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Lycenchelys ryukyuensis Shinohara & Anderson, 2007 (Japanese name: Ryukyu-hebigenge) (Figs. 36���37; Table 9) Lycenchelys ryukyuensis Shinohara & Anderson, 2007: 59, figs. 1���3, table 1 (original description, type locality: Okinawa Trough, Ryukyu Islands, East China Sea, Japan); Hatooka, 2013: 1225, 2078, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 216 (species list and etymology of scientific name). Materials examined Holotype: NSMT-P 63965, male, 159.3 mm SL, Okinawa Trough, eastern Ryukyu Islands, East China Sea (26��11.34���N, 124��54.27���E to 26��12.65���N, 124��55.47���E), 1493���1533 m depth, 26 Apr. 2002, R/V Tansei-maru, beam trawl. Paratypes (2 specimens): NSMT-P 63966���67, 2 females, 116.9���142.6 mm SL, collected with holotype. Diagnosis. Vertebrae 22���24 + 96���102 = 118���124; head length 14.1���14.4% SL; intertorbital pores and occipital pores absent; postorbital pores usually 3; suborbital pores 6 + 1���2; preoperculomandibular pores 8; vomerine teeth 2���4; palatine teeth 1���2, arranged in single row; opercular flap well developed; pelvic-fin base positioned anterior to lower edge of gill opening; lateral line complete and positioned ventrally; scales absent on pectoral fin and its base; body uniformly grayish brown when fresh. Description. Counts and proportional measurements in Table 9. ....Continued next page Body very elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 3.4���4.0 (3.4)% SL. Head moderately short, oval, dorsal profile of head gently sloping to above pectoral-fin base. Snout short, 133.7���149.7 (149.7)% of eye diameter. Eye ovoid, relatively small. Interorbital space narrow, width 19.7���22.9 (19.7)% of eye diameter. Nostril tube short, not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching to about vertical through middle of eye in adult male (holotype), reaching to below anterior margin of eye in females (2 paratypes). Labial lobe of lower jaw reduced. Teeth on jaws, vomer and palatine small and conical; upper jaw with 2���3 rows anteriorly and 1���2 rows posteriorly; lower jaw with 2���3 irregular rows anteriorly and 1���2 rows posteriorly; vomerine teeth irregularly arranged; palatine teeth in single row. Lower edge of gill opening about level with lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and triangular. Pseudobranch filaments extremely short. Lateral line deciduous. Scales small and cycloid, present on body, tail and about basal half of vertical fins. Head, nape, pectoral fin and its base lacking scales. Dorsal-fin origin above middle of pectoral fin; 1st dorsal-fin pterygiophore between neural spines of 7th and 8th vertebrae. Anal-fin origin below 15th or 16th (15th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 3rd to 5th (between 4th and 5th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 3rd to 5th (between 4th and 5th) preural vertebrae. Caudal fin with 2 epural, 4���5 (4) upper hypural and 3 lower hypural rays. Pectoral fin moderately short, reaching to about middle of abdomen; its posterior margin rounded dorsally and notched ventrally. Upper end of pectoral-fin base on about lateral midline of body. Pelvic fin short; its base posterior to lower edge of gill opening; its posterior margin reaching lower edge of gill opening. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above 1st suborbital pore (Fig. 37A, B). Postorbital pores 3 (Fig. 37A, B); distance between 1st and 2nd pores longest of those between adjacent pores. Suborbital pores 7 or 8 (7); 6 pores below eye and last 1 or 2 (1) pore(s) on ascending part of suborbital canal behind eye; 6th pore behind vertical through posterior margin of eye (Fig. 37A). Preoperculomandibular pores 8; 4 on lower jaw and 4 on preopercle; last preoperculomandibular pore posterior to lower margin of eye (Fig. 37A, C). Interorbital pores and occipital pores absent (Fig. 37B). Color in alcohol. Holotype (Fig. 36; lacking body skin) with dark brown head and pectoral fin, flesh colored body, grayish vertical fins, and pale brown margins on vertical fins. Color when fresh [based on color photograph in Shinohara & Anderson (2007)]. Head and pectoral fin blackish, body grayish, and margins of vertical fins dark brown. Distribution. East China Sea off the eastern Ryukyu Islands and in the Okinawa Trough, at depths of 991���1533 m (Shinohara & Anderson, 2007; Hatooka, 2013; this study). Size. Maximum length 17 cm TL (Hatooka, 2013). The largest specimen examined during this study measured 159.3 mm SL (161.6 mm TL). Remarks. Shinohara & Anderson (2007) compared L. ryukyuensis with Lycenchelys aratrirostris Andriashev & Permitin, 1968, Lycenchelys bellingshausenni Andriashev & Permitin, 1968, Lycenchelys folletti Anderson, 1995, Lycenchelys lonchoura Anderson, 1995 and Lycenchelys maoriensis Andriashev & Fedorov, 1986. Although L. ryukyuensis is similar to these species in sharing a combination of five characters (no interorbital pores and occipital pores, usually 3 postorbital pores, presence of pelvic fins and a single ventrally positioned lateral line; Shinohara & Anderson, 2007), it is distinguishable from L. aratrirostris, L. bellingshausenni, L. folletti and L. lonchoura in having higher numbers of dorsal- and anal-fin rays, and total vertebrae (110���114, 98���104 and 118���124, respectively, in L. ryukyuensis vs. 97���107, 79���89 and 101���110 in L. aratrirostri, 94���103, 82���91 and 102���110 in L. bellingshausenni, 97���104, 83���92 and 102���109 in L. folletti, and 103, 86 and 107 in L. lonchoura), and from L. maoriensis in having 7���8 suborbital pores and no black or brown variegations along the body and tail in males (vs. 6 suborbital pores and black or brown variegations present in L. maoriensis) (Shinohara & Anderson, 2007). On the other hand, Lycenchelys polyodon Anderson & M��ller, 2007 also has the above-mentioned combination of 5 characters, but has not been compared with L. ryukyuensis. The two are difficult to separate on the basis of numbers of dorsal- and anal fin-rays, total vertebrae and suborbital pores (113, 105, 123 and 7, respectively, and no variegations on the body and tail in L. polyodon) (Table 9). However, L. ryukyuensis is clearly distinguished from L. polyodon in having considerably lower numbers of vomerine and palatine teeth (2���4 and 1���2 in L. ryukyuensis vs. 26 and 31���33 in L. polyodon) (Table 9)., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 42-45, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Anderson, M. E. & Moller, P. R. (2007) Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere. XIII. Two new species of Lycenchelys from the Southwestern Pacific. Species Diversity, 12, 175 - 185. https: // doi. org / 10.12782 / specdiv. 12.175","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133."]}

Lycenchelys aurantiaca Shinohara & Matsuura, 1998 (Japanese name: Daidai-hebigenge) (Figs. 5���8; Table 2) Lycenchelys aurantiaca Shinohara & Matsuura, 1998: 151, figs. 1���3, table 1 (original description, type locality: off Miyagi Prefecture, Pacific coast of Honshu, Japan); Imamura, 1998: 31, fig. 8 (brief description); Hatooka, 2000: 1033, unnumbered fig. (key to species); Hatooka, 2002: 1033, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 15 (species list); Shinohara & Anderson, 2007: 64 (key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 723 (species list); Amaoka et al., 2011: 318, unnumbered fig. (brief description); Balushkin et al., 2011: 1026 (species list); Hatooka, 2013: 1227, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: NSMT-P 53147, male, 135.7 mm SL, off Miyagi Prefecture, Tohoku District, northwestern Pacific (37��58.54���N, 142��09.00���E to 37��56.84���N, 142��09.07���E), 700 m depth, 21 Apr. 1997, T/V Tanshu-maru, otter trawl. Other specimens (16 specimens, 91.6���141.3 mm SL): HUMZ 152389���90, 177032���33, 180582 ���83, 180864 ��� 67, 182661, 192779, 206876, 206878, 9 males and 5 females, 91.6���141.3 mm SL, Tohoku District, northwestern Pacific; HUMZ 192380���81, 1 male and 1 female, 102.2���123.8 mm SL, eastern Hokkaido Island, northwestern Pacific. Diagnosis. Vertebrae 19���20 + 66���70 = 85���89; head length 14.3���18.4% SL; interorbital pore 1; occipital pores absent; postorbital pores 4; suborbital pores 5 + 1; preoperculomandibular pores 7 (rarely 8); vomerine teeth 4���10; palatine teeth 8���25, arranged in single row; opercular flap well-developed; pelvic-fin base positioned anterior to lower edge of gill opening; lateral line incomplete and positioned ventrally; scales absent on pectoral fin and its base; body uniformly reddish orange when fresh. Description. Counts and proportional measurements in Table 2. ....Continued next page Body elongate, in cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 3.7���7.7 (4.0)% SL. Head short, ovoid; dorsal profile of head gently sloping to dorsal-fin origin. Cheek more swollen in large specimens (including holotype) than in small specimens. Head of adults longer in males than in females. Snout short, 67.5���111.4 (106.0)% of eye diameter. Eye ovoid, moderately large. Interorbital space narrow, width 10.9���33.7 (26.7)% of eye diameter. Nostril tube short, not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw about reaching vertical through 5th suborbital pore in adult males (including holotype), reaching below posterior margin of eye in females and juveniles. Labial lobe of lower jaw weak. Teeth on jaws, vomer and palatine small and conical; upper jaw with 2���3 rows anteriorly and single row posteriorly; lower jaw with 2���4 irregular rows anteriorly and single row posteriorly; vomerine teeth irregularly arranged; palatine teeth in single row. Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and slender (Fig. 6). Pseudobranch filaments short. Lateral line deciduous, incomplete and positioned ventrally; originating posterior to last postorbital pore and ending anterior to anus. Scales small and cycloid, present on body, pectoral axilla, tail and about 40���60% vertical fins basally. Head, nape, pectoral fin and its base without scales. Dorsal-fin origin near vertical through posterior edge of opercular flap; 1st dorsal-fin pterygiophore between neural spines of 2nd and 3rd vertebrae. Anal-fin origin below 17th or 18th (18th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 4th (between 3rd and 4th) preural vertebrae. Caudal fin with 2 epural, 3���4 (4) upper hypural and 4 lower hypural rays. Pectoral fin moderately short, reaching to middle of abdomen; its posterior margin rounded. Upper end of pectoral-fin base on about lateral midline of body. Pelvic fin short; its base anterior to lower edge of gill opening; its posterior margin reaching to or slightly beyond lower edge of gill opening. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above and slightly anterior to vertical through 1st suborbital pore (Fig. 8A, B). Postorbital pores 4; distance between 1st and 2nd pores longest of those between adjacent pores (Fig. 8A, B). Suborbital pores 6; 5 pores located below eye and 6th posterior to center of eye; 5th pore posterior to vertical through 1st postorbital pore (Fig. 8A). Preoperculomandibular pores usually 7 (including holotype); 4 on lower jaw and 3 on preopercle; 4 and 4 on left side of HUMZ 180866; last preoperculomandibular pore located posterior to lower margin of eye (Fig. 8A, C). One interorbital pore on dorsal midline between center of eyes (Fig. 8B). Occipital pores absent (Fig. 8B). Color in alcohol. Holotype (Fig. 7) with light brown head, body and vertical fins; light yellow pectoral fin and brown its ventral margin; light purple opercular region and abdomen. Head and body light brownish yellow and vertical fins whitish in HUMZ 206878. Coloration of other non-type specimens similar to holotype. Color when fresh (based on color photograph of HUMZ 152390; Fig. 5). Head, body and vertical fins uniformly reddish orange. Pectoral fin whitish, its ventral margin light brown. Opercular region and abdomen bluish black. Distribution. Off northwestern Pacific coast of eastern Hokkaido Island and off Tohoku District from Aomori to Fukushima prefectures, at depths of 500���756 m (Shinohara & Matsuura, 1998; Imamura, 1998; Hatooka, 2000, 2002, 2013; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Amaoka et al., 2011; this study). Size. The largest specimen examined during this study measured 141.3 mm SL (143.6 mm TL), exceeding the previously recorded maximum length of 135.8 mm SL (140 mm TL) (Shinohara & Matsuura, 1998; Hatooka, 2000, 2002, 2013; Amaoka et al., 2011). Remarks. Lycenchelys aurantiaca is similar to L. squamosa in having less than 100 total vertebrae, 1 interorbital pore, no occipital pores, 4 postorbital pores and no distinct spots or blotches on the body (vs. without this combination of characters in other species of Lycenchelys) (e.g., Toyoshima, 1983, 1985; Shinohara & Matsuura, 1998; Hatooka, 2000, 2002, 2013; Shinohara & Anderson, 2007; this study). Lycenchelys aurantiaca can be easily separated from L. squamosa in having lower numbers of dorsal-fin rays (82���86 vs. 88���91), anal-fin rays (68���72 vs. 73���77) and pectoral-fin rays (13���16 vs. 17���19), and total vertebrae (85���89 vs. 91���95), respectively. In addition, L. aurantiaca is distinguished from L. squamosa in having a uniformly reddish orange body when fresh and lacking scales on the pectoral fin and its base (vs. body uniformly dark brown when fresh and scales present on the pectoral fin and its base in L. squamosa) (Toyoshima, 1983, 1985; Shinohara & Matsuura, 1998; this study). Lycenchelys aurantiaca has previously been recorded only from off Tohoku District, the northwestern Pacific (Shinohara & Matsuura, 1998; Imamura, 1998; Hatooka, 2000, 2002, 2013; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Amaoka et al., 2011). Two specimens (HUMZ 192380, 192381) collected from off Kiritappu, eastern Hokkaido Island, the northwestern Pacific, and examined for this study, represent the first record of L. aurantiaca from Hokkaido waters., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 10-14, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Shinohara, G. & Matsuura, K. (1998) A new zoarcid, Lycenchelys aurantiaca, from the Pacific coast off northern Japan (Teleostei: Perciformes). Ichthyological Research, 45, 151 - 155. https: // doi. org / 10.1007 / BF 02678557","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243."]}

Lycenchelys makushok Fedorov & Andriashev, 1993 (Japanese name: Wakataka-hebigenge) (Figs. 17–20; Table 5) Lycenchelys makushok Fedorov & Andriashev, 1993: 130, figs. 1–2 (original description, type locality: off Iturup, Pacific cost of Kuril Islands); Anderson, 1994: 117 (species list); Shinohara et al., 1996: 180, fig. 2A (description); Imamura, 1997: 60 (species list); Imamura, 1998: 32, fig. 11 (brief description); Hatooka, 2000: 1032, 1590, unnumbered fig. (key to species); Hatooka, 2002: 1032, 1581, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 18 (species list); Shinohara & Anderson, 2007: 63 (key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 723 (species list); Amaoka et al., 2011: 316, unnumbered fig. (brief description); Balushkin et al., 2011: 981, 1024 (catalog of specimens); Hatooka, 2013: 1226, 2078, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: ZIN 42290, 137.1 mm SL, off Iturup, Kuril Islands, northwestern Pacific (44°39.2’N, 149°02.2’E), 800 m depth, 10 Sep. 1968, R/V Vityaz, trawl. Other specimens (40 specimens, 111.2–168.7 mm SL): HUMZ 152378, 163847, 180633, 182293, 182295, 182383–89, 182395, 182478, 182480–82, 182484–85, 182487–88, 182519, 182636–44, 182646–47, 15 males and 19 females, 111.2–147.2 mm SL, Tohoku District, northwestern Pacific; HUMZ 192427 (4), 196342, 226078, 5 males and 1 females, 133.7–168.7 mm SL, eastern Hokkaido Island, northwestern Pacific. Diagnosis. Vertebrae 24–26 + 107–113 = 132–139; head 10.0–12.9% SL; interorbital pore 1; occipital pore usually 1; postorbital pores 4; suborbital pores usually 7 + 1; preoperculomandibular pores usually 10; vomerine teeth 4–13; palatine teeth 5–12, arranged in single row; opercular flap well developed; pelvic-fin base positioned anterior to lower edge of gill opening; lateral line incomplete and positioned ventrally; scales present or absent on pectoral fin and absent on its base; body uniformly grayish brown when fresh. Description. Counts and proportional measurements in Table 5. Body strongly elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 2.1–3.6% SL (unknown for holotype). Head very short, ovoid, dorsal profile of head gently sloping from above posterior edge of eye to about above last postorbital pore. No sexual dimorphism in head recognized. Snout short, 103.3–162.8 (152.2)% of eye diameter. Eye ovoid, relatively small. Interorbital space narrow, width 10.0–43.5 (43.5)% of eye diameter. Nostril tube short, quite not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching about to about vertical through anterior margin of pupil in adult males, not reaching vertical through anterior margin of pupil in females and juveniles. Labial lobe of lower jaw weak. Teeth on jaws sharp, anterior teeth large and posterior teeth small; upper jaw with single row, sometimes having additional small teeth behind anteriormost tooth (unknown for holotype); lower jaw with 2–3 irregular rows anteriorly and single row posteriorly; vomerine teeth large and conical, arranged irregularly; palatine teeth smaller than vomerine teeth, arranged in single row. Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and triangular (Fig. 18). Pseudobranch filaments short. Lateral line deciduous, incomplete and positioned ventrally; originating posterior to 5th postorbital pore terminating its end area above about middle of anal fin. Scales small and cycloid, present on body and tail. Scales covering basal portions of dorsal and anal fins anteriorly; extent of scaled areas gradually increasing posteriorly, except at margins. Scales present or absent on pectoral axilla and basal portions of lower pectoral-fin rays (unknown for holotype). Head, nape and pectoral-fin base without scales. ....Continued next page Dorsal-fin origin above middle of pectoral fin; 1st dorsal-fin pterygiophore between neural spines of 3rd to 5th (between 3rd and 4th) vertebrae. Anal-fin origin below 20th to 22nd (20th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate or penultimate (penultimate) abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Caudal fin with 2 epurals, 4 upper hypurals and 3–5 (5) lower hypural rays. Pectoral fin moderately short, not quite reaching middle of abdomen; its posterior margin having notches. Upper end of pectoral-fin base about on lateral midline of body. Pelvic fin short; its base anterior to lower edge of gill opening; its posterior margin not reaching vertical through pectoral-fin base. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above 1st suborbital pore (Fig. 20A, B). Postorbital pores 4; distance between 1st and 2nd pores longest of those between adjacent pores (Fig. 20A, B). Suborbital pores usually 8 (8), rarely 9; 7 pores located below eye and last pore posterior to eye; when 9 pores, 8 pores below eye and 9th behind eye on left side in HUMZ 182639 and on right side in HUMZ 182293; 5th below anterior margin of pupil; last pore of those below eye located posterior to vertical posterior to margin of eye (Fig. 20A). Preoperculomandibular pores usually 10 (10), rarely 9; 5 on lower jaw, 2 at junction of lower jaw and preopercle, and 3 on preopercle; 8th and 9th pores united into 1 pore on left side of HUMZ 182479 and counted as 9; last preoperculomandibular pore located posterior to lower margin of eye (Fig. 20A, C). One interorbital pore located on dorsal midline anterior to anterior margin of pupils (Fig. 20B). Usually 1 occipital pore located on dorsal midline at middle of occiput (1); 1 additional pore present on left side of middle pore in HUMZ 182640; 2 additional pores, one on either side of middle pore in HUMZ 182642; occipital pore(s) located anterior to 3rd postorbital pore (Fig. 20B). Color in alcohol. Holotype (based on color photograph; Fig. 19) with uniformly light brown head, body and vertical fins, slightly paler pectoral fin; dark brown opercular region; purplish gray abdomen. Margin of vertical fins in non-type specimens dark brown, but coloration otherwise similar to holotype. Color when fresh (based on color photograph of HUMZ 152378; Fig. 17). Head and margin of vertical fins dark brown; body and vertical fins uniformly grayish brown; pectoral fins gray; abdomen dark purplish gray. Distribution. Off northwestern Pacific coast of the Kuril Islands, eastern Hokkaido Island and Honshu Island from Miyagi to Ibaraki prefectures, at depths of 399–1219 m (Fedorov & Andriashev, 1993; Anderson, 1994; Shinohara et al., 1996; Imamura, 1997, 1998; Hatooka, 2000, 2002, 2013; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Amaoka et al., 2011; Balushkin et al., 2011; this study). Size. Maximum length 180 mm TL (Amaoka et al., 2011; Hatooka, 2013). The largest specimen examined for this study measured 168.7 mm SL (172.2 mm TL). Remarks. Lycenchelys makushok resembles L. hippopotamus, L. melanostomias and L. rassi in having more than 100 total vertebrae, 1 interorbital pore, 1–3 occipital pores, 4 postorbital pores, a single ventral lateral line and no distinct spots or blotches on the body (vs. lacking this combination of this characters in other species of Lycenchelys) (e.g., Toyoshima, 1983, 1985; Fedorov & Andriashev, 1993; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1995; Shinohara et al., 1996; Imamura et al., 2004; Shinohara & Anderson, 2007; this study). See Remarks for L. hippopotamus for a detailed comparison of L. makushok and L. hippopotamus. Lycenchelys makushok is separable from L. melanostomias and L. rassi by the number and position of occipital pores. Lycenchelys makushok always has an occipital pore on the dorsal midline at the middle of the occiput. When 2 or 3 occipital pores are present (2 of 41 specimens observed in this study), the additional pore or two are on the lateral side or sides of the middle pore. In contrast, L. melanostomias and L. rassi usually have 2 occipital pores (1 occipital pore on right side was examined in only 1 specimen of L. melanostomias in this study), that are located on the left and right sides of the midline of the occiput. Lycenchelys makushok is further separable from L. melanostomias in having a higher number of total vertebrae (132–139, vs. 117–124, respectively), and from L. rassi in having a shorter head (head length 10.0–12.9% SL, vs. 13.4–16.4% SL, respectively).

Lycenchelys makushok Fedorov & Andriashev, 1993 (Japanese name: Wakataka-hebigenge) (Figs. 17���20; Table 5) Lycenchelys makushok Fedorov & Andriashev, 1993: 130, figs. 1���2 (original description, type locality: off Iturup, Pacific cost of Kuril Islands); Anderson, 1994: 117 (species list); Shinohara et al., 1996: 180, fig. 2A (description); Imamura, 1997: 60 (species list); Imamura, 1998: 32, fig. 11 (brief description); Hatooka, 2000: 1032, 1590, unnumbered fig. (key to species); Hatooka, 2002: 1032, 1581, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 18 (species list); Shinohara & Anderson, 2007: 63 (key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 723 (species list); Amaoka et al., 2011: 316, unnumbered fig. (brief description); Balushkin et al., 2011: 981, 1024 (catalog of specimens); Hatooka, 2013: 1226, 2078, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: ZIN 42290, 137.1 mm SL, off Iturup, Kuril Islands, northwestern Pacific (44��39.2���N, 149��02.2���E), 800 m depth, 10 Sep. 1968, R/V Vityaz, trawl. Other specimens (40 specimens, 111.2���168.7 mm SL): HUMZ 152378, 163847, 180633, 182293, 182295, 182383���89, 182395, 182478, 182480���82, 182484���85, 182487���88, 182519, 182636���44, 182646���47, 15 males and 19 females, 111.2���147.2 mm SL, Tohoku District, northwestern Pacific; HUMZ 192427 (4), 196342, 226078, 5 males and 1 females, 133.7���168.7 mm SL, eastern Hokkaido Island, northwestern Pacific. Diagnosis. Vertebrae 24���26 + 107���113 = 132���139; head 10.0���12.9% SL; interorbital pore 1; occipital pore usually 1; postorbital pores 4; suborbital pores usually 7 + 1; preoperculomandibular pores usually 10; vomerine teeth 4���13; palatine teeth 5���12, arranged in single row; opercular flap well developed; pelvic-fin base positioned anterior to lower edge of gill opening; lateral line incomplete and positioned ventrally; scales present or absent on pectoral fin and absent on its base; body uniformly grayish brown when fresh. Description. Counts and proportional measurements in Table 5. Body strongly elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 2.1���3.6% SL (unknown for holotype). Head very short, ovoid, dorsal profile of head gently sloping from above posterior edge of eye to about above last postorbital pore. No sexual dimorphism in head recognized. Snout short, 103.3���162.8 (152.2)% of eye diameter. Eye ovoid, relatively small. Interorbital space narrow, width 10.0���43.5 (43.5)% of eye diameter. Nostril tube short, quite not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching about to about vertical through anterior margin of pupil in adult males, not reaching vertical through anterior margin of pupil in females and juveniles. Labial lobe of lower jaw weak. Teeth on jaws sharp, anterior teeth large and posterior teeth small; upper jaw with single row, sometimes having additional small teeth behind anteriormost tooth (unknown for holotype); lower jaw with 2���3 irregular rows anteriorly and single row posteriorly; vomerine teeth large and conical, arranged irregularly; palatine teeth smaller than vomerine teeth, arranged in single row. Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and triangular (Fig. 18). Pseudobranch filaments short. Lateral line deciduous, incomplete and positioned ventrally; originating posterior to 5th postorbital pore terminating its end area above about middle of anal fin. Scales small and cycloid, present on body and tail. Scales covering basal portions of dorsal and anal fins anteriorly; extent of scaled areas gradually increasing posteriorly, except at margins. Scales present or absent on pectoral axilla and basal portions of lower pectoral-fin rays (unknown for holotype). Head, nape and pectoral-fin base without scales. ....Continued next page Dorsal-fin origin above middle of pectoral fin; 1st dorsal-fin pterygiophore between neural spines of 3rd to 5th (between 3rd and 4th) vertebrae. Anal-fin origin below 20th to 22nd (20th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate or penultimate (penultimate) abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Caudal fin with 2 epurals, 4 upper hypurals and 3���5 (5) lower hypural rays. Pectoral fin moderately short, not quite reaching middle of abdomen; its posterior margin having notches. Upper end of pectoral-fin base about on lateral midline of body. Pelvic fin short; its base anterior to lower edge of gill opening; its posterior margin not reaching vertical through pectoral-fin base. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above 1st suborbital pore (Fig. 20A, B). Postorbital pores 4; distance between 1st and 2nd pores longest of those between adjacent pores (Fig. 20A, B). Suborbital pores usually 8 (8), rarely 9; 7 pores located below eye and last pore posterior to eye; when 9 pores, 8 pores below eye and 9th behind eye on left side in HUMZ 182639 and on right side in HUMZ 182293; 5th below anterior margin of pupil; last pore of those below eye located posterior to vertical posterior to margin of eye (Fig. 20A). Preoperculomandibular pores usually 10 (10), rarely 9; 5 on lower jaw, 2 at junction of lower jaw and preopercle, and 3 on preopercle; 8th and 9th pores united into 1 pore on left side of HUMZ 182479 and counted as 9; last preoperculomandibular pore located posterior to lower margin of eye (Fig. 20A, C). One interorbital pore located on dorsal midline anterior to anterior margin of pupils (Fig. 20B). Usually 1 occipital pore located on dorsal midline at middle of occiput (1); 1 additional pore present on left side of middle pore in HUMZ 182640; 2 additional pores, one on either side of middle pore in HUMZ 182642; occipital pore(s) located anterior to 3rd postorbital pore (Fig. 20B). Color in alcohol. Holotype (based on color photograph; Fig. 19) with uniformly light brown head, body and vertical fins, slightly paler pectoral fin; dark brown opercular region; purplish gray abdomen. Margin of vertical fins in non-type specimens dark brown, but coloration otherwise similar to holotype. Color when fresh (based on color photograph of HUMZ 152378; Fig. 17). Head and margin of vertical fins dark brown; body and vertical fins uniformly grayish brown; pectoral fins gray; abdomen dark purplish gray. Distribution. Off northwestern Pacific coast of the Kuril Islands, eastern Hokkaido Island and Honshu Island from Miyagi to Ibaraki prefectures, at depths of 399���1219 m (Fedorov & Andriashev, 1993; Anderson, 1994; Shinohara et al., 1996; Imamura, 1997, 1998; Hatooka, 2000, 2002, 2013; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Amaoka et al., 2011; Balushkin et al., 2011; this study). Size. Maximum length 180 mm TL (Amaoka et al., 2011; Hatooka, 2013). The largest specimen examined for this study measured 168.7 mm SL (172.2 mm TL). Remarks. Lycenchelys makushok resembles L. hippopotamus, L. melanostomias and L. rassi in having more than 100 total vertebrae, 1 interorbital pore, 1���3 occipital pores, 4 postorbital pores, a single ventral lateral line and no distinct spots or blotches on the body (vs. lacking this combination of this characters in other species of Lycenchelys) (e.g., Toyoshima, 1983, 1985; Fedorov & Andriashev, 1993; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1995; Shinohara et al., 1996; Imamura et al., 2004; Shinohara & Anderson, 2007; this study). See Remarks for L. hippopotamus for a detailed comparison of L. makushok and L. hippopotamus. Lycenchelys makushok is separable from L. melanostomias and L. rassi by the number and position of occipital pores. Lycenchelys makushok always has an occipital pore on the dorsal midline at the middle of the occiput. When 2 or 3 occipital pores are present (2 of 41 specimens observed in this study), the additional pore or two are on the lateral side or sides of the middle pore. In contrast, L. melanostomias and L. rassi usually have 2 occipital pores (1 occipital pore on right side was examined in only 1 specimen of L. melanostomias in this study), that are located on the left and right sides of the midline of the occiput. Lycenchelys makushok is further separable from L. melanostomias in having a higher number of total vertebrae (132���139, vs. 117���124, respectively), and from L. rassi in having a shorter head (head length 10.0���12.9% SL, vs. 13.4���16.4% SL, respectively)., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 23-27, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Fedorov, V. V. & Andriashev, A. P. (1993) Lycenchelys makushok sp. nova (Perciformes: Zoarcidae) from the bathyal depths of the Kuril-Kamchatka Trench. Journal of Ichthyology, 33, 130 - 135.","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Shinohara, G., Endo, H. & Matsuura, K. (1996) Deep-water Fishes collection from the Pacific coast of northern Honshu, Japan. Memories of the National Science Museum, Tokyo, 29, 153 - 185.","Imamura, H. (1997) Perciform fish fauna from off Pacific of Tohoku. Researches on Basic Fishes in Tohoku District, 17, 55 - 68. [in Japanese]","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243.","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133.","Imamura, H., Machida, Y. & Ohta, S. (2004) Lycenchelys melanostomias Toyoshima, 1983, a senior synonym of L. brevimaxillaris Toyoshima, 1985 (Teleostei: Zoarcidae). Ichthyological Research, 51, 84 - 88. https: // doi. org / 10.1007 / s 10228 - 003 - 0194 - 2"]}

Lycenchelys squamosa Toyoshima, 1983 (Japanese name: Uroko-hebigenge) (Figs. 38���43; Table 10) Lycenchelys squamosus Toyoshima, 1983: 145, figs. 20���22, table 10, pl. 93 (original description, type locality: off Miyagi Prefecture, Pacific coast of Honshu Island, Japan); Toyoshima, 1984: 293, pl. 274-E (brief description); Toyoshma, 1985: 156, figs. 6���7, 12���13, 31, table 1 (description); Hatooka, 1993: 901, unnumbered fig. (key to species); Amaoka et al., 1995: 241, pl. 405 (brief description); Koyanagi, 1997: 538, fig. 5 (brief description); Imamura, 1998: 32, fig. 13 (brief description); Zama, 2001: 86, 133 (species list). Lycenchelys squamosa: Anderson, 1994: 113, 118 (species list); Shinohara et al., 1996: 182 (species list); Imamura, 1997: 60 (species list); Shinohara & Matsuura, 1998: table 1 (comparison with Lycenchelys aurantiaca); Hatooka, 2000: 1033, unnumbered fig. (key to species); Hatooka, 2002: 1033, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 20 (species list); Shiogaki et al., 2004: 71 (species list); Shinohara & Anderson, 2007: 64 (key to species); Kitagawa et al., 2008: 96, unnumbered fig. (brief description); Shinohara et al., 2009: 724 (species list); Amaoka et al., 2011: 318, unnumbered fig. (brief description); Balushkin et al., 2011: 983 (catalog of specimens); Hatooka, 2013: 1227, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: HUMZ 78464, male, 246.1 mm SL, off Miyagi Prefecture, Tohoku District, northwestern Pacific (37��49.7���N, 142��23.7���E to 37��55.4���N, 142��24.5���E), 985���1005 m depth, 8 Oct. 1978. Paratypes (2 specimens, 193.0��� 222.5 mm SL, both from Tohoku District, northwestern Pacific): HUMZ 72563, 1 female, 193.0 mm SL, off Fukushima Prefecture (37��09.4���N, 141��56E), 900���920 m depth, 19 Jan. 1978; HUMZ 78390, 1 female, 222.4 mm SL, off Aomori Prefecture (40��47.6���N, 142��16.7���E to 40��41.5���N, 142��19.2���E), 920���948 m depth, 11 Sep. 1978. Other specimens (6 specimens, 203.0��� 253.1 mm SL): HUMZ 157660, 157669, 181894���95, 192737, 2 males and 3 females, 203.0��� 253.1 mm SL, Tohoku District, northwestern Pacific; HUMZ 177036, 1 female, 241.8 mm SL, eastern Hokkaido Island, northwestern Pacific. Diagnosis. Vertebrae 20 + 71���75 = 91���95; head length 16.3���19.9% SL; interorbital pore 1; occipital pores absent; postorbital pores 4; suborbital pores 5 + 1 (rarely 5 + 2); preoperculomandibular pores 8; vomerine teeth 7���17; palatine teeth 5���25, arranged in 1 or 1���2 rows; opercular flap well-developed; pelvic-fin base positioned below lower edge of gill opening; lateral line incomplete and positioned ventrally; scales present on pectoral fin and its base; body uniformly dark brown when fresh. Description. Counts and proportional measurements in Table 10. Body elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 5.4���6.5 (5.7)% SL. Head moderately long, ovoid; dorsal profile of head gently sloping to dorsal-fin origin. Snout short, 125.8���173.2 (159.3)% of eye diameter. Eye ovoid, relatively large. Interorbital space narrow, width 15.6���35.6 (27.4)% of eye diameter. Nostril tube short, not reaching upper lip when depressed. Mouth subterminal; lower jaw extending slightly beyond upper jaw (holotype) (Fig. 39A), or upper jaw extending slightly (paratypes and non-type specimens, all females) or significantly (non-type males) beyond lower jaw (Fig. 39B, C). Posterior end of upper jaw reaching to or slightly beyond vertical through suborbital pore behind eye in males, reaching below posterior margin of eye in females. Labial lobe of lower jaw weak. Teeth on jaws, vomer and palatine small and conical; upper jaw with 2���3 rows anteriorly and single row posteriorly; lower jaw with 2���4 irregular rows anteriorly and single row posteriorly; vomerine teeth irregularly arranged; all palatine teeth in single row, or in 2 rows anteriorly and single row posteriorly (single row). Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and thin (Fig. 40). Pseudobranch filaments short. Lateral line deciduous, positioned ventrally and incomplete; its origin posterior to last postorbital pore and terminating above middle of anal fin. Scales small and cycloid, present on nape, body, pectoral axilla, about basal half of pectoral fin, pectoral-fin base, tail and vertical fins except at margin. Head without scales. ....Continued next page Dorsal-fin origin near vertical through posterior edge of opercular flap; 1st dorsal-fin pterygiophore between neural spines of 2nd and 3rd vertebrae. Anal-fin origin below 18th to 20th (19th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate abdominal vertebra or hemal spine of 1st caudal vertebra (posterior to parapophysis of ultimate abdominal vertebra). Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 4th and 5th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Caudal fin with 1���2 (2) epural, 4 upper hypural and 3���4 (4) lower hypural rays. Pectoral fin relatively long, reaching to about mid-portion of abdomen; its posterior margin rounded. Upper end of pectoral-fin base on about lateral midline of body. Pelvic fin short; its posterior margin slightly beyond lower edge of gill opening. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above and slightly anterior to vertical through 1st suborbital pore (Fig. 42A, B). Postorbital pores 4; distance between 1st and 2nd pores longest of those between adjacent pores; 3rd pore slightly higher than others (Fig. 42A, B). Suborbital pores usually 6 (including holotype), rarely 7; 5 pores below eye and last pore behind midline of eye, except for another pore just above 5th pore on left side in HUMZ 181895 and right side in HUMZ 157669; 5th pore behind vertical through 1st postorbital pore (Fig. 42A). Preoperculomandibular pores 8; 4 on lower jaw and 4 on preopercle; last preoperculomandibular pore posterior to lower margin of eye (Fig. 42A, C). One interorbital pore on dorsal midline above middle of eyes (Fig. 42B). Occipital pores absent (Fig. 42B). Color in alcohol. Holotype (Fig. 41) with light purplish brown head, body and vertical fins, color fading somewhat on body, right side of tail and vertical fins. Pectoral fin slightly lighter than body color. Head, body, vertical and pectoral fins dark brown in paratypes. Head and pectoral fin dark brown, body and vertical fins grayish brown, margins of dorsal and anal fins blackish in other specimens. Color when fresh (based on color photograph of HUMZ 192737; Fig. 38). Head and pectoral fin blackish. Body, vertical fins and pectoral-fin base uniformly dark brown. Margins of dorsal and anal fins black. Distribution. The Okhotsk Sea, off northwestern Pacific coast of eastern Hokkaido Island and in the northwestern Pacific from Aomori to Ibaraki prefectures, at depths of 310���1340 m (Toyoshima, 1983, 1984, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1994; Amaoka et al., 1995, 2011; Shinohara et al., 1996; Imamura, 1997, 1998; Koyanagi, 1997; Shinohara & Matsuura, 1998; Anderson & Fedorov, 2004; Shiogaki et al., 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Balushkin et al., 2011; this study). Size. The largest specimen examined during this study measured 253.1 mm SL (261.3 mm TL), about equal to the previously recorded maximum length (252 mm SL, 260 mm TL) (Toyoshima, 1983, 1984, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1994; Amaoka et al., 1995, 2011; Koyanagi, 1997; Shinohara & Matsuura, 1998; Shinohara & Anderson, 2007). Remarks. Lycenchelys squamosa resembles L. aurantiaca in having less than 100 total vertebrae, 1 interorbital pore, no occipital pores, 4 postorbital pores and no distinct spots or blotches on the body (vs. lacking this combination of characters in other species of Lycenchelys) (e.g., Toyoshima, 1983, 1985; Shinohara & Matsuura, 1998; Hatooka, 2000, 2002, 2013; Shinohara & Anderson, 2007; this study). See Remarks under L. aurantiaca for a detailed comparison of L. squamosa with L. aurantiaca. In Lycenchelys squamosa, the upper jaw extends forward slightly beyond the lower jaw in females (including paratypes) (Fig. 39B). In males however, the lower jaw extends slightly (holotype, 246.1 mm SL) (Fig. 39A) or considerably beyond the lower jaw (2 non-types, 217.0 and 253.1 mm SL) (Fig. 39C). According to Toyoshima (1983, 1985), the lower jaw of the holotype (male) protrudes slightly beyond the upper jaw, and the lower jaw is nearly equal to or slightly included in the upper jaw of paratypes (females). He did not mention the variation in the jaw protrusion of males because he was able to examine only 1 male specimen (= holotype). Subsequent studies have not described the variation in males of the species even though the extension of the lower jaw in males of the species as presented by Toyoshima (1985) has been repeated in some studies (e.g., Amaoka et al., 1995, 2011). This study found that the variation in males lies in the length of the upper jaw, not of the lower jaw, although the ratio of the lower jaw length seems to increase with growth (Fig. 43B). In the 3 males examined, the 246.1 mm SL specimen (holotype) with the lower jaw protrusion has a shorter upper jaw, while the largest (253.1 mm SL) and smallest (217.0 mm SL) specimens with the upper jaw protrusion have a longer upper jaw (Fig. 43A). Therefore, the length of the upper jaw may be due to intraspecific variation not associated with change in growth in males of L. squamosa. Further specimens are required to resolve the differences., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 45-50, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Toyoshima, M. (1984) Family Zoarcidae. Eelpouts. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 291 - 295. [in Japanese]","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Amaoka, K., Nakaya, K. & Yabe, M. (1995) The fishes of northern Japan. Kita-Nihon Kaiyo Center, Sapporo, 391 pp. [in Japanese]","Koyanagi, M. (1997) Zoarcidae. In: Okamura, O. & Amaoka, K. (Eds.), Sea fishes of Japan, Yama-Kei, Tokyo, pp. 536 - 538. [in Japanese]","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Zama, A. (2001) Fish fauna of Miyagi Prefecture, Japan. Sanwa Publisher, Ishinomaki, 153 pp. [in Japanese]","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Shinohara, G., Endo, H. & Matsuura, K. (1996) Deep-water Fishes collection from the Pacific coast of northern Honshu, Japan. Memories of the National Science Museum, Tokyo, 29, 153 - 185.","Imamura, H. (1997) Perciform fish fauna from off Pacific of Tohoku. Researches on Basic Fishes in Tohoku District, 17, 55 - 68. [in Japanese]","Shinohara, G. & Matsuura, K. (1998) A new zoarcid, Lycenchelys aurantiaca, from the Pacific coast off northern Japan (Teleostei: Perciformes). Ichthyological Research, 45, 151 - 155. https: // doi. org / 10.1007 / BF 02678557","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shiogaki, M., Ishito, Y., Nomura, Y. & Sugimoto, T. (2004) Revised catalog of the fishes collected from the waters of Aomori Prefecture. Bulletin of Aomori Prefectural Fisheries Research Center, (4), 39 - 80. [in Japanese with English abstract]","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243."]}

