Sebastes pachycephalus Temminck and Schlegel 1843

Kai, Yoshiaki & Nakabo, Tetsuji, 2013, Taxonomic review of the Sebastes pachycephalus complex (Scorpaeniformes: Scorpaenidae), Zootaxa 3637 (5), pp. 541-560 : 542-550

Sebastes pachycephalus Temminck and Schlegel 1843

[Japanese name: Murasoi]

( Figs. 1 , 2 A, 3A, 4; Tables 1–2)

Sebastes pachycephalus Temminck & Schlegel 1843: 47 , pl. XX-III (type locality: Nagasaki, Japan); Amaoka 1984: 312, pl. 278-E (in part, southern Hokkaido southward to Kyushu, Japan and southern Korean Peninsula); Suzuki & Kataoka 1997: 213, pl. 118-628 (Mie, Japan); Kim et al. 2004: 100 (in part, Hokkaido southward to Kyushu, Japan and southern Korean Peninsula; photo identifiable as S. nudus ); Kim et al. 2005a: 117, fig. 148 (in part, Hokkaido southward to Kyushu, Japan and southern Korean Peninsula); Kim et al. 2005b: 222 (in part, southern Hokkaido southward to Kyushu, Japan, off coasts of Korean Peninsula, and China; photo identifiable as S. nudus ).

Sebastes (Murasoius) pachycephalus , form. pachycephalus: Matsubara 1943: 239 , pl. II-1 (Chiba, Kanagawa, Aichi, Wakayama, and Kobe, Japan).

Sebastes (Murasoius) pachycephalus , form. nigricans: Matsubara 1943: 242 , pl. III-1 (Shimonoseki and Nagasaki, Japan).

Sebastes pachycephalus complex "Species P-Ni": Kai et al. 2011: 340 (Nagasaki, Shimane, Fukui, Ishikawa, Hiroshima, Wakayama, and Fukushima, Japan).

Sebastes pachycephalus pachycephalus: Matsubara 1955: 1077 , key, fig. 377 [after Matsubara (1943)] (Chiba southward to Kyushu, Japan, Busan, Korea, and China); Hiyama & Yasuda 1961: 112, pl. 148 (in part?, Hokkaido southward to Kyushu, Japan and southern Korea); Matsubara 1965: 426, unnumbered fig. [after Matsubara (1943)] (Aomori southward to Kyushu, Japan, Busan, Korea, and China); Kanayama & Kitagawa 1982: 42, unnumbered fig. (Iwate, Japan); Lindberg & Krasyukova 1987: 75, fig. 32 [after Matsubara (1943)] (Yokohama and Nagasaki, Japan and Busan, Korea); Nakabo 1993: 518, key (Chiba southward to Kyushu, Japan, Busan, Korea, and Yellow Sea); Masuda & Kobayashi 1994: 85, figs. 3–4 (Izu, Japan); Nakabo 1995: 177, color photo (Shimane, Japan); Motomura & Iwatsuki 1997: 130 (Nango, Miyazaki, Japan); Nakabo 2000: 595, key (Chiba southward to Kyushu, Japan, Busan, Korea, and Yellow Sea); Hirata et al. 2001: 48, color photo ( Kochi, Japan); Nakabo 2002a: 595, key (Chiba southward to Kyushu, Japan, Busan, Korea, and Yellow Sea); Kitagawa et al. 2008: 59, unnumbered fig. [after Kanayama & Kitagawa (1982)] (Pacific coast of Tohoku District, Northern Honshu, Japan).

Sebastes pachycephalus nigricans: Matsubara 1955: 1077 , key, fig. 378 [after Matsubara (1943)] (Shimonoseki and Nagasaki, Japan); Matsubara 1965: 427, unnumbered fig. [after Matsubara (1943)] (Shimonoseki and Nagasaki, Japan); Lindberg & Krasyukova 1987: 76, fig. 32 [after Schmidt (1931)] (Nagasaki and Misaki, Kanagawa, Japan); Nakabo 1993: 518, key (Shimonoseki and Nagasaki, Japan); Nakabo 2000: 594, key (Shimonoseki and Nagasaki, Japan); Nakabo 2002a: 595, key (Shimonoseki and Nagasaki, Japan).

