Epinephelus bruneus Bloch, 1793

Hoshino, Kouichi, Senou, Hiroshi & Nguy ễn, Quân Văn, 2024, Taxonomic Status of the Commercially Important Grouper, Epinephelus bruneus and E. moara (Osteichthys: Perciformes: Epinephelidae), with the Redescription of E. bruneus and the Description of a New Species, Species Diversity 29 (2), pp. 389-407 : 393-399

Epinephelus bruneus Bloch, 1793

[English name: Longtooth Grouper; standard Japanese name: Kue] ( Figs 1 , 4a–c , 5a, e )

Epinephelus bruneus Bloch, 1793: 15–16 , pl. 328, fig. 2 (original description; type locality: erroneously given as Norway); Bloch and Schneider 1801: 300 (description; Japan); Randall 1987: 133 (in part; description); Lee 1990: 39–40, fig. 45 (in part; description; Taiwan); Randall and Heemstra 1991: 108–111, fig. 49 (in part; description); Heemstra and Randall 1993: 119–120, pl. 9, fig. e (in part; description); Senou 1993: 613, unnumbered fig. (pictorial key); Heemstra and Randall 1999: 2490, unnumbered fig. (in part; description); Senou 2000: 718, unnumbered fig. (pictorial key); Senou 2002: 718, unnumbered fig. (pictorial key); Kim et al. 2005: 280, unnumbered fig. (diagnosis; South Korea); Yamada et al. 2007: 577–580, unnumbered fig. (description; East China Sea, Japan); Craig et al. 2011: 97–98, unnumbered figs (in part; description); Senou 2013: 789, unnumbererd fig. (pictorial key).

Serranus mo-ara Temminck and Schlegel, 1843: 10, pl. 4, fig. 2 (erroneously labelled as fig. 1) (original description; type locality: Nagasaki, Japan); Fowler 1937: 279 (in part; description; China) .

Serranus moara : Bleeker 1853: 24–25 (description; Japan); Günther 1859: 133 (description; Japan); Boeseman 1947: 27–28 (description; Japan).

Epinephelus moara : Jordan and Richardson 1910: 456 (description); Tanaka 1925: pl. 152, fig. 419 (figure; Japan); Tanaka 1927: 645–650 (description; Japan); Okada et al. 1935: 143, pl. 66 (description; Japan); Katayama 1960: 105–106, pl. 64 (description; Japan); Chyung 1961: 345, color pl. 28, fig. 139, pl. 127, fig. 617 (description; Korea); Katayama 1984: 131–132, pl. 117, figs I, J (description; Japan); Jin 1985: 19–20 (in part; description; Fujian, China); Machida 1985: 474–475, 671, fig. 254 (description; Okinawa Trough, Japan); Cheng and Zheng 1987: 290 (key); Guo et al. 2008: 107, fig. 1 (left) (description; China); M. Liu et al. 2013: 1686, 1688, 1690, table 1, fig. 2 (diagnosis; Fujian, China).

Epinephelus moara moara : Masuda 1942: 117–118, 121 (note and key; Japan).

Epinephelus brunneus : Lee 1993: 289, pl. 72-6 (in part; description; Taiwan).

