Brachypterois serrulata (Richardson 1846)

Matsunuma, Mizuki, Sakurai, Makoto & Motomura, Hiroyuki, 2013, Revision of the Indo-West Pacific genus Brachypterois (Scorpaenidae: Pteroinae), with description of a new species from northeastern Australia, Zootaxa 3693 (4), pp. 401-440 : 416-425

Brachypterois serrulata (Richardson 1846)

[English name: Sawcheek Scorpionfish]

[Standard Japanese name: Nokogiri-kasago]

( Figures 2 , 6 B, 7, 8D–F, 9D–F, 10–13, 14A, 15, 18–19, 20B; Tables 1 , 3 )

Sebastes serrulatus Richardson 1846: 215 [original locality: “Sea of China ”; type locality, based on neotype designated herein: off Dong Kang, Pingtung, southern Taiwan]; Günther 1860: 106 (type catalog of BMNH)

Scorpaenodes serrulatus: Chu 1931: 142 (in part?; Sea of China; listed)

Brachypterois serrulatus: Chu 1962: 861 , fig. 692 (South China Sea; short description); Chu et al. 1963: 462, fig. 348 (East China Sea coast of China; short description); Chen 1981: 47, figs. 9-29, 32 ( Taiwan; short description); Kanayama & Amaoka 1981: 181, figs. 1, 2 (in part; Tosa Bay, Kochi, Japan; short description); Shen 1984: 185, unnumbered fig. on p. 192 (in part?; Taiwan; key); Amaoka 1984: 302, pl. 360-G [in part?; Japan; figure taken from Fowler 1938]; Eschmeyer 1986: 465 (in part; Indo-West Pacific; key); Jean & Kuo 1988: 55 (in part?; Taiwan; listed); Wang 1993: 112 (in part?; South China Sea; listed); Ishida 1994: 5 (as phylogenetic material); Huang 1994: 739 (in part?; China, Taiwan and South China Sea; listed); Qingwen et al. 1995: 549, fig. 736 (in part?; Japan to South China Sea; short description); Allen 1997: 76, pl. 19, fig. 7 (in part; Indo-West Pacific; ecological note and diagnosis); Chen et al. 1997: 168, unnumbered fig. on top (in part?; Spratly Islands to China; short description); Poss 1999: 2299, 2310, unnumbered fig. on bottom on p. 2310 (in part; Indo-West Pacific; key and short description;); Randall & Lim 2000: 604 (in part?; South China Sea; listed); Matsuura et al. 2001: 106 (Sanya Bay, Hainan Island, China; listed; specimen examined herein); Adrim et al. 2004: 119 (in part?; Anambas Islands, South China Sea; listed)

Brachypterois serrulata: Nakabo 1993: 493 , unnumbered fig. on top [ Kochi, Japan; key; figured specimen from Gulf of Thailand (T. Nakabo, 2012, pers. comm.) examined herein]; Ishida 1997: 209, unnumbered fig. on p. 209 (Tosa Bay, Kochi, Japan; short description; Indo-West Pacific; figured specimen examined herein); Nakabo 2000: 567, unnumbered fig. on top ( Kochi, Japan; figure taken from Nakabo 1993); Murai 2001: 172, unnumbered fig. on middle (Iburi, Kochi, Japan; short description;); Shinohara et al. 2001: 314 (in part; Tosa Bay, Kochi, Japan; listed; based on Amaoka 1984 and Nakabo 1993); Mandrytsa 2001:276 (as phylogenetic material; key); Nakabo 2002: 567, unnumbered fig. on top ( Kochi, Japan; figure taken from Nakabo 1993); Ide et al. 2003: 24 (Tosa Bay, Kochi, Japan; listed); Xinbo 2006: 270, fig. 156 ( China and Taiwan; description); Shao et al. 2008: 245 (in part?; Taiwan; listed; based on Chen 1981)

Dendrochirus bellus (not of Jordan & Hubbs): Rainboth et al. 2012: 80, pl. 55, fig. 1137 (Gulf of Thailand; listed)

Neotype. ASIZP 62399 (X), 92.8 mm SL, Dong-kang, Pingtung, southern Taiwan (22°31′N, 120°22′E), 100 m, J.- H. Wu, bottom trawl, 1 Mar. 2001.