Sebastes (Sebastodes) pachycephalus: Barsukov 2003: 200 , fig.86 [after Matsubara (1943)] (Nagasaki, Japan).

Sebastes (Murasoius) pachycephalus: Jin 2006: 143 [Dalian, Yantai, and Qingdao, China; figure after Li (1955) identifiable as S. nudus ].

Sebastichthys pachycephalus : Jordan and Starks 1904: (in part, Wakanoura, Kobe, Hiroshima and Shimonoseki, Japan); Tanaka 1930: 950, fig. 516 (Wakayama, Japan): Okada et al. 1935: 206, pl. 115-1 (in part, Hokkaido southward to Kyushu, Japan and Korean Peninsula); Okada & Matsubara 1938: 306 (in part, Hokkaido southward to Kyushu, Japan and Korean Peninsula).

Sebastichthys (Pteropodus) latus Matsubara 1934: 208 (type locality: Tokyo Bay, Japan).

Sebastodes nigricaus Schmidt 1930: 461 (type locality: Japan).

Sebastodes (Acumentum) nigricans Schmidt 1931: 97 , fig. 13 (type locality: Nagasaki, Japan).

Materials examined. 105 specimens, 24.9–318.0 mm SL (one specimen, FAKU 85940 was or used for counts and measurements). Japan: FAKU 3269, 3297, 131.0– 131.8 mm SL (2 specimens), Shimonoseki, Yamaguchi; FAKU 38149, 125.3 mm SL, Oki Is., Shimane; FAKU 41897, 109552–109553, 130095, 130127, 133026, 120.8– 146.2 mm SL (6), Maizuru, Kyoto; FAKU 50088, 217.1 mm SL, Choshi, Chiba; FAKU 50168, 50169, 131.4–148.0 mm SL (2), Yawatahama, Ehime; FAKU 50279–50283, 62.6–111.5 mm SL (5), Takahama, Fukui; FAKU 59217, 143.9 mm SL, Futo, Izu, Shizuoka, coll. T. Murai; FAKU 63147–63148, 129.6–181.0 mm SL (2), Kushimoto, Wakayama; FAKU 63366–63372, 87605, 98.3–184.0 mm SL (8), Susaki, Shimoda, Shizuoka; FAKU 64210– 64213, 83.7–88.0 mm SL (4), Shirahama, Wakayama; FAKU 64290, 106.2 mm SL, Koza, Wakayama; FAKU 65097, 154.8 mm SL, Susaki, Shimoda, Shizuoka; FAKU 69927, 124.7 mm SL, Okozu, Fukui, coll. R. Doiuchi; FAKU 73673, 124.0 mm SL, Ushimado, Okayama; FAKU 82393, 97.7 mm SL, Shimabara, Nagasaki, coll. K. Nakayama; FAKU 82408–82410, 146.1– 175.1 mm SL (3), Obama, Fukui, coll. R. Doiuchi; FAKU 82524, 82525, 84104–84105, 86958–86961, 96080–96081, 130008–130009, 93.6–169.4 mm SL (16), Ushitsu, Noto, Ishikawa, coll. Y. Kai and K. Sakai; FAKU 84876, 168.3 mm SL, Onahama, Fukushima, coll. T. Sato; FAKU 85918, 86.1 mm SL, Aki, Hiroshima, coll. K. Watanabe; FAKU 85937–85941, 87654–87657, 63.9–106.6 mm SL (9), Yatsuka, Shimane, T. Morihisa; FAKU 87650–87652, 77.4–129.0 mm SL (3), Tamano, Okayama, coll. K. Nozaki; FAKU 101806, S3690, 96.1–114.7 mm SL (2), Shigoshi, Tsushima Is., Nagasaki; FAKU 104349, 67.0 mm SL, Kominato, Chiba; FAKU 133308, 274.8 mm SL, Kobe, Hyogo, coll. K. Matsubara; FAKU 133309 (holotype of S. latus ), 138.1 mm SL, Tokyo Bay, coll. S. Masuda; FAKU 133431–133432, 166.8– 169.8 mm SL (2), Miyako, Iwate, coll. Y. Kai; FAKU 134304–134305, 166.8–318.0 mm (2), Onahama, Fukushima, coll. T. Wada; KAUM–I. 3845, 40.6 mm SL, Ibusuki, Kagoshima Bay, Kagoshima, coll. KAUM Fish Team; KAUM–I. 9257, 164.7 mm SL, Sanwa, Kagoshima Bay, Kagoshima, coll. T. Ojiro; MUFS 12606–12608, 67.0– 69.4 mm SL (3), Nango, Miyazaki; NMCI–P. 1530–1532, 125.0– 145.2 mm SL (3), Ushitsu, Noto, Ishikawa; OMNH–P 3108, 68.3 mm SL, Kasumi, Hyogo, coll. Kasumi High School; OMNH–P 7929, 13607, 60.2–71.0 mm SL (2), Takeno, Hyogo; OMNH–P 9214, 37.3 mm SL, Muroto, Kochi, coll. K. Hatooka; OMNH–P 9434, 17402, 41.2–78.2 mm SL (2), Matsusaka, Mie; OMNH–P 9735, 91.7 mm SL, Kamijima, Toba, Mie, coll. K. Hatooka; OMNH–P 13415, 33.2 mm SL, Kunimi, Nagasaki, coll. K. Hatooka; OMNH–P 19307, 38367, 24.9–69.0 mm SL (2), Misaki, Osaka, coll. K. Hatooka; OMNH–P 30495, 78.3 mm SL, Kada, Wakayama, coll. K. Hatooka; RMNH D566 (lectotype of S. pachycephalus ), 261.8 mm SL, Japan (detailed collection site not given), coll. H. Bürger; ZIN 22686 (holotype of S. nigricans ), 150.0 mm SL, Nagasaki, coll. P. J. Schmidt. South Korea: FAKU 60725, 155.5 mm SL, Seogwipo, Cheju Is. Locality unknown: FAKU 13–15, 103482, 187.7– 275.2 mm SL (4).

