key: cord-0841675-9ukeuazy
authors: Chungthanawong, Sirikanya; Motomura, Hiroyuki
title: Review of the waspfish genus Neocentropogon (Tetrarogidae), with a key to genera in the family
date: 2021-02-23
journal: Ichthyol Res
DOI: 10.1007/s10228-020-00796-w
sha: 720ba6bc17923890352927b828b3c6cd67c42732
doc_id: 841675
cord_uid: 9ukeuazy

A taxonomic review of the waspfish genus Neocentropogon Matsubara 1943 (Tetrarogidae), diagnosed by the following combination of characters: body sparsely covered with small embedded cycloid scales, palatine teeth present, XIII–XVI dorsal-fin spines, the first dorsal fin originating above the orbit, five pelvic-fin soft rays, and membrane of lower four pectoral-fin rays deeply incised, resulted in the recognition of six species: Neocentropogon aeglefinus (Weber 1913), Neocentropogon affinis (Lloyd 1909a), Neocentropogon japonicus Matsubara 1943, Neocentropogon mesedai Klausewitz 1985, Neocentropogon profundus (Smith 1958), and Neocentropogon trimaculatus Chan 1966. Neocentropogon trimaculatus (anti-tropically distributed in East Asia and Australia) can be distinguished from its congeners by the presence of three dark blotches on the body (vs. absent or a single blotch); N. affinis (eastern Indian Ocean) and N. aeglefinus (Philippines to Australia) differ from other congeners in having a black blotch behind the opercle (vs. blotch absent), with the former distinguishable from the latter by dorsal rows of dark spots on the body, and pectoral and caudal fins (vs. spots absent), and 79–96 scale rows in the longitudinal series (vs. 94–137); N. mesedai (Red Sea) differs from N. profundus (southwestern Indian Ocean) and N. japonicus (northwestern Pacific Ocean) in having the lowermost four pectoral-fin rays elongated and XIII (vs. XIV–XVI) dorsal-fin spines, the latter species being separated by the symphyseal knob condition (unremarkable, N. profundus vs. pronounced, N. japonicus), dark dorsal spots on the body (vs. absent), and 5 anal-fin soft rays (vs. 6 or 7). Keys to the genera of Tetrarogidae and species of Neocentropogon are given, including taxonomic status of Vespicula Jordan and Richardson 1910 and Pseudovespicula Mandrytsa 2001.

The Indo-West Pacific family Tetrarogidae (waspfishes) has been regarded as comprising 18 valid genera, viz., Ablabys Kaup 1873 , Centropogon Günther 1860 , Coccotropsis Barnard 1927 , Cottapistus Bleeker 1876a , Glyptauchen Günther 1860 , Gymnapistes Swainson 1839 , Liocranium Ogilby 1903 , Neocentropogon Matsubara 1943 , Neovespicula Mandrytsa 2001 , Notesthes Ogilby 1903 , Ocosia Jordan and Starks 1904 , Paracentropogon Bleeker 1876b , Pseudovespicula Mandrytsa 2001 , Richardsonichthys Smith 1958 , Snyderina Jordan and Starks 1901 , Tetraroge Günther 1860 , Trichosomus Swainson 1839 , and Vespicula Jordan and Richardson 1910 , with 43 valid species overall (Fricke et al. 2020 ). The family is characterized by a compressed body, head spines, a mobile lacrimal bone, skin at the gill opening not broadly connected to the isthmus, and the lower pectoral-fin rays not separated from other pectoral-fin rays (Poss 1999 ).

This article was registered in the Official Registry of Zoological Nomenclature (ZooBank) as ECACB 520-4A7A-4DBF-AEEA-FF8F8 2A473 F0. This article waspublished as an Online First article on the online publication date shown on this page. The article should be cited by using the doi number.

