art. 31.2). Since the generic name Turrum is neuter, those specific names change to “coeruleopinnatum” and “fulvoguttatum”. On the other hand, the original specific name of “gymnostethus” need not agree in gender with the generic name because the specific name is a Greek masculine adjective (ICZN 1999, Art. 31.2.3; Kottelat, personal communication).

Craterognathus gen nov.

(Sublade G, Fig. 18)

Fig. 18

Craterognathus plagiotaenia, the species of monotypic genus Craterognathus gen. nov. (Subclade G). a₁ RMNH 26974, lectotype of Carangoides plagiotaenia, 66 mm SL, Ambon, Indonesia; a₂ FRLM 37736, 291 mm SL, Okinawa, Japan (fresh condition); a₃ ZIN 2662, syntype of Caranx brevicarinatus, 264 mm SL, Red Sea (photographed by M. Matsunuma)

Type species. Carangoides plagiotaenia Bleeker 1857 [= Craterognathus plagiotaenia (Bleeker 1857)].

Included species. Craterognathus plagiotaenia (Bleeker 1857).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, somewhat slender; dorsal contour of upper lip around symphysis gradually ascending (Arcuate type; Fig. 4g); lower jaw stout, distinctly projecting anteriorly, mouth superior; jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth band chevron-shaped or triangular (Fig. 5a, g); adipose eyelid poorly developed; breast completely scaly except for naked pectoral-fin base (Fig. 6g); curved and anterior straight parts of lateral line with cycloid scales (79–99 + 13–29), the posterior straight part with 10–20 scutes; first dorsal fin with interspinous membranes, with VIII spines, second dorsal fin with 22–24 soft rays; anal-fin soft rays 18–20; anterior second dorsal and anal fins not falcate; no finlets; both neural and hemal spines of ninth caudal vertebra expanding, Oar-shape (Fig. 7a); total gill rakers 27–40; snout short, 7.1–10.7% SL (Table 2, Fig. 10a); suborbital depth 3.4–5.0% SL (Fig. 10b, Table 2).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Snout longer than eye diameter. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 11th–16th dorsal-fin soft ray. Second dorsal-fin base slightly longer than soft portion of anal-fin base; first and second anal-fin spines developed, separated from the third, and not embedded. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen in last four or five precaudal vertebrae.

Distribution. Indo-Pacific from the Red Sea and the eastern coast of Africa to Fiji and Samoa, except Persian Gulf (Smith-Vaniz 1983, 1986, 1999).

Etymology. The name “Craterognathus” is derived from “kratero”, the Greek meaning “strong”, and “gnathos”, the Greek meaning “jaw”, in reference to the characteristic stout lower jaw.

Gender. Masculine.

Comparison. Craterognathus gen nov. is distinguishable from the former Carangoides and other genera, except for Parastromateus and two species of Selene (S. brownii and S. setapinnis) by a completely scaly breast (Fig. 6). Craterognathus also can be distinguished from Parastoromateus in having Oar-shape neural and hemal spines of ninth caudal vertebra (Fig. 7a; vs. neural and hemal spines spinous, Fig. 7c), and from Selene in having a chevron-shaped or triangular vomerine tooth patch (Fig. 5a, f; vs. diamond- or cross-shaped, Fig. 5d, e). Although Flavocaranx and Carangoides praeustus have a very small naked area anteriorly on breast, Craterognathus differs from Flavocaranx in having three pterygiophores between the first and second hemal spines (vs. four), and from Carangoides in having a chevron-shaped or triangular vomerine tooth patch (vs. anchor-shaped, Fig. 5g).

Atropus Oken 1817

(Subclade H, Fig. 19)

Fig. 19

Five included species of the genus Atropus (Subclade H). a Atropus armatus, FRLM 32079, 200 mm SL, Phuket, Thailand (fresh condition); b Atropus atropos, type species, FRLM 44041, 134 mm SL, Port Dickson, Malaysia (fresh condition); c Atropus aurochs, FRLM 14575, 356 mm SL, Ambon, Indonesia (fresh condition); d Atropus hedlandensis, FRLM 31421, 167 mm SL, Mie, Japan (fresh condition); e Atropus mentalis, FRLM 39999, 185 mm SL, Bitung, Indonesia (fresh condition)

Atropus Oken 1817: 1182 (type species: Brama atropos Bloch and Schneider 1801)

Olistus Cuvier 1829: 209 (type species: Olistus malabaricus Cuvier in Cuvier and Valenciennes 1833)

Leioglossus Bleeker 1851b: 343, 352, 367 (type species: Leioglossus carangoides Bleeker 1851b)

Ulua Jordan and Snyder 1908: 39 (type species: Ulua richardsoni Jordan and Snyder 1908)

Type species. Brama atropos Bloch and Schneider 1801 [= Atropus atropos (Bloch and Schneider 1801)].

Included species. Atropus armatus (Forsskål in Niebuhr 1775), Atropus atropos (Bloch and Schneider 1801), Atropus aurochs (Ogilby 1915), Atropus hedlandensis (Whitley 1934) and Atropus mentalis (Cuvier in Cuvier and Valenciennes 1833).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, deep, disk-like or rhomboidal; dorsal contour of upper lip around symphysis roundly convex (Round chevron type; Fig. 4h); jaws, vomer and palatines with villiform teeth forming bands except for Atropus atropos in which both jaws with conical teeth arranged biserially (or sometimes a uniserial outer row of conical teeth and an inner villiform tooth band); vomerine tooth patch chevron-shaped or triangular (Fig. 5a, f); adipose eyelid poorly developed; naked area on breast extending beyond base of last pelvic-fin soft ray and connected with naked pectoral-fin base (Fig. 6h_1–4); posterior curved part of lateral line covered by cycloid scales in Atropus armatus, Atropus hedlandensis and some Atropus mentalis, by scutes in Atropus atropos, Atropus aurochs and some Atropus mentalis; anterior straight part of lateral line covered by cycloid scales in Atropus armatus, most of Atropus hedlandensis and some Atropus mentalis, by scutes in Atropus atropos, Atropus aurochs, some Atropus hedlandensis and most of Atropus mentalis; first dorsal-fin with interspinous membranes, with VIII (rarely VII) spines, second dorsal fin with 19–23 soft rays; anal-fin soft rays 16–20; anterior second dorsal and anal fins falcate; soft rays of dorsal and anal fins elongate in some species; no finlets; both neural and hemal spines of ninth caudal vertebra expanding, Oar-shape (Fig. 7a); ultimate neural and hemal pterygiophores adjacent to neural and hemal spines of tenth caudal vertebra, respectively (Fig. 8c), except for Atropus hedlandensis; total gill rakers 20–89; snout short 7.4–10.7% SL (Table 2, Fig. 10a); suborbital narrow, 3.4–6.3% SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth superior, lower jaw strongly protruding in some species. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of fourth–14th dorsal-fin soft ray. Second dorsal-fin base slightly longer than soft part of anal-fin base; first and second anal-fin spines developed, separated from the third spine, and not embedded. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen in last four or five (Atropus hedlandensis) precaudal vertebrae.

Distribution. Indo-Pacific from the Red Sea and the eastern coast of Africa to Samoa (Smith-Vaniz 1983, 1999).

Gender. Masculine.