Diagnosis. A species of Sebastes with the following characters: cranium armed dorsally with robust preocular, supraocular, postocular, and parietal spines; interorbital space concave; lower jaw without scales, shorter than upper jaw; base of entire spinous portion of dorsal fin densely covered with minute scales; body with three dark brown saddles and lacking yellow or brownish-red markings on dorsum when fresh; soft-rayed portion of dorsal and anal fins, and caudal fin with dark spots.

Description. Measurements as percentages of SL of the specimens examined are given in Table 1. Selected counts are given in Table 2 .

Body relatively deep, moderately compressed anteriorly, progressively more compressed posteriorly. Nape and anterior body moderately convex. Tentacles absent on head and body, except for nostril tentacle. Head completely covered with ctenoid scales, except for tip of snout, maxillary, lacrimal, lower jaw, interopercle, and branchiostegal rays. Posterior part of maxillary sometimes with minute embedded scales. Body covered with ctenoid scales usually with some accessory scales in posterior field, except for pectoral fin base, prepelvic region and ventral abdominal surface. Embedded cycloid scales covering pectoral fin base and prepelvic region; remaining ventral abdominal surface completely covered with minute cycloid scales. Base of entire spinous portion of dorsal fin densely covered with minute scales, usually extending onto basal spines and membranes ( Fig. 2 A); anterior part of spinous portion of base sometimes with very narrow naked area in smaller specimens (under ca. 60 mm SL). Patches of minute scales sometimes on membrane of spinous dorsal fin. Bases of soft-rayed portion of dorsal and anal fins covered with minute scales, extending onto basal membranes.

Mouth large, slightly oblique; posterior margin of maxilla reaching or nearly reaching (reaching in lectotype) level with posterior margin of orbit. Maxilla covered with thick skin. Lower jaw shorter than upper jaw, without distinct symphysial knob. Upper and lower jaws with band of villiform teeth. Palatines and vomer with villiform teeth, latter forming V-shaped patch.

