JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology
Species Diversity, 2009, 14, 97-114
Validity of the Cottid Species Steigistrum mororane
Transferred to the Genus ICelus (Actinopterygii: Perciformes: Cottoidei), with Confirmed Records
of Steigistrum stojnageri from Japanese Waters
Osamu Tsuruokai, Takuzo Abe2 and Mamoru Yabe3
'Laboratot)) ofMttrine Biodiversily (Systematic ichtbyologJV, Graduate School of Fisheries Sciences, Ilbkicaido [Jniversity, Hctkodate, 041-Bbl1 .Jtipan Present address: IIOkkaido liakodate Fisheries High School,
2-l5-3 IVanaehama, HOkuto, 0490111 ,Jiripan E-mait: tsuru357@/gmaiLcom 2Shizugawa Alature Center, 40 Sahamoto, Minamtsanriku-cho 7bgura,
MOtqyoshi-.aun, Miyagi, 986-0781 .lapan Present address: fuculty ofFisheries Science, Hbkkaldo [lhiversity, Hakodate, 041-8611 Japan E-mail; takuzo@oshorokenJish,hohudai.ac.jp '?Laboratory ofMlxrine Biology and Biodiversity twstematic Ibhtbyology), Research Riculty ofFisheries Sciences, Hbkicaido Universits,, Htxkodate, 041-8611 Jqpan E-mail; mo,abeig2f}sh.hokudai.ac.tp
(Received 25 July 2008; Accepted 26 December 2e08)
Stetgistrum mororane Jordan and Seale, 1906, previously synon]rmized with SteCgistrum stojnegeri Jordan and Gilbert, 1898, is here redescribed as a valid species of the genus icelus. Contrary to the original description, the holotype of S, mororane has palatine teeth, which are diagnostic ef Jbeltts but absent in Steigistrum. icegus mororants comb. nov, is distinguished from al1 other species of icetus by the fo11owing combination of characters: body rela- tively short; parietal spine undeveloped; subocular and supraocular spines absent; scales absent on cheek, between dorsal scale row and lateral Iine, and below lateral line except at pectoral axMa; small and simple cirrus pres- ent on upper rim of eye; two pairs of nuchal cirri present; and four saddles present on dorsolateral surface of body, Additionally, seven specimens of Steegistrum stojnageri co}lected from the Pacific coast of eastern Hokkaido are described, representing the first confirmed records of this species in
Japanese waters. Key Words: Redescription, leelus mororanis comb. nov,, Steigistrum stoj-
negeri, confirmed record, Japan.
Introduction
The cottid genus Stelgistrunz Jerdan and Gilbert, 1898 is characterized by hav- ing four preopercular spines, teeth present on the jaws and vomer but absent on the palatines, one scale row on the lateral line and another scale row along the dor-
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98 Osamu Tsuruoka et aL
sal-fin base, a single pharyngeal tooth plate, no urogenital papilla, and a pelvic fin with one spine and three soft rays (Andriashev 1935>. Three species are currently recognized as valid in this genus: S, stojnageri Jordan and Gilbert, 1898, S. beringianum Gilbert and Burke, 1912, and S. concinnum Andriashev, 1935, All are distributed in the northern North Pacific, Bering Sea, Sea of Okhotsk, and north-
ern Sea of Japan.
In addition to those three species, Steigistrum mororane Jordan and Seale, 1906, was described on the basis of a single specimen collected from off Muroran "Mororan" (spelled in the original description), Volcano Bay, Hokkaido, Japan. Jordan et at. (1913), however, regarded S. mororane as a junior synonym of S. stoj- negeri without discussion. Andriashev (1935) indicated that S, rnororane differed from S. stojnageri jn the counts of anal-fin rays and the existence of a granulation (dermal cirrus) on the dorsal surface of the head, but he nevertheless supported the opinion of Jordan et al. (1913). Other authors followed these decisions and con- sidered S. mororane as a junior synonym of S. stojnegeri (e.g., Watanabe 1958, 1960, 1978; Lindberg and Krasyukova 1987), Recently, a large number of small cottid specimens were celleeted from the shallow marine waters of northern Japan, including near Muroran, the type local- ity of S. mororane. These specimens greatly resemble S. mororane or S. stojnegeri, but clearly belong to the genus lbelus Kreyer, 1845 on account of their having teeth on the palatines (Nelson 1984; Yabe 1985). Jordan and Seale (1906) originally de- scribed S. mororane as having ne teeth on the palatines, but our reexamination of the holotype confirmed that S. mororane does have some teeth there, Therefore, S, mororane must be placed in the genus Ibelus and is distinct from S. stojnageri. In this study, we redescribe icelus mororants on the basis of 103 specimens in- cluding the holotype and additional non-type specimens, Additionally, we re- describe SteLgistrum stojnegeri based on seven specimens newly collected from eastern Hokkaido, Japan, because the previous records of this species from Japan- ese waters are imprecise and unconfirmable,
Materials and Methods
The methods used for taking counts and measurements fbllow Hubbs and La- gler (1958) and Yabe (1991) except that interdorsal width was measured between the base of the last spine of the first dorsal fin and the origin of the second dorsal fin. Total length (TL), standard length (SL), and head length (HL) are used throughout unless indicated otherwise. All fin-ray elements were counted. Vertebrae and cau- dal-fin rays were observed and counted from radiographs. Sex was determjned by observation of the genads, The higher classification of the family Cottidae follows that presented by Imamura and Yabe (2002). Symbolic codes for institutions are those provided by Esehmeyer (1998). All materials examined in this study are given in the appendix,
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Validity of Steigistrum mororane in Ibelus 99
Taxonomy
Ibelus mororanis (Jordan and Seale, 1906) comb. nev. [New Japanese name: Usujiri-kajika] (Figs 1, 2, Tables 1, 2)
SteLgistrum mororane Jordan and Seale, 1906: 146, fig. 4 [original description, holo- type, sex unknown (31,8mm TL), in harbor at Mororan (=Muroran), Hokkaido, Japanl; BOhlke 1953: 132 (listed, type specimen in SU); Eschmeyer 1998: 1122 (validity), Steigtstrum nzororanae [sic]: Andriashev 1935: 296-297 (description, distribution, synonymy). SteLgistrunz stojnegeri (not Jordan and Gilbert, 1898): Jordan et at. 1913: 258, fig. 195 (in part, listed, Japan, distribution); Watanabe 1958: 14, 227-230, figs 77, 78, pls 5 (lower), 56 (in part, listed, Japan, diagnosis, description of internal features); 1960: 32-35, fig. 11, pl. 20 (fig, 2) (in part, description); 1976: 6 (in part, listed, Japan); 1978i 230-232, figs 77, 78, pl. 19 (seeond from top) (in part, description); 1980: 427-428, pl, 48 (description of osteologieal features); 1981: 106 (in part, de- scription of quantitative features); Lindberg and Krasyukova 1987: 195-196 (in part?, diagnosis, distribution).