Specimens examined. Japan: KPM-NI 6840 , 134.0 mm SL, Nantou, Watarai-gun, Mie Pref ., 14 August 1999; KPM-NI 26963, 120.2 mm SL, Sukumo Bay , Kochi Pref ., 27 February 2010; KPM-NI 26999, 186.3 mm SL, off Nebukawa, Odawara, Kanagawa Pref . (west of Sagami Bay ), 8 June 2010; KPM-NI 35270 [digital image; file no: KPM-NR 107343 (A)], 769 mm SL, Uji-shima Is ., Kagoshima Pref . , 21 July 2013; KPM-NI 36054 , 77.1 mm SL, Manatsuru, Kanagawa Pref . (west of Sagami Bay ), 19 April 2014; KPM-NI 45335 , 62.2 mm SL, Enoshima, Fujisawa, Kanagawa Pref . , 1 October 2017; KPM-NI 24930, 286.5 mm SL, off Nebukawa, Odawara, Kanagawa Pref . , west of Sagami Bay , 15 m deep, 23 September 2009; KPM-NI 24931, 289.3 mm SL, same area with KPM-NI 24930 , 15 m deep, 15 September 2009; KPM-NI 51186, 268.6 mm SL, Totoro-machi, Nobeoka, Miyazaki Pref . , 3 m deep, 17 December 2018; KPM-NI 53022, 177.1 mm SL, Nishi-ku, Hamamatsu, Shizuoka Pref., date unknown; KAUM–I. 50163, 277.7 mm SL, Tanega-shima Is., Kagoshima Pref., 30°37 ′ N, 131°03 ′ E, 12–13 m deep, 23 July 2012; KAUM –I . 63561, 232.5 mm SL, Tanega-shima Is., Kagoshima Pref . , 30°49 ′ N, 131°01 ′ E, 8 September 2014; KAUM–I. 76214, 202.5 mm SL, Tsuno-shima Is., Shimonoseki , Yamaguchi Pref ., 34°22 ′ N, 130°51 ′ E, 19 May 2015; KAUM –I . 80316, 224.7 mm SL, Kami-Koshiki Is., Kagoshima Pref., 31°51.48 ′ N, 129°50.23 ′ E, 7 m deep, 17 October 2015; KAUM –I . 93619, 276.0 mm SL, near Tanega-shima Is., Kagoshima Pref . , 2 m deep, 22 September 2016; KAUM–I. 110011, 292.0 mm SL, Minami Satsuma, Kagoshima Pref . , 31°25 ′ N, 130°12 ′ E, 22 November 2017; KAUM–I. 149161, 203.5 mm SL, Genkai-nada, Fukuoka Pref . , 1 December 2020; FRLM 986 , 160.0 mm SL, Kumano-nada, Hamajima-cho, Shima, Mie Pref . , 34°17.4 ′ N, 136°46.0 ′ E, 15 July 1977; FRLM 02148 , 201.6 mm SL, Zaga Is., Ago Bay, Shima-cho , Wagu, Shima, Mie Pref ., 34°16.7 ′ N, 136°48.1 ′ E to 34°16.4 ′ N, 136°48.5 ′ E, 29 July 1979; FRLM 02891 , 169.3 mm SL, same locality with FRLM 02148 , 14 December 1980; FRLM 04070 , 135.0 mm SL, same locality with FRLM 02148 , 18 July 1983; FRLM 09506 , 102.8 mm SL, same locality with FRLM 02148 , June 1989; FRLM 39231, 111.8 mm SL, same locality with FRLM 02148 , 13 July 2011; SNFR 309, 360 mm SL, edge of continental shelf, East China Sea, date unknown; SNFR 19475, 373 mm SL, Fukue Fish Market, Fukue Is ., Nagasaki Pref., East China Sea , 7 September 2013; SNFR 19488, 326 mm SL, Fukue Fish Market, Fukue Is ., Nagasaki Pref., East China Sea , 10 September 2013; SNFR 19574, 364 mm SL, Fukue Fish Market, Fukue Is ., Nagasaki Pref., East China Sea , 20 September 2013; SNFR 19589, 331 mm SL, Fukue Fish Market, Fukue Is ., Nagasaki Pref., Japan, East China Sea , 1 October 2013; ZMB 5221, 157 mm SL, Japan or China (?), lectotype of E . bruneus (dried skin of left side; digital images); RMNH PISC. D 72, 333 mm SL, Nagasaki, lectotype of S. moara (stuffed specimen; digital images); RMNH PISC. D70 and D71, Nagasaki, paralectotypes of S . moara [stuffed specimens, 21 and 31 cm SL, respectively ( Boeseman 1947), digital images; ZMB 5215, 348 mm SL, Nagasaki, paralectotype of S. moara (stuffed specimen; digital images)] .

Diagnosis. Epinephelus bruneus can be distinguished from the congeners by the combination of following characters: serrae at angle of preopercle gradually and weakly enlarged from dorsal to ventral direction, 2–11 (usually 4–7) ( Fig. 4a–c ); dorsal-fin soft rays 14–16 (usually 15) ( Table 1); anal-fin soft rays 8–9 ( Table 1); caudal fin rounded; scales on lateral body ctenoid; back and side with six irregular broad oblique transverse bars, second bar directed forward becoming nearly horizontal ventral to lateral line and reaching opercular margin only dorsal to its posterior tip ( Figs 1 , 5a ); third bar with an anterior branch connecting with second bar and reaching opercular margin ( Figs 1 , 5a ); fourth band of head broader than eye diameter ( Fig. 5e ).

Description. Meristics are in Table 1, and measurements in Table 2. Body elongated, compressed; deepest at or slightly behind dorsal-fin origin; dorsal profile gently and evenly arched. Caudal peduncle about 1/3 of body depth.