Comparison. The genus Atropus is distinguished from Alectis, Scyris and Parastromateus by having interspinous membranes of the first dorsal fin (vs. lacking the membranes), from Euprepocaranx, Paraselene and Selene by having a chevron-shaped or triangular vomerine tooth band (vs. diamond-, cross- or four-pointed-star-shaped; Fig. 5), from Craterognathus gen. nov., Parastromateus, Flavocaranx gen. nov., Ferdauia, Uraspis, Carangichthys and Carangoides by having a thoracic naked area extending posteriorly beyond pelvic-fin insertion (vs. thoracic naked area absent or not extending beyond pelvic-fin insertion; Fig. 6), from Turrum by having a shorter snout (7.4–10.7% SL vs. 10.9–14.3% SL; Fig. 10a, Table 2), and Platycaranx gen. nov. by having a wide protrusion of upper-jaw symphysis (vs. narrow protrusion; Fig. 4).

Remarks. The previous Carangoides armatus (Forsskål in Niebuhr 1775) (= Atropus armatus), Carangoides hedlandensis (Whitley 1934) (= Atropus hedlandensis), Atropus atropus and the previous Ulua mentalis (Cuvier in Cuvier and Valenciennes 1833) (= Atropus mentalis) compose a single subclade (Subclade H, Fig. 3), and they form an independent genus because they share the unique combination of morphological characters. Atropus is the oldest name among the nominal genera which were established based on the above species as type species. Atropus aurochs formerly known as Ulua aurochs (Ogilby 1915) is included in Atropus because its morphological characters agree well with the diagnosis of the genus.

The nominal genus Olistus was established by Cuvier (1829), subsequently Jordan (1917) assigned Olistus malabaricus Cuvier in Cuvier and Valenciennes 1833 as its type species. Because O. malabaricus and a different species Scomber malabaricus Bloch and Schneider 1801 have been once included in the same genus Carangoides, Carangoides malabaricus (Cuvier in Cuvier and Valenciennes 1833) became a junior secondary homonym of Carangoides malabaricus (Bloch and Schneider 1801) and the former name was subjectively invalid (ICZN 1999; Art. 57.3.1). However, the present study assigns O. malabaricus and Scomber marabaricus to different genera (Atropus for the former and Platycaranx gen. nov. for the latter), then the homonymy is canceled at present, O. malabaricus being resurrected. At the same time with O. malabaricus, Caranx ciliaris was established by Cuvier in Cuvier and Valenciennes (1833) based on a single specimen, MNHN A-5547, collected from Puducherry, India [other material, namely the specimens sent from Java, Indonesia by Kuhl and Van Hasselt and Russell’s (1803) drawing (pl. 151) are not included in its primary types as indicated by Smith-Vaniz et al. (1979)]. Because the holotype is a dried specimen, severely damaged and lacked gill arches, accurate identification of the specimen seemed difficult. Hence this nominal species has been treated as nomen dubium by Smith-Vaniz et al. (1979) or a questionable junior synonym of Carangoides armatus (= Atropus armatus) by Kottelat (2013). The present detailed observation revealed that the holotype, MNHN A-5547 (Fig. 20), is conspecific with O. malabaricus because it has round dorsal contour of head from snout to occiput, cycloid scales along anterior straight part of lateral line, 20 and 16 second dorsal- and anal-fin soft rays, respectively, and smaller eye (ED/SL ca. 8.7%). Olistus hedlandensis Whitley 1934 is considered to be conspecific with O. malabaricus by many authors (e.g., Williams et al. 1980; Gushiken 1983; Smith-Vaniz 1999). We also confirm this synonymy based on the original descriptions and figures of both nominal species. Accordingly, three nominal species, O. malabaricus, Caranx ciliaris and O. hedlandensis are the same species. The former two nominal species, however, have not been used as valid names since 1900 as far as we know, whereas the latter (as Carangoides hedlandensis) has been used as the presumed valid name by many authors over a long period of time (Smith-Vaniz et al. 1979; Williams et al. 1980; Gushiken 1983, 1984, 1988; Smith-Vaniz 1983, 1986, 1999; Gloerfelt-Tarp and Kailola 1984; Laroche et al. 1984; Sainsbury et al. 1985; Kijima et al. 1986; Allen and Swainston 1988; Randall et al. 1990; Mok 1993; Allen 1997; Rao et al. 2000; Senou 2002; Shao and Chen 2003; Rao 2004; Randall 2005; Hoese and Gates 2006; Satapoomin 2007; Springer and Smith-Vaniz 2008; Kimura 2009, 2011, 2013, 2017; White et al. 2013; Santini and Carnevale 2014; Hata 2019). Because Art. 23.9.1.1 and 23.9.1.2 of ICZN (1999) are satisfied, the youngest name, Atropus hedlandensis is valid for the species in accordance with Art. 23.9.2 of ICZN (1999).

Fig. 20

Holotype of Caranx ciliaris (a synonym of Atropus hedlandensis), MNHN A-5547, ca. 94 mm SL, Puducherry, India

Caranx citula was also described by Cuvier in Cuvier and Valenciennes (1833) based on a single specimen, and the specimen of MNHN A-6182 (Fig. 21a) can be recognized as the holotype based on body size and counts of second dorsal- and anal-fin soft rays. The present reidentification of the type series indicates that the holotype MNHN A-6182 and the paratype, MNHN A-6181 (Fig. 21b) are conspecific with Atropus armatus, the paratype MNHN A-6102 (Fig. 21c) with Atropus hedlandensis, and the paratype MNHN B-2881 (Fig. 21d) with Atropus mentalis as already stated by Smith-Vaniz et al. (1979). Therefore, Caranx citula is a junior synonym of Atropus armatus.

Fig. 21

Types of Caranx citula. a₁ MNHN A-6182, holotype, 162.3 mm SL, Red Sea (Atropus armatus); a₂ MNHN A-6181, paratype, 167 mm SL, Red Sea (Atropus armatus); b MNHN A-6102, paratype, 158 mm SL, New Guinea (Atropus hedlandensis); c MNHM B-2881, paratype of Carangoides citula, 201 mm SL, Red Sea (Atropus mentalis)

The type species of the nominal genus Ulua, Ulua richardosoni, has been considered as a junior synonym of Caranx mentalis Cuvier in Cuvier and Valenciennes 1833 (Gushiken 1983; Smith-Vaniz 1999). The morphological characters of the holotype described and figured by Jordan and Snyder (1908) agree well with those of Caranx mentalis. Therefore, those two nominal species are conspecific, Ulua being a junior synonym of Atropus.

Leioglossus carangoides, the type species of Leioglossus, was described by Bleeker (1851b) based on four syntypes of 210–370 mm in body length. However, the syntypes and four other conspecific specimens (probably Bleeker specimens) once put together into a bottle of RMNH.PISC. 6091. Of these, one specimen of 183 mm SL (ca. 235 mm TL) was extracted as RMNH.PISC. 27642 to attempt designating the lectotype. Body sizes of those eight specimens range from 73 mm to 270 mm SL (from 94 to more than 380 mm TL), and five specimens fall within the size range of the syntypes (210–370 mm in body length). According to the original description, syntypes has 18 or 19 branched pectoral-fin rays, and of those five specimens, four have 18 or 19 branched rays, but one 20 rays. As a result, the syntypes are identified into four specimens of 183 mm SL (RMNH.PISC. 27642), 270 mm SL RMNH.PISC. 6091), 187 mm SL (RMNH.PISC. 6091) and 155 mm SL (RMNH.PISC. 6091) (Fig. 22). Because those syntypes are identifiable with Atropus mentalis, Leioglossus is a junior synonym of Atropus.

