Extinct Madagascar Extinct Madagascar: Picturing the Island’s Past Steven M. Goodman and William L. Jungers Plates by Velizar Simeonovski The University of Chicago Press Chicago and London Steven M. Goodman is the MacArthur Field Biologist at the Library of Congress Cataloging-in-Publication Data Field Museum, Chicago, and based in Antananarivo, Madagas- car. William L. Jungers is distinguished teaching professor Goodman, Steven M., author. and chair of anatomical sciences at Stony Brook University Extinct Madagascar: picturing the island’s past / Steven M. School of Medicine. Velizar Simeonovski is an artist based in Goodman and William L. Jungers; plates by Velizar Simeonovski. Chicago who specializes in reconstructions of extinct species pages cm and prehistoric landscapes. Includes bibliographical references and index. ISBN 978-0-226-14397-2 (cloth) — ISBN 978-0-226-15694-1 The University of Chicago Press, Chicago 60637 (e-book) 1. Fossils—Madagascar. 2. Animals—Madagascar. The University of Chicago Press, Ltd., London 3. Geology—Madagascar. 4. Plants—Madagascar. 5. Natural © 2014 by The University of Chicago history—Madagascar. I. Jungers, William L., 1948– author. Plates © 2014 by Velizar Simeonovski II. Title. All rights reserved. Published 2014. QE757.M28G664 2014 Printed in the United States of America 560.691—dc23 2014016894 23 22 21 20 19 18 17 16 15 14 1 2 3 4 5 This paper meets the requirements of ANSI/NISO Z39.48- 1992 (Permanence of Paper). The University of Chicago Press wishes to acknowledge g ratefully the Critical Ecosystem Partnership Fund, the Ellis Goodman Family Foundation, the Field Museum of Natural History, and Association Vahatra for their generous contribu- tions toward the publication of Extinct Madagascar: Picturing the Island’s Past. ISBN-13: 978-0-226-14397-2 (cloth) ISBN-13: 978-0-226-15694-1 (e-book) DOI: 10.7208/chicago/9780226156941.001.0001 To our families (Asmina, Hesham, Vavizara, Mboty, Mahery, Mandresy, Matthew, and Jocelyn) and In memory of Madame Berthe Rakotosamimanana and Robert Dewar Contents foreword ix Paleontological and Paleoecological acknowledgments xi Evidence 45 Summary 46 PART 1: Madagascar in Perspective: Human Interactions with Now-Extinct Land Past and Present Vertebrates 47 Reptiles 47 Introduction 3 Tortoise 47 General 3 Birds 47 Aspects of Format 3 Elephant Birds 47 The Artist 5 Mammals 48 Geological Time, Dates, and Radiocarbon Lemurs 48 Dating 7 Hippos 48 What Is a Subfossil? 9 Hypotheses on What Caused the Extinctions The Types of Subfossil Sites 10 during the Holocene 52 An Overview of the Origins, Geology, Animal Extinction, Conservation, and the Future 53 Colonization, and Modern Habitats of Madagascar 16 Madagascar in Deep Time—Isolation and Origin PART 2: Case Studies of Its Plants and Animals 16 Geographical Plates History of Animal Colonization 18 Plate 1: Cap Sainte Marie—the Ecology of Elephant Geography 18 Birds and Their Interface with Humans 59 Geology 22 Plate 2: Andrahomana I—the Ecology of Extreme Vegetational Patterns 23 Southeastern Madagascar and a Barometer of Humid Forest 25 Change 65 Dry Deciduous Forest 26 Plate 3: Andrahomana II—Evidence of a Holocene Spiny Bush 27 Tsunami in the Southern Indian Ocean and Savanna and Grassland Formations 30 Predator-Prey Relations 74 A Brief History of Climatic Change on Plate 4: Tsimanampetsotsa—Rapid Ecological Shifts Madagascar since the Late Pleistocene 36 in the Face of Natural Climate Change 79 Plate 5: Taolambiby—Hypotheses Associated with History of Human Colonization of Animal Extinction and Hunting by Humans: Madagascar 38 Physical Evidence and Interpretation 87 Considerations of Language and Diff erent Types Plate 6: Ankilitelo—a Deep Pit Cave and of Introductions 38 Inferences on Recent Ecological and Faunal The Human Genetic Evidence 41 Change 94 Evidence of Pre–Iron Age Colonization of Plate 7. Ampoza I—Reconstruction of the Ecology Madagascar 42 and Fauna in a Formerly Permanent Riverine The Archaeological Record of Occupation and Habitat in the Southwest 102 Settlement 44 Plate 8. Ampoza II—Ecological Change in a Forest Community and Connecting Humid Forest Corridors to the Eastern Portion of the Island 107 Plate 9: Belo sur Mer—a Window into Diff erent Hypotheses Associated with Environmental Change: Natural versus Human-Induced 112 Plate 10: Mananjary—the Former Estuary System of Eastern Lowland Madagascar and Some Its Faunal Elements 120 Plate 11: Antsirabe Region—Ecology of Highland Marsh and Forest Habitats as a Measure of Change through Time 125 Plate 12: Ampasambazimba—Reconstruction of a Montane Woodland Habitat that No Longer Occurs on the Island 133 Plate 13: Anjohibe I—Secrets of the Past Disclosed by Careful Study of Subfossil Bone and Pollen in a Cave 144 Plate 14: Anjohibe II—Inferences Based on Cave Remains and Aspects of the Organisms Living in the Adjacent Ecosystem 150 Plate 15: Anjajavy—a Trapdoor Cave, Ecology of an