Non-type specimens. 112 specimens, 48.0– 99.3 mm SL— JAPAN (following BSKU specimens all collected by bottom trawl from Tosa Bay, Kochi): BSKU 12876, 59.2 mm SL, obtained at Usa Port, Sept. 1968; BSKU 42455, male, 84.2 mm SL, obtained at Mimase Port, 26 Nov. 1985; BSKU 52300, male, 91.8 mm SL, BSKU 52301, male, 93.2 mm SL, BSKU 52302, male, 72.9 mm SL, obtained at Irino Port, 10 July 2000; BSKU 52359, male, 73.2 mm SL, obtained at Irino Port, 2 Aug. 2000; BSKU 52411, male, 71.4 mm SL, obtained at Kochi City Central Fish Market, 19 Jan. 1995; BSKU 54828, 95.2 mm SL, obtained at Irino Port, 3 July 2000; BSKU 57572, 67.2 mm SL, obtained at Irino Port, 22 Feb. 2002: BSKU 57736, male, 69.3 mm SL, obtained at Irino Port, 12 Sept. 2001; BSKU 57874, male, 96.7 mm SL, obtained at Irino Port, 20 May 1999; BSKU 59821, female, 70.7 mm SL, obtained at Kamikawaguchi Port, 31 July 1997; BSKU 62168, 64.8 mm SL, obtained at Irino Port, 3 Oct. 2002; BSKU 62457, male, 92.0 mm SL, obtained at Susaki Port, 10 Oct. 2003; BSKU 67433, 71.5 mm SL, obtained at Susaki Port, 4 July 2002; BSKU 67592, male, 99.3 mm SL, BSKU 67593, 91.1 mm SL, obtained at Irino Port, 20 Mar. 1999; BSKU 70509, 71.3 mm SL, BSKU 70510, 86.5 mm SL, BSKU 70511, male, 65.7 mm SL, obtained at Kamikawaguchi Port, 20 Feb. 1997; BSKU 86119, male, 91.4 mm SL, BSKU 86121, female, 72.5 mm SL, obtained at Irino Port, 12 Mar. 1999; BSKU 86123, male, 73.4 mm SL, BSKU 86124, female, 67.2 mm SL, obtained at Irino Port, 12 Mar. 1999; BSKU 82651, 48.0 mm SL, 75 m, R/V Toyohata-maru, 2 Nov. 1989; BSKU 82652, 54.6 mm SL, BSKU 82653, female, 67.2 mm SL, BSKU 82654, male, 74.6 mm SL, obtained at Kamikawaguchi Port, H. Endo, 20 Mar. 1997; BSKU 86120, 76.6 mm SL, obtained at Irino Port, 12 Mar. 1999; BSKU 86150, 63.1 mm SL, obtained at Kamikawaguchi Port, 27 Apr. 1999; BSKU 86476 (2), 62.1–63.4 mm SL, obtained at Kamikawaguchi Port, 27 Apr. 1999; BSKU 89187, 52.1 mm SL, obtained at Saga Port, 27 Oct. 2006; BSKU 106470, 85.7 mm SL, BSKU 106471, 82.4 mm SL, BSKU 106472, female, 68.0 mm SL, BSKU 106473, 74.9 mm SL, obtained at Irino Port, 11 June 1999; FAKU 132868 (5), 68.9–81.2 mm SL, Irino, Tosa Bay, Kochi, 26 June 1997; *HUMZ 36292, 72.1 mm SL, HUMZ 36563, 71.7 mm SL, *HUMZ 36638, 61.3 mm SL, Tosa Bay, Kochi, K. Amaoka, trawl; KAUM–I. 6876 (X), male, 66.9 mm SL, 1 km north off Izashiki Port, Sata, Minamiosumi, Kagoshima (31°05′N, 130°41′E), 40 m, M. Yamada, set net, 27 June 2007; KAUM–I. 44569, 80.9 mm SL, KAUM–I. 44570, 89.4 mm SL, KAUM–I. 44571, 78.6 mm SL, KAUM–I. 44572, 75.6 mm SL, KAUM–I. 44573, female, 76.0 mm SL, KAUM–I. 44574, 71.2 mm SL, Tosa Bay, Kochi (obtained at Irino Port), bottom trawl, 11 June 1999; *OMNH 24502, 67.1 mm SL, Tosa Bay, Kochi, H. Hanazaki, trawl, 20 May 1997; SNFR 11585, 86.1 mm SL, Tachibana Bay, Nagasaki, 17 Nov. 1987. EAST CHINA SEA: FAKU 12094, 83.0 mm SL, 1948–1949. CHINA: MNHN A-4310 (2), 74.5–84.6 mm SL; NSMT-P 60871, 69.1 mm SL, Sanya Bay, Hainan Island, 16 m, G. Shinohara, 22 Nov. 1997. TAIWAN: ASIZP 62365, 74.0 mm SL, Dong-kang, Pingtung (22°31′N, 120°22′E), 100 m, J.-H. Wu, 16 May 2001; ASIZP 62400, 65.7 mm SL, Fong Kang, Pingtung (22°26′N, 120°38′E), 100 m, J.-H. Wu, 28 Feb. 2001; *ASIZP 62992, 60.5 mm SL, water entrance way of 1st Nuclear Power Plant, Shihmen, Taipei (25°03′N, 121°56′E), Y.-C. Liao, 1 Apr. 2005; KAUM–I. 39283 (X), female, 60.2 mm SL, KAUM–I. 39284 (X), male, 70.1 mm SL, off Tashi, Yilan, KAUM Fish Team, bottom trawl, 6 July 2011; KAUM– I. 46474, female, 75.6 mm SL, off Tashi, Yilan, H.-C. Ho, bottom trawl, 18 Nov. 2007; NTOU 20071117 -3, 76.2 mm SL, Hsiao-liou-ciou, Pingtung, 17 Nov. 2007; NMMB-P 4115, 79.3 mm SL, Fong-kang, Pingtung, J.-H. Wu, 23 Aug. 2001; NMMB-P 5181, 83.1 mm SL, Dong-kang, Pingtung, M.-C. Yu, 16 Feb. 1960; *NTUM 3854, 51.2 mm SL, off Tashi, Yilan, 18 July 1977. VIETNAM: FRLM 31479, 57.6 mm SL, FRLM 31647, 65.7 mm SL, Nha Trang, S. Kimura et al., 7 Oct. 2004; HUMZ 190502, 60.7 mm SL, HUMZ 190503, 66.3 mm SL, HUMZ 190539, 62.8 mm SL, Nha Trang, H. Imamura, 8 Oct. 2004; HUMZ 190610, 61.6 mm SL, Nha Trang, H. Imamura, 7 Oct. 2004; NMMB-P 12648 (2), 69.6–70.1 mm SL, Nha Trang, H.-C. Ho, 16 Apr. 2009; NSMT-P 68863, 57.0 mm SL, Nha Trang, G. Shinohara et al., 6 Dec. 2003; NSMT-P 69054, 55.4 mm SL, Nha Trang, K. Matsuura, 6 Dec. 2003; NSMT-P 70619, 59.7 mm SL, Nha Trang, K. Shibukawa, 7 Oct. 2004. MALAYSIA: KAUM–I. 12313, male, 77.9 mm SL, off Kota Kinabalu, Sabah, Borneo, G. Ogihara, 24 Sept. 2008; KAUM–I. 22034, male, 82.4 mm SL, off Kota Kinabalu, Sabah, Borneo, G. Ogihara, 11 Aug. 2009; KAUM–I. 49319, female, 67.1 mm SL, off Kota Kinabalu, Sabah, Borneo, H. Nishiyama, 4 Aug. 2012. THAILAND (following specimens all collected from Gulf of Thailand): KAUM–I. 23300, male, 72.5 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M. Matsunuma, trawl, 10 Sept. 2009; KAUM–I. 23436 (X), female, 62.5 mm SL, KAUM–I. 23990 (X), male, 88.2 mm SL, KAUM–I. 23991 (X), male, 86.0 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M. Matsunuma, trawl, 24 Sept. 2009; KAUM–I. 23882, male, 74.0 mm SL, KAUM–I. 23883 (X), female, 71.9 mm SL, KAUM–I. 23884 (X), female, 86.3 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M. Matsunuma, trawl, 26 Sept. 2009; KAUM–I. 24049 (X), female, 53.2 mm SL, KAUM–I. 24050 (X), female, 63.9 mm SL, KAUM–I. 24051 (X), female, 66.0 mm SL, KAUM–I. 24052 (X), female, 72.7 mm SL, KAUM–I. 24053 (X), female, 76.2 mm SL, KAUM–I. 24054 (X), female, 76.5 mm SL, KAUM–I. 24055 (X), male, 81.2 mm SL, KAUM–I. 24056 (X), male, 97.9 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, M. Matsunuma, trawl, 5 Oct. 2009; KAUM–I. 33225, male, 86.2 mm SL, obtained at fish market in Samut Sakhon Province, T. Yoshida, trawl, 30 Nov. 2010; KAUM–I. 47467, 81.5 mm SL, KAUM–I. 47468, male, 94.5 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, T. Yoshida, trawl, 20 June 2012; KAUM–I. 47472, 99.2 mm SL, KAUM–I. 47473, female, 82.5 mm SL, KAUM–I. 47474, 82.2 mm SL, KAUM–I. 47475, 93.3 mm SL, KAUM–I. 47476, 90.3 mm SL, KAUM–I. 47477, 87.2 mm SL, KAUM–I. 47478, male, 87.1 mm SL, KAUM– I. 47479, 80.0 mm SL, KAUM–I. 47480, female, 70.0 mm SL, obtained at Mahachai fish market, Samut Sakhon Province, T. Yoshida, trawl, 20 June 2012; KAUM–I. 47799, female, 68.4 mm SL, obtained at Klong Wan fish landing port, Prachuab Khirikhan Province, T. Yoshida, trawl, 24 July 2012; URM 12150, 69.4 mm SL, obtained at Song Khla fish market, H. Senou et al., 21 Oct. 1983. INDONESIA: NCIP 5002 (3), 43.2–64.0 mm SL, off Seraya Island, Natuna Islands, trawl, 16 Apr. 2011.

