TAXONOMY OF THE SUBFAMILY LYCODINAE (FAMILY ZOARCIDAE) IN JAPAN AND ADJACENT Title WATERS Author(s) TOYOSHIMA, Mitsugu Citation MEMOIRS OF THE FACULTY OF FISHERIES HOKKAIDO UNIVERSITY, 32(2), 131-243 Issue Date 1985-12 Doc URL http://hdl.handle.net/2115/21878 Type bulletin (article) File Information 32(2)_P131-243.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP 1985J TOYOSHIMA : Taxonomy of Lycodinae TAXONOMY OF THE SUBFAMilY lYCODINAE (FAMilY ZOARCIDAE) IN JAPAN AND ADJACENT WATERS BY Mitsugu TOYOSHIMA * Laboratory of Marine Zoology, Faculty of Fisheries, Hokkaido University, Hakodate, Hokkaido 041, Japan Contents page I. Introduction ............................................................ 133 II. Materials and methods. .. 134 III. Acknowledgments........................................................ 135 IV. Taxonomy of the subfamily Lycodinae .................................... 136 1. Key to genera of Lycodinae . .. 136 2. Genus Petroschmidtia . .. 137 3. Key to species of Petroschmidtia . .. 138 Petroschmidtia toyamensis Katayama .................................. 138 Petroschmidtia albonotata Taranetz et Andriashev ...................... 140 4. Genus H adropogonichthys Fedorov ......................... :.......... 142 Hadropogonichthys lindbergi Fedorov .................................. 142 5. Genus Lyconema Gilbert ............................................ 142 6. Genus Lycodonus Goode et Bean ...................................... 143 Lycodonus dorsoscutatus Oshima ...................................... 143 7. Genus Maynea Cunningham .......................................... 143 8. Genus Pachycara Zugmayer .......................................... 144 9. Genus Derepodichthys Gilbert ........................................ 144 10. Genus Taranetzella Andriashev ...................................... 144 Taranetzella lycoderma Andriashev .................................. " 145 11. Genus Lycenchelys Gill .............................................. 145 12. Key to species of Lycenchelys ........................................ 148 Lycenchelys maculatus sp. nov. ................................... " 149 Lycenchelys microporus Andriashev . .. 152 Lycenchelys volki Andriashev ........................................ 152 Lycenchelys roseus sp. nov. .......................................... 152 Lycenchelys albeolus Andriashev ...................................... 154 Lycenchelys pliciferus Andriashev ... .. 155 Lycenchelys uschakovi Andriashev .................................... 155 Lycenchelys bersteini Andriashev ...................................... 155 The present work is a portion of a dissertation that was submitted as a partial fulfillment of the requirements for Doctor's degree in Fisheries Science at Hokkaido University in 1983. * Present address: Takugin Research Institute, Doshin Bldg., Nishi 3-6, Odori, Chuo-ku, Sapporo 060, Japan. -131- Mem. Fac. Fish. Hokkaido Univ. [XXXII, 2 Lycenchelys squamosus Toyoshima .................................... 156 Lycenchelys altus sp. nov. ............................................ 158 Lycenchelys albomaculatus Toyoshima ................................ 159 Lycenchelys vitiazi Andriashev ........................................ 164 Lycenchelys camchaticus (Gilbert et Burke) ............................ 164 Lycenchelys ratmanovi Andriashev .................................... 166 Lycenchelys longirostris sp. nov. ..................................... 166 Lycenchelys hippopotamus Schmidt ......................... , .......... 169 Lycenchelys melanostomias Toyoshima ................................ 170 Lycenchelys rassi Andriashev ........................................ 173 Lycenchelys brevimaxillaris sp. nov. ................................. 174 13. Genus Embryx Jordan et Gilbert ...................................... 176 Embryx crotalinus (Gilbert) .......................................... 178 14. Genus Lycodes Reinhardt ............................................ 180 15. Key to five groups of Lycodes ........................................ 185 16. Key to species of group A . .. 185 17. Key to species of group B . .. 185 18. Key to species of group C . .. 185 19. Key to species of group D. .. 186 20. Key to species of group E . .. 187 Lycodes teraoi Katayama ............................................ 188 Lycodes sadoensis Toyoshima et Honma . .. 189 Lycodes diapteroides Taranetz et Andriashev ............................ 191 Lycodes japonicus Matsubara et Iwai .................................. 191 Lycodes tanakai Jordan et Thompson .................................. 193 Lycodes knipowitschi Popov .......................................... 195 Lycodes paucilepidotus sp. nov. ... .. 