The Origin of Higher Clades Osteology, Myology, Phylogeny and Evolution of Bony Fishes and the Rise of Tetrapods MONITORING AND EVALUATION OF SOIL CONSERVATION AND WATERSHED DEVELOPMENT PROJECTS The Origin of Higher Clades Osteology, Myology, Phylogeny and Evolution of Bony Fishes and the Rise of Tetrapods Rui Diogo Department of Anthropology The George Washington University Washington, DC USA Science Publishers Enfield (NH) Jersey Plymouth SCIENCE PUBLISHERS An imprint of Edenbridge Ltd., British Isles Post Office Box 699 Enfield, New Hampshire 03748 United States of America Website: http://www.scipub.net sales@scipub.net (marketing department) editor@scipub.net (editorial department) info@scipub.net (for all other enquiries) Library of Congress Cataloging-in-Publication Data Diogo, Rui The origin of higher clades: osteology, myology, phylogeny, and evolution of bony fishes and the rise of tetrapods/Rui Diogo p cm Includes bibliographical references and index ISBN 978-1-57808-530-9 (Paperback) Osteichthyes Evolution I Title QL618.2D56 2007 597.13’8 dc22 2007028539 ISBN 978-1-57808-530-9 (Paperback) ISBN 978-1-57808-437-1 (Hardcover) © 2007, Copyright reserved Paperback edition 2008 All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior permission This book is sold subject to the condition that it shall not, by way of trade or otherwise, be lent, re-sold, hired out, or otherwise circulated without the publisher’s prior consent in any form of binding or cover other than that in which it is published and without a similar condition including this condition being imposed on the subsequent purchaser Published by Science Publishers, Enfield, NH, USA An imprint of Edenbridge Ltd Printed in India Preface The Osteichthyes, including bony fishes and tetrapods, is a highly speciose group of animals, comprising more than 42000 living species The extraordinary taxonomic diversity of osteichthyans is associated with a remarkable variety of morphological features and adaptations to very different habitats, from the deep-sea to high mountains Osteichthyans therefore provide a very interesting case study to analyze the origin and morphological macroevolution of higher-clades In this book, I provide a new insight on the osteology, myology, phylogeny and evolution of this fascinating group, which is based on my own research and on a survey of the literature Chapters and provide a short introduction to the main aims of the book and to the methodology and methods used Chapter deals with an extensive cladistic analysis of osteichthyan higher-level interrelationships based on a phylogenetic comparison of 356 characters in 80 extant and fossil terminal taxa representing all major groups of Osteichthyes This cladistic analysis includes various terminal taxa and osteological characters, and namely a large number of myological characters, not included in previous analyses Chapter provides a general discussion on issues such as the comparative anatomy, homologies and evolution of osteichthyan cranial and pectoral muscles, the