Material examined. 103 specimens (24,O-44.7mm SL) including holotype of Stetgistrum mororane (see Appendix). Diagnosis. A species ef the genus lbelus having the fo11owing combination of characters: subocular, supraocular, parietal, and nuchal spines absent; uppermost preopercular spine unbranched; single dorsal scale row; seales of lateral line and dorsal rows with serrated margins; no scales on cheek, between dorsal scale row and lateral line, nor below lateral line except for pectoral axilla, where small scales sometimes present; small, unbranched cirrus on upper rim of eye; one pair each of supraocular and parietal cirri, and two pairs of nuchal cirri; four saddles on dorso- lateral suiface of body; soft rays of second dorsal fin 1417; anal-fin rays 1or13; pec-
toral-fin rays 15-16; vertebrae 12+23-25=35B7. Description. Proportienal measurements and counts are given in Tables 1 and 2. In the following text, data for the holotype are shown first, followed by data
for non-type specimens in parentheses. Body short, compressed (Fig, 1), body depth at origin ef first dorsal fin 1,25 (1.00-1.67) times body width. Head (Fig, 2) large, slightly compressed, length 41,7% (37.0-43.50r'o) of SL, depth at base of uppermost preopercular spine 1,25 (O.83-1.25) times head width, Caudal peduncle slender, compressed, depth 27,O% (23.8-37.0%) of its Iength, Upper profile of snout steep, length 25,69･6 (23,3-31.3[%) of HL. Nasal spine short, sharp, directed posterodorsally. Anterior nostril on outer base of nasal spine, forming short tube. Posterior nostril on anterior margin of orbit, with low rim. Mouth Iarge; lewer jaw extending ibward slightly beyond upper jaw. Maxilla extending to level of posterior margin of orbit. Small conical teeth on jaws, vomer, and palatines (palatines partly broken in holotype). Eye large, length of orbit 29.4[lh (25.0-34.5Vo> of HL, Interorbital space narrow, concave, width 25.69/k (20,8-37,O%) of orbit length. Occipital region slightly concave, without knob or ridge. Four preop- ercular spines: uppermost spine unbranched, sharp, hooked posterodorsally; sec-
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100 Osamu Tsuruoka et al.
Fig. 1. icelets mororanis (Jordan and Seale, 1906) comb. nov., HUMZ 189781, male, 37.8mm SL, colleeted from off Usujiri, Hakodate.
ond spine slightly blunted, directed posteriorly; third spine also slightiy blunted, directed ventrally; lowermost spine smal1, direeted anteroventrally, buried under skin. Numerous small scales scattered around orbit and on base of nasal spine, upper rim of eye, and dorsolateral surface of head above suborbital stay, Both jaws and ventral surface of head naked, Lateral line descending posteriorly in slight arch from upper border of supracleithrum to level of seventh or eighth (broken in holotype) ray of anal fin, then running directly posteriad along body axis, Scales on lateral 1ine large, tubular, with serrated margins dorsally and posteriorly. Mid- sized scales with serrated dorsal margins arranged along dorsal-fin base, to form dorsal scale row originating beneath base of second spine of first dorsal fin and ex- tending to caudal peduncle. Small scales with serrated margins scattered above dorsal scale row and on spines and soft rays ef dorsal fin. No scales between dersal scale row and lateral line, nor below lateral line. No (no or some small) ctenoid scales scattered in pectoral axilla. Belly naked. Four pairs of cirri on dorsal sur- face of head: pair of supraocular cirri developed as narrowly based derTnal fiaps bearing numerous short appendages along distal margin; pair of parietal cirri with short appendages along distal margins; and two pairs of small, unbranched nuchal cirri (Fig. 2). No cirri on base of nasal spine, One simple cirrus on dorsal margin of eye, One (one or two) siniple cirrus on posterier end of maxilla. Two cirri on subor- bital stay and posterior region of opercle, i.e., small cirrus with branched tip and small, simple cirrus, respectively. One or two small, simple cirri on base of each preopercular spine. Single cirrus present on several lateral-line scales. First dorsal fin originating at level of posterior margin of epercular flap. First and second dorsal fins joined by low fin-membrane, Basal length of first dorsal fin 66.7% (52,6-83.3%) of that of second dorsal fin. Second dorsal fin eriginating above 13th (11th-13th) lateraiL-line scale. Anal fin originating at level of origin of second dorsal fin, its basal length 83.3% (58.8-83.3%) of that of second dorsal fin. Pectoral fin large, length 22.7-34.5`k of SL NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Validity of Steigistrum mororane in Jtetus 101 Fig. 2. Dorsal view of head of Jeelus mororanis (Jordan and Seale 1906) comb. nov., HUMZ 189781. Scale bar 5mm. fourth or fifth ray of second