The genus Neocentropogon (type species: Paracentropogon aeglefinus Weber 1913) , one of the most poorly known genera in the family, comprises relatively small, bottom dwelling fishes, characterized by an oblique head profile, the body covered with numerous cycloid scales, palatine teeth present, the dorsal fin with 13-16 spines, its origin anterior to the orbit posterior margin, the last dorsal-fin soft ray membrane posteriorly connected to the dorsal caudal peduncle but not extending onto the upper base of the caudal fin, pelvic fin with 1 spine and 5 soft rays, and lowermost four pectoral-fin rays detached but with their basal half connected by a low membrane. Apart from numerous brief treatments in general classifications and regional faunal studies, Neocentropogon has at no time been reviewed on the basis of type and non-type materials; thus, some taxonomic confusion has resulted. Accordingly, the present review of the genus has been made on an Indo-West Pacific basis. Examination of all available type specimens and a large number of non-type specimens of Neocentropogon representing wide distributional ranges in this study resulted in six species being regarded as valid. They are redescribed here in detail. Similarly, a key to all genera of Tetrarogidae on the basis of examination of all valid species has never been published, although some authors (e.g., Poss and Rama-Rao 1984; Poss 1999 ) provided a regional key to some genera. This study aims to provide complete keys to the genera of the family and to the species of the genus, and diagnoses of six species to establish comprehensive taxonomy of Neocentropogon, based on examination of numerous specimens.

Counts and proportional measurements followed Motomura (2004a) and Motomura et al. (2008) , except scale counts followed Chungthanawong and Motomura (2018) . Standard and head lengths are expressed as SL and HL, respectively. Head spine terminology follows Randall and Eschmeyer (2002: fig. 1 ) and Motomura (2004b: fig. 1 ). Osteological characters, including vertebral counts, were observed on radiographs of Neocentropogon aeglefinus (8 specimens: CSIRO H 4032-01, MNHN 2006 -0256, 2014 , Neocentropogon affinis Lloyd 1909a (3: KAUM-I. 33280, SAIAB 65706, 2 specimens), Neocentropogon japonicus Matsubara 1943 (11: FAKU S511, 103972, KAUM-I. 20392, 20393, 30815, MNHN 1984 -0635, 2005 -0624, 2005 -0709, 2005 -1006 , 2005 -1298 , Neocentropogon profundus Smith 1958 (1: MNHN 2006 , and Neocentropogon trimaculatus Chan 1966 (11: BMNH 1965 .11.6.3, FAKU 75091, KAUM-I. 40487, 77115, 88804, MNHN 2003 -1850 , 2005 -2624 -1040 . The formula for configuration of the supraneural bones, anterior neural spines and anterior dorsal-fin pterygiophores follows Ahlstrom et al. (1976) . Swimbladder absence was confirmed by dissection of the abdomen on the right side of the body in N. aeglefinus (2: QM I. 21498, 2), N. affinis (KAUM-I. 33280), N. japonicus (8: FAKU S511, S512, 103972, KAUM-I. 20393, 30815, 81861, 114281, 114289) , N. trimaculatus (5: 77117, 88804, 97509, 97510) . Color descriptions are based on preserved specimens. The key to genera was based on specimens representing 39 species in 17 genera examined during this study (see Material examined for key to genera), and the original descriptions of four species (Bleeker 1848; Weber 1913; Poss and Eschmeyer 1975; Fricke 2017) . Comparative features for the genera are provided in Table 1 . Institutional codes follow Sabaj (2019) . 

Vespicula was originally proposed by Jordan and Richardson (1910) as a new monotypic genus for Prosopodasys gogorzae Jordan and Seale 1905, originally described on the basis of a single specimen from the Philippines. Jordan and Richardson (1910) also distinguished Vespicula from Prosopodasys Cantor 1849 on the basis of the dorsal fin with the three anteriormost spines forming an almost completely Dor (1984) and Randall (1995) (2001) for A. dracaena following comparison with A. trachinoides, which he regarded as belonging to Vespicula (but later placed in Trichosomus -see above). Although Mandrytsa (2001) did not compare P. dracaena with either V. cypho or V. zollingeri, the three species are herein regarded as belonging to a single genus due to their sharing the following major generic characters: dorsal-fin membrane between third and fourth spines deeply incised, forming a nearly separate fin; dorsal-fin origin directly above posterior margin of orbit; 5 pelvic-fin soft rays; pectoral-fin rays not detached; body with small cycloid scales, without cirri or papillae; lateral line well separated from dorsal-fin base; teeth on palatine; head profile oblique, straight; and nape flattened.

