The Evolution of Organ Systems This page intentionally left blank The Evolution of Organ Systems A. Schmidt-Rhaesa 1 3 Great Clarendon Street, Oxford OX2 6DP Oxford University Press is a department of the University of Oxford. It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York © A. Schmidt-Rhaesa 2007 The moral rights of the author have been asserted Database right Oxford University Press (maker) First published 2007 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, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization. Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by Newgen Imaging Systems (P) Ltd., Chennai, India Printed in Great Britain on acid-free paper by Antony Rowe, Chippenham ISBN 978–0–19–856668–7 978–0–19–856669–4 (Pbk.) 10 9 8 7 6 5 4 3 2 1 For my family Daniela, Jelka, Jorrit, Finn Jaro and Tjorven Finja This page intentionally left blank Contents Preface ix 1. Introduction 1 2. The phylogenetic frame 3 3. General body organization 34 4. Epidermis 54 5. Musculature 74 6. Nervous system 95 7. Sensory organs 118 8. Body cavities 148 9. Excretory systems 169 10. Circulatory systems 191 11. Respiratory systems 202 12. Intestinal systems 218 13. Reproductive organs 240 14. Gametes (Spermatozoa) 262 15. Final conclusions 293 References 294 Index 365 vii This page intentionally left blank Preface The roots of this book go back to my student which have impressed me with their dedication time, when I was joining the vivid and produc- to zoology are Vicky and John Pearse as well as tive group of Peter Ax. We studied comparative Reinhardt Møbjerg Kristensen and Claus morphology in a variety of organisms with two Nielsen. While writing on this book, hearing aims: to reconstruct phylogenetic relationships of Reinhard Rieger’s death was shocking. His and to understand the diversity of animal impact on comparative morphology and evolu- morphology, in particular with its historical, tionary biology was fundamental. evolutionary background. It was especially Some colleagues were so kind as to read Thomas Bartolomaeus’ work on body cavities chapters from this book and helped me with and excretory organs that influenced me much valuable corrections, additions, and suggestions. and his (unfortunately largely unpublished) These are Wilko Ahlrichs (Oldenburg), Thomas habilitation thesis from 1993 can be regarded as Bartolomaeus (Berlin), Thorsten Burmester a kind of spiritual ancestor of this book. Research (Hamburg), Marco Ferraguti (Milano), and like that of Thomas showed that the fascination Ronald Jenner (Bath). Many thanks to all of you. in systematics does not stop after reconstructing Other colleagues helped me by supplying a hypothesis on animal relationships, but that it images for this book or gave me valuable continues with a discussion of what the conse- information, these are (in alphabetical order): quences of this hypothesis are for the evolution Wilko Ahlrichs, Thomas Bartolomaeus, Daniela of organisms and their organ systems. Candia Carnevali, Marco Ferraguti, Peter Grobe, Therefore my first thanks go to those who Alexander Gruhl, Harald Hausen, Iben Heiner, have intimately accompanied or supervised my Holger Herlyn, Alexander Kieneke, Reinhardt scientific career, some as teachers, some as col- Møbjerg Kristensen, Juliane Kulessa, Carsten leagues and friends. These are Peter Ax, Ulrich Lüter, Georg Mayer, Günter Purschke, Björn Ehlers, Thomas Bartolomaeus, Jim Garey, and Quast, Klaus Reinhardt, Birgen Holger Rothe, Klaus Reinhold. Matthias Schaefer belongs to Thomas Stach, Andrea Tapp, Andreas Unger. this group, although never a direct teacher, he Many thanks also go to Renate Feist, who as still taught me a lot. The large group of col- technical assistant made some of the photos used leagues that joined me over the years have had in this book. an important impact, although I mention here Then I honestly thank my students who only one dear friend, Wilko Ahlrichs. joined me over the years. In offering them the One person, who has published considerable possibility to dive into comparative morphol- contributions about the evolution of organ ogy, I am not sure if I did them a favour. The systems, especially in the latest edition of his decision for comparative morphology is every- book Invertebrate Zoology, is Ed Ruppert. thing other than a road to paradise. It requires Although knowing him personally only from a an extensive amount of enthusiasm which brief ‘water-to-the-belly’ (in my case) or ‘water- Ifound admirably in most of my students. From to-the-trousers’ (in his case) meeting inside all of them I want to thank Birgen Holger Rothe, Tampa Bay, I express my deepest respect for his in particular, for his unselfish expertise and contribution and authority. Other authorities, colleagueship. ix