dorsal fin (broken in holotype); lower six (six or seven) rays slightly thickened, Pelvic fin slender, originating beneath second spine of first dorsal fin; middle ray longest; spine and outermost soft ray closely united. Caudal fin slightly rounded posteriorly; 11 rays supported by hypural plates, uppermost and lowermost two rays unbranched, middle seven rays branched; five (five to eight) upper and fbur (four to seven) lower procurrent rays. Anus located below eighth spine of first dorsal fin. Males laeking urogenital papilla. Color in alcohol. Body light brown (light brown or grayish brown), with four dark saddles dorsolaterally; area below lateral line, except for pectoral axMa and belly, light brown (light brown or grayish brown) with numerous pale spots. Head darker than body; occipital and suborbital regions especially dark. First dorsal fin transparent with large dark blotch on posterior margin. Second dorsal fin trans- parent with obscure oblique dark stripes. Pectoral fin translucent with triangular dark blotch on base, some small dark blotches on each ray. Pelvic fin pale with slightly darker base. Anal fin transparent with dark stripes. Caudal fin pale with some dark lateral stripes. Color when fresh (based on non-type materials). Ground color of body cream- white, grayish brown, or reddish brown; pectoral axilla and ventral suiface of body whitish. Four dark brown saddles across back: one below middle part of first dorsal fin, two below second dorsal fin, and one on caudal peduncle. Head below suborbital deep brewn, Some vertical brown stripes on upper jaw. Ground color of first dorsal fin transparent, bearing brown vermiculations; black blotch on poste- rior margin. Second dorsal fin transparent, bearing obscure oblique brown stripes, Black triangular blotch on base of pectoral fin. Pelvic fin white, with slightly darker spots on base. Anal fin transparent, with two or three dark brown horizon- tal stripes (discolored in holotype). Caudal fin transparent, with reddish-brown lateral stripes. Cirri darkly pigrnented except for those on lateral-line scales, Distribution. Off Muroran, Volcano Bay, Hokkaido (Jordan and Seale 1906). NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology le2 Osamu Tsuruoka et al. Fig, 3, Collection sites of the speclmens used in this study. Holotype (open circle) and non- type specimens (solid circles) of foelus mororants (Jordan and Seale, 1906) cemb. nov. and non-type specimens (solid squares) of Steigtstrum stojnqgeri Jordan and Gilbert, 1898, Off Esashi, Oumu, and Monbetsu, Okhotsk coast of Hokkaido; off Rausu on Shire- toko Peninsula, eastern Hokkaido; off Usuiiri on Oshima Peninsula, southern Hokkaido; and otT Miyagi Prefecture, Pacific coast of Honshu, Japan (Fig, 3) (this study). Rocky andfor boulder-stone areas in depths of 5-100 m. Comparison. According to Yabe (1985), Ibelus and Steigtstrum form a multiple- branching monophyletic group with Ricuzenius, Tdyrtscus, Sttengts, and lbelinus supported by a single synapomer phy: absence of the second and fourth pharyngob- ranchial bones. foelus and Steigistrum are very similar to each other in their exter- nal appearance (e,g., one dorsal scale row and one lateral-line scale row on body, pelvic fin eomposed of one spine and three soft rays), although they can be sepa- rated clearly by the presence (in lbelus) or absence (in Steigistrum) of teeth on the palatines (e,g., Jordan and Gilbert 1898; Watanabe 1958, 1960; Yabe 1985; Lindberg NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Validity of Steigistrum mororane in ketus 103 and Krasyukova 1987), Although Jordan and Seale (1906) originally described Stel- gistrum mororane as having no teeth on the palatines, the holotype of S. mororane has a few teeth en its palatines; therefore, this species should be regarded as a member of the genus Ibelas rather than the genus Steigistrum. As was mentioned above, Jbeltts mororanis dilifers from Steigistram stojnageri in having palatine teeth and, moreover, it can be distinguished also in having no scales on the cheek (vs small, scattered scales in S. stojnegeri). Additionally, dMer- ences in the number of second dorsal-fin rays, anal-fin rays, and lateral-line scales between Ibelus mororanis and Steigistrum stojnegeri are statistically significant (Mann-Whitney Utest; P NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 104 Osamu Tsumoka et al. been entrusted te NSMT, but these specimens have not been found there (G, Shino- hara pers. comm,). The ibllowing remarks are thus based wholly on Watanabe's papers cited here, since no reexamination of the material has been possible. Watanabe (1980) described the internal features of S. stojnageri based on one speci- men, no, 20147, which is reidentified herein as L mororanis because palatine teeth were illustrated in