development of zebrafish cephalic muscles and the implications for evolutionary developmental studies, the origin, homologies and evolution of one of the most peculiar and enigmatic structural complexes of osteichthyans, the Weberian apparatus, and the use of myological versus osteological characters in phylogenetic reconstructions I hope that this work may stimulate, and pave the way for, future studies on the comparative anatomy, functional morphology, phylogeny and evolution of osteichthyans and of vertebrates in general, which, as stressed throughout the book, should ideally take into account the precious information obtained from the study of muscular features Dedicated to MICHEL CHARDON, to his outstanding knowledge, to his friendship, and to his humbleness MONITORING AND EVALUATION OF SOIL CONSERVATION AND WATERSHED DEVELOPMENT PROJECTS LEE Acknowledgements First of all, I want to thank P Vandewalle and M Chardon for accepting me in the Laboratory of Functional and Evolutionary Morphology in 1998 and thus for giving me the opportunity to begin my research on the anatomy, functional morphology, phylogeny and evolution of vertebrates and of bony fishes in particular I also want to thank E Parmentier His persistence, the remarkable ability that he has to solve all types of challenges, and the courage he has to get deeply involved in different scientific areas were really inspiring for me I am also thankful to R Vari, as well as his colleagues S Weitzman, J Williams and S Jewett from the National Museum of Natural History, for accepting me in that amazing museum during two academic years and for providing numerous specimens analyzed in this work I also want to thank I Doadrio, who received me in the Museo Nacional de Ciencias Naturales de Madrid, and has made available many specimens of the vast fish collection of this museum, which is mainly the fruit of his hard work Another bright scientist who received me in his lab for several months was S Hughes, whom I thank very, very much In his lab, at the prestigious MRC Centre for Developmental Neurobiology of the King’s College of London, I took my first steps in Evolutionary Developmental Biology (“Evo-Devo”) I enjoyed much his availability, his interest, and his continuous questioning and curiosity I also want to take this occasion to thank B Wood for inviting me to continue my research at the Anthropology Department of the George Washington University, where I shall have the opportunity to expand my work to other osteichthyan groups, and particularly to primates A special thanks to the late G Teugels, as well to J Snoeks and E Vreven (Musée Royal de l’Afrique Centrale), P Laleyé (Université Nationale du Bénin), Z Peng and S He (Academy of Sciences of China at Wuhan), T Grande (Field Museum of Natural History), D Catania (California Academy of Sciences), M Stiassny (American Museum of Natural History), M Sabaj LEEE and J Lundberg (Academy of Natural Sciences of Philadelphia), W Fink, D Nelson and H Ng (Museum of Zoology, University of Michigan), R Bills and P