Eye moderately large, entirely in front of middle of head, relatively smaller in larger specimens. Interorbital space gently convex, greater than eye diameter in large (≥ 268.6 mm SL) specimens but smaller than eye diameter in small (≤ 232.5 mm SL) specimens. Snout moderately long, pointed, its length much longer than eye diameter in large (≥ 135.0 mm SL) specimens but subequal or slightly longer in small (≤ 134.0 mm SL) specimens. Nostrils small, subequal, directly in front of eye; anterior one with a flap posteriorly; posterior one located more dorsally, with a slightly elevated rim or absent.

Mouth oblique, large; upper jaw slightly to greatly extending beyond a vertical through rear edge of orbit (extending more in larger specimens); lower jaw projecting beyond upper jaw, more prominent in larger specimens. Teeth on both jaws canine-like or conical, sharply pointed, slightly curved inward. Upper-jaw teeth in two series: teeth in outer series in a row, immovable, anteriormost one enlarged to form a pair of canines; teeth in inner series in a band formed by two or three irregular rows, depressible inward, enlarged anteriorly, one or two pairs of which becoming canines, but much smaller posteriorly. Teeth in lower jaws in two series: those in outer series similar to those of upper jaw in shape and arrangement; those in inner series forming a band anteriorly, with one or two pairs enlarged canines, but in a series posteriorly (larger than corresponding teeth of outer series). Vomerine teeth small, conical, sharply pointed, curved inward, in reversed V-shaped band; palatine teeth as vomerine teeth in shape, in a narrow band. Teeth absent on tongue.

Preopercular margin rounded, serrated posteriorly; serrae gradually and weakly enlarged at angle, 2–11 (usually 4–7) in number ( Fig. 4a, b ; Table 4) but relatively smaller in largest specimen examined (769 mm SL: Fig. 4c ). Opercle tapering posteriorly, with three spines (middle one longest). Margin of interopercle smooth, or with a few to more than ten minute serrae near junction with subopercle. Gill rakers pointed, short, much shorter than gill filaments; rudimentary at uppermost and lowermost parts of gill arch.

Dorsal fin originating posterior to upper end of gill opening and anterior to end of opercular flap; base of spiny portion longer than that of soft-ray portion; dorsal-fin spines with evenly and gently curved margin, longest at third to fifth spines; last spine slightly longer than penultimate; membrane of spinous portion of dorsal fin deeply incised. Anal fin short-based, originating ventral to base of first to second dorsal-fin soft ray; first spine short; second spine slightly shorter than but much more robust than third; longest soft ray somewhat longer than that of dorsal fin. Caudal fin broadly rounded. Pectoral fin subsymmetrical, rounded posteriorly, extending to vertical through base of eighth to ninth dorsal-fin spine. Pelvic fin short, reaching or falling slightly short of vertical through posterior end of pectoral fin when depressed.

Scales small, ctenoid on body side and caudal peduncle, but cycloid in head, anterodorsal part of body, thorax, breast, abdomen, and basal part of each fin; maxillary scaled. Lateral line high, complete, gently curved following dorsal contour, straight in caudal peduncle, ending at base of caudal fin.

Body light brown when fresh, with six irregular broad oblique transverse dark brown bars on back and side, more oblique anteriorly; first bar originating anterior to dorsal-fin origin and immediately bending forward; second bar beginning at anterior portion of spiny dorsal fin and turning forward becoming nearly horizontal ventral to lateral line reaching opercular margin dorsal to its posterior end ( Figs 1 , 5a ); third bar originating at posterior portion of spiny dorsal fin, with an anterior branch that unites with second bar and reaching opercle; fourth and fifth bars extending from soft-ray portion of dorsal fin; sixth bar on caudal peduncle.

Four dark brown bands on head radiating from eye: first band from upper margin of eye to nape; second band from snout, through eye, to confluent with first bar on body; third band from posterior margin of eye, as wide as pupil at anteriormost part, gradually broadened posteriorly, bifurcated at posterior margin of preopercle, its upper branch confluent with second bar on body and anterior branch of third bar on body, its lower branch ending at opercular margin; fourth band from lower margin of eye, its width subequal to or greater than eye diameter, ending at posterior margin of subopercle and interopercle. Margins of fins white except spiny portion of dorsal fin, although white margin sometimes weak or absent in smaller (≤ 186.3 mm SL) specimens. Dorsal margin of spiny and soft dorsal-fin, and upper and lower margins of caudal fin with yellowish area in fresh specimens although color not clear in some specimens.