Fig. 22

Syntypes of Leioglossus carangoides (a junior synonym of Atropus mentalis) from Indonesia. a₁ RMNH.PISC. 27642, 183 mm SL; a₂ RMNH.PISC. 6091, 270 mm SL; a₃ RMNH.PISC. 6091, 187 mm SL; a₄ RMNH.PISC. 6091, 155 mm SL

Flavocaranx gen. nov.

(Sublade I, Fig. 23)

Fig. 23

Flavocaranx bajad, the species of monotypic genus Flavocaranx gen. nov. (Subclade I). a₁ HUMZ 165337, 183 mm SL, Red Sea, Saudi Arabia; a₂ MNHN A5752, syntype of Caranx auroguttatus, 372 mm SL, Red Sea (dry specimen)

Type species. Scomber ferdau bajad Fabricius in Niebuhr 1775 [= Flavocaranx bajad (Fabricius in Niebuhr 1775)].

Included species. Flavocaranx bajad (Fabricius in Niebuhr 1775).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, somewhat slender, oblong; dorsal contour of upper lip around symphysis roundly protruding upward (Round chevron type; Fig. 4i); jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch chevron-shaped (Fig. 5a); adipose eyelid poorly developed; very small naked thoracic area present anteriorly along ventral midline (Fig. 6i); curved part and anterior straight part of lateral line with cycloid scales (64–82 + 22–36), the posterior straight part with 23–30 scutes; first dorsal fin with interspinous membranes, with VII–VIII spines, second dorsal fin with 24–29 soft rays; anal-fin soft rays 21–24; anterior second dorsal and anal fins not falcate; no finlets; both neural and hemal spines of ninth caudal vertebra expanding, Oar-shape (Fig. 7a); six neural spines between pterygiophores supporting first and last dorsal-fin spines; four (or rarely five) pterygiophores between first and second hemal spines; total gill rakers 26–30 (25–33 in Smith-Vaniz 1999); snout length 10.6–12.6% SL (Table 2, Fig. 10a); suborbital somewhat narrow, 4.1–6.0% SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Dorsal contour of head almost straight from snout to parietal region, then roundly convex. Mouth almost terminal; snout longer than eye diameter. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of tenth–15th dorsal-fin soft ray. Second dorsal-fin base slightly longer than soft portion of anal-fin base; first and second anal-fin spines developed, separated from the third, and not embedded. Vertebrae 10 + 14; inferior foramen on last four or five precaudal vertebrae.

Distribution. Indo-West Pacific, from the Red Sea to the Arabian Gulf and Pakistan, and Thailand to New Britain (Smith-Vaniz 1999; Bogorodsky et al. 2017).

Etymology. The generic name “Flavocaranx” derived from “flavo”, the Latin meaning “yellow”, and “Caranx”, the type genus of Carangidae, in reference to yellowish body color.

Gender. Masculine.

Comparison. Flavocaranx gen nov. is distinguishable from Alectis, Scyris and Parastromateus by having interspinous membranes of the first dorsal fin (vs. lacking the membranes), from Euprepocaranx gen. nov., Paraselene, Selene and Carangoides by having a chevron-shaped vomerine tooth band (vs. diamond, cross- or four-pointed-star-shaped in Euprepocaranx, Paraselene and Selene; anchor-shaped in Carangoides; Fig. 5), from Turrum, Craterognathus gen. nov., Atropus, Platycaranx gen. nov., Parastromateus, Ferdauia, Uraspis and Carangichthys by having a narrow ventral thoracic naked area (vs. no thoracic naked area in Craterognathus and Parastromateus; broad thoracic naked area extending to or beyond pelvic-fin insertion in other genera; Fig. 6). Flavocaranx has four or five pterygiophores between the first and second hemal spines. On the Basis of this character, Flavocaranx can be distinguished from Alectis, Scyris, Euprepocaranx, Paraselene, Selene, Craterognathus, Atropus, Platycaranx, Parastromateus, Ferdauia, Uraspis, Carangichthys and Carangoides, which have three pterygiophores.

Platycaranx gen. nov.

(Subclade J, Fig. 24)

Fig. 24

Three included species of the genus Platycaranx gen. nov. (Subclade J), a Platycaranx chrysophrys, b Platycaranx malabaricus, type species and c Platycaranx talamparoides. a₁ MNHN A-560, holotype of Caranx chrysophys, 246 mm SL, Seychelles; a₂ FRLM 44037, 296 mm SL, Port Dickson, Malaysia (fresh condition); a₃ RMNH 6113, syntype of Carangoides chrysophryoides, 270 mm SL, Jakarta, Indonesia; b₁ FRLM 35763, 110 mm SL, Ambon, Indonesia (fresh condition); b₂ RMNH.PISC. 6098, lectotype of Carangoides talamparah, 137 mm SL, Jakarta, Indonesia; c₁ RMNH.PISC. 6099, lectotype of Carangoides talamparoides, 198 mm SL, Sibogha, Indonesia (right side, mirror image); c₂ FRLM 35697, 166 mm SL, Ambon, Indonesia (fresh condition)

Type species. Scomber malabaricus Bloch and Schneider 1801 [= Platycaranx malabaricus (Bloch and Schneider 1801)].

Included species. Platycaranx chrysophrys (Cuvier in Cuvier and Valenciennes 1833), Platycaranx malabaricus (Bloch and Schneider 1801) and Platycaranx talamparoides (Bleeker 1852).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, deep, rhomboidal; dorsal edge of upper lip around symphysis roundly protruding upward (Round chevron type), protrusion of upper-jaw symphysis narrow (Fig. 4j); jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch chevron-shaped or triangular (Fig. 5a, f); adipose eyelid poorly developed; naked area on breast extending beyond base of last pelvic-fin soft ray and connected with naked pectoral-fin base (Fig. 6j); curved part and usually anterior straight part of lateral line with cycloid scales (61–100 + 0–16), the posterior straight part with 16–32 scutes; first dorsal fin spines with interspinous membranes, with VII–VIII spines, second dorsal fin with 18–23 soft rays; anal-fin soft rays 15–20; no finlets; both neural and hemal spines of ninth caudal vertebra expanding, Oar-shape (Fig. 7a); ultimate neural and hemal pterygiophores adjacent to neural and hemal spines of tenth caudal vertebra, respectively (Fig. 8d); total gill rakers 21–38; snout moderate in length, 11.1–14.5% SL (Table 2, Fig. 10a); suborbital depth 5.0–8.4 % SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth almost terminal and snout tip truncate in Platycaranx chrysophrys, or mouth slightly superior, lower jaw slightly protruding in Platycaranx malabaricus and Platycaranx talamparoides. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 11th–15th dorsal-fin soft ray. Second dorsal-fin base slightly longer than anal-fin base; first and second anal-fin spines small, separated from the third spine, and not embedded in adults. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen in last five precaudal vertebrae.

Distribution. Indo-West Pacific.

Etymology. The name “Platycaranx” is derived from “platys”, the Greek meaning “flat”, and Caranx, the type genus of Carangidae, in reference to the highly compressed body.

Gender. Masculine.