Because P. gogorzae, the type species of Vespicula, has been regarded as a junior synonym of Trichosomus trachinoides, Vespicula is considered a junior synonym of Trichosomus. Therefore, V. cypho and V. zollingeri are regarded herein as species of Pseudovespicula.

Neocentropogon Matsubara 1943: 429 (type species:

Paracentropogon aeglefinus Weber 1913, by original designation)

Gadapistus de Beaufort, 1949 : 68 (type species: Paracentropogon aeglefinus) Diagnosis. A genus of the family Tetrarogidae with the following combination of characters: XIII-XVI, 6-8 dorsal-fin rays, its origin anterior to vertical through posterior margin of orbit; anteriormost dorsal-fin spines not forming separate fin; membrane of last dorsal-fin soft ray not connected posteriorly to upper caudal-fin base; I, 5 pelvic-fin rays; 13-16 pectoral-fin rays, four lowermost rays simple and detached, their basal half connected by low membrane; head and snout profile oblique; nape flattened; mouth large [37.4-49.9 (mean 45.2)% HL]; body sparsely covered with small embedded, non-imbricate cycloid scales; cirri and papillae absent on head and body; cleithral spine absent; small conical teeth on palatines; 18-26 lateral-line pores; lateral line running along upper one-third of body; tip of opercle directed backward, below lateral line.

Description. Body somewhat elongated, laterally compressed, progressively more compressed posteriorly, caudal peduncle short. Scales absent on head, pre-dorsal-fin area, dorsal-and anal-fin bases. Tentacles, cirri and skin flaps absent on head, body and fins. Lateral line straight, extending from above supracleithral spine to caudal-fin base, one lateral-line pore near caudal-fin base.

Head profile oblique with shallow concavity in front of eyes. Two nasal openings in front of orbit, subequal in diameter, tubular; anterior nostril higher than posterior nostril. Interorbital space convex; interorbital ridges weakly developed; median interorbital ridge and spines absent; ascending process of premaxilla intruding slightly into interorbital space. Nuchal, pterotic, upper posttemporal, lower posttemporal, and supracleithral forming ridges with minute spines, entirely covered with skin. Suborbital ridge weak, without spines, connected posteriorly to base of uppermost preopercular spine. Preopercle with 5 simple spines; uppermost longest, sharp, projecting from skin; second and third short, sharp, projecting from skin; fourth and fifth blunt, with broad base, hidden under skin. Opercle with smooth V-shaped crests; upper crest with minute sharp spine projecting from skin; lower crest with weak blunt spines. Cleithral bone flattened, covered with thick skin. Lacrimal with 2 simple sharp spines; anterior lacrimal spine short, directed ventrally; its tip extending well beyond dorsal margin of maxilla; posterior lacrimal spine longer, directed posteriorly. Mouth moderately large, terminal, slightly oblique; maxillary extending posteriorly to about level with middle of eye, symphyseal knob present. Lips thick; gill rakers rather short, blunt; no slit behind last gill arch.

Dorsal fin continuous; origin anterior to posterior margin of orbit; 3 anteriormost dorsal-fin spines somewhat separated from rest of fin; spinous membrane of fin deeply notched; last dorsal-fin ray membrane connected posteriorly to caudal peduncle. Anal fin continuous, III, 5-8 anal-fin rays; origin about level with origin of eleventh dorsal-fin spine; membranes of spinous portion notched; membrane of last dorsal-fin ray posteriorly connected to caudal peduncle. Pectoral fin with 13-16 rays; uppermost and lower four rays unbranched, reaching or extending beyond origin of first anal-fin spine; posterior margin of fin rounded. Origin of pelvic fin level with vertical through lower end of pectoral-fin base; posterior tip of depressed fin usually almost reaching anus. Caudal fin rounded. Vertebrae 11 + 14, including hypural. Swimbladder present. Formula for configuration of supraneural bones, anterior neural spines and anterior dorsal pterygiophores 2+1/1/1/1/1/1/1/1/1/1/1/1/ (2+1/1/1/1/1/1/1/1/1/1/1/1/1/ in N. japonicus). Epineurals usually 13 (14 in N. japonicus).