Distribution. Japanese coasts of the Pacific, Sea of Japan, and East China Sea, from Aomori Prefecture to Okinawa Island ( Senou 2013); Taiwan ( Lee 1990, 1993); South Korea ( Chyung 1961; Kim et al. 2005); and Fujian, China (M. Liu et al. 2013). The type locality and the sampling localities, as well as the reported localities in references are mapped on Fig. 6 . Based on published records that can be identified as E. bruneus , the southern limit of its range appears to be southern Fujian, in the northernmost part of the South China Sea (see Remarks).

Comparison. Based on the number of dorsal-fin soft rays ( Table 3), the size and number of serrae on the preopercular angle ( Fig. 4a–c ; Table 4), and the patterns of the body bars and the head bands (see Remarksbelow), the specimens of E. bruneus examined from Japan represent the “northern form” (= “ E. moara ” of M. Liu et al. 2013) and not the “southern form” (= “ E. bruneus ” of M. Liu et al. 2013).

Remarks. Epinephelus bruneus had long been considered as to not occur in Japan. This is partly because the type locality of E. bruneus was stated by Bloch (1793) as Norway. Although Bloch and Schneider (1801) corrected the location to Japan, this update was largely overlooked. Günther (1859) commented that Bloch (1793) noted “ Norway ” probably because he had received the fish from a friend from Norway. Paepke (1999) noted the locality as “probably China, provided by preacher Chemnitz.” Thus, the type locality is likely either Japan or China.

The name E. moara had long been applied to the grouper called “Kue” in Japan (e.g., Tanaka 1927; Okada et al. 1935; Matsubara 1955; Katayama 1960, 1984; Machida 1985), as well as in South Korea (e.g., Chyung 1961) and Taiwan (e.g., Shen 1984).

In a preliminary synopsis and subsequent revision of the Indo-Pacific groupers (the subfamily Epinephelinae ), Randall (1987) and Randall and Heemstra (1991) relegat- ed E. moara to the junior synonymy of E. bruneus . They did not provide any data supporting their conclusion, but since then, E. bruneus came to be predominantly used (e.g., Heemstra and Randall 1993, 1999; Senou 1993, 2000, 2002, 2013; Kim et al. 2005; Yamada et al. 2007; Craig et al. 2011). Randall and Heemstra (1991) proposed the English name “Longtooth grouper”, which was later adopted as an FAO name of E. bruneus ( Heemstra and Randall 1993) .

Subsequently, Guo et al. (2008) and M. Liu et al. (2013) demonstrated that there are two forms of “longtooth grouper” (“southern form” and “northern form” in this study) in China, and thus removed E. moara from the synonymy of E. bruneus applying the former name to the “northern form” and the latter name to the “southern form” but did not examine the name-bearing types (see Introduction). However, the resurrection of E. moara was followed by Fricke et al. (2024) who regarded it as valid.

In order to establish the taxonomy of “the northern form” and “the southern form”, it was necessary to examine the relevant name-bearing types. However, the lectotype of E. bruneus (ZMB 5221) is the left-side skin ( Fig. 3a–c ), and that of S. moara (RMNH D70) is a stuffed specimen ( Fig. 3d–f ), and the body colors of both specimens (important characters for identification of groupers) are lost. Thus, it was necessary to identify the lectotypes with reference also to the original illustrations of Bloch (1793) ( Fig. 2b ) and Temminck and Schlegel (1843) ( Fig. 2c ), which refer to the original coloration.

Randall and Heemstra (1991) diagnosed E. bruneus by the following combination of characters: three opercular spines, 11 dorsal-fin spines and 13–15 dorsal-fin soft rays, three anal-fin spines and eight soft rays, caudal fin round- ed, ground color grayish brown with six oblique transverse dark brown bars on the body, three or four bands radiating posteriorly from eye, absence of other conspicuous spots on body and head, and slightly or moderately enlarged serrae at the corner of the preopercle. From the lectotypes and the original illustrations of E. bruneus and E. moara , all of these characters were confirmed excluding all other groupers studied by Randall and Heemstra (1991). From the Indo-Pacific, after Randall and Heemstra (1991) as mostly covered by Parenti and Randall (2020), five additional species of Epinephelus were described (Tucker et al. 2016; Frable et al. 2019; Johnson and Wortington Wilmer 2019; Wu et al. 2020; Nakamura and Motomura 2021), and four nominal species were resurrected from their synonymies ( Randall et al. 2013; Psomadakis et al. 2015; Nakamura and Motomura 2021; Cao et al. 2022). All of these species are distinguishable from the lectotypes and the original illustrations of E. bruneus and S. moara by the same combination of characters. Accordingly, the lectotypes could be identified as E. bruneus sensu Randall and Heemstra (1991) .

