Title INTERRELATIONSHIPS OF THE FAMILY PLEURONECTIDAE (PISCES: PLEURONECTIFORMES) Author(s) SAKAMOTO, Kazuo Citation MEMOIRS OF THE FACULTY OF FISHERIES HOKKAIDO UNIVERSITY, 31(1-2), 95-215 Issue Date 1984-12 Doc URL http://hdl.handle.net/2115/21876 Type bulletin (article) File Information 31(1_2)_P95-215.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP INTERRELATIONSHIPS OF THE FAMILY PLEURONECTIDAE (PISCES: PLEURONECTIFORMES) By Kazuo SAKAMOTO * Laboratory of Marine Zoology, Faculty of Fisheries, Hokkaido University, Hakodate, Japan Contents Page I. Introduction···································································· 95 II. Acknowledgments· ................... ·.·......................................... 96 III. Materials········································································ 97 IV. Methods·····················.··· ... ····················· .......... ············· 102 V. Systematic methodology· ......................................................... 102 1. Application of numerical phenetics .............................................. 102 2. Procedures in the present study ................................................. 104 VI. Comparative morphology ........................................................ 108 1. Jaw apparatus ................................................................ 109 2. Cranium······································································ 111 3. Orbital bones . .. 137 4. Suspensorium and opercular apparatus ........................................... 140 5. Hyoid arch· ................................................................... 144 6. Branchial apparatus and gill rakers·· ............................................ 148 7. Fins·········································································· 155 8. Pectoral girdle ................................................................ 164 9. Pelvic girdle .................................................................. 169 10. Urohyal ...................................................................... 171 11. Vertebrae and their accessory bones ... .. 173 12. Caudal skeleton and fin ........................................................ 181 13. Others····················· ....... ················· ... · .......... ············· 194 VII. Interrelationships of the fishes of the Pleuronectidae ................................ 198 VIII. Summary ...................................................................... 212 IX. Literature cited ................................................................. 213 I. Introduction The fishes of the family Pleuronectidae, inhabiting the sand-muddy bottom between shallow inshore waters and continental slope, are widely distributed in the The present work was submitted as a partial fulfillment of the requirements for Doctor's degree in Fisheries Science at Hokkaido University in 1984 . • Present address: 3546 Futami, Hohoku-cho, Toyoura-gun, Yamaguchi Prefecture 759-55, Japan. - 95- Mem. Fac. Fish. Hokkaido Univ. [XXXI, 1/2 world. This family is characterized in having both eyes on the right side of the body, the optic nerve of the left eye always dorsal, the preopercle with a free margin and the fin rays without spines. The Pleuronectidae includes 114 species. While most are marine, some of them enter such fresh waters as rivers, lakes and marshes. The phylogeny of the pleuronectid fishes has been studied by several investi­ gators (Regan, 1910; Norman, 1934; Kuronuma, 1938; Hubbs, 1945; Kim, 1973; Li,1981). These authors, except Kim (1973), discussed the relation only among the subfamilies on the basis of several characters including osteology. Kim (1973) studied the interrelationships of 14 species of the Pleuronectinae (sensu Norman, 1934) based on the comparative osteology of the cranium, the urohyal, the vertebrae and the caudal skeleton. Though there are many studies on both internal and external morphology in the present family owing to their morphological peculiarities (e.g. Kyle, 1921; Hikita, 1934; Chabanaud, 1936; Yazudani, 1969; Groot, 1971 etc.), most of those studies were fragmentary and did not treat the interrelationships of the pleuronectids. Under the circumstances, it is necessary to clarify the interrelationships of the pleuronectid fishes on the basis of as many internal and external characters as possible. In order to analyse the interrelationships of the species of the Pleu­ ronectidae, numerical phenetics (Sneath and Sokal, 1973) was adopted as a system­ atic methodology. The classification of righteye flounders has been studied by many ichthyologists since the end of the nineteenth century. In 1910, Regan treated the righteye flounders as a single family Pleuronectidae for the first