Opossums This page intentionally left blank Opossums An Adaptive Radiation of New World Marsupials Robert S. Voss and Sharon A. Jansa Johns Hopkins University Press Baltimore © 2021 Johns Hopkins University Press All rights reserved. Published 2021 Printed in the United States of Ameri ca on acid- free paper 9 8 7 6 5 4 3 2 1 Johns Hopkins University Press 2715 North Charles Street Baltimore, Mary land 21218-4363 www . press . jhu . edu Library of Congress Cataloging- in- Publication Data Names: Voss, Robert S., author. | Jansa, Sharon A., author. Title: Opossums : an adaptive radiation of new world marsupials / Robert S. Voss and Sharon A. Jansa. Description: Baltimore : Johns Hopkins University Press, 2021. | Includes bibliographical references and index. Identifiers: LCCN 2020013111 | ISBN 9781421439785 (hardcover) | ISBN 9781421439792 (ebook) Subjects: LCSH: Opossums. Classification: LCC QL737.M34 V67 2021 | DDC 599.2/76—d c23 LC rec ord available at https:// lccn . loc . gov / 2020013111 A cata log rec ord for this book is available from the British Library. Special discounts are available for bulk purchases of this book. For more information, please contact Special Sales at [email protected]. Johns Hopkins University Press uses environmentally friendly book materials, including recycled text paper that is composed of at least 30 percent post- consumer waste, whenever pos si ble. For Louise Emmons This page intentionally left blank Contents Acknowle dgments ix Introduction 1 I phylogen et ic context and historical biogeography 1 Metatheria and Marsupialia 7 2 South Amer ic a, the Island Continent 14 3 The G reat American Biotic Interchange and Its Aftermath 21 II opossum classification and diversity 4 Taxonomic Accounts 29 III opossum phenotypes 5 Morphology 67 6 Physiology 101 7 Be hav ior 121 IV opossum natu ral history 8 Habitats 137 9 Diets 153 10 Parasites 169 11 Predators 186 12 Competitors and Mutualists 195 13 Population Biology 211 viii Contents V synthesis 14 Adaptive Radiation 229 Appendixes 1. A Checklist of Living Opossums (Didelphidae) 241 2. Fruit Taxa Eaten by Opossums in French Guiana 245 3. Prey Taxa Eaten by Opossums in Southeastern Brazil 247 References 249 Index 303 Acknowle dgments We are very fortunate in having colleagues who w ere willing to read early drafts of our chapters and suggest improvements. In par tic u lar, we are grateful to Robin Beck for reading Chapters 1, 2, and 3; to Vera Weisbecker for reading Chapter 5; to Christine Cooper and John Harder for reading Chapter 6; to Harry Greene for reading Chapters 9 and 11; to Lance Durden, Agustín Jiménez, and Jessica Light for reading Chapter 10; and to Gregory Adler for reading Chapters 8, 12, and 13. Any remaining errors of fact, interpretation, or emphasis are, of course, our own. Patricia J. Wynne drew all the line art for this volume with her customary skill, attention to detail, and inimitable style. Photographic material was made available by friends, colleagues, and freelance photog raphers whose generosity we gratefully acknowledge, whether or not we fi nally used their images for this volume: Víctor Acosta- Chaves, Antoine Baglan, Eduardo Beltrocco, Gerardo Ceballos, Rodrigo Conte, Mónica Díaz, Louise Emmons, Antoine Fouquet, Elí García- Padilla, Mario Gómez- Martínez, Roland Kays, Jorge La Grotteria, José Martínez, José Ochoa, Ricardo Ojeda, Pedro Peloso, Darío Podestá, Fiona Reid, Rogério Rossi, Thiago Semedo, Thiago da Silveira, Pablo Teta, and Christian Ziegler. Rights to reproduce the image in Figure 11.1 w ere purchased from Superstock, Inc. Patricia Wynne’s drawings of opossum manual osteology (Fig. 5.10) are based on micro- CT scans expertly pro cessed by Abigail Curtis for a grant proposal that was never funded. We thank Kelly Kroft for photographing the microfilariae (Fig. 10.1) in a blood smear that was originally prepared by Kevin Olival, and we thank Agustín Jiménez for identifying adult worms from the body cavity of the same specimen. We are grateful to Craig Chesek for photographing the opossum skin in Fig. 4.10. Jan Ryser very kindly sent us a copy of his 1990 doctoral disser- tation on Didelphis. Our decades- long research on opossums would have been far less productive without many substantial contributions from our students, including especially Juan Díaz- Nieto, Tom Giarla, Eliecer Gutiérrez, and Silvia Pavan. Lorissa Fujishin and Carmen Martin helped generate much of the sequence data on which our