Nasal spine simple, sharp, directed dorsally. Preocular spine robust, well developed, directed dorsoposteriorly; tip of spine reaching or extending slightly beyond level with anterior margin of pupil (ocular prosthesis in lectotype). Supraocular spine robust and well developed, directed posteriorly; tip of spine reaching or extending beyond (reaching in lectotype) level of posterior margin of orbit. Postocular spines simple, directed dorsoposteriorly. Interorbital space concave without ridge, its width about half of orbit diameter. Parietal spine well developed, somewhat divergent posteriorly in dorsal view. Supracleithral spine simple, directed posteriorly. Upper posttemporal spines flattened, with embedded base, lower posttemporal spine absent. Sphenotic, tympanic, and pterotic spines absent. Lacrimal with round lobe anteriorly and one spine posteriorly; spine flattened, blunt, directed ventroposteriorly, its tip below level of anterior margin of pupil. Suborbital without spine or ridge. Preopercle with five spines; two uppermost spines large, directed posteriorly; third moderate, directed posteriorly; forth and fifth small, blunt, directed ventroposteriorly. Opercle with two simple flattened spines directed posteriorly; upper spine somewhat larger than lower spine.

Dorsal fin with 13 spines (rarely 12 or 14; 13 in lectotype) and 10–13 (usually 12 including lectotype) soft rays; all soft rays branched (first ray weakly branched). Dorsal-fin origin above anterodorsal portion of gill slit. Dorsal fin gradually increasing in height to fifth spine, decreasing thereafter to 12th spine; 13th spine usually much larger than 12th, forming anterior support of soft-rayed portion of dorsal fin. Soft-rayed portion of dorsal fin with entire margin rounded; anterior rays longer (second or third ray longest, third in lectotype), posterior rays gradually shortening. Anal fin with three spines and six (rarely five or seven, six in lectotype) soft rays; all soft rays branched. First analfin spine slightly posterior to last dorsal-fin spine; second spine robust, longest. Anterior rays on soft-rayed portion of anal fin longest; posterior margin rounded. Posterior margin of caudal fin rounded. Pectoral fin rounded; its tip not reaching posteriorly to level with anus; ventral 10–13 (usually 11, 12 in lectotype) rays unbranched and thickened. Posterior tip of depressed pelvic fin below tip of pectoral fin, not or rarely reaching anus.

Gill rakers short, blunt, longest raker in joint between cerato- and hypobranchial, length of preceding and succeeding rakers progressively shorter; ceratobranchial rakers six–eight (usually seven), hypobranchial rakers 14– 17, often rudimentary and united, being difficult to distinguish individually (gill arches removed in lectotype).

Color when fresh ( Fig. 1 ). Head and body brown or gray dorsally and laterally, somewhat paler ventrally. Head with three indistinct dark brown bands radiating from eye. Body with three dark brown saddles, two anteriormost extending onto dorsal fin; anteriormost positioned under fifth to 11th dorsal-fin spines, second under soft-rayed portion of dorsal fin, posteriormost on caudal peduncle. Underside of head, pectoral fin base, and ventral abdominal surface with ( Figs. 1 A, B) or without dark brown spots ( Figs. 1 C–F). Dorsal fin brown or gray; dark brown spots present or absent on spinous portion, but usually present on soft-rayed portion. Anal fin brown or gray with dark brown spot on soft-rayed portion. Caudal fin brown with darker brown spots; membranes darker. Pectoral fin dark brown or grey, usually with dark spots basally; ventral half somewhat paler. Pelvic fin brown, membranes darker. In smaller specimens (under ca. 10 cm SL), dorsal, anal, caudal, and pelvic fins somewhat reddish ( Fig. 1 F).

a Lectotype of S. pachycephalus ; b Holotype of S. latus ; c Holotype of S. nigricans ; d Holotype of S. nudus ; e Holotype of S. chalcogrammus .

Color in preserved specimens ( Fig. 2 A). Head and body dark dorsally and laterally, somewhat paler ventrally. Head with three indistinct dark bands radiating from eye. Body with three dark saddles as when fresh. Underside of head, pectoral fin base, and ventral abdominal surface with or without dark brown spots. Soft-rayed portion of dorsal and anal fins, and caudal fin with dark spots.