Lycenchelys melanostomias Toyoshima, 1983 (Japanese name: Ohotsuku-hebigenge) (Figs. 21���25; Table 6) Lycenchelys sp.: Shiogaki, 1982: 23 (species list). Lycenchelys melanostomias Toyoshima, 1983: 271, 333, figs. 25���27, pl. 157 (original description, type locality: southern Okhotsk Sea, Hokkaido Island, Japan); Toyoshima, 1984: 293, pl. 274-A (brief description); Toyoshima, 1985: 170, figs. 6���7, 26���27, 31, tables 1, 4 (description); Hatooka, 1993: 902, unnumbered fig. (key to species); Anderson, 1994: 113, 117 (species list); Amaoka et al., 1995: 240, pl. 402 (brief description); Koyanagi, 1997: 538, fig. 3 (brief description); Hatooka, 2000: 1032, unnumbered fig. (key to species); Hatooka, 2002: 1032, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 18 (species list); Imamura et al., 2004: 84, figs. 1���3, table 1 (synonymy of Lycenchelys melanostomias and Lycenchelys brevimaxillaris); Shiogaki et al., 2004: 71 (species list); Imamura et al., 2005: 1, figs. 1���3, table 1 (synonymy of Lycenchelys melanostomias and Lycenchelys brevimaxillaris); Shinohara & Anderson, 2007: 64 (key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 724 (species list); Amaoka et al., 2011: 315, unnumbered fig. (brief description); Balushkin et al., 2011: 1026 (species list); Hatooka, 2013: 1226, 2078, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Lycenchelys brevimaxillaris: Toyoshima, 1985: 174, figs. 6���7, 29���30, 31, table 1 (original description, type locality: off Aomori Prefecture, Pacific coast of Honshu, Japan); Hatooka, 1993: 902, unnumbered fig. (key to species); Anderson, 1994: 117 (species list); Imamura, 1998: 31, fig. 9 (brief description); Hatooka, 2000: 1033, unnumbered fig. (key to species); Hatooka, 2002: 1033, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 16 (species list); Imamura et al., 2004: 84, figs. 1���3, table 1 (synonymy of Lycenchelys melanostomias and Lycenchelys brevimaxillaris); Shiogaki et al., 2004: 71 (species list); Imamura et al., 2005: 1, figs. 1���3, table 1 (synonymy of Lycenchelys melanostomias and Lycenchelys brevimaxillaris). Materials examined Holotype: HUMZ 77572, male, 182.3 mm SL, Kitami-Yamato Bank, Okhotsk Sea (44��19.5���N, 145��01���E), 915���925 m depth, 11 Oct. 1978; Other specimens (41 specimens, 119.3���227.6 mm SL): HUMZ 189817 (holotype of Lycenchelys brevimaxillaris), female, 185.8 mm SL, off Aomori Prefecture, Tohoku District, northwestern Pacific (41��13���N, 141��44���E), 690���750 m depth, 18 Jan. 1982; HUMZ 152380, 152384, 152404���05, 152411, 157545, 177188, 178572���75, 182565, 192724, 192804, 4 males and 10 females, 124.0��� 195.1 mm SL, Tohoku District, northwestern Pacific; HUMZ 177263, 192453, 205157, 228057, 228067���68, 228070, 228073, 228089, 5 males and 4 females, 119.3���193.2 mm SL, eastern Hokkaido Island, northwestern Pacific; HUMZ 120346, 121158, 121161, 121461, 124054���55, 124116, 126101, 126215���16, 126219���22, 126230, 126359, 126361, 10 males and 7 females, 120.6���227.6 mm SL, northeastern Hokkaido Island, Okhotsk Sea. Diagnosis. Vertebrae 22���25 + 93���102 = 117���124; head length 11.6���15.0% SL; interorbital pore 1; occipital pores usually 2; postorbital pores usually 4; suborbital pores 6���7 + 2���3; preoperculomandibular pores usually 9; vomerine teeth 3���10; palatine teeth 2���11, usually arranged in single row (sometimes 1���2 rows); opercular flap well developed; pelvic-fin base positioned posterior to lower edge of gill opening; lateral line complete and positioned ventrally; scales absent on pectoral fin and its base; body uniformly grayish-brown when fresh. Description. Counts and proportional measurements in Table 6. ...Continued next page Body very elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 3.8���7.0 (5.0)% SL. Head short, ovoid; dorsal profile of head sloping extremely gently from posterior edge of eye to above opercular flap. Head of males slightly longer than of females in adults. Snout short, 74.4���131.0% of eye diameter (eye damaged in holotype). Eye ovoid, moderately large. Interorbital space narrow, width 12.7���33.8% of eye diameter (eye damaged in holotype). Nostril tube short, not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching vertical through posterior part of eye in adult males, reaching vertical through anterior margin of pupil in females and juveniles. Labial lobe of lower jaw tending to be more developed in large males than in females and juveniles (labial lobe of lower jaw damaged in holotype). Teeth on jaws sharp; upper jaw with 2���3 rows anteriorly, 1 or 1���2 rows posteriorly (1); anteriormost teeth larger than other teeth; lower jaw with 2���5 irregular rows anteriorly, 1 or 1���2 rows posteriorly (1); vomerine and palatine teeth small and conical; vomerine teeth irregularly arranged; palatine teeth usually in single row, but sometimes 1���2 rows (no data for holotype). Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and triangular (Fig. 23). Pseudobranch filaments short. Lateral line deciduous, complete and positioned ventrally; originating posterior to last postorbital pore and terminating on tail. Scales small and cycloid, present on body and tail, except for area around pelvic fin. Scales covering basal portions of dorsal and anal fins anteriorly; extent of scaled areas gradually increasing posteriorly, except at margins. Head, nape, pectoral axilla, pectoral fin, pectoral-fin base and area around pelvic fin without scales. Dorsal-fin origin posterior to vertical through pectoral-fin base; 1st dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 2nd and 3rd) vertebrae. Anal-fin origin below 17th to 20th (18th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate or penultimate (penultimate) abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 2nd and 3rd) preural vertebrae. Last analfin pterygiophore between hemal spines of 3rd to 5th (between 3rd and 4th) preural vertebrae. Caudal fin with 1���3 (2) epural, 3���4 (4) upper hypural and 3���4 (3) lower hypural rays. Pectoral fin moderately short, not quite reaching middle of abdomen; its posterior margin rounded dorsally and having notches ventrally. Upper end of pectoral-fin base about on lateral midline of body. Pelvic fin short; its base posterior to lower edge of gill opening; its posterior margin reaching about vertical through pectoral-fin base. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above line of 1st suborbital pore (Fig. 22A, B). Postorbital pores usually 4, rarely 5 (5); when 4, distance between 1st and 2nd pores longest of those between adjacent pores; when 5, 1 additional pore present between 1st and 2nd pores (Fig. 22A, B). Suborbital pores 8���10 (unknown for holotype), 6 or 7 pores located below eye and remaining 2 or 3 pores on ascending part of suborbital canal behind eye; 5th pore below posterior margin of pupil; last pore of those below eye located posterior to vertical through posterior margin of eye (Fig. 22A). Preoperculomandibular pores usually 9 (9), rarely 8; 4 on lower jaw, 2 at junction of lower jaw and preopercle, and 3 on preopercle; 2 pores at junction of lower jaw and preopercle united into 1 pore in some specimens and preoperculomandibular pore series counted as 8; 2nd and 3rd pores united into 1 pore on right side of HUMZ 126361 and series counted as 8; last preoperculomandibular pore located posterior to lower part of eye (Fig. 22A, C). One interorbital pore located on dorsal midline anterior to middle of eyes (Fig. 22B). Occipital pores usually 2 (2), usually positioned on either side of dorsal midline; only 1 pore on right side in HUMZ 182565; occipital pore(s) located anterior to 3rd postorbital pore (Fig. 22B). Two additional unnamed pores present in interorbital space on either side in line with postorbital pores, behind posterior to posterior margin of pupil only in HUMZ 178574 (Fig. 22D). Color in alcohol. Color of holotype (Fig. 24) unknown owing to head, body, and fins lacking skin. Long preserved holotype of L. brevimaxillaris with brownish head, pectoral fin and vertical fins, uniformly grayish brown body and gray abdomen. Other recently preserved specimens with dark brown head, pectoral fin and vertical fins, uniformly pale brown body and blackish abdomen. Color when fresh (based on color photograph of HUMZ 124054; Fig. 21). Head and pectoral fin black; body and vertical fins uniformly grayish brown; margin of vertical fins slightly darker and abdomen pale purplish. Distribution. Southern Okhotsk Sea, off northwestern Pacific coast of eastern Hokkaido Island and in the northwestern Pacific from Aomori to Ibaraki prefectures, at depths of 425���1440 m (Shiogaki, 1982; Toyoshima, 1983, 1984, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1994; Amaoka et al., 1995, 2011; Koyanagi, 1997; Imamura, 1998; Anderson & Fedorov, 2004; Imamura et al., 2004, 2005; Shiogaki et al., 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; this study). Size. The largest specimen examined during this study measured 227.6 mm SL (230.8 mm TL), about equal to the previously recorded maximum length of 23 cm TL (Hatooka, 2013). Remarks. Lycenchelys melanostomias is similar to L. hippopotamus, L. makushok and L. rassi in having more than 100 total vertebrae, 1 interorbital pore, 1���2 occipital pores, 4���5 postorbital pores, a single lateral line positioned ventrally and no distinct spots or blotches on the body (vs. lacking this combination of characters in other species of Lycenchelys) (e.g., Toyoshima, 1983, 1985; Fedorov & Andriashev, 1993; Anderson, 1995; Imamura et al., 2004; Shinohara & Anderson, 2007; this study). See Remarks under accounts of L. hippopotamus, L. makushok and L. rassi for detailed comparisons of L. melanostomias with each. Two additional pores behind the posterior margin of the pupil were found in 1 specimen of L. melanostomias (Fig. 22D). These pores have not been previously described for L. melanostomias, although similar pores are known in Lycenchelys parini Fedorov, 1995 (Fig. 25). Lycenchelys melanostomias also resembles L. parini in having similar counts, proportional measurements and arrangements of head pores (e.g., dorsal-fin rays 111���120 vs. 118, and anal-fin rays 98���107 vs. 105, head length 11.6���15.0 vs. 12.2% SL, postorbital pores usually 4 vs. 4 and suborbital pores 8���10 vs. 9 in L. melanostomias respectively; Table 6) (Fedorov, 1995a; Imamura et al., 2004, 2005; this study). However, L. melanostomias is easily separable from L. parini in having the lateral line positioned ventrally rather than midlaterally on the side (Fedorov, 1995a)., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 27-32, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Shiogaki, M. (1982) A catalogue of the fishes collected from the waters of Aomori Prefecture, Japan. Bulletin of the Fisheries Experimental Station of Aomori Prefecture, 1982, 1 - 36. [in Japanese]","Toyoshima, M. (1984) Family Zoarcidae. Eelpouts. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 291 - 295. [in Japanese]","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243.","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Amaoka, K., Nakaya, K. & Yabe, M. (1995) The fishes of northern Japan. Kita-Nihon Kaiyo Center, Sapporo, 391 pp. [in Japanese]","Koyanagi, M. (1997) Zoarcidae. In: Okamura, O. & Amaoka, K. (Eds.), Sea fishes of Japan, Yama-Kei, Tokyo, pp. 536 - 538. [in Japanese]","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Imamura, H., Machida, Y. & Ohta, S. (2004) Lycenchelys melanostomias Toyoshima, 1983, a senior synonym of L. brevimaxillaris Toyoshima, 1985 (Teleostei: Zoarcidae). Ichthyological Research, 51, 84 - 88. https: // doi. org / 10.1007 / s 10228 - 003 - 0194 - 2","Shiogaki, M., Ishito, Y., Nomura, Y. & Sugimoto, T. (2004) Revised catalog of the fishes collected from the waters of Aomori Prefecture. Bulletin of Aomori Prefectural Fisheries Research Center, (4), 39 - 80. [in Japanese with English abstract]","Imamura, H., Shinohara, G. & Shiogaki, M. (2005) Rediscovery of the type specimens of Lycenchelys brevimaxillaris (Actinopterygii: Perciformes: Zoarcidae) with taxonomic notes on the species. Species Diversity, 10, 1 - 6. https: // doi. org / 10.12782 / specdiv. 10.1","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Fedorov, V. V. & Andriashev, A. P. (1993) Lycenchelys makushok sp. nova (Perciformes: Zoarcidae) from the bathyal depths of the Kuril-Kamchatka Trench. Journal of Ichthyology, 33, 130 - 135.","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133.","Fedorov, V. V. (1995 a) Lycenchelys parini sp. nova (Perciformes: Zoarcidae) from the bathyal region of the Kuril-Kamchatka trench. Journal of Ichthyology, 35, 130 - 134."]}

Lycenchelys rassi Andriashev, 1955 (Japanese name: Rasu-hebigenge) (Figs. 26���31; Table 7) Lycenchelys rassi Andriashev, 1955: 359, figs. 2, 5, 6 (original description, type locality: east coast of Sakhalin Island, Sea of Okhotsk); Andriashev, 1958:172 (description); Peden, 1973: 115, fig. 1, table 1 (description); Toyoshima, 1983: 269, 332, pl. 155 (description); Toyoshima, 1984: 293, pl. 274-B (brief description); Toyoshma, 1985: 149, 173, figs. 6���7, 28, 31, tables 1, 4 (description); Hatooka, 1993: 902, unnumbered fig. (key to species); Anderson, 1994: 113, 117 (species list); Amaoka et al., 1995: 240, pl. 403 (brief description); Anderson, 1995: 98, fig. 15 (description); Koyanagi, 1997: 538, fig. 4 (brief description); Hatooka, 2000: 1033, unnumbered fig. (keys to species); Mecklenburg et al., 2002: 703, unnumbered figs. (brief description); Hatooka, 2002: 1033, unnumbered fig. (keys to species); Anderson & Fedorov, 2004: 19 (species list); Shinohara & Anderson, 2007: 64 (key to species); Amaoka et al., 2011: 316, unnumbered fig. (brief description); Balushkin et al., 2011: 981, 1024 (catalog of specimens); Hatooka, 2013: 1226, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: ZIN 32962, female, 190.6 mm SL, off eastern Sakhalin Island, Okhotsk Sea (54��28���N, 145��21.6���E), 1500 m depth, R/V Vityaz. Other specimens (57 specimens): HUMZ 77747, 119939, 120328, 120330, 120347 ���49, 121156, 121451, 121458 ���59, 121464, 126117���22, 126185, 126187���93, 126195���96, 126198, 126200���05, 126211, 126252, 126257, 126367���75, 126377���86, 28 males and 29 females, 93.5���239.7 mm SL, northeastern Hokkaido Island, Okhotsk Sea. Diagnosis. Vertebrae 23���25 + 98���109 = 122���134; head length 13.3���16.4% SL; interorbital pore 1; occipital pores 2; postorbital pores usually 4; suborbital pores usually 7 + 1; preoperculomandibular pores usually 8; vomerine teeth 3���11; palatine teeth 3���10, usually arranged in single row (sometimes 1���2 rows); opercular flap absent; pelvic-fin base positioned below lower edge of gill opening; lateral line complete and positioned ventrally; scales present or absent on pectoral fin and absent on its base; body uniformly grayish brown when fresh. Description. Counts and proportional measurements in Table 7. Body very elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 2.5���4.8% SL (unknown for holotype). Head moderately long, ovoid; dorsal profile of head sloping extremely gently from posterior edge of eye to above about last postorbital pore. Cheek swollen in some males. Head in adults slightly longer in males than females. Snout short, 90.0���176.1 (141.3)% of eye diameter. Eye ovoid, moderately large. Interorbital space narrow, width 17.9���36.4 (19.0)% of eye diameter. Nostril tube long, reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching vertical through middle to posterior part of eye in adult males, reaching vertical through anterior margin to middle of eye in females and juveniles (middle of eye). Labial lobe of lower jaw developed. Teeth on jaws sharp; upper jaw usually with 2 rows, rarely 3 rows (unknown for holotype) anteriorly, single row posteriorly; anteriormost teeth larger than other teeth; lower jaw with 2���5 irregular rows anteriorly, 1 or 1���2 rows posteriorly (unknown for holotype); vomerine and palatine teeth small and conical; vomerine teeth irregularly arranged; palatine teeth usually in single row, sometimes 1���2 rows (no data for holotype). Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap absent (Fig. 31A, B). Gill rakers short; those on upper limb triangular, many triangular and some blunt on lower limb (Fig. 27). Pseudobranch filaments short. Lateral line deciduous, complete and positioned ventrally; originating posterior to last postorbital pore and terminating on tail. Scales small and cycloid, present on body and tail, except head, nape, pectoral-fin base and area around pelvic fin. Scales covering basal portions of dorsal and anal fins anteriorly; extent of scaled areas gradually increasing posteriorly, except at margins. Scales present or absent on pectoral axilla and basal portions of lower pectoral-fin rays (absent). ....Continued next page Dorsal-fin origin above middle of pectoral fin; 1st dorsal-fin pterygiophore between neural spines of 4th to 6th (between 5th and 6th) vertebrae. Anal-fin origin below 17th to 20th (19th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate or penultimate (penultimate) abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 3rd to 5th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 5th (between 2nd and 3rd) preural vertebrae. Caudal fin with 1���2 (2) epural, 4���5 (4) upper hypural and 3���4 (3) lower hypural rays. Pectoral fin moderately short, reaching to about middle of abdomen; its posterior margin notched. Upper end of pectoral-fin base about on lateral midline of body. Pelvic fin short; its base at about lower edge of gill opening; its posterior margin not quite reaching pectoral-fin base. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above 1st suborbital pore (Fig. 29A, B). Postorbital pores usually 4 (4), rarely 3; when 4, distance between 2nd and 3rd pores longest of those between adjacent pores; when 3, 2nd pore absent (Fig. 29A, B). Suborbital pores usually 8 (8), rarely 7 or 9; when 8, 7 pores below eye and last behind eye; when 9, 8 pores below eye and remaining pore behind eye, or 7 pores below eye and last 2 pores on ascending part of suborbital canal behind eye; 6th and 7th pores united into 1 pore on right side of HUMZ 126384 and counted as 7; 4th pore below vertical through anterior margin of eye; last pore of those below eye posterior to posterior margin of eye (Fig. 29A). Preoperculomandibular pores usually 8 (8), rarely 7 or 9; 4 on lower jaw, 1 at junction of lower jaw and preopercle, and 3 on preopercle; pore at junction of lower jaw and preopercle separated into 2 pores in some specimens and counted as 9; 1st and 2nd pores united into 1 pore on right side of HUMZ 126382, 126378 and 126211, 3rd and 4th pores united into 1 pore on left side of HUMZ 126205, and 4th and 5th pores united into 1 pore on left side of HUMZ 126382 and counted as 7; last preoperculomandibular pore posterior to lower margin of eye (Fig. 29A, C). One interorbital pore on dorsal midline anterior to middle of eyes (Fig. 29B). Occipital pores 2, positioned on either side of dorsal midline; occipital pores located anterior to 3rd postorbital pore (Fig. 29B). Color in alcohol. Holotype (based on color photograph; Fig. 28) with uniformly brown head, body and vertical fins, slightly darker pectoral fin and margin of vertical fins, dark brown opercular region and purplish gray abdomen. Most other specimens similar to holotype, and some uniformly paler than holotype. Color when fresh (based on color photograph of HUMZ 119939; Fig. 26). Head purplish brown; body and vertical fins uniformly grayish brown; margin of vertical fins dark brown; opercular region and pectoral fin blackish; abdomen purplish. Distribution. Okhotsk Sea to the eastern Bering Sea, at depths of 895���1805 m (Andriashev, 1955, 1958; Peden, 1973; Toyoshima, 1983, 1984, 1985; Anderson, 1994, 1995; Amaoka et al., 1995, 2011; Koyanagi, 1997; Hatooka, 2000, 2002, 2013; Mecklenburg et al., 2002; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Balushkin et al., 2011; this study). Size. The largest specimen examined during this study measured 239.7 mm SL (242.9 mm TL), slightly exceeding the previously recorded maximum length of 24 cm TL (Amaoka, 2011; Hatooka, 2013). Remarks. Lycenchelys rassi resembles L. hippopotamus, L. makushok and L. melanostomias in having more than 100 total vertebrae, 1 interorbital pore, 2 occipital pores, 3���4 postorbital pores, a single ventrally positioned lateral line and no distinct spots or blotches on the body (vs. lacking this combination of characters in other species of Lycenchelys) (e.g., Andriashev, 1955; Toyoshima, 1983, 1985; Fedorov & Andriashev, 1993; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1995; this study). See Remarks under accounts for L. hippopotamus and L. makushok for detailed comparisons of L. melanostomias with those two species. Although L. rassi has been previously compared with L. melanostomias (Toyoshima, 1983, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1995; Shinohara & Anderson, 2007), this study found most characters considered to be useful for separating them as not valid for doing so. For example, Anderson (1995) redescribed L. rassi based on 6 specimens and claimed that L. rassi is readily separable from L. melanostomias by the following 5 characters: 3���4 postorbital, 7 + 1 or 8 + 1 suborbital and 8 preoperculomandibular pores (vs. 5, 7 + 2 and 9 pores in L. melanostomias), dorsal-fin origin associated with 5th vertebra (vs. 2nd), and stomach pale (vs. black). However, this study found the interspecific variation in the 5 characters mentioned above to be 3���4 postorbital, 7���8 + 1���2 suborbital and 7���9 preoperculomandibular pores, and 1st dorsal-fin pterygiophore located between neural spines of 4th to 6th vertebrae in L. rassi vs. 4���5, 6���7 + 2���3 and 8���9 pores, and 2nd to 5th vertebrae respctively in L. melanostomias, and the stomach in some alcohol preserved specimens of L. melanostomias is pale. See also Imamura et al. (2004) for color of stomach in the holotype of L. melanostomias. In addition, the number of total vertebrae and the length of the head, which were described as being different between L. rassi and L. melanostomias in recently published papers (Shinohara & Anderson, 2007; Hatooka, 2013), are also insufficient to clearly separate them (total vertebrae 122���134 vs. 117���124 and head length 13.3���16.4% SL vs. 11.6���15.0% SL, respectively) (Anderson, 1995; Imamura et al., 2004, 2005; this study). Therefore, these two species cannot always be separated using previously recognized diagnostic characters. This study distinguishes L. rassi from L. melanostomias by the presence or absence of the opercular flap. All specimens of L. rassi observed in this study lack the opercular flap (Fig. 31A, B), while it is present in all specimens of L. melanostomias examined (Fig. 31C, D). The absence of the opercular flap is rare in species of Lycenchelys, and Andriashev (1955) and Toyoshima (1985) started that the absence of the opercular flap is one of the characters differing between L. rassi and its congeners [vs. L. hippopotamus in Andriashev (1955) and vs. L. brevimaxillaris in Toyoshima (1985)]. Until now, the condition of the opercular flap has not been compared between L. rassi and L. melanostomias. This study concludes that the absence of the opercular flap is very valuable for separating L. rassi from L. melanostomias because it is easy to observe and there is no intraspecific variation in both species. Lycenchelys rassi is further distinguished from L. melanostomias by their different arrangements of the postorbital pores. In specimens having 4 postorbital pores, the typical condition in both species, the distance between the 2nd and 3rd pores is much greater in L. rassi (11.8���18.2% HL) than in L. melanostomias (4.1���9.2% HL) (Fig. 30). This difference is due to the positions of the 2nd pores; the 2nd pore emanates from the sphenotic and 3rd pore from the pterotic in L. rassi, while both pores emanates from the pterotic in L. melanostomias., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 32-35, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Andriashev, A. P. (1955) A review of the fishes of the genus Lycenchelys Gill (Pisces, Zoarcidae) and related forms in the seas of the USSR and adjacent waters. Trudy Instituta Zoologii Akademia Nauk, Azerbaidzhanskoi SSR, 18, 349 - 384. [Translated from Russian in Selected taxonomic papers on northern marine fishes, OTS 61 - 31030]","Andriashev, A. P. (1958) An addition to the review of the fishes of the genus Lycenchelys Gill with descriptions of three new species from the Kuril-Kamchatka Trench. Voprosy Ikhtiologii, 8, 611 - 618. [Translated from Russian in Selected taxonomic papers on northern marine fishes, OTS 61 - 31030]","Peden, A. E. (1973) Records of eelpouts of the genus Lycenchelys and Embryx from the northeastern Pacific Ocean. Syesis, 6, 115 - 120.","Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Toyoshima, M. (1984) Family Zoarcidae. Eelpouts. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 291 - 295. [in Japanese]","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Amaoka, K., Nakaya, K. & Yabe, M. (1995) The fishes of northern Japan. Kita-Nihon Kaiyo Center, Sapporo, 391 pp. [in Japanese]","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133.","Koyanagi, M. (1997) Zoarcidae. In: Okamura, O. & Amaoka, K. (Eds.), Sea fishes of Japan, Yama-Kei, Tokyo, pp. 536 - 538. [in Japanese]","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Mecklenburg, G. W., Mecklenburg, T. A. & Thorsteinson, L. K. (2002) Fishes of Alaska. American Fisheries Society, Bethesda, xxxvii + 1037 pp.","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243.","Fedorov, V. V. & Andriashev, A. P. (1993) Lycenchelys makushok sp. nova (Perciformes: Zoarcidae) from the bathyal depths of the Kuril-Kamchatka Trench. Journal of Ichthyology, 33, 130 - 135.","Imamura, H., Machida, Y. & Ohta, S. (2004) Lycenchelys melanostomias Toyoshima, 1983, a senior synonym of L. brevimaxillaris Toyoshima, 1985 (Teleostei: Zoarcidae). Ichthyological Research, 51, 84 - 88. https: // doi. org / 10.1007 / s 10228 - 003 - 0194 - 2","Imamura, H., Shinohara, G. & Shiogaki, M. (2005) Rediscovery of the type specimens of Lycenchelys brevimaxillaris (Actinopterygii: Perciformes: Zoarcidae) with taxonomic notes on the species. Species Diversity, 10, 1 - 6. https: // doi. org / 10.12782 / specdiv. 10.1"]}

Lycenchelys remissaria Fedorov, 1995 (Japanese name: Kawari-hebigenge) (Figs. 32���35; Table 8) Lycenchelys remissaria Fedorov, 1995b: 135, figs. 1���2 (original description, type locality: off Ibaraki Prefecture, Pacific coast of Honshu Island, Japan); Shinohara et al., 1996: 181, fig. 2B (description); Imamura, 1997: 60 (species list); Imamura, 1998: 32, fig. 12 (brief description); Hatooka, 2000: 1032, 1590, unnumbered fig. (key to species); Hatooka, 2002: 1032, 1581, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 20 (species list); Shinohara & Anderson, 2007: 63, table 1 (comparison with Lycenchelys ryukyuensis and key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 724 (species list); Amaoka et al., 2011: 317, unnumbered fig. (brief description); Balushkin et al., 2011: 982 (catalog of specimens); Hatooka, 2013: 1225, 2078, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: ZIN 50586, male, 200.5 mm SL, off Miyagi Prefecture, Tohoku District, northwestern Pacific (36��54.4���N, 141��55.7���E), 1020 m depth, 4 Feb. 1981, R/V Mys Dal���niy. Other specimens (4 specimens): HUMZ 180877, NSMT-P 47278, NSMT-P 49022 (2), 1 male and 3 females, 216.1���304.4 mm SL, Tohoku District, northwestern Pacific. Diagnosis. Vertebrae 25���26 + 99���105 = 124���130; head length 14.1���15.3% SL; interorbital pores and occipital pores absent; postorbital pores 3���4; suborbital pores 7 + 1; preoperculomandibular pores 9; vomerine teeth 3���5; palatine teeth 2���4, arranged in single row; opercular flap well developed; pelvic-fin base positioned below 7th suborbital pore; two incomplete lateral lines, positioned mediolaterally and ventrally; scales present on pectoral fin and its base; body uniformly grayish when fresh. Description. Counts and proportional measurements in Table 8. ....Continued next page Body elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 4.4���5.3% SL (unknown for holotype). Head moderately long, ovoid; dorsal profile of head sloping extremely gently to dorsal-fin origin. Snout short, 95.5���106.8% of eye diameter (unknown for holotype). Eye ovoid, moderately large. Interorbital space narrow, width 14.1���21.1 (17.1)% of eye diameter. Nostril tube short, not reaching upper lip when depressed. Mouth terminal. Posterior edge of upper jaw reaching to about vertical through anterior edge of pupil or middle of eye (anterior edge of pupil); no significant sexual dimorphism recognized in upper jaw length. Labial lobe of lower jaw weak. Teeth on jaws, vomer and palatine small and conical; upper jaw with 2 rows anteriorly, and 1 or 1���2 rows posteriorly; lower jaw with 2���4 irregular rows anteriorly, and 1 or 1���2 rows posteriorly (tooth arrangement in jaws unknown for holotype); vomerine teeth irregularly arranged; palatine teeth in single row. Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap weakly developed. Gill rakers short and triangular (Fig. 33). Pseudobranch filaments short. Two lateral lines present, deciduous, incomplete; mediolateral line more deciduous, only visible above pectoral fin; ventral line originating posterior to last postorbital pore and terminating anterior to anus. Scales small and cycloid, present on nape, body, except around pelvic fin, pectoral axilla, about 10���30% of pectoral fin basally, pectoral-fin base, tail and about 70���80 % vertical fins basally (scaled areas unknown for holotype). Head and margins of vertical fins without scales. Dorsal-fin origin above middle of pectoral fin; 1st dorsal-fin pterygiophore between neural spines of 7th to 9th vertebrae (between 7th and 8th). Anal-fin origin below 16th or 17th (17th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 3rd to 5th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 3rd to 5th (between 3rd and 4th) preural vertebrae. Caudal fin with 2 epural, 3���4 (4) upper hypural and 4���5 (5) lower hypural rays. Pectoral fin moderately short, reaching to middle of abdomen; its posterior margin rounded dorsally and notched ventrally. Upper end of pectoral-fin base on about lateral midline of body. Pelvic fin relatively long; its base below 7th suborbital pore (Fig. 35A); its posterior margin reaching to or slightly beyond lower edge of gill opening (position of posterior margin of pelvic fin unknown for holotype). Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above and slightly anterior to vertical through 1st suborbital pore (Fig. 35A, B). Postorbital pores 3 (Fig. 35A, B); distance between 1st and 2nd pores longest of those between adjacent pores. Suborbital pores 8; 7 pores below eye and last on ascending part of suborbital canal behind eye; 7th pore above pelvic-fin base; 8th pore slightly higher than other 7 suborbital pores and set posterior edge of cheek (Fig. 35A). Preoperculomandibular pores 9; 5 on lower jaw and 4 on preopercle; last preoperculomandibular pore anterior to upper end of pectoral-fin base (Fig. 35A, C). Interorbital pores and occipital pores absent (Fig. 35B). Color in alcohol. Holotype (based on color photograph; Fig. 34) with dark brown head, pectoral fin and margin of vertical fins and bluish gray body, vertical fins and abdomen. Other specimens with pale chocolate brown head, pectoral fin and margins of vertical fins, pale brown body and vertical fins, and dark brown abdomen. Color when fresh (based on color photograph of HUMZ 180877; Fig. 32). Head, pectoral fin and margin of vertical fins blackish, body and vertical fins uniformly grayish, and abdomen dark gray. Distribution. Off northwestern Pacific coast of Honshu Island from Iwate to Ibaraki prefectures, at depths of 1020���2034 m (Fedorov, 1995b; Shinohara et al., 1996; Imamura, 1997, 1998; Hatooka, 2000, 2002, 2013; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Amaoka et al., 2011; Balushkin et al., 2011; this study). Size. Maximum length 32 cm TL (Hatooka, 2013; Amaoka et al., 2011). The largest specimen examined during this study measured 304.4 mm SL (311.0 mm TL). Remarks. Lycenchelys remissaria resembles Lycenchelys cicatrifer (Garman, 1899) and Lycenchelys novaezealandiae Anderson & M��ller, 2007 in having no interorbital pores and occipital pores, 3���4 postorbital pores, 2 lateral lines and scales on the nape in large specimens (vs. lacking this combination of characters in other species of Lycenchelys) (e.g., Anderson, 1995; Fedorov, 1995b; Shinohara et al., 1996; Hatooka, 2000, 2002, 2013; Anderson & M��ller, 2007; Shinohara & Anderson, 2007; this study). Lycenchelys remissaria is easily distinguished from L. novaezealandiae by its fewer numbers of dorsal-fin rays (116���123 vs. 128���132), anal-fin rays (103���107 vs. 119), and total vertebrae (124���130 vs., 139���141), respectively (Anderson & M��ller, 2007; this study). Lycenchelys remissaria is allied to L. cicatrifer in having similar counts and proportional measurements (e.g., 108���115 dorsal-fin rays, 97���104 anal-fin rays and 116���124 total vertebrae in L. cicatrifer) (Table 8). However, L. remissaria is clearly separable from L. cicatrifer by the position of the pelvic fin. In L. remissaria, the pelvic fin is positioned extremely anteriorly with its base below the the 7th suborbital pore (Fig. 35A). In contrast, the pelvic-fin base of L. cicatrifer is located posterior to the suborbital pores. The position of the pelvic-fin base is also useful for separating L. remissaria from all species of Lycenchelys (except for 6 the species without pelvic fins), which have the more posterior position of the fin., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 38-41, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Fedorov, V. V. (1995 b) Lycenchelys remissaria sp. nova (Perciformes: Zoarcidae) from the bathyal region of the ocean shores of Japan. Journal of Ichthyology, 35, 135 - 139.","Shinohara, G., Endo, H. & Matsuura, K. (1996) Deep-water Fishes collection from the Pacific coast of northern Honshu, Japan. Memories of the National Science Museum, Tokyo, 29, 153 - 185.","Imamura, H. (1997) Perciform fish fauna from off Pacific of Tohoku. Researches on Basic Fishes in Tohoku District, 17, 55 - 68. [in Japanese]","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133.","Anderson, M. E. & Moller, P. R. (2007) Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere. XIII. Two new species of Lycenchelys from the Southwestern Pacific. Species Diversity, 12, 175 - 185. https: // doi. org / 10.12782 / specdiv. 12.175"]}