Skelton (South African Institute for Aquatic Biodiversity), L Page and M Retzer (Illinois Natural History Survey), P Pruvost and G Duhamel (Museum National d’Histoire Naturelle) and R Walsh and F Slaby (George Washington University) for kindly providing a large part of the specimens analyzed I also want to acknowledge T Abreu, A Zanata, F Meunier, D Adriaens, F Wagemans, M de Pinna, P Skelton, F Poyato-Ariza, T Grande, H Gebhardt, M Ebach, A Wyss, J Waters, G Cuny, L Cavin, F Santini, J Briggs, L Gahagan, M Gayet, J Alves-Gomes, G Lecointre, L Soares-Porto, P Bockmann, B Hall, F Galis, T Roberts, G Arratia, L Taverne, E Trajano, C Ferraris, M Brito, R Reis, R Winterbottom, C Borden, B Richmond and many other colleagues for their helpful advice and assistance and for their discussions on osteichthyan anatomy, functional morphology, phylogeny and/or evolution in the last years A special thanks to V Abdala, with whom I have discussed many of the parts of this work, and with whom I hope to undertake the numerous projects we have in mind concerning vertebrate musculature, as well as to J Joss (Macquarie University) and A Gosztonyi (Centro Nacional Patagónico) for providing me the large dipnoan specimens analyzed, and to J Fernández and other people from the Museo Nacional de Ciencias Naturales de Madrid for providing me the nice salamander and lizard specimens examined My special thanks to all my friends, particularly to Pedro Brito, Claudia Oliveira, Henry Evrard and Diego Alarcon Reina Thank you very much, Alejandrita Pelito Lindo, and thanks to our amazing and adorable Tots Pelluda Very special thanks to my parents, Valter and Fatima, to my brothers, Hugo and Luis, and to my late grandfathers Raul and Amélia Thank you very much for the confidence in my work and for the close cooperation in the several projects we have together Finally, thanks to all those who have been involved in administering the various grants and other awards that I received during the last years, without which this work would really not have been possible EN List of Abbreviations* II, III, IV, V, VII, IIX, IX, X A0, A1, A1-OST A1-OST-L, A1-OST-M A2 A2-D, A2-PVM, A2-V A3', A3'’ AB-PRO AB-SUP abs AC AD-AP AD-HYO AD-OP AD-PRO AD-SUP ADM AED1 AHL, AHM am am-m AME ana ang foramens/nerves of Miles’s 1977 original drawing adductor mandibulae A0, A1 and A1-OST lateral and mesial sections of adductor mandibulae A1-OST adductor mandibulae A2 dorsal, posteroventromesial and ventral sections of adductor mandibulae adductor mandibulae A3' and A3'’ abductor profundus abductor superficialis anterior bulla of swimbladder anconaeus coracoideus adductor arcus palatini adductor hyomandibulae adductor operculi adductor profundus adductor superficialis adductor mandibulae abductor et extensor digiti I anconaeus humeralis lateralis and medialis ampulla macula of ampulla adductor mandibulae externus anterior neural arch angular *Myological structures are shown in bold !