Distribution ( Fig. 3 A). Sebastes pachycephalus is presently known from Aomori, northern Honshu Is. southward to Kagoshima Bay, southern Kyushu Is., Japan, in addition to the southern part of the Korean Peninsula, and the Yellow and Pohai Sea coasts of China. This species is solitary, inhabiting rocky reefs in shallow coastal waters.

Remarks. The present description of S. pachycephalus is based on the specimens genetically identified as “Species P-Ni” sensu Kai et al. (2011), plus additional specimens not genetically examined here, but considered as conspecific with the former due to their having minute scales below the entire dorsal-fin spine base and on the caudal fin, and lacking yellow or brownish-red markings on the dorsum.

Sebastes pachycephalus was originally described by Temminck & Schlegel (1843), based on specimens collected by H. Bürger and Ph. F. von Siebold. Later, Boeseman (1947), who reviewed the Japanese fishes collected by Bürger and von Siebold, considered four stuffed specimens (RMNH D563, D564, D565, D566) and two specimens in alcohol (RMNH 654, 655) as the type series of S. pachycephalus , designating one of them (RMNH D566) as the lectotype. The latter (RMNH D566: Fig. 4 A) had 19 pectoral-fin rays, minute scales below the entire dorsal-fin spine base, and dark spots on the belly, anal, caudal, and pelvic fins, being characteristic of specimens identified as "Species P-Ni" sensu Kai et al. (2011). Although the remaining five specimens are paralectotype of S. pachycephalus, Boeseman (1947) noted that two (RMNH 654, 655) had been identified as “ Sebasichthys elegans ” sensu Jordan & Starks (1904) owing to their having 14 dorsal-fin spines, subsequently included under Sebastes hubbsi Matsubara 1937 [see Matsubara (1936) and Matsubara (1937)]. The identity of those specimens, not examined during the present study, remains to be confirmed.

Schmidt (1930) reported Sebastodes nigricaus as a new species from Japan (detailed locality not given), providing a brief description but without designating a holotype. That species was characterized as having: D XIII, 13; A III, 7; pored lateral line scales 32; maxillary without scales; blackish coloration. Later, Schmidt (1931) described, Sebastodes (Acutomentum) nigricans on the basis of a single specimen (ZIN 22686), collected from Nagasaki, Japan. Many authors have overlooked the former name, citing only S. nigricans of Schmidt (1931) in reviews of Sebastes (e.g., Matsubara 1943; Barsukov 2003). However, Eschmeyer (2012) considered that the description of Schmidt (1930) included a definition of the species, “ nigricaus ” being a misspelling of “ nigricans ”, and thus regarded “ Sebastodes nigricans Schmidt 1930 ” as an available name. We agree that Schmidt’s (1930) description included a valid definition of the species following ICZN (1999: Art. 12). However, “ nigricaus ” is the correct original spelling following ICZN (1999: Art. 32). Although Schmidt (1930) may have misspelt his intended original name for the species, we here regard “ Sebastodes nigricaus Schmidt 1930 ” as an available name. Schmidt (1930) did not note any museum registration number for his specimen, but the counts given for dorsal- and anal-fin rays and lateral line scales were identical with those given later (Schmidt 1931), indicating that the two descriptions had been based upon the same specimen (ZIN 22686). This specimen, the holotype of S. nigricans ( Fig. 4 B), has 18 pectoral-fin rays (lower 11 rays unbranched) and minute scales below the entire dorsal-fin spine base, agreeing closely with the present specimens regarded as S. pachycephalus . Therefore, we concluded that both S. nigricaus and S. nigricans are synonyms of S. pachycephalus .