Lycenchelys albomaculata Toyoshima, 1983 (Japanese name: Shirobuchi-hebigenge) (Figs. 1���4; Table 1) Lycenchelys albomaculatus Toyoshima, 1983: 141, 269, 333, figs. 16���19, pl. 92, 156 (original description, type locality: off Kamaishi, Iwate Prefecture, Pacific coast of Honshu, Japan); Toyoshima, 1984: 293, pl. 274-C (brief description); Toyoshima, 1985: 159, figs. 6���7, 16���19, 31, table 1 (description); Hatooka, 1993: 904, unnumbered fig. (key to species); Amaoka et al., 1995: 241, pl. 404 (brief description); Imamura, 1997: 60 (species list); Koyanagi, 1997: 538, fig. 1 (brief description); Imamura, 1998: 30, fig. 7 (brief description); Zama, 2001: 86, 133 (species list). Lycenchelys albomaculata: Anderson, 1994: 112, 117 (species list); Shinohara et al., 1996: 180 (species list); Hatooka, 2000: 1035, unnumbered fig. (keys to species); Hatooka, 2002: 1035, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 15 (species list); Shiogaki et al., 2004: 71 (species list); Shinohara & Anderson, 2007: 64 (key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 723 (species list); Amaoka et al., 2011: 319, unnumbered fig. (brief description); Balushkin et al., 2011: 978 (catalog of specimens); Hatooka, 2013: 1229, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Material examined Holotype: HUMZ 59531, male, 406.6 mm SL, off Kamaishi, Iwate Prefecture, Tohoku District, northwestern Pacific, 14 Oct. 1976. Paratypes (19 specimens, 326.8���438.5 mm SL, all from Tohoku District, northwestern Pacific): HUMZ 72538��� 39, 1 male and 1 female, 356.0��� 356.4 mm SL, off Fukushima Prefecture (38��00���N, 142��10.05���E), 800���810 m depth, 6 Feb. 1978; HUMZ 72645���46, 2 males, 389.5���390.9 mm SL, off Fukushima Prefecture (37��26.5���N, 142��09.5���E), 900 m depth, 20 Jan. 1978; HUMZ 72656���57, 1 male and 1 female, 326.9���397.0 mm SL, off Miyagi Prefecture (38��02���N, 142��29���E), 1100���1150 m depth, 7 Feb. 1978; HUMZ 72708, 1 female, 326.8 mm SL, off Fukushima Prefecture (36��58.8���N, 141��47.5���E), 800 m depth, 19 Jan. 1978; HUMZ 72723, 1 male, 387.4 mm SL, off Fukushima Prefecture (37��11���N, 141��57���E), 810���820 m depth, 18 Jan. 1978; HUMZ 72734, 1 female, 358.2 mm SL, off Miyagi Prefecture (38��04���N, 142��12.2���E), 815���820 m depth, 30 Jan. 1978; HUMZ 78063, 1 male, 413.6 mm SL, off Iwate Prefecture (40��21.6���N, 142��25.6���E to 40��26.8���N, 142��22.1���E), 915���945 m depth, 25 Sep. 1978; HUMZ 78074, 1 male, 405.2 mm SL, off Iwate Prefecture (39��22.7���N, 142��36.5���E to 39��27.9���N, 142��40.2���E), 1290��� 1300 m depth, 21 Sep. 1978; HUMZ 78082���83, 2 females, 365.0��� 371.2 mm SL, off Iwate Prefecture (39��42.6���N, 142��47.5���E to 39��36.4���N, 142��44.2���E), 1120���1130 m depth, 23 Sep. 1978; HUMZ 78087, 1 female, 393.2 mm SL, off Iwate Prefecture (40��04.5���N, 142��43.5���E to 40��10.0���N, 142��39.3���E), 1095���1100 m depth, 25 Sep. 1978; HUMZ 78142, 1 female, 370.6 mm SL, off Iwate Prefecture (39��04.5���N, 142��22.7���E to 39��10.3���N, 142��25.0���E), 980���1000 m depth, 19 Sep. 1978; HUMZ 78200, 1 male, 436.0 mm SL, off Iwate Prefecture (39��40���N, 142��48.4���E to 39��45.3���N, 142��53���E), 1180���1230 m depth, 23 Sep. 1978; HUMZ 78262, 1 male, 434.5 mm SL, off Aomori Prefecture (40��48.9���N, 142��26.2���E to 40��44.4���N, 142��30.5���E), 1120���1165 m depth, 11 Sep. 1978; HUMZ 78269, 1 female, 413.0 mm SL, off Aomori Prefecture (40��47.6���N, 142��16.7���E to 40��41.5���N, 142��19.2���E), 920���948 m depth, 11 Sep. 1978; HUMZ 78322, 1 male, 438.5 mm SL, off Aomori Prefecture (41��02.4���N, 142��11.9���E to 41��08.1���N, 142��12.2���E), 1200���1205 m depth, 8 Sep. 1978. Other specimens (40 specimens, 141.4���508.0 mm SL): HUMZ 72495, 72537, 72610���11, 72648, 72704, 72733, 72738, 180859, 180870, 180879, 180883, 180900, 182282, 182314, 182316, 182318���19, 182326, 209261, 214430���31, 214626, 226944, 12 males and 12 females, 141.1���386.4 mm SL, Tohoku District, northwestern Pacific; HUMZ 119854���55, 120153, 123913, 126048, 126353���55, 4 males and 4 females, 195.3��� 508.0 mm SL, eastern Hokkaido Island, northwestern Pacific; HUMZ 133184, 196387���88, 205185, 205192, 205194, 215072, 228077, 4 males and 4 females, 397.5���454.1 mm SL, northeastern Hokkaido Island, Okhotsk Sea. Diagnosis. Vertebrae 22���25 + 99���105 = 122���128; head length 14.3���20.4% SL; interorbital pore 1; occipital pores 3; postorbital pores 4; suborbital pores 7���9 + 2���3; preoperculomandibular pores usually 9; vomerine teeth 2���10; palatine teeth 2���18, arranged in 1 or 1���2 rows; opercular flap well developed; pelvic-fin base positioned posterior to lower edge of gill opening; lateral line complete and positioned ventrally; scales present on pectoral fin and its base; body blackish brown when fresh; 6���10 white blotches present above upper edge of gill opening and on dorsal fin extending onto dorsal part of body. Description. Counts and proportional measurements in Table 1. Body elongate, cross section oval anteriorly, compressed posteriorly; its width at anal-fin origin 3.1���6.0 (5.6)% SL. Head moderately long, ovoid, dorsal profile of head sloping extremely gently to dorsal-fin origin. Cheek swol- len in large males (including holotype), more so than in females and small males. Head of adults longer in males than in females. Snout short, 72.5���219.1 (145.1)% of eye diameter. Eye ovoid, relatively large. Interorbital space relatively narrow in adults, wide in juveniles; its width 8.9���29.6 (17.5)% of eye diameter. Nostril tube short, not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching vertical through posterior margin of eye in adult males (including holotype), reaching below middle or posterior part of eye in females and juveniles. Labial lobe of lower jaw developed in adults (including holotype). Teeth on jaws, vomer and palatine small and conical; upper jaw with single row, sometimes having some additional teeth behind anterior teeth (including holotype); lower jaw with 2���4 irregular rows anteriorly and 1���2 rows posteriorly; vomerine teeth irregularly arranged; palatine teeth in 1 or 1���2 rows (1). Lower edge of gill opening below lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short; those on upper limb triangular, many triangular and some blunt on lower limb (Fig. 2). Pseudobranch filaments long. Lateral line deciduous, complete and positioned ventrally; originating posterior to last postorbital pore and terminating on tail. Scales small and cycloid, present on body, pectoral axilla, about basal half of pectoral fin, pectoral-fin base, tail and most of vertical fins except margins. Scales covering nape in adults (including holotype), but not in some juveniles. Head without scales. ....Continued next page Dorsal-fin origin above middle of pectoral fin; 1st dorsal-fin pterygiophore between neural spines of 3rd to 5th (unknown for holotype) vertebrae. Anal-fin origin below 17th to 20th (unknown for holotype) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate or penultimate abdominal vertebra (unknown for holotype). Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (unknown for holotype) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 4th (unknown for holotype) preural vertebrae. Caudal fin with 1���2 epural, 4���5 upper hypural and 3���5 lower hypural rays (unknown for holotype). Pectoral fin moderately long in juveniles, becoming relatively short in adults (including holotype); tip reaching or over middle of abdomen in juveniles, not quite reaching middle portion of abdomen in adults (including holotype); its posterior margin rounded. Upper end of pectoral-fin base on about lateral midline of body. Pelvic fin short; its base posterior to lower edge of gill opening; its posterior margin not reaching pectoral-fin base in adults (including holotype), reaching pectoral-fin base in some juveniles. Head pores small and distinct. Usually 2 nasal pores (including holotype); anterior pore in front of nostril tube, posterior pore about above 1st suborbital pore; third pore present above nostril tube on left side in HUMZ 78087 and on both sides in HUMZ 214430 (Fig. 4A, B). Postorbital pores 4; distance between 1st and 2nd pores longest of those between adjacent pores (Fig. 4A, B). Suborbital pores 9���11 (10); 7���9 (8) pores located below eye and remaining 2 or 3 (2) pores on ascending part of suborbital canal behind eye; 5th pore about below anterior margin of pupil; last pore below eye located posterior to vertical through posterior margin of eye (Fig. 4A). Preoperculomandibular pores usually 9 (including holotype), 4 on lower jaw, 2 on junction of lower jaw and preopercle, and 3 on preopercle; some specimens with 8 pores having only 1 pore on junction of lower jaw and preopercle; HUMZ 72734 having 10 pores with 4 pores on preopercle of right side; last preoperculomandibular pore located posterior to lower part of eye (Fig. 4A, C). One interorbital pore located anterior to center of eyes (Fig. 4B). Occipital pores 3; 1 on midline of occiput, and remaining 2 on left and right sides; middle pore located slightly posterior to other two; all pores located anterior to 3rd postorbital pore (Fig. 4B). Color in alcohol. Holotype (Fig. 3) with head, body and vertical fins brown, opercular region and pectoral fin darker; nine white blotches present dorsally, 1 above upper edge of gill opening and 8 on dorsal fin extending onto dorsal part of body. Paratypes and long preserved non-type specimens with coloration similar to holotype but more recently preserved specimens darker than holotype with one white blotch above upper edge of gill opening and another 5���9 on dorsal fin and dorsal part of body. Juveniles having narrower white blotches than adults. Color when fresh (based on color photograph of HUMZ 228077; Fig. 1). Head, body, pectoral-fin base and vertical fins blackish brown, pectoral fin and margin of vertical fins darker. Ten white blotches; 1 above upper edge of gill opening, 9 on dorsal fin extending onto dorsal part of body. Distribution. The Okhotsk Sea and off the northwestern Pacific coast of the Kuril Islands, and the eastern side of Hokkaido Island to the Ibaraki Prefecture, at depths of 400���1505 m (Toyoshima, 1983, 1984, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1994; Amaoka et al., 1995, 2011; Shinohara et al., 1996; Imamura, 1997, 1998; Koyanagi, 1997; Zama, 2001; Anderson & Fedorov, 2004; Shiogaki et al., 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Balushkin et al., 2011; this study). Size. The largest specimen recorded during this study was 508.0 mm SL (518.6 mm TL), exceeding the previously recorded maximum length of 50 cm TL (Amaoka et al., 1995, 2011; Koyanagi, 1997; Imamura, 1998). Remarks. Lycenchelys albomaculata is characterized by the presence of white blotches on the body. Although Lycenchelys bachmanni Gosztonyi, 1977 also has paler blotches on the body (vs. paler blotches absent in other species of Lycenchelys), its blotches are yellowish (Gosztonyi, 1977; Nakamura, 1986). In addition, L. albomaculata is easily separable from L. bachmanni in having a scaled and dark pectoral fin (vs. naked and yellowish white in L. bachimanni) (Gosztonyi, 1977; Nakamura, 1986). Furthermore, L. albomaculata has a higher number of gill rakers (1���3 + 11���14 = 13���17 in L. albomaculata vs. 2 + 8 = 10 in L. bachimanni) (Gosztonyi, 1977; Nakamura, 1986; this study). Toyoshima (1983, 1985) described the occipital pores of L. albomaculata to be ���absent���. However, this study found 3 occipital pores in all specimens of L. albomaculata, including the holotype (Fig. 4B)., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 6-10, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Toyoshima, M. (1984) Family Zoarcidae. Eelpouts. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 291 - 295. [in Japanese]","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243.","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Amaoka, K., Nakaya, K. & Yabe, M. (1995) The fishes of northern Japan. Kita-Nihon Kaiyo Center, Sapporo, 391 pp. [in Japanese]","Imamura, H. (1997) Perciform fish fauna from off Pacific of Tohoku. Researches on Basic Fishes in Tohoku District, 17, 55 - 68. [in Japanese]","Koyanagi, M. (1997) Zoarcidae. In: Okamura, O. & Amaoka, K. (Eds.), Sea fishes of Japan, Yama-Kei, Tokyo, pp. 536 - 538. [in Japanese]","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Zama, A. (2001) Fish fauna of Miyagi Prefecture, Japan. Sanwa Publisher, Ishinomaki, 153 pp. [in Japanese]","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Shinohara, G., Endo, H. & Matsuura, K. (1996) Deep-water Fishes collection from the Pacific coast of northern Honshu, Japan. Memories of the National Science Museum, Tokyo, 29, 153 - 185.","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shiogaki, M., Ishito, Y., Nomura, Y. & Sugimoto, T. (2004) Revised catalog of the fishes collected from the waters of Aomori Prefecture. Bulletin of Aomori Prefectural Fisheries Research Center, (4), 39 - 80. [in Japanese with English abstract]","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Gosztonyi, A. E. (1977) Results of the research cruises of FRV \" Walter Herwig \" to South America. XLVIII. Revision of the South American Zoarcidae (Osteichthyes, Blennioidei) with the description of three new genera and five new species. Archiv fur Fischereiwissenshaft, 27, 191 - 249.","Nakamura, I. (1986) Zoarcidae. In: Nakamura, I., Inada, T., Takeda, M. & Hatanaka, H. (Eds.), Important fishes trawled off Patagonia. Japan Marine Fishery Resources Research Center, Tokyo, pp. 232 - 243."]}

Lycenchelys hippopotamus Schmidt, 1950 (Japanese name: Hebigenge) (Figs. 9���13; Table 3) Lycenchelys hippopotamus Schmidt, 1935: 35 (nomen nudum); Taranetz, 1937: 161 (nomen nudum). Lycenchelys hippopotamus Schmidt, 1950: 106, fig 4, pl. IX (original description, type locality: east coast of Sakhalin Island, Sea of Okhotsk); Andriashev, 1955: 354, 361, figs. 2, 7���8 (description); Matsubara, 1955: 774 (key to species); Fedorov, 1976: 8, tables 3���4 (description); Toyoshima, 1983: 267, 332, pl. 154 (description); Toyoshima, 1984: 293, pl. 273-O (brief description); Toyoshma, 1985: 149, 169, figs. 6���7, 24���25, 31, table 1 (description); Hatooka, 1993: 902, unnumbered fig. (key to species); Anderson, 1994: 65, 113, 117 (osteological comments); Amaoka et al., 1995: 239, pl. 401 (brief description); Anderson, 1995: 76 (description); Koyanagi, 1997: 538, fig. 2 (brief description); Hatooka, 2000: 1032, unnumbered fig. (key to species); Hatooka, 2002: 1032, unnumbered fig. (key to species); Mecklenburg et al., 2002: 701, unnumbered figs. (brief description); Anderson & Fedorov, 2004: 17 (species list); Shinohara & Anderson, 2007: 64 (key to species); Amaoka et al., 2011: 315, unnumbered fig. (brief description); Balushkin et al., 2011: 980, 1024 (catalog of specimens); Hatooka, 2013: 1226, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Lectotype: ZIN 24826, female, 206.8 mm SL, east coast of Sakhalin Island, Okhotsk Sea (53��09.5���N, 149��52.1���E), 1150 m depth, 7 Aug. 1932. Paralectotype (2 specimens): ZIN 24826, 1 male and 1 female, 167.7���194.9 mm SL, collected with lectotype. Other specimens (29 specimens): HUMZ 77571, 77573, 77774, 119986���87, 120005, 120285, 120329, 120345, 121155, 121157, 121159 ���60, 121162, 121164, 121193, 121452, 121463, 123979, 124056, 124115, 126116, 126186, 126218, 126223, 126228, 20 males and 9 females, 117.9���211.9 mm SL, northeastern Hokkaido Island, Okhotsk Sea. Diagnosis. Vertebrae 23���24 + 105���115 = 128���138; head length 12.5���15.0% SL; interorbital pores 1���2; occipital pores 2; postorbital pores 3���4; suborbital pores usually 8 + 1; 1st suborbital pore located just below nostril tube; preoperculomandibular pores 8���9; vomerine teeth 3���7; palatine teeth 1���7, arranged in single row; opercular flap well-developed; pelvic-fin base positioned below lower edge of gill opening; lateral line complete and positioned ventrally; scales present or absent on pectoral fin and absent on its base; body uniformly dark chocolate-brown when fresh. Description. Counts and proportional measurements in Table 3. ....Continued next page Body very elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 2.3���4.6% SL (unknown for lectotype). Head moderately long, ovoid; dorsal profile of head gently sloping from posterior edge of eye to above about last postorbital pore. Head of adults generally longer in males than in females. Snout rather short, 127.2���199.8 (172.3)% of eye diameter. Eye ovoid, relatively small. Interorbital space narrow, width 20.0���37.0 (27.8)% of eye diameter. Nostril tube short, reaching 1st suborbital pore when depressed. Mouth subterminal. Posterior edge of upper jaw about reaching vertical through anterior margin of eye in adult males, not reaching vertical through anterior margin of eye in females and juveniles. Labial lobe of lower jaw weak. Teeth on jaws sharp; upper jaw with single row, anterior teeth large and posterior teeth small; 4 females, including lectotype and 1 paralectotype, with additional small teeth behind anteriormost tooth; lower jaw with 2���3 irregular rows anteriorly and single row posteriorly, anterior teeth large and those posteriorly small; vomer and palatine small and conical; vomerine teeth irregularly arranged; palatine teeth in single row. Lower edge of gill opening reaching lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short; those on upper limb triangular, many triangular and some blunt on lower limb (Fig. 10). Pseudobranch filaments relatively long. Lateral line deciduous, complete and positioned ventrally; originating posterior to last postorbital pore and terminating on tail. Scales small and cycloid, present on body, tail and about 40���80 % of vertical fins basally. Scales present or absent on basal portion of pectoral-fin rays (present in lectotype). Head, nape, pectoral axil and pectoral-fin base without scales. Dorsal-fin origin posterior to vertical through pectoral-fin base; 1st dorsal-fin pterygiophore between neural spines of 4th to 6th (between 5th and 6th) vertebrae. Anal-fin origin below 17th to 19th (17th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate to antepenultimate abdominal (penultimate) vertebra. Last dorsal-fin pterygiophore between neural spines of 2nd to 5th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 6th (between 3rd and 4th) preural vertebrae. Caudal fin with 2 epural, 4���5 (4) upper hypural and 3���4 (4) lower hypural rays. Pectoral fin moderately short, reaching to about middle of abdomen; its posterior margin rounded dorsally and having notches ventrally. Upper end of pectoral-fin base about on lateral horizontal midline of body. Pelvic fin relatively long; its base below lower edge of gill opening; its posterior margin reaching vertical through pectoral-fin base. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above 2nd suborbital pore (Fig. 11A, B). Postorbital pores usually 3 (3) (Fig. 11A, B); distance between 1st and 2nd pores longest of those between adjacent pores; 1 additional small pore present just behind 1st pore on both sides in HUMZ 121157. Suborbital pores usually 9 (9), 8 pores located below lower margin of eye and 9th behind eye; when 10 pores, 9 pores below eye and last behind eye on left side in HUMZ 77571 and 121160, and both sides in HUMZ 126194; 5th pore below vertical through anterior margin of eye; last pore of those below eye slightly posterior to vertical through 1st postorbital pore (Fig. 11A). Preoperculomandibular pores 8���9 (8); 4 on lower jaw, 1���2 (1) on junction of lower jaw and preopercle, and 3 on preopercle; last preoperculomandibular pore located posterior to lower margin of eye (Fig. 11A, C). Interorbital pores 1 or 2 (1); when 2, anterior pore located above 4th suborbital pore, and posterior pore between center or slightly posterior to center of eyes (Fig. 11B); when 1, anterior pore absent and posterior pore located slightly posterior to center of eyes. Occipital pores 2, pores on either side of dorsal midline anterior to 2nd postorbital pore (Fig. 11B). Color in alcohol. Lectotype (based on color photograph; Fig. 13) with dark brown head, pectoral fin and margin of vertical fins, slightly paler body and vertical fins; purplish gray abdomen. Non-type specimens similar to lectotype, except HUMZ 77571, 77573 and 77774, which have uniformly pale brown head and body and black dots on pectoral fin (HUMZ 121157 and 121159). Color when fresh (based on color photograph of HUMZ 124056; Fig. 9). Head and pectoral fin blackish. Body and vertical fins uniformly dark chocolate brown. Distribution. Southern Okhotsk Sea to the northwestern Bering Sea, at depths of 408���1800 m (Schmidt, 1950; Andriashev, 1955; Matsubara, 1955; Fedorov, 1976; Toyoshima, 1983, 1984, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1994, 1995; Amaoka et al., 1995, 2011; Koyanagi, 1997; Mecklenburg et al., 2002; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Balushkin et al., 2011; this study). Size. Maximum length 223 mm TL (Anderson, 1995; Mecklenburg et al., 2002). The largest specimen examined for this study was 211.9 mm SL (216.3 mm TL). Remarks. Lycenchelys hippopotamus is similar to L. makushok, L. melanostomias and L. rassi in having more than 100 total vertebrae, 1���2 interorbital pores, 2 occipital pores, 3���4 postorbital pores, single ventral lateral line and no distinct spots or blotches on the body (vs. lacking this combination of characters in other species of Lycenchelys) (e.g., Schmidt, 1950; Andriashev, 1955; Toyoshima, 1983, 1985; Fedorov & Andriashev, 1993; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1995; Shinohara et al., 1996; this study). Lycenchelys hippopotamus is distinguished from these three species by the position of the 1st suborbital pore. In L. hippopotamus, it is located just below the nostril tube (Fig. 12A), while it is located behind the nostril tube in L. makushok, L. melanostomias and L. rassi (Fig. 12B, C, D). In addition, L. hippopotamus has 1���2 interorbital pores, and when only the middle pore is present, it is located between the middle of the eyes or slightly posterior. In contrast, L. makushok, L. melanostomias and L. rassi, always have only 1 pore in the interorbital space, and it is located anterior to the middle of the eyes., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 14-19, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Schmidt, P. Y. (1950) Fishes of the Sea of Okhotsk. Transaction of the Pacific Committee of the Academy of Science of the U. S. S. R., 6, 1 - 370. [in Russian]","Schmidt, P. Y. (1935) The Sea of Okhotsk and its fish fauna. Vestnik Akademii Nauk USSR, 5, 30 - 38. [in Russian] https: // doi. org / 10.2307 / 1436107","Taranetz, A. J. (1937) Hand book for identification of fishes of the Soviet far east and adjacent waters. Bulletins of the Pacific Science Institute, 11, 1 - 200. [in Russian]","Andriashev, A. P. (1955) A review of the fishes of the genus Lycenchelys Gill (Pisces, Zoarcidae) and related forms in the seas of the USSR and adjacent waters. Trudy Instituta Zoologii Akademia Nauk, Azerbaidzhanskoi SSR, 18, 349 - 384. [Translated from Russian in Selected taxonomic papers on northern marine fishes, OTS 61 - 31030]","Matsubara, K. (1955) Fish morphology and hierarchy. Ishizaki-Shoten, Tokyo, 1605 pp. [in Japanese]","Fedorov, V. V. (1976) New data on the eel-like likods (Pisces, Zoarcidae) from the northwestern Pacific Ocean and Bering Sea. Izvestiya TINRO, 100, 3 - 18. [in Russian]","Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Toyoshima, M. (1984) Family Zoarcidae. Eelpouts. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 291 - 295. [in Japanese]","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Amaoka, K., Nakaya, K. & Yabe, M. (1995) The fishes of northern Japan. Kita-Nihon Kaiyo Center, Sapporo, 391 pp. [in Japanese]","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133.","Koyanagi, M. (1997) Zoarcidae. In: Okamura, O. & Amaoka, K. (Eds.), Sea fishes of Japan, Yama-Kei, Tokyo, pp. 536 - 538. [in Japanese]","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Mecklenburg, G. W., Mecklenburg, T. A. & Thorsteinson, L. K. (2002) Fishes of Alaska. American Fisheries Society, Bethesda, xxxvii + 1037 pp.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243.","Fedorov, V. V. & Andriashev, A. P. (1993) Lycenchelys makushok sp. nova (Perciformes: Zoarcidae) from the bathyal depths of the Kuril-Kamchatka Trench. Journal of Ichthyology, 33, 130 - 135.","Shinohara, G., Endo, H. & Matsuura, K. (1996) Deep-water Fishes collection from the Pacific coast of northern Honshu, Japan. Memories of the National Science Museum, Tokyo, 29, 153 - 185."]}

Lycenchelys tohokuensis Anderson & Imamura, 2002 (Japanese name: Kitagawa-hebigenge) (Figs. 44���49; Table 11) Lycenchelys tohokuensis Anderson & Imamura, 2002: 355, figs. 1���2 (original description, type locality: off Fukushima Prefecture, Pacific coast of Honshu Island, Japan); Anderson & Fedorov, 2004: 20 (species list); Shinohara & Anderson, 2007: 63 (key to species); Kitagawa et al., 2008: 95, unnumbered fig. (brief description); Shinohara et al., 2009: 724 (species list); Amaoka et al., 2011: 317, unnumbered fig. (brief description); Balushkin et al., 2011: 1026 (species list); Hatooka, 2013: 1227, 2078, unnumbered fig. (key to species); Yamanaka & Ito, 2014: 4 (species list); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined Holotype: HUMZ 156757, male, 269.8 mm SL, off Fukushima Prefecture, Tohoku District, northwestern Pacific (37��31.55���N, 142��12.69���E to 37��32.94���N, 142��13.31���E), 697���705 m depth, 18 Apr. 1998, T/V Tanshu-maru, otter trawl. Paratypes (4 specimens, 195.3���265.1 mm SL, all from Tohoku District, northwestern Pacific): HUMZ 152369, 1 female, 259.8 mm SL, off Aomori Prefecture (40��42.42���N, 142��09.06���E to 40��41.04���N, 142��09.95���E), 661���667 m depth, 22 Oct. 1997, R/V Wakataka-maru, otter trawl; HUMZ 171668, 1 male, 195.3 mm SL, off Fukushima Prefecture, (36��42.43���N, 141��26.10���E to 36��47.68���N, 141��27.22���E), 582���591 m depth, 11 June 2000, R/V Wakatakamaru, otter trawl; NSMT-P 64042, 1 female, 237.9 mm SL, off Fukushima Prefecture (36��36.25���N, 141��18.23���E to 36��38.91���N, 141��23.67���E), 557���599 m depth, 11 June 2000, R/V Wakataka-maru, otter trawl; HUMZ 178280, 1 female, 265.1 mm SL, off Fukushima Prefecture (37��00.94���N, 141��40.59��� E to 36��57.83���N, 141��36.70��� E), 543���546 m depth, 19 Oct. 2001, R/V Wakataka-maru, otter trawl. Other specimens (3 specimens): HUMZ 177071, 192808, 206820, 1 male and 2 females, 208.1���270.1 mm SL, Tohoku District, northwestern Pacific. Diagnosis. Vertebrae 27���29 + 88���91 = 116���119; head length 13.4���14.4% SL; interorbital pore 1; occipital pores 3; postorbital pores 5; suborbital pores 5 + 2; preoperculomandibular pores 8; vomerine teeth 7���11; palatine teeth 8���15, arranged in 2 rows anteriorly and single row posteriorly; opercular flap well developed; pelvic-fins absent; two complete lateral lines, positioned mediolaterally and ventrally; scales present on pectoral fin and its base; body uniformly brown when fresh. Description. Counts and proportional measurements in Table 11. .....Continued next page Body elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 4.2���5.6 (5.9)% SL. Head relatively short, ovoid; dorsal profile of head gently sloping to above upper edge of gill opening or dorsal-fin origin (above upper edge of gill opening). Snout short, 99.0���142.4 (99.0)% of eye diameter. Eye rounded or slightly oval (rounded), relatively small. Interorbital space moderately narrow, width 23.8���45.4 (23.8)% of eye diameter. Nostril tube long, reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching to about vertical through posterior margin of pupil. Labial lobe of lower jaw developed. Teeth on jaws, vomer and palatine small and conical; upper jaw with 2���3 rows anteriorly and single row posteriorly; lower jaw with 2���3 or 3���4 irregular rows (3���4) anteriorly and single row posteriorly; vomerine teeth irregularly arranged; palatine teeth in 2 rows anteriorly and single row posteriorly. Lower edge of gill opening reaching to or slightly above lower end of pectoral-fin base (above lower end of pectoral-fin base). Opercular flap well developed. Gill rakers, short; those on upper limb triangular, many blunt and some triangular rakers on lower limb (Fig. 45). Pseudobranch filaments relatively long. Two lateral lines, deciduous, complete; mediolateral line origin above about middle portion of abdomen and terminating on tail; ventral lateral line origin posterior to last postorbital pore and terminating on tail. Scales small and cycloid, present on body, pectoral axilla, about basal quarter of pectoral fin, pectoral-fin base, tail and most regions of vertical fins except near margins. Nape with few scales or naked (with few scales). Head without scales. Dorsal-fin origin nearly above pectoral-fin base; 1st dorsal-fin pterygiophore between neural spines of 3rd and 4th vertebrae. Anal-fin origin below 23rd to 25th (24th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate or penultimate (penultimate) abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 3rd to 5th (between 4th and 5th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 4th (between 2nd and 3rd) preural vertebrae. Caudal fin with 2 epural, 4 upper hypural and 4 lower hypural rays. Pectoral fin moderately short, reaching to anterior or middle portion of abdomen; its posterior margin rounded dorsally and notched ventrally. Upper end of pectoral-fin base slightly below lateral midline of body. Pelvic fins absent. Head pores well developed and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above 1st suborbital pore (Fig. 46A, B). Postorbital pores 5 (Fig. 46A, B); holotype with extremely small additional pore posterior to 5th pore on right side. Suborbital pores 7; 5 pores below eye and 2 pores on ascending part of suborbital canal behind eye; 5th pore behind vertical through posterior margin of eye (Fig. 46A). Preoperculomandibular pores 8; 4 on lower jaw and 4 on preopercle; last preoperculomandibular pore posterior to lower margin of eye (Fig. 46A, C). One interorbital pore on dorsal midline between anterior margins of pupils (anterior margin of pupils) (Fig. 46B). Occipital pores 3; 1 on dorsal midline of occiput, and remaining 2 on either side; 3 pores transversely aligned or middle pore slightly posterior to those on either side (middle pore located slightly behind side pores); all pores located anterior to 4th postorbital pore (Fig. 46B). Color in alcohol. Holotype (Fig. 47) with brownish head and uniformly paler body, pectoral fin and vertical fins; dark brown margins of pectoral and vertical fins and grayish abdomen. Body purplish brown in HUMZ 171668. Coloration of other specimens similar to holotype. Color when fresh (based on color photograph of HUMZ 152369; Fig. 44). Head, body and vertical fins uniformly brown, pectoral fin paler; margins of pectoral and vertical fins, and opercular region blackish; abdomen grayish. Distribution. Off northwestern Pacific coast of Tohoku District from Aomori to Fukushima prefectures, at depths of 543���905 m (Anderson & Imamura, 2002; Anderson & Fedorov, 2004; Shinohara & Anderson, 2007; Kitagawa et al., 2008; Shinohara et al., 2009; Amaoka et al., 2011; Hatooka, 2013; Yamanaka & Ito, 2014; this study). Size. The largest specimen examined during this study measured 270.1 mm SL (HUMZ 192808, 275.2 mm TL), about equal to the previously recorded maximum length (HUMZ 156757, holotype, 270 mm SL) (Anderson & Imamura, 2002). Remarks. Other than L. tohokuensis, only five species of Lycenchelys are known to lack pelvic fins: Lycenchelys fedorovi Anderson & Balanov, 2000, Lycenchelys hureaui (Andriashev, 1979), Lycenchelys monstrosa Anderson, 1982, Lycenchelys nanospinata Anderson, 1988 and Lycenchelys xanthoptera Anderson, 1991 (Anderson, 1982a, 1988, 1991; Anderson & Balanov, 2000; Anderson & Imamura, 2002). Lycenchelys tohokuensis is distinguished from the five species in having two lateral lines and palatine teeth arranged in two rows anteriorly (vs. having a single lateral line in all five species and palatine teeth arranged in a single row in L. fedorovi, L. hureaui, L. monstrosa and L. nanospinata, or lacking palatine teeth in L. xanthoptera) (Anderson, 1982a, 1988, 1991; Anderson & Balanov, 2000; Anderson & Imamura, 2002). Lycenchelys tohokuensis also differs from L. fedorovi, L. hureaui, L. monstrosa and L. nanospinata in having 116���119 total vertebrae (vs. 132���139 in L. fedorovi, 104���112 in L. hureaui, 126���132 in L. monstrosa and 105 in L. nanospinata), from L. hureaui, L. monstrosa, L. nanospinata and L. xanthoptera in having 1 interorbital pore and 3 occipital pores (vs. interorbital pores and occipital pores absent in the four), and from L. monstrosa in having vomerine teeth (vs. usually absent in L. monstrosa) (Anderson, 1982a, 1988, 1991; Anderson & Balanov, 2000; Anderson & Imamura, 2002; this study). Although the Lycodinae are recognized as having an L-shaped series of suborbital bones (Anderson, 1994), they are arranged in a semicircular series in L. tohokuensis (Fig. 45A). The latter configuration also occurs in the other three subfamilies: Lycozoarcinae Andriashev, 1939, Zoarcinae Gill, 1862 and Gymnelinae Gill, 1864 (Anderson, 1994). Lycenchelys tohokuensis clearly differs from species of Lycozoarcinae (including only Lycozoarces regani Popov, 1933) and Zoacinae (only Zoarces Cuvier, 1829 with six species) in having scales, and teeth on the vomer and palatine, and lacking the pelvic fins and dorsal-fin spines (vs. having pelvic fins and lacking scales in the Lycozoarcinae, and having pelvic fins and dorsal-fin spines, and lacking vomerine and palatine teeth in the Zoarcinae) (Anderson, 1994). When compared with genera of Gymnelinae, L. tohokuensis mostly resembles Ericandersonia Shinohara & Sakurai, 2006 [only 1 species, Ericandersonia sagamia Shinohara & Sakurai, 2006 (Fig 48)] in having similar numbers of dorsal-fin rays (112���116 vs. 113���116), anal-fin rays (93���94 vs. 93���95), pectoral-fin rays (14 vs. 14), and total vertebrae (116���119 vs.117���118), respectively (Anderson & Imamura, 2002; Shinohara & Sakurai, 2006; this study). Furthermore, numbers and arrangements of head pores in the two species are the same (2 nasal, 5 postorbital, 7 suborbital, 4 mandibular, 4 preopercular, 1 interorbital and 3 occipital pores) (Figs. 46A, B, C, 49A, B, C) (Anderson & Imamura, 2002; Shinohara & Sakurai, 2006; this study). The two are distinguishable, however, by the presence of 2 lateral lines, and a developed labial lobe on the lower jaw, and the lower edge of the gill opening set slightly above the lower end of the pectoral-fin base in L. tohokuensis (vs. lateral line single and mediolateral, labial lobe of lower jaw absent, and lower edge of the gill opening not reaching lower end of pectoral-fin base in E. sagamia) (Anderson & Imamura, 2002; Shinohara & Sakurai, 2006; this study). In addition, E. sagamia is characterized by the presence of a weak ridge (= pseudosubmental crest sensu Shinohara & Sakurai, 2006) on the ventral surface of the mandibula, which is absent in other zoarcid species (Shinohara & Sakurai, 2006; this study). The question of retaining Lycenchelys tohokuensis in Lycenchelys because of its peculiar arrangement of the suborbital bones, relative to that of other members of the subfamily needs to be tested by a phylogenetic study of the entire group., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 50-55, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Anderson, M. E. & Imamura, H. (2002) A new species of Lycenchelys (Perciformes: Zoarcidae) from the Pacific coast of northern Japan. Ichthyological Research, 49, 355 - 357. https: // doi. org / 10.1007 / s 102280200053","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Kitagawa, D., Imamura, H., Goto, T., Ishito, Y., Fujiwara, K. & Ueda, Y. (2008) Field guide of the fishes from the Tohoku District, north-eastern waters of Japan. Tokai University Press, Hadano, 141 pp. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Amaoka, K., Nakaya, K. & Yabe, M. (2011) Fishes of Hokkaido. The Hokkaido Shimbun Press, Sapporo, 482 pp. [in Japanese]","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Yamanaka, T. & Ito, K. (2014) Additional records and revisions to the revised catalog of the fishes collected from the waters of Aomori Prefecture-I. Bulletin of Aomori Prefectural Industrial Technology Research Center Fisheries Institute, 8, 1 - 10. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Anderson, M. E. & Balanov, A. A. (2000) Lycenchelys fedorovi: a new species of eelpout (Teleostei: Zoarcidae) from the northern Pacific Ocean. Copeia, 2000, 1056 - 1061. https: // doi. org / 10.1643 / 0045 - 8511 (2000) 000 [1056: LFANSO] 2.0. CO; 2","Andriashev, A. P. (1979) First finding of fishes of the families Zoarcidae and Liparididae near Kerguelen Islands. Biologiya Morya, 6, 28 - 34. [in Russian]","Anderson, M. E. (1988) Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere. I. The Antarctic and subantarctic regions. Antarctic Research Series, 47, 59 - 113. https: // doi. org / 10.1029 / AR 047 p 0059","Anderson, M. E. (1991) Studies on the Zoarcidae (Teleostei: Perciformes) of the souththern hemisphere. V. Two new species from the Weddell Sea, Antarctica. Cybium, 15, 151 - 158.","Anderson, M. E. (1982 a) A new eelpout (Teleostei: Zoarcidae) from the eastern tropical Pacific Ocean. Bulletin of Marine Science, 32 (1), 207 - 212.","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Shinohara, G. & Sakurai, H. (2006) Ericandersonia sagamia, a new genus and species of deep waters eelpouts (Perciformes: Zoarcidae) from Japan. Ichthyological Research, 53, 172 - 178. https: // doi. org / 10.1007 / s 10228 - 006 - 0333 - 7"]}

Genus Lycenchelys Gill, 1884 (Japanese name: Hebigenge-zoku) Lycenchelys Gill, 1884: 180 (type species by subsequent designation: Lycodes muraena Collett, 1878). Lycodophis Vaillant, 1888: 311 (type species by monotypy: Lycodes albus Vaillant, 1888). Embryx Jordan & Evermann, 1898: 2458 (type species by original designation: Lycodopsis crotalinus). Lyciscus Evermann & Goldsborough, 1907: 342 (type species by monotypy: Lycodopsis crotalinus Gilbert, 1890). Apodolycus Andriashev, 1979: 29 (type species by original designation: Apodolycus hureaui Andriashev, 1979). Diagnosis. Vertebrae 19���30 + 66���118 = 85���144; suborbital bones 6���10; suborbital pores 6���11; 1st dorsal-fin pterygiophores associated with vertebrae 2���21, with 0���16 free pterygiophores; palatopterygoid series weak; oral valve weak; pseudobranch, pelvic fins, and vomerine and palatine teeth usually present; scales, pyloric caeca and lateral line(s) present (Anderson, 1994; Shinohara & Matsuura, 1998; Shinohara & Anderson, 2007; this study). Distribution. Widespread in the Pacific (mainly in northwestern and eastern), Atlantic, Arctic and Southern oceans, and around the Kerguelen Islands (Anderson, 1994, 2006; Anderson & Fedorov, 2004; Anderson & M��ller, 2007; Shinohara & Anderson, 2007; Thiel et al., 2018; this study). Maximum collection depth 5320 m, in the Peru- Chile Trench (holotype of Lycenchelys atacamensis Andriashev, 1980) (Andriashev, 1980; Anderson, 1988, 1995; Anderson & Fedorov, 2004). Remarks. The genus Lycenchelys currently includes about 60 species (Anderson & Fedorov, 2004; Anderson, 2006; Anderson & M��ller, 2007; Shinohara & Anderson, 2007; Nelson et al., 2016; Thiel et al., 2018). This study recognizes 11 species of Lycenchelys in Japanese waters, in accordance with recent studies (i.e., Shinohara & Anderson, 2007; Hatooka, 2013). The diagnosis of the genus above mainly follows Anderson (1994), except for the numbers of suborbital pores and total vertebrae, variations in which were reported by Shinohara & Matsuura (1998), Shinohara & Anderson (2007) and this study. Keys to species of genus Lycenchelys from Japanese waters 1a. Interorbital pores absent............................................................................... 2 1b. Interorbital pores present............................................................................... 3 2a. Pelvic-fin base positioned below 7th suborbital pore; suborbital pores 7 + 1; preoperculomandibular pores 9; lateral lines 2; pectoral fin and its base having scales............................................................ L. remissaria 2b. Pelvic-fin base set posterior to suborbital pores; suborbital pores 6 + 1���2; preoperculomandibular pores 8; lateral line 1; pectoral fin and its base lacking scales............................................................. L. ryukyuensis 3a. Occipital pores absent................................................................................. 4 3b. Occipital pores present................................................................................ 5 4a. Total vertebrae 85���89; pectoral-fin rays 13���16; pectoral fin and its base lacking scales; body reddish orange when fresh.............................................................................................. L. aurantiaca 4b. Total vertebrae 91���95; pectoral-fin rays 17���19; pectoral fin and its base having scales; body dark brown when fresh................................................................................................. L. squamosa 5a. Distinct spots or blotches present on body.................................................................. 6 5b. Distinct spots or blotches absent on body.................................................................. 7 6a. Six to 10 white blotches present above upper edge of gill opening and on dorsal fin extending onto dorsal part of body; total vertebrae 122���128....................................................................... L. albomaculata 6b. Blackish irregular blotches on most portions of dorsal fin and dorsal part of body; total vertebrae 137���142...... L. maculata 7a. Pelvic fins present; suborbital bone arrangement L-shaped; a single lateral line.................................... 8 7b. Pelvic fins absent; suborbital bone arrangement semicircular; two lateral lines........................... L. tohokuensis 8a. First suborbital pore located just below nostril tube; interorbital pores 1���2, when 1 interorbital pore present, located between middle centers of eyes or slightly posteriorly................................................... L. hippopotamus 8b. First suborbital pore located behind nostril tube; interorbital pore 1, located anterior to center of eyes................... 9 9a. Occipital pore usually 1 (rarely 2 or 3), 1 pore always present on midline of occiput; preoperculomandibular pores 9���10; total vertebrae 132���139.......................................................................... L. makushok 9b. Occipital pores usually 2 (rarely 1), located on left and/or right sides of occiput; preoperculomandibular pores 7���9; total vertebrae 117���134....................................................................................... 10 10a. Opercular flap absent; distance between 2nd and 3rd postorbital pores long, 11.8���18.2% HL.................... L. rassi 10b. Opercular flap present; distance between 2nd and 3rd postorbital pores short, 4.1���9.2% HL............. L. melanostomias, Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 4-5, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Gill, T. N. (1884) On the anacanthine fishes. Proceedings of the Academy of Natural Sciences of Philadelphia, 36, 167 - 183.","Vaillant, L. (1888) Expeditions scientifiques du \" Travailleur \" et du \" Talisman \" pendant les annees 1880, 1881, 1882, 1883. Masson, Paris, 406 pp. https: // doi. org / 10.5962 / bhl. title. 13677","Jordan, D. S. & Evermann, B. W. (1898) The fishes of North and Middle America: A descriptive catalogue of the species of fishlike vertebrates found in the waters of North America north of the Isthmus of Panama. Bulletin of the United States National Museum, 47, 2183 - 3136. https: // doi. org / 10.5962 / bhl. title. 39716","Evermann, B. W. & Goldsborough, E. L. (1907) The fish of Alaska. Bulletin of the Bureau of Fisheries, 26, 219 - 360. https: // doi. org / 10.5962 / bhl. title. 60330","Andriashev, A. P. (1979) First finding of fishes of the families Zoarcidae and Liparididae near Kerguelen Islands. Biologiya Morya, 6, 28 - 34. [in Russian]","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Shinohara, G. & Matsuura, K. (1998) A new zoarcid, Lycenchelys aurantiaca, from the Pacific coast off northern Japan (Teleostei: Perciformes). Ichthyological Research, 45, 151 - 155. https: // doi. org / 10.1007 / BF 02678557","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Anderson, M. E. (2006) Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere. XII. Two new lycodine species from off Peru. Species Diversity, 11, 183 - 190. https: // doi. org / 10.12782 / specdiv. 11.183","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Anderson, M. E. & Moller, P. R. (2007) Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere. XIII. Two new species of Lycenchelys from the Southwestern Pacific. Species Diversity, 12, 175 - 185. https: // doi. org / 10.12782 / specdiv. 12.175","Thiel, R., Knebelsberger, T. & Eidus, I. (2018) Description and DNA barcoding of Lycenchelys lenzeni, a new species of eelpout (Perciformes: Zoarcidae) from the deep sea off the Kuril Archipelago. Zootaxa, 4370 (1), 45 - 56. https: // doi. org / 10.11646 / zootaxa. 4370.1.3","Andriashev, A. P. (1980) On the deepest occurrence of a species of the fish family Zoarcidae Lycenchelys atacamensis sp. n. from the Atacama Trench. Zoologicheskii Zhurnal, 59, 1105 - 1108. [in Russian with English summary]","Anderson, M. E. (1988) Studies on the Zoarcidae (Teleostei: Perciformes) of the southern hemisphere. I. The Antarctic and subantarctic regions. Antarctic Research Series, 47, 59 - 113. https: // doi. org / 10.1029 / AR 047 p 0059","Anderson, M. E. (1995) The eelpout genera Lycenchelys Gill and Taranetzella Andriashev (Teleostei: Zoarcidae) in the eastern Pacific, with descriptions of nine new species. Proceedings of the California Academy of Science, 49, 55 - 133.","Nelson, J. S., Grande, T. C. & Wilson, M. V. (2016) Fishes of the world, 5 th edition. John Wiley and Sons, New York, xli + 707 pp. https: // doi. org / 10.1002 / 9781119174844","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]"]}