$ Weber EH 1820 De Aure et Auditu Hominis et Animalium: Pars I—De Aure Animalium aquatilium Gehard Fleischer, Leipzig Weisel GF 1960 The osteocranium of the catostomid fish, Catostomus macrocheilus—a study in adaptation and natural relationship J Morphol 106:109-129 Weiss B, Strother W, Hartig G 1969 Auditory sensitivity in the Bullhead Catfish (Ictalurus nebulosus) Proc Nat Acad Sci US 64:552-556 Weitzman SH 1962 The osteology of Brycon meeki, a generalized characid fish, with an osteological definition of the family Stanford Ichthyol Bull 8:1-77 Weitzman SH 1964 Osteology of and relationships of South American characid fishes of subfamilies Lebiasininae and Erythrinae with special reference to subtribe Nannostomina Proc US Natl Mus 116:127-170 Weitzman SH 1967a The origin of stomiatoid fishes with comment on the classification of salmoniform fishes Copeia 1967:507-540 Weitzman SH 1967b The osteology and relationships of the Astronesthidae, a family of oceanic fishes Dana-Rep (Copenhagen) 71:154 West-Eberhard MJ 2003 Developmental Plasticity and Evolution Oxford University Press, New York Westneat MW, Thorsen DH, Walker JA, Hale ME 2004 Structure, function, and neural control of pectoral fins in fishes IEEE J Oceanic Engen 29:674683 Wiley EO 1976 The phylogeny and biogeography of fossil and recent gars (Actinopterygii: Lepisosteidae) Univ Kansas Misc Publ (Nat Hist) 64:1111 Wiley EO 1979a Ventral gill arch muscles and the interrelationships of gnathostomes, with a new classification of the Vertebrata J Linn Soc Lond (Zool) 67:149-179 Wiley EO 1979b Ventral gill arch muscles and the phylogenetic interrelationships of Latimeria Occ Pap Calif Acad Sci 134:56-67 Wilga CD, Wainwright PC, Motta PJ 2000 Evolution of jaw depression mechanisms in aquatic vertebrates: insights from Chondrichthyes Biol J Linn Soc Lond 71:165-185 Wilson MVH, Veilleux P 1982 Comparative osteology and relationships of the Umbridae (Pisces: Salmoniformes) Zool J Linn Soc Lond 76:321-325 Winterbottom R 1974 A descriptive synonymy of the striated muscles of the Teleostei Proc Acad Nat Sci (Phil) 125:225-317 Winterbottom R 1993 Myological evidence for the phylogeny of recent genera of surgeonfishes (Percomorpha, Acanthuridae), with comments on Acanthuroidei Copeia 1993:21-39 !$ Wu K-Y, Shen S-C 2004 Review of the teleostean adductor mandibulae and its significance to the systematic position of the Polymixiiformes, Lampridiformes, and Triacanthoidei Zool Stud 43:712-736 Zaragüeta-Bagils R, Lavoué S, Tillier A, Bonillo C, Lecointre G 2002 Assessment of otocephalan and protacanthopterygian concepts in the light of multiple molecular phylogenies C R Biologies 325:1191-1207 Zardoya R, Meyer A 1996 Evolutionary relationships of the coelacanth, lungfishes, and tetrapods based on the 28S ribosomal RNA gene Proc Natl Acad Sci US 93:5449-5454 Zardoya R, Cao Y, Hasegawa M, Meyer A 1998 Searching for the closest living relative(s) of tetrapods through evolutionary analysis of mitochondrial and nuclear data Mol Biol Evol 15:506-517 Zhu M, Schultze H-P 1997 The oldest sarcopterygian fish Lethaia 30:293304 Zhu M, Schultze H-P 2001 Interrelationships of