Remarks. Neocentropogon aeglefinus was originally described by Weber (1913) as a species of Paracentropogon on the basis of the following six characters: body covered with small cycloid scales, 4 or 5 preopercular spines, 8 or 9 dorsal-fin soft rays, 4-6 anal-fin soft rays, 6 branchiostegal rays, and no slit behind the last gill arch. Subsequently, Matsubara (1943) recognized that P. aeglefinus differed from Paracentropogon in eight characters: I, 5 pelvic-fin rays (vs I, 4 in the latter); symphyseal knob present (vs absent); all branchiostegal rays enlarged (vs 5 posteriormost rays only enlarged); actinosts narrow (vs wide); pyloric caeca teatlike (vs tube-like); posttemporal forked anteriorly, firmly attached to but not forming an integral part of the cranium (vs thick and strong, slightly emarginated anteriorly, immovably attached to and forming an integral part of the cranium), first pair of parapophyses on eighth vertebra (vs on fifth or sixth vertebra), vertebrae 25 including hypural (vs 24-27, usually 26), and placed the species in his new genus Neocentropogon, which was also regarded here as a valid genus.

Neocentropogon is distinguished from all other genera in the family by the combination of characters given in the Diagnosis (above). The simple four lowermost pectoral-fin rays with their basal half connected by low membrane is a unique character within the family.

De Beaufort (1949) proposed a new genus, Gadapistus, for P. aeglefinus on the basis of three characters: dorsal-fin origin anterior to posterior margin of orbit (vs above orbit in Paracentropogon); 5 pelvic-fin soft rays (vs 4), and symphyseal knob present (vs absent). However, Gadapistus is regarded as an objective junior synonym of Neocentropogon, the type species of the two genera being the same (Mandrytsa 2001; this study) .

After detailed examination of 202 specimens of Neocentropogon, this study recognizes the following six species in it: N. aeglefinus; N. affinis; N. japonicus; N. mesedai; N. profundus; N. trimaculatus. This membership is newly recognized here because a comprehensive taxonomic revision of this genus has not been performed before. 

A species of Neocentropogon with the following combination of characters: dorsal-fin rays XIV, 7 (rarely XIII or XV, 6 or 8); anal-fin soft rays 6 (rarely 5); lateral-line pores 20-24; scale rows in longitudinal series 94-137; scale rows above lateral line 8-17; symphyseal knob pronounced; postocular spine usually absent; four lowermost pectoral-fin rays not elongated; orbit diameter 11.8-15.3% (mean 13.5%) of SL; large dark blotch behind opercular margin; blotches absent on dorsal fin base; spots absent on body dorsal surface and dorsal fin; pectoral fin black.

Distribution. Currently known from the Philippines to Indonesia, Australia, the Solomon Islands, and Vanuatu in depths of 17-705 m [based on collected specimens (Fig. 3) ].

Remarks. Neocentropogon aeglefinus was originally described as Paracentropogon aeglefinus by Weber (1913) on the basis of 14 specimens from Indonesia (Halmahera, Timor and Savu seas, and Lesser Sunda Islands). The syntypes, registered as ZMA 110234 (2 specimens), 110235 (4), 110236 (5), 110237 (1), and 110240 (2) (Fricke et al. 2020) , are deposited at Naturalis Biodiversity Center, Leiden, the Netherlands. Because the Naturalis fish collection has long been inaccessible due to building renovations (and currently the covid-19 pandemic), the syntypes were unavailable for the present study. However, examination of the original description and figure (Weber 1913: 500, pl. 6, fig. 8 ) of P. aeglefinus showed it to be identical with specimens considered here as conspecific, the former having small embedded cycloid scales on the body; XIV, 8 dorsal-fin rays; 15 pectoral-fin rays, the four lowermost rays detached with their basal half connected by low membrane; I, 5 pelvic-fin rays; palatine teeth; 22 lateral-line pores; no slit behind the last gill arch; and a large dark blotch behind the opercular margin. Ontogenetic morphological change of N. aeglefinus is described in Remarks of N. trimaculatus. Iwamoto and McCosker (2014) reported five specimens (CAS 235572, 3, 65.9-78.9 mm SL; CAS 235749, 2, 78.6-80.4 mm SL) as N. affinis from between Luzon and Mindoro, Philippines. Re-examination of these specimens in this study showed them to be N. aeglefinus. (Lloyd 1909a KAUM-I. 33280, 73.9 mm SL, Thailand; b N. japonicus, KAUM-I. 30815, 85.8 mm SL, Japan; c N. trimaculatus, KAUM-I. 17716, 130 .7 mm SL, Japan lateral line 0-8; symphyseal knob pronounced; postocular spine absent; four lowermost pectoral-fin rays not elongated; orbit diameter 13.0-14.8% (mean 13.7%) of SL; a large dark blotch behind opercular margin; no blotches on dorsal-fin base; rows of dark spots scattered on dorsal body surface and dorsal, pectoral, and caudal fins.