Diagnosis. A species of Brachypterois with the following combination of characters: pectoral-fin rays 14–16 (modally 15); scale rows in longitudinal series 44–51 (47); scales above lateral line 4–6 (5); scale rows between last dorsal spine base and lateral line 4–7 (6); pre-dorsal scale rows 4–8 (5); suborbital scale rows 0–4 (2) ( Fig. 6 B); total gill rakers 15–19 (17); head length 38.3–43.1 (mean 40.2) %SL ( Fig. 7 ); orbit diameter 11.1–14.5 (12.4) %SL; upper-jaw length 18.6–21.5 (20.2) %SL; longest dorsal-fin soft ray length 18.0–23.0 (20.2) %SL; longest anal-fin soft ray length 21.0–27.0 (23.7) %SL; caudal-fin length 30.4–44.7 (35.7) %SL; posterior lacrimal spine usually directed posteriorly, not curved upward distally ( Fig. 8 D–F); spine(s) on corner of outer angular ridge not developed, size similar to other spines on ridge, directed ventrally ( Fig. 9 D–F); spines usually absent on median lateral ridge of maxilla, rarely 1–3 spines present in large adults (> 80 mm SL) ( Fig. 14 A); large number of small spots on caudal fin, 5–26 (mean 13.3) spots on longest caudal-fin ray ( Figs. 10 , 12–13 ).

