MAMMALOGY 2 MAMMALOGY Adaptation, Diversity, Ecology Fourth Edition George A. Feldhamer Lee C. Drickamer Stephen H. Vessey Joseph F. Merritt Carey Krajewski 3 © 2015 Johns Hopkins University Press All rights reserved. Published 2015 Printed in the United States of America on acid-free paper 9 8 7 6 5 4 3 2 1 Johns Hopkins University Press 2715 North Charles Street Baltimore, Maryland 21218-4363 www.press.jhu.edu Library of Congress Cataloging-in-Publication Data Mammalogy : adaptation, diversity, ecology / George A. Feldhamer [and four others]. — Fourth edition. pages cm Includes bibliographical references and index. ISBN 978-1-4214-1588-8 (hardcover : alk. paper)— ISBN 978-1-4214-1589-5 (electronic)— ISBN 1-4214-1588-7 (hardcover : alk. paper)— ISBN 1-4214-1589-5 (electronic) 1. Mammalogy. I. Feldhamer, George A. QL703.M36 2015 599—dc23 2014012644 A catalog record for this book is available from the British Library. For charts on weights & measures and conversions, see pages 748–749. Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at 410-516-6936 or [email protected]. Johns Hopkins University Press uses environmentally friendly book materials, including recycled text paper that is composed of at least 30 4 percent post-consumer waste, whenever possible. 5 For Carla, Carrie, Andy, Jenny, and Lucy – George For my siblings, Lynn Drickamer, Kurt Drickamer, Margaret Drickamer, and Priscilla Atkins – Lee For Kristin, my highly significant other – Steve In memory of my parents, Robert E. Merritt and June A. Merritt – Joe To Birdie, Jenny, Robby, and Cody—inspiration every day – Carey 6 About the Authors George A. Feldhamer is Professor Emeritus of Zoology and former Director of the Environmental Studies Program at Southern Illinois University, Carbondale. His research has focused on mammalian populations, ecology, and management; biology of introduced deer; and threatened and endangered species of rodents and bats. He is a former associate editor for Forest Biology and Ecology for the Journal of Forest Research and a former associate editor of the Wildlife Society Bulletin. He is the senior editor of Wild Mammals of North America: Biology, Management, and Conservation (2003), coauthor of Mammals of the National Parks (2005), and Deer: The Animal Answer Guide (2012), all published by Johns Hopkins University Press, and coeditor of Ecology, Behavior, and Conservation of the Golden Mouse: A Model Species for Research (2007). He has had 35 years of experience teaching an upper- division mammalogy course. In 2011, he received the Joseph Grinnell Award from the American Society of Mammalogists “to honor individuals who have made outstanding and sustained contributions to education in mammalogy” Lee C. Drickamer was Professor of Biology and Chair of the Department of Biological Sciences at Northern Arizona University, Professor of Biology at Williams College for 15 years, Professor of Zoology at Southern Illinois University, Carbondale, for 11 years, and is Regents’ Professor Emeritus of Biological Science at Northern Arizona University. He is a past president of the Animal Behavior Society, past secretary-general of the International Council of Ethologists, past chair of the Division of Animal Behavior of what is now the Society for Integrative and Comparative Biology, and former editor of Animal Behaviour. His research emphases have included social factors affecting development and reproduction in house mice and swine, behavioral ecology of house mice and deer mice, social biology of primates, intrauterine position effects on behavior and reproduction of mice and swine, the consequences of mate selection for offspring viability in house mice, and prairie dog ecology and population biology. Stephen H. Vessey is Professor Emeritus of Biological Sciences at Bowling Green State University. His research interests include the behavioral ecology of mammals, especially primates and rodents. He has 7 been studying a population of white-footed mice in northwestern Ohio for more than 40 years. He is a former associate editor of the Journal of Mammalogy and is a Fellow of the Animal Behavior Society. He taught mammalogy and animal behavior at Bowling Green for 30 years and coauthored a textbook in animal behavior with Lee Drickamer. After retiring from Bowling Green in 2000, he served as Program Director and Deputy Division Director, Division of Integrative Biology and Neurosciences, National Science Foundation. Joseph F. Merritt is Senior Mammalogist with the Illinois Natural History Survey, University of Illinois. He is the former Director of Powdermill Biological Station of the Carnegie Museum of Natural History and served as Distinguished Visiting Professor at the U.S. Air Force Academy in Colorado Springs, Colorado, during the academic year 2004– 2005. Dr. Merritt is a physiological ecologist and functional morphologist specializing in adaptations of mammals to cold. He is the author of Guide to the Mammals of Pennsylvania (1987), published by the University of Pittsburgh Press, and editor of several technical monographs on specific taxa of mammals. His book Biology of Small Mammals was published by Johns Hopkins University Press in 2010. He is currently editor in chief of the Journal of Mammalogy. Dr. Merritt teaches mammalogy at the University of Colorado Mountain Research Station, courses in mammalian biology at Illinois State University, Normal, and courses in winter ecology at the Adirondack Ecological Center, SUNY College of Environmental Science and Forestry. Carey Krajewski is Professor and Chair of Zoology at Southern Illinois University, Carbondale. His research in mammalogy has involved molecular studies of marsupial phylogeny, particularly the systematics of dasyuromorphians, as well as the evolution of gruoid birds. He was an associate editor for the Journal of Mammalogy and is currently associate editor of the Journal of Mammalian Evolution and of Molecular Phylogenetics and Evolution. From 1993 to 2010, he was an honorary research fellow in the Genetics Department at La Trobe University, Australia. He was named 2004 Outstanding Scholar in the College of Science at SIUC. 8 Brief Contents Preface PART 1 Introduction Chapter 1 The Study of Mammals Chapter 2 History of Mammalogy Chapter 3 Methods for Studying Mammals Chapter 4 Phylogeny and Diversification of Mammals Chapter 5 Evolution and Dental Characteristics Chapter 6 Biogeography PART 2 Structure and Function Chapter 7 Integument, Support, and Movement Chapter 8 Modes of Feeding Chapter 9 Control Systems and Biological Rhythms Chapter 10 Environmental Adaptations Chapter 11 Reproduction PART 3 Adaptive Radiation and Diversity 9 Chapter 12 Monotremes and Marsupials Chapter 13 Tenrecoidea, Macroscelidea, Scandentia, Dermoptera, Solenodonta, Soricomorpha, and Erinaceomorpha Chapter 14 Chiroptera Chapter 15 Primates Chapter 16 Pilosa and Cingulata (Xenarthrans), Pholidota, and Tubulidentata Chapter 17 Carnivora Chapter 18 Rodentia and Lagomorpha Chapter 19 Proboscidea, Hyracoidea, and Sirenia Chapter 20 Perissodactyla and Artiodactyla Chapter 21 Cetacea PART 4 Behavior and Ecology Chapter 22 Communication, Aggression, and Spatial Relations Chapter 23 Sexual Selection, Parental Care, and Mating Systems Chapter 24 Social Behavior Chapter 25 Dispersal, Habitat Selection, and Migration Chapter 26 Populations and Life History Chapter 27 Community Ecology PART 5 Special Topics Chapter 28 Parasites and Diseases Chapter 29 Domestication and Domesticated Mammals Chapter 30 Conservation 10