the drawings (Watanabe 1980, pl. 48). This specimen can be traced back through his earlier works. Watanabe (1978) described the external fea- tures of S. stojnageri on the basis of two specimens, nos 20146 and 20147, of whieh the latter at least is Z mororanis, as was shown above. Watanabe (1960) used the twe same specimens, according to that work's supplement (Watanabe 1981). The description of the external features of S, stojnegeri provided by Watanabe (1958) was based on one specirnen, w. 21882, and the descriptions of the internal features and quantitative characters were based on two uncatalogued specimens, These un- catalogued specimens were evidently the same as nos 20146 and 20147 used in Watanabe (1978, 1980, 1981), because the counts and measurements given in the ta- bles, figures, and plates of each study are much the same (ci Watanabe 1958, table 55, figs 77, 78. vs Watanabe 1978, table 55, figs 77, 78; Watanabe 1958, pl, 56 vs Wata- nabe 1980, pl. 48>, and Watanabe's (1958) description must, therefore, also have been partly based on at least one specimen of L mororanis. Watanabe (1976) pre- sented only S. stojnageri in his species list of the Cottoidei of Japan, but it would be appropriate to consider that the specimens involved included both l mororants and S. stojnageri, Lindberg and Krasyukova (1987) redescribed S, stojnageri on the basis of 30 specimens collected from the Sea of Okhotsk and the Sea of Japan, but some speci- mens of L mororants are likely to have been mixed in with them, because the re- ported number of dorsal-fin spines was 8-10, and dorsal scales 35--38 (L mororanis has 8-10 dorsal-fin spines and 25--38 dorsal scales, and S, stojnageri, 9-10 dorsal-fin spines and 29-36 dorsal scales; this study). Although no collection of L mororanis has been reported until now since its original description, it is likely te be widely distributed around the coast of north- ern Japan. The many specimens found in various collections and our own observa- tions while SCUBA diving demonstrate its oceurrence at least from the Okhotsk coast of Hokkaido to the Paci:fic coast of Miyagi Prefecture, Japan. Steigistrum stojnegeri Jordan and Gilbert, 1898 [English name: Furseal sculpin] [Japanese name: Ottosei-kajika] (Fig. 4, Tables 1, 2) Steigistrum stojnageri Jordan and Gilbert, 1898: 1921 [original description, holo- type, sex unknown (52mm TL), off Robben Island (-Ostrov Tyuleniy), Sea of Okhotsk off southern part of Sakhalin, Russia]; Jordan and Starks 1904: 248, fig, 7 [description, following Jordan and Gilbert (1899)]; Schmidt 1904: 117 (diag- nosis); Jordan et al. 1913: 258, fig. 195 (in part, listed, Japan, distribution); Popov 1931: 128-129 (diagnosis); Tanaka 1933: 263 (short comments on fish- eries); Okada and Matsubara 1938: 321 (key); Matsubara 1955: 1140 (key); Watanabe 1958: 14, 227-230, figs 77, 78, pls 5 (lower), 56 (in part, listed, Japan, di- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Validity of Steigtstrum mororane in Ibelus I05 agnosis, deseription of internal features); 1960: 32B5, fig. 11, pl. 20 (fig. 2) (in part, deseription); Ueno 1971: 91 (listed, Hokkaido and adjacent waters); Wata- nabe 1976: 6 (in part, listed, Japan); 1978: 230-232, figs 77, 78, pl. 19 (second from top) (in part, description); 1981: 106 (in part, description of quantitative fea- tures); Yabe 1984: 310 (diagnosis, distribution); 1985: 4 (material for phyloge- netic analysis of the superfamily Cottoidea); Nakabo 1993: 555 (key); Esch- meyer 1998: 1606 (validity); Nakabo 2000: 636 (key); Sheiko and Fedorov 2000: 29 (listed, Kamchatka and adjacent waters); Mecklenberg et aL 20e2: 447 (diag- nosis, distribution); Nakabo 2002: 636 (key); Parin et al, 2002: 86 (listed, Russia and adjacent countries, distribution); Fedorov et al. 2003: 80 (distribution). Stelgistrum steinageri [sic]: Jordan and Gilbert 1899: 456, pl. 54 (description); Solda- tov and Lindberg 1930: 182-183 (in part, diagnosis, synonymy, distribution); Popov 1933: 144 (listed, Sea of Japan); Andriashev 1935: 294-297 (in part, description, distribution, synonymy); Taranetz 1937: 109 (key); 1941: 430 (dis- cussion about Icelinae); Schmidt 1950: 127 (diagnosis, distribution); Lindberg and Krasyukova 1987: 195-196, fig, 121 (in part ?, diagnosis, distribution); Orlov 1998: 150 (listed, Pacific waters around Kuril Islands and southeastern Kam- chatka); Sokolovskaya et aL 1998: 5 (listed, Peter the Great Bay). Material examined. 