On the Wing On the Wing Insects, Pterosaurs, Birds, Bats and the Evolution of Animal Flight David E. Alexander Illustrations by Sara L. Taliaferro 1 1 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. Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States of America by Oxford University Press 198 Madison Avenue, New York, NY 10016, United States of America © Oxford University Press 2015 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, by license, or under terms agreed with the appropriate reproduction rights organization. Inquiries 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 work in any other form and you must impose this same condition on any acquirer Cataloging-in-Publication data is on file at the Library of Congress 9780199996773 9 8 7 6 5 4 3 2 1 Printed in the United States of America on acid-free paper To the late Larry Martin gone too soon CONTENTS Acknowledgments ix Note on Sources xi 1. Can’t Tell the Players without a Scorecard 1 2. Theme and Variations: Similarities and Differences among Nature’s Flyers 21 3. How to Fly? 39 4. Gliding Animals: Flight without Power 60 5. Insects: First to Fly 74 6. Birds: The Feathered Flyers 103 7. Bats: Wings in the Dark 130 8. Pterosaurs: Bygone Dragons 147 9. Pedestrians Descended from Flyers: Loss of Flight 164 10. Unifying Themes? 173 Bibliography 183 Index 197 ACKNOWLEDGMENTS This book had an amazingly long gestation. I wrote the first draft of the first chapter in 2002. Between then and now, I talked to many people about the evolution of flight, and while I may have forgotten some of the conversations or people, they nevertheless helped shape my thinking on the topic. Many of these conversations were with my colleagues at the University of Kansas, including Ron Barrett-Gonzalez, Dave Burnham, George Byers, Amanda Falk, Rudolf Jander, Matt Jones, and Bob Timm, as well as the many students in my Animal Flight Seminar over the years. Ed Wiley, Bruce Lieberman, and Michael Engel all helped this non-systematist un- derstand (and keep from embarrassing myself with) phylogenetic system- atics. Mark Robbins showed me museum specimens when I needed to see details of bird anatomy and feather structure and answered many orni- thological questions. I learned a lot about paleontology, dinosaurs, and birds from the many hours I spent in the lab of the late Larry Martin; no one enjoyed a good argument or took it less personally than Larry. Larry’s former student, Chris Bennett, patiently and thoroughly answered my many e-mails with questions about pterosaurs. I also discussed animal flight or flight evolution at various times with Roy Beckemeyer, Kristin Bishop, Sankar Chatterjee, Jeff Dawson and his grad student Ryan Chle- bak, Robert Dudley, Jimmy McGuire, the late John McMasters, Jake Socha, Sharon Swartz, Jim Usherwood, and Steve Yanoviak, and there were probably others whom I am sorry to say I have forgotten. Chris Ben- nett, Robert Dudley, Nick Longrich, and Jake Socha each read and com- mented on a chapter, and comments from two anonymous reviewers who heroically read the entire manuscript significantly improved the final ver- sion. Any mistakes or inaccuracies that may remain are, of course, my re- sponsibility. Sara Taliaferro drew most of the illustrations; hers say “(courtesy of S. T.)” in the figure legend. Also, Roy Beckemeyer and my son, Kevin Alexander, each drew a figure. (If no credit is listed with a figure, it [ ix ]
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