The description of Sebastichthys (Pteropodus) latus by Matsubara (1943) was based on a specimen collected from Tokyo Bay, Japan. Although the holotype of that species was believed lost (not at FAKU) (Eschmeyer 2012), many of Matsubara’s specimens are in fact housed at FAKU, including the holotype of Sebastes melanostictus and syntypes of Sebastes longispinis , both species having been described as new in the same paper as S. latus . Recently, a specimen (FAKU 133309; Fig. 4 C) bearing two of Matsubara’s collection numbers (577 and 651) was found that corresponded to the length and preservation condition of the type specimen of S. latus , as originally described by Matsubara (1934). According to the original description, the holotype of S. latus was 174 mm in total length and characterized by 19 pectoral-fin rays (lower 12 unbranched), a deep body (44.2% of SL), 24 gill rakers, and many dark spots on fins and underside of head. FAKU 133309 (172.5 mm in total length) had 19 pectoral-fin rays (lower 12 unbranched), deep body (42.2% of SL), and 24 gill rakers (see Tables 1–2, Fig. 4 C), completely matched the original description. In addition, Matsubara (1934) noted “frontal ridges not visible without dissection”; FAKU 133309 had been dissected around the frontal ridges. Therefore, we consider that the original description of S. latus was based on FAKU 133309, being the holotype. However, this species now is believed to be an example of S. pachycephalus , owing to the minute scales below the entire dorsal-fin spine base, and many dark spots on the fins and underside of the head and body. Thus, S. latus can be confirmed here as a junior synonym of S. pachycephalus . In fact, Matsubara (1943) had already synonymized S. latus under S. p. pachycephalus , because the diagnostic characters given in Matsubara (1934) fell within the range of the latter.

Jordan & Starks (1904) described Sebastichthys pachycephalus on the basis of specimens collected from Wakayama, Kobe (Seto Inland Sea), and Shimonoseki, Japan, referring to their color variations as follows: "golden-yellow blotches and sometimes having dark spots on breast and base of pectoral fin." “Golden-yellow blotches” is consistent with the present specimens of S. nudus , whereas “dark spots on breast and base of pectoral fin” is consistent with those of S. pachycephalus . Similarly, the description of Sebastichthys pachycephalus given by Okada et al. (1935) includes “two variations have been known; one with dark spots on the ventral body surface and the fins, and the other without such spots but with yellow or reddish-yellow markings on the dorsum.” These are consistent with presently-recognized S. pachycephalus and S. nudus respectively. However, the supporting plate matched the former in having dark spots on the ventral surface of body and the dorsal, anal and caudal fins. Okada & Matsubara (1938) also described S. pachycephalus as having variations in body coloration, following Okada et al. (1935). Thus, we consider that both species recognized here were included in their descriptions.

Matsubara (1943) recognized four forms in his Sebastes (Murasoius) pachycephalus ; viz. “ S. pachycephalus , form. pachycephalus ”, “ S. pachycephalus , form. nigricans ”, “ S. pachycephalus , form. nudus ”, and “ S. pachycephalus , form. chalcogrammus ”, the latter two being described as new. The subsequent recognition of these “forms” as subspecies by numerous authors (see below) is supported by ICZN (1999: Art. 45.6). Sebastes p. pachycephalus , based on 30 specimens (no catalog number given) collected from Chiba, Kanagawa, Aichi, Wakayama (Pacific coast of Japan) and the Seto Inland Sea, Japan, was characterized as having minute scales below the entire dorsal-fin spine base and many dark spots on the fins and underside of the head. Such characters are consistent with the present specimens regarded as S. pachycephalus . Matsubara’s (1943) description of S. p. nigricans , based on two specimens (no catalog number given), included “spinous dorsal covered by minute scales behind the second dorsal spine.” Although all of the larger specimens examined here had minute scales below the entire dorsal-fin spine base, S. p. nigricans of Matsubara (1943) was most likely to have been S. pachycephalus . Matsubara (1943) also considered S. p. pachycephalus to be distinguishable from S. p. nigricans in having “black spots on belly, breast, and underside of head, usually 19 pectoral-fin rays (17 or 18 in the latter), maxillary without scales, and accessory scales on body.” However, the presence or absence of black spots as described have been recently demonstrated as representing intraspecific variation in S. pachycephalus , based on genetic analysis (Kai et al. 2011: as "Species P-Ni"). Furthermore, the pectoral-fin ray counts fell within the present range established for S. pachycephalus , the latter two characters in fact being subject to significant individual variation in the present specimens. A discussion of Matsubara’s (1943) S. p. nudus and S. p. chalcogrammus is given under "Remarks" for S. nudus . In later papers, Matsubara (1955, 1965) similarly recognized four subspecies (not forms) under S. pachycephalus ( S. p. pachycephalus , S. p. nigricans , S. p. nudus , and S. p. chalcogrammus ), following Matsubara (1943). All of the figures included in Matsubara (1955, 1965) were modified from those in Matsubara (1943).