7 specimens collected from Japanese waters (see Appen- dix). Description. Proportional measurements and counts as given in Tables 1 and 2. Body short, compressed posteriorly (Fig. 4). Head large, slightly depressed, Caudal peduncle slender, cempressed. Upper profile of snout steep. Nasal spine short, sharp. Meuth large; lower jaw extending forward slightly beyond upper jaw. Small teeth on jaws and vomer. No teeth on palatines. Interorbital space narrow, concave. Four preopercular spines: uppermost spine unbranched, sharp, hooked posterodorsally; lower three spines slightly blunt, directed posteriorly or ven- trally. Numerous small scales scattered on fringe of nasal spine, interorbital space, suborbital region, outer rim of eye, cheek, and dorsolateral sumbce of head. Both jaws and ventral surrface of head naked. Lateral line descending in slight arch ante- riorly, Seales on lateral line large, tubular, with serrated margins dorsally and Fig. 4. Steigistrum stojnegeri Jordan and Gilbert, 1898, HUMZ 92926, female,44.2mm SL, col- lected from off Hamanaka, Pacific coast of Hokkaido. NII-Electronic Library Service Japanese SooietySociety of SystematicSystematio Zoology 106 Osamu Tsuruoka et al ． 鴇 起 卜 H 寸 10 H 一 ゆ 卜 寸 D つ Φ O 銅 一 自つ 剛 O ⊃ ［ 卜 N ． ． ．． ．． ．． ．． ．，．の ．． ．即 ，．〇 ． ，． ，． ， 勉 H 鴫 ○つ ○つ 噂 卜 寸 0つ 劬 ．00 寸 寸 ．囚 ．卜 p◎ 冖 働 り る ． H11 卜、 咽 囚 雪 6 つ 制 NI 5 つ 囚 0 ．つ じqI つ HI 卜 HI H ． ［ HI 宵 NI 9 ． − 1 IH 1 1 ー 1 ． ー 1 IQ 1 1 1 弓 」 ｛ 亠 LO マ 寸 卜 L 卜 ぼ q Ψ め 覇 一 め 9 め 一 卜 ゆ 雪 一 丶嵩 ， ． ．．て ．． ．．尊．ε． ，． ．． ，． ．， ，，耳，． ， ，，． ， 塁 Q⊃ 寸 Q つ 頃 OD O つ 寸 H 寸 H o⊃ Oつ 咽 寸 0つ の HD の ON 勉 αっ ． 囚 鬥 H H ○つ H H つ 寸 冖 ○ つ q 目 同 一 目 祠 っ 祠 ， £ ． 雑 艇 嚇 り 腿 り 」 恥 H 一 H 一 鳶 軌 ， 鬥 ．寸 ． ， ．O ，卜 ，O ，ゆ ，，寸 ，O ，α ， ， ，Φ ， 尊つ， ， ．ト ，崘 ，0 ，α ， ，N ， 閏 Q p っ H ¢ O 00 HN 寸 α つ 卜 Q つ 卜 ぜ ゆ 卜 ．OD 頃 H ，野 H ［ O つ 0CqIHO α う 頃 鋤 雑 ． 寸 つ ． 一 寸 ○つ Q つ 崘 0 つ ○つ oっ つ 目 卜 冖 一 一 目 つ NI ー 1 ー ー − 1 1 1 1 ミ 鮪 11 IO 1 ー 1 − △ △ 1 ー − 1 1 ー − IN 1 マ − 卜 例 的 〕 〕 頃 ゆ 卜 Φ α⊃ 卜 α⊃ ⇔ 醐 寸 卜 ○っ め 卜 卜 的 ゆ 寸 喝 り 障 ． ． ．． ．． ．． ．． ．． ．． ．． ．． ．． ． ．． ．． ．． 鳶 の ON α H 崘 α つ α H H 卜 阜つ 0 つ ゆ H 〇 阜つ ト ○つ ◎σ N 寸 瓠 州一 的 国 国 σ つ 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一 軸 一 Cq の ＝ 螂 口 蝦 日 祠 帽 b9 bΩ 一 祠 既 一 Φ 価 』 hO O O oり 角 qO 価 q 州 口 口 口 口 の ρ 一 一 一 喟 』 頓 娼 bQ 廃 − bo q 一 OqO 幽 一 幽 刺 ρ O 唱 唱 の 【 娼 』 cd 』 − du 邸 Oq 一 O 』 O O 自 HOQ り ρ 〇 O 』 臼 』 に 口 自 り コ ← O 自 O − 。。 d ‘ q 【 』 冖 畧 O （ 唱 喟 唱 帽 弓 」 知 鮨 （ − − O O 娼 噂 く 【 的 2 − r 〇 q q 一 「 ← 【 q 口 O 【 q 価 臼 唱 『 q 唱 q ワ σ⊃ d 49 弼 二 帽 己 の O ［ 自 『5hO一 自 「 ρ ← 口 冖［ 」 口 』 口 ‘ 眉 一 一 一 pD H ‘ 唱 頓 螂 一 唱 旨 〇 ρ5 O ∀ 茎 賦 邸 嶋 応 唱 眉 − 冒 じpd Φ 喝 応 回 一 の ∀ 親 茄 謂 一 OO 們 一 鵡 目 b9 ℃ 州 ｝ ← 一 紬 ［ 〉 ‘ 〇 而 自 の 罵 り ＝ も ρ 彡 昌 応 円 』 凵 の 〇 邸 茄 口 酢 口 コ O 旨 口 Φ 邸 口 ‘ 』 q 一 而 05q uD ← d ‘ 一 宀 り 冖 〇 二 二 d 二 宀 円 q 一 頓 qD 画 O OO O 邸 コ 日一 一 畠 O 目 口 霜 ； 一 一 お ゜ bDq ’ 制 一 一 ← 扁 鯛 鯛 ロロ ε 判 匂 鯛 璃 』 咽 鯛 閣 口 q 畠 ℃ 噂 d qO 噴 一 〇 刺 蓴 鬥 d 回 州 ［ 一 “ 州 O O 』 o O O 鯛 O O O 一 O O O 口 Φ 自o 【 O ρ O 冫 》 O ［ 旨 ⊇ 喝 芍 冒 ≧ 彡 【 も 〜 旨 而 一 ∩ ‘ 噂 ‘ 瑁 ‘ ‘ O 眉 ‘ 』 昌 ‘ 園 ▼ 』 旨 O q 一 謂 一 ｝ 眺 財 ℃ 眉 ℃ 』 』 〇 ． 邸 百 防 あ 眉 邸 q 葛茄 茄 晟 bの 細 50 茄 砧 bDq 一茄 紬 贔 例 ℃ 弓 ℃ ℃ ロコ q5 邸 Φ ［ q 唱 oり Φ ＝ 口 9d qOq［ ＝ 一 自 Qワ 9 耀 口 O O Om O Φ 口 郎 萌順 角 角 角 O Φ 琶 自 O O O 一 甲 「 冖 、 」 〔 口 「 『5 則 円 ［ ロコ 国 国 宀 山 山 山 一q 一 H 【 【 Qつ 口 H 一 q 店 q 日 司 ρ 一 鳥 qり 目 卜 一 NII-ElectronicN 工 工 Eleotronio Library Service Japanese SooietySociety of SystematicSystematio Zoology Validity of S陀 ’∫〃 召 ηz moror α ne j皿 Icelus 107 、 N ト 酬 σ ¢ α っ 頃 起 、 ｝．守 ．つ， ． ．． ．． 鋤 H 杯｝ 鴫 σつ 円 OHIOOc 頓 の 罨 叭 0 州 卜． 啅 ¢ 一． つ 頃 ． − 1 IO IH 丶§ − 酬 ， 嶂 ，σつ．〔寸 ．．つ．コ．コ．． 黛 C ◎ 国 卜 回 寸 卜 卜 O 一 鋤 囚 頃 寸 Qつ σ っ 寸 自 詑 」 qoQ ． eq，斜 ．P つ，Qつ．の ，Qつ，訓 ．卜 ． OmI D つ 冖 吩 口 OHINΦ 寸 03 爲 痔 り 0 ⊃ ○つ 調 守 ． − マ マ 劬 11 ー ．、 IO 1 ー − ー ⇔ ⊂障 O つ ゆ 卜 の 黛 ． ．． ，． ．． ．． 嵩 ゆ 摩つ ひつ D つ 寸 寸 一 り 児 寸 鴫 吩 N 寸 岬 寸 効 耜 ○ つ Φ NOQ q 自 あ 寸 ． 創っ． ．の ．H ． H ．寸 ．N ． ← 図 O ゆ N 鳴 肥⊃ 寸 ○◎ 卜 〇 薯 O 一 寸 鴫 頃 』 寸 寸 寸 〇 乞 ρ 』 Q り 厂h 描 の ． ．り ．卜 ．⇔つ．囲 ． ，H ．凶 ． 廴 邸 ． O σ つ H 卜 00 O ．寸 働 鋤 【 砺 剛 のー 四ー つ 鳴 岬 寸 の 11 ． 1 1 ． IH 1 1 − − 懃 コ Φ ゆ ゆ 寸 Φ ゆ 頃 卜 匿 曽 ． ．． ．． ． ，． ． 母 鱈 O 吩 頃 H 5 つ ○ つ 寸 寸 N 薗 畔 Q う 鄭 描 の ミ 嵩 k 掲 H ． ．の ．H ．自 つ．O ．寸 ．N ，卜 ， め 口 ゆ O 0つ 囚 〇 09 卜 C自 摘 卜 卜 0 つ 自 つ 一 賄 鴫 N 豊 「 − 1 ー ー 1 呷 マ 働 工 卜 ゆ 茎Q 円 NQ 黛 ． ． ．． ．． ．． ． 萼 ○ つ H 鴫 の 卜 寸 N り Q 牾 頃 頃 国 制 寸 寸 寸 Oq ゆ あ OD e コ の 恥 N 鴫 斜 囚 輪 ． ．，． ．、 ，， ． N 〕 鶲 O 鴫 戉 H 鴫 ¢ DN 卜 Φ た ｛ NO の 寸 訊 寸 Q っ 寸 ◎ O Po kO ［［ う 」 O ミ Q 弓 H 6 ト α 寸 卜 ， ．寸 ． ．つ．卜 ．．っ．． 寒 囚 丶 q 寸 O つ O 寸 HH ゆ α⊃ 卜 旬 O 寸 鴫 α つ αつ マ 鴫 あ 円 ゆ ヤ ー IG 10 ー ー ー 1 畠 11 O 晒 つ ゐ ゆ 一 寸 工の ロ ， ．． ．． ．， ．． O 黛 尋 ゆ 卜 Q つ 寸 卜 O ひ う 囚 寸 C ⊃ 困 寸 の 寸 口 ． 自DO ロ あ 刊− 自 ．OqHH ℃ 6e σ5 q 一あ ℃ 謂 − q （ 鯛 ⊆ 祀 （ Oq9 d 日 彦 。 O ． ［ 邸 g 瞥 ＝ 』 「 ‘ O 剃 霜 Ω ， 鶏 』 鳴 h ） 【＼、 茄 彡 防 弓 帽 網 自 〕一 d し 彡 qq 亶 邸 ［ ρ 燭 七 』 ＝ 』 』 一 瓢 茄 細 一 ＝ 口 謂 調 励 O 邸 口 日 H 鳴 二 刷 自 口 一 一 一 謁 細 泊 蝋 』 〇 ℃ 口 一 自り 一 一 C 〇 o 謂 口 Φ Φ ρ ＞ GO ∈ 一 旨 ρ O q ℃ 嚀 彡 ＝ 二 二 ← O 旨 匪 ． 旨 」 Ub （ ← 弓 づ 』 Oq ， 邸 ＝ づ Ω 祉 馬 二 自 一 而 面 O ⊆ 」彊 眺 ℃ gり ＝ 一 自 眺 O 口 ¢ O O 自DO 刊 〇 95 謁 一 、口 ． 一 固 ［〒 の ［ 同 口 一 氏 一〇 ρ 一 ≧ 〇 の Σ 」鳴 臼 ＝ ［ o ρ 一 NII-ElectronicN 工 工 Eleotronio Library ServiceServioe Japanese SooietySociety of SystematicSystematio Zoology 108 Osamu Tsuruoka et al ． c 随 回 一 lo つ 昏 1 1 Q囗 あ ， “ 輻 』 幅 煢 D っ H 馬 q二 O 昏 O 1 卜 虻 H コ 口 H 卜 自 芍 鵯 蝦 り − o馳 【 偽 あ 而 嵩 邸 旨 N 旨 O 動 ← d di に り 邸 葭 冖 卜 H くΩ 吩 tONN 馬 H N ト 』 娼 旨 ○勹 雪 o っ H 唱 H − 餌 邸 § 自ワ ． hO o ⊃ 措 〇 』 も 一 勢 u H 〇 QNN 弓 つ 寸 iH 〇 030 cuP ⊃ c つ Icu 雪 崘 H っ su 口 Q口 賄 OO Φ q 劬 一 潮 の 【 ， N ｛ 豎o 霜 葭 の 』 卜 ¢ N ー ． 