Comparison. Platycaranx gen. nov. is distinguishable from the former Carangoides and other genera, except for Euprepocaranx by round and narrow protrusion of the upper lip around symphysis (Fig. 4) and distinguishable from Euprepocaranx, by having a chevron-shaped or triangular vomerine tooth band (vs. diamond-shaped with a rounded posterior corner; Fig. 5). Platycaranx can be also distinguished from Alectis, Scyris and Parastromateus by having interspinous membranes of first dorsal fin (vs. lacking the membranes), from Paraselene and Selene by having chevron-shaped or triangular vomerine tooth patch (vs. diamond-, cross- or four-pointed-star-shaped; Fig. 5), from Craterognathus gen. nov., Parastromateus, Flavocaranx gen. nov., Ferdauia, Uraspis, Carangichthys and Carangoides by having thoracic naked area extending posteriorly beyond base of last pelvic-fin soft ray (vs. absent or not extending beyond pelvic-fin insertion; Fig. 6) and from Atropus by having a somewhat longer snout (11.1–14.2% SL vs. 7.4–10.7% SL; Fig. 10a, Table 2).

Parastromateus Bleeker 1864

(Subclade K, Fig. 25)

Fig. 25

Parastromateus niger, the species of monotypic genus Parastromateus (Subclade K). a₁ FRLM 40277, 245 mm SL, Kagoshima, Japan (fresh condition); a₂ USNM 55914, holotype of Citula hall, 49 mm SL, Luzon, Philippines (photographed by Y. Hibino)

Apolectus Cuvier in Cuvier and Valenciennes 1832: 438 (type species: Apolectus stromateus Cuvier in Cuvier and Valenciennes 1832)

Parastromateus Bleeker 1864: 174 (type species: Stromateus niger Bloch 1795)

Formio Whitley in McCulloch 1929: 193 (type species: Apolectus stromateus)

Hildebrandella Nichols 1950: 20 (type species: Citula halli Evermann and Seale 1907)

Type species. Stromateus niger Bloch 1795 [= Parastromateus niger (Bloch 1795)].

Included species. Parastromateus niger (Bloch 1795).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, deep, disk-like; dorsal contour of upper lip around symphysis gradually ascending (Arcuate type; Fig. 4k); jaws with posteriorly curved long conical teeth arranged uniserially; vomer and palatines edentate; adipose eyelid poorly developed; breast completely scaly except for naked pectoral-fin base (Fig. 6k); curved part and anterior straight part of lateral line with cycloid scales (60–69 + 6–9), the posterior straight part with 9–20 scutes; first dorsal fin spines without interspinous membranes, with IV (–V in Smith-Vaniz 1999) spines, second dorsal fin with 41–44 soft rays; anal-fin soft rays 35–39; total number of second dorsal- and anal-fin soft rays 78–83 (Table 1, Fig. 9); no pelvic fins in adults; first dorsal fin and free anal-fin spines embedded in adults; second dorsal and anal fins very deep but not falcate; no finlets; both neural and hemal spines of ninth caudal vertebra Spine-shape (Fig. 7c); eight to ten pterygiophores between first and second hemal spines; total number of gill rakers 19–22; snout relatively short, 8.9–9.6% SL (Table 2, Fig. 10a); suborbital relatively narrow, 2.7–5.3% SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth terminal. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 32nd–35th dorsal-fin soft ray. Second dorsal-fin base slightly longer than anal-fin base. Three to five neural spines between pterygiophores supporting first and last dorsal-fin spines. Vertebrae 10 + 14; inferior foramen in last three precaudal vertebrae.

Distribution. Indo-West Pacific, from the eastern coast of Africa to Japan and the northeastern coast of Australia.

Gender. Masculine.

Comparison. Parastromateus is distinguishable from the former Carangoides and other genera by numerous total number of dorsal- and anal-fin soft rays (78–83 vs. not more than 61; Table 1, Fig. 9) or no pelvic fins in adults.

Remarks. The systematic position of Parastromateus niger has been confused due to its morphological peculiarity and similar appearance to stromateids. The present species was first described by Bloch (1795) as a stromateid species, Stromateus niger. Thereafter, Bleeker (1864) established the genus Parastromateus for Stromateus niger as type species. Aside from this, Cuvier in Cuvier and Valenciennes (1832) originally described the genus Apolectus and Apolectus stromateus. However, nominal genus Apolectus Cuvier in Cuvier and Valenciennes 1832 is objectively invalid because it was preoccupied by Apolectus Bennett 1831 (junior synonym of Scomberomorus Lacepède 1801), and Whitley in McCulloch (1929) proposed Formio as a replaced name of Apolectus Cuvier in Cuvier and Valenciennes 1832 (Fricke et al. 2021). Whitley in McCulloch (1929) considered that Apolectus stromateus and Stromateus niger are conspecific, and he synonymized them under Formio niger (Bloch 1795), and established family Formidae for the species. Nichols (1950) established the genus Hildebrandella based on Citula halli (Fig. 25a₂). He doubted that this nominal species belonged to the family Carangidae because of similarity to ephippid fish Platax Cuvier 1816 in appearance. Both Apolectus stromateus (type species of Aplolectus and its replaced name Formio) and Citula halli (type species of Hildebrandella) are identifiable with juvenile Parastromateus niger from the original figures of the former two nominal species and their second dorsal- and anal-fin counts as indicated by Smith-Vaniz (1999). Therefore, both Formio and Hildebrandella are junior synonyms of Parastromateus.

After establishment of Formiidae, Parastromateus niger has been considered to belong to the family for a relatively long time, for example, Abe and Bathia (1974) placed the species in Formiidae (as Formionidae).

Smith-Vaniz (1984) demonstrated that Parastromateus niger is included in the family Carangidae because the first and second anal-fin spines are separated from the third and form free spines. Hilton et al. (2010) has conducted an osteological study on branchial arches of the species in detail, and they suggested that the affinity between Parastromateus and Hemicaranx. Based on our molecular analysis shown in Fig. 1, Hemicaranx indicates a sister relationship with Chloroscombrus of Clade 2. Therefore, there is no close relationship between Parastromateus and Hemicaranx. On the other hand, the present result of molecular phylogeny indicates that Parastromateus is closely related with the genera Ferdauia, Uraspis and Carangichthys (Fig. 3).

Ferdauia Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927

(Subclade L, Fig. 26)

Fig. 26

Two included species of the genus Ferdauia (Subclade L), a Ferdauia ferdau and b Ferdauia orthogrammus, type species. a₁ FRLM 37301, 207 mm SL, Mie, Japan (fresh condition); a₂ RMNH.PISC 6101, syntype of Carangoides hemigymnostethus, 230 mm SL, Jakarta, Indonesia; a₃ RMNH.PISC 26969, syntype or Bleeker specimen of Carangoides hemigymnostethus, 202 mm SL, Jakarta, Indonesia (or Timor); b₁ FRLM 41754, 248 mm SL, Mie, Japan (fresh condition); b₂ SAIAB 1, holotype of Carangoides nitidus, 207 mm SL, Shimoni, Kenya (photographed by H. Hata)

Ferdauia Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927: 662 (Type species: Carangoides jordani Nichols 1922)

Type species. Carangoides jordani Nichols 1922, a junior synonym of Ferdauia orthogrammus (Jordan and Gilbert 1882a).