Hiyama & Yasuda (1961) redescribed S. pachycephalus pachycephalus , commenting that the four forms were defined by differences in body coloration, thereby apparently following Matsubara (1943). It is unclear whether or not they considered the “four forms” to be included under their S. p. pachycephalus . However, the accompanying figure (pl. 148) appears to be of S. pachycephalus , owing to the dark spots on the ventral abdominal surface and dorsal, anal and caudal fins.

The status of S. pachycephalus subspecies was not addressed by Amaoka (1984), who considered S. pachycephalus simply as a single species, owing to the existence of equivocal specimens between subspecies. However, S. pachycephalus of Amaoka (1984) clearly included both S. pachycephalus and S. nudus as recognized here, one of the color photos (pl. 278-E) given by him being referable to the former in having dark spots on the dorsal, anal, and caudal fins, and two (pl. 278-F and G) to the latter in having distinct yellow or reddish markings on the dorsum.

Nakabo’s (1993, 2000, 2002a) descriptions of four subspecies, S. pachycephalus pachcephalus , S. p. nigricans , S. p. nudus , and S. p. chalcogrammus , generally followed Matsubara (1943), that of S. p. pachycephalus having been based on FAKU 40153 (unavailable for the present study) and that of S. p. nigricans , on FAKU 3297 and FAKU 109552. The latter two specimens were herein re-identified as S. pachycephalus . In addition, the diagnoses of S. p. pachycephalus proposed by Nakabo matched the specimens presently regarded as S. pachycephalus . Sebastes pachycephalus nudus and S. p. chalcogrammus of Nakabo (1993, 2000, 2002a) were both identified here as S. nudus (see "Remarks" under S. nudus ).

Lindberg & Krasyukova’s (1987) description of S. pachycephalus nigricans apparently followed the original description of S. nigricans given by Schmidt (1931). Their description of S. p. pachycephalus apparently followed that of Matsubara (1943).

In his description of Sebastes (Sebastodes) pachycephalus, Barsukov (2003) provided only pectoral-fin ray counts (usually 19) and a figure modified from that of S. p. pachycephalus of Matsubara (1943). Judging from the counts, it is most likely to have been S. pachycephalus . Barsukov (2003) also described S. (Sebastodes) nigricans as a valid species, but provided no useful diagnostic characters.

Because S. pachycephalus is common in shallow coastal waters around Japan and Korea, many records of the species exist in the literature. Tanaka (1930) reported Sebastichthys pachycephalus from Wakayama, Japan, characterized as having minute scales below the dorsal-fin spines, his supporting figure having dark spots on the dorsal, anal, caudal, and upper pectoral fins, and anterior ventral abdominal surface. Such characters agreed well with those of the present specimens regarded as S. pachycephalus . The photos of S. pachycephalus pachycephalus in Kanayama & Kitagawa (1982), Masuda & Kobayashi (1994), Nakabo (1995), Suzuki & Kataoka (1997), Hirata et al. (2001), and Kitagawa et al. (2008) are all clearly of S. pachycephalus , owing to the dark spots on the softrayed portion of the dorsal fin and caudal fin. The photo of S. pachycephalus in Kim et al. (2005a) also matched presently recognized S. pachycephalus in having dark spots on the caudal fin. Our re-examination of Motomura & Iwatsuki’s (1997) specimens of S. p. pachycephalus (MUFS 12606–12608; Miyazaki, Japan) also confirmed them as S. pachycephalus . Although the description of S. (Murasoius) pachycephalus from the Pohai and Yellow Seas given by Jin (2006) was identifiable as S. pachycephalus , as recognized here, on the basis of minute scales below the entire dorsal-fin spine base and black spots on the ventral abdominal surface, as well as on the dorsal and caudal fins, the accompanying figure was apparently modified from that of Chang et al. (1955), herein identified as S. nudus (see Remarks under S. nudus ).