Lycenchelys maculata Toyoshima, 1985 (Japanese name: Kurobuchi-hebigenge) (Figs. 14���16; Table 4) Lycenchelys sp.: Toyoshima, 1984: 293, pl. 274-D (brief description). Lycenchelys maculatus Toyoshima, 1985: 149, figs. 6���9, 31, tables 1���2 (original description, type locality: off Onahama, Pacific coast of Fukushima Prefecture, Honshu Island, Japan); Hatooka, 1993: 904, unnumbered fig. (key to species); Imamura, 1998: 31, fig. 10 (brief description); Shinohara et al., 2009: 724 (species list). Lycenchelys maculata Anderson, 1994: 117 (species list); Hatooka, 2000: 1035, unnumbered fig. (key to species); Hatooka, 2002: 1035, unnumbered fig. (key to species); Anderson & Fedorov, 2004: 17 (species list); Shinohara & Anderson, 2007: 64 (key to species); Balushkin et al., 2011: 980 (catalog of specimens); Hatooka, 2013: 1229, unnumbered fig. (key to species); Nakabo & Hirashima, 2015: 217 (species list and etymology of scientific name). Materials examined. Holotype: HUMZ 71361, male, 281.1 mm SL, off Onahama, Fukushima Prefecture, Tohoku District, northwestern Pacific, 200���300 m depth, 10 Nov. 1977. Paratypes (7 specimens, 264.4���287.5 mm SL, collection locality and depth same as holotype): HUMZ 71217, 1 male, 276.3 mm SL, 8 Nov. 1977; HUMZ 71362���64, 2 males and 1 female, 264.4���276.2 mm SL, 10 Nov. 1977; HUMZ 71391���92, 1 male and 1 female, 269.4���273.4 mm SL, 11 Nov. 1977; HUMZ 71537, 1 male, 282.9 mm SL, 13 Nov. 1977. Diagnosis. Vertebrae 28���30 + 110���112 = 137���142; head length 11.7���13.7% SL; interorbital pore 1; occipital pores 3; postorbital pores 4; suborbital pores 7 + 1���2; preoperculomandibular pores 8; vomerine teeth 1���6; palatine teeth 4���9, arranged in single row; opercular flap well developed; pelvic-fin base positioned anterior to lower edge of gill opening; lateral line incomplete and positioned ventrally; scales present on pectoral fin and absent on its base; body grayish when fresh; blackish irregular blotches on most portions of dorsal fin and dorsal part of body. Description. Counts and proportional measurements in Table 4. .....Continued next page Body very elongate, cross section oval anteriorly, compressed laterally near tail; its width at anal-fin origin 3.3���4.3 (3.3)% SL. Head short, ovoid; dorsal profile of head sloping extremely gently from above posterior edge of eye to above last postorbital pore. Snout moderately short, 127.7���179.3 (179.3)% of eye diameter. Eye ovoid, relatively small. Interorbital space moderately narrow, width 21.9���43.0 (43.0)% of eye diameter. Nostril tube short, not reaching upper lip when depressed. Mouth subterminal. Posterior edge of upper jaw reaching vertical through posterior part of eye in males (including holotype), reaching to below middle or anterior part of eye in females. Labial lobe of lower jaw moderately developed. Teeth on jaws, vomer and palatine small and conical; upper jaw with 2 rows anteriorly and single row posteriorly; lower jaw with 2 irregular rows anteriorly and single row posteriorly; vomerine teeth irregularly arranged; palatine teeth in single row. Lower edge of gill opening slightly above lower end of pectoral-fin base. Opercular flap well developed. Gill rakers short and triangular. Pseudobranch filaments long. Lateral line deciduous, incomplete and positioned ventrally; originating posterior to last postorbital pore and terminating above about middle of anal fin. Scales small and cycloid, present on body, nape, pectoral-fin rays basally, tail and about basal half of vertical fins. Head and pectoral-fin base without scales. Dorsal-fin origin above pectoral-fin base; 1st dorsal-fin pterygiophore between neural spines of 2nd to 4th (between 2nd and 3rd) vertebrae. Anal-fin origin below 23rd to 25th (25th) dorsal-fin ray; 1st anal-fin pterygiophore posterior to parapophysis of ultimate to antepenultimate (penultimate) abdominal vertebra. Last dorsal-fin pterygiophore between neural spines of 2nd to 4th (between 3rd and 4th) preural vertebrae. Last anal-fin pterygiophore between hemal spines of 2nd to 4th (between 3rd and 4th) preural vertebrae. Caudal fin with 2 epural, 5 upper hypural and 5 lower hypural rays. Pectoral fin moderately short; its posterior margin reaching about middle of abdomen, and having notches. Upper end of pectoral-fin base about on lateral midline of body. Pelvic fin short; its base anterior to lower edge of gill opening; its posterior margin just reaching to below pectoral-fin base. Head pores small and distinct. Nasal pores 2; anterior pore in front of nostril tube, posterior pore above and slightly anterior to vertical through 1st suborbital pore (Fig. 15A, B). Postorbital pores 4; distance between 1st and 2nd pores longest of those between adjacent pores (Fig. 15A, B). Suborbital pores 8���9 (9); 7 pores located below eye and 1 or 2 (2) on ascending part of suborbital canal behind eye; 4th pore below anterior margin of eye; 7th pore located posterior to posterior margin of eye (Fig. 15A). Preoperculomandibular pores 8, 4 on lower jaw and 4 on preopercle; last preoperculomandibular pore located posterior to about lower margin of eye (Fig. 15A, C). One interorbital pore located on dorsal midline anterior to eyes (Fig. 15B). Occipital pores 3; 1 on dorsal midline of occiput, remaining 2 on either side; middle pore located slightly posterior to more lateral pores; all pores located anterior to 3rd postorbital pore (Fig. 15B). Color in alcohol. Holotype (Fig. 16) with pale gray head, body, pectoral-fin base, anterior part of pectoral fin and vertical fins, slightly darker posterior part of pectoral fin; grayish irregular blotches on most portions of dorsal fin and dorsal part of body. Paratypes more uniformly brownish than holotype. Color when fresh [based on color photograph in Toyoshima (1984)]. Head, body, pectoral-fin base, anterior part of pectoral fin and vertical fins grayish, posterior part of pectoral fin and abdomen darker. Blackish irregular blotches on most portions of dorsal fin and dorsal part of body. Distribution. Off northwestern Pacific coast of the Kuril Islands, and from Fukushima to Ibaraki prefectures, at depths of 200���380 m (Toyoshima, 1983, 1984, 1985; Hatooka, 1993, 2000, 2002, 2013; Anderson, 1994; Imamura, 1997, 1998; Koyanagi, 1997; Anderson & Fedorov, 2004; Shinohara et al., 2009; Balushkin et al., 2011; this study). Size. Maximum length 30 cm TL (Hatooka, 1993, 2000, 2002, 2013). The largest specimen examined during this study was 287.5 mm SL (292.8 mm TL), and is probably the same specimen previously measured for maximum length. Remarks. Lycenchelys maculata resembles Lycenchelys kolthoffi Jensen, 1904 in having more than 100 dorsal- and anal-fin rays, and total vertebrae, pelvic fins and irregular blotches on most portions of the dorsal fin to dorsal part of the body blackish (vs. lacking this combination of characters in other species of Lycenchelys) (e.g., Jensen, 1904; Andriashev, 1955; Toyoshima, 1985; Koyanagi, 1995; this study). Lycenchelys maculata can be easily distinguished from L. kolthoffi by the numbers of postorbital pores (4 vs. 1), interorbital pore (1 vs. 0) and occipital pores (3 vs. 0), respectively (Table 4) (Koyanagi, 1995; this study). Furthermore, Lycenchelys maculata is also separable from L. kolthoffi in having higher numbers of dorsal-fin rays (134���138 vs. 114���117), anal-fin rays (115���120 vs. 102���104) and total vertebrae (137���142 vs. 116���124), respectively (Table 4) (Jensen, 1904; Andriashev, 1955; Toyoshima, 1985; Koyanagi, 1995; this study). Some previous studies described the lateral line of L. maculata as only reaching to near the posterior end of the pectoral fin (Toyoshima, 1985; Hatooka, 1993, 2000, 2002, 2013), but specimens examined for this study for the lateral line reaching to about the middle of the anal fin., Published as part of Kawarada, Shumpei, Imamura, Hisashi, Narimatsu, Yoji & Shinohara, Gento, 2020, Taxonomic revision of the genus Lycenchelys (Osteichthyes: Zoarcidae) in Japanese waters, pp. 1-66 in Zootaxa 4762 (1) on pages 19-23, DOI: 10.11646/zootaxa.4762.1.1, http://zenodo.org/record/3743698, {"references":["Toyoshima, M. (1985) Taxonomy of the subfamily Lycodinae (family Zoarcidae) in Japan and adjacent waters. Memoirs of the Faculty of Fisheries, Hokkaido University, 32, 131 - 243.","Toyoshima, M. (1984) Family Zoarcidae. Eelpouts. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The Fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 291 - 295. [in Japanese]","Hatooka, K. (1993) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. Tokai University Press, Tokyo, pp. 898 - 913 + 1343 - 1345. [in Japanese]","Imamura, H. (1998) Zoarcid fishes occurring off Pacific of Tohoku District. Researches on Basic Fishes in Tohoku District, 18, 28 - 34. [in Japanese]","Shinohara, G., Narimatsu, Y., Hattori, T., Ito, M., Takata, Y. & Matsuura, K. (2009) Annotated checklist of deep-sea fishes from the Pacific coast off Tohoku District, Japan. In: Fujita, T. (Ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. National Science Museum of Nature and Science Monographs, Tokyo, 39, pp. 683 - 735.","Anderson, M. E. (1994) Systematics and osteology of the Zoarcidae (Teleostei: Perciformes). Ichthyological Bulletin of the J. L. B. Smith Institute of Ichthyology, 60, 1 - 120.","Hatooka, K. (2000) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 2 nd edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1590 - 1593. [in Japanese]","Hatooka, K. (2002) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 1028 - 1044 + 1581 - 1583.","Anderson, M. E. & Fedorov, V. V. (2004) Family Zoarcidae Swainson, 1839 eelpouts. California Academy of Sciences, Annotated Checklists of Fishes, 34, 1 - 58.","Shinohara, G. & Anderson, M. E. (2007) Lycenchelys ryukyuensis sp. nov. (Perciformes: Zoarcidae) from the Okinawa Trough, Japan. Bulletin of the National Museum of Nature and Science, Series A (Zoology), Supplement 1, 59 - 66.","Balushkin, A. V., Sheiko, B. A. & Fedorov, V. V. (2011) Catalog of archival collection of the Zoological Institute, Russian Academy of Sciences: class Osteichthyes (bony fishes), order Perciformes, family Zoarcidae. Journal of Ichthyology, 51, 950 - 1034. https: // doi. org / 10.1134 / S 0032945211100031","Hatooka, K. (2013) Zoarcidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species. 3 rd edition. Tokai Uni- versity Press, Hadano, pp. 1220 - 1237 + 2076 - 2082. [in Japanese]","Nakabo, T. & Hirashima, Y. (2015) Scientific names of Japanese fishes: etymology. Tokai University Press, Hadano, xv + 372 pp. [in Japanese]","Toyoshima, M. (1983) Zoarcidae. In: Amaoka, K., Nakaya, K., Araya, H. & Yasui, T. (Eds.), Fishes from the north-eastern Sea of Japan and the Okhotsk Sea off Hokkaido. The intensive research of unexploited fishery resources on continental slopes. Japan Fisheries Resource Conservation Association, Tokyo, pp. 136 - 149 + 208 - 210 + 258 - 277 + 329 - 335.","Imamura, H. (1997) Perciform fish fauna from off Pacific of Tohoku. Researches on Basic Fishes in Tohoku District, 17, 55 - 68. [in Japanese]","Koyanagi, M. (1997) Zoarcidae. In: Okamura, O. & Amaoka, K. (Eds.), Sea fishes of Japan, Yama-Kei, Tokyo, pp. 536 - 538. [in Japanese]","Jensen, A. S. (1904) The North-European and Greenland Lycodinae. The Danish Ingolf-Expedition, 2 (4), 1 - 99. https: // doi. org / 10.5962 / bhl. title. 12206","Andriashev, A. P. (1955) A review of the fishes of the genus Lycenchelys Gill (Pisces, Zoarcidae) and related forms in the seas of the USSR and adjacent waters. Trudy Instituta Zoologii Akademia Nauk, Azerbaidzhanskoi SSR, 18, 349 - 384. [Translated from Russian in Selected taxonomic papers on northern marine fishes, OTS 61 - 31030]","Koyanagi, M. (1995) Zoarcidae. In: Okamura, O., Amaoka, K., Takeda, M., Yano, K., Okada, K. & Chikuni, S. (Eds.), Fishes collected by the R / V Shinkai Maru around Greenland. Japan Marine Fishery Resources Research Center, Tokyo, pp. 188 - 193 + 204 - 206."]}

Phuket Marine Biological Center Research Bulletin, 77, 1, 25-31

Imamura, Hisashi, Kimura, Katsuya, Quan, Nguyen Van (2019): First Record of Thysanophrys papillaris (Actynopterygii: Scorpaeniformes: Platycephalidae) from the Western Pacific. Species Diversity 24 (1): 17-22, DOI: 10.12782/specdiv.24.17, URL: http://dx.doi.org/10.12782/specdiv.24.17

Key to subfamilies of the family Cyclopteridae 1a. First dorsal fin inconspicuous, completely embedded under skin...................................... Liparopsinae 1b. First dorsal fin conspicuous (except for adults in some species with very low and mostly buried first dorsal fin; see Mecklen- burg et al., 2002)......................................................................................2 2a. Body generally high due to well-developed dorsal crest; first dorsal fin covered with very thick skin........ Cyclopterinae 2b. Body usually globose; first dorsal fin covered with relatively thin skin.................... Eumicrotreminae subfam. nov., Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on page 52, DOI: 10.5281/zenodo.246721

Family Cyclopteridae Bonaparte, 1831 Cyclopteridae Bonaparte, 1831: 96 (type genus: Cyclopterus Linnaeus, 1758). Diagnosis (includes autapomorphies only; see osteological features under ���Characters supporting the monophyly of Cyclopteridae ��� for other diagnostic characters). Ethmoid situated posteromesial to lateral ethmoid; cartilaginous process present between ceratohyal and epihyal; paired central cartilages present; fourth ceratobranchial widely separated from central cartilages; dorsal process on pelvis present; first two or three abdominal vertebrae each having a single pair of neural spines and lacking a neural arch; dorsal and anal stays cartilaginous; second preural centrum broad. Classification of Cyclopteridae. The following classification of the family Cyclopteridae, including three (one new) subfamilies and four genera, is proposed on the basis of the phylogenetic relationships reconstructed in the present study: Family Cyclopteridae Bonaparte, 1831 Subfamily Liparopsinae Garman, 1892 Genus Aptocyclus De La Pylaie, 1835 Subfamily Cyclopterinae Linnaeus, 1758 Genus Cyclopterus Linnaeus, 1758 Subfamily Eumicrotreminae subfam. nov. Genus Cyclopsis Popov, 1930 Genus Eumicrotremus Gill, 1862b, Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on page 52, DOI: 10.5281/zenodo.246721, {"references":["Bonaparte, C. L. (1831) Saggio di una distribuzione metodica degli animali vertebrati. Presso Antonio Boulzaler, Roma, 144 pp.","Linnaeus, C. (1758) Systema Naturae, 10 th edition. Fol. 1. Laurentii Salvii, Holmiae (= Stockholm), ii + 824 pp.","Garman, S. (1892) The Discoboli (Cyclopteridae, Liparopsidae and Liparididae). Memoirs of the Museum of Comparative Zoology, 14, 1 - 96, pls. 1 - 13.","Popov, A. M. (1930) A short review of the fishes of the family Cyclopteridae. Annals and Magazine of Natural History, Series 10, 6, 69 - 76.","Gill, T. N. (1862 b) Note on some genera of fishes of western North America. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 329 - 332."]}

Genus Cyclopterus Linnaeus, 1758 Cyclopterus Linnaeus, 1758: 260 (type species: Cyclopterus lumpus Linnaeus, 1758). Lumpus Oken (ex. Cuvier), 1817: 1182 (type species: Cyclopterus lumpus Linnaeus, 1758). Diagnosis. See ���Diagnosis��� of the subfamily Cyclopterinae. Remarks. Oken (1817) established the genus Lumpus based on ���Les Lumps��� in Cuvier (1816) (see Gill, 1903). According to subsequent studies (Lindberg, 1973; Parin et al., 2002; Mecklenburg & Sheiko, 2003), this generic name is a junior objective synonym of Cyclopterus Linnaeus, 1758. The genus includes only Cyclopterus lumpus Linnaeus, 1758., Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on page 53, DOI: 10.5281/zenodo.246721, {"references":["Linnaeus, C. (1758) Systema Naturae, 10 th edition. Fol. 1. Laurentii Salvii, Holmiae (= Stockholm), ii + 824 pp.","Gill, T. N. (1903) On some fish genera of the first edition of Cuvier's Regne animal and Oken's names. Proceedings of the United States National Museum, 26, 965 - 967.","Lindberg, G. U. (1973) Cyclopteridae. In: Hureau, J. - C. & Monod, T. (Eds.), Check-list of the fishes of the north-eastern Atlantic and of the Mediterranean, CLOFNAM, Unesco, Paris, pp. 607 - 608.","Parin, N. V., Fedorov, V. V. & Sheiko B. A. (2002) An annotated catalogue of fish-like vertebrates and fishes of the seas of Russia and adjacent countries. Part 2. Order Scorpaeniformes. Journal of Ichthyology, 42 (Supplement 1), S 60 - S 135.","Mecklenburg, C. W. & Sheiko, B. A. (2003) Family Cyclopteridae Bonaparte 1831 - lumpsuckers. California Academy of Sciences Annotated Checklists of Fishes, 6, 1 - 17."]}

Genus Cyclopsis Popov, 1930 Cyclopsis Popov, 1930: 74 (type species: Cyclopsis tentacularis Popov, 1930). Diagnosis. Head and body with many tentacles; symplectic and quadrate separated; rudimentary coronomeckelian present in tendon from adductor mandibulae section 3; posterior part of palatine covered with a cartilaginous element. Remarks. The genus is monotypic, appearing first as a nomen nudum in Soldatov & Popov (1929) and subsequently being made available in Popov (1930) and Soldatov & Lindberg (1930) (Mecklenburg & Sheiko, 2003). See Popov (1931) for a detailed description of Cyclopsis. The genus includes only Cyclopsis tentacularis Popov, 1930., Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on page 54, DOI: 10.5281/zenodo.246721, {"references":["Popov, A. M. (1930) A short review of the fishes of the family Cyclopteridae. Annals and Magazine of Natural History, Series 10, 6, 69 - 76.","Soldatov, V. & Popov, A. (1929) On the new genus Cyclopteropsis (Pisces, Cyclopteridae) from the Okhotsk Sea. Doklady Akademii Nauk SSSR, Ser. A, 1929, 239 - 242.","Soldatov, V. K. & Lindberg, G. U. (1930) A review of the fishes of the seas of the Far East. Bulletins of the Pacific Science Institute, 5, i - xlvii, 1 - 576, pls. 1 - 15. [in Russian, with English summary, and new taxa also in English]","Mecklenburg, C. W. & Sheiko, B. A. (2003) Family Cyclopteridae Bonaparte 1831 - lumpsuckers. California Academy of Sciences Annotated Checklists of Fishes, 6, 1 - 17.","Popov, A. M. (1931) Cyclopteridae of the Okhotsk Sea based on collections of the Hydrographical Expedition to the Pacific Ocean. Izvestija Akademii Nauk SSSR, 1931, 85 - 99. [In Russian, new taxa diagnoses in English]"]}

Genus Aptocyclus De la Pylaie, 1835 Aptocyclus De la Pylaie, 1835: 528 (type species: Cyclopterus ventricosus Pallas, 1769). Cyclopterichthys Steindachner, 1881: 192 [type species: Cyclopterichthys glaber Steindachner, 1881 (= junior synonym of Cyclopterus ventricosus)]. Liparops Garman, 1892: 42 [type species: Cyclopterus stelleri Pallas, 1814 (= junior synonym of Cyclopterus ventricosus)]. Elephantichthys Hubbs & Schultz, 1934: 21 [type species: Elephantichthys copeianus Hubbs & Schultz, 1934 (= junior synonym of Cyclopterus ventricosus)]. Pelagocyclus Lindberg & Legeza, 1955: 436 [type species Pelagocyclus vitiazi Lindberg & Legeza, 1955 (= junior synonym of Cyclopterus ventricosus)]. Diagnosis. See ���Diagnosis��� of the subfamily Liparopsinae. Remarks. Lindberg and Legeza (1955) established the genus Pelagocyclus, known only from the holotype of Pelagocyclus vitiazi Lindberg & Legeza, 1955, and were followed by many authors (e.g., Ueno, 1970; Kido, 1984; Nelson, 1994), until Kido & Shinohara (1996) demonstrated the conspecificity of P. vitiazi and A. ventricosus (Pallas, 1769), the latter having priority. This genus includes only A. ventricosus., Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on pages 52-53, DOI: 10.5281/zenodo.246721, {"references":["Pallas, P. S. (1769) Spicilegia Zoologica. Quibus novae imprimis et obscurae animalium species, tomus 1, fasciculos 7. Gottlieb August Lange, Berolini (= Berlin), 42 pp., 6 pls.","Steindachner, F. (1881) Sitzungsberichte der Kaiserlichen Akademie der Wissenchaften, Classe v. 83 (1. Abth.), 276 pp., 8 pls.","Garman, S. (1892) The Discoboli (Cyclopteridae, Liparopsidae and Liparididae). Memoirs of the Museum of Comparative Zoology, 14, 1 - 96, pls. 1 - 13.","Pallas, P. S. (1814) Zoographia Rosso-Asiatica, sistens omnium animalium in extenso Imperio Rossico et adjacentibus maribus observatorum recensionem, domicilia, mores et descriptiones anatomen atque icones plurimorum. Caesarea Academiae Scientiarum, Petropoli (= Saint Petersburg), vii + 428 + cxxv pp.","Hubbs, C. & Schultz L. P. (1934) Elephantichthys copeianus, a new cyclopterid fish from Alaska. Copeia, 1934, 21 - 26. https: // doi. org / 10.2307 / 1436430","Lindberg, G. U. & Legeza, M. I. (1955) A review of the genera and species of the subfamily Cyclopterinae (Pisces). Bulletin of the Zoological Institute Academy of Science, 18, 389 - 458, figs. 1 - 33. [in Russian, Ueno, T. translated in English]","Ueno, T. (1970) Fauna Japonica, Cyclopteridae (Pisces). Academic Press of Japan, Tokyo, 233 pp., 13 pls.","Kido, K. (1984) Cyclopteridae. In: Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & Yoshino, T. (Eds.), The fishes of the Japanese Archipelago. Tokai University Press, Tokyo, pp. 336 - 337.","Nelson, J. S. (1994) Fishes of the world, 3 rd edition. John Wiley & Sons, New York, xvii + 600 pp.","Kido, K. & Shinohara, G. (1996) Pelagocyclus vitiazi Lindberg & Legeza, 1955, a junior synonym of Aptocyclus ventricosus (Pallas, 1769) (Scorpaeniformes: Cyclopteridae). Ichthyological Research, 43, 175 - 177. https: // doi. org / 10.1007 / BF 02348242"]}

Genus Eumicrotremus Gill, 1862b Eumicrotremus Gill, 1862b: 330 (type species: Cyclopterus spinosus Fabricius, 1776). Cyclopteroides Garman, 1892: 37 (type species: Cyclopteroides gyrinops Garman, 1892). Lethotremus Gilbert, 1896: 449 (type species: Cyclolumpus muticus Gilbert, 1896). Cyclolumpus Tanaka, 1912: 86 (type species: Cyclopteropsis asperrimus Tanaka, 1912). Cyclopteropsis Soldatov & Popov, 1929: 240 (type species: Eumicrotremus bergi Popov, 1929). Cyclopterocottus Popov, 1930: 74 (type species: Eumicrotremus brashnikowi Schmidt, 1904). Microancanthus Voskoboinikova, 2015: 217 (type species: Microancanthus tokranovi Voskoboinikova, 2015). Georgimarinus Voskoboinikova & Nazarkin, 2015: 630 (type species: Cyclopteropsis barbatus Lindberg & Legeza, 1955). Diagnosis. Head and body lacking tentacles; some species with bony elements on head and body; one or two foramina present on anterodorsal portion of prootic; mesial process of hyomandibula receiving adductor hyomandibulae; first dorsal fin visible; end of gill slit situated above level of base of uppermost pectoral fin ray. Remarks. The synonymy of Cyclopteropsis Soldatov & Popov, 1929, Eumicrotremus Gill, 1862b, Lethotremus Gilbert, 1896, and Georgimarinus Voskoboinikova & Nazarkin, 2015 is proposed here on the basis of inferred phylogenetic relationships (see ���Ranking at subfamilial and generic levels���). Microancathus Voskoboinikova, 2015 is also regarded to be a junior synonym of Eumicrotremus owing to comparison of description of the type species of the former and synapomorphies supporting cyclopterid clades inferred here. The name Eumicrotremus is considered the valid name for the genus, having priority over the others. As redefined, Eumicrotremus is distinguished from the other three valid genera in having a visible first dorsal fin, and lacking tentacles on the head and body. Tanaka (1912) established the genus Cyclolumpus for Eumicrotremus asperrimus, although it has since been regarded as a junior synonym of Eumicrotremus (Lindberg & Legeza, 1955). The present phylogenetic analysis included E. asperrimus in clade B2, which included all examined Eumicrotremus (and Lethotremus). Accordingly, Cyclolumpus is treated as a junior synonym of Eumicrotremus, following previous studies. Garman (1892) established the genus Cyclopteroides based on Eumicrotremus gyrinops Garman, 1892. However, this genus was also regarded as a junior synonym of Eumicrotremus by Lindberg & Legeza (1955). Garman���s (1892) original description of E. gryinops included: ���Skin covered with mucus, with four series of very small, distant, one to eight spined tubercles on each side���, a description closely in agreement with a diagnostic character of Eumicrotremus [head and body (or body only) having some or many tubercles] as redefined here. In addition, although E. gyrinops was not included in the phylogenetic analysis, it was confirmed by examination of a single specimen during this study that the species has many small tubercles on the body. Therefore, the genus Cyclopteroides is treated as a junior synonym of Eumicrotremus, following previous studies. The genus Cyclopterocottus was established by Popov (1930) for Eumicrotremus brashnikowi Schmidt, 1904, although it has become regarded as a junior synonym of Cyclopteropsis (e.g., Ueno, 1970; Mecklenburg & Sheiko, 2003; Nelson, 2006; this study). Lindberg & Legeza (1955) described E. brashnikowi as having ���Spinous plates having cover side of body, nap, region of cheek���, also closely in agreement with a diagnostic character of Eumicrotremus (see above). Although E. brashnikowi was not examined during the present study, the genus Cyclopterocottus is treated as a junior synonym of Eumicrotremus. Voskoboinikova (2015) established the genus Microancanthus for M. tokranovi Voskoboinikova, 2015 (type species) and Eumicrotremus fedorovi Mandrytsa, 1991, being characterized from other cyclopterids by the following unique characters: a single pore on the infraorbital sensory canal; flat or hemispherical tubercles (as bony plaques) on the body, with numerous small spines; the opercle with a conspicuous notch on its posterior edge; and an S-sharped subopercle. However, although the above species were not available for examination during the present study, M. tokranovi, at least, has sensory tubes on the lower jaw (Voskoboinikova, 2015: fig. 5B), which supports the monophyly of clade G5 (including E. orbis, E. spinosus, ��� Lethotremus ��� awae and ��� L.��� muticus) (autapomorphic character 32-1), as recognized in this study. Accordingly, recognition of Microancanthus must result in Eumicrotremus being paraphyletic. Therefore, Microancanthus cannot be recognized from a cladistic perspective and is synonymized under Eumicrotremus in this study, although the monophyly of E. tokranovi and E. fedorovi is supportable based on the above mentioned autapomorphies. Voskoboinikova & Nazarkin (2015) established the genus Georgimarinus for Eumicrotremus barbatus (Lindberg & Legeza, 1955). They considered that the genus is characterized by unique derived characters: i.e., teeth in the outer row at the symphysis of the premaxilla fused with the premaxilla; teeth at the symphysis of the dentary fused among themselves and with dentary and forming the regular cutting edge; numerous barbs present on the head and body; and the bony plaques located in centers of connective tissue tubercles leaving the edges free. The present study examined this species and also found anterior teeth on the premaxilla and dentary fused with each other and the supporting bones, and these characters were recognized as autapomorphies of the species. However, E. barbatus was inferred to be deeply nested within clade D2 including Eumicrotremus and Cyclopleropsis, and the former genus becomes paraphyletic if Georgimarinus is regraded as valid. Therefore, following the cladistic methodology, the present study synonymizes Georgimarinus under Eumicrotremus. The genus Eumicrotremus includes the following 25 valid species, seven and two species, respectively, having been previously included in Cyclopteropsis and Lethotremus: Eumicrotremus andriashevi Perminov, 1936, Eumicrotremus awae (Jordan & Snyder, 1902), Eumicrotremus asperrimus (Tanaka, 1912), Eumicrotremus barbatus (Lindberg & Legeza, 1955), Eumicrotremus bergi (Popov, 1929), Eumicrotremus brashnikowi (Schmidt, 1904), Eumicrotremus derjugini Popov, 1926, Eumicrotremus fedorovi Mandrytsa, 1991, Eumicrotremus gyrinops (Garman, 1892), Eumicrotremus inarmatus (Mednikov & Prokhorov, 1956), Eumicrotremus jordani (Soldatov, 1929), Eumicrotremus lindbergi (Soldatov, 1930), Eumicrotremus mcalpini (Fowler, 1914), Eumicrotremus muticus (Gilbert, 1896), Eumicrotremus orbis (G��nther, 1861), Eumicrotremus pacificus Schmidt, 1904, Eumicrotremus popovi (Soldatov, 1929), Eumicrotremus phrynoides Gilbert & Burke, 1912, Eumicrotremus schmidti Lindberg & Legeza, 1955, Eumicrotremus soldatovi Popov, 1936, Eumicrotremus spinosus (Fabricius, 1776), Eumicrotremus taranetzi Perminov, 1936, Eumicrotremus tartaricus Lindberg & Legeza, 1955, Eumicrotremus terraenovae Myers & B��hlke, 1950 and Eumicrotremus tokranovi (Voskoboinikova, 2015) (see Lindberg & Legeza, 1955; Ueno, 1970; Parin et al., 2002; Mecklenburg & Sheiko, 2003)., Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on pages 54-55, DOI: 10.5281/zenodo.246721, {"references":["Gill, T. N. (1862 b) Note on some genera of fishes of western North America. Proceedings of the Academy of Natural Sciences of Philadelphia, 14, 329 - 332.","Garman, S. (1892) The Discoboli (Cyclopteridae, Liparopsidae and Liparididae). Memoirs of the Museum of Comparative Zoology, 14, 1 - 96, pls. 1 - 13.","Tanaka, S. (1912) XXXVIII. Cyclolumpus asperrimus, n. g. n. sp. (Cyclopteridae). Figures and Descriptions of the Fishes of Japan, Including the Riukiu Islands, Bonin Islands, Formosa, Kurile Islands, Korea and Southern Sakhalin, 5, 86 - 87.","Soldatov, V. & Popov, A. (1929) On the new genus Cyclopteropsis (Pisces, Cyclopteridae) from the Okhotsk Sea. Doklady Akademii Nauk SSSR, Ser. A, 1929, 239 - 242.","Popov, A. M. (1930) A short review of the fishes of the family Cyclopteridae. Annals and Magazine of Natural History, Series 10, 6, 69 - 76.","Schmidt, P. J. (1904) Fishes of the eastern seas of the Russian Empire. Scientific results of the Korea-Sakhalin Expedition of the Emperor Russian Geographical Society 1900 - 1901. Fishes of the eastern seas of the Russian Empire, Saint Petersburg, xi + 466 pp., 6 pls. [in Russian]","Voskoboinikova, O. S. & Nazarkin, M. V. (2015) Georgimarinus gen. nov. - a new genus of the family Cyclopteridae (Cottoidei). Journal of Ichthyology, 55, 630 - 635.","Lindberg, G. U. & Legeza, M. I. (1955) A review of the genera and species of the subfamily Cyclopterinae (Pisces). Bulletin of the Zoological Institute Academy of Science, 18, 389 - 458, figs. 1 - 33. [in Russian, Ueno, T. translated in English]","Ueno, T. (1970) Fauna Japonica, Cyclopteridae (Pisces). Academic Press of Japan, Tokyo, 233 pp., 13 pls.","Mecklenburg, C. W. & Sheiko, B. A. (2003) Family Cyclopteridae Bonaparte 1831 - lumpsuckers. California Academy of Sciences Annotated Checklists of Fishes, 6, 1 - 17.","Nelson, J. S. (2006) Fishes of the world, 4 th edition. John Wiley & Sons, New York, xix + 601 pp.","Mandrytsa, S. A. (1991) New species of the genus Eumicrotremus (Pisces, Cyclopteridae) from the Sea of Okhotsk. Zoological Zhurnal, 70, 148 - 151. [in Russian, with English summary]","Perminov, G. N. (1936) A review of the species of the genus Eumicrotremus Gill. Bulletin of the Far Eastern Branch of the Academy of Sciences of the USSR, 19. [in Russian, English summary]","Jordan, D. S. & Snyder, J. O. (1902) A review of the discobolous fishes of Japan. Proceedings of the United States National Museum, 24, 343 - 351.","Popov, A. M. (1926) On ichthyology of the Kara and adjacent Barents seas. Travauz de ls Societe des naturalistes de Leningrad, 56, 27 - 55. [in Russian]","Mednikov, B. M. & Prokhorov, V. G. (1956) A new species of Cyclopteropsis (Pisces, Cyclopterinae) from the Bering Sea. Doklady Akademii Nauk SSSR, Ser. A, Leningrad, 111, 717 - 719.","Soldatov, V. K. & Lindberg, G. U. (1930) A review of the fishes of the seas of the Far East. Bulletins of the Pacific Science Institute, 5, i - xlvii, 1 - 576, pls. 1 - 15. [in Russian, with English summary, and new taxa also in English]","Fowler, H. W. (1914) Fishes collected by the Peary Relief Expedition of 1899. Proceedings of the Academy of Natural Sciences of Philadelphia, 66, 359 - 366.","Gunther, A. (1861) Catalogue of the fishes in the British Museum. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Gobiidae, Discoboli, Pediculati, Blenniidae, Labyrinthici, Mugilidae, Notacanthi. Order of the trustees, London, xxv + 586 + x pp.","Gilbert, C. H. & Burke, C. V. (1912) Fishes from Bering Sea and Kamchatka. Bulletin of the Bureau of Fisheries, 30, 31 - 96.","Myers, G. S. & Bohlke, J. E. (1950) A new lump-sucker of the genus Eumicrotremus from the northwestern Atlantic. Stanford Ichthyological Bulletin, 3, 199 - 202.","Parin, N. V., Fedorov, V. V. & Sheiko B. A. (2002) An annotated catalogue of fish-like vertebrates and fishes of the seas of Russia and adjacent countries. Part 2. Order Scorpaeniformes. Journal of Ichthyology, 42 (Supplement 1), S 60 - S 135."]}

Key to subfamilies of the family Cyclopteridae 1a. First dorsal fin inconspicuous, completely embedded under skin...................................... Liparopsinae 1b. First dorsal fin conspicuous (except for adults in some species with very low and mostly buried first dorsal fin; see Mecklen- burg et al., 2002)......................................................................................2 2a. Body generally high due to well-developed dorsal crest; first dorsal fin covered with very thick skin........ Cyclopterinae 2b. Body usually globose; first dorsal fin covered with relatively thin skin.................... Eumicrotreminae subfam. nov.

Subfamily Cyclopterinae Bonaparte, 1831 Cyclopterinae Bonaparte, 1831: 96 (type genus: Cyclopterus Linnaeus, 1758). Diagnosis. Head and body with many tubercles; generally high body having developed dorsal crest; supraoccipital crest present; first dorsal fin covered with very thick skin; first dorsal proximal pterygiophore lacking spines; ventral end of gill slit reaching level with base of lowermost pectoral fin ray. Remarks. The subfamily includes only one genus, Cyclopterus., Published as part of Oku, Kanami, Imamura, Hisashi & Yabe, Mamoru, 2017, Phylogenetic relationships and a new classification of the family Cyclopteridae (Perciformes: Cottoidei), pp. 1-59 in Zootaxa 4221 (1) on page 53, DOI: 10.5281/zenodo.246721, {"references":["Bonaparte, C. L. (1831) Saggio di una distribuzione metodica degli animali vertebrati. Presso Antonio Boulzaler, Roma, 144 pp.","Linnaeus, C. (1758) Systema Naturae, 10 th edition. Fol. 1. Laurentii Salvii, Holmiae (= Stockholm), ii + 824 pp."]}

Subfamily Cyclopterinae Bonaparte, 1831 Cyclopterinae Bonaparte, 1831: 96 (type genus: Cyclopterus Linnaeus, 1758). Diagnosis. Head and body with many tubercles; generally high body having developed dorsal crest; supraoccipital crest present; first dorsal fin covered with very thick skin; first dorsal proximal pterygiophore lacking spines; ventral end of gill slit reaching level with base of lowermost pectoral fin ray. Remarks. The subfamily includes only one genus, Cyclopterus.