旨 α う O α っ 寸 Nk ← 而 魁 ［』 融 跫 や 寸 り ー 鬥 薯 H 1 p っ ヨ 1 建 嵩 粒鼠 鴇的 di Ut 鯉 Q っ 吩 一 r→ 孕つ の H 粕 一 自 っ Qり 色 の ら しっ N 邸 階 』 OkOkO o ＝ o っ 寸 10 叭． 卜 目 11 乳 口 ヨ 頓 尸 H 亠 嵐 篤 。。 ミ 自 ロ 吻 ぐ 嵩 ・， 丶 ヨ 鳥 ¢ ゜ っ コ国 黝 邸 ¢ 噂 卜 も ○⊃ H ー 沿 翫 卜 乳 寸 o っ ・・ 醗 H 嵩 コ 卜 OH 鬥 H6 起 』 碑 お ＝ 口 “ 唱− ∪○ コ ・・ o ← 屑 q 旨 OH HH ≧ ゜q コ 国 o⊃ 1 1 ⊂q 、 1 祠 090 £ 圃 唱 屑 （ （ （ 』 （ 一 一 ． O H （ ［ （ 自 卜 卜 卜 ヨ 口 亅 1ー 国 11 凵 一 黛 疑 ミ 匿 ∀ 冑 ） ） 謂 ） ） 8 bD σ5 ◎5 一 口 州 口 “ ＝ 瓸 雨 自Doo の ・。 q り ・。 蝋 自 づ 区 ⇒ 巴 Oo 而 コ → 餌 舘 餌 9 笛 ロ 自 ［d d dqO ［ 置 日 日 日 〇 （ （ （ 一 O O o o o ← o っ O つ αっ 口 O ．と む OH 亀 亀 OH118娼 煢 ρ 門 NL 鴇 鴇摘 鴇 帽 」 」 11 冂 』 鴨 O （ 亀 m （ 』竃（ 胴 冑 ぜ 黛 寸 寸 g弓 ） 懃 懃 （ ） 罍 審 （ 懃 懃 （ 一 Q 冂 勃 118OH 同 弓ミ 曽 物 黛 謎 障 舘 眉 N 黛 H 閑 給 跫 H 跫 母 母 ） 日 齪 智 ） あ 鵠 嚇 愈 N 囹 § 冂 ご 竜 § O 物 協 遣 匿 リ 的 匿 § 劬 。， 醤 ミ 口 kOkO k ） kOkO ミ） 」 k ） 勉 鋤 ミ ミ 醤 ミ ミ 鳶 象 建 ミ 鞄 離 藩 Q ミ O 5 篭 凝 謡 国 艇 o 起 σ kN kN ミ隔k Q 芝ミ爲 勉 建 如 劬 ミ 篤 物 ミ 賄 的 』 の 的 的 防 鴇 圃 り り 勇 装 燭 い の 蘿遘 の 偽 国 識 蝕 専 嵩 謡 遣 懃 専 起 窘 越 謡 ， 丶 丶丶 N ミ 丶 懃 懃 N N 嚇 嚇 勅 勉 勉 鋤 鵯 勉 曽 電 囚 N 隔 ミ N 越 ミ 効 恥 【 ミ り 的 q 酷 ミ 的 の ミ ミ 起 ゜っ ゜っ ミ 起 ρ ¢ 』 一 NII-ElectronicN 工 工 Eleotronio Library ServiceServioe Japanese SooietySociety of SystematicSystematio Zoology Validity of Stelgistrum mororane in∫lcelus 109 k IN Φ o っ 燭黔 o 誉 糧 ﾁ の っ ー 角 ρ 偽 建 」 り 一 ⊃ d ← 角 軸り ⇔q 1 』 H O 邸 娟の ∩ N 勉 自 H 酷 一 \ O 尊 寸 ○つ o ぴ っ っα っ 一 y QoQ ｿ N （的 11 〇 1 寸 寸 貸） OLO~」 O ミ 師 H 矧 一 覧 寸 （ [11 置O 1 窘 雪 、 ＋ 臼 起 −胴 輻 O OkO H 屑 拍 O 卜 ゜っ． 肖 ＞ 卜 【 Q 鳴 〉十 ＜ 鏑 畑 り 謎 1o 【 m N 寸 −@ 1L ） 嵩、 H 障 ． 而 冖 N 導 〜 寃 ﾁ o 』 11 ◎ 而 od 鋤 h 1 翰 唱 ー 一 う 懃 ミ ~ ρ 崎 目 口 IHH 楠 き ミ 一 丶N O N 的 壁 ） O （ 』 輪 特 自 （ の H O 粒 と N 偽 ＞ の 虻 「 寸 一 越 O 」 11 （ 11 【 ミ Nk （ 黛 N り的 Q 卜 価 黛 窃 恥 ） 遣 謡N 1 舒 11 冒 黝 勉 ） H1 邸 」 起 黝 陛 目 隔 ミ ， 黔 ’ 捷 謡 一 邸 鯰 じ 階 ー O ． 」 帆oD 懃 層 ． ミ 勉 ミ 轟 ） 自 勉 （ 一 蕾 踊 q ） ⊆ り ＞ 隔 描 qh リ の の H の ミ 甓 鴇 Oq O 剃 同 ミ 嵩 劬 虻 寸 コ 邸 餌 目 謡 k り 鴨 δ k 穐 蒿 cu う 日 設 防 σご 鼠 （ 隔 偽 （ ヨ O の α 』 卜 り 物 N O 自 ） ー 嵩幅 ρ 』 摘 届 勉 と 軸 11 鳴 ー 鯉＄ ミ ○ ⊃ 鴇k O 鋤 剃 ﾁ 一 動 ミ kQ 恥 （ （ り OH11 輪 』 u 鳩 O 黛 ワ N 的 （ 如 鋤 寸 11IH り q ） 凵 鬥 ＞ 』 一 口 黛 黛 一 〇 円 舘 O 口 一 つ 邸 oQ ） 〇 〇 H11 ） 鮭 」 遣 舘 q 」 【 冨 ． ミ ） ⊆ O 而 葭 （ 姶 む醤 叫 韆 【 N 輪 O 目 卜 黛 嵩 O （ 目） H の§ QN 超 ヨ 自o ら 設 恥 O 鵡 剛 齪 α GkO ミ 謡 鴇 り つ 日 （ 離 kN 唱 O 幅 凵 GD ） 唄 吻 り も k 舘 自 の k ρ と 鋤 ） 円 に ω 頓 鳩 劬 的 謡 On 一コ 麟 〇 囚 口 ＞ H α ＜ 価 卜 NII-ElectronicN N ¢ 朝 ゆ Library1 ． ， Service． o 臼 一 JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 110 Osamu Tsuruoka et aL posteriorly, Dorsal scale row originating beneath second spine of first dorsal fin and extending to caudal peduncle. Small scales scattered above dorsal scale row and en spines and soft rays of dorsal fin. No scales between dorsal scale row and lateral 1ine, nor below lateral line, One to three midsize scales scattered in pectoral axilla. Belly naked, Three pairs of cirri on dorsal surface of head. Unbranched cir- rus on dorsal margin of eye, Two simple cirri on posterior end of maxilla. Small, simple cirrus on suborbital stay, another on posterior region of opercle. One small, simple cirrus on base of each preopercular spine and on several lateral-line scales, First dorsal fin originating at level of dorsal edge of gil1 slit. First and second dorsal fins joined by low membrane; second dorsal fin originating above 12th lat- eral-line scale. Anal fin originating below origin of second dorsal fin, Pectoral fin large, rounded posteriorly. Pelvic fin slender, originating beneath second spine of first dorsal fin, Caudal fin slightly rounded. Anus located below ninth or 10th spine of first dorsal fin. Urogenital papilla of male absent. Distribution. The present specimens were collected off Chirrip, Hamanaka, and off Atoeka, Kushiro, both in eastern part of Hokkaido, Japan, the western North Pacific (Fig. 3), This species is widely distributed in the Sea of Okhotsk from Tauiskaya Bay, the Shantar Islands, and the Jonas Island via the eastern coast of Sakhalin, Terpenie Bay, and Aniwa Bay to the western coast of Kamchatka and Iturup Island (Andriashev 1935; Schmidt 1950; Lindberg and Krasyukova 1987). In the Sea of Japan, it occurs along the coast of the Primorskii from the Tatar Strait to the Peter the Great Bay (Schmidt 1950; Lindberg and Krasyukova 1987; Sokolovskaya et al, 1998). In addition, it is known from the western coast of the Kuril Islands, the western North Pacific 1998; Parin et al. 20e2), Parin et al, (Orlov- (2002) recorded this species from depths o ± 18 to 320m. Remarks. In order to eonsider the intra-specific variation of this species, non- type specimens collected from Russian Sea of Okhotsk were used as comparative material in addition to the holotype (see Appendix>. The seven specimens collected from off Ateeka and Chirrip in eastern Hokkaido are clearly assigned te Steigistrunz by the presence of four preopercular spines, teeth on jaws and vomer but not on palatines, scales