basal osteichthyans In: Ahlberg PE (ed.) Major Events in Early Vertebrate Evolution, Paleontology, Phylogeny, Genetics and Development Taylor and Francis, London, pp 289-314 Index Abductor brevis pollicis, 284 Abductor digitorum V, 284 Abductor superficialis, 148 Acipenser, 12, 17, 30, 33, 36, 38, 39, 40, 122, 148, 156, 157, 158, 159, 167, 169, 172, 173, 176, 182, 187, 189, 191, 200, 207, 215, 216, 217, 254 Acipenseridae, 3, 12, 33, 39 Acipenseriformes, 33, 34, 38, 322 Actinistia, 3, 6, 7, 12, 32, 108, 113, 114, 115, 117, 226, 236, 238, 247, 255, 260 Adductor arcus palatini, 187 Adductor mandibulae, 181, 184 Adductor operculi, 184, 245 Albula, 13, 17, 52, 60, 67, 143, 174, 176, 182, 185, 186, 195, 204, 215, 238 Albuliformes, 13, 51 Alepocephaloidea, 3, 13, 68, 73, 78, 79, 225, 226 Alepocephalus, 13, 18, 68, 69, 70, 71, 73, 74, 156, 157, 191, 203 Ambystoma, 12, 13, 15, 18, 29, 30, 32, 113, 114, 121, 125, 131, 133, 145, 150, 153, 154, 167, 168, 172, 173, 174, 175, 179, 180, 187, 188, 198, 200, 203, 207, 216, 218, 227, 251, 254, 257, 258, 281, 282, 283, 285, 286 Amia, 5, 12, 17, 32, 41, 43, 44, 45, 46, 50, 60, 61, 115, 146, 159, 169, 175, 176, 178, 184, 189, 195, 225, 226, 238, 239, 248, 249, 253 Amiidae, 3, 5, 45 Amiiformes, 45 Amniota, 131, 137, 233, 243, 257, 275, 281, 287 Amniotes, 227, 276 Amphibia, 131, 132, 227, 233, 243, 247, 255, 257, 275, 276, 281, 287 Anatophysi, 84, 100 Anconaeus, 281, 282 Anguilla, 13, 17, 54, 59, 60, 67, 168, 196, 202 Anguilliformes, 13, 51, 58, 59, 60, 61, 66, 322 Argentina, 13, 16, 18, 73, 157, 220 Argentinoidea, 3, 13, 68, 73, 226 Arrector 3, 278 Arrector dorsalis, 148, 278 Arrector ventralis, 149, 278 Articular, 217 Astronesthes, 13, 18, 74, 75, 76, 161, 167, 193, 194, 203, 323 Aulopiformes, 13, 75, 193, 212, 322 Aulopus, 13, 18, 68, 72, 74, 75, 76, 186 Autopalatines, 199 Autosphenotics, 159 Autostylic suspension, 197 Bagrus, 13, 17, 113 Barbus, 13, 17, 104, 109, 165 Basioccipital, 161 !$" Basisphenoid, 158 Bathylagus, 13, 18, 73, 205, 211 Brachyhypopomus, 13, 17, 111, 173, 192, 211 Branchiohyoideus, 243 Branchiostegal rays, 214 Brycon, 13, 17, 101, 105, 106, 110, 157 Callichthys, 13, 17, 44, 106, 112, 214 Catostomus, 13, 17, 109, 169, 211 Cervicomandibularis, 244, 247 Cetopsis, 13, 17, 112, 159, 181, 219 Chanos, 13, 17, 85, 86, 87, 88, 89, 90, 91, 92, 101, 160, 163, 211 Characiformes, 13, 84, 102, 109, 110, 288, 297, 322 Chlorophthalmus, 13, 18, 75, 76, 177, 186 Choanates, 117, 118 Chondrostei, 3, 6, 7, 12, 32, 33, 34, 36, 38, 40, 226, 227, 230, 238, 241, 256, 276, 278 Chrysichthys, 13, 17, 113 Citharinus, 13, 17, 110, 162 Cladistia, 3, 6, 7, 12, 31, 33, 226, 227, 235, 246, 259, 276 Claustrum, 294 Cleithrum, 171 Clupeocephala, 51, 66, 279, 318 Clupeoidea, 13, 79, 80, 81 Clupeoidei, 79 Clupeomorpha, 3, 73, 78, 79, 80, 288, 302 Cobitis, 13, 17, 109, 169, 186, 203 Cobitoidea, 109 Coelacanthiformes, 115 Conger, 13, 17, 54, 59, 60, 168, 196, 202 Contrahentes digitorum, 284 Contrahentium caput longum, 283 Coracobrachialis longus, 281 Coregonus, 13, 18, 74, 75, 161, 194, 215 Coronoid