time. Since then, the classification of this group has been discussed mainly at a sub familial level based on several internal and external characters (Regan, 1929; Norman, 1934; Berg, 1940; Hubbs, 1945 etc.), though some or all the subfamilies were sometimes raised to familial rank (Regan, 1920; Jordan, 1923 etc.). Nelson (1976) divided the Pleu­ ronectidae into four subfamilies and the Pleuronectinae into two tribes. In the present study, all the pleuronectid species are classified into the supraspecific taxa which are newly defined by the ranking on the basis of their phenetic interrelationships. II. Acknowledgments I wish to express my sincere gratitude to the late Dr. Takao Igarashi, the former professor of Hokkaido University, for his guidance in the course of the present study and critical reading of the manuscript. Also, I wish to thank Dr. Keikichi Hamada, professor of the same university, for critical reading of the manuscript. I wish to express my thanks to Dr. Seikichi Mishima, professor of the same university, for critical reading of the manuscript. I wish to express the deepest appreciation to Dr. Kunio Amaoka, professor of the same university, for his valuable advice and critical reading of the manuscript. I wish to thank for their help in the present study to the following persons: Mr. Koji Abe of Hokkaido Fisheries Experimental Station at Wakkanai; Dr. Gerald R. Allen of Western Australian Museum; Dr. Hirotoshi Asano of Kinki University; Dr. A.J. Bass of National Museum of New Zealand; Dr. William N. Eschmeyer of - 96- 1984J SAKAMOTO: Interrelationships of Pleuronectidae California Academy of Science; Dr. Martin Gomon of National Museum of Victoria; Dr. S.J. de Groot of Ministerie van Landbouw en Visserij; Dr. Karsten E. Hartel of Museum of Comparative Zoology; Dr. Hiroshi Hatanaka of Far Seas Fisheries Research Laboratory; Mr. Masashi Higashi of Noshappu Kanryu Aquarium; Dr. P. Alexander Hulley of South African Museum; Drs. Tamotsu Iwai, Izumi Nakamura and Tetsuji Nakabo of Kyoto University; Dr. Gavin James of Ministry of Agricul­ ture and Fisheries, New Zealand; Mr. Shinichi Kanamaru of Hokkaido Regional Fisheries Research Laboratory; Dr. Yong Uk Kim of Pusan Fisheries College; Mr. Daiji Kitagawa of Iwate Fisheries Experimental Station; Dr. Grace Klein-Mac­ Phee; Mr. Shinji Kudo of Tokai Regional Fisheries Research Laboratory; Dr. Katsuzo Kuronuma; Dr. Kenichiro Kyushin and Mr. Akihisa Iwata of the same university; Dr. Peter Last of Tasmanian Fisheries Development Authority; Dr .. Robert J. Lavenberg of Los Angels County Museum; Mr. Toshihiro Mizushima of Hokkaido Fisheries Experimental Station at Kushiro; Mr. Syuka Maruyama of Hokkaido Fisheries Experimental Station at Hakodate; Dr. Supap Monkolprasit of Kasetsart University; Dr. J~rgen Nielsen of Universiteteits Zoologiske Museum; Dr. Han Nijssen of Zoi:ilogisch Museum, Universiteit van Amsterdam; Mr. Masatoshi Nozawa of Tottori Fisheries Experimental Station; Dr. Osamu Okamura of Kochi University; Mr. Teiji Ono of Onahama Fishermen's Cooperative Association; Mr. Koichi Ouchi of Esashi Enyo Co. Ltd.; Drs. John R. Paxton and Barry C. Russell of Australian Museum; Prof. Margaret M. Smith of J.L.B. Smith Institute of Ichthyology; Mr. Umeji Suzuki; Mr. Tetsuo Takagoshi of Fukushima Fisheries Experimental Station; Drs. Toru Takai, Shyohei Nishikawa and Osame Tabeta of Shimonoseki University of Fisheries; Dr. Yoshiaki Tominaga of the University of Tokyo; Dr. Stanley H. Weitzman and Ms. Susan L. Jewett of U.S. National Museum; Dr. Mark White of Ministry of Agriculture, Fisheries and Food, England; Dr. Peter J.P. Whitehead and Mr. Alwyne Wheeler of British Museum (Natural History); Mr. Tamotsu Yamamura of Notsuke Fishermen's Cooperative Associa­ tion; Mr. Takuji Yatou. Dr. Don E. McAllister of National Museum of Canada kindly reviewed the manuscript. Mr. Nobuyoshi Kimura kindheartedly advised me on my English. I express my sincere thanks to Mr. Yuichi Kobayashi who kindly prepared a computer program for the present study. Also, I thank Dr. Yurin Sakamoto of the same university for his advice on FORTRAN program. Finally, my special thanks go to Dr. Kazuhiro Nakaya, associate professor of the same university, and the graduate students of our laboratory who have helped me in varIOUS ways. The present work was supported in part by grants from the Ito Foundation for the Advancement of Ichthyology. III. Materials The specimens examined for the present study the listed with their catalogued numbers, the numbers of the specimens and the sizes. The abbreviations prefixed to the catalogued numbers indicate the following institutions where those specimens belong: - 97- Mem. Fac. Fish. Hokkaido Univ. [XXXI, 1/2 BMNH: British Museum (Natural