Included species. Ferdauia ferdau (Fabricius in Niebuhr 1775), Ferdauia orthogrammus (Jordan and Gilbert 1882a).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, deep, elongate-oval; dorsal contour of upper lip around symphysis roundly protruding upward (Round chevron type; Fig. 4l); jaws, vomer and palatines with villiform teeth forming bands; vomerine tooth patch triangular (Fig. 5f); adipose eyelid poorly developed; naked area on breast not connecting with naked area on pectoral-fin base, reaching posteriorly to pelvic-fin insertion, but not extending beyond it (Fig. 6l_{1, 2}); curved part and anterior straight part of lateral line with cycloid scales (81–102 + 6–34), the posterior straight part with 19–41 scutes; first dorsal fin with interspinous membranes, with VI–VIII spines, second dorsal fin with 26–34 soft rays; anal-fin soft rays 21–27; total number of second dorsal- and anal-fin soft rays 55–61 (Table 1, Fig. 9); anterior second dorsal and anal fins falcate; no finlets; free anal-fin spines short, embedded in adults; caudal fin somewhat large, UCF/BD 69.8–84.2%; both neural and hemal spines of ninth caudal vertebra expanding, Blade-shape (Fig. 7b); five neural spines between pterygiophores supporting first and last dorsal-fin spines; five or six pterygiophores between first and second hemal spines; total gill rakers 24–33; snout longer than eye diameter, 8.3–11.3% SL (Table 2, Fig. 10a); suborbital relatively narrow, 3.3–5.1% SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth almost terminal, lower jaw slightly protruding. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, straight part beginning below base of 14th–23th dorsal-fin soft ray. Second dorsal-fin base slightly longer than soft portion of anal-fin base; first and second anal-fin spines small, separated from the third spine. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10 + 14; inferior foramen on last three precaudal vertebrae.

Distribution. Indo-Pacific.

Gender. Feminine.

Comparison. The genus Ferdauia is distinguishable from the former Carangoides and other genera, except for Turrum and Parastromateus by rather numerous total number of dorsal- and anal-fin soft rays (55–61; Fig. 9, Table 1), and from Alectis, Scyris and Parastromateus by having interspinous membranes of first dorsal fin (vs. lacking the membranes). Ferdauia is also distinguished from Craterognathus and Parastromateus by having a naked thoracic area (vs. no naked area; Fig. 6), from Alectis, Scyris, Euprepocaranx, Paraselene, Turrum, Atropus, Flavocaranx and Platycaranx by the naked area just reaching to pelvic-fin insertion, but not extending it (vs. naked area not reaching to pelvic-fin insertion in Flavocaranx or the area extending posteriorly beyond pelvic-fin insertion in the other genera; Fig. 6), from Selene and Carangoides by having a triangular vomerine tooth patch (cross-shaped in Selene and anchor-shaped in Carangoides; Fig. 5), and from Uraspis and Carangichthys by having villiform tooth bands on jaws (vs. small conical teeth arranged uniserially or biserially) and Blade-shape neural and hemal spines of ninth caudal vertebra (vs. Spine-shape, Fig. 7).

Remarks. The nominal genus Ferdauia has been treated as invalid after its establishment except for Whitley (1947, 1951). Gushiken (1983), Gunn (1990) and Smith-Vaniz (1999) considered the nominal genus as a junior synonym of Carangoides without explanation. In the original description, Jordan, Evermann and Wakiya in Jordan et al. (1927) stated that the genus included three species, Ferdauia jordani (Nichols 1922), Ferdauia evermanni (Nichols 1921) and Ferdauia orthogrammus, and differed from Carangoides in the naked breast. But they did not indicate the concrete difference.

Ferdauia orthogrammus and Ferdauia ferdau composing Subclade L in Fig. 3, have a naked thoracic area not extending beyond pelvic-fin insertion as stated above. This character is only seen in those two species in the previous Carangoides species except for Craterognathus plagiotaenia, Flavocaranx bajad and Carangoides praeustus. The latter three species have no or very small thoracic naked area. Therefore, the two species of Ferdauia can be distinguished from previous Carangoides species by the shape of thoracic naked area (Fig. 6).

The characteristics of Caranx orthogrammus Jordan and Gilbert 1882a, Carangoides jordani and Carangoides gymnostethoides evermanni Nichols 1921 described in their original descriptions and photographs of holotype and paratype of the latter nominal species agree well with those of Ferdauia orthogrammus, and the differences among them are small and considered to be within the range of intraspecific variation. Therefore, Carangoides jordani and Carangoides gymnostethoides evermanni can be synonymized under Caranx orthogrammus (= Ferdauia orthogrammus).

Carangoides hemigymnostethus, a junior synonym of Ferdauia ferdau, was described by Bleeker (1851b) based on three specimens (280–310 mm in body length). However, only two “types”, RMNH.PISC 6101 (Fig. 26a₂) and 26969 (extracted from RMNH.PISC 6101, Fig. 26a₃) exist. The latter specimen lacking tip of upper caudal-fin lobe can be estimated as 260 mm TL and outside the range of body length of the types. On the other hand, RMNH.PISC 6101 is estimated as ca. 294 mm in TL and agrees with the size range of the types. The label on the bottle of RMNH.PISC 6101 says “Caranx ferdau (Forskål) Carangoides hemigymnostethus Blkr, of which the lectotype, 1 ex. removed to RMNH. 26969.” But the label on the bottle of RMNH.PISC 26969 says “Caranx ferdau (Forsskål) possibly a paralectotype of Carangoides hemigymnostethus Bleeker; from RMNH 6101.” Thus, the present specimen of RMNH.PISC 26969 is probably not included in the syntypes of Carangoides hemigymnostethus.

Concerning number of pterygiophores between first and second hemal spines, Springer and Smith-Vaniz (2008) indicated that two species of Ferdauia (as Carangoides ferdau and Carangoides orthogrammus) have five or six pterygiophores. Our results are the same as theirs.

Uraspis Bleeker 1855

(Subclade M, Fig. 27)

Fig. 27

Two included species of the genus Uraspis (Subclade L), a Uraspis helvol and b Uraspis uraspis, type species. a₁ BMNH 1963.3.27.4, holotype of Scomber helvolus, 278 mm SL, Ascension Island (© the Trustees of the Natural History Museum, photographed by L. Goodayle); a₂ FRLM 39389, 221 mm SL, Mie, Japan (fresh condition); a₃ CAS-ICH 305, holotype of Leucoglossa albilinguis, 263 mm SL, Hawaii, USA (photographed by Y. Hibino); a₄ ANSP 68453, holotype of Uraspis heidi, 215 mm SL, New Jersey, USA (photographed by M. Sabaj); a₅ CAS-ICH 307, holotype of Uraspis reversa, 265 mm SL, Hawaii, USA (photographed by Y. Hibino); a₆ MCZ 16703, syntype of Caranx secundus, 259 mm SL, Cuba (photographed by H. Hata); b₁ FRLM 30714, 195 mm SL, Phuket, Thailand (fresh condition); b₂ CAS-SU 27962, holotype of Leucoglossa herklotsi, 185 mm SL, Hong Kong, China (photographed by Y. Hibino); b₃ USNM 98820, holotype of Uraspis pectoralis, 168 mm SL, Manila, Philippines (photographed by Y. Hibino)

Uraspis Bleeker 1855: 417, 418 (type species Uraspis carangoides Bleeker 1855)

Leucoglossa Jordan and Evermann in Jordan, Evermann and Tanaka 1927: 660 (type species: Leucoglossa candens Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927)

Zamora Whitley 1931: 108 (type species Caranx hullianus McCulloch 1909)

Platyuraspis Fowler 1938b: 149 (type species Uraspis heidi Fowler 1938b)

Bassetina Whitley 1941: 30 (type species Caranx hullianus McCulloch 1909)

Type species. Uraspis carangoides Bleeker 1855, a junior synonym of Uraspis uraspis (Günther 1860).