bones, 216 Coronomeckelian bone, 220 Cromeria, 13, 17, 93, 94, 96, 164, 184, 192 Cromeriini, 94 Ctenosquamata, 75 Cypriniformes, 13, 84, 108, 265, 266, 288, 290, 297, 322 Cyprinoidea, 109 Danio, 13, 17, 84, 102, 103, 104, 109, 165, 169, 219, 238, 248, 265, 266, 287 Denticeps, 13, 17, 67, 79, 80, 81, 82, 83, 189, 191, 196, 203, 208, 215, 302 Denticipitoidei, 13, 79 Depressor mandibulae, 244 Dilatator operculi, 184, 189 Diplomystes, 13, 17, 99, 112 Dipnoi, 3, 6, 12, 113, 117, 122, 123, 126, 226, 227, 233, 236, 238, 243, 247, 255, 257, 276, 280, 281, 321 Distichodus, 13, 17, 107, 110, 185, 203, 208 Dorsalis scapulae, 282 Dorsometacarpalis, 284 Elopiformes, 13, 51, 322, 323 Elopomorpha, 3, 5, 51, 59, 60, 222, 225, 318, 322 Elops, 13, 17, 32, 52, 53, 54, 55, 56, 57, 58, 60, 143, 174, 176, 208, 213, 217, 218 Engraulis, 13, 17, 69, 79, 80, 83, 155 Engrauloidea, 13, 79, 80, 81 Entopterygoid, 200, 202 Esociformes, 3, 13, 68, 74, 76, 226, 322 Esox, 13, 18, 77, 157, 158, 161, 208 Ethmalosa, 13, 17, 79, 80, 84, 155 Eurypharynx, 13, 17, 30, 44, 54, 59, 60, 118, 119, 144, 147, 159, 180, 191, 196, 321 Eurypterygii, 75, 212 Euteleostei, 3, 68, 69, 225, 318 Exoccipitals, 160 Extensor digitorum communis, 282 Extensor lateralis digiti IV, 284 Extensores digitorum breves, 284 !$# Flexor accessorius lateralis, 283 Flexor accessorius medialis, 283 Flexor digitorum communis, 282 Flexor digitorum V transversus, 284 Flexores breves profundi, 284 Flexores breves superficiales, 283 Galaxias, 13, 18, 77, 148, 172, 185, 204, 208 Galaxioidea, 3, 13, 68, 77, 221, 226 Ginglymodi, 3, 5, 6, 7, 12, 32, 39, 40, 41, 43, 44, 45, 225, 226, 227, 230, 238, 239, 241, 256, 276, 278 Gonorynchidae, 85 Gonorynchiformes, 13, 84, 85, 95, 288, 322, 323 Gonorynchus, 13, 17, 67, 85, 90, 91, 92, 145, 159, 165, 195, 196, 197, 200, 204, 208, 211, 215, 220, 305 Grasseichthys, 13, 17, 93, 94, 96, 164, 168, 192 Gymnotiformes, 13, 84, 109, 110, 111, 288, 297, 322 Gymnotus, 13, 17, 111, 112, 173, 192, 211 Halecomorphi, 3, 5, 6, 7, 12, 32, 39, 40, 41, 43, 44, 45, 46, 60, 225, 226, 227, 230, 239, 241, 256, 276, 278 Hiodon, 13, 17, 52, 61, 62, 63, 64, 156, 170, 171, 174, 200, 218 Hiodontidae, 13, 61, 303 Hyohyoideus, 146 Hyohyoideus superior, 146 Ilisha, 13, 17, 79, 80, 83, 305, 307 Intercalarium, 291 Interhyal, 211 Interhyoideus, 144 Intermandibularis, 144, 230 Intermetacarpales, 284 Interopercle, 209 Intracranial joint, 115, 162 Kneria, 13, 17, 93, 94, 164, 192, 204 Kneriidae, 85 Kneriini, 94 Lateral ethmoid, 155, 159 Latimeria, 8, 12, 13, 18, 40, 44, 46, 108, 113, 114, 115, 117, 118, 122, 162, 168, 173, 174, 175, 179, 180, 185, 188, 189, 190, 205, 207, 213, 214, 225, 226, 227, 237, 238, 240, 245, 247, 248, 249, 253, 254, 261, 264, 273, 276, 280 Latimeriidae, 115 Latimeroidei, 115 Latissimus dorsi, 282 Lepidosiren, 12, 18, 29, 30, 32, 113, 114, 122, 123, 158, 169, 170, 176, 178, 179, 180, 183, 184, 187, 198, 199, 202, 203, 204, 207, 217, 218, 225, 240, 250, 251, 252, 262 Lepidosirenidae, 12, 122 Lepisosteidae, 3, 5, 12, 40 Lepisosteus, 5, 12, 17, 29, 31, 40, 41, 43, 