Horns, Pronghorns, and Antlers George A. Bubenik Anthony B. Bubenik Editors Horns, Pronghorns, and Antlers Evolution, Morphology, Physiology, and Social Significance With 204 Illustrations Springer-Verlag New York Berlin Heidelberg London Paris Tokyo Hong Kong George A. Bubenik Anthony B. Bubenik Associate Professor Consultant in Big Game Research Department of Zoology & Management University of Guelph Thornhill, Ontario L3T 3X7 Guelph, Ontario NIG 2Wl Canada Canada Cover illustrations: Descriptions can be found in the legends to the following figures in the text Chapter I: Figure 1 (page 6); Figure 4 (page 11); Figure 26 (page 57). The illustration of the deer with antlers does not appear in this book. It was prepared by and is copyrighted by Anthony B. Bubenik. Library of Congress Cataloging-in-Publication Data Horns, pronghorns, and antlers: evolution, morphology, physiology, and social significance/George A. Bubenik, Anthony B. Bubenik, editors. p. cm. Includes bibliographical references. ISBN-13:978-1-4613-8968-2 1. Horns. 2. Antlers. I. Bubenik, George A. II. Bubenik, Anton B. QL942.H77 1990 599.73'504185-dc20 89-29975 © 1990 Springer-Verlag New York Inc. Softcover reprint of the hardcover 1st edition 1990 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag New York, Inc., 175 Fifth Avenue, New York, NY 10010, USA), except for brief excerpts in connection with reviews or scholarly analysis. Use in con nection with any form of information storage and retrieval, electronic adaptation, computer soft ware, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use of general descriptive names, trade names, trademarks, etc., in this publication, even if the former are not especially identified, is not to be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Typeset by Publishers Service of Montana, Bozeman, Montana. 9 8 765 432 1 ISBN-13:978-1-4613-8968-2 e-ISBN-13:978-1-4613-8966-8 DOl: 10.1007/978-1-4613-8966-8 Preface Since the first drawings left on walls of ancient caves, human beings have been fascinated with that unique phenomenon of the animal kingdom, the presence of horns and antlers. From the mythical ''unicorn'' exercising the power over life and death to the perceived aphrodisiacal and other medical properties of rhinoceros horns and growing antlers, these conspicuous protuberances have had a significant place in the history of mankind. Part of that ancient interest in antlers and horns was due to their value as sym bols of masculinity; this interest persists today in trophy hunting, an honorable tradition carried on for centuries in many countries of the world. This book, which deals with evolution, morphology, physiology, and behavior, has not been devised as a comprehensive review of the subject of horns, prong horns, and antlers; rather, it is a series of chapters stimulating thoughts, discus sions, and initiation of new studies. As editors, we did not interfere with the content of articles nor with the opin ions and interpretations of our contributors, and we left them to decide whether to accept the suggestions of our reviewers. Despite the fact that various aspects of cranial appendages have been studied since the end of the eighteenth century, many controversial views still exist, as witnessed in various chapters of this book. We do not consider the presence of contradictory views as a shortcoming but rather as an advantage demonstrating where our knowledge needs improve ment, where conformity exists, or where future studies should be aimed. v vi Preface Finally, both of us would like to thank all contributors to this volume for their collaboration on this rather unique project, which brings together endocrinolo gists, taxonomists, paleontologists, experimental morphologists, neuroendocri nologists, behaviorists, and physiologists. It was a great challenge, and we hope that we met it "head on." GEORGE A. BUBENIK ANTHONY B. BUBENIK Acknowledgments The scientific accuracy as well as the organization and flow of individual chap ters were greatly enhanced by the generous help of our panel of reviewers. Some of them were acknowledged by individual contributors, but others were not. Therefore, we as editors would like to express our gratitude to our following col leagues for their generous effort in reviewing various chapters of this book: E.D. Balon, C. Barrette, R.C. Bigalke, J.P. Bogart, D.E. Brown, N. Chapman, T.H. Clutton-Brock, R.J. Goss, P. Heizmann, H.A. Jacobson, D.w. Kitchen, R.L. Marchinton, C.D. Mcinnes, S.R: Scadding, D. Schams, J.M. Suttie, E. Thenius, D.E. Ullrey, A.C.V. Van Bemmel, and S.D. Webb. In addition, we would like to thank Mary Bubenik, Carolyn Pollard, Yoko Imai, and Jane Taylor for their excellent technical assistance. GEORGE A. BUBENIK ANTHONY B. BUBENIK vii Contents Preface. .. . .. . . .. . . . . . . . . . . .. . . . . . .. . .. . . . . .. . .. . . . . . .. . . . . . v Acknowledgments. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii Contributors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xl I. Evolution and Morphology 1. Epigenetical, Morphological, Physiological, and Behavioral Aspects of Evolution of Horns, Pronghorns, and Antlers . . . . . . . . . . 3 ANTHONY B. BUBENIK 2. Correlation of Reproductive and Digestive Strategies in the Evolution of Cranial Appendages. . . . . . . . . . . . . . . . . . . . . . . . .. 114 CHRISTINE M. JANIS 3. Muntiacidae....... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 134 COLIN P. GROVES AND PETER GRUBB 4. Cervidae of Southeast Asia. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 169 PETER GRUBB 5. Cranial Appendages of Giraffoidea 180 CHARLES S. CHURCHER 6. Evolution and Dispersal of African Bovidae . . . . . . . . . . . . . . . . . . .. 195 ALAN W. GENTRY II. Physiology, Genetics, and Behavior 7. The Pronghorn (Antilocapra americana) . . . . . . . . . . . . . . . . . . . . . .. 231 BART W. O'GARA ix x Contents 8. Neuroendocrine Regulation of the Antler Cycle. . . . . . . . . . . . . . . .. 265 GEORGE A. BUBENIK 9. Of Antlers and Embryos. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 298 RICHARD 1. Goss 10. Antler Regeneration: Studies with Antler Removal, Axial Tomography, and Angiography. . . . . . . . . . . . . . . . . . . . . . . . .. 313 JAMES M. SUTTlE AND PETER F. FENNESSY 11. The Role of the Nervous System in the Growth of Antlers 339 GEORGE A. BUBENIK 12. Experimental Manipulation of the Neural Control of Antler Growth 359 JAMES M. SUTTlE 13. Experimental Induction of Antler Growth. . . . . . . . . . . . . . . . . . . . .. 371 ZBIGNIEW JACZEWSKI 14. The Annual Antler Cycle of the European Roe Deer (Capreo/us capreo/us) in Relation to the Reproductive Cycle. . . . . . . . . . . . . . .. 396 ANTOINE 1. SEMPERE 15. Endocrine Mechanisms and Antler Cycles in Rusa Deer, Cervis (Rusa) timorensis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 416 SIMONE VAN MOURIK AND TEODOR STELMASIAK 16. Nutrition and Antler Development 426 ROBERT D. BROWN 17. Social Status and Antler Development in Red Deer . . . . . . . . . . . . .. 442 LUDEK BARTOS 18. Genetic Variability and Antler Development ................... 460 KIM T. SCRIBNER AND MICHAEL H. SMITH 19. The Antler as a Model in Biomedical Research 474 GEORGE A. BUBENIK References. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 488 Index of Extant and Extinct Species of Pecora . . . . . . . . . . . . . . . . . . . . .. 545 Index of General Terms . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . .. 551 Contributors LUDEK BARTOS, Group of Ethology, Institute of Animal Production, CS-104 00, Praha 10-Uhrineves, Czechoslovakia. ROBERT D. BROWN, Department of Wildlife and Fisheries, Mississippi State University, Mississippi State, Mississippi 39762-5917 U.S.A. ANTHONY B. BUBENIK, Consultant in Big Game Research & Management, 10 Stornoway Crescent, Thornhill, Ontario L3T 3X7 Canada. GEORGE A. BUBENIK, Department of Zoology, University of Guelph, Guelph, Ontario NIG 2Wl Canada. CHARLES S. CHURCHER, Department of Zoology, University of Toronto, Toronto, Ontario M5S lAl; and Department of Vertebrate Palaeontology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 Canada. PETER F. FENNESSY, MAF Technology, South Invermay Agricultural Centre, Pri vate Bag, Mosgiel, New Zealand. ALAN W. GENTRY, Department of Palaeontology, The Natural History Museum, Cromwell Road, London SW7 5BD England. RICHARD 1. Goss, Division of Biological and Medical Sciences, Brown Univer sity, Providence, Rhode Island 02912 U.S.A. COLIN P. GROVES, Department of Prehistory and Anthropology, The Australian National University, GPO Box 4, Canberra ACT 2601 Australia. PETER GRUBB, 35 Downhills Park Road, London N17 5PE England. ZBIGNIEW JACZEWSKI, Institute of Genetics and Animal Breeding, Polish Academy of Sciences, Popielno, 12-222 Wejsuny Poland. xi xii Contributors CHRISTINE M. JANIS, Division of Biology and Medicine, Brown University, Providence, Rhode Island 02912 U.S.A. BART W. O'GARA, U.S. Fish and Wildlife Service, Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, Montana 59812 U.S.A. KIM T. SCRIBNER, The University of Georgia, Savannah River Ecology Labora tory, Drawer E, Aiken, South Carolina 29802; and Department of Zoology, The University of Georgia, Athens, Georgia 30602 U.S.A. ANTOINE J. SEMPERE, Centre d'Etudes Biologiques des Animaux Sauvages (CNRS), Villiers-en-Bois, F-79360 Beauvoir-sur-Niort, France. MICHAEL H. SMITH, The University of Georgia, Savannah River Ecology Laboratory, Drawer E, Aiken, South Carolina 29802; and Department of Zool ogy, The University of Georgia, Athens, Georgia 30602 U.S.A. JAMES M. SUTTlE, MAF Technology, South Invermay Agricultural Centre, Pri vate Bag, Mosgiel, New Zealand. TEODOR STELMASIAK, Biocene International (Vic) Pty Ltd, Veterinary Precinct, VRl Building, Parkville 3052 Australia. SIMONE VAN MOURIK, University of Melbourne, School of Agriculture and Fore stry, Parkville 3052 Australia.
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