with serrated margins forming the lateral-line scale row, enlarged scales each with serrated margin form- ing one dorsal scale row, and one spine and three soft rays in the pe}vic fin (Andri- ashev 1935; Yabe 1985). Three species, S. stojnegeri, S. beringianum, and S. concin- num, are recogriized in this genus. Steigistrum stojnageri is distinguishable from S. beringianum by having the dorsal scaie row extending to the base of the caudal fin (not extending beyond the insertion of the second dorsal fin in S. beringianum), and scattered scales on the snout and cheek (vs such scales being absent or very few), It is also distinguishable from S, concinnum by having the maxilla extending nearly to the posterior edge of the eye (to the anterior edge of the eye in S. concin- num), no cirrus on the nasal spine (vs one unbranched cirrus there), and a single pair of nuchal cirri (vs two pairs>, The present seven specimens have all the diagnostic characters of S. stojnageri mentiened above, In other morphological charaeters, they are also in complete agreement with the holotype of SteLgistrum stojnageri (e.g., in the extent of scale- covered areas on the body). Their counts and proportional measurements coincide with, or are close to, those of the holotype and other specimens from the Russian side of the Sea of Okhotsk and Sea of Japan (Tables 1, 2). Thus, these seven speci- NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Validity ot' Steigistr"nz mororane in lbelus 111 mens are identified as S. stojnegeri. No significant intra-specific variation was found between the present specimens and the comparative material. Many authors have included Muroran in the distributional range of S, stoj- negeri (Okada and Matsubara 1938; Schmidt 1950; Matsubara 1955; Watanabe 1958, 1960, 1978; Ueno 1971; Yabe 1984; Nakabo 1993, 2000, 2002). Except for Watanabe, these authors merely followed Jordan et al, (1913>, who synonymized Steigistrum mororanis under S. stojnageri; their reports included no new topotypic specimens from Muroran. As was mentioned in the remarks under L mororanis, S. stojnagri might have been included in Watanabe's studies, but his specimens are unavail- able now. Thus the present report constitutes the first confirmed record of this species from Japanese waters. Acknowledgements We express our sincere thanks to Dr G, Shinohara (NSMT) for providing data on the holotype and infbrmation concerning the specimens owned by Dr Wata- nabe; Drs T. Iwamoto (CAS) and H, Imamura (Hokkaido University Museum; cur- rently HUMZ) fbr providing data on the holotype; Proi K. Nakaya (HUMZ) and Dr F, Muto (National Research Institute of Far Seas Fisheries) fbr their valuable sug- gestions on this study; and Dr H. Munehara (Usujiri Fisheries Station, Hokkaide University), Mr K. Seki (Shiretoko Diving Kikaku), and Mr N, Sato (Diving Service Grunt Sculpin) for their help tn cellecting specimens and for providing valuable infbrmation, We are also gratefu1 to Dr A, J. Orr (Alaska Fisheries Science Center, NOAA) for critically reading the manuseript. This study was partly supported by a Grant-in-Aid fbr Science Research (No. 19570076) fr'om the Japan Society fOr the Promotion of Science. Appendix: List of the Specimens Examined lt]elus mororanis: SU 9260 at CAS (holotype), harbor of Muroran, Volcano Bay; HUMZ 201287, off Menashitomari, Esashi, Sea of Okhetsk; HUMZ 190078- 190080, 194919, 201285, 201286, off Oumu, Sea of Okhotsk; HUMZ 87725, off Lake Komuke, Sea of Okhotsk; HUMZ 188575, 190126-190129, off Rausu, eastern part of Hokkaido, northwestern Pacific; HUMZ 121684, 121717, 175832, 175833, 179894-179900, 179903-179906, 179908, 179910, 179913-179917, 185295-1853e2, 185304, 185305, 186672-186675, 186722, 186723, 187659-187662, 187824-187831, 187866-187892, 187894, 189778-189783, off Usujiri, Hakodate, nerthwestern Pa- cific; HUMZ 188567-188571, eff Onagawa, Miyagi Prefecture, northwestern Pa- cific. Steigistrum stojnegeri collected t'rom Japanese waters: HUMZ 92925, 92926, off Chirrip, Hamanaka, eastern part of Hokkaido, northwestern Pacific; HUMZ 201289-201293, off Atoeka, Kushiro, eastern part of Hokkaido, northwestern Pa- cific, Comparative specimens Ibetus gilberti: ZISP 25514 (syntype), Olga Bay, Sea of Japan; ZISP 25328 (syntype), Vladimir Bay, Sea of Japan; HUMZ 87726, off Shokotsu, Sea of Okhotsk; HUMZ NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology 112 Osamu Tsuruoka et aL 140590, off Lake Saroma, Sea of Okhotsk. icelus sekii: HUMZ 179901, 179902, 187896, 187897, 188572-188574, 198892, 198893, 200258 (incl. holetype and 3 paratypes), off Rausu, eastern part of Hokkaido; HUMZ 198857, off Shari, Shiretoko Peninsula, eastern part of Hokkaido, Sea of Okhotsk; HUMZ 187634, Moekeshi Bay, Iturup Island, Sea of Okhotsk; HUMZ 187898, 188505 (2 paratypes), off