Included species. Uraspis helvola (Forster in Bloch and Schneider 1801) and Uraspis uraspis (Günther 1860).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, deep, oblong; dorsal contour of upper lip around symphysis gradually ascending (Arcuate type; Fig. 4m); jaws with small but stout conical teeth uniserially or biserially arranged; vomer and palatines edentate; adipose eyelid poorly developed; naked area on breast reaching posteriorly to pelvic-fin insertion, but not extending beyond it (Fig. 6m); whole or most curved part of lateral line covered with cycloid scales; straight part and sometimes posterior curved part of lateral line covered with strong scutes (53–75 cycloid scales and 0–10 scutes in curved part + 0–3 cycloid scales and 23–40 scutes in straight part; Table 5); first dorsal fin with interspinous membranes, with VII–VIII spines, second dorsal fin with 25–30 soft rays; anal-fin soft rays 17–23; total number of second dorsal- and anal-fin soft rays 46–51 (Table 1, Fig. 9); anterior second dorsal and anal fins not falcate; no finlets; free anal-fin spines short, embedded in adults; both neural and hemal spines of ninth caudal vertebra Spine-shape (Fig. 7c); first hemal spine distinctly crooked posteriorly just under first caudal vertebra (Fig. 2c); two to four neural spines between pterygiophores supporting first and last dorsal-fin spines; three to six pterygiophores between first and second hemal spines; total gill rakers 18–24; snout length 7.7–12.1% SL (Table 2, Fig. 10a); suborbital depth relatively narrow, 3.8–7.4% SL (Table 2, Fig. 10b); tongue, roof and floor of mouth white or pale yellow.

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth superior, lower jaw protruding. Scales on body small, cycloid, except for straight part and sometimes posterior curved part of lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 12th–17th dorsal-fin soft ray; naked area on breast connecting with naked area on pectoral-fin base in Uraspis uraspis, not connecting in Uraspis helvola. Second dorsal-fin base slightly longer than anal-fin base. Vertebrae 10 + 14; inferior foramen on last three precaudal vertebrae.

Distribution. Circumglobal in tropical and subtropical seas.

Gender. Feminine.

Comparison. The genus Uraspis is distinguishable from the former Carangoides and other genera by having a distinctly crooked first hemal spine (Fig. 2c). The genus can also be distinguished from the latter by having small conical teeth arranged uniserially or biserially on jaws except for Parastromateus, Carangichthys and Atropus atropos, by having Spine-shape neural and hemal spines on the ninth caudal vertebra (Fig. 7) except for Parastromateus, Carangichthys and Carangoides, and by having a whole straight part of lateral line covered by scutes except for Atropus and Carangichthys. Uraspis can be distinguished from Parastromateus by having a naked thoracic area (vs. no naked area; Fig. 6) and from Atropus, Carangichthys and Carangoides by lacking the vomerine tooth patch (vs. present; Fig. 5).

Remarks. Günther (1860) gave the replacement name Caranx uraspis for Uraspis carangoides because he placed this nominal species and Leioglossus carangoides in the same genus Caranx. He also gave the replacement name Caranx lioglossus Günther 1860 for Leioglossus carangoides (a synonym of Atropus mentalis). But this replacement is unneeded because Leioglossus carangoides has priority over Uraspis carangoides. Uraspis carangoides and Leioglossus carangoides are no longer in the same genus, but the former is invalid and the replacement name Uraspis uraspis is valid according to ICZN (1999, Art. 59.3). Leucoglossa candens, the type species of Leucoglossa, can be confirmed that the species belongs to Uraspis based on counts of second dorsal- and anal-fin soft rays and scutes based on its original description and posteriorly crooked first hemal spine (radiograph of the holotype; National Museum of Natural History: http://n2t.net/ark:/65665/30dfd04a2-2b52-40e3-a61b-a94e3cea57e6; accessed on 22 March 2021). Whitley (1931) established the genus Zamora based on Caranx hullianus as the type species. However, Zamora Whitley 1931 had been preoccupied by the genus of spider, Zamora Roewer 1928, and the former nominal genus was an objective homonym, being objectively invalid. Whitley (1941) gave the replacement name Bassetina Whitley 1941 for Zamora Whitley 1931. The type species of Bassetina (also Zamora Whitley 1931) is undoubtedly identifiable as a species of Uraspis from its original description and drawing. Platyuraspis was described as a subgenus of Uraspis for Uraspis heidi as the type species (Fig. 27a₃). Fowler (1938b) stated the major differences of Uraspis heidi from congeneric species were deeper contour of the body, elevated soft dorsal and anal fins with no distinct anterior lobes and with gradually longer anterior few rays, the longer pelvic fin, general coloration more or less dark neutral green with pale narrow cross bars but not extending on soft dorsal and anal fins, and all fins blackish except yellowish caudal-fin base and pale pectoral fin. He considered that those differences seemed worthy of at least subgeneric rank. However, such differences include intraspecific variations and changes associated with growth, and it is considered that the differences are not enough to divide by subgenera. Accordingly, three genus-group nominal taxa, Leucoglossa, Bassetina and Platyuraspis are junior synonyms of Uraspis.

The thoracic naked area in Wakiya’s (1924) Caranx (Uraspis) uraspis is restricted and not connected with naked pectoral-fin base from his drawing of the species (plate 31, fig. 1). Therefore, this is identifiable as Uraspis helvola.

Smith-Vaniz (1999, 2016) stated that it is virtually impossible to distinguish adults of Uraspis secunda (Poey 1860) (Fig. 27a₅) and Uraspis helvola (Fig. 27a₁) from each other, although juvenile characters involving allometric growth patterns suggest that they are different species. Moreover, Smith-Vaniz (2002) indicated that Uraspis secunda possibly a junior synonym of Uraspis helvola, in which case the latter species has a circumglobal distribution. The ML tree based on COI sequences analyzed by us (ESM Table S1) and downloaded from GenBank or BOLD (EMS Table S2) of Uraspis is shown in Fig. 28. Although there were no published COI sequences of Uraspis from Atlantic Ocean, only two species are recognized in Fig. 28, namely, only Uraspis helvola and Uraspis uraspis exist in the Indo-Pacific and the eastern Pacific Ocean. Therefore, we here treated that U. secunda is a junior synonym of Uraspis helvola.

Fig. 28

Maximum likelihood (ML) tree of the genus Uraspis based on COI sequence. Voucher specimens or accession numbers of the sequences are indicated as follows: A DBMF024-10 (Malaysia), DBMF585-10 (Malaysia), FOAH451-08 (Australia), FOAO183-14 (Australia), KAUM-I. 22155 (Malaysia), KAUM-I. 22159 (Malaysia), KU578093 (Korea), KU943790 (Taiwan), KY372378 (South China Sea) and KY372379 (South China Sea); B EU514510 (India); C JN312968 (Indonesia); D MK777195 (Vietnam); E MK777194 (Vietnam); F FOAM395-10 (Indonesia) and JN312970 (Indonesia); G KAUM-I. 22154 (Malaysia); H MN623874 (India); I KU578090 (Korea); J JX261042 (Malaysia); K FRLM 47078 (Phuket, Thailand), HQ149964 (Iran), HQ560984 (Malaysia), JN312969 (Indonesia), JX261072 (Malaysia) and MK777196 (Vietnam); L FRLM 36715 (Japan), FRLM 36902 (Japan), FRLM 38529 (Japan), FRLM 38613 (Japan), FRLM 42760 (Japan), HQ010061 (California, USA), JF494773 (South Africa), KAUM-I. 51207 (Japan), KU943818 (Taiwan), KY372377 (China) and MH638709 (South China Sea); M AMS919-08 (Australia), FRLM 39389 (Japan), GU673637 (Indonesia) and GU674099 (Indonesia); O FRLM 43488 (Carangichthys dinema, Japan, out group); P FRLM 36695 (Carangichthys oblongus, Japan, out group). Figures beside major internal branches ML bootstrap values

Figure 28 suggests existence of two genetic groups in Uraspis uraspis. Yeo and Kim (2016) also indicated the similar phylogenetic tree of Uraspis. However, we could not find clear morphological differences between those groups.