Distribution. Currently known only from the eastern Indian Ocean from the Gulf of Martaban and Andaman Sea to Nias Island, Indonesia [based on collected specimens (Fig. 3) ]. The type specimens were collected in a depth of ca. 84 m.

Remarks. Neocentropogon affinis was originally described by Lloyd (1909a) as G. affinis on the basis of seven specimens (three specimens since lost) from the Gulf of Martaban, Myanmar. Although the remaining four syntypes are no longer in good condition and with fading color, Lloyd's (1909a) description of G. affinis matched the non-type specimens considered here as conspecific with N. affinis and characterized by a greyish-brown body with a greyish blotch behind the opercular margin, two irregular rows of spots above the lateral line, and the dorsal, pectoral, and caudal fins with obscure grey spots. Because no other specimens are known, the non-type specimens listed here represent only the second and third records of the species. Those collected from Nias Island, Indonesia, indicate that the species is widely distributed in the eastern Indian Ocean. Other material examined. 73 specimens (27.5-108.5 mm SL)-JAPAN: BSKU 1923 , 86.6 mm SL, Mimase Fish Market, Kochi, 24 May 1952 BSKU 2230 , 108.5 mm SL, BSKU 2231 , 67.4 mm SL, Mimase Fish Market, Kochi, 11 Dec. 1952 BSKU 13753, 106.6 mm SL, Mimase Fish Market, Kochi, 3 Jan. 1968; BSKU 64652, 72.7 mm SL, 6 June 2003; BSKU 89660, 84.1 mm SL, Saga Fishing Port, Kuroshio, Hata, Kochi, 24 Dec. 2003; BSKU 106457, 89.8 mm SL, Irino Fishing Port, Kuroshio, Hata, Kochi, 15 Dec. 2011; FAKU S511, 98.9 mm SL, FAKU S512, 76.4 mm SL, Mimase, Kochi, 28 Dec. 1958; FAKU 103972, 94 14°00′07′′N, 120°19′04′′E, 191-197 m, RV Coriolis, 31 May 1985; MNHN 2005 -0973, 3, 77.8-89.5 mm SL, 14°00′00′′N, 120°11′24′′E, RV Coriolis, 2 June 1985 MNHN 2005 -1298 m, RV Coriolis, 2 June 1985; LOCALITY UNKNOWN: BSKU 11853, 107.0 mm SL. Diagnosis. A species of Neocentropogon with the following combination of characters: dorsal-fin rays XV, 7 (rarely XIV or XVI, 6); anal-fin soft rays 7 (rarely 6); lateral-line pores 19-24; scale rows in longitudinal series 97-139; scale rows above lateral line 10-21; symphyseal knob pronounced; postocular spine present; four lowermost pectoral-fin rays not elongated; orbit diameter 11.4-14.1% (mean 12.7%) of SL; no large blotches behind opercular margin and on dorsal-fin base; no spots on dorsal body surface or dorsal fin; pectoral fin black.

Distribution. Currently known from the Pacific coast of southern Japan (Kochi to Kagoshima prefectures) south to Taiwan, Hong Kong, the Philippines, and the South China Sea in depths of 70-440 m [based on collected specimens examined in this study (Fig. 3) ].

Remarks. Neocentropogon japonicus was originally described by Matsubara (1943) as a subspecies of N. aeglefinus on the basis of four specimens from Kochi, Japan. Subsequently, Chan (1966) regarded N. a. japonicus as a separate species, N. japonicus, because morphological differences between N. a. japonicus and N. aeglefinus, e.g., XV dorsal-fin spines in N. a. japonicus (vs XIV dorsalfin spines in N. aeglefinus) and absence of black blotch behind opercle (presence of single large black blotch), were equivalent to those of species level in this family; we also agreed with this in this study.