45, 50, 60, 159, 175, 176, 178, 184, 189, 195, 210, 225, 226, 239, 248, 253 Leptocephalus larva, 222 Levator anguli oris, 234 Levator arcus palatini, 184, 187, 189 Levator hyoideus, 244 Levator operculi, 190 Levator pterygoidei, 234 Maxilla, 191, 192, 193, 194 Megalops, 13, 17, 32, 52, 54, 59, 60, 165, 166, 174, 176, 182, 206, 208, 213, 217, 218 Mentomeckelian bones, 217 Mesethmoid, 154, 155 Mesocoracoid arch, 173, 174 Molecular studies, 29 Mormyridae, 13, 61, 303 Mormyrus, 13, 17, 52, 61, 62, 63, 65, 66, 143, 146, 170, 171, 174, 181, 200, 213, 214, 220, 238 !$$ Nematogenys, 13, 17, 112, 211, 213 Neoceratodontidae, 12, 122 Neoceratodus, 12, 14, 18, 32, 113, 121, 122, 123, 158, 169, 170, 179, 180, 183, 186, 190, 198, 199, 202, 203, 206, 218, 225, 249, 250, 251 Neopterygii, 32, 39, 40, 277, 318, 321 Neoteleostei, 3, 68, 74, 75, 193, 194, 212, 226, 318, 321 Notacanthus, 13, 17, 52, 54, 58, 60, 168, 176, 186 Notopteridae, 13, 61, 303 Omohyoideus, 258 Opercle, 208 Opsariichthys, 13, 18, 99, 109, 169, 296 Osmeroidea, 3, 13, 68, 226 Osmerus, 13, 18, 74, 77, 156, 166, 204, 220 Ostariophysi, 3, 11, 73, 78, 84, 85, 225, 226, 289, 305, 306, 318, 321 Osteichthyes, 1, 19, 28, 239, 267, 273, 274, 321 Osteoglossidae, 13, 61 Osteoglossomorpha, 3, 51, 59, 61, 63, 318 Otocephala, 3, 5, 73, 78, 225, 226, 301, 302, 305, 306, 318 Otophysi, 84, 99, 100, 102, 289, 299, 305, 306, 308, 309, 318, 321 Pantodon, 13, 17, 61, 62, 63, 65, 66, 159, 163, 171, 174, 189, 199, 200, 208, 219 Parakneria, 13, 18, 93, 94, 95, 164, 192, 204 Parietals, 160 Pectoral muscles, 149 Phractolaemus, 13, 18, 91, 92, 93, 144, 155, 156, 171, 181, 193, 197, 204, 209, 211, 212, 221 Pimelodus, 13, 18, 113, 190, 317 Plecoglossus, 13, 18, 77, 196, 204, 220 Polyodontidae, 3, 12, 33, 39 Polypteridae, 3, 12, 31 Polypterus, 12, 17, 30, 31, 32, 33, 41, 52, 148, 168, 169, 182, 184, 189, 195, 209, 210, 214, 226, 237, 238, 253, 276 Posttemporal, 171 Prearticulars, 216 Premaxillae, 194, 195 Preopercle, 208 Prevomer, 156 Pristigasteroidea, 13, 79, 80, 83 Pronator profundus, 283 Pronator teres, 282 Protacanthopterygii, 3, 5, 13 Protopterus, 12, 113, 122, 123, 170, 179, 180, 184, 225, 250, 251, 262 Protractor pterygoidei, 234 Psephurus, 12, 17, 30, 33, 36, 39, 122, 156, 159, 167, 169, 172, 173, 182, 187, 189, 192, 200, 215, 254 Pterygoids, 198 Pterygomandibularis, 234 Quadrate, 204, 206 Quadratojugals, 205 Retractor anguli oris, 234 Retroarticular, 218 Retropinna, 13, 18, 76, 77, 78, 198, 199 Rhipidistia, 117, 118, 119 Saccopharyngiformes, 13, 51, 58, 60, 61, 66, 321 Salmo, 13, 18, 68, 69, 75, 194 Salmoniformes, 3, 13, 68, 74, 76, 226, 322 Sarcopterygii, 1, 28, 29, 113, 114, 237, 254, 279, 318, 321, 325 Scaphium, 290 Searsia, 13, 18, 73, 74, 195, 220 Siluriformes, 13, 84, 109, 110, 112, 165, 288, 297, 311, 322 Silurus, 13, 18, 113, 155 Sternohyoideus, 147 Sternopygus, 13, 18, 108, 111, 192, 211, 219 !