Color of fresh specimens. Head and body pinkish to reddish-brown, darker dorsally, paler ventrally; a poorly defined large black blotch (subequal to orbit diameter) on opercle. Five dark red to black bands on head; anteriormost band from anterior margin of eye running over snout to tip of lower jaw; second extending from anteroventral margin of eye onto lacrimal and both jaws; third from just below eye to posterodorsal corner of maxilla; fourth from posteroventral margin of eye to posterior portion of opercle; fifth from posterior margin of eye to opercle, more or less continuous with blotch on opercle. Five poorly defined dark or black vertical bands on body; anteriormost band extending from between first and third dorsal-fin spine bases; second from between fourth and sixth spine bases; third from between eighth and tenth spine bases; fourth Y-shaped, upper margin of anterior branch below last spine and third soft ray bases, upper margin of posterior branch below sixth and penultimate soft ray bases, both branches connected at level of lateral line or just below dorsal-fin base; fifth on caudal peduncle just in front of caudal-fin base. Ground color of fins similar to that of head and body. Dorsal fin with a large dark red to black blotch on spinous portion between approximately sixth and ninth spines; about 4–5 more or less dotted dark red lines on soft-rayed portion. Anal fin with about 6–8 somewhat dotted dark red lines. Pectoral fin darker distally, with about 6–10 oblique narrow dark red bands. Pelvic fin with about 6–7 narrow dark red to black bands. Caudal fin membrane translucent with dark red band marginally (relatively wider in small specimens); about 60– 110 small dark red spots [maximum diameter 1.3–3.0 (mean 2.4) times in caudal-fin ray width at position where spot present on ray; based on 13 preserved specimens] on rays. Eye vivid yellow; iris dark blue.

