Functional and Evolutionary Ecology of Bats
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Functional and Evolutionary Ecology of Bats Akbar Zubaid Gary F. McCracken Thomas H. Kunz, Editors OXFORD UNIVERSITY PRESS FUNCTIONAL AND EVOLUTIONARY ECOLOGY OF BATS This page intentionally left blank FUNCTIONAL AND EVOLUTIONARY ECOLOGY OF BATS EDITED BY Akbar Zubaid, Gary F. McCracken, and Thomas H. Kunz 2006 Oxford University Press, Inc., publishes works that further Oxford University’s objective of excellence in research, scholarship, and education. 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 Copyright ß 2006 by Oxford University Press, Inc. Published by Oxford University Press, Inc. 198 Madison Avenue, New York, New York 10016 www.oup.com Oxford is a registered trademark of Oxford University Press 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 of Oxford University Press. Library of Congress Cataloging-in-Publication Data Functional and evolutionary ecology of bats / edited by Akbar Zubaid, Gary F. McCracken, and Thomas H. Kunz. p. cm. Includes bibliographical references (p. ). ISBN-13 978-0-19-515472-6 ISBN 0-19-515472-X 1. Bats—Ecology. I. Zubaid Akbar. II. McCracken, G.F. III. Kunz, Thomas H. QL737.C5F86 2005 599.4017—dc22 2004057541 987654321 Printed in the United States of America on acid-free paper Dedicated to Lim Boo Liat This page intentionally left blank Preface Research on bats during the second half of the 20th century has had a marked influence on how we now view these so-called secretive mammals. A new generation of researchers has begun to advance our knowledge about bats in disciplines including ecophysiology, functional morphology, population biology, molecular ecology, and conservation biology, with increasing num- bers of studies extending to remote corners of the Earth. Moreover, advances in molecular biology, genomics, bioinformatics, computational science, geo- graphic information systems, sophisticated imaging systems, transponders, and the miniaturization of radio-transmitters have provided new tools that were unavailable to earlier generations of bat researchers. The wealth of new technologies and information that has emerged from both field and laboratory studies has had an enormous impact on how research is now being conducted. Research on functional and evolutionary ecology of bats has emerged from largely descriptive studies to emerging emphases on hypothesis testing and experimentation in both the laboratory and the field. In part, this reflects an increased recognition that bats provide critical ecosystem services such as seed dispersal, pollination, and insect control that sustain healthy populations. This is especially true in light of growing concern over deforestation, habitat fragmentation, mining activities, and an increasing threat to natural habitats from air, water, and soil pollution and threats from global warming. This volume is the outgrowth of three symposia that were convened at the 12th International Bat Research Conference, hosted by the Universiti Kebangsaan Malaysia from August 5 to 9, 2001, held in Kuala Lumpur, Malaysia. For this conference, the editors invited conveners to organize symposia based on current research on functional and evolutionary ecology. The conveners in turn selected speakers to present reviews and analyses of their respective disciplines. Chapter authors were invited because they had distinguished themselves by expanding their research beyond traditional approaches. The conveners prepared written introductions to their respective symposia so that the overall contributions could be framed in a broad perspective. The first section on physiological ecology focuses on thermal biology of hibernation, energetics, daily heterothermy, and the evolution of basal metabolic rate. The next section on morphology includes topics ranging from form and function of dentition, ecomorphology of flower-visiting bats, form, viii Preface function, and phylogeny of quadrupedal bats, wing morphology and its implications for flight performance, and the relationships between cranial morphology and feeding ecology. The third section of the book on roosting and population biology includes topics ranging from population genetic structure, life-history traits, social behavior and relatedness, mating systems, and roosting ecology. This book would not have been possible without the dedication and enthusiasm of the conveners of each session who have helped craft the invited chapters into a synthetic whole. The convenors were not only instrumental in inviting the participants