Imamura, Hisashi (2016): Rediagnosis of Onigocia grandisquama (Actinopterygii: Perciformes: Platycephalidae) and Comparison with Congeners. Species Diversity 21: 151-159, DOI: 10.12782/sd.21.2.151

Imamura, Hisashi (2015): Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species. Zootaxa 3904 (2): 151-207, DOI: http://dx.doi.org/10.11646/zootaxa.3904.2.1

Platycephalus richardsoni Castelnau, 1872 Common English name: Tiger flathead (Figs. 23–24; Fig. 9) Platycephalus richardsoni Castelnau, 1872: 82 (type locality: Melbourne markets, Vic, Australia); McCulloch, 1929: 400; Last et al., 1983: 333, fig. 28.21; Paxton & Hanley, 1989: 469; Grant, 2004: 188, unnumbered pl.; Hoese et al., 2006: 943; Imamura, 2006: 305, tab. 1; Gomon, 2008: 520, unnumbered fig. Neoplatycephalus grandis Castelnau, 1872: 87 (type locality: Melbourne markets, Vic, Australia); McCulloch, 1929: 402. Platycephalus macrodon Ogilby, 1885: 226 (type locality: Port Jackson, NSW, Australia); McCulloch, 1929: 401. Neoplatycephalus macrodon: Whitley, 1931 b: 159. Neoplatycephalus (Colefaxia) macrodon: Whitley, 1935: 249. Neoplatycephalus richardsoni: Whitley, 1964: 57; May & Maxwell, 1986: 272, unnumbered fig.; Hutchins & Swainston, 1986: 127, fig. 197; Knapp, 1991: 29, tab. 1; Kuiter 1993: 101, unnumbered fig.; Kuiter, 1994: 518, fig. 461. Material examined. 21 specimens (91.6–461 mm SL, from southeastern Australia): AMS B. 6541, holotype of Platycephalus macrodon Ogilby, 1885, 281 mm SL, Port Jackson, NSW; AMS E. 581, 165 mm SL, Oyster Bay, Tas (42.7 °S, 148.1 °E), date unknown; AMS E.3013, 2 specimens, 174–212 mm SL, Bass Strait, between Gabo and Babel Islands, date unknown; AMS I. 23871 -002, 194 mm SL, 4 miles off Whale Beach, NSW (33 ° 37 ’S, 151 ° 22 ’E), 54 m depth, 10 Oct. 1976; AMS I. 26026 -001, 266 mm SL, off Clarence River, NSW (29 ° 26 ’S, 153 ° 26 ’E), 11 Oct. 1985; AMS I. 33329 -002, 461 mm SL, Norah Head, NSW (33 ° 17 ’S, 151 ° 35 ’E), 1984; AMS I. 40292 -001, 231 mm SL, off Bermagui, NSW (36 ° 23 ’S, 150 ° 21 ’E), 428–468 m depth, 2 May 2000; AMS IB. 507, 440 mm SL, Wineglass Bay, Tas (42 ° 10 ’S, 148 ° 18 ’E), 18 June 1940; AMS IB. 7599, 379 mm SL, AMS IB. 7600, 310 mm SL, AMS IB. 7602, 321 mm SL, Lakes Entrance, Vic (37 ° 53 ’S, 148 °00’E), 1966; CSIRO 3534 -09, 246 mm SL, south of Disaster Bay, NSW (37 ° 23 ’S, 149 ° 58 ’E – 37 ° 22 ’S, 149 ° 58 ’E), 31–36 m depth, 12 Aug. 1993; CSIRO A 1612, 91.6 mm SL, 3 miles NE of Brush Island, NSW (35 ° 32 ’S, 150 ° 25 ’E), 25 Jan 1946; CSIRO A 1614, 113 mm SL, Eden, NSW (37 °04’S, 150 ° 56 ’E), 90 m depth, 28 Apr. 1953; CSIRO CA 135, 323 mm SL, Sydney fish market, NSW, 5 July 1977; NMV A 3786, 226 mm SL, 24 km southwest of Lakes Entrance, eastern Bass Strait, Vic (38 °03’S, 147 ° 50 ’E), 25–45 m depth, 1 Oct. 1983; QM I. 22311, 307 mm SL, Eden, NSW (37 °04’S, 149 ° 55 ’E), 14 Aug. 1985; QM I. 22302, 365 mm SL, QM I. 22307, 349 mm SL, Lakes Entrance, Vic (37 ° 53 ’S, 148 °E), 14 Aug. 1985. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; caudal-fin rays 13; pored lateral-line scales 65–74; gill rakers 2–4 + 10–13 = 12–17; orbital diameter 18.4–24.1 % HL; interorbital width 10.1–15.7 % HL; postorbital length 49.7–56.2 % HL; suborbital width 4.9–9.4 % HL; occipital and lower half of suborbital regions scaled in larger specimens; margin of interopercle smooth; one or two large caniniform teeth anteromedially on upper jaw; all or most pored lateral-line scales usually with two pairs of sensory ducts and exterior openings posteriorly; fleshy sensory tubes from suborbitals and preopercle not covering cheek region; swimbladder present; abdominal cavity dark brown or black. Description. Counts and measurements shown in Table 9. Holotype of P. macrodon Non-types AMS B. 6541 n = 20 SL (mm) 281 91.6–461 Counts: D 1 I + VIII + 0 I + VII–VIII + 0–I (often I + VIII + 0) D 2 14 13–14 (usually 14) A 14 13–14 (usually 14) P 1 2 + 10 + 8 = 20 1–2 + 9–12 + 7–8 = 19–22 (usually 20) C 13 13 LLS (spines) 70 (3) 65–74 (1–2) OBS – 81–95 GR 3 + 11 = 14 2–4 + 10–13 = 12–17 As % SL: HL 33.3 34.1–36.2 PDL 34.3 34.4–37.6 LD 1 B 21.7 16.4–22.2 LD 2 B 32.5 30.0– 33.1 LAB 33.4 30.2–35.1 SNL 9.1 8.8–10.3 OD 6.7 6.4–8.6 UJL 12.3 12.6–14.3 LJL 17.0 16.5–19.1 IW 4.4 3.6–5.5 POL 17.8 17.2–19.3 SW 2.0 1.7–3.2 P 1 L 16.0 14.6 –18.0 P 2 L 23.8 21.9–26.1 CL 15.6 15.1–19.3 As % HL: SNL 27.2 26.0– 28.9 OD 20.2 18.4–24.1 UJL 36.8 36.9–39.9 LJL 51.2 48.5–53.4 IW 13.2 10.1–15.7 POL 53.4 49.7–56.2 SW 6.1 4.9–9.4 Snout, interorbit and occipital region scaled; lower half of suborbital region naked in 165 mm SL or smaller specimens, scaled in larger specimens; area anteroventral to eye naked. Interorbit narrower than orbital diameter. Upper iris lappet simple, weakly pointed; lower absent. Nasal spine usually absent, rarely one spine present in some specimens. Preorbital spine present in smallest (92 mm SL) specimen, absent in larger specimens. One suborbital spine present below posterior margin of eye in 165 mm SL or smaller specimens, absent in larger specimens (except in 379 mm SL specimen with two spines below posterior margin of eye). Three preopercular spines present; middle spine usually slightly longer than uppermost spine (more than twice length in some specimens, but not reaching opercular margin); lowermost small. Supplemental preopercular spine usually absent (present in 93 mm and 307 mm SL specimens). Interopercular flap absent; margin of interopercle smooth. Maxilla usually not reaching, or reaching just beyond anterior margin of pupil (to middle of eye in some specimens). Upper jaw with one or two large caniniform teeth anteromedially. Palatine teeth in two rows, villiform or small conical in outer row, small to moderate caniniform in inner row. Vomerine teeth in one row anteriorly, one or two rows posteriorly, becoming larger posteriorly. All or most pored lateral-line scales usually with two pairs of sensory ducts and exterior openings posteriorly, some scales sometimes with one, three or four pairs of ducts and openings. Fleshy sensory tubes from suborbitals and preopercle not covering cheek region. Posterior tip of pelvic fin reaching from anus to base of third anal-fin ray. Posterior margin of caudal fin mostly straight in smaller specimens, slightly concave in larger specimens. Swimbladder present. Color in alcohol. Ground color of head and body reddish-brown, pale or dark brown above, paler brown below. Side of body with or without a single row of purplish spots tending to form a band, or with a dark brown or purple band. First and second dorsal fins with distinct or indistinct pale brown to brown spots. Pectoral fin dusky, upper portion with brownish spots tending to form bands; lower margin of pectoral fin paler. Pelvic fin pale to dark brown; lower margin of fins paler. Anal fin pale or with melanophores along rays. Caudal fin dusky, or brown to dark brown; upper margin with or without indistinct pale brown or brown spots. Abdominal cavity dark brown or black. Distribution. Known from southeastern Australia, from off Clarence River, NSW (29 ° 26 ’S) to Investigator Strait, SA (137 ° 10 ’E), including Vic and Tas, on the continental shelf in depths from ca. 10 to 428 m (e.g., May & Maxwell, 1986; Hoese et al., 2006; Gomon, 2008; this study). Size. Maximum length 60 cm SL (Gomon, 2008). The largest specimen examined during the present study was 461 mm SL (caudal fin broken). Remarks. Platycephalus richardsoni is most similar to P. conatus, but differs in having 12–17 gill rakers in total and all or most pored lateral-line scales usually with two pairs of sensory ducts and exterior openings posteriorly (see Remarks under P. conatus). The latter character also distinguishes P. richardsoni from other species of Platycephalus characterized by pored lateral-line scales usually with a single pair of sensory ducts and exterior openings. Although Paxton & Hanley (1989), Kuiter (1994) and Hoese et al. (2006) considered P. mortoni Macleay, 1883 to be a junior synonym of P. richardsoni, Imamura (2013 a) found the former to be conspecific with Platycephalus fuscus Cuvier in Cuvier & Valenciennes, 1829. In contrast, the junior synonymy of Neoplatycephalus grandis Castelnau, 1872 under P. richardsoni, as already pointed out by several authors (Paxton & Hanley, 1989; Kuiter, 1994; Hoese et al., 2006), is reconfirmed here, both nominal taxa having “several arborescent tubes” on the lateralline scales (see original descriptions of both species by Castelnau, 1872). Unfortunately, the whereabouts of the type specimens of P. richardsoni is unknown at present and the syntypes of N. grandis were apparently lost (e.g., Hoese et al. 2006). Platycephalus macrodon (Fig. 24) was also regarded as a junior synonym of P. richardsoni by Paxton & Hanley (1989), Kuiter (1994) and Hoese et al. (2006). This synonymy is supported here due to the holotype of P. macrodon agreeing closely with the latter in pored lateral-line scale characteristics, and having no remarkable departures from the counts and proportional measurements accordied to P. richardsoni (Table 9)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 183-186, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Castelnau, F. L. (1872) Contributions to the ichthyology of Australia. No. 1. The Melbourne fish market. Proceedings of the Royal Zoological Acclimatization Society of Victoria, 1, 29 - 242.","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Ogilby, J. D. (1885) Descriptions of new fishes from Port Jackson. Proceedings of the Linnean Society of New South Wales, 10, 25 - 230.","Whitley, G. P. (1931 b) Studies in ichthyology. No. 5. Records of the Australian Museum, 18, 131 - 160. [plates. 20 - 21]","Whitley, G. P. (1935) Studies in ichthyology. No. 9. Records of the Australian Museum, 19, 215 - 250. [plates. 18]","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Kuiter, R. H. (1994) Family Platycephalidae. In: Gomon, M. F., Glover, J. C. M. & Kuiter, R. H. (Eds.), The fishes of Australia's south coast. State Print, Adelaide, pp. 514 - 522.","Macleay, W. (1883) Notes on a collection of fishes from the Burdekin and Mary rivers, Queensland. Proceedings of the Linnean Society of New South Wales, 8, 199 - 213.","Imamura, H. (2013 a) Platycephalus mortoni Macleary 1883, a junior synonym of Platycephalus fuscus Cuvier 1829 (Teleostei: Platycephalidae). Ichthyological Research, 60, 282 - 286. http: // dx. doi. org / 10.1007 / s 10228 - 013 - 0348 - 9","Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp."]}

Platycephalus angustus Steindachner, 1866 Common English name: Steindachner’s flathead (Fig. 33) Platycephalus angustus Steindachner, 1866 a: 20 (original type locality: Suriname, South America; revised type locality owing to designation of neotype: northeast of Darbilla Creek, Millingimbi, Northern Territory, Australia); Steindachner, 1866 b: 20; Steindachner, 1866 c: 312; Imamura, 2012: 5 th p. Material examined. Neotype (designated by Imamura, 2012): NTM S. 11362 -026, 345 mm SL, northeast of Darbilla Creek, Millingimbi, NT, Australia (12 °09’S, 134 ° 56 ’E), 0–2 m depth, 24 July 1984. Other specimens: 23 specimens, 53.6–378 mm SL, from northern Australia and western New Guinea, listed in Imamura (2012). Diagnosis. A species of Platycephalus with the following combination of characters: first dorsal fin with two small isolated spines anteriorly; dorsal and anal soft rays usually 13; orbital diameter 11.2–21.5 % HL; interorbital width 7.3–17.3 % HL; postorbital length 55.5–67.8 % HL; suborbital length 4.7–7.2 % HL; snout, area anteroventral to eye, interorbit, and occipital region scaled; upper iris lappet simple, triangular; a finger-like interopercular flap present; upper jaw without large caniniform teeth; teeth absent on dorsal surface of anterolateral edge of upper jaw in specimens ca. 76 mm SL or longer; palatine teeth in two rows; number of vomerine tooth rows tending to increase with growth, in two to four rows in 106–184 mm SL specimens and forming a single broad band in larger specimens; three to five dark brown to black bands and spots on caudal fin. Distribution. Known from northern Australia, from Wyndham, Kimberley region, WA (128 ° 10 ’E) to Cape Pallarenda, Townsville, Qld (146 ° 46 ’E) and Irian Jaya, western New Guinea, inhabiting creeks and rivers, river mouths, estuaries and coastal areas including mudflats influenced by fresh water, in depths of 0.5–2 m (Imamura, 2012). Size. Recorded maximum length 378 mm SL (432 mm TL) (Imamura, 2012; this study). Remarks. Platycephalus angustus had been overlooked until Imamura (2012) demonstrated its validity and provided a detailed description. Platycephalus angustus can be separated from P. cultellatus, P. indicus, P. westraliae, P. endrachtensis and Platycephalus sp. 1 and sp. 2 (sensu Nakabo, 2002), having usually 13 dorsal and anal soft rays, a finger-like interopercular flap, three to five dark brown to black bands (and spots), the first dorsal fin with two small isolated spines anteriorly, a small orbit (orbital diameter 11.2–21.5 % HL), broader interorbital space (interorbital width 7.3–17.3 % HL), longer and narrower postorbital region (postorbital length 55.5–67.8 % HL, suborbital width 4.7–7.2 % HL), and vomerine tooth rows tending to increase with growth from two to four rows in 106–184 mm SL specimens and forming a single broad band in longer specimens. Platycephalus angustus is unique in having exposed teeth on the dorsal surface of the anterolateral edge of the upper jaw in specimens ca. 76 mm SL or larger. This character is not known in any other species of Platycephalus, all of which have upper jaw teeth restricted to the anterior margin and ventral surface of the jaw (Imamura, 2012)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 197-198, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Steindachner, F. (1866 a) Fortsetzung der ichthyologische Notizen. Anzeiger der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftlichen Classe, 3, 19 - 20.","Steindachner, F. (1866 b) Ichthyologische Notizen (III). Uber einige neue Fischarten aus Sudamerika. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, 53, 208 - 214. [plates. 1 - 2]","Steindachner, F. F. (1866 c) New fishes from South America. Annals and Magazine of Natural History, 17, 311 - 312. [Series 3]","Imamura, H. (2012) Redescription of Platycephalus angustus Steindachner 1886 (Teleostei: Platycephalidae), a valid flathead in northern Australia and New Guinea. Ichthyological Research, 60, 112 - 121. http: // dx. doi. org / 10.1007 / s 10228 - 012 - 0319 - 6","Nakabo, T. (2002) Platycephalidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 601 - 602."]}

Platycephalus chauliodous Knapp, 1991 Common English name: Bigtooth flathead (Figs. 16–17; Table 6) Platycephalus chauliodous Knapp, 1991: 24, figs. 1–3 (type locality: southern WA); Hoese et al., 2006: 941; Imamura, 2006: 304, tab. 1. Neoplatycephalus sp.: Hutchins & Swainston, 1986: 127, fig. 196. Platycephalus sp.: Hutchins & Thompson, 1983: 78, fig. 115. Material examined. Holotype: AMS I. 20231 -007, 313 mm SL, Cockburn Sound, 0.5 km south of Carnac Island, WA, Australia (32 ° 10 ’S, 115 ° 40 ’E), 6–7 m depth, 27 Mar. 1978. Paratypes (5 specimens, 65.6–460 mm SL, from southwestern Australia): USNM 307398, 75.6 mm SL, off Garden Island, WA (32 ° 12 ’S, 115 ° 40 ’E), 10 Apr. 1976; WAM P. 24172 -001, 460 mm SL, Cockburn Sound, Carnac Island, 3 miles off Freemantle, WA (32 °07’S, 115 ° 40 ’E), ca. 10 m depth, 30 Dec. 1973; WAM P. 25798 -008, 65.6 mm SL, Eagle Bay, WA (33 ° 33.5 ’S, 115 °04’E), 0–4 m depth, 21 Oct. 1976; WAM P. 29879 -001, 336 mm SL, WA (31 ° 50 ’S, 115 ° 44 ’E), ca. 40 m depth, Dec. 1988; WAM P. 29944 -001, 398 mm SL, Cowaramup Beach, WA (33 ° 47 ’S, 115 °00’E), ca. 8 m depth, 19 Feb. 1989. Other specimens (3 specimens, 99.7–426 mm SL, from southwestern Australia): CSIRO CA 696, 424 mm SL, Great Australian Bight, WA (32 ° 30 ’S, 126 ° 45 ’E), 36 m depth, 13 Nov. 1996; WAM P. 28517 -005, 99.7 mm SL, Dunsborough, WA (33 ° 35 ’S, 115 °06’E), 12 Apr. 1985; WAM P. 31595 -001, 426 mm SL, Straggler’s Reef, off Fremantle, WA, Jan. 1999. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal-fin rays usually 14; anal-fin rays usually 13; caudal-fin rays 9–11 (usually 10–11); pored lateral-line scales 59–68; orbital diameter 11.3–21.8 % HL; interorbital width 9.3–17.5 % HL; postorbital length 50.0– 61.1 % HL; snout, area anteroventral to eye, interorbit, occipital and lower half of suborbital naked; one or two large caniniform teeth anteromedially on upper jaw; margin of interopercle scalloped; fleshy sensory tubes from suborbitals and preopercle slightly developed, partly covering dorsal and ventral margins of cheek region. Description. Counts and measurements shown in Table 6. Data for all specimens presented first, followed by holotype condition in parentheses. Snout, area anteroventral to eye, interorbit, and occipital and lower half of suborbital regions naked. Interorbit narrower than orbital diameter in 99.7 mm SL or smaller specimens, wider in larger specimens (including holotype). Upper iris lappet broad, bilobed; lower weakly convex. Nasal, preorbital, preocular, suborbital and supplemental preopercular spines absent. Lower preopercular spine slightly shorter than upper in 99.7 mm SL or smaller specimens, longer than upper, not reaching opercular margin in larger specimens (including holotype). Interopercular flap absent; margin of interopercle scalloped. Maxilla usually reaching to or slightly beyond anterior margin of pupil in 99.7 mm SL or smaller specimens, to middle of eye in larger specimens [up to 426 mm SL (including holotype)] and to posterior margin of eye in largest (460 mm SL). Upper jaw with one or two (two) large caniniform teeth anteromedially. Palatine teeth in two rows, villiform in outer row, moderate or large caniniform (large caniniform) in inner row. Vomerine teeth usually in one row (including holotype), becoming larger posteriorly; three larger teeth positioned medially to tooth row in 398 mm SL specimen. Fleshy sensory tubes from suborbitals and preopercle slightly developed, partly covering dorsal and ventral margins of cheek region. Posterior tip of pelvic fin reaching from anus to base of third anal-fin ray (base of second anal-fin ray). Posterior margin of caudal fin mostly straight or slightly rounded (mostly straight). Color in alcohol. Ground color of head and body of holotype (Fig. 17) pale brown above, slightly paler below. Dorsal surface of head and body with many small brown and white spots. First and second dorsal fins with whitish and pale brownish spots along fin rays. Pectoral fin with distinct whitish and pale brownish spots anterodorsally; other portions of pectoral fin with indistinct paler spots tending to form bands. Pelvic fin with moderate pale brown spots. Anal fin pale. Caudal fin with small pale brown spots, small whitish spots basally. Distribution. Known only from southwestern Australia, from Great Australian Bight (149 ° 16 ’E), WA to near Perth (31 ° 50 ’S), infrequently taken on shallow coastal reefs in depths from 4 to ca. 40 m (e.g., Hutchins & Swainston, 1986; Knapp, 1991; Hoese et al., 2006; this study). Size. Previously known maximum length 46.5 cm (Hutchins & Swainston, 1986). One specimen examined during the present study measured 460 mm SL (541 mm TL). TABLE 6. Comparison of counts and proportional measurements of Platycephalus chauliodous. *Usually 10, nine in smallest specimen. Remarks. As pointed out by Knapp (1991), P. chauliodous has one fewer anal-fin rays (13–14, usually 13) than second dorsal-fin rays (14–15, usually 14), an unusual characteristic in Platycephalus (usually the same number of second dorsal- and anal-fin rays). Also unusual is a scalloped interopercular margin that is found elsewhere only in P. orbitalis among species of Platycephalus. Apart from the above characters, P. chauliodous resembles P. aurimaculatus, P. conatus and P. richardsoni in having one or two large caniniform teeth anteromedially on the upper jaw (large caniniform teeth absent on upper jaw in other species of Platycephalus). The first-mentioned can be easily distinguished from the other three species in having 9–11 caudal-fin rays (usually 10–11) (vs. 12 or 13), 59–68 pored lateral-line scales (75–90 in P. aurimaculatus, 70–81 in P. conatus and 65–74 in P. richardsoni), the occipital region naked (vs. scaled), and fleshy sensory tubes from the suborbitals and preopercle slightly developed, partly covering the dorsal and ventral margins of the cheek region (vs. tubes not covering cheek region). The orbital diameter/ interorbital width relationship is also valuable for separating P. chauliodous from the above three species; viz. P. chauliodous has a narrower orbit and broader interorbit, the former dimension being greater than the latter in smaller specimens but less in 312 mm SL or larger specimens, whereas the orbit diameter is consistently greater than the interorbital width in all examined specimens of the other species (orbital diameter 11.3–21.8 % HL and interorbital width 9.3–17.5 % HL in P. chauliodous vs. 16.3–22.4 % and 7.4–11.7 % in P. aurimaculatus, 16.9–22.6 % and 7.4–14.1 % in P. conatus, and 18.4–24.1 % and 10.1–15.7 % in P. richardsoni) (Fig. 18 A). In addition, P. chauliodous has a greater postorbital length (50.0– 61.1 % HL vs. 49.1–56.9 %, 49.7–56.2 % and 49.7–56.2 %, respectively) (Fig. 18 B)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 174-176, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp."]}

Platycephalus aurimaculatus Knapp, 1987 Common English name: Toothy flathead (Figs. 19���20; Table 7) Platycephalus sp.: Last et al., 1983: 334, fig. 28.23. Neoplatycephalus sp. 1: May & Maxwell, 1986: 273, unnumbered fig. Platycephalus (Neoplatycephalus) aurimaculatus Knapp, 1987: 53 (type locality: Bass Strait, east of King Island, Tas, Australia). Neoplatycephalus aurimaculatus: Knapp, 1991: 29, tab. 1; Kuiter 1993: 102, unnumbered fig. Platycephalus aurimaculatus: Kuiter, 1994: 516, fig. 459; Hoese et al., 2006: 940; Imamura, 2006: 304, tab. 1; Gomon, 2008: 518, unnumbered fig. Platycephalus conatus (not Waite & McCulloch, 1915): Imamura, 2006: 304, tab. 1 (in part). Material examined. Holotype: NMV A 1404, 279 mm SL, 60 km east-northeast of Bold Head, King Island, Central Bass Strait, Tas, Australia (39 �� 56 ��� 25 ���S ��� 39 �� 53 ��� 52 ���S, 144 �� 48 ���03���E ��� 144 �� 47 ��� 58 ���E), 29.4-49.4 m depth, 3 Feb. 1981. Paratypes (4 specimens, 211���322 mm SL, from southeastern Australia): AMS I. 26328 -001 (ex. NMV A 1389), 322 mm SL, off Cape Otway, Bass Strait, Vic (39 ��05���S, 143 �� 26 ���E), 82 m depth, 31 Jan. 1981; NMV A3733, 2 specimens, 211���229 mm SL, 3.2 km west of Sandringham, Port Phillip Bay, Vic (37 �� 57 ���S, 144 �� 57 ���E), 30 Mar. 1971; USNM 280181, 315 mm SL, Bass Strait, NSW, Feb. 1981. Other specimens (17 specimens, 121���489 mm SL, from southern Australia): AMS I. 21309 -001, 377 mm SL, southeast of Bass Strait, Tas (40 �� 25 ���S, 147 �� 35 ���E), 46 m depth, 16 Oct. 1979; AMS E. 750, 233 mm SL, Marsden Point, Kangaroo Island, SA (35 �� 34 ���S, 137 �� 38 ���E), 19 Aug. 1909; AMS IB. 2118, 296 mm SL, Geelong, Vic (38 �� 10 ���S, 144 �� 21 ���E), 11 Apr. 1948; CSIRO 3698, 380 mm SL, south of Point Cuvier, Great Australian Bight, WA (33 �� 37 ���S, 124 �� 48 ���E ��� 33 �� 35 ���S, 124 �� 47 ���E), 50 m depth, 29 Nov. 1981; CSIRO H 3796 -01, 489 mm SL, Disaster Bay, NSW (37 �� 17 ������ 37 �� 18 ���S, 149 �� 59 ���E), 26���33 m depth, 13 Nov. 1996; NMV A1424, 2 specimens, 176���191 mm SL, NMV A 24057, 181 mm SL, 5 km south-southeast of Cape Woolomai, Phillip Island, Vic (38 �� 43 ���S, 145 �� 34 ���E), 46 m depth, 17 Jan. 1980; NMV A 1568, 245 mm SL, 46 km east-northeast of North Point, Flinders Island, Eastern Bass Strait, Tas (39 �� 31 ��� 21 ������ 39 �� 34 ��� 19 ���S, 148 �� 24 ���01������ 148 �� 24 ���07���E), 19.5���36.5 m depth, 8 Feb. 1981; NMV A1581, 3 specimens, 121���132 mm SL, Vic, 13 Nov. 1979; QM I. 22300, 285 mm SL, NSMT-P 112680, 401 mm SL, Great Australian Bight, SA (32 �� 21.3 ���S, 130 �� 2.1 ���E), 62 m depth, 30 Nov. 1975; QM I. 22304, 316 mm SL, Lakes Entrance, Vic (37 �� 53 ���S, 148 ��E), 14 Aug. 1984; QM I. 22309, 390 mm SL, Western Port, Vic (38 �� 26 ���S, 145 ��08���E), 14 Aug. 1984; WAM P. 30105 -002, 464 mm SL, Capel, WA (33 �� 25 ���S, 115 �� 21 ���E), 6 Sep. 1990. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; caudal-fin rays 12���13; pored lateral-line scales 75���90; gill rakers 1���2 + 6���7 = 7���9; orbital diameter 16.3���22.4 % HL; interorbital width 7.4���11.7 % HL; postorbital length 49.1���56.9 % HL; suborbital width 6.7 ���9.0% HL; occipital region scaled; lower half of suborbital region naked; margin of interopercle smooth; one to three large caniniform teeth anteromedially on upper jaw; all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly; fleshy sensory tubes from suborbitals and preopercle not covering cheek region; swimbladder absent; abdominal cavity pale, slightly silvery. Description. Counts and measurements shown in Table 7. Data for all specimens presented first, followed by holotype condition in parentheses. SL (mm) 279 211���322 121���489 Counts: Snout, area anteroventral to eye, anterior one-fourth to entire area of interorbit (anterior half) and lower half of suborbital region naked; occipital region scaled. Interorbit narrower than orbital diameter. Upper iris lappet simple, moderately long; lower weakly convex, or absent (absent). Nasal, preorbital and supplemental preopercular spines absent. One suborbital spine present below posterior margin of eye in smallest 121 mm SL specimen, absent in others. Lower preopercular spine longer than upper, not reaching opercular margin. Interopercular flap absent; margin of interopercle smooth. Maxilla extending beyond anterior margin of pupil; some specimens with maxilla reaching to middle of eye (slightly beyond anterior margin of pupil in holotype). Anteromedial portion of upper jaw with one to three (one) large caniniform teeth. Palatine teeth in two rows, villiform or small conical (small conical) in outer row, small to large caniniform (moderately caniniform) in inner row. Vomerine teeth in a single row. Fleshy sensory tubes from suborbitals and preopercle not covering cheek region. Posterior tip of pelvic fin reaching from anus to base of third anal-fin ray (base of second anal-fin ray). Posterior margin of caudal fin mostly straight, or slightly rounded (mostly straight). Swimbladder absent. Color in alcohol. In holotype (Fig. 20), ground color of head and body pale brown above, slightly paler below. Dorsal surface of head and body with scattered small white spots. Side of body with greenish-gray band. First and second dorsal fins with melanophores along fin rays. Pectoral fin pale with brown spots tending to form bands; lower margin of pectoral fin paler. Pelvic fin pale brown; its lower margin paler. Anal fin with melanophores along rays. Caudal fin with pale brown spots and bands posteriorly. In other specimens (Fig. 19), first and second dorsal fins with melanophores or small indistinct pale brown spots along fin rays. Caudal fin with dark brown spots, or pale brown spots, bands and/or indistinct markings. Abdominal cavity pale, slightly silvery. Distribution. Known only from southern Australia, from to Disaster Bay, NSW (149 �� 59 ���E) to Capel, WA (115 �� 21 ���E), including Vic, Tas and SA, in bays and coastal waters in depths from 10 to 160 m (e.g., Knapp, 1987; Hoese et al., 2006; Gomon, 2008; this study). Size. Previously reported maximum length 55 cm SL (Gomon, 2008). One specimen examined during the present study measured 489 mm SL (565 mm TL). Remarks. Platycephalus aurimaculatus is most similar to P. conatus and P. richardsoni in having usually 14 anal-fin rays, 12 or 13 caudal-fin rays, the occipital region scaled, the margin of the interopercle smooth and large caniniform teeth anteromedially on the upper jaw, the combination of such characters not occurring in other species of Platycephalus. The present species is easily distinguished from P. conatus and P. richardsoni in lacking a swimbladder (vs. present), and having the lower half of the suborbital region naked (vs. scaled, but scales not developed in 165 mm SL or smaller specimens of P. richardsoni) and the abdominal cavity pale and slightly silvery (vs. dark brown or black). Postorbital length and suborbital width at any given length may also help to separate P. aurimaculatus and P. conatus (postorbital length 49.1���56.9 % HL and suborbital width 6.7 ���9.0% HL in the former vs. 49.7���56.2 % HL and 5.4���9.7 % HL, respectively, in the latter), although broad overlaps occur in both proportions (Fig. 21). Platycephalus aurimaculatus is distinguishable from P. richardsoni in having 75���90 pored lateral-line scales (vs. 65���74), 7���9 total gill rakers (vs. 12���17), a narrower interorbit (7.4���11.7 % HL in P. aurimaculatus vs. 10.1���15.7 % HL in P. richardsoni) (Fig. 18 A) and all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly (vs. most scales with two pairs of ducts and openings)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 177-180, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Knapp, L. W. (1987) New Australian fishes. Part 13. Two new species of Platycephalidae. Memoirs of Museum Victoria, 48, 53 - 55.","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp.","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Kuiter, R. H. (1994) Family Platycephalidae. In: Gomon, M. F., Glover, J. C. M. & Kuiter, R. H. (Eds.), The fishes of Australia's south coast. State Print, Adelaide, pp. 514 - 522.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Waite, E. R. & McCulloch, A. R. (1915) The fishes of the South Australian Government Trawling Cruise, 1914. Transactions Royal Society of South Australia, 9, 455 - 476. [plates. 12 - 15]"]}

Platycephalus westraliae (Whitley, 1938) Common English name: Yellowtail flathead (Figs. 31���32; Table 12) Trudis bassensis westraliae Whitley, 1938: 199 (type locality: Swan River estuary, WA, Australia); Whitley, 1964: 57. Platycephalus endrachtensis (not Quoy & Gaimard, 1825): Taylor, 1964: 287, pl. 65; Hutchins & Thompson, 1983: 78, fig. 110; Gloerfelt-Tarp & Kailola, 1984: 113, fig. 12, unnumbered fig.; Sainsbury et al. 1985: 112, unnumbered fig.; Hutchins & Swainston, 1986: 127, fig. 205; Allen & Swainston, 1988: 52, fig. 278; Paxton & Hanley, 1989: 468; Knapp, 1991: 29, tab. 3; Allen, 1997: 80, pl. 21 - 17; Knapp, 1999: 2408, unnumbered fig.; Hoese et al., 2006: 941; Imamura, 2006: 305, tab. 1. Platycephalus westraliae: Imamura, 2008: 405, figs. 1 a, 6 a, 7. Platycephalus bassensis (not Cuvier in Cuvier & Valenciennes, 1829): Paxton & Hanley, 1989: 468 (in part); Hoese et al., 2006: 940 (in part). Platycephalus indicus (not Linnaeus, 1758): Grant, 2004: 192, pl. 87. Material examined. Holotype: AMS IA. 7189, 98.7 mm SL, Swan River estuary, WA, Australia. Paratype: AMS IA. 7190, 85.8 mm SL, collected with holotype. Other specimens (31 specimens, 63.2���553 mm SL, from western to southeastern Australia): 23 specimens listed in Imamura (2008); 8 additional specimens: AMS I. 20751 -005, 389.3 mm SL, 3 miles northwest of Lizard Island, Qld (14 �� 38 ���S, 145 �� 24 ���E), 25 m depth, 8 Feb. 1979; CSIRO CA 767, 200 mm SL, Thursday or Prince of Wales Islands, Torres Strait, Qld (10 �� 40 ���S, 142 �� 15 ���E); CSIRO CA 1162, 375 mm SL, east of Barrow Island, WA, date unknown; NTM S. 11255 -003, 330 mm SL, Caiman Creek, Port Essington, NT (11 �� 14 ���S, 132 �� 12 ���E), 13 May 1983; NTM S. 12941 -001, 369 mm SL, Arafura Sea, NT (11 �� 29 ���S, 133 �� 36 ���E), 22 Oct. 1990; NTM S. 16160 -041, 169 mm SL, Port Patterson, Quail Island, NT (12 �� 31.36 ���S, 130 �� 25.59 ���E), 7 Dec. 2002; QM I.28582, 1 of 20 specimens, Pine River mouth, Qld (27 �� 17 ���S, 153 ��03���E), 27 Oct. 1997; WAM P. 30645 -001, 553 mm SL, Burswood Casino, Perth, WA. Diagnosis. A species of Platycephalus with the following combination of characters: first dorsal fin with a single small isolated spine anteriorly; second dorsal- and anal-fin rays usually 13; interorbital width 6.3 ���17.0% HL; postorbital length 51.2���60.9 % HL; snout, area anteroventral to the eye, interorbit, and occipital region scaled; upper iris lappet broad, usually slightly bilobed; a finger-like interopercular flap present; upper jaw without large caniniform teeth; teeth absent on dorsal surface of anterolateral edge of upper jaw; palatine teeth in two rows; vomerine teeth in one or two (usually one) rows anteriorly, one to four (usually one or two) posteriorly; three or four longitudinal dark bands on caudal fin; upper lobe of caudal fin with yellow blotch when fresh. Description. Counts and measurements shown in Table 12. Data for all specimens presented first, followed by holotype condition in parentheses. SL (mm) 98.7 85.8 63.2���553 Counts: *Abnormal condition. Snout, area anteroventral to eye, interorbit and occipital region scaled; lower half of suborbital region naked. Interorbit narrower than orbital diameter in 369 mm SL or smaller specimens, wider than latter in larger specimens. Upper iris lappet broad, usually slightly bilobed (including holotype), rarely pointed or mostly straight in some specimens; lower weakly convex. Nasal spine present in smaller specimens, tending to disappear with growth, absent in 330 mm SL or larger specimens (present in holotype). Preorbital spine absent. One suborbital spine usually present below posterior margin of eye in 149 mm SL or smaller specimens (including holotype), usually absent in larger specimens; one additional suborbital spine sometimes present below and slightly anterior to middle of eye (absent in holotype). Lower preopercular spine slightly longer than upper, not reaching opercular margin. Supplemental preopercular spine usually present in 123 mm SL or smaller specimens (including holotype), absent in larger specimens. Finger-like interopercular flap present; margin of interopercle smooth. Maxilla reaching from just before anterior margin of pupil to posterior margin of eye (reaching to near anterior margin of pupil). Upper jaw with moderate or large conical or small caniniform teeth anteromedially (large conical teeth). Palatine teeth in two rows, villiform in outer row, moderate conical in inner row. Vomerine teeth usually in one (including holotype), rarely two rows (in 375 mm SL specimen) anteriorly, usually one or two (two) rows, rarely three (in 330 mm SL specimen) posteriorly, tending to become larger posteriorly. Fleshy sensory tubes from suborbitals and preopercle not covering cheek region. Posterior tip of pelvic fin reaching from anus to base of fourth anal-fin ray (base of third anal-fin ray). Posterior margin of caudal fin slightly rounded, mostly straight, or slightly concave (mostly straight). Color in alcohol. Holotype discolored, but retaining brownish spots on side of body, and first and second dorsal, pectoral and pelvic fins, melanophores between ninth and last anal-fin rays, and three black longitudinal bands on caudal fin (Fig. 32). In other specimens (Fig. 31), ground color of head and body pale brown to dark brown above, paler below. Dorsal surface of head and body with small dark spots. Side of body with or without gray or brown band, or spots. Body below second dorsal fin with or without distinct or indistinct darker bands. First and second dorsal, pectoral and pelvic fins with small pale to dark brown spots. Anal fin pale in 291 mm SL or smaller specimens, with melanophores on membranes between 11 th to last rays in 330 to 389 mm SL specimens, dusky in largest (553 mm SL) specimen. Caudal fin with three or rarely four (in a few specimens) brown to black longitudinal bands. Upper lobe of caudal fin with yellow blotch when fresh (e.g., Gloerfelt-Tarp & Kailola, 1984; Sainsbury et al. 1985; Hutchins & Swainston 1986). Distribution. Known from Australia (except Vic, Tas and SA), from Fremantle, WA (32 ��03���S) across northern Australia to Port Hacking, NSW (32 ��04���S), including NT and Qld, and also from the southern coasts of Papua New Guinea and Java (e.g., Hutchins & Swainston 1986; Knapp 1999; Imamura, 2008; this study). Size. Maximum length 1 m (Hutchins & Swainston, 1986). The largest specimen examined during the present study was 553 mm SL (635 mm TL). Remarks. Platycephalus westraliae had been considered as a subspecies (Whitley, 1938, 1964) or junior synonym of P. bassensis (Paxton & Hanley, 1989; Hoese et al., 2006) until Imamura (2008) showed it to be a valid species in its own right. In addition, up to that time, specimens had often been identified as ��� P. endrachtensis ���. Of the species of Platycephalus with usually 13 second dorsal- and anal-fin rays, and two or more brown to black longitudinal bands on the caudal fin, P. westraliae differs from P. angustus and P. cultellatus in having the first dorsal fin with a small isolated spine anteriorly [usually two in the latter (but rarely one in P. angustus)] and a shorter postorbital region at a given length (51.2���60.9 % HL in P. westraliae vs. 55.5���67.8 % HL in P. angustus and 51.8���66.8 % HL in P. cultellatus) (Fig. 30 B). The former character is also useful for separating P. westraliae from Platycephalus sp. 1 and 2 (sensu Nakabo, 2002). Platycephalus westraliae also differs from P. angustus in having teeth absent on the dorsal surface of the anterolateral edge of the upper jaw (teeth present in P. angustus), from P. australis sp. nov. (described below) in having a shorter postorbital region at a given length (postorbital length 51.2���60.9 % HL in P. westraliae vs. 51.6���63.6 % HL in P. australis sp. nov.) (Fig. 30 B) and from P. indicus in having a narrower interorbit at a given length (interorbital width 6.3 ���17.0% HL in P. westraliae vs. 7.2���18.4 % HL in P. indicus) (Fig. 30 A). Platycephalus westraliae is separable from the above mentioned six species in having a broad bilobed upper iris lappet [vs. usually simple triangular, although sometimes broad bilobed in P. australis sp. nov. and P. i n di c us (see Remarks under P. endrachtensis)]. Finally, in fresh specimens the yellow blotch on the upper portion of the caudal fin is useful for distinguishing P. westraliae, although P. australis sp. nov. and P. indicus both have a yellow band on the middle of the caudal fin when fresh (e.g., Knapp, 1999; Imamura, 2009; this study)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 194-197, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Whitley, G. P. (1938) Studies in ichthyology. No. 11. Records of the Australian Museum, 20, 195 - 199. [plates. 21]","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","Quoy, J. R. C. & Gaimard, J. P. (1825) Chapter IX. Description des Poissons. In: Freycinet, L. de. Voyage autour du Monde … execute sur les corvettes de L. M. \" L'Uranie \" et \" La Physicienne, \" pendant les annees 1817, 1818, 1819 et 1820. Par is, pp. 192 - 401. [plates. 43 - 65]","Taylor, W. R. (1964) Fishes of Arnhem Land. In: Specht, R. L. (Ed.), Records of the American-Australian scientific expedition to Arnhem Land, Vol. 4. Melbourne University Press, Melbourne, pp. 45 - 307.","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp.","Gloerfelt-Tarp, T. & Kailola, P. J. (1984) Trawled fishes of southern Indonesia and northern Australia. The Australian Development Assistance Bureau, the Directorate General of fisheries, Indonesia and the German Agency for Technical Cooperation, Jakarta, xvi + 2 plates. + 406 pp.","Sainsbury, K. J., Kailola, P. J. & Leyland, G. G. (1985) Continental shelf fishes of northern and north-western Australia. Clouston and Hall and Peter Pownall Fisheries Information Service, Canberra, vii + 375 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Allen, G. R. & Swainston, R. (1988) The marine fishes of north-western Australia. A field guide for anglers and divers. Western Australian Museum, Perth, vi + 201 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Allen, G. R. (1997) Marine fishes of tropical Australia and south-east Asia. A field guide for anglers and divers. Western Australian Museum, Perth, 292 pp.","Knapp, L. W. (1999) Platycephalidae. In: Carpenter, K. E. & Niem, V. H. (Eds.), FAO species identification guide for fishery purposes. The living marine resources of the western Central Pacific. Vol. 4. Bony fish. Part 2 (Mugilidae to Carangidae). FAO, Rome, pp. 2385 - 2421.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Imamura, H. (2008) Synonymy of two species of the genus Platycephalus and validity of Platycephalus westraliae (Pisces: Scorpaeniformes). Ichthyological Research, 55, 399 - 406. http: // dx. doi. org / 10.1007 / s 10228 - 008 - 0046 - 1","Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp.","Linnaeus, C. (1758) Systema naturae, 10 th edition. Vol. 1. Laurentii Salvii, Holmiae (= Stockholm), ii + 824 pp.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Nakabo, T. (2002) Platycephalidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 601 - 602.","Imamura, H. & Knapp, L. W. (2009) Platycephalus orbitalis, a new species of flathead (Teleostei: Platycephalidae) collected from western Australia. Zootaxa, 2271, 57 - 63."]}