Color of preserved specimens. Head and body creamy-white to brown, darker dorsally, paler ventrally; formerly reddish markings brownish and indistinct. Eye dark blue to gray; iris white to dark blue.

Distribution. Brachypterois serrulata is distributed in the northwestern Pacific, from the South China Sea north to southern Japan, including the Natuna Islands, Borneo, the Gulf of Thailand, Ng Tran Bay, Vietnam and Taiwan. The northernmost records from southern Japan are from Tosa Bay, Shikoku Island ( Fig. 11 ). The 5 specimens were collected at depths of 40–100 m (most from 100 m).

Remarks. Meristics and morphometrics of specimens of B. serrulata examined are shown in Tables 1 , 3 . Brachypterois serrulata was originally described by Richardson (1846) as Sebastes serrulatus on the basis of a single specimen (type locality stated as “Sea of China ”) which is apparently lost (see Introduction). This species is regarded herein as the valid name for this common species in the northwestern Asian waters based on our specimen examinations in museum collections and field surveys in Japan, Taiwan, Thailand and Malaysia. The original description by Richardson (1846) did not include distinct diagnostic characters, although the detailed description of head spines and ridges were given. However, he did not mention any spine on the median lateral ridge of the maxilla which might suggest an absence of spines on the ridge in the type specimen of S. serrulatus , meeting the condition of B. serrulata recognized herein. According to our examination, specimens of B. serrulifer with similar size (4 inches length) to the holotype of S. serrulatus have 0–12 (mean 3.1) spines on the median lateral ridge on the maxilla [33 specimens examined of ca. 10 cm total length (65.4–75.2 mm SL)].

......continued on the next page a Morphometric data from 78 specimens.

b Upper unbranched ray + branched ray + lower unbranched rays = total rays ton left side of body/ total rays on right side. c Rakers on upper limb + rakers on lower limb = total rakers. d At vertical midline of eye.

e At posterior end of preocular spine base.

DS and LL indicate dorsal-fin spine base and lateral line, respectively.

Because the similarity of S. serrulatus to its congeners and the confusion in the literature between the identities of S. serrulatus and B. serrulifer , a neotype for S. serrulatus is necessary to represent the species. A specimen (ASIZP 62399, 92.8 mm SL; Fig. 12 C) is therefore proposed as the neotype for S. serrulatus . Accordingly, southern Taiwan becomes the type locality of S. serrulatus under the Article 76.3 of the International Code on Zoological Nomenclature (ICZN, 1999).

Although Kanayama & Amaoka (1981) first recorded B. serrulata from Japan based on four specimens collected from Tosa Bay, Kochi, the figured specimen (HUMZ 36733) was identified here as B. serrulifer . Another specimen (HUMZ 35083) examined by them, not examined in this study, was examined and dissected by Ishida (1994) during a phylogenetic study. The remaining two specimens (BSKU 2270 and BSKU 7262) are apparently missing (H. Endo, 2012, pers. comm.).

Rainboth et al. (2012) reported B. serrulata from the Gulf of Thailand but provided a figure (pl. 55, fig. 1136) taken from Gloerfelt-Tarp & Kailola (1984: unnumbered fig. on p. 108). The photographed specimen (NTM S. 11016-003), collected from Indonesia, was identified as B. serrulifer by us. On the other hand, a figure identified as Dendrochirus bellus ( Jordan & Hubbs 1925) ( Scorpaenidae ) in Rainboth et al. (2012: pl. 55, fig. 1137) is clearly a B. serrulata .