196 Lycodes raridens Taranetz et Andriashev .............................. 198 Lycodes mucosus Richardson. .. 200 Lycodes turneri Bean ................................................ 202 Lycodes heinemani Soldatov . .. 202 Lycodes bathybius Schmidt .......................................... 203 Lycodes sigmatoides Lindberg et Krasyukova .......................... 204 Lycodes macrolepis Taranetz et Andriashev ............................ 204 Lycodes semenovi Popov. .. 205 Lycodes uschakovi Popov ............................................ 205 Lycodes macrochir Schmidt .......................................... 205 Lycodes soldatovi Taranetz et Andriashev .............................. 207 Lycodes yamatoi sp. nov. ............................................ 208 Lycodes ygreknotatus Schmidt ........................................ 210 Lycodes microlepidotus Schmidt ...................................... 210 Lycodes ochotensis Schmidt .......................................... 211 Lycodes jenseni Taranetz et Andriashev ................................ 211 Lycodes palearis Gilbert .............................................. 212 Lycodes fasciatus (Schmidt) .......................................... 214 Lycodes schmidti Grazianov .......................................... 214 Lycodes multiJasciatus Schmidt ...................................... 214 Lycodes fulvus sp. nov ............................................... 216 Lycodes matsubarai sp. nov ........................................... 218 Lycodes caudimaculatus Matsubara .................................... 220 Lycodes microporus Toyoshima ........................................ 221 Lycodes ocellatus sp. nov. ............................................ 223 -132- 1985J TOYOSHIMA: Taxonomy of Lycodinae Lycodes nakamurai Tanaka .......................................... 224 Lycodes hubbsi Matsubara. .. 226 Lycodes pectoralis sp. nov. ......................................... 228 Lycodes diapterus Gilbert ............................................ 230 Lycodes albolineatus Andriashev ...................................... 232 Lycodes brevipes Bean .............................................. 233 Lycodes brunneofasciatus Suvorov .................................... 234 Lycodes andriashevi Fedorov . .. 235 Lycodes obscurus sp. nov. ............................................ 235 21. Genus Aprodon Gilbert .............................................. 237 Aprodon cortezianus Gilbert .......................................... 237 22. Genus Lycodopsis Collett ............................................ 238 V. Summary .. .. 238 VI. Literature cited. .. 239 I. Introduction The family Zoarcidae is comprised of three subfamilies, Zoarcinae (including Hadropareinae and Gymnelinae), Lycodinae, and Bothrocarinae (including Lyco­ gramminae, Melanostigmatinae and Lycodapodinae). The Lycodinae is the largest subfamily of zoarcids, and known predominantly from the temperate to polar seas of the Hemisphere. The fishes of the subfamily are bottom dwellers usually captured from the continental shelves and slopes. The taxonomic study of the lycodine fishes from the waters north of Okhotsk Sea has been carried out mainly by the Russian and American ichthyologists (Collett, 1880; Jordan and Evermann, 1898; Jensen, 1904; Knipowitschi, 1906; Taranetz, 1937; Schmidt, 1950; Andriashev, 1954, 1958, 1959). However, their studies were not always satisfactory because they examined small samples with limited geographic representation and because they established many subspecies based on modest differences. On the other hand, the Japanese species had been studied mainly by American ichthyologists (Jordan and Fowler, 1902; Jordan and Hubbs, 1925, etc.). On the bases of these studies, Okada and Matsubara (1938), without using subfamilies, grouped the 24 known Japanese species, including the Okhotsk species described by the Russian ichthyologists (Schmidt, 1904; Soldatov, 1917; Soldatov and Lind­ berg, 1930, etc.), in 13 genera. Matsubara (1955), also without adopting subfamilies, presented an analytical key to 48 species of 17 genera hitherto known from the Sea of Japan, Okhotsk Sea and the Pacific coast of Japan. However, his key can not be easily used for identification of Japanese zoarcids because they were mixed with many Russian species that he was unable to study. These circumstances have resulted in considerable confusion in the taxonomy of the Lycodinae as well as in other subfamilies of Zoarcidae. It is the principal aim of this paper to resolve these problems and clarify the relationships amongst species and genera. This paper discusses alllycodine genera from the Northern Hemisphere and all species known from the Sea of Japan, Okhotsk Sea, western Pacific Ocean and Bering Sea. -133-