A photograph reported by Mohsin and Ambak (1996: 572, fig. 924 ) as N. aeglefinus was re-identified here as N. japonicus on the basis of having XV, 7 dorsal-fin rays and 7 anal-fin soft rays, and lacking a dark blotch behind the opercular margin. Iwamoto and McCosker (2014) reported N. aeglefinus from between Luzon and Mindoro as the first record from the Philippines. However, since a photograph of a fresh specimen (Iwamoto and McCosker 2014: 287, pl. 16, fig. 91) showed no dark blotch behind the opercular margin, it was herein identified as N. japonicus.

Two specimens (BSKU 17283, 89.8 mm SL; BSKU 17284, 95.8 mm SL) from the South China Sea, midway between Indochina and Borneo, represent the southernmost distribution records of N. japonicus. Klausewitz 1985 fig. 1 ) due to holotype completely faded (Fig. 1d) ]. Distribution. Currently known only from the Red Sea (Fig. 3) , the type specimens having been collected from the central area at a depth of 363-383 m, and an additional specimen ) from the Gulf of Aqaba, northern Red Sea, at a depth of 300-350 m.

Remarks. Neocentropogon mesedai was originally described by Klausewitz (1985) on the basis of five specimens from Mismaris-Trough, southwest of Jeddah, Saudi Arabia, Red Sea. The dorsal-fin spine number (XIII) in N. mesedai is unique in the genus.

Neocentropogon profundus (Smith 1958 Distribution. Currently known only from the western Indian Ocean (Mozambique and Réunion) (Fig. 3) . The specimens examined in this study were collected in depths of 146-227 m. Eleven specimens of N. profundus were collected on the sea surface after the eruption of Piton de la Fournaise, Réunion (Quéro et al. 2011) .

Remarks. Neocentropogon profundus was originally described (as Paracentropogon profundus) by Smith (1958) on the basis of a single specimen taken from the stomach of a rosy snapper, Pristipomoides microlepis (Bleeker 1869) [currently Pristipomoides filamentosus (Valenciennes in Cuvier and Valenciennes 1830)], caught in 146 m off Mozambique, western Indian Ocean. The species has been regarded as belonging to Neocentropogon due to the lack of cirri on the posterior end of the interorbital ridge (Poss and Rama-Rao 1984; Klausewitz 1985) . Fig. 4 Relationships of a, d posterior lacrimal spine length; b, e pectoral-fin length; c, f first dorsal-fin spine length (all percentages of standard length) to standard length (mm) in Neocentropogon aeglefinus (red circles) and N. trimaculatus (green squares). Arrowheads indicate holotype 7 (rarely 6 or 8); lateral-line pores 21-26; scale rows in longitudinal series 92-143; scale rows above lateral line 1-19; symphyseal knob unremarkable; postocular spine present; four lowermost pectoral-fin rays elongated; orbit diameter 10.5-14.7% (mean 12.7%) of SL; head with brownish stripes radiating from pupil; large dark blotch behind opercular margin; 2 large dark blotches on dorsal-fin base; spots absent on dorsal body surface; poorly defined blotches on dorsal fin; pectoral fin whitish with dark blotches.

Distribution. Currently known from southern Japan (including East China Sea) to Taiwan, Hong Kong, northern and eastern Australia, New Caledonia, and the Tonga Trench, from depths of 27-500 m [based on collected specimens (Fig. 3) ], N. trimaculatus is considered to have an anti-equatorial distribution.

Remarks. Neocentropogon trimaculatus, originally described by Chan (1966) on the basis of a single specimen from waters off Hong Kong, is allopatrically distributed in the Northern and Southern hemispheres (Fig. 3) . Although gene flow between the two hemispheres is considered unlikely, comparisons of specimens did not show any significant (Tables 2, 4 ). Accordingly, the northern and southern populations are regarded here as a single species. Although the relative length of the pectoral fin shortens with growth (Fig. 4e) , the lower four pectoral-fin rays remained elongated in larger specimens (Fig. 6) . Analyses of 35 measurements taken from 38 specimens (24.5-145.6 mm SL) of N. trimaculatus and 77 specimens (29.6-116.4 mm SL) of N. aeglefinus indicated similar proportional changes with growth between the two species (selected characters in Fig. 4 ). In addition, the relative lengths of fin rays of N. trimaculatus decreased remarkably with growth compared with those of N. aeglefinus (Figs. 4b, c, e, f, 5, 6) . Analyses of ontogenetic morphological changes in the other species of Neocentropogon could not be made because of the limited number of available specimens.