Biological notes. Among specimens with sex determined, an individual of 60.2 mm SL (KAUM–I. 39283), collected from off southern Taiwan in July 2011, was the smallest female examined in the present study that possessed ripe eggs approaching terminal stage (about 0.4 mm in diameter). Two specimens (BSKU 52359, 73.2 mm SL; BSKU 59821, 70.7 mm SL) from Tosa Bay, Kochi, Japan and three specimens (KAUM–I. 23436, 62.5 mm SL; KAUM–I. 23883, 71.9 mm SL; KAUM–I. 23884, 86.3 mm SL) from the Gulf of Thailand, all five specimens with developed gonads were collected between July and September, inclusive ( Fig. 15 A). Two males (KAUM–I. 23882, 74.0 mm SL, Gulf of Thailand; KAUM–I. 12313, 77.9 mm SL, off Borneo) collected in September had developed testes at the late spermatogenic stage ( Fig. 15 B). These data appear to indicate a breeding season of B. serrulata between July and September. Stomach content analyses (four specimens only) indicated that B. serrulata feeds on small shrimps and crabs.

Geographical variation. Some measurement variations were noted between specimens from Indonesia (Natuna Islands) and Thailand (Gulf of Thailand), and those from other localities, including Malaysia (Borneo) and north of Vietnam (see list of specimens examined). The former tended to have slightly greater proportions for head length [38.7–43.1 (mean 40.7)% SL in Indonesia and Thailand specimens vs. 38.3–41.9 (40.0)% SL in other specimens], and the greatest length of the longest dorsal-fin soft ray [19.9–23.0 (21.1)% SL vs. 18.1–21.4 (19.8)% SL], longest anal-fin soft ray [23.7–27.0 (24.9)% SL vs. 21.0–25.9 (23.3)% SL], longest pectoral-fin ray [49.6– 62.8 (55.0)% SL vs. 44.0–56.7 (50.2)% SL], pelvic-fin spine [14.9–19.3 (17.3)% SL vs. 13.5–19.2 (16.0)% SL], longest pelvic-fin soft ray [28.7–32.0 (30.5)% SL vs. 25.7–32.4 (29.1)% SL] and longest caudal-fin ray [34.9–44.7 (37.7)% SL vs. 30.4–38.7 (34.9)% SL] ( Table 3 ). However no other significant morphological differences were found between the two groups. In addition, all of the above characters change significantly with growth in Brachypterois (see under Morphological changes with growth). Such minor morphometric differences were therefore regarded here as representing geographic variations of B. serrulata .

Figures

FIGURE 2. Diagram of mandible of Brachypterois serrulata, showing series of spines and ridges. SMS spines around mandibular symphysis; SODR spines on outer dentary ridge; SIDR spines on inner dentary ridge; SCDR spines on central dentary ridge; SOAR spines on outer angular ridge; SIAR spines on inner angular ridge; SP sensory pore. Left to anterior. Arrow shows position of lower lip attachment to dentary.

FIGURE 6. Relationships of number of suborbital scale rows to standard length in (A) Brachypterois curvispina sp. nov., (B) B. serrulata and (C) B. serrulifer.

FIGURE 7. Relationships of: (A) head length; (B) orbit diameter; (C) upper jaw length; (D) longest dorsal-fin soft ray length; (E) longest anal-fin soft ray length; and (F) caudal-fin length to standard length in Brachypterois curvispina sp. nov. (stars, solid lines), B. serrulata (circles, dotted lines) and B. serrulifer (triangles, chain lines). H 1 holotype of B. curvispina; H 2 holotype of B. serrulifer; N neotype of Sebastes serrulatus.

