The Origin of Higher Clades Osteology, Myology, Phylogeny and Evolution of Bony Fishes and the Rise of Tetrapods (cid:1) (cid:8) (cid:8)(cid:11) (cid:8) (cid:2)(cid:3)(cid:4)(cid:5)(cid:2)(cid:6)(cid:4)(cid:3)(cid:7) (cid:9)(cid:3)(cid:10) (cid:12)(cid:9)(cid:13)(cid:14)(cid:9)(cid:5)(cid:4)(cid:2)(cid:3) (cid:2)(cid:15) (cid:16) (cid:8)(cid:17) (cid:8) (cid:2)(cid:4)(cid:13) (cid:2)(cid:3)(cid:16)(cid:11)(cid:6)(cid:12)(cid:9)(cid:5)(cid:4)(cid:2)(cid:3) (cid:9)(cid:3)(cid:10) (cid:18) (cid:8)(cid:10) (cid:8)(cid:20) (cid:9)(cid:5)(cid:11)(cid:6)(cid:16)(cid:19)(cid:11)(cid:10) (cid:11)(cid:12)(cid:11)(cid:13)(cid:2)(cid:20)(cid:1)(cid:11)(cid:3)(cid:5) (cid:6)(cid:2)(cid:21)(cid:11)(cid:17)(cid:5)(cid:16) 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 [email protected] (marketing department) [email protected] (editorial department) [email protected] (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) 1. Osteichthyes--Evolution. I. Title. QL618.2D562007 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. (cid:1)(cid:2)(cid:3)(cid:4)(cid:5)(cid:6)(cid:3) The Osteichthyes, including bony fishes and tetrapods, is a highly speciose group of animals, comprising more than 42000 living species. The extraordi- nary 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 macroevolu- tion 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 1 and 2 provide a short introduction to the main aims of the book and to the method- ology and methods used. Chapter 3 deals with an extensive cladistic analysis of osteichthyan higher-level interrelationships based on a phyloge- netic comparison of 356 characters in 80 extant and fossil terminal taxa representing all major groups of Osteichthyes. This cladistic analysis in- cludes various terminal taxa and osteological characters, and namely a large number of myological characters, not included in previous analyses. Chapter 4 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 implica- tions 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 (cid:1) (cid:8) (cid:8)(cid:11) (cid:8) (cid:2)(cid:3)(cid:4)(cid:5)(cid:2)(cid:6)(cid:4)(cid:3)(cid:7) (cid:9)(cid:3)(cid:10) (cid:12)(cid:9)(cid:13)(cid:14)(cid:9)(cid:5)(cid:4)(cid:2)(cid:3) (cid:2)(cid:15) (cid:16) (cid:8)(cid:17) (cid:8) (cid:2)(cid:4)(cid:13) (cid:2)(cid:3)(cid:16)(cid:11)(cid:6)(cid:12)(cid:9)(cid:5)(cid:4)(cid:2)(cid:3) (cid:9)(cid:3)(cid:10) (cid:18) (cid:8)(cid:10) (cid:8)(cid:20) (cid:9)(cid:5)(cid:11)(cid:6)(cid:16)(cid:19)(cid:11)(cid:10) (cid:11)(cid:12)(cid:11)(cid:13)(cid:2)(cid:20)(cid:1)(cid:11)(cid:3)(cid:5) (cid:6)(cid:2)(cid:21)(cid:11)(cid:17)(cid:5)(cid:16) (cid:1)(cid:2)(cid:2) (cid:7)(cid:6)(cid:8)(cid:9)(cid:10)(cid:11)(cid:12)(cid:3)(cid:13)(cid:14)(cid:3)(cid:15)(cid:3)(cid:9)(cid:16)(cid:17) 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 (cid:1)(cid:2)(cid:2)(cid:2) 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. (cid:2)(cid:3) (cid:18)(cid:19)(cid:17)(cid:16)(cid:20)(cid:10)(cid:4)(cid:20)(cid:7)(cid:21)(cid:21)(cid:2)(cid:3)(cid:22)(cid:19)(cid:5)(cid:16)(cid:19)(cid:10)(cid:9)(cid:17)(cid:23) II, III, IV, V, VII, IIX, IX, X foramens/nerves of Miles’s 1977 original drawing A0, A1, A1-OST adductor mandibulae A0, A1 and A1-OST A1-OST-L, A1-OST-M lateral and mesial sections of adductor mandibulae A1-OST A2 adductor mandibulae A2 A2-D, A2-PVM, A2-V dorsal, posteroventromesial and ventral sections of adductor mandibulae A3', A3'’ adductor mandibulae A3' and A3'’ AB-PRO abductor profundus AB-SUP abductor superficialis abs anterior bulla of swimbladder AC anconaeus coracoideus AD-AP adductor arcus palatini AD-HYO adductor hyomandibulae AD-OP adductor operculi AD-PRO adductor profundus AD-SUP adductor superficialis ADM adductor mandibulae AED1 abductor et extensor digiti I AHL, AHM anconaeus humeralis lateralis and medialis am ampulla am-m macula of ampulla AME adductor mandibulae externus ana anterior neural arch ang angular *Myological structures are shown in bold