Platycephalus grandispinis Cuvier in Cuvier & Valenciennes, 1829 Common English name: Longspine flathead (Figs. 6–7) Platycephalus grandispinis Cuvier in Cuvier & Valenciennes, 1829: 242 (type locality: unknown); Imamura, 2013 b: 187, figs. 1 –3, 5–6, 8– 9. Platycephalus longispinis Macleay, 1884: 170 (type locality: outside Port Jackson, NSW, Australia); McCulloch, 1929: 401; Coleman, 1980: 108, unnumbered color fig.; Hutchins & Thompson, 1983: 78, fig. 111; Hutchins & Swainston, 1986: 127, fig. 199; May & Maxwell, 1986: 276, unnumbered fig.; Paxton & Hanley, 1989: 469; Knapp, 1991: 29, tab. 3; Kuiter 1993: 102, unnumbered fig.; Hoese et al., 2006: 942; Imamura, 2006: 305, tab. 1. Longitrudis longispinis: Whitley, 1931 a: 327; Whitley, 1931 b: 159; Whitley, 1964: 57. Material examined. Holotype: MNHN 6844, 212 mm SL, locality unknown [see Imamura (2013 b) who inferred the holotype as having been collected from the area between Shark Bay (ca. 25 °S) and Doubtful Islands (ca. 120 °E)]. Other specimens: 44 specimens, including AMS I. 16355 -001, 233 mm SL, holotype of Platycephalus longispinis Macleay, 1884, 120– 286 mm SL, from southeastern to southwestern Australia, listed in Imamura (2013 b). Diagnosis. A species of Platycephalus with the following combination of characters: 14 second dorsal- and anal-fin rays; 73–82 pored lateral-line scales; gill rakers 4–7 + 14–19 = 19–26; interorbit becoming broader with growth, shorter than orbital diameter; a distinct interopercular flap absent (sometimes a small flap or weak convexity present); supraoccipital with a ridge, usually ending in a spine; lower preopercular spine much longer than upper, usually closely approaching posterior margin of opercle, ratio of lower/upper spines 1.9–3.3, tending to become smaller with growth; upper jaw without large caniniform teeth; head and body without small dark dots; posteroventral portion of caudal fin with a blackish or dark brownish marking. Description. A full description was given by Imamura (2013 b). Distribution. Known from southeastern to southwestern Australia, from Fraser Island, Qld (ca. 25 °S), across NSW, Vic and SA to Shark Bay, WA (ca. 25 °S) in depths from ca. 17 to 91 m (e.g., Hutchins and Thompson, 1983; Kuiter, 1993; Hoese et al., 2006; Imamura, 2013 b; this study). Size. Maximum length 38 cm (May & Maxwell, 1986; Kuiter, 1993). The largest specimen examined during the present study was 286 mm SL (332 mm TL). Remarks. The name Platycephalus grandispinis had been largely forgotten until Imamura (2013 b) showed it to have priority over a well-known species, P. longispinis, following detailed examinations of holotypes of the two nominal species (Figs. 7) and non-type specimens. See Imamura (2013 b) for a discussion of P. grandispinis. A comparison of P. grandispinis with P. bassensis is provided herein under the latter., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 162-163, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp.","Imamura, H. (2013 b) Validity of Platycephalus grandispinis Cuvier, 1829, with priority over Platycephalus longispinis Macleay, 1884 (Actinopterygii: Scorpaeniformes: Platycephalidae). Species Diversity, 18, 183 - 192. http: // dx. doi. org / 10.12782 / sd. 18.2.183","Macleay, W. (1884) Notices of new fishes. Proceedings of the Linnean Society of New South Wales, 9, 170 - 172.","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Whitley, G. P. (1931 a) New names for Australian fishes. Australian Zoologist, 6, 310 - 334. [plates. 25 - 27]","Whitley, G. P. (1931 b) Studies in ichthyology. No. 5. Records of the Australian Museum, 18, 131 - 160. [plates. 20 - 21]","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127."]}

Key to the species of Australian Platycephalus The following key to the species of Platycephalus from Australia has been prepared for adults and young specimens with adult-like characters (e.g., specimens of P. f us cu s ca. 100 mm SL or smaller differ from adults in caudal fin color pattern and would not key comfortably to that species); see individual species descriptions for identification of smaller specimens. 1 a. Head and body subcylindrical, slightly depressed; upper preopercular spine longer than lower................ P. laevigatus 1 b. Head and body strongly depressed; lower preopercular spine longer than upper..................................... 2 2 a. One to three large caniniform teeth present anteromedially on upper jaw.......................................... 3 2 b. Large caniniform teeth absent on upper jaw................................................................. 6 3 a. Anal-fin rays usually 13; branched caudal-fin rays 9–11; occipital region naked; margin of interopercle scalloped................................................................................................... P. chauliodous 3 b. Anal-fin rays usually 14; branched caudal-fin rays 12–13; occipital region scaled; margin of interopercle smooth.......... 4 4 a. Pectoral-fin rays usually 19; swimbladder absent; abdominal cavity pale.............................. P. aurimaculatus 4 b. Pectoral-fin rays usually 20 or more; swimbladder present; abdominal cavity dark brown or black...................... 5 5 a. Pored lateral-line scales usually with one pair of sensory ducts with posterior external openings; total gill rakers 7–11.................................................................................................... P. conatus 5 b. Pored lateral-line scales usually with two pairs of sensory ducts with posterior external openings (some scales sometimes with one, three or four pairs of ducts); total gill rakers 12–17............................................. P. richardsoni 6 a. Distinct interopercular flap absent (small flap sometimes present)............................................... 7 6 b. Finger-like interopercular flap present..................................................................... 8 7 a. Total gill rakers> 17; caudal fin with 1–2 blackish or dark brown markings or spots posteroventrally.................... 9 7 b. Total gill rakers 18................................................................... P. caeruleopunctatus 8 b. Total gill rakers P. bassensis 9 b. Ridge present on supraoccipital, usually ending in a spine; small distinct dark dots absent from dorsal surface of head and body......................................................................................... P. grandispinis 10 a. Head and body marbled with irregular dark brown, brown and pale bands and spots...................... P. marmoratus 10 b. Head and body without distinct spots and bands..................................................... P. orbitalis 11 a. Caudal fin with several dark spots dorsally or posteroventrally, but without bands.................................. 12 11 b. Caudal fin with several dark bands....................................................................... 13 12 a. First dorsal fin with a single small isolated spine anteriorly; second dorsal- and anal-fin rays usually 14; caudal fin with several dark spots posteroventorally................................................................... P. speculator 12 b. First dorsal fin usually with two small isolated spines anteriorly; second dorsal- and anal-fin rays usually 13; caudal fin with several dark spots dorsally........................................................................ P. fuscus 13 a. Teeth present on dorsal surface of anterolateral edge of upper jaw....................................... P. angustus 13 b. Teeth absent on dorsal surface of anterolateral edge of upper jaw................................................ 14 14 a. Upper iris lappet usually broad, bilobed; caudal fin with yellow marking posterodorsally when fresh........... P. westraliae 14 b. Upper iris lappet triangular, simple; caudal fin either lacking yellow marking, or not situated as above................. 15 15 a. Total gill rakers 10–15 (tending to decrease with growth); caudal fin usually with four or more dark bands, lacking a yellow marking when fresh........................................................................ P. endrachtensis 15 b. Total gill rakers 4–10 (tending to decrease with growth); caudal fin with three or four dark bands, a yellow marking on midline when fresh............................................................................ P. australis sp. nov., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on page 154, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552

Platycephalus laevigatus Cuvier in Cuvier & Valenciennes, 1829 Common English name: Rock flathead (Figs. 1–2; Table 1) Platycephalus laevigatus Cuvier in Cuvier & Valenciennes, 1829: 248 [type locality: Western Port (as Port Western), Vic, Australia]; Quoy & Gaimard, 1834: 684, fig. 4; Castelnau, 1872: 84; McCulloch, 1929: 402; Coleman, 1980: 114, unnumbered fig.; Scott et al., 1980: 169, unnumbered fig.; Hutchins & Thompson, 1983: 78, fig. 114; Last et al., 1983: 332, fig. 28.20; Hutchins & Swainston, 1986: 127, fig. 195; Paxton & Hanley, 1989: 469; Knapp, 1991: 29, tab. 3; Kuiter 1993: 105, unnumbered fig.; Kuiter, 1994: 521, fig. 463; Grant, 2004: 192, unnumbered pl.; Hoese et al., 2006: 942; Imamura, 2006: 305, tab. 1; Gomon, 2008: 520, unnumbered fig. Platycephalus proximus Castelnau, 1872: 85 (type locality: Melbourne market, Vic, Australia); McCulloch, 1929: 400. Laeviprora laevigata (misspelling of Leviprora): Whitley, 1964: 57. Laeviprora proxima (misspelling of Leviprora): Whitley, 1964: 57. Material examined. Holotype: MNHN 6866, 302 mm SL, Western Port (as Port Western), Vic, Australia. Other specimens (21 specimens, 66.7–484 mm SL, from southern Australia): AMS I. 2834, 257 mm SL, Albany, WA (35 °01’S, 117 ° 53 ’E); AMS I. 7543, 276 mm SL, AMS I. 7544, 271 mm SL, Melbourne market, Vic, Nov. 1905; AMS I. 17610 -012, 66.7 mm SL, 16 km north of Pt. Vincent, Pine Point, SA (34 ° 46 ’S, 137 ° 48 ’E), 23 Dec. 1973; AMS I. 19676 -001, 388 mm SL, AMS I. 19676 -002, 398 mm SL, Merimbula estuary, NSW (36 ° 53 ’S, 149 ° 56 ’E), 18 May 1976; AMS IB. 1961, 401 mm SL, Greenwell Point, NSW (34 ° 50 ’S, 150 ° 45 ’E), 1947; AMS IB. 2112, 260 mm SL, Port Albert, Vic (38 ° 45 ’S, 146 ° 41 ’E), 1948; AMS IB. 3554, 431 mm SL, Nowra, NSW (34 ° 53 ’S, 150 ° 36 ’E), 1956; AMS IB. 7589, 484 mm SL, AMS IB. 7592, 287 mm SL, Tooradin, Vic (38 ° 13 ’S, 145 ° 23 ’E); CSIRO CA 596, 281 mm SL, St. Kilda, SA, 24 Jan. 1975; CSIRO T 1644, 145 mm SL, North East River, Flinders Island, Tas; CSIRO T 1678, 78.5 mm SL, Norfolk Bay, Tas (43 °00’S, 147 ° 45 ’E), 21 Apr. 1978; CSIRO T 1719, 113.6 mm SL, North East River, Flinders Island, Tas (39 ° 47 ’S, 147 ° 58 ’E), 31 Jan. 1978; MNHN A 4285 (holotype of Platycephalus proximus Castelnau, 1872), 341 mm SL, Melbourne market, Vic; NMV A 9294, 97.2 mm SL, sea-grass flats, Price Creek entrance, Yorke Peninsula, SA (35 ° 33 ’S, 128 °08’E), 3 Dec. 1986; NMV A 18576, 85.8 mm SL, mudflat, south of boat ramp, Stony Point, Western Port, Vic (38 ° 22 ’ 36 ”S, 145 ° 13 ’ 24 ”E), 0.5 m depth, 9 Dec. 1996; NMV A 26195 -002, 132 mm SL, subtidal reef, Port Julia, Yorke Peninsula, Gulf of St. Vincent, SA (34 ° 39 ’S, 137 ° 52 ’E), 15 Dec. 1994; NMV A 29380 -003, 159 mm SL, Corner Inlet, Port Franklin, Vic (38 ° 41 ’S, 146 ° 16 ’E), 0–1 m depth, 1 Mar. 2005; WAM P. 27752 -001, 329 mm SL, Fremantle, WA (32 °03’S, 115 ° 44 ’E), Aug. 1982. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; interorbital width 3.4–7.8 % HL; postorbital length 51.4–65.2 % HL; head and body subcylindrical, elongate and only slightly depressed; occipital region naked; dorsal and lateral surfaces of head mostly smooth, with preopercular (2), opercular (2) and supracleithral (1) spines; upper preopercular spine longer than lower; upper jaw without large caniniform teeth; head with two dark brown bands from anterior of snout to posterior margin of opercle on both sides, area between bands whitish-brown, pale brown or brown, area below each band pale brown in specimens 159 mm SL or smaller. Description. Counts and measurements shown in Table 1. Data for all specimens presented first, followed by holotype condition in parentheses. Head and body subcylindrical, elongate, only slightly depressed. Snout, area anteroventral to eye, interorbit, and occipital and lower half of suborbital regions naked. Snout short, subequal or slightly longer (subequal in holotype) than orbital diameter. Interorbit much narrower than orbital diameter. Upper iris lappet simple, triangular; lower simple, weakly convex. Dorsal and lateral surfaces of head mostly smooth, but with preopercular (2) and opercular spines (2). One supracleithral spine usually present (absent on left side, present on right side). Upper preopercular spine longer than lower, not reaching opercular margin. Small or finger-like interopercular flap usually present (finger-like flap present); margin of interopercle smooth. Maxilla reaching slightly beyond anterior margin of pupil to slightly beyond middle of eye (reaching to middle of eye). Anteromedial portion of upper jaw with moderate to large conical or small caniniform (moderate conical) teeth. Palatine teeth irregularly arranged in two to four (three) rows, tending to become larger medially. Vomerine teeth irregularly arranged in one to four (three) rows anteriorly, two to six (four) rows posteriorly, becoming larger posteriorly. Fleshy sensory tubes from suborbitals and preopercle slightly developed, partly covering dorsal and ventral margins of cheek region. Posterior tip of pelvic fin usually reaching just short of anus (including holotype), or to origin of anal fin. Posterior margin of caudal fin rounded (not confirmed in holotype). Color in alcohol. Color of holotype considerably faded, but retaining small brownish spots on sides of head and body, first and second dorsal, pectoral, pelvic and caudal fins (Fig. 2 A). In other specimens (Fig. 1 A, B), ground color of head and body dark brown above, paler below. Side of head and body with dark brown spots. First and second dorsal, pectoral, pelvic and caudal fins with many small brownish, dark brownish and blackish spots; those on pectoral and pelvic fins tending to form bands. Anal fin without melanophores in 132 mm SL or smaller specimens, with melanophores along rays by 159 mm SL, with dark spots posteroventrally by 257 mm SL and with spots ventrally in larger specimens. In 159 mm SL or smaller specimens, head with dark brown band from anterior part of snout to posterior margins of opercles on both sides; area between bands whitish-brown, pale brown or brown; areas below bands pale brown (Fig. 1 C). Holotype Holotype of P. proximus Non-types MNHN 6866 MNHN A 4285 n = 20 SL (mm) 477 341 66.7– 484 Counts: Distribution. Known from southern Australia, from NSW (31 °01’S), across SA, Vic and Tas, to Local Waters, WA (32 °03’S), occurring on weed-covered reefs and sea-grass beds in depths from 0.5 to 20 m (e.g., Last et al., 1983; Kuiter, 1993, 1994; Hoese et al., 2006; this study). Size. Maximum length 80.8 cm (Hutchins & Swainston, 1986). The largest specimen examined during this study was 484 mm SL (573 mm TL) (Fig. 1 A, B). Remarks. Platycephalus laevigatus is easily separable from congeneric species in having the head and body subcylindrical, elongate and only slightly depressed (well depressed in others), occipital region naked (vs. scaled, except in P. chauliodous), dorsal and lateral surfaces of the head mostly smooth, only two distinct preopercular, two opercular and one supracleithral spine [vs. some or many other spines and ridges, including lachrymal and preocular spines (but preocular spine rarely absent in P. chauliodous)] and the upper preopercular spine longer than the lower [vs. lower spine longer than upper, except in some smaller specimens of P. caeruleopunctatus and P. chauliodous (upper spine about equal to or longer than lower)]. Although P. laevigatus and P. chauliodous share several other characters setting them apart from other species, P. laevigatus is further distinguished from P. chauliodous in having the upper jaw without large caniniform teeth (such teeth medially on the anterior part of the upper jaw in the latter). In addition, the head coloration in 159 mm SL or smaller specimens of P. laevigatus is unique, thereby allowing distinction from other members of the genus (Fig. 1 C). Although interorbital width becomes relatively broader with growth in all species of Platycephalus (Imamura, 2012, 2013a), P. laevigatus has the narrowest interorbit of all at comparable sizes (interorbital width 3.4–7.8 % HL in P. laevigatus). Comparison of interorbital width in all examined specimens of P. laevigatus and six species of Platycephalus is characterized by interorbital width narrower than orbital diameter, as shown in Fig. 3 A (viz. 7.4–11.7 % HL in P. aurimaculatus, 7.2–12.8 % in P. bassensis, 7.4–14.1 % in P. conatus, 6.9–12.3 % in P. grandispinis, 5.8–14.6 % in P. speculator and 10.1–15.7 % in P. richardsoni; in all other species of Platycephalus, interorbital width becomes greater than orbital diameter with growth). In addition, P. laevigatus is also separable from the above same six species in having a longer postorbital region (postorbital length 51.4–65.2 % HL in P. laevigatus vs. 49.1–56.9 %, 49.6–57.1 %, 49.7–56.2 %, 48.5–54.4 %, 51.2–59.1 % and 49.7–56.2 %, respectively) (Fig. 3 B). The synonymy of P. laevigatus and P. proximus Castelnau, 1872 (Fig. 2 B) has been recognized by Paxton & Hanley (1989), Kuiter (1994) and Hoese et al. (2006), but without justification. Platycephalus proximus, known only from the holotype from Melbourne market, agrees well with P. laevigatus, having the above mentioned characters, and other counts and proportional measurements similar, without any remarkable differences (Table 1). The present study confirms the priority of P. laevigatus over P. proximus., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 154-158, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp.","Quoy, J. R. C. & Gaimard, J. P. (1834) Voyage de decouvertes de \" L'Astrolabe \" axecute par ordre du Roi, pendant les annees 1826 - 29, sous le commandement de M. J. Dumont d'Urville, vol. 3. Pilet Aine, Paris.","Castelnau, F. L. (1872) Contributions to the ichthyology of Australia. No. 1. The Melbourne fish market. Proceedings of the Royal Zoological Acclimatization Society of Victoria, 1, 29 - 242.","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Scott, T. D., Glover, C. J. M. & Southcott, R. V. (1980) The marine and freshwater fishes of south Australia, 2 nd edition. DJ Woolman, Adelaide, 392 pp.","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp.","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Kuiter, R. H. (1994) Family Platycephalidae. In: Gomon, M. F., Glover, J. C. M. & Kuiter, R. H. (Eds.), The fishes of Australia's south coast. State Print, Adelaide, pp. 514 - 522.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","Imamura, H. (2012) Redescription of Platycephalus angustus Steindachner 1886 (Teleostei: Platycephalidae), a valid flathead in northern Australia and New Guinea. Ichthyological Research, 60, 112 - 121. http: // dx. doi. org / 10.1007 / s 10228 - 012 - 0319 - 6","Imamura, H. (2013 a) Platycephalus mortoni Macleary 1883, a junior synonym of Platycephalus fuscus Cuvier 1829 (Teleostei: Platycephalidae). Ichthyological Research, 60, 282 - 286. http: // dx. doi. org / 10.1007 / s 10228 - 013 - 0348 - 9"]}

Platycephalus orbitalis Imamura & Knapp, 2009 Common English name: Western-Australian flathead (Fig. 15) Platycephalus orbitalis Imamura & Knapp, 2009: 58, figs. 1���5 (type locality: northwest of Rottnest Island, WA, Australia). Platycephalus marmoratus (not Stead, 1908): Hutchins & Thompson, 1983: 78, fig. 112; Hutchins & Swainston, 1986: 127, fig. 204 (in part); Paxton & Hanley, 1989: 469 (in part); Knapp, 1991: 29, tab. 3 (in part); Hutchins, 2001: 28; Hoese et al., 2006: 943 (in part). Material examined. Holotype: CSIRO H 6349 -04, 267 mm SL, northwest of Rottnest Island, WA, Australia (31 �� 52.56 ���S, 115 �� 18.30 ���E ��� 31 �� 52.60 ���S, 115 �� 18.49 ���E), 100���102 m depth, 10 Apr. 2006. Paratypes: 6 specimens, 236���333 mm SL, from western Australia, listed in Imamura & Knapp (2009). Other specimen (1 specimen, 337 mm SL, from northwestern Australia): WAM P. 27606 -001, 337 mm SL, WA (32 ��03���S, 115 �� 44 ���E), 28 March 1982. Diagnosis. A species of Platycephalus with the following combination of characters: dorsal- and anal-fin rays 13; pored lateral-line scales 65���68, all lacking spines; gill rakers 2 + 6���7 = 8���10; snout and interorbit naked; interopercular flap absent; margin of interopercle scalloped; upper jaw without large caniniform teeth; fleshy sensory tubes from infraorbitals and preopercle well developed, covering cheek region, except anteroventrally; head and body lacking distinct large spots and bands dorsally; caudal fin blackish with a pale posterior margin. Description. A full description was given by Imamura & Knapp (2009). Distribution. Known only from western Australian waters, from Cape Leeuwin (34 ��S) north to Cape Cuvier (24 ��S) (e.g., Hutchins, 2001; Imamura & Knapp, 2009; this study). Size. Known maximum length 337 mm SL (400 mm TL) (Imamura & Knapp, 2009; this study). Remarks. Although Imamura & Knapp (2009) described the holotype as having 2 + 4 = 6 gill rakers, reexamination has confirmed the number as 2 + 6 = 8. The present species is mostly similar to P. marmoratus (see Remarks for P. marmoratus)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 173-174, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Imamura, H. & Knapp, L. W. (2009) Platycephalus orbitalis, a new species of flathead (Teleostei: Platycephalidae) collected from western Australia. Zootaxa, 2271, 57 - 63.","Stead, D. G. (1908) New fishes from New South Wales (No. 1). Department of Fisheries, New South Wales, Sydney, 12 pp. [5 plates]","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Hutchins, J. B. (2001) Checklist of the fishes of Western Australia. Records of the Western Australian Museum, Supplement, 63, 9 - 50.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948."]}

Platycephalus fuscus Cuvier in Cuvier & Valenciennes, 1829 Common English name: Dusky flathead (Figs. 25–26; Table 10) Platycephalus fuscus Cuvier in Cuvier & Valenciennes, 1829: 241 (type locality: Port Jackson, NSW, Australia); Quoy & Gaimard, 1834: 681, fig. 1; Castelnau, 1872: 86; McCulloch, 1929: 400; Coleman, 1980: 110, unnumbered color fig.; Scott et al., 1980: 167, unnumbered fig.; Hutchins & Swainston, 1986: 127, fig. 196; Paxton & Hanley, 1989: 468; Knapp, 1991: 29, tab. 3; Kuiter 1993: 104, two unnumbered figs.; Kuiter, 1994: 522, fig. 464; Grant, 2004: 189, pl. 86; Hoese et al., 2006: 941; Imamura, 2006: 305, tab. 1; Gomon, 2008: 519, unnumbered fig. Platycephalus cinereus Günther, 1872: 661 (type locality: South Australia). Platycephalus mortoni Macleay, 1883: 206 (type locality: lower Burdekin River, Qld, Australia); McCulloch, 1929: 401. Planiprora fusca: Whitley, 1931 a: 237; Whitley, 1964: 57. Planiprora cinerea: Whitley, 1964: 57. Material examined. Holotype: MNHN 6867, 346 mm SL, Port Jackson, NSW, Australia. Other specimens (21 specimens, 102–750 mm SL, from eastern Australia): AMS I. 6741 -001, 241 mm SL, Lane Cove River, NSW (33 ° 50 ’S, 151 ° 11 ’E), 5 Sep. 1972; AMS I. 17178 -010, 2 specimens, 252–317 mm SL, Sydney Harbour, NSW (33 ° 50 ’S, 151 ° 16 ’E), 13 Aug. 1972; AMS I. 21149 -032, 212 mm SL, Bohle River estuary, Townsville, Qld (19 ° 13 ’S, 146 ° 48 ’E), 8 Oct. 1965; AMS I. 22474 -001, 120 mm SL, Tuross estuary, Reedy Island, NSW (36 °05’S, 150 °07’E), 1981; AMS I. 30344 -001, 210 mm SL (dissected by Imamura, 1996), Apple Tree Bay, Ku-ring-gai Chase National Park, NSW (33 ° 39 ’S, 151 °01’E), 1 Oct. 1974; AMS I. 34342 -001, 292 mm SL, northeast end of West Flat, Port Clinton, Qld (22 ° 33.28 ’S, 150 ° 45.05 ’E), 0–1 m depth, 25 Sep. 1993; AMS I. 36105 -001, 183 mm SL, Botany Bay, NSW (33 ° 59 ’S, 151 °09’E), 0–5 m depth, 9 Apr. 1992; AMS I. 41262 -024, 218 mm SL, Lennox Head Beach, NSW (28 ° 48.11 ’S, 153 ° 35.83 ’E), 0–1 m depth, 19 Mar. 2002; AMS I. 41264 - 0 20, 2 of 5 specimens, 164–189 mm SL, north side of Prospect Bridge, North Creek, NSW (28 ° 50.87 ’S, 153 ° 34.28 ’E), 0–0.5 m depth, 19 Mar. 2002; AMS I. 41287 -020, 186 mm SL, Yamba, NSW (38 °03’S, 147 ° 50 ’E), 0–0.5 m depth, 11 Dec. 2002; AMS I. 41874 -011, 204 mm SL, Hastings Point, just inside mouth of Cudgen Creek, NSW (28 ° 21.68 ’S, 153 ° 34.53 ’E), 0–1.5 m depth, 11 Dec. 2002; AMS IB. 8204, 579 mm SL, Eden, NSW (37 °04’S, 149 ° 55 ’E), 9 June 1968; BMNH 1870.12. 27.25, holotype of Platycephalus cinereus Günther, 1872, 330 mm SL, South Australia; CSIRO CA 126, 277 mm SL, fish market, Brisbane, Qld, 6 June 1977; CSIRO H 3986 -01, 410 mm SL, east of Newcastle, NSW (32 ° 54 ’S, 152 °00’E), 79 m depth, 29 Mar. 1995; NMV A 1688, 750 mm SL, Snowy River mouth, Mario, Vic (37 ° 48 ’S, 148 ° 32 ’E), 27 Nov. 1980; NMV A 25513 -001, 457 mm SL, Gippsland Lakes, Vic (37 ° 59 ’S, 147 ° 43 ’E), 20 Nov. 2003; QM I. 16513, 373 mm SL, Murray River mouth, north of Cardwell, Qld (18 °05’S, 146 °02’E), date unknown; QM I. 20085, 102 mm SL, Russell River mouth, Qld (17 ° 14 ’S, 145 ° 58 ’E), Nov. 1982. Diagnosis. A species of Platycephalus with the following combination of characters: first dorsal fin usually with two small isolated spines anteriorly; second dorsal- and anal-fin rays usually 13; interorbital width 8.2–17.1 % HL; postorbital length 6.5 –10.0% HL; a finger-like interopercular flap present; upper jaw without large caniniform teeth; teeth absent on dorsal surface of anterolateral edge of upper jaw; palatine teeth in two rows; vomerine teeth in one row anteriorly, two to three posteriorly; in larger specimens, caudal fin brownish, usually with a single black spot near upper posterior margin, upper half of fin to margin with or without small pale brown, brown or black spots. Description. Counts and measurements shown in Table 10. Data for all specimens presented first, followed by holotype condition in parentheses. Snout, area anteroventral to eye, interorbit and occipital region scaled; lower half of suborbital region naked. Interorbit narrower than orbital diameter in 346 mm SL (holotype) or smaller specimens, equal to or wider in larger specimens. Upper iris lappet simple, triangular; lower simple, weakly convex. Nasal spine present in smallest 102 mm SL specimen, absent in larger specimens. Preorbital spine absent. One suborbital spine present below eye or absent in 330 mm SL or smaller specimens, absent in larger specimens (including holotype). Lower preopercular spine slightly longer than upper, not reaching opercular margin. Supplemental preopercular spine present or absent (absent); often present in smaller specimens. Finger-like interopercular flap present; margin of interopercle smooth. Maxilla reaching from anterior to near posterior margins of pupil (middle of eye) [slightly beyond posterior margin of eye in largest (750 mm SL) specimen]. Upper jaw with some moderate or large conical, or small or moderate caniniform (small caniniform) teeth anteromedially. Palatine teeth in two rows, villiform in outer row, moderately conical in inner row. Vomerine teeth villiform, in one row anteriorly, two or three (two) posteriorly, tending to become larger posteriorly; posterior row number tending to increase with growth. Fleshy sensory tubes from suborbitals and preopercle not covering cheek region. Posterior tip of pelvic fin reaching from anus to base of third anal-fin ray (base of second anal-fin ray). Posterior margin of caudal fin slightly concave or mostly straight (mostly straight). Color in alcohol. Color of holotype mostly faded, retaining only one brownish marking on posterior portion of caudal fin (Fig. 26 A). In other specimens (Fig. 25), ground color of head and body pale to dark brown above, paler below. Dorsal surface of head and body with small darker spots; two darker bands below second dorsal fin. Side of body with or without gray or brown spots or bands. First and second dorsal, pectoral and pelvic fins with small pale brown or brown spots. Anal fin pale or with melanophores along rays. In 120 mm SL or larger specimens, caudal fin brownish, usually with single black spot near upper posterior margin; upper half of fin to margin otherwise with or without small pale brown, brown, or black spots. Smallest (102 mm SL) specimen with three distinct blackish bands on posterior portion of caudal fin. Posterior and ventral margins of caudal fin paler. Largest (750 mm SL) specimen with many small dark brown spots on entire caudal fin. Distribution. Known from eastern Australia, fromRussell River mouth, Qld (17 ° 14 ’S) to Port Phillip Bay, Vic (ca. 38 ° 59 ’S, 145 °E), including NSW, on shallow sand and mud substrate to 75 m depth, preferring bays, estuaries and quiet inlets (e.g., Kuiter, 1993; Hoese et al., 2006; Gomon, 2008; this study). Size. Maximum length 1.2 m (e.g., Kuiter, 1993; Gomon, 2008). The largest specimen examined during the present study was 750 mm SL (874 mm TL). Remarks. Platycephalus fuscus is similar to P. endrachtensis, P. w e s t r al i a e, P. angustus, P. australis, P. cultellatus, P. indicus and two undescribed species (Platycephalus sp. 1 and sp. 2 sensu Nakabo, 2002) in having usually 13 second dorsal- and anal-fin rays, the snout, area anteroventral to eye, interorbit and occipital region scaled, large caniniform teeth absent anteromedially on the upper jaw, a finger-like interopercular flap and palatine teeth in two rows. Although P. f us cus mostly resembles P. angustus and P. cultellatus in having the first dorsal fin usually with two small isolated spines anteriorly (one in other Australian species), it can be distinguished from them by the narrower interorbit and shorter postorbital region (interorbital width 8.2–17.1 % HL and postorbital length 6.5 –10.0% HL in P. fuscus vs. 7.3–17.3 % HL and 55.5–67.8 % HL in P. angustus, and 6.5 –19.0% HL and 51.8–66.8 % HL in P. cultellatus) (Fig. 26). Platycephalus fuscus also differs from P. angustus in lacking teeth on the dorsal surface of the anterolateral edge of the upper jaw (teeth present in P. angustus specimens ca. 76 mm SL or longer) and having vomerine teeth in one row anteriorly, two to three posteriorly (number of vomerine tooth rows tending to increase with growth (from two to four rows in 106–184 mm SL specimens and a single broad band of teeth in larger specimens). Caudal fin coloration clearly distinguishes P. f us cus from the above eight species (except for the smallest specimen examined). See Imamura (2013 a) for a detailed discussion of the synonymy of P. f us c u s and Platycephalus mortoni Macleay, 1883, originally described from a single specimen from lower Burdekin River, Qld, Australia. Paxton & Hanley (1989) and Hoese et al. (2006) regarded P. f u s c u s as a senior synonym of Platycephalus cinereus Günther, 1872 (Fig. 26 B), but gave no reasons. This synonymy was reconfirmed here., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 186-190, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp.","Quoy, J. R. C. & Gaimard, J. P. (1834) Voyage de decouvertes de \" L'Astrolabe \" axecute par ordre du Roi, pendant les annees 1826 - 29, sous le commandement de M. J. Dumont d'Urville, vol. 3. Pilet Aine, Paris.","Castelnau, F. L. (1872) Contributions to the ichthyology of Australia. No. 1. The Melbourne fish market. Proceedings of the Royal Zoological Acclimatization Society of Victoria, 1, 29 - 242.","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Scott, T. D., Glover, C. J. M. & Southcott, R. V. (1980) The marine and freshwater fishes of south Australia, 2 nd edition. DJ Woolman, Adelaide, 392 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Kuiter, R. H. (1994) Family Platycephalidae. In: Gomon, M. F., Glover, J. C. M. & Kuiter, R. H. (Eds.), The fishes of Australia's south coast. State Print, Adelaide, pp. 514 - 522.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Gunther, A. (1872) Report on several collections of fishes recently obtained for the British Museum. Proceedings of the Zoological Society of London, 1871, 652 - 675. [plates. 53 - 70]","Macleay, W. (1883) Notes on a collection of fishes from the Burdekin and Mary rivers, Queensland. Proceedings of the Linnean Society of New South Wales, 8, 199 - 213.","Whitley, G. P. (1931 a) New names for Australian fishes. Australian Zoologist, 6, 310 - 334. [plates. 25 - 27]","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","Imamura, H. (1996) Phylogeny of the family Platycephalidae and related taxa (Pisces: Scorpaeniformes). Species Diversity, 1, 123 - 233.","Nakabo, T. (2002) Platycephalidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 601 - 602.","Imamura, H. (2013 a) Platycephalus mortoni Macleary 1883, a junior synonym of Platycephalus fuscus Cuvier 1829 (Teleostei: Platycephalidae). Ichthyological Research, 60, 282 - 286. http: // dx. doi. org / 10.1007 / s 10228 - 013 - 0348 - 9"]}

Platycephalus marmoratus Stead, 1908 Common English name: Marbled flathead (Figs. 13–14; Table 5) Platycephalus marmoratus Stead, 1908: 9, pls. 3–5 (type locality: Port Stephens, NSW, Australia); McCulloch 1929: 402; Coleman, 1980: 109, unnumbered fig.; Hutchins & Thompson 1983: 78, fig. 112 (in part); Hutchins & Swainston 1986: 127, fig. 204 (in part); Paxton & Hanley, 1989: 469 (in part); Knapp, 1991: 29, tab. 3 (in part); Kuiter 1993: 104, two unnumbered figs. (in part); Hutchins 2001: 28; Grant 2004: 196, pl. 89, unnumbered pl.; Hoese et al., 2006: 942 (in part); Imamura, 2006: 296, figs. 1–6. Planiprora marmorata: Whitley, 1964: 57. Material examined. Holotype: AMS I. 15260, 310 mm SL, Port Stephens, NSW, Australia, June 1904. Other specimens (18 specimens, 152–472 mm SL, from southeastern Australia): 7 specimens listed in Imamura (2006) plus 11 additional specimens: AMS I. 18756 -001, 331 mm SL, NSW (33 ° 38 ’S, 151 ° 27 ’E), 56–59 m depth, 7 Apr. 1975; AMS I. 21570 -001, 292 mm SL, east of Bulli, NSW (34 ° 20 ’S, 150 ° 55 ’E), Feb. 1980; AMS I. 26271 -012, 362 mm SL, northeast of Sandon Bluffs, NSW (29 ° 39 ’S, 153 ° 23 ’E), 36–43 m depth, 10 Oct. 1985; AMS I. 43814 -005, 228 mm SL, Newcastle, NSW (32 ° 51 ’S, 151 ° 53 ’E), 26 May 1993; CSIRO H 3985 -01, 354 mm SL, south of Newcastle, NSW (33 ° 111 ’S, 151 ° 43 ’E), 57 m depth, 29 March 1995; CSIRO H 6836 -06, 262 mm SL, H 6836 -07, 298 mm SL, east of Terrigal, NSW (32 ° 25 ’S, 151 ° 32 ’E – 33 ° 23 ’S, 151 ° 35 ’E), 48–55 m depth, 13 May 2006; NMV A 11848, 335 mm SL, off Bermagui, NSW (36 ° 26 ’S, 150 °05’E), Apr. 1992; QM I.17021, 2 of 3 specimens, 193–222 mm SL, 7 mile north-northwest of Cape Moreton, Moreton Island, Qld (26 ° 55 ’S, 153 ° 25 ’E), 109.7 m depth, 27 Feb. 1975; QM I. 2842, 399 mm SL, Caloundra Bank, Qld (26 ° 48 ’S, 153 ° 12 ’E). Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays 13; pored lateral-line scales 63–70, all lacking spines; gill rakers 2 + 7–11 = 9–13; snout and interorbit naked; interopercular flap absent; margin of interopercle smooth; upper jaw without large caniniform teeth; fleshy sensory tubes from suborbitals and preopercle slightly developed, partially covering dorsal and ventral margins of cheek region; body and head marbled with dark brown, brown and pale irregular bands and spots; caudal fin blackish with pale posterior margin. Holotype Non-types AMS I. 15260 n = 18 SL (mm) 310 151–472 Counts: D 1 I + VII + 0 I + VI–VIII + 0–I (often I + VII + 0) D 2 13 13 A 13 13 P 1 2 + 12 + 7 = 21 1–2 + 11–16 + 5–8 = 19–22 Description. A full description was given by Imamura (2006), which examined only specimens from southeastern Australia (thus specimens of P. orbitals, known from western Australia and previously confused with P. marmoratus, are not included). Counts and measurements are shown in Table 5. Distribution. Known only from eastern Australia, from off Caloundra Bank, Qld (26 ° 48 ’S) to Flinders Island, Tas (40 °01’S), including NSW and Vic, on sand near low rocky reefs or on rubble in coastal bays to deep offshore reefs in depths from 20 to 80 m (e.g., Paxton & Hanley, 1989; Kuiter, 1993; Hoese et al., 2008; this study). Size. Maximum length 56 cm (Grant, 2004). The largest specimen examined during the present study was 472 mm SL (550 mm TL). Remarks. Platycephalus marmoratus had been confused with a second species, known only from western Australia, until the latter was described (as Platycephalus orbitalis) by Imamura & Knapp (2009). The former closely resembles P. orbitalis in having 13 second dorsal- and anal-fin rays, 63–70 pored lateral-line scales (65–68 in P. orbitalis), the snout and interorbit naked, interopercular flap absent, fleshy sensory tubes from the infraorbitals and preopercle covering the cheek region, and the caudal fin blackish with a pale posterior margin. However, it is separable from P. orbitalis in having a smooth interopercle margin (scalloped in P. orbitalis), fleshy sensory tubes slightly developed, partly covering the dorsal and ventral margins of the cheek region (vs. tubes well developed, mostly covering cheek region, except anteroventrally) and the body and head marbled with dark brown, brown, and pale irregular bands and spots (vs. body and head lacking large distinct spots and bands dorsally). Although overlapping, some head proportions also aid the distinction of P. marmoratus from P. orbitalis, viz. smaller orbit and broader interorbit (orbital diameter 15.2–20.1 % HL and interorbital width 13.6–21.4 % HL in P. marmoratus vs. 17.5–20.3 % HL and 11.9–14.6 % HL in P. orbitalis) (see Imamura & Knapp, 2009: fig. 4). Gill raker numbers (9–13 in total in P. marmoratus vs. 8–10 in P. orbitalis) may help distinguish between the species., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 171-173, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Stead, D. G. (1908) New fishes from New South Wales (No. 1). Department of Fisheries, New South Wales, Sydney, 12 pp. [5 plates]","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Hutchins, J. B. (2001) Checklist of the fishes of Western Australia. Records of the Western Australian Museum, Supplement, 63, 9 - 50.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","Imamura, H. & Knapp, L. W. (2009) Platycephalus orbitalis, a new species of flathead (Teleostei: Platycephalidae) collected from western Australia. Zootaxa, 2271, 57 - 63."]}

