Title Phylogeny of the Order Gadiformes (Teleostei, Paracanthopterygii) Author(s) ENDO, Hiromitsu Citation MEMOIRS OF THE GRADUATE SCHOOL OF FISHERIES SCIENCES, HOKKAIDO UNIVERSITY, 49(2), 75-149 Issue Date 2002-12 Doc URL http://hdl.handle.net/2115/22016 Type bulletin (article) File Information 49(2)_P75-149.pdf Instructions for use Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP Mem. Grad. Sch. Fish. Sci. Hokkaido Univ. Phylogeny of the Order Gadiformes (Teleostei, Paracanthopterygii) Hiromitsu END0 1)2) Contents I. Introduction .............................................................................................................................. ·········76 II. Materials and methods .................................................................................................................. ······77 III. Monophyly of Gadiformes ..................................................................................................................... 78 IV. Relatives of Gadiformes ............................................................................................................ ············82 V. Interrelationships of lower gadiforms ..................................................................................................... ·83 1. Characters used in the first analysis ............................................................................................ ·83 2. Relationships·········· ................................................................................................................ ·95 VI. Intrarelationships of higher gadoids ........................................................................................................ ·97 1. Characters used in the second analysis ....................................................................................... ···97 2. Relationships ....................................................................................................................... ·107 VII. Cladistic classification of Gadiformes ...................................................................................................... 110 VIII. Comparison with previous works '" ................................................................................................... ···112 I. Suborders .......................................................................................................................... ····112 2. Family Melanonidae ............................................................................................................ ···115 3. Family Steindachneriidae ...................................................... ···················································116 4. Family Macrouridae ............................................................................................................... 116 5. Family Euclichthyidae ............................................................................................................ 116 6. Family Moridae ............................................................................................................ ·········117 7. Family Macruronidae ............................................................................................................ ···117 8. Family Merlucciidae ............................................................................................................ ···117 9. Family Ranicipitidae ............................................................................................................ ···118 10. Family Bregmacerotidae ............................................................................................................ 118 11. Family Muraenolepididae ........................................................................................................... ·118 12. Family Gadidae ............................................................................................................ ············118 IX. General considerations ............................................................................................................ ············120 1. Character evolution of supernumerary fin-rays ........................................................................... ···120 1-1. Character states .............................................................................................................. ·122 1-2. Order and polarity ............................................................................................................ 123 1-3. Dorsal supernumerary fin-rays ...................................................... ·······································124 2. Biogeography ..................................................................................................................... ···126 2-1. Phylogenetic dispersal of Merlucciidae ................................................................................ ·127 2-2. Phylogenetic dispersal of Gadidae ...................................................... ·································127 2-3. Phylogenetic dispersal of Gadinae ...................................................................................... ·129 3. Phylogenetic position of extinct gadoids ...................................................................................... ·132 3-1. Genus t Rhinocephalus ..................................................................................................... ·132 3-2. Genus t Palaeogadus ........................................................................................................ ·134 3-3. Geological times of gadoid fossils ...................................................... ·································135 4. Heterochronic evolution ...................................................... ·········.·········································136 4-1. Paedomorphoses in Macrouridae .......................................................................................... 137 4-2. Progenesis in Bregrnacerotidae ............................................................................................ ·139 The present work was submitted as a partial fulfillment of the requirements for Doctor's degree in Fisheries Science at Hokkaido University in 1995. 1) Laboratory of Marine Biodiversity (Systematic Ichthyology), Graduate School of Fisheries Sciences, Hokkaido University, 3- 1-1 MinatD-cho, Hakodate, Hokakido 041-8611, Japan 2) Present address: Laboratory of Marine Biology, Faculty of Science, Kochi University, 2-5-1 Akebono-cho, Kochi 780-8520, Japan (e-mail: [email protected]) - 75 - Mem. Grad. Sch. Fish. Sci. Hokkaido Univ. 49(2), 2002. 4-3. Neoteny in Muraenolepididae ...................................................... ·······································140 4-4. Possible heterochrony in other higher gadoids ....................................................................... ·140 5. Conclusions ............................................................................................................ ············141 X. Acknowledgements··············································································································· ................. ·142 XI. Literature cited ............................................................................................................ ·····················144 XII. Appendix ............................................................................................................ ······························148 Abstract Intrarelationships of the order Gadiformes (sensu Nelson, 1994), focused on the higher gadoid taxa, were hypothesized cladistically on the basis of 49 morphological transformations. These indicated that gadiforms should be classified into three suborders and eleven families: Melanonidae (Melanonoidei), forming the sister group of all other gadiforms, Macrouridae and Steindachneriidae (Macrouroidei), and Euclichthyidae, Moridae, Macrur­ onidae, Merlucciidae, Ranicipitidae, Gadidae, Bregmacerotidae and Muraenolepididae (Gadoidei). In addition, four subfamilies were recognized in both Macrouridae (Bathygadinae, Macrourinae, Macrouroidinae and Trachyrin­ cinae) and Gadidae (Group I: Gaidropsarinae and Phycinae; Group II: Lotinae and Gadinae). Three clades were recognized within gadids: Gadiculini (Gadiculus), Gadini (Merlangius, Pollachius, Theragra, Gadus, Melanogrammus, Eleginus and Microgadus), and Trisopterini (Trisopterus, Micromesistius, Arctogadus and Boreogadus). The evolution of dorsal supernumerary fin-ray characters, gadid biogeography, phylogenetic pos­ itions of extinct gadoid genera and possible heterochrony among gadiforms are discussed. Key words: Phylogeny, Gadiformes, cladistics, character evolution, biogeography, fossils, heterochrony I. Introduction Howes (1990, 1991a) reached the following consensus: bregmacerotids and muraenolepidids belong to The order Gadiformes (sensu Nelson, 1994) comprises advanced gadoids, and each of the "Gadidae" and about 500 species included in 85 genera representing a "Merlucciidae" (sensu Nelson, 1984) are polyphyletic. number of morphologically divergent families: These conclusions also differed from the traditional view Ranicipitidae, Euclichthyidae, Macrouridae, Steindach­ that the Muraenolepididae was the most primitive family neriidae, Moridae, Melanonidae, Macruronidae, Breg­ among gadiforms (e.g., Svetovidov, 1948; Cohen, 1984). macerotidae, Muraenolepididae, Phycidae, Merlucciidae However, in spite of these cladistic studies, gadiform and