Previous comparisons among species of Neocentropogon were based on major meristics (such as numbers of dorsal-and anal-fin rays) and coloration, mostly only following original descriptions. In this study, additional characters (i.e., condition of head spines and symphyseal knob, numbers of pores, and some morphometrics) based on the examination of numerous specimens were used for comparisons for the first time in this genus.

Although N. trimaculatus (Fig. 1f ) resembled N. aeglefinus (Fig. 1a) and N. affinis (Fig. 1b) in sharing 13-15 (mode 14) dorsal-fin spines and a dark blotch behind the opercular margin above the pectoral fin, it could be easily distinguished from the latter two species by the two blotches on the dorsal-fin base extending up to the fin (vs. absent), head with brownish stripes radiating from the pupil (vs. absent), lowermost four pectoral-fin rays elongated (vs. not elongated), postocular spine present (vs. absent), 6-8 (mode 7) anal-fin soft rays [vs. 5 or 6 (6)], and symphyseal knob unremarkable (vs. pronounced). Neocentropogon affinis is clearly separated from N. aeglefinus, the former having rows of dark spots on the dorsal body surface, dorsal fin, pectoral fin, and caudal fin (vs. spots absent), 79-96 scale rows in the longitudinal series (vs. 94-137) , and 0-8 scale rows above the lateral line (vs. 8-17) . Neocentropogon mesedai (Fig. 1d) is similar to N. profundus (Fig. 1e) and N. japonicus (Fig. 1c)   Fig. 6 Preserved specimens of Neocentropogon trimaculatus at different growth stages. a KAUM-I. 77119, 24.5 mm SL, East China Sea; b KAUM-I. 09519, 29.6 mm SL, Japan; c KAUM-I. 22469, 39.8 mm SL, Japan; d BMNH 1965.11.6.3, holotype, 80.8 mm SL, South China Sea; e KAUM-I. 77117, 116.0 mm SL, East China Sea ▸ in lacking a dark blotch behind the opercular margin. However, it differs from the latter two species in having the four lowermost pectoral-fin rays elongated (vs. not elongated), 13 dorsal-fin spines (vs. 14-16), and 18 lateral-line pores (vs. 19-24) . Neocentropogon profundus can be distinguished from N. japonicus by the unremarkable symphyseal knob (vs. pronounced), postocular spine absent (vs. present), 5 anal-fin soft rays (vs. 6-7), 71-76 scale rows in the longitudinal series (vs 97-139), 5-8 scale rows above the lateral line (vs. 10-21), greater orbit diameter [15.4-17.0% (mean 15.9%) of SL vs. 11.4-14.1% (12.7%)], and irregular dark spots present on the dorsal body surface (vs. absent).

Ablabys binotatus (Peters 1855) : 3 specimens (90.6-95.2 mm SL), including holotypes of Apistus binotatus and Amblyapistus marleyi Regan 1919, listed in Chungthanawong and Motomura (2018) . Ablabys taenianotus (Cuvier 1829) : 36 specimens (16.4-100.9 mm SL), including holotype of Amblyapistus slacksmithi Whitley 1958 listed in Chungthanawong and Motomura (2018) . Ablabys gymnothorax Chungthanawong and Motomura 2018: 4 specimens (47.9-82.8 mm SL), including holotype of A. gymnothorax, listed in Chungthanawong and Motomura (2018) . Ablabys macracanthus (Bleeker 1852) : 5 specimens (57.2-70.2 mm SL), listed in Chungthanawong and Motomura (2018 

Other material examined. 37 specimens (24.5-145.6 mm SL)-AUSTRALIA: AMS I. 33448-001, 70.5 mm SL

CSIRO H 580-10, 3 specimens, 80.6-92.4 mm SL

CSIRO H 1358-08, 136.1 mm SL, Dunk Island, Qld, 18°06.2′S, 147°08.05′E, 200 m, lobster trawl