but also arranged for reviewers, assisted in the editing process, and wrote an introduction to their respective sessions. Each manuscript was thoroughly read by at least two anonymous reviewers, and we thank our colleagues who reviewed one or more of the chapter manuscripts. These individuals include Nadia Ayoub, Robert Barclay, Andy Biewener, Wieslaw Bogdanowicz, Mark Brigham, Tamsin Burland, Brock Fenton, Rod Foster, Trish Freeman, Fritz Geiser, Erin Gillim, John Hermanson, Anthony Herrel, Bruce Jayne, Kate Jones, Gerald Kerth, Barry Lovegrove, Alex Menzel, Collin O’Donnell, Steve Rossiter, Amy Russell, Randy Small, John Speakman, Tim Strickler, Carl Thulin, Don Thomas, Annie Tibbels, Peter Ungar, Maarten Vonhof, Jerry Wilkinson, Craig Willis, and John Winkelmann. We also thank Bethany Bernasconi, Kristine Faloon, and Erin Ruppert of Boston University’s Center for Ecology and Conservation Biology for assisting in the final preparation of the book manuscript. We are grateful to Kirk Jenson, who expressed enthusiasm for this project from the onset and for his patience and forbearance. We especially thank Peter Prescott who assumed responsibility for shepherding the final stages of this book to completion. We also thank Kaity Cheng of the editorial and production staff at Oxford University Press for her assistance in preparing the final product. Finally, we thank our wives, Lisa Syarma Addini, Jamey Dobbs, and Margaret Kunz, for their patience, tolerance, understanding, love, and support. Contents Contributors, xiii PART I Physiological Ecology Donald W. Thomas & John R. Speakman 1 Energetics, Thermal Biology, and Torpor in Australian Bats, 5 Fritz Geiser 2 Temperature, Hibernation Energetics, and the Cave and Continental Distributions of Little Brown Myotis, 23 Murray M. Humphries, John R. Speakman, & Donald W. Thomas 3 Daily Heterothermy by Temperate Bats Using Natural Roosts, 38 Craig K.R. Willis 4 Exploring the Evolution of the Basal Metabolic Rate in Bats, 56 Ariovaldo P. Cruz-Neto & Kate E. Jones PART II Functional Morphology Elizabeth R. Dumont & William A. Schutt, Jr. 5 Quantifying Relationships between Form and Function and the Geometry of the Wear Process in Bat Molars, 93 Alistair R. Evans ix x Contents 6 Dynamic Complexity of Wing Form in Bats: Implications for Flight Performance, 110 Sharon M. Swartz, Kristin Bishop, & Maryem-Fama Ismael Aguirre 7 Performance Analysis as a Tool for Understanding the Ecological Morphology of Flower-Visiting Bats, 131 Christopher W. Nicolay & York Winter 8 Quadrupedal Bats: Form, Function, and Evolution, 145 William A. Schutt, Jr., & Nancy B. Simmons 9 The Correlated Evolution of Cranial Morphology and Feeding Behavior in New World Fruit Bats, 160 Elizabeth R. Dumont PART III Roosting Ecology and Population Biology Gary F. McCracken, Linda F. Lumsden, & Thomas H. Kunz 10 Social and Population Structure in the Brown Long-Eared Bat, Plecotus auritus, 185 Tamsin M. Burland, Abigail C. Entwistle, & Paul A. Racey 11 Relatedness, Life History, and Social Behavior in the Long-Lived Bechstein’s Bat, Myotis bechsteinii, 199 Gerald Kerth 12 Causes and Consequences of Genetic Structure in the Greater Horseshoe Bat, Rhinolophus ferrumequinum, 213 Stephen J. Rossiter, Gareth Jones, Roger D. Ransome, & Elizabeth M. Barratt 13 Population Genetic Structure of Very Large Populations: The Brazilian Free-Tailed Bat, Tadarida brasiliensis, 227 Amy L. Russell & Gary F. McCracken Contents xi 14 Evolutionary Dynamics of the Short-Nosed Fruit Bat, Cynopterus sphinx (Pteropodidae): Inferences from the Spatial Scale of Genetic and Phenotypic Differentiation, 248 Jay F. Storz, Hari R. Bhat, Johnson Balasingh, P. Thiruchenthil Nathan, & Thomas H. Kunz 15 Conflicts and Strategies in the Harem-Polygynous Mating System of the Sac-Winged Bat, Saccopteryx bilineata, 269 Christian C. Voigt, Gerald Heckel, & Otto von Helversen 16 Flexibility and Specificity in the Roosting Ecology of the Lesser Long-Eared Bat, Nyctophilus geoffroyi: A Common and Widespread Australian Species, 290 Linda F. Lumsden & Andrew F. Bennett 17 Causes and Consequences of Tree-Cavity Roosting in a Temperate Bat, Chalinolobus tuberculatus, from New Zealand, 308 Colin F.J. O’Donnell & Jane A. Sedgeley Index, 329 This page intentionally left blank Contributors Akbar Zubaid is Professor of Biological Sciences at the Universiti Kebangsaan Malaysia. His research focuses mostly on population biology, community ecology, and conservation biology of bats. He is the author or coauthor of over 60 publications on bats. He is the coeditor (with Zainal Abidin Abu