Data comparing the specimens examined with the lectotypes of E. bruneus and S. moara , including data from other references, are presented in Tables 3, 4. In both of the lectotypes, there are 15 dorsal-fin soft rays ( Fig. 3c, f ; Table 3) and 5 (but 4 on the left side in the S. moara lectotype) serrae at the preopercular angle that are gradually and weakly enlarged compared with other serrae ( Fig. 3b, e ; Table 4). These characters agree with specimens of the “northern form” (= “ E. moara ” of M. Liu et al. 2013) and differ from those in the “southern form” (= “ E. bruneus ” of M. Liu et al. 2013) that has 13–14 dorsal-fin rays and 2–4 abruptly and remarkably enlarged serrae at the preopercular angle ( Fig. 4d–f ).

In addition, in the original illustrations, the second bar on the body is directed anteriorly and becomes nearly horizontal ventral to the lateral line, reaching the opercle wholly dorsal to the posterior tip of the opercle ( Fig. 5c, d ) agreeing with the “northern form” (M. Liu et al. 2013: fig. 2a, b; Fig. 5a ). However, in the “southern form” the bar is directed ventrally and steeply oblique, reaching the opercle dorsal and ventral to its posterior tip (M. Liu et al. 2013: fig. 1b; Fig. 5b ). The anterior branch of the third bar on the body is depicted in the published illustration and its original illustration of E. bruneus in Bloch (1793) ( Figs 2a, c , 5c ) (not illustrated in the original illustration of S. moara where the area is hidden by the pectoral fin; Fig. 5d ), which agrees with the “northern form” (M. Liu et al. 2013: fig. 2a, b; Fig. 5a ), whereas it is absent in the “southern form” (M. Liu et al. 2013: fig. 1b; Fig. 5b ). The fourth band on the head is broader than the eye diameter in the original illustration of E. bruneus ( Fig. 5g ) (not illustrated in that of E. moara ; Fig. 2c ) and in the “northern form” (M. Liu et al. 2013: fig. 2a, b; Fig. 5e ), whereas it is much narrower than the pupil in the “southern form” (M. Liu et al. 2013: fig. 1b; Fig. 5f ). All these characters in the lectotypes and the original illustrations of E. bruneus and S. moara agree with the “northern form” (= “ E. moara ” of M. Liu et al. 2013). Accordingly, the two lectotypes are considered conspecific and both of the scientific names Epinephelus bruneus and Serranus moara are applicable to the “northern form”. Its valid name is E. bruneus Bloch, 1793 , and thus E. moara ( Temminck and Schlegel, 1843) is a junior synonym based on the Principle of Priority (Article 23 of the International Code of Zoological Nomenclature; International Commission on Zoological Nomenclature 1999). This conclusion supports the proposal of Randall (1987) and Randall and Heemstra (1991). The “southern form” is considered distinct from all its known congeners and is therefore described below as a new species.

The original illustration of E. bruneus ( Fig. 2b ) agrees with the lectotype in artificially small head length (35.1% of SL; 36.9% in the lectotype when measured with a caliper on the image printed on paper), whereas 39.1%–43.2% in the examined specimens of E. bruneus ( Table 2). Also, the original illustration agrees with the lectotype in the count of dorsal-fin soft rays (15; Table 3), and five slightly enlarged serrae of the preopercular angle. These indicate that the original illustration was modeled after the lectotype, which had been already prepared as the left-side skin.

“ Epinephelus moara ” and “ E. bruneus ” reported from Japan in Jordan and Richardson (1910), Tanaka (1925, 1927), Katayama (1960), Machida (1985), and Yamada et al. (2007) can be identified as E. bruneus from the following characters: 14–15 dorsal-fin soft-rays, preopercular serrae at angle not remarkably enlarged [e.g., described as “slightly enlarged” by Tanaka (1927) and “somewhat enlarged” by Katayama (1960)], scales mostly ctenoid, and the patterns of the body bars and/or the head bands as depicted in the picture or illustration. Based on the patterns of the body bars and the head bands, the pictures of “ E. moara ” or “ E. bruneus ” in various references ( Okada et al. 1935: pl. 66; Masuda et al. 1975: pl. 48; Katayama 1984: pl. 117, fig. I; Masuda and Kobayashi 1994: figs 7, 8 on p. 112; Senou 1997: unnumbered figs; Kuriiwa 2018: unnumbered fig.; Sakurai 2019: unnumbered fig.; Shimose 2021: unnumbered fig.) are E. bruneus . Similarly, the records from Taiwan of “ E. moara ” in Shen (1984) and “ E. brunneus ” in Lee (1990, 1993) and those from South Korea as “ E. moara ” in Chyung (1961) and as E. bruneus in Kim et al. (2005) can be clearly identified as E. bruneus , from the patterns of bars on the body and the head bands in the figures.