Carangichthys Bleeker 1853

(Subclade N, Fig. 29)

Fig. 29

Three included species of the genus Carangichthys (Subclade M), a Carangichthys dinema, type species, b Carangichthys humerosus and c Carangichthys oblongus. a₁ FRLM 38625, 349 mm SL, Okinawa, Japan (fresh condition); a₂ Carangichthys dinema, FRLM 36756, 150 mm SL, Mie, Japan (fresh condition); a₃ RMNH.PISC 26973, holotype of Carangichthys typus, 86 mm SL, Kema, Indonesia; a₄ NMW 60929, holotype of Caranx longipes, 59 mm SL, Samoa (© Naturhistorisches Museum, Wien, photographed by A. Naseka); b Carangichthys humerosus, FRLM 25724, 150 mm SL, Bitung, Indonesia (fresh condition); c₁ MNHN A-6172, holotype of Caranx oblongus, 261 mm SL, Vanikoro, Solomon Islands; c₂ FRLM 37150, 169 mm SL, Mie, Japan (fresh condition); c₃ USNM 51951, holotype of Caranx deani, 111 mm SL, Negros, Philippines

Carangichthys Bleeker 1853: 760 (type species: Carangichthys typus Bleeker 1853)

Type species. Carangichthys typus Bleeker 1853, a junior synonym of Carangichthys dinema (Bleeker 1851b).

Included species. Carangichthys dinema (Bleeker 1851b), Carangichthys humerosus (McCulloch 1915) and Carangichthys oblongus (Cuvier in Cuvier and Valenciennes 1833).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, deep, oval or elongate-oval; dorsal contour of upper lip around symphysis roundly protruding upward (Round chevron type; Fig. 4n); upper jaw with conical-tooth band and lower jaw with an outer row of slightly large conical teeth and an inner band of villiform teeth; vomerine tooth patch triangular (Fig. 5f); palatines with small conical teeth, outer teeth larger in juveniles; adipose eyelid poorly developed; naked area on breast not connecting with naked area on pectoral-fin base, either reaching posteriorly to pelvic-fin insertion in Carangichthys dinema and Carangichthys oblongus (Fig. 6n_{1, 3}) or extending beyond pelvic-fin insertion in Carangichthys humerosus; entire or most curved part of lateral line covered with cycloid scales; straight part and usually posterior curved part of lateral line covered with scutes (52–64 cycloid scales and 0–9 scutes in curved part + 0–4 cycloid scales and 17–40 scutes in straight part, Table 5); posterior and ventral margins of preopercle serrated in juveniles; first dorsal fin with interspinous membranes, with VII–VIII spines, second dorsal fin with 17–22 soft rays; anal-fin soft rays 15–19; anterior second dorsal and anal fins falcate with elongate rays in adults; caudal fin long, UCF/BD 62.6–88.4%; no finlets; both neural and hemal spines of ninth caudal vertebra Spine-shape (Fig. 7c); total gill rakers 24–32; snout short, 8.5–10.7% SL (Table 2, Fig. 10a); suborbital relatively narrow, 4.1–6.3% SL (Table 2, Fig. 10a).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth almost terminal, lower jaw very slightly projecting. Scales on body small, cycloid, except for straight part and sometimes posterior curved part of lateral line; single lateral line with curved anterior and straight posterior parts, straight part beginning below base of 9th–14th dorsal-fin soft ray. Second dorsal-fin base merely longer than soft portion of anal-fin base; first and second anal-fin spines small, separated from the third spine. Six neural spines between pterygiophores supporting first and last dorsal-fin spines; three pterygiophores between first and second hemal spines. Vertebrae 10+14; inferior foramen on last three or four precaudal vertebrae.

Distribution. Indo-West Pacific, from the eastern coast of Africa to Fiji (Smith-Vaniz 1999).

Gender. Masculine.

Comparison. The genus Carangichthys is distinguishable from the former Carangoides and other genera by having small conical teeth arranged uniserially or biserially on jaws except for Parastromateus, Uraspis and Atropus atropos, by having Spine-shape neural and hemal spines on the ninth caudal vertebra (Fig. 7) except for Parastromateus, Uraspis and Carangoides, and by having whole straight part of lateral line usually covered by scutes except for Atropus and Uraspis. Carangichthys can be distinguished from Parastromateus by having a naked thoracic area (vs. no naked area; Fig. 6) and from Uraspis and Carangoides by having a triangular vomerine tooth patch (vs. lacking vomerine tooth patch in Uraspis and anchor-shaped in Carangoides).

Remarks. The genus Carangichthys has been treated as a valid genus (e.g., Gushiken 1983, 1984; Senou 2002, 2013) or as a junior synonym of the previous Carangoides (e.g., Smith-Vaniz 1999). Both Gushiken (1983, 1984) and Senou (2002, 2013) characterized the genus by whole straight part of lateral line covered by scutes. In the original description of the genus, Bleeker (1853) placed particular emphasis on serrated preopercular margin. Although this feature is unique to this genus, it is a characteristic only in juveniles and the preopercular margins become smooth with growth. As stated in “Diagnosis”, Carangichthys has unique characteristics and should be regarded as an independent genus. The phylogenetic tree indicated in Fig. 3 also supports this result. Carangichthys humerosus is included in the genus based on its morphological characteristics although the species is not included in the present molecular analysis.

Carangoides dinema was described without a type specimen. But its original description agrees well with our results of examination on the holotype of Carangichthys typus (RMNH.PISC 26973, Fig. 29a₃), the type species of the genus. Therefore, the latter species can be synonymized under the former, as already stated by Gushiken (1983) and Smith-Vaniz (1999).

Caranx deani (Fig. 29c₃) was considered as a junior synonym of Carangoides dinema (see Smith-Vaniz 1999). However, the holotype of this nominal species (USNM 51951) has longer straight part of lateral line than curved part and 40 scutes (44 in the original description). Therefore, this nominal species is conspecific with Carangichthys oblongus and as a junior synonym of the latter.

The holotype of Caranx longipes, NMW 60929 (Fig. 29a₄) is identifiable with Carangichthys dinema by general body shape and having 19 second dorsal-fin soft rays, straight part of lateral line with 25 scutes, shorter than curved part, and naked thoracic area just reaching to pelvic-fin insertion.

Carangoides Bleeker 1851b

(Clade 2, Fig. 30)

Fig. 30

Two included species of the genus Carangoides, a Carangoides ire and b Carangoides praeustus, type species. a MNHN 0000-5847, holotype of Caranx ire, 131 mm SL, Puducherry, India; b FRLM 35478, 129 mm SL, Ha Long, Vietnam (fresh condition)

Carangoides Bleeker 1851b: 352 (type species: Caranx praeustus Anonymous [Bennett] 1830)

Type species. Caranx praeustus Anonymous [Bennett] 1830 [= Carangoides praeustus (Anonymous [Bennett] 1830)].