FIGURE 8. Spines on lacrimal bones of Brachypterois curvispina sp. nov. (A – C), B. serrulata (D – F) and B. serrulifer (G – I). (A) CSIRO H 7204 - 02, male, 78.5 mm SL, holotype of B. curvispina; (B) CSIRO H 6784 - 02, female, 75.2 mm SL, paratype of B. curvispina; (C) CSIRO H 6750 - 01, sex undetermined, 57.4 mm SL; (D) KAUM – I. 24056, male, 97.9 mm SL; (E) KAUM – I. 24054, female, 76.5 mm SL; (F) KAUM – I. 24050, female, 63.9 mm SL; (G) KAUM – I. 22033, male, 83.0 mm SL; (H) CAS 60396 (1 of 19 specimens), female, 67.9 mm SL; (I) CAS 49547 (1 of 14), sex undetermined, 57.6 mm SL. LLS lateral lacrimal spine; ALS anterior lacrimal spine; PLS posterior lacrimal spine. Left to anterior. Bars indicate 5 mm.

FIGURE 9. Spines on angular bones of Brachypterois curvispina sp. nov. (A – C), B. serrulata (D – F) and B. serrulifer (G – I). (A) CSIRO H 7204 - 02, male, 78.5 mm SL, holotype of B. curvispina; (B) CSIRO H 6784 - 02, female, 75.2 mm SL, paratype of B. curvispina; (C) CSIRO H 6750 - 01, sex undetermined, 57.4 mm SL; (D) KAUM – I. 24056, male, 97.9 mm SL; (E) KAUM – I. 24054, female, 76.5 mm SL; (F) KAUM – I. 24050, female, 63.9 mm SL; (G) KAUM – I. 22033, male, 83.0 mm SL; (H) CAS 60396 (1 of 19 specimens), female, 67.9 mm SL; (I) CAS 49547 (1 of 14), sex undetermined, 57.6 mm SL. Arrows indicate spine (s) on posterior corner of outer angular ridge. Left to anterior. Bars indicate 1 mm.

FIGURE 10. Relationships of number of spots on longest caudal-fin ray to standard length in Brachypterois curvispina sp. nov. (stars), B. serrulata (circles) and B. serrulifer (triangles; dotted symbols indicate specimens from the Indian Ocean, remaining specimens from the Pacific Ocean). H holotype of B. serrulifer. Note: counts unable to be made in holotype of B. curvispina and neotype of Sebastes serrulatus.

FIGURE 11. Distributional records of Brachypterois curvispina sp. nov. (stars), B. serrulata (circles) and B. serrulifer (triangles) based on collected specimens examined in this study. Closed symbols indicate type localities.

FIGURE 12. Preserved specimens of Brachypterois serrulata. (A) NCIP 5002 (1 of 3 specimens), 43.2 mm SL; (B) KAUM – I. 44571, 78.6 mm SL; (C) ASIZP 62399, 92.8 mm SL, neotype of Sebastes serrulatus.

FIGURE 13. Fresh specimens of Brachypterois serrulata. (A) NCIP 5002 (1 of 3 specimens), 55.9 mm SL (photo: Fahmi); (B) FRLM 31479, 57.6 mm SL (photo: S. Kimura); (C) ASIZP 62992, 60.5 mm SL (photo: Y. - C. Liao); (D) KAUM – I. 6876, male, 66.9 mm SL; (E) KAUM – I. 23882, male, 74.0 mm SL; (F) KAUM – I. 24053, female, 76.2 mm SL; (G) BSKU 42455, male, 84.0 mm SL (photo: BSKU); (H) KAUM – I. 47468, 94.5 mm SL.

FIGURE 14. Relationships of number of spines on median lateral ridge of maxilla to standard length in (A) Brachypterois serrulata and (B) B. serrulifer. H holotype of B. serrulifer. Note: all specimens examined, including type series, of B. curvispina sp. nov. and neotype of Sebastes serrulatus lacked spines.

FIGURE 15. Photographs of sections from gonads of Brachypterois serrulata. (A) ovary from KAUM – I. 24053, 76.2 mm SL; (B) testis from KAUM – I. 12313, 77.9 mm SL.

Tables

TABLE 3. Meristics and morphometrics, expressed as percentages of standard length, of Brachypterois serrulata.

	Neotype ASIZP 62399	Non-types from Non-types from Malaysia and north Thailand and of Vietnam Indonesia (n = 82 a) (n = 32)
SL (mm)	92.8	48.0–99.3 43.2–99.2
Meristics		Modes
Dorsal-fin rays	XIII, 11	XII–XIV, 9–11 XII, 9–11 XIII, 10
Anal-fin rays	III, 5	III, 4–6 III, 5–6 III, 5