Reproductive Biology of Bats This Page Intentionally Left Blank Reproductive Biology of Bats
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Reproductive Biology of Bats This Page Intentionally Left Blank Reproductive Biology of Bats Edited by Elizabeth G. Crichton Henry Doorly Zoo Omaha, Nebraska and Philip H. Krutzsch University of Arizona Tucson, Arizona ACADEMIC PRESS A Harcourt Science and Technology Company San Diego San Francisco New York Boston London Sydney Tokyo This book is printed on acid-free paper Copyright © 2000 Academic Press All Rights Reserved. No part of this publication may be reproduced or transmitted in any form or by any means electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Academic Press A Harcourt Science and Technology Company 32 Jamestown Road London NW1 7BY http://www.academicpress.com Academic Press A Harcourt Science and Technology Company 525 B Street, Suite 1900, San Diego, California 92101-4495, USA http://www.academicpress.com Library of Congress Catalog Card Number: 99-066843 A catalogue record for this book is available from the British Library ISBN 0-12-195670-9 Typeset by Phoenix Photosetting, Chatham, Kent Printed in Great Britain at the University Press, Cambridge 00 01 02 03 04 05 9 8 7 6 5 4 3 2 1 Contents Preface ix 1 Endocrinology of Reproduction in Bats: Central Control 1 Edythe L.P. Anthony 1.1 Introduction 1 1.2 The hypothalamic-pituitary complex 2 1.3 GnRH and portal mechanisms 4 1.4 GnRH perikarya and seasonal dynamics of the GnRH system 9 1.5 The nervus terminalis 12 1.6 Pituitary gonadotropins 12 1.7 Prolactin 17 1.8 Pineal melatonin and the suprachiasmatic nucleus 19 1.9 Summary and future perspectives 21 Acknowledgments 22 References 22 2 Endocrine Regulation of Reproduction in Bats: the Role of Circulating Gonadal Hormones 27 Len Martin and Ric T.F. Bernard 2.1 Introduction 27 2.2 Peripheral hormone levels and their evaluation 28 2.3 Source and synthesis of hormones in the Chiroptera 29 2.4 Peripheral hormone concentrations, binding proteins and hormone action in male bats 34 2.5 Peripheral hormone concentrations, binding proteins and hormone action in female bats 45 2.6 Unilateral endometrial growth (UEG) in Pteropus spp.; implications for systemic hormone levels 56 2.7 General discussion 57 References 58 3 Peripheral Endocrines in Bat Reproduction 65 Gary G. Kwiecinski and David A. Damassa 3.1 Introduction 65 3.2 Peripheral endocrines in bats 66 3.3 Sex hormone-binding globulin (SHBG) 77 3.4 Summary 85 References 86 vi Contents 4 Anatomy, Physiology and Cyclicity of the Male Reproductive Tract 91 Philip H. Krutzsch 4.1 Introduction 92 4.2 Puberty 93 4.3 Reproductive organ anatomy 93 4.4 Penis 101 4.5 Spermatozoa 105 4.6 Reproductive patterns 108 4.7 Conclusions and future directions 128 Acknowledgments 129 References 137 5 Anatomy and Physiology of the Female Reproductive Tract 157 John J. Rasweiler IV and Nilima K. Badwaik 5.1 Introduction 157 5.2 Anatomy of the reproductive tract 158 5.3 Functional asymmetries of the female tract 173 5.4 Nonpregnant reproductive cycles 186 5.5 Adaptive significance of reproductive asymmetries and menstruation 199 5.6 Acknowledgments 208 References 208 6 Pregnancy 221 Nilima K. Badwaik and John J. Rasweiler IV 6.1 Introduction 221 6.2 Preimplantation development 222 6.3 Implantation of the blastocyst 228 6.4 Development of placental organs and foetal membranes 242 6.5 Highly invasive trophoblast 269 6.6 Litter size and gestation period 273 6.7 The endocrinology of pregnancy 275 6.8 Postpartum oestrus 278 Acknowledgments 280 References 281 7 Sperm Storage and Fertilization 295 Elizabeth G. Crichton 7.1 Introduction: The what, when, where and why of sperm storage 295 7.2 The history of sperm storage in Chiroptera 297 7.3 The occurrence of sperm storage in Chiroptera 298 7.4 The how of sperm storage 300 7.5 Preparation of sperm for fertilization 311 7.6 Conclusions and future directions 311 Acknowledgments 312 References 312 Contents vii 8 Bat Mating Systems 321 Gary F. McCracken and Gerald S. Wilkinson 8.1 Introduction 321 8.2 Mating systems in bats and other mammals 322 8.3 Single male/multi-female polygynous groups 330 8.4 Multi-male/multi-female groups 341 8.5 Single male/single female groups 351 8.6 Ecological determinants of mating systems in bats 353 References 357 9 Life-history and Reproductive Strategies of Bats 363 Paul A. Racey and Abigail C. Entwistle 9.1 Introduction 364 9.2 Costs of reproduction 367 9.3 Seasonality and reproduction 367 9.4 Reproductive patterns of bats 371 9.5 Reproductive delays 379 9.6 Adaptive significance of reproductive delays 386 9.7 What limits reproduction? 