Urakawa, Hokkaido, northwestern Pacific; HUMZ 179892, 190688, 190689, 192091, 192092, off Usujiri, Hakodate, northwest- ern PacMc. SteLgistrum stojnageri: USNM 48236 (holotype), off Robben Island, Sea of Okhotsk; HUMZ 141143, off Feklistova Island, Sea of Okhotsk; HUMZ 140178, off Bolshoi Shantar Island, Sea of Okhotsk; HUMZ 140715, Sevemy Bay, Sakhalin, Sea of Okhotsk; SU 7577 at CAS, Aniwa Bay, Sakhalin, Sea of Okhotsk; SU 22324 at CAS (incl. 5 specimens), off western coast of Kamchatka, Sea of Okhotsk; ZISP 44166, Aniwa Bay, Sakhalin, Sea of Okhotsk. References Andriashev, A. P, 1935, Ubersjcht der Gattung Steigtstrum Jordan und Gilbert (Pisces, Cotti- dae) nebst Beschreibung einer neuen Art aus dem Beringmeer. Zoologischer Anzeiger 111: 289-297. BOhlke, J. E. 1953. A catalogtie of the type specimens of recent fishes in the Natural History Museum ofthe Stanford University. Stanfbrd Ichthyological Bulletin 5: 1-168, Eschmeyer, W. N. (Ed.) 1998. dntalQg ofFishes, Vbls 1-3. Califbrnia Aeademy of Sciences, San Francisco, 2905 pp. Fedorov, V. V,, Chereshnev, I. A., Nazarkin, M. V,, Shestakov, A, V, and Volobuev, V. V. 2003. CZitatog ofMtzrine and 1;)'eshtvater Ftshes of the Aiorthern Part of the Sea of Okhotsk. Dal- nauka, Vladivostok, 204 pp, LIn Russian] Hubbs, C. L. and Lagler, K. F. 1958, Fishes of the Great Lakes region. Bulletin of Cranbroek Institute of Science 26: 1-213. Imamura, H, and Yabe, M. 2002. Demise of the Scoxpaeniformes (Actinopterygii: Percomor- pha): an alternative phylogenetic hypothesis. Bulletin of Fisheries Seiences, Hokkaido University 53: 107-128. Jordan, D. S. and Gilbert, C. H. 1898. Steigistrum, new genus. Pp. 1921-1922. in: Jordan, D, S, and Evermann, B. W. (Eds) The fishes of North and Middle Americai a descriptive cata- logue of the species of fish-like vertebrates fbund in the waters of North America, north of the Isthmus of Panama. Part II. Bulletin of the United States National Museum 47: 1-xxlv+1241-2183. Jordan, D. S. and Gilbert, C. H. 1899. The fishes of Bering Sea. Pp. 433-492, pls 43-85. in: Jor- dan, D. S. (Ed.) F2tr Seats, anci Fletr-seat lslands of the IVOrth Pacptc Ocean. Pttrt llI] Gov- erriment Printing Office, Washington, D.C. Jordan, D. S. and Seale, A. 1906. Descriptions of sjx new species of fishes from Japan. Pro- ceedings of the United States National Museum 30: 143-148, Jordan, D. S. and Starks, E. C. I904, A review ot' the Cottidae or sculpins fbund in the waters of Japan, Proceedings of the United States National Museum 27: 231-335. Jordan, D. S,, Tanaka, S. and Snyder, J. O, 1913, A catalogue of the fishes of Japan, Journal of the College of Science, Imperial University of Tokyo 33: 1-497. Lindberg, G, U, and Krasyukova, Z. V. 1987. Fishes of the Sea of Jtzpan and Adjacent Ftirts of NII-Electronic Library Service JapaneseJapaneseSociety Society ofSystematicof Systematic Zoology Validjty of Steigtstrum mororane iii ibetus 113 Okhotsk and Yiztlow Sea, Part 5. 71eleostomi, Osteichtdyes, Actinoptetlygii, 30 Scorpaeni- formes (J76. lhm. Scorpaenidae-194. Fam. Liparididaoj, Nauka SSSR, Leningrad, 526pp, [In Russian] Matsubara, K. 1955. Fish Aclbrphology and Hierarcby. Part JIL Ishizaki-shoten, Tokyo, xi-v+ 791-1377pp. [In Japanese] Mecklenburg, C. W., Mecklenburg, T. A. and Thorsteinson, L. K. 2002. Ftshes ofAlaska. Amer- ican Fisheries Society, Bethesda, Maryland, xxxvii p 1037pp. Nakabo, T. 1993. Cottidae. Pp. 548-567, 1300-1303. ini Nakabo, T. Jbelus. Occasional Papers of the California Academy of Seiences 138: 1-52. Okada, Y. and Matsubara, K. 1938. Kigys to the Fishes and Fish-like Animals of .Jtlpan. San- seido, Tokyo, xl+584pp., 113pls. [In Japanese] Orlov, A, M, 1998, Demersal ichthyofauna of Pacific waters around the Kuril Islands and southeastern Kamchatka. Russian Journal of Marine Biology 24: 144-160. Parin, N. N., Fedorov, V. V, and Sheiko, B, A. 2002. An annotated catalogue of fish-like verte- brates and fishes of the sea of Russia and adjacent countries. Part I. Order Seorpaeni- formes. Journal of Ichthyology 42 (supplement l): 60-135. Popov, A. M. 1931. Zur Fischfauna des ochotskischen Meeres. Issledovanija Morey SSSR 14: 121-154. rln Russian with Geuman summary] Popov, A. M. 1933, Beitrage zur Ichthyofauna des japanischen Meeres. Issledovanlja Morey SSSR 19: 139-155. [In Russian with Geuman summary] Schmidt, P. J. 1904. Pisces .vatxriunz Orientaltum imperii Rossici [Fishes of the Eastern Russian Empire]. Geograficheskogo Obshchestva, St. Petersburg, xiT466pp,, 6pls. [In Russian] Schmidt, P, Y. 1950, Ryby Okhotskogo Moray [Fishes of the Sea of Okhotsk]. Trudy Tikhookeanskogo Komiteta 6: 1-370, pls 1-20. [In Russian] Sheiko, B. A, and Fedorov, V. V, 2000, Chapter L CIass Cephalaspidomorphi-Lampreys. Class Chondrichthyes-Cartilaginous fishes. 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