Extinct Lemur, and Untold Extinct and Extant Biodiversity 157 Plate 16: Ankarana I—Ecological Change of a Forest Community, a View from the Ground Up 162 Plate 17: Ankarana II—Ecological Change of a Forest Community, a Bird’s-Eye View from the Forest Canopy 168 Plate 18: Ankarana III—Tragedy and How the Bone Remains of an Extinct Lemur Can Help Piece Together How It Lived and the Former Local Forest Ecology 173 Species Plates Plate 19: Cryptoprocta spelea—an Extinct Mega- Predator and Aspects of How It May Have Lived and Hunted 177 Plate 20: Stephanoaetus mahery—a Presumed Primate Specialist and Its Role in the Evolution of Behavioral Aspects of Living and Extinct Lemurs 182 references 187 index to malagasy place-names 201 index to scientific names 203 a color gallery of plates 1–20 follows p. 132. Foreword Madagascar, like Africa, is not a place for the faint- what sort of stuff to watch out for around there, and hearted. If you are prone to complain a lot, you will hiring local camp help and guides—I asked the chief fi nd lots to complain about in this challenging land, if any other vazaha had come out to see his cave in with its climate extremes, bad roads, complex bu- recent years. His answer was similar to one I have reaucracy, and, for a foreigner, just plain strange- heard in other remote villages: “Yes, there was one a ness. It is thus no surprise that, again like Africa, the few years ago—he looked like Jesus.” Well, it would kind of vazaha (foreigners) who take to Madagascar not be much hyperbole to say that if nature in Mada- tend to be extraordinary people. Steve Goodman and gascar has a savior, Steve is probably it. Bill Jungers are two of the fi rst examples to come to But I have never really worked much with Steve, mind, and their remarkable resolve, thoroughness, except on a few papers about interesting fossils I have and persistence is in evidence on every page of this found that he took an interest in, and contributions book. These are not timid people, and I have never to a couple of his excellent edited volumes. By the had a boring moment with either of them. time he was hitting full stride there, I was looking at Steve was somebody whom I did not actually run places with interesting similarities and diff erences into in Madagascar until well into my many years elsewhere around the world. I sincerely wish we there. I was down in one of the most arid parts of could have worked together more, and, in fact, we are southwest Madagascar, by that remarkable hyper- making plans for an upcoming project in one of the saline playa lake with the unpronounceable name: Tsi- most remote places in western Madagascar, where I manampetsotsa (for understandable reasons, it often really look forward to getting to know this “unforget- turns up in the literature with alternate spellings). table character” better. I admit to having been a little skeptical at fi rst. My recollections of early Steve Goodman are scant Here was this scrawny, hirsute white guy, like my- and perhaps tattered by too many hard decades in self, but who also had, like me, plenty of training remote places and confused by too many other sto- for Madagascar in Africa. He was traveling alone, ries not yet written down. Bill Jungers, on the other checking out sites for what would eventually be a hand, I know a lot better; I count him among my best uniquely stellar career on this island nation. Steve friends, and on refl ection he has probably saved my was a neophyte on the island and lacked the refl ex life indirectly at least once or twice, simply by being that comes through experience, but he had a certain there with his always casual and friendly demeanor— something—a determination and quickness of wit— and his imposing size. Tall and stocky, Billi-be, as that is essential for success en brousse in a place like Malagasy often call him (“Big Bill”), is a true giant of a Madagascar. I came away from our fi rst encounter, man in every way. Like Steve, he has a CV full of envi- after hearing about his optimistic agenda, thinking: able accomplishments that just go on and on, decade This guy will go far in Madagascar—or die trying. after decade. He is also truly full of what we out here Now, a quarter century later, Steve Goodman is the in Hawai‘i call “Aloha,” a kind of radiant positive at- name on the lips of more Malagasy people out in ev- titude that overpowers the negative forces so often ery remote place in Madagascar than any other vazaha arrayed on all sides in our kind of complicated en- who has anything to do with nature. In this juncture, deavors. He is at once both a dead-serious academic I remember visiting one particularly remote cave in administrator and a top-notch scientist in his several Madagascar. At the nearest village, while doing my chosen fi elds. At the same time, he is more fun to be visite de courtoisie—showing my papers, fi nding out around than anybody of his academic stature whom I ix