392 9.8 Limits to reproductive output 398 Acknowledgments 401 References 401 10 Parental Care and Postnatal Growth in the Chiroptera 415 Thomas H. Kunz and Wendy R. Hood 10.1 Introduction 415 10.2 Parental care 416 10.3 Size and developmental state at birth 441 10. Postnatal growth 442 10.5 Conclusions, predictions and future directions 453 Acknowledgments 454 References 454 11 Environmental Regulation of Reproduction 469 Paul D. Heideman 11.1 Introduction 469 11.2 Ultimate factors 470 11.3 Mechanisms of environmental regulations of reproduction in bats 473 11.4 Insights from field studies 491 11.5 What should we do next? 493 11.6 Conclusion 494 Acknowledgments 494 References 495 Species Index 501 Subject Index 507 This Page Intentionally Left Blank Preface Some 20 years ago we discussed with Dr William A. Wimsatt the possibility of bringing together a volume devoted to the reproductive biology of Chiroptera. Our idea, at that time, was to present such a work as Volume 4 in the Biology of Bats. Many events interfered with the initiation of such a review, and finally the untimely death of Dr Wimsatt almost permanently set it aside. It is to Bill’s vision and memory that we gratefully and lovingly dedicate this volume. He is sorely missed not only for his contributions to this science but for his deep and sincere friendship and support. The great and ever growing interest in these unique and highly specialized mammals and the many new findings about their reproduction caused us to realize the importance of bringing together the available data concerning the reproductive aspect of the life history of these mammals. Knowledge of reproductive events and their controlling mechanisms may ultimately be central to the management of vulnerable populations. Perhaps the application of assisted reproductive technologies will become critical. The current knowledge of the reproductive biology of bats is fragmentary, appears in a wide range of scientific literature, and only concerns a small fraction of the 960+ species of Chiroptera. This mammalian order is worldwide in distribution and comprises nearly 25 per cent of the existing living mammal fauna. Volumes that synoptically consider aspects of bat reproduction include: Slaughter and Walton (1970), Kunz (1982, 1984), Hill and Smith (1984) and Altringham (1996). A comprehensive volume reviewing this critically significant area of chiropteran biology, however, is lacking and leaves us without an in-depth compilation of pertinent literature, which is worldwide, in many languages and often difficult to obtain. Our aim is to summarize (in one volume) the current knowledge of reproductive biology in bats. Perhaps this will provide a stimulus to students and/or established investigators to find answers to some of the many unknowns. Authors contributing to this volume were selected on their contributions to the subject field that they were to review. Their reviews were to bring forth and summarize data on specific topics. Each chapter was to include and summarize primarily reviews of the published literature and, less often, original work in progress. Contributors were encouraged to freely develop their ideas and presentations which resulted in some, but meaningful, redundancy. We are forever grateful to a number of individuals who helped make this immense task possible. We wish to thank our spouses Dorothy and Harold for their special support and understanding. We also owe special thanks to David Piper who took the time from his demanding schedule to advise upon and provide computer support, and David Karabinus, a trusted colleague, for his unselfish help in the x Preface mechanics of word processing. Secretarial assistance was graciously given by Lori Taylor and Susan Eastman of the Department of Cell Biology and Anatomy, University of Arizona. Robert S. McCuskey, Department head, is thanked for his support in the use of many department facilities and services. We are especially indebted to Dr. Ted Fleming for his contribution in the preparation of the Subject Index. In addition, we acknowledge the following colleagues who freely provided their professional insight and expertise in anonymously reviewing one or more of the chapter manuscripts: B.B. Benson, R.T.F. Bernard, J. Bradbury, G. McCracken, M. Cukierski, M.B. Fenton, A.W. Gustafson, Mac E. Hadley, P.D. Heideman, P. Hoyer, C. Pow, P.A. Racey, E.H. Studier, E.L. Cockrum and E.D. Widmaier. Elizabeth G. Crichton Philip H. Krutzsch Contributors Anthony, Edythe L.P., Department of Biology, Rhode Island College, Providence, RI 02908-1991, USA Badwaik, Nilima K., Department of Obstetrics and Gynecology, Cornell University Medical College, 1300 York Avenue, New York, NY 10021, USA Bernard, Ric T.F., Department of Zoology and Entomology, Rhodes University, P.O. Box 94 Grahamstown, 6140 South Africa Crichton, Elizabeth G., Center for Conservation and Research, Henry Doorly Zoo, Omaha, NE 68107, USA Damassa, David A., Department of Anatomy and Cellular Biology, Tufts University Schools of Medicine, Dental Medicine and Veterinary Medicine, 136 Harrison Avenue, Boston, MA 02111, USA Entwistle, A.C., Fauna and Flora International, Great Eastern House, Tenison Road, Cambridge, CB1 2DT, United Kingdom Heideman, Paul D., Biology Department, The College of William and Mary, P.O.