Many authors, including Jordan & Starks (1904), Tanaka (1930), and Matsubara (1943), have cited Richardson’s (1854) report of Sebastes pachycephalus from China (no specific locality record given). However, “several large, pale or bluish round spots” on the body sides in Richardson’s description do not occur in either S. pachycephalus or S. nudus , as recognized here. Cheng’s (1997) S. nigricans was characterized as having reddish markings above the opercle and between the soft-rayed portions of the dorsal and anal fins, and vertical stripes on the caudal fin. Such characters agree rather more with Sebastes longispinis (Matsubara 1934) (see Nakabo 2002a). Sebastes pachycephalus of Steindachner & Döderlein (1884) is likely to have been S. hubbsi , owing to the dorsalfin spine count (14).

The following reports cannot now be attributed with certainty to either of their two species of Sebastes recognized here because of the lack of definitive characters: Jordan & Snyder (1900) as “ Sebastodes pachycephalus ” (Misaki, Kanagawa, Japan), Jordan & Snyder (1901) as “ Sebastodes pachycephalus ” (Yokohama, Japan), Jordan & Metz (1913) as “ Sebastichthys pachycephalus ” (Busan, Korea), Jordan et al. (1913) as “ Sebasichthys pachycephalus ” (Wakayama southward to Nagasaki, Japan), Jordan & Thompson (1914) as “ Sebastichthys pachycephalus ” (Misaki, Kanagawa, Japan), Jordan & Hubbs (1925) as “ Sebastichthys pachycephalus ” (Otaru of Hokkaido, Yokohama, and Toyama, Japan), Ueno (1971) as “ Sebastes pachycephalus ” (Hokkaido and northern Honshu Is., Japan), Shiogaki et al. (2004) as “ Sebastes pachycephalus pachycephalus , S. p. chalcogrammus , S. p. nudus ” (Aomori, Japan), Shinohara & Williams (2006) as “ Sebastes pachycephalus ” (Sagami Sea, Japan).

Figures

FIGURE 1. Sebastes pachycephalus in life (A) and when fresh (B – F). (A) FAKU 130127, 129.6 mm SL, Maizuru, Kyoto, Japan; (B) FAKU 73673, 124.0 mm SL, Ushimado, Okayama, Japan; (C) FAKU 133431, 169.8 mm SL, Miyako, Iwate, Japan; (D) FAKU 133432, 166.8 mm SL, Miyako, Iwate, Japan; (E) FAKU 84110, 141.9 mm SL, Noto, Ishikawa, Japan; (F) OMNH- P 9214, 37.3 mm SL, Muroto, Kochi, Japan. A – E photograph by Y. Kai, F photograph by K. Hatooka (OMNH).

FIGURE 2. Schematic illustrations of Sebastes pachycephalus (A) and S. nudus (B), indicating the squamation pattern below the dorsal-fin spines. (A) FAKU 65097, 154.8 mm SL, Susaki, Shimoda, Shizuoka, Japan; (B) FAKU 84093, 158.3 mm SL, Miyako, Iwate, Japan.

FIGURE 3. Distribution of Sebastes pachycephalus (A) and S. nudus (B). Closed circles based on specimens examined in this study; open circles based on literature records.

FIGURE 4. Type specimens of nominal species related to Sebastes pachycephalus. (A) Holotype of S. pachycephalus, RMNH- D 566, 261.8 mm SL; (B) holotype of S. nigricans, ZIN 22686, 150.0 mm SL; (C) holotype of S. latus, FAKU 133309, 138.1 mm SL.

Tables

TABLE 2. Frequency distribution of selected counts from specimens of Sebastes pachycephalus and S. nudus.

	Pectoral-fin rays (left side)			Pectoral-fin rays (right side)
	16 17 18 19	20		16 17 18 19	20
S. pachycephalus	1 18 64 17a, b	2		- 23 60c 18b	2
S. nudus	9 67d 12e -	-		6 70d, e 9 -	-
	Pored lateral-line scales
	27 28 29 30	31	32	33 34 35
S. pachycephalus	1 1 3 15	39c	24b	13 5 2
S. nudus	1 1 9 13	23	26d, e	9 1 3