$% Stokellia, 13, 18, 76, 77, 78, 198, 199 Stomias, 13, 18, 74, 75, 76, 191, 194, 211, 323 Stomiiformes, 13, 76, 193, 212, 322, 323 Subopercle, 208 Supracleithrum, 166, 167 Supramaxillae, 193 Surangulars, 216 Swimbladder, 165 Symplectic, 205 158, 162, 168, 173, 174, 175, 178, 179, 180, 181, 185, 187, 188, 189, 198, 200, 203, 205, 207, 212, 214, 215, 216, 218, 227, 234, 239, 251, 257, 258, 281, 283, 285, 286, 317 Trichomycterus, 13, 18, 106, 112 Tripus, 295 Teleostei, 3, 5, 6, 7, 9, 11, 32, 39, 40, 41, 44, 45, 46, 50, 61, 63, 69, 80, 227, 230, 238, 241, 256, 274, 275, 276, 277, 278, 303, 310, 318, 321 Teleostomi, 1, 28 Tetrapoda, 4, 5, 7, 11, 12, 113, 117, 128, 318, 321 Thryssa, 13, 17, 79, 80, 83, 155, 307 Thymallus, 13, 18, 75, 157, 161, 194 Timon, 12, 13, 18, 32, 113, 114, 120, 121, 131, 133, 144, 145, 150, 152, 153, 154, Weberian apparatus, 11, 84, 98, 105, 224, 288, 289, 290, 291, 292, 293, 297, 300, 301, 302, 305, 308, 309, 310 Umbra, 13, 18, 77, 161 Urohyal, 213 Xenocharax, 13, 18, 99, 110 Xenodermichthys, 13, 18, 68, 71, 73, 74, 209 Xenomystus, 13, 17, 61, 62, 65, 67, 143, 144, 149, 170, 171, 199, 200, 219, 223 Zebrafish, 230, 241, 256, 267, 275, 278 MONITORING AND EVALUATION OF SOIL CONSERVATION AND WATERSHED DEVELOPMENT PROJECTS SH LEV-OP HH-SUP INTE LEV-AP HH-INF DIL-OP AD-HYO PH 0.1 mm INTM-P INTM-A ADM OM Plate Ventrolateral view of the cephalic muscles of a 4-day-old zebrafish larva (3.0 mm TL) BRM LEV-5 (BRM) AD-OP CMYK CMYK CMYK CMYK SH HH-SUP HH-INF PH LEV-AP INTE AD-OP AD-HYO DIL-OP LEV-OP 0.1 mm INTM-P INTM-A ADM AD-AP Plate Ventrolateral view of the cephalic muscles of a 4-day-old zebrafish larva (3.0 mm TL) PR-PEC (BRM) LEV-5 (BRM) CMYK CMYK CMYK CMYK LEV-OP DIL-OP HH-INF AD-OP HH-AB AD-HYO LEV-AP AD-AP 0.1 mm PR-H INTM-A AW A2 A0/A1-OST Plate Ventral view of the cephalic muscles of a 24-day-old zebrafish larva (6.0 mm TL) LEV-5 (BRM) SH HH-AD CMYK CMYK CMYK CMYK LEV-AP 0.1 mm A2 AW A0/A1-OST AD-AP Plate Dorsal view of the cephalic muscles of a 24-day-old zebrafish larva (6.0 mm TL) AD-OP LEV-5 AD-HYO (BRM) DIL-OP LEV-OP CMYK CMYK CMYK CMYK AD-HYO HH-AB HH-INF DIL-OP AD-OP PR-H-V 0.1 mm A2 A1-OST A0 AD-AP PR-H-D LEV-AP Plate Ventrolateral view of the cephalic muscles of a 35-day-old zebrafish larva (7.4 mm TL) HH-AD LEV-OP CMYK CMYK CMYK CMYK HH-AB HH-INF CMYK PR-H-V PR-H-D 0.1 mm Plate Ventral view of the cephalic muscles of a 35-day-old zebrafish larva (7.4 mm TL) HH-AD CMYK CMYK CMYK AD-OP AD-HYO DIL-OP AD-HYO LEV-AP LEV-AP 0.1 mm AD-AP Plate Dorsal view of the cephalic muscles of a 4-day-old zebrafish larva (3.0 mm TL) LEV-OP LEV-OP AD-OP DIL-OP CMYK CMYK CMYK CMYK ... concerning the relationships of higher clades One of our aims here is thus to discuss whether or not the results of the cladistic analysis of the present work, which is precisely focused on the higher- level... are the closest living relatives of teleosts, the amiids of the genus Amia, or both these fishes and the lepisosteids of the genera Lepisosteus and Atractosteus, that is, the members of the three... works on the interrelationships among the various subunits of these two groups These results can also help to clarify the origin and homologies of certain structures found in the members of these