Platycephalus conatus Waite & McCulloch, 1915 Common English name: Deepwater flathead (Fig. 22; Table 8) Platycephalus (Neoplatycephalus) conatus Waite & McCulloch, 1915: 466, pl. 12, figs. 1 ���3, 5 (type locality: Great Australian Bight, SA, Australia). Neoplatycephalus conatus: May & Maxwell, 1986: 271, unnumbered fig.; Knapp, 1991: 29, tab. 1; Kuiter, 1994: 517, fig. 460. Platycephalus conatus: McCulloch, 1929: 401; Last et al., 1983: 332, fig. 28.19; Paxton & Hanley, 1989: 468; Hoese et al., 2006: 941; Imamura, 2006: 304, tab. 1; Gomon, 2008: 519, unnumbered fig. Platycephalus speculator (not Klunzinger, 1872): Coleman, 1980: 112, unnumbered color fig. Neoplatycephalus speculator (not Klunzinger, 1872): Scott et al., 1980: 167, unnumbered fig. Material examined. Paratypes (2 specimens, 243���269 mm SL, from southern Australia): AMS I. 13649, 243 mm SL, Great Australian Bight, 146���219 m depth, Apr. 1913; AMS I. 13650, 269 mm SL, Great Australian Bight, 145���220 m depth, Apr. 1913. Other specimens (17 specimens, 158���607 mm SL, from southwestern and southern Australia): AMS I. 12393, 236 mm SL, AMS I. 2394, 227 mm SL, Investigator Strait, SA (35 �� 25 ���S, 137 �� 22 ���E), 1912; AMS I. 18710 -007, 2 specimens, 158���174 mm SL, Great Australian Bight, WA (33 �� 20.8 ���S, 127 �� 29.7 ���E), 300���310 m depth, 27 Feb. 1976; AMS I. 25551 -001, 303 mm SL, Great Australian Bight, SA (37 �� 45 ���S, 139 �� 45 ���E), 11 Nov. 1984; CSIRO H 1919 -05, 486 mm SL, Great Australian Bight, SA, 130 m depth, Feb. 1988; CSIRO H 4133 -04, 550 mm SL, Great Australian Bight, WA (32 ��S, 129 ��E��� 33 ��S, 131 ��E), 130 m depth, Jan. 1996; CSIRO T 529, 337 mm SL, Great Australian Bight, WA (32 �� 32 ���S, 129 ��05���E ��� 33 �� 22 ���S, 128 �� 57 ���E), 235 m depth, 22 May 1983; CSIRO T 532, 179 mm SL, Great Australian Bight, SA (33 �� 22 ���S, 128 �� 57 ���E), 222 m depth, 22 May 1983; NMV A 2916, 291 mm SL (dissected by Imamura, 1996), 29 km south-southwest of Lakes Entrance, Vic (38 ��S, 147 �� 59 ���E), 53 m depth, date unknown; NMV A 9033, 166 mm SL, 120 km south-southwest of Middini Beach, Great Australian Bight, WA (33 �� 13 ���S, 127 ��02���E), 98���147 m depth, 14 Feb. 1990; NMV A 9488, 410 mm SL, 95 km southwest of Geraldton, WA (29 �� 21 ��� 48 ������ 29 �� 20 ��� 24 ���S, 113 �� 46 ��� 36 ������ 113 �� 45 ��� 54 ���E), 530���942 m depth, 6 June 1991; NMV A 24526, 191 mm SL, Great Australian Bight, WA (33 ��S, 127 ��E), 160 m depth, 7 Dec. 2001; NMV A 25185 -001, 365 mm SL, Port Lincoln, Spencer Gulf, SA (34 �� 44 ���S, 135 �� 52 ���E), 28 July 2003; NSMT-P 112677, 607 mm SL, Great Australian Bight, SA (33 �� 18.8 ���S, 130 �� 8.7 ���E), 157 m depth, 30 Nov. 1975; QM I. 22822, 237 mm SL, off Portland, Vic (38.2 ��S, 141.4 ��E), 140 m depth, date unknown; WAM P. 28617 -005, 432 mm SL, SA (35 ��00���S, 117 �� 52 ���E), 1975. Paratypes Non-types n = 2 n = 17 SL (mm) 243���269 73.0��� 504 Counts: Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; caudal-fin rays 12���13; pored lateral-line scales 70���81; gill rakers 1���2 + 6���9 = 7���11; orbital diameter 16.9���22.6 % HL; interorbital width 7.4���14.1 % HL; postorbital length 49.7���56.2 % HL; suborbital width 5.4���9.7 % HL; occipital and lower half of suborbital regions scaled; margin of interopercle smooth; one or two large caniniform teeth present anteromedially on upper jaw; all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly; swimbladder present; abdominal cavity dark brown or black. Description. Counts and measurements shown in Table 8. Snout, interorbit, and occipital and lower half of suborbital regions scaled; area anteroventral to eye naked. Interorbit narrower than orbital diameter. Upper iris lappet simple, weakly pointed; lower absent. Nasal, preorbital and supplemental preopercular spines absent. One suborbital spine present below and slightly posterior to posterior margin of eye in 337 mm SL or smaller specimens, absent in larger specimens; one additional spine also present below middle of eye on right side of 191 mm SL specimen. Lower preopercular spine longer than upper, not reaching opercular margin; some specimens with lower spine more than twice length of upper. Interopercular flap absent; margin of interopercle smooth. Maxilla not usually reaching anterior margin of pupil or beyond, although it may reach to middle of eye in some specimens. Upper jaw with one or two large caniniform teeth anteromedially. Palatine teeth in two rows, villiform or small conical in outer row, large conical to large caniniform in inner row. Vomerine teeth in one or two rows. Fleshy sensory tubes from suborbitals and preopercle not covering cheek region. Posterior tip of pelvic fin reaching origin of anal fin to base of third anal-fin ray. Posterior margin of caudal fin slightly concave. Swimbladder present. Color in alcohol. Ground color of head and body pale brown above, paler below. Side of body with single row of purplish gray spots. First and second dorsal fins with indistinct pale brown spots. Pectoral and pelvic fins pale brown, with or without indistinct brownish bands; lower margin of fins paler. Anal fin with melanophores along rays. Caudal fin usually dusky; NMV A 9033 with brown spots anteriorly, brown bands posteriorly. Abdominal cavity dark brown or black. Distribution. Known from southern Australia, from Lakes Entrance, Vic (147 �� 59 ���E) to Geraldton, WA (ca. 29 �� 21 ���S), including SA, on continental shelf and upper slopes in depths from 53 to 530 m (e.g., May & Maxwell, 1986; Hoese et al., 2006; Gomon, 2008; this study). Size. Maximum length 70 cm SL (Gomon, 2008). The largest specimen examined during the present study was 607 mm SL (701 mm TL). Remarks. Platycephalus conatus most resembles P. richardsoni in having the lower half of the suborbital region scaled, a swimbladder present and the abdominal cavity dark brown or black; this combination of characters does not occur in other species of Platycephalus. Platycephalus conatus can be separated from P. richardsoni in having 7���11 total gill rakers (vs. 12���17 in P. richardsoni), and all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly (vs. usually two pairs of sensory ducts and exterior openings)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 181-183, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Waite, E. R. & McCulloch, A. R. (1915) The fishes of the South Australian Government Trawling Cruise, 1914. Transactions Royal Society of South Australia, 9, 455 - 476. [plates. 12 - 15]","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1994) Family Platycephalidae. In: Gomon, M. F., Glover, J. C. M. & Kuiter, R. H. (Eds.), The fishes of Australia's south coast. State Print, Adelaide, pp. 514 - 522.","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Klunzinger, C. B. (1872) Zur fischfauna von sud-Australien. Archiv fur Naturgeschichte, 38, 17 - 47. [plates 2]","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Scott, T. D., Glover, C. J. M. & Southcott, R. V. (1980) The marine and freshwater fishes of south Australia, 2 nd edition. DJ Woolman, Adelaide, 392 pp.","Imamura, H. (1996) Phylogeny of the family Platycephalidae and related taxa (Pisces: Scorpaeniformes). Species Diversity, 1, 123 - 233."]}

Platycephalus conatus Waite & McCulloch, 1915 Common English name: Deepwater flathead (Fig. 22; Table 8) Platycephalus (Neoplatycephalus) conatus Waite & McCulloch, 1915: 466, pl. 12, figs. 1 –3, 5 (type locality: Great Australian Bight, SA, Australia). Neoplatycephalus conatus: May & Maxwell, 1986: 271, unnumbered fig.; Knapp, 1991: 29, tab. 1; Kuiter, 1994: 517, fig. 460. Platycephalus conatus: McCulloch, 1929: 401; Last et al., 1983: 332, fig. 28.19; Paxton & Hanley, 1989: 468; Hoese et al., 2006: 941; Imamura, 2006: 304, tab. 1; Gomon, 2008: 519, unnumbered fig. Platycephalus speculator (not Klunzinger, 1872): Coleman, 1980: 112, unnumbered color fig. Neoplatycephalus speculator (not Klunzinger, 1872): Scott et al., 1980: 167, unnumbered fig. Material examined. Paratypes (2 specimens, 243–269 mm SL, from southern Australia): AMS I. 13649, 243 mm SL, Great Australian Bight, 146–219 m depth, Apr. 1913; AMS I. 13650, 269 mm SL, Great Australian Bight, 145–220 m depth, Apr. 1913. Other specimens (17 specimens, 158–607 mm SL, from southwestern and southern Australia): AMS I. 12393, 236 mm SL, AMS I. 2394, 227 mm SL, Investigator Strait, SA (35 ° 25 ’S, 137 ° 22 ’E), 1912; AMS I. 18710 -007, 2 specimens, 158–174 mm SL, Great Australian Bight, WA (33 ° 20.8 ’S, 127 ° 29.7 ’E), 300–310 m depth, 27 Feb. 1976; AMS I. 25551 -001, 303 mm SL, Great Australian Bight, SA (37 ° 45 ’S, 139 ° 45 ’E), 11 Nov. 1984; CSIRO H 1919 -05, 486 mm SL, Great Australian Bight, SA, 130 m depth, Feb. 1988; CSIRO H 4133 -04, 550 mm SL, Great Australian Bight, WA (32 °S, 129 °E– 33 °S, 131 °E), 130 m depth, Jan. 1996; CSIRO T 529, 337 mm SL, Great Australian Bight, WA (32 ° 32 ’S, 129 °05’E – 33 ° 22 ’S, 128 ° 57 ’E), 235 m depth, 22 May 1983; CSIRO T 532, 179 mm SL, Great Australian Bight, SA (33 ° 22 ’S, 128 ° 57 ’E), 222 m depth, 22 May 1983; NMV A 2916, 291 mm SL (dissected by Imamura, 1996), 29 km south-southwest of Lakes Entrance, Vic (38 °S, 147 ° 59 ’E), 53 m depth, date unknown; NMV A 9033, 166 mm SL, 120 km south-southwest of Middini Beach, Great Australian Bight, WA (33 ° 13 ’S, 127 °02’E), 98–147 m depth, 14 Feb. 1990; NMV A 9488, 410 mm SL, 95 km southwest of Geraldton, WA (29 ° 21 ’ 48 ”– 29 ° 20 ’ 24 ”S, 113 ° 46 ’ 36 ”– 113 ° 45 ’ 54 ”E), 530–942 m depth, 6 June 1991; NMV A 24526, 191 mm SL, Great Australian Bight, WA (33 °S, 127 °E), 160 m depth, 7 Dec. 2001; NMV A 25185 -001, 365 mm SL, Port Lincoln, Spencer Gulf, SA (34 ° 44 ’S, 135 ° 52 ’E), 28 July 2003; NSMT-P 112677, 607 mm SL, Great Australian Bight, SA (33 ° 18.8 ’S, 130 ° 8.7 ’E), 157 m depth, 30 Nov. 1975; QM I. 22822, 237 mm SL, off Portland, Vic (38.2 °S, 141.4 °E), 140 m depth, date unknown; WAM P. 28617 -005, 432 mm SL, SA (35 °00’S, 117 ° 52 ’E), 1975. Paratypes Non-types n = 2 n = 17 SL (mm) 243–269 73.0– 504 Counts: Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; caudal-fin rays 12–13; pored lateral-line scales 70–81; gill rakers 1–2 + 6–9 = 7–11; orbital diameter 16.9–22.6 % HL; interorbital width 7.4–14.1 % HL; postorbital length 49.7–56.2 % HL; suborbital width 5.4–9.7 % HL; occipital and lower half of suborbital regions scaled; margin of interopercle smooth; one or two large caniniform teeth present anteromedially on upper jaw; all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly; swimbladder present; abdominal cavity dark brown or black. Description. Counts and measurements shown in Table 8. Snout, interorbit, and occipital and lower half of suborbital regions scaled; area anteroventral to eye naked. Interorbit narrower than orbital diameter. Upper iris lappet simple, weakly pointed; lower absent. Nasal, preorbital and supplemental preopercular spines absent. One suborbital spine present below and slightly posterior to posterior margin of eye in 337 mm SL or smaller specimens, absent in larger specimens; one additional spine also present below middle of eye on right side of 191 mm SL specimen. Lower preopercular spine longer than upper, not reaching opercular margin; some specimens with lower spine more than twice length of upper. Interopercular flap absent; margin of interopercle smooth. Maxilla not usually reaching anterior margin of pupil or beyond, although it may reach to middle of eye in some specimens. Upper jaw with one or two large caniniform teeth anteromedially. Palatine teeth in two rows, villiform or small conical in outer row, large conical to large caniniform in inner row. Vomerine teeth in one or two rows. Fleshy sensory tubes from suborbitals and preopercle not covering cheek region. Posterior tip of pelvic fin reaching origin of anal fin to base of third anal-fin ray. Posterior margin of caudal fin slightly concave. Swimbladder present. Color in alcohol. Ground color of head and body pale brown above, paler below. Side of body with single row of purplish gray spots. First and second dorsal fins with indistinct pale brown spots. Pectoral and pelvic fins pale brown, with or without indistinct brownish bands; lower margin of fins paler. Anal fin with melanophores along rays. Caudal fin usually dusky; NMV A 9033 with brown spots anteriorly, brown bands posteriorly. Abdominal cavity dark brown or black. Distribution. Known from southern Australia, from Lakes Entrance, Vic (147 ° 59 ’E) to Geraldton, WA (ca. 29 ° 21 ’S), including SA, on continental shelf and upper slopes in depths from 53 to 530 m (e.g., May & Maxwell, 1986; Hoese et al., 2006; Gomon, 2008; this study). Size. Maximum length 70 cm SL (Gomon, 2008). The largest specimen examined during the present study was 607 mm SL (701 mm TL). Remarks. Platycephalus conatus most resembles P. richardsoni in having the lower half of the suborbital region scaled, a swimbladder present and the abdominal cavity dark brown or black; this combination of characters does not occur in other species of Platycephalus. Platycephalus conatus can be separated from P. richardsoni in having 7–11 total gill rakers (vs. 12–17 in P. richardsoni), and all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly (vs. usually two pairs of sensory ducts and exterior openings).

Platycephalus caeruleopunctatus McCulloch, 1922 Common English name: Bluespotted flathead (Figs. 8–9; Table 3) Platycephalus caeruleopunctatus McCulloch, 1922: 120 (type locality: Port Jackson, NSW, Australia); McCulloch, 1929: 400; Coleman, 1980: 113, unnumbered color fig.; Hutchins & Swainston, 1986: 127, fig. 201; May & Maxwell, 1986: 275, unnumbered fig.; Paxton & Hanley, 1989: 468; Knapp, 1991: 29, tab. 3; Kuiter 1993: 103, unnumbered fig.; Grant, 2004: 196; Hoese et al., 2006: 940; Imamura, 2006: 304, tab. 1. Trudis caeruleopunctatus: Whitley, 1931 b: 158; Whitley, 1964: 57. Material examined. Lectotype: AMS I. 3163, 291 mm SL, Port Jackson, NSW, Australia, designated by Whitley (1931 b). Other specimens (20 specimens, 73.0– 504 mm SL, from southeastern Australia): AMS I. 3947, 266 mm SL, off Barrenjoey, NSW (33 ° 35 ’S, 151 ° 21 ’E), 46–51 m depth, 22 Feb. 1898; AMS I. 9567, 206 mm SL, near Sydney, NSW (33 ° 50 ’S, 151 ° 20 ’E). 1908; AMS I. 10329, 193 mm SL, off Murray River mouth, SA (35 °00’S, 138 °00’E), 37 m depth, 17 Aug. 1909; AMS I. 10655, 278 mm SL, Lord Howe Island, NSW (31 ° 31 ’S, 159 °05’E), 1910; AMS I. 14928, 242 mm SL, Bellinger River, NSW (30 ° 27 ’S, 152 ° 54 ’E), 10 Nov. 1913; AMS I. 15202, 217 mm SL, Lake Macquarie, NSW (33 °02’S, 151 ° 36 ’E), 18 Nov. 1913; AMS I. 19832 -002, 338 mm SL, 6 km off Whale Beach, Sydney, NSW (33 ° 39 ’S, 151 ° 23 ’E), 55 m depth, 10 Oct. 1976; AMS I. 19893 -013, 2 specimens, 73.0– 95.1 mm SL, north end of Black Point, south of Nadgee River mouth, NSW (37 ° 30 ’S, 149 ° 58 ’E), 5 m depth, 26 Aug. 1976; AMS I. 20721 -001, 275 mm SL, Coffs Harbour, NSW (30 ° 18 ’S, 153 °08’E), 13 Aug. 1978; AMS I. 20723 -001, 127 mm SL, Red Rock, NSW (29 ° 59 ’S, 153 ° 14 ’E), 11 Mar. 1978; AMS I. 37964 -001, 111 mm SL, Hordern’s Bay, Port Hacking, NSW (34 °05’S, 151 °09’E), 10 Aug. 1999; AMS I. 39988 -003, 133 mm SL, Sydney Harbour, NSW (33 ° 51 ’S, 151 ° 15 ’E), 2 Sep. 1999; AMS I. 44633 -022, 87.5 mm SL, Mimosa Rocks National Park, Moon Bay, NSW (36 ° 41 ’ 44 ”S, 149 ° 59 ’ 23 ”E), 3–5 m depth, 10 Apr. 2008; AMS IA. 5700, 166 mm SL, Maroubra, Sydney, NSW (33 ° 51 ’S, 151 ° 16 ’E), 1933; AMS IB. 6292, 142 mm SL, off Ballina, NSW (28 °S, 153 °E), 1962; CSIRO H 4494 -01, 504 mm SL, Lakes Entrance, Vic (37 ° 55 ’S, 148 °01’E), 5 March 1997; NMV A 2903, 419 mm SL, 40 km south of Lakes Entrance, Vic (38 ° 10 ’S, 148 °E), 53 m depth; NMV A 2908, 460 mm SL, 29 km south-southeast of Lakes Entrance, Vic (38 °S, 147 ° 59 ’E), 53 m depth; QM I. 22310, 330 mm SL, Lakes Entrance, Vic (37.53 °S, 148 °E), 14 Aug. 1985. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; gill rakers 4–6 + 13–15 = 19–21; interorbital width 5.9–16.6 % HL; suborbital width 7.6–11.8 % HL; a finger-like interopercular flap present; upper jaw without large caniniform teeth; caudal fin with three to six dark brown to black bars and/or spots posteriorly. Description. Counts and measurements shown in Table 3. Data for all specimens presented first, followed by lectotype condition in parentheses. Snout, area anteroventral to eye, interorbit and occipital region scaled; lower half of suborbital region naked. Snout shorter than orbital diameter in 88 mm SL or smaller specimens, longer in larger specimens (including lectotype). Interorbit narrower than orbital diameter in 419 mm SL or smaller specimens, wider in larger specimens (including lectotype). Upper iris lappet simple, triangular; lower simple, weakly convex. Nasal spine usually absent (including lectotype), rarely one spine present in one specimen (323 mm SL). One suborbital spine usually present below posterior margin of eye in 242 mm SL or smaller specimens, absent in larger specimens (including lectotype); one additional suborbital spine present below and slightly anterior to middle of eye in 133 mm and 166 mm SL specimens. Lower preopercular spine about equal in length to upper spine in smallest (73.0 mm SL) specimen, or slightly longer than upper but not reaching opercular margin in larger specimens (including lectotype). A supplemental preopercular spine present in some specimens, 278 mm SL or smaller, absent in larger specimens (including lectotype). Finger-like interopercular flap present; margin of interopercle smooth. Maxilla reaching to or just beyond anterior margin of pupil, but not to middle of eye (beyond anterior margin of pupil). Anterior portion of upper jaw with some large conical or small caniniform (small caniniform) teeth medially. Palatine teeth irregularly arranged in one (rarely, in 87.5 mm SL specimen), or more usually two to four (ca. four) rows. Vomerine teeth irregularly arranged in one or two (one) rows anteriorly, two to five (three) posteriorly, becoming larger posteriorly; posterior row number tending to increase with growth. Fleshy sensory tubes from suborbitals not covering cheek region; those from preopercle not covering cheek region in 142 mm SL or smaller specimens; not covering it or slightly developed and partially covering its ventral margin in larger specimens (not covering cheek region). Posterior tip of pelvic fin not quite reaching anal-fin origin, or reaching up to base of fourth anal-fin ray (not reaching anal-fin origin). Posterior margin of caudal fin tending to change with growth; slightly rounded or mostly straight by 166 mm SL, mostly straight or slightly concave by 338 mm SL, and slightly concave in larger specimens (mostly straight). Lectotype Non-types AMS I. 3163 n = 20 SL (mm) 291 73.0– 504 Counts: Color in alcohol. Color of lectotype mostly faded, retaining brownish spots approximating bands on pectoral fin and six black bars on posterior portion of caudal fin (Fig. 9). In other specimens (Fig. 8), ground color of head and body pale brown above, paler below. Dorsal surface of body with several indistinct brown or distinct dark brown bands, and/or scattered small brownish spots; scattered small pale spots present dorsally in 461 mm SL specimen. Side of body sometimes with dark brown, gray, or purplish gray band. First and second dorsal fins with pale brownish elliptical spots along rays. Pectoral fin with several pale brown bands, with small pale brown spots tending to form bands, or with large brown spots not forming bands; lower margin of pectoral fin pale. Pelvic fin pale brown or brown, with or without indistinct spots; outer margin of pelvic fin paler. Anal fin pale or with melanophores on posterior two rays, or somewhat dusky in some larger specimens. Caudal fin with three to six dark brown to black bars and/or spots posteriorly, tending to become longer ventrally; other caudal fin regions with several pale brown to brown spots. Distribution. Known from southeastern Australia, from off Calliope River, Qld (24 °01’S) to outside Murray River mouth, SA (138 °E), occurring in coastal bays and estuaries to offshore sand flats in depths from 5 to 100 m (e.g., Last et al., 1983; Kuiter, 1993; Hoese et al., 2006; this study). Size. Maximum length recorded 68 cm (Hutchins & Swainston, 1986). The largest specimen examined during the present study was 504 mm SL (573 mm TL) (Fig. 8). Remarks. Platycephalus caeruleopunctatus bears most similarities to P. speculator in having usually 14 second dorsal- and anal-fin rays, a finger-like interopercular flap, and the caudal fin with three to six dark brown to black bars and/or spots posteriorly (two to four black spots irregularly positioned posteroventrally in 155 mm SL or smaller specimens, or with three or four black spots serially arranged posteroventrally along caudal fin margin in larger specimens in P. speculator). This combination of characters was not apparent in other species of Platycephalus. However, P. caeruleopunctatus has 19–21 gill rakers in total (11–14 in P. speculator), plus a broader interorbit (width 5.9–16.6 % HL in vs. 5.8–14.6 % HL in P. speculator) and suborbital region (suborbital width 7.6–11.8 % HL vs. 6.1 –10.0% HL), which largely separates the two species (Fig. 10)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 164-166, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["McCulloch, A. R. (1922) Check list of the fish and fish-like animals of New South Wales. Part III. Australian Zoologist, 2, 86 - 130. [plates. 25 - 43]","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Whitley, G. P. (1931 b) Studies in ichthyology. No. 5. Records of the Australian Museum, 18, 131 - 160. [plates. 20 - 21]","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp."]}

Genus Platycephalus Bloch, 1795 Platycephalus Bloch, 1795: 96 (type species: Platycephalus spathula Bloch, 1795, a junior synonym of Callionymus indicus Linnaeus, 1758, according to de Beaufort & Briggs, 1962). Calliomorus Lacépède, 1800: 343 (type species: Callionymus indicus). Neoplatycephalus Castelnau, 1872: 87 [type species: Neoplatycephalus grandis Castelnau, 1872, a junior synonym of Platycephalus richardsoni Castelnau, 1872 (see Remarks under P. richardsoni)]. Cacumen Whitley, 1931 a: 326 (type species: Platycephalus speculator Klunzinger, 1872). Planiprora Whitley, 1931 a: 327 (type species: Platycephalus fuscus Cuvier in Cuvier & Valenciennes, 1829). Trudis Whitley, 1931 a: 327 (type species: Platycephalus bassensis Cuvier in Cuvier & Valenciennes, 1829). Longitrudis Whitley, 1931 a: 327 (type species: Platycephalus longispinis Macleay, 1884, a junior synonym of Platycephalus grandispinis Cuvier in Cuvier & Valenciennes, 1829, according to Imamura, 2013 b). Colefaxia Whitley, 1935: 249 [type species: Platycephalus macrodon Ogilby, 1885, a junior synonym of Platycephalus richardsoni (see Remarks under P. richardsoni), established as a subgenus of Neoplatycephalus]. Diagnosis. A genus of the family Platycephalidae with the following characters: head and body usually well depressed (cylindrical and only slightly depressed in Platycephalus laevigatus), covered with ctenoid scales dorsally, cycloid scales ventrally; postorbital region and opercle scaled; snout usually longer than orbital diameter (shorter in some P. laevigatus and smaller P. caeruleopunctatus); eye becoming relatively narrower with growth; interorbit becoming relatively broader with growth; upper surface of eye without papillae or ocular flaps; upper iris lappet usually simple, triangular (broad, usually bilobed in P. westraliae and P. chauliodous, and some P. i nd i c u s and P. australis sp. nov.; simple, weakly pointed in P. conatus and P. richardsoni; broad, well expanded, simple in P. marmoratus and P. orbitalis), lower iris lappet usually broad, simple, weakly convex (absent in P. conatus and P. richardsoni); usually one or two lachrymals (absent in P. laevigatus), one preocular (absent in P. l a e vi ga t us and P. chauliodous), two distinct preopercular and two opercular spines present [sometimes small third (= lowermost) preopercular spine present below others, except in P. richardsoni, in which small third spine always present]; one or two suborbital spines present or absent in smaller specimens (if present, disappearing with growth); some supraorbital, one (sometimes two or more) pterotic, one parietal, one supratemporal, one posttemporal and one supracleithral spine usually present in smaller specimens, tending to disappear with growth (except for supracleithral, these spines absent even in smallest examined specimen of P. laevigatus); teeth on upper and lower jaws, palatine and vomer; tooth band on upper jaw without a distinct notch medially; a single tooth patch on vomer, arranged in shallow V-shape or crescentic; lip margins without papillae; all or most pored lateral-line scales with one pair of sensory ducts and exterior openings posteriorly (P. richardsoni only with all or most scales usually with two pairs of ducts and openings); some anterior pored lateral-line scales sometimes with a small spine; first dorsal fin originating slightly posterior to opercular margin; first and second dorsal fins narrowly separated; pectoral fin rounded posteriorly; pelvic fin with one spine and five soft rays; all soft pelvic-fin rays branched, fourth soft pelvic-fin ray longest. Remarks. The present study recognizes the following 16 species of Platycephalus from Australian waters: P. angustus, P. aurimaculatus, P. bassensis, P. caeruleopunctatus, P. chauliodous, P. conatus, P. endrachtensis, P. fuscus, P. grandispinis, P. laevigatus, P. marmoratus, P. orbitalis, P. richardsoni, P. speculator, P. westraliae and P. australis sp. nov. In addition to these, Platycephalus cultellatus Richardson, 1846, Platycephalus indicus (Linnaeus, 1758) and two undescribed species [Platycephalus sp. 1 and sp. 2 (sensu Nakabo, 2002)] are known elsewhere in the Indo-western Pacific (e.g., Knapp, 1986; Nakabo, 2002; Imamura et al., 2006). Imamura (1996) regarded Neoplatycephalus Castelnau, 1872, Cacumen Whitley, 1931 a, Planiprora Whitley, 1931 a, Trudis Whitley, 1931 a and Longitrudis Whitley, 1931 a to be junior synonyms of Platycephalus based on the phylogenetic relationships of seven species of Platycephalus, or morphological comparisons among the genus. At that time, Imamura (1996) did not address the validity or otherwise of Calliomorus Lacépède, 1800 and Colefaxia Whitley, 1935. However, the former is a synonym of Platycephalus, due to Platycephalus and Calliomorus having been based on P. i ndicus and P. spathula, respectively, the latter being a junior synonym of the former according to de Beaufort & Briggs (1962). Colefaxia was established as a subgenus of Neoplatycephalus on the basis of Platycephalus macrodon, a junior synonym of P. richardsoni (see Remarks under P. richardsoni). Because Neoplatycephalus, a junior synonym of Platycephalus, was itself based on P. richardsoni, Colefaxia is a junior synonym of the former and is invalid [see ICZN (1999: Art. 61.3.3)]., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on page 153, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Bloch, M. E. (1795) Naturgeschichte der auslandischen Fische. 9. J. Morino, Berlin, ii + 192 pp.","Linnaeus, C. (1758) Systema naturae, 10 th edition. Vol. 1. Laurentii Salvii, Holmiae (= Stockholm), ii + 824 pp.","de Beaufort, L. F. & Briggs, J. C. (1962) The fishes of the Indo-Australian Archipelago. Vol. 11. Scleroparei, Hypostomides, Pediculati, Plectognathi, Opisthomi, Discocephali, Xenopterygii. Brill, E. J., Leiden, vi + 481 pp.","Lacepede, B. G. E. (1800) Histoire naturelle des poisons. Vol. 2. Plassan, Imprimeur-Librairie, Paris, lxiv + 632 pp. [20 plates]","Castelnau, F. L. (1872) Contributions to the ichthyology of Australia. No. 1. The Melbourne fish market. Proceedings of the Royal Zoological Acclimatization Society of Victoria, 1, 29 - 242.","Whitley, G. P. (1931 a) New names for Australian fishes. Australian Zoologist, 6, 310 - 334. [plates. 25 - 27]","Klunzinger, C. B. (1872) Zur fischfauna von sud-Australien. Archiv fur Naturgeschichte, 38, 17 - 47. [plates 2]","Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp.","Macleay, W. (1884) Notices of new fishes. Proceedings of the Linnean Society of New South Wales, 9, 170 - 172.","Imamura, H. (2013 b) Validity of Platycephalus grandispinis Cuvier, 1829, with priority over Platycephalus longispinis Macleay, 1884 (Actinopterygii: Scorpaeniformes: Platycephalidae). Species Diversity, 18, 183 - 192. http: // dx. doi. org / 10.12782 / sd. 18.2.183","Whitley, G. P. (1935) Studies in ichthyology. No. 9. Records of the Australian Museum, 19, 215 - 250. [plates. 18]","Ogilby, J. D. (1885) Descriptions of new fishes from Port Jackson. Proceedings of the Linnean Society of New South Wales, 10, 25 - 230.","Richardson, J. (1846) Report on the ichthyology of the seas of China and Japan. Report of the British Association for the Advancement of Science 15 th Meeting, 1845, 187 - 320.","Nakabo, T. (2002) Platycephalidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 601 - 602.","Knapp, L. W. (1986) Family No. 155: Platycephalidae. In: Smith, M. M. & Heemstra, P. C. (Eds.), Smiths ' sea fishes. Springer- Verlag, Macmillan, pp. 482 - 486.","Imamura, H. (1996) Phylogeny of the family Platycephalidae and related taxa (Pisces: Scorpaeniformes). Species Diversity, 1, 123 - 233.","ICZN (International Commission on Zoological Nomenclature) (1999) International code of zoological nomenclature, 4 th edition. The International Trust for Zoological Nomenclature, London, xxiv + 306 pp."]}