NTM S. 12927-010, 2, 58.3-80.5 mm SL, north of Bathurst Island, Arafura Sea, NT, 09°59′S, 130°10′E

QM I. 22111, 106.8 mm SL, Swain Reefs, Qld, 22°00′S, 153°31′E, 270 m, trawl

QM I. 38768, 91.6 mm SL, Surfers Paradise, Qld, 28°00′S, 153°42′E, 102 m, trawl

Diagnosis. A species of Neocentropogon with the following combination of characters: dorsal-fin rays usually XIV, 8 (rarely XIII or XV

2-54.1 mm SL, Australia; QM I. 365, 67.2 mm SL, Australia; QM I. 13106, 2, 40.6-47.9 mm SL, Australia; QM I. 13367, 4, 13.6-31.5 mm SL, Australia; QM I. 14305, 50.2 mm SL, Australia; QM I. 20635, 63.2 mm SL, Australia; QM I. 32241, 17.2 mm SL, Australia; QM I. 32455, 17.1 mm SL, Australia. Coccotropsis gymnoderma

2681, paratype of P. scorpio, 62.4 mm SL, Australia; QM I. 1578, paratype of P. scorpio, 68.6 mm SL, Australia; KAUM-I. 17161, 54.6 mm SL, Malaysia. Glyptauchen panduratus (Richardson 1850): 8 specimens (37.3-150.9 mm SL)-AMS B. 5786, holotype of Glyptauchen insidiator mirandus Whitley 1931, 150.9 mm SL, Australia; AMS IA. 4634, holotype of Glyptauchen insidiator Whitley 1931, 100.9 mm SL, Australia

Gymnapistes marmoratus (Cuvier in Cuvier and Valenciennes 1829): 5 specimens (68.6-115.1 mm SL) -MNHN 6523, 2 syntypes of Apistus marmoratus, 98.1-115.1 mm SL, Indonesia; AMS I. 26833-009, 69.3 mm SL, Australia; CAS 028249, 68.6 mm SL, Australia; CAS-SU 31909, 87.4 mm SL, Australia. Liocranium pleurostigma

mm SL)-QM I. 509, paralectotype of L. praepositum, 81.1 mm SL, Australia

mm SL)-NMW 78424, syntype of Centropogon troschelii Steindachner 1866, 186.6 mm SL, Australia; NMW 12094, syntype of C. troschelii, 202.3 mm SL, Australia; QM I. 954, holotype of Centropogon nitens De Vis 1884b

18496-001, paratype of O. apia, 49.5 mm SL

mm SL)-USNM 99513, holotype of Ocosia gracile Fowler 1943, 36.4 mm SL, Japan; NSMT-P 8374, 88.0 mm SL, Japan; NSMT-P 61707, 42.8 mm SL, Japan; NSMT-P 64365, 2, 35.6-39.8 mm SL, Japan; NSMT-P 101405, 32.1 mm SL, Japan; USNM 122289, 33.6 mm SL, Japan; USNM 135658, 31.0 mm SL, Japan; USNM 135663, 34.3 mm SL, locality unknown

7 mm SL, Japan; NSMT-P 117610, 63.4 mm SL, Japan; NSMT-P 117611, 71.6 mm SL, Japan. Ocosia zaspilota Poss and Eschmeyer 1975: 6 specimens (64.6-87.4 mm SL) -CAS 34024, 68.9 mm SL, Philippines; CAS 235825, 64.6 mm SL, Philippines

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providing photographs of fresh specimens of N. trimaculatus; volunteers and students of KAUM for curatorial assistance and collection of specimens; and G. Hardy (Ngunguru, New Zealand) for reading the manuscript and providing help with English. This study was supported in part by JSPS KAKENHI Grant Numbers JP23580259, JP26450265, and 20H03311; the JSPS Core-to-Core Program: B Asia-Africa Science Platforms; the "Biological Properties of Biodiversity Hotspots in Japan" project of the National Museum of Nature and Science, Tsukuba, Japan; and "Establishment of Glocal Research and Education Network in the Amami Islands" project of Kagoshima University adopted by the Ministry of Education, Culture, Sports, Science and Technology, Japan.