Included species. Carangoides ire (Cuvier in Cuvier and Valenciennes 1833) and Carangoides praeustus (Anonymous [Bennett] 1830).

Diagnosis. A genus of the family Carangidae defined by the following combination of characters: body compressed, elongated-oval, somewhat lower; dorsal contour of upper lip around symphysis roundly protruding upward (Round chevron type; Fig. 4o); upper jaw with a band of small conical teeth, lower jaw with conical teeth forming a band anteriorly and uniserially arranged posteriorly; vomer and palatines with villiform teeth; vomerine tooth patch anchor-shaped (Fig. 5g); adipose eyelid poorly developed; naked thoracic area somewhat small and not connecting with naked area on pectoral-fin base; curved part and anterior straight part of lateral line with cycloid scales (56–72 + 2–11), the posterior straight part with 17–26 scutes; first dorsal fin with interspinous membranes, with VIII spines, second dorsal fin with 21–25 soft rays; anal-fin soft rays 18–21; anterior second dorsal fin not falcate, slightly higher than the first; anterior anal fin not falcate; no finlets; both neural and hemal spines of ninth caudal vertebra not expanding, Spine-shape (Fig. 7c); two pterygiophores between first and second hemal spines; total gill rakers 32–47; snout length 8.6–11.3% SL (Table 2, Fig. 10a); suborbital narrow, 2.4–3.2% SL (Table 2, Fig. 10b).

Description. Characters given in the diagnosis are not repeated. Counts and measurements are shown in Tables 1, 2, 5 and 6. Mouth terminal; snout longer than eye diameter. Scales on body small, cycloid, except for posterior lateral line; single lateral line with curved anterior and straight posterior parts, the straight part beginning below base of 11th–13th dorsal-fin soft ray; naked thoracic area extending just beyond pelvic-fin insertion in Carangoides ire (Fig. 6o₁) or very small on jugular in Carangoides praeustus (Fig. 6o₂). Second dorsal-fin base slightly longer than anal-fin base; first and second anal-fin spines developed, separated from the third spine, and not embedded. Six neural spines between pterygiophores supporting first and last dorsal-fin spines. Tip of second dorsal fin jet black.

Distribution. Indo-West Pacific, from the Persian Gulf (Smith-Vaniz 1983) to Indonesia.

Comparison. The genus Carangoides is distinguishable from the former Carangoides and other genera in having an anchor-shaped vomerine tooth patch (Fig. 5). The genus is also distinguished from other carangid genera in having irregular row of small conical teeth on both jaws, poorly developed eyelids, cycloid scales on curved and anterior straight parts of the lateral line, scutes on the posterior straight portion of lateral line, and no finlets.

Remarks. Smith-Vaniz (1999) stated that the former Carangoides praeustus consisted of two geographic populations (Western Central Pacific and Indian Ocean from the Arabian/Persian Gulf to Bay of Bengal) and they differed in total gill-raker counts (32–37 in the former population vs. 40–47 in the latter) and patterns of naked area on breast (usually completely scaly or nearly so vs. breast typically with a narrow naked midventral area extending entire length of breast); if future studies showed that these two “geographic races” were actually represented by different sister species, the oldest available name for the Indian Ocean species is Carangoides ire (Cuvier). Bogorodsky et al. (2017: fig. 15) indicated the obvious differentiation of their Carangoides praeustus from Indian Ocean and western Pacific Ocean. Carangoides ire was originally described based on a single specimen MNHN 0000-5847 from Puducherry, India (Fig. 30a). The holotype of this nominal species has a distinct thoracic naked area (Fig. 6o₁) and somewhat more gill-raker count (Table 5). As stated above, Indian-Ocean and western Pacific populations are genetically differentiated clearly (Bogorodsky et al. 2017). From these facts, we here resurrect Carangoides ire as a valid species from the Indian Ocean which should be redescribed.

Key to the genera of the species formerly belonging to Carangoides and its related genera

• 1a. Interspinous membranes of first dorsal fin absent … 2

• 1b. Interspinous membranes of first dorsal fin present … 3

• 2a. Dorsal edge of upper lip around symphysis gradually ascending (Fig. 4k); total number of second dorsal- and anal-fin soft rays 78–83; no pelvic fins in adults; suborbital relatively narrow, 2.7–5.3% SL … Parastromateus Bleeker 1864

• 2b. Dorsal edge of upper lip around symphysis steeply protruding upward (Fig. 4b); total number of second dorsal- and anal-fin soft rays 32–40; pelvic fins present in adults; suborbital broad, 12.5–17.3% SL … Scyris Cuvier 1829

• 2c. Dorsal edge of upper lip around symphysis roundly protruding upward (Fig. 4a); total number of second dorsal- and anal-fin soft rays 33–35; pelvic fins present in adults; suborbital moderately broad, 7.0–9.0% SL … Alectis Rafinesque 1815

• 3a. Dorsal edge of upper lip around symphysis gradually ascending (Fig. 4g); suborbital narrow, 3.4–5.0% SL … Craterognathus gen. nov.

• 3b. Dorsal edge of upper lip around symphysis steeply protruding upward (Fig. 4d, e); suborbital very broad, 13.1–22.5% SL … 4

• 3c. Dorsal edge of upper lip around symphysis roundly protruding upward (Fig. 4c, f, h–j, l–o); suborbital narrow, not more than 8.6% SL … 5

• 4a. Breast completely scaly, or narrow naked area present behind operculum not connected to naked pectoral-fin base (Fig. 6e); vomerine tooth patch diamond- or cross-shaped with a long posterior extension (Fig. 5d, e) … Selene Lacepède 1802

• 4b. Naked area on breast connected with naked pectoral-fin base (Fig. 6d); vomerine tooth patch shaped four-pointed-star (Fig. 5c) … Paraselene gen. nov.

• 5a. Vomerine tooth patch absent … Uraspis Bleeker 1855

• 5b. Vomerine tooth patch present … 6

• 6a. Vomerine tooth patch anchor-shaped (Fig. 5g) … Carangoides Bleeker 1851b

• 6b. Vomerine tooth patch diamond-shaped with round posterior corner (Fig. 5b) … Euprepocaranx gen. nov.

• 6c. Vomerine tooth patch chevron-shaped or triangular (Fig. 5a, f) … 7

• 7a. Jaws with small conical teeth uniserially or biserially arranged … Carangichthys Bleeker 1853

• 7b. Jaws with villiform teeth forming bands … 8

• 8a. Naked area on breast very small along anterior ventral midline (Fig. 6i) … Flavocaranx gen. nov.

• 8b. Naked area on breast reaching to or extending beyond pelvic-fin insertion (Fig. 6f, h, j, l) … 9

• 9a. Naked area on breast not extending beyond pelvic-fin insertion (Fig. 6l) … Ferdauia Jordan, Evermann and Wakiya in Jordan, Evermann and Tanaka 1927

• 9b. Naked area on breast extending beyond pelvic-fin insertion (Fig. 6f, h, j) … 10

• 10a. Dorsal edge of upper lip around symphysis roundly protruding upward, protrusion of upper-jaw symphysis narrow (Fig. 4j) … Platycaranx gen. nov.

• 10b. Dorsal edge of upper lip around symphysis roundly protruding upward, protrusion of upper-jaw symphysis wide (Fig. 4f, h) … 11

• 11a. Snout long, 10.9–14.3% SL … Turrum Whitley 1932

• 11b. Snout short, 7.4–10.7% SL … Atropus Oken 1817