Notes on fishery, aquaculture, and conservation. In Japan and China, E. bruneus is highly valued in the market (e.g., Nakagawa et al. 2015; Gao et al. 2018). In Japan, the nation-wide landing statistics are not available, but in Nagasaki Prefecture (westernmost part of Japanese mainland), approximately 150 tons (probably including artificially reared and released fish) were landed in 2017 ( Agari 2018). Also in Japan, the technology for the mass production of larvae and juveniles is established ( Teruya and Yoseda 2006), and the annual number of artificially reared juveniles that are released to the wild has increased from 90000 individuals in 2007 to 274000 in 2016 ( Japan Fisheries Research and Education Agency 2018). Note that the now invalid scientific name E. moara is used in some recent aquaculture studies (e.g., Miyaki et al. 2005; Guo et al. 2009, 2014; Tian et al. 2017; Gao et al. 2018; Zhang et al. 2020), whereas the valid name E. bruneus is used in others (e.g., Q. Liu et al. 2013; Inoue et al. 2016). In the IUCN Red List, this species is assessed as DD (data deficient) ( Liu 2018) under the name of E. moara .

Figures

Fig. 1. Epinephelus bruneus. a, KAUM–I. 80316, 224.7mm SL, Japan; b, KPM-NI 26963, 120.2mm SL, Japan; c, KPM-NI 35270, 769mm SL, Japan.

Fig. 4. Variation of serrae of the preopercular angle in Epinephelus bruneus (= “northern form”; a–c) and E. randalli sp. nov. (= “southern form”; d–f). a, KPM-NI 26999, 186.3mm SL, Japan; b, KPM-NI 26963, 120.2mm SL, Japan; c, KPM-NI 35270, 769mm SL, Japan; d, NSMT- P 66257, 143.3mm SL, Vietnam, holotype; e, NSMT-P 68341, 164.4mm SL, Vietnam, paratype; f, NSMT-P 68411, 334mm SL, Vietnam, paratype.

Fig. 5. Second and third bars of body (a–d) and fourth bar of head (e–g) of Epinephelus bruneus (= “northern form”) and E. randalli sp. nov.(= “southern form”). a, e, Epinephelus bruneus, KAUM–I. 80316, 224.7mm SL, Japan; b, f, E. randalli sp. nov., NSMT-P 66257, 143.3mm SL, Vietnam, holotype; c, g, original illustration for description of E. bruneus (Bloch 1793); d, original illustration for description of Serranus moara (Temminck and Schlegel 1843). Downward arrow, second bar of body ventral to lateral line; upward arrow, anterior branch of third bar of body; rectangle, fourth bar of head.

Fig. 6. Distribution of Epinephelus bruneus and E. randalli sp. nov. Type locality of Serranus moara (= E. bruneus, solid circle); localities of E. bruneus specimens examined in this study (large open circles on magnified map); localities of E. bruneus in references (small open circles on wide area map); type locality of E. randalli sp. nov. (solid square); localities of paratypes of E. randalli sp. nov. (large open squares); localities of E. randalli sp. nov. in references (small open squares).

Fig. 3. Lectotypes of Epinephelus bruneus Bloch, 1793 (a–c: ZMB 5221, 157mm SL) and Serranus moara Temminck and Schlegel, 1843 (d–f: RMNH PISC. D72, 333mm SL). a, d, Left side; b, e, serrae at preopercular angle (indicated by arrows); c, f, dorsal-fin spines (indicated by Roman numerals) and soft-rays (indicated by Arabic numerals) seen from right side.

Fig. 2. Illustrations of Epinephelus bruneus Bloch, 1793 and Serranus moara Temminck and Schlegel, 1843. a, Published illustration in original description of E. bruneus; b, its original illustration deposited in ZMB; c, original illustration for description of S. moara deposited in RMNH.