Platycephalus bassensis Cuvier in Cuvier & Valenciennes, 1829 Common English name: Southern sand flathead (Figs. 4–5; Table 2) Platycephalus bassensis Cuvier in Cuvier & Valenciennes, 1829: 247 [type locality: Western Port (as Port Western), Vic, Australia]; Quoy & Gaimard, 1834: 683, fig. 3; Castelnau, 1872: 83; McCulloch, 1929: 400; Coleman, 1980: 111, unnumbered fig.; Last et al., 1983: 331, fig. 28.18; Hutchins & Swainston, 1986: 127, fig. 198; May & Maxwell, 1986: 274, unnumbered fig.; Paxton & Hanley, 1989: 468 (in part, including P. westraliae; see Remarks); Knapp, 1991: 29, tab. 3; Kuiter 1993: 103, two unnumbered figs.; Kuiter, 1994: 519, fig. 462; Grant, 2004: 189, unnumbered pl.; Hoese et al., 2006: 940 (in part, including P. westraliae); Imamura, 2006: 304, tab. 1; Gomon, 2008: 520, unnumbered fig. Platycephalus tasmanius Richardson, 1842: 96 (name also on p. 72) (type locality: Port Arthur, Tas, Australia). Trudis bassensis: Whitley, 1931 a: 327; Whitley, 1931 b: 157; Whitley, 1964: 57. Platycephalus bassensis bassensis: Scott et al., 1980: 168, unnumbered fig. Material examined. Holotype: MNHN 1437, 233 mm SL, Western Port, Vic, Australia. Other specimens (26 specimens, 114–418 mm SL, from southeastern Australia): 7 specimens were listed in Imamura (2008); 14 additional specimens: AMS A. 16458, 242 mm SL, AMS A. 16459, 274 mm SL, Hobsons Bay, Vic (37 ° 51 ’S, 144 ° 56 ’E), 1883; AMS E. 4953, 267 mm SL, Hobart Wharf, Tas (42 ° 50 ’S, 147 ° 15 ’E), 1914; AMS E. 5457, 358 mm SL, Recherche Bay, Tas (43 ° 33 ’S, 146 ° 54 ’E), 15 m depth, 24 July 1914; AMS I. 6274, 272 mm SL, Tamar River Heads, Tas (41 ° 20 ’S, 147 °02’E), 1903; AMS I. 7541, 401 mm SL, data unknown; AMS I. 7542, 324 mm SL, Derwent Estuary, Tas (43 °02’S, 147 ° 22 ’E), 1913; AMS I. 20194 -050, 2 specimens, 213–218 mm SL, Investigator Strait, SA (35 ° 20 ’S, 137 ° 50 ’E), 20 m depth, 14 May 1978; BMNH 1855.9.19.56– 57 and BMNH 1855.9.19.67– 68, included in a single jar, 4 syntypes of Platycephalus tasmanius Richardson, 1842 (loc. Port Arthur, Tas, Australia) and one non-type (loc. unknown), all specimens lacking individual labels; one specimen with damaged snout (SL unmeasured), remaining four specimens 170–303 mm SL; CSIRO A 741, 137 mm SL, Kettering, D’Entrecasteaux Channel, Tas (43 °09’S, 147 ° 16 ’E), 1 May 1951; CSIRO CA 3690, 283 mm SL, Great Australian Bight, SA (32 ° 12 ’S, 131 ° 25 ’E – 32 ° 13 ’S, 131 ° 24 ’E), 60 m depth, 8 Dec. 1981; NSMT-P 112668, 418 mm SL, northwest coast of Tas, 28 m depth, Dec. 17 1975; NMV A 29215 -018, 215 mm SL, off Plank Point, Spencer Gulf, SA (33 ° 25 ’ 42 ”S, 137 ° 29 ’ 36 ”E), 22–29 m depth, 13 Oct. 2005; QM I. 22312, 368 mm SL, Lakes Entrance, Vic (37 ° 53 ’S, 148 °02’E), 14 Aug. 1985. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; pored lateral-line scales 70–82; gill rakers 4–6 + 13–16 = 17–22; a distinct interopercular flap absent; ratio of lower/upper preopercular spines 1.6–2.2; ridge and spine absent on supraoccipital; upper jaw without large caniniform teeth; many small dark dots on dorsal surface of head and body; caudal fin with a single large irregular blackish or dark brown marking, or two longitudinal dark brown or black spots posteroventrally. Description. Counts and measurements shown in Table 2. Data for all specimens presented first, followed by holotype condition in parentheses. Interorbit and occipital region scaled; snout and area anteroventral to eye scaled or naked (unclear in holotype), lower half of suborbital region naked. Interorbit narrower than orbital diameter. Upper iris lappet simple, triangular; lower simple, weakly convex. Nasal, preorbital and supplementary preopercular spines absent. Suborbit al spines usually absent; one suborbital spine present below posterior margin of eye in some specimens 176 mm SL or smaller. Ridges and spines absent on supraoccipital. Lower preopercular spine longer than upper, not reaching opercular margin; ratio of lower/upper spines 1.6–2.2 (> 2), tending to become larger with growth (lower spine becoming relatively longer than upper). Distinct interopercular flap absent, although small flap sometimes present (on left side only); margin of interopercle smooth. Maxilla reaching beyond anterior margin of eye. Anterior portion of upper jaw with some large conical or small caniniform (small caniniform) teeth medially. Palatine teeth irregularly arranged in three to nine (ca. five) rows, becoming larger medially; row number tending to increase with growth. Vomerine teeth irregularly arranged in one or two (two) rows anteriorly, two to six (three) posteriorly, becoming larger posteriorly; anterior and posterior row numbers tending to increase with growth. Fleshy sensory tubes on suborbitals and preopercle usually not covering cheek region (including holotype); those from preopercle slightly developed and partially covering ventral margin of cheek region in some specimens. Posterior tip of pelvic fin reaching anal-fin origin or second anal-fin ray (damaged in holotype). Posterior margin of caudal fin usually slightly concave, straight in 137 and 155 mm SL specimens (damaged in holotype). Color in alcohol. Color of holotype mostly faded, retaining a dusky spot on posteroventral portion of caudal fin (Fig. 5 A). In other specimens (Fig. 4), ground color of head and body pale brown to dark brown above, paler below. Dorsal surface of head and body with many small dark dots, that of body with two to four indistinct darker bands. Head with or without brownish band below eye. Side of body with gray or pale purple spots tending to form single band, and dark brown spots. First and second dorsal, and pectoral fins with small brownish spots; those on pectoral fin tending to form bands. Ground color of pectoral fin pale brown or brown, with paler margin ventrally and darker spots often tending to form bands. Pelvic fin brownish or dark brownish with pale outer margin but without spots, or its ground color pale brownish with brownish spots. Anal fin with melanophores on basal portion of rays. Caudal fin with single large irregular blackish or dark brown marking, or two to four longitudinal dark brown to black bands or spots posteroventrally, and several small pale brown to black spots elsewhere. Distribution. Known from southeastern Australia, from Red Rock, NSW (29 ° 59 ’S), to the Great Australian Bight, SA (131 ° 24 ’– 25 ’E), including Vic and Tas, occurring in bays and inlets and on the continental shelf in depths from 15 to ca. 100 m (e.g., Last et al., 1983; May & Maxwell, 1986; Hoese et al., 2006; this study). Size. The largest specimen recorded during the present study was 418 mm SL (483 mm TL), exceeding the previously reported maximum length of 401 mm SL (460 mm TL) (May & Maxwell, 1986). Holotype 5 specimens, including Non-types MNHN 1437 4 syntypes of P. tasmanius n = 21 SL (mm) 233 170–303 * 137– 418 Counts: * Including one specimen with damaged snout and SL unmeasured. Remarks. Among the other species of Platycephalus usually characterized by 14 second dorsal- and anal-fin rays (P. laevigatus, P. grandispinis, P. speculator, P. caeruleopunctatus, P. conatus, P. aurimaculatus and P. richardsoni), P. bassensis is most similar to P. grandispinis in having a somewhat longer lower preopercular spine (extremely long in P. grandispinis), the caudal fin with a single large irregular blackish or dark brown marking, or two longitudinal dark brown or black spots posteroventrally (a single irregular blackish or dark brownish marking in P. grandispinis), and in lacking a distinct interopercular flap, the combination of these characters not occurring in the remaining six species. The present species is easily separable from P. grandispinis in lacking a ridge and spine on the supraoccipital (vs. ridge present on supraoccipital, usually ending in a spine) and having many small dark dots on the dorsal surface of the head and body (vs. dots absent). The ratio of lower/upper preopercular spines may also help distinguish P. bassensis from P. grandispinis (ratio 1.6–2.2 in P. bassensis vs. 1.9–3.3 in P. grandispinis; see also Imamura, 2013 b: fig. 10). Although P. bassensis is also similar to some specimens of P. conatus in having a somewhat longer lower preopercular spine (some specimens of P. conatus with lower spine more than twice length of upper spine), it is separable from the latter in having the upper jaw without large caniniform teeth, as well as from P. aurimaculatus and P. richardsoni (upper jaw with large caniniform teeth anteromedially in latter two species and P. c o na t u s). Platycephalus bassensis is distinguishable from P. caeruleopunctatus in having 70–82 pored latera-line scales (83–93 in P. caeruleopunctatus) and P. speculator in having 17–22 gill rakers in total (11–14 in P. speculator) (pored lateral-line scale data for P. caeruleopunctatus taken from Knapp, 1991 and the present study). Western Australia has been included in the distribution of P. bassensis by some authors, including Paxton & Hanley (1989) and Hoese et al. (2006). They considered Trudis bassensis westraliae Whitley, 1938, originally described on the basis of two specimens from the Swan River estuary, WA, to be a junior synonym of P. bassensis. However, Imamura (2008) showed the validity of the former (as Platycephalus westraliae) and referred that name to the species previously recognized as “ Platycephalus endrachtensis ” (with three black bars on the caudal fin) by authors including Taylor (1964), Gloerfelt-Tarp & Kailola (1984), Allen & Swainston (1988) and Knapp (1999). In addition, as no specimens of P. bassensis from WA were found during the present study, that area is omitted from the distribution of the species recognized here. The revised westernmost record of P. bassensis is the Great Australian Bight, SA (ca. 131 °E), represented by CSIRO CA 3690. The taxonomic status of Platycephalus tasmanius (Fig. 5 B), described from four syntypes from Tasmania by Richardson (1842), has not previously been clearly demonstrated, although Paxton & Hanley (1989), Kuiter (1994) and Hoese et al. (2006) considered it a junior synonym of P. bassensis, without any reasons given. After a comparison of the type specimens of both species, the synonymy was confirmed, owing to the close similarity of the former and non-types of P. bassensis in both the taxonomic characters mentioned above, and in counts and proportional measurements (Table 2)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 159-162, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Cuvier, G. & Valenciennes, A. (1829) Histoire Naturelle des Poissons, vol. 4. F. G. Levault, Paris-Strasbourg, xxvi + 2 + 518 pp.","Quoy, J. R. C. & Gaimard, J. P. (1834) Voyage de decouvertes de \" L'Astrolabe \" axecute par ordre du Roi, pendant les annees 1826 - 29, sous le commandement de M. J. Dumont d'Urville, vol. 3. Pilet Aine, Paris.","Castelnau, F. L. (1872) Contributions to the ichthyology of Australia. No. 1. The Melbourne fish market. Proceedings of the Royal Zoological Acclimatization Society of Victoria, 1, 29 - 242.","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Kuiter, R. H. (1994) Family Platycephalidae. In: Gomon, M. F., Glover, J. C. M. & Kuiter, R. H. (Eds.), The fishes of Australia's south coast. State Print, Adelaide, pp. 514 - 522.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Richardson, J. (1842) Description of Australian fish. Transactions of the Zoological Society of London, 3, 69 - 131. [plates. 4 - 6]","Whitley, G. P. (1931 a) New names for Australian fishes. Australian Zoologist, 6, 310 - 334. [plates. 25 - 27]","Whitley, G. P. (1931 b) Studies in ichthyology. No. 5. Records of the Australian Museum, 18, 131 - 160. [plates. 20 - 21]","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127.","Scott, T. D., Glover, C. J. M. & Southcott, R. V. (1980) The marine and freshwater fishes of south Australia, 2 nd edition. DJ Woolman, Adelaide, 392 pp.","Imamura, H. (2008) Synonymy of two species of the genus Platycephalus and validity of Platycephalus westraliae (Pisces: Scorpaeniformes). Ichthyological Research, 55, 399 - 406. http: // dx. doi. org / 10.1007 / s 10228 - 008 - 0046 - 1","Imamura, H. (2013 b) Validity of Platycephalus grandispinis Cuvier, 1829, with priority over Platycephalus longispinis Macleay, 1884 (Actinopterygii: Scorpaeniformes: Platycephalidae). Species Diversity, 18, 183 - 192. http: // dx. doi. org / 10.12782 / sd. 18.2.183","Whitley, G. P. (1938) Studies in ichthyology. No. 11. Records of the Australian Museum, 20, 195 - 199. [plates. 21]","Taylor, W. R. (1964) Fishes of Arnhem Land. In: Specht, R. L. (Ed.), Records of the American-Australian scientific expedition to Arnhem Land, Vol. 4. Melbourne University Press, Melbourne, pp. 45 - 307.","Gloerfelt-Tarp, T. & Kailola, P. J. (1984) Trawled fishes of southern Indonesia and northern Australia. The Australian Development Assistance Bureau, the Directorate General of fisheries, Indonesia and the German Agency for Technical Cooperation, Jakarta, xvi + 2 plates. + 406 pp.","Allen, G. R. & Swainston, R. (1988) The marine fishes of north-western Australia. A field guide for anglers and divers. Western Australian Museum, Perth, vi + 201 pp.","Knapp, L. W. (1999) Platycephalidae. In: Carpenter, K. E. & Niem, V. H. (Eds.), FAO species identification guide for fishery purposes. The living marine resources of the western Central Pacific. Vol. 4. Bony fish. Part 2 (Mugilidae to Carangidae). FAO, Rome, pp. 2385 - 2421."]}

Key to the species of Australian Platycephalus The following key to the species of Platycephalus from Australia has been prepared for adults and young specimens with adult-like characters (e.g., specimens of P. f us cu s ca. 100 mm SL or smaller differ from adults in caudal fin color pattern and would not key comfortably to that species); see individual species descriptions for identification of smaller specimens. 1 a. Head and body subcylindrical, slightly depressed; upper preopercular spine longer than lower................ P. laevigatus 1 b. Head and body strongly depressed; lower preopercular spine longer than upper..................................... 2 2 a. One to three large caniniform teeth present anteromedially on upper jaw.......................................... 3 2 b. Large caniniform teeth absent on upper jaw................................................................. 6 3 a. Anal-fin rays usually 13; branched caudal-fin rays 9–11; occipital region naked; margin of interopercle scalloped................................................................................................... P. chauliodous 3 b. Anal-fin rays usually 14; branched caudal-fin rays 12–13; occipital region scaled; margin of interopercle smooth.......... 4 4 a. Pectoral-fin rays usually 19; swimbladder absent; abdominal cavity pale.............................. P. aurimaculatus 4 b. Pectoral-fin rays usually 20 or more; swimbladder present; abdominal cavity dark brown or black...................... 5 5 a. Pored lateral-line scales usually with one pair of sensory ducts with posterior external openings; total gill rakers 7–11.................................................................................................... P. conatus 5 b. Pored lateral-line scales usually with two pairs of sensory ducts with posterior external openings (some scales sometimes with one, three or four pairs of ducts); total gill rakers 12–17............................................. P. richardsoni 6 a. Distinct interopercular flap absent (small flap sometimes present)............................................... 7 6 b. Finger-like interopercular flap present..................................................................... 8 7 a. Total gill rakers> 17; caudal fin with 1–2 blackish or dark brown markings or spots posteroventrally.................... 9 7 b. Total gill rakers 8 a. Total gill rakers> 18................................................................... P. caeruleopunctatus 8 b. Total gill rakers 9 a. Ridge and spines absent on supraoccipital; many small dark dots scattered on dorsal surface of head and body... P. bassensis 9 b. Ridge present on supraoccipital, usually ending in a spine; small distinct dark dots absent from dorsal surface of head and body......................................................................................... P. grandispinis 10 a. Head and body marbled with irregular dark brown, brown and pale bands and spots...................... P. marmoratus 10 b. Head and body without distinct spots and bands..................................................... P. orbitalis 11 a. Caudal fin with several dark spots dorsally or posteroventrally, but without bands.................................. 12 11 b. Caudal fin with several dark bands....................................................................... 13 12 a. First dorsal fin with a single small isolated spine anteriorly; second dorsal- and anal-fin rays usually 14; caudal fin with several dark spots posteroventorally................................................................... P. speculator 12 b. First dorsal fin usually with two small isolated spines anteriorly; second dorsal- and anal-fin rays usually 13; caudal fin with several dark spots dorsally........................................................................ P. fuscus 13 a. Teeth present on dorsal surface of anterolateral edge of upper jaw....................................... P. angustus 13 b. Teeth absent on dorsal surface of anterolateral edge of upper jaw................................................ 14 14 a. Upper iris lappet usually broad, bilobed; caudal fin with yellow marking posterodorsally when fresh........... P. westraliae 14 b. Upper iris lappet triangular, simple; caudal fin either lacking yellow marking, or not situated as above................. 15 15 a. Total gill rakers 10–15 (tending to decrease with growth); caudal fin usually with four or more dark bands, lacking a yellow marking when fresh........................................................................ P. endrachtensis 15 b. Total gill rakers 4–10 (tending to decrease with growth); caudal fin with three or four dark bands, a yellow marking on midline when fresh............................................................................ P. australis sp. nov.

Platycephalus speculator Klunzinger, 1872 Common English name: Southern bluespotted flathead (Figs. 11���12; Table 4) Platycephalus speculator Klunzinger, 1872: 28 (type locality: Hobson���s Bay, Vic, Australia); McCulloch, 1929: 400; Hutchins & Thompson, 1983: 78, fig. 109; Last et al., 1983: 334, fig. 28.22; Hutchins & Swainston, 1986: 127, fig. 200; May & Maxwell, 1986: 277, unnumbered fig.; Paxton & Hanley, 1989: 470; Knapp, 1991: 29, tab. 3; Kuiter 1993: 103, unnumbered fig.; Grant, 2004: 194, unnumbered pl.; Hoese et al., 2006: 943; Imamura, 2006: 305, tab. 1; Gomon, 2008: 521, unnumbered fig. Platycephalus castelnaui Macleay, 1881: 587 (type locality: King George Sound, WA, Australia); McCulloch, 1929: 401; Coleman, 1980: 115, unnumbered color fig. Cacumen speculator: Whitley, 1931 a: 326. Planiprora melsoni Whitley, 1945: 38 (type locality: beach at Geraldton, WA, Australia). Planiprora castelnaui: Whitley, 1952: 32; Whitley, 1964: 57. Neoplatycephalus speculator: Whitley, 1964: 57. Material examined. Holotype: SMNS 1570, 256 mm SL, Hobson's Bay, Vic, Australia. Other specimens (20 specimens, 117���493 mm SL, from southern Australia): AMS I. 16354 -001, holotype of Platycephalus castelnaui Macleay, 1881, 281 mm SL, King George Sound, WA; AMS I. 17611 -010, 117 mm SL, York Peninsula, SA (35 ��00���S, 137 �� 44 ���E), 1 m depth, 23 Dec. 1973; AMS I. 17615 -010, 144 mm SL, 16 km south of Tumby Bay, SA (34 �� 24 ���S, 136 ��08���E), 27 Dec. 1973; AMS I. 20180 -048, 3 specimens, 152���157 mm SL, Penneshaw, Kangaroo Island, SA (35 �� 44 ���S, 138 �� 58 ���E), 3���5 m depth, 9 March 1978; AMS IB. 1367, holotype of Planiprora melsoni Whitley, 1945, 118 mm HL (head only), beach at Geraldton, WA; AMS IB. 2119, 287 mm SL, Port Arlington, Vic (38 ��07���S, 144 �� 40 ���E), 1947; AMS IB. 2120, 243 mm SL, Port Arlington, Vic (38 ��07���S, 144 �� 40 ���E), 7 Sep. 1947; AMS IB. 7593, 339 mm SL, Port Phillip Bay, Vic (38 ��09���S, 144 �� 52 ���E), 1966; CSIRO H 4317 -02, 451 mm SL, CSIRO H 4317 -04, 315 mm SL, fish market, from Port Phillip Bay, Vic, March 1997; NMV 29242 -001, 232 mm SL, off Wallaroo, Spencer Gulf, SA (33 �� 49 ��� 51 ���S, 137 �� 32 ���01���E), 22���29 m depth, 7 Oct. 2005; NMV A21467, 2 specimens, 144���179 mm SL, Vic, 13 Nov. 1979; NMV A 3509, 220 mm SL, off Point Ormand, just south of Hobsons Bay, Port Phillip Bay, Vic (37 �� 53 ���S, 144 �� 59 ���E), 29 Oct. 1980; NMV A 19854, 255 mm SL, 3.2 km west of Sandringham, Port Phillip Bay, Vic (37 �� 57 ���S, 144 �� 57 ���E), 30 March 1971; WAM P. 24537 - 0 0 1, 404 mm SL, Mandurah, WA (32 �� 32 ���S, 115 �� 43 ���E), 1973; WAM P. 25720 -001, 493 mm SL, Five Fathom Bank, WA (32 �� 12 ���S, 115 �� 40 ���E), Dec. 1976; WAM P. 28294 -002, 212 mm SL, Lucky Bay, WA (34 ��00���S, 122 �� 14 ���E), 12 Apr. 1984. Diagnosis. A species of Platycephalus with the following combination of characters: second dorsal- and analfin rays usually 14; gill rakers 2���3 + 9���11 = 11���14; interorbital width 5.8���14.6 % HL; suborbital width 6.1 ���10.0% HL; a finger-like interopercular flap present; upper jaw without large caniniform teeth; caudal fin with two to four black spots irregularly positioned posteroventrally in 155 mm SL or smaller specimens, three or four black spots serially arranged posteroventrally along caudal-fin margin in larger specimens. Description. Counts and measurements shown in Table 4. Data for all specimens presented first, followed by holotype condition in parentheses. Snout, area anteroventral to eye, interorbit and occipital region scaled; lower half of suborbital region naked. Interorbit narrower than orbital diameter. Upper iris lappet simple, triangular; lower simple, weakly convex. Nasal spine absent. Preorbital spine usually absent (including holotype), rarely present in some specimens. One suborbital spine usually present below posterior margin of eye in 221 mm SL or smaller specimens, absent in larger specimens (including holotype); one additional suborbital spine present below and slightly anterior to middle of eye in 144 mm SL specimen. Lower preopercular spine slightly longer than upper, not reaching opercular margin. Supplemental preopercular spine usually absent, present only on right side in 152 mm SL specimen. Finger-like interopercular flap present; margin of interopercle smooth. Maxilla reaches or beyond anterior margin of eye (beyond it). Anterior portion of upper jaw with some large conical or small to large caniniform (small caniniform) teeth medially. Palatine teeth irregularly arranged in usually two, sometimes three or four rows (two); row number tending to increase with growth. Vomerine teeth irregularly arranged in one or two (one) rows anteriorly, one to four (one) posteriorly, becoming larger posteriorly; posterior row number tending to increase with growth. Fleshy sensory tubes from suborbitals not covering cheek region; those from preopercle not covering cheek region in 144 mm SL and smaller specimens, usually partly or mostly covering ventral margin of cheek region in larger specimens (partly covering), rarely tubes not covering cheek region in some specimens. Posterior tip of pelvic fin reaching anal-fin origin to base of fourth anal-fin ray (reaching base of second anal-fin ray). Posterior margin of caudal fin tending to change with growth, slightly rounded or mostly straight by 212 mm SL, mostly straight or slightly concave by 339 mm SL, slightly concave in larger specimens (damaged in holotype). Color in alcohol. Color of holotype completely faded (Fig. 12 A). In other specimens (Fig. 11), ground color of head and body pale brown to dark brown above, paler below. Dorsal surface of head and body with many darker dots or small white spots, or without dots and spots. Some specimens with two distinct or indistinct darker bands below second dorsal fin. Side of body with or without gray band. First and second dorsal, and pectoral fins with small pale to dark brownish spots; those on pectoral fin tending to form bands. Pelvic fin pale brown or brown, with or without darker spots tending to form bands; outer margin of pelvic fin paler. Anal fin pale in smaller specimens, with melanophores along rays posteriorly in 221 mm SL or larger specimens; melanophores scattered over most of anal fin in 287 mm SL or larger specimens. Caudal fin with two to four black spots irregularly arranged posteroventrally in 155 mm SL or smaller specimens, with three or four black spots serially arranged posteroventrally along caudal fin margin in larger specimens; other areas of caudal fin with several small pale brown or brown spots or with indistinct irregular pale to dark brown markings. Distribution. Known from southern Australia, from Cape Everard, Vic (149 �� 16 ���E) to Kalbarri, WA (27 �� 42 ���S), including Tas and SA, mainly on shallow sand patches among seagrasses in bays and large estuaries in depths from 1 to 30 m (e.g., Last et al., 1983; Hoese et al., 2006; Gomon, 2008; this study). Size. Maximum length 90 cm (Last et al., 1983). The largest specimen examined in the present study was 493 mm SL (577 mm TL) (Fig. 11). Remarks. The holotype of P. speculator has 12 second dorsal-fin rays, whereas other specimens possessed 13���14 (usually 14). Because all specimens conformed closely with the other diagnostic characters, incuding other counts and proportional measurements (Table 4), the occurrence of 12 second dorsal-fin rays in the holotype is regarded as a rare intraspecific variation. Platycephalus speculator most closely resembles P. caeruleopunctatus (see Remarks under P. caeruleopunctatus for a comparison of the two species). The former is also similar to P. f uscus, P. endrachtensis, P. westraliae, P. angustus and P. australis sp. nov. (described below) from Australia, and P. indicus and P. cultellatus from the Indo-West Pacific in having a finger-like interopercular flap, but separable from them in having usually 14 second dorsal- and anal-fin rays (vs. usually 13 rays). Platycephalus castelnaui Macleay, 1881 (Fig. 12 B) and Planiprora melsoni Whitley, 1945 (Fig. 12 C) were regarded as junior synonyms of P. speculator by Paxton & Hanley (1989) and Hoese et al. (2006), but without justification. However, the synonymy of P. speculator and P. castelnaui is supported in this study on the basis of sharing 14 dorsal- and anal-fin rays, 13 gill rakers in total, a narrower interorbit (10.5 % HL) and suborbital width (8.2 % HL), and a finger-like interopercular flap present (see also Table 4). Although only the head (118 mm in length) of the holotype of Planiprora melsoni was retained, comparison with a non-type specimen of P. speculator with a similar head length (WAM P. 24537, 404 mm SL, 120 mm HL) showed the head proportional measurements to agree closely (SNL 26.3 % HL, OD 16.6 %, UJL 32.3 %, LJL 49.5 %, IW 12.4 %, POL 57.6 %, SW 9.0% in WAM P. 24537; see Table 4), confirming their conspecificity., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 167-170, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Klunzinger, C. B. (1872) Zur fischfauna von sud-Australien. Archiv fur Naturgeschichte, 38, 17 - 47. [plates 2]","McCulloch, A. R. (1929) Check list of the fishes recorded from Australia. Part 3. Memoirs of the Australian Museum, 5, 329 - 436. http: // dx. doi. org / 10.3853 / j. 0067 - 1967.5.1929.475","Hutchins, B. & Thompson, M. (1983) The marine and estuarine fishes of south-western Australia. Western Australian Museum, Perth, 103 pp.","Last, P. R., Scott, E. O. G. & Talbot, H. F. (1983) Fishes of Tasmania. Tasmanian Fisheries Development Authority, Hobart, viii + 563 pp.","Hutchins, B. & Swainston, R. (1986) Sea fishes of southern Australia. Complete field guide for anglers and divers. Swainston Publishing, Perth, 180 pp.","May, J. L. & Maxwell, J. G. H. (1986) Field guide to trawl fish from temperate waters of Australia, revised edition. CSIRO Division of Fisheries Research, Hobart, 492 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1991) Platycephalus chauliodous, a new flathead fish from the eastern Indian Ocean (Teleostei: Platycephalidae). Proceedings of the Biological Society of Washington, 104, 23 - 29.","Kuiter, R. H. (1993) Coastal fishes of south-eastern Australia. Crawford House Press Pty Ltd, Bathurst, New South Wales, xxxi + 437 pp.","Grant, E. M. (2004) Grant's guide to fishes, 10 th edition. E. M. Grant Pty. Ltd., Scarborough, Queensland, 880 pp.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Gomon, M. F. (2008) Family Platycephalidae. In: Gomon, M. F., Bray, D. J. & Kuiter, R. H. (Eds.), Fishes of Australia's southern coast. Reed New Holland, Chatswood, pp. 515 - 521.","Macleay, W. (1881) Descriptive catalogue of the fishes of Australia. Part II. Proceedings of the Linnean Society of New South Wales, 5, 510 - 629. [plates. 13 - 14]","Coleman, N. (1980) Australian Sea Fishes. South of 30 ° S. Doubleday Australia Pty. Ltd., Lane Cove, New South Wales, 302 pp.","Whitley, G. P. (1931 a) New names for Australian fishes. Australian Zoologist, 6, 310 - 334. [plates. 25 - 27]","Whitley, G. P. (1952) Some noteworthy fishes from eastern Australia. Proceedings of the Royal Zoological Society of New South Wales, 1950 - 51, 27 - 32.","Whitley, G. P. (1964) A survey of Australian ichthyology. Proceedings of the Linnean Society of New South Wales, 89, 11 - 127."]}

Platycephalus orbitalis Imamura & Knapp, 2009 Common English name: Western-Australian flathead (Fig. 15) Platycephalus orbitalis Imamura & Knapp, 2009: 58, figs. 1–5 (type locality: northwest of Rottnest Island, WA, Australia). Platycephalus marmoratus (not Stead, 1908): Hutchins & Thompson, 1983: 78, fig. 112; Hutchins & Swainston, 1986: 127, fig. 204 (in part); Paxton & Hanley, 1989: 469 (in part); Knapp, 1991: 29, tab. 3 (in part); Hutchins, 2001: 28; Hoese et al., 2006: 943 (in part). Material examined. Holotype: CSIRO H 6349 -04, 267 mm SL, northwest of Rottnest Island, WA, Australia (31 ° 52.56 ’S, 115 ° 18.30 ’E – 31 ° 52.60 ’S, 115 ° 18.49 ’E), 100–102 m depth, 10 Apr. 2006. Paratypes: 6 specimens, 236–333 mm SL, from western Australia, listed in Imamura & Knapp (2009). Other specimen (1 specimen, 337 mm SL, from northwestern Australia): WAM P. 27606 -001, 337 mm SL, WA (32 °03’S, 115 ° 44 ’E), 28 March 1982. Diagnosis. A species of Platycephalus with the following combination of characters: dorsal- and anal-fin rays 13; pored lateral-line scales 65–68, all lacking spines; gill rakers 2 + 6–7 = 8–10; snout and interorbit naked; interopercular flap absent; margin of interopercle scalloped; upper jaw without large caniniform teeth; fleshy sensory tubes from infraorbitals and preopercle well developed, covering cheek region, except anteroventrally; head and body lacking distinct large spots and bands dorsally; caudal fin blackish with a pale posterior margin. Description. A full description was given by Imamura & Knapp (2009). Distribution. Known only from western Australian waters, from Cape Leeuwin (34 °S) north to Cape Cuvier (24 °S) (e.g., Hutchins, 2001; Imamura & Knapp, 2009; this study). Size. Known maximum length 337 mm SL (400 mm TL) (Imamura & Knapp, 2009; this study). Remarks. Although Imamura & Knapp (2009) described the holotype as having 2 + 4 = 6 gill rakers, reexamination has confirmed the number as 2 + 6 = 8. The present species is mostly similar to P. marmoratus (see Remarks for P. marmoratus).

Platycephalus australis sp. nov. Proposed common English name: Australian bartail flathead (Figs. 34 ���35, 36 A, 37; Tables 13���14) Platycephalus indicus (not Linnaeus, 1758): Paxton & Hanley, 1989: 469; Knapp, 1999: 2409, unnumbered fig. (in part); Hoese et al., 2006: 942. Material examined. Holotype: WAM P. 29859 -001, 477 mm SL, Exmouth Gulf, WA, Australia (22 �� 28 ���S, 114 �� 13 ���E), 22 May 1988. *Tending to decrease with growth. **Nine or 10 caudal-fin rays in specimens Diagnosis. A species of Platycephalus with the following combination of characters: first dorsal fin with a single small isolated spine anteriorly; second dorsal-fin rays 13���14, usually 13: anal-fin rays 13; pectoral-fin rays 19���21, usually 20; gill rakers 1���2 + 3���8 = 4���10 (tending to decrease with growth); postorbital length 51.6���63.6 % HL; snout, area anteroventral to eye, interorbit, and occipital region scaled; upper iris lappet usually simple, triangular; a finger-like interopercular flap present; upper jaw without large caniniform teeth; teeth absent on dorsal surface of anterolateral edge of upper jaw; palatine teeth in two rows; vomerine teeth usually in one row; caudal fin with a yellow marking on midline when fresh. Description. Counts and measurements shown in Table 13. Data for all specimens, including both non-types and paratypes, presented first, followed by holotype in parentheses. Body greatly depressed, mostly covered with ctenoid scales, but some cycloid scales on undersurface. Head greatly flattened, length 2.8���3.2 (3.2) in SL; scales covering snout, a small area anteroventral to eye, interorbit, occipital region, nape, and postorbital and opercular regions; suborbital region naked. Snout robust, longer than orbital diameter, length 3.6 ���4.0 (3.8) in HL. Upper surface of eye without papillae. Upper iris lappet simple, triangular dorsally; lower weakly convex (unobservable in holotype) (Fig. 35 B). Interorbital width 5.5���15.3 (6.3) in HL, increasing with growth; orbital diameter 5.4 ���8.0 (8.4) in HL, decreasing with growth; interorbit narrower than orbital diameter in smaller specimens, becoming equal to or wider than orbital diameter by 232 mm SL (including holotype). Spines and ridges on top and side of head weakly developed (Fig. 35 B). Nasal usually with a single distinct spine in 123 mm SL or smaller specimens (absent in 106 mm SL specimen), a small or rudimentary spine, or spine absent in larger specimens (absent in holotype). Lachrymal with two embedded antrorse spines. Single preocular spine present. Single preorbital spine present or absent in 109 mm SL or smaller specimens, absent in larger specimens (including holotype). Suborbital ridge usually with a spine below and slightly posterior to posterior margin of eye, often with a spine below and slightly anterior to middle of eye in 125 mm or smaller specimens (lacking both spines on right side of 123 mm SL specimen); with or without a spine below and slightly posterior to posterior margin of eye in 175 to 270 mm SL specimens; and without spines in 273 mm SL or larger specimens (including holotype). Supraorbital ridge serrated posteriorly, with one to seven small spines (two on left and one on right in holotype). Single postocular spine present. Pterotic with one to six spines (one). Frontal and supraoccipital with entirely smooth ridges. Parietal with one or two spines (rudimental spine on left, spine absent on right). Supratemporal usually with one spine (sometimes two or zero) in 294 mm SL or smaller specimens (spines absent in 415 mm SL specimen and holotype). Posttemporal usually with one spine; rarely absent in some specimens in 125 mm SL or larger (including holotype). Supracleithrum usually with one spine (spine absent only in holotype). Preopercle with two distinct spines; lower spine slightly longer than upper (including holotype) or spines subequal, not reaching posterior margin of opercle; upper spine with supplementary spine in 147 mm SL or smaller specimens, usually without in 184 mm SL or larger specimens (including holotype) (rudimental supplementary spine in 294 mm SL specimen). Opercle with two spines, lacking prominent ridge. Finger-like interopercular flap present; margin of interopercle not scalloped. Maxilla reaching beyond anterior margin of orbit, length 2.6���2.8 (2.6) in HL, tending to extend posteriorly with growth [just below posterior margin of orbit in largest examined specimen (holotype)]. Teeth in bands on jaws and palatine, a single shallow V-shaped or crescentic patch on vomer (shallow crescentic patch in holotype); tooth band on upper jaw lacking distinct notch mesially. Upper jaw with several large conical or small caniniform (large conical in holotype) teeth anteromedially, villiform teeth anterolaterally, teeth tending to be larger medially; remainder of jaw with small conical and villiform teeth; teeth absent on dorsal surface of anterolateral edge of upper jaw. Lower jaw usually with one villiform tooth row laterally (a narrow villiform band in holotype) and one small to moderate conical tooth row medially, teeth tending to become larger posteriorly; lateral villiform teeth forming two or three rows anteriorly in several paratypes and non-types. Palatine with two tooth rows; inner row with small conical teeth, outer row with small villiform teeth. Vomer with single villiform to moderate conical (moderate conical in holotype) tooth row; some specimens 147 mm SL or larger with additional villiform teeth anteriorly and/or posteriorly (teeth anteriorly in holotype). Lip margins without papillae. Fleshy sensory tubes on suborbitals and preopercle not covering cheek region. Pored scales in lateral line each with a pair of sensory ducts and exterior openings posteriorly. First dorsal fin originating posterior to opercular margin. First and second dorsal fin narrowly separated. Pectoral fin rounded posteriorly, length 5.6���8.1 (7.1) in SL. Posterior tip of pelvic fin reaching between anus and base of fourth anal-fin ray (reaching anal-fin origin), length 4.0��� 5.1 (4.5) in SL. Caudal fin usually slightly rounded or mostly straight posteriorly (slightly concave in holotype), length 5.5���6.8 (7.2) in SL. Color in alcohol. In holotype (Fig. 34), body and head brown above, pale below. Head with single indistinct, dark brown band crossing occipital and anterior opercular regions. Side of head with many small dark brown spots. Body with two dark brown bands below second dorsal fin; anterior band broad, posterior band narrow. First and second dorsal, pectoral and pelvic fins with small, dark brownish spots along rays. Pectoral and pelvic fins with paler outer margins. Anal fin pale, with melanophores along membrane between seventh to last fin rays; melanophores thicker posteriorly. Caudal fin with the following markings: a long dark brown band on both the upper and lower portions; short blackish band above long dark brown band on upper portion; short dark brown band below long dark brown band on lower portion; small brownish spots on dorsal portion. In paratypes and non-types, caudal fin usually with three dark brown or black bands; middle band paler than dorsal and ventral bands. In CSIRO H 3916 -01 (415 mm SL, paratype) caudal fin with four bands; middle two bands brownish, and dorsal and ventral two bands dark brown. Color when fresh (based on color photographs; Figs. 36 A, 37). Caudal fin with yellow marking on midline; its size variable, small (Fig. 36 A), medium (Fig. 37 A) or large (Fig. 37 A). Other coloration similar to preserved condition. Distribution. Known from northern Australia, from Exmouth Gulf, WA (22 �� 28 ���S) to south of Moreton Bay, Qld (27 �� 25 ���S) in depths from at least 0.5���36 m (this study). Size. Recorded maximum length 477 mm SL (554 mm TL) (this study; Fig. 34). Etymology. The specific name australis derived from Latin is proposed in reference to its type locality, Australia. Remarks. Platycephalus australis sp. nov. has previously been misidentified as Platycephalus indicus, both species being characterized by usually 13 second dorsal- and anal-fin rays, the snout, area anteroventral to the eye, interorbit and occipital region scaled, large caniniform teeth absent on the upper jaw, a finger-like interopercular flap, palatine teeth arranged in two rows and the caudal fin with a yellow marking on the middle when fresh. In fact, the yellow mid-caudal fin marking occurs only in the above two species. However, P. australis sp. nov. is separable from P. i n di c us in having fewer total gill rakers (4���10 in P. australis vs. 7���10 in P. i n di c us, tending to decrease with growth in both species) and a longer postorbital region (51.6���63.6 % HL vs. 51.4���61.6 % HL) than the latter at a comparable size, although a considerable overlap occurs (Fig. 38). In addition, P. australis sp. nov. differs from P. indicus in having a greater number of pectoral-fin rays (19���21, usually 20 vs. 18-20, usually 19) (Table 14). The validity of P. australis sp. nov. has also been demonstrated from genetically reconstructed phylogenetic relationships of Platycephalus, showing that ��� P. indicus ��� from Australia, including a specimen with a yellow marking on the mid-caudal fin (thus probably P. australis sp. nov.) and P. i nd i c u s from the Indian Ocean and Indonesia are included in different monophyletic clades, the degree of speciation between the two species being similar to that of P. conatus and P. richardsoni (sister species) (W. White, personal communication, 20 June 2013). A detailed comparison of P. australis sp. nov. and P. i ndicus is presented in Table 13. Among other species with usually 13 second dorsal- and anal-fin rays, P. australis sp. nov. is easily distinguished from P. endrachtensis in having a broader interorbit and longer postorbital region (interorbital width 6.5���18.1 % HL and postorbital length 51.6���63.6 % HL in P. australis sp. nov. vs. 7.7 ���12.0% HL and 50.7���56.9 % HL in P. endrachtensis) (Fig. 30), from P. angustus, P. cultellatus, P. f u s cu s, and Platycephalus sp. 1 and sp. 2 (sensu Nakabo, 2002) in having the first dorsal fin with a single small isolated spine anteriorly (usually two in the other species), and from P. westraliae in having a simple triangular upper iris lappet (usually broad and bilobed in P. westraliae)., Published as part of Imamura, Hisashi, 2015, Taxonomic revision of the flathead fish genus Platycephalus Bloch, 1785 (Teleostei: Platycephalidae) from Australia, with description of a new species, pp. 151-207 in Zootaxa 3904 (2) on pages 199-203, DOI: 10.11646/zootaxa.3904.2.1, http://zenodo.org/record/233552, {"references":["Linnaeus, C. (1758) Systema naturae, 10 th edition. Vol. 1. Laurentii Salvii, Holmiae (= Stockholm), ii + 824 pp.","Paxton, J. R. & Hanley, J. E. (1989) Platycephalidae. In: Paxton, J. R., Hoese, D. F., Allen, G. R. & Hanley, J. E. (Eds.), Zoological catalogue of Australia. Vol. 7. Pisces. Petromyzontidae to Carangidae. Australian Government Publishing Service, Canberra, pp. 465 - 472.","Knapp, L. W. (1999) Platycephalidae. In: Carpenter, K. E. & Niem, V. H. (Eds.), FAO species identification guide for fishery purposes. The living marine resources of the western Central Pacific. Vol. 4. Bony fish. Part 2 (Mugilidae to Carangidae). FAO, Rome, pp. 2385 - 2421.","Hoese, D. F., Bray, D. J., Paxton, J. R. & Allen, G. R. (2006) Fishes. In: Beesley, P. L. & Wellas, A. (Eds.), Zoological catalogue of Australia. Vol. 35, part 2. ABRS & CSIRO Publishing, Collingwood, pp. 934 - 948.","Nakabo, T. (2002) Platycephalidae. In: Nakabo, T. (Ed.), Fishes of Japan with pictorial keys to the species, English edition. Tokai University Press, Tokyo, pp. 601 - 602."]}

Knapp, Leslie W., Imamura, Hisashi (2014): Grammoplites vittatus (Valenciennes), (Scorpaeniformes, Platycephalidae), removed from synonymy with Grammoplites scaber (Linnaeus). Zootaxa 3846 (3): 447-450, DOI: http://dx.doi.org/10.11646/zootaxa.3846.3.8

Key to the Species of Grammoplites 1a Uppermost preopercular spine not reaching opercular margin; all LL scales bearing a spine; no discrete black blotch at rear of first dorsal fin; anal fin rays 12..................................................................................................................... 2 1b Uppermost preopercular spine reaching to or past opercular margin; posterior LL scales may not bear spines; rear of first dorsal fin with prominent black blotch; anal-fin rays 13................................................................... G. suppositus 2a Total GR count usually 6; dorsum brown with 3���4 darker bands; anal fin dusky, often with dark ray tips................ 3 2b Total GR count 7���8; dorsum gray or light tan with 5���6 indistinct bands; anal fin whitish........................... G. vittatus 3a Interorbital width 9.8���13.1 % of HL; caudal fin usually with broad dark band at rear edge .......................... G. scaber 3b Interorbital width 6.4���8.4 % of HL; caudal fin yellowish.. ............................................................................ G. knappi Remarks. Grammoplites suppositus (Troschel, 1840) is found from the Red Sea to the Gulf of Aden, Persian Gulf and the Bay of Bengal. The range of G. vittatus appears to be restricted to the Persian Gulf and the northern Arabian Sea. Grammoplites scaber is more widely distributed, from the Bay of Bengal to the Gulf of Thailand and Indonesia. Grammoplites knappi was reported by Imamura and Amaoka (1994) from the Gulf of Thailand, Hainan Island, China. and from off Sarawak, Borneo. Additional collecting will hopefully determine more exact range limits for these species., Published as part of Knapp, Leslie W. & Imamura, Hisashi, 2014, Grammoplites vittatus (Valenciennes), (Scorpaeniformes, Platycephalidae), removed from synonymy with Grammoplites scaber (Linnaeus), pp. 447-450 in Zootaxa 3846 (3) on page 450, DOI: 10.11646/zootaxa.3846.3.8, http://zenodo.org/record/4928695

Ho, Hsuan-Ching, Heemstra, Phillip C., Imamura, Hisashi (2014): A new species of the sandperch genus Parapercis from the western Indian Ocean (Perciformes: Pinguipedidae). Zootaxa 3802 (3): 335-345, DOI: http://dx.doi.org/10.11646/zootaxa.3802.3.3